Effects of clothianidin on Bombus impatiens (Hymenoptera: Apidae) colony health and foraging ability

We conducted laboratory experiments to investigate the lethal and sublethal effects of clothianidin on bumble bee, Bombus impatiens Cresson, colony health and foraging ability. Bumble bee colonies were exposed to 6 ppb clothianidin, representing the highest residue levels found in field studies on pollen, and a higher dose of 36 ppb clothianidin in pollen. Clothianidin did not effect pollen consumption, newly emerged worker weights, amount of brood or the number of workers, males, and queens at either dose. The foraging ability of worker bees tested on an artificial array of complex flowers also did not differ among treatments. These results suggest that clothianidin residues found in seed-treated canola and possibly other crops will not adversely affect the health of bumble bee colonies or the foraging ability of workers.     

Consistent pollen nutritional intake drives bumble bee (Bombus impatiens) colony growth and reproduction across different habitats

Foraging behavior is a critical adaptation by insects to obtain appropriate nutrients from the environment for development and fitness. Bumble bees (Bombus spp.) form annual colonies which must rapidly increase their worker populations to support rearing reproductive individuals before the end of the season. Therefore, colony growth and reproduction should be dependent on the quality and quantity of pollen resources in the surrounding landscape. Our previous research found that B. impatiens foraging preferences to different plant species were shaped by pollen protein:lipid nutritional ratios (P:L), with foragers preferring pollen species with a ~5:1 P:L ratio. In this study, we placed B. impatiens colonies in three different habitats (forest, forest edge, and valley) to determine whether pollen nutritional quality collected by the colonies differed between areas that may differ in resource abundance and diversity. We found that habitat did not influence the collected pollen nutritional quality, with colonies in all three habitats collecting pollen averaging a 4:1 P:L ratio. Furthermore, there was no difference in the nutritional quality of the pollen collected by colonies that successfully reared reproductives and those that did not. We found however, that “nutritional intake,” calculated as the colony‐level intake rate of nutrient quantities (protein, lipid, and sugar), was strongly related to colony growth and reproductive output. Therefore, we conclude that B. impatiens colony performance is a function of the abundance of nutritionally appropriate floral resources in the surrounding landscape. Because we did not comprehensively evaluate the nutrition provided by the plant communities in each habitat, it remains to be determined how B. impatiens polylectic foraging strategies helps them select among the available pollen nutritional landscape in a variety of plant communities to obtain a balance of key macronutrients.     

Colony response to graded resource changes: an analytical model of the influence of genotype,environment, and dominance

Successful social groups must respond dynamically to environmental changes. However, a flexible group response requires the coordination of many individuals. Here we offer a static analytical model that integrates variation in environment-based cues for performance of a task with genetically and environmentally based variation in individual responses, and predicts the resultant colony behavior for that task. We also provide formulae for computing effective number of alleles in a haplo-diploid colony founded by any number of parents. Variable colony resources combined with variation among worker phenotypes generate known patterns of colony flexibility, allowing us to explicitly test how the number of loci, dominance/codominance, and the phenotype's environment influences group response. Our model indicates that the number of loci strongly influences colony behavior. For one or two loci, the proportion of workers foraging for pollen remain constant over vast increases in colony pollen stores, but then drops dramatically when the pollen stores increase past a specific threshold. As the number of loci controlling pollen foraging increases, graded increases in pollen stores result in a graded drop in the proportion of the worker population foraging for pollen. The effect of number of alleles is less strong, a result we discuss in light of the fact that a low number of effective alleles are expected in a colony. Comparisons of our model with empirical honey bee (Apis mellifera) data indicate that worker foraging response to pollen stores is driven by one or two loci, each with dominant allelic effects. The growing body of evidence that genotype has strong effects on task performance in social insect colonies, and the variation in within-colony genetic diversity across social insect taxa, make our model broadly applicable in explaining social group coordination.     

