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and Atta colombica in Corcovado, Costa Rica; see also MEA Whitehouse, K Jaffe 1996, Ant wars: Combat strategies, territoriality and nest defence in the leaf-cutting ant Atta laevigata, Anim. Behav. 51: 1207–1217. Such confrontations may go largely unseen in the canopy. But leaf cutting is likely to reduce the productivity of the trees on which aggressive canopy ants of other species depend, and it is known that some of these dominant ants fight with leafcutters; see JK Wetterer 1994, Attack by Paraponera clavata prevents herbivory by the leaf-cutting ant, Atta cephalotes, Biotropica 26: 462–465; and n. 1.

79. A queen can rear workers if her garden fails, though it’s unknown whether her workers can forage for a replacement fungus; see H Fernández-Marín, WT Wcislo 2005, Production of minima workers by gynes of Atta colombica that lack a fungal pellet, J. Kansas Entomol. Soc. 78: 290–292.

80. MB Dijkstra, DR Nash, JJ Boomsma 2005, Self-restraint and sterility in workers of Acromyrmex and Atta leafcutter ants, Insectes Soc. 52: 67–76.

81. AS Yang 2007, Thinking outside the embryo: The superorganism as a model for evodevo studies, Biol. Theory 2: 398–408.

82. EO Wilson 1983, Caste and division of labor in leaf-cutter ants, IV: Colony ontogeny of A. cephalotes, Behav. Ecol. Sociobiol. 14: 55–60.

83. JK Wetterer 1994, Ontogenetic changes in forager polymorphism and foraging ecology in the leaf-cutting ant Atta cephalotes, Oecologia 98: 235–238.

84. A Powell, S Shennan, MG Thomas 2009, Late Pleistocene demography and the appearance of modern human behavior, Science 324: 1298–1301; Elman R. Service, Origins of the State and Civilization (New York: W. W. Norton, 1975).

85. Overlap in generations, the reproductive division of labor, and cooperative care of the young are considered the essential attributes of the most complex, or eusocial, insect societies, which include all ant species except possibly a few that lack distinct queens.

15. The Origins of Agriculture

1. I recommend the review by TR Schultz, UG Mueller, CR Currie, SA Rehner, Reciprocal illumination: A comparison of agriculture in humans and in fungus-growing ants, in Insect-Fungal Associations: Ecology and Evolution, ed. FE Vega, M Blackwell (Oxford: Oxford University Press), pp. 149–190. Versions of the slovenly ant hypothesis have involved fungus growth on animal matter, vegetable matter, fecal matter, or plant roots; see UG Mueller, TR Schultz, CR Currie, RMM Adams, D Malloch 2001, The origin of the attine ant-fungus mutualism, Q. Rev. Biol. 76: 169–197.

2. Stephen B. Brush and Monica L. Smith, personal communications; Stephen B. Brush, Farmers’ Bounty: Locating Crop Diversity in the Contemporary World (New Haven: Yale Univ. Press, 2004); Stephen Budiansky, The Covenant of the Wild: Why Animals Chose Domestication (New Haven: Yale University Press, 1999); and David Rindos, The Origins of Agriculture (New York: Academic Press, 1984).

3. One group of ants feeds the pellets to its larvae, such that the workers “combine the contents of the dust-bin and garbage-can and serve up the mixture as appropriate food for their young—a truly remarkable example of food-conservation”; see WM Wheeler, IW Bailey 1920, The feeding habits of pseudomyrmine and other ants, Trans Am. Philos. Soc. 22: 235–279.

4. With their lumpy bodies, Proatta also look like Atta leafcutters, which is how they got their name; see MW Moffett 1986, Behavior of Malayan group-predatory ant Proatta butteli: An Old-World relative of attine ants, Insectes Soc. 33: 444–457. The closest living relative of fungus-growing ants has untidy habits similar to Proatta; see C Rabeling, M Verhaagh, UG Mueller 2006, Behavioral ecology and natural history of Blepharidatta brasiliensis, Insectes Soc. 53: 300–306; and JLM Diniz, CRF Brandão, CI Yamamoto 1998, Biology of Blepharidatta ants, the sister group of the Attini: A possible origin of fungus-ant symbiosis, Naturwissenschaften 85: 270–274.

5. RP Coppinger, CK Smith 1983, The domestication of evolution, Environ. Conserv. 10: 283–292.

6. Feeding on fungi is