The regulation of foraging activity in red harvester ant colonies

Behavioral plasticity in social insects is intriguing because colonies adjust to environmental change through the aggregated responses of individuals. Without central control, colonies adjust numbers of workers allocated to various tasks. Individual decisions are based on local information from the environment and other workers. This study examines how colonies of the seed-eating ant Pogonomyrmex barbatus adjust the intensity of foraging in an arid environment where conspecific neighbors compete for foraging area. The main question is how foragers decide whether to leave the nest. Patrollers search the area before foragers emerge. Removal experiments show that the return of the patrollers stimulates the onset of foraging, and later, the rate at which foragers return affects the rate at which foragers continue to leave the nest. Foraging activity is less sensitive to changes in the rate of returning foragers than to changes in the rate of returning patrollers. These results suggest that whether a colony forages at all on a given day depends on conditions detected early by patrollers but that once foraging begins, the intensity of foraging does not track, on an hourly timescale, how quickly foragers can find food.     

Colony variation in the collective regulation of foraging by harvester ants

This study investigates variation in collective behavior in a natural population of colonies of the harvester ant, Pogonomyrmex barbatus. Harvester ant colonies regulate foraging activity to adjust to current food availability; the rate at which inactive foragers leave the nest on the next trip depends on the rate at which successful foragers return with food. This study investigates differences among colonies in foraging activity and how these differences are associated with variation among colonies in the regulation of foraging. Colonies differ in the baseline rate at which patrollers leave the nest, without stimulation from returning ants. This baseline rate predicts a colony's foraging activity, suggesting there is a colony-specific activity level that influences how quickly any ant leaves the nest. When a colony's foraging activity is high, the colony is more likely to regulate foraging. Moreover, colonies differ in the propensity to adjust the rate of outgoing foragers to the rate of forager return. Naturally occurring variation in the regulation of foraging may lead to variation in colony survival and reproductive success.     

Interactions with Combined Chemical Cues Inform Harvester Ant Foragers' Decisions to Leave the Nest in Search of Food

Social insect colonies operate without central control or any global assessment of what needs to be done by workers. Colony organization arises from the responses of individuals to local cues. Red harvester ants (Pogonomyrmex barbatus) regulate foraging using interactions between returning and outgoing foragers. The rate at which foragers return with seeds, a measure of food availability, sets the rate at which outgoing foragers leave the nest on foraging trips. We used mimics to test whether outgoing foragers inside the nest respond to the odor of food, oleic acid, the odor of the forager itself, cuticular hydrocarbons, or a combination of both with increased foraging activity. We compared foraging activity, the rate at which foragers passed a line on a trail, before and after the addition of mimics. The combination of both odors, those of food and of foragers, is required to stimulate foraging. The addition of blank mimics, mimics coated with food odor alone, or mimics coated with forager odor alone did not increase foraging activity. We compared the rates at which foragers inside the nest interacted with other ants, blank mimics, and mimics coated with a combination of food and forager odor. Foragers inside the nest interacted more with mimics coated with combined forager/seed odors than with blank mimics, and these interactions had the same effect as those with other foragers. Outgoing foragers inside the nest entrance are stimulated to leave the nest in search of food by interacting with foragers returning with seeds. By using the combined odors of forager cuticular hydrocarbons and of seeds, the colony captures precise information, on the timescale of seconds, about the current availability of food.     

How patrollers set foraging direction in harvester ants

Recruitment to food or nest sites is well known in ants; the recruiting ants lay a chemical trail that other ants follow to the target site, or they walk with other ants to the target site. Here we report that a different process determines foraging direction in the harvester ant Pogonomyrmex barbatus. Each day, the colony chooses from among up to eight distinct foraging trails; colonies use different trails on different days. Here we show that the patrollers regulate the direction taken by foragers each day by depositing Dufour's secretions onto a sector of the nest mound about 20 cm long and leading to the beginning of a foraging trail. The patrollers do not recruit foragers all the way to food sources, which may be up to 20 m away. Fewer foragers traveled along a trail if patrollers had no access to the sector of the nest mound leading to that trail. Adding Dufour's gland extract to patroller-free sectors of the nest mound rescued foraging in that direction, while poison gland extract did not. We also found that in the absence of patrollers, most foragers used the direction they had used on the previous day. Thus, the colony's 30-50 patrollers act as gatekeepers for thousands of foragers and choose a foraging direction, but they do not recruit and lead foragers all the way to a food source.     

