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20 - Primate Empathy

A Flexible and Multi-Componential Phenomenon

Published online by Cambridge University Press:  28 July 2022

Bennett L. Schwartz
Florida International University
Michael J. Beran
Georgia State University
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Empathy, the capacity to share and understand others’ states, is crucial for facilitating enduring social relationships and managing ingroup and outgroup dynamics. Despite being at the center of much scrutiny and debate in human research, the evolutionary foundations of empathy remain relatively opaque. Moreover, inconsistencies remain regarding definitions and theoretical models, leading to discrepancies in how to systematically represent and address empathy and understand its evolutionary basis. As a complex, multidimensional phenomenon, certain components of empathy are likely to be evolutionarily ancient whereas others may be more derived. As our closest living relatives, nonhuman primates provide an opportunity to explore the evolutionary origins of empathy and its subcomponents. Due to the rich diversity of primate societies, we can comparatively study evidence of affective responding and empathic behavior within the context of different social dynamics and organization. Although studies have been conducted on individual primate species, especially the great apes, direct species comparisons are rare. Here we examine the literature investigating evidence for empathy among primates focusing on its underlying affective and cognitive components. In reviewing the literature, we also highlight species that need more coverage to enhance our overall understanding of how empathy has evolved within the primate order.

Publisher: Cambridge University Press
Print publication year: 2022

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Adriaense, J. E. C., Koski, S. E., Huber, L., & Lamm, C. (2020). Challenges in the comparative study of empathy and related phenomena in animals. Neuroscience & Biobehavioral Reviews, 112, 6282.Google Scholar
Amici, F., Aureli, F., & Call, J. (2014). Response facilitation in the four great apes: Is there a role for empathy? Primates, 55, 113118.Google Scholar
Amici, F., Aureli, F., Visalberghi, E., & Call, J. (2009). Spider monkeys (Ateles geoffroyi) and capuchin monkeys (Cebus apella) follow gaze around barriers: Evidence for perspective taking? Journal of Comparative Psychology, 123, 368374.Google Scholar
Amici, F., Visalberghi, E., & Call, J. (2014). Lack of prosociality in great apes, capuchin monkeys and spider monkeys: Convergent evidence from two different food distribution tasks. Proceedings of the Royal Society B: Biological Sciences, 281, 20141699.Google Scholar
Arre, A. M., Clark, C. S., & Santos, L. R. (2020). Do young rhesus macaques know what others see? A comparative developmental perspective. American Journal of Primatology, 82, e23054.Google Scholar
Aureli, F., Das, M., Verleur, D., & van Hooff, J. A. R. A. M. (1994). Postconflict social interactions among barbary macaques (Macaca sylvanus). International Journal of Primatology, 15, 471485.Google Scholar
Baenninger, R. (1997). On yawning and its functions. Psychonomic Bulletin & Review, 4, 198207.Google Scholar
Baker, M. A., Cronin, M. J., & Mountjoy, D. G. (1976). Variability of skin temperature in the waking monkey. The American Journal of Physiology, 230, 449455.Google Scholar
Batson, C. D. (2011). Empathy-induced altruism: Friend or foe of the common good? In Forsyth, D. R. & Hoyt, C. L. (Eds.), For the greater good of all: Perspectives on individualism, society, and leadership (pp. 2947). Palgrave Macmillan US.Google Scholar
Batson, C. D., Early, S., & Salvarani, G. (1997). Perspective taking: Imagining how another feels versus imaging how you would feel. Personality and Social Psychology Bulletin, 23, 751758.Google Scholar
