Animal Weapons
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9. Wonderful accounts of the behavior of dung beetles are provided in Gonzalo Halffter and Eric G. Matthews, The Natural History of Dung Beetles of the Subfamily Scarabaeinae (Coleoptera, Scarabaeidae) (Palermo, Italy: Medical Books di G. Cafaro, 1966); Gonzalo Halffter and William David Edmonds, The Nesting Behavior of Dung Beetles (Scarabaeinae): An Ecological and Evolutive Approach (Mexico, D.F.: Instituto de Ecologia, 1982); and Leigh W. Simmons and James T. Ridsdill-Smith, Ecology and Evolution of Dung Beetles (Oxford: Blackwell Publishing, 2011). I also recommend papers by Hiroaki Sato, such as H. Sato and M. Imamori, “Nesting Behaviour of a Subsocial African Ball-Roller Kheper platynotus (Coleoptera, Scarabaeidae),” Ecological Entomology 12 (1987): 415–25; and H. Sato, “Two Nesting Behaviours and Life History of a Subsocial African Dung Rolling Beetle, Scarabaeus catenatus (Coleoptera: Scarabaeidae),” Journal of Natural History 31 (1997): 457–69.
10. Keith Philips and I used a phylogenetic tree depicting the branching relationships among dung beetle species to test for an association between evolutionary gains or losses of male horns and the tunneling versus rolling behavior of each species. We found that tunneling behavior strongly predicted the evolution of horns, and when species switched from tunneling to ball-rolling behavior, they subsequently lost their male horns. D. J. Emlen and T. K. Philips, “Phylogenetic Evidence for an Association Between Tunneling Behavior and the Evolution of Horns in Dung Beetles (Coleoptera: Scarabaeidae: Scarabaeinae),” in Coleopterists Society Monographs 5 (2006): 47–56.
11. D. J. Emlen, “Alternative Reproductive Tactics and Male Dimorphism in the Horned Beetle Onthophagus acuminatus,” Behavioral Ecology and Sociobiology 41 (1997): 335–41; A. P. Moczek and D. J. Emlen, “Male Horn Dimorphism in the Scarab Beetle Onthophagus taurus: Do Alternative Tactics Favor Alternative Phenotypes?” Animal Behaviour 59 (2000): 459–66.
6. Duels
1. For a biography of Lanchester see P. W. Kingsford, F. W. Lanchester: A Life of an Engineer (London: Edward Arnold, 1960).
2. Frederick W. Lanchester, Aircraft in Warfare: The Dawn of the Fourth Arm (London: Constable, 1916).
3. Phillip M. Morse and George E. Kimball, Methods of Operations Research (New York: John Wiley and Sons, 1951); and James G. Taylor, Lanchester Models of Warfare (Arlington, VA: Operations Research Society of America, 1983).
4. For example, see P. R. Wallis, “Recent Developments in Lanchester Theory,” Operations Research 19 (1968): 191–95, which reports on the Operational Research Conference on Recent Developments in Lanchester Theory, sponsored by the NATO Science Committee and held in Munich in July 1967.
5. For examples, see P. Morse and G. Kimball, Methods of Operations Research (Cambridge, MA: Technology Press of MIT, 1951), or Frederick S. Hillier and Gerald J. Lieberman, Introduction to Operations Research, 9th ed. (Boston: McGraw Hill, 2009). In 1956, the journal Operations Research dedicated an issue to the memory of Frederick Lanchester: Joseph McCloskey, “Of Horseless Carriages, Flying Machines, and Operations Research: A Tribute to Frederick Lanchester,” 4 (1956): 141–47. To this day, the Institute for Operations Research and Management Sciences (INFORMS) names its highest prize after Lanchester.
6. For a superb explanation of the logic of Lanchester’s models see John W. R. Lepingwell, “The Laws of Combat? Lanchester Re-examined,” International Security 12 (1987): 89–134.
