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de Waal, F. B. (1996). Macaque social culture: development and perpetuation of affiliative networks. J Comp Psychol, 110(2), 147–154.
Abstract: Maternal affiliative relations may be transmitted to offspring, similar to the way in which maternal rank determines offspring rank. The development of 23 captive female rhesus monkeys (Macaca mulatta) was followed from the day of birth until adulthood. A multivariate analysis compared relations among age peers with affiliative relations, kinship, and rank distance among mothers. Maternal relations were an excellent predictor of affiliative relations among daughters, explaining up to 64% of the variance. Much of this predictability was due to the effect of kinship. However, after this variable had been controlled, significant predictability persisted. For relations of female subjects with male peers, on the other hand, maternal relations had no significant predictive value beyond the effect of kinship. One possible explanation of these results is that young rhesus females copy maternal social preferences through a process of cultural learning.
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de Waal, F. B., & Luttrell, L. M. (1986). The similarity principle underlying social bonding among female rhesus monkeys. Folia Primatol (Basel), 46(4), 215–234.
Abstract: Twenty adult female rhesus monkeys (Macaca mulatta) were observed over a three-year period. They lived in a mixed captive group with kinship relations known for three generations. The study's aim was to test Seyfarth's [J. theor. Biol. 65: 671-698, 1977] model of rank-related grooming and to investigate two other possible determinants of social bonding, i.e. relative age and the group's stratification into two social classes. Data on affiliation, coalitions, and social competition were collected by means of both focal observation and instantaneous time sampling. Whereas certain elements of the existing model were confirmed, its explanatory principles were not. Social competition did not result in more contact among close-ranking females (the opposite effect was found), and the relation between affiliative behavior and coalitions was more complex than predicted. Based on multivariate analyses and a comparison of theoretical models, we propose a simpler, more encompassing principle underlying interfemale attraction. According to this 'similarity principle', rhesus females establish bonds with females whom they most resemble. The similarity may concern genetical and social background, age, hierarchical position and social class. Effects of these four factors were independently demonstrated. The most successful model assumed that similarity factors influence female bonding in a cumulative fashion.
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Dugatkin, L. A., & Godin, G. J. (1992). Predator inspection, shoaling and foraging under predation hazard in the Trinidadian guppy,Poecilia reticulata. Environmental Biology of Fishes, 34(3), 265–276.
Abstract: Guppies,Poecilia reticulata, living in stream pools in Trinidad, West Indies, approached a potential fish predator (a cichlid fish model) in a tentative, saltatory manner, mainly as singletons or in pairs. Such behavior is referred to as predator inspection behavior. Inspectors approached the trunk and tail of the predator model more frequently, more closely and in larger groups than they approached the predator's head, which is presumably the most dangerous area around the predator. However, guppies were not observed in significantly larger shoals in the stream when the predator model was present. In a stream enclosure, guppies inspected the predator model more frequently when it was stationary compared to when it was moving, and made closer inspections to the posterior regions of the predator than to its head. Therefore, the guppies apparently regarded the predator model as a potential threat and modified their behavior accordingly when inspecting it. Guppies exhibited a lower feeding rate in the presence of the predator, suggesting a trade-off between foraging gains and safety against predation. Our results further suggest that predator inspection behavior may account for some of this reduction in foraging. These findings are discussed in the context of the benefits and costs of predator inspection behavior.
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Dugatkin, L. A., & Mesterton-Gibbons, M. (1996). Cooperation among unrelated individuals: reciprocal altruism, by-product mutualism and group selection in fishes. Biosystems, 37(1-2), 19–30.
Abstract: Cooperation among unrelated individuals can evolve not only via reciprocal altruism but also via trait-group selection or by-product mutualism (or some combination of all three categories). Therefore the (iterated) prisoner's dilemma is an insufficient paradigm for studying the evolution of cooperation. We replace this game by the cooperator's dilemma, which is more versatile because it enables all three categories of cooperative behavior to be examined within the framework of a single theory. Controlled studies of cooperation among fish provide examples of each category of cooperation. Specifically, we describe reciprocal altruism among simultaneous hermaphrodites that swap egg parcels, group-selected cooperation among fish that inspect dangerous predators and by-product mutualism in the cooperative foraging of coral-reef fish.
