established is Lyme disease, caused by a spirochete that we get from the bite of ticks carried by mice and deer. Although the first known human cases in the United States appeared only as recently as 1962, Lyme disease is already reaching epidemic proportions in many parts of our country. The future of AIDS, derived from monkey viruses and first documented in humans around 1959, is even more secure (from the virus's perspective). LETHALGIFT OF LIVESTOCK • Z O 9 The final stage of this evolution is represented by the major, long-established epidemic diseases confined to humans. These diseases must have been the evolutionary survivors of far more pathogens that tried to make the jump to us from animals—and mostly failed. What is actually going on in those stages, as an exclusive disease of animals transforms itself into an exclusive disease of humans? One transformation involves a change of intermediate vector: when a microbe relying on some arthropod vector for transmission switches to a new host, the microbe may be forced to find a new arthropod as well. For example, typhus was initially transmitted between rats by rat fleas, which sufficed for a while to transfer typhus from rats to humans. Eventually, typhus microbes discovered that human body lice offered a much more efficient method of traveling directly between humans. Now that Americans have mostlv deloused themselves, typhus has discovered a new route into us: by infecting eastern North American flying squirrels and then transferring to people whose attics harbor flying squirrels. In short, diseases represent evolution in progress, and microbes adapt by natural selection to new hosts and vectors. But compared with cows' bodies, ours offer different immune defenses, lice, feces, and chemistries. In that new environment, a microbe must evolve new ways to live and to propagate itself. In several instructive cases doctors or veterinarians have actually been able to observe microbes evolving those new ways. The best-studied case involves what happened when myxomatosis hit Australian rabbits. The myxo virus, native to a wild species of Brazilian rabbit, had been observed to cause a lethal epidemic in European domestic rabbits, which are a different species. Hence the virus was intentionally introduced to Australia in 1950 in the hopes of ridding the continent of its plague of European rabbits, foolishly introduced in the nineteenth century. In the first year, myxo produced a gratifying (to Australian farmers) 99.8 percent mortality rate in infected rabbits. Unfortunately for the farmers, the death rate then dropped in the second year to 90 percent and eventually to 25 percent, frustrating hopes of eradicating rabbits completely from Australia. The problem was that the myxo virus evolved to serve its own interests, which differed from ours as well as from those of the rabbits. The virus changed so as to kill fewer rabbits and to permit lethally infected ones to live longer before dying. As a result, a less lethal myxo virus spreads baby viruses to more rabbits thaa did the original, highly virulent 2. I O •GUNS,GERMS, AND STEEL For a similar example in humans, we have only to consider the surprising evolution of syphilis. Today, our two immediate associations to syphilis are genital sores and a very slowly developing disease, leading to the death of many untreated victims only after many years. However, when syphilis was first definitely recorded in Europe in 1495, its pustules often covered the body from the head to the knees, caused flesh to fall off people's faces, and led to death within a few months. By 1546, syphilis had evolved into the disease with the symptoms so well known to us today. Apparently, just as with myxomatosis, those syphilis spirochetes that evolved so as to keep their victims alive for longer were thereby able to transmit their spirochete offspring into more victims. The importance of lethal microbes in human history is well illustrated by Europeans' conquest and depopulation of the New World. Far more Native Americans died in bed from Eurasian germs than on the battlefield from European guns and swords. Those germs undermined Indian resistance by killing most Indians and their leaders and by sapping the survivors' morale. For instance, in 1519 Cortes landed on the coast of Mexico with 600 Spaniards, to conquer the fiercely militaristic Aztec Empire with a population of many millions. That Cortes reached the Aztec capital of Tenochtitlan, escaped with the loss of 'only' two-thirds of his force, and managed to fight his way back to the coast demonstrates both Spanish military advantages and the initial naivete of the Aztecs. But when Cortes's next onslaught came, the Aztecs were no longer naive and fought street by street with the utmost tenacity. What gave the Spaniards a decisive advantage was smallpox, which reached Mexico in 1520 with one infected slave arriving from Spanish Cuba. The resulting epidemic proceeded to kill nearly half of the Aztecs, including Emperor Cuitlahuac. Aztec survivors were demoralized by the mysterious illness that killed Indians and spared Spaniards, as if advertising the Spaniards' invincibility. By 1618, Mexico's initial population of about 20 million had plummeted to about 1.6 million. Pizarro had similarly grim luck when he landed on the coast of Peru in 1531 with 168 men to conquer the Inca Empire of millions. Fortunately for Pizarro and unfortunately for the Incas, smallpox had arrived overland around 1526, killing much of the Inca population, including both the emperor Huayna Capac and his designated successor. As we saw in Chap- LETHALGIFT OF LIVESTOCK • III ter 3 the result of the throne's being left vacant was that two other sons of Huayna Capac, Atahuallpa and Huascar, became embroiled in a civil war that Pizarro exploited to conquer the divided Incas. When we in the United States think of the most populous New World societies existing in 1492, only those of the Aztecs and the Incas tend to come to our minds. We forget that North America also supported populous Indian societies in the most logical place, the Mississippi Valley, which contains some of our best farmland today. In that case, however, conquis-tadores contributed nothing directly to the societies' destruction; Eurasian germs, spreading in advance, did everything. When Hernando de Soto became the first European conquistador to march through the southeastern United States, in 1540, he came across Indian town sites abandoned two years earlier because the inhabitants had died in epidemics. These epidemics had been transmitted from coastal Indians infected by Spaniards visiting the coast. The Spaniards' microbes spread to the interior in advance of the Spaniards themselves. De Soto was still able to see some of the densely populated Indian towns lining the lower Mississippi. After the end of his expedition, it was a long time before Europeans again reached the Mississippi Valley, but Eurasian microbes were now established in North America and kept spreading. By the time of the next appearance of Europeans on the lower Mississippi, that of French settlers in the late 1600s, almost all of those big Indian towns had vanished. Their relics are the great mound sites of the Mississippi Valley. Only recently have we come to realize that many of the mound-building societies were still largely intact when Columbus reached the New World, and that they collapsed (probably as a result of disease) between 1492 and the systematic European exploration of the Mississippi. When I was young, American schoolchildren were taught that North America had originally been occupied by only about one million Indians. That low number was useful in justifying the white conquest of what could be viewed as an almost empty continent. However, archaeological excavations, and scrutiny of descriptions left by the very first European explorers on our coasts, now suggest an initial number of around 20 million Indians. For the New World as a whole, the Indian population decline in the century or two following Columbus's arrival is estimated to have been as large as 95 percent. The main killers were Old World germs to which Indians had never een exposed, and against which they therefore had neither immune nor ill• GUNS, GERMS, AND STEEL genetic resistance. Smallpox, measles, influenza, and typhus competed for top rank among the killers. As if these had not been enough, diphtheria, malaria, mumps, pertussis, plague, tuberculosis, and yellow fever came up close behind. In countless cases, whites were actually there to
