Anatomical constraints generate honesty: acoustic cues to age and weight in the roars of red deer stags.

Early work on loud calling in mammals emphasized the importance of dynamic characteristics such as calling rate as cues to fitness and fighting ability. In contrast, little is known of the potential for fine-scaled acoustic cues to provide receivers with direct information on fitness. Fundamental frequency has typically been considered a good potential indicator of body size in the literature, but resonance frequencies (formants), which should be constrained by the length of the vocal tract, have received less attention. We conducted a detailed acoustic analysis on an extensive database of roars from red deer stags, Cervus elaphus, in a free-ranging population to investigate which variables provided honest information on age, body weight and reproductive success. Although fundamental frequency was higher in young stags than in adults, it did not decrease with body weight within adults and source cues (i.e. those generated by the larynx) in general did not provide clear information on fitness-related characteristics. In contrast, minimum formant frequencies, reached during the part of the roar when the mobile larynx is most fully retracted towards the sternum, decreased with body weight and age and were strongly negatively correlated with our index of reproductive success. Such production-related acoustic cues to body size and fitness, rendered honest by an anatomical constraint limiting the downward movement of the larynx, provide receivers with accurate information that could be used to assess rivals and choose mates.