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Griffin, D. R. (2001). Animals know more than we used to think (Vol. 98).
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Hampton, R. R. (2001). Rhesus monkeys know when they remember. Proc. Natl. Acad. Sci. U.S.A., 98(9), 5359–5362.
Abstract: Humans are consciously aware of some memories and can make verbal reports about these memories. Other memories cannot be brought to consciousness, even though they influence behavior. This conspicuous difference in access to memories is central in taxonomies of human memory systems but has been difficult to document in animal studies, suggesting that some forms of memory may be unique to humans. Here I show that rhesus macaque monkeys can report the presence or absence of memory. Although it is probably impossible to document subjective, conscious properties of memory in nonverbal animals, this result objectively demonstrates an important functional parallel with human conscious memory. Animals able to discern the presence and absence of memory should improve accuracy if allowed to decline memory tests when they have forgotten, and should decline tests most frequently when memory is attenuated experimentally. One of two monkeys examined unequivocally met these criteria under all test conditions, whereas the second monkey met them in all but one case. Probe tests were used to rule out “cueing” by a wide variety of environmental and behavioral stimuli, leaving detection of the absence of memory per se as the most likely mechanism underlying the monkeys' abilities to selectively decline memory tests when they had forgotten.
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Hauser, M. D., Kralik, J., Botto-Mahan, C., Garrett, M., & Oser, J. (1995). Self-recognition in primates: phylogeny and the salience of species-typical features. Proc. Natl. Acad. Sci. U.S.A., 92(23), 10811–10814.
Abstract: Self-recognition has been explored in nonlinguistic organisms by recording whether individuals touch a dye-marked area on visually inaccessible parts of their face while looking in a mirror or inspect parts of their body while using the mirror's reflection. Only chimpanzees, gorillas, orangutans, and humans over the age of approximately 2 years consistently evidence self-directed mirror-guided behavior without experimenter training. To evaluate the inferred phylogenetic gap between hominoids and other animals, a modified dye-mark test was conducted with cotton-top tamarins (Saguinus oedipus), a New World monkey species. The white hair on the tamarins' head was color-dyed, thereby significantly altering a visually distinctive species-typical feature. Only individuals with dyed hair and prior mirror exposure touched their head while looking in the mirror. They looked longer in the mirror than controls, and some individuals used the mirror to observe visually inaccessible body parts. Prior failures to pass the mirror test may have been due to methodological problems, rather than to phylogenetic differences in the capacity for self-recognition. Specifically, an individual's sensitivity to experimentally modified parts of its body may depend crucially on the relative saliency of the modified part (e.g., face versus hair). Moreover, and in contrast to previous claims, we suggest that the mirror test may not be sufficient for assessing the concept of self or mental state attribution in nonlinguistic organisms.
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Proops, L., McComb, K., & Reby, D. (2009). Cross-modal individual recognition in domestic horses (Equus caballus). Proc. Natl. Acad. Sci. U.S.A., 106(3), 947–951.
Abstract: Individual recognition is considered a complex process and, although it is believed to be widespread across animal taxa, the cognitive mechanisms underlying this ability are poorly understood. An essential feature of individual recognition in humans is that it is cross-modal, allowing the matching of current sensory cues to identity with stored information about that specific individual from other modalities. Here, we use a cross-modal expectancy violation paradigm to provide a clear and systematic demonstration of cross-modal individual recognition in a nonhuman animal: the domestic horse. Subjects watched a herd member being led past them before the individual went of view, and a call from that or a different associate was played from a loudspeaker positioned close to the point of disappearance. When horses were shown one associate and then the call of a different associate was played, they responded more quickly and looked significantly longer in the direction of the call than when the call matched the herd member just seen, an indication that the incongruent combination violated their expectations. Thus, horses appear to possess a cross-modal representation of known individuals containing unique auditory and visual/olfactory information. Our paradigm could provide a powerful way to study individual recognition across a wide range of species.
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Hollenhorst, H., Weil, S., & Krueger, K. (2015). Innovative behavour in horses (Equus caballus). In , & K. Krueger (Ed.), Proceedings of the 3. International Equine Science Meeting. Proc. 3. Int. Equine. Sci. Mtg. Wald: Xenophon Publishing.
