Introduction

Silent, insidious, devastating. This is how an entomological attack is likely to unfold today. But to imagine modern uses of insects as weapons, we must look to the past. History reveals an unholy trinity of strategies—transmission of pathogenic microbes, destruction of livestock and crops, and direct attacks on humans—through which six-legged soldiers have wreaked havoc on human society.

The woman nervously checks her watch. In an hour, the human tide of the New York City rush hour will begin to pour underground. But, for the moment, there are few people who can see her reach into a shopping bag and take out a soda can. The woman peels off a strip of tape covering the opening and rolls the can beneath a bench on the subway platform. She heads to the escalator a bit more quickly than she and her fellow terrorists had been trained to move, but she's anxious to complete the other deposits. The woman and the five hundred hungry fleas make their respective escapes. She can only guess how many commuters will find red lumps on their legs in the morning, but whoever is bitten will be wracked by fever within days. Swelling lymph nodes might tip off a perceptive physician, but most of the victims will succumb to the ravages of bubonic plague. She knows that only a few hundred Americans will die, but millions will panic when they realize their vulnerability.

Bacteria-laden fleas spread throughout a subway system would echo the most terrible military use of insects in human history—disease vectors. The most devastating entomological attack took place in 1343, when Janibeg, the last Mongol khan, unwittingly allied with insect-borne disease in attempting to take the city of Kaffa. The Asian leader never suspected the role that fleas played in the ensuing pandemic that killed 25 million people. But neither did Europe's foremost military leader understand that his greatest defeats were caused by insects.

In 1799, Napoleon Bonaparte's campaign against the Ottoman Empire was defeated by flea-borne plague, and three years later his bid to establish a stepping stone into North America was crushed by yellow fever mosquitoes in Haiti. Napoleon's worst defeat by insects came in 1812. Rather than taking Russia, his Grande Armée lost 200,000 men to louse-borne typhus—a disaster that was replayed a hundred years later.

If the Austrians had established a western front against Russia, the course of World War I might have changed dramatically. But typhus kept the Central Powers from invading Serbia. In the Second World War insects were weap-onized by the Japanese. General Ishii Shiro's Unit 731 produced hundreds of millions of infected insects and dispersed them across China (and attempted to infiltrate the United States). By the end of the war, Ishii's fleas and flies were responsible for more deaths than the atomic bombs dropped on Japan.

Not to be outdone, scientists at Fort Detrick, Maryland, developed entomological weapons and conducted secret, open-air trials with (uninfected) vectors over U.S. populations in the 1950s. During the Korean War, the North Koreans and Chinese assembled a massive dossier to support the accusation that the U.S. military released an entomological potpourri infected with a microbial menagerie. The Americans passionately denied the charges, as they did when accused of using insects to spread disease in Cuba and Vietnam. But governments often disavow politically problematic knowledge.

In recent years, a troubling theory was quashed by federal agencies: the possibility that bioterrorists were responsible for the outbreak of West Nile virus. Saddam Hussein's minions had the motive, means, and opportunity, but the evidence is too circumstantial to accept—and too intriguing to ignore. We were virtually unable to check this mosquito-borne disease, and West Nile virus was a case of the sniffles compared to what would happen should Rift Valley fever be introduced. But an entomological attack need not target humans to inflict a terrible toll.

A white cardboard tent hangs in a tree like a Lilliputian bivouac for an elfin tree-climber. But the structure has no fanciful function, as the printed warnings make clear. Striding officiously beneath the luxuriant canopy of the orange grove, a man stops and removes the tent. The badge affixed to his breast pocket affirms that he is empowered to disturb federal property. He scowls upon seeing a series of cabalistic symbols scrawled over the warning label. The man peers inside, then rips open the tent and utters a mixed curse and prayer. The sticky interior of the tent has trapped dozens of flies. Their wings look like tiny stained-glass windows crafted in amber tones. These are the only Mediterranean fruit flies he's ever seen outside of the tedious training sessions in Sacramento. By the end of summer, $100 million in fruit will fall from the trees and rot in the California sun.

The second major use of insects as weapons is as assailants of agriculture. Starving one's enemy or crippling his economy by unleashing insects to destroy crops or livestock is a relatively recent innovation. Although farmers have battled pests for millennia, not until we mastered the industrial-scale mass production of insects could inducing hunger and poverty through entomological warfare become a military strategy.

In 1938, a British scientist warned: "It would be very surprising, for example, if insect pests . . . were not [dispersed] by hostile aeroplanes in the course of a future war." And by the summer of 1944, Germany had stockpiled 30 million secret weapons: the Colorado potato beetle. Whether an insectan Blitzkrieg landed on Britain's farms is a matter of debate, but when the Second World War devolved into the Cold War, using insects as covert weapons against an enemy's agriculture became very tempting.

As Cuba was poised to start a nuclear Armageddon in 1962, the U.S. Army was prepared to release planthoppers to destroy the Cuban sugarcane crop and cripple the nation's export economy. Neither missiles nor insects were launched that October, but for years Castro accused the Americans of infesting Cuban farms with aphids, beetles, moths, and mites. And in 1997 Cuba formally charged the U.S. State Department with releasing thrips to decimate the island nation's agriculture. The United Nations concluded that the pest outbreak "most likely" arose from an accidental introduction.

