21. A. J. Moore and P. Wilson, “The Evolution of Sexually Dimorphic Earwig Forceps: Social Interactions Among Adults of the Toothed Earwig, Vostox apicedentatus,” Behavioral Ecology 4 (1993): 40–48; J. L. Tomkins and L. W. Simmons, “Female Choice and Manipulations of Forceps Size and Symmetry in the Earwig Forficula auricularia L.,” Animal Behaviour 56 (1998): 347–56.
22. A. Malo, E. R. S. Roldan, J. Garde, A. J. Soler, and M. Gomendio, “Antlers Honestly Advertise Sperm Production and Quality,” Proceedings of the Royal Society of London, Series B 272 (2005): 149–57.
23. A. Balmford, A. M. Rosser, and S. D. Albon, “Correlates of Female Choice in Resource-Defending Antelope,” Behavioral Ecology and Sociobiology 31 (1992): 107–14.
24. N. A. M. Rodger, The Command of the Ocean—A Naval History of Britain 1649–1815 (W. W. Norton, 2005).
25. R. L. O’Connell, Of Arms and Men: A History of War, Weapons, and Aggression (Oxford: Oxford University Press, 1989); O’Connell, Soul of the Sword: An Illustrated History of Weaponry and Warfare from Prehistory to the Present (New York: Free Press, 2002); R. Gardiner and B. Lavery, The Line of Battle: The Sailing Warship 1650–1840 (London: Conway Maritime Press, 2004).
26. Gardiner and Lavery, Line of Battle.
27. O’Connell, Of Arms and Men; O’Connell, Soul of the Sword.
28. Ibid.
29. Gardiner and Lavery, Line of Battle.
30. Ibid.
31. D. Miller and L. Peacock, Carriers: The Men and the Machines (New York: Salamander Press, 1991).
32. Wikipedia, s.v. “Boeing F/A-18E/F Super Hornet.”
33. Miller and Peacock, Carriers.
10. Sneaks and Cheats
1. D. J. Emlen, “Alternative Reproductive Tactics and Male-Dimorphism in the Horned Beetle Onthophagus acuminatus (Coleoptera: Scarabaeidae),” Behavioral Ecology and Sociobiology 41 (1997): 335–41.
2. Ibid.
3. The fact that small males “switch off” horn growth provided an unusual opportunity to study developmental mechanisms associated with horn development. Genetically similar males could be raised under conditions that either triggered or suppressed horn growth, and these animals could then be compared in terms of hormone levels, patterns of cell growth, and gene expression. Like a “toe in the door,” horn dimorphism afforded rare glimpses into details of insect development. For papers examining how developmental hormones appear to regulate horn growth, see D. J. Emlen and H. F. Nijhout, “Hormonal Control of Male Horn Length Dimorphism in the Dung Beetle Onthophagus taurus (Coleoptera: Scarabaeidae),” Journal of Insect Physiology 45 (1999): 45–53; D. J. Emlen and H. F. Nijhout, “Hormonal Control of Male Horn Length Dimorphism in Onthophagus taurus (Coleoptera: Scarabaeidae): A Second Critical Period of Sensitivity to Juvenile Hormone,” Journal of Insect Physiology 47 (2001): 1045–54; and A. P. Moczek and H. F. Nijhout, “Developmental Mechanisms of Threshold Evolution in a Polyphenic Beetle,” Evolution and Development 4 (2002): 252–64. For a comparative study examining the evolution of horn dimorphism in dung beetles, see D. J. Emlen, J. Hunt, and L. W. Simmons, “Evolution of Sexual Dimorphism and Male Dimorphism in the Expression of Beetle Horns: Phylogenetic Evidence for Modularity, Evolutionary Lability, and Constraint,” American Naturalist 166 (2005): S42–S68; and for more recent studies examining patterns of gene expression in developing horns, see A. P. Moczek and L. M. Nagy, “Diverse Developmental Mechanisms Contribute to Different Levels of Diversity in Horned Beetles,” Evolution and Development 7 (2005): 175–85; A. P. Moczek and D. J. Rose, “Differential Recruitment of Limb Patterning Genes During Development and Diversification of Beetle Horns,” Proceedings of the National Academy of Sciences 106 (2009): 8992–97; T. Kijimoto, J. Costello, Z. Tang, A. P. Moczek, and J. Andrews, “EST and Microarray Analysis of Horn Development in Onthophagus Beetles,” BMC Genomics 10 (2009): 504; E. C. Snell-Rood, A. Cash, M. V. Han, T. Kijimoto, J. Andrews, and A. P. Moczek, “Developmental Decoupling of Alternative Phenotypes: Insights From the Transcriptomes of Horn-Polyphenic Beetles,” Evolution 65 (2011): 231–45.