The short-term regulation of foraging in harvester ants

In the seed-eating ant Pogonomyrmex barbatus, the return ofsuccessful foragers stimulates inactive foragers to leave thenest. The rate at which successful foragers return to the nestdepends on food availability; the more food available, the morequickly foragers will find it and bring it back. Field experimentsexamined how quickly a colony can adjust to a decline in therate of forager return, and thus to a decline in food availability,by slowing down foraging activity. In response to a brief, 3-to 5-min reduction in the forager return rate, foraging activityusually decreased within 2–3 min and then recovered within5 min. This indicates that whether an inactive forager leavesthe nest on its next trip depends on its very recent experienceof the rate of forager return. On some days, colonies respondedmore to a change in forager return rate. The rapid colony responseto fluctuations in forager return rate, enabling colonies toact as risk-averse foragers, may arise from the limited intervalover which an ant can track its encounters with returning foragers.     

The Regulation of Ant Colony Foraging Activity without Spatial Information

Many dynamical networks, such as the ones that produce the collective behavior of social insects, operate without any central control, instead arising from local interactions among individuals. A well-studied example is the formation of recruitment trails in ant colonies, but many ant species do not use pheromone trails. We present a model of the regulation of foraging by harvester ant (Pogonomyrmex barbatus) colonies. This species forages for scattered seeds that one ant can retrieve on its own, so there is no need for spatial information such as pheromone trails that lead ants to specific locations. Previous work shows that colony foraging activity, the rate at which ants go out to search individually for seeds, is regulated in response to current food availability throughout the colony's foraging area. Ants use the rate of brief antennal contacts inside the nest between foragers returning with food and outgoing foragers available to leave the nest on the next foraging trip. Here we present a feedback-based algorithm that captures the main features of data from field experiments in which the rate of returning foragers was manipulated. The algorithm draws on our finding that the distribution of intervals between successive ants returning to the nest is a Poisson process. We fitted the parameter that estimates the effect of each returning forager on the rate at which outgoing foragers leave the nest. We found that correlations between observed rates of returning foragers and simulated rates of outgoing foragers, using our model, were similar to those in the data. Our simple stochastic model shows how the regulation of ant colony foraging can operate without spatial information, describing a process at the level of individual ants that predicts the overall foraging activity of the colony.     

Interactions Increase Forager Availability and Activity in Harvester Ants

Social insect colonies use interactions among workers to regulate collective behavior. Harvester ant foragers interact in a chamber just inside the nest entrance, here called the ''entrance chamber''. Previous studies of the activation of foragers in red harvester ants show that an outgoing forager inside the nest experiences an increase in brief antennal contacts before it leaves the nest to forage. Here we compare the interaction rate experienced by foragers that left the nest and ants that did not. We found that ants in the entrance chamber that leave the nest to forage experienced more interactions than ants that descend to the deeper nest without foraging. Additionally, we found that the availability of foragers in the entrance chamber is associated with the rate of forager return. An increase in the rate of forager return leads to an increase in the rate at which ants descend to the deeper nest, which then stimulates more ants to ascend into the entrance chamber. Thus a higher rate of forager return leads to more available foragers in the entrance chamber. The highest density of interactions occurs near the nest entrance and the entrances of the tunnels from the entrance chamber to the deeper nest. Local interactions with returning foragers regulate both the activation of waiting foragers and the number of foragers available to be activated.     

How does colony growth influence communication in ants?

Summary. As Lasius niger societies grow from incipient nests to mature colonies, their foraging strategies shift from the individual exploitation of food sources to mass recruitment. Colony size instead of age is the key factor that shapes the exploratory and foraging responses of Lasius niger: a drastic reduction (or increase) of the population elicits an activity profile similar to that observed in younger (or older) societies of the same size. As a colony grows, the proportion of patrollers significantly decreases while the proportion of conveyors remains rather constant. As regards the energetic return, it increases with incipient nest size due to the replacement of minims by ordinary workers of larger crop capacity. We also demonstrate that minims of incipient nests modulate their trail-laying behaviour according to the social context, in this case the colony size. During their ontogenesis, L. niger colonies exhibit a progressive integration of individual foragers into a network of communication, the adaptive significance of which is discussed.     

