Spatial foraging patterns and colony energy status in the African honey bee,Apis mellifera scutellata

The relationship between changes in foraging patterns (inferred from waggle dance activity) and colony energy status (inferred from brood rearing activity, food storage, and colony weight) was examined for the African honey bee during a period of relative resource abundance and resource dearth. When resources were more abundant mean foraging distances (about 400 m) and foraging areas (4–5 km²) were small, and colonies recruited to 12–19 different sites per day. Colony foraging ranges and sites visited increased slightly during the dearth period, yet foraging continued to be concentrated within less than 10 km². The degree to which fluctuations in foraging patterns were correlated with colony energy status varied with the availability of floral resources. During periods of relative forage abundance, increases in foraging range and number of sites visited were significantly correlated with increases in brood rearing and colony weight. In contrast, colonies examined during periods of resource dearth exhibited no correlations between foraging areas, foraging distances, and fluctuations in brood rearing, food storage, or colony weight. Thus, during dearth periods colonies may not be able to coordinate foraging patterns with changes in colony energy status.     

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.     

Assessment of Chronic Sublethal Effects of Imidacloprid on Honey Bee Colony Health

Here we present results of a three-year study to determine the fate of imidacloprid residues in hive matrices and to assess chronic sublethal effects on whole honey bee colonies fed supplemental pollen diet containing imidacloprid at 5, 20 and 100 μg/kg over multiple brood cycles. Various endpoints of colony performance and foraging behavior were measured during and after exposure, including winter survival. Imidacloprid residues became diluted or non-detectable within colonies due to the processing of beebread and honey and the rapid metabolism of the chemical. Imidacloprid exposure doses up to 100 μg/kg had no significant effects on foraging activity or other colony performance indicators during and shortly after exposure. Diseases and pest species did not affect colony health but infestations of Varroa mites were significantly higher in exposed colonies. Honey stores indicated that exposed colonies may have avoided the contaminated food. Imidacloprid dose effects was delayed later in the summer, when colonies exposed to 20 and 100 μg/kg experienced higher rates of queen failure and broodless periods, which led to weaker colonies going into the winter. Pooled over two years, winter survival of colonies averaged 85.7, 72.4, 61.2 and 59.2% in the control, 5, 20 and 100 μg/kg treatment groups, respectively. Analysis of colony survival data showed a significant dose effect, and all contrast tests comparing survival between control and treatment groups were significant, except for colonies exposed to 5 μg/kg. Given the weight of evidence, chronic exposure to imidacloprid at the higher range of field doses (20 to 100 μg/kg) in pollen of certain treated crops could cause negative impacts on honey bee colony health and reduced overwintering success, but the most likely encountered high range of field doses relevant for seed-treated crops (5 μg/kg) had negligible effects on colony health and are unlikely a sole cause of colony declines.     

Division of Labor Associated with Brood Rearing in the Honey Bee: How Does It Translate to Colony Fitness?

Division of labor is a striking feature observed in honey bees and many other social insects. Division of labor has been claimed to benefit fitness. In honey bees, the adult work force may be viewed as divided between non-foraging hive bees that rear brood and maintain the nest, and foragers that collect food outside the nest. Honey bee brood pheromone is a larval pheromone that serves as an excellent empirical tool to manipulate foraging behaviors and thus division of labor in the honey bee. Here we use two different doses of brood pheromone to alter the foraging stimulus environment, thus changing demographics of colony division of labor, to demonstrate how division of labor associated with brood rearing affects colony growth rate. We examine the effects of these different doses of brood pheromone on individual foraging ontogeny and specialization, colony level foraging behavior, and individual glandular protein synthesis. Low brood pheromone treatment colonies exhibited significantly higher foraging population, decreased age of first foraging and greater foraging effort, resulting in greater colony growth compared to other treatments. This study demonstrates how division of labor associated with brood rearing affects honey bee colony growth rate, a token of fitness.     

