Kompass im Kopf

Ant compass – how desert ants learn to navigate Successful spatial orientation is a daily challenge for many animals. Cataglyphis desert ants are famous for their navigational performances. They return to the nest after extensive foraging trips without any problems. How do ants take their navigational systems into operation? After conducting different tasks in the dark nest for several weeks, they become foragers under bright sun light. This transition requires both a drastic switch in behavior and neuronal changes in the brain. Experienced foragers mainly rely on visual cues. They use a celestial compass and landmark panoramas. For that reason, naïve ants perform stereotype learning walks to calibrate their compass systems and acquire information about the nest's surroundings. During their learning walks, the ants frequently look back to the nest entrance to learn the homing direction. For aligning their gazes, they use the earth's magnetic field as a compass reference. This magnetic compass in Cataglyphis ants was previously unknown.     

Context-dependent navigation in a collectively foraging species of ant, Messor cephalotes

More than 100 years of scientific research has provided evidence for sophisticated navigational mechanisms in social insects. One key role for navigation in ants is the orientation of workers between food sources and the nest. The focus of recent work has been restricted to navigation in individually foraging ant species, yet many species do not forage entirely independently, instead relying on collectively maintained information such as persistent trail networks and/or pheromones. Harvester ants use such networks, but additionally, foragers often search individually for food either side of trails. In the absence of a trail, these ‘off-trail’ foragers must navigate independently to relocate the trail and return to the nest. To investigate the strategies used by ants on and off the main trails, we conducted field experiments with a harvester ant species, Messor cephalotes, by transferring on-trail and off-trail foragers to an experimental arena. We employed custom-built software to track and analyse ant trajectories in the arena and to quantitatively compare behaviour. Our results indicate that foragers navigate using different cues depending on whether they are travelling on or off the main trails. We argue that navigation in collectively foraging ants deserves more attention due to the potential for behavioural flexibility arising from the relative complexity of journeys between food and the nest.     

Learning and time‐dependent cue choice in the desert ant,Melophorus bagoti

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.     

Interactions Increase Forager Availability and Activity in Harvester Ants

Social insect colonies use interactions among workers to regulate collective behavior. Harvester ant foragers interact in a chamber just inside the nest entrance, here called the ''entrance chamber''. Previous studies of the activation of foragers in red harvester ants show that an outgoing forager inside the nest experiences an increase in brief antennal contacts before it leaves the nest to forage. Here we compare the interaction rate experienced by foragers that left the nest and ants that did not. We found that ants in the entrance chamber that leave the nest to forage experienced more interactions than ants that descend to the deeper nest without foraging. Additionally, we found that the availability of foragers in the entrance chamber is associated with the rate of forager return. An increase in the rate of forager return leads to an increase in the rate at which ants descend to the deeper nest, which then stimulates more ants to ascend into the entrance chamber. Thus a higher rate of forager return leads to more available foragers in the entrance chamber. The highest density of interactions occurs near the nest entrance and the entrances of the tunnels from the entrance chamber to the deeper nest. Local interactions with returning foragers regulate both the activation of waiting foragers and the number of foragers available to be activated.     

Mapping the navigational knowledge of individually foraging ants,Myrmecia croslandi

Ants are efficient navigators, guided by path integration and visual landmarks. Path integration is the primary strategy in landmark-poor habitats, but landmarks are readily used when available. The landmark panorama provides reliable information about heading direction, routes and specific location. Visual memories for guidance are often acquired along routes or near to significant places. Over what area can such locally acquired memories provide information for reaching a place? This question is unusually approachable in the solitary foraging Australian jack jumper ant, since individual foragers typically travel to one or two nest-specific foraging trees. We find that within 10 m from the nest, ants both with and without home vector information available from path integration return directly to the nest from all compass directions, after briefly scanning the panorama. By reconstructing panoramic views within the successful homing range, we show that in the open woodland habitat of these ants, snapshot memories acquired close to the nest provide sufficient navigational information to determine nest-directed heading direction over a surprisingly large area, including areas that animals may have not visited previously.     

Wüstennavigatoren en miniature

Desert navigators en miniature Cataglyphis, a strictly diurnal, heat‐tolerant, high‐speed desert ant, employs a path integrator as its main navigational means. By continually measuring directions steered and distances covered the path integrator computes a navigation vector, which can lead the ant directly back to its central place, the nest, and to any point which it has visited before. The path integration vector receives compass information from the pattern of polarized light in the sky (via a set of specialized photoreceptors at the dorsal rim of the eye), and derives information about travel distance from a stride integrator (pedometer) and an optic‐flow meter exploiting self‐induced image motion across the ventral retina. The path integrator is fully functional already at the beginning of the ant's foraging life. Later it keeps running whenever the ant is on a foraging excursion irrespective of whether other navigational tools are at work as well. Finally it provides a scaffold for landmark learning. View‐based landmark information is acquired by taking panoramic “snapshots” at certain places and routes. By comparing this memorized visual information with the actual one received during later journeys the ants are able to return to familiar places and to follow familiar routes even without the aid of the path integrator. The ant's navigational performances known to date can be simulated by designing a decentralized network, in which the individual tools are interconnected in flexible and context dependent ways.     

