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Powell, D. (2008). Female–female competition or male mate choice? Patterns of courtship and breeding behavior among feral horses (Equus caballus) on Assateague Island. J. Ethol., 26(1), 137-144.
Abstract: Previous research on the Assateague horses found that high-ranking females had more surviving offspring than low-ranking females. Variance in reproductive success may be the result of a variety of proximate processes that affect sexual behavior such as mate choice and mate competition. A study was done to determine whether patterns of courtship, social, and sexual behavior could be identified that would suggest mate choice and/or mate competition. Behavioral data were collected from approximately 40 sexually mature mares living in harem bands. Stallions showed more interest in the eliminations of dominant mares than subordinate mares. Males also engaged in significantly more high-intensity (e.g., mounts and copulations) sexual behavior with dominant mares than subordinate mares, and there was a trend for males to engage in more low-intensity (e.g., flehmen and ano-genital sniffing) sexual behavior with dominant mares than subordinate mares. There was no effect of mare rank on spatial relationships with the stallion; however, dominant mares did attempt to restrict reproductive access to the stallion by harassing and disrupting copulations. Higher foaling rates among dominant mares on Assateague Island could therefore be the result of rank-related mate choice by stallions and direct female competition for mating opportunities.
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Prud`homme, J., & Chapais, B. (1996). Development of intervention behavior in Japanese macaques: Testing the targeting hypothesis. Int. J. Primatol., 17(3), 429–443.
Abstract: Matrilineal dominance systems, which characterize several species of cercopithecines, are determined largely by the patterning of third-party aggressive interventions in conflicts. Although the role of interventions in structuring rank relations has received much attention, very few studies have dealt specifically with the development of intervention behavior. In other words,most studies have focused on the interventions received and their effect on the recipients rather than on the interventions performed and the goals of the interveners. We analyzed the intervention behavior of 10 juvenile females in a colony of 40 Japanese macaques (Macaca fuscata)housed at the University of Montreal Laboratory of Behavioral Primatology. The analysis of 749 interventions performed by the juveniles over their first 4 years and 2425 interventions received over the same period reveals that (1) juvenile females intervened selectively against females ranking below their mother, (2) they began to intervene at about the same time that they began to challenge the latter females in dyadic contests, (3) they sided with females as well as with males against these females, (4) juvenile interveners incurred little risks in terms of aggressive retaliation from their targets, (5) they derived immediate benefits in terms of conflicts won over stronger targets, (6) interventions often did not take place when the possible recipients needed support, and (7) interveners did not conform to a pattern of mutually preferential support. These results support the view that interventions by juveniles are selfish (vs altruistic) and constitute a low-cost and effective means to target and to outrank prospectively subordinate females.
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Ramos-Fernández, G., Boyer, D., Aureli, F., & Vick, L. (2009). Association networks in spider monkeys (Ateles geoffroyi). Behav. Ecol. Sociobiol., 63(7), 999-1013.
Abstract: We use two novel techniques to analyze association patterns in a group of wild spider monkeys (Ateles geoffroyi) studied continuously for 8 years. Permutation tests identified association rates higher or lower than chance expectation, indicating active processes of companionship and avoidance as opposed to passive aggregation. Network graphs represented individual adults as nodes and their association rates as weighted edges. Strength and eigenvector centrality (a measure of how strongly linked an individual is to other strongly linked individuals) were used to quantify the particular role of individuals in determining the network's structure. Female–female dyads showed higher association rates than any other type of dyad, but permutation tests revealed that these associations cannot be distinguished from random aggregation. Females formed tightly linked clusters that were stable over time, with the exception of immigrant females who showed little association with any adult in the group. Eigenvector centrality was higher for females than for males. Adult males were associated mostly among them, and although their strength of association with others was lower than that of females, their association rates revealed a process of active companionship. Female–male bonds were weaker than those between same-sex pairs, with the exception of those involving young male adults, who by virtue of their strong connections both with female and male adults, appear as temporary brokers between the female and male clusters of the network. This analytical framework can serve to develop a more complete explanation of social structure in species with high levels of fission–fusion dynamics.
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Ransom, J. I., Cade, B. S., & Hobbs, N. T. (2010). Influences of immunocontraception on time budgets, social behavior, and body condition in feral horses. Appl. Anim. Behav. Sci., 124(1-2), 51–60.
