Behavioral mechanisms of collective nest-site choice by the ant Temnothorax curvispinosus

This paper examines the individual behavior underlying collective choice among nest sites by the ant Temnothorax (formerly Leptothorax) curvispinosus. Colonies can actively compare options, rejecting a mediocre site when it is paired with a good one, but accepting the same mediocre design if it is instead paired with a worse site. This ability emerges from the behavior of an active minority of workers who organize emigrations. When one of these finds a promising site, she recruits nest mates to it, but only after a delay that varies inversely with site quality. Ants first recruit fellow active ants via slow tandem runs, but eventually switch to speedier transports of the colony’s passive majority. Later transports grow faster still, as ants improve their speed with experience. An ant’s choice of recruitment type is governed by a quorum rule, such that her likelihood of starting to transport increases with the population of the new site. The size of the quorum depends on experience, with ants demanding a larger population to launch immediately into transport than they do to switch to transport after first leading a few tandem runs. Perception of quorum attainment requires direct contact between ants. The ants’ behavior qualitatively matches that of T. albipennis, where models have shown that decentralized choice of the best site depends on quality-dependent recruitment delays, amplified by a quorum rule for initiating transport. Parameter estimates for an agent-based model show significant quantitative differences between the species, and suggest that T. albipennis may place relatively greater emphasis on emigration speed. Received 11 February 2005; revised 10 May 2005; accepted 20 May 2005.     

Quorum sensing by encounter rates in the ant Temnothorax albipennis

Emigrating colonies of the ant Temnothorax (formerly Leptothorax)albipennis can choose the best of several nest sites, even whenthe active ants organizing the move do not compare sites. Thiscollective ability depends on a quorum rule used by ants assessinga candidate site. Only when the site's population has surpasseda threshold do they switch from slow recruitment of fellow activeants by tandem runs to rapid transport of the majority of thecolony. Here, I show that ants perceive the achievement of aquorum through their rate of direct encounters with nest matesat the site. When ants in a crowded site were prevented fromtactile contact with nest mates, they recruited by tandem runs,as though to an empty nest. Furthermore, when the encounterrate was raised independent of population, by reducing the sizeof the candidate nest, ants started to transport at a significantlylower population. The switch occurred at the same encounterrate regardless of nest size, whether the rate was measuredas the mean over the entire visit or as the inverse of the latencyuntil the first encounter. Because encounter rate reflects thedensity of nest mates and thus varies with nest size as wellas population, the ants' collective decision-making algorithmmay be robust to the exact population at which the switch totransport occurs. Ants cease monitoring quorum presence afterswitching to transport, coincident with an abrupt shorteningof visit duration by approximately 2 min, which may be interpretedas the time required for quorum detection.     

Speed versus accuracy in collective decision making

We demonstrate a speed versus accuracy trade-off in collective decision making. House-hunting ant colonies choose a new nest more quickly in harsh conditions than in benign ones and are less discriminating. The errors that occur in a harsh environment are errors of judgement not errors of omission because the colonies have discovered all of the alternative nests before they initiate an emigration. Leptothorax albipennis ants use quorum sensing in their house hunting. They only accept a nest, and begin rapidly recruiting members of their colony, when they find within it a sufficient number of their nest-mates. Here we show that these ants can lower their quorum thresholds between benign and harsh conditions to adjust their speed-accuracy trade-off. Indeed, in harsh conditions these ants rely much more on individual decision making than collective decision making. Our findings show that these ants actively choose to take their time over judgements and employ collective decision making in benign conditions when accuracy is more important than speed.     

Colony size affects collective decision-making in the ant Temnothorax albipennis

Social insects are well-known for their ability to achieve robust collective behaviours even when individuals have limited information. It is often assumed that such behaviours rely on very large group sizes, but many insect colonies start out with only a few workers. Here we investigate the influence of colony size on collective decision-making in the house-hunting of the ant Temnothorax albipennis. In experiments where colony size was manipulated by splitting colonies, we show that worker number has an influence on the speed with which colonies discover new nest sites, but not on the time needed to make a decision (achieve a quorum threshold) or total emigration time. This occurred because split colonies adopted a lower quorum threshold, in fact they adopted the same threshold in proportion to their size as full-size colonies. This indicates that ants may be measuring relative quorum, i.e. population in the new nest relative to that of the old nest, rather than the absolute number. Experimentally reduced colonies also seemed to gain more from experience through repeated emigrations, as they could then reduce nest discovery times to those of larger colonies. In colonies of different sizes collected from the field, total emigration time was also not correlated with colony size. However, quorum threshold was not correlated with colony size, meaning that individuals in larger colonies adopted relatively lower quorum thresholds. Since this is a different result to that from size-manipulated colonies, it strongly suggests that the differences between natural small and large colonies were not caused by worker number alone. Individual ants may have adjusted their behaviour to their colony’s size, or other factors may correlate with colony size in the field. Our study thus shows the importance of experimentally manipulating colony size if the effect of worker number on the emergence of collective behaviour is to be studied. Received 13 December 2005; revised 9 May 2006; accepted 15 May 2006.     

