Researchers have begun to explore animals' capacities for uncertainty monitoring and metacognition. This exploration could extend the study of animal self-awareness and establish the relationship of self-awareness to other-awareness. It could sharpen descriptions of metacognition in the human literature and suggest the earliest roots of metacognition in human development. We summarize research on uncertainty monitoring by humans, monkeys, and a dolphin within perceptual and metamemory tasks. We extend phylogenetically the search for metacognitive capacities by considering studies that have tested less cognitively sophisticated species. By using the same uncertainty-monitoring paradigms across species, it should be possible to map the phylogenetic distribution of metacognition and illuminate the emergence of mind. We provide a unifying formal description of animals' performances and examine the optimality of their decisional strategies. Finally, we interpret animals' and humans' nearly identical performances psychologically. Low-level, stimulus-based accounts cannot explain the phenomena. The results suggest granting animals a higher-level decision-making process that involves criterion setting using controlled cognitive processes. This conclusion raises the difficult question of animal consciousness. The results show that animals have functional features of or parallels to human conscious cognition. Remaining questions are whether animals also have the phenomenal features that are the feeling/knowing states of human conscious cognition, and whether the present paradigms can be extended to demonstrate that they do. Thus, the comparative study of metacognition potentially grounds the systematic study of animal consciousness.

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The authors asked whether animals and humans would use similarly an uncertain response to escape indeterminate memories. Monkeys and humans performed serial probe recognition tasks that produced differential memory difficulty across serial positions (e.g., primacy and recency effects). Participants were given an escape option that let them avoid any trials they wished and receive a hint to the trial's answer. Across species, across tasks, and even across conspecifics with sharper or duller memories, monkeys and humans used the escape option selectively when more indeterminate memory traces were probed. Their pattern of escaping always mirrored the pattern of their primary memory performance across serial positions. Signal-detection analyses confirm the similarity of the animals' and humans' performances. Optimality analyses assess their efficiency. Several aspects of monkeys' performance suggest the cognitive sophistication of their decisions to escape.

The authors asked whether animals, like humans, use an uncertain response adaptively to escape indeterminate stimulus relations. Humans and monkeys were placed in a same-different task, known to be challenging for animals. Its difficulty was increased further by reducing the size of the stimulus differences, thereby making many same and different trials difficult to tell apart. Monkeys do escape selectively from these threshold trials, even while coping with 7 absolute stimulus levels concurrently. Monkeys even adjust their response strategies on short time scales according to the local task conditions. Signal-detection and optimality analyses confirm the similarity of humans' and animals' performances. Whereas associative interpretations account poorly for these results, an intuitive uncertainty construct does so easily. The authors discuss the cognitive processes that allow uncertainty's adaptive use and recommend further comparative studies of metacognition.

Rhesus monkeys (Macaca mulatta) have learned, like humans, to use an uncertain response adaptively under test conditions that create uncertainty, suggesting a metacognitive process by which human and nonhuman primates may monitor their confidence and alter their behavior accordingly. In this study, 4 rhesus monkeys generalized their use of the uncertain response, without additional training, to 2 familiar tasks (2-choice discrimination learning and mirror-image matching to sample) that predictably and demonstrably produce uncertainty. The monkeys were significantly less likely to use the uncertain response on trials in which the answer might be known. These results indicate that monkeys, like humans, know when they do not know and that they can learn to use a symbol as a generalized means for indicating their uncertainty.

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There was a strong consensus in the commentaries that animals' performances in metacognition paradigms indicate high-level decisional processes that cannot be explained associatively. Our response summarizes this consensus and the support for the idea that these performances demonstrate animal metacognition. We amplify the idea that there is an adaptive advantage favoring animals who can – in an immediate moment of difficulty or uncertainty – construct a decisional assemblage that lets them find an appropriate behavioral solution. A working consciousness would serve this function well. This explains why animals may have the functional equivalent of human declarative consciousness. However, like other commentators who were friendly to this equivalence, we approach carefully the stronger claims that animals' metacognitive performances imply full-blown self-awareness or phenomenal consciousness.We discuss the commentators' interesting ideas for future research, as well as their intriguing ideas about the evolution and development of metacognition and its relation to theory of mind. We also discuss residual confusions about existing research and remaining methodological issues.

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