Path integration controls nest-plume following in desert ants

The desert ant Cataglyphis fortis is equipped with sophisticated navigational skills for returning to its nest after foraging. The ant's primary means for long-distance navigation is path integration, which provides a continuous readout of the ant's approximate distance and direction from the nest. The nest is pinpointed with the aid of visual and olfactory landmarks. Similar landmark cues help ants locate familiar food sites. Ants on their outward trip will position themselves so that they can move upwind using odor cues to find food. Here we show that homing ants also move upwind along nest-derived odor plumes to approach their nest. The ants only respond to odor plumes if the state of their path integrator tells them that they are near the nest. This influence of path integration is important because we could experimentally provoke ants to follow odor plumes from a foreign, conspecific nest and enter that nest. We identified CO(2) as one nest-plume component that can by itself induce plume following in homing ants. Taken together, the results suggest that path-integration information enables ants to avoid entering the wrong nest, where they would inevitably be killed by resident ants.     

Interactions with Combined Chemical Cues Inform Harvester Ant Foragers' Decisions to Leave the Nest in Search of Food

Social insect colonies operate without central control or any global assessment of what needs to be done by workers. Colony organization arises from the responses of individuals to local cues. Red harvester ants (Pogonomyrmex barbatus) regulate foraging using interactions between returning and outgoing foragers. The rate at which foragers return with seeds, a measure of food availability, sets the rate at which outgoing foragers leave the nest on foraging trips. We used mimics to test whether outgoing foragers inside the nest respond to the odor of food, oleic acid, the odor of the forager itself, cuticular hydrocarbons, or a combination of both with increased foraging activity. We compared foraging activity, the rate at which foragers passed a line on a trail, before and after the addition of mimics. The combination of both odors, those of food and of foragers, is required to stimulate foraging. The addition of blank mimics, mimics coated with food odor alone, or mimics coated with forager odor alone did not increase foraging activity. We compared the rates at which foragers inside the nest interacted with other ants, blank mimics, and mimics coated with a combination of food and forager odor. Foragers inside the nest interacted more with mimics coated with combined forager/seed odors than with blank mimics, and these interactions had the same effect as those with other foragers. Outgoing foragers inside the nest entrance are stimulated to leave the nest in search of food by interacting with foragers returning with seeds. By using the combined odors of forager cuticular hydrocarbons and of seeds, the colony captures precise information, on the timescale of seconds, about the current availability of food.     

Coupled oscillators and activity waves in ant colonies

We investigated the phenomenon of activity cycles in ants, taking into account the spatial structure of colonies. In our study species, Leptothorax acervorum, there are two spatially segregated groups in the nest. We developed a model that considers the two groups as coupled oscillators which can produce synchronized activity. By investigating the effects of noise on the model system we predicted how the return of foragers affects activity cycles in ant colonies. We tested these predictions empirically by comparing the activity of colonies under two conditions: when foragers are and are not allowed to return to the nest. The activity of the whole colony and of each group within the colony was studied using image analysis. This allowed us to reveal the spatial pattern of activity wave propagation in ant colonies for the first time.     

Garden ant homing behavior in a maze task based on local visual cues

Although it has been shown that visual cues play an essential role in navigation by the garden ant Lasius niger, no previous studies have addressed the way in which information from local visual cues is acquired and utilized in navigation. We found that in the absence of pheromone trails, ants whose homing motivation was triggered by feeding returned to the nest following local visual cues. In our experiments, the ants travelled through a maze to reach a feeder. They explored the maze and sometimes became trapped in its dead ends. We found that the ants more effectively used visual cues during their homeward journey if they experienced a dead end during their outward journey. This result suggested that the ants used the information acquired from visual cues during the outward journey to avoid a dead end on their return journey.     

