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Call, J., Agnetta, B., & Tomasello, M. (2000). Cues that chimpanzees do and do not use to find hidden objects. Anim. Cogn., 3(1), 23–34.
Abstract: Chimpanzees follow conspecific and human gaze direction reliably in some situations, but very few chimpanzees reliably use gaze direction or other communicative signals to locate hidden food in the object-choice task. Three studies aimed at exploring factors that affect chimpanzee performance in this task are reported. In the first study, vocalizations and other noises facilitated the performance of some chimpanzees (only a minority). In the second study, various behavioral cues were given in which a human experimenter either touched, approached, or actually lifted and looked under the container where the food was hidden. Each of these cues led to enhanced performance for only a very few individuals. In the third study – a replication with some methodological improvements of a previous experiment – chimpanzees were confronted with two experimenters giving conflicting cues about the location of the hidden food, with one of them (the knower) having witnessed the hiding process and the other (the guesser) not. In the crucial test in which a third experimenter did the hiding, no chimpanzee found the food at above chance levels. Overall, in all three studies, by far the best performers were two individuals who had been raised in infancy by humans. It thus seems that while chimpanzees are very good at “behavior reading” of various sorts, including gaze following, they do not understand the communicative intentions (informative intentions) behind the looking and gesturing of others – with the possible exception of enculturated chimpanzees, who still do not understand the differential significance of looking and gesturing done by people who have different knowledge about states of affairs in the world.
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Call, J., Aureli, F., & de Waal, F. B. M. (2002). Postconflict third-party affiliation in stumptailed macaques. Anim. Behav., 63(2), 209–216.
Abstract: Stumptailed macaques, Macaca arctoides, are characterized by high levels of postconflict affiliative contacts between opponents. We investigated the occurrence of postconflict affiliative contacts between opponents and third parties that were not involved in the original conflict. We collected 10-min focal observations during postconflict and control periods in which we recorded all aggressive and affiliative behaviours between opponents and third parties. We distinguished three types of third parties depending on the relationship with the focal animal: own kin, opponent's kin and individuals unrelated to both opponents. We analysed the interactions with third parties separately, while distinguishing two classes of affiliative behaviours: (1) allogrooming and contact sitting and (2) sociosexual behaviours (e.g. genital inspection). The macaques showed differences between postconflict and control periods in their affiliative contacts with third parties. Aggressors received more postconflict grooming and contact sitting from their opponents' kin, received more sociosexual behaviour from their own kin and unrelated individuals, and directed more sociosexual behaviour to unrelated individuals. Victims received and directed less postconflict grooming from and towards their own kin. They received more postconflict sociosexual behaviour from all partners except their own kin and directed more sociosexual behaviour to all partners except the opponent's kin. This study establishes the occurrence of multiple postconflict triadic affiliation in stumptailed macaques, and is the first to show that victims receive contacts from third parties in a cercopithecine species, a behaviour previously described only in chimpanzees. It also highlights the importance of analysing the different affiliative behaviours separately in postconflict situations. Otherwise, many of the patterns we report, especially those involving victims, would have been missed.
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Call, J., Aureli, F., & de Waal, F. B. M. (1999). Reconciliation patterns among stumptailed macaques: a multivariate approach. Anim. Behav., 58(1), 165–172.
Abstract: This study focused on two aspects of the dynamics of reconciliation in stumptailed macaques, Macaca arctoides. First, we investigated the combined effects of multiple variables (i.e. sex, age, rank, conflict intensity, outcome, or number of participants, interopponent distance, kinship and friendship) on the occurrence of reconciliation. Second, we investigated whether opponents used different behaviour patterns in their postconflict reunions depending on the characteristics of their conflicts or their relationship with their opponents. We studied a multimale, multifemale group of 38 stumptailed macaques housed in a large outdoor compound. Three types of data were collected: (1) instantaneous scan sampling of contact sitting to infer 'friendship'; (2) ad libitum data on bared-teeth and teeth-chattering displays to infer dominance rank; (3) 10-min focal observations during postconflict (PC) and matched control (MC) periods in which we recorded interopponent distance at the beginning of the observation and all aggressive and affiliative behaviours between former opponents. Our study confirmed the high conciliatory tendency of stumptailed macaques previously reported for other groups. A stepwise logistic regression revealed that initial interopponent distance in PC, friendship and kinship were the only factors that independently contributed to explain the occurrence of reconciliation. Two main clusters of postconflict behavioural patterns emerged: allogrooming+contact sitting and sociosexual behaviours (e.g. hold-bottom). It is hypothesized that postconflict allogrooming and contact sitting may be used for the maintenance of valuable relationships, whereas sociosexual behaviours may be used more indiscriminately by any pair of opponents as a buffering mechanism to prevent immediate recurrence of aggression. Copyright 1999 The Association for the Study of Animal Behaviour.
