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Schwartz, L. P., Silberberg, A., Casey, A. H., Kearns, D. N., & Slotnick, B. (2017). Does a rat release a soaked conspecific due to empathy? Anim. Cogn., 20(2), 299–308.
Abstract: In Experiment 1, rats choosing in an E maze preferred to release a rat standing in a pool of water to dry ground over a rat already standing on dry ground. Five additional experiments showed that the choosing rat's preference for releasing the wet rat was maintained by two separable outcomes: (1) the social contact offered by the released rat and (2) the reinforcing value of proximity to a pool of water. These results call into question Sato et al.'s (Anim Cogn 18:1039-1047, 2015) claim to have demonstrated that a rat's releasing of a wet rat to dry ground is empathically motivated.
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Giljov, A., Malashichev, Y., & Karenina, K. (2019). What do wild saiga antelopes tell us about the relative roles of the two brain hemispheres in social interactions? Anim. Cogn., .
Abstract: Two brain hemispheres are unequally involved in the processing of social stimuli, as demonstrated in a wide range of vertebrates. A considerable number of studies have shown the right hemisphere advantage for social processing. At the same time, an approach-withdrawal hypothesis, mainly based on experimental evidence, proposes the involvement of both brain hemispheres according to approach and withdrawal motivation. The present study aimed to test the relative roles of the two hemispheres in social responses displayed in a natural context. Visual biases, implicating hemispheric lateralization, were estimated in the social interactions of saiga antelope in the wild. In individually identified males, the left/right visual field use during approach and withdrawal responses was recorded based on the lateral head/body position, relative to the conspecific. Lateralized approach responses were investigated in three types of interactions, with left visual field bias found for chasing a rival, no bias--for attacking a rival, and right visual field bias--for pursuing a female. In two types of withdrawal responses, left visual field bias was found for retreating after fighting, while no bias was evident in fight rejecting. These findings demonstrate that neither the right hemisphere advantage nor the approach-withdrawal distinction can fully explain the patterns of lateralization observed in social behaviour. It is clear that both brain hemispheres play significant roles in social responses, while their relative contribution is likely determined by a complex set of motivational and emotional factors rather than a simple dichotomous distinction such as, for example, approach versus withdrawal motivation.
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Bernauer, K., Kollross, H., Schuetz, A., Farmer, K., & Krueger, K. (2020). How do horses (Equus caballus) learn from observing human action? Anim. Cogn., 23, 1–9.
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.
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Baragli, P., Scopa, C., Maglieri, V., & Palagi, E. (2021). If horses had toes: demonstrating mirror self recognition at group level in Equus caballus. Anim. Cogn., .
Abstract: Mirror self-recognition (MSR), investigated in primates and recently in non-primate species, is considered a measure of self-awareness. Nowadays, the only reliable test for investigating MSR potential skills consists in the untrained response to a visual body mark detected using a reflective surface. Here, we report the first evidence of MSR at group level in horses, by facing the weaknesses of methodology present in a previous pilot study. Fourteen horses were used in a 4-phases mirror test (covered mirror, open mirror, invisible mark, visible colored mark). After engaging in a series of contingency behaviors (looking behind the mirror, peek-a-boo, head and tongue movements), our horses used the mirror surface to guide their movements towards their colored cheeks, thus showing that they can recognize themselves in a mirror. The analysis at the group level, which 'marks' a turning point in the analytical technique of MSR exploration in non-primate species, showed that horses spent a longer time in scratching their faces when marked with the visible mark compared to the non-visible mark. This finding indicates that horses did not see the non-visible mark and that they did not touch their own face guided by the tactile sensation, suggesting the presence of MSR in horses. Although a heated debate on the binary versus gradualist model in the MSR interpretation exists, recent empirical pieces of evidence, including ours, indicate that MSR is not an all-or-nothing phenomenon that appeared once in phylogeny and that a convergent evolution mechanism can be at the basis of its presence in phylogenetically distant taxa.
