Contextual imitation of intransitive body actions in a Beluga whale (Delphinapterus leucas): A “do as other does” study
Open accessAbstract
Cetaceans are remarkable for exhibiting group-specific behavioral traditions or cultures in several behavioral domains (e.g., calls, behavioral tactics), and the question of whether they can be acquired socially, for example through imitative processes, remains open. Here we used a "Do as other does" paradigm to experimentally study the ability of a beluga to imitate familiar intransitive (body-oriented) actions demonstrated by a conspecific. The participant was first trained to copy three familiar behaviors on command (training phase) and then was tested for her ability to generalize the learned "Do as the other does" command to a different set of three familiar behaviors (testing phase). We found that the beluga (1) was capable of learning the copy command signal "Do what-the-other-does"; (2) exhibited high matching accuracy for trained behaviors (mean = 84% of correct performance) after making the first successful copy on command; (3) copied successfully the new set of three familiar generalization behaviors that were untrained to the copy command (range of first copy = 12 to 35 trials); and (4) deployed a high level of matching accuracy (mean = 83%) after making the first copy of an untrained behavior on command. This is the first evidence of contextual imitation of intransitive (body-oriented) movements in the beluga and adds to the reported findings on production imitation of sounds in this species and production imitation of sounds and motor actions in several cetaceans, especially dolphins and killer whales. Collectively these findings highlight the notion that cetaceans have a natural propensity at skillfully and proficiently matching the sounds and body movements demonstrated by conspecifics, a fitness-enhancing propensity in the context of cooperative hunting and anti-predatory defense tactics, and of alliance formation strategies that have been documented in these species' natural habitats. Future work should determine if the beluga can also imitate novel motor actions.
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Belugas have the capacity for contextual imitation of intransitive and novel body actions.
“They have also demonstrated the ability for behavioral imitation during play [44] and the capacity for contextual imitation of intransitive and novel body actions [45,46].”
From Evidence for mirror self-recognition in beluga whales (Delphinapterus leucas) (2026) · cites this paper as [45]
The Discussion reports that one captive beluga copied familiar intransitive body movements on command, supporting the contextual-imitation portion. It explicitly says that whether belugas can imitate novel intransitive actions remains to be established, so this source does not support the novel-actions portion. (LLM, conf=0.94)
Experimental — accuracy not yet measuredBelugas show the capacity for both vocal and behavioral imitation.
“This species shows the capacity for both vocal and behavioral imitation.”
From Evidence for mirror self-recognition in beluga whales (Delphinapterus leucas) (2026) · cites this paper as [45]
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Cites
The faculty of language: what is it, who has it, and how did it evolve?
Hauser, M. D., Chomsky, N., & Fitch, W. T. (2002) · Science · doi:10.1126/science.298.5598.1569 · Cited by 5,189 · see citing works
Tomasello, M., Kruger, A. C., & Ratner, H. H. (1993) · Behavioral and Brain Sciences · doi:10.1017/s0140525x0003123x · Cited by 1,891 · see citing works
SOCIAL LEARNING IN ANIMALS: CATEGORIES AND MECHANISMS
Heyes, C. M. (1994) · Biological reviews/Biological reviews of the Cambridge Philosophical Society · doi:10.1111/j.1469-185x.1994.tb01506.x · Cited by 1,023 · see citing works
Culture in whales and dolphins
Rendell, L., & Whitehead, H. (2001) · Behavioral and Brain Sciences · doi:10.1017/s0140525x0100396x · Cited by 858 · see citing works
Causes and consequences of imitation
Heyes, C. M. (2001) · Trends in Cognitive Sciences · doi:10.1016/s1364-6613(00)01661-2 · Cited by 620 · see citing works
Advances in the Study of Behavior
(2016) · Advances in the study of behavior · doi:10.1016/s0065-3454(16)30011-0 · Cited by 614 · see citing works
(2013) · Psychology Press eBooks · doi:10.4324/9781315801889 · Cited by 601 · see citing works
Imitation in Animals and Artifacts
Dautenhahn, K., & Nehaniv, C. (2002) · The MIT Press eBooks · doi:10.7551/mitpress/3676.001.0001 · Cited by 586 · see citing works
The different roles of social learning in vocal communication
Janik, V. M., & Slater, P. J. B. (2000) · Animal Behaviour · doi:10.1006/anbe.2000.1410 · Cited by 541 · see citing works
Imitation in a home-raised chimpanzee.
