Abstract: Living in complex social systems requires perceptual and cognitive capacities for the recognition of group membership and individual competitors. Olfaction is one means by which this can be achieved. Many animals can identify individual proteins in urine, skin secretions, or saliva by scent. Additionally, marking behaviour in several mammals and especially in horses indicates the importance of sniffing conspecifics’ faeces for olfactory recognition. To test this hypothesis, we conducted two separate experiments: Experiment 1 addressed the question of whether horses can recognise the group membership of other horses by sniffing their faeces. The horses were presented with four faecal samples: (1) their own, (2) those of other members of their own group, (3) those of unfamiliar mares, and (4) those of unfamiliar geldings. Experiment two was designed to assess whether horses can identify the group member from whom a faecal sample came. Here, we presented two groups of horses with faecal samples from their group mates in random distribution. As controls, soil heaps and sheep faecal samples were used. In experiment one, horses distinguished their own from their conspecifics’ faeces, but did not differentiate between familiarity and sex. In experiment two, the horses from both groups paid most attention to the faeces of the horses from which they received the highest amount of aggressive behaviours. We therefore suggest that horses of both sexes can distinguish individual competitors among their group mates by the smell of their faeces.

Abstract: Differentiating between individuals with different knowledge states is an important step in child development and has been considered as a hallmark in human evolution. Recently, primates and corvids have been reported to pass knower–guesser tasks, raising the possibility of mental attribution skills in non-human animals. Yet, it has been difficult to distinguish ‘mind-reading’ from behaviour-reading alternatives, specifically the use of behavioural cues and/or the application of associatively learned rules. Here, I show that ravens (Corvus corax) observing an experimenter hiding food are capable of predicting the behaviour of bystanders that had been visible at both, none or just one of two caching events. Manipulating the competitors' visual field independently of the view of the test-subject resulted in an instant drop in performance, whereas controls for behavioural cues had no such effect. These findings indicate that ravens not only remember whom they have seen at caching but also take into account that the other's view was blocked. Notably, it does not suffice for the birds to associate specific competitors with specific caches. These results support the idea that certain socio-ecological conditions may select for similar cognitive abilities in distantly related species and that some birds have evolved analogous precursors to a human theory-of-mind.

Abstract: Horses are likely to exhibit aggression when meeting for the first time. Therefore, this study compared 3 methods for mixing horses to evaluate their effectiveness in reducing aggressive interactions: (1) mixing pairs of horses in a paddock (P, 10 minutes, 15 tests), (2) introducing 1 unfamiliar horse to a pair of familiar, resident horses in a paddock (PP, 10 minutes, 15 tests), (3) allowing limited physical contact between pairs of horses for a short period of pre-exposure in neighboring boxes (B, 5 minutes, 16 tests) before mixing them in a paddock (BP, 10 minutes 16 tests). A total of 16 Swedish Standardbred mares, aged 6-18 years (mean age ± SD: 11 ± 4.4), were included in the study. Half of the horses were familiar with each other (resident horses, n = 8), whereas the other half were bought in from a variety of sources (unfamiliar horses, n = 8). Social interactions, consisting of behaviors from the sender, the receiver, and the subsequent sender's response, were recorded continuously as frequencies. There were no differences in the frequencies of aggressive behaviors between the 3 mixing methods, including those aggressive behaviors in which physical contact had been attempted (kick, strike). Although resident horses were overall more aggressive (median number of aggressive behaviors per horse, 62; Q1, 36; Q3, 68.5) than unfamiliar horses (median per horse, 4; Q1, 2; Q3, 12.5) during all tests (U = 97, P = 0.003), none of the 62 tests needed to be terminated. Unfamiliar horses did not receive more aggression from resident horses in PP (mean per test ± SD: 5.1 ± 3.1) than in P (mean per test ± SD: 6.4 ± 4.9) (t = 0.63, P = 0.544). However, the behavior “attack” was more frequent in PP (median per test, 2; Q1, 0; Q3, 5) than in P (median per test, 0; Q1, 0; Q3, 1) (U = 282, P = 0.042), and “flee” was more frequent in PP (median per test, 6; Q1, 4; Q3, 8) than in P (median per test, 1; Q1, 0; Q3, 6) (U = 290, P = 0.018). Pre-exposure in boxes did not reduce aggression in BP (median per test, 7; Q1, 4.3; Q3, 11.8) as compared with P (median per test, 6; Q1, 2; Q3, 16) (U = 264, P = 0.767), but during pre-exposure in B tests, horses exchanged more nonaggressive (median per test, 2; Q1, 0.3; Q3, 4) than aggressive (median frequency of aggressive behavior, 0; Q1, 0; Q3, 1) (W = 71, P = 0.013) and mixed interactions (median per test, 0; Q1, 0; Q3, 1) (W = 92, P = 0.016) through the opening. Results suggest mixing an unfamiliar horse with 2 resident horses at the same time instead of one by one may be preferable. In this way, the total aggression received by the unfamiliar horse will potentially be less, even though aggressive interactions may be more intense.

Abstract: In animals, behavioural properties such as aggressive propensity are often consistent over a life span, and they may form part of a behavioural syndrome. We studied how aggressive propensity influences dominance, contest behaviour and growth in the cooperatively breeding cichlid fish Neolamprologus pulcher. We tested whether intrinsic aggressive propensity (1) influences dominance in paired contests, (2) causes different aggression levels in contests with partners matched for aggressive propensity compared to unmatched partners, and how it (3) affects growth rate in groups that were either matched or unmatched for aggressive propensity. Intrinsic aggressive propensity was first scored with a mirror test and classified as high, medium or low. Thereafter we tested fish with either high or low aggressive propensity with partners matched for size and either matched or unmatched for aggressive type in a paired contest for a shelter. We scored dominance, aggression and submission. As predicted, (1) dominance was more clearly established in unmatched than in matched contests and (2) individuals with high aggressive propensity launched more attacks overall than fish with low intrinsic aggressiveness, suggesting a higher propensity to escalate independently of winning or losing the paired contest. However, contrary to expectation, (3) individuals with low aggressiveness grew faster than aggressive ones in unmatched groups, whereas the opposite occurred in matched groups. This suggests that individuals with low aggressive propensity may benefit from conflict evasion, which might allow them to gain dominance in the future owing to larger body size.

Abstract: More than half a century has now elapsed since coalition or alliance formation theory (CAFT) was first developed. During that time, researchers have amassed a vast amount of detailed and high-quality data on coalitions or alliances among primates and other animals. But models have not kept pace, and more relevant theory is needed. In particular, even though CAFT is primarily an exercise in polyadic game theory, game theorists have devoted relatively little attention to questions that motivate field research, and much remains largely unexplored. The state of the art is both a challenge and an opportunity. In this review we describe a variety of game-theoretic and related modelling approaches that have much untapped potential to address the questions that field biologists ask.