Compound Eye Adaptations for Diurnal and Nocturnal Lifestyle in the Intertidal Ant,Polyrhachis sokolova

The Australian intertidal ant, Polyrhachis sokolova lives in mudflat habitats and nests at the base of mangroves. They are solitary foraging ants that rely on visual cues. The ants are active during low tides at both day and night and thus experience a wide range of light intensities. We here ask the extent to which the compound eyes of P. sokolova reflect the fact that they operate during both day and night. The ants have typical apposition compound eyes with 596 ommatidia per eye and an interommatidial angle of 6.0°. We find the ants have developed large lenses (33 µm in diameter) and wide rhabdoms (5 µm in diameter) to make their eyes highly sensitive to low light conditions. To be active at bright light conditions, the ants have developed an extreme pupillary mechanism during which the primary pigment cells constrict the crystalline cone to form a narrow tract of 0.5 µm wide and 16 µm long. This pupillary mechanism protects the photoreceptors from bright light, making the eyes less sensitive during the day. The dorsal rim area of their compound eye has specialised photoreceptors that could aid in detecting the orientation of the pattern of polarised skylight, which would assist the animals to determine compass directions required while navigating between nest and food sources.     

Light intensity limits foraging activity in nocturnal and crepuscular bees

A crepuscular or nocturnal lifestyle has evolved in bees severaltimes independently, probably to explore rewarding pollen sourceswithout competition and to minimize predation and nest parasites.Despite these obvious advantages, only few bee species are nocturnal.Here we show that the sensitivity of the bee apposition eyeis a major factor limiting the ability to forage in dim light.We present data on eye size, foraging times, and light levelsfor Megalopta genalis (Augochlorini, Halictidae) in Panama,and Lasioglossum (Sphecodogastra) sp. (Halictini, Halictidae)in Utah, USA. M. genalis females forage exclusively during twilight,but as a result of dim light levels in the rain forest, theyare adapted to extremely low intensities. The likely factorlimiting their foraging activity is finding their nest entranceon return from a foraging trip. The lowest light intensity atwhich they can do this, both in the morning and the evening,is 0.0001 cd m^–2. Therefore, they leave the nest at dimmerlight levels in the morning than in the evening. Lasioglossum(Sphecodogastra) foraging is limited by light intensity in theevening, but probably by temperature in the morning in the temperateclimate of Utah. We propose that the evolution of nocturnalityin bees was favored by the large variance in the size of females.     

Field study on the foraging characteristics of a ponerine ant,Hagensia havilandi Forel

Summary A field study of the foraging strategy used by the ponerine ant,Hagensia havilandi is reported. They have permanent nests in the leaf litter of coastal forests.H. havilandi is a diurnal forager and collects a variety of live and dead arthropods. These predatory ants exhibit individual foraging with no cooperation in the search for or retrieval of food items. Three colonies were observed and showed similar temporal and spatial foraging patterns. The paths of individual ants were followed and the results showed that the foragers exhibit area fidelity, and return to the nest via a direct route on finding on prey item. Several foragers did not return to the nest at dusk but returned the following morning. Occasionally a limited amount of tandem recruitment was displayed.     

Use of Long-Term Stored Vector Information in the Neotropical Ant <Emphasis Type="BoldItalic">Gigantiops destructor</Emphasis>

We investigated how the formicine ant Gigantiops destructor can use vector information to navigate within the cluttered environment of the rain forest. Displaced foragers use skylight information to move in the theoretical feeder-to-nest direction, whether they are prevented from updating their path-integrator during foraging or captured at the departure from their nest, i.e. with a current accumulator state very close to zero. Only ants that have collected food are able to download a long-term stored reference vector pointing in the nest direction, irrespective of the current accumulator state of their path-integrator stored in a working memory and independent of familiar landmarks. Depending on the release sites, ants that became lost at a maximum distance of 50 cm could still hit and recognize their familiar route, or they engaged in a systematic search for it centered on the release sites. In contrast to Cataglyphis desert ants, Gigantiops ants do not rely primarily on the current accumulator state of their egocentric path integrator. Such a long-term vector-based navigation primed by food capture is well adapted for a tropical ant foraging during periods spanning several hours. This could prevent the numerous cumulative errors in the evaluation of the angles steered that might result from a continuously running path-integrator operating during complex foraging patterns performed at ground or arboreal levels and during passive displacement in response to heavy rain.     

Navigation without vision: bumblebee orientation in complete darkness

In flight cages, worker bumblebees (Bombus impatiens) spontaneously explored the surroundings of their nest and foraged in complete darkness, by walking instead of flying, from feeders up to 150 cm away from the nest. This behaviour was wholly unexpected in these classically visual foragers. The finding provides a controlled system for dissecting possible non-visual components of navigation used in daylight. It also allows us to isolate navigation mechanisms used in naturally dark situations, such as in the nest. Using infrared video, we mapped walking trails. We found that bumblebees laid odour marks. When such odour cues were eliminated, bees maintained correct directionality, suggesting a magnetic compass. They were also able to assess travel distance correctly, using an internal, non-visual, measure of path length. Path integration was not employed. Presumably, this complex navigational skill requires visual input in bees.     

