Supra-regular recursion in primate vocal sequences

Nitschke, R.; Krapp, L. S.; Cathcart, C.; Gallot, Q.; Bosshard, A. B.; Burkart, J. M.; Berthet, M.; Townsend, S. W.; Surbeck, M.; Zuberbühler, K.; Bickel, B.

doi:10.64898/2026.09.19.752870

bioRxiv · 10.64898/2026.09.19.752870

Supra-regular recursion in primate vocal sequences

Nitschke, R.·Krapp, L. S.·Cathcart, C.·Gallot, Q.·Bosshard, A. B.·Burkart, J. M.·Berthet, M.·Townsend, S. W.·Surbeck, M.·Zuberbühler, K.·Bickel, B.

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Abstract

Complex syntax is the widely accepted demarcation between animal and human communication. Although animal vocal systems occasionally contain pockets of syntax-like structures, their grammars are thought to be at best regular, whereas human language is supra-regular, i.e., including at least context-free computational complexity. Leveraging a new likelihood method for probabilistic formal grammars, we show that the call sequences of three phylogenetically diverse nonhuman primates (bonobos, olive colobus monkeys, common marmosets) include parts that are most likely generated by supra-regular recursive grammars, similar to human language. These findings challenge syntax as the major transition in hominid evolution and opens up prospects for alternative evolutionary scenarios.

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Nitschke, R., Krapp, L. S., Cathcart, C., Gallot, Q., Bosshard, A. B., Burkart, J. M., Berthet, M., Townsend, S. W., Surbeck, M., Zuberbühler, K., Bickel, B.. 2026-09-21. Supra-regular recursion in primate vocal sequences. https://doi.org/10.64898/2026.09.19.752870

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