Cataglyphis Fortis Research Articles

Vector-based and landmark-guided navigation in desert ants inhabiting landmark-free and landmark-rich environments

Two species of desert ants - the North African Cataglyphis fortis and the central Australian Melophorus bagoti - differ markedly in the visual complexity of their natural habitats: featureless salt pans and cluttered, steppe-like terrain, respectively. Here we ask whether the two species differ in their navigational repertoires, in particular, whether in homing they place different emphasis on their vector-based and landmark-based routines. In trying to answer this question, we applied the same experimental paradigms to individual foragers of either species on either continent: training and/or testing with and/or without artificial landmark arrays. We found that the open-terrain species C. fortis runs off its (path integration) home vector much more readily even in unfamiliar landmark settings than the cluttered-terrain species M. bagoti. These data support the hypothesis that C. fortis has a higher propensity to rely on vector-mediated navigation, whereas in the same experimental situations M. bagoti more easily switches to landmark-guided behaviour. In the actual navigational performances, such species-specific propensities are most likely shaped by environment-dependent individual experiences.

Antennal-Lobe Organization in Desert Ants of the Genus Cataglyphis

Desert ants of the genus Cataglyphis possess remarkable visual navigation capabilities. Although Cataglyphis species lack a trail pheromone system, Cataglyphis fortis employs olfactory cues for detecting nest and food sites. To investigate potential adaptations in primary olfactory centers of the brain of C. fortis, we analyzed olfactory glomeruli (odor processing units) in their antennal lobes and compared them to glomeruli in different Cataglyphis species. Using confocal imaging and 3D reconstruction, we analyzed the number, size and spatial arrangement of olfactory glomeruli in C. fortis, C.albicans, C.bicolor, C.rubra, and C.noda. Workers of all Cataglyphis species have smaller numbers of glomeruli (198–249) compared to those previously found in olfactory-guided ants. Analyses in 2 species of Formica – a genus closely related to Cataglyphis – revealed substantially higher numbers of olfactory glomeruli (c. 370), which is likely to reflect the importance of olfaction in these wood ant species. Comparisons between Cataglyphis species revealed 2 special features in C. fortis. First, with c. 198 C. fortis has the lowest number of glomeruli compared to all other species. Second, a conspicuously enlarged glomerulus is located close to the antennal nerve entrance. Males of C. fortis possess a significantly smaller number of glomeruli (c. 150) compared to female workers and queens. A prominent male-specific macroglomerulus likely to be involved in sex pheromone communication occupies a position different from that of the enlarged glomerulus in females. The behavioral significance of the enlarged glomerulus in female workers remains elusive. The fact that C. fortis inhabits microhabitats (salt pans) that are avoided by all other Cataglyphis species suggests that extreme ecological conditions may not only have resulted in adaptations of visual capabilities, but also in specializations of the olfactory system.

Visual associative learning in two desert ant species

Visual cues are important navigational tools for many solitary foraging insects. In addition to information provided by path integration, desert ants learn and use visual cues for homing back to their nest. In this study, we compared the visually based learning of two desert ant species: the North African Cataglyphis fortis and the Australian Melophorus bagoti, each of which lives in ecologically similar but visually different environments. In our experiment, ants’ choice performance was measured by training foragers in a channel system. We used a decision box with two visual stimuli during their homebound trips, with one of the stimuli always being the correct one that provided thoroughfare. To determine any habitat effects on learning, we examined intraspecific comparisons in C. fortis with different nest surroundings. The intraspecific comparison in C. fortis revealed no differences in learning the task. In general, C. fortis showed little learning in our task and the results were similar for ants from barren and cluttered environments. Overall, M. bagoti learned the task faster and had a higher level of accuracy than C. fortis. One explanation for this species-specific difference could be that the cluttered habitat of M. bagoti favours the evolution of visual associative learning more so than the plain habitat of C. fortis.