Colony migration in the tropical honey beeApis dorsata F. (Hymenoptera: Apidae)

Summary To obtain insights into the organization and adaptive significance of seasonal migration by colonies of the giant honey bee,Apis dorsata, we monitored the arrivals and departures of colonies in a rain forest habitat in northeastern Thailand, compared patterns of honey bee abundance with other measures of habitat variability, and observed the role of dance communication in organizing the migratory departure of a colony. Colonies arrived in the area during the end of the dry season, reproduced, and then departed early in the rainy season. During the immigration phase, early-arriving colonies stayed only temporarily, as if assessing habitat quality. Colonies departing after a long stay always left barren combs behind, suggesting that they had left in response to deteriorating resource quality. These observations support the idea that migration allows colonies to track seasonally varying resources in different regions. Our observations of a colony preparing for migration revealed that the dance language is involved in organizing the colony's departure, but that dancers signal only the direction to be taken, rather than, as in dances to feeding sites, both the direction and distance of a particular location.     

Survival of honeybees in cold climates: the critical timing of colony growth and reproduction

Abstract. 1. The adaptive significance of the timing of growth and reproduction by honeybee, Apis mellifera L., colonies in cold climates was studied by describing the seasonal patterns of food storage, brood rearing, and swarming, and then observing the consequences of experimentally perturbing the seasonal cycles of brood rearing and swarming.
2. Colonies consume large amounts of food over winter (20+ kg of honey), but have only a brief period (about 14 weeks) for food collection each year.
3. The honeybee's striking habits of starting brood rearing in midwinter and swarming in late spring evidently help colonies achieve maximum use of the short summer season. Colonies whose onset of-brood rearing was experimentally postponed until early spring showed greatly retarded colony growth and swarming. Other experiments demonstrated that late swarms starve more often during winter than do early swarms.
4. We conclude that the timings of colony growth and reproduction are essential elements in the honeybee's suite of adaptations for winter survival.     

Preparation for disturbance-induced absconding of Cape honeybee colonies (Apis mellifera capensis Esch.)

African honeybees, Apis mellifera, are characterised by frequent disturbance-induced absconding. However, the effectiveness in preparation before such disturbance-induced absconding has not been rigorously quantified yet. We investigated the effectiveness of preparation for disturbance-induced absconding by evaluating colony phenotypes prior to and after absconding in ten colonies of the Cape honeybee, A. m. capensis. Seven non-absconding colonies at the same apiary were used as controls. While seven absconded colonies left neither stores nor brood behind, three colonies abandoned only a small area of honey, pollen, open or capped brood. At the end of the observations, the control colonies still had pollen and honey stores and brood. The mean reduction rate between a major disturbance and the absconding event was 0.052 ± 0.018 cm² stores and open brood per worker per day. Our results demonstrate that disturbance-induced absconding can also occur with preparation, if the disturbance is not highly destructive and enough time for preparation is available. We conclude that Cape honeybee colonies can show a considerable high effectiveness in their preparation before disturbance-induced absconding, which limits the loss of colony resources. In light of the general high mobility of African honeybee colonies such an efficient behaviour is probably adaptive. Received 22 December 2004; revised 3 June 2005; accepted 13 June 2005.     

A Four-Year Field Program Investigating Long-Term Effects of Repeated Exposure of Honey Bee Colonies to Flowering Crops Treated with Thiamethoxam

Neonicotinoid residues in nectar and pollen from crop plants have been implicated as one of the potential factors causing the declines of honey bee populations. Median residues of thiamethoxam in pollen collected from honey bees after foraging on flowering seed treated maize were found to be between 1 and 7 µg/kg, median residues of the metabolite CGA322704 (clothianidin) in the pollen were between 1 and 4 µg/kg. In oilseed rape, median residues of thiamethoxam found in pollen collected from bees were between <1 and 3.5 µg/kg and in nectar from foraging bees were between 0.65 and 2.4 µg/kg. Median residues of CGA322704 in pollen and nectar in the oilseed rape trials were all below the limit of quantification (1 µg/kg). Residues in the hive were even lower in both the maize and oilseed rape trials, being at or below the level of detection of 1 µg/kg for bee bread in the hive and at or below the level of detection of 0.5 µg/kg for hive nectar, honey and royal jelly samples. The long-term risk to honey bee colonies in the field was also investigated, including the sensitive overwintering stage, from four years consecutive single treatment crop exposures to flowering maize and oilseed rape grown from thiamethoxam treated seeds at rates recommended for insect control. Throughout the study, mortality, foraging behavior, colony strength, colony weight, brood development and food storage levels were similar between treatment and control colonies. Detailed examination of brood development throughout the year demonstrated that colonies exposed to the treated crop were able to successfully overwinter and had a similar health status to the control colonies in the following spring. We conclude that these data demonstrate there is a low risk to honey bees from systemic residues in nectar and pollen following the use of thiamethoxam as a seed treatment on oilseed rape and maize.     