Tandem Recruitment and Foraging in the Ponerine Ant <Emphasis Type="Italic">Pachycondyla harpax</Emphasis> (Fabricius)

Tandem running is a common recruitment strategy in ant species with small colony sizes. During a tandem run, an informed leader guides a usually naïve nestmate to a food source or a nest site. Some species perform tandem runs only during house hunting, suggesting that tandem running does not always improve foraging success in species known to use tandem running as a recruitment strategy, but more natural history information on tandem running under natural conditions is needed to better understand the adaptive significance of tandem recruitment in foraging. Studying wild colonies in Brazil, we for the first time describe tandem running in the ponerine ant Pachycondyla harpax (Fabricius). We asked if foragers perform tandem runs to carbohydrate- (honey) and protein-rich (cheese) food items. Furthermore, we tested whether the speed and success rate of tandem runs depend on the foraging distance. Foragers performed tandem runs to both carbohydrate food sources and protein-rich food items that exceed a certain size. The probability to perform a tandem run and the travelling speed increase with increasing foraging distances, which could help colonies monopolize more distant food sources in a competitive environment. Guiding a recruit to a food source is costly for leaders as ants are ~66% faster when travelling alone. If tandem runs break up (~23% of all tandem runs), followers do not usually discover the food source on their own but return to the nest. Our results show that tandem running to food sources is common in P. harpax, but that foragers modify their behaviour according to the type of food and its distance from the nest. Competition with other ants was intense and we discuss how tandem running in P. harpax might help colonies to build-up a critical number of ants at large food items that can then defend the food source against competitors.     

How collective comparisons emerge without individual comparisons of the options

Collective decisions in animal groups emerge from the actions of individuals who are unlikely to have global information. Comparative assessment of options can be valuable in decision-making. Ant colonies are excellent collective decision-makers, for example when selecting a new nest-site. Here, we test the dependency of this cooperative process on comparisons conducted by individual ants. We presented ant colonies with a choice between new nests: one good and one poor. Using individually radio-tagged ants and an automated system of doors, we manipulated individual-level access to information: ants visiting the good nest were barred from visiting the poor one and vice versa. Thus, no ant could individually compare the available options. Despite this, colonies still emigrated quickly and accurately when comparisons were prevented. Individual-level rules facilitated this behavioural robustness: ants allowed to experience only the poor nest subsequently searched more. Intriguingly, some ants appeared particularly discriminating across emigrations under both treatments, suggesting they had stable, high nest acceptance thresholds. Overall, our results show how a colony of ants, as a cognitive entity, can compare two options that are not both accessible by any individual ant. Our findings illustrate a collective decision process that is robust to differences in individual access to information.     

Improving Decision Speed,Accuracy and Group Cohesion through Early Information Gathering in House-Hunting Ants

Background

Successful collective decision-making depends on groups of animals being able to make accurate choices while maintaining group cohesion. However, increasing accuracy and/or cohesion usually decreases decision speed and vice-versa. Such trade-offs are widespread in animal decision-making and result in various decision-making strategies that emphasize either speed or accuracy, depending on the context. Speed-accuracy trade-offs have been the object of many theoretical investigations, but these studies did not consider the possible effects of previous experience and/or knowledge of individuals on such trade-offs. In this study, we investigated how previous knowledge of their environment may affect emigration speed, nest choice and colony cohesion in emigrations of the house-hunting ant Temnothorax albipennis, a collective decision-making process subject to a classical speed-accuracy trade-off.

Methodology/Principal Findings

Colonies allowed to explore a high quality nest site for one week before they were forced to emigrate found that nest and accepted it faster than emigrating naïve colonies. This resulted in increased speed in single choice emigrations and higher colony cohesion in binary choice emigrations. Additionally, colonies allowed to explore both high and low quality nest sites for one week prior to emigration remained more cohesive, made more accurate decisions and emigrated faster than emigrating naïve colonies.

