Abstract

Sporadic large outbreaks of herbivorous insects alter forest ecosystems via a top-down effect, and a high frequency of outbreaks led to the death of the trees on which the insects fed. The top-down effect of predators, such as other insects and birds, often regulates the outbreaks. Predator insects exert effective top-down control on outbreaks of herbivorous insects, but the effect of bird predation varies because birds have a broader diet range and higher mobility than insects. It was hypothesized that when large insect outbreaks occur during the breeding season of birds, most bird species would not respond owing to a fixed home range during this season and would, therefore, not exert top-down effects. To test this hypothesis, the relationship between bird and beech sawfly abundance was investigated for over nine years in the Tanzawa Mountains of central Japan, where temperate beech forests and large sporadic outbreaks of woody herbivorous insects occur. In this study, 28 649 female beech sawfly adults were captured, and 2515 individuals were identified from 50 bird species. The abundance of most bird species foraging on beech sawfly larvae, except Sitta europaea, did not increase proportionally with the number of beech sawfly adults. Many bird species did not respond to the unpredictable and temporary abundance of food resources in a way that would enhance their density, supporting the hypothesis. These results can be explained by examining the nesting and foraging guilds of the birds. S. europaea is less restricted in nesting sites and nests earlier in the season than other species; therefore, they could select nests in areas where beech sawfly larvae were abundant. Regarding foraging, S. europaea has a unique foot shape that allows it to move down the tree trunk freely and respond rapidly to fluctuations in beech sawfly abundance. Bird species that exert top-down effects on insects show promise for use in an integrated pest management approach to manage sporadic outbreaks of herbivorous insects in forest ecosystems.

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