Abstract
The best known of the conflicts occurring in
eusocial Hymenoptera is queen-worker conflict over sex ratio. So far,
sex ratio theory has mostly focused on optimal investment in the
production of male versus female sexuals, neglecting the investment in
workers. Increased investment in workers decreases immediate sexual
productivity but increases expected future colony productivity. Thus, an
important issue is to determine the queen's and workers' optimal
investment in each of the three castes (workers, female sexuals,
and male sexuals), taking into account a possible trade-off between
production of female sexuals and workers (both castes developing from
diploid female eggs). Here, we construct a simple and general kin
selection model that allows us to calculate the evolutionarily stable
investments in the three castes, while varying the identity of the party
controlling resource allocation (relative investment in workers,
female sexuals, and male sexuals). Our model shows that queens and
workers favor the investment in workers that maximizes lifetime colony
productivity of sexual males and females, whatever the colony kin
structure. However, worker production is predicted to be at this optimum
only if one of the two parties has complete control over resource
allocation, a situation that is evolutionarily unstable because it
strongly selects the other party to manipulate sex allocation in
its favor. Queens are selected to force workers to raise all the
males by limiting the number of eggs they lay, whereas workers should
respond to egg limitation by raising a greater proportion of the female
eggs into sexual females rather than workers as a means to attain a
more female-biased sex allocation. This tug-of-war between queens
and workers leads to a stable equilibrium where sex allocation is
between the queen and worker optima and the investment in workers is
below both parties' optimum. Our model further shows that, under
most conditions, female larvae are in strong conflict with queens
and workers over their developmental fate because they value their
own reproduction more than that of siblings. With the help of our
model, we also investigate how variation in queen number and number
of matings per queen affect the level of conflict between queens,
workers, and larvae and ultimately the allocation of resource in the
three castes. Finally, we make predictions that allow us to test
which party is in control of sex allocation and caste determination.
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