Modulation of dear enemy effects by male dusky warblers (Phylloscopus fuscatus) at different reproductive stages.

Territorial animals show less aggression during intrusion by neighbours than by strangers, a phenomenon known as the "dear enemy effect". Recent studies have shown that the dear enemy effect is variable in some birds throughout the breeding season. However, there is limited research on changes in dear enemy effects in songbirds during the breeding season. In this study, we monitored dusky warblers (Phylloscopus fuscatus) throughout their breeding stages, conducting playback experiments simulating incursions by neighbours and strangers during their egg-laying and incubation/nestling periods, and recording physical responses and song responses from territory owners. The results showed that male dusky warblers responded similarly to neighbours and strangers during the female egg-laying period, but there was a significantly stronger response to strangers than to neighbours during the incubation/nestling period. This suggests that male dusky warblers adjust the intensity of their defences against neighbours according to the reproductive status of their own mate. This may be because neighbouring males threaten the paternity of territorial males during the egg-laying period, so territorial males demote the 'dear enemy' friendly relationships with their neighbours to defend their mate and paternity. In contrast, during the non-reproductive period of females, when neighbours are less of a threat to the paternity of territorial males, lowering defences against neighbouring males allow them to devote more time and energy to activities that improve their fitness, such as foraging and seeking extra-pair mating.