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Abstract Objectives Recent advances in bio-inspired navigation have sparked interest in the phenomenon of skylight polarization. This interest stems from the potential of skylight-based orientation sensors, which performance can be simulated using physical models. However, the effectiveness of machine learning algorithms in this domain relies heavily on access to large datasets for training. Although there are several databases of simulated images in literature, there remains a lack of publicly available annotated real-world color polarimetric images of the sky across various weather conditions. Data description We present here a dataset obtained from a long-term experimental setup designed to collect polarimetric images from a stand-alone camera. The setup utilizes a Division-of-Focal-Plane polarization camera equipped with a fisheye lens mounted on a rotative telescope mount. Furthermore, we obtained the sensor’s orientation within the East-North-Up (ENU) frame from a geometrical calibration and an algorithm provided with the database. To facilitate further research in this area, the present sample dataset spanning two months has been made available on a public archive with manual annotations as required by deep learning algorithms. The images were acquired at 10 min intervals and were taken with various exposure times ranging from 33µs to 300ms.

Objectives : Recent advances in bio-inspired navigation have sparked interest in the phenomenon of skylight polarization. This interest stems from the potential of skylight-based orientation sensors, which performance can be simulated using physical models. However, the effectiveness of machine learning algorithms in this domain relies heavily on access to large datasets for training. Although there are several databases of simulated images in literature, there remains a lack of publicly available annotated real-world color polarimetric images of the sky across various weather conditions. Data description: We present here a dataset obtained from a long-term experimental setup designed to collect polarimetric images from a stand-alone camera. The setup utilizes a Division-of-Focal-Plane polarization camera equipped with a fisheye lens mounted on a rotative telescope mount. Furthermore, we obtained the sensor's orientation within the East-North-Up (ENU) frame from a geometrical calibration and an algorithm provided with the database. To facilitate further research in this area, the present sample dataset spanning two months has been made available on a public archive with manual annotations as required by deep learning algorithms. The images were acquired at 10 min intervals and were taken with various exposure times ranging from 33µs to 300ms. [ABSTRACT FROM AUTHOR]

This review article aims to address common research questions in passive polarized vision for robotics. What kind of polarization sensing can we embed into robots? Can we find our geolocation and true north heading by detecting light scattering from the sky as animals do? How should polarization images be related to the physical properties of reflecting surfaces in the context of scene understanding? This review article is divided into three main sections to address these questions, as well as to assist roboticists in identifying future directions in passive polarized vision for robotics. After an introduction, three key interconnected areas will be covered in the following sections: embedded polarization imaging; polarized vision for robotics navigation; and polarized vision for scene understanding. We will then discuss how polarized vision, a type of vision commonly used in the animal kingdom, should be implemented in robotics; this type of vision has not yet been exploited in robotics service. Passive polarized vision could be a supplemental perceptive modality of localization techniques to complement and reinforce more conventional ones. [ABSTRACT FROM AUTHOR]

Many animal species rely on the Earth's magnetic field during navigation, but where in the brain magnetic information is processed is still unknown. To unravel this, we manipulated the natural magnetic field at the nest entrance of Cataglyphis desert ants and investigated how this affects relevant brain regions during early compass calibration. We found that manipulating the Earth's magnetic field has profound effects on neuronal plasticity in two sensory integration centers. Magnetic field manipulations interfere with a typical look-back behavior during learning walks of naive ants. Most importantly, structural analyses in the ants' neuronal compass (central complex) and memory centers (mushroom bodies) demonstrate that magnetic information affects neuronal plasticity during early visual learning. This suggests that magnetic information does not only serve as a compass cue for navigation but also as a global reference system crucial for spatial memory formation. We propose a neural circuit for integration of magnetic information into visual guidance networks in the ant brain. Taken together, our results provide an insight into the neural substrate for magnetic navigation in insects. [ABSTRACT FROM AUTHOR]