4. A. P. Moczek and D. J. Emlen, “Male Horn Dimorphism in the Scarab Beetle, Onthophagus taurus: Do Alternative Reproductive Tactics Favour Alternative Phenotypes?” Animal Behaviour 59 (2000): 459–66; R. Madewell and A. P. Moczek, “Horn Possession Reduces Maneuverability in the Horn-Polyphenic Beetle, Onthophagus nigriventris,” Journal of Insect Science 6 (2006): 21.
5. L. W. Simmons, J. L. Tomkins, and J. Hunt, “Sperm Competition Games Played by Dimorphic Male Beetles,” Proceedings of the Royal Society of London, Series B 266 (1999): 145–50.
6. For an overview of alternative reproductive tactics in animals, see R. F. Oliveira, M. Taborsky, and H. J. Brockmann, eds., Alternative Reproductive Tactics: An Integrative Approach (Cambridge: Cambridge University Press, 2008).
7. J. T. Hogg and S. H. Forbes, “Mating in Bighorn Sheep: Frequent Male Reproduction via a High-Risk ‘Unconventional’ Tactic,” Behavioral Ecology and Sociobiology 41 (1997): 33–48; D. W. Coltman, M. Festa-Bianchet, J. T. Jorgenson, and C. Strobeck, “Age-Dependent Sexual Selection in Bighorn Rams,” Proceedings of the Royal Society of London, Series B 269 (2002): 165–72.
8. M. R. Gross and E. L. Charnov, “Alternative Male Life Histories in Bluegill Sunfish,” Proceedings of the National Academy of Sciences 77 (1980): 6937–40; W. J. Dominey, “Maintenance of Female Mimicry as a Reproductive Strategy in Bluegill Sunfish (Lepomis macrochirus),” Environmental Biology of Fishes 6 (1981): 59–64; M. R. Gross, “Disruptive Selection for Alternative Life Histories in Salmon,” Nature 313 (1985): 47–48; C. J. Foote, G. S. Brown, and C. C. Wood, “Spawning Success of Males Using Alternative Mating Tactics in Sockeye Salmon, Oncorhynchus nerka,” Canadian Journal of Fisheries and Aquatic Sciences 54 (1997): 1785–95.
9. J. G. van Rhijn, “On the Maintenance and Origin of Alternative Strategies in the Ruff Philomachus pugnax,” Ibis 125 (1983): 482–98; D. B. Lank, C. M. Smith, O. Hanotte, T. Burke, and F. Cooke, “Genetic Polymorphism for Alternative Mating Behaviour in Lekking Male Ruff Philomachus pugnax,” Nature 378 (1995): 59–62.
10. J. Jukema and T. Piersma, “Permanent Female Mimics in a Lekking Shorebird,” Biology Letters 2 (2006): 161–64.
11. Ibid.
12. S. M. Shuster and M. J. Wade, “Female Copying and Sexual Selection in a Marine Isopod Crustacean, Paracerceis sculpta,” Animal Behaviour 41 (1991): 1071–78; S. M. Shuster, “The Reproductive Behaviour of α-, β-, and γ-Male Morphs in Paracerceis sculpta, a Marine Isopod Crustacean,” Behaviour 121 (1992): 231–58; S. M. Shuster and M. J. Wade, “Equal Mating Success Among Male Reproductive Strategies in a Marine Isopod,” Nature 350 (1991): 608–10.
13. Ibid.
14. R. T. Hanlon, M.-J. Naud, P. W. Shaw, J. T. Havenhand, “Behavioural Ecology: Transient Sexual Mimicry Leads to Fertilization,” Nature 433 (2005): 212.
15. Ibid.
16. Sun Tzu, The Art of War, trans. Samuel B. Griffith (New York: Oxford University, 1963); Mark McNeilly, Sun Tzu and the Art of Modern Warfare (Oxford: Oxford University Press, 2001).
