Abstract: Individual recognition has been attributed a crucial role in the evolution of complex social systems such as helping behaviour and cooperation. A classical example for interspecific cooperation is the mutualism between the cleaner fish Labroides dimidiatus and its client reef fish species. For stable cooperation to evolve, it is generally assumed that partners interact repeatedly and remember each other's past behaviour. Repeated interactions may be achieved by site fidelity or individual recognition. However, as some cleaner fish have more than 2,300 interactions per day with various individuals per species and various species of clients, basic assumptions of cooperation theory might be violated in this mutualism. We tested the cleaner L. dimidiatus and its herbivorous client, the surgeon fish Ctenochaetus striatus, for their ability to distinguish between a familiar and an unfamiliar partner in a choice experiment. Under natural conditions, cleaners and clients have to build up their relationship, which is probably costly for both. We therefore predicted that both clients and cleaners should prefer the familiar partner in our choice experiment. We found that cleaners spent significantly more time near the familiar than the unfamiliar clients in the first 2 minutes of the experiment. This indicates the ability for individual recognition in cleaners. In contrast, the client C. striatus showed no significant preference. This could be due to a sampling artefact, possibly due to a lack of sufficient motivation. Alternatively, clients may not need to recognise their cleaners but instead remember the defined territories of L. dimidiatus to achieve repeated interactions with the same individual.

Abstract: The ability to reliably detect what others are attending to seems important for social species to interact with their partners. Domestic horses (Equus caballus) have lived with humans for over five thousand years, hence they might have developed sensitivity to human attention. In the present study, we investigated whether horses would discriminate the situation in which a human experimenter could see them from the situation in which she could not. Specifically, we tested whether horses understand the role of eyes in human attentional states, produce more visual gestures when the experimenter can see their begging behaviors and produce more auditory or tactile gestures when she can not. We used with a slight modification the paradigm that previously yielded support for chimpanzee understanding of human attention (Hostetter et al. 2007). Twelve horses were offered food by the experimenter who showed various attentional states in front of them. We scored frequency of begging behaviors by the horses. In experiment 1, we set three kinds of condition: hand over the eyes, hand over the mouth and away. In the last condition there was only a food in front of horses, which was a control condition. The results showed that horses produced more auditory or tactile begging behaviors when the experimenter“s eyes were not visible than when her eyes were visible, but there was no difference in visual begging behaviors. In experiment 2, we set two kinds of condition: eyes closed and eyes open. The horses also produced more auditory or tactile begging behaviors when the experimenter”s eyes were closed than when they were open. However, there was no difference in visual begging behaviors. These results show that horses discriminate the situation in which humans can see from that in which humans can not. Of special interest, horses increased only auditory or tactile behaviors, not all types of communicative behaviors, when the experimenter could not see their begging behaviors. This result suggests that horses are sensitive to human attentional states. Moreover, horses may do recognize the eyes as an important indicator of whether or not humans will respond to their behavior and they may be able to behave flexibly depending upon human attentional states

Abstract: An altruistic individual has to gamble on cooperation to a stranger because it does not know whether the stranger is trustworthy before direct interaction. Nowak and Sigmund (Nature 393 (1998a) 573; J. Theor. Biol. 194 (1998b) 561) presented a new theoretical framework of indirect reciprocal altruism by image scoring game where all individuals are informed about a partner's behavior from its image score without direct interaction. Interestingly, in a simplified version of the image scoring game, the evolutionarily stability condition for altruism became a similar form of Hamilton's rule, i.e. inequality that the probability of getting correct information is more than the ratio of cost to benefit. Since the Hamilton's rule was derived by evolutionarily stable analysis, the evolutionary meaning of the probability of getting correct information has not been clearly examined in terms of kin and group selection. In this study, we applied covariance analysis to the two-score model for deriving the Hamilton's rule. We confirmed that the probability of getting correct information was proportional to the bias of altruistic interactions caused by using information about a partner's image score. The Hamilton's rule was dependent on the number of game bouts even though the information reduced the risk of cooperation to selfish one at the first encounter. In addition, we incorporated group structure to the two-score model to examine whether the probability of getting correct information affect selection for altruism by group selection. We calculated a Hamilton's rule of group selection by contextual analysis. Group selection is very effective when either the probability of getting correct information or that of future interaction, or both are low. The two Hamilton's rules derived by covariance and contextual analyses demonstrated the effects of information and group structure on the evolution of altruism. We inferred that information about a partner's behavior and group structure can produce flexible pathways for the evolution of altruism.

Abstract: For species in which group membership frequently changes, it has been a challenge to characterize variation in individual interactions and social structure. Quantifying this variation is necessary to test hypotheses about ecological determinants of social patterns and to make predictions about how group dynamics affect the development of cooperative relationships and transmission processes. Network models have recently become popular for analyzing individual contacts within a population context. We use network metrics to compare populations of Grevy's zebra (Equus grevyi) and onagers (Equus hemionus khur). These closely related equids, previously described as having the same social system, inhabit environments differing in the distribution of food, water, and predators. Grevy's zebra and onagers are one example of many sets of coarsely similar fission-fusion species and populations, observed elsewhere in other ungulates, primates, and cetaceans. Our analysis of the population association networks reveals contrasts consistent with their distinctive environments. Grevy's zebra individuals are more selective in their association choices. Grevy's zebra form stable cliques, while onager associations are more fluid. We find evidence that females associate assortatively by reproductive state in Grevy's zebra but not in onagers. The current approach demonstrates the utility of network metrics for identifying fine-grained variation among individuals and populations in association patterns. From our analysis, we can make testable predictions about behavioral mechanisms underlying social structure and its effects on transmission processes.