6 The Life-cycle The true Bumblebees are social insects, in the sense that they show a division of labour between the reproductive males and females, and a caste of 'worker' bees. The latter are (usually) infertile, and their lives are devoted to foraging for food and tending the larvae produced by the single female ('queen') which establishes the nest. Unlike the honey bees, Bumblebee nests last no longer than one season. Typically, a fertilised female emerges from hibernation in early spring, and commences feeding on nectar and pollen from available flowers (Salix spp., Lamium album, and several early-flowering garden plants are very important for these early-flying females). Then begins a more-or-less prolonged search for suitable nesting sites (which vary considerably from one species to another). Death-rates among these over-wintered females are very high indeed, and their dead bodies arc easy to find on the ground around food-sources, roadside verges and elsewhere. Once a nest is established, the female deposits a clump of pollen in it, and builds a wax cell into which she lays her first batch of eggs. The resulting larvae are nurtured exclusively by the female, who can be observed during the spring collecting nectar and pollen for this first brood. Soon this task is taken over by the newly emerged workers, and the female then remains in the nest, continuing to lay eggs. During the late spring and early summer the number of workers produced in each nest continues to grow until a remarkable transition takes place. Instead of workers, a brood of fertile males and females emerges from the nest. These mate, and the males continue to live 'solitary' existences outside the nest, foraging only for their own subsistence. Meanwhile, the fertilized females build up an internal store of nutrients to take them through their winter hibernation, and begin searching for suitable hibernation sites (usually burrowing a few centimeters into a north- facing bank). However, in favourable seasons the females of some species establish nests in mid-summer and complete a second full life- cycle in the same year. Actively foraging workers of some species can often be observed well into October. Whilst the above features of the life-cycle are common to all the true Bumblebees, there are some interesting variations in the details. In some species (Bombus terrestris, for example), stores of pollen are maintained in wax cells or empty pupal cocoons. The larvae of this group of bees spin separate compartments in the nest, and are fed individually by the queen or workers with a mixture of nectar and pollen from the pollen stores. This tends to produce workers more-or- less equal in size. Bumblebees of this group arc known as 'pollen storers'. Alternatively, in such species as Bombus hortorum and Bombus pascuorum, the larvae stay together in a common brood chamber. Wax 'pockets' are built in this, and foraging workers or queens deposit their loads of pollen into them. The larvae feed themselves from the resulting mass. These bees are termed 'pocket makers', and because of competition between larvae for access to food in the brood chamber, there are said to be bigger differences in the size of the resulting adult workers than with the pollen storcrs. More research is needed into the processes involved in the seasonal shift from production of workers to fertile males and females. It seems likely that the mechanisms vary considerably from species to species, but two main groups of species are distinguished. In so-called 'simple' species (e.g. Bombus pratorum) the queen appears not to influence the transition to production of new queens. Such factors as the ratio of workers to larvae, the overall density of bees in the nest and temperature arc thought to lead to some larvae feeding more and for longer than others, and these eventually emerge as fertile females. In the case of 'complex' species (e.g. Bombus terrestris) it seems that queens can regulate the timing of the production of new queens by emitting pheromoncs which influence worker- activity. In these species there is a more clear distinction in size between workers and queens. Essex Naturalist (New Series) 16 (1999)