Visual experience and age affect synaptic organization in the mushroom bodies of the desert ant Cataglyphis fortis

Desert ants of the genus Cataglyphis undergo an age-related polyethism from interior workers involved in brood care and food processing to short-lived outdoor foragers with remarkable visual navigation capabilities. The quick transition from dark to light suggests that visual centers in the ant's brain express a high degree of plasticity. To investigate structural synaptic plasticity in the mushroom bodies (MBs)-sensory integration centers supposed to be involved in learning and memory-we immunolabeled and quantified pre- and postsynaptic profiles of synaptic complexes (microglomeruli, MG) in the visual (collar) and olfactory (lip) input regions of the MB calyx. The results show that a volume increase of the MB calyx during behavioral transition is associated with a decrease in MG numbers in the collar and, less pronounced, in the lip. Analysis of tubulin-positive profiles indicates that presynaptic pruning of projection neurons and dendritic expansion in intrinsic Kenyon cells are involved. Light-exposure of dark-reared ants of different age classes revealed similar effects. The results indicate that this structural synaptic plasticity in the MB calyx is primarily driven by visual experience rather than by an internal program. This is supported by the fact that dark-reared ants age-matched to foragers had MG numbers comparable to those of interior workers. Ants aged artificially for up to 1 year expressed a similar plasticity. These results suggest that the high degree of neuronal plasticity in visual input regions of the MB calyx may be an important factor related to behavior transitions associated with division of labor.

Do desert ants smell the scenery in stereo?

Desert ants, Cataglyphis fortis, navigate individually in the inhospitable saltpans of Tunisia using path integration for long-distance navigation, and visual and olfactory landmarks for fine-scale orientation in the vicinity of the nest entrance. Here, we show in a field experiment that the ants are able to locate the nest entrance within a two-dimensional olfactory array. Ants were trained to forage in an open channel and to memorize the nest entrance relative to four odours that were applied at the corners of an invisible quadratic array. In a test situation, the ants pinpointed the fictive nest only when the odours were present at their learned positions. Our results suggest that the ants had learned the olfactory scenery around their nest. Furthermore, unilaterally antennectomized ants could not pinpoint the nest within a two-dimensional array. Hence, this kind of orientation depends on the simultaneous input of both antennae, that is, on a stereo sense of smell. Until now, insects and mammals, including humans, have only been known to use bilateral sensory input to follow a concentration gradient of an odour. Our evidence suggests that desert ants require a stereo sense of smell to make use of the olfactory scenery around their nest for homing.

Desert ants use foraging distance to adapt the nest search to the uncertainty of the path integrator

Path integration enables desert ants to return to their nest on a direct path. However, the mechanism of path integration is error prone and the ants often miss the exact position of the nest entrance in which case they engage in systematic search behavior. The pattern produced by this search behavior is very flexible and enables the ants to take the errors into account that have been accumulated during foraging and homing. Here, we assess which parameter the desert ant Cataglyphis fortis uses to adapt its systematic search behavior to the uncertainty of its path integrator when deprived of additional external cues. We compared groups of ants that had covered the same distance between their nest and a food source but differed in the overall length of their foraging excursions. Our results show that the width of the ants’ search density profile depends on the distance the ants have ventured out from the nest, that is, the length of the home vector, but not on the tortuousness of their outbound path, that is, the number of steps made during foraging. This distance value is readily available through the path integrator and obviously sufficient to calibrate the ants’ systematic search patterns. Key words: Cataglyphis, density profile, foraging distance, path integrator, systematic search, uncertainty. [Behav Ecol]

Comparing inclined locomotion in a ground-living and a climbing ant species: sagittal plane kinematics

Formicine ants are able to detect slopes in the substrates they crawl on. It was assumed that hair fields between the main segments of the body and between the proximal leg segments contribute to graviception which triggers a change of posture in response to substrate slopes. The sagittal kinematics of two ant species were investigated and compared on different slopes. Cataglyphis fortis, a North African desert ant, is well known for its extraordinary sense of orientation in texturally almost uniform habitats, while Formica pratensis, a common central-European species, primarily uses landmarks and pheromone traces for orientation. A comparison of these two species reveals differences in postural adaptations during inclined locomotion. Only minor slope-dependent angular adjustments were observed. The largest is a 25 degrees head rotation for Cataglyphis, even if the slope is changed by 150 degrees, suggesting dramatic changes in the field of vision. The trunk's pitch adjustment towards the increasing slope is low in both species. On all slopes Cataglyphis achieves higher running speeds than Formica and displays greater slope-dependent variation in body height. This indicates different strategies for coping with changing slopes. These specific aspects have to be reflected in the ants' respective mode of slope perception.