7. John Keegan, The Face of Battle: A Study of Agincourt, Waterloo, and the Somme (London: Penguin Books, 1983).
8. Lanchester, Aircraft in Warfare; Lepingwell, “Laws of Combat?”: 89–134.
9. In fact, at Agincourt, the advantage worked the other way. The French marched into battle using traditional close-range tactics of knights in armor, but the English combined the strength of their knights with a new type of weapon, the longbow, wielded by thousands of archers. The English were able to concentrate arrow fire in a way that the French could not, and, despite being drastically outnumbered at the outset of the battle, the English won the day. For reasons we come back to in later chapters, this battle and others like it (for example, the Battle of Crécy) marked significant turning points in the nature of battle, spelling the beginning of the end for knights resplendent in suits of expensive armor.
10. For examples, see J. H. Engel, “A Verification of Lanchester’s Law,” Operations Research 2 (1954): 163–71; Thomas W. Lucas and Turker Turkes, “Fitting Lanchester’s Equations to the Battles of Kursk and Ardennes,” Naval Research Logistics 54 (2003): 95–116; and Taylor, Lanchester Models of Warfare.
11. Lepingwell, “Laws of Combat?,” 89–134.
12. For explanations of this logic and application to military history, see R. L. O’Connell, Of Arms and Men: A History of War, Weapons, and Aggression (Oxford: Oxford University Press, 1990).
13. Many social insects do fight battles as armies, with swarms from one colony engaging swarms from another. Several authors have applied Lanchester’s linear and square laws to these battles. For example, see N. R. Franks and L. W. Partridge, “Lanchester Battles and the Evolution of Combat in Ants,” Animal Behaviour 45 (1993): 197–99; T. P. McGlynn, “Do Lanchester’s Laws of Combat Describe Competition in Ants?” Behavioral Ecology 11 (2000): 686–90; Martin Pfeiffer and Karl E. Linsenmair, “Territoriality in the Malaysian Giant Ant Camponotus gigas (Hymenoptera/Formicidae),” Journal of Ethology 19 (2001): 75–85; and Nicola J. R. Plowes and Eldridge S. Adams, “An Empirical Test of Lanchester’s Square Law: Mortality During Battles of the Fire Ant Solenopsis invicta,” Behavioral Ecology 272 (2005): 1809–14.
14. Jon M. Hastings, “The Influence of Size, Age, and Residency Status on Territory Defense in Male Western Cicada Killer Wasps (Sphecius grandis, Hymenoptera: Sphecidae),” Journal of the Kansas Entomological Society 62 (1989): 363–73.
15. Cicada-killer wasps may lack big weapons for a second reason, too. For many insects that fight in the air, agility and maneuverability matter even more than strength or size. Big weapons may hinder mobility in these fights in much the same way that weapons impede speed in predators. Many wasps, dragonflies, damselflies, and butterflies fight vicious, acrobatic airborne battles, and almost all of these species lack elaborate weapons. Part of this, no doubt, is due to unpredictability arising from scrambles; the rest is likely balancing selection arising from the need for agility. For example papers of fights in these types of insects, see Greg F. Grether, “Intrasexual Competition Alone Favors a Sexually Selected Dimorphic Ornament in the Rubyspot Damselfly Hetaerina americana,” Evolution 50 (1996): 1949–57; D. J. Kemp and C. Wiklund, “Fighting Without Weaponry: A Review of Male-Male Contest Competition in Butterflies,” Behavioral Ecology and Sociobiology 49 (2001): 429–42; J. Contreras-Garduño, J. Canales-Lazcana, and A. Córdoba-Aguilar, “Wing Pigmentation, Immune Ability, Fat Reserves, and Territorial Status in Males of the Rubyspot Damselfly, Hetaerina americana,” Journal of Ethology 24 (2006): 165–73; M. A. Serrano-Meneses, A. Córdoba-Aguilar, V. Méndez, S. J. Layen, and T. Székely, “Sexual Size Dimorphism in the American Rubyspot: Male Body Size Predicts Male Competition and Mating Success,” Animal Behaviour 73 (2007): 987–97.
