Leaf kiss-squeaks showed a slightly lower maximum frequency than kiss-squeaks unaided produced by the twelve individuals from whom there were recordings of both variants, but this difference did not reach statistical significance (Wilcoxon signed-ranks test: T = -1.609, N = 12, P = 0.108; Figure 4) .
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Population-specific use of the same tool-assisted alarm call between two wild orangutan populations (Pongo pygmaeus wurmbii) indicates functional arbitrariness [corrected].
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