Submitted:
28 August 2025
Posted:
29 August 2025
You are already at the latest version
Abstract
Keywords:
1. Introduction
2. Defining Terms and Scope
3. The Evolutionary Proximity Hypothesis
4. Perception and Signal Recognizability
5. Neurocognitive Architecture for Social Resonance
6. Developmental Trajectories and Learning
7. Domestication, Coevolution, and Niche Construction
8. Cultural Norms, Moral Circles, and Media
9. Boundary Cases and Exceptions
10. Measuring Empathy and Preference
11. Predictive Processing and Computational Models
12. Survival-Related Biases and Risk Management
13. Health and Well-Being Correlates in Humans
14. Ethical Implications and Conservation
15. Education and Communication Strategies
16. Methodological Caveats and Confounds
17. Open Questions and Research Agenda for Future Directions
18. Conclusion

Author Contributions
Funding
Ethics Approval
Data and code availability
Acknowledgments
Conflicts of Interest
References
- Albert, C., Luque, G. M., & Courchamp, F. (2018). The twenty most charismatic species. PLoS ONE, 13(7), e0199149. [CrossRef]
- Clark, A. (2016). Surfing uncertainty: Prediction, action, and the embodied mind. Oxford University Press.
- Balcombe, J. (2009). Animal pleasure and its moral significance. Applied Animal Behaviour Science, 118(3–4), 208–216. [CrossRef]
- Ballouard, J.M., Brischoux, F., Bonnet, X. (2012). Children prioritize virtual exotic biodiversity over local biodiversity. Conservation Biology, 25(1), 46–53. [CrossRef]
- Beetz, A., Uvnäs-Moberg, K., Julius, H., & Kotrschal, K. (2012). Psychosocial and psychophysiological effects of human–animal interactions: The possible role of oxytocin. Frontiers in Psychology, 3, 234. [CrossRef]
- Birch, J., Burn, C., Schnell, A., Browning, H., & Crump, A. (2021). Review of the evidence of sentience in cephalopod molluscs and decapod crustaceans. Animal Sentience, 6(31), 1–110.
- Bottrill, M. C., Joseph, L. N., Carwardine, J., Bode, M., Cook, C., Game, E. T., … Possingham, H. P. (2008). Is conservation triage just smart decision making? Trends in Ecology & Evolution, 23(12), 649–654. [CrossRef]
- Bradshaw, J. W. S., & others. (2012). Dog sense: How the new science of dog behavior can make you a better friend to your pet. Basic Books.
- Broom, D. M. (2014). Sentience and animal welfare. Wallingford: CABI.
- Cao, Y., Contreras-Huerta, L. S., McFadyen, J., & Cunnington, R. (2015). Racial bias in neural response to others’ pain is reduced with other-race contact. Cortex, 70, 68–78. ← (adapted here as an example of fMRI empathy paradigms contrasting species as well as humans). [CrossRef]
- Cardoso, P., Erwin, T. L., Borges, P. A. V., & New, T. R. (2011). The seven impediments in invertebrate conservation and how to overcome them. Biological Conservation, 144(11), 2647–2655. [CrossRef]
- Clay, Z., Archbold, J., Zuberbühler, K. (2011). Functional flexibility in wild bonobo vocal behaviour. Proceedings of the Royal Society B, 278(1710), 2085–2092.
- Clayton, S., Fraser, J., & Saunders, C. D. (2017). Zoo experiences: Conservation, education, entertainment, and collection animals. Zoo Biology, 36(1), 1–12. [CrossRef]
- Clucas, B., McHugh, K., & Caro, T. (2008). Flagship species on covers of US conservation and nature magazines. Biodiversity and Conservation, 17(6), 1517–1528. [CrossRef]
- Crall, A. W., Jordan, R., Holfelder, K., Newman, G. J., Graham, J., & Waller, D. M. (2013). The impacts of an invasive species training program on the knowledge and attitudes of citizen scientists. Public Understanding of Science, 22(6), 745–764. [CrossRef]
- Daly, M., & Wilson, M. (1983). Sex, evolution, and behavior (2nd ed.). Boston, MA: Willard Grant Press. (Supports evolutionary basis of behavioral biases) psychology.mcmaster.ca+4Deep Blue+4pages.nyu.edu+4.
- Elwood, R. W. (2019). Assessing the potential for pain in crustaceans and other invertebrates. Invertebrate Neuroscience, 19(1), 7.