16. The most comprehensive work on mating behavior and sexual selection in horseshoe crabs has been conducted by Jane Brockmann and her students at the University of Florida. For example, see H. Jane Brockmann and Dustin Penn, “Male Mating Tactics in the Horseshoe Crab, Limulus polyphemus,” Animal Behaviour 44 (1992): 653–65.
17. O’Connell, Of Arms and Men.
18. J. H. Christy and M. Salmon, “Ecology and Evolution of Mating Systems of Fiddler Crabs (Genus Uca),” Biological Reviews 59 (1984): 483–509; N. Knowlton and B. D. Keller, “Symmetric Fights as a Measure of Escalation Potential in a Symbiotic, Territorial Snapping Shrimp,” Behavioral Ecology and Sociobiology 10 (1982): 289–92; M. D. Jennions and P. R. Y. Backwell, “Residency and Size Affect Fight Duration and Outcome in the Fiddler Crab Uca annulipes,” Biological Journal of the Linnean Society 57 (1996): 293–306.
19. The most co
mprehensive work to date on these bizarre wasps has been done by Robert Longair of the University of Calgary. His field studies in the Ivory Coast showed that males used their long tusks in fights with rival males over mud burrow–like nests on the undersides of leaves containing newly emerging females. See, for example, Robert W. Longair, “Tusked Males, Male Dimorphism, and Nesting Behavior in a Subsocial Afrotropical Wasp, Synagris cornuta, and Weapons and Dimorphism in the Genus (Hymenoptera: Vespidae: Eumeninae),” Journal of the Kansas Entomological Society 77 (2004): 528–57.
20. An early study from 1931 showed that the rhinoceros beetle Diloboderus fights over burrows in the soil: J. B. Daguerre, “Costumbres Nupciales del Diloboderus abderus Sturm,” Rev. Soc. Entomologia Argentina 3 (1931): 253–56. Several studies by William Eberhard have looked at fighting behavior of rhinoceros beetles inside hollowed-out plant stems. See his book chapter “The Function of Horns in Podischnus agenor (Dynastinae) and Other Beetles” in Sexual Selection and Reproductive Competition in Insects, ed. M. S. Blum and N. A. Blum (New York: Academic Press, 1979), 231–59, and his paper “Use of Horns in Fights by the Dimorphic Males of Ageopsis nigricollis Coleoptera Scarabeidae, Dynastinae,” Journal of the Kansas Entomological Society 60 (1987): 504–9.
21. Tusked frogs are a truly bizarre group of amphibian species. For more on their morphology and behavior I particularly recommend Sharon Emerson, “Courtship and Nest-Building Behavior of a Bornean Frog, Rana blythi,” Copeia 1992 (1992): 1123–27; Kaliope Katsikaros and Richard Shine, “Sexual Dimorphism in the Tusked Frog, Adelotus brevis (Anura: Myobatrachidae): the Roles of Natural and Sexual Selection,” Biological Journal of the Linnean Society 60 (1997): 39–51; and Hiroshi Tsuji and Masafumi Matsui, “Male-Male Combat and Head Morphology in a Fanged Frog (Rana kuhlii) from Taiwan,” Journal of Herpetology 36 (2002): 520–26.
22. S. S. B. Hopkins, “The Evolution of Fossoriality and the Adaptive Role of Horns in the Mylagaulidae (Mammalia: Rodentia),” Proceedings of the Royal Society of London Series B, Biological Sciences 272 (2005): 1705–13.
