The role of internal and external information in foraging decisions of Melipona workers (Hymenoptera: Meliponinae)

Social insect foragers have to make foraging decisions based on information that may come from two different sources: information learned and memorised through their own experience (“internal” information) and information communicated by nest mates or directly obtained from their environment (“external” information). The role of these sources of information in decision-making by foragers was studied observationally and experimentally in stingless bees of the genus Melipona. Once a Melipona forager had started its food-collecting career, its decisions to initiate, continue or stop its daily collecting activity were mainly based upon previous experience (activity on previous days, the time at which foraging was initiated the day(s) before, and, during the day, the success of the last foraging flights) and mediated through direct interaction with the food source (load size harvested and time to collect a load). External information provided by returning foragers advanced the start of foraging of experienced bees. Most inexperienced bees initiated their foraging day after successful foragers had returned to the hive. The start of foraging by other inexperienced bees was stimulated by high waste-removal activity of nest mates. By experimentally controlling the entries of foragers (hence external information input) it was shown that very low levels of external information input had large effect on the departure of experienced foragers. After the return of a single successful forager, or five foragers together, the rate of forager exits increased dramatically for 15 min. Only the first and second entry events had large effect; later entries influenced forager exit patterns only slightly. The results show that Melipona foragers make decisions based upon their own experience and that communication stimulates these foragers if it concerns the previously visited source. We discuss the organisation of individual foraging in Melipona and Apis mellifera and are led to the conclusion that these species behave very similarly and that an information-integration model (derived from Fig. 1) could be a starting point for future research on social insect foraging. Received: 16 April 1997 / Accepted after revision: 30 August 1997     

Does a polyandrous honeybee queen improve through patriline diversity the activity of her colony’s scouting foragers?

Recent studies indicate that the foraging success of a honeybee colony is enhanced when it has numerous genetically diverse patrilines because of queen polyandry. We determined whether foraging is improved in part because patriline diversity generates more responsive populations of scouting foragers. Scouts search for new food sources and advertise them with waggle dances to inform other foragers about unexploited discoveries. We moved multiple-patriline and single-patriline colonies to unfamiliar locations so that colonies relied heavily on successful scouts to initiate recruitment and then compared the development of foraging effort between the two types of colonies. More waggle dance signals were produced during the incipient stages of foraging in multiple-patriline colonies compared to single-patriline colonies because scouts reported food discoveries with longer dances. Scouts also returned to multiple-patriline colonies at rates that were two thirds higher than those of single-patriline colonies, although return rates for general forager populations were not significantly different between colony types. The distance of reported food sources from hives increased with time for all colonies, but by the end of their first day in an unfamiliar environment, maximal foraging reach was greater if colonies had multiple patrilines. Most scouts in multiple-patriline colonies came from a minority of scout-rich patrilines that were generally not those from which general forager populations were derived; the presence of such scout-rich patrilines was correlated with the extent of recruitment signaling in colonies. We show how a honeybee colony’s scouting effort is (and is not) enhanced when extremely polyandrous queens produce genetically diverse colonies.     

Dominance and polyethism in the eusocial wasp Mischocyttarus mastigophorus (Hymenoptera: Vespidae)

Dominance interactions affected patterns of non-reproductive division of labor (polyethism) in the eusocial wasp Mischocyttarus mastigophorus. Socially dominant individuals foraged for food (nectar and insect prey) at lower rates than subordinate individuals. In contrast, dominant wasps performed most of the foraging for the wood pulp used in nest construction. Social dominance also affected partitioning of materials collected by foragers when they returned to the nest. Wood pulp loads were never shared with nest mates, while food loads, especially insect prey, were often partitioned with other wasps. Dominant individuals on the nest were more likely to take food from arriving foragers than subordinate individuals. The role of dominance interactions in regulating polyethism has evolved in the eusocial paper wasps (Polistinae). Both specialization by foragers and task partitioning have increased from basal genera (independent-founding wasps, including Mischo-cyttarus spp.) to more derived genera (swarm-founding Epiponini). Dominance interactions do not regulate forager specialization or task partitioning in epiponines. I hypothesize that these changes in polyethism were enabled by the evolution of increased colony size in the Epiponini. Received: 8 December 1997 / Accepted after revision: 28 March 1998     

