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Eusociality. Conflicts over reproduction Definition and occurrence Explanations for worker sterility Routes to sociality. Definition. Overlapping generations Reproductive division of labor including nonreproductive (or sterile) workers Caste polymorphism (in some cases)
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Eusociality • Conflicts over reproduction • Definition and occurrence • Explanations for worker sterility • Routes to sociality
Definition • Overlapping generations • Reproductive division of labor including nonreproductive (or sterile) workers • Caste polymorphism (in some cases) • Parental care in a permanent nest
Social spiders Communal, but no sterile caste
Why worker sterility? • Kin selection • Parental manipulation
Why worker sterility? Kin selection • Because of haplodiploidy, workers are more closely related to sisters(r = 3/4) than to offspring (r = 1/2), assuming females mate once • But, they are more closely related to male offspring (r = 1/2) and nephews (r = 3/8) than to brothers (r = 1/4). Therefore, workers should lay unfertilized eggs if mothers mate singly (bumblebees, stingless bees) • If mothers are polyandrous, then female workers may be more closely related to brothers than half-nephews (r = 1/8). Expect workers to kill unfertilized eggs laid by other workers in polyandrous species, e.g. honeybees.
Worker policing in honeybees “Queenright” “Queenless”
Why worker sterility? Parental manipulation • If parents can prevent their young from reproducing, then it may be better to help • A mother that traded eggs with her daughters would trade grandkids (r = 1/4) for offspring (r = 1/2) and gain a 2-fold advantage. Daughter trades offspring for siblings and loses nothing (r =1/2 for both).
Sex ratio investment • Offspring control: expect 3:1 investment sex ratios because females are related to sisters by 3/4 and to brothers by 1/4 • Parental control: expect 1:1 sex ratios • Data fit 3:1, rejects parental manipulation • Only slave-makers are 1:1
Factors promoting eusociality • Genetics • High relatedness of workers to reproductives • Ecology • High cost of dispersal • Need for assistance in nest construction • Reproductive success is proportionately higher for family than individual • Need to defend a multi-generation nest
Routes to sociality Predict r < 0.5 Predict r > 0.5
Inbreeding-outbreeding cycles in termites Termites are diploid. Both sexes help. In some species, multiple reproductives occur. If these are produced by inbreeding over multiple generations, then termites within a colony could be very closely related