Abstract: Abstract In recent years, behavioral ecologists have embraced social network analysis (SNA) in order to explore the structure of animal societies and the functional consequences of that structure. We provide a conceptual introduction to the field that focuses on historical developments, as well as on novel insights generated by recent work. First, we discuss major advances in the analysis of nonhuman societies, culminating in the use of SNA by behavioral ecologists. Next, we discuss how network-based approaches have enhanced our understanding of social structure and behavior over the past decade, focusing on: (1) information transmission, (2) collective behaviors, (3) animal personality, and (4) cooperation. These behaviors and phenomena possess several features—e.g., indirect effects, emergent properties—that network analysis is well equipped to handle. Finally, we highlight recent developments in SNA that are allowing behavioral ecologists to address increasingly sophisticated questions regarding the structure and function of animal sociality.

Abstract: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.

Abstract: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.

Abstract: Delayed-responses have been traditionally employed to investigate the temporal characteristics of animals“ ability to represent and recall objects that have disappeared. In the typical condition, the animal, usually a mammal, observes the experimenter hiding an interesting goal (e.g. some food) in a certain location. A delayed-response task (DRT) was administered to 4 female Esperia pony (2 years old) coming from a free-range breed (Frosinone, Italy) and to 7 female Amiata donkeys (4.2±2 years old) coming from a conservation stock (University of Pisa, Italy). The DRT's apparatus was located in a square fence. A single ”U-shaped“ screen (330x160x140 cm) made by wood shavings blocks was positioned in the centre of the fence. A gap (40x50 cm) on the ground was in the middle of the central side of the U-shaped-screen and served to make the food-attractor disappear. The food-attractor consisted in cereal flakes and fresh grass for ponies and cereal flakes for donkeys. A bucket full of food was placed on a dolly tied on a rope which could be pulled by an experimenter. In a preliminary training each animal was allowed to eat food from the bucket and, while the animal was eating, the dolly was gently pulled away from the animal, and beyond the screen through the gap. The subjects needed to move around of the screen in order to retrieve the food. As a reinforcement, they were allowed to eat some food from the bucket once behind the screen. From trial to trial, the bucket was presented farther and farther (starting with a distance of 1 m in front of the screen to reach 7 m). Therefore subjects were tested in the DRT requiring them to rejoin the bucket with the goal-food disappearing behind the screen as in the preliminary training but following a 10 s delay. For the DRT, the bucket was placed 7 m in front of the screen, 3 m away from the animal's starting area. Then the dolly was pulled away from the animal. Ten seconds after the disappearance of the dolly behind the screen the animal was released from the starting area. The DRT ended when the subject had reached the attractor behind the screen on 3 consecutive trials. Results showed that all animals were able to rejoin the food behind the screen after 10 s delay. The mean time of the delayed-response (mean±sd, in s) in the ponies (1st: 19.8±8; 2nd: 10.8±2.2; 3rd: 12.8±2.8) and in the donkeys (1st: 28.4±10; 2nd: 26.9±13; 3rd: 24.3±16.6) showed a trend to decrease from first trial to third. These preliminary results suggest that like other mammals our ponies and donkeys can maintain a working memory trace of the location where biologically attractive objects have been seen to disappear. In conclusion, this study paves the way to set up a viable model system for the investigation of the more sophisticated aspects of Equids” cognitive abilities such as working memory.

Abstract: An animal that has stepped over an obstacle with its forelimbs uses a memory of the obstacle to guide the hind limbs so that they also clear the obstacle, even in situations in which long pauses are introduced between forelimb and hind limb stepping. To further clarify the features of hind limb obstacle clearance memory, the present study examined hind limb obstacle clearance in the horse. A rider guided horses over obstacles and paused the horse over obstacles in tests that examined the relationship between forelimb and hind limb stepping, with the following results. First, the horses displayed memory for an obstacle as measured by hind limb lifting over the obstacle for durations lasting as long as 15 min. The response was not dependent upon ongoing visualization of the obstacle, as limb lifting was unaffected by visual occlusion with blinders, a blindfold, or by removing the obstacle during the pause. Second, previous experience of stepping over an obstacle led to pause-related hind limb lifting at the object's previous location even on trials for which there was no obstacle and so no preceding forelimb lifting. Third, whereas a horse would lift its hind limbs to clear two successively presented obstacles, replacing an obstacle before the horse after the forelimbs had cleared the obstacle prevented subsequent hind limb lifting at the obstacle's previous location. Taken together the results show that hind limb obstacle clearance is guided by a place-object memory. The results are discussed in relation to the differential sensory and memonic control of forelimb and hind limb stepping with the suggestion that place-object memory can guide hind stepping as well as overshadow working memory from front leg stepping.