Berthier, J. M., & Semple, S. (2018). Observing grooming promotes affiliation in Barbary macaques. Proceedings of the Royal Society B, 285, 20181964.Google Scholar
Bezerra, B. M., Keasey, M. P., Schiel, N., & da Silva Souto, A. (2014). Responses towards a dying adult group member in a wild New World monkey. Primates, 55, 185188.Google Scholar
Boesch, C. (1992). New elements of a theory of mind in wild chimpanzees. Behavioral and Brain Sciences, 15, 149150.Google Scholar
Bradley, M. M., Miccoli, L., Escrig, M. A., & Lang, P. J. (2008). The pupil as a measure of emotional arousal and autonomic activation. Psychophysiology, 45, 602607.Google Scholar
Bray, J., Krupenye, C., & Hare, B. (2014). Ring-tailed lemurs (Lemur catta) exploit information about what others can see but not what they can hear. Animal Cognition, 17, 735744.Google Scholar
Buchanan, T. W., Bagley, S. L., Stansfield, R. B., & Preston, S. D. (2012). The empathic, physiological resonance of stress. Social Neuroscience, 7, 191201.Google Scholar
Burkart, J. M., & Heschl, A. (2007). Understanding visual access in common marmosets, Callithrix jacchus: Perspective taking or behaviour reading? Animal Behaviour, 73, 457469.Google Scholar
Burkart, J. M., & van Schaik, C. (2013). Group service in macaques (Macaca fuscata), capuchins (Cebus apella) and marmosets (Callithrix jacchus): A comparative approach to identifying proactive prosocial motivations. Journal of Comparative Psychology, 127, 212225.CrossRefGoogle ScholarPubMed
Burkett, J. P., Andari, E., Johnson, Z. V., Curry, D. C., de Waal, F. B. M., & Young, L. J. (2016). Oxytocin-dependent consolation behavior in rodents. Science, 351, 375378.CrossRefGoogle ScholarPubMed
Butovskaya, M. L., Meishvili, N. V., & Chalyan, V. G. (2015). Redirection of aggression and consolation in hamadryas baboons. Neuroscience and Behavioral Physiology, 45, 417422.Google Scholar
Call, J., Aureli, F., & de Waal, F. B. M. (2002). Postconflict third-party affiliation in stumptailed macaques. Animal Behaviour, 63, 209216.Google Scholar
Call, J., & Tomasello, M. (2008). Does the chimpanzee have a theory of mind? 30 years later. Trends in Cognitive Sciences, 12, 187192.CrossRefGoogle ScholarPubMed
Campbell, L. A. D. (2019). Fostering of a wild, injured, juvenile by a neighbouring group: Implications for rehabilitation and release of Barbary macaques confiscated from illegal trade. Primates, 60, 339345.CrossRefGoogle ScholarPubMed
Campbell, L. A. D., Tkaczynski, P. J., Mouna, M., Qarro, M., Waterman, J., & Majolo, B. (2016). Behavioral responses to injury and death in wild Barbary macaques (Macaca sylvanus). Primates; Journal of Primatology, 57, 309315.Google Scholar
Campbell, M. W., & de Waal, F. B. M. (2011). Ingroup-outgroup bias in contagious yawning by chimpanzees supports link to empathy. PLoS ONE, 6, e18283.CrossRefGoogle ScholarPubMed
Canteloup, C., Piraux, E., Poulin, N., & Meunier, H. (2016). Do Tonkean macaques (Macaca tonkeana) perceive what conspecifics do and do not see? PeerJ, 4, e1693.Google Scholar
Chapman, C. A., & Chapman, L. J. (1987). Social responses to the traumatic injury of a juvenile spider monkey (Ateles geoffroyi). Primates, 28, 271275.Google Scholar
Chartrand, T. L., & Bargh, J. A. (1999). The chameleon effect: The perception–behavior link and social interaction. Journal of Personality and Social Psychology, 76, 893910.Google Scholar
Chotard, H., Ioannou, S., & Davila‐Ross, M. (2018). Infrared thermal imaging: Positive and negative emotions modify the skin temperatures of monkey and ape faces. American Journal of Primatology, 80, e22863.CrossRefGoogle ScholarPubMed
Claidière, N., Whiten, A., Mareno, M. C., Messer, E. J. E., Brosnan, S. F., Hopper, L. M., Lambeth, S. P., Schapiro, S. J., & McGuigan, N. (2015). Selective and contagious prosocial resource donation in capuchin monkeys, chimpanzees and humans. Scientific Reports, 5, 7631.Google Scholar
