Abstract: To determine whether wild Indo-Pacific bottlenose dolphins (Tursiops aduncus) at Mikura Island, Japan, show asymmetry of eye or flipper use during a social behaviour, we investigated the laterality of flipper-to-body (F-B) rubbing, in which one dolphin (“rubber”) rubs the body of another (“rubbee”) with its flipper. We analysed 382 episodes of video-recorded F-B rubbings performed by identified individuals (N = 111 rubbers). F-B rubbing was conducted significantly more frequently with the left flipper than with the right flipper. The duration of F-B rubbings was also significantly longer with the left flipper than with the right flipper. Of 20 dolphins, nine individuals showed significant left-side bias as the rubber in this behaviour, whereas no dolphins showed significant right-side bias. The results indicate a population-level left-side bias of the rubber in F-B rubbing. An analysis of the swimming configurations during this behaviour suggests that the asymmetry in F-B rubbing was caused not only by the laterality of the rubber, but by a preference for use of the left eye in both dolphins during this behaviour. Dolphins used the left eye significantly more frequently than the right eye during the inquisitive behaviour, while they showed no significant bias in flipper use during the object-carrying behaviour. These facts also suggest that the asymmetry of F-B rubbing is caused by the preference for using the left eye. Significant left-side bias was observed only in F-B rubbings initiated by the rubbee, in which the rubbee determined its position during this behaviour. This suggests that this behavioural asymmetry was enhanced by the rubbees choosing the left side of the rubber to ensure better and longer rubs.

Abstract: Crib-biting (wind-sucking) might be a coping response of the horses to the challenges of
uncontrolled environmental events. Prevention of this stereotypic behaviour evokes
physiological responses consistent with increased stress. Reducing the incidence of cribbiting,
however, is important in order to prevent undesirable physical and behavioural
consequences (tooth erosion, altered gut function, gastric inflammation/ulceration, colic, etc.).
Common treatment of crib-biting is the application of a cribbing collar, which limits the
flexion of the neck making this stereotypic movement uncomfortable and difficult. Another
method, the modified Forssell"s operation, is becoming more and more popular amongst the
horse owners. It is based on the removal of the muscles used in crib-biting (m.omohyoideus,
m.sternohyoideus, m.sternothyrohyoideus) and the ventral branches of the spinal accessory
nerves. Surveys on the success of this surgical procedure have revealed inconsistent results,
and, contrary to the cribbing collar, its effect on the stress level have not been studied either.
The aim of our study was to determine whether the modified Forssell"s procedure is superior
to the cribbing collar treatment.
Differences in stress management was tested by a crib-biting provoking test, in which
surgically treated horses, crib-biting horses, crib-biting horses with cribbing collar, and
normal horses (those showing no stereotypies), altogether 56 horses were compared. In this
test, a food bucket had been placed out of the reach of the animal, from which titbits were
given 3 times. Behaviour and heart rate variability (HRV) of the horses were recorded and
analysed throughout the test. Hypotheses were tested by linear mixed model.
According to our results, both prevention methods (collar or surgery) inhibited crib-biting
successfully though not totally. Regarding behaviour and heart rate variability, horses
prevented from crib-biting (by collar or surgery) differed significantly from crib-biting and
normal horses but not from each other.
Normal horses were usually trying to reach the food-bucket while present and were standing
still afterwards, whereas the other three groups had not really made efforts to reach the
bucket, spent less time with resting, and performed or tried crib-biting. During the stress-test,
normal and crib-biting horses had shown good stress-adaptation to the challenge since their
HRV, after an initial increase, returned to the basal value by the end. On the contrary, HRV of
the two prevented groups remained elevated and showed large oscillations throughout. They
had not found a successful coping behaviour either.
Our results suggest that since prevention may significantly increase distress, the treatment in
itself, without changing the motivation of the horse to perform the replacement behaviour – it
seems to be unsatisfactory and insufficient. After prevention the motivation of the horse to
perform crib-biting should be addressed. In addition, considering that prevention by collar and
surgery had not resulted in any significant behavioural or physiological differences, the
superiority of the modified Forssell"s procedure might be questioned. However, the surgery
might be recommended if treatment with collar is ineffective.

