The transformation of domestic plants and animals � from age-old selective breeding to today�s genetic engineering � has had an enormous impact on all living things on the planet.

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SIX

THE FIRST 10 MILLION YEARS

The Recovery Fauna

Sitting high atop a hill with an unbroken view to the west, Seattle�s Harborview Hospital enjoys a commanding vista of the vast city sprawling along the shores of the inland waterway known as Puget Sound. On clear days the rugged Olympic Mountains float above the far western horizon, seemingly trying to snare the setting sun with their jagged rocky tops, while to the south Mt. Rainier looms like some upstart Baskin Robbins flavor of the month. Perhaps no other public hospital in the world enjoys such a magnificent setting.
Such bucolic pleasures are largely lost on Harborview�s clientele, however. Most who come here are beyond caring about such things, for those passing

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through these doors usually do so only as a last resort. And they do not come alone. The transients, drug addicts, gunshot victims, and uninsured who make up a large percentage of Harborview�s patients often bring with them exotic new strains of microbes, organisms that are assuredly only recently evolved.
Beginning in the middle part of the twentieth century, scientists and doctors waged a campaign of eradication against bacterial illnesses, using the then newly developed antibiotic drugs. The result was a mass extinction of bacteria � a concentrated interval of death resulting in the loss of uncountable individual microbes, and for all intents and purposes, the extinction of whole species. Smallpox, rabies, typhoid, rubella, cholera: the ancient microbial scourges of humankind were wiped out. The bacteria causing these ancient plagues were faced with two alternatives: evolve or die. Most died. But a few evolved forms resistant to antibiotic drugs. Fifty years after their invention, these �miracle� drugs have unleashed a diversity of new drug-resistant species that would never have evolved but for the influence of humanity. These are surely but the beginning of a host of new microbial species.
And so too with the biosphere, except that the �antibiotic� is us. As a result of our antibiotic influence, and the current pulse of extinction it has engendered, many currently living species will die. Some, however, will survive and thrive, becoming the rootstock of a new biota. Some have already done so, for one of the precepts of this book is that significant portions of the �recovery fauna� that follows any mass extinction are already with us, and dominating terrestrial habitats, in the form of domesticated plants and animals. Evolution obviously continues, but much of it is now �directed� for human purposes, or occurs as a by-product of human activities.

Charles Darwin began his On the Origin of Species with a chapter on domestication. Before introducing any other data or argument, he pointed to the many varieties of domesticated animals and plants as one of the clearest proofs of the existence of organic evolution � in this case, the evolution of new types of animals and plants bred to serve as food or as companions to humanity.
As with most of his conclusions, Darwin was right about this point. But we can take it one step further. Domesticated animals and plants are the dominant members of what can be called the �recovery fauna� accruing, directly or indirectly,

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from the extinction of megamammals (and that extinction�s impetus toward the development of agriculture). That many of these animals are taking the functional place of extinct or endangered megamammal species is no accident. The cows, pigs, sheep, horses, and other familiar domesticated animals now covering the world�s grasslands rapidly replaced the many species of extinct or endangered large wild grazers. The stimulus to that evolutionary changeover has, of course, been the stern hand of humanity.

Characteristics of Domestication

Although most animal species can be �tamed�, or to some extent habituated to the presence of humans if raised by them from a young age, domestication goes far beyond this simple behavior modification. Domesticating a species requires not only a concerted effort over extended periods of time, but certain pre-existing features of the species in question. In the past this effort was made only for reasons such as enhanced food yield, transportation, or protection from predators. Domesticated animals are the evolutionary results of human-induced �unnatural selection�.
Very few large mammals have been domesticated. Biologist Jared Diamond of UCLA showed that of the 150 species of terrestrial, noncarnivorous mammals larger than about 30 kilograms, only 14 have been domesticated. All but one of these originated in the Eurasian region; the only New World exception to this rule is the llama. All of the domesticated species are derived from wild species with similar characteristics: all seem to show rapid growth to maturity, an ability to breed in captivity, little tendency to panic when startled, an amenable and tractable disposition, and a social structure and hierarchy that permits domestication. All of these characteristics were further selected for by a brutal form of �natural selection�: those individuals that showed the favored characters were allowed to breed; those that did not were killed.
Interestingly, neurologist Terry Deacon has noted a further characteristic: all domesticated animals appear to have undergone a loss of intelligence compared with their wild ancestors. This observation makes one wonder whether humans, compared with their ancestors, have also been �domesticated�, and undergone a similar reduction in intelligence.
The first domesticated species may well have been the dog. All modern dog species seem to be derived from the Asiatic wolf. Although the first anatomically

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Foremost among the survivors of the next millennium will be those species that humans have had a hand in developing: domesticated plants and animals.

