The costs and benefits of cooperation between the Australian lycaenid butterfly,Jalmenus evagoras,and its attendant ants

Summary The larvae and pupae of the Australian lycaenid butterfly, Jalmenus evagoras associate mutualistically with ants in the genus Iridomyrmex. Four ant exclusion experiments in three field sites demonstrated that predation and parasitism of J. evagoras are so intense that individuals deprived of their attendant ants are unlikely to survive. Larvae and pupae of J. evagoras aggregate, and the mean number of attendant ants per individual increases with larval age and decreases with group size. Field observations showed that young larvae could gain more attendant ants per individual by joining the average size group of about 4 larvae than by foraging alone. Aggregation behaviour is influenced by ant attendance: young larvae and pupating fifth instars aggregated significantly more often on plants with ants than on plants where ants had been excluded. In return for tending and protecting the larvae, ants were rewarded by food secretions that can amount to as much as 409 mg dry biomass from a single host plant containing 62 larvae and pupae of J. evagoras over a 24 h period. Larval development in the laboratory lasted approximately a month, and larvae that were tended by ants developed almost 5 days faster than larvae that were not tended. However, tended individuals, particularly females, pupated at a significantly lower weight than their untended counterparts, and the adults that eclosed from these pupae were also lighter and smaller. On average, pupae that were tended by ants lost 25% more weight than untended pupae, and in contrast with larvae, they took longer to eclose than pupae that were not tended. These experimental results are discussed in terms of costs and benefits of association for both partners, and of aggregation for the lycaenids.     

Self-organization in collective honeybee foraging: emergence of symmetry breaking,cross inhibition and equal harvest-rate distribution

De Vries and Biesmeijer described in 1998 an individual-oriented model that simulates the collective foraging behaviour of a colony of honeybees. Here we report how this model has been expanded and show how, through self-organization, three colony-level phenomena can emerge: symmetry breaking, cross inhibition and the equal harvest-rate distribution. Symmetry breaking is the phenomenon that the numbers of foragers visiting two equally profitable food sources will diverge after some time. Cross inhibition is the phenomenon that, by increasing the profitability of one of two equal food sources, the number of foragers visiting the other source will decrease. In some circumstances, the bees foraging on two sources of different profitabilities will be distributed between these sources such that the two average energy harvest rates are equal. We will refer to this phenomenon as the equal harvest-rate distribution. For each of these three phenomena, we show what the necessary behavioural rules to be followed by the individual forager bees are, and what the necessary circumstances are (that is, what values the model parameters should take) in order for these phenomena to arise. It seems that patch size and forager group size largely determine when each of these phenomena will arise. Experimenting with two types of currency, net gain rate and net gain efficiency, revealed that only gain rate may result in an equal harvest-rate distribution of foragers visiting different food sources.     

Regulation of pollen foraging in honeybee colonies: effects of young brood, stored pollen, and empty space

Pollen storage in a colony of Apis mellifera is actively regulated by increasing and decreasing pollen foraging according to the “colony's needs.” It has been shown that nectar foragers indirectly gather information about the nectar supply of the colony from nestmates without estimating the amount of honey actually stored in the combs. Very little is known about how the actual colony need is perceived with respect to pollen foraging. Two factors influence the need for pollen: the quantity of pollen stored in cells and the amount of brood. To elucidate the mechanisms of perception, we changed the environment within normal-sized colonies by adding pollen or young brood and measured the pollen-foraging activity, while foragers had either direct access to them or not. Our results show that the amount of stored pollen, young brood, and empty space directly provide important stimuli that affect foraging behavior. Different mechanisms for forager perception of the change in the environment are discussed. Received: 13 June 1998 / Accepted after revision: 25 October 1998     

Energetic and time costs of foraging in harvester ants,Pogonomyrmex occidentalis

Summary Western harvester ants, Pogonomyrmex occidentalis, preferentially utilize low vegetational cover pathways. Energetic costs for foraging ants were less than 0.1% of caloric rewards of harvested seeds, suggesting that reduction of energetic cost is not a major benefit of this preference. Walking speed was significantly faster on lower cover routes, increasing net return rates from equidistant artificial food sources. Undisturbed foragers on low cover routes traveled farther, increasing their total foraging area without increasing foraging time. These results suggest that in animals with low costs of locomotion relative to energetic rewards, time costs are more important than direct energetic costs in influencing foraging decisions. In baited experiments with equidistant food sources, preferential use of low cover routes resulted in a large increase in net energetic gain rate, but only a slight increase in energetic efficiency. Under natural conditions, net energetic gain rates were constant for foragers using low and high vegetational cover routes, but foragers using low cover paths had lower efficiencies. This suggests that net energetic gain rate is a more important currency than energetic efficiency for foraging harvester ants.     

