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.     

Cooperative prey-retrieving in the ant Cataglyphis floricola: an unusual short-distance recruitment

Cataglyphis ants are mostly scavengers adapted to forage individually in arid environments. Although they are widely thought to have lost the capacity of recruitment, we provide evidence that C. floricola foragers that find a large prey near their nest are able to solicit the help of nestmates to carry it cooperatively. After discovering a non-transportable prey, these ants readily return to their nest and stimulate the exit of several recruits. This rudimentary form of recruitment, which is absent in the sympatric species C. rosenhaueri, is only employed when the prey is sufficiently close to the nest entrance (<1 m) and does not allow the food location to be communicated. Instead, C. floricola recruits search for the prey in all directions until they discover it and transport it cooperatively to their nest.     

Landmark cues can change the motivational state of desert ant foragers

Desert ants of the genus Cataglyphis rely on path integration vectors to return to the nest (inbound runs) and back to frequently visited feeding sites (outbound runs). If disturbed, e.g., experimentally displaced on their inbound runs, they continue to run off their home-bound vector, but if disturbed in the same way on their outbound runs, they do not continue their feeder-based vector, but immediately switch on the home-bound state of their path integration vector and return to the nest. Here we show that familiar landmarks encountered by the ants during their run towards the feeder can change the ants’ motivational state insofar that the ants even if disturbed continue to run in the nest-to-feeder direction rather than reverse their courses, as they do in landmark-free situations. Hence, landmark cues can cause the ants to change their motivational state from homing to foraging.     

The hidden spiral: systematic search and path integration in desert ants,Cataglyphis fortis

The main navigational mechanism used by foraging desert ants of the genus Cataglyphis is path integration (dead reckoning). Any such egocentric system of navigation is prone to cumulative navigational errors. Hence, while homing Cataglyphis might have reset its path integration system and yet not arrived at the start of its foraging excursion, the nest entrance. Then it resorts to piloting or performs a systematic search for the nest. The search pattern consists of a system of loops of ever increasing size centred about the origin, i.e. the start of the search. Here we show that underlying the system of loops is a spiral search programme that gets transformed into the observed pattern of loops by the ant's idiosyncratic path-integration algorithm. The ant starts to follow a spiral course, then breaks off this course and walks towards the centre, i.e. to what its path-integration system has computed to be the origin of the search. This reset episode is followed by another spiral course, which is terminated by the next reset, and so forth. After each reset, the spiral gets wider, so that the whole pattern expands. Futhermore, every now and then the spiral might change its sign. Computer simulations based on these simple rules lead to search patterns of the kind actually recorded in Cataglyphis ants. These patterns ensure that those parts of the area in which the target (nest entrance) is most likely to be located are searched most heavily; in other words: the search density profile is adapted to the probability density function of the target.     

Parallel evolution of thermophilia: daily and seasonal foraging patterns of heat‐adapted desert ants: Cataglyphis and Ocymyrmex species*

In the deserts of northern and southern Africa, respectively, ants of the genera Cataglyphisf oerster (Formicinae) and Ocymyrmexe mery (Myrmicinae) occupy the same ecological niche, which comprises that of a strictly diurnal thermophilic scavenger. Their daily foraging activities exhibit a bimodal pattern in summer and unimodality or complete inactivity in winter. The present study investigates whether these overall patterns are a result of endogenous annual activity rhythms of the colony or are triggered directly by the prevailing ambient temperatures. By exploiting various seasonal temperature regimes and, in particular, by creating near‐nest winter conditions experimentally in summer, it is shown that the latter hypothesis is generally true. However, there are daily and annual variations in the temperature set points at which foraging activities start and finish. These temperatures are lower in the winter than in the summer months and, in summer, they are lower in the morning than in the afternoon. The level of foraging activity in the afternoon reaches maximum values at surface temperatures of 60–63 °C. This means that, in summer months, these thermophilic ants concentrate their foraging activities into a period of almost lethal temperature regimes, during which they have to devote a substantial portion of their time outside the nest to respite (i.e. cooling‐off) behaviour.     

