No abstract

Insects possess small brains but exhibit sophisticated behaviour, specifically their ability to learn to navigate within complex environments. To understand how they learn to navigate in a cluttered environment, we focused on learning and visual scanning behaviour in the Australian nocturnal bull ant, Myrmecia midas, which are exceptional visual navigators. We tested how individual ants learn to detour via a gap and how they cope with substantial spatial changes over trips. Homing M. midas ants encountered a barrier on their foraging route and had to find a 50-cm gap between symmetrical large black screens, at 1m distance towards the nest direction from the centre of the releasing platform in both familiar (on-route) and semi-familiar (off-route) environments. Foragers were tested for up to 3 learning trips with the changed conditions in both environments. Results showed that on the familiar route, individual foragers learned the gap quickly compared to when they were tested in the semi-familiar environment. When the route was less familiar, and the panorama was changed, foragers were less successful at finding the gap and performed more scans on their way home. Scene familiarity thus played a significant role in visual scanning behaviour. In both on-route and off-route environments, panoramic changes significantly affected learning, initial orientation and scanning behaviour. Nevertheless, over a few trips, success at gap finding increased, visual scans were reduced, the paths became straighter, and individuals took less time to reach the goal.

Many animals navigate in a structurally complex environment, which requires them to detour around the physical barriers that they encounter. Although many studies in animal cognition suggest that they are able to adeptly avoid obstacles, it is unclear whether a new route is learned to navigate around these barriers and, if so, what sensory information may be used to do so. We investigated detour learning in traveling up a tree in the Australian bull ant, Myrmecia midas, which primarily uses visual landmarks. We first placed a barrier on the ants' upward path. Initially, 46% of foragers were unsuccessful in detouring the obstacle. On subsequent trips, the ants became more successful and established a new route. We observed up to eight successful foraging trips detouring around the barrier. We then tested the same foragers in a series of manipulations, including changing the position of the barrier, making a new gap in the middle of the obstacle, or removing the barrier altogether. The ants mostly showed the same learned motor routine, detouring with a similar path as in the initial trials, suggesting that foragers were not relying on barrier cues and therefore learned a new route around the obstacle. When foragers encountered new olfactory or tactile cues, or the visual environment was blocked; however, their navigation was profoundly disrupted. These results suggest that changing sensory information drastically affects the foragers' navigational performance.

How do ants resolve conflicts between different sets of navigational cues during navigation? When two cue sets point to diametrically opposite directions, theories predict that animals should pick one set of cues or the other. Here we tested how nocturnal bull ants Myrmecia midas adjust their paths along established routes if route following does not lead to their entry into their nest. During testing, foragers were repeatedly placed back along their homeward route up to nine times, a procedure called rewinding. This procedure produced an accumulating path integrator, or vector, in diametric opposition to the learned landmark views of the route. Repeated rewinding made some individuals head initially in the nest-to-feeder vector direction, but all ants ended up using the visual scene for homing, demonstrating the importance of view-based homing in this species. Repeated rewinding, however, led to path deteriorations; with increased path meander and scanning, results also found in desert ants. After nine rewinding trips, ants were displaced off their route in further manipulations, to a site near the nest, an unfamiliar site, or with the terrestrial surround entirely covered. The results show that a change in visual conditions diminished the weight accorded to path integration: the off-route ants no longer headed off in the vector direction as they did on the immediately preceding trial. They relied on celestial compass cues in other ways for homing. Experiment 2 showed the effects of rewinding in the unaltered natural habitat were not view-specific in these bull ants.

Many insects orient by comparing current panoramic views of their environment to memorised views. We tested the navigational abilities of night-active Myrmecia midas foragers while we blocked segments of their visual panorama. Foragers failed to orient homewards when the front view, lower elevations, entire terrestrial surround, or the full panorama was blocked. Initial scanning increased whenever the visual panorama was blocked but scanning only increased along the rest of the route when the front, back, higher, or lower elevations were blocked. Ants meandered more when the front, the back, or the higher elevations were obscured. When everything except the canopy was blocked, the ants were quick and direct, but moved in random directions, as if to escape. We conclude that a clear front view, or a clear lower panorama is necessary for initial homeward headings. Furthermore, the canopy is neither necessary nor sufficient for homeward initial heading, and the back and upper segments of views, while not necessary, do make finding home easier. Discrepancies between image analysis and ant behaviour when the upper and lower views were blocked suggests that ants are selective in what portions of the scene they attend to or learn.