Dispersed central place foraging in Australian meat ants

Summary Australian meat ants often inhabit colonies with widely dispersed nest holes, and this study examines how resource is harvested and distributed in a colony ofIridomyrmex sanguineus Smith (Formicidae: Dolichoderinae). The three principal types of foragers (tenders, honeydew transporters, scavengers) exhibited nest hole fidelity, where harvested resource was consistently delivered to the same nest hole by each foraging individual. Australian meat ants thus use a harvesting system based on dispersed central place foraging. Evidence of frequent larval transport among nest holes, age polyethism developing in the direction of foraging, and the tendency for nest-associated workers to accept new nest holes more readily than foragers, suggests that workers develop fidelity to the particular nest hole in which they eclose. Coupled with larval transport, nest hole fidelity may allow a colony with widely dispersed nest holes to adjust its structure to more efficiently harvest a resource distributed unevenly in space or time.     

The effects of colony characteristics on life span and foraging behavior of individual wasps (Polybia occidentalis,Hymenoptera: Vespidae)

Summary We studied the effects of intrinsic colony characteristics and an imposed contingency on the life span and behavior of foragers in the swarm-founding social waspPolybia occidentalis. Data were collected on marked, known-age workers introduced into four observation colonies.To test the hypothesis that colony demographic features affect worker life span, we examined the relationships of colony age and size with worker life span using survivorship analysis. Colony age and size had positive relationships with life span; marked workers from two larger, older colonies had longer life spans (¯X = 24.7 days) than those from two smaller, younger colonies (¯X = 20.1 days).We quantified the effects of experimentally imposed nest damage on forager behavior, to determine which of three predicted behavioral responses by foragers to this contingency (increased probability of foraging for building material, increased rate of foraging, or decrease in age of onset of foraging) would be employed. Increasing the colony level of need for materials used in nest construction (wood pulp and water) by damaging the nests of two colonies did not cause an increase in either the proportion of marked workers that gathered nest materials or in foraging rates of marked individuals, when compared with introduced workers in two simultaneously observed control colonies. Instead, nest damage caused a decrease in the age at which marked workers first foraged for pulp and water. The response to an increase in the need for building materials was an acceleration of behavioral development in some workers.     

Foraging decisions of individual workers vary with colony size in the greenhead ant Rhytidoponera metallica (Formicidae, Ectatomminae)

Summary. The ability of worker ants to adapt their behaviour depending on the social environment of the colony is imperative for colony growth and survival. In this study we use the greenhead ant Rhytidoponera metallica to test for a relationship between colony size and foraging behaviour. We controlled for possible confounding ontogenetic and age effects by splitting large colonies into small and large colony fragments. Large and small colonies differed in worker number but not worker relatedness or worker/brood ratios. Differences in foraging activity were tested in the context of single foraging cycles with and without the opportunity to retrieve food. We found that workers from large colonies foraged for longer distances and spent more time outside the nest than foragers from small colonies. However, foragers from large and small colonies retrieved the first prey item they contacted, irrespective of prey size. Our results show that in R. metallica, foraging decisions made outside the nest by individual workers are related to the size of their colony.Received 23 March 2004; revised 3 June 2004; accepted 4 June 2004.     

Can alloethism in workers of the bumblebee, Bombus terrestris, be explained in terms of foraging efficiency?

Bumblebee workers vary greatly in size, unlike workers of most other social bees. This variability has not been adequately explained. In many social insects, size variation is adaptive, with different-sized workers performing different tasks (alloethism). Here we established whether workers of the bumblebee, Bombus terrestris (L.) (Hymenoptera; Apidae), exhibit alloethism. We quantified the size of workers engaging in foraging compared to those that remain in the nest, and confirmed that it is the larger bees that tend to forage (X±SE thorax widths 4.34±0.01 mm for nest bees and 4.93±0.02 mm for foragers). We then investigated whether large bees are better suited to foraging because they are able to transport heavier loads of food back to the nest. Both pollen and nectar loads of returning foragers were measured, demonstrating that larger bees do return with a heavier mass of forage. Foraging trip times were inversely related to bee size when collecting nectar, but were unrelated to bee size for bees collecting pollen. Overall, large bees brought back more nectar per unit time than small bees, but the rate of pollen collection appeared to be unrelated to size. The smallest foragers had a nectar foraging rate close to zero, presumably explaining why foragers tend to be large. Why might larger bees be better at foraging? Various explanations are considered: larger bees are able to forage in cooler conditions, may be able to forage over larger distances, and are perhaps also less vulnerable to predation. Conversely, small workers are presumably cheaper to produce and may be more nimble at within-nest tasks. Further research is needed to assess these possibilities. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Information needs at the beginning of foraging: grass-cutting ants trade off load size for a faster return to the nest

Background

Acquisition of information about food sources is essential for animals that forage collectively like social insects. Foragers deliver two commodities to the nest, food and information, and they may favor the delivery of one at the expenses of the other. We predict that information needs should be particularly high at the beginning of foraging: the decision to return faster to the nest will motivate a grass-cutting ant worker to reduce its loading time, and so to leave the source with a partial load.

