Humans can spontaneously create rules that allow them to efficiently generalize what they have learned to novel situations. An enduring question is whether rule-based generalization is uniquely human or whether other animals can also abstract rules and apply them to novel situations. In recent years, there have been a number of high-profile claims that animals such as rats can learn rules. Most of those claims are quite weak because it is possible to demonstrate that simple associative systems (which do not learn rules) can account for the behavior in those tasks. Using a procedure that allows us to clearly distinguish feature-based from rule-based generalization (the Shanks–Darby procedure), we demonstrate that adult humans show rule-based generalization in this task, while generalization in rats and pigeons was based on featural overlap between stimuli. In brief, when learning that a stimulus made of two components (“AB”) predicts a different outcome than its elements (“A” and “B”), people spontaneously abstract an opposites rule and apply it to new stimuli (e.g., knowing that “C” and “D” predict one outcome, they will predict that “CD” predicts the opposite outcome). Rats and pigeons show the reverse behavior—they generalize what they have learned, but on the basis of similarity (e.g., “CD” is similar to “C” and “D”, so the same outcome is predicted for the compound stimulus as for the components). Genuinely rule-based behavior is observed in humans, but not in rats and pigeons, in the current procedure. Electronic supplementary materialThe online version of this article (doi:10.1007/s10071-015-0895-8) contains supplementary material, which is available to authorized users.

Pigeons were trained to discriminate photographs of cat faces from dog faces. They were then presented with test stimuli involving high- and low-pass spatial frequency filtering. Discrimination was maintained with both types of filtered stimuli, though it was increasingly impaired the more information was filtered out, and high-pass filtering impaired discrimination more than low-pass filtering. The pigeons were then exposed to hybrid stimuli in which high-pass filtered dog faces were combined with low-pass filtered cat faces, and vice versa. Response to hybrid stimuli was determined more by the low spatial frequency content than by the high-frequency content, whereas humans viewing the same stimuli at corresponding viewing distance respond more strongly to the high-frequency content. These results are unexpected given that, compared with humans, pigeons' behavior tends to be controlled by the local details of visual stimuli rather than their global appearance.

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