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.     

Combined use of pheromone trails and visual landmarks by the common garden ant Lasius niger

This study investigated the relative importance of pheromone trails and visual landmarks on the ability of Lasius niger foragers to relocate a previously used food source. Colonies formed foraging trails to a 1-M sucrose feeder. Sections of this trail were then presented back to the same colony after variable time intervals. Individual outgoing foragers were observed to determine if they walked for 15 cm in the direction of the feeder or not. On newly established pheromone trails formed by 500 ant passages, 77% of the foragers walked in the correct direction vs 31% for control foragers (no trail pheromone). Pheromone trails decayed to the control levels in 20–24 h. Trails formed with fewer ant passages (125 or 30) decayed quicker. The use of visual landmarks was investigated by using trails with outgoing foragers from the colony that established the trail, either in the same room or in a different room, with different visual landmarks, to that used during trail establishment. Approximately 20% more ants walked in the correct direction in the same room vs the different room. This difference decreased to around 10% 2 h after trail establishment, indicating that the ants in the different room were learning the new visual cues to navigate by. Our results show that visual landmarks and pheromone trails are approximately equally useful in initially guiding L. niger foragers to food locations and that these two information sources have a complementary function.     

Predator recognition and evasive behavior by sweat bees, <Emphasis Type="Italic"> Lasioglossum umbripenne </Emphasis>(Hymenoptera: Halictidae), in response to predation by ants, <Emphasis Type="Italic"> Ectatomma ruidum </Emphasis>(Hymenoptera: Formicidae)

Ectatomma ruidum is an abundant soil-nesting Neotropical ant, which displays extensive behavioral flexibility during foraging activities. We studied here one unusual element of their behavioral repertoire: ambush predation. A worker of E. ruidum waits near a nest of a social sweat bee, Lasioglossum umbripenne, lunging at incoming bees, or less frequently, at departing bees. However, bees detected ambushing ants and modified their behavior. Dead ants placed at bees' nest entrances significantly decreased bee activity, indicating that bees recognized dead ants as potential predators. Neither simple black models (square and rectangle) nor olfactory cues had any effect on overall bee activity. A returning bee usually approached her entrance and immediately entered, but if an ant was waiting at the nest, a bee was significantly more likely to abort the first approach flight and then to re-approach the nest on the side opposite the ant's position. As models became increasingly ant-like, returning bees more frequently aborted their first approach flight, expressing other behaviors before entering nests. These behaviors included withdrawal followed by an approach from a different direction; zigzagging flights, either from a distance or close to the entrance or even a close inspection; landing a short distance from the nest, then approaching on foot or waiting for several seconds before entering. Ants responded with effective counter-behaviors. Behavioral flexibility in nest entering/exiting by L. umbripenne and in hunting strategy by E. ruidum shows the complexity of this predator-prey relationship, and illustrates the importance of information processing by both species involved in determining the outcome of the interspecific interaction.     

A South East Asian ponerine ant of the genus Leptogenys (Hym., Form.) with army ant life habits

Summary The emigration and raiding behavior of the SE Asian ponerine ant Leptogenys sp. 1, which resembles L. mutabilis, were observed in the field (Ulu Gombak, Malaysia). The ants formed monogynous colonies that consisted of up to 52 100 workers. The bivouac sites of this species were found in leaf litter, rotten logs, ground cavities, etc., and were rarely modified by the ants. The colonies stayed in these temporary nests for several hours to 10 days; afterwards, they moved to a new nest site. The emigration distances ranged from 5–58 m. Since nest changing takes place at irregular intervals, and pupae and larvae are always present in the nest relocations of Leptogenys sp. 1, the emigration behavior is not linked to a synchronized brood development. Leptogenys sp. 1 is a nocturnal forager; in our study, up to 42 600 workers participated in each raid. The ants move forward on a broad front; behind the swarm a fan-shaped network of foraging columns converges to form a main trunk trail. A new system of foraging trails is developed in each raid. The workers search for their prey collectively; they attack and retrieve the booty together. The diet of Leptogenys sp. 1 consists mainly of arthropods. Army ant behavior is characterized by (1) formation of large monogynous colonies, (2) frequent emigrations, and (3) mass raids in which all foraging activities are carried out collectively. Since Leptogenys sp. 1 performs these typical army ant behavior patterns, this species represents the army ant ecotype. However, this species differs considerably from army ant species that have synchronized broods and huge colonies with dichthadiiform queens.Dedicated to Professor Dr. M. Lindauer on the occasion of his 70th birthday     