Estimation of homing distance in desert ants,Cataglyphis fortis, remains unaffected by disturbance of walking behaviour

Desert ants, Cataglyphis fortis, use a stride integrator as a distance gauge in their well-studied path integration system (while a skylight compass provides the direction gauge). To further scrutinize the mechanisms of the ant odometer, we tried to disturb the stride integrator by interfering with normal walking behaviour. First, legs that contribute to one of the two leg tripods alternately used in normal walking were selectively amputated. This prevented the normal tripod gait and should interfere with both the normal walking programme controlled by the central nervous system, and normal sensory feedback from the legs. Second, manipulation of the walking substrate in the form of regular corrugations was observed to interfere with normal walking behaviour, at least for corrugation wavelengths (12-25 mm) in the range of normal stride lengths. The animals fell and stumbled, or footfall patterns were entrained to the corrugation wavelength. The relationship between stride length and stride frequency was altered in several situations. Surprisingly, distance estimation and homing performance remained virtually unaffected even by the most severe interferences with walking behaviour. This demonstrates a remarkable robustness of walking behaviour and homing, and it suggests that stride length is determined by robust signals of leg sense organs.

Desert ants do not rely on sky compass information for the perception of inclined path segments

On their foraging excursions, desert ants (Cataglyphis fortis) navigate by means of path integration, exhibiting high precision even in undulating terrain. To avoid errors in their home vector the ants must measure the slopes of ascents and descents in their itinerary. This information is necessary to convert the distances actually walked on the slopes into the ground distance, which is the crucial parameter for vector navigation. However, it is as yet unknown how Cataglyphis perceives the slopes of inclines. We asked whether the ants gauge slopes by means of the changes in the sky's polarization pattern or the sun's position imposed by the unusual head posture when walking on inclines. During training on a ramp, we manipulated the sky compass cues available for the ants. If both sky compass cues, polarization pattern and sun position, were excluded during ramp training, the ants behaved in subsequent tests as if they had never encountered a slope. However, the presence of either of the two compass cues was sufficient for the ants to memorize the sloped path segment. In a third experiment, the supposed polarization cue was excluded by covering the training and test channels with a linear polarization filter. In these tests the ants clearly showed that they had perceived and memorized an ascent also without a change in the polarization pattern. Hence, changes in polarization perception can be ruled out as the crucial cue for gauging slopes, and we conclude that slopes are monitored by a, still unknown, proprioceptive mechanism.

The learning and maintenance of local vectors in desert ant navigation

The desert ant Cataglyphis fortis has at least three types of navigational strategy that can guide it between its nest and a familiar food site. The initial strategy after first finding a food site is based on a path integration memory of the position of the food site with respect to the nest. A second strategy is based on visual snapshot memories of features viewed from near or on the way to the food site. A third strategy uses local vector memories of the direction and length of habitual route segments. We show here that while such local vectors encode sufficient information to guide an individual along both the direction and distance of a route segment, its acquisition and long-term maintenance requires support from the other two strategies. We trained ants along an L-shaped route, designed to show that ants can learn local vectors on the way to a food site. The sharp turn appears to present particular difficulties for the ants. When low bushes 20-30 m from the route were removed, local vectors were briefly unaffected, but then deteriorated. The vectors improved again once the missing bushes were replaced by artificial landmarks. The fragility of local vector memories may permit an ant the flexibility to adapt its route to fluctuations in the distribution of its resources.

Local and global navigational coordinate systems in desert ants

While foraging, the desert ant Cataglyphis fortis keeps track of its position with respect to its nest through a process of path integration (PI). Once it finds food, it can then follow a direct home vector to its nest. Furthermore, it remembers the coordinates of a food site, and uses these coordinates to return to the site. Previous studies suggest, however, that it does not associate any coordinates remembered from previous trips with familiar views such that it can produce a home vector when displaced to a familiar site. We ask here whether a desert ant uses any association between PI coordinates and familiar views to ensure consistent PI coordinates as it travels along a habitual route. We describe an experiment in which we manipulated the PI coordinates an ant has when reaching a distinctive point along a habitual route on the way to a feeder. The subsequent home vectors of the manipulated ants, when displaced from the food-site to a test ground, show that also when a route memory is evoked at a significant point on the way to a food site, C. fortis does not reset its PI coordinates to those it normally has at that point. We use this result to argue that local vector memories, which encode the metric properties of a segment of a habitual route, must be encoded in a route-based coordinate system that is separate from the nest-based global coordinates. We propose a model for PI-based guidance that can account for several puzzling observations, and that naturally produces the route-based coordinate system required for learning and following local vectors.

