Hummingbird responses to gender-biased nectar production: are nectar biases maintained by natural or sexual selection?

Pollinators mediate the evolution of secondary floral traits through both natural and sexual selection. Gender-biased nectar, for example, could be maintained by one or both, depending on the interactions between plants and pollinators. Here, I investigate pollinator responses to gender-biased nectar using the dichogamous herb Chrysothemis friedrichsthaliana (Gesneriaceae) which produces more nectar during the male floral phase. Previous research showed that the hummingbird pollinator Phaethornis striigularis visited male-phase flowers more often than female-phase flowers, and multiple visits benefited male more than female fecundity. If sexual selection maintains male-biased rewards, hummingbirds should prefer more-rewarding flowers independent of floral gender. If, however, differential rewards are partially maintained through natural selection, hummingbirds should respond to asymmetry with visits that reduce geitonogamy, i.e. selfing and pollen discounting. In plants with male biases, these visit types include single-flower visits and movements from low to high rewards. To test these predictions, I manipulated nectar asymmetry between pairs of real or artificial flowers on plants and recorded foraging behaviour. I also assessed maternal costs of selfing using hand pollinations. For plants with real flowers, hummingbirds preferred more-rewarding flowers and male-phase morphology, the latter possibly owing to previous experience. At artificial arrays, hummingbirds responded to extreme reward asymmetry with increased single-flower visits; however, they moved from high to low rewards more often than low to high. Finally, selfed flowers did not produce inferior seeds. In summary, sexual selection, more so than geitonogamy avoidance, maintains nectar biases in C. friedrichsthaliana, in one of the clearest examples of sexual selection in plants, to date.     

Testing dynamic variance-sensitive foraging using individual differences in basal metabolic rates of zebra finches

Social foragers can alternate between searching for food (producer tactic), and searching for other individuals that have located food in order to join them (scrounger tactic). Both tactics yield equal rewards on average, but the rewards generated by producer are more variable. A dynamic variance-sensitive foraging model predicts that social foragers should increase their use of scrounger with increasing energy requirements and/or decreased food availability early in the foraging period. We tested whether natural variation in minimum energy requirements (basal metabolic rate or BMR) is associated with differences in the use of producer–scrounger foraging tactics in female zebra finches Taeniopygia guttata . As predicted by the dynamic variance-sensitive model, high BMR individuals had significantly greater use of the scrounger tactic compared with low BMR individuals. However, we observed no effect of food availability on tactic use, indicating that female zebra finches were not variance-sensitive foragers under our experimental conditions. This study is the first to report that variation in BMR within a species is associated with differences in foraging behaviour. BMR-related differences in scrounger tactic use are consistent with phenotype-dependent tactic use decisions. We suggest that BMR is correlated with another phenotypic trait which itself influences tactic use decisions.     

Observational Learning in the White-Eared Hummingbird (Hylocharis leucotis): Experimental Evidence

The adoption of new food resources can be facilitated by the ability to learn through observation of other individuals who use them. This behavior, termed observational learning, applies to any problem solving in which a naive individual who has observed an experienced individual learns a behavior faster than another who has not. Hummingbirds consume nectar from flowers of a large number of plant species, which are very diverse in morphology and color. During their local or migratory movements, they can observe the use of floral resources by conspecifics and heterospecifics which may change their foraging preferences. Although there is evidence that hummingbirds can use observational learning to exploit new floral resources, it is necessary to generate additional information by studying different hummingbird species. In this work, the learning performance of White‐eared hummingbirds (Hylocharis leucotis) was studied in the presence or absence of a knowledgeable tutor. In a first experiment, naïve hummingbirds learned to feed on arrays of artificial flower of two colors: red (previously known resource) and yellow (novel resource), where only one color had nectar. Naive hummingbirds visited red flowers faster and more often than rewarded yellow flowers. Individuals with the best performance on each color were further trained to ensure that they only visited flowers of a specific color, and were then used as tutors in the next experiment, in which new naive hummingbirds, caged individually, were allowed to observe them foraging on the artificial arrays. These naïve individual were then exposed alone to the same array used by their tutor. Tutored hummingbirds learned to feed faster and more frequently from nectar‐containing flowers of the array than naive individuals. Likewise, all tutored individuals only visited flowers of the color that had been previously visited by their tutors. This study provides experimental evidence that hummingbirds taken directly from the field can use observational learning as an efficient strategy to access new floral resources.     