Hayes, K. J., & Hayes, C. (1952) · Journal of Comparative and Physiological Psychology · doi:10.1037/h0053609 · Cited by 456 · see citing works
The Cultural Lives of Whales and Dolphins
Whitehead, H., & Rendell, L. E. (2014) · University of Chicago Press eBooks · doi:10.7208/chicago/9780226187426.001.0001 · Cited by 359 · see citing works
Cetaceans have complex brains for complex cognition
Marino, L., Connor, R. C., Fordyce, R. E., Herman, L. M., Hof, P. R., Lefebvre, L., Lusseau, D., McCowan, B., Nimchinsky, E. A., Pack, A. A., et al. (2007) · PLoS Biology · doi:10.1371/journal.pbio.0050139 · Cited by 339 · see citing works
Chapter 3 Social Processes Influencing Learning in Animals: A Review of the Evidence
Hoppitt, W., & Laland, K. N. (2008) · Advances in the study of behavior · doi:10.1016/s0065-3454(08)00003-x · Cited by 327 · see citing works
Connor, R. C. (2007) · Philosophical Transactions of the Royal Society B: Biological Sciences · doi:10.1098/rstb.2006.1997 · Cited by 289 · see citing works
Selective Imitation in Domestic Dogs
Range, F., Virányi, Z., & Huber, L. (2007) · Current Biology · doi:10.1016/j.cub.2007.04.026 · Cited by 281 · see citing works
Primate Culture and Social Learning
Whiten, A. (2000) · Cognitive Science · doi:10.1207/s15516709cog2403_6 · Cited by 277 · see citing works
(1995) · Choice Reviews Online · doi:10.5860/choice.32-5060 · Cited by 270 · see citing works
Can Young Chimpanzees (Pan Troglodytes) Imitate Arbitrary Actions? Hayes & Hayes (1952) Revisited
Custance, D. M., Whiten, A., & Bard, K. A. (1995) · Behaviour · doi:10.1163/156853995x00036 · Cited by 244 · see citing works
What Is the Significance of Imitation in Animals?
Heyes, C. M., & Ray, E. D. (2000) · Advances in the study of behavior · doi:10.1016/s0065-3454(08)60106-0 · Cited by 228 · see citing works
Social evolution in toothed whales
Connor, R. C., Mann, J., Tyack, P. L., & Whitehead, H. (1998) · Trends in Ecology & Evolution · doi:10.1016/s0169-5347(98)01326-3 · Cited by 227 · see citing works
Cetacean vocal learning and communication
Janik, V. M. (2014) · Current Opinion in Neurobiology · doi:10.1016/j.conb.2014.06.010 · Cited by 211 · see citing works
Reproducing human actions and action sequences: “Do as I Do!” in a dog
Topál, J., Byrne, R. W., Miklósi, Á., & Csányi, V. (2006) · Animal Cognition · doi:10.1007/s10071-006-0051-6 · Cited by 158 · see citing works
Vocal, Social, and Self-Imitation by Bottlenosed Dolphins
Herman, L. M. (2002) · The MIT Press eBooks · doi:10.7551/mitpress/3676.003.0004 · Cited by 157 · see citing works
Imitation of novel complex actions: What does the evidence from animals mean?
