Long-term dynamics of the network structures in seed dispersal associated with fluctuations in bird migration and fruit abundance patterns.

In temperate zones, seed-dispersal networks by migratory birds are formed on long time scale. In mid-October from 2005 to 2016, to explore the dynamics of the network structures, we examined interannual variability of fruit abundance, bird migration, and seed-dispersal networks in central Japan. For 12 years, the fruit abundance exhibited a remarkable fluctuation across years, with the number of fruiting plants and matured fruits fluctuating repeatedly every other year, leading to the periodic fluctuations. The abundance of migratory birds was also fluctuated. According to the abundance of fruits and migratory birds, the 12 years was classified into three types: frugivores and fruits were abundant, frugivores were abundant but fruits were scarce, and frugivores were scarce. The seed-dispersal networks were investigated by collecting faeces and vomits of migrants. Of the 6652 samples collected from 15 bird species, 1671 (25.1%) included seeds from 60 plant species. Main dispersers were composed of Turdus pallidus, T. obscurus, and Zosterops japonicus. The network structures were almost nested for 12 years. Specifically, the nested structure was developed in years when fruit abundance was low. GLM analyses showed the abundance of migrants, particularly T. pallidus and T. obscurus, had strong positive effects on nested structure. It may be caused by the fact the two Turdus species were more frequently functioning as generalist dispersers when fruit abundance was lower. Our study suggested fruit abundance and foraging behaviour of frugivores determine the network structures of seed dispersal on long time scale.