Relationships between densities of previous and simultaneous foragers and the foraging behaviour of three bumblebee species

1. The local insect composition may be as important as the local floral composition for bumblebees' foraging behaviour. However, little is known about how the local abundance of insects affects the foraging patterns of individuals. 2. Using field observations, we studied the relationships between the local density of previous and simultaneous foragers and the local foraging behaviour of Bombus pascuorum, Bombus lucorum/B. terrestris and Bombus lapidarius while pollinating Centaurea jacea. 3. The number of bumblebees foraging in the plots was positively related to the number of new individuals arriving at these plots. The number of inflorescences contacted and the duration of visits were negatively related to the number of simultaneous foragers, but only in B. lucorum/B. terrestris. The effects of previous foragers on the behaviour of other bumblebees were species‐specific and variable in their direction. Such contrasting effects can be explained in terms of bumblebee species' abundances and functional similarity. In some cases, the effect of previous foragers increased with Centaurea density. 4. The local abundance of previous and simultaneous foragers affected the foraging behaviour of particular bumblebee individuals in complex ways. Future studies on local foraging behaviour might benefit from including the abundance of co‐foragers.     

Corticosterone and foraging behavior in a pelagic seabird

Because endocrine mechanisms are thought to mediate behavioral responses to changes in the environment, examining these mechanisms is essential for understanding how long-lived seabirds adjust their foraging decisions to contrasting environmental conditions in order to maximize their fitness. In this context, the hormone corticosterone (CORT) deserves specific attention because of its major connections with locomotor activities. We examined for the first time the relationships between individual CORT levels and measurements of foraging success and behavior using satellite tracking and blood sampling from wandering albatrosses (Diomedea exulans) before (pretrip CORT levels) and after (posttrip CORT levels) foraging trips during the incubation period. Plasma CORT levels decreased after a foraging trip, and the level of posttrip CORT was negatively correlated with individual foraging success, calculated as total mass gain over a foraging trip. Pretrip CORT levels were not linked to time spent at sea but were positively correlated with daily distance traveled and maximum range at sea. In this study, we were able to highlight the sensitivity of CORT levels to variation in energy intake, and we showed for the first time that individual CORT levels can be explained by variation in foraging success. Relationships between pretrip CORT levels and daily distance traveled and maximum range were independent of pretrip body mass, suggesting that slight elevations in pretrip CORT levels might facilitate locomotor activity. However, because both foraging behavior and pretrip CORT levels could be affected by individual quality, future experimental studies including manipulation of CORT levels are needed to test whether CORT can mediate foraging decisions according to foraging conditions.     

Risk‐averse inflorescence departure in hummingbirds and bumble bees: could plants benefit from variable nectar volumes?

Most hermaphroditic, many-flowered plants should suffer reduced fitness from within-plant selfing (geitonogamy). Large inflorescences are most attractive to pollinators, but also promote many flower visits during a single plant visit, which may increase selfing and decrease pollen export. A plant might avoid the negative consequences of attractiveness through modification of the floral display to promote fewer flower visits, while retaining attractiveness. This report shows that increasing only the variance in nectar volume per flower results in fewer flower visits per inflorescence by wild hummingbirds ( Selasphorus rufus ) and captive bumble bees ( Bombus flavifrons ) foraging on artificial inflorescences. Inflorescences were either constant (all flowers contained the same nectar volume) or variable (half the flowers were empty, the other half contained twice as much nectar as in the constant flowers). Both types of inflorescence were simultaneously available to foragers. Risk-averse foraging behaviour was expressed as a patch departure preference: birds and bees visited fewer flowers on variable inflorescences, and this preference was expressed when resource variability could be determined only by concurrent sampling. When variance treatments were clearly labelled with colour and offered to hummingbirds, the departure effect was maintained; however, when preference was measured by inflorescence choice, birds did not consistently prefer to visit constant inflorescences. The reduced visitation lengths on variable inflorescences by both birds and bees documented in this study imply that variance in nectar production rates within inflorescences may represent an adaptive trait to avoid the costs of geitonogamy.     