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Call, J., & Carpenter, M. (2001). Do apes and children know what they have seen? Anim. Cogn., 3(4), 207–220.
Abstract: Chimpanzees and young children understand much about what other individuals have and have not seen. This study investigates what they understand about their own visual perception. Chimpanzees, orangutans, and 2.5-year-old children were presented with a finding game in which food or stickers were hidden in one of two or three tubes. We varied whether subjects saw the baiting of the tubes, whether subjects could see through the tubes, and whether there was a delay between baiting and presentation of the tubes to subjects. We measured not only whether subjects chose the correct tube but also, more importantly, whether they spontaneously looked into one or more of the tubes before choosing one. Most apes and children appropriately looked into the tubes before choosing one more often when they had not seen the baiting than when they had seen the baiting. In general, they used efficient search strategies more often than insufficient or excessive ones. Implications of subjects' search patterns for their understanding of seeing and knowing in the self are discussed.
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Call, J., Carpenter, M., & Tomasello, M. (2005). Copying results and copying actions in the process of social learning: chimpanzees (Pan troglodytes) and human children (Homo sapiens). Anim. Cogn., 8(3), 151–163.
Abstract: There is currently much debate about the nature of social learning in chimpanzees. The main question is whether they can copy others' actions, as opposed to reproducing the environmental effects of these actions using their own preexisting behavioral strategies. In the current study, chimpanzees (Pan troglodytes) and human children (Homo sapiens) were shown different demonstrations of how to open a tube-in both cases by a conspecific. In different experimental conditions, demonstrations consisted of (1) action only (the actions necessary to open the tube without actually opening it); (2) end state only (the open tube, without showing any actions); (3) both of these components (in a full demonstration); or (4) neither of these components (in a baseline condition). In the first three conditions subjects saw one of two different ways that the tube could open (break in middle; caps off ends). Subjects' behavior in each condition was assessed for how often they opened the tube, how often they opened it in the same location as the demonstrator, and how often they copied the demonstrator's actions or style of opening the tube. Whereas chimpanzees reproduced mainly the environmental results of the demonstrations (emulation), human children often reproduced the demonstrator's actions (imitation). Because the procedure used was similar in many ways to the procedure that Meltzoff (Dev Psych 31:1, 1995) used to study the understanding of others' unfulfilled intentions, the implications of these findings with regard to chimpanzees' understanding of others' intentions are also discussed.
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Call, J., Hare, B. A., & Tomasello, M. (1998). Chimpanzee gaze following in an object-choice task. Anim. Cogn., 1(2), 89–99.
Abstract: Many primate species reliably track and follow the visual gaze of conspecifics and humans, even to locations above and behind the subject. However, it is not clear whether primates follow a human's gaze to find hidden food under one of two containers in an object-choice task. In a series of experiments six adult female chimpanzees followed a human's gaze (head and eye direction) to a distal location in space above and behind them, and checked back to the human's face when they did not find anything interesting or unusual. This study also assessed whether these same subjects would also use the human's gaze in an object-choice task with three types of occluders: barriers, tubes, and bowls. Barriers and tubes permitted the experimenter to see their contents (i.e., food) whereas bowls did not. Chimpanzees used the human's gaze direction to choose the tube or barrier containing food but they did not use the human's gaze to decide between bowls. Our findings allowed us to discard both simple orientation and understanding seeing-knowing in others as the explanations for gaze following in chimpanzees. However, they did not allow us to conclusively choose between orientation combined with foraging tendencies and understanding seeing in others. One interesting possibility raised by these results is that studies in which the human cannot see the reward at the time of subject choice may potentially be underestimating chimpanzees' social knowledge.
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Cameron EZ, L. W. L. (2000). Individual mares bias investment in sons and daughters in relation to their condition. Anim. Behav., 60(3), 359–367.