Byrne, R. W. (2002) · Advances in the study of behavior · doi:10.1016/s0065-3454(02)80006-7 · Cited by 143 · see citing works
Huber, L., Range, F., Voelkl, B., Szucsich, A., Virányi, Z., & Miklósi, Á. (2009) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2009.0060 · Cited by 140 · see citing works
Three Sources of Information in Social Learning
Call, J., & Carpenter, M. (2002) · The MIT Press eBooks · doi:10.7551/mitpress/3676.003.0010 · Cited by 137 · see citing works
IMITATION IN ANIMALS: EVIDENCE, FUNCTION, AND MECHANISMS
Zentall, T. R. (2001) · Cybernetics & Systems · doi:10.1080/019697201300001812 · Cited by 131 · see citing works
Animal Behaviour: Evolution and Mechanisms
Kappeler, P. M. (2010) · Springer eBooks · doi:10.1007/978-3-642-02624-9 · Cited by 124 · see citing works
THE EVOLUTION OF INTELLIGENCE (2):
(2010) · Harvard University Press eBooks · doi:10.2307/j.ctvjf9xvk.24 · Cited by 121 · see citing works
BODY IMITATION IN AN ENCULTURATED ORANGUTAN (PONGO PYGMAEUS)
Call, J. (2001) · Cybernetics & Systems · doi:10.1080/019697201300001821 · Cited by 105 · see citing works
Intelligence and rational behaviour in the bottlenosed dolphin
Herman, L. M. (2006) · Oxford University Press eBooks · doi:10.1093/acprof:oso/9780198528272.003.0020 · Cited by 91 · see citing works
Social learning and culture in animals
van Schaik, C. P. (2010) · Springer eBooks · doi:10.1007/978-3-642-02624-9_20 · Cited by 86 · see citing works
Spontaneous human speech mimicry by a cetacean
Ridgway, S. H., Carder, D. A., Jeffries, M., & Todd, M. A. (2012) · Current Biology · doi:10.1016/j.cub.2012.08.044 · Cited by 79 · see citing works
Simon says: The development of imitation in an enculturated orangutan.
Miles, H. L., Mitchell, R. W., & Harper, S. E. (1996) · Cited by 63 · see citing works
What Can Imitation Do for Cooperation?
(2013) · The MIT Press eBooks · doi:10.7551/mitpress/9033.003.0020 · Cited by 56 · see citing works
Fugazza, C., & Miklósi, Á. (2015) · Applied Animal Behaviour Science · doi:10.1016/j.applanim.2015.08.033 · Cited by 55 · see citing works
Cultural evolution of killer whale calls: background, mechanisms and consequences
Filatova, O. A., Samarra, F. I. P., Deecke, V. B., Ford, J. K. B., Miller, P. J. O., & Yurk, H. (2015) · Behaviour · doi:10.1163/1568539x-00003317 · Cited by 51 · see citing works
Trained Motor Imitation by Bottlenose Dolphins ( Tursiops Truncatus )
Bauer, G. B., & Johnson, C. M. (1994) · Perceptual and Motor Skills · doi:10.2466/pms.1994.79.3.1307 · Cited by 51 · see citing works
Experimental evidence for action imitation in killer whales (Orcinus orca)
Zamorano-Abramson, J., Hernández‐Lloreda, M. V., Call, J., & Colmenares, F. (2012) · Animal Cognition · doi:10.1007/s10071-012-0546-2 · Cited by 49 · see citing works
Galef, B. G. (2013) · Behavioural Processes · doi:10.1016/j.beproc.2013.07.026 · Cited by 49 · see citing works
Dissecting the imitation faculty: The multiple imitation mechanisms (MIM) hypothesis
Subiaul, F. (2009) · Behavioural Processes · doi:10.1016/j.beproc.2009.12.002 · Cited by 43 · see citing works
Imitation: what animal imitation tells us about animal cognition
Bates, L. A., & Byrne, R. W. (2010) · Wiley Interdisciplinary Reviews Cognitive Science · doi:10.1002/wcs.77 · Cited by 43 · see citing works
Animal behaviour: evolution and mechanisms
(2010) · Choice Reviews Online · doi:10.5860/choice.48-1464 · Cited by 37 · see citing works
Mitchell, R. W., & Anderson, J. R. (1993) · Primates · doi:10.1007/bf02382625 · Cited by 36 · see citing works
Dogs, Canis familiaris, fail to copy intransitive actions in third-party contextual imitation tasks
Tennie, C., Glabsch, E., Tempelmann, S., Bräuer, J., Kaminski, J., & Call, J. (2009) · Animal Behaviour · doi:10.1016/j.anbehav.2009.03.008 · Cited by 30 · see citing works
Blindfolded Imitation in a Bottlenose Dolphin ( Tursiops truncatus )
