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Virtual reality (VR) offers an appealing experimental framework for studying visual performances of insects under highly controlled conditions. In the case of the honeybee Apis mellifera, this possibility may fill the gap between behavioural analyses in free-flight and cellular analyses in the laboratory. Using automated, computer-controlled systems, it is possible to generate virtual stimuli or even entire environments that can be modified to test hypotheses on bee visual behaviour. The bee itself can remain tethered in place, making it possible to record neural activity while the bees is performing behavioural tasks. Recent studies have examined visual navigation and attentional processes in VR on flying or walking tethered bees, but experimental paradigms for examining visual learning and memory are only just emerging. Behavioural performances of bees under current experimental conditions are often lower in VR than in natural environments, but further improvements on current experimental protocols seem possible. Here we discuss current developments and conclude that it is essential to tailor the specifications of the VR simulation to the visual processing of honeybees to improve the success of this research endeavour.

Ants are known to use the terrestrial visual panorama in navigation. Recent evidence has accumulated for the use of the currently perceived visual panorama to determine a direction to head in. The pattern of the height of the terrestrial surround, the skyline, is one key cue for the Central Australian red honey ant Melophorus bagoti in determining a direction of travel. But ants might also possess some mechanism to match the skyline heights encountered during training, which functions to steer away from regions whose skyline is too high and towards regions whose skyline is too low. We made an initial test of this hypothesis by training ants to visit a feeder centred between two experimentally constructed walls of black cloth. Trained ants were then tested for their initial homing direction with the walls retaining their heights as encountered in training (controls), with one of the walls lowered or raised in height, or with one wall lowered and the opposite wall raised. Wall-height manipulations deflected the initial headings of ants towards the lower wall, with combined wall lowering and wall raising changing the initial headings by ~30° when compared with controls. The results suggest that the ants combined the dictates of the panorama in determining the best direction of travel (a heading towards the nest) with some attractor mechanism that functions to establish the skyline heights of training conditions (a heading towards the lower wall).

The Central Australian desert ant Melophorus bagoti is known to use celestial cues for compass orientation. We manipulated the available celestial cues for compass orientation for ants that had arrived at a feeder, were captured and then released at a distant test site that had no useful terrestrial panoramic cues. When tested in an enclosed transparent box that blocked some or most of the ultraviolet light, the ants were still well oriented homewards. The ants were again significantly oriented homewards when most of the ultraviolet light as well as the sun was blocked, or when the box was covered with tracing paper that eliminated the pattern of polarised light, although in the latter case, their headings were more scattered than in control (full-cue) conditions. When the position of the sun was reflected 180° by a mirror, the ants headed off in an intermediate direction between the dictates of the sun and the dictates of unrotated cues. We conclude that M. bagoti uses all available celestial compass cues, including the pattern of polarised light, the position of the sun, and spectral and intensity gradients. They average multiple cues in a weighted fashion when these cues conflict.

The Australian desert ant Melophorus bagoti (Formicidae) is a thermophilic, solitary foraging ant that inhabits the semi-arid regions of Australia. In recent years, it has become a model species for the study of navigation. However, its ecological traits are not well understood, especially on the level of the entire colony. Here, we investigated this species daily activity schedule and diet composition, and examined its foraging behaviour. Foraging activity is confined to a window of roughly 50-70°C soil surface temperature, and foragers reacted quickly to temperature changes. Consequently, the pattern of daily outbound traffic during summer is unimodal on warm days and bimodal on very hot days. Foragers are opportunistic scavengers; dead insects make up a large proportion of food items, but grass seeds are also occasionally brought back to the nest in large amounts. Diet composition changes with the seasonal availability of certain food groups. Melophorus bagoti foragers have the ability to recruit nestmates to profitable food sources. Recruitment seems to function without the use of pheromone trails, but the exact mechanism requires further investigation.