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

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

The relationship between comb age and performance of honey bee (Apis mellifera) colonies

A study on the relationship between the age of comb and the activity of the hybrid Carniolan honey bee colonies in collecting pollen activity, worker brood production, colony strength, and honey yield was conducted. In comparison to colonies with combs aged 4-years, colonies with combs aged 1, 2 and 3-years significantly exceeded in the number returning workers, number returning workers with pollen loads, rate of storing pollen, rate of worker brood production, and size of colony population. Colonies with combs aged 1, 2 and 3-years produced significantly more honey than colonies with combs aged 4-years (5.25, 4.90 and 4.65 kg/colony vs. 4.45 kg/colony, respectively). It can be concluded that the foraging rate, gathering and storing pollen, brood production, colony population size, and honey yield significantly depended on the age of combs. Beekeepers can replace old combs with new ones to increase brood and honey production.     

蜜蜂花粉采集行为的调控机制

蜜蜂Apisspp.是一种社会性昆虫。社会性昆虫在对它们群体自身数量和巢穴环境的调节方面表现出明显的稳态特点,Emerson将这种稳态调节称为社会性稳态。蜂群中花粉的储存量就具有稳态的特点。蜂群的花粉采集行为是由蜂群对花粉的需要决定的。关于蜜蜂花粉采集行为的调控机制,目前的研究主要集中于是哪些信息以及蜂群是如何识别这些信息从而调控其采粉行为,主要形成了直接识别和间接识别2种假说。对这2种假说进行综述。     

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

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

Genotype and rearing environment affect honeybee perception and foraging behaviour

We tested the effects of larval and preforaging rearing environment on the foraging behaviour and sucrose response thresholds of honeybees, Apis mellifera L., derived from high and low pollen-hoarding strains. Bees were reared as larvae and as preforaging adults in colonies containing high and low pollen-hoarding strains, then cofostered in unrelated common wild-type colonies from which to forage. Genotype, but not rearing environment, had strong effects on the likelihood to forage for pollen or nectar, the size of pollen or nectar load, and the concentration of sugar in the nectar they collected. Genotype and rearing environment affected adult wet weights and sucrose concentration response threshold, as measured with the proboscis extension response assay. Bees from the high pollen-hoarding strain were more sensitive to conditions of the rearing environment than were bees of the low strain. High- and low-strain bees produced different colony environments that affected developmental, behavioural and physical traits of the individuals they reared. This demonstrates how genotype and colony environment correlate and affect phenotype. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Multiple stressors: using the honeybee model BEEHAVE to explore how spatial and temporal forage stress affects colony resilience

The causes underlying the increased mortality of honeybee Apis mellifera colonies observed over the past decade remain unclear. Since so far the evidence for monocausal explanations is equivocal, involvement of multiple stressors is generally assumed. We here focus on various aspects of forage availability, which have received less attention than other stressors because it is virtually impossible to explore them empirically. We applied the colony model BEEHAVE, which links within‐hive dynamics and foraging, to stylized landscape settings to explore how foraging distance, forage supply, and “forage gaps”, i.e. periods in which honeybees cannot find any nectar and pollen, affect colony resilience and the mechanisms behind. We found that colony extinction was mainly driven by foraging distance, but the timing of forage gaps had strongest effects on time to extinction. Sensitivity to forage gaps of 15 days was highest in June or July even if otherwise forage availability was sufficient to survive. Forage availability affected colonies via cascading effects on queen's egg‐laying rate, reduction of new‐emerging brood stages developing into adult workers, pollen debt, lack of workforce for nursing, and reduced foraging activity. Forage gaps in July led to reduction in egg‐laying and increased mortality of brood stages at a time when the queen's seasonal egg‐laying rate is at its maximum, leading to colony failure over time. Our results demonstrate that badly timed forage gaps interacting with poor overall forage supply reduce honeybee colony resilience. Existing regulation mechanisms which in principle enable colonies to cope with varying forage supply in a given landscape and year, such as a reduction in egg‐laying, have only a certain capacity. Our results are hypothetical, as they are obtained from simplified landscape settings, but they are consistent with existing empirical knowledge. They offer ample opportunities for testing the predicted effects of forage stress in controlled experiments.     