Clay, Z., & de Waal, F. B. M. (2013). Bonobos respond to distress in others: Consolation across the age spectrum. PLoS ONE, 8, e55206.Google Scholar
Clay, Z., Webb, C. E, Romero, T., & de Waal, F. B. M. (2021). Comparative perspectives of empathy development: Insights from chimpanzees and bonobos. In Dukes, D., Samson, A.C., Walle, E. A. (Eds.), The Oxford handbook of emotional development. Oxford University Press.Google Scholar
Cordoni, G., Palagi, E., & Tarli, S. B. (2006). Reconciliation and consolation in captive western gorillas. International Journal of Primatology, 27, 13651382.CrossRefGoogle Scholar
Cronin, K. A., Schroeder, K. K. E., Rothwell, E. S., Silk, J. B., & Snowdon, C. T. (2009). Cooperatively breeding cottontop tamarins (Saguinus oedipus) do not donate rewards to their long-term mates. Journal of Comparative Psychology, 123, 231241.Google Scholar
Cronin, K. A., van Leeuwen, E. J. C., Vreeman, V., & Haun, D. B. M. (2014). Population-level variability in the social climates of four chimpanzee societies. Evolution and Human Behavior, 35, 389396.Google Scholar
Cuff, B. M. P., Brown, S. J., Taylor, L., & Howat, D. J. (2016). Empathy: A review of the concept. Emotion Review, 8, 144153.Google Scholar
Davila-Ross, M., Menzler, S., & Zimmermann, E. (2008). Rapid facial mimicry in orangutan play. Biology Letters, 4, 2730.Google Scholar
Davis, J., Davis, , Redshaw, J., Suddendorf, T., Nielsen, M., Kennedy-Costantini, S., Oostenbroek, J., & Slaughter, V. (2021). Does neonatal imitation exist? Insights from a meta-analysis of 336 effect sizes. Perspectives on Psychological Science, 16, 13731397.Google Scholar
de Marco, A., Cozzolino, R., Dessì‐Fulgheri, F., & Thierry, B. (2011). Collective arousal when reuniting after temporary separation in Tonkean macaques. American Journal of Physical Anthropology, 146, 457464.CrossRefGoogle ScholarPubMed
de Waal, F. B. M. (1988). The communicative repertoire of captive bonobos (Pan paniscus), compared to that of chimpanzees. Behaviour, 106, 183251.Google Scholar
de Waal, F. B. M. (2007). The “Russian doll” model of empathy and imitation. In Bråten, S. (Ed.), Advances in consciousness research: Vol. 68. On being moved: From mirror neurons to empathy (pp. 4969). John Benjamins Publishing Company.Google Scholar
de Waal, F. B. M. (2008). Putting the altruism back into altruism: The evolution of empathy. Annual Review of Psychology, 59, 279300.Google Scholar
de Waal, F. B. M. (2019). Mama’s last hug: Animal emotions and what they tell us about ourselves. W. W. Norton & Company.Google Scholar
de Waal, F. B. M., & Aureli, F. (1996). Consolation, reconciliation, and a possible cognitive difference between macaques and chimpanzees. In Russon, A. E., Bard, K. A., & Parker, S. T. (Eds.), Reaching into thought: The minds of the great apes (pp. 80110). Cambridge University Press.Google Scholar
de Waal, F. B. M., & Preston, S. D. (2017). Mammalian empathy: Behavioural manifestations and neural basis. Nature Reviews Neuroscience, 18, 498509.CrossRefGoogle ScholarPubMed
de Waal, F. B. M., & van Roosmalen, A. (1979). Reconciliation and consolation among chimpanzees. Behavioral Ecology and Sociobiology, 5, 5566.CrossRefGoogle Scholar
Decety, J., & Svetlova, M. (2012). Putting together phylogenetic and ontogenetic perspectives on empathy. Developmental Cognitive Neuroscience, 2, 124.Google Scholar
Demuru, E., & Palagi, E. (2012). In bonobos yawn contagion is higher among kin and friends. PLoS ONE, 7, e49613.Google Scholar
Dettmer, A. M., Kaburu, S. S. K., Simpson, E. A., Paukner, A., Sclafani, V., Byers, K. L., Murphy, A. M., Miller, M., Marquez, N., Miller, G. M., Suomi, S. J., & Ferrari, P. F. (2016). Neonatal face-to-face interactions promote later social behaviour in infant rhesus monkeys. Nature Communications, 7, 11940.Google Scholar
Dezecache, G., Zuberbühler, K., Davila-Ross, M., & Dahl, C. D. (2017). Skin temperature changes in wild chimpanzees upon hearing vocalizations of conspecifics. Royal Society Open Science, 4, 160816.Google Scholar
