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: This study evaluates the horse (Equus caballus) use of human local enhancement cues and reaction to human attention when making feeding decisions. The superior performance of dogs in observing human states of attention suggests this ability evolved with domestication. However, some species show an improved ability to read human cues through socialization and training. We observed 60 horses approach a bucket with feed in a three-way object-choice task when confronted with (a) an unfamiliar or (b) a familiar person in 4 different situations: (1) squatting behind the bucket, facing the horse (2) standing behind the bucket, facing the horse (3) standing behind the bucket in a back-turned position, gazing away from the horse and (4) standing a few meters from the bucket in a distant, back-turned position, again gazing away from the horse. Additionally, postures 1 and 2 were tested both with the person looking permanently at the horse and with the person alternating their gaze between the horse and the bucket. When the person remained behind the correct bucket, it was chosen significantly above chance. However, when the test person was turned and distant from the buckets, the horses’ performance deteriorated. In the turned person situations, the horses approached a familiar person and walked towards their focus of attention significantly more often than with an unfamiliar person. Additionally, in the squatting and standing person situations, some horses approached the person before approaching the correct bucket. This happened more with a familiar person. We therefore conclude that horses can use humans as a local enhancement cue independently of their body posture or gaze consistency when the persons remain close to the food source and that horses seem to orientate on the attention of familiar more than of unfamiliar persons. We suggest that socialization and training improve the ability of horses to read human cues.

Abstract: Horses are a highly social species living in complex social systems which should require them to memorise and generalise social experiences and distinguish between familiar and unfamiliar conspecifics. In the main part of my thesis I concentrated on the specific conflict situation of a horse being introduced into a new social group, and investigated its behaviour and stress level. Horses were either introduced (1) immediately, (2) after an observation period, or (3) together with an integration horse after an observation period. Additionally, in the second part of my thesis I arranged several experiments to elaborate additional aspects which could affect the behaviour of horses during introductions. In this study I could describe a simplified method for measuring stress through the analysis of faecal GCMs in horses. An enzyme immunoassay (EIA) for 11-oxoaetiocholanolone using 11-oxoaetiocholanolone-17-CMO: BSA (3?,11-oxo-A EIA) as antigen showed high amounts of immunoreactive substances. The new assay increases the accuracy of the test and lowers the expenses per sample; also storing of samples at room temperature after collection is less critical. This is a big advantage both in the field of wildlife management of equids and in the field of equestrian sports (chapter 1). Comparing the different introduction techniques, the introduction with an integration horse led to significantly less total interactions and lower levels of aggression than the introduction of single horses, both immediately and after several days of observing the new group. Additionally, by observing the behaviour of the horses during everyday sociality I could develop a formula describing the interrelationship between expected aggression level and enclosure size per horse. The curve takes an exponential shape. Starting from a space allowance of 300 m2 and more per horse, the amount of aggressions per hour approaches zero. For the reduction of aggression levels and injury risks in socially kept horses I recommend an enclosure size of at least 300 m2 per horse (chapter 2). I further investigated the stress level of the introduced animals. Horses which were immediately introduced did not show elevated faecal GCMs. In contrast, horses which were introduced after an observation period had slightly elevated values 2 and 3 days after the introduction. For horses introduced together with an integration horse faecal GCMs were significantly above the baseline value on the day of introduction and 1 day after it. These differences between introduction techniques indicate that the introduction event itself is not as stressful as previously assumed. Rather standing together with an integration horse and not being able to integrate immediately into the complete group elicits stress in horses (chapter 3). In the commentary of chapter 4 several studies are discussed which failed to demonstrate social learning in horses. It is argued that they did not consider important aspects which could have an influence, such as the dominance status or the social background of the horses (chapter 4). In chapter 5 a social feeding situation was investigated. The social rank as well as the position of conspecifics affected the feeding strategy of horses. Domestic horses used social cognition and strategic decision making in order to decide where to feed. When possible they tended to return to the same, continuously supplied feeding site and switched to an ?avoidance tendency? in the presence of dominant horses or when another horse was already feeding there (chapter 5). One possibility to recognize group members is through olfactory recognition. In chapter 6 it is shown that horses are able to distinguish their own from their conspecifics? faeces. In addition, they paid most attention to the faeces of those group members from which they received the highest amount of aggressive behaviour (chapter 6). Horses show cognitive abilities because they are able to use humans as local enhancement cues when searching for food, independently of their body posture or gaze consistency when the persons face them. Moreover, they seem to orientate on the attention of familiar persons more than of unfamiliar persons (chapter 7). Altogether, the results of this thesis provide further support for the view that horses show good conflict resolution strategies. They are perfectly able to deal with the conflict situation of being introduced to new group members, and the introduction event itself is not as stressful as previously assumed. It is rather suggested that standing together with an integration horse and not being able to integrate immediately into the complete group elicits stress in horses. All additional experimental set-ups could demonstrate that horses are well capable of social cognition.

Abstract: <sec> <title>Background</title> <p>Animals' ability for cross-modal recognition has recently received much interest. Captive or domestic animals seem able to perceive cues of human attention and appear to have a multisensory perception of humans.</p> </sec> <sec> <title>Methodology/Principal Findings</title> <p>Here, we used a task where horses have to remain immobile under a vocal order to test whether they are sensitive to the attentional state of the experimenter, but also whether they behave and respond differently to the familiar order when tested by a familiar or an unknown person. Horses' response varied according to the person's attentional state when the order was given by an unknown person: obedience levels were higher when the person giving the order was looking at the horse than when he was not attentive. More interesting is the finding that whatever the condition, horses monitored much more and for longer times the unknown person, as if they were surprised to hear the familiar order given by an unknown voice.</p> </sec> <sec> <title>Conclusion/Significance</title> <p>These results suggest that recognition of humans may lie in a global, integrated, multisensory representation of specific individuals, that includes visual and vocal identity, but also expectations on the individual's behaviour in a familiar situation.</p> </sec>

Abstract: Reasons for performing study: In horses, serum cortisol concentration is considered to provide an indirect measurement of stress. However, it includes both free and bound fractions. The sampling method is also invasive and often stressful. This is not the case for salivary cortisol, which is collected using a more welfare-friendly method and represents a part of the free cortisol fraction, which is the biologically active form. Objectives: To compare salivary and serum cortisol assays in horses, in a wide range of concentrations, using an adrenocorticotropic hormone (ACTH) stimulation test, in order to validate salivary cortisol for stress assessment in horse. Methods: In 5 horses, blood samples were drawn using an i.v. catheter. Saliva samples were taken using swabs. Cortisol was assayed by radioimmunoassay. All data were treated with a regression method, which pools and analyses data from multiple subjects for linear analysis. Results: Mean ± s.d. cortisol concentrations measured at rest were 188.81 ± 51.46 nmol/l in serum and 1.19 ± 0.54 nmol/l in saliva. They started increasing immediately after ACTH injection and peaks were reached after 96 ± 16.7 min in serum (356.98 ± 55.29 nmol/l) and after 124 ± 8.9 min in saliva (21.79 ± 7.74 nmol/l, P<0.05). Discharge percentages were also different (225% in serum and 2150% in saliva, P<0.05). Correlation between serum and salivary cortisol concentrations showed an adjusted r2= 0.80 (P<0.001). The strong link between serum and salivary cortisol concentrations was also estimated by a regression analysis. Conclusions: The reliability of both RIAs and regression found between serum and salivary cortisol concentrations permits the validation of saliva-sampling as a noninvasive technique for cortisol level assessment in horses.