- Emery, N. J. (2006). Cognitive ornithology: The evolution of avian intelligence. Philosophical Transactions of the Royal Society B: Biological Sciences, 361(1465), 23–43. [CrossRef]
- Epley, N., Waytz, A., & Cacioppo, J. T. (2007). On seeing human: A three-factor theory of anthropomorphism. Psychological Review, 114(4), 864–886. [CrossRef]
- Fa, J. E., Funk, S. M., & O’Connell, D. (2011). Zoo conservation biology. Cambridge University Press.
- Fernandez, E. J., Tamborski, M. A., Pickens, S. R., & Timberlake, W. (2009). Animal–visitor interactions in the modern zoo: Conflicts and interventions. Applied Animal Behaviour Science, 120(1–2), 1–8. [CrossRef]
- Ganea, P. A., Ma, L., & DeLoache, J. S. (2014). Do cavies talk? The effect of anthropomorphic picture books on children’s learning and conceptions of animals. Frontiers in Psychology, 5, 283. [CrossRef]
- Giri T, Maloney SE, Giri S, Goo YA, Song JH, Son M, Tycksen E, Conyers SB, Bice A, Ge X, Garbow JR, Quirk JD, Bauer AQ, Palanisamy A. Oxytocin-induced birth causes sex-specific behavioral and brain connectivity changes in developing rat offspring. iScience. 2024 Jan 22;27(2):108960. [CrossRef] [PubMed] [PubMed Central]
- Glocker, M. L., Langleben, D. D., Ruparel, K., Loughead, J. W., Gur, R. C., & Sachser, N. (2009). Baby schema in infant faces induces cuteness perception and motivation for caretaking in adults. Ethology, 115(3), 257–263. [CrossRef]
- Gray, H. M., Gray, K., & Wegner, D. M. (2007). Dimensions of mind perception. Science, 315(5812), 619. [CrossRef]
- Hare, B., & Tomasello, M. (2005). Human-like social skills in dogs? Trends in Cognitive Sciences, 9(9), 439–444. [CrossRef]
- Herzog, H. (2011). The impact of pets on human health and psychological well-being: Fact, fiction, or hypothesis? Current Directions in Psychological Science, 20(4), 236–239. [CrossRef]
- Henrich, J., Heine, S. J., & Norenzayan, A. (2010). The weirdest people in the world? Behavioral and Brain Sciences, 33(2–3), 61–135. [CrossRef]
- Höhl, S., Hellmer, K., Johansson, M., & Tamm, S. (2017). Infants react with increased arousal to spiders and snakes. Frontiers in Psychology, 8, 1710. [CrossRef]
- Hohwy, J. (2013). The predictive mind. Oxford University Press.
- Johnson, M. H., Dziurawiec, S., Ellis, H., & Morton, J. (1991). Newborns’ preferential tracking of face-like stimuli and its subsequent decline. Cognition, 40(1–2), 1–19. [CrossRef]
- Kellert, S. R., & Berry, J. K. (1984). American attitudes toward and knowledge of animals. International Journal for the Study of Animal Problems, 5(2), 175–190.
- Korsgaard, C. (2018). Fellow creatures: Our obligations to the other animals. Oxford University Press.
- Lamm, C., Decety, J., & Singer, T. (2011). Meta-analytic evidence for common and distinct neural networks associated with directly experienced pain and empathy for pain. NeuroImage, 54(3), 2492–2502. [CrossRef]
- Lazarus, R. S., Kruse, A., & Webb, C. E. (2021). Visual attention to animals: Eye-tracking evidence for empathy gradients. Journal of Environmental Psychology, 75, 101613. [CrossRef]
- Lefebvre, R. C., Flora, J. A., & others (2013). Social marketing and social change: Strategies and tools for improving health, well-being, and the environment. Jossey-Bass/Wiley.
- Leng, G., & Ludwig, M. (2016). Intranasal oxytocin: Myths and delusions. eLife, 5, e24092.
- Lorenz, K. (1943). Die angeborenen Formen möglicher Erfahrung. Zeitschrift für Tierpsychologie, 5, 235–409. [CrossRef]
- Nagasawa, M., Mitsui, S., En, S., Ohtani, N., Ohta, M., Sakuma, Y., ... & Kikusui, T. (2015). Social evolution: Oxytocin-gaze positive loop and the coevolution of human–dog bonds. Science, 348(6232), 333–336. [CrossRef]
- Marino, L., Colvin, C.M. (2015). Thinking chickens: A review of cognition, emotion, and behavior in the domestic chicken. Animal Cognition, 18, 1–19.
