Pallante V., Johann A., Huffman M.A., Palagi E. (2022). Stone handling in geladas (Theropithecus gelada): implications for spontaneous drawing-like activity as a playful behavior. Journal of Ethology,

Nolfo A.P., Casetta G., Palagi E. 2022. Visual communication in social play of a hierarchical carnivore species: the case of wild spotted hyenas. Current Zoology 68(4), 411–422.

Maglieri V., Zanoli A., Mastrandrea F., Palagi E. 2022. The relaxed open mouth is a true signal in dogs: demonstrating Tinbergen’s ritualisation process. Animal Behaviour 188, 65-74.

Palagi E., Celeghin A., Tamietto M., Winkielman P., Norscia, I. 2022. Disentangling attentional and affective contribution to contagious yawning. Neuroscience and Biobehavioural Reviews 132, 892-893. doi:10.1016/j.neubiorev.2021.10.050

Palagi E, Caruana F, de Waal F.B.M. 2022 The naturalistic approach to laughter in humans and other animals: toward a unified theory. Philosophical transactions of the Royal Society of London. Series B, 377(1863), pp. 2021017520210175.

Bresciani C., Cordoni G.*, Palagi E.* (2022) Playing together, laughing together: rapid facial mimicry and social sensitivity in lowland gorillas, Current Zoology, 68(5): 560–569. *Senior authorship shared

Zanoli A., Gamba M., Lemasson A., Norscia I.*, Palagi E.* Inter-sexual multimodal communication during mating in wild geladas: the leading role of females. Current Zoology, 68(5) 570–580. *Senior authorship shared

Casetta G., Nolfo A.P., Palagi E. (2022). Yawning informs behavioural state changing in wild spotted hyaenas. Behavioral Ecology and Sociobiology. 76, 152.

Giuntini S., Ancillotto L., Falaschi M., Viviano A., Palagi E., Mori E. 2022. Alien parakeets as a potential threat to the common noctule Nyctalus noctula. Biodiversity and Conservation.

Zonato A., Gagliardo A., Bandoli F.*, Palagi E.* 2022. Reaching versus catching: flexible hand preference in ring-tailed lemurs. Ethology Ecology & Evolution 10.1080/03949370.2022.2098382. *Senior authorship shared

Gallo A., Zanoli A., Caselli M., Norscia I.*, Palagi E.* 2022. The face never lies: facial expressions and mimicry modulate playful interactions in wild geladas. Behavioral Ecology and Sociobiology 76(1),19. doi:10.1007/s00265-022-03126-4

Bertini M., Annicchiarico G., Bresciani C., Cordoni G., Palagi E. 2022. Playful interactions and facial mimicry in infant bonobos (Pan paniscus), Ethology Ecology & Evolution, 34: 344-359, DOI:10.1080/03949370.2021.1988723

Davila-Ross M*., Palagi E*. 2022. Laughter, play faces and mimicry in animals: evolution and social functions. Philosophical Transactions of the Royal Society of London Series B, 377 (1863). doi:10.1098/rstb.2021-0177). *Shared contribution.

Scopa C., Maglieri V., Baragli P., Palagi E. 2022. Getting rid of blinkers: the case of mirror self-recognition in horses (Equus caballus). Animal Cognition 25, 711–716.

Pedruzzi L., Aychet J., Le Vern L., Maglieri V., Rossard A., Lemasson A.*, Palagi E.* 2022. Familiarity modulates both intra- and interspecific yawn contagion in red-capped mangabeys. Scientific Reports 12, 11138. *Senior authorship shared

Caruana, F., Palagi, E., de Waal, F.B.M. 2022. Cracking the laugh code: laughter through the lens of biology, psychology and neuroscience. Philosophical Transactions of the Royal Society of London. Series B, 377(1863), pp. 20220159.

Palagi, E., Pellis S.M (2022). Play behavior: a comparative perspective. In: Routledge Handbook of Comparative Psychology (Freeberg, T.M., Ridley, A.R., & d’Ettorre, P. eds.). pp 202-212


Casetta G., Nolfo P.A., Palagi E. 2021. Yawn contagion promotes motor synchrony in wild lions, Panthera leo. Animal Behaviour 174, 149-159.

Maglieri V., Riccobono M.G., Giunchi D., Palagi E. 2021 Navigating from live to virtual social interactions: looking at but not manipulating smartphones provokes a spontaneous mimicry response in the observers. Journal of Ethology 39(3), 287-296.