Colony energy requirements affect the foraging currency of bumble bees

Summary This study examines whether the foraging behavior of worker bumble bees (Bombus: Apidae) collecting nectar on inflorescences of seablush (Plectritis congesta: Valerianaceae) is affected by colony energetic requirements, which were experimentally manipulated either by adding sucrose solution to honey pots or by removing virtually all available nectar from the pots. The competing hypotheses tested were: (1) no change; energetic requirements do not affect behavior, since there is a single best way to collect food in a given environment; (2) energetic currency; the energetic currency maximized by foragers changes according to colony energetic condition, with nectar-depletion causing a shift from maximizing long-term productivity to maximizing immediate energetic gain, thereby de-emphasizing energetic costs; and (3) predation; foragers devalue risk of predation as risk of starvation increaes, with colony nectar-depletion causing foragers to be less predation riskaverse in order to increase immediate energetic gain. Relative to when their colony energy reserves were enhanced, foragers from nectar-depleted colonies selected smaller inflorescences, visited fewer flowers per inflorescence, probed flowers at a higher rate while on each inflorescence, and walked between inflorescences less often, thereby spending a greater proportion of their foraging trip in flight. These behaviors increased a bee's energetic costs while foraging, and should also have increased its immediate energetic gains, allowing rejection of the no change hypothesis. Predictions of the predation hypothesis were generally not supported, and our results best support the energetic currency hypothesis. Foraging currency of bumble bees therefore appears to be a function of colony energetic state. Offprint requests to: R.V. Cartar     

Division of foraging labor in ants can mediate demands for food and safety

Solitary foragers can balance demands for food and safety by varying their relative use of foraging patches and their level of vigilance. Here, we investigate whether colonies of the ant, Formica perpilosa, can balance these demands by dividing labor among workers. We show that foragers collecting nectar in vegetation near their nest are smaller than are those collecting nectar at sites away from the nest. We then use performance tests to show that smaller workers are more likely to succumb to attack from conspecifics but feed on nectar more efficiently than larger workers, suggesting a size-related trade-off between risk susceptibility and harvesting ability. Because foragers that travel away from the nest are probably more likely to encounter ants from neighboring colonies, this trade-off could explain the benefits of dividing foraging labor among workers. In a laboratory experiment, we show that contact with aggressive workers results in an increase in the mean size of recruits to a foraging site: this increase was not the result of more large recruits, but rather because fewer smaller ants traveled to the site. These results suggest that workers particularly susceptible to risk avoid dangerous sites, and suggest that variation in worker size can allow colonies to exploit profitably both hazardous and resource-poor patches.Communicated by L. Sundström     

The use of waggle dance information by honey bees throughout their foraging careers

We studied the extent to which worker honey bees acquire information from waggle dances throughout their careers as foragers. Small groups of foragers were monitored from time of orientation flights to time of death and all in-hive behaviors relating to foraging were recorded. In the context of a novice forager finding her first food source, 60% of the bees relied, at least in part, on acquiring information from waggle dances (being recruited) rather than searching independently (scouting). In the context of an experienced forager whose foraging has been interrupted, 37% of the time the bees resumed foraging by following waggle dances (being reactivated) rather than examining the food source on their own (inspecting). And in the context of an experienced forager engaged in foraging, 17% of the time the bees initiated a foraging trip by following a waggle dance. Such dance following was observed much more often after an unsuccessful than after a successful foraging trip. Successful foragers often followed dances just briefly, perhaps to confirm that the kind of flowers they had been visiting were still yielding forage. Overall, waggle dance following for food discovery accounted for 12–25% of all interactions with dancers (9% by novice foragers and 3–16% by experienced foragers) whereas dance following for reactivation and confirmation accounted for the other 75–88% (26% for reactivation and 49–62% for confirmation). We conclude that foragers make extensive use of the waggle dance not only to start work at new, unfamiliar food sources but also to resume work at old, familiar food sources.     

Individually different foraging methods in the desert ant Cataglyphis bicolor (Hymenoptera,Formicidae)

