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Adventures Among Ants - Mark W. Moffett [179]

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with small colonies) survive the queen’s death by having the workers take over reproduction; see the conclusion to this book and Bert Hölldobler and Edward O. Wilson, The Superorganism (New York: W.W. Norton, 2008), pp. 356–426.

42. A Buschinger, U Maschwitz, Defensive behavior and defensive mechanisms in ants, in Defensive Mechanisms in Social Insects, ed. Henry R. Hermann (New York: Praeger Scientific, 1984), pp. 95–150; the quote appears on p. 124.

12. Slaves of Sagehen Creek

1. I focus here on breviceps, but there is excellent work on the other Polyergus species.

2. Donato Grasso, personal communication. Another fascinating possibility is that the Formica bring up some brood to make them accessible to raiding parties in a tactic that diverts the Polyergus from more critical parts of the nest, in a kind of “lizard loses its tail” sacrifice.

3. Apparently colony identity odors can be detected not just at contact but also over distances of a few millimeters; see AS Brandstaetter, A Endler, CJ Kleineidam 2008, Nestmate recognition in ants is possible without tactile interaction, Naturwissenschaften 95: 601–608.

4. Propaganda substances have so far been studied only in the European Amazon ant; see R. Visicchio, A Mori, DA Grasso, C Castracani, F Le Moli 2001. Glandular sources of recruitment, trail, and propaganda semiochemicals in the slave-making ant Polyergus rufescens, Ethol. Ecol. Evol. 13: 361–372.

5. Those Formica species that have a shorter history of conflict with the Amazons tend to fight back more, and as a consequence they suffer higher mortality from raids. The species raided can vary from place to place, even from colony to colony; see, e.g., JM Bono, R Blatrix, MF Antolin, JM Herbers 2007, Pirate ants (Polyergus breviceps) and sympatric hosts (Formica occulta and Formica sp. cf. argentea): Host specificity and coevolutionary dynamics, Biol. J. Linn. Soc. 91: 565–572.

6. JM Herbers, S Foitzik 2002, The ecology of slavemaking ants and their hosts in north temperate forests, Ecology 83: 148–163; and RJ Stuart 1988, Collective cues as a basis for nestmate recognition in polygynous leptothoracine ants, Proc. Natl. Acad. Sci. 85: 4572–4575.

7. TO Richardson, PA Sleeman, JM McNamara, AL Houston, NR Franks 2007, Teaching with evaluation in ants, Curr. Biol. 17: 1520–1526; and E. Leadbeater, N Raine, L Chittka 2006, Social learning: Ants and the meaning of teaching, Curr. Biol. 16: R323–R325.

8. RJ Stuart 1986, An early record of tandem running in leptothoracine ants: Gottfrid Adlerz 1896, Psyche 93: 103–106; and TM Alloway 1979, Raiding behaviour of two species of slave-making ants, Harpagoxenus americanus and Leptothorax duloticus, Anim. Behav. 27: 202–210.

9. J Beibl, RJ Stuart, J Heinze, S Foitzik 2005, Six origins of slavery in formicoxenine ants, Insectes Soc. 52: 291–297.

10. MW Moffett 1989, Life in a nutshell, National Geographic 175: 783–796.

11. A Lenoir, P D’Ettorre, C Errard 2001, Chemical ecology and social parasitism in ants, Annu. Rev. Entomol. 46: 573–599. Slavemaker colonies act more like predators than parasites, which feed on a single host; see M Brandt, S Foitzik, B Fischer-Blass, J Heinze 2005, The coevolutionary dynamics of obligate ant social parasite systems: Between prudence and antagonism, Biol. Rev. Camb. Philos. Soc. 80: 251–267. We’ll see in the next chapter that the Amazons collect some of the colony’s food, in that the slaves eat some of the raided brood.

12. John Lubbock, Ants, Bees, and Wasps (New York: Appleton, 1883). Thief ants belong to Solenopsis subgenus Diplorhoptrum.

13. Pierre Huber, Recherches sur les moeurs des fourmis indigènes (Geneva: J. J. Paschoud, 1810), pp. 219–224.

14. Charles Darwin, Origin of Species (London: John Murray, 1859), pp. 219–224.

15. Milton Meltzer, Slavery: A World History (Chicago: Da Capo Press, 1993); and Orlando Patterson, Slavery and Social Death: A Comparative Study (Cambridge, MA: Harvard University Press, 1982).

16. A Achenbach, S Foitzik 2009, First evidence for slave rebellion: Enslaved ant workers systematically kill the

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