Dimberg, U., & Thunberg, M. (2012). Empathy, emotional contagion, and rapid facial reactions to angry and happy facial expressions. PsyCh Journal, 1, 118127.Google Scholar
Dindo, M., Whiten, A., & de Waal, F. B. M. (2009). In-group conformity sustains different foraging traditions in capuchin monkeys (Cebus apella). PLoS ONE, 4, e7858.CrossRefGoogle ScholarPubMed
Ebisch, S. J., Aureli, T., Bafunno, D., Cardone, D., Romani, G. L., & Merla, A. (2012). Mother and child in synchrony: Thermal facial imprints of autonomic contagion. Biological Psychology, 89, 123129.Google Scholar
Eisenberg, N., Guthrie, I. K., Murphy, B. C., Shepard, S. A., Cumberland, A., & Carlo, G. (1999). Consistency and development of prosocial dispositions: A longitudinal study. Child Development, 70, 13601372.CrossRefGoogle ScholarPubMed
Feneran, A. N., O’Donnell, R., Press, A., Yosipovitch, G., Cline, M., Dugan, G., Papoiu, A. D. P., Nattkemper, L. A., Chan, Y. H., & Shively, C. A. (2013). Monkey see, monkey do: Contagious itch in nonhuman primates. Acta Dermato-Venereologica, 93, 2729.Google Scholar
Ferrari, P. F., Visalberghi, E., Paukner, A., Fogassi, L., Ruggiero, A., & Suomi, S. J. (2006). Neonatal imitation in rhesus macaques. PLOS Biology, 4, e302.Google Scholar
Fraser, O. N., & Bugnyar, T. (2010). Do ravens show consolation? Responses to distressed others. PLoS ONE, 5, e10605.Google Scholar
Fraser, O. N., Koski, S. E., Wittig, R. M., & Aureli, F. (2009). Why are bystanders friendly to recipients of aggression? Communicative & Integrative Biology, 2, 285291.Google Scholar
Fraser, O. N., Stahl, D., & Aureli, F. (2008). Stress reduction through consolation in chimpanzees. Proceedings of the National Academy of Sciences, 105, 85578562.Google Scholar
Fuhrmann, D., Ravignani, A., Marshall-Pescini, S., & Whiten, A. (2014). Synchrony and motor mimicking in chimpanzee observational learning. Scientific Reports, 4, 17.CrossRefGoogle ScholarPubMed
Fujisawa, K. K., Kutsukake, N., & Hasegawa, T. (2006). Peacemaking and consolation in Japanese preschoolers witnessing peer aggression. Journal of Comparative Psychology, 120, 4857.Google Scholar
Hare, B., Call, J., Agnetta, B., & Tomasello, M. (2000). Chimpanzees know what conspecifics do and do not see. Animal Behaviour, 59, 771785.Google Scholar
Hare, B., Melis, A. P., Woods, V., Hastings, S., & Wrangham, R. (2007). Tolerance allows bonobos to outperform chimpanzees on a cooperative task. Current Biology, 17, 619623.Google Scholar
Hatfield, E., Cacioppo, J. T., & Rapson, R. L. (1993). Emotional contagion. Current Directions in Psychological Science, 2, 96100.Google Scholar
Hepach, R., Vaish, A., & Tomasello, M. (2013). A new look at children’s prosocial motivation. Infancy, 18, 6790.CrossRefGoogle Scholar
Herrmann, E., Hare, B., Call, J., & Tomasello, M. (2010). Differences in the cognitive skills of bonobos and chimpanzees. PLoS ONE, 5, e12438.CrossRefGoogle ScholarPubMed
Hess, U., & Blairy, S. (2001). Facial mimicry and emotional contagion to dynamic emotional facial expressions and their influence on decoding accuracy. International Journal of Psychophysiology, 40, 129141.Google Scholar
Hirata, S. (2009). Chimpanzee social intelligence: Selfishness, altruism, and the mother–infant bond. Primates, 50, 311.Google Scholar
Hopper, L. M., Lambeth, S. P., Schapiro, S. J., & Whiten, A. (2008). Observational learning in chimpanzees and children studied through “ghost” conditions. Proceedings of the Royal Society B: Biological Sciences, 275, 835840.Google Scholar
Ioannou, S., Ebisch, S., Aureli, T., Bafunno, D., Ioannides, H. A., Cardone, D., Manini, B., Romani, G. L., Gallese, V., & Merla, A. (2013). The autonomic signature of guilt in children: A thermal infrared imaging study. PLoS ONE, 8, e79440.Google Scholar