Foragers of Melophorus bagoti often return to previously rewarding sites to search for more food items. They are opportunistic scavengers that exploit both protein and carbohydrate food sources. Under natural foraging conditions, protein food items are distributed sparsely and randomly, whereas carbohydrates come in patches that are often renewable. This makes for vastly different foraging scenarios that a single forager is confronted with. In theory, foraging performance can be greatly improved if foragers are able to adjust their strategy to different food item distributions. This could be achieved through individual foraging experience or by employing pre-existing, intrinsic foraging strategies. We investigated this by offering both kinds of food with the same distribution: as a renewable food source at a fixed location. After removal of the food source, outbound foragers displayed an area-restricted search centered on its location. Searches for protein had a greater spread than those for carbohydrates, which matches the natural distribution pattern of these food types. However, searches for both kinds of food follow the same general strategy, which is best described as a Brownian-like walk. We suggest that the observed adaptive behavior is a result of differential learning effort.

Ants use the panoramic skyline in part to determine a direction of travel. A theoretically elegant way to define where terrestrial objects meet the sky is to use an opponent-process channel contrasting green wavelengths of light with ultraviolet (UV) wavelengths. Compared with the sky, terrestrial objects reflect relatively more green wavelengths. Using such an opponent-process channel gains constancy in the face of changes in overall illumination level. We tested the use of UV wavelengths in desert ants by using a plastic that filtered out most of the energy below 400 nm. Ants, Melophorus bagoti, were trained to home with an artificial skyline provided by an arena (experiment 1) or with the natural panorama (experiment 2). On a test, a homing ant was captured just before she entered her nest, and then brought back to a replicate arena (experiment 1) or the starting point (the feeder, experiment 2) and released. Blocking UV light led to deteriorations in orientation in both experiments. When the artificial skyline was changed from opaque to transparent UV-blocking plastic (experiment 3) on the other hand, the ants were still oriented. We conclude that UV wavelengths play a crucial role in determining direction based on the terrestrial surround.

Two species of Australian desert ants, one inhabiting a visually rich environment (Melophorus bagoti) and one inhabiting a visually barren environment (as yet unnamed and dubbed Melophorus sp.), were tested on path integration and subsequent search. For each species, prominent landmarks were placed near the nest, and ants were trained to visit a feeder. After training over 2 days, an ant was captured at the feeder when it had grabbed some food and tested, just once individually, in homing with the training landmarks either present or absent. Their subsequent search was also recorded on gridded paper according to a grid marked at the test site. Both species headed initially in the feeder-nest direction, but directional scatter was larger when training landmarks were absent. Melophorus bagoti ran a shorter distance before starting to search on tests with the landmarks absent, but Melophorus sp. ran a similar distance in both conditions. In both species, the variance in distance run was larger when landmarks were absent. In searching, both species expanded their search pattern more when landmarks were absent than when landmarks were present. Contrary to predictions, the distribution of the lengths of search segments was best described as a single exponential function on tests with landmarks absent and as a double exponential function with landmarks present. Two species of Australian desert ants of the same genus but occupying visually different habitats were studied in similar experiments. One inhabits a barren saltpan while the other inhabits a semi-arid environment filled with grass, shrubs and trees. The ants were compared in their return home after a short outbound trip (2 m) to a feeder and the subsequent search for the nest. When the ants were trained in an environment with some prominent artificial landmarks which were subsequently removed in tests, ants living in the visually cluttered habitat cut short their initial run, starting to engage in search earlier than ants living on the salt pan. Search characteristics were, on the other hand, similar in the two species, both expanding their search loops over time, more so when the familiar scene was changed. Visual ecology drives some aspects of navigation, while common task requirements drive other aspects.

Australian desert ants, Melophorus bagoti, return home after foraging by means of path integration and visual navigation. If these mechanisms do not deliver them exactly at the inconspicuous nest entrance, they engage in a systematic search. Here we describe the structure of this search pattern in detail. Trained ants ran home from a feeder in the natural visual setting where they navigated very accurately, and were then captured for tests on a distant test field after they had almost reached their nest. The search pattern consisted of loops and was centred on the position where the nest was most likely to be located. At first, it covered a rather small area, but then gradually extended outwards to cover a larger area. The search density was also adapted to the preceding outbound foraging distance, with longer distances leading to flatter, wider search distributions. Since the visual surround at the time of capture was similar for ants with all outbound distances, we suggest this is an adaptation to the cumulative error of the homing vector. The frequency distribution of segment lengths in the paths of searching ants does not show characteristics of a Levy walk strategy. Instead, it is well described by a double exponential model, lending support to a theoretically optimal strategy that consists of a mixture of two random walks, as in the composite Brownian walk strategy.