Individual variation in baseline and stress-induced corticosterone and prolactin levels predicts parental effort by nesting mourning doves

Endocrine systems have an important mechanistic role in structuring life-history trade-offs. During breeding, individual variation in prolactin (PRL) and corticosterone (CORT) levels affects behavioral and physiological processes that drive trade-offs between reproduction and self-maintenance. We examined patterns in baseline (BL) and stress induced (SI; level following a standard capture-restraint protocol) levels of PRL and CORT for breeding mourning doves (Zenaida macroura). We determined whether the relationship of adult condition and parental effort to hormone levels in wild birds was consistent with life-history predictions. Both BL PRL and BL CORT level in adults were positively related to nestling weight at early nestling ages, consistent with the prediction of a positive relationship of hormone levels to current parental effort of adults and associated increased energy demand. Results are consistent with the two hormones acting together at baseline levels to limit negative effects of CORT on reproduction while maintaining beneficial effects such as increased foraging for nestling feeding. Our data did not support predictions that SI responses would vary in response to nestling or adult condition. The magnitude of CORT response in the parents to our capture-restraint protocol was negatively correlated with subsequent parental effort. Average nestling weights for adults with the highest SI CORT response were on average 10–15% lighter than expected for their age in follow-up visits after the stress event. Our results demonstrated a relationship between individual hormone levels and within population variation in parental effort and suggested that hormonal control plays an important role in structuring reproductive decisions for mourning doves.     

Seasonal changes in juvenile hormone titers and rates of biosynthesis in honey bees

Honey bee colonies can respond to changing environmental conditions by showing plasticity in age related division of labor, and these responses are associated with changes in juvenile hormone. The shift from nest taks to foraging has been especially well characterized; foraging is associated with high juvenile hormone titers and high rates of juvenile hormone biosynthesis, and can be induced prematurely in young bees by juvenile hormone treatment or by a shortage of foragers. However, very few studies have been conducted that study plasticity in division of labor under naturally occurring changes in the environment. To gain further insight into how the environment and juvenile hormone influence foraging behavior, we measured juvenile hormone titers and rates of biosynthesis in workers during times of the year when colony activity in temperate climates is reduced: late fall, winter, and early spring. Juvenile hormone titers and rates of biosynthesis decreased in foragers in the fall as foraging diminished and bees became less active. This demonstration of a natural drop in juvenile hormone confirms and extends previous findings when bees were experimentally induced to revert from foraging to within-hive tasks. In addition, endocrine changes in foragers in the fall are part of a larger seasonally related phenomenon in which juvenile hormone levels in younger, pre-foraging bees also decline in the fall and then increase the following spring as colony activity increases. The seasonal decline in juvenile hormone in foragers was mimicked in summer by placing a honey bee colony in a cold room for 8 days. This suggests that seasonal changes in juvenile hormone are not related to photoperiod changes, but rather to changes in temperature and/or colony social structure that in turn influence endocrine and behavioral development. We also found that active foragers in the late winter and early spring had lower juvenile hormone levels than active foragers in late spring. In light of recent findings of a possible link between juvenile hormone and neuroanatomical plasticity in the bee brain, these results suggest that bees can forage with low juvenile hormone, after previous exposure to some threshold level of juvenile hormone leads to changes in brain structure.     

Simple foraging rules in competitive environments can generate socially structured populations

Social vertebrates commonly form foraging groups whose members repeatedly interact with one another and are often genetically related. Many species also exhibit within‐population specializations, which can range from preferences to forage in particular areas through to specializing on the type of prey they catch. However, within‐population structure in foraging groups, behavioral homogeneity in foraging behavior, and relatedness could be outcomes of behavioral interactions rather than underlying drivers. We present a simple process by which grouping among foragers emerges and is maintained across generations. We introduce agent‐based models to investigate (1) whether a simple rule (keep foraging with the same individuals when you were successful) leads to stable social community structure, and (2) whether this structure is robust to demographic changes and becomes kin‐structured over time. We find the rapid emergence of kin‐structured populations and the presence of foraging groups that control, or specialize on, a particular food resource. This pattern is strongest in small populations, mirroring empirical observations. Our results suggest that group stability can emerge as a product of network self‐organization and, in doing so, may provide the necessary conditions for the evolution of more sophisticated processes, such as social learning. This taxonomically general social process has implications for our understanding of the links between population, genetic, and social structures.     