23. My first attempt at dissertation research (before I ended up in Panama) was a study of horns in the giant rhinoceros beetle Golofa porteri in Ecuador and southern Colombia, where these beetles fight to defend new plant shoots of a bamboo-like plant high up on cloud forest ridges. My attempts failed miserably, as I was unable to locate large populations to study. But that project was inspired by the beautiful paper by William Eberhard entitled “Fighting Behavior of Male Golofa porteri Beetles (Scarabaeidae: Dynastinae),” Psyche 83 (1978): 292–98.
24. Most work on the evolution of enlarged male hind legs in leaf-footed bugs has been done by Takahisa Miyatake of the University of Ryukyus in Japan. For example, see his paper “Territorial Mating Aggregation in the Bamboo Bug, Notobitus meleagris, Fabricius (Heteroptera: Coreidae),” Journal of Ethology, 13 (1995): 185–89, or his paper “Functional Morphology of the Hind Legs as Weapons for Male Contests in Leptoglossus australis (Heteroptera: Coreidae),” Journal of Insect Behavior 10 (1997): 727–35. Also see the paper by William Eberhard, “Sexual Behavior of Acanthocephala declivis guatemalana (Hemiptera: Coreidae) and the Allometric Scaling of their Modified Hind Legs,” Annals of the Entomological Society of America 91 (1998): 863–71.
25. Horned chameleons, despite their notoriety, remain almost completely unstudied in the wild. One early study was conducted by Stanley Rand, “A Suggested Function of the Ornamentation of East African Forest Chameleons,” Copeia 1961 (1961): 411–14. Another was conducted by Stephen Parcher, “Observations on the Natural Histories of Six Malagasy Chamaeleontidae,” Zeitschrift für Tierpsychologie 34 (1974): 500–23.
26. Tadatsugu Hosoya and Kunio Araya have a beautiful paper entitled “Phylogeny of Japanese Stag Beetles (Coleoptera: Lucanidae) Inferred from 16S mtrRNA Gene Sequences, with Reference to the Evolution of Sexual Dimorphism of Mandibles,” Zoological Science 22 (2005): 1305–18. In this paper the authors trace the evolutionary history of enlarged male mandibles in stag beetles. Their data suggest that huge weapons arose independently in at least two separate lineages of these beetles and that, once there, enlarged male mandibles were lost several different times. They are able to interpret these evolutionary losses of male weapons in the context of the natural history and behavior of the beetles.
27. These little flies have proven to be very difficult to study. They have yet to be successfully reared in the laboratory, and most species live in remote parts of New Guinea and surrounding islands. One species makes it into tropical northern Australia. Systematic study and taxonomy of these amazing flies has been conducted largely by David McAlpine, including “A Systematic Study of Phytalmia (Diptera, Tephritidae) with Description of a New Genus,” Systematic Entomology 3 (1978): 159–75. The first field study of these flies that I am aware of is the paper by M. S. Moulds, “Field Observations on the Behavior of a North Queensland Species of Phytalmia (Diptera: Tephritidae),” Journal of the Australian Entomological Society 16 (1978): 347–52. More recently, the behavior of these flies has been studied by Gary Dodson, “Resource Defense Mating System in Antlered Flies, Phytalmia spp. (Diptera: Tephritidae),” Annals of the Entomological Society of America 90 (1997): 496–504.
28. The classic papers on stalk-eyed fly behavior are by Dietrich Burkhardt and Ingrid de la Motte, including their papers “Big ‘Antlers’ are Favoured: Female Choice in Stalk-Eyed Flies (Diptera, Insecta), Field Collected Harems and Laboratory Experiments,” Journal of Comparative Physiology A 162 (1988): 649–52; and “Signalling Fitness: Larger Males Sire More Offspring: Studies of the Stalk-Eyed Fly Cyrtodiopsis whitei (Diopsidae, Diptera),” Journal of Comparative Physiology A 174 (1994): 61–4.