Brood pheromone regulates foraging activity of honey bees (Hymenoptera: Apidae)

Brood pheromone modulated the foraging behavior of commercial honey bee, Apis mellifera L., colonies pollinating a 10-ha market garden of cucumber, Cucurbita pepo L., and zucchini, Cucumis saticus L., in Texas in late autumn. Six colonies were randomly selected to receive 2000 larval equivalents of brood pheromone and six received a blank control. The ratio of pollen to nonpollen foragers entering colonies was significantly greater in pheromone-treated colonies 1 h after treatment. Pheromone-treated foragers returned with pollen load weights that were significantly heavier than controls. Pollen returned by pheromone-treated foragers was 43% more likely to originate from the target crop. Number of pollen grains washed from the bodies of nonpollen foragers from pheromone-treated colonies was significantly greater than controls and the pollen was 54% more likely to originate from the target crop. Increasing the foraging stimulus environment with brood pheromone increased colony-level foraging and individual forager efforts. Brood pheromone is a promising technology for increasing the pollination activity and efficiency of commercial honey bee colonies.     

Changing colony growth rates inCamponotus floridanus as a behavioral response to conspecific presence (Hymenoptera: Formicidae)

Previous work with the antCamponotus floridanus demonstrated that perception of competition can be clearly differentiated from effects of mortality and decreased resources. That is, brood biomass in ant colonies decreases as a consequence of a behavioral decision(s) rather than because of limited food availability or reduced numbers of brood tenders. The experiments presented here extend that work. Under experimental conditions, colony growth inC. floridanus is modified by distance between brood and unrelated conspecifics and by worker age distribution. When nonnestmates are encountered at the nest versus at a separate foraging site, less brood is maintained by a colony. Although colonies with older workers maintain a brood biomass similar to that of colonies with younger workers, that biomass is concentrated in fewer, larger, more rapidly maturing larvae. These effects seem to be due entirely to worker control.     

Communication between the fungus garden and workers of the leaf-cutting ant, Atta sexdens rubropilosa, regarding choice of substrate for the fungus

Bait made from orange peel, containing the fungicide cycloheximide, was initially harvested by workers of Atta sexdens rubropilosa (Forel) and incorporated into the fungus garden as substrate for the fungus. The bait was subsequently rejected by the worker ants days later. Exposure of the fungus to cycloheximide, in laboratory sub-colonies, resulted in the fungus being ‘stressed’. By interchanging normal fungus garden with ‘stressed’ fungus garden, a change in the foraging behaviour of the workers was evident. –Two hypotheses to explain the behavioural changes were tested: a volatile semiochemical is produced by the fungus which affects the foragers directly, or contact between workers (and fungus garden) is necessary for information regarding fungal substrate to be transmitted through the worker force. When pairs of sub-colonies were connected (one colony of each pair exposed to cycloheximide in the bait) and workers were initially prevented from passing from one colony to the other, one colony continued to forage on orange bait while the other did not. When both colonies were allowed to make full contact then both colonies failed to accept orange bait. This discounted the first hypothesis, but supported the second, as a highly volatile chemical should be able to diffuse between colonies. When large foragers were prevented from making contact with the second colony, the information may be communicated by smaller workers.     

Non-lethal sampling of DNA from bumble bees for conservation genetics

Summary Non-lethal sampling of DNA from individuals in wild populations will often be required for studies of the conservation genetics of social insects, since it avoids destroying members of scarce or declining species. We investigated the effectiveness and consequences of methods of non-lethal sampling of DNA from bumble bee workers. In an experiment with two captive and confined Bombus terrestris colonies, we found that, unlike sampling haemolymph, sampling the terminal portion of the tarsus of a mid-leg of a worker reliably yielded amplifiable microsatellite DNA and did not significantly reduce worker survivorship. In a further experiment with four B. terrestris colonies whose workers were allowed to forage freely at flowers in the external environment, tarsal sampling of either a mid-leg or a hind-leg had no significant effects on worker survivorship, the mean body mass of foraging workers, the frequency or duration of foraging trips, mass of pollen loads or mass of nectar loads. We therefore suggest that tarsal sampling of either a mid-leg or a hind-leg is an effective and acceptable means of non-lethally sampling DNA from workers in wild populations of bumble bees, because effects on individual and colony performance are likely to be absent or minimal.Received 2 December 2002; revised 10 April 2003; accepted 25 April 2003.     