Visual cue learning and odometry in guiding the search behavior of desert ants, Melophorus bagoti, in artificial channels

Terrestrial panoramic cues, path integration and search behavior are the main navigational strategies used by ants to locate food and find their way back to the nest. Searching becomes important when the other navigational cues are either not available or cannot provide sufficient information to pinpoint the goal. When searching in one-dimensional channels Melophorus bagoti ants exhibit a systematic drift in the starting-point-to-goal direction as they turn back and forth, sometimes past the goal location ( Narendra et al., 2008). Here we show that this drift in channels is not a stereotypical part of the search behavior in these ants. It rather depends on the conditions of training. In experiments in which the nest entrance is located not at the end but at the side of the channel, forward drift is not always part of the nest search. Experiments on food searches showed that with the food source at the end of the channel, ants performed a linear drift in the starting-point-to-food direction. With food at the side of the channel, they showed a less pronounced drift toward the food source. In this constrained environment, especially with the goal at the end of the channel, ants seem to learn a routine such as ‘run along the channel’, and mix this routine with their usual strategy of turning back and forth in search.     

Foraging tempo in two woodland and species

Running speeds of Myrmica punctiventris and Aphaenogaster rudis workers were measured, and a good correspondence between laboratory and field behaviour was obtained. In the laboratory, foraging tempo and foraging efficiency were calculated for two colony sizes and five patterns of prey availability. Running speeds were strongly dependent on colony size for both species; when retrieving prey, foragers from large colonies ran significantly faster than those from small colonies. In addition, ants searching for prey ran more slowly than those returning to the nest with prey. Efficiency, measured as the propensity to return to the nest in a straight line, was most strongly a function of distance from the nest. Finally, no relationship between an ant's speed and its efficiency of return was found.     

Colony variation in the collective regulation of foraging by harvester ants

This study investigates variation in collective behavior in a natural population of colonies of the harvester ant, Pogonomyrmex barbatus. Harvester ant colonies regulate foraging activity to adjust to current food availability; the rate at which inactive foragers leave the nest on the next trip depends on the rate at which successful foragers return with food. This study investigates differences among colonies in foraging activity and how these differences are associated with variation among colonies in the regulation of foraging. Colonies differ in the baseline rate at which patrollers leave the nest, without stimulation from returning ants. This baseline rate predicts a colony's foraging activity, suggesting there is a colony-specific activity level that influences how quickly any ant leaves the nest. When a colony's foraging activity is high, the colony is more likely to regulate foraging. Moreover, colonies differ in the propensity to adjust the rate of outgoing foragers to the rate of forager return. Naturally occurring variation in the regulation of foraging may lead to variation in colony survival and reproductive success.     

Ants versus spiders: interference competition between two social predators

Interference competition can profoundly influence the outcome of species interactions and may lead to either coexistence or exclusion. Our understanding of how interference can lead to coexistence remains incomplete, particularly when interference fails to result in resource partitioning. I document a novel form of interference competition between an ant (Myrmecia pyriformis) and a social spider (Delena cancerides) with similar foraging patterns. Of 120 nest boxes occupied by D. cancerides in the field, 7 (6 %) were invaded by M. pyriformis ants over a 2-month period. After eliminating spiders from the nest boxes, the ants proceeded to fill the boxes with debris, rendering them useless to the spiders. The ants do not occupy the nest boxes; thus, interference occurs at a resource that is necessary to one species, but not the other. I discuss how further research into this system may improve our understanding of how interference competition can support coexistence. I also suggest modeling exploitation and interference competition on multiple resources to align with this and other empirical examples where different forms of competition occur for different resources.     

Visual Navigation during Colony Emigration by the Ant Temnothorax rugatulus

Many ants rely on both visual cues and self-generated chemical signals for navigation, but their relative importance varies across species and context. We evaluated the roles of both modalities during colony emigration by Temnothorax rugatulus. Colonies were induced to move from an old nest in the center of an arena to a new nest at the arena edge. In the midst of the emigration the arena floor was rotated 60°around the old nest entrance, thus displacing any substrate-bound odor cues while leaving visual cues unchanged. This manipulation had no effect on orientation, suggesting little influence of substrate cues on navigation. When this rotation was accompanied by the blocking of most visual cues, the ants became highly disoriented, suggesting that they did not fall back on substrate cues even when deprived of visual information. Finally, when the substrate was left in place but the visual surround was rotated, the ants'' subsequent headings were strongly rotated in the same direction, showing a clear role for visual navigation. Combined with earlier studies, these results suggest that chemical signals deposited by Temnothorax ants serve more for marking of familiar territory than for orientation. The ants instead navigate visually, showing the importance of this modality even for species with small eyes and coarse visual acuity.