17. Andrew Mack, “Why Big Nations Lose Small Wars: The Politics of Asymmetric Conflict,” World Politics 27 (1975): 175–200; Ivan Arreguin-Toft, “How the Weak Win Wars: A Theory of Asymmetric Conflict,” International Security 26 (2001): 93–128.
18. Ibid.
19. Ibid.
20. Raphael Perl and Ronald O’Rourke, “Terrorist Attack on USS Cole: Background and Issues for Congress,” in Emerging Technologies: Recommendations for Counter-Terrorism, eds. Joseph Rosen and Charles Lucey (Hanover, NH: Institute for Security Technology Studies, Dartmouth University, 2001): 52–58.
21. Trevor N. Dupuy, The Evolution of Weapons and Warfare (New York: Dacapo Press, 1984).
22. Brian Mazanec, “The Art of (Cyber) War,” Journal of International Security Affairs 16 (2009): 3–19; Jason Fritz, “How China Will Use Cyber Warfare to Leapfrog in Military Competitiveness,” Culture Mandala: The Bulletin of the Centre
for East-West Cultural and Economic Studies 8 (2008): 28–80.
23. Ibid.
24. Mark Clayton, “Chinese Cyberattacks Hit Key US Weapons Systems: Are They Still Reliable?” Christian Science Monitor, May 28, 2013; Ewen MacAskill, “Obama to Confront Chinese President Over Spate of Cyber-Attacks on US,” Guardian, May 28, 2013; Fritz, “How China Will Use Cyber Warfare to Leapfrog in Military Competitiveness,” 28–80.
25. Ibid.
26. Leyla Bilge and Tudor Dumitras, “Before We Knew It: An Empirical Study of Zero-Day Attacks in the Real World,” Proceedings of the 2012 ACM Conference on Computer and Communications Security, 2012: 833–44.
11. End of the Race
1. F. Kottenkamp, History of Chivalry and Ancient Armour (London: Willis and Sotheran Publishers, 1857); G. Duby, The Chivalrous Society, trans. Cynthia Poston (Berkeley: University of California Press, 1977); R. L. O’Connell, Of Arms and Men: A History of War, Weapons, and Aggression (Oxford: Oxford University Press, 1989); J. France, Western Warfare in the Age of the Crusades, 1000-1300 (Ithaca, NY: Cornell University Press, 1999).
2. Trevor N. Dupuy, The Evolution of Weapons and Warfare (New York: Da Capo Press, 1984).
3. Ibid.
4. Ibid.
5. Dupuy, Evolution of Weapons and Warfare; O’Connell, Of Arms and Men; R. L. O’Connell, Soul of the Sword: An Illustrated History of Weaponry and Warfare from Prehistory to the Present (New York: The Free Press, 2002).
6. Ibid.
7. Ibid.
8. Ibid.
9. Ibid.
10. Trevor Dupuy provides a great overview of the battle of Crécy in his book The Evolution of Weapons and Warfare. For more comprehensive treatments of the battle see Henri de Wailly, Crécy 1346: Anatomy of a Battle (Poole, NY: Blandford Press, 1987); and A. Ayton, P. Preston, F. Autrand, and B. Schnerb, The Battle of Crécy, 1346 (Woodbridge, Suffolk, UK: Boydell Press, 2005).
11. Ibid.
12. Ibid.
13. A vivid description of the soldier’s perspective of the battle of Agincourt is provided by John Keegan in his book The Face of Battle: A study of Agincourt, Waterloo, and the Somme (London: Penguin, 1983). For more comprehensive coverage of this battle, see J. Barker, Agincourt: The King, the Campaign, the Battle (London: Little Brown, 2005); and A. Curry, Agincourt: A New History (London: Tempus Publishing, 2005).