Thermoregulatory brood transport in the fire ant, <Emphasis Type="Italic">Solenopsis invicta</Emphasis>

Nest structure in ants is often designed to optimize the colony’s ability to thermoregulate, and this specialization is most highly developed in mound-building ant species. Solenopsis invicta invest a large amount of energy in building mounds and transporting their brood up and down in their nests as a means of thermoregulation. Because few ant species build true mounds, we wanted to determine the effectiveness of these mounds in harvesting solar heat as well as to distinguish what factors (temperature vs. circadian rhythm) govern where fire ants place their brood in the mound and when they place it. We measured temperature patterns in the mound over several days at different depths and under different conditions (under direct sunlight or shade), and then conducted a series of field experiments to manipulate the orientation and time of heating. On cool mornings in spring or fall, surface temperatures of the mound rise at the fastest rate on the side receiving the most direct sunlight (usually the south side). This heating causes a temperature gradient through different depths in the mound, and shows little difference from outside ground temperature at a depth greater than ~40 cm inside the nest. In the morning, fire ants move their brood up into the mound on the side most directly heated, and when temperatures exceed optimal (~32°C) they move their brood down the temperature gradient to lower depths in the nest. In addition to this, mound temperature does not only increase due to direct sunlight, but temperature also increases higher than ground temperatures when the mound is in the shade due to its low specific heat. Experiments in which sunlight was mirrored to the normally shaded side of the mound, or when mounds were heated at night, revealed that S. invicta primarily track temperature patterns and do not rely on behavioral habits or circadian rhythms for the thermoregulatory transport of their brood. When mounds were shaded, S. invicta brood was evenly distributed directly under the surface of the mound rather than aggregating towards a specific side. The fire ant mound is important for thermoregulation because, compared to moundless subterranean nests, it absorbs heat more rapidly both in direct sunlight and shady conditions. Temperature tracking within the nest is key to understanding thermoregulatory placement of fire ant brood, as well as insight into the production of sexual brood and reproduction. Received 9 August 2007; revised 31 January 2008; accepted 7 February 2008.     

Centrifugal waste disposal and the optimization of ant nest craters

We considered how ant, Tapinoma simrothi, colonies dispose of the material generated by their nest excavations in nest entrance craters. We developed a model for optimum crater formation on both flat and sloping ground (although we have not solved it in full generality for sloping ground). We found that the ants, when working on flat ground, show a close approximation to this least costly waste disposal. Craters on slopes may not be optimal even though they will be cheaper to make than ones of a similar volume on level ground. We further tested the model with a manipulative experiment, which also suggested the simple rule of thumb that ants might use. In response to having one-quarter of their crater removed, the ants focused most of their further waste disposal in that quarter. This suggests that their rule of thumb may be to deposit material at the nearest point with an angle of elevation less than that of the optimal angle. This should generate symmetrical craters on flat ground. The model also makes certain additional and testable predictions about the fine structure of craters.     

Hypoxic conditions and oxygen supply in nests of the mangrove ant, Camponotus anderseni, during and after inundation

The small ant Camponotus anderseni lives exclusively in twigs of the mangrove tree Sonneratia alba, and during inundation, the entrance hole is blocked with a soldier’s head which effectively prevents flooding. The nests can be very crowded, with the ants and coccids filling up to 50% of the volume, and due to their metabolic activity, the conditions in the nests during inundation become hypercapnic and hypoxic. Each nest has only one entrance, and the opening is quite small (1.56 ± 0.03 mm). The mean diameter of the galleries is 2.31 ± 0.23 mm, independent of the thickness of the twig and length of the nest. During normal conditions with open nests, the oxygen depletion is substantial in the part of the nest most distant from the opening, and in a 120 mm long nest the oxygen concentration can be as low as 15.7%. During simulated inundation, in which the nest entrances were blocked, the oxygen concentration dropped to very low levels (<0.5%) after one hour. After opening the nest entrance, the oxygen concentration increased again, but for a 100 mm long nest it took nearly 20 minutes before the concentration was back to the normal depressed level. Mathematical modelling of the steady-state oxygen concentrations in the innermost part of the nests shows a lower O₂ concentration than calculated. The time for equilibration of oxygen after inundation is longer than expected for small nests, presumably because the passive diffusion is obstructed by the nest contents. The “dilemma” faced by C. anderseni is to avoid drowning without suffering anoxia or hypercapnia, and they show a remarkable ability to adapt to the extreme conditions in the mangrove and exploit a niche where the density of other ants is insignificant. Received 13 December 2007; revised 30 July 2008; accepted 6 September 2008.     

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.     