Net reproductive rate of unmanaged honeybee colonies, (Apis mellifera L.)

Summary The net reproductive rate of unmanaged honeybee colonies has never been fully determined for honey bees in temperate climates. In this study, five overwintered colonies in Kansas, USA, were allowed to swarm naturally (Winston. 1980). These colonies and their swarms were studied over the winter (i.e. one generation). The net reproductive rateR ₀ was estimated to be 2.18. Afterswarms were found to contribute substantially (41.2%) to this net reproductive rate. The autumn and spring food reserves and brood areas of established colonies and colonies established from prime swarms and afterswarms are compared. Winter survival of afterswarms was related to autumn honey stores, and the brood areas of surviving afterswarms were smaller than those of prime swarms or established colonies.     

The provision of supplementary food to hive bees

Sucrose syrup is less satisfactory than honey as a winter and spring food for bees: a mixture of approximately equal parts of honey and syrup is of almost the same value as honey alone.
Feeding confined to early autumn induced the strongest spring development of colonies; syrup feeding in the spring may retard colony development, and food supplied at this time is apparently wasted. Feeding with syrup and pollen is advantageous only when a colony is lacking in stores of carbohydrate and protein.
A total of four British Standard brood combs full of pollen provides sufficient protein for a colony on B.S. equipment from autumn until April: the best results were obtained by providing colonies on eleven B.S. combs, in early autumn, with 35-40 lb. of honey, or honey and concentrated sucrose syrup, and four brood combs full of pollen.     

Seasonal patterns of foraging activity in colonies of the African honey bee,Apis mellifera scutellata,in Africa

Summary Seasonal foraging patterns were investigated using six observation colonies maintained in the Okavango Delta, Botswana. Pollen collection, flight from the hive, and recruitment for pollen and nectar sources occurred throughout the 11 months of the study. However, the distribution of foraging activity throughout the day changed seasonally. Colonies emphasized recruitment for pollen sites throughout most of the year. Brood production occurred in all months except May, and there was a significant, positive correlation between the proportion of recruitment activity devoted to pollen sources and the amount of brood comb in the colonies. The seasonal foraging patterns ofscutellata in the Okavango were similar to those of Africanized honey bees in the neotropics. The extended foraging season and emphasis on pollen collection may be associated with the high swarming rates and migrational movements of tropical honey bees.     

Regulation of nectar collection in relation to honey storage levels by honey bees, Apis mellifera

Honey bees collect distinct nutrient sources in the form ofnectar (energy) and pollen (nitrogen). We investigated the effectof varying energy stores on nectar and pollen foraging. We foundno significant changes in nectar foraging in response to changesin honey storage levels within colonies. Individual foragersdid not vary activity rates or nectar load sizes in responseto changes in honey stores, and colonies did not increase nectarintake rates when honey stores within the hive were decreased.This result contrasts with pollen foraging behavior, which isextremely sensitive to colony state. Our data show that individualforaging decisions during nectar collection and colony regulationof nectar intake are distincdy different from pollen foraging.The behavior of honey bees illustrates that foraging strategy(and therefore foraging models) can incorporate multiple currencies,including both energy and protein intake.[Behav Ecol 7: 286–291(1996)]     

The effects of land use on honey bee (Apis mellifera) population density and colony strength parameters in the Eastern Cape, South Africa

Livestock farming in the Eastern Cape, South Africa is a common land use practice that has affected the biodiversity of plants and animals in the region negatively. Indigenous populations of wild honey bee (Apis mellifera) colonies also may suffer. Recently, farmers in the Eastern Cape have been converting their farms to game reserves as ecotourism attractions and nature conservation lands. Consequently, the goal of this research was to determine if land use habits (livestock farming and conversion to game reserves) in the Eastern Cape affect honey bee colony population density and colony strength parameters. A series of indices were developed to compare the relative population densities of colonies in two or more areas by counting the number of foraging bees and number of bee lines established at feeding stations. Wild colonies on farms and reserves were located and sampled to determine land use effects on colony strength parameters including total area of comb in the colony, the area of comb containing stored honey, pollen, and brood, adult bee population, weight per bee, and the colony nest cavity volume ratio. When viewed collectively, the data indicated that land use practices have affected honey bee nesting dynamics in the Eastern Cape. Trends in the data suggested that colonies nesting on the reserves may occur in greater densities than those nesting on livestock farms, though they do not appear to be healthier. Hopefully, this work will be continued since honey bee conservation in areas where they are native is crucial to the health of agriculture and whole ecosystems globally.     

