Adventures Among Ants - Mark W. Moffett [164]
9. CJ Lumsden, B Hölldobler 1983, Ritualized combat and intercolony communication in ants, J. Theor. Biol. 100: 81–98.
10. NR Franks, LW Partridge 1993, Lanchester battles and the evolution of combat in ants, Anim. Behav. 45: 197–199; and SD Porter, CD Jorgensen 1981, Foragers of the harvester ant, Pogonomyrmex owyheei: A disposable caste? Behav. Ecol. Sociobiol. 9: 247–256. A sad fact of human military history is that putting the most expendable soldiers in the front lines often pays off. Although the modern military does what it can to protect its soldiers (conducting saturation bombing before sending them in, for example), it is no accident that at the time of this writing, the last war in which a U.S. general died in combat was the Vietnam War—and it was caused by a helicopter crash in dense fog, not enemy fire.
11. Similar shifts back and forth between nests also occur in army ants; see WH Gotwald Jr. 1978, Emigration behavior of the East African driver ant Dorylus (Anomma) molestus, J.N.Y. Entomol. Soc. 86: 290.
12. RS Savage 1847, On the habits of the “drivers” or visiting ants of West Africa, Trans. R. Entomol. Soc. Lond. 5: 1–15; the quotation appears on p. 4.
13. See, e.g., C. Schöning, WM Njagi, NR Franks 2005, Temporal and spatial patterns in the emigrations of the army ant Dorylus (Anomma) molestus in the montane forest of Mt. Kenya, Ecol. Entomol. 30: 532–540; and EO Wilson 1958, The beginnings of nomadic and group-predatory behavior in the ponerine ants, Evolution 12: 24–31.
14. H Topoff, J Mirenda 1980, Army ants on the move: Relation between food supply and emigration frequency, Science 207: 1099–1100. Certain Leptogenys ants that, like diversus, are now known to raid like army ants are thought to have become nomadic prior to evolving into mass foragers, but apparently for a different reason: species of this genus move due to the frequent disturbances they experience from nesting in the leaf litter; see V Witte, U Maschwitz 2002, Coordination of raiding and emigration in the ponerine army ant Leptogenys distinguenda: A signal analysis, J. Insect Behav. 15: 195–217.
15. See, e.g., JM Leroux 1982, Ecologie des populations de dorylines Anomma nigricans dans la région de Lamto (Côte d’Ivoire), Publications du Laboratoire de Zoologie, no. 22, Ecole Normale Supérieure, Paris. One New World army ant has even been recorded encamped at one site for at least eight months; see HG Fowler 1979, Notes on Labidus praedator in Paraguay, J. Nat. Hist. 13: 3–10.
16. J Smallwood 1982, Nest relocations in ants, Insectes Soc. 29: 138–147. Even some large, entrenched colonies can migrate often; see, e.g., HG Fowler 1981, On the emigration of leaf-cutting ant colonies, Biotropica 13: 316.
17. V Witte and U Maschwitz 2008, Mushroom harvesting ants in the tropical rain forest, Naturwissenschaften 95: 1049–1054.
18. In a variant of fission called budding, far less than half the colony departs with a newly crowned queen to start their own colony elsewhere, leaving the original queen with her nest. Terry McGlynn thinks that budding and fission may be widespread among ants with small nests in rainforest leaf litter, where competition can be too intense for queens to start colonies alone. See TP McGlynn 2006, Ants on the move: Resource limitation of a litter-nesting ant community in Costa Rica, Biotropica 38: 419–427. Over much of pre-agricultural human history, hunter-gatherer groups arose in much the same way. Bands split as they grew beyond a few dozen individuals and either began experiencing discord or had trouble searching widely for meals as a foraging group; without infrastructure and stockpiles, such divisions were easy. These groups also fused more readily than do ant colonies. See FW Marlowe 2005, Hunter-gatherers and human evolution, Evol. Anthropol. 14: 54–67.
19. Though specialized egg-carrying by young queens suggests that they are able to start colonies