Summary Observations and field experiments on the foraging behaviour of individual workers of Cataglyphis bicolor in a Southern Tunisian shrub desert are reported. The workers search singly for their food (mostly animal carcasses) and are singleprey loaders. The individuals differ to a great extent in their persistence to re-search the place of a find on a previous foraging excursion. The differences range continuously from thoroughly researching a place to just walking by. If, in an experiment, the same reward is offered farther from the nest, each ant persists more in re-searching the place than if food is offered close to the nest. In a further experiment, some individuals persisted less in searching near the former finding site if they had collected a fly than after collecting a piece of cheese. There is, however, evidence that individuals do not differ in their food preference. Persistent individuals, which re-search the place of a former find, are faster than non-persistent ones in retrieving food that is experimentally arranged in an aggregated manner. The experiment failed to demonstrate the (reverse) superiority of non-persistent individuals foraging on homogeneously distributed food. The observations of unmanipulated foraging excursions in the field suggest such an advantage for non-persistent foragers under natural conditions where food in general occurs widely dispersed. The colony as a whole retrieves more food within the same time from an experimental lay-out that is homogeneous than from an aggregated one. The behavioural differences between individuals could be caused by a training bias of the short-lived foragers, leading to a different assessment of the profitability of a searching method which implies returning to a formerly rewarding place. Thus, each worker uses the most promising behaviour according to its individual experience. Alternatively, the individually different searching methods could mainly contribute to the welfare of the colony as a whole rather than leading to a maximal short-term efficiency of each individual. In particular, the colony, disposing of only a few highly persistent foragers, could quickly exploit occasional short-lived, but unpredictible, clumps of food within its foraging range.     

Foraging in the seed-harvester ant genus Pogonomyrmex: are energy costs important?

Energy intake and expenditure on natural foraging trips were estimated for the seed-harvester ants, Pogonomyrmex maricopa and P. rugosus. During seed collection, P. maricopa foraged individually, whereas P. rugosus employed a trunk-trail foraging system. Energy gain per trip and per minute were not significantly different between species. There was also no interspecific difference in energy cost per trip, but energy cost per minute was lower for P. maricopa foragers because they spent on average 7 min longer searching for a load on each trip. Including both unsuccessful and successful foraging trips, average energy gain per trip was more than 100 times the energy cost per trip for both species. Based on this result, we suggest that time cost incurred during individual foraging trips is much more important than energy cost in terms of maximizing net resource intake over time. In addition, because energy costs are so small relative to gains, we propose that energy costs associated with foraging may be safely ignored in future tests of foraging theory with seed-harvesting ant species.     

Vocal mediation of foraging competition in the cooperatively breeding green woodhoopoe (<Emphasis Type="Italic">Phoeniculus purpureus</Emphasis>)

Many group-living species produce frequent vocalisations when foraging, but the function of these food-associated calls is often difficult to divine. I investigated the kek call of the cooperatively breeding green woodhoopoe (Phoeniculus purpureus), a species in which individuals have preferred foraging techniques dependent on their bill size. Individuals called at a greater rate (1) in foraging compared to non-foraging situations, and (2) in groups containing potential foraging competitors (i.e. individuals that foraged using the same preferred techniques). I therefore asked whether the kek call is used to recruit conspecific foragers or whether it acts as a vocal signal of foraging niche and mediates foraging competition. Foragers that were vocalising were no more likely to be approached than those that were silent, and individuals gained no foraging advantage from the close proximity of another group member. Thus, keks are unlikely to be used to recruit conspecifics. Instead, they appear to regulate spacing between potential foraging competitors. Although an individual forager was equally likely to be closely approached by all other group members, it increased its calling rate only in response to potential foraging competitors. This increase in calling rate resulted in the approaching individual moving away, thus maintaining some separation between individuals that forage in the same way. Maintenance of such spacing is important because the success rate of an individual decreased when a foraging competitor was close by.Communicated by M. Leonard     

Global optimization from suboptimal parts: foraging sensu lato by leaf-cutting ants

Central-place foraging theory has been unable to explain the load selection behavior of leaf-cutting ants (Atta spp., Attini: Formicidae). We suggest that this is due to incomplete consideration of the sequence of behaviors involved in resource acquisition by these ants. Unlike most central-place foragers, leaf-cutting ants do not return to their nests with food. Instead, the leaf fragments they gather must be processed within the nest to convert them to substrate for fungal gardens. We have shown previously that leaf fragment size affects the rate of distribution and processing of leaf tissue inside laboratory nests of Atta colombica. Including these tasks in the calculation of foraging rate may help explain load selection and other features of central-place foraging by Atta colonies. Here we develop a mathematical model of the complete sequence of external and internal tasks that lead to addition of substrate to fungal gardens. Using realistic parameter values, the leaf fragment sizes predicted to maximize a colony's rate of foraging in this broad sense correspond well with the mean fragment sizes actually collected by Atta colonies in the field. The optimal fragment size for global performance in the model is below the size that would maximize the delivery rate by above-ground foragers. The globally optimal size also fails to maximize the rate of either fragment distribution or fragment processing within the nest. Our results show how maximum collective performance of an ensemble of linked tasks may require behavior that would appear suboptimal in a piecemeal analysis of tasks.     

