Abstract: Following massive persecution and eradication, strict legal protection facilitated a successful reestablishment of wolf packs in Germany, which has been ongoing since 2000. Here, we describe this recolonization process by mitochondrial DNA control-region sequencing, microsatellite genotyping and sex identification based on 1341 mostly non-invasively collected samples. We reconstructed the genealogy of German wolf packs between 2005 and 2015 to provide information on trends in genetic diversity, dispersal patterns and pack dynamics during the early expansion process. Our results indicate signs of a founder effect at the start of the recolonization. Genetic diversity in German wolves is moderate compared to other European wolf populations. Although dispersal among packs is male-biased in the sense that females are more philopatric, dispersal distances are similar between males and females once only dispersers are accounted for. Breeding with close relatives is regular and none of the six male wolves originating from the Italian/Alpine population reproduced. However, moderate genetic diversity and inbreeding levels of the recolonizing population are preserved by high sociality, dispersal among packs and several immigration events. Our results demonstrate an ongoing, rapid and natural wolf population expansion in an intensively used cultural landscape in Central Europe.

Abstract: A previous study demonstrated that horses can learn socially from observing humans, but could not draw any conclusions about the social learning mechanisms. Here we develop this by showing horses four different human action sequences as demonstrations of how to press a button to open a feed box. We tested 68 horses aged between 3 and 12 years. 63 horses passed the habituation phase and were assigned either to the group Hand Demo (N = 13) for which a kneeling person used a hand to press the button, Head Demo (N = 13) for which a kneeling person used the head, Mixed Demo (N = 12) for which a squatting person used both head and hand, Foot Demo (N = 12) in which a standing person used a foot, or No Demo (N = 13) in which horses did not receive a demonstration. 44 horses reached the learning criterion of opening the feeder twenty times consecutively, 40 of these were 75% of the Demo group horses and four horses were 31% of the No Demo group horses. Horses not reaching the learning criterion approached the human experimenters more often than those who did. Significantly more horses used their head to press the button no matter which demonstration they received. However, in the Foot Demo group four horses consistently preferred to use a hoof and two switched between hoof and head use. After the Mixed Demo the horses' actions were more diverse. The results indicate that only a few horses copy behaviours when learning socially from humans. A few may learn through observational conditioning, as some appeared to adapt to demonstrated actions in the course of reaching the learning criterion. Most horses learn socially through enhancement, using humans to learn where, and which aspect of a mechanism has to be manipulated, and by applying individual trial and error learning to reach their goal.

Abstract: Animals can indirectly gather meaningful information about other individuals by eavesdropping on their third-party interactions. In particular, eavesdropping can be used to indirectly attribute a negative or positive valence to an individual and to adjust one's future behavior towards that individual. Few studies have focused on this ability in nonhuman animals, especially in nonprimate species. Here, we investigated this ability for the first time in domestic horses (Equus caballus) by projecting videos of positive and negative interactions between an unknown human experimenter (a “positive” experimenter or a “negative” experimenter) and an actor horse. The horses reacted emotionally while watching the videos, expressing behavioral (facial expressions and contact-seeking behavior) and physiological (heart rate) cues of positive emotions while watching the positive video and of negative emotions while watching the negative video. This result shows that the horses perceived the content of the videos and suggests an emotional contagion between the actor horse and the subjects. After the videos were projected, the horses took a choice test, facing the positive and negative experimenters in real life. The horses successfully used the interactions seen in the videos to discriminate between the experimenters. They touched the negative experimenter significantly more, which seems counterintuitive but can be interpreted as an appeasement attempt, based on the existing literature. This result suggests that horses can indirectly attribute a valence to a human experimenter by eavesdropping on a previous third-party interaction with a conspecific.

Abstract: This paper describes a method in which horses learn to communicate by touching different neutral visual symbols, in order to tell the handler whether they want to have a blanket on or not. Horses were trained for 10-15min per day, following a training program comprising ten steps in a strategic order. Reward based operant conditioning was used to teach horses to approach and touch a board, and to understand the meaning of three different symbols. Heat and cold challenges were performed to help learning and to check level of understanding. At certain stages, a learning criterion of correct responses for 8-14 successive trials had to be achieved before proceeding. After introducing the free choice situation, on average at training day 11, the horse could choose between a “no change” symbol and the symbol for either “blanket on” or “blanket off” depending on whether the horse already wore a blanket or not. A cut off point for performance or non-performance was set to day 14, and 23/23 horses successfully learned the task within this limit. Horses of warm-blood type needed fewer training days to reach criterion than cold-bloods (P<0.05). Horses were then tested under differing weather conditions. Results show that choices made, i.e. the symbol touched, was not random but dependent on weather. Horses chose to stay without a blanket in nice weather, and they chose to have a blanket on when the weather was wet, windy and cold (χ2=36.67, P<0.005). This indicates that horses both had an understanding of the consequence of their choice on own thermal comfort, and that they successfully had learned to communicate their preference by using the symbols. The method represents a novel tool for studying preferences in horses.