Social behavior and brood decline in reproductive-phase colonies ofBelonogaster petiolata (Degeer) (Hymenoptera: Vespidae)

Colonies ofBelonogaster petiolata in Gauteng (South Africa) produced reproductive offspring (gynes and males) in late January and early February of each nesting season; their appearance was associated with a decline in worker and brood numbers. Brood decline could commence in the presence of a dominant, reproductively active queen, and loss or removal of the queen was not followed directly by cessation of nest growth and brood care. An older worker usually took over the α-position in queenless colonies. Several factors appear to contribute to brood decline and, ultimately, termination of the colony cycle in this species. These include (1) cessation of the supply of solid food to colonies (and particularly their larvae) during the reproductive phase, (2) a decrease in the worker/larva ratio during the latter phase due to the progressive loss of workers, (3) increasing number of gynes and males, and (4) an adult priority over food reception from foragers.     

Evaluation of drone brood removal for management of Varroa destructor (Acari: Varroidae) in colonies of Apis mellifera (Hymenoptera: Apidae) in the northeastern United States

The efficacy of drone brood removal for the management of Varroa destructor Anderson & Trueman in colonies of the honey bee, A. mellifera L., was evaluated. Colonies were treated with CheckMite+ in the fall of 2002. The following spring, quantities of bees and brood were equalized, but colonies were not retreated. The brood nest of each colony consisted of 18 full-depth worker combs and two full-depth drone combs. Each worker comb had <12.9 cm2 of drone cells. Standard management practices were used throughout the season. Colonies were randomly assigned to one of two groups. In the control group, drone combs remained in place throughout the season. In the treatment group, drone combs were removed on 16 June, 16 July, 16 August, and 16 September and replaced with empty drone combs (16 June) or with drone combs removed on the previous replacement date. In the early fall, the average mite-to-bee ratio in the control group was significantly greater than the corresponding ratio in the treatment group. Drone brood removal did not adversely affect colony health as measured by the size of the worker population or by honey production. Fall worker populations were similar in the two groups. Honey production in treatment colonies was greater than or similar to production in control colonies. These data demonstrate that drone brood removal can serve as a valuable component in an integrated pest management program for V. destructor and may reduce the need for other treatments on a colony-by-colony basis.     

Spatial foraging patterns of the African honey bee,Apis mellifera scutellata

Recruitment patterns were investigated for the African honey bee in the Okavango River Delta, Botswana. The waggle dances of two observation colonies maintained in the field were monitored and used to construct maps of daily recruitment activity. These maps revealed that the African colonies frequently adjusted the allocation of recruits among food patches, recruited for 16–17 different food sites/day over areas of 55–80 km ² ,and concentrated the majority of recruitment within 1 km of the hives (median foraging distances for the two colonies were 295 and 563 m). In both colonies pollen foragers were more abundant than nectar foragers, and pollen sources indicated by waggle dancers were significantly closer to the hives than nectar sources. Compared to the recruitment patterns of temperate climate colonies, the African colonies had smaller recruitment areas, smaller mean recruitment distances, and a greater emphasis on pollen foraging. These differences may be related to the contrasting survival strategies followed by tropical-versus temperate-climate honey bees.     

Seasonal cycles of growth,development and movement of the African honey bee,Apis mellifera scutettata,in Africa

Summary The relationship between the annual colony cycle and seasonal patterns of forage availability was investigated for the African honey bee,Apis mellifera scutellata, in the Okavango River Delta, Botswana. The annual cycle occurred in three distinct periods. The swarming season occurred from October-November, following two to three months of intense brood production, and coincided with the end of peak forage abundance. The migration season occurred from November-May and coincided with reduced and variable floral resources. During the migration season, brood production and food storage were generally low but quite variable from month to month, and swarms passing over the study area at this time traveled in an easterly direction. The migration season was followed by the establishment period (June-September), in which large numbers of new colonies traveling from the west moved into the study area. The establishment period coincided with, and slightly preceded, the period of peak forage abundance, and colonies devoted resources collected at this time almost entirely to brood rearing, which culminated in swarm production. The data suggest that honey bee colonies in the Okavango are mobile and gear their reproduction and movement to seasonally shifting resource pattern.     

