Both sheep and goats can solve inferential by exclusion tasks.

Despite the domestication of sheep and goats by humans for several millennia, we still lack comparative data on their cognitive capacity. Comparing the cognitive skills of farm animals can help understand the evolution of cognition. In this study, we compared the performances of sheep and goats in inference by exclusion tasks. We implemented two tasks, namely a cup task and a tube task, to identify whether success in solving the task could be attributed to either low-level mechanisms (avoiding the empty location strategy) or to deductive reasoning (if two possibilities A and B, but not A, then it must be B). In contrast to a previous study comparing goats and sheep in a cup task, we showed that both species solved the inferential condition with high success rates. In the tube task, performances could not be explained by alternative strategies such as avoiding the empty tube or preferring the bent tube. When applying a strict set of criteria concerning responses in all conditions and controlling for the potential effects of experience, we demonstrate that two individuals, a goat and a sheep, fulfil these criteria. This suggests that sheep and goats are able to make inferences based on deductive reasoning.

How does the expressiveness of leaders affect followership in domestic horses (Equus ferus caballus)?

In collective movements, some individuals are more effective and attractive leaders than others. Parameters such as social network, personality, and physiologic needs failed to explain why group members follow one leader more than another. In this study in the domestic horse, we propose to focus on the leader's attitude and its impact to the followers' recruitment during two conditions: spontaneous group departures or experimentally induced departures. We postulate that the expressiveness of the leader could enhance its attractiveness and thus produce a successful followership. We found that a high expressiveness level is associated with a high curiosity score and with a low social status. This propensity to initiate while being expressive was higher in the experimental condition than in the spontaneous condition which was expected, since the experimental procedure sought to increase the motivation of initiators with a highly appetent reward. Moreover, the more intense the leader's expressiveness is, the faster the followers will join the movement regardless of the condition. This positive and dynamic attitude can be perceived as an attractive feature by the followers. Finally, we evidenced collective arousal in followers initiated by the leader that could be essential for group cohesion. The present study is the first to investigate the expressiveness level in animals, and provides new insights into the emergence of leadership.

The influence of phylogeny, social style, and sociodemographic factors on macaque social network structure.

Among nonhuman primates, the evolutionary underpinnings of variation in social structure remain debated, with both ancestral relationships and adaptation to current conditions hypothesized to play determining roles. Here we assess whether interspecific variation in higher-order aspects of female macaque (genus: Macaca) dominance and grooming social structure show phylogenetic signals, that is, greater similarity among more closely-related species. We use a social network approach to describe higher-order characteristics of social structure, based on both direct interactions and secondary pathways that connect group members. We also ask whether network traits covary with each other, with species-typical social style grades, and/or with sociodemographic characteristics, specifically group size, sex-ratio, and current living condition (captive vs. free-living). We assembled 34-38 datasets of female-female dyadic aggression and allogrooming among captive and free-living macaques representing 10 species. We calculated dominance (transitivity, certainty), and grooming (centrality coefficient, Newman's modularity, clustering coefficient) network traits as aspects of social structure. Computations of K statistics and randomization tests on multiple phylogenies revealed moderate-strong phylogenetic signals in dominance traits, but moderate-weak signals in grooming traits. GLMMs showed that grooming traits did not covary with dominance traits and/or social style grade. Rather, modularity and clustering coefficient, but not centrality coefficient, were strongly predicted by group size and current living condition. Specifically, larger groups showed more modular networks with sparsely-connected clusters than smaller groups. Further, this effect was independent of variation in living condition, and/or sampling effort. In summary, our results reveal that female dominance networks were more phylogenetically conserved across macaque species than grooming networks, which were more labile to sociodemographic factors. Such findings narrow down the processes that influence interspecific variation in two core aspects of macaque social structure. Future directions should include using phylogeographic approaches, and addressing challenges in examining the effects of socioecological factors on primate social structure.

Ecology of information: social transmission dynamics within groups of non-social insects.

While many studies focus on how animals use public information, the dynamics of information spread and maintenance within groups, i.e. the 'ecology of information', have received little attention. Here we use fruitflies trained to lay eggs on specific substrates to implement information into groups containing both trained and untrained individuals. We quantify inter-individual interactions and then measure the spread of oviposition preference with behavioural tests. Untrained individuals increase their interactive approaches in the presence of trained individuals, and the oviposition preference transmission is directly proportional to how much trained and untrained individuals interact. Unexpectedly, the preference of trained individuals to their trained oviposition substrate decreases after interactions with untrained individuals, leading to an overall informational loss. This shows that social learning alone is not enough to support informational stability.