Social information,conformity and the opportunity costs paid by foraging fish

Animals pay opportunity costs when pursuing one of several mutually exclusive courses of action. We quantified the opportunity costs of conforming to the behaviour of others in foraging sticklebacks (Pungitius pungitius), using an arena in which they were given the option of shoaling in one area or searching for food in another. Fish foraging in the absence of stimulus conspecifics found the prey patch sooner and spent longer exploiting it than those in trials where a stimulus shoal was present. Furthermore, in trials where the stimulus shoal exhibited feeding cues, subjects approached them sooner and spent more time shoaling with them, exploring less of the arena than in trials where the stimulus shoal exhibited no such cues. This suggests sensitivity not only to the mere presence of conspecifics, but also to the social information that they produce. We also saw that groups of focal fish, compared to single individuals, were less influenced by the stimulus shoal and explored more of the arena, a behaviour that may be attributed to facilitation, competition or both. Such opportunity costs are likely to be offset by benefits such as reduced predation risk, and we discuss this in terms of the trade-offs associated with living in groups.     

Costs and benefits of pocket gopher foraging: linking behavior and physiology

Animals can attain fitness benefits by maintaining a positive net energy balance, including costs of movement during resource acquisition and the profits from foraging. Subterranean rodent burrowing provides an excellent system in which to examine the effects of movement costs on foraging behavior because it is energetically expensive to excavate burrows. We used an individual-based modeling approach to study pocket gopher foraging and its relationship to digging cost, food abundance, and food distribution. We used a unique combination of an individual-based foraging-behavior model and an energetic model to assess survival, body mass dynamics, and burrow configurations. Our model revealed that even the extreme cost of digging is not as costly as it appears when compared to metabolic costs. Concentrating digging in the area where food was found, or area-restricted search (ARS), was the most energetically efficient digging strategy compared to a random strategy. Field data show that natural burrow configurations were more closely approximated by the animals we modeled using ARS compared to random diggers. By using behavior and simple physiological principles in our model, we were able to observe realistic body mass dynamics and recreate natural movement patterns.     

How habitat affects the benefits of communication in collectively foraging honey bees

Honey bees (Apis mellifera) use the dance language to symbolically convey information about the location of floral resources from within the nest. To figure out why this unique ability evolved, we need to understand the benefits it offers to the colony. Previous studies have shown that, in fact, the location information in the dance is not always beneficial. We ask, in which ecological habitats do honey bee colonies actually benefit from the dance language, and what is it about those habitats that makes communication useful? In this study, we examine the effects of floral distribution patterns on the benefits of dance communication across five different habitats. In each habitat, we manipulated colonies' ability to communicate and measured their foraging success, while simultaneously characterizing the naturally occurring floral distribution. We find that communication is most beneficial when floral species richness is high and patches contain many flowers. These are ecological features that could have helped shape the evolution of the honey bee dance language.     

Ecological costs and benefits of communal behavior in a presocial spider

Summary Philoponella oweni (Uloboridae) is a facultatively communal spider found in the southwestern United States. Solitary females and communal groups of females coexist in the same habitats. Communal females are known to produce more eggs than solitary females. Communal females might acquire the energy for the production of additional eggs by one of three means: they might have a longer life span, and thus more time in which to acquire energy; web maintenance might be more efficient in a communal group; or communal females might spend more time feeding. Field data from a marked population in Arizona showed the third alternative was true.Two possible causes for the greater time spent feeding by communal females were investigated; either the communal web or group is a more efficient trap for insects, or the communal groups are located at sites where insects are more abundant. Analysis of feeding rates and sticky trap data confirmed the second alternative.Potential web attachment sites are limited in some habitats, and mortality during dispersal is very high. When potential web attachment sites are scarce more of the population is found in communal groups; when they are abundant more of the population is solitary. The pattern of dispersal and overwintering is this species suggests that females renewing a colony in the spring could be related.     