True north can be determined on Earth by three means: magnetic compasses, stars, and via the global navigation satellite systems (GNSS), each of which has its own drawbacks. GNSS are sensitive to jamming and spoofing, magnetic compasses are vulnerable to magnetic interferences, and the stars can be used only at night with a clear sky. As an alternative to these methods, nature-inspired navigational cues are of particular interest. Celestial polarization, which is used by insects such as Cataglyphis ants, can provide useful directional cues. Migrating birds calibrate their magnetic compasses by observing the celestial rotation at night. By combining these cues, we have developed a bioinspired optical method for finding the celestial pole during the daytime. This method, which we have named SkyPole, is based on the rotation of the skylight polarization pattern. A polarimetric camera was used to measure the degree of skylight polarization rotating with the Sun. Image difference processes were then applied to the time-varying measurements in order to determine the north celestial pole’s position and thus the observer’s latitude and bearing with respect to the true north. [ABSTRACT FROM AUTHOR]

The Milky Way is used by nocturnal flying and walking insects for maintaining heading while navigating. In this study, we have explored the feasibility of the method for machine vision systems on autonomous vehicles by measuring the visual features and characteristics of the Milky Way. We also consider the conditions under which the Milky Way is used by insects and the sensory systems that support their detection of the Milky Way. Using a combination of simulated and real Milky Way imagery, we demonstrate that appropriate computer vision methods are capable of reliably and accurately extracting the orientation of the Milky Way under an unobstructed night sky. The technique presented achieves angular accuracy of better then ± 2 ° under moderate light pollution conditions but also demonstrates that higher light pollution levels will adversely effect orientation estimates by systems depending on the Milky Way for navigation. [ABSTRACT FROM AUTHOR]

The behaviours and cognitive mechanisms animals use to orient, navigate, and remember spatial locations exemplify how cognitive abilities have evolved to suit a number of different mobile lifestyles and habitats. While spatial cognition observed in vertebrates has been well characterised in recent decades, of no less interest are the great strides that have also been made in characterizing and understanding the behavioural and cognitive basis of orientation and navigation in invertebrate models and in particular insects. Insects are known to exhibit remarkable spatial cognitive abilities and are able to successfully migrate over long distances or pinpoint known locations relying on multiple navigational strategies similar to those found in vertebrate models—all while operating under the constraint of relatively limited neural architectures. Insect orientation and navigation systems are often tailored to each species' ecology, yet common mechanistic principles can be observed repeatedly. Of these, reliance on visual cues is observed across a wide number of insect groups. In this review, we characterise some of the behavioural strategies used by insects to solve navigational problems, including orientation over short-distances, migratory heading maintenance over long distances, and homing behaviours to known locations. We describe behavioural research using examples from a few well-studied insect species to illustrate how visual cues are used in navigation and how they interact with non-visual cues and strategies. [ABSTRACT FROM AUTHOR]

The Milky Way is used by nocturnal flying and walking insects for maintaining heading while navigating. In this study, we have explored the feasibility of the method for machine vision systems on autonomous vehicles by measuring the visual features and characteristics of the Milky Way. We also consider the conditions under which the Milky Way is used by insects and the sensory systems that support their detection of the Milky Way. Using a combination of simulated and real Milky Way imagery, we demonstrate that appropriate computer vision methods are capable of reliably and accurately extracting the orientation of the Milky Way under an unobstructed night sky. The technique presented achieves angular accuracy of better then ±2° under moderate light pollution conditions but also demonstrates that higher light pollution levels will adversely effect orientation estimates by systems depending on the Milky Way for navigation.

Navigation is comprised of a variety of strategies which rely on multiple external cues to shape a navigator's behavioral output. Here, we explored in the ant Veromessor pergandei, the interactions between the information provided by the pheromone trail and the home vector guided by the celestial compass. We found that a cross sensory interaction between the pheromone cue and the path integrator underlies correct orientation during the inbound journey. The celestial compass provides directional information, while the presence of the trail pheromone acts as a critical context cue, triggering distinct behaviors (vector orientation, search, and backtracking). While exposed to the pheromone, foragers orient to the vector direction regardless of vector state, while in the pheromone's absence, the current remaining vector determines the forager's navigational behavior. This interaction also occurs in foragers with no remaining path integrator, relying on the activation of a celestial compass-based memory of the previous trip. Such cue interactions maximize the foragers' return to the nest and inhibit movement off the pheromone trail. Finally, our manipulations continuously rotated foragers away from their desired heading, yet foragers were proficient at counteracting these changes, steering to maintain a correct compass heading even at rotational speeds of ~ 40°/s. [ABSTRACT FROM AUTHOR]

