Army ants overwhelm their prey with strength in numbers. Soldiers have big heads with strong jaws, but they still need to be mobile since they march along with columns of workers on raids.
Above: Termite soldiers have even bigger heads than army ant soldiers, in part because, unlike ants, they do not need to run long distances. Instead, they position themselves at the entrances to tunnels, biting anything that attempts to enter. Below: Termites rely on fortresses to defend themselves against attacks by army ants. By restricting access to a small number of tunnels, termites remove the numerical advantage of the army ants, forcing their soldiers to confront termite soldiers in duels the termites are likely to win.
Castles rely on heavily guarded, tiny entrances, too. Gatehouses work like tunnels of termite nests, stripping away the numerical advantage of attacking armies.
Above: Fortification styles evolved in tandem with ever more effective artillery. Early structures relied on square-sided, protruding towers to provide flanking fire along the walls, but corners proved vulnerable to flying boulders. Below: Later castles had round towers to deflect artillery. Even when struck directly, these towers were less likely to shatter.
Cannons destroyed even the most magnificent castles, shattering towers, breaching walls, and collapsing the arms race, until a new style of fortress emerged. Star forts sit low to the ground and rely on extensive earthworks to absorb the impact of cannons, and angled, pointy walls to deflect cannonballs fired from any direction.
Exploding artillery spelled the end for even these forts. From WWII onward the safest place to hide has been in dispersed bunkers deep belowground, like Cheyenne Mountain, headquarters of NORAD during the Cold War.
Medieval knights fought as individuals in duels with rival knights. Above: The most common battles were tournaments—structured spectacles with strict rules—that tended to favor the more heavily armed and better trained contestor. Below: Even in full battle, knights confronted other knights in hand-to-hand combat that often unfolded as duels.
Acknowledgments
A project of this scope would not have been possible without support from colleagues, family, and friends. First and foremost, I thank my family: my wife, Kerry, and my children, Cory and Nicole, for bearing with me as I became ever more consumed by this process. I thank my colleagues, collaborators, and students, who picked up slack while I hid in the stacks of the library writing, especially Cerisse Allen, Laura Corley-Lavine, Annika Duke, Ian Dworkin, Hiroki Gotoh, Erin McCullough, Devin O’Brien, Jema Rushe, Jennifer Smith, Ian Warren, and Robbie Zinna. I thank the National Science Foundation, especially Zoe Eppley, Irwin Forseth, Dianna Padilla, Adam Summers, Kimberlyn Williams, and William Zamer, for funding my research program. The NSF plays an absolutely vital role in supporting basic research. They are the lifeblood of scholarship in the United States, and none of my work would have been possible without them.
Writing a book of this nature required a great deal of “deprogramming.” I had to strip away all the stylistic rules I teach my students so assiduously. Twenty years of scripting grant proposals and academic papers for technical journals proved to be a hindrance, rather than an asset. I had to start over—learn to write anew. This was a refreshing (and exhausting) experience, and I could not possibly have managed it without the patience and critical feedback of my editor, Gillian Blake, her assistant, Caroline Zancan, and my colleague and friend Carl Zimmer, all of whom took me to task on multiple occasions.
This work benefitted greatly from feedback from many additional people. For critiquing all or parts of this manuscript in its various forms, I thank Brett Addis, Harrison Ambrose III, Harrison Ambrose IV, Katharine Ambrose, Tina Bennett, Alexis Billings, Kelly Bright, Kerry Bright, Ray Bright, Kristen Crandell, Annika Duke, Cory Emlen, Natalia Demong Emlen, Stephen Emlen, Daphne Fairbairn, Harry Greene, Melissa Hamre, Matthew Herron, Erin Kuiper, Tara Maginnis, Christine Miller, Devin O’Brien, Alison Perkins, Mike Ryan, David Tuss, and Carl Zimmer. Katharine Ambrose, Kerry Bright, Stephen Emlen, Erin McCullough, Alison Perkins, and David Tuss, in particular, worked through the entire manuscript more than once, providing feedback that dramatically improved the final text.
