Differential effects of moving versus stationary territorial intruders on territory defence in a songbird.

1. In territorial contests, not only acoustic or other signals, but also the movements of a territorial intruder are likely to influence the response of a resident. 2. We tested this movement hypothesis by simulating moving vs. stationary intruders into the territories of winter wrens Troglodytes troglodytes, using the same non-interactive song playbacks in both treatments. 3. Male winter wrens showed a different long-term singing reaction in response to a moving than to a stationary intruder. 4. One day after experiencing an intruder that was switching between three locations, residents started to sing earlier before sunrise, and they sang more and longer songs at dawn than before the intrusion. 5. Residents receiving the same playback from one location only reacted by starting to sing later relative to sunrise, and by singing fewer and shorter songs than before the intrusion. 6. We could not discriminate between the treatments when examining the short-term singing reactions during and immediately after the playbacks. However, our results clearly demonstrate an effect of the spatial behaviour of territorial intruders on the long-term territory defence of residents at dawn, about 24 h after an intrusion. 7. We argue that spatial behaviour of territorial intruders should be an integral part of the study of animal territory defence behaviour. Investigating long-term changes in territory defence at dawn is a sensitive tool for discriminating between different types of intruders.