Abstract: The Trivers-Willard hypothesis (TWH) predicts that a mother will treat a son or daughter differently depending on her ability to invest and the impact of her investment on offspring reproductive success. Although many studies have investigated the hypothesis, few have definitively supported or refuted it because of confounding factors or an inappropriate level of analysis. We studied maternal investment in sons and daughters in feral horses, Equus caballus, which meet the assumptions of the TWH with a minimum of confounding variables. Population level analyses revealed no differences in maternal behaviour towards sons and daughters. When we incorporated mare condition, we found that sons were more costly to mares in good condition, whereas daughters were more costly to mares in poor condition, although no differences in maternal behaviour were found. However, since the TWH makes predictions about individual mothers, we examined investment by mares who reared both a son and a daughter in different years of the study. Mares in good condition invested more in their sons in terms of maternal care patterns, costs to maternal body condition and costs to future reproduction. Conversely, mares in poor condition invested more in daughters. Therefore, with an appropriate level of analysis in a species in which confounding variables are minimal, the predictions of the Trivers-Willard hypothesis are supported. Copyright 2000 The Association for the Study of Animal Behaviour.
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Cameron, E. Z. (1998). Is suckling behaviour a useful predictor of milk intake? A review. Anim. Behav., 56(3), 521–532.
Abstract: In studies on mammalian parental investment, time spent suckling is often used as a predictor of the milk transferred from mother to infant. It is assumed that the rate of milk transfer is positively correlated with the time spent suckling. However, this assumption has not been tested and empirical studies show conflicting results. Nevertheless, in species in which suckling can readily be observed, time spent suckling is still used to measure milk transfer, although an increasing number of workers recognize that the measure is potentially inaccurate. A meta-analysis on studies that have correlated measures of time spent suckling with milk intake estimates based on weight gain revealed a weak positive relationship and significant heterogeneity between studies. Isotope-labelling techniques for the measurement of milk transfer independent of behaviour have been in use since the 1970s, particularly in studies of species in which suckling is difficult to observe. Only one study has attempted to correlate behavioural measures with independent isotope measures, and it found no relationship between the two measures. I suggest that researchers have avoided such a test as it is unlikely that a strong relationship will be found between milk transfer and suckling behaviour, and I discuss the various factors that confound the relationship and contribute to high heterogeneity between studies. Consequently, the assumption that milk transfer can be measured by time spent suckling has inadequate empirical foundation, and needs to be tested using isotope-labelling methods. Copyright 1998 The Association for the Study of Animal Behaviour
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Cameron, E. Z., Linklater, W. L., Stafford, K. J., & Minot, E. O. (2008). Maternal investment results in better foal condition through increased play behaviour in horses. Anim. Behav., 76(5), 1511–1518.
Abstract: Play behaviour is widespread in mammals, but benefits to play have been difficult to demonstrate. Physical training is one of the many proposed hypotheses, suggesting that males and females should play differently, that increased maternal investment should lead to increases in play, and that increases in play should result in physical advantages. In a population of feral horses, Equus caballus, males and females did not differ in their play behaviour except that males initiated more of their play bouts than females. Maternal condition influenced play behaviour only in males, with sons of mothers in good condition playing more. However, when we controlled for maternal effects by comparing a son and a daughter of the same mother, daughters played more when their mother was in poor condition and sons played more when their mother was in good condition. Mothers of foals that played more lost more condition. Therefore, the difference in play behaviour could not be explained by offspring sex or maternal condition alone, but play behaviour mirrored variation in maternal investment. In addition, those individuals that played more survived better and had better body condition as yearlings despite weaning earlier. Since increased activity has been linked to enhanced musculoskeletal development in domestic horses, we suggest that play provides a link between increased maternal investment, increased body condition and future reproductive success in feral horses, and probably in other species.
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Cameron, E. Z., Stafford, K. J., Linklater, W. L., & Veltman, C. J. (1999). Suckling behaviour does not measure milk intake in horses, Equus caballus. Anim. Behav., 57(3), 673–678.
Abstract: Studies of parental investment in mammals have frequently used suckling behaviour to estimate energy transfer from mother to offspring, and consequently to measure maternal input. Such studies assume that the more an offspring sucks, the more milk it will receive. This assumption has been questioned, and a review of the literature found little support for it. To test if suckling behaviour provided an accurate index of milk or energy intake we used a radioactive isotope technique to label the milk of thoroughbred mares and to measure milk transfer to foals. We found no significant linear relationship between usual measures of suckling behaviour and milk or energy intake. No behaviours associated with suckling nor with characteristics of mares and foals improved the relationship; only the number of butts associated with each suck episode even approached significance. If we had used suckling behaviour to test theories on differential maternal investment our conclusions would have been in error. For example, female foals tended to suck for longer than males did but there was no difference in the amount of milk transferred. Consequently, we show that measures of suckling behaviour do not adequately predict milk intake in the domestic horse and we suggest that conclusions about differential maternal investment in mammals based on suckling behaviour are likely to be in error. Copyright 1999 The Association for the Study of Animal Behaviour.
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