Ioannou, S., Gallese, V., & Merla, A. (2014). Thermal infrared imaging in psychophysiology: Potentialities and limits: Thermal infrared imaging in psychophysiology. Psychophysiology, 51, 951963.Google Scholar
Isern-Mas, C., & Gomila, A. (2019). Making sense of emotional contagion. Journal of Philosophical Studies, 12, 71100.Google Scholar
Kano, F., Hirata, S., Deschner, T., Behringer, V., & Call, J. (2016). Nasal temperature drop in response to a playback of conspecific fights in chimpanzees: A thermo-imaging study. Physiology & Behavior, 155, 8394.Google Scholar
Kapitány, R., & Nielsen, M. (2017). Are yawns really contagious? A critique and quantification of yawn contagion. Adaptive Human Behavior and Physiology, 3, 134155.Google Scholar
Kappeler, P. M. (1993). Reconciliation and post-conflict behaviour in ringtailed lemurs, Lemur catta and redfronted lemurs, Eulemur fulvus rufus. Animal Behaviour, 45, 901915.Google Scholar
Kim, Y., Martinez, L., Choe, J. C., Lee, D.-J., & Tomonaga, M. (2015). Orangutans (Pongo spp.) do not spontaneously share benefits with familiar conspecifics in a choice paradigm. Primates: Journal of Primatology, 56, 193200.Google Scholar
Koski, S. E., & Sterck, E. H. M. (2007). Triadic postconflict affiliation in captive chimpanzees: Does consolation console? Animal Behaviour, 73, 133142.CrossRefGoogle Scholar
Kret, M. E., Tomonaga, M., & Matsuzawa, T. (2014). Chimpanzees and humans mimic pupil-size of conspecifics. PLoS ONE, 9, e104886.Google Scholar
Krupenye, C., & Call, J. (2019). Theory of mind in animals: Current and future directions. WIREs Cognitive Science, 10, e1503.Google Scholar
Kuraoka, K., & Nakamura, K. (2011). The use of nasal skin temperature measurements in studying emotion in macaque monkeys. Physiology & Behavior, 102, 347355.Google Scholar
Laméris, D. W., Berlo, E. van, Sterck, E. H. M., Bionda, T., & Kret, M. E. (2020). Low relationship quality predicts scratch contagion during tense situations in orangutans (Pongo pygmaeus). American Journal of Primatology, 82, e23138.Google Scholar
Lamm, C., Batson, C. D., & Decety, J. (2007). The neural substrate of human empathy: Effects of perspective-taking and cognitive appraisal. Journal of Cognitive Neuroscience, 19, 4258.Google Scholar
Leone, A., Ferrari, P. F., & Palagi, E. (2014). Different yawns, different functions? Testing social hypotheses on spontaneous yawning in Theropithecus gelada. Scientific Reports, 4, 4010.CrossRefGoogle ScholarPubMed
Lindegaard, M. R., Liebst, L. S., Bernasco, W., Heinskou, M. B., Philpot, R., Levine, M., & Verbeek, P. (2017). Consolation in the aftermath of robberies resembles post-aggression consolation in chimpanzees. PLoS ONE, 12, e0177725.Google Scholar
MacLean, E. L., Sandel, A. A., Bray, J., Oldenkamp, R. E., Reddy, R. B., & Hare, B. A. (2013). Group size predicts social but not nonsocial cognition in lemurs. PLoS ONE, 8, e66359.Google Scholar
MacLean, P. D. (1985). Brain evolution relating to family, play, and the separation call. Archives of General Psychiatry, 42, 405417.Google Scholar
Mallavarapu, S., Stoinski, T. S., Bloomsmith, M. A., & Maple, T. L. (2006). Postconflict behavior in captive western lowland gorillas (Gorilla gorilla gorilla). American Journal of Primatology, 68, 789801.Google Scholar
Mancini, G., Ferrari, P. F., & Palagi, E. (2013). In play we trust: Rapid facial mimicry predicts the duration of playful interactions in geladas. PLoS ONE, 8, e66481.Google Scholar
Martin, J. S., Koski, S. E., Bugnyar, T., Jaeggi, A. V., & Massen, J. J. M. (2021). Prosociality, social tolerance and partner choice facilitate mutually beneficial cooperation in common marmosets, Callithrix jacchus. Animal Behaviour, 173, 115136.Google Scholar
Massen, J. J. M., & Gallup, A. C. (2017). Why contagious yawning does not (yet) equate to empathy. Neuroscience & Biobehavioral Reviews, 80, 573585.Google Scholar
Massen, J. J. M., Šlipogor, V., & Gallup, A. C. (2016). An observational investigation of behavioral contagion in common marmosets (Callithrix jacchus): Indications for contagious scent-marking. Frontiers in Psychology, 7.Google Scholar
