Avian diving, respiratory physiology and the marginal value theorem

Abstract

Behavioural studies of diving birds have reported that the ratio of dive duration to the duration of the subsequent period on the surface displays a characteristic relation to dive duration. For short dives, the dive to surface ratio increases with dive duration, whereafter the relation peaks, and for longer dives decreases with increasing dive duration. Such a relationship is not a general prediction of existing marginal value models which have been used to predict optimal diving behaviour. This may be because the smooth curve used to describe the oxygen gain rate of individuals after surfacing is not a good reflection of the respiratory physiology of birds. Here we argue that on physiological grounds, the oxygen gain curve for avian divers will not be smooth, but will have two distinct regions (representing oxygen recovery in the respiratory tract, and in haemoglobin and myoglobin, respectively). Modifying two of the classical diving models by incorporating such a kinked curve causes them to predict the humped relationship between dive to surface ratio and dive duration under many circumstances. We also present data on the duration of dives and surface periods from three species of diving seabirds: the shag,Phalacrocorax aristotelis, the black guillemot,Cepphus grylleand the common guillemot,Uria aalge. All three species showed a humped relationship for dive to surface ratio as a function of dive duration. In line with the predictions of our model, when oxygen stores on surfacing were greatly depleted, the dive to surface ratio peaked at short dive durations.