The influence of ants on host plant selection by Jalmenus evagoras,a myrmecophilous lycaenid butterfly

Summary (1) Females of the myrmecophilous lycaenid butterfly, Jalmenus evagoras are far more likely to lay eggs on plants that contain their attendant ants, Iridomyrmex sp. 25 than on plants without ants, although the clutch sizes of individual egg masses laid in either situation is the same. (2) Ovipositing females respond to the presence or absence of ants before they alight on a potential food plant. Once they have landed, they are equally likely to ley eggs whether or not they encounter ants. (3) Ovipositing females prefer to lay eggs on plants that contain ant tended homopterans than on plants that contain only a few foraging ants. The presence of ant tended homopterans can act as a strong stimulus to induce females to lay eggs on plant species that differ from their original host species. (4) Ant dependent oviposition behavior has been described or suggested in 46 species of lycaenid and one riodinid. In general, the more dependent a species is upon ants for either food or protection, the more likely it is to use ants as cues in oviposition. Prominent characteristics of lycaenids that have ant dependent oviposition are described and discussed. (5) Myrmecophilous lycaenids that may use ants as cues in oviposition feed on a significantly wider range of plants than non-myrmecophilous lycaenids. Possible reasons for this pattern and its ecological significance are discussed.     

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.     

Encounter rate and task allocation in harvester ants

As conditions change, social insect colonies adjust the numbers of workers engaged in various tasks, such as foraging and nest work. This process of task allocation operates without central control; individuals respond to simple, local cues. This study investigates one such cue, the pattern of an ant's interactions with other workers. We examined how an ant's tendency to perform midden work, carrying objects to and sorting the refuse pile of the colony, is related to the recent history of the ant's brief antennal contacts, in laboratory colonies of the red harvester ant, Pogonomyrmex barbatus. The probability that an ant performed midden work was related to its recent interactions in two ways. First, the time an ant spent performing midden work was positively correlated with the number of midden workers that ant had met while it was away from the midden. Second, ants engaged in a task other than midden work were more likely to begin to do midden work when their rate of encounter per minute with midden workers was high. Cues based on interaction rate may enable ants to respond to changes in worker numbers even though ants cannot count or assess total numbers engaged in a task. Received: 1 July 1998 / Accepted: 15 November 1998     

Homopteran chemical signatures reduce aggression of tending ants

Summary. Because generalist ants are aggressive towards foreign insects, the recognition of homopterans by tending ants is critical in ant/homopteran trophobiosis. Herein we report experimental evidence indicating that Argentine ant, Linepithema humile (Mayr) (Hymenoptera: Formicidae) learn to associate the production of honeydew with the chemical characteristics of homopteran cuticle, suppressing ant aggression and allowing the ants to tend homopterans. Although chemically-mediated associative learning is well understood in honeybee foraging, to our knowledge, it has not been reported before in ant/homopteran trophobiosis.     

Effects of an Alien Ant Invasion on Abundance,Behavior, and Reproductive Success of Endemic Island Birds

Abstract: Biological invaders can reconfigure ecological networks in communities, which changes community structure, composition, and ecosystem function. We investigated whether impacts caused by the introduced yellow crazy ant (Anoplolepis gracilipes), a pantropical invader rapidly expanding its range, extend to higher‐order consumers by comparing counts, behaviors, and nesting success of endemic forest birds in ant‐invaded and uninvaded rainforest on Christmas Island (Indian Ocean). Point counts and direct behavioral observations showed that ant invasion altered abundances and behaviors of the bird species we examined: the Island Thrush (Turdus poliocephalus erythropleurus), Emerald Dove (Chalcophaps indica natalis), and Christmas Island White‐eye (Zosterops natalis). The thrush, which frequents the forest floor, altered its foraging and reproductive behaviors in ant‐invaded forest, where nest‐site location changed, and nest success and juvenile counts were lower. Counts of the dove, which forages exclusively on the forest floor, were 9–14 times lower in ant‐invaded forest. In contrast, counts and foraging success of the white‐eye, a generalist feeder in the understory and canopy, were higher in ant‐invaded forest, where mutualism between the ant and honeydew‐secreting scale insects increased the abundance of scale‐insect prey. These complex outcomes involved the interplay of direct interference by ants and altered resource availability and habitat structure caused indirectly by ant invasion. Ecological meltdown, rapidly unleashed by ant invasion, extended to these endemic forest birds and may affect key ecosystem processes, including seed dispersal.     