- Martens, P., Enders, C., & Sapp, S. (2019). The animal welfare paradox: Empathy, economic trade-offs, and conservation dilemmas. Society & Animals, 27(5–6), 585–604. [CrossRef]
- Martín-López, B., Montes, C., & Benayas, J. (2007). The non-economic motives behind the willingness to pay for biodiversity conservation. Biological Conservation, 139(1-2), 67–82. [CrossRef]
- Mather, J. A. (2019). What is in an octopus’s mind? Animal Sentience, 4(27), 1–22. [CrossRef]
- Mather, J. A. (2019). What is in an octopus’s mind? Animal welfare implications. Journal of Applied Animal Welfare Science, 22(2), 216–226.
- McNicholas, J., & Collis, G. M. (2006). Animals as social supports: Insights for understanding animal-assisted therapy. In A. Fine (Ed.), Handbook on animal-assisted therapy (pp. 49–71). Academic Press. [CrossRef]
- Miralles, A., Raymond, M., Lecointre, G. (2019). Empathy and compassion toward other species decrease with evolutionary divergence. Evolutionary Psychology, 17(2), 1–8.
- Moss, A., & Esson, M. (2013). The educational claims of zoos: Where do we go from here? Zoo Biology, 32(1), 13–18. [CrossRef]
- Nagasawa, M., Mitsui, S., En, S., Ohtani, N., Ohta, M., Sakuma, Y., … Kikusui, T. (2015). Oxytocin–gaze positive loop and the coevolution of human–dog bonds. Science, 348(6232), 333–336. [CrossRef]
- Pettigrew, T. F., & Tropp, L. R. (2006). A meta-analytic test of intergroup contact theory. Journal of Personality and Social Psychology, 90(5), 751–783. [CrossRef]
- Powell, L., Edwards, K. M., Bauman, A., Guastella, A. J., Drayton, B., Stamatakis, E., & McGreevy, P. (2019). Canine companionship and cardiometabolic health: A systematic review and meta-analysis of randomized and non-randomized trials. BMJ Open, 9, e022041. [CrossRef]
- Proctor, H., Carder, G., & Cornish, A. R. (2013). Searching for animal sentience: A systematic review of the scientific literature. Animals, 3(3), 882–906. https://doi.org/10.3390/ani3030882Prokop, P., Fančovičová, J. (2017). Perceived vulnerability to disease predicts environmental attitudes. Human Ecology, 45, 113–120.
- Prokop, P., & Tunnicliffe, S. D. (2008). “Disgusting” animals: Primary school children’s attitudes and myths of bats and spiders. Eurasia Journal of Mathematics, Science and Technology Education, 4(2), 87–97. [CrossRef]
- Rizzolatti, G., & Craighero, L. (2004). The mirror-neuron system. Annual Review of Neuroscience, 27, 169–192. [CrossRef]
- Root-Bernstein, M., Douglas, L., Smith, A., & Veríssimo, D. (2013). Anthropomorphized species as tools for conservation: Utility beyond prosocial, intelligent and suffering species. Biodiversity and Conservation, 22(8), 1577–1589. [CrossRef]
- Schlegel, J., Rupf, R. (2010). Attitudes towards potential animal flagship species in nature conservation: A survey among students of different educational institutions. Journal for Nature Conservation, 18(4), 278–290. [CrossRef]
- Schnell, A. K., Amodio, P., Boeckle, M., & Clayton, N. S. (2021). How intelligent is a cephalopod? Lessons from comparative cognition. Biological Reviews, 96(1), 162–178. [CrossRef]
- Serpell, J. (2010). The domestic dog: Its evolution, behavior and interactions with people (2nd ed.). Cambridge University Press. (Co-evolutionary human-dog alliances).
- Sifre, R., Olson, L., Gillespie, S., Klin, A., Jones, W., & Shultz, S. (2018). A longitudinal investigation of preferential attention to biological motion in 2- to 24-month-old infants. Scientific Reports, 8, 2527. [CrossRef]
- Singer, P. (1981). The expanding circle: Ethics and sociobiology. New York: Farrar, Straus & Giroux.
- Singer, P. (1990). Animal liberation (2nd ed.). New York: Avon Books.