Jaakkola, K., Guarino, E., & Rodriguez, M. (2010) · International Journal of Comparative Psychology · doi:10.46867/ijcp.2010.23.04.08 · Cited by 29 · see citing works
McDonald, F. J. (2014) · Philosophical Psychology · doi:10.1080/09515089.2014.945519 · Cited by 28 · see citing works
Observational Learning in Wild and Captive Dolphins
Yeater, D. B., & Kuczaj, S. A. (2010) · International Journal of Comparative Psychology · doi:10.46867/ijcp.2010.23.03.04 · Cited by 26 · see citing works
Byrne, R. W. (2009) · Current Biology · doi:10.1016/j.cub.2008.11.027 · Cited by 24 · see citing works
Switching strategies: a dolphin’s use of passive and active acoustics to imitate motor actions
Jaakkola, K., Guarino, E., Rodriguez, M., & Hecksher, J. (2013) · Animal Cognition · doi:10.1007/s10071-013-0605-3 · Cited by 19 · see citing works
Zentall, T. R. (2011) · Interaction Studies Social Behaviour and Communication in Biological and Artificial Systems · doi:10.1075/is.12.2.03zen · Cited by 19 · see citing works
Preliminary Study of Object Labeling Using Sound Production in a Beluga
Murayama, T., Fujii, Y., Hashimoto, T., Shimoda, A., Iijima, S., Hayasaka, K., Shiroma, N., Koshikawa, M., Katsumata, H., Soichi, M., et al. (2012) · International Journal of Comparative Psychology · doi:10.46867/ijcp.2012.25.03.04 · Cited by 12 · see citing works
Cetacean Cognitive Specializations
Jaakkola, K. (2012) · Oxford University Press eBooks · doi:10.1093/oxfordhb/9780199738182.013.0009 · Cited by 11 · see citing works
Imitation, Social Learning, and Cultural Traditions
Dean, L. G., Vale, G. L., & Whiten, A. (2018) · The International Encyclopedia of Anthropology · doi:10.1002/9781118924396.wbiea1622 · Cited by 7 · see citing works
Flagged first, then by citations
1 of the 55 references couldn’t be matched to a known paper.
Research works printed but not linked: 2
- [46]Topál J, Byrne RW, Miklósi A, Csányi V. Reproducing human actions and action sequences:“Do as I Do!” in a dog. Anim Cog. 2006; 9: 355– 367. 17024511
- [50]Xitco MJ. Mimicry of modeled behaviours by bottlenose dolphins. Unpublished Master of Science thesis, University of Hawaii, Honolulu; 1988.
Cited by
Sorted by citations
Imitation of novel conspecific and human speech sounds in the killer whale ( Orcinus orca )
Zamorano-Abramson, J., Hernández‐Lloreda, M. V., García, L., Colmenares, F., Aboitiz, F., & Call, J. (2018) · Proceedings of the Royal Society B Biological Sciences · doi:10.1098/rspb.2017.2171 · Cited by 39 · see citing works
The development of socio-sexual behavior in belugas (Delphinapterus leucas) under human care
Lilley, M. K., Ham, J. R., & Hill, H. M. M. (2019) · Behavioural Processes · doi:10.1016/j.beproc.2019.104025 · Cited by 34 · see citing works
Ham, J. R., Lilley, M. K., Lelekach, J., Miller, M. R., Robeck, T. R., Pellis, S. M., & Hill, H. M. M. (2022) · Behavioural Processes · doi:10.1016/j.beproc.2022.104695 · Cited by 22 · see citing works
Orcas remember what to copy: a deferred and interference-resistant imitation study
Zamorano-Abramson, J., Hernández‐Lloreda, M. V., Colmenares, F., & Call, J. (2023) · Animal Cognition · doi:10.1007/s10071-023-01756-3 · Cited by 17 · see citing works
Cetacean Brain, Cognition, and Social Complexity
Marino, L. (2022) · Ethology and behavioral ecology of marine mammals · doi:10.1007/978-3-030-98100-6_4 · Cited by 9 · see citing works
Multimodal imitative learning and synchrony in cetaceans: A model for speech and singing evolution
Zamorano-Abramson, J., Michon, M., Hernández‐Lloreda, M. V., & Aboitiz, F. (2023) · Frontiers in Psychology · doi:10.3389/fpsyg.2023.1061381 · Cited by 8 · see citing works
Hill, H. M. M., Lilley, M. K., Ham, J. R., & Robeck, T. R. (2024) · Theriogenology Wild · doi:10.1016/j.therwi.2024.100071 · Cited by 7 · see citing works
Marino, L. (2020) · Advances in neuroethics · doi:10.1007/978-3-030-31011-0_10 · Cited by 7 · see citing works
Odontocetes (‘toothed whales’): cognitive science and moral standing–are dolphins persons?