Massen, J. J. M., Vermunt, D. A., & Sterck, E. H. M. (2012). Male yawning is more contagious than female yawning among chimpanzees (Pan troglodytes). PLoS ONE, 7(7), e40697.Google Scholar
Matheson, M. D. (1999). Social contact following severe aggression in rhesus macaques (Macaca mulatta): A new test of the consolation hypothesis. International Journal of Primatology, 20, 961975.Google Scholar
McFarland, R., & Majolo, B. (2012). The occurrence and benefits of postconflict bystander affiliation in wild Barbary macaques, Macaca sylvanus. Animal Behaviour, 84, 583591.Google Scholar
Melis, A. P., Call, J., & Tomasello, M. (2006). Chimpanzees (Pan troglodytes) conceal visual and auditory information from others. Journal of Comparative Psychology, 120, 154162.Google Scholar
Meltzoff, A. N., & Moore, M. K. (1977). Imitation of facial and manual gestures by human neonates. Science, 198, 7578.Google Scholar
Myowa, M. (1996). Imitation of facial gestures by an infant chimpanzee. Primates, 37, 207213.Google Scholar
Myowa‐Yamakoshi, M., Tomonaga, M., Tanaka, M., & Matsuzawa, T. (2004). Imitation in neonatal chimpanzees (Pan troglodytes). Developmental Science, 7, 437442.Google Scholar
Nakayama, K., Goto, S., Kuraoka, K., & Nakamura, K. (2005). Decrease in nasal temperature of rhesus monkeys (Macaca mulatta) in negative emotional state. Physiology & Behavior, 84, 783790.Google Scholar
Norscia, I., Demuru, E., & Palagi, E. (2016). She more than he: Gender bias supports the empathic nature of yawn contagion in Homo sapiens. Royal Society Open Science, 3, 150459.Google Scholar
Ostner, J., Wilken, J., & Schülke, O. (2021). Social contagion of affiliation in female macaques. Royal Society Open Science, 8, 201538.Google Scholar
Overduin-de Vries, A. M., Spruijt, B. M., & Sterck, E. H. M. (2014). Long-tailed macaques (Macaca fascicularis) understand what conspecifics can see in a competitive situation. Animal Cognition, 17, 7784.Google Scholar
Palagi, E. (2018). Not just for fun! Social play as a springboard for adult social competence in human and non-human primates. Behavioral Ecology and Sociobiology, 72, 90.Google Scholar
Palagi, E., Bertini, M., Annicchiarico, G., & Cordoni, G. (2020). Mirror replication of sexual facial expressions increases the success of sexual contacts in bonobos. Scientific Reports, 10, 18979.Google Scholar
Palagi, E., Cordoni, G., & Tarli, S. B. (2006). Possible roles of consolation in captive chimpanzees (Pan troglodytes). American Journal of Physical Anthropology, 129, 105111.Google Scholar
Palagi, E., Dall’Olio, S., Demuru, E., & Stanyon, R. (2014). Exploring the evolutionary foundations of empathy: Consolation in monkeys. Evolution and Human Behavior, 35, 341349.Google Scholar
Palagi, E., Leone, A., Mancini, G., & Ferrari, P. F. (2009). Contagious yawning in gelada baboons as a possible expression of empathy. Proceedings of the National Academy of Sciences, 106, 1926219267.Google Scholar
Palagi, E., Norscia, I., & Cordoni, G. (2019). Lowland gorillas (Gorilla gorilla gorilla) failed to respond to others’ yawn: Experimental and naturalistic evidence. Journal of Comparative Psychology, 133, 406416.Google Scholar
Palagi, E., Norscia, I., & Demuru, E. (2014). Yawn contagion in humans and bonobos: Emotional affinity matters more than species. PeerJ, 2, e519.Google Scholar
Palagi, E., Norscia, I., Pressi, S., & Cordoni, G. (2019). Facial mimicry and play: A comparative study in chimpanzees and gorillas. Emotion, 19, 665681.Google Scholar
Palagi, E., Paoli, T., & Tarli, S. B. (2004). Reconciliation and consolation in captive bonobos (Pan paniscus). American Journal of Primatology, 62, 1530.Google Scholar
Parr, L. A. (2001). Cognitive and physiological markers of emotional awareness in chimpanzees (Pan troglodytes). Animal Cognition, 4, 223229.Google Scholar