Conclusions/Significance

These results show that colonies gather and store information about available nest sites while their nest is still intact, and later retrieve and use this information when they need to emigrate. This improves colony performance. Early gathering of information for later use is therefore an effective strategy allowing T. albipennis colonies to improve simultaneously all aspects of the decision-making process – i.e. speed, accuracy and cohesion – and partly circumvent the speed-accuracy trade-off classically observed during emigrations. These findings should be taken into account in future studies on speed-accuracy trade-offs.     

Ants move to improve: colonies of Leptothorax albipennis emigrate whenever they find a superior nest site

A high-quality home can be a major factor determining fitness. However, when house hunting becomes necessary, animals might often face a speed-versus-accuracy trade-off and therefore be unable to survey their environment extensively for the optimum site. We found that the ant Leptothorax albipennis was able to correct errors made in such a hurried decision by continuing to survey even after a colony had settled in a nest site. Colonies moved from intact undisturbed nests to another nest site whenever the new nest site presented a sufficient improvement in nest quality. Thus, scout ants must be able to judge and compare the quality of the new, empty nest site with the one currently inhabited by the colony. Emigrations from intact nests were initiated by high numbers of ants recruited by tandem runs. This evidence may explain how a small number of scouts can motivate an entire colony to move when there is no immediate need to do so. Compared with their behaviour in emigrations from destroyed nests, the ants favoured even more strongly accuracy over speed, because they waited for a larger number of scouts to agree on one site before starting the emigration. They could do this without increased risk because the rest of the colony remained safely in the old nest.     

A simple threshold rule is sufficient to explain sophisticated collective decision-making

Decision-making animals can use slow-but-accurate strategies, such as making multiple comparisons, or opt for simpler, faster strategies to find a 'good enough' option. Social animals make collective decisions about many group behaviours including foraging and migration. The key to the collective choice lies with individual behaviour. We present a case study of a collective decision-making process (house-hunting ants, Temnothorax albipennis), in which a previously proposed decision strategy involved both quality-dependent hesitancy and direct comparisons of nests by scouts. An alternative possible decision strategy is that scouting ants use a very simple quality-dependent threshold rule to decide whether to recruit nest-mates to a new site or search for alternatives. We use analytical and simulation modelling to demonstrate that this simple rule is sufficient to explain empirical patterns from three studies of collective decision-making in ants, and can account parsimoniously for apparent comparison by individuals and apparent hesitancy (recruitment latency) effects, when available nests differ strongly in quality. This highlights the need to carefully design experiments to detect individual comparison. We present empirical data strongly suggesting that best-of-n comparison is not used by individual ants, although individual sequential comparisons are not ruled out. However, by using a simple threshold rule, decision-making groups are able to effectively compare options, without relying on any form of direct comparison of alternatives by individuals. This parsimonious mechanism could promote collective rationality in group decision-making.     

Information flow,opinion polling and collective intelligence in house-hunting social insects

The sharing and collective processing of information by certain insect societies is one of the reasons that they warrant the superlative epithet ''super-organisms'' (Franks 1989, Am. Sci. 77, 138-145). We describe a detailed experimental and mathematical analysis of information exchange and decision-making in, arguably, the most difficult collective choices that social insects face: namely, house hunting by complete societies. The key issue is how can a complete colony select the single best nest-site among several alternatives? Individual scouts respond to the diverse information they have personally obtained about the quality of a potential nest-site by producing a recruitment signal. The colony then deliberates over (i.e. integrates) different incoming recruitment signals associated with different potential nest-sites to achieve a well-informed collective decision. We compare this process in honeybees and in the ant Leptothorax albipennis. Notwithstanding many differences - for example, honeybee colonies have 100 times more individuals than L. albipennis colonies - there are certain similarities in the fundamental algorithms these societies appear to employ when they are house hunting. Scout honeybees use the full power of the waggle dance to inform their nest-mates about the distance and direction of a potential nest-site (and they indicate the quality of a nest-site indirectly through the vigour of their dance), and yet individual bees perhaps only rarely make direct comparisons of such sites. By contrast, scouts from L. albipennis colonies often compare nest-sites, but they cannot directly inform one another of their estimation of the quality of a potential site. Instead, they discriminate between sites by initiating recruitment sooner to better ones. Nevertheless, both species do make use of forms of opinion polling. For example, scout bees that have formerly danced for a certain site cease such advertising and monitor the dances of others at random. That is, they act without prejudice. They neither favour nor disdain dancers that advocate the site they had formerly advertised or the alternatives. Thus, in general the bees are less well informed than they would be if they systematically monitored dances for alternative sites rather than spending their time reprocessing information they already have. However, as a result of their lack of prejudice, less time overall will be wasted in endless debate among stubborn and potentially biased bees. Among the ants, the opinions of nest-mates are also pooled effectively when scouts use a threshold population of their nest-mates present in a new nest-site as a cue to switch to more rapid recruitment. Furthermore, the ants'' reluctance to begin recruiting to poor nest-sites means that more time is available for the discovery and direct comparison of alternatives. Likewise, the retirement of honeybee scouts from dancing for a given site allows more time for other scouts to find potentially better sites. Thus, both the ants and the bees have time-lags built into their decision-making systems that should facilitate a compromise between thorough surveys for good nest-sites and relatively rapid decisions. We have also been able to show that classical mathematical models can illuminate the processes by which colonies are able to achieve decisions that are relatively swift and very well informed.     

