orker honey bees are marvelous somewhat of a chronological order, as long task – after all, they cannot feed larvae with- creatures. These females could as one remembers (1) workers may skip out being able to produce brood food. Given Whave been queens, had their older tasks, (2) workers perform many tasks at that worker honey bees perform the vast ma- sisters simply fed them more food while they each age, and (3) workers spend more time jority of essential tasks in the colony, their were developing as larvae. The amount and resting and patrolling the nest than they do activity ensures that the colony has all of its quality of diet they were fed while young anything else. needs met. The local environment around were the only things separating them from Emphasizing point 3 above, I find it quite the colony also likely plays a role in worker royalty. If they had been a queen, they could interesting that workers bees are even called task allocation, given that colony require- have laid eggs, maintained the colony’s ho- worker bees at all. They spend the vast ments vary based on a variety of environ- meostasis, and lived up to a few years. Yet, majority of their lives resting in the nest. mental conditions and that workers respond they were born workers, destined to work Resting, of course, grants a worker needed to colony needs by doing jobs that must be their lives away in just a few short weeks. respite. After all, who does not benefit from done to ensure colony homeostasis. Many worker bee equals in other social a little R&R? Things are not always as they I am going to spend the rest of the article insect colonies, such as ants or termites, are seem, though, as bees are not always resting introducing you to the tasks performed by born into a caste system, with a designated when they appear to be. Workers may remain worker honey bees. I will do my best to task that they perform their entire lives. Not immobile while secreting wax, producing present the tasks to you in the order that they so for worker honey bees. Worker honey brood food, etc. Thus, the period of immo- typically are performed, though admittedly bees subscribe to temporal polyethism, or bility may simply be times of preparation I found this really hard to do given the mas- “time related” (temporal) “many” (poly) to perform a given task. Worker bees also sive variation in the timing and duration of “behaviors” (ethism). This simply means spend quite a lot of time patrolling the nest. tasks that workers perform. Keep in mind, that worker honey bees perform a number The performance of this behavior seems to bee progression through these tasks varies; of tasks over time and the performance of be a worker’s attempt to find out what needs only very loose conclusions can be drawn these tasks is age dependent. Beekeepers to be done in a colony, with the idea that they about the order and performance of worker called this phenomenon age-related division are looking for something to do. tasks. Much of what I share will be based on of labor. The end result is simple: worker Though there is considerable overlap in the information provided by Winston (1980) honey bees are not born into a task in which the tasks that workers perform, there is an and Seeley (1982). The books they wrote are they remain their entire lives. Instead, they overall pattern to where and when they per- masterpieces of bee biology and ecology. progress through a somewhat predictable form the tasks. Young worker bees engage They are worth being in every beekeeper’s series of tasks as they march toward the day in nest-based tasks while older worker bees library. I provide full citations in the refer- that they can work no longer. perform tasks near the colony entrance or ence section of this manuscript in the event Interestingly enough, not all workers outside of the colony. Generally speaking, that you wish to explore these books fur- perform all tasks. Some workers may skip the tasks follow this natural progression: ther. The order that I discuss the tasks are certain tasks altogether as they mature. Fur- (1) cell cleaning and capping, (2) brood and the approximate order in which they are pre- thermore, the task system is not rigid. Old queen tending, (3) comb building, cleaning formed, as noted by Winston (1980). I list bees can perform young bee tasks and vice and food handling, and (4) outside tasks. the common age ranges and the mean age of versa. This can be exacerbated further given Worker progression through their tasks workers performing each task. the fact that a worker bee usually performs probably is regulated physiologically, hor- multiple tasks at any one age. Regardless, monally and by colony need. For example, AGE-RELATED TASKS enough of a pattern exists in worker task per- certain glands in workers must develop be- Cell preparation (about 2-16 days of formance that we can discuss these tasks in fore the workers are able to perform a given age, mean age about 8 days old) – This October 2015 1077 ABJ_October_2015.indb 1077 8/31/2015 9:09:44 AM is the first task that workers perform when cell inspections lead a worker to feed the de- wait close to the nest entrance, ready for they emerge from their cells as adult bees. veloping larvae. A cell may be visited many nectar-laden foragers to return. The nectar This task can be split into two subtasks: thousands (5000+) of times while the larvae receivers are quick to unload the incoming polishing cells and cleaning cells, with the grows, with less than half of those visits foragers so that the foragers can return to the latter happening later in the bee’s life. For (1,000 – 2,000) resulting in larval feeding field. A single returning forager may regur- the former, the workers begin cleaning a cell events. Interestingly, workers bees may be gitate her crop contents to 2-3 receiver bees. from the back of the cell, working their way able to determine the age of the developing The receiver bees, in turn, transport the nec- up the walls of the cell and to the entrance larvae because they are able to adjust the pro- tar into the nest as they search for a cell in of the cell (Figure 1). They will remove the portion of glandular secretions in the diet in which they can deposit their nectar load. remains of cocoons and excreta left by pre- response to the age of the immature bee (dif- Before they deposit the load, workers will vious cell occupants. They will cover any ferent age larvae eat slightly different mix- spit out the nectar and suck it back into their remaining adulteration in the cell with a thin tures of brood food). A single nurse bee rears mouths over and over. This behavior helps layer of wax. the equivalent of 2-3 larvae during its life. start the process of nectar dehydration and conversion into honey. After blowing nectar bubbles, the worker bees deposit the nectar into available cells (Figure 5). Figure 1 – Two worker bees cleaning cells shortly after their emergence as Figure 3 – A worker bee feeding a adults. Photograph by Mike Bentley. larva. Photograph by Mike Bentley. Capping brood (about 3-10 days of Tending the queen (about 3-14 days Figure 5 – A worker depositing nec- age, mean age about 8 days old) – Young of age, mean age about 11 days old) – tar into a cell. Photograph by Mike worker bees are able to secrete wax in small Worker bees tend the queen about the same Bentley. amounts. This allows them to cap the cells time period that they perform nurse bee of larvae that are ready to begin transform- tasks. Workers tending the queen lick her, Cleaning debris (11-15 days of age, ing into pupae (Figure 2). They also may examine her with their antennae and fore- mean age about 14 days old) – Workers seal cells of ripe honey while at this age. A legs, groom her and feed her with glandular performing this task are the waste manage- cell can take 20 minutes to 6 hours to cap. secretions and honey (Figure 4). The queen ment force of the hive. They will remove may be fed by workers every 20-30 minutes debris from the nest. This debris includes for about 2-3 minutes per feeding. Worker moldy pollen, old cell cappings, wax debris, bees performing this task often form a circle dead brood or adult bees, etc. The removal around the queen, with the circling bees of dead or diseased brood is called hygienic being referred to as the queen’s attendants behavior and is a special type of cleaning or retinue. The licking behavior is quite behavior that helps the colony battle pests important. Licking the queen is how the and diseases. With this behavior, the work- workers acquire and distribute the queen’s ers can detect dead or diseased brood in pheromones around the nest. This spreading cells, even capped cells, uncap the cells, and of her pheromones maintains nest homeo- remove the dead or sick bees from the nest. stasis. It lets all workers know she is present. This behavior is the subject of bee selection programs aimed at reducing disease and pest pressures in the nest. Handling pollen (about 10-19 days of age, mean age about 15 days old) – Unlike nectar foragers, pollen foragers deposit their pollen loads directly into cells (Figure 6).