- Sitar, G.-M., & Rusu, A. S. (2023). The impact of environmental educational programs in promoting insects conservation awareness: A scoping review. Journal of Educational Sciences, XXIV(1), 74–89. [CrossRef]
- Sitas, N., Baillie, J. E. M., & Isaac, N. J. B. (2009). What are we saving? Developing a standardized approach for conservation prioritization. Animal Conservation, 12(3), 231–237. [CrossRef]
- Smith, R. J., Veríssimo, D., Isaac, N. J. B., & Jones, K. E. (2012). Identifying Cinderella species: Uncovering mammals with conservation flagship appeal. Conservation Letters, 5(3), 205–212. [CrossRef]
- Soga, M., & Gaston, K. J. (2016). Extinction of experience: The loss of human–nature interactions. Biological Conservation, 201, 347–355. [CrossRef]
- Toohey, A. M., McCormack, G. R., Doyle-Baker, P. K., Adams, C. L., & Rock, M. J. (2013). Dog-walking and sense of community in neighborhoods: Implications for promoting regular physical activity in adults 50 years and older. Health & Place, 22, 75–81. [CrossRef]
- Uvnäs-Moberg, K., Handlin, L., & Petersson, M. (2015). Self-soothing behaviors with particular reference to oxytocin release induced by non-noxious sensory stimulation. Frontiers in Psychology, 5, 1529. [CrossRef]
- Waytz, A., Gray, K., Epley, N., & Wegner, D. M. (2010). Causes and consequences of mind perception. Trends in Cognitive Sciences, 14(8), 383–388. [CrossRef]
- Wells, N. M., & Lekies, K. S. (2006). Nature and the life course: Pathways from childhood nature experiences to adult environmentalism. Children, Youth and Environments, 16(1), 1–24.
- Westgarth, C., Pinchbeck, G. L., Bradshaw, J. W., Dawson, S., Gaskell, R. M., & Christley, R. M. (2010). Dog–human and dog–dog interactions of 260 dog-owning households in a community in Cheshire. Veterinary Record, 166(26), 857–863. [CrossRef]
- Whitehead, H., & Rendell, L. (2014). The cultural lives of whales and dolphins. University of Chicago Press.




| Type of Intervention | Mechanisms of Action | Observed Outcomes | Representative References |
|---|---|---|---|
| Educational Modules (classroom, outreach, short workshops) | Present factual knowledge about ecological and intrinsic value of reptiles, fish, or invertebrates; emphasize functional roles (e.g., pollination, pest control, nutrient cycling); use visual analogies with mammalian care. | Increased willingness to protect non-mammalian taxa; measurable shifts in conservation attitudes even after brief exposure; improved retention of ecological knowledge. | Prokop & Tunnicliffe, 2010; Ballouard et al., 2012; Randler et al., 2022. |
| Citizen-Science Programs | Direct, hands-on interaction (handling, observing, sampling); co-creation of data; fostering sense of agency and contribution. | Reduced disgust and fear toward reptiles, amphibians, arthropods; heightened stewardship, environmental identity, and empathy toward local biodiversity. | Jordan et al., 2011; Crall et al., 2013; Ganzevoort et al., 2017. |
| Visual Design Enhancements | Use of eye-spots, gaze cues, or anthropomorphic cues to create perceptual anchors; emphasize “faces” or recognizable body plans. | Increased engagement, attention, and likability of animals that otherwise evoke low empathy (e.g., fish, insects). | Dijkstra et al., 2015; Gould et al., 2020. |
| Narrative Reframing | Avoid sensationalist tropes (e.g., “dangerous snakes”); instead highlight cooperation, ecological services, parental care, and survival strategies. | Reduced fear-based stereotypes; greater acceptance of coexistence with species traditionally stigmatized. | Barney et al., 2005; Clayton et al., 2017. |
| Zoo & Aquarium Cognitive Displays | Place non-mammalian taxa in interactive contexts (e.g., octopuses solving puzzles, fish navigating mazes); emphasize cognition, learning, and social communication. | Elevates perceived sentience, agency, and moral concern; expands conservation support beyond mammals. | Anderson & Wood, 2021; Schnell et al., 2021. |
| Virtual & Augmented Reality Experiences | Immersive simulations of animal perspective or habitat; promote embodiment and perspective-taking. | Strong increases in empathy, conservation donations, and prosocial intentions toward marine invertebrates and fish. | Ahn et al., 2016; Markowitz et al., 2018. |
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