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Rands, S. A., Cowlishaw, G., Pettifor, R. A., Rowcliffe, J. M., & Johnstone, R. A. (2008). The emergence of leaders and followers in foraging pairs when the qualities of individuals differ. BMC Evol Biol, 8, 51.
Abstract: BACKGROUND: Foraging in groups offers animals a number of advantages, such as increasing their likelihood of finding food or detecting and avoiding predators. In order for a group to remain together, there has to be some degree of coordination of behaviour and movement between its members (which may in some cases be initiated by a decision-making leader, and in other cases may emerge as an underlying property of the group). For example, behavioural synchronisation is a phenomenon where animals within a group initiate and then continue to conduct identical behaviours, and has been characterised for a wide range of species. We examine how a pair of animals should behave using a state-dependent approach, and ask what conditions are likely to lead to behavioural synchronisation occurring, and whether one of the individuals is more likely to act as a leader. RESULTS: The model we describe considers how the energetic gain, metabolic requirements and predation risks faced by the individuals affect measures of their energetic state and behaviour (such as the degree of behavioural synchronisation seen within the pair, and the value to an individual of knowing the energetic state of its colleague). We explore how predictable changes in these measures are in response to changes in physiological requirements and predation risk. We also consider how these measures should change when the members of the pair are not identical in their metabolic requirements or their susceptibility to predation. We find that many of the changes seen in these measures are complex, especially when asymmetries exist between the members of the pair. CONCLUSION: Analyses are presented that demonstrate that, although these general patterns are robust, care needs to be taken when considering the effects of individual differences, as the relationship between individual differences and the resulting qualitative changes in behaviour may be complex. We discuss how these results are related to experimental observations, and how the model and its predictions could be extended.
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Rogers, L. J. (2000). Evolution of hemispheric specialization: advantages and disadvantages. Brain Lang, 73(2), 236–253.
Abstract: Lateralization of the brain appeared early in evolution and many of its features appear to have been retained, possibly even in humans. We now have a considerable amount of information on the different forms of lateralization in a number of species, and the commonalities of these are discussed, but there has been relatively little investigation of the advantages of being lateralized. This article reports new findings on the differences between lateralized and nonlateralized chicks. The lateralized chicks were exposed to light for 24 h on day 19 of incubation, a treatment known to lead to lateralization of a number of visually guided responses, and the nonlateralized chicks were incubated in the dark. When they were feeding, the lateralized chicks were found to detect a stimulus resembling a raptor with shorter latency than nonlateralized chicks. This difference was not a nonspecific effect caused by the light-exposed chicks being more distressed by the stimulus. Instead, it appears to be a genuine advantage conferred by having a lateralized brain. It is suggested that having a lateralized brain allows dual attention to the tasks of feeding (right eye and left hemisphere) and vigilance for predators (left eye and right hemisphere). Nonlateralized chicks appear to perform these dual tasks less efficiently than lateralized ones. Reference is made to other species in discussing these results.
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Krueger, K., & Flauger, B. (2008). Social feeding decisions in horses (Equus caballus). Behav. Process., 78(1), 76–83.
Abstract: Like many other herbivores, in a natural environment equids feed on rather evenly distributed resources. However, the vegetation in their vast habitats constantly changes. If food is plentiful only little competition occurs over food, and in non-competitive situations domestic horses tend to return to the same feeding site until it is overgrazed. In contrast, they compete over limited food for which the social status of the individuals appears to be important. Especially in ruminants several studies have proved an influence of social organisations, rank, sex and the depletion of feeding sites on the feeding behaviour of individuals. However, it is not yet understood whether and how social aspects affect horses“ feeding decisions. Curiosity about the influence of social rank on the horses” feeding decisions between two, equally with high-quality surplus food-filled buckets placed in different social feeding conditions, led us to create the test below. The observer horses were alternately tested with a dominant and a subordinate demonstrator placed in one of the three different positions. We conclude that domestic horses use social cognition and strategic decision making in order to decide where to feed in a social feeding situation. When possible they tend to return to the same, continuously supplied feeding site and switch to an “avoidance tendency” in the presence of dominant horses or when another horse is already feeding there. Thus, the social rank and the position of conspecifics affect the feeding strategy of horses.
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Flauger, B. (2011). The introduction of horses into new social groups with special regard to their stress level. Ph.D. thesis, , .
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.
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Pluhácek, J., Bartos, L., Bartosová, J., & Kotrba, R. (2010). Feeding behaviour affects nursing behaviour in captive plains zebra (Equus burchellii). Appl. Anim. Behav. Sci., 128(1-4), 97–102.
