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Klingel, H. (1968). Sozial Organisation und Verhaltensweisen von Hartmann- und Bergzebras (Equus zebra hartmannae und E. z. zebra). Z. Tierpsychol., 25, 76–88.
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Robbins, M. M., Robbins, A. M., Gerald-Steklis, N., & Steklis, H. D. (2005). Long-term dominance relationships in female mountain gorillas: strength, stability and determinants of rank. Behaviour, 142(6), 779–809.
Abstract: A common practice in studies of social animals is to rank individuals according to dominance status, which has been shown to influence access to limited resources and stability of social relationships, and may in turn correlate with reproductive success. According to the socioecological model for primates, most female dominance relationships are either nepotistic or virtually undetectable (egalitarian), with nepotistic species being philopatric, and dispersing females being egalitarian. Female mountain gorillas (Gorilla beringei beringei) disperse, and they have been characterized as being egalitarian, but previous studies have not examined their dominance relationships from a long-term perspective. We evaluated 15 matrices of displacement/supplantation interactions that spanned 30 years of observations in the Virunga Volcanoes region, and included 51 female mountain gorillas in six groups. Only 4% of displacements were directed against higher ranking females, and when matrices had less than 5% unknown dyads, linearity indices were consistently greater than 0.95. Therefore, previous results suggesting undetectable dominance relationships may have reflected an insufficient quantity of data for this species, rather than actual nonlinearity in its hierarchies. Dominance depended on age and group tenure rather than nepotism, yet some females maintained a high ranking for most of adulthood (15-25 years). Most rank shifts occurred through changes in group composition, rather than switches in established relationships. These results fit within growing evidence for linear individualistic hierarchies in some primates, often coupled with dispersal, as commonly found in ungulates. In light of these results, we propose that the dominance relationships of female mountain gorilla are best characterized as “Dispersal-Individualistic” instead of the previously suggested “Dispersal-Egalitarian”.
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Kroodsma, D. E., & Miller, E. H. (Eds.). (1996). Ecology and evolution of acoustic communication in birds. Ithaca: Cornell University Press.
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Stahl, J., Tolsma, P. H., Loonen, M. J. J. E., & Drent, R. H. (2001). Subordinates explore but dominants profit: resource competition in high Arctic barnacle goose flocks.61(1), 257–264.
Abstract: Social dominance plays an important role in assessing and obtaining access to patchy or scarce food sources in group-foraging herbivores. We investigated the foraging strategies of individuals with respect to their social position in the group in a flock of nonbreeding, moulting barnacle geese, Branta leucopsis, on high Arctic Spitsbergen. We first determined the dominance rank of individually marked birds. The dominance of an individual was best described by its age and its sex-specific body mass. Mating status explained the large variation in dominance among younger birds, as unpaired yearlings ranked lowest. In an artificially created, competitive situation, subordinate individuals occupied explorative front positions in the flock and were the first to find sites with experimentally enriched vegetation. Nevertheless, they were displaced quickly from these favourable sites by more dominant geese which were able to monopolize them. The enhanced sites were subsequently visited preferentially by individuals that succeeded in feeding there when the exclosures were first opened. Data on walking speed of foraging individuals and nearest-neighbour distances in the group suggest that subordinates try to compensate for a lower energy intake by exploring and by lengthening the foraging bout. Observations of our focal birds during the following breeding season revealed that females that returned to the study area were significantly more dominant in the previous year than those not seen in the area again.
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Tebbich, S., Taborsky, M., & Winkler, H. (1996). Social manipulation causes cooperation in keas.52(1), 1–10.
Abstract: Abstract. This study assessed whether keas,Nestor notabilis, are able to cooperate in an instrumental task. Seven birds of a captive group were tested in group situations and in dyads. At least two individuals had to manipulate an apparatus to obtain food but only one participant was rewarded. One bird had to push down a lever to enable another one to collect food from a box. The distribution of the two different roles was clearly dependent on hierarchy. The higher ranking individual always obtained the reward and each bird changed its role according to dominance status. Owing to the non-linear hierarchy in the group, each bird participating in cooperative interactions had at least one submissive partner. Therefore, in group situations the reward was distributed symmetrically and cooperation was persistent. In dyadic test situations, three individual keas aggressively manipulated their respective subordinate partners to open the apparatus. Their dominance status enabled them to force cooperation.
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Vervaecke, H., Stevens, J., Vandemoortele, H., Sigurjönsdöttir, H., & De Vries, H. (2007). Aggression and dominance in matched groups of subadult Icelandic horses (Equus caballus). J. Ethol., 25(3), 239–248.
Abstract: Abstract We studied sex differences in the nature of aggression and dominance behaviour in two newly formed groups of 1-year-old Icelandic horses. One herd contained nine geldings, the other nine mares. The groups were matched with regard to dominance-determining traits such as age, weaning age, composition of native herd, social experience, genetic origin, body condition and maternal dominance status. High-ranking individuals of both sexes were more aggressive, high-ranking males were older, and high-ranking females had a better body condition. Frequencies of aggressions were similar in both groups. The mares reacted significantly more by showing submission upon an aggression rather than by not responding or by escalating the aggression. For the geldings, this difference was not observed due to a lower tendency to submit. A linear dominance hierarchy was found in both groups. David`s scores provided additional information regarding cardinal rank distances and were used to calculate steepness of hierarchies. The female hierarchy was somewhat steeper compared to the male hierarchy, suggesting somewhat lower despotism among males. This was mainly a consequence of the lower unidirectionality in male submission. Male contests occurred predominantly between dyads at top and mid positions, suggesting a low degree of acceptance of the hierarchy.
