TableĀ 1.

Challenges in applying inclusive fitness theory to the alternative individual strategies of reproduction versus altruism in insect societies.

challenge in applying inclusive fitness theoryempirical example in relation to reproduction versus altruismselected references
must consider both direct and indirect effects of kinshipin queenless colonies, only the direct effect applies in relation to worker egg laying (production of males). In queenright colonies, indirect effects, via coercion (e.g. killing of worker-laid eggs, aggression of egg laying workers), also occur but vary in importance among speciesWenseleers & Ratnieks (2006b)
must consider both reproducing and coercing individualsworker policing (coercion) is more strongly selected for when relatedness among female offspring is low due to multiple mating by the queen. Self-restraint/acquiescence is more strongly selected when relatedness is high or when coercion is effectiveRatnieks (1988), Wenseleers et al. (2004b)
must take into account all relevant partiesegg policing, for example, can be carried out by the queen or by workers. Worker policing of eggs may be carried out either by egg-laying workers (selfish policing) or by non-reproducing workersWenseleers & Ratnieks (2006a)
must consider idiosyncracies that affect power relations among colony membersbrood-rearing method of stingless bees versus honeybees. When each larva is reared in a sealed cell (mass provisioning) individual larvae have more power over their own caste fate than in the honeybee, in which each larva is reared progressively in an open cellBourke & Ratnieks (1999), Wenseleers & Ratnieks (2004)
must consider informationegg policing requires policing individuals to be able to discriminate between queen-laid and worker-laid eggsBeekman & Ratnieks (2003)
evasionindividuals have an incentive to evade social coercion. Dwarf queens in trigonine stingless bees develop in a worker cell. Honeybee workers may lay eggs that evade worker policing, or enter a nearby queenless colony in which worker policing has been switched offBeekman & Oldroyd (2008), Ribeiro et al. (2006)
must consider interactions with other conflictsworker policing of worker-laid eggs can be selected for on sex allocation grounds. This may account for the occurrence of worker policing in species with queens mated to a single maleFoster & Ratnieks (2001b)
must consider also benefits and costspolicing on colony efficiency grounds. Insurance and head start benefits in the origin of eusociality. Ecological factors that affect the ease or difficulty of founding a nest independentlyRatnieks (1988), Queller (1989)