Abstract: Equids spend more than half of the day feeding. Lactation is a very demanding form of maternal investment. In an environment such as a zoo, where no grass but several feeding sites are present, conflict between suckling behaviour of the foal and feeding behaviour of the lactating mother should occur. We observed 20 foals of captive plains zebra, Equus burchellii, at the Dvur Králové Zoo, Czech Republic, and collected data concerning suckling events during 17 months of observation. First, we examined whether feeding by the mother while nursing affected suckling behaviour. We found that when the mother was feeding, the proportion of suckling bouts she terminated decreased with increasing age of the foal, whereas it did not change when she was not feeding. This result supported the trade-off between suckling and feeding behaviour which has been reported in other ungulates. Second, we examined what affected interruptions of feeding behaviour of the mother during the suckling bout. The proportion of interruptions of feeding by the mother during nursing increased with increasing age of her foal. This coincides with declining time spent nursing. In addition, younger mothers interrupted their feeding behaviour during suckling bouts more often than older ones. Mothers interrupted feeding during the suckling bout more often when they nursed a daughter than when they nursed a son. The results of our study show that feeding while suckling could reduce parent-offspring conflict and improve welfare of captive foals and mares.
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Hoffmann, G., Bentke, A., Rose-Meierhöfer, S., Berg, W., Mazetti, P., & Hardarson, G. H. (2012). Influence of an active stable system on the behavior and body condition of Icelandic horses. animal, 6(10), 1684–1693.
Abstract: Horses are often stabled in individual boxes, a method that does not meet their natural needs and may cause psychical and
musculoskeletal diseases. This problem is particularly evident in Iceland, where horses often spend the long winter periods in cramped
boxes. The aim of this study was to analyze the suitability of a group housing system in Iceland, but the results are also applicable to
horses of other regions. Eight Icelandic horses were observed in an active stable system, and their behavior and time budget were
recorded. Movement and lying behavior were studied with ALT (Activity, Lying, Temperature detection) pedometers. The effect of an
automatic concentrate feeding station (CFS) on the horses’ behavior was examined. In the first period of investigation, the horses
were fed concentrates manually, and in the second period, they were fed with the CFS. Additional behavioral observations and a
determination of social hierarchy occurred directly or by video surveillance. The physical condition of the horses was recorded by body
weight (BW) measurement and body condition scoring (BCS). The results showed a significant increase between the first and second
trial periods in both the activity (P,0.001) and the lying time (P50.003) of the horses with use of the CFS. However, there was no
significant change in BW during the first period without the CFS (P50.884) or during the second period with the CFS (P50.540).
The BCS of the horses was constant at a very good level during both trial periods, and the horses showed a low level of aggression, a
firm social hierarchy and behavioral synchronization. This study concludes that group housing according to the active stable principle is
a welfare-friendly option for keeping horses and is a suitable alternative to conventional individual boxes.
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Baumgartner, M., Boisson, T., Erhard, M. H., & Zeitler-Feicht, M. H. (2020). Common Feeding Practices Pose A Risk to the Welfare of Horses When Kept on Non-Edible Bedding. Animals, 10, 441.
Abstract: During the evolution of the horse, an extended period of feed intake, spread over the entire 24-h period, determined the horses� behaviour and physiology. Horses will not interrupt their feed intake for more than 4 h, if they have a choice. The aim of the present study was to investigate in what way restrictive feeding practices (non ad libitum) affect the horses� natural feed intake behaviour. We observed the feed intake behaviour of 104 horses on edible (n = 30) and non-edible bedding (n = 74) on ten different farms. We assessed the duration of the forced nocturnal feed intake interruption of horses housed on shavings when no additional roughage was available. Furthermore, we comparatively examined the feed intake behaviour of horses housed on edible versus non-edible bedding. The daily restrictive feeding of roughage (2 times a day: n = 8; 3 times a day: n = 2), as it is common in individual housing systems, resulted in a nocturnal feed intake interruption of more than 4 hours for the majority (74.32%, 55/74) of the horses on shavings (8:50 ± 1:25 h, median: 8:45 h, minimum: 6:45 h, maximum: 13:23 h). In comparison to horses on straw, horses on shavings paused their feed intake less frequently and at a later latency. Furthermore, they spent less time on consuming the evening meal than horses on straw. Our results of the comparison of the feed-intake behaviour of horses on edible and non-edible bedding show that the horses� ethological feeding needs are not satisfied on non-edible bedding. If the horses accelerate their feed intake (also defined as �rebound effect�), this might indicate that the horses� welfare is compromised. We conclude that in addition to the body condition score, the longest duration of feed intake interruption (usually in the night) is an important welfare indicator of horses that have limited access to roughage.
