Dominance and fertility in a functionally monogynous ant

Summary Colonies of some leptothoracine ants may contain several inseminated but sterile females in addition to a single, fertile queen (functional monogyny). We here report the first observations on the behavior of these supernumerary females in the nearctic ant Leptothorax sp.A, a species belonging to the L. muscorum complex.In four colonies, each with up to eight intermorphic females, ritualized or openly aggressive interactions between individuals were observed, similar to those among workers of some other leptothoracine ants. The responses of individual females during encounters with nestmates apparently reflect the existence of linear dominance hierarchies. In each colony, the highest ranking individual was fed and groomed significantly more often than other females, and was the only one to oviposit after hibernation and to become fully physogastric during the first weeks of spring. When these -females were removed from three colonies, several other females started to lay eggs. However, in each colony only the highest ranking individual remained fertile; the others either were pushed out of the nest and finally killed by the workers, or their ovaries degenerated again.Aggressive interactions among females may also play an important role in the foundation of new colonies either by inducing intermorphs to leave the maternal nest to found new colonies solitarily or by inducing colony fission. Offprint requests to: J. Heinze at his new address     

Individual experience-based foraging can generate community territorial structure for competing ant species

To gain additional territory while defending existing territory, animals must acquire and use information regarding resource characteristics and competitive pressure. For social organisms like ants, individual workers have experiences to acquire information, but territory establishment is a colony level behavior. Colony behavior, in turn, affects community structure. Here, I investigate how an individual ant’s previous experience affects its future foraging behavior and how individual behaviors can scale up to community territorial structure for two coexisting Formica species. To do this, I combine a field survey, a multi-agent computer simulation, and a manipulation experiment. The field survey shows that workers of both species co-occur on many trees early in the season, but ants on trees become segregated by species as the season progresses. The simulation demonstrates how this segregated spatial distribution can result from ants using a foraging strategy in which individuals show a preference for foraging sites based on previous experience. The experiment suggests that these ants are indeed capable of experience-based foraging behavior; ants preferentially return to sites where they have had positive experiences and avoid sites where they have had negative experiences. Results from this study suggest that spatially explicit information can be collected and stored by individuals to facilitate colony territorial structure, and that future investigations of community territory formation should include effects of individual previous experience.     

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.     

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.

Division of labour in a crisis: task allocation during colony emigration in the ant Leptothorax unifasciatus (Latr.)

Division of labour during colony emigration is widespread in ants. An important problem is how tasks are allocated during colony movement from one nest site to another. The generally favoured view is that emigrations are organised by a minority group of individuals, which either work unusually hard at tasks (elites) or have the exclusive task of carrying out the emigration (moving specialists). Five consecutive emigrations of a Leptothorax unifasciatus (Latr.) colony showed that the number of transporters, i.e. the individuals that took an active part in the emigration by transporting brood and ants, was smaller than it would have been if allocation of this task was random during each emigration. However, single emigrations of another three colonies, for which the spatial distribution and behaviour of the workers had been observed for a week prior to the emigration, demonstrated that the transporters did not form a homogeneous group. They differed in their spatial positions and tasks before the emigration. There was also no evidence that transporters worked harder or less hard than their nestmates before the emigration. Therefore, the individuals which carry out emigrations in L. unifasciatus colonies appear to be neither moving specialists nor elites. We propose that task allocation during emigrations of L. unifasciatus colonies is based on a feedback mechanism that involves learning.     

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.     

The superseded female's dilemma: ultimate and proximate factors that influence guarding behaviour of the carpenter bee Xylocopa pubescens

Both solitary and primitively social nests of the facultatively social carpenter bee Xylocopa pubescens can be found throughout most of the breeding season. In social nests there is reproductive division of labour between a dominant forager and a guarding female. Two types of guarding females can be discerned: the young pre-reproductive guards, and older, formerly reproductive guards. The latter type of guard is found when, after a take-over of reproductive dominance either by a nestmate (mostly a daughter) or an intruder, the defeated female stays in the nest instead of leaving to try and found or usurp another nest. She is then manipulated into the role of a guard. The dominant female profits from the presence of the guard since she protects the nest against pollen robbery by conspecifics (Hogendoorn and Velthuis 1993). We have studied why superseded females might prefer to remain as a guard, rather than try their luck somewhere else. The hypotheses investigated pertain to (1) the difficulty for the defeated female of finding a new nest and of restarting reproductive activities due to (a) ecological constraints (nest and pollen shortage) and (b) the effect of age and wear on the defeated female; (2) the effects of guarding in terms of inclusive fitness. We found that superseded females remained as guards significantly more often when a nestmate (not necessarily close kin) took over reproductive dominance than when an intruder did so. Other factors associated with the decision of the defeated female to stay or leave were her age and the number of her own young still present after the supersedure. The probability of finding or constructing a new nest was lower for old than for young females. After finding a nest, old females produced less brood than young foundresses. As a result of these two factors old superseded females gained, in terms of inclusive fitness, by staying as guards, whereas young females profited from leaving the nest. We interpret these results as an indication that guarding behaviour has evolved due to kin selection. However, kin discrimination apparently did not occur. Therefore we conclude that in this species kin selection is not, in the proximate frame of reference, based on kin recognition and preference for helping kin. Correspondence to: K. Hogendoorn     