The desert harvester ant (Veromessor pergandei) employs a mixture of social and individual navigational strategies at separate stages of their foraging trip. Individuals leave the nest along a pheromone-based column, travelling 3–40 m before spreading out to forage individually in a fan. Foragers use path integration while in this fan, accumulating a direction and distance estimate (vector) to return to the end of the column (column head), yet foragers' potential use of path integration in the pheromone-based column is less understood. Here we show foragers rely on path integration both in the foraging fan and while in the column to return to the nest, using separate vectors depending on their current foraging stage in the fan or column. Returning foragers displaced while in the fan oriented and travelled to the column head location while those displaced after reaching the column travel in the nest direction, signifying the maintenance of a two-vector system with separate fan and column vectors directing a forager to two separate spatial locations. Interestingly, the trail pheromone and not the surrounding terrestrial cues mediate use of these distinct vectors, as fan foragers briefly exposed to the pheromone cues of the column in isolation altered their paths to a combination of the fan and column vectors. The pheromone acts as a contextual cue triggering both the retrieval of the column-vector memory and its integration with the forager's current fan-vector. [ABSTRACT FROM AUTHOR]

The polarisation pattern of skylight serves as an orientation cue for many invertebrates. Solitary foraging ants, in particular, rely on polarised light to orient along with a number of other visual cues. Yet it is unknown, if this cue is actively used in socially foraging species that use pheromone trails to navigate. Here, we explore the use of polarised light in the presence of the pheromone cues of the foraging trail. The desert harvester ant, Veromessor pergandei, relies on pheromone cues and path integration in separate stages of their foraging ecology (column and fan, respectively). Here, we show that foragers actively orient to an altered overhead polarisation pattern, both while navigating individually in the fan and while on the pheromone-based column. These heading changes occurred during twilight, as well as in the early morning and late afternoon before sunset. Differences in shift size indicate that foragers attend to both the polarisation pattern and the sun's position when available, yet during twilight, headings are dominated by the polarisation pattern. Finally, when the sun's position was experimentally blocked before sunset, shift sizes increased similar to twilight testing. These findings show that celestial cues provide directional information on the pheromone trail. [ABSTRACT FROM AUTHOR]

Maintaining positional estimates of goal locations is a fundamental task for navigating animals. Diverse animal groups, including both vertebrates and invertebrates, can accomplish this through path integration. During path integration, navigators integrate movement changes, tracking both distance and direction, to generate a spatial estimate of their start location, or global vector, allowing efficient direct return travel without retracing the outbound route. In ants, path integration is accomplished through the coupling of pedometer and celestial compass estimates. Within path integration, it has been theorized navigators may use multiple vector memories for way pointing. However, in many instances, these navigators may instead be homing via view alignment. Here, we present evidence that trail-following ants can attend to segments of their global vector to retrace their non-straight pheromone trails, without the confound of familiar views. Veromessor pergandei foragers navigate to directionally distinct intermediate sites via path integration by orienting along separate legs of their inbound route at unfamiliar locations, indicating these changes are not triggered by familiar external cues, but by vector state. These findings contrast with path integration as a singular memory estimate in ants and underscore the system's ability to way point to intermediate goals along the inbound route via multiple vector memories, akin to trapline foraging in bees visiting multiple flower patches. We discuss how reliance on non-straight pheromone-marked trails may support attending to separate vectors to remain on the pheromone rather than attempting straight-line shortcuts back to the nest., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

Foraging ants are known to use multiple sources of information to return to the nest. These cue sets are employed by independent navigational systems including path integration in the case of celestial cues and vision-based learning in the case of terrestrial landmarks and the panorama. When cue sets are presented in conflict, the Australian desert ant species, Melophorus bagoti, will choose a compromise heading between the directions dictated by the cues or, when navigating on well-known routes, foragers choose the direction indicated by the terrestrial cues of the panorama against the dictates of celestial cues. Here, we explore the roles of learning terrestrial cues and delays since cue exposure in these navigational decisions by testing restricted foragers with differing levels of terrestrial cue experience with the maximum (180°) cue conflict. Restricted foragers appear unable to extrapolate landmark information from the nest to a displacement site 8 m away. Given only one homeward experience, foragers can successfully orient using terrestrial cues, but this experience is not sufficient to override a conflicting vector. Terrestrial cue strength increases with multiple experiences and eventually overrides the celestial cues. This appears to be a dynamic choice as foragers discount the reliability of the terrestrial cues over time, reverting back to preferring the celestial vector when the forager has an immediate vector, but the forager's last exposure to the terrestrial cues was 24 hr in the past. Foragers may be employing navigational decision making that can be predicted by the temporal weighting rule. [ABSTRACT FROM AUTHOR]

