Abstract: In several lek-breeding populations of birds and mammals, females arriving on leks tend to join males that already have females in their territories. This might occur either because females have an evolved preference for mating with males that are attractive to other females, or because they join groups of other females to obtain greater safety from predation or dangerous harassment by males. We have previously used controlled experiments to show that oestrous fallow deer females join males with established harems because they are attracted to female groups rather than to the males themselves. Here we demonstrate that the preference for males with females over males without females is specific to oestrous females and weak or absent in anoestrous ones, and that it is not associated with a preference for mating with males that have previously been seen to mate with other females. Furthermore, oestrous females given the choice between males that do not already have females with them show no significant preference for antlered over deantlered males or for older males over younger ones. We conclude that female attraction to other females on the lek is likely to be an adaptation to avoiding harassment in mixed-sex herds. In this situation, a male's ability to maintain the cohesion of his harem may be the principal cause of variation in mating success between males.

Abstract: In a number of mammalian species, males appear to have superior spatial abilities to females. The favoured explanations for this cognitive difference are hormonal, with higher testosterone levels in males than females leading to better spatial performance, and evolutionary, where sexual selection has favoured males with increased spatial abilities for either better navigational skills in hunting or to enable an increased territory size. However, an alternative explanation for this sex difference focuses on the role of varying levels of oestrogen in females in spatial cognition (the 'fertility and parental care' hypothesis). One possibility is that varying oestrogen levels result in variation in spatial learning and memory so that, when tested across the oestrous cycle, females perform as well as males on days of low oestrogen but more poorly on days of high oestrogen. If day in the oestrous cycle is not taken into account then, across an experiment, any sex differences found would always produce male superiority. We used a spatial working memory task in a Morris water maze to test the spatial learning and memory abilities of male and female rats. The rats were tested across a number of consecutive days during which the females went through four oestrous cycles. We found no overall sex differences in latencies to reach a submerged platform in a Morris water maze but, on the day of oestrus (low oestrogen), females took an extra swim to learn the platform's location (a 100% increase over the other days in the cycle). Female swim speed also varied across the oestrous cycle but females were no less active on the day of oestrus. These results oppose the predictions of the fertility and parental care hypothesis.

Abstract: Animal communication generally occurs in the environment of a network of several potential signallers and receivers. Within a network environment, it is possible to gain relative information about conspecifics by eavesdropping on signalling interactions. We presented male great tits with the opportunity to gain such information by simulating singing interactions using two loudspeakers. Interactions were presented so that relevant information was not available in the absolute singing behaviour of either individual, only in the relative timing of their songs in the interaction as a whole. We then assayed the information extracted by focal males by subsequently introducing one of the 'interactants' (i.e. loudspeakers) into the territory of the focal male. Focal males responded with a reduced song output to males that had just 'lost' an interaction. Focal males did not respond significantly differently to 'winners' as compared with intruders recently involved in an interaction that contained no consistent information. Focal males also responded by switching song types more often when encountering males that had recently been involved in a low-intensity interaction. These results provide the clearest evidence yet that male songbirds extract information from signal interactions between conspecifics in the field.

Abstract: Animals frequently use signals that travel further than the spacing between individuals. For every intended recipient of a given signal there are likely to be many other individuals that receive information. Eavesdropping on signalling interactions between other individuals provides a relatively cost-free method of assessing future opponents or mates. Male great tits (Parus major) extract relative information from such interactions between individuals unknown to them. Here, we show that male great tits can take information gathering a stage further and obtain more information about a previously unencountered intruder, by the hitherto unknown capability of combining information gathered by eavesdropping with that derived from their own direct interaction with an individual. Prior experience with an intruder (A) was achieved by subjecting a focal male to different levels of intrusion simulated using interactive playback. This intruder (A) then took part in a simulated interaction with an unknown male (B) outside the territorial boundary of the focal males. In response to subsequent intrusion by the second male (B), focal males showed low song output in response to males that had lost to a male that the subject was able to beat. Males of known high quality, or those about which information was ambiguous, elicited a high level of song output by focal males. We discuss the implications of this finding for the evolution of communication and social behaviour.

Abstract: Behaviour under conditions of mild stress shows consistent patterns in all vertebrates: exploratory behaviour, boldness, aggressiveness covary in the same way. The existence of highly consistent individual variation in these behavioural strategies, also referred to as personalities or coping styles, allows us to measure the behaviour under standardized conditions on birds bred in captivity, link the standardized measurements to the behaviour under natural conditions and measure natural selection in the field. We have bred the great tit (Parus major), a classical model species for the study of behaviour under natural conditions, in captivity. Here, we report a realized heritability of 54 +/- 5% for early exploratory behaviour, based on four generations of bi-directional artificial selection. In addition to this, we measured hand-reared juveniles and their wild-caught parents in the laboratory. The heritability found in the mid-offspring-mid-parent regression was significantly different from zero. We have thus established the presence of considerable amounts of genetic variation for personality types in a wild bird.