Dwindling pollen resources trigger the transition to broodless populations of long-lived honeybees each autumn

Abstract.  1. Each autumn in northern regions, honeybee colonies shift from populations of short-lived workers that actively rear brood to broodless populations of long-lived winter bees. To determine if dwindling pollen resources trigger this transition, the natural disappearance of external pollen resources was artificially accelerated or delayed and colonies were monitored for effects on the decline in brood-rearing activity and the development of populations of long-lived winter bees.
2. Delaying the disappearance of pollen resources postponed the decline in brood rearing in colonies. Colonies with an extended supply of pollen reared workers longer into October before brood rearing ended than control colonies or colonies for which pollen supply was cut short artificially in autumn.
3. Colonies with extended pollen supply produced more workers throughout autumn than colonies with less pollen, but the development of the population of long-lived winter bees was delayed until relatively later in autumn. Colonies produced similar numbers of winter bees, regardless of the timing of the disappearance of pollen resources.
4. Mean longevity of autumn-reared workers was inversely related to the amount of brood remaining to be reared in colonies when workers eclosed. Consequently, long-lived workers did not appear in colonies until brood rearing declined, which in turn was controlled by the availability of pollen.
5. Dwindling pollen resources provide a powerful cue that initiates the transition to populations of broodless winter bees because it directly affects the brood-rearing capacity of colonies and indirectly indicates deteriorating environmental conditions associated with the approach of winter.     

Rates of brood development in a social wasp: effects of colony size and parasite infection

Summary Colonies of eusocial insect species are most vulnerable during the founding stage. Many species have evolved means to minimize the length of the founding, or pre-emergence, stage by accelerating the rate of development of the first worker offspring. Other things being equal, the sooner a colony can begin producing workers, the less the risk of colony failure, the steeper the growth curve of the colony during the ergonomic stage, and the larger the colony will be at reproductive maturity. Swarm-founding species, whose founding units consist of hundreds or thousands of workers, may face less selection pressure to minimize the duration of the founding stage than independent-founding species. However, swarm size varies within species, and small swarms face greater risk of extinction during the founding stage than large swarms. This consideration predicts that within a species, small swarms should have shorter founding stages than large swarms, likely by rearing a small group of precocious brood. On the other hand, evidence that large social groups organize colony labor more efficiently, gather resources more predictably, and homeostatically maintain physical conditions inside the nest within narrower ranges, predicts that larger groups should rear all brood more rapidly and therefore have shorter founding stages. To test whether small or large swarms have shorter founding stages in colonies of Polybia occidentalis, a Neotropical swarm-founding wasp, we measured brood development rates in colonies collected after 28 days of development, just short of the minimum egg-to-adult development time. We found that as size increased across colonies, pre-emergence times decreased, mean age of pupae in the nest increased, and median age of the brood (larvae + pupae) increased. That is, brood developed significantly faster in large colonies than in small ones. Using these same measures, we also found that infection by a gregarine parasite increased brood development time, independently of colony size.Received 10 April 2003; revised 29 October 2003; accepted 21 November 2003.     

Effects of brood pheromone (SuperBoost) on consumption of protein supplement and growth of honey bee (Hymenoptera: Apidae) colonies during fall in a northern temperate climate

Honey bee, Apis mellifera L. (Hymenoptera: Apidae), nutrition is vital for colony growth and maintenance of a robust immune system. Brood rearing in honey bee colonies is highly dependent on protein availability. Beekeepers in general provide protein supplement to colonies during periods of pollen dearth. Honey bee brood pheromone is a blend of methyl and ethyl fatty acid esters extractable from cuticle of honey bee larvae that communicates the presence of larvae in a colony. Honey bee brood pheromone has been shown to increase protein supplement consumption and growth of honey bee colonies in a subtropical winter climate. Here, we tested the hypothesis that synthetic brood pheromone (SuperBoost) has the potential to increase protein supplement consumption during fall in a temperate climate and thus increase colony growth. The experiments were conducted in two locations in Oregon during September and October 2009. In both the experiments, colonies receiving brood pheromone treatment consumed significantly higher protein supplement and had greater brood area and adult bees than controls. Results from this study suggest that synthetic brood pheromone may be used to stimulate honey bee colony growth by stimulating protein supplement consumption during fall in a northern temperate climate, when majority of the beekeepers feed protein supplement to their colonies.     