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modern dogs may date back as far as 12,000 years ago, bones of canids recognizable as �dogs� first appear around human campsites between 7000 and 6000 B.C. Even though wolves can still interbreed with dogs, implying that true biological speciation has not occurred, behavioral differences make such interbreeding rare; thus the modern dog varieties as we known them are functionally distinct from wolves. The domestication of the dog also led to distinct anatomical changes. Compared with wolves, dogs have smaller skulls, shorter jaws, smaller teeth, and pronounced variation in coat color. Dogs also appear to be less intelligent than wolves. Most dog varieties now recognizable were produced in the eighteenth and nineteenth centuries; prior to that, dogs generally were used for hunting (hounds) or tending flocks (sheepdogs).
The bones of domesticated livestock first appear in the fossil record slightly later than those of dogs. Sheep and goats came first; the earliest evidence for their domestication, dated at around 8000 B.C., comes from various sites in southwestern Asia (the areas now composed of Israel, Iran, Jordan, and Syria). Cattle were derived from an entirely extinct species of wild cowlike creatures. Domesticated pigs also date back to about 8000 B.C. Four thousand years later, the domesticated horse was developed from wild horses in Eastern Europe. (The ancestor of the domestic horse, called Przewalski�s horse, still exists, in small numbers, in reserves in Poland.) Donkeys, water buffaloes, and llamas were domesticated at about the same time, while chickens and camels were not brought into the menagerie until about 2500 B.C. A wide variety of smaller �pets� have been domesticated as well: house cats, guinea pigs, rabbits, white rats, hamsters, and various birds. All are the result of human effort.
In almost every case, the transformation of a wild species into a domestic species involves substantial physical and behavioral modification. It has long been thought that this process occurred in stages, beginning with �taming� and progressing to gradual genetic transformation. How such great genetic change was forced in such a short period of time has always been a puzzle. New research by geneticists may have provided an answer. There appears to be a master gene controlling a complex of genes that in turn affect tameness, reaction to stress, coat color, facial morphology, and social interactions. By changing a single gene rather than an entire complex, domestication could proceed relatively quickly.

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Domesticated Crop Species

While it may be argued that the near coincidence of the beginning of agriculture and the end of the Age of Megamammals is just that � coincidence � a strong argument also can be made for cause and effect. The extinction of many of the larger animals upon which humans depended for food, coming as it did with a major climate change (affecting plant and small animal resources also used for food) may not have been mere chance.
While cereals such as wild wheat and barley were harvested as much as 12,000 years ago, it seems that the first domestication of plants took place about 10,000 years ago, at the time when the last mammoths, mastodons, and many other larger animal species were dying out in North America and had just disappeared in Europe and Asia. This was the time when food-gathering peoples began to collect the seeds of wild plants and replant them in the ground. The domestication process appears to have involved the natural hybridization of several wild species, followed by selection by humans for desired characteristics. Thus �domestication� of plants, like that of animals, involved the genetic modification of the wild species through a very rough form of natural selection: those plants with usable traits were kept; those without were killed. Since the trend in plant modification has been toward an increase in the size of the edible or usable parts, most plant species have lost the ability to disperse widely, and protective mechanisms such as thorns have generally been lost as well.
The number of domesticated plant species is relatively quite small. There are more than two hundred thousand species of angiosperms, or flowering plants, yet only ten of these provide the vast majority of human food. Among these ten are grasses and cereals such as wheat, rice, and maize, which are all characterized by seeds rich in starch and protein. Cereals are planted on 70% of the world�s cropland and produce about 50% of the calories consumed by humanity. Other plants in the top ten include sugarcane, yams, potatoes, bananas, soybeans, and manioc. Worldwide, about three thousand species of plants are used as human food, but only about two hundred of these have become domesticated.