Orientation and Communication in the Neotropical Ant Odontomachus bauri Emery (Hymenoptera,Formicidae, Ponerinae)

The Neotropical species Odontomachus bauri employs canopy orientation during foraging and homing. An artificial canopy pattern above the ants is much more effective as an orientation cue than horizontal landmarks or chemical marks. However, both horizontal visual cues and chemical marks on the ground can serve in localizing the nest entrance. Successful O. bauri foragers recruit nestmates to leave the nest and search for food. However, the recruitment signals do not contain directional information. Antennation bouts and pheromones from the pygidial gland most likely serve as stimulating recruitment signals. Secretions from the mandibular and poison gland elicit alarm and attack behavior.     

Synchronization of the behaviour within nests of the antLeptothorax acervorum (Fabricius)—II. Modelling the phenomenon and predictions from the model

Observations on activity of ants of the speciesLeptothorax acervorum show that ants within the nest are inactive for about 72% of their time (Frankset al., 1990.Bull. math. Biol.,52, 597–612). By examination of the activity of individual ants it is demonstrated that activity bouts of individuals are highly synchronized. The bursts of activity detected by Frankset al. occurred three to four times per hour. In this paper we develop a model to describe the phenomenon. As a result of the interdependence of the number of active ants within the nest and the high level of community activity some predictions are made, which are supported by experimental data in a quantitative way. In case of starvation the number of active ants will increase and no rhythms should occur. When proportionally more brood is present the rhythms should occur with a higher frequency. Eventually the rhythm breaks down and a stable equilibrium is reached.     

Time-courses of memory decay in vector-based and landmark-based systems of navigation in desert ants, Cataglyphis fortis

In foraging and homing, desert ants of the genus Cataglyphis employ two different systems of navigation: a vector-based or dead-reckoning mechanism, depending on angles steered and distances travelled, and a landmark-based piloting mechanism. In these systems the ants use either celestial or terrestrial visual information, respectively. In behavioural experiments we investigated how long these types of information are preserved in the ant's memory, i.e. how long the ants are able to orient properly in either way. To answer this question, ants were tested in specific dead-reckoning and piloting situations, whereby the two vector components, direction and distance, were examined separately. The ability to follow a particular vector course vanishes rapidly. Information about a given homing direction is lost from the 6th day on (the time constant of the exponential memory decay function is τ = 4.5 days). The homing distances show a significantly higher dispersion from the 4th day on (τ = 2.5 days). Having learned a constellation of landmarks positioned at the corners of an equidistant triangle all ants were oriented properly after 10 days in captivity, and 64% of the ants exhibited extremely precise orientation performances even when tested after 20 days. Thus, the memory decay functions have about the same short time-course for information on distance and direction, i.e. information used for dead-reckoning. In contrast, landmark-based information used in pinpointing the nest entrance is stored over the entire lifetime of a Cataglyphis forager. Accepted: 18 January 1997     

Thermoregulatory trade-offs result from vegetation removal by a harvester ant

Abstract.  1. Analyses of ecological trade-offs help to explain how organisms balance competing demands. Harvester ants ( Pogonomyrmex occidentalis ), are conspicuous residents of shortgrass prairie in western North America; worker P. occidentalis actively clear all vegetation from the immediate vicinity of their large gravel mounds. This study is based on the prediction that vegetation clearing yields a thermal trade-off by increasing soil temperatures; during cool periods the resulting increase in soil temperature opens new time windows for activity, while during hot periods the soil temperature is more likely to exceed the maximum thermal tolerance for this species. To test the hypothesis that daily and seasonal trade-offs in ant activity result from vegetation removal, the effects of experimentally applied shade on activity patterns were measured.
2. Harvester ant activity correlated highly with ground temperature; experimental shading of ant mounds shifted daily activity patterns by lowering ground temperature. Shading in the morning significantly delayed the onset of ant activity by preventing solar warming of the mound. In contrast, mid-day experimental shading prevented elevation of ground temperatures to above 50 °C and allowed ants to remain active when lethally high temperatures would normally force them inside the mound.
3. A model derived from field data predicted surface ground temperature (and therefore ant activity) based on air temperature and solar radiation, under conditions of sun and shade. For each of six seasons modelled, shade removal yielded a net gain of activity time. These results indicate that vegetation removal by harvester ants produces an advantageous thermoregulatory effect by helping to maximise activity time.     

