Sound production in Pituophis melanoleucus (Serpentes: Colubridae) with the first description of a vocal cord in snakes

Abstract

The pine, gopher, or bull snake (Pituophis melanoleucus) makes two different defensive sounds. Hisses are characterized by lack of frequency and amplitude modulation; bellows have a brief initial period of high-amplitude, broad-frequency sound followed by a longer period of lower-amplitude, constant-frequency sound. Both defensive sounds contain distinct harmonic elements. The modulation and harmonic nature of these sounds seems to be unique among snakes. The larynx of Pituophis is unusual in having an epiglottal keel, a dorsal expansion of the cricoid cartilage, previously proposed to contribute to sound production; however, this study shows that it plays only a small role in increasing the amplitude of bellows. Within the larynx of Pituophis is a "vocal cord," the laryngeal septum, which is a flexible, horizontal shelf of tissue that divides the anterior portion of the larynx. Removal of the laryngeal septum alters the defensive sounds and eliminates their harmonic elements. The laryngeal septum is unique among previously described vertebrate vocal cords or folds because it is supported by the cricoid (as opposed to arytenoid) cartilage and is a single (as opposed to bilaterally paired) structure.