The costs and benefits of predator inspection behaviour in Thomson's gazelles

Summary When Thomson's gazelles (Gazella thomsoni) detect stalking predators, such as cheetahs (Acinonyx jubatus) and lions (Panthera leo), they often approach and follow the predator for up to 72 min (average 14 min). Coursing predators are rarely approached. Gazelle groups were more likely to approach cheetahs if the groups were larger, if the vegetation was low, or if the cheetahs came closer to the group. Immature gazelles were more likely to approach than adults, and a higher proportion of group members participated in inspection behaviour in small groups than in large ones. Gazelles approached closer in less risky situations: if they were in larger groups or if the vegetation was low. Inspection behaviour caused cheetahs to move further between rests and between hunting attempts. Approaching cheetahs was risky, particularly for younger gazelles (probability of being killed while inspecting a cheetah was 1 in 5000 approaches for adults and 1 in 417 approaches for half-grown/adolescent gazelles), and the risks were higher than monitoring cheetahs from a distance. The time costs of predator inspection were also considerable (less than 4.2% of daylight time budget), suggesting that the benefits must be substantial to offset these costs. The results suggested that inspection behaviour was multifunctional, causing stalking predators to move out of the vicinity, enabling gazelles to monitor the predators' movements, and providing an opportunity, particularly for younger animals, to learn about predators. By approaching, gazelles also inform predators that they have been detected and alert other gazelles to the predators' presence.     

The costs and benefits of territory sharing for the Caribbean coral reef fish,Scarus iserti

Summary Along the Caribbean coast of Panama, groups of unrelated female striped parrotfish, Scarus iserti, co-defend a common feeding territory. Field manipulations of group size and composition were performed to examine the benefits and costs accrued by dominant fish within these shared territories. Dominant fish benefit from the presence of relatively large subordinates because they share in the defense of the territory. Removals of these fish caused increases in defense time and decreases in feeding time for dominant group members. Dominants benefit from the presence of small subordinates because they increase the foraging efficiency of dominants. Removals of smaller subordinates caused reductions in the feeding time of dominant fish, although no changes in defense time occurred. Concurrently, dominant fish reduce costs of resource depletion by displacing subordinate group members from good food patches. Dominance interactions within a group reduce the amount of time subordinates spend feeding (subordinate individuals fed at higher rates following the removal of a dominant) and limit a subordinate's access to high quality resources. This combination of benefits and reduced costs ultimately contributes to the economic defensibility of a striped parrotfish territory and has led to the evolution of group territorial behavior in the absence of kin selection and cooperative parental care.     

Sexual conflict in a polygynous primate: costs and benefits of a male-imposed mating system

Fundamental reproductive interests dictate that females generally benefit most from mate selectivity and males from mate quantity. This can create conflict between the sexes and result in sexual coercion: male use of aggression to garner mating success at a cost to females. Potential fitness costs of sexual coercion, however, can be difficult to measure. Here, we demonstrate benefits to males and costs to females of female defense polygyny in wild hamadryas baboons, cercopithecoid primates in which females are coercively transferred among social units by males, restricting both female choice and bonding among female kin. Of all coercive transfers (takeovers) of females with young infants, 67 % were followed by infant mortality, which was significantly more likely to occur after takeovers than at other times. As expected, infant mortality decreased time to subsequent conception but lengthened intervals between surviving infants. Following infant survival, whether a female had experienced a takeover after the previous birth was a significant predictor of subsequent interbirth interval, with interbirth intervals of females remaining with the same male between births being significantly shorter than those of females incurring takeovers between births. Together, these results reveal that takeovers increase the chance of infant mortality while delaying subsequent conception. Male-driven female defense polygyny in this species is thus costly to females in two ways. These results demonstrate that reproductive strategies benefitting males can evolve despite substantial costs to females. These costs may be mitigated over the long term, however, by female counterstrategies and protective behavior by males.     

Learning during competitive positioning in the nest: do nestlings use ideal free ‘foraging’ tactics?

Begging behaviour of nestling birds may involve more than a simple, honest source of information for parents to use in provisioning. Many aspects of begging behaviour relate instead to sibling competition for food items within the nest, and we might expect evidence of adaptive learning and behavioural adjustment in response to experience of the competitive environment. In this study, we consider begging in different locations within the nest as analogous to foraging in different patches, varying in food availability. Using hand-feeding trials, we created zones of differing profitability within an artificial nest by adjusting either the prey size or number of items delivered, and allowed only indirect competition between pairs of southern grey shrike (Lanius meridionalis) nestlings. Nestlings demonstrated the ability to detect differences in zone profitability and position themselves accordingly. By the end of both the prey size and delivery rate trials nestlings had increased the amount of time spent in the high quality zone. Such movement in response to differences in load quality, as well as frequency, demonstrates the ability of nestlings to learn about their environment and to facultatively adjust their begging in order to maximise energetic rewards.     