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Trösch, M., Pellon, S., Cuzol, F., Parias, C., Nowak, R., Calandreau, L., et al. (2020). Horses feel emotions when they watch positive and negative horse-human interactions in a video and transpose what they saw to real life. Anim. Cogn., 23(4), 643–653.
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.
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Genov, P. W., & Kostava, V. (1993). Untersuchungen zur zahlenmäßigen Stärke des Wolfes und seiner Einwirkung auf die Haustierbestände in Bulgarien. Zeitschrift für Jagdwissenschaft, 39(4), 217–223.
Abstract: Die Untersuchung wurde in der Zeitspanne von 1984 bis 1988 durchgeführt. Es wurden die Protokolle des Staatlichen Versicherungsinstituts benutzt, die Angaben für Raubüberfälle von Wölfen auf Haustiere beinhalten (Tabelle 1). Außerdem wurden Angaben über die während dieser Zeitspanne erlegten Wölfe zusammengefaßt. Die Abschußzahlen lauten: 1984 – 163, 1985 – 147, 1986 – 179, 1987 – 211 und 1988 – 220 Tiere. Die Anzahl der in den einzelnen Gebirgen lebenden Wölfe wurde nach einer Umfrage festgestellt. Für die in Betracht kommenden Gebirge werden folgende Bestandszahlen angenommen: Rhodopen -- 60-80 Individuen, 189 bis 264 km2 pro Tier, Rila- und Piringebirge -- 60-80 Tiere, 109 bis 145 km2 pro Tier, Ossogowo-Belassiza Gebirgssystem -- 40-50 Individuen, 57-70 km2 pro Tier, West- und Mittelbalkan -- 35-38 Wölfe, 200 km2 pro Tier. Dazu kommen noch 10-15 Wölfe im Flußbecken von Beli Lom und etwa 20 Exemplare in Strandscha- und Sakargebirge. Insgesamt lebten in Bulgarien im Jahre 1988 etwa 260-330 Wölfe (Abb. 1).
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Mori, E., Benatti, L., Lovari, S., & Ferretti, F. (2016). What does the wild boar mean to the wolf? European Journal of Wildlife Research, 63(1), 9.
Abstract: Generalist predators are expected to shape their diets according to the local availability of prey species. In turn, the extent of consumption of a prey would be influenced by the number of alternative prey species. We have tested this prediction by considering the wild boar and the grey wolf: two widespread species whose distribution ranges overlap largely in Southern Europe, e.g. in Italy. We have reviewed 16 studies from a total of 21 study areas, to assess whether the absolute frequency of occurrence of wild boar in the wolf diet was influenced by (i) occurrence of the other ungulate species in diet and (ii) the number of available ungulate species. Wild boar turned out to be the main prey of the wolf (49% occurrence, on average), followed by roe deer (24%) and livestock (18%). Occurrence of wild boar in the wolf diet decreased with increasing usage of roe deer, livestock, and to a lower extent, chamois and red deer. The number of prey species did not influence the occurrence of wild boar in the wolf diet. The wild boar is a gregarious, noisy and often locally abundant ungulate, thus easily detectable, to a predator. In turn, the extent of predation on this ungulate may not be influenced so much by the availability of other potential prey. Heavy artificial reductions of wild boar numbers, e.g. through numerical control, may concentrate predation by wolves on alternative prey (e.g. roe deer) and/or livestock, thus increasing conflicts with human activities.
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Ronnenberg, K., Habbe, B., Gräber, R., Strauß, E., & Siebert, U. (2017). Coexistence of wolves and humans in a densely populated region (Lower Saxony, Germany). Basic. Appl. Ecol., 25, 1–14.
