Watching the ducks on a neighborhood pond can be an entertaining and rewarding pastime. I myself, along with my nine-year-old co-investigator, have taken daily opportunities to feed some ducks on a nearby pond over the past several months. In doing so, we not only had fun but also managed to conduct some urban science that led us to a new scientific discovery: Mallards (Anas platyrhynchos L.) engage in indirect reciprocity with humans. Scientists have known for decades, of course, that indirect reciprocity was critical to the evolution of human social interaction in large-scale societies, but we believe we are the first to identify indirect reciprocity at work in human-duck social interaction.
Here’s how we made this discovery.
On random days, we take a soda cracker along with us to feed to a single lucky duck. On the other days, we take our walks without a cracker. What my young co-investigator and I have noticed is that on cracker days, after we’ve fed the cracker to the first duck that approaches us (the “focal duck,” which we also call “the recipient”), other ducks (which we call “entertainment ducks,” or “indirect reciprocators”) appear to take notice of our generosity toward the recipient. Almost immediately, the indirect reciprocators start to perform all sorts of entertaining behaviors: They swim toward us eagerly, they waddle up to us enthusiastically, they stare at us with their dead, obsidian eyes, they quack imploringly. It’s all very amusing and my co-investigator and I have a great time. Take note of the fact that we always bring only a single cracker with us on cracker days. As a result, the indirect reciprocators have absolutely nothing to gain from the entertainment they provide. In fact, they actually incur costs (in the form of energy expended and lost foraging time) when they do so. Thus, their indirect reciprocity behavior is altruistic.
Our experience with the indirect reciprocators is very different on non-cracker days. If a focal duck comes up to us on a non-cracker day, there’s just no cracker to be had, no matter how charming or insistent the request. Dejected, the focal duck typically waddles or paddles away within a few seconds. Now, what do you suppose the entertainment ducks do after we refuse to feed the focal duck? That’s right. They withhold their entertainment behaviors. This pattern, of course, is exactly as one would expect if the entertainment ducks were regulating their entertainment behaviors according to the logic of indirect reciprocity.
Theorists typically assume that the computational demands for indirect reciprocity to evolve are quite extensive. For instance, indirect reciprocators need to possess computational machinery that enables them to acquire information about the actions of donors—either through direct sensory experience of donor-recipient interactions, or (more rarely) language-based gossip, or (even more rarely) social information stored in an external medium, such written records or the reputational information that’s often available in online markets. Indirect reciprocators also need be able to tag donors’ actions toward recipients as either “beneficial” or “non-beneficial,” store that social information in memory, and then feed that information to motivational systems that can produce the indirect reciprocity behaviors that will serve as rewards to donors. However, the indirect reciprocity we’ve identified in our mallards suggests that those computational requirements may be fulfilled in vertebrates more commonly than theorists originally thought.
Neither of us could figure out for sure whether the focal ducks were transmitting information about our generosity/non-generosity to the indirect reciprocators through verbal (or non-verbal) communication, but we think it is unlikely. Instead, we suspect that the indirect reciprocators were directly observing our behavior and then using that sensory information to regulate their indirect reciprocity behavior.
In support of this interpretation, we note that on several cracker days, it was not only other ducks that engaged us as indirect reciprocators, but individuals from two different species of turtles (which we believe to be Rachemys scripta and Apalone ferox) as well. The turtles’ indirect reciprocity behaviors, of course, were different from those of the ducks, due to differences in life history and evolutionary constraints: The turtles didn’t reward our generosity through waddle-based or quack-based rewarding, but rather, by (a) rooting around in the mud where the focal duck had received the cracker earlier, and (b) trying to grab the focal duck by the leg and drag it to a gruesome, watery death. The fact that turtles engaged in their own forms of indirect reciprocity suggests that they, at least, were obtaining information about our generosity via direct sensory experience, rather than through duck-turtle communication or written or electronic records: It is widely accepted, after all, that turtles don’t understand Mallardese or use eBay.
The involvement of turtles as indirect reciprocators also suggests that indirect reciprocity might be even more prevalent–and more complex–than even we originally suspected. Not only does indirect reciprocity evolve to regulate interactions within species (viz., Homo sapiens), and between species (viz., between Homo sapiens and Anas platyrhynchos L., as we have documented here), but also among species (Homo sapiens as donors, Anas platyrhynchos L. as recipients, and Rachemys scripta and Apalone ferox as indirect reciprocators).
Finally, we should point out that although our results are consistent with the indirect reciprocity interpretation that we have proffered here, other interpretations are possible as well. We look forward to new work that can arbitrate between these two accounts (and perhaps others). We also see excellent opportunities for simulation studies that can shed light on the evolution of indirect reciprocity involving interactions between two or even three different species, which my co-Investigator thinks she might pursue after she has mastered long division.
h/t Eric P.