Performance evaluation of indigenous and exotic honey bee (Apis mellifera L.) races in Assir region,southwestern Saudi Arabia

This study was conducted in the Assir region of southwestern Saudi Arabia to compare the activities of honeybee colonies of indigenous Apis mellifera jemenitica (AMJ) and imported Apis mellifera carnica (AMC) during the late summer and autumn of 2009 and 2010. The results showed that the workers of the two races exhibited relatively similar forage timings throughout the period of study (August–November). The highest numbers of foraged workers were recorded at 6:00 am, 10:00 am and 6:00 pm, while the lowest numbers were recorded at 8:00 am, 12:00 pm and 4:00 pm. Although foraging activity was negatively affected by decreased temperature, AMJ was more resistant to cold than AMC. In the first season, the smallest amount of worker brood rearing was recorded in August, and the highest amount of rearing occurred in November in both races. In the second season, the smallest amount of brood was observed in October, and the largest amount of brood was observed in November. Brood rearing and pollen collecting was significantly (P < 0.05) higher in AMJ compared with AMC, while AMC stored significantly (P < 0.05) more honey than AMJ during the tested periods. In AMJ colonies, a positive significant correlation was observed between the area of the sealed worker brood and stored pollen, while a negative but nonsignificant correlation was observed between the area of the sealed worker brood and surplus honey. In the AMC colonies, a positive significant correlation was observed between the area of the sealed brood and the stored pollen and surplus honey.     

Comparison of the activity and productivity of Carniolan (Apis mellifera carnica Pollmann) and Yemeni (Apis mellifera jemenitica Ruttner) subspecies under environmental conditions of the Al-Ahsa oasis of eastern Saudi Arabia

This study was conducted at the apiary of the Agricultural and Veterinary Training and Research Station of King Faisal University in the Al-Ahsa oasis of eastern Saudi Arabia. We performed a comparison between Carniolan (Apis mellifera carnica Pollmann) and Yemeni (Apis mellifera jemenitica Ruttner) honeybee races to determine the monthly fluctuations in foraging activity, pollen collection, colony growth and honey yield production under the environmental conditions of the Al-Ahsa oasis of eastern Saudi Arabia. We found three peaks in the flight activity of the two races, and the largest peaks occurred during September and October. Compared to Carniolan bee colonies, the performance of Yemeni bee colonies was superior in terms of stored pollen, worker and drone brood rearing, and the adult population size. The Carniolan bee colonies produced 27.77% and 27.50% more honey than the Yemeni bee colonies during the flow seasons of alfalfa and sidir, respectively, with an average increase of 27.64%. It could be concluded that the race of bees is an important factor affecting the activity and productivity of honeybee colonies. The Yemeni bee race produced more pollen, a larger brood and more bees, which exhibited a longer survival. The imported Carniolan bees can be reared in eastern Saudi Arabia, but the Yemeni bee race is still better.     

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.     

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.     

The impact of caging the queens during the flow season on some biological activities of honeybee colonies

This study was achieved in a private apiary located in a banana farm in Sa El Hagar, Basioun, Gharbia, Egypt from August 15, 2019 to May 25, 2020, including the banana (Musa sp., Musaceae) flow season (August and September) and extend to Egyptian clover (Trifolium alexandrinum L., Fabaceae) flow season (May). The study aimed to evaluate the effect of confining the queen during the banana flow season on the brood rearing, honey yield, and activation of worker's ovaries. Also, we determined the negative impact of caging the queen during the banana flow season on the activity of the colony in brood rearing, storing pollen, and honey yield after releasing the queen on 5 October, extending to the next flow season in May. The obtained results showed that the honeybee colonies with the caged queen produced significantly more honey yield and less brood production than the free queen ones during the banana flow season. Also, the caging of the queen did not affect the colony strength after releasing the queen despite the partial development of the ovaries of some workers, but they did not lay eggs. In addition, releasing the queens suppressed the ovaries of the laying workers. It can be concluded that caging the queen during the banana flow season helps the colonies to produce more honey yield without effect on the colony strength after releasing the queen despite the ovaries development of few workers without egg-laying.     

Influence of virgin queen honeybees (Apis mellifera) on queen rearing and foraging

ABSTRACT. Virgin queens are as effective as mated laying queens at inhibiting colonies from rearing queens but not from producing queen cell cups. Colonies without brood produce fewer queen cell cups than similar colonies that have brood. Colonies without queens forage much less and collect less pollen than with either a mated or virgin queen. Colonies with virgin queens forage as much as those with mated queens but collected less pollen.     

THE INFLUENCE OF INFECTION WITH NOSEMA APIS ON THE ACTIVITIES AND LONGEVITY OF THE WORKER HONEYBEE

Infection of the adult worker honeybee with Nosema apis reduces or obviates brood feeding and causes her to commence foraging earlier than a healthy bee. The length of foraging activity and the total length of life of infected bees is reduced.
In colonies infected with N. apis the rate of brood rearing is severely depressed during April, May and June, the degree of depression being proportional to the percentage infection.
Infection decreases during July, August and September, and consequently the rate of brood rearing increases, but the resulting addition in foraging population is usually too late to increase the honey crop.     