Shape and efficiency of wood ant foraging networks

We measured the shape of the foraging trail networks of 11 colonies of the wood ant Formica aquilonia (Formica rufa group). We characterized these networks in terms of their degree of branching and the angles between branches, as well as in terms of their efficiency. The measured networks were compared with idealized model networks built to optimize one of two components of efficiency, total length (i.e., total amount of trail) and route factor (i.e., average distance between nest and foraging site). The analysis shows that the networks built by the ants obtain a compromise between the two modes of efficiency. These results are largely independent of the size of the network or colony size. The ants’ efficiency is comparable to that of networks built by humans but achieved without the benefit of centralized control.     

Behavioral response of a generalist predator to chemotactile cues of two taxonomically distinct prey species

Chemotactile cues unintentionally left by animals can play a major role in predator–prey interactions. Specialized predators can use them to find their prey, while prey individuals can assess predation risk. However, little is known to date about the importance of chemotactile cues for generalist predators such as ants. Here, we investigated the response of a generalized predatory ant, Formica polyctena, to cues of two taxonomically distinct prey: a spider (Pisaura mirabilis) and a cricket (Nemobius sylvestris). In analogy, we studied whether crickets and spiders showed antipredator behavior in response to ant cues. When confronted with cues of the two prey species, Formica polyctena workers showed increased residence time and reduced movement speed, which suggests success-motivated searching behavior and thus increased foraging effort. The ants’ response did not differ between cues of the two prey species, coinciding with similar aggression and consumption rates of dead prey. However, the cuticular hydrocarbons, which likely resemble part of the potential cues, differed strongly between the species, with only few methyl-branched alkanes in common. This suggests that ants respond to multiple compounds left by other organisms with prey-search behavior. The two prey species, in turn, showed no detectable antipredator behavior in response to ant cues. Our study shows that ants can detect and respond to chemotactile cues of taxonomically and ecologically distinct prey species, probably to raise their foraging success. Using such chemotactile cues for prey detection may drastically increase their foraging efficiency and thus contribute to the high ecological success of ants.     

Radio tagging reveals the roles of corpulence,experience and social information in ant decision making

Ant colonies are factories within fortresses (Oster and Wilson 1978). They run on resources foraged from an outside world fraught with danger. On what basis do individual ants decide to leave the safety of the nest? We investigated the relative roles of social information (returning nestmates), individual experience and physiology (lipid stores/corpulence) in predicting which ants leave the nest and when. We monitored Temnothorax albipennis workers individually using passive radio-frequency identification technology, a novel procedure as applied to ants. This method allowed the matching of individual corpulence measurements to activity patterns of large numbers of individuals over several days. Social information and physiology are both good predictors of when an ant leaves the nest. Positive feedback from social information causes bouts of activity at the colony level. When certain social information is removed from the system by preventing ants returning, physiology best predicts which ants leave the nest and when. Individual experience is strongly related to physiology. A small number of lean individuals are responsible for most external trips. An individual’s nutrient status could be a useful cue in division of labour, especially when public information from other ants is unavailable.     

Predatory behavior of Polybia sericea (Olivier), a tropical social wasp (Hymenoptera: Vespidae)

Summary Polybia sericea (Olivier) (Hymenoptera: Vespidae) foragers were trained to visit experimental foraging plots and tests were conducted to determine the role of visual, olfactory, and chemotactile cues in prey location. Foragers approached prey from downwind and hovered downwind of visual and olfactory stimuli. Olfactory cues were more likely to elicit landing than were visual cues. Biting of potential prey was most consistently elicited by a combination of visual, tactile, and chemotactile cues. Foragers encountering large prey carried a piece back to the nest and returned for prey remains. These returning foragers used visual cues to direct intensive aerial search; olfactory prey cues elicited landing.     