Three compass systems based on global cues known to exist in migrating birds are reviewed. Two of these systems are based on celestial cues, a time-dependent sun compass and time-independent, i.e. not involving the internal clock, star compass. The third system is the magnetic compass, based on a separate sensory modality, which currently attracts much attention from behavioural ecologists, physiologists and physicists. The complex pattern of hierarchy and interactions between these compass systems is briefly discussed. It is argued that rules of integration of information from different compass cues are likely dependent on ecological and geographic conditions the birds are facing during their journey, so it is likely that no single set of rules is shared by all migrating birds. [ABSTRACT FROM AUTHOR]

Cataglyphis desert ants are famous navigators. Like all central place foragers, they are confronted with the challenge to return home, i.e. relocate an inconspicuous nest entrance in the ground, after their extensive foraging trips. When leaving the underground nest for the first time, desert ants perform a striking behavior, so-called learning walks that are well structured. However, it is still unclear how the ants initially acquire the information needed for sky- and landmark-based navigation, in particular how they calibrate their compass system at the beginning of their foraging careers. Using high-speed video analyses, we show that different Cataglyphis species include different types of characteristic turns in their learning walks. Pirouettes are full or partial rotations (tight turns about the vertical body axis) during which the ants frequently stop and gaze back in the direction of the nest entrance during the longest stopping phases. In contrast, voltes are small walked circles without directed stopping phases. Interestingly, only Cataglyphis ant species living in a cluttered, and therefore visually rich, environment (i.e. C. noda and C. aenescens in southern Greece) perform both voltes and pirouettes. They look back to the nest entrance during pirouettes, most probably to take snapshots of the surroundings. In contrast, C. fortis inhabiting featureless saltpans in Tunisia perform only voltes and do not stop during these turns to gaze back at the nest - even if a set of artificial landmarks surrounds the nest entrance. [ABSTRACT FROM AUTHOR]

Solitary foraging ants have a navigational toolkit, which includes the use of both terrestrial and celestial visual cues, allowing individuals to successfully pilot between food sources and their nest. One such celestial cue is the polarization pattern in the overhead sky. Here, we explore the use of polarized light during outbound and inbound journeys and with different home vectors in the nocturnal bull ant, Myrmecia midas. We tested foragers on both portions of the foraging trip by rotating the overhead polarization pattern by ±45°. Both outbound and inbound foragers responded to the polarized light change, but the extent to which they responded to the rotation varied. Outbound ants, both close to and further from the nest, compensated for the change in the overhead e-vector by about half of the manipulation, suggesting that outbound ants choose a compromise heading between the celestial and terrestrial compass cues. However, ants returning home compensated for the change in the e-vector by about half of the manipulation when the remaining home vector was short (1−2 m) and by more than half of the manipulation when the remaining vector was long (more than 4 m). We report these findings and discuss why weighting on polarization cues change in different contexts.

Summary Ants can navigate over long distances between their nest and food sites using visual cues [ 1, 2 ]. Recent studies show that this capacity is undiminished when walking backward while dragging a heavy food item [ 3–5 ]. This challenges the idea that ants use egocentric visual memories of the scene for guidance [ 1, 2, 6 ]. Can ants use their visual memories of the terrestrial cues when going backward? Our results suggest that ants do not adjust their direction of travel based on the perceived scene while going backward. Instead, they maintain a straight direction using their celestial compass. This direction can be dictated by their path integrator [ 5 ] but can also be set using terrestrial visual cues after a forward peek. If the food item is too heavy to enable body rotations, ants moving backward drop their food on occasion, rotate and walk a few steps forward, return to the food, and drag it backward in a now-corrected direction defined by terrestrial cues. Furthermore, we show that ants can maintain their direction of travel independently of their body orientation. It thus appears that egocentric retinal alignment is required for visual scene recognition, but ants can translate this acquired directional information into a holonomic frame of reference, which enables them to decouple their travel direction from their body orientation and hence navigate backward. This reveals substantial flexibility and communication between different types of navigational information: from terrestrial to celestial cues and from egocentric to holonomic directional memories. Video Abstract [ABSTRACT FROM AUTHOR]

