Rules of supply and demand regulate recruitment to food in an ant society

The process by which ant scouts move a group of nestmates toward a newly discovered food site is called recruitment. In this paper, I report on the interactions between scouts and nestmates that result in a graded recruitment response to graded food quality in the fire ant, Solenopsis invicta. Twelve experimental groups composed of 100 fire ant workers and 50 fire ant larvae were established (three experimental groups per colony × four stock colonies). Each experimental group was placed in a shallow, artificial nest with a glass cover. After a 48-h period of food deprivation, experimental groups were exposed to one of three concentrations of sugar water. Behavioral interactions between scouts and nestmates in each group were videotaped at 10× magnification for 20 min. Detailed behavioral data on a total of 120 scouts (10 scouts per experimental group) and ~1,000 nestmates (~90 nestmates per experimental group) were transcribed from the videotapes using standard play and frame-by-frame techniques. Throughout the recruitment process, scouts employed six discrete behaviors to inform nestmates of the location and quality of a food site. Scouts laid incoming trails, waggled their heads, increased walking tempo, stroked nestmates with their antennae, advertised with a brief food display, and led groups of nestmates to the food site by laying outgoing trails. In turn, nestmates assessed the food sample with antennae, then responded to or resisted recruitment based on the quality of food advertised, their employment status and their level of hunger. In summary, recruitment was an emergent property based on competent supply and demand decisions made face-to-face inside the nest rather than on the trail or at the food site.Communicated by J. Heinze     

Decision making in ant foragers (<Emphasis Type="Italic">Lasius niger</Emphasis>) facing conflicting private and social information

Foragers of many ant species use pheromone trails to guide nestmates to food sources. During foraging, individual workers can also learn the route to a food source. Foragers of the mass-recruiting ant Lasius niger use both pheromone trails and memory to locate a food source. As a result, an experienced forager can have a conflict between social information (trail pheromones) and private information (route memory) at trail bifurcations. We tested decision making in L. niger foragers facing such an informational conflict in situations where both the strength of the pheromone trail and the number of previous visits to the food source varied. Foragers quickly learned the branch at a T bifurcation that leads to a food source, with 74.6% choosing correctly after one previous visit and 95.3% after three visits. Pheromone trails had a weaker effect on choice behaviour of naïve ants, with only 61.6% and 70.2% choosing the branch that had been marked by one or 20 foragers versus an unmarked branch. When there was a conflict between private and social information, memory overrides pheromone after just one previous visit to a food source. Most ants, 82–100%, chose the branch where they had collected food during previous foraging trips, with the proportion depending on the number of previous trips (1 v. 3) but not on the strength of the pheromone trail (1 v. 20). In addition, the presence of a pheromone trail at one branch in a bifurcation had no effect on the time it took an experienced ant to choose the correct branch (the branch without pheromone). These results suggest that private information (navigational memory) dominates over social information (chemical tail) in orientation decisions during foraging activities in experienced L. niger foragers.     

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     

Chemical trail communication in the amblyoponine species Mystrium rogeri Forel (Hymenoptera, Formicidae, Ponerinae)

Summary. Workers of the amblyoponine species Mystrium rogeri employ trail communication during recruitment to food sources and new nest sites. The trail pheromone originates from a hitherto unknown sternal gland located in the 7th abdominal sternite. The recruiting ant deposits the gland secretions by a special gaster-dragging behavior. The recruitment behavior can be complemented by a rapid vertical body shaking performed by some recruiting ants inside the nest. M. rogeri workers possess a large pygidial gland, the secretion of which elicits a repellent response in other ant species. Received 25 May 1998; accepted 15 June 1998.     

Graded recruitment in a ponerine ant

Summary (1) The giant tropical ant, Paraponera clavata, exhibits graded recruitment responses, depending on the type, quantity, and quality of a food source. More ants are initially recruited to a large prey or scavenge item than to a large quantity of sugar water. (2) Individual ants encountering prey items gauge the size and/or unwieldiness of the item, regardless of the weight, when determining whether to recruit. (3) The trail pheromone of this species is often used as an orientation device by individual ants, independent of recruitment of nestmates. (4) It is proposed that the foraging behavior of P. clavata represents one of the evolutionary transitions from the independent foraging activities of the primitive ants to the highly coordinated cooperative foraging activities of many higher ants.     

Orientation and recruitment behavior in the ponerine ant Pachycondyla tesserinoda (Emery): laying of individual-specific trails during tandem running

Summary Tandem leaders of Pachycondyla tesserinoda mark their way during nest moving. While scouting and foraging for food in an unknown area, chemical orientation is used as well. The origin of the trail substance could not be identified. Secretions of gastral exocrine glands did not induce trail following behavior. Nor do these secretions elicit tandem-following reactions; rather the latter is released by the general body surface odor. The trail substance and the substance used for marking the nest entrance are colony-specific. Moreover, individual tandem leaders recognize and show a preference for their own trails. This extraordinary effect is independent of the age of the trail. P. tesserinoda workers search individually for food and new nest sites. Targets which are important for the colony are directly shown to nestmates by tandem running. Due to this type of foraging and scouting individual-specific trails may be advantageous for this ant.     