How flexible is the systematic search behaviour of desert ants?

Foraging desert ants, Cataglyphis fortis , rely on path integration for navigation. If the path integrator has not led them to the exact position of the point of departure, the nest entrance, they start a systematic search for the nest. Previous studies have shown that the longer the preceding outbound run has been the less confidence an ant has in its path integrator. This lower confidence is expressed by wider search loops. In this study, we investigated whether additional cues influence the systematic search patterns of desert ants. We captured ants that were trained to a feeder, either at different points during their inbound journeys or when they were about to enter the nest. They were then transferred to an unfamiliar test area, within which their paths were recorded. Most of the ants captured along their inbound path reeled off the remaining part of their runs and then commenced their nest search, whereas those captured at the nest entrance started searching for the nest entrance immediately. The latter group of ants had far narrower search patterns than ants that were captured during their inbound runs, irrespective of search path length (20, 40 or 50 m). This indicates that the ants' systematic search behaviour is more flexible than assumed hitherto. We compare the results with two previous studies and discuss different potential cues that ants could use to adapt their search patterns.

Smells like home: Desert ants, Cataglyphis fortis, use olfactory landmarks to pinpoint the nest

BackgroundCataglyphis fortis ants forage individually for dead arthropods in the inhospitable salt-pans of Tunisia. Locating the inconspicuous nest after a foraging run of more than 100 meters demands a remarkable orientation capability. As a result of high temperatures and the unpredictable distribution of food, Cataglyphis ants do not lay pheromone trails. Instead, path integration is the fundamental system of long-distance navigation. This system constantly informs a foraging ant about its position relative to the nest. In addition, the ants rely on visual landmarks as geocentric navigational cues to finally pinpoint the nest entrance.ResultsApart from the visual cues within the ants' habitat, we found potential olfactory landmark information with different odour blends coupled to various ground structures. Here we show that Cataglyphis ants can use olfactory information in order to locate their nest entrance. Ants were trained to associate their nest entrance with a single odour. In a test situation, they focused their nest search on the position of the training odour but not on the positions of non-training odours. When trained to a single odour, the ants were able to recognise this odour within a mixture of four odours.ConclusionThe uniform salt-pans become less homogenous if one takes olfactory landmarks into account. As Cataglyphis ants associate environmental odours with the nest entrance they can be said to use olfactory landmarks in the vicinity of the nest for homing.

Surface structure helps desert ants return to known feeding sites

Desert ants, Cataglyphis fortis, return to their nest when they are disturbed during their foraging trips. Training them to a landmark corridor enabled us to induce ants that were captured immediately after leaving the nest and transferred to an unknown area to start their foraging trips.1 However, most of the ants never traveled the entire foraging distance to the feeder, but aborted their runs after the landmark corridor was no longer visible. Therefore, apart from landmark information and path integrator, there are additional cues that determine the ants’ foraging behavior. Considering the reduced straightness of the outbound runs, I argue that surface structure might have a remarkable impact on foraging desert ants.

Repeated training does not improve the path integrator in desert ants

Desert ants, Cataglyphis fortis, return to their nest by means of path integration vectors. By using the reversal of these vectors, they approach previously visited feeding sites again. They adjust these vectors whenever outbound and inbound vector are set into conflict or when they make use of external cues. Here, we examine the influence of repeated training on the accuracy, precision, and straightness of outbound and inbound vectors. We trained desert ants to forage to and fro between their nest and a feeder and made sure that they relied exclusively on their path integrator. Neither the ants’ outbound nor their inbound runs, which, in general, are straighter than the outbound runs, become more accurate, precise, or straighter during repeated training. Hence, repeated training does not improve the path integrator in desert ants.

Three-dimensional orientation in desert ants: context-independent memorisation and recall of sloped path segments

Desert ants (Cataglyphis fortis) navigate by a combination of path integration and landmark-based route memories. Their ability to correct sloped path segments to their ground distances enables them to orientate accurately even in undulating terrain. In this study, we tested whether or not ants incorporate vertical components of an itinerary into their route memory in similar ways as they do with visual landmarks and horizontal changes of direction. In two separate experiments, we trained desert ants to walk over artificial hills and later tested their acceptance of slopes within novel contexts. In the first paradigm, ants had to traverse a hill only on their outbound run, but not on their homebound trip. In a follow-up experiment, we confronted ramp-trained animals with descents in a completely new temporal and spatial context. The animals transferred their newly acquired acceptance of slopes from the outbound to the homebound run as well as to novel foraging trips. Cataglyphis obviously dissociates the experience of sloped path segments from the original context in which they appeared, thus reducing their significance as a navigational aid.

