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Bentley-Condit, V., & Smith, E. O. (2010). Animal tool use: current definitions and an updated comprehensive catalog. Behaviour, 147(2), 185–32.
Abstract: Despite numerous attempts to define animal tool use over the past four decades, the definition remains elusive and the behaviour classification somewhat subjective. Here, we provide a brief review of the definitions of animal tool use and show how those definitions have been modified over time. While some aspects have remained constant (i.e., the distinction between 'true' and 'borderline' tool use), others have been added (i.e., the distinction between 'dynamic' and 'static' behaviours). We present an updated, comprehensive catalog of documented animal tool use that indicates whether the behaviours observed included any 'true' tool use, whether the observations were limited to captive animals, whether tool manufacture has been observed, and whether the observed tool use was limited to only one individual and, thus, 'anecdotal' (i.e., N = 1). Such a catalog has not been attempted since Beck (1980). In addition to being a useful reference for behaviourists, this catalog demonstrates broad tool use and manufacture trends that may be of interest to phylogenists, evolutionary ecologists, and cognitive evolutionists. Tool use and tool manufacture are shown to be widespread across three phyla and seven classes of the animal kingdom. Moreover, there is complete overlap between the Aves and Mammalia orders in terms of the tool use categories (e.g., food extraction, food capture, agonism) arguing against any special abilities of mammals. The majority of tool users, almost 85% of the entries, use tools in only one of the tool use categories. Only members of the Passeriformes and Primates orders have been observed to use tools in four or more of the ten categories. Thus, observed tool use by some members of these two orders (e.g., Corvus, Papio) is qualitatively different from that of all other animal taxa. Finally, although there are similarities between Aves and Mammalia, and Primates and Passeriformes, primate tool use is qualitatively different. Approximately 35% of the entries for this order demonstrate a breadth of tool use (i.e., three or more categories by any one species) compared to other mammals (0%), Aves (2.4%), and the Passeriformes (3.1%). This greater breadth in tool use by some organisms may involve phylogenetic or cognitive differences � or may simply reflect differences in length and intensity of observations. The impact that tool usage may have had on groups' respective ecological niches and, through niche-construction, on their respective evolutionary trajectories remains a subject for future study.
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Cooper, J. J., & Albentosa, M. J. (2005). Behavioural adaptation in the domestic horse: potential role of apparently abnormal responses including stereotypic behaviour. Livest. Prod. Sci., 92(2), 177–182.
Abstract: Classically, biologists have considered adaptation of behavioural characteristics in terms of long-term functional benefits to the individual, such as survival or reproductive fitness. In captive species, including the domestic horse, this level of explanation is limited, as for the most part, horses are housed in conditions that differ markedly from those in which they evolved. In addition, an individual horse's reproductive fitness is largely determined by man rather than its own behavioural strategies. Perhaps for reasons of this kind, explanations of behavioural adaptation to environmental challenges by domestic animals, including the capacity to learn new responses to these challenges, tend to concentrate on the proximate causes of behaviour. However, understanding the original function of these adaptive responses can help us explain why animals perform apparently novel or functionless activities in certain housing conditions and may help us to appreciate what the animal welfare implications might be. This paper reviews the behavioural adaptation of the domestic horse to captivity and discusses how apparently abnormal behaviour may not only provide a useful practical indicator of specific environmental deficiencies but may also serve the animal as an adaptive response to these deficiencies in an “abnormal” environment.
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Boissy, A. (1995). Fear and Fearfulness in Animals. The Quarterly Review of Biology, 70(2), 165–191.
Abstract: Persistence of individual differences in animal behavior in reactions to various environmental challenges could reflect basic divergences in temperament, which might be used to predict details of adaptive response. Although studies have been carried out on fear and anxiety in various species, including laboratory, domestic and wild animals, no consistent definition of fearfulness as a basic trait of temperament has emerged. After a classification of the events that may produce a state of fear, this article describes the great variability in behavior and in physiological patterns generally associated with emotional reactivity. The difficulties of proposing fearfulness-the general capacity to react to a variety of potentially threatening situations-as a valid basic internal variable are then discussed. Although there are many studies showing covariation among the psychobiological responses to different environmental challenges, other studies find no such correlations and raise doubts about the interpretation of fearfulness as a basic personality trait. After a critical assessment of methodologies used in fear and anxiety studies, it is suggested that discrepancies among results are mainly due to the modulation of emotional responses in animals, which depend on numerous genetic and epigenetic factors. It is difficult to compare results obtained by different methods from animals reared under various conditions and with different genetic origins. The concept of fearfulness as an inner trait is best supported by two kinds of investigations. First, an experimental approach combining ethology and experimental psychology produces undeniable indicators of emotional reactivity. Second, genetic lines selected for psychobiological traits prove useful in establishing between behavioral and neuroendocrine aspects of emotional reactivity. It is suggested that fearfulness could be considered a basic feature of the temperament of each individual, one that predisposes it to respond similarly to a variety of potentially alarming challenges, but is nevertheless continually modulated during development by the interaction of genetic traits of reactivity with environmental factors, particularly in the juvenile period. Such interaction may explain much of the interindividual variability observed in adaptive responses.
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Carson, K., & Wood-Gush, D. G. M. (1983). Equine behaviour: I. A review of the literature on social and dam--Foal behaviour. Applied Animal Ethology, 10(3), 165–178.
