Experimental manipulation of energy expenditure has long been recognized as an effective means for identifying causative effects and avoiding confounded interpretations arising from spurious correlations. This approach has been successfully applied mainly to studies on birds, particularly on reproducing adults, whereas manipulations in mammals have proved more problematic. Here, we tested the hypothesis that shaving off 50% of the dorsal pelage should effectively increase energy expenditure in wild root voles (Microtus oeconomus) in their natural environment. We measured daily energy expenditure (DEE), using doubly labelled water in shaved and unshaved voles at the beginning of winter. The difference in DEE (corrected for body mass and year effects) between experimental and control group fluctuated from 11.5% to 17.3%. Probability of recapture over the 3 day DEE assay was strongly dependent on body mass, but did not differ between shaved and unshaved animals; however, a prevalence of larger (heavier) shaved individuals was observed. Shaved animals lost more weight between the first and second trapping. Shaving therefore appears to be an effective method of increasing the cost of total DEE in wild endotherms in their natural environment.
KEY WORDS: Body mass, Doubly labelled water, Fur insulation, Mammal, Metabolic rate, Thermoregulation
INTRODUCTIONThe idea to overload an animal's energy expenditure to increase its total energy budget came from classical studies on parental effort (Drent and Daan, 1980). Experimental manipulations of animal energetics have mainly been successfully applied in studies on reproducing birds. This is because of a relative ease of implementing such manipulations by changing clutch size (Knowles et al., 2010) or the costs of flight (by feather clipping) (Carrascal and Polo, 2006; Barron et al., 2013). Experimental manipulation of energy expenditure outside the breeding season, particularly in mammals, is far more difficult and is mostly limited to laboratory conditions, which allows for manipulation via control of the ambient temperature (Selman et al., 2008; Chappell et al., 2007), food quality or quantity (Cao et al., 2009; Gutowski et al., 2011), litter size (Simons et al., 2011Zhao et al., 2013) or the composition of respired air (Rosenmann and Morrison, 1974; Cheviron et al., 2013). The lack of effective means of manipulation of energy expenditure in free ranging, non-reproducing mammals is reflected in the scarcity of such studies, which mainly rely on natural, uncontrolled variation of environmental factors (Nagy et al., 1999;Nagy, 2005), and therefore, are inherently correlative in nature. The aim of this study was to test under natural conditions the hypothesis that shaving, as a method of increasing costs of thermoregulation, increases total daily energy expenditure (DEE). Measurements using pelt-covered, internally heated metal casts of animals indicated that naked skin increases thermal conductance (Jofré and CaviedesVidal, 2003) and by contrast, long and thick f...