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.     

Individual and collective foraging decisions: a field study of worker recruitment in the gypsy ant <Emphasis Type="Italic">Aphaenogaster senilis</Emphasis>

In social insects, the decision to exploit a food source is made both at the individual (e.g., a worker collecting a food item) and colony level (e.g., several workers communicating the existence of a food patch). In group recruitment, the recruiter lays a temporary chemical trail while returning from the food source to the nest and returns to the food guiding a small group of nestmates. We studied how food characteristics influence the decision-making process of workers changing from individual retrieving to group recruitment in the gypsy ant Aphaenogaster senilis. We offered field colonies three types of prey: crickets (cooperatively transportable), shrimps (non-transportable), and different quantities of sesame seeds (individually transportable). Colonies used group recruitment to collect crickets and shrimps, as well as seeds when they were available in large piles, while small seed piles rarely led to recruitment. Foragers were able to “measure” food characteristics (quality, quantity, transportability), deciding whether or not to recruit, accordingly. Social integration of individual information about food emerged as a colony decision to initiate or to continue recruitment when the food patch was rich. In addition, group recruitment allowed a fast colony response over a wide thermal range (up to 45°C ground temperature). Therefore, by combining both advantages of social foraging (group recruitment) and thermal tolerance, A. senilis accurately exploited different types of food sources which procured an advantage against mass-recruiting and behaviorally dominant species such as Tapinoma nigerrimum and Lasius niger.     

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     

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.     

Both evaluation of resource quality and speed of recruited leaf-cutting ants (Acromyrmex lundi) depend on their motivational state

Summary During recruitment, running velocity of both outbound and laden workers of the leaf-cutting ant Acromyrmex lundi depended on the information about resource quality they received from the first successful recruiter. In independent assays, single scout ants were allowed to collect sugar solutions of different concentrations and to recruit nestmates. Recruited workers were presented with standardized paper discs rather than the sugar solution given to the original recruiting ant. Outbound recruited workers were observed to run faster the more concentrated the solution found by the recruiter. Speed of disc-laden workers also depended on the concentration of the solution found by the recruiter, i.e. on the information about food quality they received, since they had no actual contact with the sugar solution. Disc-laden workers ran, as intuitively expected, slower than outbound workers. The reduction in speed, however, could not be attributed to the effects of the load itself, because workers collecting discs of the same weight, but with added sugar, ran as rapidly as outbound, unladen workers. Workers collecting standardized sugared discs reinforced the chemical trail on their way to the nest. The percentage of trail-layers was higher when workers were recruited to 10% than to 1% sugar solution, even though they collected the same kind of discs at the source. Their evaluation of resource quality, therefore, depended on their motivational state, which was modulated by the information they received during recruitment. Using previously published data on energetics of locomotion in leaf-cutting ants, travel costs of A. lundi workers recruited to sugar solutions of different concentration could be estimated. For workers recruited to the more concentrated solution, both speed and oxygen consumption rate increased by a roughly similar factor. Therefore, although workers ran faster to the high-quality resource, their actual energy investment per trip remained similar to that made by workers recruited to the low-quality resource. It is suggested that the more motivated workers reduced travel time without increasing energy costs during the trip. The adaptive value of these responses seems to be related to a rapid transmission of information about a newly discovered food source.     

Synergy between pheromone trails and quorum thresholds underlies consensus decisions in the ant Myrmecina nipponica

Coordination of group actions in social organisms is often a self-organised process lacking central control. These collective behaviours are driven by mechanisms of positive feedback generated through information exchange. Understanding how different methods of communication generate positive feedback is an essential step in comprehending the functional mechanisms underlying complex systems. The Japanese small-colony ant, Myrmecina nipponica uses both pheromone trails and an apparent quorum response during consensus decisions over a new home. Both of these mechanisms have been shown to generate positive feedback and are effective means of selecting among mutually exclusive courses of action. In this study, I investigate how pheromone trails and quorum thresholds contribute to consensus decisions during house-hunting in this species through experimental manipulations of pheromone trails, colony size and environmental context. Results demonstrate that (1) providing colonies with pre-established pheromone trails increased the number of ants finding the new site and led to higher quorum thresholds and more rapid relocations, (2) experimentally halving colony size resulted in a proportional decrease in quorum thresholds and (3) colonies relocating long distances had higher quorums than those relocating short distances. Taken together, these data indicate that pheromone trails are important for recruitment and navigation during nest site selection, but that decision making is contingent on a quorum response. Such synergy between mechanisms of positive feedback may be a common means of optimising collective behaviours.     