Octopamine influences honey bee foraging preference

Colony condition and differences in individual preferences influence forage type collected by bees. Physiological bases for the changing preferences of individual foragers are just beginning to be examined. Recently, for honey bees octopamine is shown to influence age at onset of foraging and probability of dance for rewards. However, octopamine has not been causally linked with foraging preference in the field. We tested the hypothesis that changes in octopamine may alter forage type (preference hypothesis). We treated identified foragers orally with octopamine or its immediate precursor, tyramine, or sucrose syrup (control). Octopamine-treated foragers switched type of material collected; control bees did not. Tyramine group results were not different from the control group. In addition, sugar concentrations of nectar collected by foragers after octopamine treatment were lower than before treatment, indicating change in preference. In contrast, before and after nectar concentrations for bees in the control group were similar. These results, taken together, support the preference hypothesis.     

Plant and habitat use by Black‐breasted Pufflegs (Eriocnemis nigrivestis), a critically endangered hummingbird

Black‐breasted Pufflegs (Eriocnemis nigrivestis) are hummingbirds endemic to Ecuador and are considered critically endangered because of their limited distribution, a population estimated at fewer than 1000 individuals, and ongoing habitat degradation. From November 2013 to June 2016, we examined the foraging preferences of these hummingbirds using a combination of direct observations, time‐lapse cameras, and motion‐detection software. We first identified 21 species of ornithophilous plants distributed among five sites in the northwestern flanks of the Pichincha volcano in northwest Ecuador. We then monitored these plant species using time‐lapse cameras and recorded 144 visits by Black‐breasted Pufflegs to seven of the 21 species. Most visits (128 of 144 visits, 89%) were to just two species of plants, Macleania rupestris and Palicourea fuchsioides, the latter of which is also an endemic and threatened species. In addition, Black‐breasted Pufflegs were only observed in the most pristine habitats. Given the potential negative effects of climate change for species in the tropical Andes plus the possible loss and degradation of habitat resulting from human activities, efforts are needed to conserve habitats currently used by Black‐breasted Pufflegs, recover degraded habitats, and connect isolated patches of suitable habitat. Our results concerning species of flowering plants used most by Black‐breasted Pufflegs (P. fuchsioides and M. rupestris) should help guide any habitat restoration initiatives.     

Density‐Dependent Foraging and Interference Competition by Common Gartersnakes are Temperature Dependent

The ideal free distribution (IFD) predicts that optimal foragers will select foraging patches to maximize food rewards and that groups of foragers should thus be distributed between food patches in proportion to the availability of food in those patches. Because many of the underlying mechanisms of foraging are temperature dependent in ectotherms, the distribution of ectothermic foragers between food patches may similarly depend on temperature because the difference in fitness rewards between these patches may change with temperature. We tested the hypothesis that the distribution of Common Gartersnakes (Thamnophis sirtalis) between food patches can be explained by an IFD, but that conformance to an IFD weakens as temperature departs from the optimal temperature because fitness rewards, interference competition and the number of individuals foraging are highest at the optimal temperature. First, we determined the optimal temperature for foraging. Second, we examined group foraging at three temperatures and three density treatments. Search time was optimized at 27°C, handling time at 29°C and digestion time at 32°C. Gartersnakes did not match an IFD at any temperature, but their distribution did change with temperature: snakes at 20°C and at 30°C selected both food patches equally, while snakes at 25°C selected the low food patch more at low density and the high food patch more at high density. Food consumption and competition increased with temperature, and handling time decreased with temperature. Temperature therefore had a strong impact on foraging, but did not affect the IFD. Future work should examine temperature‐dependent foraging in ectotherms that are known to match an IFD.     

An empirical demonstration of risk-sensitive foraging preferences

We report laboratory experiments with yellow-eyed juncos (Junco phaeonotus) revealing that the birds' foraging preferences for variable rewards respond not only to the mean, but also the variance, of food rewards. The nature of their preferences for variable rewards is related to their expected daily energy budget. We summarize the birds' preferences in utility functions for energetic rewards. Since mean reward size is inadequate to predict their behaviour, we believe that foraging models should consider environmental stochasticity and an animal's response to this variation.     

