
392 17. Ant Algorithms
specialization, in such a way that there is a fitness gain as a consequence of such
specialization [774].
Division of labor has been observed in a number of eusocial insects [774, 850], including
ants [328, 348, 558, 693, 882], wasps [298, 624], and the honey bee [393, 625, 733]. Here
eusociality refers to the presence of [214]:
• cooperation in caring for the young,
• reproductive division of labor, with more or less sterile individuals working on
behalf of those individuals involved with reproduction, and
• overlap of generations.
Within these insect colonies, a number of tasks are done, including reproduction,
caring for the young, foraging, cemetery organization, waste disposal, and defense.
Task allocation and coordination occurs mostly without any central control, especially
for large colonies. Instead, individuals respond to simple local cues, for example
the pattern of interactions with other individuals [328], or chemical signals. Also
very interesting, is that task allocation is dynamic, based on external and internal
factors. Even though certain groups of individuals may specialize in certain tasks,
task switching occurs when environmental conditions demand such switches. This
section provides an overview of the division of labor in insect colonies. Section 17.3
gives a summary of different types of division of labor, and gives an overview of a
few mechanisms that facilitate task allocation and coordination. A simple model of
division of labor based on response thresholds is discussed in Section 17.3.2. This
model is generalized in Section 17.3.3 to dynamic task allocation and specialization.
17.3.1 Division of Labor in Insect Colonies
Social insects are characterized by one fundamental type of division of labor i.e. re-
productive division of labor [216, 882]. Reproduction is carried out by a very small
fraction of individuals, usually by one queen. In addition to this primary division of
labor into reproductive and worker castes, a further division of labor exists among the
workers. Worker division of labor can occur in the following forms [77]:
• Temporal polyethism, also referred to as the age subcaste [77, 216, 393, 625].
With temporal polyethism, individuals of the same age tend to perform the
same tasks, and form an age caste. Evidence suggests that social interaction is a
mechanism by which workers assess their relative ages [625]. As an example, in
honey bee colonies, younger bees tend to the hive while older bees forage [393].
Temporal polyethism is more prominent in honey bees and eusocial wasps [625].
• Worker polymorphism, where workers have different morphologies [77, 558,
882]. Workers of the same morphological structure belong to the same morpho-
logical caste, and tend to perform the same tasks. For example, minor ants care
for the brood, while major ants forage [77]. As another example, the termite
species Termes natalensis has soldier ants that differ significantly in morpho-
logical structure from the worker termites [558]. Morphological castes are more
prominent in ant and termite species [625].