Abstract: In most cases, the social organisation of each of the seven species of Equidae existing today outside captivity is either territorial or non-territorial. The striking differences found between these two types of organisation in the social grouping and bonds, mating behaviour, leadership and dominance hierarchies of the animals are examined. It is thought that the non-territorial species show a less primitive type of organisation than the territorial animals. Infant Equidae are precocious animals and are able to follow their dams soon after birth. They stay close by their dams and travel with the herd from an early age and are therefore classified as “followers”, in contrast to the species which have a period of hiding after birth. Dams recognise their foals immediately after birth, whereas it takes 2 or 3 days for a foal to form an attachment to its dam. Being in close proximity to their dams, foals are able to nurse frequently and, unless artificially weaned, a foal will nurse until its dam foals again. Foals start to graze during their first week and as they grow older they spend more time grazing and less time nursing and resting. It is normal for foals to be corprophagic until one month old, and this provides them with bacteria essential for the digestion of fibre. Play behaviour is solitary in very young foals, but after 4 weeks of age, foals play together, with male foals playing more than females and showing more aggressive, fighting movements in play.
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Holzapfel, M., Wagner, C., & Kluth, G. et al. (2011). Zur Nahrungsökologie der Wölfe (Canis lupus) in Deutschland. Beiträge zur Jagd- und Wildforschung, 36, 117–128.
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Gaunitz, C., Fages, A., Hanghøj, K., Albrechtsen, A., Khan, N., Schubert, M., et al. (2018). Ancient genomes revisit the ancestry of domestic and Przewalski's horses. Science, 360(6384), 111–114.
Abstract: The Eneolithic Botai culture of the Central Asian steppes provides the earliest archaeological evidence for horse husbandry, ~5,500 ya, but the exact nature of early horse domestication remains controversial. We generated 42 ancient horse genomes, including 20 from Botai. Compared to 46 published ancient and modern horse genomes, our data indicate that Przewalski's horses are the feral descendants of horses herded at Botai and not truly wild horses. All domestic horses dated from ~4,000 ya to present only show ~2.7% of Botai-related ancestry. This indicates that a massive genomic turnover underpins the expansion of the horse stock that gave rise to modern domesticates, which coincides with large-scale human population expansions during the Early Bronze Age.
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McBride, S. D., Parker, M. O., Roberts, K., & Hemmings, A. (2017). Applied neurophysiology of the horse; implications for training, husbandry and welfare. Appl. Anim. Behav. Sci., 190, 90–101.
Abstract: Understanding the neural circuits underlying equine behaviour has the potential to help optimise strategies of husbandry and training. This review discusses two areas of neurophysiological research in a range of species and relates this information to the horse. The first discussion focuses on mechanisms of learning and motivation and assesses how this information can be applied to improve the training of the horse. The second concerns the identification of the equine neurophysiological phenotype, through behavioural and genetic probes, as a way of improving strategies for optimal equine husbandry and training success. The review finishes by identifying directions for future research with an emphasis on how neurophysiological systems (and thus behaviour) can be modified through strategic husbandry. This review highlights how a neurophysioloigical understanding of horse behaviour can play an important role in attaining the primary objectives of equitation science as well as improving the welfare of the horse.
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Versace, E., Morgante, M., Pulina, G., & Vallortigara, G. (2007). Behavioural lateralization in sheep (Ovis aries). Behav. Brain. Res., 184(1), 72–80.
Abstract: This study investigates behavioural lateralization in sheep and lambs of different ages. A flock was tested in a task in which the animals were facing an obstacle and should avoid it on either the right or left side to rejoin flock-mates (adult sheep) or their mothers (lambs). A bias for avoiding the obstacle on the right side was observed, with lambs apparently being more lateralized than sheep. This right bias was tentatively associated with the left-hemifield laterality in familiar faces recognition which has been documented in this species. Differences between adult sheep and lambs were likely to be due to differences in social reinstatement motivation elicited by different stimuli (flock-mates or mothers) at different ages. Preferential use of the forelegs to step on a wood-board and direction of jaw movement during rumination was also tested in adult animals. No population bias nor individual-level lateralization was observed for use of the forelegs. At the same time, however, there was a large number of animals showing individual-level lateralization for the direction of jaw movement during rumination even though there was no population bias. These findings highlight that within the same species individual- and population-level lateralization can be observed in different tasks. Moreover, the results fit the general hypothesis that population-level asymmetries are more likely to occur in tasks that require social coordination among behaviourally asymmetric individuals.
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Mejdell, C. M., Buvik, T., Jørgensen, G. H. M., & Bøe, K. E. (2016). Horses can learn to use symbols to communicate their preferences. Appl. Anim. Behav. Sci., 184, 66–73.
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.
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Briard, L., Deneubourg, J. - L., & Petit, O. (2017). How stallions influence the dynamic of collective movements in two groups of domestic horses, from departure to arrival. Behav. Process., 142, 56–63.
Abstract: Abstract The role of leader in polygynous species has been solely attributed to the male for some time, but recent studies shown decision making to be distributed within the group. However, the specific reproductive strategy and behavioural repertoire of males in polygynous species such as horses may mean that these individuals still have the potential to play a specific role during decision-making. To investigate this subject, we thoroughly studied the behaviour of two domestic stallions during collective movements of their group. We found that they initiated rarely and sometimes failed to recruit the entire group. When departing as followers, they did not accelerate the joining process. Both stallions preferentially occupied the rear position and exhibited numerous monitoring behaviours. Herding behaviours were performed by only one stallion and mostly occurred outside movement context. Finally, we removed this herding stallion from its group to evaluate how the group dynamic changed. As a result, half of the collective movements were five times slower and mares were more dispersed in comparison when the stallion was in the group. Overall, our results suggest that, the two stallions maintained their role of group monitors from departure to arrival. Their influence on the movement dynamic was indirect and did not play a specific role in the process of decision making.
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