Exploitation of carbohydrate food sources in <Emphasis Type="Italic">Polybia occidentalis</Emphasis>: social cues influence foraging decisions in swarm-founding wasps

An efficient exploitation of carbohydrate food sources would be beneficial for social wasp species that store nectar within their nest. In the swarm-founding polistine wasp Polybia occidentalis, we now demonstrate that the decisions of when and where to forage are influenced by information from conspecifics. Only when foragers had been trained to collect at artificial carbohydrate feeders did newcomers (food-source-naive individuals) continuously arrive at these feeders during 2 h of experiment. In control tests, in which no forager had been trained, not a single newcomer alighted at any of the offered carbohydrate food sources. This indicates that, during the foraging process, a nest-based input provided by successful foragers must have stimulated nestmates to search for food. Once activated, the newcomers’ choice on where to collect was strongly influenced by field-based social information. The mere visual presence of accumulated conspecifics (wasp dummies placed on one of the feeders) attracted newcomers to the food sources. Interestingly, however, visual enhancement was not the only decision-biasing factor at the feeding site. In an experimental series where searching wasps had to choose between the experimental feeder at which 3 foragers continuously collected and the control feeder with nine wasp dummies, only 40% of the wasps chose the visually enhanced feeder. This points to the existence of additional mechanisms of local enhancement. The possibility that, in social wasps, recruitment is involved in the exploitation of carbohydrate food sources is discussed.     

The regulation of pollen foraging by honey bees: how foragers assess the colony's need for pollen

Summary The honey bee colony presents a challenging paradox. Like an organism, it functions as a coherent unit, carefully regulating its internal milieu. But the colony consists of thousands of loosely assembled individuals each functioning rather autonomously. How, then, does the colony acquire the necessary information to organize its work force? And how do individuals acquire information about specific colony needs, and thus know what tasks need be performed? I address these questions through experiments that analyze how honey bees acquire information about the colony's need for pollen and how they regulate its collection. The results demonstrate features of the colony's system for regulating pollen foraging: (1) Pollen foragers quickly acquire new information about the colony's need for pollen. (2) When colony pollen stores are supplemented, many pollen foragers respond by switching to nectar foraging or by remaining in the hive and ceasing to forage at all. (3) Pollen foragers do not need direct contact with pollen to sense the colony's change of state, nor do they use the odor of pollen as a cue to assess the colony's need for pollen. (4) Pollen foragers appear to obtain their information about colony pollen need indirectly from other bees in the hive. (5) The information takes the form of an inhibitory cue. The proposed mechanism for the regulation of pollen foraging involves a hierarchical system of information acquisition and a negative feedback loop. By taking advantage of the vast processing capacity of large numbers of individuals working in parallel, such a system of information acquisition and dissemination may be ideally suited to promote efficient regulation of labor within the colony. Although each individual relies on only limited, local information, the colony as a whole achieves a finely-tuned response to the changing conditions it experiences.     

Division of labor between scouts and recruits: genetic influence and mechanisms

Every recruitment system in social insects requires some individuals that serve as scouts, foragers that search independently for food sources. It is not well understood which factors influence whether an individual becomes a scout or a recruit, nor how the division of labor between the two forager groups is regulated. It is shown here for honeybees (Apis mellifera), using two different molecular techniques, that there is a genetically based difference in the probability that individuals will scout independently for food. In contrast to earlier suggestions, experimental tests showed that the age of a bee does not seem to influence its probability of becoming a scout or a recruit. Furthermore, scout bees do not search opportunistically for either pollen or nectar but, rather, individuals have preferences that are genetically based. These findings are discussed in the framework of foraging regulation by specialization in honeybees and the adaptive significance of polyandry. Received: 23 October 1997 / Accepted after revision: 10 April 1998     

Brood pheromone stimulates pollen foraging in honey bees (Apis mellifera)