Principal Findings

Field results showed that at the initial foraging phase, most grass-cutting ant foragers (Acromyrmex heyeri) returned unladen to the nest, and experienced head-on encounters with outgoing workers. Ant encounters were not simply collisions in a probabilistic sense: outgoing workers contacted in average 70% of the returning foragers at the initial foraging phase, and only 20% at the established phase. At the initial foraging phase, workers cut fragments that were shorter, narrower, lighter and tenderer than those harvested at the established one. Foragers walked at the initial phase significantly faster than expected for the observed temperatures, yet not at the established phase. Moreover, when controlling for differences in the fragment-size carried, workers still walked faster at the initial phase. Despite the higher speed, their individual transport rate of vegetable tissue was lower than that of similarly-sized workers foraging later at the same patch.

Conclusions/Significance

At the initial foraging phase, workers compromised their individual transport rates of material in order to return faster to the colony. We suggest that the observed flexible cutting rules and the selection of partial loads at the beginning of foraging are driven by the need of information transfer, crucial for the establishment and maintenance of a foraging process to monopolize a discovered resource.     

The allocation of foragers in red wood ants

Abstract. 1. We studied how colonies of the red wood ant, Formica polyctena , adjust the numbers of foragers allocated to different foraging trails. In a series of field experiments, foragers were marked and transferred from one nest to another, related nest, where they joined the foraging force. Transferred workers acted as a reserve of uncommitted, available foragers.
2. Previous work shows that each individual forager habitually uses one trail. We found that for an uncommitted forager, the influence of recruitment initially is stronger than that of directional fidelity. Transferred workers were likely to use trails leading to new food sources. When transferred to a new nest, foragers were not likely to use a trail in the same direction as their original trail in the donor nest.
3. After a week, transferred foragers tended to develop route fidelity. Even after bait was no longer present, they continued to use the trail that had formerly led to a bait source.
4. We examined how colonies adjust numbers on a trail by experimentally depleting some trails. Colonies usually did not compensate for depletion: foragers were not recruited to depleted trails.
5. In general, the dynamics of foraging in this species facilitate a consistent foraging effort rather than rapid adjustments of forager allocation.     

Colony‐level behavioural variation correlates with differences in expression of the foraging gene in red imported fire ants

Among social insects, colony‐level variation is likely to be widespread and has significant ecological consequences. Very few studies, however, have documented how genetic factors relate to behaviour at the colony level. Differences in expression of the foraging gene have been associated with differences in foraging and activity of a wide variety of organisms. We quantified expression of the red imported fire ant foraging gene (sifor) in workers from 21 colonies collected across the natural range of Texas fire ant populations, but maintained under standardized, environmentally controlled conditions. Colonies varied significantly in their behaviour. The most active colonies had up to 10 times more active foragers than the least active colony and more than 16 times as many workers outside the nest. Expression differences among colonies correlated with this colony‐level behavioural variation. Colonies with higher sifor expression in foragers had, on average, significantly higher foraging activity, exploratory activity and recruitment to nectar than colonies with lower expression. Expression of sifor was also strongly correlated with worker task (foraging vs. working in the interior of the nest). These results provide insight into the genetic and physiological processes underlying collective differences in social behaviour. Quantifying variation in expression of the foraging gene may provide an important tool for understanding and predicting the ecological consequences of colony‐level behavioural variation.     

An exploration of the relationship between recruitment communication and foraging in stingless bees

Social information is widely used in the animal kingdom and can be highly adaptive. In social insects, foragers can use social information to find food, avoid danger, or choose a new nest site. Copying others allows individuals to obtain information without having to sample the environment. When foragers communicate information they will often only advertise high-quality food sources, thereby filtering out less adaptive information. Stingless bees, a large pantropical group of highly eusocial bees, face intense inter- and intra-specific competition for limited resources, yet display disparate foraging strategies. Within the same environment there are species that communicate the location of food resources to nest-mates and species that do not. Our current understanding of why some species communicate foraging sites while others do not is limited. Studying freely foraging colonies of several co-existing stingless bee species in Brazil, we investigated if recruitment to specific food locations is linked to 1) the sugar content of forage, 2) the duration of foraging trips, and 3) the variation in activity of a colony from 1 day to another and the variation in activity in a species over a day. We found that, contrary to our expectations, species with recruitment communication did not return with higher quality forage than species that do not recruit nestmates. Furthermore, foragers from recruiting species did not have shorter foraging trip durations than those from weakly recruiting species. Given the intense inter- and intraspecific competition for resources in these environments, it may be that recruiting species favor food resources that can be monopolized by the colony rather than food sources that offer high-quality rewards.     