Social effects on foraging behavior and success depend on local environmental conditions

In social groups, individuals' dominance rank, social bonds, and kinship with other group members have been shown to influence their foraging behavior. However, there is growing evidence that the particular effects of these social traits may also depend on local environmental conditions. We investigated this by comparing the foraging behavior of wild chacma baboons, Papio ursinus, under natural conditions and in a field experiment where food was spatially clumped. Data were collected from 55 animals across two troops over a 5‐month period, including over 900 agonistic foraging interactions and over 600 food patch visits in each condition. In both conditions, low‐ranked individuals received more agonism, but this only translated into reduced foraging performances for low‐ranked individuals in the high‐competition experimental conditions. Our results suggest one possible reason for this pattern may be low‐ranked individuals strategically investing social effort to negotiate foraging tolerance, but the rank‐offsetting effect of this investment being overwhelmed in the higher‐competition experimental environment. Our results also suggest that individuals may use imbalances in their social bonds to negotiate tolerance from others under a wider range of environmental conditions, but utilize the overall strength of their social bonds in more extreme environments where feeding competition is more intense. These findings highlight that behavioral tactics such as the strategic investment of social effort may allow foragers to mitigate the costs of low rank, but that the effectiveness of these tactics is likely to be limited in certain environments.     

Foraging errors play a role in resource exploration by bumble bees (Bombus terrrestris)

If the cognitive performance of animals reflects their particular ecological requirements, how can we explain appreciable variation in learning ability amongst closely related individuals (e.g. foraging workers within a bumble bee colony)? One possibility is that apparent ‘errors’ in a learning task actually represent an alternative foraging strategy. In this study we investigate the potential relationship between foraging ‘errors’ and foraging success among bumble bee (Bombus terrestris) workers. Individual foragers were trained to choose yellow, rewarded flowers and ignore blue, unrewarded flowers. We recorded the number of errors (visits to unrewarded flowers) each bee made during training, then tested them to determine how quickly they discovered a more profitable food source (either familiar blue flowers, or novel green flowers). We found that error prone bees discovered the novel food source significantly faster than accurate bees. Furthermore, we demonstrate that the time taken to discover the novel, more profitable, food source is positively correlated with foraging success. These results suggest that foraging errors are part of an ‘exploration’ foraging strategy, which could be advantageous in changeable foraging environments. This could explain the observed variation in learning performance amongst foragers within social insect colonies.     

Elevated stress hormone diminishes the strength of female preferences for acoustic signals in the green treefrog

Mate selection can be stressful; time spent searching for mates can increase predation risk and/or decrease food consumption, resulting in elevated stress hormone levels. Both high predation risk and low food availability are often associated with increased variation in mate choice by females, but it is not clear whether stress hormone levels contribute to such variation in female behavior. We examined how the stress hormone corticosterone (CORT) affects female preferences for acoustic signals in the green treefrog, Hyla cinerea. Specifically, we assessed whether CORT administration affects female preferences for call rate — an acoustic feature that is typically under directional selection via mate choice by females in most anurans and other species that communicate using acoustic signals. Using a dual speaker playback paradigm, we show that females that were administered higher doses of CORT were less likely to choose male advertisement calls broadcast at high rates. Neither CORT dose nor level was related to the latency of female phonotactic responses, suggesting that elevated CORT does not influence the motivation to mate. Results were also not related to circulating sex steroids (i.e., progesterone, androgens or estradiol) that have traditionally been the focus of studies examining the hormonal basis for variation in female mate choice. Our results thus indicate that elevated CORT levels decrease the strength of female preferences for acoustic signals.     

Social Environment Affects Peripheral Oxytocin and Cortisol during Stress Responses in Guinea-Pigs

Numerous studies have demonstrated interactions between oxytocin (OT) secretion, adrenal activity, an animal's social environment, and stress responses. In the present study, we hypothesized that partner preference and pair bonding cause increased peripheral OT, which down‐regulates the adrenal and behavioral response in stress. In addition, we tested whether these interactions depended on sex, the social environment of the individual, or the type of stressor. Experiments were carried out on guinea‐pigs held in sexual‐pairs or stressed by isolation. Among the paired individuals choice tests were carried out to document partner preference. Female preference for a male was expressed by spatial cohesion. Stress responses to abiotic stimuli were examined and compared between isolated and cohabited animals with or without preference. Results show that OT of cohabited animals was significantly higher than during isolation. OT levels were further increased in males preferred by females. Cortisol (CORT) levels were elevated in isolated animals. There were no significant differences between pairs with and without female preference. Behaviorally, partner preferences were expressed by high amounts of tactile contact. The stress‐induced behavioral immobility response after exposure to a noise stressor was significantly reduced in preferred males and to a lesser extent in their female partners. Only females with preferences showed an endocrine stress response. Their levels of OT increased. There was no consistent post‐stressor release of CORT. The data indicate that the social environment of an individual, here expressed as preference or isolation, influences peripheral OT secretion. Behavioral stress responses were similarly affected by social factors without a clear involvement of peripheral CORT or OT.     