Inferences about food location in three cercopithecine species: an insight into the socioecological cognition of primates.

Many animal species use a variety of cognitive strategies to locate food resources. One strategy is to make inferences by exclusion, i.e., perceiving the absence of reward as a cue that another location should be investigated. The use of such advanced cognitive strategies may be more prominent in species that are known to frequently solve social challenges, and inferential reasoning has mainly been investigated in social species such as corvids, dogs, dolphins and non-human primates. In this paper, we investigate how far social intricacy may explain the disparity of reasoning performances observed in three cercopithecine species that differ in the density of their social network and the diversity of their social partners. We used standard reasoning tasks, testing the volume concept and inference by exclusion using visual and auditory modalities. We showed that Old World monkeys can infer the location of invisible food by exclusion. In addition, Tonkean macaques and olive baboons had greater performances in most tasks compared to rhesus macaques. These responses are consistent with the social complexity displayed by these three species. We suggest that the cognitive strategies required to navigate through a demanding social world are involved in the understanding of the physical domain.

Less bins, less baboons: reducing access to anthropogenic food effectively decreases the urban foraging behavior of a troop of chacma baboons (Papio hamadryas ursinus) in a peri-urban area.

In South Africa, chacma baboons (Papio hamadryas ursinus) living near peri-urban areas may forage on anthropogenic food. Baboons have been recorded to damage crops, scatter waste from trash bins, and damage homes. A number of methods have been tested over the past 20 years to solve these problems, but none proved successful over the long-term or involved considerable costs. An efficient management system requires a detailed knowledge of how baboon troops proceed and organize during these urban foraging actions. This study examines the response of a troop of baboons to an experimental reduction of anthropogenic food sources in a peri-urban environment, the George campus of Nelson Mandela University (SA). We gradually suppressed access to waste food in trash cans, reducing the amount of anthropogenic food available. This change in food availability led baboons to modify their urban foraging strategy. They compensated for the lack of anthropogenic food by spending more time foraging on natural food and less time in urban areas. However, the troop still exploited waste-free areas during the experiment and even more when the conditions were normal again. Overall, these results show the ability of baboons to adapt to changes in anthropogenic food availability but also that they are highly dependent on this type of resource. Limiting its access is a mitigation strategy that humans must absolutely develop for reaching a high level of coexistence with baboons.

Does a high social status confer greater levels of trust from groupmates? An experimental study of leadership in domestic horses.

In collective movements, specific individuals may emerge as leaders. In this study on the domestic horse (Equus ferus caballus), we conducted experiments to establish if an individual is successfully followed due to its social status (including hierarchical rank and centrality). We first informed one horse about a hidden food location and recorded by how many it was followed when going back to this location. In this context, all horses lead their groupmates successfully. In a second step, we tested whether group members would trust some leaders more than others by removing the food before the informed individual led the group back to the food location. In addition, two control initiators with intermediate social status for which the food was not removed were tested. The results, confirmed by simulations, demonstrated that the proportions of followers for the unreliable initiator with highest social status are greater than the ones of the unreliable initiator with lowest social status. Our results suggest an existing relationship between having a high social status and a leadership role. Indeed, the status of a leader sometimes prevail at the detriment of the accuracy of the information, because an elevated social status apparently confers a high level of trust.

Group size, grooming and fission in primates: a modeling approach based on group structure.

In social animals, fission is a common mode of group proliferation and dispersion and may be affected by genetic or other social factors. Sociality implies preserving relationships between group members. An increase in group size and/or in competition for food within the group can result in decrease certain social interactions between members, and the group may split irreversibly as a consequence. One individual may try to maintain bonds with a maximum of group members in order to keep group cohesion, i.e. proximity and stable relationships. However, this strategy needs time and time is often limited. In addition, previous studies have shown that whatever the group size, an individual interacts only with certain grooming partners. There, we develop a computational model to assess how dynamics of group cohesion are related to group size and to the structure of grooming relationships. Groups' sizes after simulated fission are compared to observed sizes of 40 groups of primates. Results showed that the relationship between grooming time and group size is dependent on how each individual attributes grooming time to its social partners, i.e. grooming a few number of preferred partners or grooming equally or not all partners. The number of partners seemed to be more important for the group cohesion than the grooming time itself. This structural constraint has important consequences on group sociality, as it gives the possibility of competition for grooming partners, attraction for high-ranking individuals as found in primates' groups. It could, however, also have implications when considering the cognitive capacities of primates.

