Substitution of heat from exercise and digestion by ducks diving for mussels at varying depths and temperatures

Diving birds can lose significant body heat to cold water, but costs can be reduced if heat from exercising muscles or the heat increment of feeding (HIF) can substitute for thermogenesis. Potential for substitution depends jointly on the rate of heat loss, the rate of heat produced by exercise, and the level of HIF. To explore these interactions, we measured oxygen consumption by lesser scaup ducks (Aythya affinis) diving to depths of 1.2 and 2 m at thermoneutral (23 degrees C) and subthermoneutral (18 and 8 degrees C) temperatures. Birds dove while fasted and when feeding on blue mussels (Mytilus edulis). Substitution occurred if HIF or costs of diving above resting metabolic rate (RMR) were lower at 18 or 8 degrees C than at 23 degrees C, indicating reduction in the thermoregulatory part of RMR. For fasted scaup diving to 1.2 m, substitution from exercise heat was not apparent at either 18 or 8 degrees C. At 2 m depth, dive costs above RMR were reduced by 5% at 18 degrees C and by 40% at 8 degrees C, indicating substitution. At 1.2 m depth (with voluntary intake of only 1.4-17% of maintenance requirements), HIF did not differ between temperatures, indicating no substitution. However, at 2 m (intake 13-25% of maintenance), substitution from HIF was 23% of metabolizable energy intake at 18 degrees C and 22% at 8 degrees C. These results show that even with low HIF due to low intake rates, substitution from HIF can add to substitution from the heat of exercise.