Distances and directions are computed separately by honeybees in landmark-based search

Honeybees were trained with two landmarks at some angle (e.g., 120°) apart from the target. On crucial unrewarded tests, only a single landmark was present. If distances and directions to landmarks are computed separately (independent averaging), the search distance to the landmark should equal the landmark-target distance found in training. If entire vectors are averaged, the search distance should be much shorter. Three experiments with short target-landmark distances showed results in between the predictions of the two hypotheses. A fourth experiment used longer target-landmark distances and isolated double peaks on single-landmark tests: one predicted by the independent averaging hypothesis, and one very close to the landmark. The near peak is interpreted as arising from approach and exploration of a landmark in a new location, and not from searching.