Mating system evolution in response to search costs in the speckled wood butterfly, Pararge aegeria

A general and intuitive prediction from models of mate preference is that when the cost of searching for mates increases, individuals should become less choosy. Here, we test this prediction by comparing the mating propensity of females in two populations of the butterfly Pararge aegeria. The populations originated from southern Sweden and Madeira and due to different adult emergence patterns throughout the year, the average density of males per female is likely to be lower on Madeira. Therefore, we expected that the cost of searching should be greater on Madeira and, consequently, that the Madeiran females should be less choosy. In line with predictions, the Madeiran females mated significantly sooner after the first interaction with males than did females from southern Sweden. This difference may reflect a weaker preference for territorial males over non-territorial patrollers in the Madeiran population, because of the greater costs of searching. The Madeiran females also showed a shorter time lag between mating and the start of oviposition. We discuss this unexpected result and propose that the same mechanism could also explain this population difference, i.e. different costs of searching for suitable host plants. Both search processes are fundamental for female reproductive success and we find it plausible that they can be generalised into the same theory of optimal search behaviour. Received: 14 May 1998 / Accepted after revision: 13 December 1998     

Set-up costs and the existence of competitive equilibrium when extraction capacity is limited

Although set-up costs are prevalent and substantial in natural resource extraction, it is known that a Walrasian competitive equilibrium cannot exist in simple extraction models with set-up costs. This paper demonstrates that this result is sensitive to the assumption of unlimited extraction capacity and derives sufficient conditions for existence. An equilibrium exists if extraction is limited such that each firm earns sufficient surplus to cover its set-up costs or if firms choose extraction capacity subject to non-increasing returns. The resulting competitive equilibrium price either grows at the rate of interest when total extraction is below industry capacity or is constant when industry capacity is fully utilized. In the equilibrium, identical deposits are opened simultaneously, and set-up costs for new deposits are incurred when the industry has excess capacity rather than when capacity is fully utilized.     

Effects of sex-associated competitive asymmetries on foraging group structure and despotic distribution in Andean condors

Phenotype-limited interference models assume competitive asymmetries among conspecifics and unequal sharing of resources. Their main prediction is a correlation between dominance status and patch quality: dominant individuals should preferentially exploit better-quality habitats. We tested assumptions and predictions of the phenotype-limited interference model in Andean condors (Vultur gryphus), a New World vulture with strong sexual size dimorphism (males are 30–40% heavier than females). We recorded searching birds in habitats differing in quality: mountains and plains. We also observed scavenging behaviour at 20 sheep carcasses, and videotaped 5 of them. Intraspecific hierarchy at carcasses was based on size: males dominated females and, within each sex, older birds dominated younger ones. Adult males and juvenile females occupied extreme positions in the feeding hierarchy. Aggression was directed at those individuals belonging to lower hierarchical levels. In high-quality areas (mountains), more condors arrived at carcasses. Juvenile females were more often observed searching in low-quality areas (plains), far from breeding areas and main roost sites. GLM analyses of individual behaviour showed that the hierarchy did not influence time of arrival, but low-ranking individuals spent more time at carcasses, especially if the number of condors at arrival was high. Additionally, low-ranking condors spent less time feeding at carcasses when individuals of higher hierarchical levels were present. On the other hand, the number of condors present had a positive effect on feeding rates of dominant individuals, probably because of a reduction in individual vigilance. These results support most of the assumptions and predictions of the phenotype-limited distribution model, although a spatial truncated distribution between phenotypes was not observed. Asymmetric feeding pay-off, unequal parental roles and sexual selection constraints could favour sexual divergence in body size in Andean condors. Received: 6 April 1998 / Accepted after revision: 11 July 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.     

Effects of costs and benefits of brood care on filial cannibalism in the sand goby

This study tested experimentally whether clutch size and the cost of care affect filial cannibalism in the sand goby, Pomatoschistus minutus. Evolutionary models of filial cannibalism suggest that egg eating has evolved as a way for the male parent to prolong his breeding season. These models assume that eggs function as an alternative energy source for the constrained parent. I manipulated clutch size by allowing males to mate with either one or two females, representing a small and a large clutch, respectively. The addition of a small male shore crab, a common egg predator, increased the cost of care. I quantified fat reserves as a measure of the condition of guarding males. Males who did not build nests had lower fat reserves than males who built nests, suggesting that males with low energy reserves do not start breeding. Males with small clutches lost their nest to the crab more often than males with large clutches. Neither filial cannibalism nor the amount of eggs eaten were affected by the treatments. Males who consumed eggs had a higher fat percentage than males who did not eat eggs. The result that males with small clutches lost their nests to the crabs supports the idea that eggs are defended only if the benefit from continued care will outweigh the cost and that males therefore are sensitive to the trade-off between present and future reproductive success. Received: 15 May 1997 / Accepted after revision: 15 November 1997