Effects of territorial intrusions on eavesdropping neighbors: communication networks in nightingales

Animal communication often occurs in communication networks in which multiple signalers and receivers are within signaling range of each other. In such networks, individuals can obtain information on the quality and motivation of territorial neighbors by eavesdropping on their signaling interactions. In songbirds, extracting information from interactions involving neighbors is thought to be an important factor in the evolution of strategies of territory defense. In a playback experiment with radio-tagged nightingales Luscinia megarhynchos we here demonstrate that territorial males use their familiar neighbors' performance in a vocal interaction with an unfamiliar intruder as a standard for their own response. Males were attracted by a vocal interaction between their neighbor and a simulated stranger and intruded into the neighbor's territory. The more intensely the neighbor had interacted with playback, the earlier the intrusions were made, indicating that males eavesdropped on the vocal contest involving a neighbor. However, males never intruded when we had simulated by a second playback that the intruder had retreated and sang outside the neighbor's territory. These results suggest that territorial males use their neighbors' singing behavior as an early warning system when territorial integrity is threatened. Simultaneous responses by neighboring males towards unfamiliar rivals are likely to be beneficial to the individuals in maintaining territorial integrity.