Division of labor inPonera pennsylvannica (Formicidae: Ponerinae)

Summary We examined division of labor and colony demography in the antPonera pennsylvannica. Observation of three colonies with individually marked workers revealed a high degree of interindividual behavioral variation and a rough but consistent division of labor between brood tenders and foragers. This division was present both in colonies consisting entirely of workers produced in the previous summer and in colonies containing freshly eclosed ants. Two colonies showed typical age-based polyethism, with young ants focusing on brood care and overwintered ants on foraging. No such age basis was detected in the third colony. This difference may relate to variability in brood production schedules. Colonies showing temporal polyethism had two peaks of brood production and thus had relatively large brood populations when the first young workers eclosed, while the third colony had only one peak and little brood for the young workers to tend. Even if young ants have a lower threshold for brood care, it may have been concealed in the latter situation. Demographic data indicate that natural colonies produce one brood per year and that workers typically eclose into colonies with relatively low brood care demands. This suggests that overwintered workers do most of a colony's work and that the division of labor among overwintered ants is the more important one under natural conditions. The basis of this division is as yet unknown. These results also suggest that small colony size, univoltine brood schedule and a close association between foraging and brood care do not preclude division of labor among specialized castes, as has been suggested for another ponerine species (Traniello 1978).     

The distribution of Paenibacillus larvae spores in adult bees and honey and larval mortality, following the addition of American foulbrood diseased brood or spore-contaminated honey in honey bee (Apis mellifera) colonies

Within colony transmission of Paenibacillus larvae spores was studied by giving spore-contaminated honey comb or comb containing 100 larvae killed by American foulbrood to five experimental colonies respectively. We registered the impact of the two treatments on P. larvae spore loads in adult bees and honey and on larval mortality by culturing for spores in samples of adult bees and honey, respectively, and by measuring larval survival. The results demonstrate a direct effect of treatment on spore levels in adult bees and honey as well as on larval mortality. Colonies treated with dead larvae showed immediate high spore levels in adult bee samples, while the colonies treated with contaminated honey showed a comparable spore load but the effect was delayed until the bees started to utilize the honey at the end of the flight season. During the winter there was a build up of spores in the adult bees, which may increase the risk for infection in spring. The results confirm that contaminated honey can act as an environmental reservoir of P. larvae spores and suggest that less spores may be needed in honey, compared to in diseased brood, to produce clinically diseased colonies. The spore load in adult bee samples was significantly related to larval mortality but the spore load of honey samples was not.     

The relationship between population size,amount of brood,and individual foraging behaviour in the honey bee,Apis mellifera L.

This study experimentally examines the relationship between colony state and the behaviour of individual pollen and nectar foragers in the honey bee, Apis mellifera L. In the first experiment we test the prediction that individual pollen foragers from colonies with higher brood quantities should exhibit a greater work effort for pollen resources than individual pollen foragers from colonies with low brood quantities. Eight colonies were assigned into two treatment groups; HIGH brood colonies were manipulated to contain 9600±480 cm² brood area; LOW brood colonies were manipulated to contain 1600±80 cm² brood area. We measured colony brood levels over the course of the experiment and collected individual pollen loads from returning pollen foragers. We found that, while colonies remained significantly different in brood levels, individual pollen foragers from HIGH brood colonies collected larger loads than individuals from LOW brood colonies. In the second experiment we investigated the influence of colony size on the behaviour of individual nectar foragers. We assigned eight colonies to two treatment groups; LARGE colonies were manipulated to contain 35000±1700 adult workers with 3500±175 cm² brood area, and SMALL colonies were manipulated to contain 10000±500 adult workers with 1000±50 cm² brood area. We observed foraging trips of individually marked workers and found that individuals from LARGE colonies made longer foraging trips than those from SMALL colonies (LARGE: 1666.7±126.4 seconds, SMALL: 1210.8±157.6 seconds), and collected larter nectar loads (LARGE: 19.2±1.0 l, SMALL: 14.6±0.8 l). These results indicate that individual nectar foragers from LARGE colonies tend to work harder than individuals from SMALL colonies. Both experiments indicate that the values of nectar and pollen resources to a colony change depend on colony state, and that individual foragers modify their behaviour accordingly.