Cordoni G., Favilli E., Palagi E. 2021. Earlier than previously thought: yawn contagion in preschool children. Developmental Psychobiology 63(5), 931-944.

Baragli P., Scopa C., Maglieri V., Palagi E. 2021. If horses had toes: demonstrating mirror self-recognition at group level in Equus caballus. Animal Cognition 24, 1099–1108.

Aychet J., Blois-Heulin C., Palagi E., Lemasson A. 2021. Facial displays in red-capped mangabeys (Cercocebus torquatus): Repertoire, social context, and potential intentionality. Journal of Comparative Psychology 135(1), 98-113.

Gallo A., Caselli M., Norscia I.*, Palagi E.* 2021. Let’s unite in play! Play modality and group membership in wild geladas. Behavioural Processes 184, 104338. *Senior authorship and corresponding shared.

Norscia I.*, Agostini L., Moroni A., Caselli M., Micheletti-Cremasco M., Vardé C., Palagi E.* 2021. Yawning is more contagious to pregnant than nulliparous women: naturalistic and experimental evidence. Human Nature 32(2):301-325. doi: 10.1007/s12110-021-09404-w. Corresponding shared.

Zannella A., Stanyon R., Maglieri V., Palagi E. 2021. Not all yawns tell the same story: the case of Tonkean macaques. American Journal of Primatology 83: e23263. doi: 10.1002/ajp.23263.

Zanoli A., Gamba M., Lemasson A., Palagi E.*, Norscia I.* 2021 Looking into each other’s eyes makes it better: Eye-to-Eye Contact enhances sexual interactions in geladas. Animal Behaviour 177, 269-276. *Senior authorship and corresponding shared.

Anzà, S., Demuru, E., Palagi, E. 2021 Sex and grooming as exchange commodities in female bonobos’ daily biological market. Sciemtific Reports Rep 11, 19344.

Caselli M., Zanoli A., Dagradi C., Gallo A., Yazezew D., Tadesse A., Capasso M., Ianniello D., Rinaldi L., Palagi E.*, Norscia I.* 2021. Wild geladas (Theropithecus gelada) in crops – more than in pasture areas – reduce aggression and affiliation. Primates, 62(4), 571-584. *Senior authorship and corresponding shared.

Nolfo A.P., Casetta G., Palagi E. 2021 Play fighting in wild spotted hyenas: like a bridge over the troubled water of a hierarchical society. Animal Behaviour 180, 363-373.

Gallo A., Zanoli A., Caselli M., Palagi E.*, Norscia I.* 2021. First evidence of yawn contagion in a wild monkey species. Scientific Reports 11, 17957. *Senior authorship and corresponding shared

Caselli M., Zanoli A, Palagi E.*, Norscia I.* 2021 Infant handling increases grooming towards mothers in wild geladas (Theropithecus gelada). Behavioural Processes 104501, *Senior authorship and corresponding shared

Palagi E., Bergman T.J. 2021. Bridging captive and wild studies: behavioural plasticity and social complexity in Theropithecus gelada. Animals 2021, 11, 3003.

Bresciani C., Cordoni G.*, Palagi E.* 2021. Playing together, laughing together: rapid facial mimicry and social sensitivity in lowland gorillas. Current Zoology, zoab092.*Senior authorship shared

Zanoli A., Gamba M., Lemasson A., Norscia I.*, Palagi E.* 2021. Inter-sexual multimodal communication during mating in wild geladas: the leading role of females. Current Zoology, zoab093, *Senior authorship and corresponding shared

Palagi E., Celeghin A., Tamietto M., Winkielman P., Norscia, I. 2021. Disentangling attentional and affective contribution to contagious yawning. S0149-7634(21)00495-4. doi: 10.1016/j.neubiorev.2021.10.050 Neuroscience and Biobehavioural Reviews.


Palagi, E., Celeghin A., Tamietto M., Winkielman P., Norscia I. 2020. The neuroethology of spontaneous mimicry and emotional contagion in human and non-human animals. Neuroscience and Biobehavioral Reviews 111, 149–165.

Palagi E., Bertini M, Annicchiarico G., Cordoni G. 2020. Mirror replication of sexual facial expressions increases the probability of successful sexual contacts in bonobos. Scientific Reports 10, 18979.