Differences in nutrient requirements imply a non-linear emergence of leaders in animal groups.

Collective decision making and especially leadership in groups are among the most studied topics in natural, social, and political sciences. Previous studies have shown that some individuals are more likely to be leaders because of their social power or the pertinent information they possess. One challenge for all group members, however, is to satisfy their needs. In many situations, we do not yet know how individuals within groups distribute leadership decisions between themselves in order to satisfy time-varying individual requirements. To gain insight into this problem, we build a dynamic model where group members have to satisfy different needs but are not aware of each other's needs. Data about needs of animals come from real data observed in macaques. Several studies showed that a collective movement may be initiated by a single individual. This individual may be the dominant one, the oldest one, but also the one having the highest physiological needs. In our model, the individual with the lowest reserve initiates movements and decides for all its conspecifics. This simple rule leads to a viable decision-making system where all individuals may lead the group at one moment and thus suit their requirements. However, a single individual becomes the leader in 38% to 95% of cases and the leadership is unequally (according to an exponential law) distributed according to the heterogeneity of needs in the group. The results showed that this non-linearity emerges when one group member reaches physiological requirements, mainly the nutrient ones - protein, energy and water depending on weight - superior to those of its conspecifics. This amplification may explain why some leaders could appear in animal groups without any despotism, complex signalling, or developed cognitive ability.

Social network modeling: a powerful tool for the study of group scale phenomena in primates.

Social Network Analysis is now a valuable tool to study social complexity in many animal species, including primates. However, this framework has rarely been used to implement quantitative data on the social structure of a group within computer models. Such approaches allow the investigation of how social organization constrains other traits and also how these traits can impact the social organization in return. In this commentary, we discuss the powerful potential of social network modeling as a way to study group scale phenomena in primates. We describe the advantages of using such a method and we focus on the specificity of this approach in primates, given the particularities of their social networks compared with those of other taxa. We also give practical considerations and a list of examples as for the choice of parameters that can be used to implement the social layer within the models.

How can social network analysis improve the study of primate behavior?

When living in a group, individuals have to make trade-offs, and compromise, in order to balance the advantages and disadvantages of group life. Strategies that enable individuals to achieve this typically affect inter-individual interactions resulting in nonrandom associations. Studying the patterns of this assortativity using social network analyses can allow us to explore how individual behavior influences what happens at the group, or population level. Understanding the consequences of these interactions at multiple scales may allow us to better understand the fitness implications for individuals. Social network analyses offer the tools to achieve this. This special issue aims to highlight the benefits of social network analysis for the study of primate behaviour, assessing it's suitability for analyzing individual social characteristics as well as group/population patterns. In this introduction to the special issue, we first introduce social network theory, then demonstrate with examples how social networks can influence individual and collective behaviors, and finally conclude with some outstanding questions for future primatological research.

Measurement of Endurance of Untrained Five-Year-Old and Six-Year-Old Horses Raised in Multi-age Herds on Pasture.

The objective of this study was to explore how the endurance of five-year-old horses, kept out on pasture all their life and ridden for the first time well into their fifth year of age, developed within one year and compared to that of six-year-old horses raised under the same conditions and to other horses. Horses were submitted to a standardized exercise test (SET) to calculate their v4 (velocity run under defined conditions inducing 4 mmol/L of blood lactate concentration (LA)) and v180 (velocity run under defined conditions inducing a heart rate of 180 beats/min). The test consisted of up to five consecutive intervals at increasing speed until the blood LA of a horse increased above 4 mmol/L. The blood LA measured after each interval was plotted exponentially against running speed to derive v4 from the blood lactate-running speed relationship, and the mean heart rate during the intervals was plotted linearly against running speed to derive v180 from the heart rate-running speed relationship. The following were examined: (1) the development of v4 and v180 of five-year-old horses within one year through measurements in September and in the following July and September; the comparison of endurance variables (2) between five-year-olds and six-year-olds; (3) between six-year-olds on consecutive years; and 4) between six-year-olds and foreign horses. The results showed that: (1) there were no changes of either variable within one year (repeated measures ANOVA P > .05); (2) there were no significant differences between five-year-olds and six-year-olds (one-way ANOVA P > .05); (3) no significant differences between six-year-old groups (one-way ANOVA; P > 05); and (4) foreign horses had higher v4 and v180 values than six-year-olds (one-way ANOVA; P = .0001 and P = .003, respectively). There was no significant relationship between v4 and v180 (P > .05; r2 = 0.02). In conclusion, one additional year on pasture in multiage herds did not increase the endurance variables of five-year-old horses. Thus, the endurance appeared to be consolidated in these horses at the age of five years, and additional training seems to be necessary to increase it.

