The Rise and Fall of the Dinosaurs The Untold Story of a Lost World (Steve Brusatte)

The Dawn of the Dinosaurs


Chapter Title art by Todd Marshall

“BINGO,” MY FRIEND GRZEGORZ NIEDZWIEDZKI shouted, pointing at a knife-thin separa­tion between a slim strip of mudstone and a thicker layer of coarser rock right above it. The quarry we were exploring, near the tiny Polish village of Zachelmie, was once a source of sought-after limestone but had long been abandoned. The surrounding landscape was littered with decaying smoke­stacks and other remnants of central Poland’s industrial past. The maps de­ceitfully told us we were in the Holy Cross Mountains, a sad patch of hills once grand but now nearly flattened by hundreds of millions of years of erosion. The sky was gray, the mosquitoes were biting, heat was bouncing off the quarry floor, and the only other people we saw were a couple of wayward hikers who must have made a tragically wrong turn.

“This is the extinction,” Grzegorz said, a big smile creasing the unshaven stubble of many days of fieldwork. “Many footprints of big reptiles and mammal cousins below, but then they disappear. And above, we see nothing for awhile, and then dinosaurs.”

We may have been peering at some rocks in an overgrown quarry, but what we were really looking at was a revolution. Rocks record history; they tell stories of deep ancient pasts long before humans walked the Earth. And the narrative in front of us, written in stone, was a shocker. That switch in the rocks, detectable perhaps only to the overtrained eyes of a scientist, documents one of the most dra­matic moments in Earth history. A brief instance when the world changed, a turning point that hap­pened some 252 million years ago, before us, before woolly mammoths, before the dinosaurs, but one that still reverberates today. If things had unfolded a little differently back then, who knows what the modern world would be like? It’s like wondering what might have happened if the archduke was never shot.

IF WE’D BEEN standing in this same spot 252 million years ago, during a slice of time geologists call the Permian Period, our surroundings would have been barely recognizable. No ruined factories or other signs of people. No birds in the sky or mice scurrying at our feet, no flowery shrubs to scratch us up or mos­quitoes to feed on our cuts. All of those things would evolve later. We still would have been sweating, though, because it was hot and unbearably humid, proba­bly more insufferable than Miami in the middle of the summer. Raging rivers would’ve been draining the Holy Cross Mountains, which were actually proper mountains back then, with sharp snowy peaks jutting tens of thousands of feet into the clouds. The rivers wound their way through vast forests of conifer trees —early relatives of today’s pines and junipers—emptying into a big basin flank­ing the hills, dotted with lakes that swelled in the rainy season but dried out when the monsoons ended.

These lakes were the lifeblood of the local ecosystem, watering holes that provided an oasis from the harsh heat and wind. All sorts of animals flocked to them, but they weren’t animals we would know. There were slimy salamanders bigger than dogs, loitering near the water’s edge and occasion­ally snapping at a passing fish. Stocky beasts called pareiasaurs waddled around on all fours, their knobby skin, front-heavy build, and general brutish appearance making them seem like a mad reptil­ian offensive lineman. Fat little things called dicynodonts rummaged around in the muck like pigs, using their sharp tusks to pry up tasty roots. Lording over it all were the gorgonopsians, bear-size mon­sters who reigned at the top of the food chain, slicing into pareiasaur guts and dicynodont flesh with their saberlike canines. This cast of oddballs ruled the world right before the dinosaurs.

Then, deep inside, the Earth began to rumble. You wouldn’t have been able to feel it on the sur­face, at least when it kicked off, right around 252 million years ago. It was happening fifty, maybe even a hundred, miles underground, in the mantle, the middle layer of the crust-mantle-core sandwich of Earth’s structure. The mantle is solid rock that is so hot and under such intense pressure that, over long stretches of geological time, it can flow like extra-viscous Silly Putty. In fact, the mantle has cur­rents just like a river. These currents are what drive the conveyor-belt system of plate tectonics, the forces that break the thin outer crust into plates that move relative to each other over time. We wouldn’t have mountains or oceans or a habitable surface without the mantle currents. However, every once in a while, one of the currents goes rogue. Hot plumes of liquid rock break free and start snaking their way upward to the surface, eventually bursting out through volcanoes. These are called hot spots. They’re rare, but Yellowstone is an example of an active one today. The constant supply of heat from the deep Earth is what powers Old Faithful and the other geysers.

This same thing was happening at the end of the Permian Period, but on a continent-wide scale. A massive hot spot began to form under Siberia. The streams of liquid rock rushed through the mantle into the crust and flooded out from volcanoes. These weren’t ordinary volcanoes like the ones we’re most used to, the cone-shaped mounds that sit dormant for decades and then occasionally explode with a bunch of ash and lava, like Mount Saint Helens or Pinatubo. They wouldn’t have erupted with the vigor of those vinegar-and-baking-soda contraptions so many of us made as science fair experi­ments. No, these volcanoes were nothing more than big cracks in the ground, often miles long, that continuously belched out lava, year after year, decade after decade, century after century. The erup­tions at the end of the Permian lasted for a few hundred thousand years, perhaps even a few million. There were a few bigger eruptive bursts and quieter periods of slower flow. All in all, they expelled enough lava to drown several million square miles of northern and central Asia. Even today, more than a quarter billion years later, the black basalt rocks that hardened out of this lava cover nearly a million square miles of Siberia, about the same land area as Western Europe.

Imagine a continent scorched with lava. It’s the apocalyptic disaster of a bad B movie. Suffice it to say, all of the pareiasaurs, dicynodonts, and gorgonopsians living anywhere near the Siberian area code were finished. But it was worse than that. When volcanoes erupt, they don’t expel only lava, but also heat, dust, and noxious gases. Unlike lava, these can affect the entire planet. At the end of the Permian, these were the real agents of doom, and they started a cascade of destruction that would last for millions of years and irrevocably change the world in the process.

Dust shot into the atmosphere, contaminating the high-altitude air currents and spreading around the world, blocking out the sun and preventing plants from photosynthesizing. The once lush conifer forests died out; then the pareiasaurs and dicynodonts had no plants to eat, and then the gorgonop- sians had no meat. Food chains started to collapse. Some of the dust fell back through the atmos­phere and combined with water droplets to form acid rain, which exacerbated the worsening situation on the ground. As more plants died, the landscape became barren and unstable, leading to massive erosion as mudslides wiped out entire tracts of rotting forest. This is why the fine mudstones in the Zachelmie quarry, a rock type indicative of calm and peaceful environments, suddenly gave way to the coarser boulder-strewn rocks so characteristic of fast-moving currents and corrosive storms. Wildfires raged across the scarred land, making it even more difficult for plants and animals to survive.

But those were just the short-term effects, the things that happened within the days, weeks, and months after a particularly large burst of lava spilled through the Siberian fissures. The longer-term ef­fects were even more deadly. Stifling clouds of carbon dioxide were released with the lava. As we know all too well today, carbon dioxide is a potent greenhouse gas, which absorbs radiation in the at­mosphere and beams it back down to the surface, warming up the Earth. The CO2 spewed out by the Siberian eruptions didn’t raise the thermostat by just a few degrees; it caused a runaway greenhouse effect that boiled the planet. But there were other consequences as well. Although a lot of the carbon dioxide went into the atmosphere, much of it also dissolved into the ocean. This causes a chain of chemical reactions that makes the ocean water more acidic, a bad thing, particularly for those sea creatures with easily dissolvable shells. It’s why we don’t bathe in vinegar. This chain reaction also draws much of the oxygen out of the oceans, another serious problem for anything living in or around water.

Descriptions of the doom and gloom could go on for pages, but the point is, the end of the Per­mian was a very bad time to be alive. It was the biggest episode of mass death in the history of our planet. Somewhere around 90 percent of all species disappeared. Paleontologists have a special term for an event like this, when huge numbers of plants and animals die out all around the world in a short time: a mass extinction. There have been five particu larly severe mass extinctions over the past 500 million years. The one 66 million years ago at the end of the Cretaceous period, which wiped out the dinosaurs, is surely the most famous. We’ll get to that one later. As horrible as the end-Cretaceous extinction was, it had nothing on the one at the end of the Permian. That moment of time 252 million years ago, chronicled in the swift change from mudstone to pebbly rock in the Polish quarry, was the closest that life ever came to being completely obliterated.

Then things got better. They always do. Life is resilient, and some species are always able to make it through even the worst catastrophes. The volcanoes erupted for a few million years, and then they stopped as the hot spot lost steam. No longer blighted by lava, dust, and carbon dioxide, ecosystems were gradually able to stabilize. Plants began to grow again, and they diversified. They provided new food for herbivores, which provided meat for carnivores. Food webs reestablished themselves. It took at least five million years for this recovery to unfold, and when it did, things were better but now very different. The previously dominant gorgonopsians, pareiasaurs, and their kin were never to stalk the lakesides of Poland or anywhere else while the plucky survivors had the whole Earth to themselves. A largely empty world, an uncolonized frontier. The Permian had transitioned into the next interval of ge­ological time, the Triassic, and things would never be the same. Dinosaurs were about to make their entrance.

AS A YOUNG paleontologist, I yearned to understand exactly how the world changed as a result of the end-Permian extinction. What died and what sur­vived, and why? How quickly did ecosystems recover? What new types of never- before-imagined creatures emerged from the post-apocalyptic blackness? What aspects of our modern world were first forged in the Permian lavas?

There’s only one way to start answering these questions. You need to go out and find fossils. If a murder has been committed, a detective begins by studying the body and the crime scene, looking for fingerprints, hair, clothing fibers, or other clues that might tell the story of what unfolded, and lead to the culprit. For paleontologists, our clues are fossils. They are the currency of our field, the only records of how long-extinct organisms lived and evolved.

Fossils are any sign of ancient life, and they come in many forms. The most familiar are bones, teeth, and shells—the hard parts that form the skeleton of an animal. After being buried in sand or mud, these hard bits are gradually replaced by minerals and turned to rock, leaving a fossil. Some­times soft things like leaves and bacteria can fossilize as well, often by making impressions in the rock. The same is sometimes true of the soft parts of animals, like skin, feathers, or even muscles and internal organs. But to end up with these as fossils, we need to be very lucky: the animal needs to be buried so quickly that these fragile tissues don’t have time to decay or get eaten by predators.

Everything I describe above is what we call a body fossil, an actual part of a plant or animal that turns into stone. But there is another type: a trace fossil, which records the presence or behavior of an organism or preserves something that an organism produced. The best example is a footprint; others are burrows, bite marks, coprolites (fossilized dung), and eggs and nests. These can be particularly valuable, because they can tell us how extinct animals interacted with each other and their environ­ment—how they moved, what they ate, where they lived, and how they reproduced.

The fossils that I’m particularly interested in belong to dinosaurs and the animals that came imme­diately before them. Dinosaurs lived during three periods of geological history: the Triassic, Jurassic, and Cretaceous (which collectively form the Mesozoic Era). The Permian Period—when that weird and wonderful cast of creatures was frolicking alongside the Polish lakes—came right before the Triassic. We often think of the dinosaurs as ancient, but in fact, they’re relative newcomers in the history of life.

The Earth formed about 4.5 billion years ago, and the first microscopic bacteria evolved a few hun­dred million years later. For some 2 billion years, it was a bacterial world. There were no plants or ani­mals, nothing that could easily be seen by the naked eye, had we been around. Then, some time around 1.8 billion years ago, these simple cells developed the ability to group together into larger, more complex organisms. A global ice age—which covered nearly the entire planet in glaciers, down to the tropics—came and went, and in its aftermath the first animals got their start. They were simple at first—soft sacs of goo like sponges and jellyfish, until they invented shells and skeletons. Around 540 million years ago, during the Cambrian Period, these skeletonized forms exploded in diversity, became extremely abundant, started eating one another, and began forming complex ecosystems in the oceans. Some of these animals formed a skeleton made of bones—these were the first vertebrates, and they looked like flimsy little minnows. But they, too, continued to diversify and eventually some of them turned their fins into arms, grew fingers and toes, and emerged onto the land, about 390 million years ago. These were the first tetrapods, and their descendants include all vertebrates that live on land today: frogs and salamanders, crocodiles and snakes, and then later, dinosaurs and us.

We know this story because of fossils—thousands of skeletons and teeth and footprints and eggs found all over the world by generations of paleontologists. We’re obsessed with finding fossils and no­torious for going to great (and sometimes stupid) lengths to discover new ones. It could be a lime­stone pit in Poland or maybe a bluff behind a Walmart, a dump pile of boulders at a construction site, or the rocky walls of a ripe landfill. If there are fossils to be found, then at least some swashbuckling (or stupid) paleontologist will brave whatever heat, cold, rain, snow, humidity, dust, wind, bug, stench, or war zone stands in the way.

That’s why I started going to Poland. I first visited in the summer of 2008, a twenty-four-year-old in between finishing my master’s and starting my PhD; I went to study some intriguing new reptile fossils that had been found a few years earlier in Silesia, the sliver of southwestern Poland that for years was fought over by Poles, Germans, and Czechs. The fossils were kept in a museum in Warsaw, treasures of the Polish state. I remember the buzz as I approached the capital’s central station on a delayed train from Berlin, night shadows covering the hideous Stalin-era architecture of a city rebuilt from ruins after the war.

As I stepped off the train, I scanned the crowd. Somebody was supposed to be there holding a sign with my name. I arranged my visit through a series of formal e-mails with a very senior Polish profes­sor, who badgered one of his graduate students into meeting me at the station and guiding me to the small guestroom where I would stay at the Polish Institute of Paleobiology, just a few stories above where the fossils were kept. I had no idea whom I was looking for, and because the train had been more than an hour late, I figured the student had escaped back to the lab, leaving me on my own to navigate a foreign city in the twilight, with the few words of Polish on the glossary page of my guide­book.

Just as I was starting to panic, I saw a sheet of white paper flapping in the wind, my name hastily scrawled across it. The man holding it was young, with a close-cropped military hairstyle, his hairline just starting to recede like mine. His eyes were dark, and he was squinting. A thin veneer of stubble covered his face, and he seemed to be a little darker than most of the Poles I knew. Tanned, almost. There was something vaguely sinister about him, but that changed in an instant when he recognized me coming toward him. He broke into a huge smile, grabbed my bag, and gripped my hand firmly. “Welcome to Poland. My name is Grzegorz. How about some dinner?”

We were both tired, I from the long train journey, Grzegorz from working the whole day describing a new batch of fossil bones that he and his crew of undergraduate assistants had just found in south­eastern Poland a few weeks before, hence the field tan he was sporting. But we ended up knocking back several beers and talking for hours about fossils. This guy had the same raw enthusiasm for di­nosaurs that I had, and he was full of iconoclastic ideas about what happened after the end-Permian extinction.

Grzegorz and I became fast friends. For the rest of that week, we studied Polish fossils together, and then during the following four summers, I came back to Poland to do fieldwork with Grzegorz, often joined by the third musketeer in our band, the young British paleontologist Richard Butler. Dur­ing that time we found a lot of fossils and came up with some new ideas about how dinosaurs got their evolutionary start in those heady days after the end-Permian extinction. Over the course of those years, I saw Grzegorz transition from an eager, but still somewhat meek, graduate student into one of Poland’s leading paleontologists. A few years before turning thirty, he discovered, in a different corner of the Zachelmie quarry, a trackway left by one of those first fishy creatures to walk out of the water and onto land, some 390 million years ago. His discovery was published on the cover of Nature, one of the world’s leading scientific journals. He was invited to a special audience with Poland’s prime minister and gave a TED talk. His steely face—not his fossil discoveries, him—graced the cover of the Polish version of National Geographic.

He had become something of a scientific celebrity, but more than anything else Grzegorz enjoyed heading out into nature and looking for fossils. He called himself a “field animal,” explaining that he loved camping and hacking through brush much more than the genteel ways of Warsaw. He couldn’t help it. He grew up around Kielce, the main city of the Holy Cross Mountains region, and started col­lecting fossils as a child. He developed a particular talent for finding a type that many paleontologists ignore: trace fossils. Footprints, hand impressions, tail drags: the marks dinosaurs and other animals left when they moved across mud or sand, going about their daily business of hunting, hiding, mating, socializing, feeding, and loitering. He was absolutely enamored of tracks. An animal has only one skeleton, but it can leave millions of footprints, he would often remind me. Like an intelligence opera­tive, he knew all the best places to find them. This was his backyard, after all. It was quite the back­yard to grow up in, too, because it turned out that those animal-infested seasonal lakes that covered the area during the Permian and Triassic were perfect environments for preserving tracks.

For four summers we indulged Grzegorz’s love of tracks. Richard and I tagged along as he led us to many of his secret sites, which were mostly abandoned quarries, bits of rock poking out of streams, and rubbish piles along the ditches of the many new roads that were being built in the area, where workmen would dump the slabs of stone they cut through when laying asphalt. We found a lot. Or rather, Grzegorz did. Both Richard and I developed an eye for the often small hand- and footprints left by lizards, amphibians, and early dinosaur and crocodile relatives, but we could never compete with the master.

The thousands of tracks that Grzegorz found over his two decades of collecting, plus the pittance of new ones that Richard and I stumbled upon, ended up telling quite a story. There were many types of tracks, belonging to a whole slew of different creatures. And they didn’t come from just one mo­ment in time, but from a sequence of tens of millions of years, beginning in the Permian, continuing across the great extinction into the Triassic, and even reaching the next stage of geological time, the Jurassic Period, which began about 200 million years ago. When the seasonal lakes dried up, they left vast mud flats that animals walked across, leaving their marks. The rivers would continuously bring in new sediment to cover up the mud flats, burying them and turning them to stone. The cycle repeated year after year after year, so there is now layer upon layer upon layer of tracks in the Holy Cross Moun­tains. For paleontologists this is a bonanza: an opportunity to see how animals and ecosystems were changing over time, particularly after the cataclysmic end-Permian extinction.

