Chapter 13 Opener: Weaver ants form superbly cooperative societies This chapter is about the evolution of social behavior. C:\Figures\Chapter13\high-res\Alcock8e-ChOpener-13.jpg

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13.1 The energy budget of “helpers” in Neolamprologus pulcher (Part 2) This species of fish is a type of cichlid (not the variety discussed earlier, but very similar), and will display a wide range of biparental care behaviors. C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-01-2.jpg In the above scenario, we are seeing an even more complex grouping whereby some non-reproductive fish become “helpers” at the nest site.

13.1 The energy budget of “helpers” in Neolamprologus pulcher (Part 1) The role of the helper is primarily to perform “submissive” behaviors around the nesting site. C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-01-1.jpg

13.2 Effect of removal of the top-ranked subordinate helper in a cooperatively breeding group In this, another cichlid species, the change in patterns seen in helper fish where the head subordinate (HS) is removed is shown. The HS removal changes dramatically the boundaries traveled by the other five “helper” fish. C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-02-0.jpg

13.3 Reproductive interference in a social animal The acorn woodpecker is a bird species that lives in a communal nesting site. Here we have REPRODUCTIVE INTERFERENCE being shown where the bird removes a nestmate’s egg. C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-03-0.jpg

13.5 Effect of parasites on cliff swallow nestlings C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-05-0.jpg From a nest infested with swallow bugs… note small size. From a nest treated with insecticide.

13.6 Social living with defensive benefits? Schooling of fish (these are striped catfish) can reduce predation pressures and increase survivability of individuals. C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-06-0.jpg

13.8 The different categories of helping behavior Fitness consequences of different helping behaviors. C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-08-0.jpg

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13.10 Cooperative courtship of the long-tailed manakin C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-10-0.jpg Alpha male mates with all females. Beta male mates with none. Other subordninates mate with none.

13.11 Cooperation with an eventual payoff C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-11-0.jpg

13.12 A meerkat sentinel on the alert for approaching predators Reciprocity hypothesis – the idea that there is reciprocal altruism in which a helpful action is repaid at a later date by the recipient of the assistance. C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-12-0.jpg

13.13 Reciprocity in a social primate Grooming is one example of a relatively low cost form of reciprocity. C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-13-0.jpg

13.14 Experimental demonstration of reciprocity in cotton-top tamarins (Part 1) Tamarin helper (right) can drag food to its partner. Or the Tamarin helper may be non-responsive. C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-14-1.jpg

13.15 The prisoner’s dilemma Aka: Game Theory C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-15-0.jpg

Direct fitness – fitness gained through reproduction Indirect fitness – fitness gained by helping “kin” Inclusive fitness – the combination of direct and indirect fittness Hamilton’s rule – a rare altruistic allele can become more common in a population only if the indirect fitness gained by the altruist is greater than the direct fitness it loses as a result of the self-sacrificing actions.

13.18 Altruism and relatedness in pied kingfishers Primary helpers are related to the breeders whereas the secondary helpers are not. C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-18-0.jpg

13.23 Haplodiploid sex determination in Hymenoptera (Part 2) C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-23-2.jpg

13.23 Haplodiploid sex determination in Hymenoptera (Part 3) C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-23-3.jpg

13.24 Conflict within ant colonies over reproduction The center ant is covered with the sting scent of the queen to identify her as becoming reproductive. The three worker ants hold her in position to prevent reproduction. C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-24-0.jpg A single ant iimpobolizes another ant whose ovaries have begun to mature.

13.27 Eusocial insects have sterile castes C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-27-0.jpg Many social insects like honeybees and ants have a sterile caste.

13.28 Suicidal sacrifice by a worker bee C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-28-0.jpg

13.30 Haplodiploidy and the evolution of eusociality in the Hymenoptera (Part 1) C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-30-1.jpg

13.30 Haplodiploidy and the evolution of eusociality in the Hymenoptera (Part 2) C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-30-2.jpg

Note the larger forelimbs on the sterile worker. 13.31 A sterile thrips soldier (right) next to a reproductive foundress female (left) Note the larger forelimbs on the sterile worker. C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-31-0.jpg

13.34 A mammal with an effectively sterile caste C:\Figures\Chapter13\high-res\Alcock8e-Fig-13-34-0.jpg