Abstract: Managers concerned with shrinking habitats and limited resources for wildlife seek effective tools for limiting population growth in some species. Fertility control is one such tool, yet little is known about its impacts on the behavioral ecology of wild, free-roaming animals. We investigated influences of the immunocontraceptive porcine zona pellucida (PZP) on individual and social behavior in bands of feral horses (Equus caballus) in three discrete populations and used 14 hierarchical mixed effect models to gain insight into the influences of PZP treatment on feral horse behavior. A model of body condition was the strongest predictor of feeding, resting, maintenance, and social behaviors, with treated females allocating their time similarly to control females. Time spent feeding declined 11.4% from low condition to high condition females (F1,154 = 26.427, P < 0.001) and was partially reciprocated by a 6.0% increase in resting (F1,154 = 7.629, P = 0.006), 0.9% increase in maintenance (F1,154 = 7.028, P = 0.009), and 1.8% increase in social behavior (F1,154 = 15.064, P < 0.001). There was no difference detected in body condition of treated versus control females (F1,154 = 0.033, P = 0.856), but females with a dependent foal had lower body condition than those without a foal (F1,154 = 4.512, P = 0.038). Herding behavior was best explained by a model of treatment and the interaction of band fidelity and foal presence (AICc weight = 0.660) which estimated no difference in rate of herding behavior directed toward control versus treated females (F1,102 = 0.196, P = 0.659), but resident females without a dependent foal were herded 50.9% more than resident females with a foal (F3,102 = 8.269, P < 0.001). Treated females received 54.5% more reproductive behaviors from stallions than control mares (F1,105 = 5.155, P = 0.025), with the model containing only treatment being the most-supported (AICc weight = 0.530). Treated and control females received harem-tending behaviors from stallions equally (F1,105 = 0.001, P = 0.969) and agonistic behaviors from stallions equally (F1,105 < 0.001, P = 0.986). Direct effects of PZP treatment on the behavior of feral horses appear to be limited primarily to reproductive behaviors and most other differences detected were attributed to the effects of body condition, band fidelity, or foal presence. PZP is a promising alternative to traditional hormone-based contraceptives and appears to contribute few short-term behavioral modifications in feral horses.
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Reddon, A. R., & Hurd, P. L. (2009). Individual differences in cerebral lateralization are associated with shy-bold variation in the convict cichlid. Anim. Behav., 77(1), 189–193.
Abstract: Cerebral lateralization, the preferential use of one hemisphere of the brain to perform certain cognitive functions, is a widespread and evolutionarily ancient adaptation. Lateralization appears to enhance cognitive capacity, yet substantial individual variation in the strength cerebral lateralization is apparent in all species studied so far. It is puzzling that cerebral lateralization, a seemingly advantageous trait, has not been driven to fixation. It has been suggested that variation in lateralization may be linked to individual variation in behaviour, which itself may be subject to disruptive selection. We examined the relation between cerebral lateralization and individual variation in boldness in the convict cichlid, Archocentrus nigrofasciatus. We show that convict cichlids that are more strongly lateralized when exploring a familiar environment, but not a novel one, are quicker to emerge from a shelter in a test for boldness. The possibility that cerebral lateralization is linked to life history strategy is discussed.
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Silk, J. B. (2002). Kin Selection in Primate Groups. Int. J. Primatol., 23(4), 849–875.
Abstract: Altruism poses a problem for evolutionary biologists because natural selection is not expected to favor behaviors that are beneficial to recipients, but costly to actors. The theory of kin selection, first articulated by Hamilton (1964), provides a solution to the problem. Hamilton's well-known rule (br > c) provides a simple algorithm for the evolution of altruism via kin selection. Because kin recognition is a crucial requirement of kin selection, it is important to know whether and how primates can recognize their relatives. While conventional wisdom has been that primates can recognize maternal kin, but not paternal kin, this view is being challenged by new findings. The ability to recognize kin implies that kin selection may shape altruistic behavior in primate groups. I focus on two cases in which kin selection is tightly woven into the fabric of social life. For female baboons, macaques, and vervets maternal kinship is an important axis of social networks, coalitionary activity, and dominance relationships. Detailed studies of the patterning of altruistic interactions within these species illustrate the extent and limits of nepotism in their social lives. Carefully integrated analyses of behavior, demography, and genetics among red howlers provide an independent example of how kin selection shapes social organization and behavior. In red howlers, kin bonds shape the life histories and reproductive performance of both males and female. The two cases demonstrate that kin selection can be a powerful source of altruistic activity within primate groups. However, to fully assess the role of kin selection in primate groups, we need more information about the effects of kinship on the patterning of behavior across the Primates and accurate information about paternal kin relationships.