The Transgenic Revolution: Building Weeds

The genetic engineering our ancestors used to introduce new characters into their agricultural crops and domestic animals was crude but effective: save the favored

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varieties and let them breed; kill off the others. But in the twentieth century a new type of genetic engineering appeared � one that alters the genomes themselves. This new way of introducing novelty is sweeping the agricultural regions of the Earth, and will surely have unintended consequences. It may be that the transgenic revolution will bring novelty into the biotic world in ways almost unimaginable � and not all of them desirable. It appears, for example, to be on the verge of creating �superweeds�.
Modern genetic technology allows the transfer of genetic material from one species to another. This new genetic information is permanently integrated into the genome of the second species, conferring new traits upon it. For all intents and purposes, a new type of organism is let loose into the biosphere each time this is done. The organism thus transformed is called a transgenic plant, animal, or microbe. These transgenic creatures have not arisen through the natural processes of evolution, but they are among the most portentous developments for the future of evolution on this planet.
Transgenic organisms are possible because of the existence of certain genes capable of �jumping� from one chromosome to another. The first discovery of jumping genes was made by American geneticist Barbara McClintock in the 1940s. McClintock was studying the genetics of maize (corn), and observed that certain genes, such as those responsible for seed color, were capable of moving from one chromosome to another. The significance of this discovery was largely overlooked until the 1970s, when it was independently rediscovered by other researchers examining the ways in which certain bacteria develop resistance to antibiotics. The genes, or sections of DNA, coding for these specific characters in bacteria do not actually �jump�; instead, they produce copies of themselves, which are inserted at other points either on chromosomes or in genetic code-carrying organelles called plasmids.
The discovery of these jumping genes, technically called transposons, unleashed a torrent of research in the 1980s and into the 1990s. These peculiar strings of DNA have the ability to repeatedly cut and paste themselves into different parts of an organism�s genetic code. What made them famous � and may eventually make them infamous � is that the transposons of one organism can be used to paste new genetic information into the DNA of entirely unrelated organisms.
Much of the research using transposons was conducted on fruit flies. The fruit fly Drosophila is one of the stalwarts of experimental genetics, since it breeds

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quickly and its genetic code is well known. In the early 1980s Gerald Rubin and Alan Spradling of the Carnegie Institute discovered a transposon in Drosophila that could be used to incorporate new genetic information into these flies. They succeeded in changing the genetic code of the transposon and re-inserting it into the fly. With this operation, they had succeeded in creating a fly with a new genetic code that could be passed on to a succeeding generation. They had created a transgenic species � one entirely new to the world, a species with a genetic code created not by nature, but by science.
These early experiments altered very little in the new fly. The majority of its genetic code was the same as that of the unaltered species. But certain characters, such as color or eye type, could be changed. Further work showed that certain fruit fly transposons were not only useful in changing the genetic code in fruit flies, but could be placed into entirely unrelated species. A method had been developed that allowed true genetic engineering of insects.
The goals of genetic alteration of insects are laudable. Insects create havoc in human society in two ways: they serve as vectors of diseases (e.g., malaria, yellow fever, and some types of sleeping sickness), and they consume a large proportion of human crops. Genetic engineering is attempting to mitigate both of these problems. Nevertheless, the results of this work have been slow. In certain disease-spreading mosquitoes, geneticists have so far been able to change eye color, but they have yet to alter the structures involved in spreading disease-causing microbes. To further aid this process, geneticists have infected the target insects with viruses that act like transposons. The virus, once in the body of the insect host, can alter the way a disease is passed on. Some crop pests, such as the Mediterranean fruit fly and the screwworm, have been successfully targeted using transgenic or other genetic techniques (e.g., producing sterile members of a species that spread among the viable members of the population).
While transgenic techniques are just coming into use in controlling insect pests, such tools are already used widely in crop plants. Genetic engineering has succeeded in adding new genes to the DNA of various crops, whereas conventional breeding only adds variants to an already existing genetic complex. So, while domestication has enhanced the valued characteristics of many plant species, transgenic research has added entirely new characteristics, such as greater tolerance of heat and drought, greater resistance to insect predation and disease, and greater yield.