Honeybees (Apis mellifera) are regularly faced with the task of navigating back to their hives from remote food sources. They have evolved several methods to do this, including compass-directed “vector” flights and the use of landmarks. If these hive-centered mechanisms are disrupted, bees revert to searching for the hive, using an optimal Levy flight searching strategy. The same strategy is adopted when a food source at a known location ceases to be available. Here, we show that the programming for this Levy strategy does not need to be very sophisticated or clever on the bee’s part, as Levy flight patterns can be derived from the Weber–Fechner law in a bee’s odometer. Odometry errors of a different kind occur in desert ants (Cataglyphis spp., Melophorus bagoti). The searching behaviors of these ants are very similar in overall structure to that of honeybees but do not display any Levy flight characteristics. We suggest that errors in the estimation of distance can be implicitly involved in shaping the structure of systematic search behavior and should not be regarded as merely deficiencies in the odometer.

Summary Many animals - including insects - navigate visually through their environment. Solitary foraging desert ants are known to acquire visual information from the surrounding panorama and use it to navigate along habitual routes or to pinpoint a goal like the nest. Returning foragers that fail to find the nest entrance engage in searching behaviour, during which they continue to use vision. The characteristics of searching behaviour have typically been investigated in unfamiliar environments. Here we investigated in detail the nest searching behaviour of Melophorus bagoti foragers within the familiar visual environment of their nest. First, by relating search behaviour to the information content of panoramic (360°) images, we found that searches were more accurate in visually cluttered environments. Second, as observed in unfamiliar visual surrounds, searches were dynamic and gradually expanded with time, showing that nest-pinpointing is not rigidly controlled by vision. Third, contrary to searches displayed in unfamiliar environments, searches observed here could be modelled as a single exponential search strategy, which is similar to a Brownian walk, and there was no evidence of a Lévy walk. Overall, our results revealed that searching behaviour is remarkably flexible and varies according to the relevance of information provided by the surrounding visual scenery.

The Australian desert ant Melophorus bagoti often follows stereotypical routes through a cluttered landscape containing both distant panoramic views and obstacles (plants) to navigate around. We created an artificial obstacle course for the ants between a feeder and their nest. Landmarks comprised natural objects in the landscape such as logs, branches, and tussocks. Many ants travelled stereotypical routes home through the obstacle course in training, threading repeatedly the same gaps in the landmarks. Manipulations altering the relations between the landmarks and the surrounding panorama, however, affected the routes in two major ways. Both interchanging the positions of landmarks (transpositions) and displacing the entire landmark set along with the starting position of the ants (translations) (1) reduced the stereotypicality of the route, and (2) increased turns and meanders during travel. The ants might have used the entire panorama in view-based travel, or the distal panorama might prime the identification and use of landmarks en route. Despite the large data set, both options (not mutually exclusive) remain viable.

Over the expanse of the salt lakes of the Lake Eyre basin lives a species of ant of the genus Melophorus (as yet unnamed), both on the edges of the salt-pan and on the salt-pan itself. Most of the foragers are small (~3.0–5.0 mm), but ~10% of the ants seen foraging are much larger (~7.0 mm) and may form a soldier caste. Foragers are thermophilic and show diurnal activity, displaying a single-peaked activity profile across the day, with activity time limited to 2–6 h at most each day (during the late-summer study period, 28 February to 28 March 2012). They forage largely for dead arthropods, but also occasionally bring home plant materials. Foraging success (not considering possible liquid food intake) is ~20%, resembling the success rate of their congener Melophorus bagoti, which inhabits cluttered environments. When displaced with food from a feeder, the ants head systematically and precisely in the feeder-to-nest direction, thus exhibiting path integration abilities involving celestial compass cues. The study of this species provides an interesting comparative perspective in contrasting desert ants of the same genus and thus genetic heritage inhabiting habitats differing in complexity of panoramic terrestrial cues as well as comparing ecologically similar species inhabiting the same type of habitat (in the present case, salt-pans) but differing in their phylogenetic relationships.