Choosing a home: how the scouts in a honey bee swarm perceive the completion of their group decision making

This study considers the mystery of how the scout bees in a honey bee swarm know when they have completed their group decision making regarding the swarm's new nest site. More specifically, we investigated how the scouts sense when it is appropriate for them to begin producing the worker piping signals that stimulate their swarm-mates to prepare for the flight to their new home. We tested two hypotheses: "consensus sensing," the scouts noting when all the bees performing waggle dances are advertising just one site; and "quorum sensing," the scouts noting when one site is being visited by a sufficiently large number of scouts. Our test involved monitoring four swarms as they discovered, recruited to, and chose between two nest boxes and their scouts started producing piping signals. We found that a consensus among the dancers was neither necessary nor sufficient for the start of worker piping, which indicates that the consensus sensing hypothesis is false. We also found that a buildup of 10–15 or more bees at one of the nest boxes was consistently associated with the start of worker piping, which indicates that the quorum sensing hypothesis may be true. In considering why the scout bees rely on reaching a quorum rather than a consensus as their cue of when to start preparing for liftoff, we suggest that quorum sensing may provide a better balance between accuracy and speed in decision making. In short, the bees appear to begin preparations for liftoff as soon as enough of the scout bees, but not all of them, have approved of one of the potential nest sites.

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.     

False feedings at the nests of carrion crows<Emphasis Type="Italic"> Corvus corone corone</Emphasis>

False feedings, when individuals visit the nest but refrain from feeding the chicks, occur in some cooperative species and have been interpreted in the white-winged chough (Corcorax melanorhamphos) as active deception by helpers towards the rest of the group. In a cooperatively breeding population of carrion crows (Corvus corone corone) 81.5% of the individuals that provided nestling care showed various kinds of false feedings: arriving at the nest with no food, consuming part or all the food brought to the nest, or taking back from a chicks gape the food that had just been delivered. False feedings occurred on average during 16.3% of nest visits, with some individuals performing them at very high rates (up to 64% of nest visits). False feedings occurred at similar rates in unassisted pairs and groups with helpers, and breeding females showed false feeding at significantly higher rates than other group members. Furthermore, individuals showed false feedings regardless of whether they were alone on the nest or in the presence of other group members, and false feedings did not provoke aggression by the rest of the group. False feedings are not likely to represent deceptive help in the carrion crow. We suggest that crows evaluate the chicks condition during nest visits and that false feedings occur as result of a trade-off between their own hunger and the chicks needs.Communicated by W.A. Searcy     

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     

Less is more: social learning of predator recognition requires a low demonstrator to observer ratio in Arctic charr (Salvelinus alpinus)

Reintroduction programs for conservation purposes suffer from low post-release survival of captive-bred animals, especially so with fish reintroductions. High mortality is most often due to weakened antipredator skills, which could be enhanced through direct experience with predators and social learning. As yet, the relative efficiency of these learning methods has not been tested, and the significance of the factors determining how efficiently antipredator responses are transmitted socially from experienced (demonstrator) to predator naive (observer) individuals is largely unknown. Using hatchery-bred stock of the endangered Saimaa Arctic charr (Salvelinus alpinus), we show for the first time that: (1) contrary to theoretical predictions, acquired predator recognition was only socially transmitted from predator-experienced to predator-naive fish in groups with a smaller proportion of experienced than naive individuals, and that (2) socially acquired avoidance of the predator odors was as strong as in those fish conditioned with a combination of odors and predator exposure. We conclude that using social transmission procedures in training fish for reintroduction programs may provide considerable ecological, economical and ethical advantages.Communicated by J. Krause     

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.     

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.     

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.     