Abstract: Contrary to the widely-spread assumption that horses just have restricted cognitive capacities and are not very flexible in their behaviors, we showed that horses display innovative behavior and even make use of tools (Krueger 2015, Krueger et al. 2015). These findings derive from a database (http://innovative-behaviour.org/) the Equine behavior team managed in the past two years. Some horses did not only show single innovations, but several different innovations. The number of innovations per individual varied from 1 to 10. 20 % of all innovative horses in the database showed more than one innovation. These individuals can be called the ‘true innovators’. Moreover innovations were dependent on age. Young horses were more innovative than older ones, whereby horses at the age of five to nine years were the most innovative. When considering the housing system innovative horses in a single housing (inside box, outside box, paddock box) had a slight majority towards horses in group housing (open stable, active stable, pasture day and night). But given the fact that ratings on housing system frequencies state 95% of the horses to be kept in individual housing, innovations in individual housing are rare. Nevertheless, horses kept in inside boxes without a window, opened doors more often than all other horses. Aside from this effect, housing systems did not trigger the frequency of innovative behavior. Innovations for gaining freedom and innovations in general were widespread among horses with daily access to pasture and daily contact with conspecifics. Innovations for gaining food were not more likely to occur in horses that were fed little amounts of roughage. In conclusion, the housing of horses does not seem to be the primary catalyst for developing innovative behavior in horses. What makes a “true innovator” in horses, in addition to age, remains to be seen.
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Hori, Y., Takimoto, A., & Fujita, K. (2012). Are there breed difference in referential behavior in horses (Equus caballus)? In K. Krueger (Ed.), Proceedings of the 2. International Equine Science Meeting (Vol. in press). Wald: Xenophon Publishing.
Abstract: Domesticated animals are characterized by variability of breeds. There is a great diversity in body size and/or coat color between different breeds. However, there are few scientific researches about difference in cognition and behavior between breeds. Comparison of behavior between breeds may be useful for the study of genetics behind the diversity of cognition and behavior. In the present study, we investigated behavioral differences between horse breeds. We tested two different breeds which have different histories, thoroughbreds and creoles. Thoroughbreds are racing horses which have been exposed to strict selection toward racing performance for about 300 years. Creoles are descendents of horses which were brought to South America by Spanish people in 15th century and used by native cowboys for riding. We compared the behavior in a difficult situation by using an “unsolvable task”. The experimenter put a food reward into a transparent box and closed it firmly so that horses could not take the reward. We compared the referential behavior (gazing behavior toward the experimenter) between thoroughbreds and creoles. We analyzed referential behavior by using generalized linear models (GLM) and model selection by Akaike’s information criterion (AIC). There were no effect of breed in the frequency and the duration of the referential behavior. But the latency before looking at the experimenter tended to be shorter in thoroughbreds than in creoles. This result suggests that there may be breed differences in horses’ social cognition and behavior. However, the effect of sex was also seen. Furthermore, we could not exclude the environmental effect (e. g. feeding environments, trainings) in this study. So we cannot explain the variation in referential behavior by breed effect only. We need to replicate the result by controlling environmental effects.
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Dalla Costa, E., Rabolini, A., Scelsa, A., Canali, E., & Minero, M. (2012). A study on inter-observer reliability of castration pain assessment in horses. In K. Krueger, & (Ed.), Proceedings of the 2. International Equine Science Meeting (Vol. in press). Wald: Xenophon Publishing.