Sociality and sex allocation in an Australian allodapine bee Braunsapis protuberans

Abstract  Herein the first detailed study of sociality and life history in an Australian species of the allodapine bee genus Braunsapis is provided. Colonies of B. protuberans were sampled from the Great Sandy National Park in Queensland from late winter until later summer. Single-female colonies appear to have suffered substantially higher rates of total brood loss, or failure to produce brood, than multifemale colonies, but rearing efficiency in colonies with at least some brood did not differ between these two colony types. There was marked reproductive skew in multifemale colonies and ovary size was strongly linked to relative body size. However, females with smaller body size and smaller ovaries also had lower levels of wing wear, suggesting that they do not specialise in foraging. These patterns are similar to an African species, Braunsapis vitrea , and suggest that colonies may principally comprise a dominant reproductive female who provides food for her juvenile offspring, along with a variable number of subordinate females who may be waiting to inherit their natal nest in the event of their mother's death. However, it is likely that in such events, the remaining female(s) will care for orphaned brood as some single-female colonies contained substantial older brood, even though the sole female had little or no wing wear. Such alloparental care in the case of orphaning may help explain the high level of female bias in sex ratios.     

Larger laboratory colonies consume proportionally less energy and have lower per capita brood production in Temnothorax ants

Colony size can affect individual- and colony-level behavioral and physiological traits in social insects. Changes in behavior and physiology in response to colony growth and development can affect productivity and fitness. Here, we used respirometry to study the relationship between colony size and colony energy consumption in Temnothorax rugatulus ants. In addition, we examined the relationship between colony size and worker productivity measured as per capita brood production. We found that colony metabolic rate scales with colony size to the 0.78 power and the number of brood scales with the number of workers to the 0.49 power. These regression analyses reveal that larger ant colonies use proportionally less energy and produce fewer brood per worker. Our findings provide new information on the relationships between colony size and energetic efficiency and productivity in a model ant genus. We discuss the potential mechanisms giving rise to allometric scaling of metabolic rate in ant colonies and the influence of colony size on energy consumption and productivity in general.     

Strong constraints to independent nesting in a facultatively social bee: quantifying the effects of enemies-at-the-nest

Constraints to independent nesting play a key role in the understanding of social evolution in insects, but the source and the magnitude of such constraints are poorly known for many key taxa. In allodapine bees it is known that solitary nesting females have low rates of successful brood rearing and that this drives selection for cooperative nesting. It has been hypothesized that these constraints are due to the presence of enemies-at-the-nest, such as ants, but no direct link has been demonstrated between such enemies and colony failure. We set up an experiment in which solitary founded nests of an Australian allodapine bee, Exoneura nigrescens, were either protected from non-flying predators or left unprotected, and compared the resulting colony survival and brood production rates. We found that protected colonies have much higher rates of survival and that the constraints to independent nesting are extreme, with a mean of less than one offspring per nest at the end of the brood rearing period. This means that cooperative nesting is essential for this species to persist in its habitat. Received 6 July 2007; revised 5 November 2007; accepted 12 November 2007.     

Effects of Brood Pheromone Modulated Brood Rearing Behaviors on Honey Bee (Apis mellifera L.) Colony Growth

A hallmark of eusociality is cooperative brood care. In most social insect systems brood rearing labor is divided between individuals working in the nest tending the queen and larvae, and foragers collecting food outside the nest. To place brood rearing division of labor within an evolutionary context, it is necessary to understand relationships between individuals in the nest engaged in brood care and colony growth in the honey bee. Here we examined responses of the queen, queen-worker interactions, and nursing behaviors to an increase in the brood rearing stimulus environment using brood pheromone. Colony pairs were derived from a single source and were headed by open-mated sister queens, for a total of four colony pairs. One colony of a pair was treated with 336 μg of brood pheromone, and the other a blank control. Queens in the brood pheromone treated colonies laid significantly more eggs, were fed longer, and were less idle compared to controls. Workers spent significantly more time cleaning cells in pheromone treatments. Increasing the brood rearing stimulus environment with the addition of brood pheromone significantly increased the tempo of brood rearing behaviors by bees working in the nest resulting in a significantly greater amount of brood reared.     

