Microhabitat variables explain patch switching by wintering Bewick's swans through giving-up net energy intake rates

Microhabitat variables are determinants for animals to select forage patches and evaluate the cost/benefit tradeoffs of habitat switching. Optimal foragers would weigh habitat quality by giving-up net energy intake rate (GUN), which includes the energy intake rate and cost rate. The GUNs, energy intake rate, and cost rate can be influenced by variations in different microhabitat factors and interactive effects. In this study, we assessed the GUN patterns of wintering Bewick's swans and the effects of microhabitat factors on their foraging strategy in three different habitats: foxnut ponds, paddy fields, and shallow lake. The foraging behaviors and microhabitat variables of the swans were investigated during the winters of 2016-2018 and 2017/2018 at Huangpi and Shengjin Lakes in Anhui Province, southeastern China. The results showed that the percentage of disturbance time and the giving-up food density in shallow water had significant negative effects on GUNs. In contrast, water depth and the giving-up food density in deep water showed positive effects on GUNs. 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 GUNs may mitigate energy demand and environmental stress by switching foraging habitat while in subprime foraging habitat if alternative habitats that offered higher net rates of energy gain were available.