The Bold and the Variable: Fish with High Heterozygosity Act Recklessly in the Vicinity of Predators

Variation in the innate behavioral response to predation threat is often assumed to reflect genetic differences among the prey individuals. To date, no published results, however, exist that would offer explanation for the origin of this behavioral variation within populations. Using microsatellites as markers, we estimated the genetic variability of juvenile brown trout (Salmo trutta) individuals whose behavior had been individually recorded in a trade‐off situation where both predator chemical cues and food were present. Mean overall heterozygosity and the internal relatedness of fish associated significantly with their activity and foraging, so that the genetically more variable individuals showed more risk‐prone behavior under predation risk. No association between genetic variability and behavior was found in trials where predator odors were not present. These results were consistent over the three study populations of brown trout with different backgrounds, suggesting that the phenomenon is of general nature in this species. Of the possible mechanisms suggested to enable the existence of the positive association between neutral microsatellite variation and fitness‐related trait, the local effect hypothesis gained more support from our data than the general effect hypothesis.     

Baseline glucocorticoids are drivers of body mass gain in a diving seabird

Life‐history trade‐offs are influenced by variation in individual state, with individuals in better condition often completing life‐history stages with greater success. Although resource accrual significantly impacts key life‐history decisions such as the timing of reproduction, little is known about the underlying mechanisms driving resource accumulation. Baseline corticosterone (CORT, the primary avian glucocorticoid) mediates daily and seasonal energetics, responds to changes in food availability, and has been linked to foraging behavior, making it a strong potential driver of individual variation in resource accrual and deposition. Working with a captive colony of white‐winged scoters (Melanitta fusca deglandi), we aimed to causally determine whether variation in baseline CORT drives individual body mass gains mediated through fattening rate (plasma triglycerides corrected for body mass). We implanted individuals with each of three treatment pellets to elevate CORT within a baseline range in a randomized order: control, low dose of CORT, high dose of CORT, then blood sampled and recorded body mass over a two‐week period to track changes in baseline CORT, body mass, and fattening rates. The high CORT treatment significantly elevated levels of plasma hormone for a short period of time within the biologically relevant, baseline range for this species, but importantly did not inhibit the function of the HPA (hypothalamic–pituitary–adrenal) axis. Furthermore, an elevation in baseline CORT resulted in a consistent increase in body mass throughout the trial period compared to controls. This is some of the first empirical evidence demonstrating that elevations of baseline CORT within a biologically relevant range have a causal, direct, and positive influence on changes in body mass.     

Monitoring Flower Visitation Networks and Interactions between Pairs of Bumble Bees in a Large Outdoor Flight Cage

Pollinators, such as bees, often develop multi-location routes (traplines) to exploit subsets of flower patches within larger plant populations. How individuals establish such foraging areas in the presence of other foragers is poorly explored. Here we investigated the foraging patterns of pairs of bumble bees (Bombus terrestris) released sequentially into an 880m² outdoor flight cage containing 10 feeding stations (artificial flowers). Using motion-sensitive video cameras mounted on flowers, we mapped the flower visitation networks of both foragers, quantified their interactions and compared their foraging success over an entire day. Overall, bees that were released first (residents) travelled 37% faster and collected 77% more nectar, thereby reaching a net energy intake rate 64% higher than bees released second (newcomers). However, this prior-experience advantage decreased as newcomers became familiar with the spatial configuration of the flower array. When both bees visited the same flower simultaneously, the most frequent outcome was for the resident to evict the newcomer. On the rare occasions when newcomers evicted residents, the two bees increased their frequency of return visits to that flower. These competitive interactions led to a significant (if only partial) spatial overlap between the foraging patterns of pairs of bees. While newcomers may initially use social cues (such as olfactory footprints) to exploit flowers used by residents, either because such cues indicate higher rewards and/or safety from predation, residents may attempt to preserve their monopoly over familiar resources through exploitation and interference. We discuss how these interactions may favour spatial partitioning, thereby maximising the foraging efficiency of individuals and colonies.