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The genetic engineering techniques used in agriculture are widespread and sophisticated. Genetic engineers can move genes from virtually any biological source into crop species. Genes have been added to engineered crops from organisms as diverse as chickens, hamsters, fireflies, and fish, as well as from a slew of plant and microbial species. The new transgenic plants are genuinely novel organisms, some of them containing the genetic codes of plants, animals, and microbes in a single species.
The addition of new genes to various plant species has yielded spectacular dividends in terms of crop yield. But this new technology also poses substantial risks and clearly will affect the future of evolution on this planet. The creation of new plant types could affect the biosphere in various ways. The most important of these is the possibility that newly inserted genes will jump to other, nonengineered species (such as weeds) or move out of the agricultural fields altogether into wild native plant populations. It is this potential intermixing of new genes with those of already established plant species beyond the boundaries envisioned or designed by agricultural scientists that could have the most interesting � and potentially biosphere-altering � effects. Under rare circumstances, if new traits of transgenic species escape into the wild, weeds adaptively superior to native plants could be created. Since most of the traits being transferred into transgenic crops, such as hardiness, resistance to pests, and growth rate, increase their fitness relative to the original species, there is great potential for transgenics to become, or to help produce, new weedy species.
There are several avenues by which the genes of transgenic crops can become established in the wild. First and simplest, the transgenic crop itself can escape and become a weed. Second, the transgenic crop can release pollen into the wild, which can be incorporated into a wild relative of the original transgenic host. The incorporation of the new genes into the wild plant creates a new transgenic weed.
Weeds have many definitions, which are often colored by human values. In agriculture, they are plants that occur in the wrong place at the wrong time (some plants are weeds in certain situations and favored crops in others � lawn grass, for example). A more human-specific definition of a weed is any plant that is objectionable to or interferes with the activities or welfare of humans. Nevertheless, weedy species have a number of characteristics:

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From this list, we can see that the characters of weeds are also highly desirable in crop plants. One goal of transgenic technology, therefore, has been to confer characteristics of weeds on crop species. Transgenes spliced into crops may alter traits such as seed germination ability, seed dormancy, or tolerance of either biotic or abiotic factors such as pests, drought, heat, or disease, creating a more persistent or resistant species in the process. Such new traits may enhance the new crop�s ability to invade other habitats. Genes affecting seedling growth rates, root growth rates, and drought tolerance are currently being developed.
Jane Rissler and Margaret Mellon of the Union of Concerned Scientists have studied the ecological risks of transgenic crops in thorough detail. One of their most important concerns relates to the transformation of nonweedy crop species into weeds through genetic engineering. They note that a widely held notion is that changing a non-weed into a weed involves the conversion of many genes, not just the two or three currently used in transgenic crops. Changing corn from a crop to a weed, for example, would involve a number of genetic changes, since corn is one of the most human-dependent (and thus intolerant) plant species on Earth. Other crops, however, already possess many weedy traits, and thus one to three new traits could indeed create a new weedy species. Examples include alfalfa and other forages, barley, lettuce, rice, blackberries, radishes, raspberries, and sunflowers.
Transgenic species may produce secondary effects as well. The invasion of transgenic plants into new habitats affects not only the invaded plant populations, but the entire ecosystem, including the suite of animals living within that ecosystem. Perhaps more dangerous than the conversion of crop plants into weeds is the escape and transference of new genes into already existing weeds, making them �superweeds�. The transfer of disease resistance or pest resistance to established

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weedy species may change a familiar weed into an even more formidable pest. Weeds that have developed resistance to herbicides because of escaped or transferred transgenes are already appearing in some parts of the world.
Agribusiness, which counts transgenic technology as the jewel in its technological crown, is in the business of feeding the world�s humans � and making profits in doing so, of course. One of the great fears of those producing transgenic crops is that farmers will simply take the seeds from the first crop and use them ever after, and will not need to buy new seeds from the corporation that produced them in the first place. Like the software industry, which fears the copying of its products above all else, the major biotechnology companies dealing in transgenic crops have been searching for some way to stem the illegal use of their products after the first purchase. The solution is something known as the �terminator gene�.
The first terminator gene was produced by the large American biotechnology firm Monsanto, and was engineered to protect Monsanto�s patent rights on several types of transgenic crops. It is a genetic modification that prevents seeds from germinating after the season in which they were sold. In the works are genes that will allow but a single crop and not produce future seeds � a bit like the seedless watermelon, but more efficient.
The great fear is that such terminator genes will jump to unmodified varieties of crop plants. If the terminator gene in a tomato plant jumped to other varieties of tomatoes, there a is real potential of plants never producing the crop they were intended to produce.

Does Evolution Have a Future?