Parr, L., & Waller, B. M. (2006). Understanding chimpanzee facial expression: Insights into the evolution of communication. Social Cognitive and Affective Neuroscience, 1, 221228.Google Scholar
Paukner, A., & Anderson, J. R. (2006). Video-induced yawning in stumptail macaques (Macaca arctoides). Biology Letters, 2, 3638.Google Scholar
Paukner, A., Suomi, S. J., Visalberghi, E., & Ferrari, P. F. (2009). Capuchin monkeys display affiliation toward humans who imitate them. Science, 325, 880883.Google Scholar
Pérez‐Manrique, A., & Gomila, A. (2018). The comparative study of empathy: Sympathetic concern and empathic perspective-taking in non-human animals. Biological Reviews, 93, 248269.Google Scholar
Platek, S. M., Critton, S. R., Myers, T. E., & Gallup, G. G. (2003). Contagious yawning: The role of self-awareness and mental state attribution. Cognitive Brain Research, 17, 223227.Google Scholar
Plotnik, J. M., & de Waal, F. B. M. (2014). Asian elephants (Elephas maximus) reassure others in distress. PeerJ, 2, e278.Google Scholar
Premack, D., & Woodruff, G. (1978). Does the chimpanzee have a theory of mind? Behavioral and Brain Sciences, 1, 515526.Google Scholar
Preston, S. D., & de Waal, F. B. M. (2002). Empathy: Its ultimate and proximate bases. Behavioral and Brain Sciences, 25, 120.Google Scholar
Prochazkova, E., Prochazkova, L., Giffin, M. R., Scholte, H. S., Dreu, C. K. W. D., & Kret, M. E. (2018). Pupil mimicry promotes trust through the theory-of-mind network. Proceedings of the National Academy of Sciences, 115, E7265E7274.Google Scholar
Provine, R. R. (1986). Yawning as a stereotyped action pattern and releasing stimulus. Ethology, 72, 109122.Google Scholar
Provine, R. (1992). Contagious laughter: Laughter is a sufficient stimulus for laughs and smiles. Bulletin of the Psychonomic Society, 30, 14.Google Scholar
Pruetz, J. D. (2011). Targeted helping by a wild adolescent male chimpanzee (Pan troglodytes verus): Evidence for empathy? Journal of Ethology, 29, 365368.Google Scholar
Reddy, R. B., Krupenye, C., MacLean, E. L., & Hare, B. (2016). No evidence for contagious yawning in lemurs. Animal Cognition, 19, 889898.Google Scholar
Reimert, I., Fong, S., Rodenburg, T. B., & Bolhuis, J. E. (2017). Emotional states and emotional contagion in pigs after exposure to a positive and negative treatment. Applied Animal Behaviour Science, 193, 3742.Google Scholar
Romero, T., & de Waal, F. B. M. (2010). Chimpanzee (Pan troglodytes) consolation: Third-party identity as a window on possible function. Journal of Comparative Psychology, 124, 278286.Google Scholar
Rymarczyk, K., Żurawski, Ł., Jankowiak-Siuda, K., & Szatkowska, I. (2018). Neural correlates of facial mimicry: Simultaneous measurements of EMG and BOLD responses during perception of dynamic compared to static facial expressions. Frontiers in Psychology, 9, 52.Google Scholar
Samuni, L., Preis, A., Mielke, A., Deschner, T., Wittig, R. M., & Crockford, C. (2018). Social bonds facilitate cooperative resource sharing in wild chimpanzees. Proceedings of the Royal Society B: Biological Sciences, 285, 20181643.Google Scholar
Sandel, A. A., MacLean, E. L., & Hare, B. (2011). Evidence from four lemur species that ringtailed lemur social cognition converges with that of haplorhine primates. Animal Behaviour, 81, 925931.Google Scholar
Santos, L. R., Nissen, A. G., & Ferrugia, J. A. (2006). Rhesus monkeys, Macaca mulatta, know what others can and cannot hear. Animal Behaviour, 71, 11751181.Google Scholar
Sato, Y., Hirata, S., & Kano, F. (2019). Spontaneous attention and psycho-physiological responses to others’ injury in chimpanzees. Animal Cognition, 22, 807823.Google Scholar
Scerif, G., Gomez, J.-C., & Byrne, R. W. (2004). What do Diana monkeys know about the focus of attention of a conspecific? Animal Behaviour, 68, 12391247.Google Scholar
Schino, G., & Marini, C. (2012). Self-protective function of post-conflict bystander affiliation in mandrills. PLoS ONE, 7, e38936.Google Scholar