Foraging tempo in two woodland and species

Running speeds of Myrmica punctiventris and Aphaenogaster rudis workers were measured, and a good correspondence between laboratory and field behaviour was obtained. In the laboratory, foraging tempo and foraging efficiency were calculated for two colony sizes and five patterns of prey availability. Running speeds were strongly dependent on colony size for both species; when retrieving prey, foragers from large colonies ran significantly faster than those from small colonies. In addition, ants searching for prey ran more slowly than those returning to the nest with prey. Efficiency, measured as the propensity to return to the nest in a straight line, was most strongly a function of distance from the nest. Finally, no relationship between an ant's speed and its efficiency of return was found.     

Genetic diversity within honeybee colonies increases signal production by waggle-dancing foragers

Recent work has demonstrated considerable benefits of intracolonial genetic diversity for the productivity of honeybee colonies: single-patriline colonies have depressed foraging rates, smaller food stores and slower weight gain relative to multiple-patriline colonies. We explored whether differences in the use of foraging-related communication behaviour (waggle dances and shaking signals) underlie differences in foraging effort of genetically diverse and genetically uniform colonies. We created three pairs of colonies; each pair had one colony headed by a multiply mated queen (inseminated by 15 drones) and one colony headed by a singly mated queen. For each pair, we monitored the production of foraging-related signals over the course of 3 days. Foragers in genetically diverse colonies had substantially more information available to them about food resources than foragers in uniform colonies. On average, in genetically diverse colonies compared with genetically uniform colonies, 36% more waggle dances were identified daily, dancers performed 62% more waggle runs per dance, foragers reported food discoveries that were farther from the nest and 91% more shaking signals were exchanged among workers each morning prior to foraging. Extreme polyandry by honeybee queens enhances the production of worker-worker communication signals that facilitate the swift discovery and exploitation of food resources.     

The size–distance relationship in the wood ant Formica rufa

1. The size–distance relationship among honeydew‐collecting foragers of the red wood ant Formica rufa was investigated. Within the colony territory, the size (as measured by head width) and fresh weight of samples of foragers were determined for ants ascending and descending trees near, and farther from, the central nest mound. 2. The mean size of the ants was significantly higher at far trees than at near trees in six out of the seven colonies investigated, confirming the general presence of the size–distance relationship. 3. In three colonies, a load–distance relationship was also found. For a given head width, honeydew‐carrying ants descending far trees were significantly heavier than those descending near trees (i.e. they were carrying heavier loads from trees farther away from the central nest mound). 4. This is the first time that both load–distance and size–distance relationships have been reported in foraging workers from the same ant colony. 5. The combined effects of these characteristics suggest that colony foraging efficiency is enhanced by far trees being visited by the larger workers that then return with heavier loads of honeydew.     

Nest demographics and foraging behavior of <Emphasis Type="Italic">Apterostigma collare</Emphasis> Emery (Hymenoptera,Formicidae) provide evidence of colony independence

Apterostigma collare Emery is a highly derived fungus-growing ant within the Tribe Attini whose small, fungal nests are found in tropical rain forests. This study focuses on determining the colony structure of A. collare, specifically searching for evidence of polydomy or independence. We surveyed and observed nests in the field, and performed foraging bioassays and dissected nests in the laboratory. We determined the size and contents of nests in field populations. Nests found near other nests were not statistically different in size compared to nests found alone. There was also no statistical difference between near and lone nests regarding the presence of a queen in the nest. Most nests contained one queen with brood and workers, regardless of their proximity to other nests. Observations also were made of foraging and trail-marking behaviors. Foraging activity observed in the field revealed that workers left the nest area and followed trails upwards into the canopy, but they did not interact with foragers from other nearby nests. In a laboratory foraging arena, foragers marked a trail to a food source by dragging the gaster. Bioassays showed that A. collare workers preferred their own foraging trails, but not those of other conspecific colonies. All results suggest that each nest represents an independent colony, supporting a previous report that nests found in close proximity do not constitute a polydomous colony. Received 19 July 2006; revised 23 March 2007; accepted 6 June 2007.