Influence of pollen diet in spring on development of honey bee (Hymenoptera: Apidae) colonies

The effects of changes in spring pollen diet on the development of honey bee, Apis mellifera L. (Hymenoptera: Apidae), colonies were examined in a 3-yr study (2002-2004). Pollen-supplemented and pollen-limited conditions were created in colonies every spring, and brood rearing and honey yields were subsequently monitored throughout the summer. In all 3 yr, colonies that were supplemented with pollen or a pollen substitute in the spring started rearing brood earlier than colonies in other treatment groups and produced the most workers by late April or early May. In 2002, these initial differences were reflected by a two-fold increase in annual honey yields by September for colonies that were pollen-supplemented during the spring compared with pollen-limited colonies. In 2003 and 2004, differences between treatment groups in the cumulative number of workers produced by colonies disappeared by midsummer, and all colonies had similar annual honey yields (exception: in one year, productivity was low for colonies supplemented with pollen before wintering). Discrepancies between years coincided with differences in spring weather conditions. Colonies supplemented with pollen or a substitute during the spring performed similarly in all respects. These results indicate that an investment in supplementing the pollen diet of colonies would be returned for situations in which large spring populations are important, but long-term improvement in honey yields may only result when spring foraging is severely reduced by inclement weather. Beekeepers should weigh this information against the nutritional deficiencies that are frequently generated in colonies by the stresses of commercial management.     

Environmental influences on flight activity of USDA-ARS Russian and Italian stocks of honey bees (Hymenoptera: Apidae) during almond pollination

Differences in flight activity and in the percentages of pollen foragers between commercially managed honey bees, Apis mellifera L. (Hymenoptera: Apidae), of two stocks (USDA-ARS Russian, n = 41 colonies; and Italian, n = 43 colonies) were evaluated in an almond, Prunus dulcis (Miller) D. A. Webb, orchard in Kern Co., CA, during February and March 2002. Flight activity was measured by taking 1-min counts of bees exiting colonies on each of 9 d. Flight activity was best predicted with a model containing the effects of colony size (populations of adult bees and sealed brood), temperature, time of day, the interaction of adult bee population with temperature, and the interaction of adult bee population with time of day. Flight increased linearly with adult bee and brood population, had a quadratic relationship with temperature (increasing, but less so at higher temperatures), and had a quadratic relationship with time of day (decreasing, but less so at later times). Larger colonies had more response to changing temperatures and less response to different times of day than small colonies. Bee type had no direct influence on flight activity at any given colony size, temperature, or time of observation or when evaluated using a reduced data set retaining 34 Italian colonies and 32 Russian colonies whose mean sizes were equal. Overall, however, Russian colonies were less populous by about one-fourth and so fielded on average 71% of the foragers that Italian colonies did. Pollen collection was measured by capturing returning foragers on 4 d. The percentages of foragers with pollen were not different for the bee types.     

The influence of social structure on brood survival and development in a socially polymorphic ant: insights from a cross‐fostering experiment

Animal societies vary in the number of breeders per group, which affects many socially and ecologically relevant traits. In several social insect species, including our study species Formica selysi, the presence of either one or multiple reproducing females per colony is generally associated with differences in a suite of traits such as the body size of individuals. However, the proximate mechanisms and ontogenetic processes generating such differences between social structures are poorly known. Here, we cross‐fostered eggs originating from single‐queen (= monogynous) or multiple‐queen (= polygynous) colonies into experimental groups of workers from each social structure to investigate whether differences in offspring survival, development time and body size are shaped by the genotype and/or prefoster maternal effects present in the eggs, or by the social origin of the rearing workers. Eggs produced by polygynous queens were more likely to survive to adulthood than eggs from monogynous queens, regardless of the social origin of the rearing workers. However, brood from monogynous queens grew faster than brood from polygynous queens. The social origin of the rearing workers influenced the probability of brood survival, with workers from monogynous colonies rearing more brood to adulthood than workers from polygynous colonies. The social origin of eggs or rearing workers had no significant effect on the head size of the resulting workers in our standardized laboratory conditions. Overall, the social backgrounds of the parents and of the rearing workers appear to shape distinct survival and developmental traits of ant brood.     

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.     

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).     

不同人工代花粉对蜂群群势和生产性能的影响

早春将群势、蜂王年龄和质量基本一致的25群意大利蜜蜂Apis melliferaL.,随机分为5个处理,分别饲喂纯花粉和4种不同的人工代用花粉,观测它们对蜂群群势和生产性能的影响。结果表明,含有添加剂的各代用花粉组与纯花粉组的蜂群的蜂子数量差异均不显著,但均显著高于纯豆粕组(P<0.01);各组产蜜量随着纯花粉所占比例的减少而呈现递减的趋势,但饲喂含有60%花粉+40%豆粕+添加剂1组与纯花粉组没有明显的差异;含有添加剂的各代用花粉组的王台接受率均显著高于纯花粉组(P<0.01);饲喂含有添加剂的各代用花粉组与饲喂纯花粉组相比,王浆产量及蜂王浆中的10-HDA、水分、蛋白质的含量及酸度均没有差异。