Foraging and the mechanisms that regulate the quantity of food collected are important evolutionary and ecological attributes for all organisms. The decision to collect pollen by honey bee foragers depends on the number of larvae (brood), amount of stored pollen in the colony, as well as forager genotype and available resources in the environment. Here we describe how brood pheromone (whole hexane extracts of larvae) influenced honey bee pollen foraging and test the predictions of two foraging-regulation hypotheses: the indirect or brood-food mechanism and the direct mechanism of pollen-foraging regulation. Hexane extracts of larvae containing brood pheromone stimulated pollen foraging. Colonies were provided with extracts of 1000 larvae (brood pheromone), 1000 larvae (brood), or no brood or pheromone. Colonies with brood pheromone and brood had similar numbers of pollen foragers, while those colonies without brood or pheromone had significantly fewer pollen foragers. The number of pollen foragers increased more than 2.5-fold when colonies were provided with extracts of 2000 larvae as a supplement to the 1000 larvae they already had. Within 1 h of presenting colonies with brood pheromone, pollen foragers responded to the stimulus. The results from this study demonstrate some important aspects of pollen foraging in honey bee colonies: (1) pollen foragers appear to be directly affected by brood pheromone, (2) pollen foraging can be stimulated with brood pheromone in colonies provided with pollen but no larvae, and (3) pollen forager numbers increase with brood pheromone as a supplement to brood without increasing the number of larvae in the colony. These results support the direct-stimulus hypothesis for pollen foraging and do not support the indirect-inhibitor, brood-food hypothesis for pollen-foraging regulation. Received: 5 March 1998 / Accepted after revision: 29 August 1998     

Determinants of multiple central-place territory use in wild young-of-the-year Atlantic salmon (<Emphasis Type="Italic">Salmo salar</Emphasis>)

Patterns of space use provide key insights into how animals exploit local resources and are linked to both the fitness and distribution of individuals. We studied territory size, mobility, and foraging behavior of young-of-the-year Atlantic salmon Salmo salar in relation to several key environmental factors in Catamaran Brook, New Brunswick, Canada. The 50 study fish were all multiple central-place foragers (i.e., alternated among several sit-and-wait foraging stations) and showed great variability in territory size and the total distance traveled within the territories. Territory size increased with the mean distance traveled between consecutive foraging stations, the number of stations visited, and the mean foraging radius. Fish also varied greatly in how much of the total travel distance was associated with foraging at a station (14.8–91.8%) versus switching among stations (4.6–84.3%). As predicted, fish in slow-flowing waters, where drifting prey were scarce, used larger multiple central-place territories than individuals in faster, more productive waters. Interestingly, however, the most mobile fish did not inhabit slow-running waters as predicted but were found at intermediate (optimal) water current velocities. Hence, our study suggests that among some multiple central-place foragers, increased mobility may not only serve to increase prey encounter rate but may reflect an attempt to patrol territories in favorable habitats. Further studies are needed to determine the generality and the ultimate benefits of multiple central-place space use among stream-dwelling fish and other animals.     

Avian flocking reduces starvation risk: an experimental demonstration

Summary Theory suggests that variance in individual food intake is lower during group foraging. Consequently, group foraging can at times reduce starvation risk. In aviary experiments using green-finches we demonstrate how intake variability decreases during group foraging because individuals use feeding by flock mates as a cue to locate food (local enhancement). Flocking preferences of greenfinches responded to variance in energy gain as predicted by theoretical models for foragers attempting to reduce starvation risk. While energy budget was positive the greenfinches were risk averse and foraged socially. Their preference shifted towards more risk prone solitary foraging when kept on a negative energy budget. We conclude that time or energy net gains are not necessary for foraging groups to form, but reductions in starvation risk may be sufficient.     

Social context influences cue-mediated recruitment in an invasive social wasp

Social insects often serve as model systems for communication and recruitment studies, and yet, it remains controversial whether social vespid wasps can reliably communicate resource information to nestmates. In this study, I present empirical evidence that foraging strategies depend on the initial assessment of resource size and potential competition by foraging yellowjackets. The context dependent foraging behavior of Vespula pensylvanica provides a potential explanation for the inconsistent reports of the existence of recruitment communication in vespid wasps. Furthermore, life history traits may influence yellowjacket foraging behavior; annual V. pensylvanica colonies, whose foragers routinely patrol near the nest, exhibited increased bait visitation in response to the return of successful foragers, whereas perennial colonies did not. These behavioral disparities provide insight into how foraging strategies and search patterns may shift with colony size and longevity. In experiments that investigate the effects of visual cues of conspecifics and bait dispersion, foraging decisions corresponded with expectations of yellowjackets integrating resource quantity and access into a perception of demand. When resource competition could be assessed as high, V. pensylvanica foragers quickly exploited the bait closest to their colony regardless of occupation by other wasps; however, foragers preferred visiting unoccupied baits in situations where competition could be perceived as low. Moreover, a meta-analysis revealed that context-dependent, cue-mediated recruitment was widespread in Vespidae, where such foraging behaviors changed with habitat and the potential for resource competition. Such plastic foraging strategies may contribute to the invasion success of some vespid wasps.     