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Slobodchikoff, C., Paseka, A., & Verdolin, J. (2009). Prairie dog alarm calls encode labels about predator colors. Anim. Cogn., 12(3), 435–439.
Abstract: Some animals have the cognitive capacity to differentiate between different species of predators and generate different alarm calls in response. However, the presence of any addition information that might be encoded into alarm calls has been largely unexplored. In the present study, three similar-sized human females walked through a Gunnison’s prairie dog ( Cynomys gunnisoni ) colony wearing each of three different-colored shirts: blue, green, and yellow. We recorded the alarm calls and used discriminant function analysis to assess whether the calls for the different-colored shirts were significantly different. The results showed that the alarm calls for the blue and the yellow shirts were significantly different, but the green shirt calls were not significantly different from the calls for the yellow shirt. The colors that were detected, with corresponding encoding into alarm calls, reflect the visual perceptual abilities of the prairie dogs. This study suggests that prairie dogs are able to incorporate labels about the individual characteristics of predators into their alarm calls, and that the complexity of information contained in animal alarm calls may be greater than has been previously believed.
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Truppa, V., Garofoli, D., Castorina, G., Piano Mortari, E., Natale, F., & Visalberghi, E. (2010). Identity concept learning in matching-to-sample tasks by tufted capuchin monkeys (Cebus apella). Anim. Cogn., 13(6), 835-848.
Abstract: The abstract concept of equivalence is considered one of the bases of higher-order cognition, and it has been the subject of considerable research in comparative cognition. This study examined the conditions under which tufted capuchin monkeys (Cebus apella) are able to acquire an identity concept. Six capuchin monkeys were trained to solve a visual matching-to-sample (MTS) task on the basis of perceptual identity. The acquisition of the identity rule was inferred from the subject’s ability to solve transfer tests with novel stimuli. We evaluated the ability of the capuchin monkeys to match the shape of novel stimuli after training with both several small stimulus sets (Experiment 1) and a large stimulus set (Experiment 2). Moreover, we examined the ability of capuchins to transfer the concept to novel visual dimensions, such as colour and size and to transfer to novel spatial arrangements of the stimuli (Experiment 2). We demonstrated that the ability of capuchins to match novel stimuli was improved by increasing the number of stimuli used during training (Experiments 1 and 2) and that after a widely applicable identity concept based on the stimulus shape was acquired, the capuchins were able to match stimuli according to an identity rule based on both the colour and size of the stimuli and when the spatial arrangement of the stimuli was varied (Experiment 2). This study is the first to demonstrate that the size of the training set affects the acquisition of an abstract identity concept in an MTS task in non-human primates.
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van Schaik, C. P. (2010). Social learning and culture in animals. In P. Kappeler (Ed.), Animal Behaviour: Evolution and Mechanisms (pp. 623–653). Springer Berlin Heidelberg.
Abstract: Most animals must learn some of the behaviours in their repertoire, and some must learn most. Although learning is often thought of as an individual exercise, in nature much learning is social, i.e. under the influence of conspecifics. Social learners acquire novel information or skills faster and at lower cost, but risk learning false information or useless skills. Social learning can be divided into learning from social information and learning through social interaction. Different species have different mechanisms of learning from social information, ranging from selective attention to the environment due to the presence of others to copying of complete motor sequences. In vertical (or oblique) social learning, naïve individuals often learn skills or knowledge from parents (or other adults), whereas horizontal social learning is from peers, either immatures or adults, and more often concerns eavesdropping and public information use. Because vertical social learning is often adaptive, maturing individuals often have a preference for it over individual exploration. The more cognitively demanding social learning abilities probably evolved in this context, in lineages where offspring show long association with parents and niches are complex. Because horizontal learning can be maladaptive, especially when perishable information has become outdated, animals must decide when to deploy social learning. Social learning of novel skills can lead to distinct traditions or cultures when the innovations are sufficiently rare and effectively transmitted socially. Animal cultures may be common but to date taxonomic coverage is insufficient to know how common. Cultural evolution is potentially powerful, but largely confined to humans, for reasons currently unknown. A general theory of culture is therefore badly needed.
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VanDierendonck, M. C. (2006). Diff erences in social behaviour between late pregnant, post-partum and barren mares in a herd of Icelandic horses (Vol. Chapter 5). Ph.D. thesis, , Universiteit Utrecht.
Abstract: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.
Keywords: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.
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