David Tuss was a pleasure to work with, patiently bouncing illustrations back and forth with me, reworking and, at times, completely redrawing figures until we both were thrilled with the outcome. For sharing details of their research experiences, I thank John Christy, Gerald Wilkinson, and David Zeh; for guiding me into the world of publishing, I thank Ben Roberts; and for first suggesting that I delve into the literature on human weapons, I thank Alison Kalett.
Military history turned out to be a vast and overwhelming literature, and a few authors stood out for me. The writings of Robert O’Connell, in particular, were transformative as I stepped, tentatively at first, and then with ever greater tenacity, out of the world of biology and into that of military history. In many ways, O’Connell has done in his books the same thing that I attempt here, but in reverse. His expertise is in military history, yet he reaches into biology as needed to provide perspective. I write about biology, dipping into military history for precisely the same reason. For readers interested in this interface of biology and history, I highly recommend his books Of Arms and Men: A History of War, Weapons, and Aggression (Oxford: Oxford University Press, 1989) and Soul of the Sword: An Illustrated History of Weapons and Warfare from Prehistory to the Present (New York: Free Press, 2002). I also thank him for generously taking the time to read through this manuscript, correcting many of the details of my coverage of military history, particularly material related to the Cold War.
I also highly recommend Trevor Dupuy’s book The Evolution of Weapons and Warfare (New York: Da Capo Press, 1984), which helped put the major transitions in military technology into perspective for me; John Keegan’s The Face of Battle: A Study of Agincourt, Waterloo, and the Somme (London: Penguin Books, 1983), which brought the realities of ancient battle to life with startling vividness and relevance; and David Hoffman’s The Dead Hand: The Untold Story of the Cold War Arms Race and Its Dangerous Legacy (New York: Doubleday, 2009)—brilliant, poignant, and terrifying in its portrayal of the aftermath of history’s deadliest arms race.
I thank Caffé Dolce and its wonderful staff for the perfect place to think and write, as well as for the best coffee in Missoula. Molly Bloom, Michelle Daniel, and Meryl Levavi, my Henry Holt production team, were thorough and patient—exactly what I needed. Finally, I’d like to thank my agent, Tina Bennett, and her assistant, Svetlana Katz, for unflinching support and guidance through all of the stages of this process. I could not have done this without them.
ALSO BY DOUGLAS J. EMLEN
Evolution: Making Sense of Life (WITH CARL ZIMMER)
A Handbook of Biological Investigation (WITH HARRISON W. AMBROSE III, KATHARINE PECKHAM AMBROSE, AND KERRY L. BRIGHT)
About the Author
DOUGLAS J. EMLEN is a professor of biology at the University of Montana. He is the recipient of the Presidential Early Career Award for Scientists and Engineers from the Office of Science and Technology Policy at the White House, multiple research awards from the National Science Foundation, including its five-year CAREER award, and a Young Investigators Award and the E. O. Wilson Naturalist Award from the American Society of Naturalists. His research has been featured in outlets including the New York Times and National Public Radio’s Fresh Air.
ANIMAL WEAPONS: THE EVOLUTION OF BATTLE
Copyright © 2014 by Douglas J. Emlen. All rights reserved. For information, address Henry Holt and Co., 175 Fifth Avenue, New York, N.Y. 10010.
www.henryholt.com
Jacket art by David J. Tuss
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The Library of Congress has cataloged the print edition as follows:
Emlen, Douglas J
ohn, 1967–
Animal weapons: the evolution of battle / by Douglas J. Emlen; illustrated by David J. Tuss.—First edition.
pages cm
Includes bibliographical references and index.
ISBN 978-0-8050-9450-3 (hardcover)—ISBN 978-1-4299-4739-8 (electronic book)
1. Animal weapons. 2. Animal defenses. 3. Defensive (Military science). I. Title.
QL940.E45 2014
591.47—dc23 2014004772
e-ISBN 978-1-4299-4739-8
First Edition: November 2014
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