14. Dupuy, Evolution of Weapons and Warfare; O’Connell, Of Arms and Men; O’Connell, Soul of the Sword.
15. Surprisingly few studies have actually measured the strength and nature of selection on extreme weapons—it’s incredibly labor-intensive to do. But those that have often find that as weapons get bigger and bigger the success of males rises up to a point, beyond which success begins to drop. Males with the very largest weapons tend to do worse than males with slightly smaller weapons. If males with the biggest weapons of all fared the best, selection would be “open-ended” and directional. The fact that the very biggest males did slightly worse shows us that selection in these populations is stabilizing, and it suggests these populations may be at or close to their balance point. Examples of this type of selection can be found in harlequin beetles, amphipod crustaceans, and red deer. D. W. Zeh, J. A. Zeh, and G. Tavakilian, “Sexual Selection and Sexual Dimorphism in the Harlequin Beetle Acrocinus longimanus,” Biotropica (2002): 86–96; G. A. Wellborn, “Selection on a Sexually Dimorphic Trait in Ecotypes Within the Hyalella azteca Species Complex (Amphipoda: Hyalellidae),” American Midland Naturalist 143 (2000): 212–25; L. E. B. Kruuk, J. Slate, J. M. Pemberton, S. Brotherstone, F. Guinness, and T. Clutton-Brock, “Antler Size in Red Deer: Heritability and Selection but no Evolution,” Evolution 56 (2002): 1683–95.
16. To use the same basic equation I allude to in the text, the benefits of having big weapons (B), representing the offspring produced by the females they guard, are offset by production, maintenance, and fighting costs (C). When B − C > 0, then selection drives the evolution of weapons toward ever greater size. What’s not shown is our assumption that males are gleaning 100 percent of the fertilizations of the females they defend: (1 * B) − C > 0. Cheating males erode the rewards to dominant males, so that they reap only a fraction of the sired offspring that they otherwise would have. In the hypothetical beetle example where sneak males, on average, steal one-fourth of the fertilizations, the new equation would be: (0.75 * B) − C > 0. Benefits are devalued by the proportion of fertilizations stolen by cheaters (1 − 0.25 = 0.75). At some point, if cheaters steal enough, the realized benefits may be low enough that they no longer exceed the costs, even for the most successful guarding males.
17. R. Moen, J. Pastor, and Y. Cohen, “Antler Growth and Extinction of Irish Elk,” Evolutionary Ecology Research 1 (1999): 235–49.
18. Ibid.
19. R. Baker and G. Wilkinson, “Phylogenetic Analysis of Sexual Dimorphism and Eye Stalk Allometry in Stalk-Eyed Flies (Diopsidae),” Evolution 55 (2001): 1373–85; M. Kotrba, “Baltic Amber Fossils Reveal Early Evolution of Sexual Dimorphism in Stalk-Eyed Flies (Diptera: Diopsidae),” Organisms, Diversity and Evolution 2004 (2004): 265–75.
20. T. Hosoya and K. Araya, “Phylogeny of Japanese Stag Beetles (Coleoptera: Lucanidae) Inferred from 16s mtrRNA Gene Sequences, with References to the Evolution of Sexual Dimorphism of Mandibles,” Zoological Science 22 (2005): 1305–18.
21. M. Tabana and N. Okuda, “Notes on Nicagus japonicus Nagel,” Gekkan-Mushi 292 (1992): 17–21; K. Katovich and N. L. Kriska, “Description of the Larva of Nicagus obscurus (LeConte) (Coleoptera: Lucanidae: Nicaginae), with Comments on Its Position in Lucanidae and Notes on Adult Habitat,” Coleopterists Bulletin 56 (2002): 253–58.
22. D. J. Emlen, J. Marangelo, B. Ball, and C. W. Cunningham, “Diversity in the Weapons of Sexual Selection: Horn Evolution in the Beetle Genus Onthophagus (Coleoptera: Scarabaeidae),” Evolution 59 (2005): 1060-84.
23. T. M. Caro, C. M. Graham, C. J. Stoner, and M. M. Flores, “Correlates of Horn and Antler Shape in Bovids and Cervids,” Behaviorial Ecology and Sociobiology 55 (2003): 32–41; J. Bro-Jørgensen, “The Intensity of Sexual Selection Predicts Weapon Size in Male Bovids,” Evolution 61 (2007): 1316–26.
24. J. L. Coggeshall, “The Fireship and Its Role in the Royal Navy” (dissertation, Texas A&M University, 1997); P. Kirsch, Fireship: The Terror Weapon of the Age of Sail, trans. John Harland (Barnsley, UK: Seaforth Publishing, 2009).