Tandem running by foraging Pachycondyla striata workers in field conditions vary in response to food type,food distance,and environmental conditions

Ants show collective and individual behavioral flexibility in their response to immediate context, choosing for example between different foraging strategies. In Pachycondyla striata, workers can forage solitarily or recruit and guide nestmates to larger food sources through tandem running. Although considered more ancestral and less efficient than pheromone trail-laying, this strategy is common especially in species with small colony size. What is not known is how the decision to recruit or follow varies according to the immediate context. That is, how fine adjustments in information transfer affect immediate foraging decisions at the colony level. Here, we studied individually marked workers and evaluated their foraging decisions when food items varied in nature (protein versus carbohydrate), size, and distance from the nest at different temperatures and humidity levels. Our results show that tandem run leaders and potential followers adjust their behavior according to a combination of external factors. While 84.2% of trips were solitary, most ants (81%) performed at least 1 tandem run. However, tandem runs were more frequent for nearby resources and at higher relative humidity. Interestingly, when food items were located far away, tandem runs were more successful when heading to protein sources (75%) compared with carbohydrate sources (42%). Our results suggest that the social information transfer between leaders and followers conveys more information than previously thought, and also relies on their experience and motivation.     

Coordination of Raiding and Emigration in the Ponerine Army Ant Leptogenys distinguenda (Hymenoptera: Formicidae: Ponerinae): A Signal Analysis

Several glandular sources of trail pheromones have been discovered in army ants in general. Nevertheless, at present the understanding of the highly coordinated behavior of these ants is far from complete. The importance of trail pheromone communication for the coordination of raids and emigrations in the ponerine army ant Leptogenys distinguenda was examined, and its ecological function is discussed. The secretions of at least two glands organize the swarming activities of L. distinguenda. The pygidial gland is the source of an orientation pheromone holding the group of raiding workers together. The same pheromone guides emigrations to new nest sites. In addition, the poison sac contains two further components: one with a weak orientation effect and another which produces strong, but short-term attraction and excitement. The latter component is important in prey recruitment and characterizes raid trails. This highly volatile recruitment pheromone allows the extreme swarm dynamic characteristic of this species. Emigration trails lack the poison gland secretion. Due to their different chemical compositions, the ants are thus able to distinguish between raid and emigration trails. Nest emigration is not induced chemically, but mechanically, by the jerking movements of stimulating workers.     

Do ants make direct comparisons?

Many individual decisions are informed by direct comparison of the alternatives. In collective decisions, however, only certain group members may have the opportunity to compare options. Emigrating ant colonies (Temnothorax albipennis) show sophisticated nest-site choice, selecting superior sites even when they are nine times further away than the alternative. How do they do this? We used radio-frequency identification-tagged ants to monitor individual behaviour. Here we show for the first time that switching between nests during the decision process can influence nest choice without requiring direct comparison of nests. Ants finding the poor nest were likely to switch and find the good nest, whereas ants finding the good nest were more likely to stay committed to that nest. When ants switched quickly between the two nests, colonies chose the good nest. Switching by ants that had the opportunity to compare nests had little effect on nest choice. We suggest a new mechanism of collective nest choice: individuals respond to nest quality by the decision either to commit or to seek alternatives. Previously proposed mechanisms, recruitment latency and nest comparison, can be explained as side effects of this simple rule. Colony-level comparison and choice can emerge, without direct comparison by individuals.     