Abstract: Since the first sporadic occurrences of grey wolves (Canis lupus) west of the Polish border in 1996, wolves have shown a rapid population recovery in Germany. Wolves are known to avoid people and wolf attacks on humans are very rare worldwide. However, the subjectively perceived threat is considerable, especially as food-conditioned habituation to humans occurs sporadically. Lower Saxony (Germany) has an exceedingly higher human population density than most other regions with territorial wolves; thus, the potential for human-wolf conflicts is higher. Using hunters' wildlife survey data from 455 municipalities and two years (2014-2015) and data from the official wolf monitoring (557 confirmed wolf presences and 500 background points) collected between 2012-2015, grey wolf habitat selection was modelled using generalized additive models with respect to human population density, road density, forest cover and roe deer density. Moreover, we tested whether habitat use changed in response to human population and road density between 2012/2013 and 2014/2015. Wolves showed a preference for areas of low road density. Human population density was less important as a covariate in the model of the survey data. Areas with higher prey abundance (5-10 roe deer/km2) and areas with >20% forest cover were preferred wolf habitats. Wolves were mostly restricted to areas with the lowest road and human population densities. However, between the two time periods, avoidance of human density decreased significantly. Recolonization of Germany is still in its early stages and it is unclear where this process will halt. To-date authorities mainly concentrate on monitoring measures. However, to avoid conflict, recolonization will require more stringent management of wolf populations and an improved information strategy for rural populations.
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Heyes, C. M. (1994). Social learning in animals: categories and mechanisms. Biol. Rev., 69(2), 207–231.
Abstract: There has been relatively little research on the psychological mechanisms of social learning. This may be due, in part, to the practice of distinguishing categories of social learning in relation to ill-defined mechanisms (Davis, 1973; Galef, 1988). This practice both makes it difficult to identify empirically examples of different types of social learning, and gives the false impression that the mechanisms responsible for social learning are clearly understood. It has been proposed that social learning phenomena be subsumed within the categorization scheme currently used by investigators of asocial learning. This scheme distinguishes categories of learning according to observable conditions, namely, the type of experience that gives rise to a change in an animal (single stimulus vs. stimulus-stimulus relationship vs. response-reinforcer relationship), and the type of behaviour in which this change is detected (response evocation vs. learnability) (Rescorla, 1988). Specifically, three alignments have been proposed: (i) stimulus enhancement with single stimulus learning, (ii) observational conditioning with stimulus-stimulus learning, or Pavlovian conditioning, and (iii) observational learning with response-reinforcer learning, or instrumental conditioning. If, as the proposed alignments suggest, the conditions of social and asocial learning are the same, there is some reason to believe that the mechanisms underlying the two sets of phenomena are also the same. This is so if one makes the relatively uncontroversial assumption that phenomena which occur under similar conditions tend to be controlled by similar mechanisms. However, the proposed alignments are intended to be a set of hypotheses, rather than conclusions, about the mechanisms of social learning; as a basis for further research in which animal learning theory is applied to social learning. A concerted attempt to apply animal learning theory to social learning, to find out whether the same mechanisms are responsible for social and asocial learning, could lead both to refinements of the general theory, and to a better understanding of the mechanisms of social learning. There are precedents for these positive developments in research applying animal learning theory to food aversion learning (e.g. Domjan, 1983; Rozin & Schull, 1988) and imprinting (e.g. Bolhuis, de Vox & Kruit, 1990; Hollis, ten Cate & Bateson, 1991). Like social learning, these phenomena almost certainly play distinctive roles in the antogeny of adaptive behaviour, and they are customarily regarded as 'special kinds' of learning (Shettleworth, 1993).(ABSTRACT TRUNCATED AT 400 WORDS)
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Schino, G., & Aureli, F. (2016). Reciprocity in group-living animals: partner control versus partner choice. Biol Rev, 92(2), 665–672.
Abstract: ABSTRACT Reciprocity is probably the most debated of the evolutionary explanations for cooperation. Part of the confusion surrounding this debate stems from a failure to note that two different processes can result in reciprocity: partner control and partner choice. We suggest that the common observation that group-living animals direct their cooperative behaviours preferentially to those individuals from which they receive most cooperation is to be interpreted as the result of the sum of the two separate processes of partner control and partner choice. We review evidence that partner choice is the prevalent process in primates and propose explanations for this pattern. We make predictions that highlight the need for studies that separate the effects of partner control and partner choice in a broader variety of group-living taxa.
Keywords: cooperation; reciprocity; partner control; partner choice; proximate mechanisms
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