Palagi, E., Cordoni, C. 2020.  Intraspecific motor and emotional alignment in dogs and wolves: the basic building blocks of dog–human affective connectedness. Animals, 10: 241; doi:10.3390/ani10020241

Norscia I., Zanoli A., Gamba M., Palagi E. 2020. Auditory contagious yawning is highest between friends and family members: support to the emotional bias hypothesis. Frontiers in Psychology 11: 442; doi:10.3389/fpsyg.2020.00442

Maglieri, V., Bigozzi, F., Riccobono, M. G., Palagi, E. 2020. Levelling playing field: synchronization and rapid facial mimicry in dog-horse play. Behavioural Processes 174:

Palagi E., Barbuti R., Norscia I., 2020. Aquarium cichlid fish Tropheus moorii flexibly adjust hierarchy when tank fish species composition changes: A pilot study. Journal of Zoo and Aquarium Research 8(2): 1-8.

Annicchiarico G., Bertini M., Cordoni G., Palagi E. 2020. Look at me while having sex! Eye-to-eye contact affects homosexual behaviour in bonobo females. Behaviour 157:949–970.

Cangiano M., Palagi E. 2020. First evidence of stone handling in geladas: from simple to more complex forms of object play. Behavioural Processes 180: 180:104253.


Palagi E., Guillén-Salazar F., Llamazares-Martín C. 2019. Spontaneous yawning and its potential functions in South American sea lions (Otaria flavescens). Scientific Reports 9, 17226.

Cordoni C., Palagi E. 2019. Back to the future: a glance over wolf social behaviour to understand dog-human relationship. Animals, 9, 991; doi:10.3390/ani9110991

Maglieri V., Prato-Previde E., Tomasi E., Palagi E. 2019. Wolf-like or dog-like? A comparison of gazing behaviour across three dog breeds tested in their familiar environments. R. Soc. open sci. 6:190946.

Palagi E., Marchi E., Cavicchio P., Bandoli F. 2019. Sharing playful mood: rapid facial mimicry in Suricata suricatta. Animal Cognition, 22(5), 719-732. (doi:10.1007/s10071-019-01269-y)

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(3), 406-416. (doi: 10.1037/com0000175)

Smaldino P.E, Palagi E., Burghardt G.M., Sergio Pellis S.M. 2019. The evolution of two types of play. Behavioral Ecology, 30(5), 1388–1397. (doi:10.1093/beheco/arz090)

Pallante V., Ferrari P.F., Gamba M., Palagi E. 2019. Embracing in a female bonded monkey species (Theropithecus gelada). Journal of Comparative Psychology (on line doi:10.1037/com0000173)

Palagi E., Norscia I. 2019. Etologia dell’emozione animale: studio e interpretazione. Sistemi intelligenti, 1: 11-32. (doi:10.1422/92933)

Palagi E., Norscia I., Pressi S., Cordoni G. 2019. Facial mimicry and play: a comparative study in chimpanzees and gorillas. Emotion 19(4):665-681. (doi: 10.1037/emo0000476)


Palagi E. 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 (doi:10.1007/s00265-018-2506-6)

Palagi E., Norscia I., Pressi S., Cordoni G. 2018. Facial mimicry and play: a comparative study in chimpanzees and gorillas. Emotion (

Cordoni, G., Norscia, I., Bobbio, M., Palagi, E. 2018. Differences in play can illuminate differences in affiliation: a comparative study on chimpanzees and gorillas. PLoS ONE 13(3): e0193096.

Pallante, V., Stanyon, R., Palagi, E. Calming an aggressor through spontaneous post-conflict triadic contacts: appeasement in Macaca tonkeana. Aggressive Behavior, 44: 406-415 (

Demuru E., Ferrari PF., Palagi E. Is birth attendance a uniquely human feature? New evidence suggests that bonobo females protect and support the parturient. Evolution and Human Behavior, 39 (5), 502-510 (

Palagi E., Leone A., Demuru E., Ferrari PF. 2018 High-ranking geladas protect and comfort others after conflicts. Scientific Reports 8, 15291 (

Scopa C., Palagi E., Sighieri C.m, Baragli P. 2018 Physiological outcomes of calming behaviors support the resilience hypothesis in horses. Scientific Reports 8, 17501 (doi:10.1038/s41598-018-35561-7)


Palagi, E., Scopa C. 2017. Integrating Tinbergen’s inquiries: Mimicry and play in humans and other social mammals. Learning & Behavior, 45(4), 378-389 (doi:10.3758/s13420-017-0278-x).

