The organisation of collective foraging in the harvester termite Hodotermes mossambicus (Isoptera)

Summary The process by which the foragers build up a foraging system between the nest hole and the food source, starting from an initial random distribution, was investigated under conditions of both light and dark in the laboratory.The extent to which trail pheromone is used can be determined by observing the abdominal position of workers moving outside the nest and can be classified into three categories, namely no trail-laying, marking and directional trail-laying.When workers of this species emerge from the nest hole into light emitted from a certain direction they orientate photo-menotactically. First, they mark the area around the nest hole, moving in small loops; they then make larger exploratory loops without trail-laying. Once they find a source of food they return directly to the immediate vicinity of the nest hole. As the area around the hole has already been marked it is easy for the termites to locate the hole. When a piece of food has been deposited by the hole the termites display directional trail-laying behaviour by leaving a trail in the direction of the source of food, this trail being stronger nearer the nest and decreasing as it approaches its goal. Inexperienced explorers are biased in this direction with a statistically higher frequency. If successful they join in the foraging and repeatedly contribute to the trail, which then gradually brings together most of the foraging population.In darkness exploratory loops are made with continuous marking, always returning to the nest hole, whereby a scent gradient decreasing away from the hole is formed. Once a source of food has been discovered the termites are neither able to return directly to the nest hole nor to return along their outward marking trail. They leave marking trails making exploratory loops from the food source with successively increasing radii. Another scent gradient, decreasing outwards from the food, is created. If the termite reaches the increasing scent around the nest hole from the decreasing scent around the food source it will locate the hole chemotactically. As other food finders repeatedly cross from one field to the other the scents will combine and become concentrated, thus forming a foraging path.     

Energetics,reproductive suppression and obligate communal breeding in carnivores

Summary Leptogenys processionalis Jerdon forages on termites and other arthropods by raiding in branched trails. Growth and topology of these search trails were studied using Horton's (1945) technique orginally developed to analyze the branching pattern of river systems. Branching was always a bifurcating process and branches emerged symmetrically on either side of the main trail. Branching coefficients (R _b ) were similar to those of a few biological branching systems, such as lungs, that are considered to be non-random in their branching pattern. The R _b values indicated that the rate of branching and growth of trails remained constant within each foraging bout. The length of trails became shorter as they grew out and branched. The branching process was a function of the spatial separation of food patches in the terminal search field. Ants in the terminal search field send signals on encountering prey. The recruits cannot discriminate between these signals if they arise from two food patches situated <40 cm from each other, and hence converge on them in a single trail. However, discrimination is possible when food patches are >40 cm apart and hence recruits congregate on them separately in two trails, resulting in branching. Thus, the branching process is a result of independent decision by the ants conforming to certain simple rules and not a collective decision of the whole colony. We argue that mass recruiting ants selected to forage by branching pattern of trails because of its efficiency over other topologies (Stevens 1973) in minimizing the cost of travel, both from the nest to the food patches and between food patches. Further, the branch angles appear to be a trade-off to minimize travel cost and the resistance to the flow of ants comprising the column.Offprint requests to: T. Veena     

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.     

Pheromone trails in the Brazilian ant Pheidole oxyops: extreme properties and dual recruitment action

Communication of feeding locations is widespread in social animals. Many ants use pheromone trails to guide nestmates to food sources, but trail properties and how they are used vary. The ant Pheidole oxyops retrieves prey cooperatively using multiple workers. The recruited workers are guided to the prey by a pheromone trail laid by the initial discoverer. In comparison to other ants, this trail has extreme properties. Despite being laid by just one ant, freshly laid trails are followed very accurately (84.4?% correct choices at a bifurcation), but decay in only 5–7?min. This extreme accuracy and short duration probably reflect adaptations to underlying differences in feeding ecology. In particular, P. oxyops needs to rapidly recruit nestmates to a precise location in a competitive environment. Rapid decay combined with a natural walking speed of 1.4?m/min should set an upper limit of 4?m (an 8-m round trip) on recruitment range. However, experimentally placed food items up to 8?m from the nest entrance were cooperatively retrieved. This greater range is due to the trail having a dual recruitment role. It not only recruits from the nest but also intercepts ants already outside the nest, causing them to join the trail. Seventy-five per cent of ants joining the trail then followed it towards the food item. Even when direct recruitment from the nest was prevented, this secondary recruitment action resulted in seven times as many ants locating a food source than by chance discovery and in items being moved 46?% sooner.     

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.     

