Abstract
Alternative reproductive tactics (ARTs), discrete phenotypic variations evolved to maximize fitness, may entail different cost-benefit trade-offs. In large mammals, differences in costs associated with ARTs-including energy expenditure and parasite infection-are typically greatest during the breeding season. Nonetheless, physiological and behavioral differences between ARTs can manifest throughout the year, possibly involving costs that may contribute to maintain ARTs within populations. Using the number of nematode larvae per gram of feces (LPG) as a proxy, we explored the temporal changes in lung parasite infection in territorial and nonterritorial male chamois Rupicapra in the Gran Paradiso National Park (Italy), between 2011 and 2012. We aimed to identify which tactic-specific physiological and behavioral features (including age, hormonal levels, inter- and intrasexual interactions, and space use) or climatic factors (temperature and precipitation) best explained yearly variation in parasite infection within and between ARTs. Generalized additive mixed models showed that the fecal larval counts of lung nematodes underwent strong temporal changes in both male types. Differences between ARTs (with higher LPG values in territorial than nonterritorial males) were greatest during the rut and-to a lesser extent-in spring, respectively, at the peak and at the onset of territoriality. The difference in LPG between tactics was largely explained by the greater levels of hormone metabolites in territorial males during the rut. The other variables did not contribute significantly to explain the different shedding of larvae within and between ARTs. Our analysis suggests that different values of LPG between territorial and nonterritorial males are largely a result of tactic-specific differences in the secretion of hormone metabolites, but only during the rut. To clarify whether rut-related parasitism contributes to the maintenance of ARTs, tactic-specific life history trade-offs, for example, between reproduction and parasite-related mortality, must be investigated.