Llamazares-Martín, C., Scopa, C., Guillén-Salazar, F., Palagi, E. Strong competition does not always predict play asymmetry: the case of South American sea lions (Otaria flavescens). Ethology, 123 (4): 270–282. doi:10.1111/eth.12594

Zannella, A., Stanyon, R., Palagi E., 2017. Yawning and social styles: different functions in tolerant and despotic macaques (Macaca tonkeana and Macaca fuscata). Journal of Comparative Psychology. 131(3): 179-188

Norscia, I., Palagi, E. When do you scratch that itch? The relative impact of different factors on scratching depends on the selection of time scale and timing. Royal Society Open Science. 4: 170106.

Llamazares-Martín, C., Scopa, C., Guillén-Salazar, F., Palagi, E. Relaxed open mouth reciprocity favours playful contacts in south american sea lions (Otaria flavescens). Behavioural Processes 140:87-95. doi:10.1016/j.beproc.2017.04.007

Baragli, P., Demuru, E., Scopa, C., Palagi, E. Are horses capable of mirror self-recognition? A pilot study. PLoS ONE 12(5): e0176717.

Baragli, P., Vitale, V., Sighieri, C., Lanatà, A., Palagi, E., Reddon, A. 2017. Consistency and flexibility in solving spatial tasks: different horses show different cognitive styles. Scientific Reports 7: 16557. (doi:10.1038/s41598-017-16729-z)


Palagi E., Burghardt G.M., Smuts B., Cordoni G., Dall’Olio S., Fouts H.N., Řeháková-Petrů, Siviy S. M; Pellis S.M. 2016. Rough-and-tumble play as a window on animal communication. Biological Reviews 91: 311–327 (doi:10.1111/brv.12172)

Palagi, E., Fouts, H.N. 2016. Motivation of play: from ethological to neurological perspectives. Behaviour 153: 655–662.

Palagi, E., Cordoni, G., Demuru, E., Bekoff, M. 2016. Fair play and its connection with social tolerance, reciprocity and the ethology of peace. Behaviour, 153: 1195–1216

Norscia, I., Demuru, E., Palagi E. She more than he: gender bias supports the empathic nature of yawn contagion in Homo sapiens. Royal Society Open Science, 3. (

Norscia, I., Demuru, E., Palagi E. Difference in contagious yawning between susceptible men and women: why not? Royal Society Open Science 3 160477. (doi:10.1098/rsos.160477).

Arrigoni Ferrari, G., Nicolini, Y., Demuru, E., Tosato, C., Hussein, M., Scesa, E., Romei, L., Boerci, M., Iappini, E., Dalla Rosa Prati, G., Palagi, E., Ferrari P. F. 2016. Ultrasonographic investigation of human fetus responses to maternal communicative and non-communicative stimuli. Frontiers in Psychology 7: (doi: 10.3389/fpsyg.2016.00354)

Scopa, C., Palagi, E. Mimic me while playing! Social tolerance and rapid facial mimicry in macaques (Macaca tonkeana and Macaca fuscata). Journal of Comparative Psychology 130(2):153-61. (doi:10.1037/com0000028).

Cordoni, G., Palagi E. Aggression and hierarchical steepness inhibit social play in adult wolves. Behaviour 153: 749 – 766. (doi:10.1163/1568539X-00003326)

Cordoni, G., Nicotra, V., Palagi, E. Unveiling the “secret” of play in dogs (Canis lupus familiaris): asymmetry and signals. Journal of Comparative Psychology 130: 278-287. (doi: 10.1037/com0000035).

Verbeek, P., Palagi, E. Looking at natural peace through Niko Tinbergen’s lens. Behaviour, 153: 1005–1011.

Pallante, V., Stanyon, R., Palagi E., 2016. Agonistic support towards victims buffers aggression in geladas (Theropithecus gelada). Behaviour, 153: 1217–1243.

Cordoni, G., Demuru, E., Ceccarelli, E., Palagi E. Play, aggressive conflict and reconciliation in pre-school children: what matters? Behaviour, 153: 1075–1102.


Palagi, E., Norscia I., 2015. The season for peace: reconciliation in a despotic species (Lemur catta). PLOS ONE 10(11): e0142150. doi:10.1371/journal.pone.0142150

Cordoni, G., Palagi E. Being a victim or an aggressor: Different functions of triadic post-conflict interactions in wolves (Canis lupus lupus). Behaviour 41: 526-36. doi: 10.1002/ab.21590.

