Quantifying the free living energy exchanges of Arctic ungulates with stable isotopes

Abstract

When natural diets meet an animal's requirement for energy, other essential nutrients will usually be supplied in amounts at least sufficient for survival. Knowledge of the energy requirements of free ranging species under typical conditions are important in assessing both their nutritional needs and their ecological impact. The doubly labelled water (DLW) method is currently the most promising objective field methodology for estimating free living energy expenditure but expenditure is only equal to the energy requirement when an animal is in energy balance. Reproduction and seasonal cycles of fat deposition and utilization represent significant components of the energy budget of arctic ungulates but the information gained in the course of a typical DLW study may be used to estimate processes such as milk output and fat storage and mobilization in order to predict requirements from expenditure. The DLW method has been exhaustively validated under highly controlled conditions and the introduction of innovations such as faecal sampling for the estimation of body water isotopic enrichment, the availability of appropriate correction factors and stoichiometrics for known sources of error, and iterative calculation of unknown parameters, have produced a methodology suitable for use in truly free ranging species. The few studies carried out so far in arctic ungulates indicate that previous predictions have generally underestimated the true level of expenditure, that there is considerable between animal variation in the level of expenditure and that this is largely determined by physical activity. The disadvantages of the DLW methodology are that it remains expensive and the isotope analysis is technically demanding. Furthermore, although DLW can provide an accurate value for free living energy expenditure, it is often important to have information on the individual components of expenditure, for example the relative contribution of physical activity and thermoregulatory thermogenesis, in order to interpret the values for overall expenditure. For these reasons the most valuable use of the DLW method in the field may be to validate factorial models and other approaches so that they may be used with confidence. Additional important information on the energy exchanges of free ranging animals may be obtained from other stable isotope methodologies. In addition to the use of the isotopes 2H and lsO in the DLW method, natural variations in the abundance of "C and 15N in the arctic environment may be exploited to study diet selection in truly free living arctic ungulates.