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Dyer, J. R. G., Johansson, A., Helbing, D., Couzin, I. D., & Krause, J. (2009). Leadership, consensus decision making and collective behaviour in humans. Phil. Trans. Biol. Sci., 364(1518), 781–789.
Abstract: This paper reviews the literature on leadership in vertebrate groups, including recent work on human groups, before presenting the results of three new experiments looking at leadership and decision making in small and large human groups. In experiment 1, we find that both group size and the presence of uninformed individuals can affect the speed with which small human groups (eight people) decide between two opposing directional preferences and the likelihood of the group splitting. In experiment 2, we show that the spatial positioning of informed individuals within small human groups (10 people) can affect the speed and accuracy of group motion. We find that having a mixture of leaders positioned in the centre and on the edge of a group increases the speed and accuracy with which the group reaches their target. In experiment 3, we use large human crowds (100 and 200 people) to demonstrate that the trends observed from earlier work using small human groups can be applied to larger crowds. We find that only a small minority of informed individuals is needed to guide a large uninformed group. These studies build upon important theoretical and empirical work on leadership and decision making in animal groups.
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Dyson, H. J., & Beattie, J. K. (1982). Spin state and unfolding equilibria of ferricytochrome c in acidic solutions. J Biol Chem, 257(5), 2267–2273.
Abstract: Equilibrium, stopped flow, and temperature-jump spectrophotometry have been used to identify processes in the unfolding of ferricytochrome c in acidic aqueous solutions. A relaxation occurring in approximately 100 microseconds involves perturbation of a spin-equilibrium between two folded conformers of the protein with methionine-80 coordinated or dissociated from the heme iron. The protein unfolds more slowly, in milliseconds, with dissociation and protonation of histidine-18. These two transitions appear cooperative in equilibrium measurements at low (0.01 M) ionic strength, but are separated at higher (0.10 M) ionic strength. They are resolved under both conditions in the dynamic measurements. The spin-equilibrium description permits a unified explanation of a number of properties of ferricytochrome c in acidic aqueous solutions.
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Faria, J. J., Dyer, J. R. G., Tosh, C. R., & Krause, J. (2010). Leadership and social information use in human crowds. Anim. Behav., 79(4), 895–901.
Abstract: One of the big challenges for group-living animals is to find out who in a group has pertinent information (regarding food or predators) at any moment in time, because informed individuals may not be obviously recognizable to other group members. We found that individuals in human groups were capable of identifying those with information, and this identification increased group performance: the speed and accuracy of groups in reaching a target. Using video analysis we found how informed individuals might have been identified by other group members by means of inadvertent social cues (such as starting order, time spent following and group position). Furthermore, we were able to show that at least one of these cues, the group position of informed individuals, was indeed correlated with group performance. Our final experiment confirmed that leadership was even more efficient when the group members were given the identity of the leader. We discuss the effect of information status regarding the presence and identity of leaders on collective animal behaviour.
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Fischhoff, I. R., Sundaresan, S. R., Cordingley, J., Larkin, H. M., Sellier, M. - J., & Rubenstein, D. I. (2007). Social relationships and reproductive state influence leadership roles in movements of plains zebra, Equus burchellii. Anim. Behav., 73(5), 825–831.
Abstract: In animal groups, collective movements emerge from individual interactions. Biologists seek to identify how characteristics of actors in these groups, and their relationships, influence the decision-making process. We distinguished two basic factors determining leadership in group choices: identity and state. We hypothesized that identity is more important to leadership in groups with stable relationships, which permit the development of habitual roles. In groups with fluid membership, particular individuals or subgroups are less likely to emerge as consistent leaders. Instead, we predicted that movement initiation in unstable groups depends on individual state at the time of the decision. We characterized how identity and reproductive state influenced leadership patterns in the movements of plains zebra. As in many other mammals, lactation in this species significantly alters water and energy needs. We investigated leadership in tightly knit harems and loosely bonded herds of multiple harems. Harem females tended to have habitual roles in the initiation of harem movement. In herds, however, we found no consistent leaders among harems. At both levels of social organization, lactation was a key determinant of leadership. In harems, lactating females were more likely to initiate movement than nonlactating females. In turn, harems containing lactating females were more likely to lead herd movements. Thus, we conclude that social relationships and reproductive state together shape the interactions that produce group behaviours. One benefit to lactating females of leading herd movements is preferential access to scarce water. Thus, leadership roles in group decisions may have fitness consequences.