Recruitment and Other Communication Behavior in the Ponerine Ant Ectatomma ruidum

The ponerine ant Ectatomma ruidum, though previously reported to possess only rudimentary recruitment ability, was found to lay chemical trails for mass recruitment to rich or difficult food sources. The pheromone originates from the Dufour's gland, a new source of trail pheromones in the primitive ant subfamily Ponerinae. During nest emigrations, E. ruidum practices stereotyped social carrying in the myrmicine mode. The discovery of this form of social carrying and of a recruitment pheromone in the Dufour's gland secretions support the hypothesis that the subfamily Myrmicinae is derived from an ectatommine ancestor. Other communication behaviors exhibited by E. ruidum include exchange of liquid food carried between the mandibles, chemical alarm communication, nest entrance marking, and an additional social carrying posture previously unknown in ants.     

Coupled adult-brood transport augments relocation in the Indian queenless ant Diacamma indicum

Colony relocation is an important aspect in the lives of social insects. In ants, the process of relocation is further complicated as brood, in addition to adults, have to be transported to the new nest. Here, we have investigated brood transport in the Indian ponerine ant Diacamma indicum, which uses tandem running—a primitive mode of recruitment—for the entire colony to relocate. We have found that there were no brood transport specialists and most of the brood was transported in the mandibles of followers that were being tandem run. Therefore, in a single tandem run, one adult and one brood item was effectively transported by tandem leaders augmenting the relocation process.     

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.     

Nest architecture and traffic flow: large potential effects from small structural features

1. Research on human pedestrian dynamics predicts that seemingly small architectural features of the surroundings can have large effects on the behaviour of crowds and the flow of pedestrian traffic, particularly when a crowd is panicked. This theoretical framework might usefully be applied to the study of collective movement within subterranean nests of social insects. 2. We examined the rate of egress from artificial nests by alarmed Linepithema humile ants. In accord with model predictions, but counter to intuition, we found that a partially obstructed exit enhanced the average rate of escape from the nest. 3. The study of traffic flows in subterranean nests is almost non‐existent, but it would be worth studying the effect of nest design elements on collective movements, given the great variety of nest forms among ants and termites.     

Anti-predator behaviour in the ant Pheidole desertorum: The importance of multiple nests

The relationship between emigrations and anti-predator behaviour in the ant Pheidole desertorum was investigated in the field. Frequent emigrations in P. desertorum result in the colony having multiple nests, of which only one is occupied at a time. A field procedure demonstrated that the ants quickly find and enter the unoccupied nests after a nest evacuation caused by the army ant Neivamyrmex nigrescens. When colonies were denied access to their unoccupied nests after an evacuation, survival of the brood and alates was significantly lower than in colonies allowed access to all their nests. These results suggest that emigrations in P. desertorum are part of a defence strategy against army ants.     

The ‘truck‐driver’ effect in leaf‐cutting ants: how individual load influences the walking speed of nest‐mates

The foraging behaviour of social insects is highly flexible because it depends on the interplay between individual and collective decisions. In ants that use foraging trails, high ant flow may entail traffic problems if different workers vary widely in their walking speed. Slow ants carrying extra‐large loads in the leaf‐cutting ant Atta cephalotes L. (Hymenoptera: Formicidae) are characterized as ‘highly‐laden’ ants, and their effect on delaying other laden ants is analyzed. Highly‐laden ants carry loads that are 100% larger and show a 50% greater load‐carrying capacity (i.e. load size/body size) than ‘ordinary‐laden’ ants. Field manipulations reveal that these slow ants carrying extra‐large loads can reduce the walking speed of the laden ants behind them by up to 50%. Moreover, the percentage of highly‐laden ants decreases at high ant flow. Because the delaying effect of highly‐laden ants on nest‐mates is enhanced at high traffic levels, these results suggest that load size might be adjusted to reduce the negative effect on the rate of foraging input to the colony. Several causes have been proposed to explain why leaf‐cutting ants cut and carry leaf fragments of sizes below their individual capacities. The avoidance of delay in laden nest‐mates is suggested as another novel factor related to traffic flow that also might affect load size selection The results of the presennt study illustrate how leaf‐cutting ants are able to reduce their individual carrying performance to maximize the overall colony performance.