Pettit, S. G., & McCulloch, S. P. (2023) · Journal of Applied Animal Ethics Research · doi:10.1163/25889567-bja10033 · Cited by 5 · see citing works
Minding the Minds: A Primer on Cognitive Challenge for Marine Mammals in Human Care
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Zamorano-Abramson, J., & Hernández-Lloreda, M. V. (2023) · Animals · doi:10.3390/ani13243763 · Cited by 2 · see citing works
Self-citationThird-party imitation is not restricted to humans
Haldar, E., Sanchez, A. H., Tennie, C., Ortiz, S. T., Vos, J., Valbert, M., & von Bayern, A. V. (2025) · bioRxiv · doi:10.1038/s41598-025-11665-9 · Cited by 2 · see citing works
Vocal Imitation, A Specialized Brain Function That Facilitates Cultural Transmission in Songbirds
Tanaka, M. (2023) · Springer eBooks · doi:10.1007/978-981-99-0831-8_5 · Cited by 1 · see citing works
O’Sullivan, E., & Caldwell, C. A. (2018) · Springer eBooks · doi:10.1007/978-3-319-47829-6_1871-1 · Cited by 1 · see citing works
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“They have also demonstrated the ability for behavioral imitation during play [44] and the capacity for contextual imitation of intransitive and novel body actions [45,46].”
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Third-party imitation is not restricted to humansPreprint
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- Authors
- Abramson, J. Z., Hernández-Lloreda, M. V., Esteban, J.-A., Colmenares, F., Aboitiz, F., & Call, J.
- Affiliations
- Universidad Complutense de Madrid (ES)
- Pontificia Universidad Católica de Chile (CL)
- University of St Andrews (GB)
- Venue
- PLoS ONE
- Publisher
- Public Library of Science (PLoS)
- Published
- 2017-06-21
- Volume
- 12
- Issue
- 6
- Pages
- e0178906
- ISSN
- 1932-6203 (ISSN-L)
- PMC
- PMC5479519
- OpenAlex
- W2660320799
- Cited by
- 20
- References
- 55
- Citations by year
- 2026: 2 · 2025: 3 · 2024: 2 · 2023: 5 · 2022: 3 · 2020: 1 · 2019: 2 · 2018: 2
- FWCI
- 1.03
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Licence
- Locations
- PLoS ONE (publishedVersion, cc-by)
- Europe PMC (PubMed Central) (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- MPG.PuRe (Max Planck Society) (submittedVersion, cc-by)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- Max Planck Digital Library (submittedVersion, other-oa)
- Figshare (submittedVersion, cc-by)
- St Andrews Research Repository (St Andrews Research Repository) (submittedVersion, cc-by)
- Funders
- Fondo Nacional de Desarrollo Científico y Tecnológico (3140580)
- Ministerio de Economía y Competitividad (PSI2011-29016-C02-01, PSI2014-51890-C2-1-P)
- Fundación Banco Santander (UCM-BSCH GR3/14-940813)
- Fondo Nacional de Desarrollo Científico y Tecnológico (CL) (1160258)
- European Research Council under the European Union's Seventh Framework Programme ((FP7/2007-2013) / ERC grant SOMICS agreement n° 609819)
- Topics
- Physical Sciences › Environmental Science › Ecology › Marine animal studies overview
- Social Sciences › Psychology › Social Psychology › Primate Behavior and Ecology
- Social Sciences › Cultural Studies › Language and cultural evolution
- Keywords
- beluga whale, imitation, social learning, behavioral generalization, animal cognition, cetacean social behavior
- MeSH terms
- Animals, Auditory Perception, Behavior, Animal, Female, Imitative Behavior, Learning, Male, Motor Activity, Psychomotor Performance, Beluga Whale
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