proportionately than any human flying machine. “The U.S. Air Force,” one commentator has written, “has put them in wind tunnels to see how they do it, and despaired.” They, too, gorged on the rich air. In Carboniferous forests dragonflies grew as big as ravens. Trees and other vegetation likewise attained outsized proportions. Horsetails and tree ferns grew to heights of fifty feet, club mosses to a hundred and thirty.

The first terrestrial vertebrates-which is to say, the first land animals from which we would derive-are something of a mystery. This is partly because of a shortage of relevant fossils, but partly also because of an idiosyncratic Swede named Erik Jarvik whose odd interpretations and secretive manner held back progress on this question for almost half a century. Jarvik was part of a team of Scandinavian scholars who went to Greenland in the 1930s and 1940s looking for fossil fish. In particular they sought lobe-finned fish of the type that presumably were ancestral to us and all other walking creatures, known as tetrapods.

Most animals are tetrapods, and all living tetrapods have one thing in common: four limbs that end in a maximum of five fingers or toes. Dinosaurs, whales, birds, humans, even fish-all are tetrapods, which clearly suggests they come from a single common ancestor. The clue to this ancestor, it was assumed, would be found in the Devonian era, from about 400 million years ago. Before that time nothing walked on land. After that time lots of things did. Luckily the team found just such a creature, a three-foot-long animal called an Ichthyostega. The analysis of the fossil fell to Jarvik, who began his study in 1948 and kept at it for the next forty-eight years. Unfortunately, Jarvik refused to let anyone study his tetrapod. The world’s paleontologists had to be content with two sketchy interim papers in which Jarvik noted that the creature had five fingers in each of four limbs, confirming its ancestral importance.

Jarvik died in 1998. After his death, other paleontologists eagerly examined the specimen and found that Jarvik had severely miscounted the fingers and toes-there were actually eight on each limb-and failed to observe that the fish could not possibly have walked. The structure of the fin was such that it would have collapsed under its own weight. Needless to say, this did not do a great deal to advance our understanding of the first land animals. Today three early tetrapods are known and none has five digits. In short, we don’t know quite where we came from.

But come we did, though reaching our present state of eminence has not of course always been straightforward. Since life on land began, it has consisted of four megadynasties, as they are sometimes called. The first consisted of primitive, plodding but sometimes fairly hefty amphibians and reptiles. The best-known animal of this age was the Dimetrodon, a sail-backed creature that is commonly confused with dinosaurs (including, I note, in a picture caption in the Carl Sagan book Comet). The Dimetrodon was in fact a synapsid. So, once upon a time, were we. Synapsids were one of the four main divisions of early reptilian life, the others being anapsids, euryapsids, and diapsids. The names simply refer to the number and location of small holes to be found in the sides of their owners’ skulls. Synapsids had one hole in their lower temples; diapsids had two; euryapsids had a single hole higher up.

Over time, each of these principal groupings split into further subdivisions, of which some prospered and some faltered. Anapsids gave rise to the turtles, which for a time, perhaps a touch improbably, appeared poised to predominate as the planet’s most advanced and deadly species, before an evolutionary lurch let them settle for durability rather than dominance. The synapsids divided into four streams, only one of which survived beyond the Permian. Happily, that was the stream we belonged to, and it evolved into a family of protomammals known as therapsids. These formed Megadynasty 2.

Unfortunately for the therapsids, their cousins the diapsids were also productively evolving, in their case into dinosaurs (among other things), which gradually proved too much for the therapsids. Unable to compete head to head with these aggressive new creatures, the therapsids by and large vanished from the record. A very few, however, evolved into small, furry, burrowing beings that bided their time for a very long while as little mammals. The biggest of them grew no larger than a house cat, and most were no bigger than mice. Eventually, this would prove their salvation, but they would have to wait nearly 150 million years for Megadynasty 3, the Age of Dinosaurs, to come to an abrupt end and make room for Megadynasty 4 and our own Age of Mammals.