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Vervaecke, H., Vries, H. D., & Elsacker, L. V. (1999). An Experimental Evaluation Of The Consistency Of Competitive Ability And Agonistic Dominance In Different Social Contexts In Captive Bonobos. Behaviour, 136(4), 423–442.
Abstract: Bonobos have been described as a relatively egalitarian and female dominant species. The exact nature and quality of their dominance relationships and the existence of female dominance are current topics of dispute. We investigated the consistency across social contexts, the stability in time, and the degree of expression of the competitive feeding ability and agonistic dominance in a captive group of bonobos. First, we examined whether the competitive feeding ranks and agonistic ranks differed in different dyadic contexts, triadic contexts and the whole group context. For some pairs of animals the dominance relationships with respect to competitive feeding altered with different group compositions. The agonistic dominance relationships changed accordingly. The competitive feeding ranks and agonistic ranks in the experiments correlated strongly with each other. The alpha position was occupied by a female, but not all females outranked all males. We suggest that females can profit from each others presence to gain inter-sexual dominance. Second, although the agonistic rank order in the whole group remained the same over at least five years, some dyadic competitive feeding ranks changed over time, resulting in a stronger female intersexual dominance. Third, the degree of expression of the behaviors used to quantify dyadic competitive and agonistic dominance was not high, in line with the popular 'egalitarian' epithet. Notwithstanding its low consistency across contexts, the dominance hierarchy in the whole group has a strong predictive value for other social relationships such as grooming. Given this strong effect of rank on other behaviours and given the strong dependency of rank on social context, the choice of the right party members may be a crucial factor in the fission-fusion processes of free-ranging bonobos.
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Barette, C., & Vandal, D. (1986). Social rank, dominance, antler size, and access to food in snow-bound wild woodland caribou. Behaviour, 97(1-2), 118–146.
Abstract: We spent two winters studying the social behaviour of wild woodland caribou (Rangifer tarandus caribou) at a time when their main food (ground lichens; Cladina sp.) is available only at snow craters dug by the animals. The competition for access to such craters was severe, the animals constantly trying to take over the craters of others. During a two-month period when a group maintained a constant size (20) and composition (all age-sex classes represented), we could rank the animals in a rather linear dominance hierarchy (Landau's index = 0.87). Rank was correlated with access to resources, percent of time spent active, and percent of time feeding in craters. It was also correlated with age and antler size. However, rank is not an attribute of individuals, but of a relationship between individuals. As such it is only an intervening variable between physical attributes and access to resources, a variable whose value has meaning only within a given group. Among the three attributes studied (age, sex, antler size), the latter was by far the best predictor of the occurrence and outcome of interactions. Between two individuals within any of the three age-sex classes studied (adult and yearling males and adult females), the one with larger antlers initiated significantly more often, escalated its aggression (to the point of hitting the target) less often, and enjoyed a higher success rate in obtaining resources. When their antlers were larger than those of an adult male target (i.e. males that had shed their antlers), adult females won almost all their interactions with adult males even though they escalated only one fourth of them. This clarifies the long-standing speculation that female caribou have antlers and shed them later than males, in order to overcome their sexual handicap in competition for food in the winter. We conclude that the link between rank and dominance of an individual on one hand, and some of its attributes on the other (e.g. sex, age, weight, antler size) is fundamentally realized by the animal itself through its active preference for targets it is likely to beat, i.e. targets with smaller antlers.
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Shettleworth, S. J. (2003). Memory and hippocampal specialization in food-storing birds: challenges for research on comparative cognition. Brain Behav Evol, 62(2), 108–116.
Abstract: The three-way association among food-storing behavior, spatial memory, and hippocampal enlargement in some species of birds is widely cited as an example of a new 'cognitive ecology' or 'neuroecology.' Whether this relationship is as strong as it first appears and whether it might be evidence for an adaptive specialization of memory and hippocampus in food-storers have recently been the subject of some controversy [Bolhuis and Macphail, 2001; Macphail and Bolhuis, 2001]. These critiques are based on misconceptions about the nature of adaptive specializations in cognition, misconceptions about the uniformity of results to be expected from applying the comparative method to data from a wide range of species, and a narrow view of what kinds of cognitive adaptations are theoretically interesting. New analyses of why food-storers (black-capped chickadees, Poecile Atricapilla) respond preferentially to spatial over color cues when both are relevant in a memory task show that this reflects a relative superiority of spatial memory as compared to memory for color rather than exceptional spatial attention or spatial discrimination ability. New studies of chickadees from more or less harsh winter climates also support the adaptive specialization hypothesis and suggest that within-species comparisons may be especially valuable for unraveling details of the relationships among ecology, memory, and brain in food-storing species.
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