Energy expenditure during flight in relation to body mass: effects of natural increases in mass and artificial load in Rose Coloured Starlings.

Rose Coloured Starlings (Sturnus roseus) flew repeatedly for several hours in a wind tunnel while undergoing spontaneous variation in body mass. The treatments were as follows: flying unrestrained (U), with a control harness of 1.2% of their body mass (C), or with a harness of 7.4% of their body mass, which was either applied immediately before the flight (LS) or at least 9 days in advance (LL). Energy expenditure during flight (ef in W) was measured with the Doubly Labelled Water method. Flight costs in L(S) and LL were not significantly different and therefore were pooled (L). The harness itself did not affect ef, i.e. U and C flights were not different. ef was allometrically related with body mass m (in g). The slopes were not significantly different between the treatments, but ef was increased by 5.4% in L compared to C flights (log10(ef) = 0.050 + 0.47 x log10(m) for C, and log10(ef) = 0.073 + 0.47 x log10(m) for L). The difference in ef between C, LS and LL was best explained by taking the transported mass m transp (in g) instead of m into account (log10(ef) = -0.08 + 0.54 x log10(m transp)). Flight costs increased to a lesser extent than expected from interspecific allometric comparison or aerodynamic theory, regardless of whether the increase in mass occurred naturally or artificially. We did not observe an effect of treatment on breast muscle size and wingbeat frequency. We propose that the relatively low costs at a high mass are rather a consequence of immediate adjustments in physiology and/or flight behaviour than of long-term adaptations.

Energy expenditure and wing beat frequency in relation to body mass in free flying Barn Swallows (Hirundo rustica).

Many bird species steeply increase their body mass prior to migration. These fuel stores are necessary for long flights and to overcome ecological barriers. The elevated body mass is generally thought to cause higher flight costs. The relationship between mass and costs has been investigated mostly by interspecific comparison and by aerodynamic modelling. Here, we directly measured the energy expenditure of Barn Swallows (Hirundo rustica) flying unrestrained and repeatedly for several hours in a wind tunnel with natural variations in body mass. Energy expenditure during flight (e (f), in W) was found to increase with body mass (m, in g) following the equation e (f) = 0.38 x m (0.58). The scaling exponent (0.58) is smaller than assumed in aerodynamic calculations and than observed in most interspecific allometric comparisons. Wing beat frequency (WBF, in Hz) also scales with body mass (WBF = 2.4 x m (0.38)), but at a smaller exponent. Hence there is no linear relationship between e (f) and WBF. We propose that spontaneous changes in body mass during endurance flights are accompanied by physiological changes (such as enhanced oxygen and nutrient supply of the muscles) that are not taken into consideration in standard aerodynamic calculations, and also do not appear in interspecific comparison.