Gray tree frogs, Hyla versicolor, give lower-frequency aggressive calls in more escalated contests

As animal contests escalate, variation in the performance of aggressive signaling behaviors can give important insights into contest dynamics. In anuran amphibians, males of numerous species utilize distinctive aggressive vocalizations during disputes over calling spaces. Little is known, however, about the causes and consequences of variation in aggressive-call characteristics. We analyzed recordings of calls made during staged aggressive interactions between male gray tree frogs, Hyla versicolor, to determine how variation in a key aggressive-call characteristic, dominant frequency, was affected by increasing contest escalation. We found that dominant frequencies of aggressive calls were lower than those of advertisement calls used to attract females. Furthermore, we found that males lowered their aggressive-call frequencies with increases in escalation. Winners tended to have lower-frequency aggressive calls than losers. We conclude that aggressive calls in H. versicolor are similar to the graded aggressive calls that have been described in several other species. This gradation may allow males to balance the energetic costs of producing lower-frequency calls with the benefits of efficiently repelling rival males. Other processes related to motivation and the physiological effects of participating in contests may also be responsible for the observed variation in aggressive-call frequency with contest escalation. Our results demonstrate that detailed experimental studies of aggressive calling behavior in anurans, which to this point have rarely been performed, are feasible and generate important insights relating to general problems in animal contest behavior and animal communication.