Landscape complementation and food limitation of large herbivores: Habitat-related constraints on the foraging efficiency of wild pigs

Abstract

1. The effect that the proximity of habitats containing essential resources has on animal abundance at large spatial scales is called landscape complementation. Landscape complementation can influence interaction between large herbivores and their food resources where the proximity of habitats containing essential resources constrains their foraging or demographic efficiency. 2. We tested the effect that the proximity of a thermal refuge (riverine woodlands) had on interaction between wild pigs and their food resources (pasture). The numerical response of pigs (Sus scrofa L.) (estimated as r quarter-1) to pasture biomass was contrasted between four sites that were progressively more isolated from a major floodplain containing extensive areas of riverine woodland. To test whether proximity to riverine woodlands affected the numerical response of pigs through a constraint on foraging efficiency, we contrasted the numerical response of pigs between the four areas as pasture biomass declined. To test whether pigs exploited riverine woodlands primarily as a thermal refuge, we contrasted the numerical response of pigs between the four areas in different seasons. 3. We found that although pasture biomass was similar in the four areas, r was lower than expected for given pasture biomass on the two areas further away from riverine woodlands. We also found that while the effect that proximity to riverine woodlands had on the numerical response of pigs became more pronounced when pasture biomass was low, it was not significantly affected by season. 4. These results suggest that the need to access riverine woodlands compromises the foraging efficiency of wild pigs when the distance to this habitat is relatively high, but that the need to access this habitat may not be purely related to thermoregulation. 5. We developed a simple mechanistic model that allows the effects of landscape complementation on herbivore foraging and demographic efficiency to be estimated, and used the model to predict the effect that proximity to riverine woodlands would have on variation in pig density. The model suggests that wild pigs cannot persist in areas more then 10 km from extensive riverine woodlands, unless those areas are periodically recolonized. This suggests that at the margin of their range around inland river systems, given locations that can be occupied by wild pigs will vary temporally between being sources, pseudosinks and sinks.