Comparative analysis of morphological and acoustic correlates of bush-cricket tympanic membranes

Abstract Bush-crickets exhibit remarkable auditory diversity, having the ears in the forelegs, where the tympana could be exposed or covered by cuticular flaps known as auditory pinnae. These tympanic membranes receive sound either directly or internally via an acoustic tracheal tube. Despite extensive research on auditory tuning, the relationships between tympanal structure, body size, and acoustic traits remain unclear. In this study, µCT reconstructions with AI-assisted segmentation and phylogenetically informed regressions across 18 bush-cricket species were used to investigate how tympanal surface area and mean cross-sectional thickness relate to body size, carrier frequency, and the presence of auditory pinnae. Tympanal surface area scaled positively with body size, whereas thickness was independent of size. Carrier frequency decreased with increasing body size but showed no direct association with tympanal properties. Among 14 species with auditory pinnae, tympanal dimensions showed no correlation with peak cavity resonance frequency, indicating semi-independent evolution of pinna cavity and tympanal traits. Across species, pinnae did not alter tympanal surface area, although unilateral pinnae were linked to thicker tympana. Within these unilateral species, the pinna-covered tympanum remained consistently larger in area but thinner than the exposed side. Overall, these findings indicate that tympanal evolution reflects a balance of scaling constraints and localised effects of auditory pinnae. Summary Statement The tympanal morphology in bush-crickets reflects a balance of scaling and pinna effects. Surface area scales with body size and unilateral pinnae generate consistent asymmetries. Competing Interest Statement The authors have declared no competing interest. Footnotes Section 3.2 revised with 14 specimen. Section 3.3 revised with 10 bilateral, 4 unilateral and 4 exposed specimen. Section 3.4 revised with 4 unilateral specimen Figure 3 - 5 revised. Table 2 - 3 revised. Discussion and conclusion updated based on changes in results, figures and tables. Supplementary file updated.