Signals use by leaders in Macaca tonkeana and Macaca mulatta: group-mate recruitment and behaviour monitoring.

Animals living in groups have to make consensus decisions and communicate with each other about the time, or the direction, in which to move. In some species, the process relies on the proposition of a single individual, i.e. a first individual suggests a movement and the other group members decide whether or not to join this individual. In Tonkean (Macaca tonkeana) and rhesus macaques (Macaca mulatta), it has been observed that this first individual displays specific signals at departure. In this paper, we aimed to explore the function of such behaviours, i.e. if these behaviours were recruitment signals or only cues about the motivation of the first departed individual. We carried out temporal analyses and studied the latencies of the first departed individual's behaviours and of the joining of other group members. We also assessed whether the social style of a species in terms of dominance and kinship relationships influenced the patterns of signal emissions. We then analyzed how the first departed individual decided to make a pause or to stop it according to the identities of group members that joined the collective movement. Results showed that Tonkean macaques and rhesus macaques seemed to use back-glances to monitor the joining of other group members and pauses to recruit such individuals. This was especially the case for highly socially affiliated individuals in Tonkean macaques and kin-related individuals in rhesus macaques. Moreover, back-glances and pauses disappeared when such individuals joined the first departed individual. From these results, we suggested that such behaviour could be considered intentional. Such findings could not be highlighted without temporal analyses and accurate observations on primate groups in semi-free ranging conditions.

Hamsters in the city: A study on the behaviour of a population of common hamsters (Cricetus cricetus) in urban environment.

Animals in urban environments face challenging situations and have to cope with human activities. This study investigated the ecology and behaviour of a population of European hamsters (Cricetus cricetus) living in the city centre of Vienna (Austria). We recorded the surface activities of 35 hamsters in May 2015. Each focal animal was observed for 15 minutes, and a total of 66 focal samples were analysable. As a prey species in an environment teeming with human activities, we predicted a high level of vigilance by the hamsters. The results show that while animals dedicated a lot of time to vigilance, most of their time was spent foraging. The study also explores whether the frequency of vigilance behaviours differ between males and females. We found that vigilance behaviours were expressed in a different manner by males and females. Finally, we investigated the distribution of the burrows on green spaces depending on proximity to trees and on noise levels. We found a biased distribution of burrows, with a spatial preference for location protected by the vegetation and distant to noise sources. Although burrows were located preferentially under vegetation cover, levels of noise did not determine their positions. Moreover, this species does not respond to disturbances like daily urban noises, probably due to habituation. The common hamster is an endangered species; our results lead to a greater knowledge of its behaviour in a persistent urban population.

Captive-reared European hamsters follow an offensive strategy during risk-assessment.

Understanding whether captive-reared animals destined to reintroduction are still able to discriminate predators has important implications for conservation biology. The endangered European hamster benefits from conservation programs throughout Europe, in which several thousand individuals are released into the wild every year. Despite this, the anti-predator strategy of hamsters and their ability to maintain predator discrimination in captivity remain to be investigated. Here, we explore the predator discrimination behaviour of captive-reared European hamsters and their response to different predation cues. When first exposed to the urine of cats and goats in a Y-maze test, hamsters spent more time close to the cat scent rather than to the goat scent. In a second experiment, during which hamsters were exposed to a non-mobile European ferret (inside a cage), hamsters significantly increased the time spent close to the ferret's cage and displayed aggressive behaviour towards the ferret. Furthermore, they did not take refuge inside an anti-predation tube (APT), a device designed to upgrade wildlife underpasses and reconnect wild hamster populations. Finally, when exposed to a mobile ferret (but without physical contact), hamsters displayed mobbing and aggressive behaviours towards the ferret, before taking refuge inside the APT. Taken together, our results show that captive-reared hamsters are still able to detect and react to predation cues, but that they initially adopt an offensive strategy (grunting, spitting, mobbing) during the risk-assessment phase. After risk assessment, however, hamsters used the APT as a refuge. Our study provides important insights into the anti-predator behaviour of hamsters. Testing the efficacy of the APT, a device that will allow upgrading wildlife underpasses for the hamster and other rodents, is also of great importance and is instrumental in conservation efforts for these species.

How many for dinner? Recruitment and monitoring by glances in capuchins.