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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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Flauger, B., & Krueger, K. (2013). Aggressionslevel und Platzangebot bei Pferden (Equus caballus) [ Aggression level and enclosure size in horses (Equus caballus)]. Pferdeheilkunde, 29(4), 495–504.
Abstract: Viele Pferdebesitzer bevorzugen aus Angst vor aggressiven Interaktionen und Verletzungsgefahr der Tiere untereinander die Einzelhaltung, obwohl von Tierschutzorganisationen die Gruppenhaltung für Pferde empfohlen wird. In dieser Studie beobachteten wir während des alltäglichen Soziallebens als auch bei der Eingliederung von neuen Gruppenmitgliedern das Sozialverhalten, insbesondere das Aggressionsverhalten, von elf Gruppen domestizierter Pferde (Equus caballus) verschiedener Größe und Zusammensetzung. Während des alltäglichen Soziallebens hatten die Gruppe und der Paddock-Typ (Gras / kein Gras) keinen Einfluss auf die Verhaltensweisen, wohingegen die Paddockgröße unter 10000 m2 einen signifikanten Einfluss auf die submissiven Verhaltensweisen (GzLM; n=56; t=-2.061, P=0.044) und einen nicht signifikanten Einfluss auf die aggressiven Verhaltensweisen (GzLM; n=56; t=-1.782, P=0.081) hatte. Allerdings verringerten sich sowohl die aggressiven als auch die submissiven Verhaltensweisen mit steigendem Platzangebot bis zu 10000 m2 (Spearman rank Korrelation; n=56; aggressive Verhaltensweisen: r = -0.313, P = 0.019; submissive Verhaltensweisen: r = -0.328, P = 0.014). Während den Eingliederungen reduzierten sich die Aggressionen pro Stunde mit der Vergrößerung des Platzangebotes (Spearman rank Korrelation; n=28; r=-0.402, P=0.034). Dies zeigte sich noch deutlicher, wenn Beobachtungen mit einem Platzangebot von über 10000 m2 ausgeschlos- sen wurden (Spearman rank Korrelation; n=23; r=-0.549, P=0.007). Während des alltäglichen Soziallebens näherte sich der Aggressionslevel der Nulllinie an, wenn das Platzangebot pro Pferd mehr als 331 m2 betrug. Deshalb empfehlen wir zur Reduzierung des Aggressionslevels und des Verletzungsrisikos von sozial gehaltenen Pferdegruppen ein Platzangebot von mindestens 331 m2 pro Pferd.
[Even though animal welfare organisations propose group housing for horse welfare, many owners stable horses individually for fear of aggressive interactions and injury risks. In the present study we observed social behaviour, and especially aggressiveness, in eleven domestic horse groups (Equus caballus) of different size and composition, in basic social situations and when new group members were introduced. During basic social situations, the group and the type of paddock (grass / no grass) had no effect on any of the behaviours, where- as the enclosure size below 10,000 m2 had a significant effect on submissive behaviour (GzLM; n=56; t=-2.061, P=0.044) and an insignificant effect on aggressive behaviour (GzLM; n=56; t=-1.782, P=0.081). However, aggressive and submissive behaviour dimi- nished with the increase of enclosure sizes up to 10,000 m2 (Spearman rank correlation; n = 56; aggressive behaviour: r = -0.313, P=0.019; submissive behaviour: r=-0.328, P=0.014). During introductions, aggression levels per hour decreased with any increase of enclosure size (Spearman rank correlation; n=28; r=-0.402, P=0.034) and even more when enclosure sizes above 10,000 m2 were excluded (Spearman rank correlation; n=23; r=-0.549, P=0.007). During basic social situations the aggression level approached zero when the space allowance was more than 331 m2 per horse. We therefore recommend keeping horse groups in an enclosure with at least 331 m2 per horse to reduce aggression and injuries.]
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