Larynx morphology and sound production in three species of Testudinidae

Although the ability to vocalize is widespread among tortoises, the mechanisms of sound production in chelonians remain undescribed. In this study, we analyze the morphology and histology of the larynx of three species of Testudinidae (Testudo hermanni, T. graeca, and T. marginata) in order to ascertain the presence of vibrating acoustic structure, and based on our findings we propose a general model for phonation in tortoises. The structure of the larynx of the three tortoises analyzed is simple: three cartilages (the cricoid and two arytenoids) form the skeleton of the larynx, while two pairs of muscles (the dilators and constrictors) control the widening and closing of the glottis. The larynx is supported in the oral cavity by the hyoid cartilage, which in tortoises assumes the same functions of the thyroid cartilage of mammals. Two bands of elastic fibers are inserted in the lateral walls of the larynx just upstream of the glottis, and can be stretched away from the hyoid by the movements of the arytenoids. Their position and structure suggest that these bands are capable of vibrating during exhalation, and therefore may be considered vocal cords. The cricoid of T. marginata and T. graeca hold two diverticula, not previously reported, which might function as a low-frequency resonating chamber, improving the harmonic structure of tortoise calls. The structure of the larynx is compared with that of other vertebrates and the relationships between morphology and phonation are discussed. This is the first detailed description of anatomical structures possibly devoted to vocalization in chelonians.