Seyfarth, R. M., & Cheney, D. L. (2013). Affiliation, empathy, and the origins of theory of mind. Proceedings of the National Academy of Sciences, 110, 1034910356.Google Scholar
Sonnby-Borgström, M. (2002). Automatic mimicry reactions as related to differences in emotional empathy. Scandinavian Journal of Psychology, 43, 433443.Google Scholar
Speakman, J. R., & Ward, S. (1998). Infrared thermography: Principles and applications. Zoology, 101, 224232.Google Scholar
Tan, J., Ariely, D., & Hare, B. (2017). Bonobos respond prosocially toward members of other groups. Scientific Reports, 7, 111.Google Scholar
Tan, J., & Hare, B. (2013). Bonobos share with strangers. PLoS ONE, 8, e51922.CrossRefGoogle ScholarPubMed
Tan, J., Kwetuenda, S., & Hare, B. (2015). Preference or paradigm? Bonobos show no evidence of other-regard in the standard prosocial choice task. Behaviour, 152, 521544.Google Scholar
van Berlo, E., Díaz-Loyo, A. P., Juárez-Mora, O. E., Kret, M. E., & Massen, J. J. M. (2020). Experimental evidence for yawn contagion in orangutans (Pongo pygmaeus). Scientific Reports, 10, 22251.Google Scholar
van den Berg, L. M., Reader, S. M., Goossens, B. M. A., & Sterck, E. H. M. (2012). An analysis of gaze following to a hidden location in long-tailed macaques (Macaca fascicularis). Behaviour, 149, 13191337.Google Scholar
van der Schalk, J., Fischer, A., Doosje, B., Wigboldus, D., Hawk, S., Rotteveel, M., & Hess, U. (2011). Convergent and divergent responses to emotional displays of ingroup and outgroup. Emotion, 11, 286298.Google Scholar
van Schaik, C. P., Deaner, R. O., & Merrill, M. Y. (1999). The conditions for tool use in primates: Implications for the evolution of material culture. Journal of Human Evolution, 36, 719741.Google Scholar
Verbeek, P., & de Waal, F. B. M. (1997). Postconflict behavior of captive brown capuchins in the presence and absence of attractive food. International Journal of Primatology, 18, 703725.Google Scholar
Want, S. C., & Harris, P. L. (2002). How do children ape? Applying concepts from the study of non-human primates to the developmental study of “imitation” in children. Developmental Science, 5, 114.Google Scholar
Watts, D. P. (1995). Post-conflict social events in wild mountain gorillas (Mammalia, Hominoidea): II. Redirection, side directed behaviour and consolation. Ethology, 100, 158174.Google Scholar
Watts, D. P. (2002). Reciprocity and interchange in the social relationships of wild male chimpanzees. Behaviour, 139, 343370.Google Scholar
Webb, C. E., Romero, T., Franks, B., & de Waal, F. B. M. (2017). Long-term consistency in chimpanzee consolation behaviour reflects empathetic personalities. Nature Communications, 8, 18.Google Scholar
Wittig, R. M., & Boesch, C. (2003). The choice of post-conflict interactions in wild chimpanzees (Pan troglodytes). Behaviour, 140, 15271559.CrossRefGoogle Scholar
Yamamoto, C., Morisaka, T., Furuta, K., Ishibashi, T., Yoshida, A., Taki, M., Mori, Y., & Amano, M. (2015). Post-conflict affiliation as conflict management in captive bottlenose dolphins (Tursiops truncatus). Scientific Reports, 5, 14275.Google Scholar
Yamamoto, S. (2015). Non-reciprocal but peaceful fruit sharing in wild bonobos in Wamba. Behaviour, 152, 335357.Google Scholar
Yamamoto, S., Humle, T., & Tanaka, M. (2009). Chimpanzees help each other upon request. PLoS ONE, 4, e7416.Google Scholar
Yamamoto, S., Humle, T., & Tanaka, M. (2012). Chimpanzees’ flexible targeted helping based on an understanding of conspecifics’ goals. Proceedings of the National Academy of Sciences, 109, 35883592.Google Scholar
Zahn-Waxler, C., & Radke-Yarrow, M. (1990). The origins of empathic concern. Motivation and Emotion, 14, 107130.Google Scholar
Zentall, T. R. (2003). Imitation by animals: How do they do it? Current Directions in Psychological Science, 12, 9195.CrossRefGoogle Scholar

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