Palagi, E., Demuru E. 2015. Play fair, game over: le regole “biologicamente” tacite del regno animale. Sistemi Intelligenti 27: 181-190.

Demuru E., Ferrari P.F., Palagi E. 2015. Emotionality and intentionality in bonobo playful communication. Animal Cognition 18: 333-344

Pellis S.M., Burghardt G.M., Palagi E., Mengel M. 2015. Modeling play: distinguishing between origins and current functions. Adaptive Behavior 23: 331–339 doi:10.1177/1059712315596053

Zannella, A., Norscia I., Stanyon, R., Palagi E., 2015. Testing yawning hypotheses in wild populations of two strepsirrhine species: Propithecus verreauxi and Lemur catta. American Journal of Primatology, 77: 1207–1215. doi: 10.1002/ajp.22459

Norscia I., Palagi E. The socio-matrix reloaded: from hierarchy to dominance profile in wild lemurs. PeerJ 3:e729. DOI 10.7717/peerj.729

Palagi E., Stanyon, R.R., Demuru, E. 2015. Play to learn. Teach by play. Behavioral and Brain Sciences, 38: doi:10.1017/S0140525X14000557

Palagi, E., Nicotra, V., Cordoni, G., 2015. Rapid mimicry and emotional contagion in domestic dogs. Royal Society Open Science, 2: 150505. doi:10.1098/rsos.150505


Palagi E., Dall’Olio S., Demuru E., Stanyon R. 2014. Exploring the evolutionary foundations of empathy: consolation in monkeys. Evolution and Human Behavior 35: 341-349

Palagi E., Norscia I., Spada G. 2014. Relaxed open mouth as a playful signal in wild ring-tailed lemurs. American Journal of Primatology 76: 1074-1083.

Palagi E., Norscia I., Demuru, E. 2014. Yawn contagion in humans and bonobos: emotional affinity matters more than species. 2014; 2: e519. (doi:10.7717/peerj.519).

Leone A., Ferrari PF., Palagi E. Different yawns, different functions? Testing social hypotheses on spontaneous yawning in Theropithecus gelada. Scientific Reports 4: 4010 (DOI:doi:10.1038/srep04010)


Palagi E, Norscia I 2013. Bonobos protect and console friends and kin. PLoS ONE 8(11): e79290. doi:10.1371/journal.pone.0079290

Mancini G., Ferrari P.F., Palagi E. Rapid facial mimicry in geladas. Scientific Reports 3: 1527 (doi: 10.1038/srep01527)

Mancini G., Ferrari P.F., Palagi E. In play we trust. Rapid facial mimicry predicts the duration of playful interactions in geladas. PLoS ONE 8(6):e66481. (doi: 10.1371/journal.pone.0066481)


Palagi, E., Cordoni, G. 2012. The right time to happen: play developmental divergence in the two Pan —PLoS ONE 7(12), e52767.

Addessi E., Borgi M., Palagi E. Is Primatology an equal-opportunity discipline? PLoS ONE 7(1): e30458. doi:10.1371/journal.pone.0030458

Sclafani V., Norscia I., Antonacci D., Palagi E. Scratching around mating: factors affecting anxiety in wild Lemur catta. Primates 53, 247-254. (doi:10.1007/s10329-012-0294-6)

Dall’Olio S, Norscia I, Antonacci D, Palagi E. Sexual signalling in Propithecus verreauxi: male ‘‘chest badge’’ and female mate choice. PLoS ONE 7(5):e37332. (doi:10.1371/journal.pone.0037332).

Ciani F., Dall’Olio S., Stanyon R., Palagi E. Social tolerance and adult play in macaque societies: a comparison with different human cultures. Animal Behaviour 84, 1313-1322.

Demuru E., Palagi E. In bonobos yawn contagion is higher among kin and friends. PLoS ONE 7(11): e49613. (doi:10.1371/journal.pone.0049613).


Palagi E., Norscia I. 2011. Scratching around stress: hierarchy and reconciliation make the difference in prosimians. Stress 14: 93-97.

Norscia I., Palagi, E. Fragment quality and distribution of the arboreal primate Propithecus verreauxi in the spiny forest of South Madagascar. Journal of Tropical Ecology 27: 103-106.

Palagi E., Mancini G. 2011. Playing with the face: playful facial chattering and its modulation in a monkey species. Journal of Comparative Psychology 125: 11-21.

Norscia I., Palagi, E. Do wild brown lemurs reconcile? Not always. Journal of Ethology 29:181–185.