Locomotor impairment and defense in gravid lizards (Eumeces laticeps): behavioral shift in activity may offset costs of reproduction in an active forager

Summary Female Eumeces laticeps experience a substantial decrease in running speed (ca. 25%) and an even greater loss of endurance (slightly over 50%) while gravid. Because some widely foraging lizards, including E. laticeps, rely primarily on running to escape predators, the decreases in speed and stamina may contribute to an increased risk of predation. However, observations suggest that gravid females become less active or conspicuous on the surface. Ambush foraging lizards rely relatively more on crypsis associated with immobility to avoid predation and thus can have greater average relative clutch mass (RCM) than active foragers. Behavioral compensation for locomotor impairment by becoming less active or conspicuous may allow some species the advantages inherent in both high relative clutch mass when gravid and the increased energetic profitability of active foraging when not gravid. As females gain weight during the breeding season, they may forage actively until the risk due to increasing locomotor impairment becomes too great and then change defensive strategy to greater reliance on crypsis. Without such a shift, widely foraging squamate reptiles may be less able than ambush foragers to exploit life-historical strategies demanding high current investment in reproduction.     

Potential mechanisms for the communication of height and distance by a stingless bee, Melipona panamica

This study investigates the recruitment communication mechanisms of a stingless bee, Melipona panamica, whose foragers can evidently communicate the three-dimensional location of a good food source. To determine if the bees communicate location information inside or outside the nest, we conducted removal experiments by training marked foragers to one of two identical feeders and then separating these experienced foragers from potential recruits as they left the nest. The feeders were positioned to test the communication of each dimension. The results show that recruits do not simply follow experienced foragers to the food source. Height and distance are communicated within the nest, while direction is communicated outside the nest. We then examined the pulsed sounds produced by recruiting foragers. While unloading food, recruiting foragers produced several short pulses and one or more very long pulses. On average, the longest unloading pulse per performance was 31–50% longer (P ≤ 0.018) for bees foraging on the forest floor than for bees foraging at the top of the forest canopy (40 m high). While dancing, recruiting foragers produced sound pulses whose duration was positively correlated with the distance to the food source (P < 0.001). Dancing recruiters also produced several short sound pulses followed by one or more long pulses. The longest dance pulse per performance was 291 ± 194 ms for a feeder 25 m from the nest and 1858 ± 923 ms for a feeder 360 m away from the nest. The mechanism of directional communication remains a mystery. However, the direction removal experiment demonstrates that newcomers cannot use forager-deposited scent marks for long-distance orientation (>100 m from the nest). Received: 25 September 1997 / Accepted after revision: 31 May 1998     

The organization of work in Polybia occidentalis: costs and benefits of specialization in a social wasp

Summary Nest construction, a complex social activity requiring the coordination of 3 tasks (Fig.2), was compared in large (<350 adults) and small (<50 adults) colonies of Polybia occidentalis. The 3 tasks—water foraging, pulp foraging, and building—are performed by 3 separate groups of workers (Fig.4). Of the 8 acts comprising the 3 tasks, 5 regularly involve the transfer of water or pulp from one worker to another on the nest.Small colonies required nearly twice as long (35.4 worker-min) as large colonies (20.1 workermin) to complete a unit amount of construction work. Behavioral acts involving material transfer among workers were responsible for most of the increase in small colonies. In other words, the waiting times experienced by material donors and recipients were greater in small colonies. In small colonies workers switched among the three tasks more frequently than in large colonies (Fig. 4). This was the result of more frequent switching by generalists (workers that performed 2 or 3 of the tasks), rather than by a decrease in the proportion of specialists (workers performing only 1 task type) (Fig. 3).The series-parallel system by which Polybia occidentalis organizes nest construction has a major advantage over the series operation of solitary wasps. Pulp foragers collect and carry loads that are 6.1 times as large as builders can work with at the nest, and water foragers bring in loads that appear to be limited only by crop capacity and that provide all the moisture necessary for the complete processing of 0.74 of a foraged pulp load. As a result P. occidentalis can collect and process a given amount of nest material using 2.6 times fewer foraging trips than would be required by the series system. This in turn means that P. occidentalis not only achieves an energy saving that probably more than offsets the increased costs of material handling at the nest, but it reduces the exposure of its foragers to predators in the field.