Patterns of reproductive effort in male ungulates

In ungulates, males and females have contrasting life histories, as usually only the females raise the young. How reproductive effort in males varies with individual level and population level characteristics has received little attention in the literature. Using published information on direct (weight loss during the rut) and indirect (rut-related changes in activity budgets, fighting frequency, etc.) measures of reproductive effort, we tested whether effort in males increased with (H-1) increasing age, (H-2) increasing body size, (H-3) decreasing population density, (H-4) increasingly female-biased sex ratio and with a younger male age structure. Consistent with H-1, reproductive effort was consistently higher in prime-aged than in younger males in a large number of studies. Among younger males, sub-adult males had an equal or higher effort than yearling males. Prime-aged males had more typical rutting behaviour (e.g. roaring, tending, fighting and chasing frequencies) and they often lowered their intake of forage during the rut. However, reproductive effort usually declined for very old age classes. Reproductive effort increased with size (H-2) also after accounting for age. Data on effort vs (H-3) density and (H-4) sex ratio/male age structure were inconclusive. Clearly, far more studies relating effort to population characteristics are needed before the rutting ecology of male ungulates can be understood.