29. Gerald Wilkinson at the University of Maryland has studied the behavior and genetics of stalk-eyed flies for almost twenty years. He and many of his doctoral students and postdoctoral research fellows have constructed phylogenies for this family of flies, conducted multigeneration experiments with flies in the laboratory, and studied several species in the field. Some of my favorites of their papers are Patrick Lorch, Gerald Wilkinson, and Paul Reillo, “Copulation Duration and Sperm Precedence in the Stalk-Eyed Fly, Cyrtodiopsis whitei (Diptera: Diopsidae),” Behavioral Ecology and Sociobiology 32 (1993): 303–11; Gerald Wilkinson and Gary Dodson, “Function and Evolution of Antlers and Eye Stalks in Flies,” in The Evolution of Mating Systems in Insects and Arachnids, ed. J. Choe and B. Crespi (Cambridge: Cambridge University Press, 1997), 310–28; and Tami Panhuis and Gerald Wilkinson, “Exaggerated Male Eye Span Influences Contest Outcome in Stalk-Eyed Flies,” Behavioral Ecology and Sociobiology 46 (1999): 221–27. I also recommend Rick Baker and Gerald Wilkinson, “Phylogenetic Analysis of Eye Stalk Allometry and Sexual Dimorphism in Stalk-Eyed Flies (Diopsidae),” Evolution 55 (2001): 1373–85.
30. Kevin Fowler and Andrew Pomiankowski head a stalk-eyed fly research group at the University College London. Their group has combined field studies of sexual selection in these flies with laboratory studies of eyestalk development. I recommend Patrice David, Andrew Hingle, D. Greig, A. Rutherford, Andrew Pomiankowski, and Kevin Fowler, “Male Sexual Ornament Size but not Asymmetry Reflects Condition in Stalk-Eyed Flies,” Proceedings of the Royal Society B: Biological Sciences 265 (1998): 2211–16; Andrew Hingle, Kevin Fowler, and Andrew Pomiankowski, “Size-Dependent Mate Preference in the Stalk-Eyed Fly Cyrtodiopsis dalmanni,” Animal Behaviour 61 (2001): 589–95; and Jen Small, Sam Cotton, Kevin Fowler, and Andrew Pomiankowski, “Male Eyespan and Resource Ownership Affect Contest Outcome in the Stalk-Eyed Fly, Teleopsis dalmanni,” Animal Behaviour 78 (2009): 1213–20.
31. For eloquent treatments of this spectacular period of naval warfare, see W. Murray, The Age of the Titans: The Rise and Fall of the Great Hellenistic Navies (Oxford: Oxford University Press, 2012); and John D. Grainger, Hellenistic and Roman Naval Wars 336BC–31BC (South Yorkshire, UK: Pen and Sword Books, 2011). The classic book on this period is Lionel Casson’s Ships and Seamanship
in the Ancient World (Baltimore: Johns Hopkins University Press, 1995), and I also recommend his book The Ancient Mariners, 2nd ed. (Princeton, NJ: Princeton University Press, 1991). For a particularly well-illustrated account of the ships and this period see Robert Gardiner, ed., The Age of the Galley: Mediterranean Oared Vessels Since Pre-Classical Times (London: Book Sales Publishing, 2000).
32. John Morrison and John Coates, Greek and Roman Oared Warships 399–30BC (Oxford: Oxbow Books, 1997).
33. My favorite account of this arms race, including especially the idea that the battering ram changed the nature of combat by causing ships to interact like individuals, thus fulfilling Lanchester’s linear law, is provided by Robert L. O’Connell in his book Of Arms and Men, and in his book Soul of the Sword: An Illustrated History of Weaponry and Warfare from Prehistory to the Present (New York: Free Press, 2002).
34. John Morrison and John Coates, Greek and Roman Oared Warships.
35. John Morrison and Roderick Williams, Greek Oared Ships (Cambridge: Cambridge University Press, 1968).
36. Gardiner, Age of the Galley.
37. Casson, Ships and Seamanship in the Ancient World; Gardiner, Age of the Galley; and O’Connell, Soul of the Sword.