Norscia I., Palagi E. When play is a family business: adult play, hierarchy, and possible stress reduction in common marmosets. Primates 52: 101-104.

Cordoni G., Palagi E. Ontogenetic trajectories of chimpanzee social play: what similarities with humans? PLoS ONE 6(11): e27344. doi:10.1371/journal.pone.0027344.

Norscia I., Palagi E. Yawn contagion and empathy in Homo sapiens. PLoS ONE 6(12): e28472. (doi:10.1371/journal.pone.0028472).


Leone A., Palagi E. Reconciling conflicts in a one-male society: the case of geladas. Primates 51:203-212.

Leone A., Mignini M., Mancini G., Palagi E. Aggression does not increase friendly contacts among bystanders in geladas (Theropithecus gelada). Primates 51:299-305.

Antonacci D., Norscia I., Palagi E. Stranger to familiar: wild strepsirhines manage xenophobia by playing. PLoS ONE 5(10): e13218. doi:10.1371/journal.pone.0013218.


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 of the United States of America 106: 19262-19267.

Palagi E., Norscia, I. 2009. Multimodal signaling in wild Lemur catta: economic design and territorial function of urine marking. American Journal of Physical Anthropology 139:182 – 192.

Palagi E., Cordoni G. 2009. Postconflict third-party affiliation in Canis lupus: do wolves share similarities with the great apes? Animal Behaviour 78: 979-986.

Palagi E. Adult play fighting and potential role of tail signals in ringtailed lemurs (Lemur catta). Journal of Comparative Psychology 123: 1-9.

Norscia I., Antonacci A., Palagi E. 2009 Mating first, mating more: biological market fluctuation in a wild prosimian. PLoS ONE 4(3): e4679 (doi:10.1371/journal.pone.0004679).

Tacconi G., Palagi E. Play behavioural tactics under space reduction: social challenges in bonobos (Pan paniscus). Animal Behaviour 78: 469–476.

Mancini G., Palagi E. Play and social dynamics in a captive herd of gelada baboons (Theropithecus gelada). Behavioural Processes 82: 286-292


Palagi E. 2008 Sharing the motivation to play: the use of signals in adult bonobos. Animal Behaviour 75: 887-896.

Palagi E., Antonacci D., Norscia I. 2008. Peacemaking on treetops: first evidence of reconciliation from a wild prosimian (Propithecus verreauxi). Animal Behaviour 76: 737-747.

Cordoni G., Palagi E. 2008 Reconciliation in wolves (Canis lupus): new evidence for a comparative perspective. Ethology 114: 298-308.

Palagi E., Chiarugi E., Cordoni G. 2008. Peaceful post-conflict interactions between aggressors and bystanders in captive lowland gorillas (Gorilla gorilla gorilla). American Journal of Primatology 70: 949-955.

Norscia I., Palagi E. Berenty 2006: census of Propithecus verreauxi and possible evidence of population stress. International Journal of Primatology 29: 1099-1115.


Palagi E., Antonacci D., Cordoni G. 2007. Fine-tuning of social play in juvenile lowland gorillas (Gorilla gorilla gorilla). Developmental Psychobiology 49 (4): 433-445.

Palagi E., Paoli T. 2007. Play in adult bonobos (Pan paniscus): modality and potential meaning. American Journal of Physical Anthropology 134 (2): 219-225.

Palagi E. Play at work: revisiting data focussing on chimpanzees (Pan troglodytes). Journal of Anthropological Sciences 85: 153-164.

Palagi E., Dapporto L. 2007. Females do it better. Individual recognition experiments reveal sexual dimorphism in Lemur catta (Linnaeus, 1758) olfactory motivation and territorial defence. The Journal of Experimental Biology 210: 2700-2705.

Dapporto L., Cini A., Palagi E., Simonti A., Morelli M., Turillazzi S. 2007. Behaviour and chemical signature of pre-hibernating females of Polistes dominulus infected by the strepsipteran Xenos vesparum. Parasitology 134: 545-552.

Paoli T., Tacconi G., Borgognini Tarli S., Palagi E. 2007. Influence of feeding and short-term crowding on the sexual repertoire of captive bonobos (Pan paniscus). Annales Zoologici Fennici 44: 81-88.

Cordoni G., Palagi E. 2007 Response of captive lowland gorillas (Gorilla gorilla gorilla) to different housing conditions: testing the aggression/density and coping models. Journal of Comparative Psychology 121 (2): 171-180.