Encouraged by their ally's success in drawing world attention to the U.S. entomological warfare program, the North Vietnamese reported that Americans had loosed "killer insects" on the countryside (the dead crops probably succumbed to Agent Orange). Developing countries might be vulnerable, but most military strategists believe that modern pest-management practices make it impossible to starve an industrial nation via entomological warfare. Economic losses are another matter.

The Asian longhorned beetle that was accidentally introduced to the United States in 1995 will, if not stopped, inflict an economic toll exceeding the cost of the attack on the World Trade Center—an entomological scenario not lost on bioterrorists. In 1989, a covert group of environmental radicals threatened to release Medflies—voracious pests of valuable fruit crops—unless insecticide spraying was halted in California. Had they succeeded in establishing this pest, losses could have reached $13.4 billion.

Today's international terrorists are keenly aware of the potential of entomological weapons to inflict staggering economic losses and social turmoil. And the United States is far more vulnerable than the government would like to admit. Perhaps a demented terrorist might even consider adapting the oldest use of insects as weapons, a strategy that makes the suffering rather more immediate.

An 88-year-old woman was checking on some vacant property she owned when she noticed a door to a shed that was normally locked was open. She entered the shed to check inside and was immediately attacked by a large swarm of bees. After getting a neighbor's attention, she was able to walk about 45 m (150 ft) from where the attack began and then collapsed in the yard. . . . Upon arrival in the emergency department, the patient [was] moaning and complaining of pain but alert and cooperative. Her lungs were clear to auscultation. Swelling of the tongue was noted. The patient was given morphine sulfate 4 mg intravenously for pain. . . . The patient was transferred to the ICU in stable but guarded condition. . . . Blood pressure at the time of transfer was 190/110 mm Hg, heart rate was 105/min with sinus tachycardia and occasional premature atrial contractions, and respirations were 22/min. The emergency department staff estimated that the patient had sustained approximately 1000 bee stings.1

This case study of an Arizona woman attacked by Africanized bees does not have a happy ending. She died less than 96 hours after the attack. Killer bees were not introduced into the United States as an act of war, but there are plenty of insects capable of inflicting debilitating pain, even death, that could be weaponized. Indeed, humans have used stinging insects as weapons for thousands of years.

The oldest tactic in biological warfare was the heaving of beehives and wasp nests at an entrenched enemy. So effective were stinging creatures that, when a people couldn't find insects, they conscripted the next best thing. In the second century, the Middle Eastern stronghold of Hatra forced Rome's finest legions to hightail it back home—an apropos image for an army that had been stung into submission by scorpions dropped from the city walls.

Stinging insects can be unreliable combatants, being unable to follow orders. So the ancients developed poisoned-tipped projectiles (legend has it that Hercules invented poisoned arrows by emulating wasps). The Romans yearned to obtain a mysterious poison from India. They never found the source, but scientists believe that a beetle excreted the poison—a chemical 15 times more potent than cobra venom. But there is another way to overcome the problem of one's proximity to untrustworthy conscripts.

Bees, wasps, and hornets were catapulted for centuries in battles stretching across Europe. But few history books reveal that King Richard was both Lion-Hearted and Bee-Armed, that stinging projectiles ensured the defeat of Henry I by the Duke of Lorraine, or that the entomological predecessor of the Gatling gun was a windmill-like device that propelled straw beehives from the ends of its rapidly rotating arms. And there are even fewer accounts of how insects were used as instruments of torture.

Although the "Great Game" in which England and Russia vied for control of Central Asia in the 1800s is an important chapter in history, the chilling tale of the Bug Pit and its victims—including a pair of unfortunate British officers—is rarely told. Created by the emir of Bukhara, the torture chamber was stocked with assassin bugs that slowly ate their victims alive. But modern militaries don't use insects to inflict agony, right? Wrong. The Vietcong wired boxes of scorpions to trip wires as booby traps in the subterranean tunnels of Cu Chi. And for their part, the Americans figured out how to issue chemical commands to order one of the world's biggest and meanest species of bees to attack the enemy.

The three major forms of entomological warfare (disease vectors, agricultural pests, and direct attacks) capture much of human ingenuity in conscripting these creatures for military use. However, insects are far too diverse and humans far too clever to stay within these neat categories. Today, scientists are designing insect-machine hybrids—tiny cyborgs to infiltrate enemy positions, gather military intelligence, and assassinate key individuals. The poisons devised to control insect pests have been transformed into deadly chemical weapons, and arthropod toxins serve as molecular models for the next generation of poisons. And as we delve deeper into genetics and become able to create new life forms, the possibilities may soon be limited only by our desire to relieve—or inflict—suffering. Imagine the power that would come to a terrorist organization whose genetic engineers altered the biochemistry of a common species of mosquito in the United States so that the blood-feeding insects were capable of transmitting AIDS.

Such modern possibilities are rooted in a remarkable story of human inquiry, ingenuity, and brutality. This is the tale of one of history's most potent military alliances: the intelligence of humans and the power of insects.

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STINGING DEFEATS AND VENOMOUS VICTORIES

The insects fell into the Romans' eyes and the exposed parts of their bodies . . . digging in before they were noticed, they bit and stung the soldiers, causing severe injuries.

—Herodian, a historian of ancient Antioch, describing the defense of Hatra in 199 ce

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