The presence of non-territorial males in Willow Warbler Phylloscopus trochilus populations–a removal study

A total of six (unmarked) and 24 (individually colour-ringed) male Willow Warblers Phylloscopus trochilus were removed from their territories in two study areas: subalpine birch and lowland mixed forest, respectively. The removals were made during three separate periods throughout the breeding season: (1) pair-formation and egg-laying; (2) early incubation; and (3) middle and second half of the incubation period. During Period 1 all of the males removed ( n = 10) were quickly replaced by new males, while during Period 2 eight of the 11 males removed were replaced. The new males defended basically the same territorial borders as the males they replaced. During Period 3 only one of the nine males removed was replaced. Since the replacements occurred within a few hours of the removals, it is suggested that the new settlers were non-territorial males that were already present in the respective areas before the experimental removals were made. The mean wing-lengths of the two groups of males indicated that the initial occupants of the territories were 2 years old or older in both study areas, whereas the new settlers in the subalpine birch forest (mean Willow Warbler density: 55 territories/km²) were 1-year-old birds, and two years old, or older, in the mixed forest area (mean Willow Warbler density: 133 territories/km²).