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GUNs were significantly different among the three habitats. GUNs also decreased as winter progressed. Swans would decrease their GUNs under unfavorable foraging conditions such as more disturbances; however, GUNs would increase with water depth and food availability in patches with deep water. The swans demonstrated diverse GUN patterns in different foraging habitats, driven by the tradeoffs between energy intake rates and energy cost rates that were influenced by microhabitat variables. It implies that waterbirds exposed to decreasing