Desert ants adjust their approach to a foraging site according to experience

In the habitat of desert ants, Cataglyphis fortis, a constant wind is usually blowing during the daytime. When visiting a familiar food source, the ants steer some distance downwind of the feeder, rather than attempting a direct approach that might miss small food sources, in particular. In the downwind area, the ants pick up the odor plume emanating from the food and follow it upwind to the prey. This strategy saves considerable walking distance and time. The additional path necessitated by the downwind strategy is only about 0.75 to 2 m, depending on nest–feeder distance, while missing the food on the upwind side results in much longer search trajectories. During the initial three to five visits to a feeding site, downwind distance and length of the approach path are shortened notably, and the approach trajectory is straightened. Desert ants further exhibit considerable short-term flexibility in their approach. Experienced individuals are evidently able to decide upon leaving the nest which direction to choose toward the feeder, depending on current wind direction (that fluctuates slightly during the day). Notable changes in wind direction occur primarily overnight. For larger nest–feeder distances, the animals adjust their approach en route to the altered wind direction during their first foraging trip in the morning.

Desert ants do not acquire and use a three-dimensional global vector

BackgroundDesert ants (Cataglyphis fortis) are central place foragers that navigate by means of path integration. This mechanism remains accurate even on three-dimensional itineraries. In this study, we tested three hypotheses concerning the underlying principles of Cataglyphis' orientation in 3-D: (1) Do the ants employ a strictly two-dimensional representation of their itineraries, (2) do they link additional information about ascents and descents to their 2-D home vector, or (3) do they use true 3-D vector navigation?ResultsWe trained ants to walk routes within channels that included ascents and descents. In choice tests, ants walked on ramps more frequently and at greater lengths if their preceding journey also included vertical components. However, the sequence of ascents and descents, as well as their distance from nest and feeder, were not retraced. Importantly, the animals did not compensate for an enforced vertical deviation from the home vector.ConclusionWe conclude that Cataglyphis fortis essentially represents its environment in a simplified, two-dimensional fashion, with information about vertical path segments being learnt, but independently from their congruence with the actual three-dimensional configuration of the environment. Our findings render the existence of a path integration mechanism that is functional in all three dimensions highly unlikely.

Wind and sky as compass cues in desert ant navigation

While integrating their foraging and homing paths, desert ants, Cataglyphis fortis, depend on external compass cues. Whereas recent research in bees and ants has focused nearly exclusively on the polarization compass, two other compass systems--the sun compass and the wind (anemo) compass--as well as the mutual interactions of all these compass systems have received little attention. In this study, we show that of the two visual compass systems, it is only the polarization compass that invariably outcompetes the wind compass, while the sun compass does so only under certain conditions. If the ants are experimentally deprived of their polarization compass system, but have access simultaneously to both their sun compass and their wind compass, they steer intermediate courses. The intermediate courses shift the more towards the wind compass course, the higher the elevation of the sun is in the sky.

Hair plate mechanoreceptors associated with body segments are not necessary for three-dimensional path integration in desert ants,Cataglyphis fortis

In formicine ants, the hair fields associated with the neck and the petiole (alitrunk-petiole and petiole-gaster joints) have long been established to function in graviception. Here, we examine a possible role of these hair receptors in three-dimensional (3-D) path integration of the (formicine) desert ant, Cataglyphis fortis. Cataglyphis judge the ground distance when travelling over hills, allowing correct homing even in (unpredictably) uneven terrain. We eliminated the function of these hair sensors in graviception either by shaving the hairs or by immobilising the joints monitored by the hair plates. With that major component of their sense of graviception eliminated, one would expect the ants to disregard, or at least misgauge, the ascents and descents performed across hills during outbound journey. The ants should thus consider the (much longer) actual walking trajectory, instead of the base distance, when calculating their homing distance. Surprisingly, neither shaving nor immobilisation of the hair sensillae affected correct path integration, across both uneven terrain (3-D) and level surface. If anything, the ants underestimated homing distance, which may reflect a general, safety-oriented navigation strategy. Animals that had performed the outbound journey with their gaster fixed in a horizontal position underestimated their homing so dramatically that this latter explanation cannot hold.