Group members present physical and physiological differences according to their age, sex or social status, which could generate motivation differences among individuals during travel. In spite of these divergences of interest among individuals, the group succeeds more often than not in making a collective decision about departure time and which direction to take. To reach a consensus decision, animals should exchange information relating to characteristics of group movement through different communication channels. The main purpose of this study is to understand the function of behaviour patterns displayed during movements of white-faced capuchins (Cebus capucinus). We designed experiments in which we provoked collective movements involving a binary choice. During experiments, a video camera recorded the behaviour of each capuchin, which enabled us to determine which individuals displayed a behavioural pattern during travel and how this behaviour influenced the other group members. We found that looking backwards seemed to permit the recruitment of group mates during collective movement. This behaviour also seemed to allow the quantification of the number of followers, since the emitter modified its locomotion speed according to this number. In this preliminary study, we showed that visual behaviour was used to recruit and monitor group mates during collective movements and provided information on mechanisms involved in maintaining cohesion and coordination among group members during travel.

Connecting paths between juvenile and adult habitats in the Atlantic green turtle using genetics and satellite tracking.

Although it is commonly assumed that female sea turtles always return to the beach they hatched, the pathways they use during the years preceding their first reproduction and their natal origins are most often unknown, as it is the case for juvenile green turtles found in Martinique waters in the Caribbean. Given the oceanic circulation of the Guiana current flowing toward Martinique and the presence of important nesting sites for this species in Suriname and French Guiana, we may assume that a large proportion of the juvenile green turtles found in Martinique are originating from the Suriname-French Guiana beaches. To confirm this hypothesis, we performed mixed stock analysis (MSA) on 40 green turtles sampled in Martinique Island and satellite tracked 31 juvenile green turtles tagged in Martinique to (a) assess their natal origin and (b) identify their destination. Our results from MSA confirm that these juveniles are descendant from females laying on several Caribbean and Atlantic beaches, mostly from Suriname and French Guiana, but also from more southern Brazilian beaches. These results were confirmed by the tracking data as the 10 turtles leaving Martinique headed across the Caribbean-Atlantic region in six different directions and 50% of these turtles reached the Brazilian foraging grounds used by the adult green turtles coming from French Guiana. One turtle left the French Guianan coast to perform the first transatlantic migration ever recorded in juvenile green turtles, swimming toward Guinea-Bissau, which is the most important nesting site for green turtles along the African coast. The extensive movements of the migrant turtles evidenced the crossing of international waters and more than 25 exclusive economic zones, reinforcing the need for an international cooperative network to ensure the conservation of future breeders in this endangered species.

Post-hatching parental care behaviour and hormonal status in a precocial bird.

In birds, the link between parental care behaviour and prolactin release during incubation persists after hatching in altricial birds, but has never been precisely studied during the whole rearing period in precocial species, such as ducks. The present study aims to understand how changes in parental care after hatching are related to circulating prolactin levels in mallard hens rearing ducklings. Blood was sampled in hens over at least 13 post-hatching weeks and the behaviour of the hens and the ducklings was recorded daily until fledging. Contacts between hens and the ducklings, leadership of the ducklings and gathering of them steadily decreased over post-hatching time. Conversely, resting, preening and agonistic behaviour of hens towards ducklings increased. Plasma prolactin concentrations remained at high levels after hatching and then fell after week 6 when body mass and structural size of the young were close to those of the hen. Parental care behaviour declined linearly with brood age, showed a disruption of the hen-brood bond at week 6 post-hatching and was related to prolactin concentration according to a sigmoid function. Our results suggest that a definite threshold in circulating prolactin is necessary to promote and/or to maintain post-hatching parental care in ducks.

How stallions influence the dynamic of collective movements in two groups of domestic horses, from departure to arrival.

The role of leader in polygynous species has been solely attributed to the male for some time, but recent studies have shown decision making to be distributed within the group. However, the specific reproductive strategy and behavioural repertoire of males in polygynous species such as horses may mean that these individuals still have the potential to play a specific role during decision making. To investigate this subject, we thoroughly studied the behaviour of two domestic stallions during collective movements of their group. We found that they initiated rarely and sometimes failed to recruit the entire group. When departing as followers, they did not accelerate the joining process. Both stallions preferentially occupied the rear position and exhibited numerous monitoring behaviours. Herding behaviours were performed by only one stallion and mostly occurred outside movement context. Finally, we removed this herding stallion from its group to evaluate how the group dynamic changed. As a result, half of the collective movements were five times slower and mares were more dispersed in comparison when the stallion was in the group. Overall, our results suggest that, the two stallions maintained their role of group monitors from departure to arrival. Their influence on the movement dynamic was indirect and did not play a specific role in the process of decision making.