The function of polydomy: the ant <Emphasis Type="Italic">Crematogaster torosa</Emphasis> preferentially forms new nests near food sources and fortifies outstations

Many ant species are polydomous, forming multiple spatially segregated nests that exchange workers and brood. However, why polydomy occurs is still uncertain. We investigated whether colonies of Crematogaster torosa form new polydomous nests to better exploit temporally stable food resources. Specifically, we tested the effect of food presence or absence and distance on the likelihood that colonies would form a new nest. Because this species also forms little-known structures that house only workers without brood (outstations), we also compared the function of this structure with true nests. Laboratory-reared colonies were connected to a new foraging arena containing potential nest sites with or without food for 4 months. When food was present, most colonies formed polydomous nests nearby and the remainder formed outstations. When food was absent, the behavior of colonies differed significantly, frequently forming outstations but never polydomous nests. Distance had no effect on the type of structure formed, but when food was present, a larger proportion of the workforce moved shorter distances. Workers often fortified the entrances to both structures and used them for storage of dried insect tissue (“jerky”). In an investigation of spatial fidelity, we found that workers on the between-nest trail were associated with the original nest, whereas workers collecting food were more likely to be associated with the new nest or outstation. C. torosa appears to have a flexible colony structure, forming both outstations and polydomous nests. Polydomous nests in this species were associated with foraging and were only formed near food resources.     

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     

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.     

Costs of trail construction and maintenance in the leaf-cutting ant Atta columbica

Leaf-cutting ants of the genus Atta use trunk trails during foraging which may persist for months or years. The time and energy costs of trail construction and maintenance were estimated for colonies of Atta columbica on Barro Colorado Island, Panama, to determine if these costs are likely to constrain new trail construction and promote persistence of existing trails. Large workers 2.2-2.9 mm in headwidth participated in trail-clearing significantly more frequently than typical leaf-carriers, indicating that they may form a distinctive task group within the foraging force. Small litter items were carried off trails, while large ones were cut up before removal, greatly increasing the costs of removing large litter items. The average time cost of removing a kilogram of litter was estimated at 3,359 ant-hours, and energy costs at 4.6 kJ. Colonies maintained trail systems 267 m in length and 16.5 m² in area, and built an estimated 2.7 km of trail with an area of 134 m² during a year. Based on litter standing crop and estimates of litterfall rates, total costs to colonies averaged 11,000 ant-days of work and the energy equivalent of 8,000 leaf burdens. These costs are small relative to the number of available workers and rates of mass harvest, suggesting that costs do not significantly constrain trail construction. Instead, trails may persist because they provide access to high-quality resources or because only a few trails are required to fully exploit the foraging territory.     

Trail discrimination signal of Lasius japonicus (Hymenoptera: Formicidae)

Summary. Trail-following behavior of Lasius japonicus was colony-specific in the field, while trail pheromone activity was not. We found that the footprint substance caused colony-specific trail-following behavior only when working in conjunction with the trail pheromone. The footprint substance alone did not lead the workers to follow trails. The substance consisted mainly of hydrocarbons with composition almost identical to that of cuticular hydrocarbons, except for the absence of n-alkanes. Nestmate workers shared footprint hydrocarbon profiles as well as cuticular hydrocarbons, but the profiles differed among colonies. We therefore consider that the footprint hydrocarbon profiles serve as the trail discrimination signal in L. japonicus.     

High social density increases foraging and scouting rates and induces polydomy in Temnothorax ants

Many organisms live in crowded groups where social density affects behavior and fitness. Social insects inhabit nests that contain many individuals where physical interactions facilitate information flow and organize collective behaviors such as foraging, colony defense, and nest emigration. Changes in nest space and intranidal crowding can alter social interactions and affect worker behavior. Here, I examined the effects of social density on foraging, scouting, and polydomy behavior in ant colonies—using the species Temnothorax rugatulus. First, I analyzed field colonies and determined that nest area scaled isometrically with colony mass—this indicates that nest area changes proportionally with colony size and suggests that ants actively control intranidal density. Second, laboratory experiments showed that colonies maintained under crowded conditions had greater foraging and scouting activities compared to the same colonies maintained at a lower density. Moreover, crowded colonies were significantly more likely to become polydomous. Polydomous colonies divided evenly based on mass between two nests but distributed fewer, heavier workers and brood to the new nests. Polydomous colonies also showed different foraging and scouting rates compared to the same colonies under monodomous conditions. Combined, the results indicate that social density is an important colony phenotype that affects individual and collective behavior in ants. I discuss the function of social density in affecting communication and the organization of labor in social insects and hypothesize that the collective management of social density is a group level adaptation in social insects.     

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     

Dance precision of <Emphasis Type="Italic">Apis florea</Emphasis>—clues to the evolution of the honeybee dance language?