Identifying what animals made which particular track is relatively straightforward. You compare the shape of the track to the shape of hands and feet. How many fingers or toes are there? Which ones are longest? Which way do they face? Do only the fingers and toes make an impression, or does the palm of the hand and arch of the foot also leave a mark? Are the left and right tracks really close to­gether, as the trackmaker was walking with its limbs right under its body, or are they far apart, made by a creature with limbs sprawled out to the side? By following this checklist, you can usually figure out which general group of animals left the tracks in question. Pinpointing an exact species is almost impossible, but distinguishing the tracks of reptiles from amphibians, or dinosaurs from crocodiles, is easy enough.

The Permian tracks from the Holy Cross Mountains are a diverse lot, and most were made by am­phibians, small reptiles, and early synapsids, progenitors of mammals that are often annoyingly, and incorrectly, described as mammal-like reptiles (although they are not actually reptiles) in kids’ books and museum exhibits. Gorgonopsians and dicynodonts are two types of these primitive synapsids. By all accounts these latest Permian ecosystems were strong—there were many varieties of animals, some small and others more than ten feet long and weighing over a ton, living together, thriving in the arid climate along the seasonal lakes. There are, however, no signs of dinosaur or crocodile tracks in the Permian layers, or even any tracks that look like precursors to these animals.

Everything changes at the Permian-Triassic boundary. Following the tracks across the extinction is like reading an arcane book in which a chapter of English follows one written in Sanskrit. The latest Permian and earliest Triassic seem to be two different worlds, which is remarkable because the tracks were all left in the identical place, in the same exact environment and climate. Southern Poland didn’t stop being a humid lakeland fed by raging mountain streams as the Permian ticked over into the Tri­assic. No, it was the animals themselves that changed.

I get the creeps when looking at the earliest Triassic tracks. I can sense the long-distant specter of death. There are hardly any tracks at all, just a few small prints here and there, but a lot of burrows jutting deep into the rock. It seems the surface world was annihilated and whatever creatures inhab­ited this haunted landscape were hiding underground. Almost all of the tracks belong to small lizards and mammal relatives, probably not much larger than a groundhog. Many of the diverse tracks of the Permian are gone, particularly those made by the larger proto-mammal synapsids, and they never reappear.

Things gradually start to improve as you follow the tracks up through time. More track types ap­pear, some of the prints get larger, and burrows become rarer. The world was clearly recovering from the shock of end-Permian volcanoes. Then, about 250 million years ago, just a couple of million years after the extinction, a new type of track starts showing up. They’re small, just a few centimeters long, about the size of a cat’s paw. They are arranged in narrow trackways, the five-fingered handprints po­sitioned in front of the slightly larger footprints, which have three long central toes flanked by a tiny toe on each side. The best place to find them is near a tiny Polish village called Stryczowice, where you can park your car at a bridge, scramble your way through thorns and bramble, and poke around the banks of a narrow stream littered with track-covered rock slabs. Grzegorz discovered the site when he was young and proudly took me there once, on a miserable July day of obscene humidity, bugs, rain, and thunder. After a few minutes of hacking through the weeds, we were soaked, my field notebook warping as ink started to run off the pages.

The tracks found here go by the scientific name of Prorotodactylus. Grzegorz wasn’t quite sure what to make of them. They were certainly different from the other tracks found alongside them, and all of the tracks from the Permian. But what kind of animal made them? Grzegorz had a hunch they could have something to do with dinosaurs, because an elderly paleontologist named Hartmut Haubold had reported similar tracks from Germany in the 1960s and had argued that they were made by early dinosaurs or close cousins. But Grzegorz wasn’t sold on the idea. He had spent most of his young career studying tracks and hadn’t spent much time with actual dinosaur skeletons, so it was difficult for him to match the prints to a trackmaker. That’s where I came in. For my master’s degree, I constructed a family tree of Triassic reptiles, a genealogy showing how the first dinosaurs were related to the other animals of the time. I spent months in museum collections studying fossil bones, so I knew the anatomy of the first dinosaurs quite well. As did Richard, who wrote a PhD thesis on early di­nosaur evolution. The three of us put our heads together to figure out what culprit was responsible for the Prorotodactylus tracks, and we did indeed conclude that it was a very dinosaurlike animal. We an­nounced our interpretation in a scientific paper we published in 2010.

The clues, of course, are in the details of the tracks. When I look at the Prorotodactylus trackways, the first thing that jumps out at me is that they are very narrow. There is only a little bit of space be­tween the left and right tracks in the sequence, just a few centimeters. There’s only one way for an an­imal to make tracks like this: by walking upright, with the arms and legs right underneath the body. We walk upright, so when we leave footprints on the beach, the left and right ones are very close to­gether. Same with a horse—take a look at the pattern of horseshoe impressions left by a galloping horse next time you’re on a farm (or wagering a few bucks at the track), and you’ll see what I mean. But this style of walking is actually quite rare in the animal kingdom. Salamanders, frogs, and lizards move in a different way. Their arms and legs stick out sideways from the body. They sprawl. That means their trackways are much wider, with big separation between the left and right tracks made by their spread-eagle limbs.

The Permian world was dominated by sprawlers. After the extinction, however, one new group of reptiles evolved from these sprawlers but developed an upright posture—the archosaurs. This was a landmark evolutionary event. Sprawling is all well and good for cold-blooded critters that don’t need to move very fast. Tucking your limbs under your body, however, opens up a new world of possibilities. You can run faster, cover greater distances, track down prey with greater ease, and do it all more effi­ciently, wasting less energy as your columnar limbs move back and forth in an orderly fashion rather

than twisting around like those of a sprawler.

Grzegorz Niedzwiedzki examines a life-size model of the Prorotodactylus trackmaker: a proto-di­nosaur very similar to the ancestor that gave rise to dinosaurs.

Courtesy of Grzegorz Niedzwiedzki._______________________________________________________

A handprint overlapping a footprint of Prorotodactylus, from Poland. For scale, the handprint is about 1 inch long.

Photo courtesy of the author

We may never know exactly why some of these sprawlers started walking upright, but it probably was a consequence of the end-Permian extinction. It’s easy to imagine how this new getup gave ar- chosaurs an advantage in the postextinction chaos, when ecosystems were struggling to recover from the volcanic haze, temperatures were unbearably hot, and empty niches abounded, waiting to be filled by whatever mavericks could evolve ways to endure the hellscape. Walking upright, it seems, was one of the ways in which animals recovered—and indeed, improved—after the planet was shocked by the volcanic eruptions.

Not only did the new upright-walking archosaurs endure, but they thrived. From their humble ori­gins in the traumatic world of the Early Triassic, they later diversified into a staggering variety of species. Very early, they split into two major lineages, which would grapple with each other in an evo­lutionary arms race over the remainder of the Triassic. Remarkably, both of these lineages survive today. The first, the pseudosuchians, later gave rise to crocodiles. As shorthand, they are usually re­ferred to as the crocodile-line archosaurs. The second, the avemetatarsalians, developed into pterosaurs (the flying reptiles often called pterodactyls), dinosaurs, and by extension the birds that, as we shall see, descended from the dinosaurs. This group is called the bird-line archosaurs. The Proroto- dactylus tracks from Stryczowice are some of the first signs of archosaurs in the fossil record, traces of the great-great-great-grandmother of this whole menagerie.

Exactly what kind of archosaur was Prorotodactylus? Some peculiarities in the footprints hold im­portant clues. Only the toes make an impression, not the metatarsal bones that form the arch of the foot. The three central toes are bunched very close together, the two other toes are reduced to nub­bins, and the back end of the print is straight and razor-sharp. These may seem like anatomical minu­tiae, and in many ways they are. But as a doctor is able to diagnose a disease from its symptoms, I can recognize these features as hallmarks of dinosaurs and their very closest cousins. They link to unique features of the dinosaur foot skeleton: the digitigrade setup, in which only the toes make con­tact with the ground when walking, the very narrow foot in which the metatarsals and toes are bunched together, the pathetically atrophied outer toes, the hinge-like joint between the toes and the metatarsals, which reflects the characteristic ankle of dinosaurs and birds, which can move only in a back-and-forth direction, without even the slightest possibility of twisting.

The Prorotodactylus tracks were made by a bird-line archosaur very closely related to the di­nosaurs. In scientific parlance, this makes Prorotodactylus a dinosauromorph, a member of that group that includes dinosaurs and the handful of their very closest cousins, those few branches just below the bloom of dinosaurs on the family tree of life. After the evolution of the upright-walking ar- chosaurs from the sprawlers, the origin of dinosauromorphs was the next big evolutionary event. Not only did these dinosauromorphs stand proudly on their erect limbs, but also they had long tails, big leg muscles, and hips with extra bones connecting the legs to the trunk, all of which allowed them to move even faster and more efficiently than other upright-walking archosaurs.

As one of the first dinosauromorphs, Prorotodactylus is something of a dinosaur version of Lucy, the famous fossil from Africa that belongs to a very humanlike creature but is not quite a true human, a member of our species, Homo sapiens. In the same way that Lucy looks like us, Prorotodactylus would have appeared and behaved very much like a dinosaur, but it’s simply not considered a true di­nosaur by convention. That’s because scientists decided long ago that a dinosaur should be defined as any members belonging to that group including the plant-eating Iguanodon and the meat-eating Megalosaurus (two of the first dinosaurs found by scientists in the 1820s) and all descendants of their common ancestor. Because Prorotodactylus did not evolve from this common ancestor, but slightly before it, it is not a true dinosaur by definition. But that’s just semantics.

In Prorotodactylus we’re looking at traces left behind by the type of animal that evolved into di­nosaurs. It was about the size of a house cat and would have been lucky to tip the scales at ten pounds. It walked on all fours, leaving handprints and footprints. Its limbs must have been quite long, judging from the big gaps between successive prints of the same hands and feet. The legs must have been particularly long and skinny, because the footprints often are positioned in front of the hand­prints, a sign that its feet were overstepping its hands. The hands were small and would have been good at grabbing things, whereas the long, compressed feet were perfect for running. The Proroto- dactylus animal would have been gangly looking, with the speed of a cheetah but the awkward propor­tions of a sloth, perhaps not the type of animal you would expect the great Tyrannosaurus and Bron­tosaurus to ultimately evolve from. And it wasn’t very common either: less than 5 percent of all the tracks found at Stryczowice belong to Prorotodactylus, an indication that these proto-dinosaurs were not especially abundant or successful when they first arose. Instead, they were far outnumbered by small reptiles, amphibians, and even other types of primitive archosaurs.

These rare, weird, not-quite-true-dinosaur dinosauromorphs continued to evolve as the world healed in the Early and Middle Triassic. The Polish track sites, stacked orderly in time sequence like the pages of a novel, document it all. Sites like Wióry, Palẹgi, and Baranow yield an equally unfamiliar array of dinosauromorph tracks—Rotodactylus, Sphingopus, Parachirotherium, Atreipus—which diver­sify over time. More and more track types show up; they get larger; they develop a greater diversity of shape, some even losing their outer toes entirely so that the center toes are all that remain. Some of the trackways stop showing impressions of the hand—these dinosauromorphs were walking on only their hind legs. By about 246 million years ago, dinosauromorphs the size of wolves were racing around on two legs, grabbing prey with their clawed hands, acting a whole lot like a pint-size version of a T. rex. They weren’t living only in Poland; their footprints are also found in France and Germany and the southwestern United States, and their bones start showing up in eastern Africa and later Argentina and Brazil. Most of them ate meat, but some of them turned vegetarian. They moved quickly, grew fast, had high metabolisms, and were active, dynamic animals compared to the lethargic amphibians and reptiles they were cohabitating with.

At some point, one of these primitive dinosauromorphs evolved into true dinosaurs. It was a radi­cal change in name only. The boundary between nondinosaurs and dinosaurs is fuzzy, even artificial, a by-product of scientific convention. The same way that nothing really changes as you cross the bor­der from Illinois into Indiana, there was no profound evolutionary leap as one of these dog-size di- nosauromorphs changed into another dog-size dinosauromorph that was just over that dividing line on the family tree that denotes dinosaurs. This transition involved the development of only a few new fea­tures of the skeleton: a long scar on the upper arm that anchored muscles to move the arms in and out, some tablike flanges on the neck vertebrae that supported stronger muscles and ligaments, and an open-window-like joint where the thighbone meets the pelvis. These were minor changes, and to be honest, we don’t really know what was driving them, but we know that the dinosauromorph-dinosaur transition wasn’t a major evolutionary jump. A far bigger evolutionary event was the origin of the swift­running, strong-legged, fast-growing dinosauromorphs themselves.

The first true dinosaurs arose some time between 240 and 230 million years ago. The uncertainty reflects two problems that continue to cause me headaches but are ripe to be solved by the next gen­eration of paleontologists. First, the earliest dinosaurs are so similar to their dinosauromorph cousins that it is hard to tell their skeletons apart, never mind their footprints. For instance, the puzzling Nyasasaurus, known from part of an arm and a few vertebrae from approximately 240-million-year-old rocks in Tanzania, may be the world’s oldest dinosaur. Or it may be just another dinosauromorph on the wrong side of the genealogical divide. The same is true of some of the Polish footprints, particu­larly the larger ones made by animals walking on their hind legs. Maybe some of these were made by real, true, honest-to-goodness dinosaurs. We just don’t have a good way of telling apart the tracks of the earliest dinosaurs and their closest nondinosaur relatives, because their foot skeletons are so similar. But maybe it doesn’t matter too much, as the origin of true dinosaurs was much less impor­tant than the origin of dinosauromorphs.

The other, much more glaring issue is that many of the fossil-bearing rocks of the Triassic are very poorly dated, particularly those from the early to middle parts of the period. The best way to figure out the age of rocks is to use a process called radiometric dating, which compares the percentages of two different types of elements in the rock—say, potassium and argon. It works like this. When a rock cools from a liquid into a solid, minerals form. These minerals are made up of certain elements, in our case including potassium. One isotope (atomic form) of potassium (potassium-40) is not stable, but slowly undergoes a process called radioactive decay, in which it changes into argon-40 and expels a small amount of radiation, causing the beeps you’d hear on a Geiger counter. Beginning the moment a rock solidifies, its unstable potassium starts changing into argon. As this process continues, the accumu­lating argon gas becomes trapped inside the rock where it can be measured. We know from lab exper­iments the rate at which potassium-40 changes into argon-40. Knowing this rate, we can take a rock, measure the percentages of the two isotopes, and calculate how old the rock is.

Radiometric dating revolutionized the field of geology in the middle of the twentieth century; it was pioneered by a Brit named Arthur Holmes, who once occupied an office a few doors down from mine at the University of Edinburgh. Today’s labs, like the ones run by my colleagues at New Mexico Tech and the Scottish Universities Environmental Research Centre near Glasgow, are high-tech, ultramod­ern facilities where scientists in white lab coats use multi million-dollar machines bigger than my old Manhattan apartment to date microscopic rock crystals. The techniques are so refined that rocks hun­dreds of millions of years old can be precisely dated to a small window of time, within a few tens or hundreds of thousands of years. These methods are so fine-tuned that independent labs routinely cal­culate the same dates for samples of the same rocks analyzed blindly. Good scientists check their work this way, to make sure their methodology is sound, and test after test has shown that radiomet­ric dating is accurate.

But there is one major caveat: radiometric dating works only on rocks that cool from a liquid melt, like basalts or granites that solidify from lava. The rocks that contain dinosaur fossils, like mudstone and sandstone, were not formed this way, but rather from wind and water currents that dumped sedi­ment. Dating these types of rocks is much more difficult. Sometimes a paleontologist is lucky and finds a dinosaur bone sandwiched between two layers of datable volcanic rocks that provide a time envelope for when that dinosaur must have lived. There are other methods that can date individual crystals found in sandstones and mudstones, but these are expensive and time-consuming. This means that it’s often difficult to date dinosaurs accurately. Some parts of the dinosaur fossil record have been well dated—when there are enough interspersed volcanic rocks to give a timeline or the in­dividual-crystal technique has been successful—but not the Triassic. There are just a handful of well- dated fossils, so we are not entirely confident of what order certain dinosauromorphs appeared in (es­pecially when trying to compare the ages of species found in distant parts of the world) or when true dinosaurs emerged out of the dinosauromorph stock.

ALL UNCERTAINTIES ASIDE, we do know that by 230 million years ago, true di­nosaurs had entered the picture. The fossils of several species with unques­tionable signature features of dinosaurs are found in well-dated rocks of that age. They’re found in a place far from where the earliest dinosauromorphs were cavorting in Poland—the mountainous canyons of Argentina.

Ischigualasto Provincial Park, in the northeastern part of Argentina’s San Juan Province, is the type of place that just looks as though it should be bursting with dinosaurs. It’s also called Valle de la Luna —the Valley of the Moon—and you could easily imagine its being on some other planet, full of wind- sculpted hoodoos, narrow gullies, rust-covered cliffs, and dusty badlands. To the northwest are the towering peaks of the Andes, and far to the south are the dry plains that cover most of the country, where cows graze on the grass that makes Argentine beef so delicious. For centuries Ischigualasto has been an important crossing for livestock making their way from Chile to Argentina, and today many of the few people who live in the area are ranchers.

This stunning landscape also happens to be the best place in the world for finding the oldest di­nosaurs. That’s because the red, brown, and green rocks that have been carved and eroded into such magical shapes were formed in the Triassic, in an environment both full of life and perfect for preserv­ing fossils. In many ways, this landscape was similar to the Polish lakelands that preserved the tracks of Prorotodactylus and other dinosauromorphs. The climate was hot and humid, although perhaps a little more arid and not pounded by such strong seasonal monsoons. Rivers snaked their way into a deep basin, occasionally bursting their banks during rare storms. Over a period of 6 million years, the rivers built up repeating sequences of sandstone, formed in the river channels, and mudstone, formed from the finer particles that escaped the river and settled out on the surrounding floodplains. Many dinosaurs frolicked on these plains, along with a wealth of other animals—big amphibians, piglike dicynodonts whose ancestors managed to make it through the end-Permian extinction, beaked plant-eating reptiles called rhynchosaurs (primitive cousins of the archosaurs) and furry little cyn- odonts that looked like a cross between a rat and an iguana. Floods would occasionally interrupt this paradise, killing the dinosaurs and their friends and burying their bones.

