Abstract: ABSTRACT Reciprocity is probably the most debated of the evolutionary explanations for cooperation. Part of the confusion surrounding this debate stems from a failure to note that two different processes can result in reciprocity: partner control and partner choice. We suggest that the common observation that group-living animals direct their cooperative behaviours preferentially to those individuals from which they receive most cooperation is to be interpreted as the result of the sum of the two separate processes of partner control and partner choice. We review evidence that partner choice is the prevalent process in primates and propose explanations for this pattern. We make predictions that highlight the need for studies that separate the effects of partner control and partner choice in a broader variety of group-living taxa.

Abstract: How animals use space has important consequences for feeding ecology, social organization, mating strategies and conservation management. In white rhinoceros, female home ranges are much larger than male territories, suggesting that movement patterns are influenced by factors other than resource distribution. In this study we placed radiotransmitters on 15 female white rhinoceros, recording 1758 locations and collecting behavioural data during 1671 observation sessions, making this the largest data set of its kind in this species. We investigated how habitat variables and male territories influenced female movement and reproductive behaviour. Female home ranges were approximately 20 km2 and core areas were 5 km2, with male territories roughly the same size as female core areas. Female range size did not vary with season, but the pattern of space use did vary. Females used grassland habitat preferentially, utilizing these areas significantly more than expected based on availability. Findings relevant to the mating strategy include: (1) the amount of grassland in a male's territory predicted female use of the territory; (2) the time that a female spent in a male's territory was a significant predictor of reproductive activity with the male, indicating that females probably mate with the most familiar male; and (3) the temporal pattern of female space use suggests that females did not increase mate sampling behaviour nor did they become more choosy about which males they visited when reproductively active. These findings suggest that males may maximize reproductive success by defending areas containing more grassland habitat.

Abstract: The function of olfactory signalling in social species is less well understood than in asocial species. Consequently, we examined olfactory communication in the ringtailed lemur, a socially complex primate that retains a functional vomeronasal organ, has well-developed scent glands and shows a suite of scent-marking behaviour. To assess the information content of different types of scent gland secretions, we decoupled olfactory cues from the visual and behavioural modalities with which scent marking is normally associated. We presented male and female subjects (signal receivers) with a series of choice tests between odours derived from conspecific donors (signal senders) varying by sex, age, social status and reproductive condition. We additionally examined the influence of the receivers' reproductive state and familiarity with the signaller. The reproductive condition, social status and familiarity of senders and receivers affected signal transmission; specifically, male receivers attended most to the odours of conspecifics in breeding condition and to the odours of familiar, dominant animals. By contrast, females varied their responses according to both their own reproductive state and that of the sender. Based on male and female patterns of countermarking, we suggest that scent marking serves a function in intergroup spacing and intrasexual competition for both sexes, as might be expected in a female-dominant species. By contrast, minimal female interest in male odours counters a female mate choice function for scent marking in this species. Nevertheless, scent marks are critical to male-male competition and, therefore, may be subject to sexual selection.

Abstract: The responsiveness of 10 horses and 10 ponies to environmental challenge (represented by an open field test) was assessed using a qualitative approach based on free choice profiling methodology (FCP), which gives observers complete freedom to choose their own descriptive terms. Data were analysed with generalised Procrustes analysis (GPA), a multivariate statistical technique associated with FCP. A cross-validation of the outcomes of this approach to data recorded through quantitative behaviour analysis, and through a questionnaire given to the animals' owner/riding instructor, was also performed using principal component analysis (PCA). Twelve undergraduate students generated their own descriptive vocabularies, by watching 20 horse/pony video clips lasting 2.5 min each. GPA showed that the consensus profile explained a high percentage of variation among the 12 observers, and differed significantly from the mean randomised profile (p < 0.001). Two main dimensions of the consensus profile were identified, explaining 60% and 5.2% of the variation between animals, respectively. The 12 observer word charts interpreting these dimensions were semantically consistent, as they all converged towards the same meaning, albeit using different terms. The most used term to describe the positive end of axis 1 was “quiet”, whereas “attentive” was the best positive descriptor of axis 2. The most frequently used descriptors for the negative ends of axes 1 and 2 were “nervous” and “bored”, respectively. Thus, axis 1 was labelled as “quiet/nervous” and axis 2 was named as “attentive/bored”. A marked effect of animal category was observed on the scores of the animals on the first dimension (p < 0.001). Horses received significantly higher scores, and were thus assessed as more quiet and calm, than ponies. Conversely, ponies tended to receive lower scores on the second dimension (p < 0.12), therefore they appeared less curious and attentive. The results of the PCA showed that the variables from different types of measurement clearly had meaningful relationships. For instance, the variables with the highest loading on the positive end of axis 1 were all indicative of tractable and docile animals, whereas axis 2 showed high loadings on the positive end for variables indicating attentive animals. Qualitative behaviour assessment proved to be an appropriate methodology for the study of horse behavioural responsiveness, in that it provided a multifaceted characterisation of horse behavioural expression that was in agreement with other quantitative and subjective assessments of the animals' behaviour.

Abstract: A group of animals can only move cohesively, if group members “somehow” reach a consensus about the timing (e.g., start) and the spatial direction/destination of the collective movement. Timing and spatial decisions usually differ with respect to the continuity of their cost/benefit distribution in such a way that, in principle, compromises are much more feasible in timing decision (e.g. median preferred time) than they are in spatial decisions. The consequence is that consensus costs connected to collective timing decisions are usually less skewed amongst group members than are consensus costs connected to spatial decisions. This, in turn, influences the evolution of decision sharing: sharing in timing decisions is most likely to evolve when conflicts are high relative to group cohesion benefits, while sharing in spatial decisions is most likely to evolve in the opposite situation. We discuss the implications of these differences for the study of collective movement decisions.