Palagi E., Cordoni G., Borgognini Tarli S. 2006. Possible roles of consolation in captive chimpanzees (Pan troglodytes). American Journal of Physical Anthropology 127: 105-111.

Palagi E. 2006. Social play in bonobos (Pan paniscus) and chimpanzees (Pan troglodytes): Implications for natural social systems and inter-individual relationships. American Journal of Physical Anthropology 129:418-426.

Paoli T., Palagi E., Tacconi G., Borgognini Tarli S. 2006 Perineal swelling, inter-menstrual cycle, and female sexual behavior in bonobos (Pan paniscus). American Journal of Primatology 68: 333-347.

Paoli T., Palagi E., Borgognini Tarli S.M. 2006. Reevaluation of dominance hierarchy in bonobos (Pan paniscus). American Journal of Physical Anthropology 130: 116-122.

Palagi E., Dapporto L. 2006. Beyond odor discrimination: demonstrating individual recognition in Lemur catta. Chemical Senses 31: 437-443.

Palagi E., Dapporto L. 2006 Urine marking and urination in Lemur catta: a comparison of design features. Annales Zoologici Fennici 43: 280-284.

Dapporto L., Palagi E., Cini A., Turillazzi S. 2006. Pre-hibernating aggregations of Polistes dominulus: an occasion to study early dominance assessment in social insects. Naturwissenschaften 93: 321-324.

Koyama N., Palagi E. 2006. Managing conflict: evidence from wild and captive primates. International Journal of Primatology 27: 1235-1240.

Palagi E., Paoli T., Borgognini Tarli S. 2006. Short-term benefits of play behavior and conflict prevention in Pan paniscus. International Journal of Primatology 27: 1257-1270.

Cordoni G., Palagi E., Borgognini Tarli S. 2006. Reconciliation and consolation in captive western gorillas. International Journal of Primatology 27: 1365-1382.

Dapporto L., Turillazzi S., Palagi E. Dominance interactions in young adult foundresses of a paper wasp: a play-like behaviour? Journal of Comparative Psychology 120: 394-400.

Dapporto, L., Palagi, E. Wasps in the shadow: Looking at the pre-hibernating clusters of Polistes dominulus. Annales Zoologici Fennici 43: 583–594.


Palagi E., Paoli T., Borgognini Tarli S.M. 2005. Aggression and reconciliation in two captive groups of Lemur catta. International Journal of Primatology 26: 279-294.

Palagi E., Dapporto L., Borgognini-Tarli S., 2005 The neglected scent: On the marking function of urine in Lemur catta. Behavioural Ecology and Sociobiology, 58: 437-445.

Dapporto L., Palagi E., Turillazzi S. 2005. Sociality outside the nest: helpers in pre-hibernating clusters of Polistes dominulus. Annales Zoologici Fennici 42: 135-139.


Palagi E., Paoli T., Borgognini Tarli S.M. 2004. Reconciliation and consolation in captive bonobos (Pan paniscus). American Journal of Primatology 62: 15-30.

Palagi E., Telara S., Borgognini Tarli S.M. 2004. Reproductive strategies in Lemur catta: balance among sending, receiving, and counter-marking scent signals. International Journal of Primatology 25: 1019-1031.

Dapporto L., Palagi E., Turillazzi S. 2004. Cuticular hydrocarbons of Polistes dominulus as a biogeographic tool: a study of populations from the Tuscan Archipelago and surrounding areas. Journal of Chemical Ecology 30: 2139-2151.

Palagi E., Cordoni G., Borgognini Tarli S.M. 2004. Immediate and delayed benefits of play behaviour: new evidences from chimpanzees (Pan troglodytes). Ethology 110: 949-962.

Dapporto L., Palagi E., Cini A., Cotoneschi C., Paoli T., Turillazzi S. 2004. Do nest fidelity and tolerance decrease at the end of the colony cycle in Polistes dominulus (Christ.). Redia 87: 227-229.


Palagi E., Telara S., Borgognini Tarli S.M., 2003. Sniffing behavior in Lemur catta: seasonality, sex, and rank. International Journal of Primatology 24: 335-350.

Palagi E., Gregorace A., Borgognini Tarli S.M. 2002. Development of olfactory behavior in captive ring-tailed lemurs (Lemur catta). International Journal of Primatology 23: 587-599.