The area is so heavily eroded today, and so little disturbed by buildings and roads and other human nuisances that cover up fossils, that the dinosaurs are relatively easy to find, at least com­pared to so many other parts of the world where we hike around for days just praying to find anything, even just a tooth. The very first discoveries here were made by cowpokes or other locals, and it wasn’t until the 1940s that scientists began to collect, study, and describe fossils from Ischigualasto, then still another few decades until intensive expeditions were launched.

The first major collecting trips were led by one of the giants of twentieth-century paleontology, the Harvard professor Alfred Sherwood Romer, the man who wrote the textbook that I still use to teach my graduate students in Edinburgh. During his first trip, in 1958, Romer was already sixty-four years old and regarded as a living legend, yet there he was driving a rickety car through the badlands because he had a hunch that Ischigualasto would be the next big frontier. On that trip he found part of a skull and skeleton of a “moderately large” animal, as he so modestly put it in his field notebook. He brushed away as much rock as he could, coated the bones in newspaper, applied a coat of plaster that would harden and protect the bones, and chiseled them out of the ground. He sent the bones back to Buenos Aires, where they would be loaded on a ship to the United States, so he could care­fully clean and study them in his lab. But the fossils took a detour. They were impounded for two years at the port in Buenos Aires before customs officials finally gave the go-ahead. By the time the fossils arrived at Harvard, Romer had occupied himself with other things, and it was only years later that other paleontologists recognized that the master had found the very first good dinosaur from Is- chigualasto.

Some Argentines weren’t so happy that a Norteamericano had come down to their neighborhood to collect fossils, which were being removed from Argentina and studied in the United States. That spurred a pair of up-and-coming homegrown scientists, Osvaldo Reig and José Bonaparte, to organize their own expeditions. They assembled a team and set out for Ischigualasto in 1959, and then again three times during the early 1960s. It was during the 1961 field season that Reig and Bonaparte’s crew met a local rancher and artist named Victorino Herrera, who knew the hills and crevasses of Is- chigualasto the way an Inuit knows snow. He recalled seeing some bones crumbling out of the sand­stone and led the young scientists to the spot.

Herrera had found bones all right, lots of them, and clearly they were part of the back end of a di­nosaur skeleton. After a few years of study, Reig described the fossils as a new species of dinosaur that he called Herrerasaurus in the rancher’s honor, a mule-size creature that could sprint on its hind legs. Later detective work showed that Romer’s impounded fossils belonged to the same animal, and future discoveries revealed that Herrerasaurus was a fierce predator with an arsenal of sharp teeth and claws, a primitive version of T. rex or Velociraptor. Herrerasaurus was one of the very first thero- pod dinosaurs—a founding member of that dynasty of smart, agile predators that would later ascend to the top of the food chain and ultimately evolve into birds.

You might think this discovery would have encouraged paleontologists from throughout Argentina to flock to Ischigualasto in some kind of mad dinosaur rush. But it didn’t happen. After Reig and Bona­parte’s expeditions ended, things got quiet. The late 1960s and 1970s were not a prime time for di­nosaur research. There was little funding and, believe it or not, little public interest. Things picked up again in the late 1980s, when a thirty-something paleontologist from Chicago named Paul Sereno put together a joint Argentine-American team of other ambitious young guns, mostly graduate students and junior professors. They set out in the footsteps of Romer, Reig, and Bonaparte, the latter meeting with the group for a few days to guide them to some of his favorite sites. The trip was a rousing suc­cess: Sereno found another skeleton of Herrerasaurus and many other dinosaurs, proving that Is- chigualasto still had plenty of fossils to give up.

Three years later, Sereno was at it again, bringing much of the same crew back to Ischigualasto to explore new territory. One of his assistants was a wisecracking student named Ricardo Martinez. While out prospecting one day, Martinez picked up a fist-size hunk of rock covered in a gnarly frosting of iron minerals. Just another piece of junk, he thought, but as he reached back to toss it aside, Martinez noticed something pointy and shiny sticking out of the cobble. They were teeth. Glancing back at the ground, dumbfounded, he realized that he had plucked the head off the nearly complete skeleton of a dinosaur, a long-legged, lightly built speed demon about the size of a golden retriever. They named it Eoraptor. Those teeth poking out from the skull turned out to be highly unusual: the ones in the back of the jaw were sharp and serrated like a steak knife, surely to slice through flesh, but the ones at the tip of the snout were leaf-shaped with coarse projections called denticles, the same type of tooth that some long-necked, potbellied sauropod dinosaurs would later use to grind plants. This hinted that Eoraptor was an omnivore and possibly a very early member of the sauropod lineage, a primitive cousin of Brontosaurus and Diplodocus.

I met Ricardo Martinez many years later, around the time that I first laid eyes on the gorgeous skeleton of Eoraptor. I was an undergraduate student at the University of Chicago, training in Paul Sereno’s lab, when Ricardo came to work on a clandestine project, later announced as yet another new dinosaur from Ischigualasto, the terrier-size primitive theropod Eodromaeus. I took a liking to Ri­cardo right away. Paul was running an hour late, stuck in traffic on Lake Shore Drive, and Ricardo was literally twiddling his thumbs, hunched in the corner of the lab office. It was an incongruously disen­gaged posture from a man who very quickly revealed himself to be the very type of hot blooded, fast­talking, fossil-loving typhoon that I longed to be. He kind of looked like the Dude from The Big Lebowski: wild tangled hair, beard thick around the mouth, interesting fashion sense. He regaled me with stories of working in the wilds of Argentina, recounting with theatrical hand gestures how his hun­gry crew would sometimes hunt down stray cattle on their ATVs, delivering killing blows with the busi­ness end of their geological rock hammers. He could tell I was developing a romantic attraction to Ar­gentina and told me to look him up if I ever came to visit.

Five years later, I took him up on the offer when I attended the hardest-rocking scientific confer­ence I’ve ever had the pleasure of speaking at. Usually conferences are fairly stale affairs, held in Marriotts and Hyatts in cities like Dallas and Raleigh, where scientists gather to listen to each other speak in cavernous banquet halls that usually host weddings, drinking overpriced hotel beer while catching up on field stories. The conference that Ricardo and his colleagues hosted in the city of San Juan was anything but. The dinner on the last evening was legendary, like one of those hedonistic house parties in a rap video. A local politician adorned with a sash opened the proceedings, managing to make an outrageous quip about some of the foreigners in attendance. The main course was a phonebook-size slab of grass-fed beef, washed down with copious amounts of red wine. After dinner was dancing, for hours, fueled by an open bar with hundreds of bottles of vodka, whiskey, brandy, and a local firewater whose name I can’t remember. At about three a.m., there was a break in the pro­ceedings while a make-your-own taco bar was assembled outside, a tasty change from the humidity of the dance floor. We staggered back to our hotels as dawn broke. Ricardo was right. I would love Ar­gentina.

Before the debauchery of that evening, I spent several days in the collections of Ricardo’s mu­seum, the Instituto y Museo de Ciencias Naturales in the lovely city of San Juan. Most of the riches of Ischigualasto are kept here, Herrerasaurus, Eoraptor, and Eodromaeus among them, but also many other dinosaurs. There’s Sanjuansaurus, a close cousin of Herrerasaurus that was also a fierce preda­tor. In another drawer is Panphagia, similar to Eoraptor in being a primitive miniature cousin of the later colossal sauropods, and Chromogisaurus, a larger Brontosaurus relative that grew up to a cou­ple of meters long and was something of a middle-of-the-food-chain plant-eater. There are also the scrappy fossils of a dinosaur called Pisanosaurus, a dog-size animal that shares some features of the teeth and jaws with the ornithischian dinosaurs—the group that would later diversify into a vast range of plant-eating species, from the horned Triceratops to the duck-billed hadrosaurs. And they’re still finding new dinosaurs in Ischigualasto, so who knows what new characters will be added if you are lucky enough to visit.

As I was pulling open the specimen cabinet doors, carefully removing the fossils to measure and photograph them, I felt like something of an historian, one of those scholars who spends dark hours in the archives, scrutinizing ancient manuscripts. The analogy is deliberate, because the Ischigualasto fossils are indeed historical artifacts, primary-source objects that help us tell the story of deep prehis­toric pasts, millions of years before monks started writing on parchment. The bones that Romer, Reig, and Bonaparte, and then later Paul, Ricardo, and their many colleagues, have pried from the lunar landscape of Ischigualasto are the very first records of true dinosaurs, living, evolving, and beginning their long march to dominance.

These first dinosaurs weren’t quite dominant yet, overshadowed by the larger and more diverse amphibians, mammal cousins, and crocodile relatives that they lived alongside on those dry, occa­sionally flooded plains of the Triassic. Even Herrerasaurus probably wasn’t at the top of the food chain, ceding that title to the murderous twenty-five-foot-long crocodile-line archosaur Saurosuchus. But the dinosaurs had arrived on the scene. The three major groups—the meat-eating theropods, long­necked sauropods, and herbivorous ornithischians—had already diverged from each other on the fam­ily tree, siblings setting out to form their own broods.

The skull of Eoraptor and the hand of Herrerasaurus, two of the oldest dinosaurs. Photo courtesy of the author

The dinosaurs were on the march.

Dinosaurs Rise Up


Chapter Title art by Todd Marshall

IMAGINE A WORLD WITH NO BORDERS. I’m not channeling John Lennon. What I mean is, envision a version of Earth where all of the land is connected together—no patchwork of continents separated by oceans and seas, just a single expanse of dry ground stretching from pole to pole. Given enough time and a good pair of shoes, you could walk from the Arctic Circle across the equator to the South Pole. If you ventured too far inland, you would find yourself many thousands of miles—tens of thousands, even—from the closest beach. But if you fancied a swim, you could take a dip in the vast ocean surrounding the big slab of land you called home and, theoretically at least, paddle from one coast all the way around the planet to the other coast without having to dry off.

It may sound fanciful, but this is the world the dinosaurs grew up in.

When the very first dinosaurs, like Herrerasaurus and Eoraptor, evolved from their cat-size di- nosauromorph ancestors some 240 to 230 million years ago, there were no individual continents—no Australia or Asia or North America. There was no Atlantic Ocean separating the Americas from Europe and Africa, no Pacific Ocean on the flip side of the globe. Instead, there was just one huge solid unbro­ken mass of land—what geologists refer to as a supercontinent. It was surrounded by a single global ocean. Geography class would have been easy in those days: the supercontinent we call Pangea, and the ocean we call Panthalassa.

The dinosaurs were born into what we would see as a totally alien world. What was it like to live in such a place?

First, let’s think about the physical geography. The supercontinent spanned an entire hemisphere of the Triassic Earth from North Pole to South. It looked something like a gigantic letter C, with a big in­dentation in the middle where an arm of Panthalassa cut into the land. Towering mountain ranges snaked across the landscape at odd angles, marking the sutures where smaller blocks of crust had once collided to build the giant continent, the pieces of a jigsaw puzzle. This puzzle wasn’t put to­gether very easily or very quickly. For hundreds of millions of years, heat deep inside the planet pushed and tugged on the many smaller continents that were home to generations of animals long before the dinosaurs, until all of the land was globbed together into one sprawling kingdom.

And what about the climate? No better way to put it: the earliest dinosaurs lived in a sauna. The Earth was a whole lot warmer back in the Triassic Period than it is today. In part, that’s because there was more carbon dioxide in the atmosphere, so more of a greenhouse effect, more heat radiating across the land and sea. But the geography of Pangea exacerbated things. On one side of the globe, dry land extended from pole to pole, but on the other side, there was open ocean. That meant that currents could travel unimpeded from the equator to the poles, so there was a direct path for water baked in the low-latitude sun to heat up the high-latitude regions. This prevented ice caps from form­ing. Compared to today, the Arctic and Antarctic were balmy, with summer temperatures similar to those of London or San Francisco, and winter temperatures that barely inched below freezing. They were places that early dinosaurs and the other creatures with whom they shared the earth could eas­ily inhabit.

If the poles were that warm, then the rest of the world must have been a hothouse. But it’s not as though the entire planet was a desert. Once again the geography of Pangea made things much more complex. Because the supercontinent was basically centered on the equator, half the land was always scorching in the summer while the other half was cooling down in the winter. The marked temperature differences between north and south caused violent air currents to regularly stream across the equa­tor. When the seasons changed, these currents shifted direction. That kind of thing happens today in some parts of the world, particularly India and Southeast Asia. It’s what drives the monsoons, the al­ternation of a dry season with a prolonged deluge of rain and nasty storms. You’ve probably seen im­ages in the newspaper or on the nightly news: floods drowning homes, people fleeing from raging tor­rents, mudslides burying villages. The modern monsoons are localized, but the Triassic ones were global. They were so severe that geologists have invented a hyperbolic term to describe them: mega­monsoons.

Many a dinosaur was probably swept away by floodwaters or entombed by mud avalanches. But the megamonsoons also had another effect. They helped divide Pangea into environmental provinces, characterized by different amounts of precipitation, varying severity of the monsoonal winds, and dif­ferent temperatures. The equatorial region was extremely hot and humid, a tropical hell that would make summer in today’s Amazon seem a trip to Santa’s workshop by comparison. Then there were vast stretches of desert, extending about 30 degrees of latitude on either side of the equator—like the Sahara, only covering a much broader swath of the planet. Temperatures here were well into the hun­dreds (over 35 degrees Celsius), probably all year long, and the monsoonal rains that pounded other parts of Pangea were absent here, offering little more than a trickle of precipitation. But the mon­soons exerted a great impact in the midlatitudes. These areas were slightly cooler but much more wet and humid than the deserts, far more hospitable to life. Herrerasaurus, Eoraptor, and the other Is- chigualasto dinosaurs lived in such a setting, smack in the middle of the midlatitude humid belt of southern Pangea.

Pangea may have been a united landmass, but its treacherous weather and extreme climates gave it a dangerous unpredictability. It wouldn’t have been a particularly safe or pleasant place to call home. But the very first dinosaurs had no choice. They entered a world still recovering from the terri­ble mass extinction at the end of the Permian, a land subject to the violent whims of storms and the blight of blistering temperatures. So did many other new types of plants and animals that were getting their start after the mass extinction cleared the planet. All of these newbies were thrust onto an evolu­tionary battlefield. It was far from certain that dinosaurs were going to emerge triumphant. After all, they were small and meek creatures, nowhere near the top of the food chain during their earliest years. They were hanging around with lots of other species of small-to-midsize reptiles, early mam­mals, and amphibians in the middle of the food pyramid, fearful of the crocodile-line archosaurs, who held the throne. Nothing was handed to the dinosaurs. They were going to have to earn it.

DURING MANY SUMMERS, I’ve journeyed deep into the subtropical arid belt of northern Pangea, on the hunt for fossils. Of course, the supercontinent itself is long gone, having gradually fractured into our modern continents during the more than 230 million years since the primeval dinosaurs started their evolu­tionary march. What I’ve been exploring is a remnant of old Pangea that can be found in the sunny Algarve region of Portugal, at the very southwestern corner of Europe. During those formative years when dinosaurs were navigating the megamonsoons and boiling heat waves of the Triassic, this part of Portugal was only 15 or 20 degrees north of the equator, about the same latitude as Central America today.

As with so many adventures in paleontology, it was a random clue that put Portugal on my radar. After our first jaunt together in Poland, visiting Grzegorz and studying fossils of some of the dinosauro- morph ancestors of dinosaurs, my British buddy Richard Butler and I developed something of an ad­diction. We became obsessed with the Triassic Period. We wanted to understand what the world was like when dinosaurs were still young and vulnerable. So we scoured the map of Europe looking for other places where there were accessible rocks of Triassic age, the type of sediments that could con­ceivably contain the fossils of dinosaurs and other animals living alongside them. Richard came across a short paper in an obscure scientific journal, describing some scraps of bone from southern Portugal that were collected by a German geology student in the 1970s. The student had been in Por­tugal to make a map of the rock formations, a rite of passage for all undergraduate geology majors. He had little interest in fossils, so he threw the specimens in his rucksack and hauled them back to Berlin, where they languished in a museum for nearly three decades until some paleontologists recog­nized them as skull pieces of ancient amphibians. Triassic amphibians. That was enough to get us ex­cited. There were Triassic fossils in a beautiful part of Europe and nobody had been looking for them for decades. We had to go.

That tip brought Richard and me to Portugal in the late summer of 2009, the hottest part of the year. We teamed up with another friend, Octávio Mateus, who wasn’t even thirty-five years old at the time but was already regarded as Portugal’s leading dinosaur hunter. Octavio grew up in a little town called Lourinhã, on the windy Atlantic coast north of Lisbon. His parents were amateur archaeologists and historians who spent weekends exploring the countryside, which just so happened to be strewn with Jurassic dinosaur fossils. The Mateus family and their ragtag band of local enthusiasts collected so many dinosaur bones, teeth, and eggs that they needed a place to put them, so when Octávio was nine years old, his parents started their own museum. Today, the Museu da Lourinhã houses one of the most important collections of dinosaurs in the world, many of which have been collected by Octávio—who went on to study paleontology and become a professor in Lisbon—and by his ever-ex­panding army of students, volunteers, and homegrown helpers.