All honeybee species make use of the waggle dance to communicate the direction and distance to both food sources and potential new nest sites. When foraging, all species face an identical problem: conveying information about profitable floral patches. However, profound differences in nesting biology (some nest in cavities while others nest in the open, often on a branch or a cliff face) may mean that species have different requirements when dancing to advertise new nest sites. In cavity nesting species, nest sites are a precise location in the landscape: usually a small opening leading to a cavity in a hollow tree. Dances for cavities therefore need to be as precise as possible. In contrast, when the potential nest site comprises a tree or perhaps seven a patch of trees, precision is less necessary. Similarly, when a food patch is advertised, dances need not be very precise, as floral patches are often large, unless they are so far away that recruits need more precise information to be able to locate them. In this paper, we study the dance precision of the open-nesting red dwarf bee Apis florea. By comparing the precision of dances for food sources and nest sites, we show that A. florea workers dance with the same imprecision irrespective of context. This is in sharp contrast with the cavity-nesting Apis mellifera that increases the precision of its dance when advertising a potential new home. We suggest that our results are in accordance with the hypothesis that the honeybees’ dance communication initially evolved to convey information about new nest sites and was only later adapted for the context of foraging.     

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.     

A stingless bee (Melipona panamica) indicates food location without using a scent trail

The study of location specification in recruitment communication by bees has focused on two dimensions: direction and distance from the nest. Yet the third dimension, height above ground, may be significant in the tall and dense forest habitats of stingless bees. Foragers of the stingless bee Scaptotrigona postica recruit to a specific three-dimensional location by laying a scent trail. Stingless bees in the genus Melipona are thought to have a more sophisticated recruitment system that communicates distance through sounds inside the nest and direction through pointing zig-zag flights outside the nest. However, prior research on Melipona has not examined height communication or even established that foragers can recruit newcomers to a specific location. We used identical paired feeders to investigate recruitment to food in M panamica on Barro Colorado Island, Panama. We trained foragers from an observation hive to one feeder and monitored both feeders for the subsequent arrival of newcomers. We changed the relative positions of the feeders to test for correct direction, distance, and canopy-level communication. A 40-m canopy tower located inside the forest enabled us to examine canopy-level communication. We found that M. panamica foragers can recruit to a specific (1) direction, (2) distance, and (3) canopy level. To test the possibility that foragers accomplish this by means of a scent trail, we placed the colony on one shore of a small cove and trained bees over 116 m of open water to a feeder located on the opposite shore. We also placed a second feeder on this shore, equidistant from the colony but 20 m from the first feeder. Significantly more newcomers consistently arrived at the feeder visited by the foragers. Thus foragers evidently do not need a scent trail to communicate direction. Inside the nest, a forager produces pulsed sounds while visibly vibrating her wings after returning from a good food source. She is attended by other bees who cluster and hold their antennae around her, following her as she rapidly spins clockwise and counterclockwise. Locational information may be encoded in this behavior. However, foragers may also directly lead newcomers to the food source. Further experiments are planned to test for such piloting and other communication mechanisms.     

Forager specialization and the control of nest repair in Polybia occidentalis Olivier (Hymenoptera : Vespidae)

We measured patterns of individual forager specialization and colony-wide rates of material input during periods of response to experimental nest damage and during control periods in three colonies of the tropical social wasp Polybia occidentalis.

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.     

The chemosensory ability of the predatory leech Whitmania laevis (Arhynchobdellida: Haemopidae) for prey searching

Although prey-detecting and searching abilities of predatory leeches of rhynchobdellid or the Erpobdelliformes of arhynchobdellid species have been studied in the past, hirudiniformes leeches are rarely mentioned. In this study, we investigated the chemosensory ability for prey-detecting and searching in Whitmania laevis, a hirudiniformes species that mainly preys on freshwater snails, and examined if such ability aided in their prey selection. Five sympatric snail species, i.e., apple snail Pomacea canaliculata, thiarid snail Thiara tuberculata, viviparid snail Sinotaia quadrata, ear pond snail Radix auricularia swinhoei and tadpole snail Physa acuta were used as prey. Our results showed that W. laevis has the chemosensory ability to detect the waterborne odors of snails. However, they follow the snails by their mucus trails, and not by the odor that the snails leave in the water. Of these five snail species, W. laevis only followed the trails of the thiarid snails, ear pond snails and tadpole snails, and did not show a different response to the trails produced by snails of different sizes. Our results suggest that W. laevis can use waterborne odors to detect the existence of prey. They rely on mucus trails to follow their preferred prey, but do not distinguish between snails of a preferred size by their mucus trails. In addition, when following the trail of a preferred snail, W. laevis exhibits a newly described searching behavior, i.e., head tapping, and may use it to locate a snail trail and increase its probability of finding the trail-laying snail nearby.