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.     

Blinkered teaching: tandem running by visually impaired ants

During tandem runs, one ant worker recruits another to an important resource. Here, we begin to investigate how dependent are tandem leaders and followers on visual cues by painting over their compound eyes to impair their vision. There are two ways in which Temnothorax albipennis might use vision during tandem running. First, the follower might track the movements of the leader by keeping it in sight. Our results suggest that the ants do not use vision in this way. For example, in all four classes of tandem run (those with either leader or follower, both, or neither of their participants with visual impairments) progress was most smooth at about 3 mm/s. This suggests that communication between leaders and followers during tandem runs is not based on vision and is purely tactile and pheromonal. Second, the leader and the follower might be using vision to navigate and our results support this possibility but also suggest that these ants have other methods of navigation. Ants with visual impairments were more likely to follow than to lead, but could occupy either role, even though they had many fully sighted nestmates. This might help to explain why the ants did not focus grooming on their most visually impaired nestmates. Wild-type tandem runs, with both participants fully sighted and presumably taking time to learn landmarks, were overall significantly slower, smoother, and a little less tortuous, than the other treatments. All four classes of tandem run significantly increased mean instantaneous speeds and mean absolute changes in instantaneous acceleration over their journeys. Moreover, tandems with sighted followers increased their speed with time more than the other treatments. In general, our findings suggest that eyesight is used for navigation during tandem running but that these ants also probably use other orientation systems during such recruitment and to learn how to get to new nest sites. Our results suggest that the ants’ methods of teaching and learning are very robust and flexible.     

Information transfer during recruitment in the ant<Emphasis Type="Italic"> Lasius niger</Emphasis> L. (Hymenoptera: Formicidae)

In this paper, we used the food-correlated search behavior observed in foraging ants returning to a previously rewarding site to study information transfer during recruitment in the ant Lasius niger. We hypothesized that, if information about the characteristics of the food is conveyed during recruitment, food-correlated search tactics should also be observed in recruited workers. Our results show that the characteristics of the trajectories of recruited workers are comparable to those of scout ants returning to a site or prior food find and depend more on the type (prey/sugar) than on the quality (sugar concentration) of the food discovered by the scouts. Independent of sugar concentration, workers recruited to a source of sugar search with a greater sinuosity than workers recruited to a prey. Experimental manipulation of the recruitment signals (chemical trail and contact between ants) shows that the trail pheromone laid down by recruiting ants does not play a role in the modification of trajectory sinuosity. This change appears to be most likely triggered by a direct perception of the residue of sugar smeared on the body of the recruiting workers coming back to the nest.Communicated by J. Heinze     

The origin of the chemical profiles of fungal symbionts and their significance for nestmate recognition in <Emphasis Type="Italic">Acromyrmex</Emphasis> leaf-cutting ants

Cuticular hydrocarbon profiles are essential for nestmate recognition in insect societies, and quantitative variation in these recognition cues is both environmentally and genetically determined. Environmental cues are normally derived from food or nest material, but an exceptional situation may exist in the fungus-growing ants where the symbiotic fungus garden may be an independent source of recognition compounds. To investigate this hypothesis, we quantified the chemical profiles of the fungal symbionts of 18 sympatric colonies of Acromyrmex echinatior and Acromyrmex octospinosus and evaluated the quantitative variation of the 47 compounds in a multivariate analysis. Colony-specific chemical profiles of fungal symbionts were highly distinct and significantly different between the two ant species. We also estimated the relative genetic distances between the fungal symbionts using amplified fragment length polymorphism (AFLP) and correlated these with the overall (Mahalanobis) chemical distances between the colony-specific profiles. Despite the standardized laboratory conditions, the correlations were generally weak, but a statistically significant portion of the total variation in chemical profiles could be explained by genetic differences between the fungal symbionts. However, there was no significant effect of ant species in partial analyses because genetic differences between symbionts tend to coincide with being reared by different ant species. However, compound groups differed significantly with amides, aldehydes, and methyl esters contributing to the correlations, but acetates, alkanes, and formates being unrelated to genetic variation among symbionts. We show experimentally that workers that are previously exposed to and fed with the fungal symbiont of another colony are met with less aggression when they are later introduced into that colony. It appears, therefore, that fungus gardens are an independent and significant source of chemical compounds, potentially contributing a richer and more abundant blend of recognition cues to the colony “gestalt” than the innate chemical profile of the ants alone. Freddie-Jeanne Richard and Michael Poulsen contributed equally to this work.     