It was fitting that Octavio, Richard, and I set out in the August heat, because we were chasing the fossils of animals that lived in the very hottest and driest sector of Pangea. But it wasn’t very good strategy on our part. For several days, we hiked through the sun-baked hills of the Algarve, our sweat soaking the geological maps that we hoped would lead us to our treasure. We checked out nearly every speck of Triassic-age rock on the maps and relocated the site where the geology student had collected his amphibian bones, but all we saw were fossil crumbs. As our week in the field drew to a close, we were hot and exhausted, and staring down the barrel of failure. On the verge of defeat, we thought we should take one more hike in the area where the geology student made his discovery. It was a scorcher of a day, the thermometer on our handheld GPS units reaching 120 degrees Fahren­heit (50 Celsius).

After an hour or so of prospecting together, we decided to split up. I stayed near the base of the hills, scrutinizing the fragments of bone scattered across the ground in a desperate attempt to trace them to their source. I had no luck. But then I heard an excited voice scream from somewhere up on the ridge. I detected a hint of a lyrical Portuguese accent, so it must have been Octávio. I rushed to­ward where I thought the voice was coming from, but now there was nothing but silence. Maybe I was imagining things, the heat playing tricks on my brain. Eventually I saw Octávio in the distance, rubbing his eyes like someone woken up by a phone call in the middle of the night. He was stumbling, giving off a bit of a zombie vibe. It was weird.

When Octávio saw me, he gathered himself and burst into song. “I found it, I found it, I found it,” he repeated over and over. He was holding a bone. What he didn’t have was a water bottle. And sud­denly it made sense. He had forgotten his water in the car, a bad thing for such a hot day, but he had happened upon the layer where the amphibian bones were coming out. The combination of exhilara­tion and dehydration had caused him to pass out for a moment. But now he was back into conscious­ness, and a few moments later, Richard had scrambled his way through the brush to join us. After ex­changing excited hugs and high fives, we celebrated further by rehydrating with beers at a small café down the road.

What Octávio had found was a half-meter-thick layer of mudstone full of fossil bones. We returned several times over the next few years to meticulously excavate the site, which turned out to be a chore because the bone layer seemed to extend infinitely into the hillside. I had never seen so many fossils concentrated together in one area. It was a mass graveyard. Countless skeletons of amphibians called Metoposaurus—supersize versions of today’s salamanders that were the size of a small car— were jumbled together in a chaotic mess. There must have been hundreds of them. Some 230 million years ago, a flock of these slimy, ugly monsters suddenly died when the lake they were living in dried up, collateral damage of the capricious Pangean climate.

Giant amphibians like Metoposaurus were leading actors in the story of Triassic Pangea. They prowled the shores of rivers and lakes over much of the supercontinent, particularly the subtropical arid regions and midlatitude humid belts. If you were a frail little primitive dinosaur like Eoraptor, you would want to avoid the shorelines at all costs. It was enemy territory. Metoposaurus was there wait­ing, lurking in the shallows, ready to ambush anything that ventured too close to the water. Its head was the size of a coffee table, and its jaws were studded with hundreds of piercing teeth. Its big, broad, almost flat upper and lower jaws were hinged together at the back and could snap shut like a toilet seat to gobble up whatever it wanted. It would only take a few bites to finish off a delicious di­nosaur supper.

Salamanders bigger than humans seem like a mad hallucination. As bizarre as they were, though, Metoposaurus and its kin were not aliens. These terrifying predators were the ancestors of today’s frogs, toads, newts, and salamanders. Their DNA flows through the veins of the frog hopping around your garden or the one you dissected in high school biology class. As a matter of fact, many of today’s most recognizable animals can be traced back to the Triassic. The very first turtles, lizards, crocodiles, and even mammals came into the world during this time. All of these animals—so much a fabric of the Earth we call home today—rose up alongside the dinosaurs in the harsh surroundings of prehistoric Pangea. The apocalypse of the end-Permian extinction left such an empty playing field that there was space for all sorts of new creatures to evolve, which they did unabated during the 50 million years of the Triassic. It was a time of grand biological experimentation that changed the planet forever and re­verberates still today. It’s no wonder many paleontologists refer to the Triassic as the “dawn of the modern world.”

Excavating the Metoposaurus bone bed in Algarve, Portugal, with Octávio Mateus, Richard Butler, and our team.

Photo courtesy of the author

If you could put yourself into the tiny feet of our furry, mouse-size Triassic mammalian ancestors, you would be looking up at a world that was starting to show whispers of today. Yes, the physical planet itself was completely different—a supercontinent, marked by intense heat and violent weather. Nevertheless, the parts of the land not engulfed by desert were covered in ferns and pine trees. There were lizards darting around in the forest canopy, turtles paddling in the rivers, amphibians running amok, many familiar types of insects buzzing around. And there were dinosaurs, mere bit characters in this ancient scene but destined for greater things to come.

AFTER SEVERAL YEARS of excavating the supersalamander mass grave in Portugal, we’ve collected a lot of bones of Metoposaurus, enough to fill the workshop in Octávio’s museum. But we’ve also found other animals that died when the pre­historic lake evaporated. We dug up part of the skull of a phytosaur, a long­snouted relative of crocodiles that hunted on land and in the water. We’ve scooped up many teeth and bones of various fishes, which were probably the primary source of food for Metoposaurus. Other small bones hint at a badger­size reptile.

What we haven’t found yet are any signs of dinosaurs.

It’s strange. We know dinosaurs were living south of the equator, in the humid river valleys of Is- chigualasto, at the same general time that Metoposaurus was terrorizing the lakes of Triassic Portu­gal. We also know that many different types of dinosaurs were commingling in Ischigualasto: all of those creatures that I studied in Ricardo Martfnez’s museum in Argentina. Meat-eating theropods like Herrerasaurus and Eodromaeus, primitive long-necked sauropod precursors like Panphagia and Chro- mogisaurus, early ornithischians (cousins of the horned and duck-billed dinosaurs). No, they weren’t at the top of the food pyramid. Yes, they were outnumbered by the jumbo amphibians and crocodile relatives, but they were at least beginning to make their mark.

So why don’t we see them in Portugal? It could be, of course, that we just haven’t found them yet. Absence of evidence is not always evidence of absence, as all good paleontologists must continually remind themselves. Next time we go back into the scrublands of the Algarve and carve out another section of the bone bed, maybe we’ll find ourselves a dinosaur. However, I’m willing to bet against that, because a pattern is starting to emerge as paleontologists discover more and more Triassic fos­sils from around the world. Dinosaurs seem to be present and starting to slowly diversify in the tem­perate humid parts of Pangea, particularly in the southern hemisphere, during a slice of time from about 230 to 220 million years ago. Not only do we find their fossils in Ischigualasto, but also in parts of Brazil and India that were once in the Pangean humid zone. Meanwhile, in the arid belts closer to the equator, dinosaurs were absent or extremely rare. Just as in Portugal, there are great fossil sites in Spain, Morocco, and along the eastern coast of North America where you can find plenty of amphib­ians and reptiles, but nary a dinosaur. All of these places were in the parched arid sector of Pangea during those 10 million years when dinosaurs were beginning to blossom in the more bearable humid regions. It seems these first dinosaurs couldn’t handle the desert heat.

It’s an unexpected story line. Dinosaurs didn’t just sweep across Pangea the moment they origi­nated, like some infectious virus. They were geographically localized, held in place not by physical bar­ricades but by climates they couldn’t endure. For many millions of years, it looked as if they might re­main provincial rubes, stuck in one zone in the south of the supercontinent, unable to break free—an aging high school football hero of faded dreams, who could have been something if only he’d been able to get out of his tiny hometown.

Underdogs—that’s what these first humidity-loving dinosaurs were. They wouldn’t have been a very impressive bunch. Not only were they trapped by the deserts, but even where they were able to eke out a living, they were barely getting by, at least at first. True, there were several dinosaur species in Ischigualasto, but these made up only about 10 to 20 percent of the total ecosystem. They were vastly outnumbered by early mammal relatives, like the pig-mimic dicynodonts that ate roots and leaves, and by other types of reptiles, most notably rhynchosaurs, which chopped plants with their sharp beaks, and crocodile cousins like the mighty apex predator Saurosuchus. At the same time but slightly to the east, in what is now Brazil, the story was much the same. There were a few different types of di­nosaurs closely related to species in Ischigualasto: the carnivorous Staurikosaurus was a cousin of Herrerasaurus, and the small long-necked creature Saturnalia was very similar to Panphagia. But they were quite rare, again overwhelmed by masses of proto-mammals and rhynchosaurs. Even farther to the east, where the humid zone continued into what is now India, there were a handful of primitive long-necked sauropod relatives, like Nambalia and Jaklapallisaurus, but once again they were role players in ecosystems ruled by other species.

Then, when it appeared that dinosaurs were never going to escape their rut, two important things happened that gave them an opening.

First, in the humid belt, the dominant large plant-eaters, the rhynchosaurs and dicynodonts, be­came less common. In some areas they disappeared entirely. We don’t yet fully understand why, but the consequences were unmistakable. The fall of these herbivores gave the plant-eating primitive sauropod cousins like Panphagia and Saturnalia an opportunity to seize a new niche in some ecosys­tems. Before long they were the main herbivores in the humid regions of both the southern and north­ern hemispheres. In the Los Colorados Formation of Argentina, a unit of rock laid down from about 225 to 215 million years ago that was formed directly after the Ischigualasto dinosaurs left their fos­sils, the sauropod antecedents are the most common vertebrates. There are more fossils of these cow-to-giraffe-size plant-guzzlers—among them Lessemsaurus, Riojasaurus, and Coloradisaurus—than any other type of animal. In all, dinosaurs comprise about 30 percent of the ecosystem, while the once dominant mammal relatives dip below 20 percent.

It wasn’t only a southern Pangean story. Across the equator in primeval Europe, then part of the Northern Hemisphere humid sector, other long-necked dinosaurs were also thriving. As in Los Col- orados, they were the most common large plant-eaters in their habitats. One of these species, Pla- teosaurus, has been found at over fifty sites throughout Germany, Switzerland, and France. There are even mass graves like the Metoposaurus bone bed in Portugal, where dozens (or more) of Pla- teosauruses died together when the weather turned rough, a sign of just how many of these dinosaurs were flocking across the landscape.

The second major breakthrough, around 215 million years ago, was that the first dinosaurs began arriving in the subtropical arid environments of the Northern Hemisphere, then about 10 degrees

above the equator, now part of the American Southwest. We don’t know exactly why dinosaurs were now able to migrate out of their safe humid homes and into the harsh deserts. It probably had some­thing to do with climate change-shifts in the monsoons and the amount of carbon dioxide in the at­mosphere made the differences between the humid and arid regions less stark, so dinosaurs could move more easily between them. Whatever the reason, at long last dinosaurs were making inroads into the tropics, expanding into parts of the world that had previously eluded them.

The best records of desert-living Triassic dinosaurs come from areas that are once again deserts today. Across much of the postcard-pretty landscape of northern Arizona and New Mexico are hoodoos, badlands, and canyons carved out of colorful red and purple rocks. These are the sand­stones and mudstones of the Chinle Formation, a third-of-a-mile-thick rock sequence formed from the ancient sand dunes and oases of tropical Pangea during the last half of the Triassic, from about 225 to 200 million years ago. Petrified Forest National Park, which should be on the itinerary of any dino- loving tourist visiting the southwestern states, has one of the best exposures of the Chinle Formation, full of thousands of enormous fossilized trees that were uprooted and buried in flash floods right around the time that dinosaurs were starting to settle in the area.

Some of the most exciting paleontological fieldwork over the past decade has targeted the Chinle Formation. New discoveries have painted a striking new image of what the first desert-dwelling di­nosaurs were like and how they fit into their broader ecosystems. Leading the charge is a remarkable group of young researchers, who were all graduate students when they began exploring the Chinle. The core of the group is the four-man band of Randy Irmis, Sterling Nesbitt, Nate Smith, and Alan Turner. Irmis is a bespectacled introvert but a beast of a field geologist; Nesbitt is an expert on fossil anatomy who’s always wearing a baseball cap and quoting television comedy shows; Smith is a smooth-dressing Chicagoan who likes to use statistics to study dinosaur evolution; and Turner, an ex­pert on building family trees of extinct groups, is affectionately called Little Jesus because of his flow­ing locks, bushy beard, and moderate stature.

The quartet is a half generation ahead of me on the career path. They were working on their PhDs when I was starting to do research as an undergraduate. As a young student, I was in awe of them, as if they were a paleontology Rat Pack. They traveled in a herd at research conferences, often with other friends of theirs who worked in the Chinle: Sarah Werning, a specialist on how dinosaurs and other reptiles grew; Jessica Whiteside, a brilliant geologist who studied mass extinctions and ecosystem changes in deep time; Bill Parker, the paleontologist at Petrified Forest National Park and an expert on some of the close crocodile relatives who lived with early dinosaurs; Michelle Stocker, who studied some of the other proto-crocodiles (and whom Sterling Nesbitt later convinced to marry him-propos- ing on a field trip, no less-forming a different sort of Triassic dream team). They were the hotshot young scientists whom I looked up to, the type of researchers I wanted to become.

For many years, the Chinle Rat Pack has been spending summers in northern New Mexico, in the pastel drylands near the tiny hamlet of Abiquiú. In the mid-1800s, this outpost was an important stop on the Old Spanish Trail, a trade route that linked nearby Santa Fe with Los Angeles. Today only a few hundred people remain, making the area feel like a remote backwater within the world’s most indus­trialized country. Some people like that kind of seclusion, though. One of them was Georgia O’Keeffe, the modernist American artist famous for her paintings of flowers that were intimate to the point of abstraction. O’Keeffe was also drawn to sweeping landscapes, and she was moved by the striking beauty and incomparable hues of natural light in the Abiquiú area. She bought a house nearby, on the sprawling grounds of a desert retreat called Ghost Ranch. There she could explore nature and experi­ment with new painting styles without being bothered by anyone. The red cliffs and colorful candy- striped canyons of the ranch, bathed in sparkling sunbursts, are common motifs in the work she pro­duced here.

After O’Keeffe died, in the mid-1980s, Ghost Ranch became a pilgrimage site for art lovers hoping to catch some of that desert spark that so inspired the old master. Few of these cultured travelers probably realize that Ghost Ranch is also bursting with dinosaur bones.

But the Rat Pack knew.

They understood that in 1881 a scientific mercenary named David Baldwin had been sent to northern New Mexico by the Philadelphia paleontologist Edward Drinker Cope, with the singular mis­sion to find fossils that Cope could stick in the face of his Yale rival, Othniel Charles Marsh. The two Easterners were engaged in a bitter feud known to history as the Bone Wars (of which, more later), but by this stage of their careers, neither of them particularly liked to brave the elements and Native American war parties—Geronimo would continue raiding New Mexico and Arizona until 1886. Rather than look for fossils themselves, they relied on a network of hired guns. Baldwin was the type of char­acter they often employed: a mysterious loner who would jump on his mule and head deep into the badlands for months at a time, even during the bleak winter, and eventually emerge loaded up with di­nosaur bones. In fact, Baldwin had worked for both of the pugnacious paleontologists: he was once a trusted confidant of Marsh’s, but now his loyalties were with Cope. Thus it was Cope who was the lucky recipient of the collection of small, hollow dinosaur bones that Baldwin pried out of the desert near Ghost Ranch. These bones belonged to a totally new type of dog-size, lightweight, fast-running, sharp-toothed, primitive Triassic dinosaur Cope later called Coelophysis. Like Herrerasaurus from Ar­gentina, which would be found many decades later, it was one of the earliest members of the thero- pod dynasty that would eventually produce T. rex, Velociraptor, and birds.

The Chinle Rat Pack also knew that a half century after Baldwin’s discovery, another East Coast paleontologist, Edwin Colbert, took a liking to the Ghost Ranch area. He was a much more pleasant in­dividual than Cope or Marsh. When Colbert set out for Ghost Ranch in 1947, he was in his early for­ties, already ensconced in one of the top jobs in the field: curator of vertebrate paleontology at New York City’s American Museum of Natural History. That summer, while O’Keeffe was painting mesas and rock sculptures only a few miles away, Colbert’s field assistant George Whitaker made an as­tounding discovery. He came across a Coelophysis graveyard, hundreds of skeletons in all, a pack of predators buried by a freak flood. I can imagine he must have felt something similar to our unbridled joy when we found our Metoposaurus bone bed in Portugal. Overnight Coelophysis became the quin­tessential Triassic dinosaur, the creature that immediately came to mind when people envisioned what the earliest dinosaurs looked like, how they behaved, and what environments they lived in. For years the American Museum crew kept digging and digging, hacking out blocks of the bone bed, which were distributed to museums around the world. Odds are, if you go see a big dinosaur exhibit today, you’ll see a Ghost Ranch Coelophysis.

The Chinle Rat Pack also knew of one final, and perhaps most important, clue. Because so many Coelophysis skeletons were found together, excavating the mass grave site diverted everyone’s atten­tion for decades. It sucked up most of the money for fieldwork, most of the time and energy of the field crews. But it was merely a single site in the expanse of Ghost Ranch, tens of thousands of acres covered by fossil-rich Chinle rocks. More must have been out there. So it was no surprise to them when, in 2002, a retired forest manager named John Hayden discovered some bones while hiking less than half a mile from the main gate of Ghost Ranch.

A few years later, the team of Irmis, Nesbitt, Smith, and Turner returned to the spot, got out their tools, and started digging. It took a lot of time and a lot of sweat. Once, when I was catching up with the quartet in a New York City Irish pub, Nate Smith turned to me, cocked his head up toward the ceil­ing, and said with a hint of cheeky machismo, “The amount of rock we removed that summer, yeah, it would fill up this bar.”

But the toil was worth it. The crew confirmed that there were indeed fossil bones at the site. Then they kept finding more and more of them, hundreds, thousands. It turned out to be a river channel de­posit, where currents had dumped the skeletons of many unlucky creatures swept into the water some 212 million years ago. With the right cocktail of good detective work and a drive to make their own discoveries even though they were still students, the Rat Pack had unearthed a treasure trove of Triassic fossils. The site—nicknamed the Hayden Quarry after the sharp-eyed forester who noticed the first fossil eroding out of the ground—has become one of the world’s most important Triassic fossil lo­calities.

