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Our results show that hammerstone selection and transport differ between Ta and FBV, suggesting that these important cognitive actions are pre-cognitive and thus independent of learned tool use. Recent work has shown that adult capuchin monkeys are sensitive to the hardness of their tools during nut-cracking [ 25 ], and that with experience, they can select and transport stones of varying hardness with a precision and magnitude that appear to be superior to those of chimpanzees [ 48 ], as found in this study. How do capuchin monkeys perceive the hardness of their tools? It is tempting to imagine that they use similar mechanisms as human adults and children. For example, in adults, the sensitivity of the temporal bone to the sound pressure generated by the striking of hard and soft objects varies according to the degree of the difference between the two stimuli; this phenomenon has been called a binaural unmasking effect [ 49 ]. In infants, if the temporal bone is sensitive to the difference between two sounds, this is called a binaural unmasking effect. Although the link between binaural unmasking and percussive tool use has not been demonstrated, the sensitivity of the human ear to the pitch and frequency of sounds generated by a hammer (e.g. F2 and F3) and a lighter stone (e.g. F1 and F2) does appear to be altered by experience [ 50 ], so it is possible that Sapajus are similarly sensitive to the sound pressure generated by hammer stones and lighter stones.
Sapajus are able to produce a great variety of percussive sounds because they possess a very powerful lever system. Their lever is called the masseter and it is activated by the contraction of a group of muscles (among them, the zygomaticus) with the jaw bone as a pivot. Compared to humans, the size of this lever is 30 times greater in capuchins. The masseter is a powerful lever that is able to exert a force of approximately 10 times the body weight [ 51 ]. It is the neck muscles that function as the rope and the tail bone as the fulcrum. With a powerful lever system, the Sapajus can generate a wide range of sounds according to the shape of their skull and the size of the masseter [ 52 ], which they can modify at any time. As a result, Sapajus are able to modulate their percussive actions according to the resistance of the nut they are trying to crack. A study of their gestures and vocalizations confirmed that this is the case [ 25 ]. We suppose that capuchins, particularly adult females, have internalized this concept and are able to assess the resistance of their nut-cracking tools by the sound pressure generated when they strike them. By contrast, the social information transmitted by bodily gestures and facial expressions of capuchins during the percussive action of nut-cracking is limited. Capuchins use food and tools with the same hands, which do not change according to the evolution of social learning, and only subtle changes to the head and body in the manner of shaking a nut in the air before cracking it can occur [ 25 ]. Finally, the morphology and behaviour of Pan and Sapajus show that either species can produce a range of percussive sounds, allowing them to crack many different species of nuts.