Foraging loads of red wood ants: Formica aquilonia (Hymenoptera: Formicidae) in relation to tree characteristics and stand age

Background. Foraging efficiency is critical in determining the success of organisms and may be affected by a range of factors, including resource distance and quality. For social insects such as ants, outcomes must be considered at the level of both the individual and the colony. It is important to understand whether anthropogenic disturbances, such as forestry, affect foraging loads, independent of effects on the quality and distribution of resources. We asked if ants harvest greater loads from more distant and higher quality resources, how individual efforts scale to the colony level, and whether worker loads are affected by stand age. Methods. First, we performed a fine-scale study examining the effect of distance and resource quality (tree diameter and species) on harvesting of honeydew by red wood ants, Formica aquilonia, in terms of crop load per worker ant and numbers of workers walking up and down each tree (ant activity) (study 1). Second, we modelled what the combination of load and worker number responses meant for colony-level foraging loads. Third, at a larger scale, we asked whether the relationship between worker load and resource quality and distance depended on stand age (study 2). Results. Study 1 revealed that seventy percent of ants descending trees carried honeydew, and the percentage of workers that were honeydew harvesters was not related to tree species or diameter, but increased weakly with distance. Distance positively affected load mass in both studies 1 and 2, while diameter had weak negative effects on load. Relationships between load and distance and diameter did not differ among stands of different ages. Our model showed that colony-level loads declined much more rapidly with distance for small diameter than large diameter trees. Discussion. We suggest that a negative relationship between diameter and honeydew load detected in study 1 might be a result of crowding on large diameter trees close to nests, while the increase in honeydew load with distance may result from resource depletion close to nests. At the colony level, our model suggests that very little honeydew was harvested from more distant trees if they were small, but that more distant larger trees continued to contribute substantially to colony harvest. Although forestry alters the activity and foraging success of red wood ants, study 2 showed that it does not alter the fundamental rules determining the allocation of foraging effort.