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: Behavioural innovations have become central to our thinking about how animals adjust to changing environments. It is now well established that animals vary in their ability to innovate, but understanding why remains a challenge. This is because innovations are rare, so studying innovation requires alternative experimental assays that create opportunities for animals to express their ability to invent new behaviours, or use pre-existing ones in new contexts. Problem solving of extractive foraging tasks has been put forward as a suitable experimental assay. We review the rapidly expanding literature on problem solving of extractive foraging tasks in order to better understand to what extent the processes underpinning problem solving, and the factors influencing problem solving, are in line with those predicted, and found, to underpin and influence innovation in the wild. Our aim is to determine whether problem solving can be used as an experimental proxy of innovation. We find that in most respects, problem solving is determined by the same underpinning mechanisms, and is influenced by the same factors, as those predicted to underpin, and to influence, innovation. We conclude that problem solving is a valid experimental assay for studying innovation, propose a conceptual model of problem solving in which motor diversity plays a more central role than has been considered to date, and provide recommendations for future research using problem solving to investigate innovation. This article is part of a Special Issue entitled: Cognition in the wild.

Abstract: The interaction of horses with humans is a dynamic state, but it is not clearly understood how horses perceive humans. Nervousness is transmissible from humans to horses indicated by increased horse heart rate (HR), however no studies have investigated whether horses can differentiate between humans who are physiologicallystressed (eg. after exercising) as opposed to psychologically-stressed (eg. feeling nervous/afraid). Horses (N=10) were randomly subjected to each of four treatments: 1) no human [control], 2) a calm human comfortable around horses [CALM; N=2 humans], 3) a physically-stressed human [PHYS; human exercised to reach 70% of maximum HR; N=2 humans], and 4) a psychologically-stressed human [PSYCH; human who was nervous around horses; N=14 humans]. Humans ranked themselves on a scale of 1-10 for their nervousness around horses. Both humans and horses were equipped with a HR monitor. Behavioural observations of the horses [gait, head position relative to the withers, distance from human, orientation toward human] were recorded live. Horses were allowed to wander loose in a round pen for 5 minutes of baseline recordings, at which time the human subject entered the round pen, stood in the centre and placed a blindfold over his/her eyes. The human remained in the centre of the round pen for an additional 5 minutes. Horse HR during control did not differ from when the human was present in the CALM and PSYCH treatment, and was lower during the PHYS treatment (51a vs 54a vs 55a vs 45b bpm for control, CALM, PSYCH and PHYS respectively; a,b differ p<0.0001). Over the 5 minute test period, horse HR decreased in PHYS and PSYCH (p<0.01) whereas it increased in CALM (p<0.0001). Horse HR decreased with increasing human rank of nervousness around horses (p=0.0156), and horses stood nearer to the human when they faced the human (p<0.0001) regardless of treatment. Horses moved at a faster gait in the control treatment, and their gait was slowest in the PSYCH treatment (p<0.0001), and the horse’s head position was lower in the PHYS and PSYCH treatments compared to CALM or baseline (p< 0.0001). A lower horse head position was positively correlated to a lower horse HR (p<0.0001) and negatively correlated to horse age (p<0.0001). Human HR was affected by treatment, with PHYS having the highest HR (p<0.0001). Human HR increased when the horse was facing away from the human, even though the human was blindfolded (p=0.0395). Overall, horses appear to be influenced by the physiological and psychological state of a human without any direct contact. Horses’ posture does reflect their physiological state. Understanding how horses react to human physiological and psychological states is especially important in equine-assisted activities, where the response of the horse has specific implications for the human participant.