A mechanical signal biases caste development in a social wasp

Understanding the proximate mechanisms of caste development in eusocial taxa can reveal how social species evolved from solitary ancestors. In Polistes wasps, the current paradigm holds that differential amounts of nutrition during the larval stage cause the divergence of worker and gyne (potential queen) castes. But nutrition level alone cannot explain how the first few females to be produced in a colony develop rapidly yet have small body sizes and worker phenotypes. Here, we provide evidence that a mechanical signal biases caste toward a worker phenotype. In Polistes fuscatus, the signal takes the form of antennal drumming (AD), wherein a female trills her antennae synchronously on the rims of nest cells while feeding prey-liquid to larvae. The frequency of AD occurrence is high early in the colony cycle, when larvae destined to become workers are being reared, and low late in the cycle, when gynes are being reared. Subjecting gyne-destined brood to simulated AD-frequency vibrations caused them to emerge as adults with reduced fat stores, a worker trait. This suggests that AD influences the larval developmental trajectory by inhibiting a physiological element that is necessary to trigger diapause, a gyne trait.     

Perception of the pollen need by foragers in a honeybee colony

Honeybees, Apis mellifera, adjust their pollen foraging activity according to the need for pollen within the colony, determined by the amount of stored pollen and young brood present in the hive. To clarify how pollen foragers detect the supply of pollen, we followed individual honeybees while they were returning with pollen. Pollen foragers deposited their loads on the frame where most of the unsealed brood was, independent of the position of this frame within the hive. They also inspected more cells on that frame and spent most of their time there, indicating that pollen foragers may individually evaluate the pollen requirements of the colony. In 18 normal-sized colonies we also tested whether olfactory cues provided by a frame of hungry young brood or an additional pollen frame covered by cages affect foraging activity. These experiments showed that olfactory stimulation within the colony is insufficient to increase or decrease the foraging effort, but suggest that foragers must have direct contact with the brood and pollen area to regulate their foraging activity according to the conditions in the colony. The different mechanisms by which foragers may gather the information about pollen supply are discussed. Copyright 2000 The Association for the Study of Animal Behaviour.     

不同饲料对小峰熊蜂工蜂群发育的影响

糖和花粉对熊蜂的生长发育和繁殖起重要作用。本研究测定和分析了2种糖源（白砂糖、蜂蜜）和5种花粉（杏花粉、油菜花粉、向日葵花粉、玉米花粉、茶花粉）的饲料组合对小峰熊蜂Bombus hypocrita的无王工蜂群寿命、产卵前期时间、产卵量、幼虫拖出数、卵杯数、雄蜂出房时间和子代雄蜂数量的影响。结果表明： 只饲喂糖类食物时, 工蜂寿命显著短于有花粉的饲料组, 而且工蜂不产卵。含有花粉的各饲料组之间工蜂寿命存在显著差异, 饲喂玉米花粉的工蜂寿命小于其他4种花粉组。在工蜂产卵前期时间和卵杯数量方面, 10种含有花粉的饲料组之间差异不显著; 但在产卵量、幼虫拖出数、雄蜂出房时间和子代雄蜂数量等方面, 各饲料组之间差异较大。白砂糖和杏花粉组的产卵量最高; 杏花粉组和茶花粉组的幼虫拖出数量显著低于其他花粉组; 油菜花粉组和杏花粉组的雄蜂出房时间显著短于其他花粉组; 杏花粉组的子代雄蜂数量显著高于其他饲料组。结果显示不同饲料组合对小峰熊蜂工蜂群的发育影响很大。因此建议在熊蜂的人工繁育过程中, 在不同的发育阶段应给予不同的饲料配方。     

Do bumblebees always forage as much as they could?