The skull of Coelophysis, the primitive theropod found in abundance at Ghost Ranch.

Courtesy of Larry Witmer.

The quarry provides a snapshot of an ancient ecosystem, one of the first deserts that dinosaurs were able to live in. It wasn’t the picture the Chinle Rat Pack was expecting. When the young maver­icks started digging in the mid-2000s, the prevailing wisdom was that dinosaurs conquered the deserts soon after they arrived in the Late Triassic. Other scientists had collected a wealth of fossils from similar-age rock units in New Mexico, Arizona, and Texas, which seemed to belong to more than a dozen species of dinosaurs, ranging from stocky apex predators and smaller meat-eaters to many different types of plant-munching ornithischians, the ancestors of Triceratops and the duckbills. It seemed that dinosaurs were everywhere. But that wasn’t the case in the Hayden Quarry. There were monster amphibians closely related to our Portuguese Metoposaurus, primitive crocodiles and some of their long-snouted and armored relatives, skinny reptiles with short legs called Vancleavea, which looked like scaly dachshunds, and even funny little reptiles that hung from the trees like chameleons, called drepanosaurs. Those are the common animals in the quarry. Dinosaurs were anything but. The Rat Pack found only three types of dinosaurs: a fleet-footed predator very similar to Baldwin’s Coelo- physis, another swift carnivore called Tawa, and a somewhat larger and stockier meat-eater called Chindesaurus, which was closely related to the Argentine Herrerasaurus. Each is represented by only a few fossils.

It was a great surprise to the team. Dinosaurs were rare in the tropical deserts of the Late Triassic, and it was only the meat-eaters that seemed to be hanging about. There were no plant-eating di­nosaurs, none of the ancestral long-necked species that were so common in the humid zones, none of the ornithischian forebears of Triceratops. It’s a meager bunch of dinosaurs surrounded by all sorts of bigger, meaner, more common, more diverse animals.

What, then, to make of the dozens of Triassic dinosaur species that other scientists had identified from all over the American Southwest? Irmis, Nesbitt, Smith, and Turner scrutinized all of the evidence they could find, traveling to every small-town museum where researchers had deposited their fossils. They saw that most of these specimens were isolated teeth and scraps of bone, not the best founda­tion for naming new species. But that wasn’t the shocker. The more they found at Hayden Quarry, the better search image the crew developed in their heads. They became able to tell a dinosaur from a crocodile from an amphibian almost by instinct. In a series of eureka moments, they realized that most of those supposed dinosaur fossils collected by others weren’t dinosaurs at all, but primitive di- nosauromorph cousins of dinosaurs or, in some cases, early crocodiles and their kin that just so hap­pened to look like dinosaurs.

So not only were dinosaurs rare in the Late Triassic deserts, but they were still living alongside their archaic relatives, the same types of animals that were leaving their tiny footprints in Poland nearly 40 million years earlier. It was a jarring realization. Up until then, almost everyone thought that the primitive dinosauromorphs were an uninteresting ancestral stock whose only destiny was to give birth to the mighty dinosaurs. Once that job was done, they could quietly fade away to extinction. But here they were, all over Late Triassic North America, even a new poodle-size species called Dromo- meron in the Hayden Quarry, living alongside proper dinosaurs for some 20 million years.

Probably the only person not surprised by the findings was another student, an Argentine named Martín Ezcurra. Independently of the four American grad students, Martín was starting to doubt the identifications of some of the supposed North American “dinosaurs” collected by the older generation of paleontologists, but he didn’t have the resources to go study them, because he was from South America and still learning English.

That, and he was a teenager.

One thing he did have, however, was access to the tremendous collections of Ischigualasto di­nosaurs from his home country, thanks to the generosity of Ricardo Martínez and other curators who responded positively to the unusual request of a high schooler wanting to visit their museums. Martín gathered photos of many of the mysterious North American specimens and carefully compared them to the Argentine dinosaurs, and recognized that there were key differences. One North American species in particular, a skinny carnivore called Eucoelophysis, which was supposed to be a theropod, was actually a primitive dinosauromorph. He published this result in a scientific journal in 2006, the year before Irmis, Nesbitt, Smith, and Turner published their first findings. Martín was seventeen years old when he wrote his paper.

It’s hard to fathom why dinosaurs were doing so poorly in the deserts while so many other animals, including their dinosauromorph precursors, were having a better go at it. To get to the bottom of the question, Chinle’s Rat Pack collaborated with the skilled geologist Jessica Whiteside, who was also part of our excavation teams in Portugal. Jessica is a maestro at reading the rocks. Better than any­one I’ve ever known, she can look at a sequence of rocks and tell you how old they are, what the envi­ronments were like when they formed, how hot it was, even how much rain there was. Set her loose at a fossil site, and she’ll come back with a story from the distant past of changing climates, shifting weather, evolutionary explosions, and great extinctions.

Jessica put her sixth sense to use at Ghost Ranch and determined that the animals of the Hayden Quarry did not have an easy life. They lived in an environment that wasn’t always a desert, but one in which seasonal climates dramatically fluctuated. It was bone-dry for much of the year, but wetter and cooler during other times—hyperseasonality, as Jessica and the Rat Pack call it. The culprit was car­bon dioxide. Jessica’s measurements show that there were somewhere around 2,500 molecules of carbon dioxide per every million molecules of air in the tropical regions of Pangea back when the Hay­den Quarry animals were alive. That’s more than six times the amount of carbon dioxide today. Let that sink in for a minute—just think about how quickly temperatures are rising now and how anxious we are about future climate change, even though there is much less carbon dioxide in today’s atmos­phere. The high concentration of carbon dioxide in the Late Triassic started a chain reaction: huge fluctuations in temperature and precipitation, raging wildfires during parts of the year but humid spells in others. Stable plant communities had a difficult time establishing themselves.

It was a chaotic, unpredictable, unstable part of Pangea. Some animals could deal with that better than others. Dinosaurs seem to have been able to cope a little bit, but not able to truly thrive. The smaller meat-eating theropods were able to manage, but the larger, fast-growing plant-eaters, which required a steadier diet, could not. Even some 20 million years after they had originated, even after they had taken over the large-herbivore niche in humid ecosystems and started to colonize the hotter tropics, dinosaurs were still having trouble with the weather.

IF YOU WERE standing on safe ground during a Late Triassic flood, watching the animals eventually buried at Hayden Quarry get swept up by the seasonal river that drowned them, you might have had a hard time telling some of the corpses apart as they floated by. Sure, it would be easy to recognize one of the giant su­persalamanders or some of those weird chameleon-mimic reptiles. But you might not be able to distinguish dinosaurs like Coelophysis and Chindesaurus from some of the crocodiles and their kin. Even if you were able to watch these animals alive, going about their business of eating and moving and interacting with each other, you still might have trouble.

Why the confusion? It’s the same reason that the previous generation of paleontologists working in the American Southwest so often misidentified crocodile fossils as dinosaurs, and why other scien­tists in Europe and South America made the same mistakes. During the Late Triassic, there were many other animals that really, really looked and behaved like dinosaurs. In evolutionary biology speak, this is called convergence: different types of creatures resembling each other because of simi­larities in lifestyle and environment. It’s why birds and bats, which both fly, each have wings. It’s why snakes and worms, which both squirm through underground burrows, are both long, skinny, and leg­less.

The convergence between dinosaurs and crocodiles is surprising, shocking even. The alligators that prowl the Mississippi delta and the crocodiles that lurk in the Nile may appear vaguely prehis­toric, but they don’t look anything like a T. rex or a Brontosaurus. During the Late Triassic, however, crocodiles were very different.

Recall that dinosaurs and crocodiles are both archosaurs—members of that large group of upright­walking reptiles that started to blossom after the end-Permian mass extinction, which proliferated be­cause they could move much faster and more efficiently than the sprawling animals of the time. Early in the Triassic, archosaurs split into two major clans: the avemetatarsalians, which led to dinosauro- morphs and dinosaurs, and the pseudosuchians, which gave rise to crocodiles. During the dizzying splurge of postextinction evolution, the pseudosuchian tribe also produced a number of other sub­groups that diversified in the Triassic but then went extinct. Because they don’t survive today—unlike the crocodiles and dinosaurs (in the guise of birds) —these groups have largely been forgotten about, considered oddities from a distant past, evolutionary dead ends that never rose to the top. That stereotype is wrong, though, because for much of the Triassic these crocodile-line archosaurs were thriving.

Most of the major types of Late Triassic pseudosuchians can be found at Hayden Quarry. There is a phytosaur called Machaeroprosopus, a member of that group of long-snouted, semiaquatic ambush predators whose bones we also found in Portugal. They were bigger than a motorboat and snatched fish—and the occasional passing dinosaur—with the hundreds of spiky teeth in their stretched jaws. It was neighbors with Typothorax, a plant-eater built like a tank with armor covering its body and spikes sticking out from its neck. It belongs to a group called the aetosaurs, a hugely successful family of mid-tier herbivores that closely resembled the armored ankylosaur dinosaurs that evolved millions of years later. They were good diggers and may have even cared for their young by building and guarding nests. Then there are proper crocodiles, but nothing like the ones we’re familiar with today. These primitive Triassic species—the ancestral breed that modern crocs evolved from—looked like grey­hounds: they were about the same size, stood on four legs, had the emaciated build of a supermodel, and could sprint like champions. They fed on bugs and lizards and were most certainly not top preda­tors. That title went to the rauisuchians, a ferocious bunch that grew up to twenty-five feet long, bigger than the largest saltwater crocodiles today. We met one of them previously, Saurosuchus, the top gun in the Ischigualasto ecosystem that would have haunted the nightmares of the very first dinosaurs. Imagine a slightly smaller version of a T. rex walking around on four legs, with a muscular skull and neck, railroad-spike teeth, and a bone-breaking bite.

There’s also another type of crocodile-line archosaur found at Ghost Ranch—not in the Hayden Quarry itself, but in the nearby Coelophysis graveyard. It was found in 1947, not long after Whitaker discovered the bone bed, during those first few weeks of excavation. The American Museum team was digging up so many Coelophysis skeletons that, after a while, the excitement wore off and they got a little bored. Everything they saw started to look like Coelophysis. So they didn’t notice that one of the skeletons they collected was similar in size to Coelophysis, and had the same long legs and light build, but was a little different in other ways—notably, it had a beak instead of an arsenal of sharp teeth. The technicians back in New York didn’t notice either. They started to remove the specimen from the block of rock it was embedded in, but were all too keen to stop once they determined it was just another Coelophysis. It could go in the storehouse with the rest of them.

The fossil stayed in the bowels of the museum, unconserved and unloved, until 2004. That’s when one of the Ghost Ranch quartet, Sterling Nesbitt, started his PhD at Columbia University in New York. Because he was planning a project on Triassic dinosaurs, he went back through all of the fossils col­lected by Colbert, Whitaker, and their teams in the 1940s. Many were still encased in plaster, so they would have to remain on the shelves. But that one block from 1947 had been opened and partially prepared by the conservators, so Sterling could study it. With an excited pair of eyes and an enthusi­asm that escaped the weary field hands a half century earlier, Sterling recognized that he wasn’t look­ing at any old Coelophysis. He saw that it had a beak; he realized that its body proportions were differ­ent, that its arms were tiny. And then he noticed features of the ankle that were nearly identical to those of crocodiles. He wasn’t looking at a dinosaur at all; he was looking at a pseudosuchian that was heavily convergent on dinosaurs.

The fierce predator Batrachotomus, one of the crocodile-line archosaurs (rauisuchians) that preyed on early dinosaurs.

Photo courtesy of the author

This was the sort of discovery that young scientists dream about when secluded, alone with their thoughts, trawling through the drawers of museum collections. Since Sterling discovered it, he got to name it, and he chose the evocative moniker Effigia okeeffeae: the first name being the Latin word for ghost, in reference to Ghost Ranch, and the second paying homage to the ranch’s most famous resi­dent. Effigia made international headlines: the media loved this awkward-looking, toothless, stub­armed ancient crocodilian creature trying to pretend that it was a dinosaur. Stephen Colbert even de­voted a segment of his show to the new discovery, complaining in jest that it should have been named after Edwin Colbert (who coincidentally shared a surname with the comedian) and not the feminist artist. I remember seeing that segment during the last year of my undergraduate studies, right around the time I was starting to plot out my own graduate-school future, and being in awe that a young grad student’s work could make such an impact.

It also motivated me. Up until that point, I had been studying only dinosaurs, but I started to grasp that Effigia and the other dino-imitating pseudosuchians were critical in understanding how dinosaurs ascended to power. I started to read many of the classic studies in dinosaur paleontology, works by gi­ants like Robert Bakker and Alan Charig, which were effusive in arguing that dinosaurs were special. They were so well endowed with superior speed, agility, metabolism, and intelligence that they out- competed all of the other Triassic animals—the giant salamanders, the early mammal-like synapsids, and the crocodile-line pseudosuchians. Dinosaurs were the chosen ones. It was their manifest destiny to take on the weaker species, best them, and establish a global empire. There was almost a religious feel to some of these writings, perhaps not a surprise, given that Bakker also dabbles as an ecumeni­cal Christian preacher and is renowned for his high-energy lectures, delivered in the style of an evan­gelist testifying to his congregation.

Dinosaurs outmaneuvering their foes on the Late Triassic battlefield. It was a good story, but it didn’t sit well with me. New discoveries seemed to be upending the narrative, and a lot of that had to do with the pseudosuchians. So many of these crocodile-line archosaurs were dead ringers for di­nosaurs. Or maybe it was the other way around: maybe Triassic dinosaurs were trying to be pseudo- suchians. Regardless, if the two groups were similar in so many ways, then how could you argue that dinosaurs were a superior race? And it wasn’t only the convergence between dinosaurs and pseudo- suchians that threw up a red flag. There were more pseudosuchians than dinosaurs in the Late Trias­sic: more species and a greater abundance of these species in individual ecosystems. The menagerie of croc cousins from Ghost Ranch—phytosaurs, aetosaurs, rauisuchians, Effigia-like animals, true crocodiles—was not a local phenomenon. These were diverse groups that prospered throughout much of the world.

But, as scientists often like to say when trying to critique each other with subtlety, this all sounded a little arm-wavy. Could we somehow compare explicitly how dinosaurs and pseudosuchians were evolving in the Late Triassic? Was there a way to test whether one group was more successful than the other and whether that was changing over time? I buried myself in literature on statistics, unfamil­iar territory for somebody who was consumed by dinosaurs but not yet very aware of other fields and techniques. I was a bit embarrassed to realize that invertebrate paleontologists—our redheaded stepsiblings, who study fossils like clams and corals, which don’t have bones—had come up with a method two decades earlier, one that had been ignored by dinosaur workers. It was something called morphological disparity.

Morphological disparity sounds like a fancy term, but it is simply a measure of diversity. You can measure diversity in many ways. Counting up the number of species is one tack: you can say that South America is more diverse than Europe because there are more animal species there. Or you can compute diversity based on abundance: insects are more diverse than mammals because there are more insects in any given ecosystem. What morphological disparity does is measure diversity based on features of the anatomy. Thinking this way, you can consider birds to be more diverse than jellyfish, because birds have a much more complex body with lots of different parts, whereas jellyfish are just sacs of goo. This type of diversity measure can give great insight into evolution, because so many as­pects of animal biology, behavior, diet, growth, and metabolism are controlled by anatomy. If you re­ally want to know how a group is changing over time or how two groups compare in diversity, I would argue that morphological disparity is the most powerful way to do so.

Counting the number of species or the abundance of individuals is easy. All you need are a good set of eyes and a calculator. But how to measure morphological disparity? How to take all the com­plexity of the animal body and turn it into a statistic? I followed the approach pioneered by the inverte­brate paleontologists. It went something like this. I first came up with a list of all of the Triassic di­nosaurs and pseudosuchians, as these were the animals that I wanted to compare. I then spent months studying the fossils of these species and made a list of hundreds of features of the skeleton in which they vary. Some have five toes, others have three. Some walk on four legs, others on two. Some have teeth, others do not. I encoded these features in a spreadsheet as zeros and ones, just as a computer programmer would. Herrerasaurus walks on two legs, state 0. Saurosuchus walks on four legs, state 1. At the end of nearly a year of work, I had a database with seventy-six Triassic species, each assessed for 470 features of the skeleton.

With the long slog of data collection done, it was time for the math. The next step was to make what is called a distance matrix. It quantifies how different each species is from every other species, based on the database of anatomical characteristics. If two species share all features, then their dis­tance score is 0. They are identical. If two other species share no characteristics, then their distance score is 1. They are completely different. For the in-between cases, let’s say that Herrerasaurus and Saurosuchus share 100 characteristics but differ in the 370 others. Their distance score would be 0.79: the 370 features they differ in divided by the 470 total features in the data set. The best way to envision this is to think of those tables in a road atlas, which give distances between different cities. Chicago is 180 miles from Indianapolis. Indianapolis is 1,700 miles from Phoenix. Phoenix is 1,800 miles from Chicago. That table is a distance matrix.

Here’s the neat trick about a distance matrix in an atlas. You can take that table of road distances between cities, stick it into a statistics software program, run what is called a multivariate analysis, and the program will spit out a plot. Each city will be a point on that plot, and the points will be sepa­rated by distance, in perfect proportion. In other words, the plot is a map—a geographically correct map with all of the cities in the right places and distances relative to each other. So what happens if we instead input the distance matrix that encapsulates the skeletal differences of Triassic dinosaurs and pseudosuchians? The statistics program will also produce a plot in which each species is repre­sented by a point, a plot that scientists call a morphospace. But really it is just a map. It visually shows the spread of anatomical diversity among the animals in question. Two species close together have very similar skeletons, just as Chicago and Indianapolis are comparatively near geographically. Two species at far corners of the graph have very different anatomies, like the longer distance between Chicago and Phoenix.