The visual world is rich in linearly polarized light stimuli, which are hidden from the human eye. But many invertebrate species make use of polarized light as a source of valuable visual information. However, exploiting light polarization does not necessarily imply that the electric (e)-vector orientation of polarized light can be perceived as a separate modality of light. In this Review, I address the question of whether invertebrates can detect specific e-vector orientations in a manner similar to that of humans perceiving spectral stimuli as specific hues. To analyze e-vector orientation, the signals of at least three polarization-sensitive sensors (analyzer channels) with different evector tuning axes must be compared. The object-based, imaging polarization vision systems of cephalopods and crustaceans, as well as the water-surface detectors of flying backswimmers, use just two analyzer channels. Although this excludes the perception of specific e-vector orientations, a two-channel system does provide a coarse, categoric analysis of polarized light stimuli, comparable to the limited color sense of dichromatic, 'color-blind' humans. The celestial compass of insects employs three or more analyzer channels. However, that compass is multimodal, i.e. e-vector information merges with directional information from other celestial cues, such as the solar azimuth and the spectral gradient in the sky, masking evector information. It seems that invertebrate organisms take no interest in the polarization details of visual stimuli, but polarization vision grants more practical benefits, such as improved object detection and visual communication for cephalopods and crustaceans, compass readings to traveling insects, or the alert 'water below!' to water-seeking bugs. [ABSTRACT FROM AUTHOR]

Ants use multiple cues for navigating to a food source or nest location. Directional information is derived from pheromone trails or visual landmarks or celestial objects. Some ants use the celestial compass information along with an 'odometer' to determine the shortest distance home, a strategy known as path integration. Some trail-following ants utilise visual landmark information whereas few of the solitary-foraging ants rely on both path integration and visual landmark information. However, it is unknown to what degree trail-following ants use path integration and we investigated this in a trunk-trail-following ant, Iridomyrmex purpureus. Trunk-trail ants connect their nests to food sites with pheromone trails that contain long-lasting orientation information. We determined the use of visual landmarks and the ability to path integrate in a trunk-trail forming ant. We found that experienced animals switch to relying on visual landmark information, and naïve individuals rely on odour trails. Ants displaced to unfamiliar locations relied on path integration, but, surprisingly, they did not travel the entire homebound distance. We found that as the homebound distance increased, the distance ants travelled relying on the path integrator reduced. [ABSTRACT FROM AUTHOR]

Many animal species, including some social hymenoptera, use the visual system for navigation. Although the' insect compound eyes have been well studied, less is known about the second visual system in some insects, the ocelli. Here we demonstrate navigational functions of the ocelli in the visually guided Australian desert ant Melophorus bagoti. These ants are known to rely on both visual landmark learning and path integration. We conducted experiments to reveal the role of ocelli in the perception and use of celestial compass information and landmark guidance. Ants with directional information from their path integration system were tested with covered compound eyes and open ocelli on an unfamiliar test field where only celestial compass cues were available for homing. These full-vector ants, using only their ocelli for visual information, oriented significantly towards the fictive nest on the test field, indicating the use of celestial compass information that is presumably based on polarised skylight, the sun's position or the colour gradient of the sky. Ants without any directional information from their path-integration system (zerovector) were tested, also with covered compound eyes and open ocelli, on a familiar training field where they have to use the surrounding panorama to home. These ants failed to orient significantly in the homeward direction. Together, our results demonstrated that M. bagoti could perceive and process celestial compass information for directional orientation with their ocelli. In contrast, the ocelli do not seem to contribute to terrestrial landmark-based navigation in M. bagoti. [ABSTRACT FROM AUTHOR]