Summary Bumblebees must forage to provide food to the colony. However, foraging is costly as worker longevity is inversely related to foraging effort. Given this trade-off, workers from colonies with abundant food supplies could either maintain foraging to increase reserves for future use or forage less to avoid the associated costs. We tested these hypotheses over one summer, using 13 pairs of field colonies of Bombus impatiens. Half of the colonies were provided with a sucrose solution ad libitum and pollen at regular intervals throughout their entire development, while the other half served as controls. We measured the forager activity rates in colonies with infra-red motion detectors fit in nest box entrances. However, due to reasons beyond our control (loss of the queen, usurpation by Psithyrus, debris in the entrance tunnel, etc.), we could use data from only two pairs of colonies for the analysis. Food supplemented colonies had a forager activity rate per worker 25% lower than controls which supports the hypothesis that workers reduce risks when given the opportunity.Received 15 May 2003; revised 15 January 2004; accepted 19 February 2004.     

How average relatedness affects the frequency of trophallaxis between workers in an experimental colony of the polygynous ant,Camponotus yamaokai

To determine whether workers engage in trophallactic behavior preferentially toward highly related nestmates, we examined these behaviors between workers in colonies consisting of both kin and non-kin (mixed colony) and of only kin (pure colony). These behaviors were observed under sufficiently fed and in both starvation and normal feeding conditions. Workers in mixed colonies exhibited trophallactic behaviors equally toward kin and non-kin in both experimental conditions. The frequency of trophallaxis between workers in mixed colonies, however, tended to be lower than that in pure colonies, is especially under starvation conditions. The low frequency of trophallaxis among workers discussed in terms of the low degree of genetic relatedness among nestmates in mixed colonies.     

Major Quantitative Trait Loci Affecting Honey Bee Foraging Behavior

We identified two genomic regions that affect the amount of pollen stored in honey bee colonies and influence whether foragers will collect pollen or nectar. We selected for the amount of pollen stored in combs of honey bee colonies, a colony-level trait, and then used random amplified polymorphic DNA (RAPD) markers and interval mapping procedures with data from backcross colonies to identify two quantitative trait loci (pln1 and pln2, LOD 3.1 and 2.3, respectively). Quantitative trait loci effects were confirmed in a separate cross by demonstrating the cosegregation of marker alleles with the foraging behavior of individual workers. Both pln1 and pln2 had an effect on the amount of pollen carried by foragers returning to the colony, as inferred by the association between linked RAPD marker alleles, D8-.3f and 301-.55, and the individual pollen load weights of returning foragers. The alleles of the two marker loci were nonrandomly distributed with respect to foraging task. The two loci appeared to have different effects on foraging behavior. Individuals with alternative alleles for the marker linked to pln2 (but not pln1) differed with respect to the nectar sugar concentration of their nectar loads.     

Size variation and foraging rate in bumblebees (Bombus terrestris)

Summary: Size polymorphism is an important life history trait in bumblebees with strong impact on individual behavior and colony organization. Within a colony larger workers tend to serve as foragers, while smaller workers fulfill in-hive tasks. It is often assumed that size-dependent division of labor relates to differences in task performance. In this study we examined size-dependent interindividual variability in foraging, i.e. whether foraging behavior and foraging capability of bumblebee workers are affected by their size. We observed two freely foraging Bombus terrestris colonies and measured i) trip number, ii) trip time, iii) proportion of nectar trips, and iv) nectar foraging rate of different sized foragers. In all observation periods large foragers exhibited a significantly higher foraging rate than small foragers. None of the other three foraging parameters was affected by worker size. Thus, large foragers contributed disproportionately more to the current nectar influx of their colony. We provide a detailed discussion of the possible proximate mechanisms underlying the differences in foraging rate.