Simple rules based on pile slope are used in the self organization of sand pile formation by <Emphasis Type="Italic">Pheidole oxyops</Emphasis> ants

We tested the hypothesis that slope influences where worker ants deposit excavated soil on piles near the nest entrance. We predicted that ants will deposit their load near the top of a pile where the slope changes from upward to downward, to prevent material rolling back towards the entrance. We tested this hypothesis by studying five natural colonies of Pheidole oxyops ants at a field site at S?o Sim?o, Brazil. At this site, each colony was dumping sandy soil excavated from its underground nest in a crescent-shaped pile c. 13 cm from and perpendicular to the nest entrance. Each nest was given an experimental sand pile of symmetrical curved cross section on a plywood platform that could be tilted 15 degrees up or down. From videos, the locations where individual ants dumped their soil loads were measured in relation to the inner (position = 0) and outer (position = 1) edges of the pile. When the platform was tilted down the ants deposited their loads significantly closer to the inner edge (0.458 ± 0.007) than when not tilted (0.530 ± 0.006). When the platform was tilted up the ants deposited their loads significantly further from the inner edge (0.626 ± 0.006) than when not tilted (0.522 ± 0.006). These results support the hypothesis that ants use pile slope in deciding where to dump their load. A similar rule is probably used in other ant species that place excavated soil in steep piles near the nest entrance. Received 5 February 2007; revised 10 June 2007; accepted 9 October 2007.     

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.     

Search strategies of ants in landmark-rich habitats

Search is an important tool in an ant’s navigational toolbox to relocate food sources and find the inconspicuous nest entrance. In habitats where landmark information is sparse, homing ants travel their entire home vector before searching systematically with ever increasing loops. Search strategies have not been previously investigated in ants that inhabit landmark-rich habitats where they typically establish stereotypical routes. Here we examine the search strategy in one such ant, Melophorus bagoti, by confining their foraging in one-dimensional channels to determine if their search pattern changes with experience, location of distant cues and altered distance on the homebound journey. Irrespective of conditions, we found ants exhibit a progressive search that drifted towards the fictive nest and beyond. Segments moving away from the start of the homeward journey were longer than segments heading back towards the start. The right tail distribution of segment lengths was well fitted by a power function, but slopes less than −3 on a log-log plot indicate that the process cannot be characterized as Lévy searches that have optimal slopes near –2. A double exponential function fits the distribution of segment lengths better, supporting another theoretically optimal search pattern, the composite Brownian walk.     

Gaseous templates in ant nests

We apply a diffusion model to the atmosphere of ant nests. With particular reference to carbon dioxide (CO2), we explore analytically and numerically the spatial and temporal patterns of brood- or worker-produced gases in nests. The maximum concentration within a typical one-chamber ant nest with approximately 200 ants can reach 12.5 times atmospheric concentration, reaching 95% of equilibrium concentrations within 15 min. Maximum concentration increases with increasing number of ants in the nest (or production rate of the gas), distance between the centre of the nest ants and the nest entrance, entrance length, wall thickness, and with decreasing entrance width, wall permeability and diffusion coefficient. The nest can be divided into three qualitatively distinct regions according to the shape of the gradient: a plateau of high concentration in the back half of the nest; an intermediate region of increasingly steep gradient towards the entrance; and a steep linear gradient in the entrance tunnel. These regions are robust to changes in gas concentrations, but vary with changes in nest architecture. The pattern of diffusing gases contains information about position and orientation relative to gas sources and sinks, and about colony state, including colony size, activity state and aspects of nest architecture. We discuss how this diffusion pattern may act as a "dynamic template", providing local cues which trigger behavioural acts appropriate to colony needs, which in turn may feed back to changes in the gas template. In particular, wall building occurs along lines of similar concentration for a variety of nest geometries; there is surprising convergence between the period of cycles of synchronously active ants and the time taken for CO2 levels to equilibrate; and the qualitatively distinct regions of the "dynamic template" correspond to regions occupied by different groups of ants.     

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.     

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.     

Complex Nesting Behavior by Two Neotropical Species of the Ant Genus Stenamma (Hymenoptera: Formicidae)1

Two Neotropical ant species, Stenamma expolitum Smith and S. alas new species (described here), exhibit three unusual nesting behaviors: (1) they build architecturally sophisticated nest entrances that elevate the nest opening away from the surface; (2) they maintain multiple identical nests but occupy only one of them; and (3) they keep a round “door pebble” at the nest entrance, with which they plug the opening in response to army ants. Adaptive hypotheses for these behaviors are discussed, including the possibility that there are multiple lines of defense against army ant predation.