Although several studies have examined how honeybees gauge and report the distance and direction of a food source to their nestmates, relatively little is known about how this information is combined to obtain a representation of the position of the food source. In this study we manipulate the amount of celestial compass information available to the bee during flight, and analyse the encoding of spatial information in the waggle dance as well as in the navigation of the foraging bee. We find that the waggle dance encodes information about the total distance flown to the food source, even when celestial compass cues are available only for a part of the journey. This stands in contrast to how a bee gauges distance flown when it navigates back to a food source that it already knows. When bees were trained to find a feeder placed at a fixed distance in a tunnel in which celestial cues were partially occluded and then tested in a tunnel that was fully open to the sky, they searched for the feeder at a distance that corresponds closely to the distance that was flown under the open sky during the training. Thus, when navigating back to a food source, information about distance travelled is disregarded when there is no concurrent input from the celestial compass. We suggest that bees may possess two different odometers ― a 'community' odometer that is used to provide information to nestmates via the dance, and a 'personal' odometer that is used by an experienced individual to return to a previously visited source. [ABSTRACT FROM AUTHOR]

Migratory birds use multiple sources of compass information for orientation, including the geomagnetic field, the sun, skylight polarization patterns and star patterns. In this paper we review the results of cue-conflict experiments designed to determine the relative importance of the different compass mechanisms, and how directional information from these compass mechanisms is integrated. We focus on cue-conflict experiments in which the magnetic field was shifted in alignment relative to natural celestial cues. Consistent with the conclusions of earlier authors, our analyses suggest that during the premigratory season, celestial information is given the greatest salience and used to recalibrate the magnetic compass by both juvenile and adult birds. Sunset polarized light patterns from the region of the sky near the horizon appear to provide the calibration reference for the magnetic compass. In contrast, during migration, a majority of experiments suggest that birds rely on the magnetic field as the primary source of compass information and use it to calibrate celestial compass cues, i.e. the relative saliency of magnetic and celestial cues is reversed. An alternative possibility, however, is suggested by several experiments in which birds exposed to a cue conflict during migration appear to have recalibrated the magnetic compass, i.e. their response is similar to that of birds exposed to cue conflicts during the premigratory season. The general pattern to emerge from these analyses is that birds exposed to the cue conflict with a view of the entire sunset sky tended to recalibrate the magnetic compass, regardless of whether the cue conflict occurred during the premigratory or migratory period. In contrast, birds exposed to the cue conflict in orientation funnels and registration cages that restricted their view of the region of sky near the horizon (as was generally the case in experiments carried out during the migratory season) did not recalibrate the magnetic compass but, instead, used the magnetic compass to calibrate the other celestial compass systems. If access to critical celestial cues, rather than the timing of exposure to the cue conflict (i.e. premigratory vs migratory), determines whether recalibration of the magnetic compass occurs, this suggests that under natural conditions there may be a single calibration reference for all of the compass systems of migratory birds that is derived from sunset (and possibly also sunrise) polarized light cues from the region of sky near the horizon. In cue-conflict experiments carried out during the migratory season, there was also an interesting asymmetry in the birds' response to magnetic fields shifted clockwise and counterclockwise relative to celestial cues. We discuss two possible explanations for these differences: (1) lateral asymmetry in the role of the right and left eye in mediating light-dependent magnetic compass orientation and (2) interference from the spectral and intensity distribution of skylight at sunset with the response of the light-dependent magnetic compass. [ABSTRACT FROM AUTHOR]

To investigate how visual cues are integrated into a navigational strategy for homing in the Australian sleepy lizard ( Tiliqua rugosa), lizards were displaced beyond their home range, either with full access to visual cues or with no access to visual cues during the displacement. Homeward orientation was significantly worse when lizards were denied visual cues during the displacement than when they were not. However when lizards were displaced with their field of view restricted to the sky, their homeward orientation was equally as good as that of lizards displaced with no visual restriction. These experiments suggest that sleepy lizards use celestial cues to determine the compass bearing of the outward journey, and reverse this bearing to orient in the homeward direction (course reversal). In a subsequent experiment, lizards oriented randomly with respect to home when the parietal eye was entirely covered with a patch during the displacement and return, while control lizards fitted with a sham parietal eye patch were well oriented towards home. In both groups, the lateral eyes were unobstructed and had complete access to visual cues including celestial cues and landmarks. These results suggest that the parietal eye plays a highly significant role in sleepy lizard homing, perhaps mediating a sky polarization compass sense. [ABSTRACT FROM AUTHOR]