Our species has learned how to circumvent the normal rules of evolutionary change: we have learned how to build new species. Have we also achieved the ability to alter those rules? Norman Myers of Oxford asks this question in his prescient and disturbing 1998 paper, �The Biodiversity Crisis and the Future of Evolution�. Myers makes a subtle but important point: humans pose �pronounced threats to certain basic processes of evolution such as natural selection, speciation, and origination�. Myers has cried wolf before, but the wolf was always there, dining happily on flocks of the world�s species. Is he being alarmist in this case? Although many have warned of a biodiversity crisis, Myers alone warns of an evolution crisis. He bases his conclusion on two perceptions: first, that we have entered a new phase of mass extinction, and second, that the normal recovery

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period from mass extinction will not pertain to this one; in fact, the recovery will be considerably delayed.
Myers cites three aspects of this particular mass extinction that will affect its evolutionary outcome (and which make it different from any mass extinction of the past):

The current mass extinction may be unique not only in what it kills, but in how its recovery proceeds. In past times the tropical regions of the world have served as storehouses for recovery. Because they have always held the greatest diversity of species on the planet, they have long served as �evolutionary powerhouses� � areas that seem to spawn new species and new types of species at a higher rate than other parts of the world. Paleontologist David Jablonski of the University of Chicago has shown that innovation can be related to geography. Innovation in evolution is the appearance of evolutionary novelty, and the tropical regions seem to be home to more innovation than other regions. Yet the tropics are now the sites of the highest densities of humans and, the greatest human population increases. This pattern may curtail the evolution not only of new species, but also of new types of species.
The current crisis in biodiversity may also substantially reduce the number of new species evolving a large body size. Megamammals need very large habitat areas to survive; it may also be true that they need equally large areas to speciate. With the reduction of wild habitat, and especially free rangeland, virtually everywhere on Earth, there may be no way for large mammals and other vertebrates to produce new species. Therefore, a consequence of human population growth and habitat disturbance may be not only the extinction of large mammals, reptiles, and birds, but the inability of new large species to take their place, simply because the mechanism of speciation for large body sizes has been derailed by environmental fragmentation.

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Implications for Conservation Planners

A large and vibrant community of conservationists, scientists, politicians, and laypeople are actively engaged in intensive efforts to preserve biodiversity. One of the most important such efforts is habitat preservation. Yet even the most Herculean of efforts will save only patches of habitat in a sea of agricultural fields and spreading human landscapes. As long as humanity rules, it is doubtful that hundreds of thousands of miles of unfenced, unimpeded native habitat will be available to replace the species already lost since the end of the Ice Age. This fact has led Norman Myers to pose the following questions:

This last recommendation is heresy by the rules of modern conservation. It has long been argued that endemic centers � those regions that contain species found

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nowhere else � are among the most important places to save. But Myers�s point is that endemic centers exist because they have not produced large numbers of successful species. Endemic centers are often living museums of ancient species that do not have much potential for future evolution.

The Weeds of Humanity

The vast human enterprise has created a new recovery fauna, and will continue to provide opportunities for new types of species that possess weedy qualities and have the ability to exploit the new anthropogenic world. Chief among these will be those species best preadapted for dealing with humanity: flies, rats, raccoons, house cats, coyotes, fleas, ticks, crows, pigeons, starlings, English sparrows, and intestinal parasites, among others. These and our domesticated vassals will dominate the recovery fauna. Among plants, the equivalents will be the weeds. According to many seers, this group of new flora and fauna will be with us for an extended period of time � a time span measured in the millions of years. And if humanity continues to exist and thrive (as I believe it will), this recovery biota may dominate any new age of organisms on Earth.
A sense of how long the recovery fauna may last was estimated in a disturbing paper published in the prestigious journal Nature in the spring of 2000. The authors, James Kirchner and Ann Weil, posed a question: how quickly does biodiversity rebound after a mass extinction? How long will the world exist at a very low biodiversity? The answer, it turned out, was far longer than anyone had heretofore estimated. By analyzing the fossil record of all recoverable organisms (compiled by the late Jack Sepkoski of the University of Chicago), Kirchner and Weil found that fully 10 million years passed by, on average, before the biodiversity of the world recovered to its pre-extinction values. Even more surprising than this long lag period between extinction and full recovery was their finding that it occurred whether the extinction was small or large. We paleontologists had assumed that the time to recovery would somehow correlate with the magnitude of the extinction � that after a small extinction, the biosphere would recover quickly, and that it was only after the greatest of the mass extinctions that a long recovery period was necessary. But to the surprise of us all, Kirchner and Weil found this not to be the case � 10 million years was necessary even after the smaller extinctions. They concluded their paper with the following passage:

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It appears that our return to a new biota will take a long time after the mass extinction is finished. And what might that new fauna and flora look like? Some predictions can be made � and such predictions are the subject of the next chapter.

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CONTENTS