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Stevens, J., Vervaecke, H., De Vries, H., & Van Elsacker, L. (2006). Social structures in Pan paniscus: testing the female bonding hypothesis. Primates, 47(3), 210–217.
Abstract: Abstract Based on previous research in captivity, bonobos, Pan paniscus, have been called a female-bonded species. However, genetic and behavioural data indicate that wild females migrate. Bonding between these unrelated females would then be in contradiction with socio-ecological models. It has been argued that female bonding has been overemphasized in captive bonobos. We examine patterns of proximity, grooming and support behaviour in six well established captive groups of bonobos. We find that female bonding was not a typical characteristic of all captive bonobo groups. In only two groups there was a trend for females to prefer proximity with other females over association with males. We found no evidence that following or grooming between females was more frequent than between males and unrelated females or between males. Only in coalitions, females supported each other more than male-female or male-male dyads. We also investigated five mother-son pairs. Grooming was more frequent among mothers and sons than in any other dyad, but sons did not groom their mothers more than males groomed unrelated females. Mothers groomed their sons, or provided more support to them than females groomed or supported unrelated males. Thus, while bonds between females were clearly present, intersexual relations between males and either unrelated females or their mothers are of more, or equal importance.
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Stevens, J., Vervaecke, H., de Vries, H., & Van Elsacker, L. (2005). The influence of the steepness of dominance hierarchies on reciprocity and interchange in captive groups of bonobos (Pan paniscus). Behaviour, 142(7), 941–960.
Abstract: Biological market models explain variability in reciprocity and interchange between groups. In groups with a shallow dominance gradient, grooming will be mostly exchanged for itself (i.e. exchange will occur). In groups with steep dominance hierarchies, interchange is expected: individuals will groom higher ranking individuals to get access to limited resources or commodities such as support in conflicts, and grooming will be traded for these commodities. We examine patterns of reciprocity in grooming and support, and of interchange of grooming for support or for tolerance in six captive groups of bonobos. We test whether differences between groups in patterns of reciprocity and interchange can be attributed to differences in a measure of steepness of dominance hierarchies, which is based on dyadic agonistic interactions. We found that grooming was reciprocal in some, but not all groups. Support was highly reciprocal, but this was a side effect of dominance in most groups. Interchange between grooming and support was observed in some groups. Corroborating earlier findings, this was a side effect of individuals preferring high ranking individuals as grooming and support partners, possibly because these high-ranking individuals provide more efficient support in conflicts. There was no evidence for interchange of grooming for tolerance. Variability in grooming reciprocity was explained by differences in steepness of dominance hierarchies, as predicted by the biological market models. In groups with a shallow dominance hierarchy, grooming was more reciprocal. This was not true for reciprocity in support. There was some evidence that individuals groomed dominants more frequently in groups with a steep dominance hierarchy. The variation in interchange relations between grooming and support did not depend on the steepness of dominance hierarchies. We suggest that grooming in itself is a valuable commodity in bonobos, especially under captive conditions, which can be exchanged reciprocally. Bonobos may interchange grooming for another value equivalent, with food sharing as a very likely candidate. This interchange effects seem more dependent on potential to monopolise food than on steepness of dominance hierarchies per se.
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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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Vervaecke, H., de Vries, H., & van Elsacker, L. (2000). The Pivotal Role Of Rank In Grooming And Support Behavior In A Captive Group Of Bonobos (Pan Paniscus). Behaviour, 137(11), 1463–1485.
Abstract: We investigated dyadic grooming relationships in a captive group of bonobos (Pan paniscus) and questioned what social function grooming fulfils in the 'market of services and favors'. Hereto we examined which of two theoretical models – grooming for support (Seyfarth, 1977, 1980) or grooming according to the similarity principle (de Waal & Luttrell, 1986) – best accounted for the observed grooming distribution. Similarity in traits did not correlate with increased grooming or close proximity among the individuals. Therefore, the similarity hypothesis was rejected. Seyfarth's model of rank-related grooming was largely confirmed. The animals distributed their grooming according to the rank of the receivers. We found an exchange between grooming and receipt of support. There was more grooming up than down the hierarchy. However, not all predictions about rank-related competition over grooming were confirmed. We found that dyadic grooming reciprocity indeed increased with decreasing rank distance. Yet, there was no increase of grooming within the dyad with decreasing rank distance and high ranking individuals were not competed over at the highest rates. The observed correlation between grooming and support received represents an important fit with Seyfarth's prediction, but does not allow for conclusions about underlying causal processes. Other causal explanations, besides the 'groom to receive support' hypothesis, that could explain a similar correlation are discussed.
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