Palagi E., Lunardini A., 1998. The influence of sexual swelling on the behaviour of captive female chimpanzees (Pan troglodytes). Folia Primatologica 69: 28-34.



Norscia I. & Palagi E. 2016. The Missing Lemur Link: An Ancestral Step in the Evolution of Human Behaviour. Cambridge Studies in Biological and Evolutionary Anthropology. Cambridge University Press. Pp. 1-289.



Llamazares-Martín, C., Palagi, E. 2021. Playing on the edge of the sea: a comparative analysis in Otariids and Odobenids. In: Ethology and Behavioral Ecology of Otariids and the Odobenid (Campagna, C. & Harcourt, R., eds.) Springer Heidelberg, Germany. pp. 391-412.

Palagi, E., Norscia, I. 2018. Emotional contagion. In: Encyclopedia of Animal Cognition and Behavior (Editors Jennifer Vonk, Todd K. Shackelford).

Clay, Z., Palagi E., de Waal, FBM. 2018. Ethological approaches to empathy in primates. In: Neuronal Correlates of Empathy (Editors Ksenia Z. Meyza and Ewelina Knapska). San Diego (USA): Elsevier Science Publishing Co Inc. ( pp. 53-66.

Palagi, E., Demuru, E. 2017. Pan paniscus or Pan ludens? Bonobos, playful attitude and social tolerance. In: Bonobos – Unique in Mind, Brain and Behaviour. Editors Brian Hare and Shinya Yamamoto: Oxford University Press. © Oxford University Press. (pp 65- 77) DOI: 10.1093/acprof:oso/ 9780198728511.003.0005

Palagi E. Humans, other animals, and the biology of morality. In: Beastly Morality: Animals as Moral Agents. (Editor Jonathan Crane) Columbia University Press. pp. 90-103.

Cordoni G., Palagi E. Play and post-conflict mechanisms in wolves: a comparison with human and non-human primates. In: Wolves: Biology, Behavior and Conservation (Editor Nadya Gotsiridze-Columbus). Nova Science Publishers. Pp.133-156.

Cordoni G., Palagi E. Modelli sociali a confronto: forme di celebrazione e riappacificazione nelle antropomorfe africane. In M. Celentano, B. de Mori, P. Zecchinato, editori, Etologia ed Etica pp. 91-106.

Palagi E. Playing alone and with others – A lesson from animals. In: A Handbook of Solitude: Psychological Perspectives on Social Isolation, Social Withdrawal, and Being Alone (Editors Robert J. Coplan & Julie Bowker). Wiley Blackwell. pp. 463-482

Palagi E. Playing at every age: modalities and potential functions in non-human primates. In: The Oxford Handbook of the Development of Play (Editor Anthony Pellegrini). Oxford University Press. (ISBN10: 0195393007/ISBN13: 978-0-19-539300-2) Pp. 70-82.

Palagi E., Paoli T. 2008. Social play in bonobos: not only an immature matter. In: Bonobos: Behaviour, Ecology, and Conservation (Editors Takeshi Furuichi & Jo Thompson). Springer Springer-Verlag New York. pp. 55-74. (ISBN-10: 0387747850/ISBN-13: 9780387747859).

Paoli T., Palagi E. What does agonistic dominance imply in bonobos? In: Bonobos: Behaviour, Ecology, and Conservation (Editors Takeshi Furuichi & Jo Thompson). Springer-Verlag New York, pp 35-54. (ISBN-10: 0387747850/ISBN-13: 9780387747859).

Palagi E., Cordoni G. 2007. The art of peace-keeping. Different tactics in the great apes. In: Issues in the Psychology of Motivation. (Editor Paula R. Zelick). Nova Science Publishers. pp. 157-171. (ISBN: 1-60021-631-5/ISBN 13: 978-1-60021-631-2).


 Palagi E., Fouts H.N. 2016. Motivation of play: from ethological to neurological perspectives. Behaviour, Vol. 153(6-7).

Verbeek P., Palagi E. 2016. Obstacles and catalysts of peaceful behavior. Behaviour, Vol. 153(9-11).

Gregorace A., Palagi E., Borgognini Tarli S.M., 2001. Proceedings of the XIV Congress of the Italian Primatological Society (Calci, 9-11 October, 2000). Folia Primatologica 72: 128-152.

Cordoni G., Norscia I., Palagi E. 2008. Proceedings of the XVIII Congress of the Italian Primatological Society (Calci, 16-18 May, 2007). Folia Primatologica 79:123-159.