This map of Triassic dinosaurs and pseudosuchians allows us to measure morphological disparity. We can group the animals in the plot by which great tribe they belong to—dinosaurs or pseudosuchi- ans—and calculate which is occupying a larger swath of that map and therefore more anatomically di­verse. In the same vein, we can further group the animals by time—Middle Triassic versus Late Trias­sic, let’s say—and see if dinosaurs or pseudosuchians were becoming more or less anatomically di­verse as the Triassic progressed. We did that and came up with a startling result that we published in 2008 in a study that helped launch my career. All throughout the Triassic, the pseudosuchians were significantly more morphologically diverse than dinosaurs. They filled a larger spread of that map, meaning they had a greater range of anatomical features, which indicated that they were experiment­ing with more diets, more behaviors, more ways of making a living. Both groups were becoming more diverse as the Triassic unfolded, but the pseudosuchians were always outpacing the dinosaurs. Far from being superior warriors slaying their competitors, dinosaurs were being overshadowed by their crocodile-line rivals during the 30 million years they coexisted in the Triassic.

PUT YOURSELF BACK in the tiny furry feet of our Triassic mammalian ancestors, surveying the Pangean scene as the Triassic was drawing to a close 201 mil­lion years ago. You would be seeing dinosaurs, but you wouldn’t be surrounded by them. Depending on where you were, you might not have noticed them at all. They were relatively diverse in the humid regions, where protosauropods got as large as giraffes and were the most abundant plant-eaters, but there carnivorous theropods and herbivorous to omnivorous ornithischians were con­siderably smaller and less common. In the more arid zones, there were only small meat-eaters, the herbivores and larger species being unable to tolerate the hyperseasonal weather and megamonsoons. There were no dinosaurs re­motely approaching a Brontosaurus or T. rex in size, and all across the super­continent they were living under the thumb of their much more diverse, much more successful pseudosuchian adversaries. You would probably consider the dinosaurs a fairly marginal group. They were doing OK, but so were many other newly evolved types of animals. If you were of a gambling persuasion, you would probably have bet on one of these other groups, most likely those pesky crocodile-line archosaurs, as the ones that would eventually become dominant, grow to massive sizes, and conquer the world.

Some 30 million years after they originated, the dinosaurs had yet to mount a global revolution.


Dinosaurs Become Dominant

Scottish sauropod

Chapter Title art by Todd Marshall

SOME TIME AROUND 240 MILLION YEARS ago, the Earth began to crack. True dinosaurs hadn’t quite evolved yet, but their cat-size dinosauromorph ancestors would have been there to experience the cracking—except there wasn’t much to expe­rience, at least not yet. There may have been some minor earthquakes, but these probably wouldn’t have even registered with the dinosauromorphs, who were busy with more important things like fending off the supersalamanders and surviving the megamonsoons. As these dinosauromorphs gave rise to di­nosaurs, the fracturing continued, many thousands of feet underground. Imper­ceptible on the surface, these fissures were slowly moving, growing, merging to­gether, a hidden danger lurking under the feet of Herrerasaurus, Eoraptor, and the other first dinosaurs.

The very foundation of Pangea was splitting, and with the blissful ignorance of homeowners who don’t realize there’s a creeping crack in their basement until their house comes tumbling down, the di­nosaurs had no inkling that their world was going to dramatically change.

As these earliest dinosaurs were evolving in fits and starts during the final 30 million years of the Triassic, great geological forces were tugging on Pangea from both the east and west. These forces—a planet-scale cocktail of gravity, heat, and pressure—are strong enough to make continents move over time. Because the pull was coming from two opposite directions, Pangea began to stretch and gradu­ally become thinner, each small earthquake causing another tear. Imagine Pangea as a giant pizza, being torn apart by two hungry friends at opposite ends of the table: the crust becomes thinner until there is a rupture and it breaks into two. The same thing happened with the supercontinent. After a few tens of millions of years of slow and steady tug-of-war, east versus west, the cracks reached the surface, and the giant landmass began to unzip down its middle.

It’s because of that ancient divorce between east and west Pangea that the seaboard of North America is separated from western Europe and South America sits apart from Africa. It’s why there is now an Atlantic Ocean, which didn’t exist until seawater rushed in to fill the gap between the separat­ing tracts of land. Those forces and fractures over 200 million years ago shaped our modern geogra­phy. But there was more to it than that, because continents don’t just split up and call it a day. As with human relationships, things can get really nasty when a continent breaks up. And the dinosaurs and other animals growing up on Pangea were about to be changed forever by the aftereffects of their home being ripped in two.

The problem boils down to this: as a continent tears, it bleeds lava. It’s nothing more than basic physics. The Earth’s outer crust is pulled apart and thins, decreasing pressure on the deeper parts of the Earth. As pressure lessens, magma from the deeper Earth rises to the surface and erupts through volcanoes. If there is only a little rip in the crust—two small bits of a continent separating from each other, let’s say—then the effects aren’t too bad. You might get a few volcanoes, some lava and ash, some local destruction, and then eventually it stops. That kind of thing is happening in eastern Africa today, and it’s far from catastrophic. But if you’re slashing apart an entire supercontinent, then you approach the realm of apocalypse.

At the very end of the Triassic, 201 million years ago, the world was violently remade. For 40 mil­lion years, Pangea had been gradually splintering apart, and magma had been welling underground. Now that the supercontinent had finally cracked, the magma had somewhere to go. Like a hot-air bal­loon rising through the sky, the liquid-rock reservoir rushed upward, broke through the shattered sur­face of Pangea, and gushed out onto the land. As with the volcanoes that had erupted at the end of the Permian Period some 50 million years earlier, causing the extinction that allowed dinosaurs and their archosaur cousins to get their start, these end-Triassic eruptions were different from any that hu­mans have ever witnessed. We’re not talking Pinatubo here, with hot clouds of ash bursting into the sky. Instead, over a period of some six hundred thousand years, there were four big pulses of drama, when enormous amounts of lava would surge out of the Pangean rift zone like tsunamis from hell. I’m hardly exaggerating: some of the flows were, added up together, up to three thousand feet thick; they could have buried the Empire State Building twice over. In all, some three million square miles of cen­tral Pangea were drowned in lava.

It goes without saying that this was a bad time to be a dinosaur, or for that matter, any other type of animal. These were some of the largest volcanic eruptions in Earth history. Not only did lava smother the land, but noxious gases that rode up with the lava poisoned the atmosphere and caused runaway global warming. These things triggered one of the biggest mass extinctions in the history of life, a mass die-off that claimed over 30 percent of all species and maybe much more. Paradoxically, however, it was also a mass extinction that helped dinosaurs break out of their early-life slump and become the enormous, dominant animals that stoke our imaginations.

IF YOU’RE WALKING down Broadway in New York City and happen to catch a gap between the skyscrapers, you can see straight across the Hudson River to New Jersey. You’ll notice that the Jersey side of the river is defined by a steep cliff of drab brown rock, about a hundred feet high, studded with vertical cracks. Lo­cals refer to it as the Palisades. During the summer it can be almost unrecog­nizable, engulfed by a dense forest of trees and bushes that somehow cling to the sheer slopes. Commuter towns like Jersey City and Fort Lee are perched on top of the cliff, and the western end of the George Washington Bridge is built deep into it, an ideal anchor for the world’s busiest overwater crossing. If you wanted to, you could walk along the Palisades for about fifty miles, from where it begins in Staten Island and extends along the Hudson to where it juts into up­state New York.

Millions of people look at this cliff every week. Hundreds of thousands of people live on it. Few re­alize that it is a remnant of those ancient volcanic eruptions that tore apart Pangea and ushered in the Age of Dinosaurs.

The Palisades is what geologists refer to as a sill—an intrusion of magma that pokes its way in be­tween two layers of rock far underground, but then hardens into stone before it can erupt as lava. Sills are part of the internal plumbing system of volcanoes. Before they harden into rock, they are pipes, which transport magma underground. Sometimes they are conduits that bring magma to the surface; other times they are dead-end extensions of the volcanic system, cul-de-sacs that magma can’t es­cape from. The Palisades sill formed at the end of the Triassic, as Pangea was rupturing along what would become the eastern coast of North America, just a few miles from what is now New York City. It formed from those very magmas that were coursing up from the deep earth as the supercontinent broke into two.

The magma that became the Palisades sill never made it to the surface. It never got to be part of those three-thousand-foot-thick lava sheets flushed out of the Pangean rift, the ones that engulfed ecosystems and belched out the carbon dioxide that would doom much of the planet. About twenty miles to the west the magmas did erupt, however, and the basalt rocks that formed from them can be seen in a low range of hills called the Watchung Mountains in northern New Jersey. Calling them mountains is generous—they’re just a few hundred feet high, and they cover a tiny area about forty miles north to south—but they are a beloved oasis of natural beauty within one of the most urbanized parts of the world.

In the middle of the mountains is Livingston, a bedroom community of about thirty thousand peo­ple. In 1968 some folks discovered dinosaur footprints a couple of miles north of the town, in an abandoned quarry where red shales, formed in rivers and lakes near the ancient volcanoes, were being mined. There was a blurb in the local newspaper, which caught the eye of a mother, who told her fourteen-year-old-son, Paul Olsen, who was gobsmacked to learn that dinosaurs once lived so close to his home. He rounded up his friend, Tony Lessa, and they hopped on their bicycles and sped to the old quarry. It was no more than an overgrown, rock-strewn hole in the ground, but the discovery had caused a local sensation and several amateur collectors were already there, on the hunt for more tracks. Olsen and Lessa befriended some of the amateurs, who taught them the basics of fossil col­lecting: how to identify dinosaur footprints, how to remove them from the rock, and how to study them.

The two teenagers became obsessed. They kept coming back to the quarry, and before long they were working late into the night, removing slabs of dinosaur footprints by firelight, even in the dead of winter. They had to go to school during the day, so the night was their only option. For over a year they toiled, outlasting the other rockhounds, who began to trickle away once the excitement of the discov­ery died down. The boys collected hundreds of tracks left by all kinds of creatures, including meat-eat­ing dinosaurs similar to Coelophysis from Ghost Ranch, plant-eating dinosaurs, and some of the scaly and furry creatures that lived alongside. But the more they collected, the more they became dis­mayed: during their nighttime excavations, they were constantly interrupted by trucks illegally dump­ing trash, and while they were at school, unscrupulous collectors would often sneak into the quarry and poach footprints the boys hadn’t yet been able to remove.

So what’s a 1960s teenager to do when his favorite fossil site is being destroyed? Paul Olsen skipped the middlemen and went right to the top. He began writing letters to Richard Nixon, the newly elected president who had yet to disgrace himself. Lots of letters. He begged Nixon to use his presi­dential powers to get the quarry preserved as a protected park, and even sent a fiberglass cast of a theropod track to the White House. Olsen led a media campaign, too, and was profiled in an article in Life magazine. His brazen persistence paid off: in 1970 the company that owned the quarry donated the land to the county, which made it into a dinosaur park called the Riker Hill Fossil Site. The next year, the site was granted official national landmark status and Olsen received a presidential com­mendation for his work. Little did he know it, but he was also an inch away from a White House visit. Some of Nixon’s image-conscious aides thought a photo op with a young science enthusiast would be great PR for the jowly president, but it was killed at the last minute by Nixon’s advisor John Ehrlich- man, later one of the key villains of Watergate.

It was a great accomplishment for a kid—collecting a haul of dinosaur tracks, getting his site pre­served for posterity, becoming pen pals with the president. But Paul Olsen didn’t stop. He went to col­lege to study geology and paleontology, completed a PhD at Yale, and was hired as a professor at Co­lumbia University, across the Hudson from Riker Hill. He became one of the leading academic paleon­tologists in the world and was elected to the National Academy of Sciences, one of the greatest hon­ors for any American scientist. He also had the burden of being a member of my PhD committee, a far lesser honor, when I did my doctorate, in New York. During that time, he became one of my most trusted mentors, a brilliant sounding board for whatever crazy research ideas I had. For two years, I assisted him as he taught his popular undergraduate course on dinosaurs at Columbia, always over­subscribed by nonmajor students, who were seduced by the eminent scientist with a white Geraldo moustache prancing around with the enthusiasm that comes from several preclass energy drinks. Much of my ebullient, wildly animated lecturing style comes from watching Paul.

Paul Olsen made his career by continuing what he started as a teenager. Much of his work has fo­cused on those events that were occurring around the time dinosaurs were leaving footprints in New Jersey: the breakup of Pangea at the very end of the Triassic, the unimaginable volcanic eruptions, the mass extinction, and the rise of dinosaurs to global dominance as the Triassic transitioned into the subsequent Jurassic Period.

Although he had no idea when he first cycled up to that quarry as a kid, Paul grew up in the best place in the world for studying the Late Triassic and Early Jurassic. His boyhood stomping grounds are within a geological structure called the Newark Basin, a bowl-like depression filled with Triassic and Jurassic rocks. It is one of many such structures—called rift basins, because they formed as Pangea rifted apart—extending for over a thousand miles down the eastern coast of North America. The Bay of Fundy, up north in Canada, laps onto one of these basins. Farther south is the Hartford Basin, which cuts through much of central Connecticut and Massachusetts. Then the Newark Basin, followed by the Gettysburg Basin, site of the famous Civil War battle, the topography of the rocks so instrumental in shaping military strategy that depended on securing bits of high ground. South of Gettysburg are many smaller basins that pepper the backcountry of Virginia and North Carolina, finally culminating in the huge Deep River Basin of the Carolina interior.

These rift basins follow the fracture between east and west Pangea. They are the dividing line, the frontier, the place where the supercontinent tore up. As those east-west tugging forces started to pull Pangea apart, faults formed deep within the crust, cutting through what used to be solid rock. Each bit of tugging would cause an earthquake, which would cause the rocks on either side of the fault to move a little bit relative to each other. Over millions of years the faults reached the surface, and as one side continued to fall, a basin was formed: a depression on the downward side of the fault rimmed by a high mountain range on the upward side. Each of the eastern North American rift basins formed this way, the result of more than 30 million years of pressure, tension, and tremors.

This is exactly what is occurring in eastern Africa today, as Africa is pulling away from the Middle East at the rate of about one centimeter per year. The two landmasses used to be connected about 35 million years ago, but now they are separated by the long and skinny Red Sea, which continues to get wider year by year and will one day turn into an ocean. To the south, on the African mainland, there is a north-south band of basins, each growing wider and deeper with every earthquake that is yanking Africa and Arabia farther apart. Some of the deepest lakes in the world, like the nearly mile- deep Lake Tanganyika, fill some of these basins. Others are crisscrossed by raging rivers, which rush down from the mountains above, irrigating great tropical ecosystems lush with some of Africa’s most familiar plants and animals. Sprinkled throughout, poking up in random places, are volcanoes like Mount Kilimanjaro, escape valves for the magma building up underground as the land fractures. Oc­casionally one of these goes off and buries the basins, and their inhabitants, in lava and ash.

Paul Olsen’s Newark Basin, and the many others lining the eastern coast of North America, under­went a similar process of evolution. They were formed gradually by earthquakes, were flushed with rivers that supported diverse ecosystems, eventually became so deep and full of water that the rivers turned into lakes, and then, depending on the quirks of climate, the lakes would dry up, rivers would form again, and the whole process would start over. Cycle after cycle after cycle. Dinosaurs, pseudo- suchian cousins of crocodiles, supersalamanders, and early relatives of mammals thrived along the rivers’ edges, and blooms of fish choked the lakes. These animals left their fossils—the footprints Paul Olsen started to collect as a teenager, as well as bones—in the thousands of feet of sandstones, mud­stones, and other rocks deposited by the rivers and lakes. And then, when Pangea had been stretched to its limit, the crust burst and volcanoes started to erupt, burying the basins and the creatures that lived within them.

The first eruptions didn’t occur in the Newark Basin area. They happened in what is now Morocco, which at that time was nudged up against what would become eastern North America, just a few hun­dred miles or so from modern New York City. Then lava began pouring out in other places where Pangea was splitting: in the Newark Basin, in what is now Brazil, in those same lake environments where we found the supersalamander graveyard in Portugal—all along that zipper line, which, many millions of years later, would transform into the Atlantic Ocean. The lava came in four waves, each scorching the once verdant rift basins, each spreading toxic fumes all over the planet, each making a bad situation worse and worse. In only about half a million years—a blink of an eye in geological terms —the eruptions stopped, but they transformed the Earth forever.

The dinosaurs, pseudosuchian crocodile-line archosaurs, big amphibians, and early mammal rela­tives living in the rift basins were blissfully unaware of what was about to happen. Things went sour quickly.

The initial eruptions in Morocco released clouds of carbon dioxide, a powerful greenhouse gas, which rapidly warmed the planet. It got so hot that strange ice formations buried within the seafloor, called clathrates, melted in unison all throughout the world’s oceans. Clathrates are unlike the solid blocks of ice we’re used to, the ones we put in our drinks or carve into fancy sculptures at parties. They are a more porous substance, a latticework of frozen water molecules that can trap other sub­stances inside it. One of those substances is methane, a gas that seeps up constantly from the deep Earth and infiltrates the oceans but is caged in the clathrates before it can leak into the atmosphere. Methane is nasty: it’s an even more powerful greenhouse gas than carbon dioxide, packing an earth­warming punch over thirty-five times as great. So when that first torrent of volcanic carbon dioxide in­creased global temperatures and melted the clathrates, all of that once-trapped methane was sud­denly released. This initiated a runaway train of global warming. The amount of greenhouse gas in the atmosphere approximately tripled within a few tens of thousands of years, and temperatures in­creased by 3 or 4 degrees Celsius.

