Multi-level combinatoriality in magpie non-song vocalizations.

Comparative studies conducted over the past few decades have provided important insights into the capacity for animals to combine vocal segments at either one of two levels: within- or between-calls. There remains, however, a distinct gap in knowledge as to whether animal combinatoriality can extend beyond one level. Investigating this requires a comprehensive analysis of the combinatorial features characterizing a species' vocal system. Here, we used a nonlinear dimensionality reduction analysis and sequential transition analysis to quantitatively describe the non-song combinatorial repertoire of the Western Australian magpie ( Gymnorhina tibicen dorsalis ). We found that (i) magpies recombine four distinct acoustic segments to create a larger number of calls, and (ii) the resultant calls are further combined into larger call combinations. Our work demonstrates two levels in the combining of magpie vocal units. These results are incongruous with the notion that a capacity for multi-level combinatoriality is unique to human language, wherein the combining of meaningless sounds and meaningful words interactively occurs across different combinatorial levels. Our study thus provides novel insights into the combinatorial capacities of a non-human species, adding to the growing evidence of analogues of language-specific traits present in the animal kingdom.