Abstract: Pain evaluation is a key issue for equine welfare and it is often cause of concern because it is difficult to determine its intensity and duration. This is essential when different people are looking after the animals and they need to decide when or not giving analgesics to guarantee the welfare of the subject. The most widely used technique to determine pain in horses is identifying pain related behaviors. The aim of this study was to determine inter-observer reliability of two different assessors evaluating pain related behaviors in horses undergoing castration. 8 stallions of different breed, aged between 2 and 4 years, were included in the study. All the subjects underwent routine castration (closed technique in general anesthesia). The subjects were placed in an observation box for 5 days and their behavior was recorded for 15 minutes before the surgery and 4, 8, 16, 24 and 40 hours after intervention. Two blind observers, using a given ethogram of horse pain related behaviors modified from literature (for a review Ashley, 2005), analyzed horses behavior at each interval. Descriptive statistics and K Kendall test were performed. Observers agreed significantly assessing agitation, reluctance to move, kicking the abdomen, lethargy, rolling, attention and curiosity (P<0.05), however agreement was low for head movements, stretching, flank watching, lowered head carriage, weight shifting, abnormal movement, fixed stare. Our results show that assessing pain in horses should be a cause of concern, because different pain related behaviors are difficult to identify and to have agreement between two observers. Training of care takers of horses on identification of specific behaviors is needed to standardize pain assessment. Acknowledgements: The authors wish to thank the EU VII Framework programme (FP7-KBBE-2010-4) for financing the Animal Welfare Indicators (AWIN) project and for providing funds for Emanuela Dalla Costa and Michela Minero to present this paper.
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Bonnie, K. E., & de Waal, F. B. M. (2006). Affiliation promotes the transmission of a social custom: handclasp grooming among captive chimpanzees. Primates, 47(1), 27–34.
Abstract: Handclasp grooming is a unique social custom, known to occur regularly among some, but not all populations of chimpanzees (Pan troglodytes). As with other cultural behaviors, it is assumed that this distinctive grooming posture is learned socially by one individual from another. However, statistical comparisons among factors thought to influence how a behavior spreads within a group have never, to our knowledge, been conducted. In the present study, the origination and spread of handclasp grooming in a group of captive chimpanzees was followed throughout more than 1,500 h of observation over a period of 12 years. We report on the frequency, bout duration, and number and demography of performers throughout the study period, and compare these findings to those reported for wild populations. We predicted that dyads with strong affiliative ties, measured by time spent in proximity to and grooming one another, were likely to develop a handclasp grooming partnership during the study period. A quadratic assignment procedure was used to compare correlations among observed frequencies of grooming and proximity with handclasp grooming in all possible dyads within the group. As predicted, the formation of new handclasp grooming dyads was positively correlated with the rate of overall grooming and proximity within a dyad. In addition, in nearly all dyads formed, at least one individual had been previously observed to handclasp groom. We concluded that affiliation and individual experience determines the transmission of handclasp grooming among captive chimpanzees.
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Brosnan, S. F., & de Waal, F. B. M. (2005). Responses to a simple barter task in chimpanzees, Pan troglodytes. Primates, 46(3), 173–182.
Abstract: Chimpanzees (Pan troglodytes) frequently participate in social exchange involving multiple goods and services of variable value, yet they have not been tested in a formalized situation to see whether they can barter using multiple tokens and rewards. We set up a simple barter economy with two tokens and two associated rewards and tested chimpanzees on their ability to obtain rewards by returning the matching token in situations in which their access to tokens was unlimited or limited. Chimpanzees easily learned to associate value with the tokens, as expected, and did barter, but followed a simple strategy of favoring the higher-value token, regardless of the reward proffered, instead of a more complex but more effective strategy of returning the token that matched the reward. This response is similar to that shown by capuchin monkeys in our previous study. We speculate that this response, while not ideal, may be sufficient to allow for stability of the social exchange system in these primates, and that the importance of social barter to both species may have led to this convergence of strategies.
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Assersohn, C., Whiten, A., Kiwede, Z. T., Tinka, J., & Karamagi, J. (2004). Use of leaves to inspect ectoparasites in wild chimpanzees: a third cultural variant? Primates, 45(4), 255–258.
Abstract: We report 26 cases of using leaves as tools with which wild chimpanzees (Pan troglodytes schweinfurthii) in the Sonso community, Budongo Forest, Uganda, appeared to inspect objects removed during grooming. Careful removal of potential ectoparasites and delicate lip or manual placement on leaves followed by intense visual examination characterised this behaviour. It appears to be done to judge whether either ingestion or discarding is most appropriate, the former occurring in most cases. This behaviour may represent a third variant of ectoparasite handling, different from those described at Tai and Gombe, yet sharing features with the latter. These two East African techniques may thus have evolved from leaf grooming.
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