Identification of trail pheromone compounds from the labial glands of the stingless bee Geotrigona mombuca

Foragers of several species of stingless bees deposit pheromone spots in the vegetation to guide recruited nestmates to a rich food source. Recent studies have shown that Trigona and Scaptotrigona workers secrete these pheromones from their labial glands. An earlier report stated that species within the genus Geotrigona use citral from their mandibular glands for scent marking. Since convincing experimental proof for this conjecture is lacking, we studied the glandular origin of the trail pheromone of Geotrigona mombuca. In field bioassays, newly recruited bees were diverted by artificial scent trails that branched off from the natural scent trail deposited by their nestmates only when they were baited with extracts from the foragers’ labial glands. Compounds extracted from the mandibular glands, however, did not release trail following behavior. This demonstrates that the trail pheromone of G. mombuca is produced in the labial glands, as in Trigona and Scaptotrigona. Furthermore, in chemical analyses citral was identified exclusively in the foragers’ mandibular glands, which disproves its supposed role as a trail pheromone. The labial glands contained a series of terpene- and wax type esters, with farnesyl butanoate as major constituent. We, therefore, postulate that the trail pheromone of G. mombuca is composed of a blend of esters.     

Polyethism and the importance of context in the alarm reaction of the grass-cutting ant, Atta capiguara

Leaf-cutting ants exhibit an aggressive alarm response. Yet in most alarm reactions, not all of the ants encountering a disturbance will respond. This variability in behaviour was investigated using field colonies of Atta capiguara, a grass-cutting species. Crushed ant heads were applied near foraging trails to stimulate alarm reactions. We found that minor workers were disproportionately likely to respond. Only 34.7DŽ.8% of ants travelling along foraging trails were minor workers, but 82.1Lj.1% of ants that responded were minors. Workers transporting grass did not respond at all. The alarm response was strongest at the position and time where minors were most abundant. Ants were more likely to respond when they were travelling along trails with low rather than high traffic. Minor workers followed a meandering route along the trail, compared with the direct route taken by foragers. We argue that an important function of minor workers on foraging trails is to patrol the trail area for threats, and that they then play the key role in the alarm reaction.     

A trail pheromone component of the African stink ant, Pachycondyla (Paltothyreus) tarsata Fabricius (Hymenoptera: Formicidae: Ponerinae)

Summary. The African stink ants (Pachycondyla tarsata) lay recruitment trails with secretions from sternal glands. The glandular secretions consist of 10 compounds, 9 of which have been chemically identified. One of the substances, 9-heptadecanone, elicits trail following behavior in P. tarsata workers that have before been stimulated by a sucessful scout ant. Received 7 August 1998; accepted 24 November 1998.     

Quorum sensing, recruitment, and collective decision-making during colony emigration by the ant Leptothorax albipennis

When its nest is damaged, a colony of the ant Leptothorax albipennis skillfully emigrates to the best available new site. We investigated how this ability emerges from the behaviors used by ants to recruit nestmates to potential homes. We found that, in a given emigration, only one-third of the colony's workers ever recruit. At first, they summon fellow recruiters via tandem runs, in which a single follower is physically led all the way to the new site. They later switch to recruiting the passive majority of the colony via transports, in which nestmates are simply carried to the site. After this switch, tandem runs continue sporadically but now run in the opposite direction, leading recruiters back to the old nest. Recruitment accelerates with the start of transport, which proceeds at a rate 3 times greater than that of tandem runs. The recruitment switch is triggered by population increase at the new site, such that ants lead tandem runs when the site is relatively empty, but change to transport once a quorum of nestmates is present. A model shows that the quorum requirement can help a colony choose the best available site, even when few ants have the opportunity to compare sites directly, because recruiters to a given site launch the rapid transport of the bulk of the colony only if enough active ants have been "convinced" of the worth of the site. This exemplifies how insect societies can achieve adaptive colony-level behaviors from the decentralized interactions of relatively poorly informed insects, each combining her own limited direct information with indirect cues about the experience of her nestmates.     

Graded recruitment and hunting strategies linked to prey weight and size in the ponerine ant Ectatomma ruidum

According to the weight and size of their prey, Ectatomma ruidum workers can employ different recruitment systems (solitary hunting, cooperative hunting and group hunting with recruitment) when mastering and retrieving prey items from short distances from the nest. Prey size determined the backwards entry typically adopted by this species, while prey weight determined the predatory strategy selected. After a common initial sequence (search for prey, detection, localization), predatory sequences varied in terms of the type of approach, the site of seizure, the reaction after stinging and the type of transport. Nevertheless, irrespective of prey weight and size, seizure was preferentially oriented towards the head and prey were always stung. Short-range recruitment and mass recruitment without trail laying were elicited by a large range of heavy prey (> 2.5 times the weight of an individual worker). According to the mortality risk associated with each prey, hunters exhibited a “prudent” stinging posture associated with an increase in the duration of the subsequent phase of waiting for prey immobilization. The overall time of capture was positively correlated with the weight of the prey. When collective hunting strategies were involved, E. ruidum colonies matched the number of recruited hunters to the size and weight of the prey. Compared to solitary hunting strategies, for short food–nest distances, this graded recruitment appeared to enhance the energetic benefits derived by this species from the use of recruitment systems: the higher the number of workers involved in the recruitment process, the greater the energetic benefits obtained. The exhibition or absence of trail laying behavior in the recruitment responses displayed by E. ruidum workers is discussed in relation to their involvement in scavenging or predatory behavior. Received: 27 June 1996 / Accepted after revision: 3 March 1997