Ecosystems on land and in the oceans couldn’t cope with such rapid change. The much hotter temperatures made it impossible for many plants to grow, and indeed upwards of 95 percent of them went extinct. Animals that fed on the plants found themselves without food, and many reptiles, am­phibians, and early mammal relatives died out, like dominoes falling up the food chain. Chemical chain reactions made the ocean more acidic, decimating the shelly organisms and collapsing food webs. Climate became dangerously variable, with episodes of intense heat followed by cooler periods. This enhanced the temperature differences between northern and southern Pangea, causing the meg­amonsoons to become more severe, the coastal regions to become even wetter, and the continental interiors to grow much drier. Pangea had never been a particularly hospitable place, but those early di­nosaurs that already were constrained by the monsoons, the deserts, and their pseudosuchian rivals were now in even worse shape.

So how did these dinosaurs, still at such a relatively young stage in their evolution, deal with a world that was changing so quickly? The clues are in the footprints that Paul Olsen has been studying now for nearly fifty years. The quarry that Paul explored in New Jersey is one of more than seventy places where dinosaur footprints have been found along the eastern seaboard of the US and Canada. These sites are positioned one on top of the other, in geological sequence, stretching over 30 million years, from around the time the first dinosaurs were originating in what is now South America (but still absent in modern-day North America), through the Late Triassic, across the volcanic extinction, and into the ensuing Jurassic Period. Generations of dinosaurs and other animals left their traces in those cyclical beds of sandstones and mudstones deposited in the rift basins, and by studying them in suc­cession, you can see how these creatures were evolving.

The rocks tell a remarkable story. During the Late Triassic, beginning about 225 million years ago, when the rift basins were just beginning to form, dinosaurs started to leave their marks in the form of rare footprints. There are three-toed tracks called Grallator, ranging from about two to six inches (five to fifteen centimeters) long, made by small, fast-running, meat-eating dinosaurs that stood on two legs like the Ghost Ranch Coelophysis. There’s a second type of track called Atreipus, which are about the same size as Grallator but include small handprints next to the three-toed footprints, a sign that the trackmaker was walking on all fours. They were probably made by primitive ornithischian di- nosaurs—the oldest cousins of Triceratops and the duck-billed dinosaurs—or perhaps by close di- nosauromorph cousins to dinosaurs. These dinosaur tracks are vastly outnumbered by the prints left by pseudosuchians, large amphibians, proto-mammals, and small lizards. Dinosaurs were there, but they still remained role players in the rift basin ecosystems, right up until the end of the Triassic.

But then the volcanoes kicked into gear. Suddenly the diversity of non-dinosaur tracks drops dra­matically in those first Jurassic rocks above the lava flows. Many non-dinosaur tracks abruptly disap­pear, including some of the most conspicuous prints left by crocodile-cousin pseudosuchians, which had previously been more abundant and diverse than the dinosaurs. Whereas dinosaurs made up only about 20 percent of all tracks before the volcanoes, right afterward half of all footprints belong to dinosaurs. A variety of totally new dinosaur tracks enter the record: a handprint-footprint duo called Anomoepus probably made by an ornithischian, a large four-toed print called Otozoum made by the very first long-necked proto-sauropods to live in the rift valleys, and a three-toed track called Eu- brontes that belonged to another type of swift predator. These Eubrontes tracks are a little over a foot long (about thirty-five centimeters), a big size increase over the Grallator prints left by similar but much smaller carnivores during the prevolcano days of the Triassic.

It’s probably not what you were expecting. After some of the largest volcanic eruptions in Earth his­tory desecrated ecosystems, dinosaurs became more diverse, more abundant, and larger. Completely new dinosaur species were evolving and spreading into new environments, while other groups of ani­mals went extinct. As the world was going to hell, dinosaurs were thriving, somehow taking advantage of the chaos around them.

When the volcanoes ran out of lava and their six-hundred-thousand-year reign of terror was over, the world was a very different place than it had been in the Late Triassic. It was much warmer, storms were more intense, and wildfires ignited with ease; new types of ferns and ginkgos had replaced the once abundant broadleaf conifers; and many of the most charismatic Triassic animals were gone. The piglike mammal-relative dicynodonts and the beaked plant-munching rhynchosaurs were both extinct; the supersalamander amphibians, almost completely knocked out. What about the pseudosuchians, those crocodile-line archosaurs that were overshadowing, outmuscling, and seemingly outcompeting the dinosaurs during the final 30 million years of the Triassic? Nearly every species bit the dust. The long-snouted phytosaurs, the tanklike aetosaurs, the apex-predator rauisuchians, and the weird Effi- gia-like critters that resembled dinosaurs—none of them were ever to be heard from again. The only pseudosuchians that made it through the great Pangean breakup were a few types of primitive croco­diles, a handful of battle-worn stragglers that would eventually evolve into the modern alligators and crocodiles but would never enjoy the same success they had in the Late Triassic, when they seemed primed to take over the world.

Somehow dinosaurs were the victors. They endured the Pangean split, the volcanism, and the wild climate swings and fires that vanquished their rivals. I wish I had a good answer for why. It’s a mystery that quite literally has kept me up at night. Was there something special about dinosaurs that gave them an edge over the pseudosuchians and other animals that went extinct? Did they grow faster, re­produce quicker, have a higher metabolism, or move more efficiently? Did they have better ways of breathing, hiding, or insulating their bodies during extreme heat and cold snaps? Maybe, but the fact that so many dinosaurs and pseudosuchians looked and behaved so similarly makes such ideas tenu­ous at best. Maybe dinosaurs were just lucky. Perhaps the normal rules of evolution are ripped up when such a sudden, devastating, global catastrophe happens. It could be that the dinosaurs simply were the ones that walked away from the plane crash unscathed, saved by good fortune, when so many others died.

Whatever the answer, it’s a riddle waiting for the next generation of paleontologists to figure out.

THE JURASSIC PERIOD marks the beginning of the Age of Dinosaurs proper. Yes, the first true dinosaurs entered the scene at least 30 million years before the Jurassic began. But as we’ve seen, these earlier Triassic dinosaurs had not even a remote claim to being dominant. Then Pangea began to split, and the dinosaurs emerged from the ashes and found themselves with a new, much emptier world, which they proceeded to conquer. Over the first few tens of mil­lions of years of the Jurassic, dinosaurs diversified into a dizzying array of new species. Entirely new subgroups originated, some of which would persist for an­other 130-plus million years. They got larger and spread around the globe, colo­nizing humid areas, deserts, and everything in between. By the middle part of the Jurassic, the major types of dinosaurs could be found all over the world. That quintessential image, so often repeated in museum exhibits and kids’ books, was real life: dinosaurs thundering across the land, at the top of the food chain, ferocious meat-eaters comingling with long-necked giants and ar­mored and plated plant-eaters, the little mammals and lizards and frogs and other non-dinosaurs cowering in fear.

Here are some of the familiar dinosaurs that start to show up after the Pangean rift volcanoes ush­ered in the Jurassic. There were meat-eating theropods like Dilophosaurus, with a weird double-mo­hawk crest on its skull; at around twenty feet long, it was much larger than the mule-size Coelophysis and most other Triassic carnivores. Plant-eating ornithischians covered in armor plates, like Sceli- dosaurus and Scutellosaurus, would soon after give rise to the familiar tanklike ankylosaurs and back- plated stegosaurs. Small, fast-moving, probably omnivorous ornithischians like Heterodontosaurus and Lesothosaurus, were early members of that lineage that would eventually produce the horned and duck-billed dinosaurs. Other familiar dinosaurs that had been around in the Triassic but restricted to only a few environments, like the long-necked proto-sauropods and the most primitive ornithischi- ans, finally began to migrate around the planet.

Nothing in this inventory of growing diversity encapsulates the newfound dominance of dinosaurs quite like the sauropods. They are those unmistakable long-necked, column-limbed, potbellied, plant­devouring, small-brained behemoths. Some of the most famous dinosaurs of all are sauropods: Bron­tosaurus, Brachiosaurus, Diplodocus. They show up in almost all museum exhibits and are stars of Jurassic Park; Fred Flintstone used one to mine slate, and a green cartoon sauropod has been the logo of Standard Oil for decades. Along with T. rex, they are the iconic dinosaurs.

Sauropods evolved from an ancestral stock, what I’ve been calling the proto-sauropods, in the lat­est Triassic. These protospecies were the dog-to-giraffe-size plant-eaters with fairly long necks that were among that first wave of dinosaurs to appear in Ischigualasto about 230 million years ago. They then became the main herbivores in the humid parts of Triassic Pangea but were kept from achieving their full potential by their inability to settle the deserts. That changed in the early part of the Jurassic, when sauropods were able to break free of their environmental restrictions and move about the globe, evolving their characteristic noodle-necked bodies and growing to monstrous sizes in the process.

The skull of Plateosaurus, one of the proto-sauropods, the ancestral stock that gave rise to the


Photo courtesy of the author

Fossils of some of the first truly gigantic sauropods—ones that weighed over ten tons, were over fifty feet long, and had necks that could stretch several stories into the sky—have started turning up in Scotland over the past few decades, on a beautiful island off the west coast called the Isle of Skye. The clues have been meager—a stocky limb bone here, a tooth or a tail vertebra there—but they hint at an animal of enormous size living about 170 million years ago, far enough into the Jurassic that the Pangean split and volcanic apocalypse were distant memories but still during that time when di­nosaurs were putting the final flourishes on their rise to dominance.

The sauropod fossils from Skye piqued my interest when I moved to Scotland in 2013 to take up my new position at the University of Edinburgh, fresh out of my PhD in New York and bounding with the excitement of starting my own research lab. During my first few weeks on the job, I began to hang out with two scientists in my department: Mark Wilkinson, a hardened field geologist whose ponytail and scruffy beard give him the look of a hippie, and Tom Challands, a redheaded packhorse of man who also had a doctorate in paleontology, albeit on microscopic fossils from over 400 million years ago. Tom had recently finished a stint in the real world, putting his geological skills to use working for an energy company on the search for oil. For part of that time, he lived in a custom-made camper van, fitted with a bed and small kitchen, which he would park near whatever sites he was surveying. His new bride put the kibosh on that lifestyle after their wedding, but the van still came in handy for field­work travel, and Tom would often spend his weekends driving along the misty coasts of Scotland look­ing for whatever fossils he could find. Both Tom and Mark had done some geological work on Skye and knew the terrain well, so we made a pact to hunt for better fossils of the mysterious giant sauropods.

The more we read about Skye, the more one name kept popping up: Dugald Ross. It was a name I wasn’t familiar with. He wasn’t a paleontologist or a geologist or a scientist of any kind. Yet he had dis­covered and described many of the dinosaur fossils found on Skye. Dugald was a local boy who grew up in the tiny hamlet of Ellishadder on the far northeastern arm of the island, a rugged landscape of craggy peaks, green hills, peat-colored streams, and windswept shores that looks like something out of a fantasy novel—very Tolkienesque. He was raised in a household that spoke Gaelic, the native lan­guage of the Scottish Highlands, which is spoken by only about fifty thousand people today but which still has a presence on the road signs and in the schools on remote islands like Skye. When Dugald was fifteen years old, he found a cache of arrow points and Bronze Age artifacts near his family’s home, and this sparked an obsession with the history of his native island that continued into adult­hood, as he carved out a career as a builder and crofter (a Scottish Highlands term for a small-scale farmer and sheepherder).

I got in touch with Dugald and told him about our dreams of finding huge dinosaurs on his island. It was one of the most fortunate e-mails I’ve ever sent, because it struck up a friendship and a re­markable scientific collaboration. Dugald—or Dugie, as he prefers to be called—invited us to visit him when we came up to his island a few months later. He instructed us to drive up the main two-lane road that snakes along the coast of northeastern Skye and meet him at a long ranch-style building, made up of a collage of different-size gray stones and a black tile roof, with antique farming instru­ments strewn across the lawn outside. There was a sign out front that said TAIGH-TASGAIDH—the Gaelic word for museum. Dugie emerged from his big red work van with a set of oversize skeleton keys, made his introductions, and proudly led us inside. In his soft-spoken lyrical accent—a charming combi­nation of Sean Connery-style Scots and an Irish brogue—he explained how he had taken the ruins of a one-room schoolhouse and built the structure we were standing in, the Staffin Museum. He founded the museum when he was nineteen. Today, this single room—without a café, a big gift shop, or other expensive trappings of big-city museums, or even electricity—contains many of the dinosaurs he’s found on Skye, along with artifacts that trace the history of the island’s human inhabitants. It’s a sur­real experience: big dinosaur bones and footprints displayed next to old mill wheels, iron rods for pick­ing turnips, and antique mole traps once used by Highland farmers.


The enchanting landscape of the Isle of Skye, Scotland.

Photo courtesy of the author

For the rest of that week, Dugie led us to many of his favorite hunting spots. We found a lot of

Jurassic fossils—the jaw of a dog-size crocodile, the teeth and backbones of reptiles called
ichthyosaurs, which resembled dolphins and lived in the oceans when dinosaurs started to dominate the land—but no giant sauropods. Over the next few years, we kept coming back.
Dugie Ross removing a dinosaur bone from a boulder on the Isle of Skye. Photo courtesy of the author







































The dinosaur dance floor of sauropod tracks that I discovered with Tom Challands on the Isle of


Photo courtesy of the author

Finally, in the spring of 2015, we found what we set out for, although we didn’t even realize it at first. We spent most of the day on our hands and knees, looking for tiny fish teeth and scales embed­ded in a platform of Jurassic rocks that stretched into the icy waters of the North Atlantic, right below the ruins of a fourteenth-century castle. This was Tom’s idea: he now studies fossil fish, and in ex­change for his help finding dinosaurs, I promised to assist him in collecting fishy bits. We had been squinting at the rocks for hours, bundled up in three layers of waterproof clothing but still freezing. The tide was coming in, the late afternoon light was going down, and dinner was beckoning. So Tom and I packed up our gear and our bags of fish teeth and started to stroll back to his tricked-out van parked on the other side of the beach. That’s when something caught our eyes. It was a malformed depression in the rock, about the size of a car tire. We had missed it earlier because our eyes were fo­cused on the much smaller fish bones, our search image totally unsuitable for noticing something so big.

As we continued to walk, we started to notice many other similar depressions, now visible in the low-angle afternoon light. They were all about the same size, and the closer we looked, the more we saw that they stretched in every direction around us. They seemed to show a pattern. Individual holes were lined up in two long rows, in something of a zigzag arrangement: left-right, left-right, left-right. Ribbons of them were crisscrossing much of the rock platform that we had been working on all day.

Tom and I looked at each other. It was the kind of knowing glance between brothers, a nonverbal connection based on years of shared experience. We had seen these types of things before, not in Scotland, but in places like Spain and western North America. We knew what they were.

The holes in front of us were fossilized tracks, huge ones. Dinosaur tracks, no doubt. As we looked closer, we could see that there were both handprints and footprints, and some of them had finger and toe marks. They had the telltale shape of tracks left by sauropods. We had found a 170-million-year- old dinosaur dance floor, records left by colossal sauropods that were about fifty feet long and weighed as much as three elephants.

The tracks were made in an ancient lagoon, an environment not commonly associated with sauropods. We usually envision these monstrous dinosaurs stampeding across the land, causing a small earthquake with each step. And they did. But by the middle part of the Jurassic, the sauropods had become so diverse that they started branching out into other ecosystems, always searching for the vast quantities of leafy food needed to fuel their giant bodies. Our trackway site in Skye has at least three different layers of footprints, made by different generations of sauropods wading through a salty lagoon, living with smaller plant-eating dinosaurs, the occasional pickup-truck-size carnivore, and many types of crocodiles, lizards, and swimming mammals with flat tails like beavers. Scotland was much warmer back then, a land of swamps and sandy beaches and rolling rivers on an island in the middle of the growing Atlantic Ocean, perched between North American and European landmasses that moved farther and farther apart as Pangea continued to split. Thoroughly ruling this land were the sauropods and other dinosaurs, which had now—finally—become a global phenomenon.

THERE’S REALLY NO better way to say it: the sauropods that made their marks in that ancient Scottish lagoon were awesome creatures. Awesome in the literal sense of the word—impressive, daunting, inspiring awe. If I was handed a blank sheet of paper and a pen and told to create a mythical beast, my imagination could never match what evolution created in sauropods. But they were real: they were born, they grew, they moved and ate and breathed, they hid from predators, they slept, they left footprints, they died. And there’s absolutely noth­ing like sauropods around today—no animals with a similar long-necked and swollen-gut body type, no creatures on land that even remotely approach them in size.

Sauropods are so mind-twistingly big that, when their first fossil bones were discovered in the 1820s, scientists found themselves in a bind. Some of the first dinosaurs were being found around the same time, like the meat-eating Megalosaurus and the beaked herbivore Iguanodon. These were big animals, no doubt, but nowhere near the size of the creatures that left the gigantic sauropod bones. So scientists didn’t make the connection with dinosaurs. Instead, they considered the sauro- pod bones to belong to the one type of thing they knew could get so huge: whales. It was a few decades before that mistake was corrected. Amazingly, later discoveries would show that many sauropods got even bigger than most whales. They were the largest animals that ever walked the land, and they push the limit for what evolution can achieve.

This raises a question that has fascinated paleontologists for over a century: how did sauropods become so large?

It’s one of the great puzzles of paleontology. But before trying to solve it, we first need to come to grips with a more fundamental issue: how big did sauropods get? How long were they, how high could they stretch their necks, and most important, how much did they weigh? These turn out to be difficult questions to answer, particularly when it comes to weight, because you can’t just stick a dinosaur on a scale and weigh it. A trade secret among paleontologists is that many of the fantastical numbers you see in books and museum exhibits—Brontosaurus weighed a hundred tons and was bigger than a plane!—are pretty much just made up. Educated guesses or, in some cases, barely that. Recently, however, paleontologists have come up with two different approaches to more accurately predict the weight of a dinosaur based on its fossil bones.