Each of these massive transformations, as well as many smaller ones between and since, was dependent on that paradoxically important motor of progress: extinction. It is a curious fact that on Earth species death is, in the most literal sense, a way of life. No one knows how many species of organisms have existed since life began. Thirty billion is a commonly cited figure, but the number has been put as high as 4,000 billion. Whatever the actual total, 99.99 percent of all species that have ever lived are no longer with us. “To a first approximation,” as David Raup of the University of Chicago likes to say, “all species are extinct.” For complex organisms, the average lifespan of a species is only about four million years-roughly about where we are now.

Extinction is always bad news for the victims, of course, but it appears to be a good thing for a dynamic planet. “The alternative to extinction is stagnation,” says Ian Tattersall of the American Museum of Natural History, “and stagnation is seldom a good thing in any realm.” (I should perhaps note that we are speaking here of extinction as a natural, long-term process. Extinction brought about by human carelessness is another matter altogether.)

Crises in Earth’s history are invariably associated with dramatic leaps afterward. The fall of the Ediacaran fauna was followed by the creative outburst of the Cambrian period. The Ordovician extinction of 440 million years ago cleared the oceans of a lot of immobile filter feeders and, somehow, created conditions that favored darting fish and giant aquatic reptiles. These in turn were in an ideal position to send colonists onto dry land when another blowout in the late Devonian period gave life another sound shaking. And so it has gone at scattered intervals through history. If most of these events hadn’t happened just as they did, just when they did, we almost certainly wouldn’t be here now.

Earth has seen five major extinction episodes in its time-the Ordovician, Devonian, Permian, Triassic, and Cretaceous, in that order-and many smaller ones. The Ordovician (440 million years ago) and Devonian (365 million) each wiped out about 80 to 85 percent of species. The Triassic (210 million years ago) and Cretaceous (65 million years) each wiped out 70 to 75 percent of species. But the real whopper was the Permian extinction of about 245 million years ago, which raised the curtain on the long age of the dinosaurs. In the Permian, at least 95 percent of animals known from the fossil record check out, never to return. Even about a third of insect species went-the only occasion on which they were lost en masse. It is as close as we have ever come to total obliteration.

“It was, truly, a mass extinction, a carnage of a magnitude that had never troubled the Earth before,” says Richard Fortey. The Permian event was particularly devastating to sea creatures. Trilobites vanished altogether. Clams and sea urchins nearly went. Virtually all other marine organisms were staggered. Altogether, on land and in the water, it is thought that Earth lost 52 percent of its families-that’s the level above genus and below order on the grand scale of life (the subject of the next chapter)-and perhaps as many as 96 percent of all its species. It would be a long time-as much as eighty million years by one reckoning-before species totals recovered.

Two points need to be kept in mind. First, these are all just informed guesses. Estimates for the number of animal species alive at the end of the Permian range from as low as 45,000 to as high as 240,000. If you don’t know how many species were alive, you can hardly specify with conviction the proportion that perished. Moreover, we are talking about the death of species, not individuals. For individuals the death toll could be much higher-in many cases, practically total. The species that survived to the next phase of life’s lottery almost certainly owe their existence to a few scarred and limping survivors.

In between the big kill-offs, there have also been many smaller, less well-known extinction episodes-the Hemphillian, Frasnian, Famennian, Rancholabrean, and a dozen or so others-which were not so devastating to total species numbers, but often critically hit certain populations. Grazing animals, including horses, were nearly wiped out in the Hemphillian event about five million years ago. Horses declined to a single species, which appears so sporadically in the fossil record as to suggest that for a time it teetered on the brink of oblivion. Imagine a human history without horses, without grazing animals.

In nearly every case, for both big extinctions and more modest ones, we have bewilderingly little idea of what the cause was. Even after stripping out the more crackpot notions there are still more theories for what caused the extinction events than there have been events. At least two dozen potential culprits have been identified as causes or prime contributors: global warming, global cooling, changing sea levels, oxygen depletion of the seas (a condition known as anoxia), epidemics, giant leaks of methane gas from the seafloor, meteor and comet impacts, runaway hurricanes of a type known as hypercanes, huge volcanic upwellings, catastrophic solar

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