Defense against parasites by hitchhikers in leaf-cutting ants: a quantitative assessment

Summary A curious behavior in leaf-cutting ants in the genus Atta is the hitchhiking of small minim workers on leaf fragments carried by larger workers. Two functions of these hitchhikers have been proposed: (1) defense of leaf carriers against parasitic flies in the family Phoridae (ant protection hypothesis; Eibl-Eibesfeldt and Eibl-Eibesfeldt 1967) and (2) reduction of transport costs of small minims that collect plant fluids (energy conservation hypothesis; Stradling 1978).We studied hitchhiking behavior in colonies of Atta colombica on Barro Colorado Island, Panama, and found strong evidence in favor of the ant protection hypothesis. Females of Apocephalus attophilus (Diptera: Phoridae) attack leaf carriers of A. colombica and deposit eggs in the head capsules of these ants. Our observations indicate that parasites require leaf fragments to stand on during oviposition, and, as a result, only leaf carriers are susceptible to parasitic attack. The presence of hitchhikers reduces significantly both the time parasites spend on leaf fragments and the probability that they will land in the first place. Results of experimental introductions of parasites and a year of biweekly censuses at ten colonies indicate that leaf-cutting ants adjust the level of hitchhiking to accommodate both daily and seasonal changes in the abundance of parasites.We found little evidence in support of the energy conservation hypothesis. If it is assumed that all minim workers hitchhike back to the nest, our calculations indicate that total transport costs along a foraging trail are reduced by 10% or less. However, our observations indicate that only 50% of returning minim workers hitchhike, and therefore energy savings are actually considerably less than 10%. Leaf-cutting ants in the genera Atta and Acromyrmex are attacked by over 20 species of parasitic phorids. In the discussion we review what is known about these associations and suggest that these parasites have influenced the ecology and evolution of polyethism in leaf-cutting ants. Offprint requests to: D.H. Feener (at his present address)     

Patterns of participation and free riding in territorial conflicts among ringtailed lemurs (<Emphasis Type="Italic">Lemur catta</Emphasis>)

Cooperation in animal social groups may be limited by the threat of free riding, the potential for individuals to reap the benefits of other individuals actions without paying their share of the costs. Here we investigate the factors that influence individual contributions to group-level benefits by studying individual participation in territorial defense among female ringtailed lemurs (Lemur catta). To control for potentially confounding factors, particularly group size, we studied two semi-free-ranging groups at the Duke University Primate Center. First, we used a combination of experimental and observational methods to investigate the costs and benefits of territorial defense for individual lemurs. We found three indications of costs: physical contact occurred during inter-group encounters, participation in territorial defense was negatively correlated with ambient temperature, and rates of self-directed behaviors increased during encounters. Benefits were more difficult to quantify, but observational and experimental tests suggested that individuals shared the gains of territorial defense by foraging in defended territories. Thus, during experiments in which one of the groups was prevented from defending its territory, the free-ranging group made more frequent incursions into the other groups territory. Second, we examined variation in participation in territorial defense. Individuals varied significantly in their rates of aggression and genital marking during inter-group encounters. The extensive variation documented among individuals was partially accounted for by dominance rank, kinship and patterns of parental care. However, we found no evidence to suggest that participation was enforced through punishment (policing) or exchange of benefits involving grooming. In conclusion, this study provides further insights into cooperative behavior in mammalian social groups by revealing how the costs and benefits of territoriality influence patterns of individual participation in the context of shared (collective) goods.Communicated by P. Kappeler     

Behavioral and life-history components of division of labor in honey bees (Apis mellifera L.)

Variability exists among worker honey bees for components of division of labor. These components are of two types, those that affect foraging behavior and those that affect life-history characteristics of workers. Variable foraging behavior components are: the probability that foraging workers collect (1) pollen only; (2) nectar only; and (3) pollen and nectar on the same trip. Life history components are: (1) the age the workers initiate foraging behavior; (2) the length of the foraging life of a worker; and (3) worker length of life. We show how these components may interact to change the social organization of honey bee colonies and the lifetime foraging productivity of individual workers. Selection acting on foraging behavior components may result in changes in the proportion of workers collecting pollen and nectar. Selection acting on life-history components may affect the size of the foraging population and the distribution of workers between within nest and foraging activities. We suggest that these components define possible sociogenic pathways through which colony-level natural selection can change social organization. These pathways may be analogous to developmental pathways in the morphogenesis of individual organisms because small changes in behavioral or life history components of individual workers may lead to major changes in the organizational structure of colonies. Correspondence to: R.E. Page, Jr.