25. O’Connell, Of Arms and Men; Coggeshall, “The Fireship and Its Role in the Royal Navy”; O’Connell, Soul of the Sword; R. Gardiner and B. Lavery, The Line of Battle: The Sailing Warship 1650–1840 (London Conway Maritime Press, 2004); Kirsch, Fireship.
26. Ibid.
27. James Coggeshall provides a nice description of the role of fireships in the English attack on the Spanish Armada in his dissertation “The Fireship and Its Role in the Royal Navy.” More thorough treatments of this battle are provided in Michael Lewis, The Spanish Armada (New York: T. Y. Crowell, 1968); Colin Martin and Geoffrey Parker, The Spanish Armada (New York: Penguin Books, 1999).
28. O’Connell, Of Arms and Men; Coggeshall, “The Fireship and Its Role in the Royal Navy”; O’Connell, Soul of the Sword; Gardiner and Lavery, Line of Battle; Robert Jackson, Sea Warfare: From World War I to the Present (San Diego: Thunder Bay Press, 2008).
29. Gardiner and Lavery, Line of Battle.
30. O’Connell, Of Arms and Men; O’Connell, Sacred Vessels: The Cult of the Battleship and the Rise of the U.S. Navy (Oxford: Oxford University Press, 1991); O’Connell, Soul of the Sword; Gardiner and Lavery, Line of Battle; Jackson, Sea Warfare.
31. O’Connell, Of Arms and Men; O’Connell, Sacred Vessels; O’Connell, Soul of the Sword; R. K. Massie, Dreadnought: Britain, Germany, and the Coming of the Great War (New York: Random House, 2007); Massie, Castles of Steel: Britain, Germany and the Winning of the Great War at Sea (New York: Random House, 2008).
32. Ibid.
33. Ibid.
34. Ibid.
35. Ibid.
36. Massie, Dreadnought; Massie, Castles of Steel; Jackson, Sea Warfare.
37. Ibid.
38. Jackson, Sea Warfare.
39. Ibid.
40. Ibid.
41. Ibid.
42. Tony Bridgland, Sea Killers in Disguise: The Story of the Q-Ships and Decoy Ships in the First World War (Annapolis, MD: Naval Institute Press, 1999).
43. Ibid.
44. Jackson, Sea Warfare.
45. Ibid.
12. Castles of Sand and Stone
1. For excellent treatments of the biology of African army ants, or driver ants, see W. H. Gotwald Jr., The Army Ants: The Biology of Social Predation (Ithaca, NY: Cornell University Press, 1995); and B. Hölldobler and Edward O. Wilson, The Ants (Cambridge, MA: Belknap Press of Harvard University Press, 1990). For vivid and highly readable accounts of ant battles, I recommend works by Mark Moffett, including Adventures Among Ants: A Global Safari with a Cast of Trillions (Berkeley: University of California Press, 2010), and “Ants and the Art of War,” Scientific American, December 2011, 84–9.
2. Hölldobler and Wilson, Ants.
3. Caspar Schöning and Mark W. Moffett, “Driver Ants Invading a Termite Nest: Why Do the Most Catholic Predators of All Seldom Take This Abundant Prey?” Biotropica 39 (2007): 663–67.
4. W. H. Gotwald Jr., “Predatory Behavior and Food Preferences of Driver Ants in Selected African Habitats,” Annals of the Entomological Society of America 67 (1974): 877–86; Gotwald, Army Ants.
5. For comprehensive treatments of the biology of termites, I highly recommend Takuya Abe, David Edward Bignell, and Masahiko Higashi, Termites: Evolution, Sociality, Symbioses, Ecology (Boston: Kluwer Academic Publishers, 2000); and David Edward Bignell, Yves Roisin, and Nathan Lo, The Biology of Termites: A Modern Synthesis (New York: Springer, 2011).
6. Johanna P. E. C. Darlington, “Populations in Nests of the Termite Macrotermes subhyalinus in Kenya,” Insectes Sociaux 37 (1990): 158–68.
7. Charles Noirot and Johanna P. E. C. Darlington, “Termite Nests: Architecture, Regulation, and Defence,” chap. 6 in Abe, Bignell, and Higashi, Termites; Judith Korb, “Termite Mound Architecture, from Function to Construction,” chap. 13 in Bignell, Roisin, and Lo, Biology of Termites.
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