• Novomessor cockerelli foragers attend to both path-integration and visual panorama cues. • When in conflict, foragers initially oriented only to the path integrator, yet cue weighting was dynamic. • At unfamiliar sites, foragers travelled ∼85 % of their path integrator before the onset of search. • Panorama analysis showed panorama memories were useful only over short distances from known sites. • Heavy path integration reliance is consistent with a habitat that contains few prominent terrestrial cues. Foraging ants use multiple navigational strategies, including path integration and visual panorama cues, which are used simultaneously and weighted based upon context, the environment and the species' sensory ecology. In particular, the amount of visual clutter in the habitat predicts the weighting given to the forager's path integrator and surrounding panorama cues. Here, we characterize the individual cue use and cue weighting of the Sonoran Desert ant, Novomessor cockerelli, by testing foragers after local and distant displacement. Foragers attend to both a path-integration-based vector and the surrounding panorama to navigate, on and off foraging routes. When both cues were present, foragers initially oriented to their path integrator alone, yet weighting was dynamic, with foragers abandoning the vector and switching to panorama-based navigation after a few meters. If displaced to unfamiliar locations, experienced foragers travelled almost their full homeward vector (∼85 %) before the onset of search. Through panorama analysis, we show views acquired on-route provide sufficient information for orientation over only short distances, with rapid parallel decreases in panorama similarity and navigational performance after even small local displacements. These findings are consistent with heavy path integrator weighting over the panorama when the local habitat contains few prominent terrestrial cues. [ABSTRACT FROM AUTHOR]

• Backtracking inconsistent with established criteria in solitary foraging species. • Backtracking mediated by the foraging column rather than panorama cues. • Backtracking direction is dictated by the sun's position. • Veromessor pergandei do not retain panorama information around nest. Diverse species may adopt behaviourally identical solutions to similar environmental challenges. However, the underlying mechanisms dictating these responses may be quite different and are often associated with the specific ecology or habitat of these species. Foraging desert ants use multiple strategies in order to successfully navigate. In individually foraging ants, these strategies are largely visually-based; this includes path integration and learned panorama cues, with systematic search and backtracking acting as backup mechanisms. Backtracking is believed to be controlled, at least in solitary foraging species, by three criteria: 1) foragers must have recent exposure to the nest panorama, 2) the path integrator must be near zero, and 3) the ant must be displaced to an unfamiliar location. Instead of searching for the nest, under these conditions, foragers head in the opposite compass direction of the one in which they were recently travelling. Here, we explore backtracking in the socially foraging desert harvester ant (Veromessor pergandei) , which exhibits a foraging ecology consisting of a combination of social and individual cues in a column and fan structure. We find that backtracking in V. pergandei , similar to solitary foraging species, is dependent on celestial cues, and in particular on the sun's position. However, unlike solitary foraging species, backtracking in V. pergandei is not mediated by the same criteria. Instead the expression of this behaviour is dependent on the presence of the social cues of the column and the proportion of the column that foragers have completed prior to displacement. [ABSTRACT FROM AUTHOR]

Summary Desert ants ( Cataglyphis ) are famous insect navigators. During their foraging lives, the ants leave their underground colonies for long distances and return to their starting point with fair accuracy [ 1, 2 ]. Their incessantly running path integrator provides them with a continually updated home vector [ 3–5 ]. Directional input to their path integrator is provided by a visual compass based on celestial cues [ 6, 7 ]. However, as path integration is prone to cumulative errors, the ants additionally employ landmark guidance routines [ 8–11 ]. At the start of their foraging lives, they acquire the necessary landmark information by performing well-structured learning walks [ 12, 13 ], including turns about their vertical body axes [ 14 ]. When Cataglyphis noda performs these pirouettes, it always gazes at the nest entrance during the longest of several short stopping phases [ 14 ]. As the small nest entrance is not visible, the ants can adjust their gaze direction only by reading out their path integrator. However, recent experiments have shown that, for adjusting the goal-centered gaze directions during learning walks, skylight cues are not required [ 15 ]. A most promising remaining compass cue is the geomagnetic field, which is used for orientation in one way or the other by a variety of animal species [ 16–25 ]. Here, we show that the gaze directions during the look-back-to-the-nest behavior change in a predictable way to alterations of the horizontal component of the magnetic field. This is the first demonstration that, in insects, a geomagnetic compass cue is both necessary and sufficient for accomplishing a well-defined navigational task. [ABSTRACT FROM AUTHOR]