Division of foraging labor in ants can mediate demands for food and safety

Solitary foragers can balance demands for food and safety by varying their relative use of foraging patches and their level of vigilance. Here, we investigate whether colonies of the ant, Formica perpilosa, can balance these demands by dividing labor among workers. We show that foragers collecting nectar in vegetation near their nest are smaller than are those collecting nectar at sites away from the nest. We then use performance tests to show that smaller workers are more likely to succumb to attack from conspecifics but feed on nectar more efficiently than larger workers, suggesting a size-related trade-off between risk susceptibility and harvesting ability. Because foragers that travel away from the nest are probably more likely to encounter ants from neighboring colonies, this trade-off could explain the benefits of dividing foraging labor among workers. In a laboratory experiment, we show that contact with aggressive workers results in an increase in the mean size of recruits to a foraging site: this increase was not the result of more large recruits, but rather because fewer smaller ants traveled to the site. These results suggest that workers particularly susceptible to risk avoid dangerous sites, and suggest that variation in worker size can allow colonies to exploit profitably both hazardous and resource-poor patches.Communicated by L. Sundström     

Predatory behavior and chemical communication in two Metapone species (Hymenoptera:Formicidae)

Summary. Colonies of two species of Metapone (M. madagascarica, M. new species.) were collected in Madagascar and established in laboratory nests. It could be demonstrated that both species are specialist predators of termites (Cryptotermes kirbyi). During hunting the ants sting the termites and thereby paralyze and preserve the prey alive. In this way prey can be stored in the ant nest for extended periods. During foraging and colony emigrations the ants lay chemical trails with poison gland secretions. Among the seven compounds identified in the venom only methyl pyrrole-2-carboxylate elicits trail following behavior in both Metapone species. Received 11 February 2002, accepted 23 February 2002.     

Induced responses to herbivory in the Neotropical ant-plant association between Azteca ants and Cecropia trees: response of ants to potential inducing cues

Plant defense against herbivores often involves constitutive and inducible mechanisms of resistance. Obligate ant-plants, which provide food and housing for ants, are thought to primarily rely on ants for defense against herbivores. This form of plant defense has largely been viewed as static. We have been investigating the dynamic nature of Azteca ants as an inducible defense of Cecropia trees. Ants rapidly recruit to and patrol sites of foliar damage. We propose that Azteca ants can be viewed as an inducible defense for Cecropia trees because of their sensitivity to cues associated with herbivory, their rapid and aggressive recruiting ability, and their reclaimable and redeployable nature as a plant defense. In this study, we examine ant behavior following plant damage, and the potential cues that indude ant recruitment. We found that ants present on leaves when the plant is damaged leave the damaged leaf and recruit other ants to it, presumably by laying recruitment trails. Volatile leaf cues associated with herbivory were important in eliciting an induced response in two experiments. However, we found that cues associated with a congeneric plant elicited a much stronger ant response than conspecific cues. Although the type of leaf damage (gaping wounds versus leaf edge wounds) did not affect the level of ant recruitment, the extent of damage did. Leaves with one hole punched showed a 50% increase in ants, while leaves with five holes punched in them elicited a 100% increase in ant numbers. In sum, it appears that multiple plant-related cues associated with herbivory are involved in induction of ant recruitment in the Cecropia-Azteca system. We discuss the generality of ant responses to herbivory in obligate ant-plant systems, and in facultative ant-plant associations, which may be more common. Received: 23 March 1998 / Accepted after revision: 5 July 1998