Abstract
The group augmentation hypothesis states that individuals gain benefits by increasing group size and is rarely studied because it often overlaps with other evolutionary mechanisms like reciprocity. When rescuing a fellow prey from death, the mere presence of that member in the group can yield different benefits that can be passive like risk dilution or active like delayed reciprocity. We were able to separate the effects of passive and active group augmentation benefits by experimentally manipulating the behaviour of prey (altruist vs. non-altruist) in a video game where the costs and benefits of altruism were easily measured. The game pits four players-prey that must acquire resources to survive, while avoiding getting captured by a fifth player-predator. We instructed half of the prey players to avoid rescuing other prey, and the other half to rescue when possible. Our data showed that increasing the frequency of altruistic prey in a group increased survival of all group members. Maintaining group size yielded passive benefits like improving resource acquisition and facilitating future rescues. These passive benefits had a higher impact on survival than the active benefits (e.g., being rescued in return through reciprocity). Thus, we were able to support the importance of passive benefits from anti-predator altruistic behaviour using a biologically relevant online video game.
Significance statement
Altruism can generate different benefits at once. Moreover, these benefits, like reciprocity or group augmentation, are often intertwined so distinguishing them in empirical data is complex. We were able to measure different benefits related to altruism and its effect on group size, using data from a multiplayer video game where players could rescue each other from predation and elimination. Rescuing others was risky but was beneficial to the survival of all the other group members. Benefits from belonging in a larger group, facilitated resource acquisition and rescues, were the most important benefits of altruism in our system, whereas reciprocity, a largely studied mechanism, was very rare. Our study adds important evidence of a mechanism, group augmentation, that is often overlooked or ignored when studying the selection of anti-predatory cooperative behaviours.





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Data availability
The datasets generated during the experiment and analyzed during the current study are available in an OSF repository under a Attribution-NonCommercial-NoDerivs 4.0 International license.
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Acknowledgements
We thank Behaviour Interactive for their collaborative partnership in which they generously provide data and logistical support. We also thank the Player Insight team at Behaviour Interactive and the participants of our study. Finally, we are thankful to the members of GREEIB at UQAM and multiple anonymous reviewers for their feedback.
Funding
This research was possible due to an internship at Behaviour Interactive that was supported by grants from Mitacs (ref. IT06306) and Behaviour Interactive. Funding was also provided by a Natural Sciences and Engineering Research Council of Canada (NSERC) Discovery Grant to CDK.
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Céré, J., Kelly, C.D. & Montiglio, PO. Untangling the contribution of active and passive group augmentation benefits to the multilevel selection of altruism using a video game. Behav Ecol Sociobiol 78, 66 (2024). https://doi.org/10.1007/s00265-024-03463-6
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DOI: https://doi.org/10.1007/s00265-024-03463-6