The first is really quite simple and relies on basic physics: heavier animals require stronger limb bones to support their weight. This logical principle is reflected in how animals are built. Scientists have measured the limb bones of many living animals, and it turns out that the thickness of the main bone in each limb that supports the animal—the femur (thighbone) for those that walk on two legs only or the femur plus the humerus (upper arm bone) for those that stand on all fours—is strongly sta­tistically correlated with the weight of the animal. In other words, there is a basic equation that works for almost all living animals: if you can measure limb-bone thickness, you can then calculate body weight with a small but recognized margin of error—simple algebra you can do with a basic calculator.

The second method is more intensive but a lot more interesting. Scientists are starting to build three-dimensional digital models of dinosaur skeletons, add on the skin and muscles and internal or­gans in animation software, and use computer programs to calculate body weight. It’s a method pio­neered by a number of young British paleontologists—Karl Bates, Charlotte Brassey, Peter Falking- ham, and Susie Maidment—and their network of collaborators, who include everyone from biologists specializing in living animals to computer scientists and programmers.

A few years ago, when I was finishing my PhD, Karl and Peter invited me to take part in a study of sauropod body size and proportions using digital models. It was an ambitious goal: make detailed computer animations of all sauropods with complete enough skeletons and figure out how big these animals were and how their bodies changed as they grew into truly titanic sizes. I was invited for
purely practical reasons: some of the best sauropod skeletons in the world are on display at the Amer­ican Museum of Natural History in New York City, where I was based at the time, and they needed data for one of them in particular, a Late Jurassic species called Barosaurus. They instructed me how to gather the information to build the model, and I was surprised that all it required was a normal digi­tal camera, a tripod, and a scale bar. I took about a hundred photos of the Barosaurus skeletal mount from all possible angles, keeping my camera steady on the tripod and making sure to include a ruler in most of the images. Then Karl and Peter input the images into a computer program that matches equivalent points on the photographs, works out the distances between them based on the scale, and does this continuously until a three-dimensional model is built from the original 2-D images.

Brontosaurus at the American Museum of Natural History in New York, with a human skeleton for scale.

American Museum of Natural History Library


A digital computer model of the skeleton of the sauropod Giraffatitan, which helps scientists calcu­late the weight of the animal.

Courtesy of Peter Falkingham and Karl Bates.

The technique is called photogrammetry, and it’s revolutionizing how we study dinosaurs. The su­per-accurate models it creates can be measured in precise detail. Or they can be loaded into anima­tion software and made to run and jump, in order to determine what kinds of motions and behaviors dinosaurs were capable of. They can even be used to animate movies or television documentaries, ensuring that the most realistic dinosaurs appear on screen. These models are bringing dinosaurs to life.

Our computer modeling study and more traditional studies based on limb-bone measurements come to the same conclusion: sauropod dinosaurs were really, really big. The primitive proto- sauropods like Plateosaurus began to experiment with relatively large sizes in the Triassic, as some of them got up to about two or three tons in weight. That’s roughly equivalent to a giraffe or two. But after Pangea started to split, the volcanoes erupted, and the Triassic turned into the Jurassic, the true sauropods got much larger. The ones that left tracks in the Scottish lagoon weighed about ten to twenty tons, and later in the Jurassic, famous beasties like Brontosaurus and Brachiosaurus ex­panded to more than thirty tons. But that was nothing compared to some supersize Cretaceous species like Dreadnoughtus, Patagotitan, Argentinosaurus—members of an aptly named subgroup called the titanosaurs—which weighed in excess of fifty tons, more than a Boeing 737.

The biggest and heaviest land animals today are elephants. Their sizes vary, depending on where they live and which species they belong to, but most weigh about five or six tons. Apparently the largest one ever recorded was around eleven tons. They have nothing on sauropods. Which circles back to the money question: how were these dinosaurs able to attain sizes so completely out of scale with anything else evolution has ever produced?

The first thing to consider is what animals require to become really big. Perhaps most obvious, they need to eat a lot of food. Based on their sizes and the nutritional quality of the most common Jurassic foodstuffs, it’s estimated that a big sauropod like Brontosaurus probably needed to eat around a hundred pounds of leaves, stems, and twigs every day, maybe more. So they needed a way to gather and digest such vast quantities of grub. Secondly, they need to grow fast. Growing bit by bit, year by year is all well and good, but if it takes you over a century to get big, that’s many opportunities for a predator to eat you, or a tree to fall on you during a storm, or a disease to take you out long be­fore you grow into your full-size adult body. Third, they must be able to breathe very efficiently, so they can take in enough oxygen to power all of the metabolic reactions in their immense bodies. Fourth, they need to be constructed in a way that their skeleton is strong and sturdy, but also not so bulky that it can’t move. Finally, they need to shed excess body heat, because in hot weather it is very easy for a big creature to overheat and die.

Sauropods must have been able to do all of these things. But how? Many scientists who started to ponder this riddle decades ago went for the easiest answer: maybe there was something different in the physical environment back in the Triassic, Jurassic, and Cretaceous. Perhaps gravity was weaker, so heftier animals could move and grow more easily back then. Or maybe there was more oxygen in the atmosphere, so the hulking sauropods could breathe, and therefore grow and metabolize, more efficiently. These speculations might sound convincing, but on closer scrutiny they don’t check out. There is no evidence gravity was substantially different during the Age of Dinosaurs, and oxygen levels back then were about the same as today, or maybe even slightly lower.

That leaves only one plausible explanation: there was something intrinsic about sauropods that al­lowed them to break the shackles that constrained all other land animals—mammals, reptiles, am­phibians, even other dinosaurs—to much smaller sizes. The key seems to be their unique body plan, which is a mixture of features that evolved piecemeal during the Triassic and earliest Jurassic, culmi­nating in an animal perfectly adapted for thriving at large size.

It all starts with the neck. The long, spindly, slinky-shaped neck is probably the single most distinc­tive feature of sauropods. A longer-than-normal neck started to evolve in the very oldest Triassic proto- sauropods, and it got proportionally longer over time, as sauropods both added more vertebrae—the individual bones in the neck—and stretched each individual vertebra ever further. Like Iron Man’s armor, the long necks conferred a kind of superpower: they allowed sauropods to reach higher in the trees than other plant-eating animals, giving them access to a whole new source of food. They could also park themselves in one area for several hours and extend their necks up and down and all around like a cherry picker, gobbling up plants while expending very little energy. That meant they were able to eat more food, and thus take in energy more efficiently, than their competitors. That’s adaptive advantage number one: their necks permitted them to eat the huge meals necessary to put on excessive weight.

Then there’s the way that they grew. Recall that the dinosauromorph ancestors of dinosaurs devel­oped higher metabolisms, faster growth rates, and a more active lifestyle than many of the amphib­ians and reptiles that were also diversifying in the earliest Triassic. They weren’t lethargic, and it didn’t take them aeons to grow into adults like an iguana or a crocodile. This was also true of all of their di­nosaur descendants. Studies of bone growth indicate that most sauropods matured from guinea-pig­size hatchlings to airplane-size adults in only about thirty or forty years, an incredibly short period of time for such a remarkable metamorphosis. That’s advantage two: sauropods obtained the fast growth essential to reach large size from their distant, cat-size ancestors.

Sauropods also retained something else from their Triassic ancestors: a highly efficient lung. The lungs of sauropods were very similar to those of birds and very different from ours. While mammals have a simple lung that breathes in oxygen and exhales carbon dioxide in a cycle, birds have what is called a unidirectional lung: air flows across it in one direction only, and oxygen is extracted during both inhalation and exhalation. The bird-style lung is extra efficient, sucking up oxygen with each breath in and each exhalation. It’s an astounding feature of biological engineering, made possible by a series of balloonlike air sacs connected to the lung, which store some of the oxygen-rich air taken in during inhalation, so that it can be passed across the lung during exhalation. Don’t worry if it sounds confusing: it is such a strange lung that it took biologists many decades to figure out how it works.

We know that sauropods had such a birdlike lung because many bones of the chest cavity have big openings, called pneumatic fenestrae, where the air sacs extended deep inside. They are exactly the same structures in modern birds, and they can only be made by air sacs. So that’s adaptation three: sauropods had ultra-efficient lungs that could take in enough oxygen to stoke their metabolism at huge size. Theropod dinosaurs had the same bird-style lungs, which could have been one factor that allowed tyrannosaurs and other giant hunters to get so large, but the ornithischian dinosaurs did not. This is why duck-billed dinosaurs, stegosaurs, horned species, and armored dinosaurs were never able to grow as huge as sauropods.

It turns out that air sacs also have another function. Aside from storing air in the breathing cycle, they also lighten the skeleton when they invade bone. In effect, they hollow out the bone, so that it still has a strong outer shell but is much more lightweight, the way an air-filled basketball is lighter than a rock of similar size. Want to know how sauropods could hold up their long necks without top­pling over like an unbalanced seesaw? It’s because all of the vertebrae were so engulfed by air sacs that they were little more than honeycombs, featherweight but still strong. And that’s advantage four: the air sacs allowed sauropods to have a skeleton that was both sturdy and light enough to move around. Without air sacs, mammals, lizards, and ornithischian dinosaurs had no such luck.

And what about the fifth special adaptation, being able to expel excess body heat? The lungs and air sacs helped with this too. There were so many air sacs, and they extended throughout so much of the body, snaking their way into bones and between internal organs, that they provided a large sur­face area for dissipating heat. Each hot breath would be cooled by this central air conditioning system.

Putting it all together, that’s how you can build a supergiant dinosaur. If sauropods had lacked any one of these features—the long neck, the fast growth rates, the efficient lung, the system of skeleton­lightening and body-cooling air sacs—then they probably would not have been capable of becoming such behemoths. It wouldn’t have been biologically possible. But evolution assembled all of the pieces, put them together in the right order, and when the kit was finally assembled in the post-vol­canic world of the Jurassic, sauropods suddenly found themselves able to do something no other ani­mals, before or since, have been able to do. They became biblically huge and swept around the world; they became dominant in the most magnificent way—and they would remain so for another hundred million years.


Dinosaurs and Drifting Continents


Chapter Title art by Todd Marshall

NESTLED WITHIN THE LEAFY STREETS of New Haven, Connecticut, on the northern fringes of the Yale University campus, there is a shrine. The Great Hall of Di­nosaurs at Yale’s Peabody Museum may not bill itself as a place of spiritual pil­grimage, but that’s sure what it feels like to me. I get a shiver, as when I walked into Catholic mass as a child. It’s not a normal shrine—no statues of deities, flickering candles, or the hint of incense. It’s also not particularly magnificent, at least from the outside, tucked away inside a fairly nondescript brick building that blends in with the rest of the university’s lecture halls. But it houses relics that, to me, are as sacred as those you’ll find in most any religious shrine: di­nosaurs. To me, there is nowhere better, anywhere on the planet, to go and im­merse yourself in the wonder of the prehistoric world.

The Great Hall was originally built in the 1920s to house Yale’s incomparable dinosaur collection, assembled over many decades by roughnecks who fanned across the American West and, for the right fee, sent fossil treasures eastward to be studied by the Ivy League elite. Coming up on its centen­nial, the gallery retains all of its original charm. This isn’t some New Age exhibit space with flashing computer screens and dinosaur holograms and a roaring soundtrack in the background. It’s a temple of science, where skeletons of some of the most iconic dinosaurs stand in solemn vigil, lights down low, in the sort of silence you really do expect in a church.

Covering the entire east wall is a mural that stretches more than a hundred feet long and sixteen feet high. Taking four and a half years to complete, it was painted by a man named Rudolph Zallinger, who was born in Siberia, moved to the United States, and took up illustration professionally during the Great Depression. If he were around today, Zallinger would probably be working for an animation stu­dio as a storyboard artist. He was a master at setting scenes and incorporating diverse sets of charac­ters, telling grandiose stories with the stroke of his brush. His most famous work is undoubtedly The March of Progress—that often satirized timeline of human evolution in which a knuckle-walking ape gradually morphs into a spear-carrying man. More people have probably come to understand, or mis­understand, the theory of evolution through that one image than through all of the textbooks, school lectures, and museum exhibits the world over.

But before he was painting humans, Zallinger was obsessed with dinosaurs. His mural inside the Great Hall—called The Age of Reptiles—is the crowning achievement of that stage of his career. It’s been on US postage stamps, was featured in a Life magazine series, and is either reproduced or pla­giarized on all sorts of dinosaur paraphernalia. It’s the Mona Lisa of paleontology, surely the single most talked-about piece of dinosaur artwork that has ever been created. But really, it’s more akin to the Bayeux Tapestry, because it tells an epic tale of conquest. It’s the saga of how fishy creatures first emerged onto land, colonized a new environment, and diversified into reptiles and amphibians; then, of how these reptiles split off into the mammal and lizard lines, the proto-mammals having their day and the lizards following, eventually producing the dinosaurs.

As the mural nears its end, some sixty feet and 240 million years from where it started, after a long journey through alien landscapes of primeval scaly beasts, the painting finally becomes engulfed in dinosaurs. It kind of sneaks up on you, as the transition from the lizards and proto-mammals to the dinosaurs unfolds incrementally across the canvas. Now it’s dinosaurs everywhere, of all shapes and sizes, some enormous and others blending into the background. Suddenly, the mural has taken on the feel of something quite different—of a Soviet propaganda poster with Stalin gesticulating before a crowd of peasants, or one of those hilariously self-aggrandizing frescoes in Saddam’s palaces. One glance at the dinosaurs and I feel the power. Strength, control, dominance. The dinosaurs were in command, and this was their world.

The theropod Deinonychus stands guard over the Zallinger mural at the Peabody Museum, Yale


Photo courtesy of the author

This part of Zallinger’s mural beautifully encapsulates what it was like when dinosaurs had as­cended to the peak of their evolutionary success. A monstrous Brontosaurus lounges in a swamp in the foreground, munching away on the ferns and evergreen trees surrounding the water. Off to the side, a bus-size Allosaurus rips into a bloodied carcass with its teeth and claws, its massive feet stomping on its prey for a little extra insult. Keeping a safe distance is a peaceful grazing Stegosaurus, which displays its full arsenal of bony plates and spikes just in case the carnivore has other ideas. Far in the background, where the swamp disappears into a wall of snowcapped moun­tains, another sauropod uses its long neck to vacuum shrubs off of the ground. Meanwhile, two pterosaurs—those flying reptiles closely related to dinosaurs, often called pterodactyls—chase each other overhead, dipping and diving through the tranquil blue sky.

Odds are, this is the type of image that many of us think of when we think of dinosaurs. These are dinosaurs at their pinnacle.

ZALLINGER’S MURAL IS not fiction. Like any good art, it takes a few liberties here and there, but it is largely rooted in fact. It’s based on those very same di­nosaurs that stand in front of it in the Great Hall: familiar names like Bron­tosaurus, Stegosaurus, and Allosaurus. These dinosaurs lived during the Late Jurassic Period, about 150 million years ago. By this time, dinosaurs had al­ready become the dominant force on land. Their victory over the pseudosuchi- ans was 50 million years in the rearview mirror, and it had been a good 20 mil­lion years since some of the first giant long-necked species were splashing through the lagoons of Scotland. Nothing was holding back the dinosaurs any­more.

We know a lot about the dinosaurs of the Late Jurassic. That’s because there are abundant fossils from this time, in many parts of the world. It’s just one of those quirks of geology: some time periods are better represented in the fossil record than others. It’s usually because more rocks were being formed during that time, or rocks of that age have better survived the rigors of erosion, flooding, vol­canic eruptions, and all of the other forces that conspire to make fossils difficult to find. When it comes to the Late Jurassic, we enjoy two lucky breaks. First, there were hugely diverse communities of dinosaurs living alongside rivers, lakes, and seas all around the world—the perfect places to bury fossils in sediments that later turned to rock. Second, these rocks are today exposed in places conve­nient for paleontologists—in sparsely populated and dry regions of the United States, China, Portugal, and Tanzania, where annoyances like buildings, highways, forests, lakes, rivers, and oceans don’t cover up the fossil booty.

The most famous Late Jurassic dinosaurs—those in Zallinger’s mural—come from a thick rock de­posit that pokes out all across the western United States. Its technical term is the Morrison Formation, named for a small town in Colorado where there are some beautiful exposures of its colorful mud­stones and beige-tinged sandstones. The Morrison Formation is a monster: it can be found in thirteen states today, covering nearly four hundred thousand square miles (a million square kilometers) of the American scrublands. It is easily sculpted into low hills and undulating badlands, the sort of classic backdrop you see in Western films. It’s also the source rock for some of the country’s most important uranium ore deposits. And, yes, it’s a hotbed of dinosaurs, ones whose uranium-infused bones make Geiger counters sing.


Paul Sereno in Wyoming.
Photo courtesy of the author
























Excavating sauropod bones in the Morrison Formation near Shell, Wyoming. At the center back is

Sara Burch, who later became an expert on T. rex arms (see Chapter 6).

I worked in the Morrison Formation for two summers as an undergraduate. It’s where I cut my teeth excavating dinosaur skeletons. I was apprenticing in the lab of the University of Chicago’s Paul Sereno, whom we last met leading the expeditions to Argentina that turned up some of the world’s very oldest dinosaurs, the Triassic-age Herrerasaurus, Eoraptor, and Eodromaeus. But Paul seemed to study everything and do fieldwork everywhere: he had also found bizarre fish-eating and long­necked dinosaurs in Africa, he’d explored China and Australia, and he’d even described important fos­sils of crocodiles, mammals, and birds.

In addition, like any academic paleontologist, Paul also had to spend time in the classroom. Each year he taught a popular undergraduate class called Dinosaur Science, which combined theory with practice. Because you can’t find dinosaurs anywhere near Chicago, the class would take a ten-day field trip each summer to Wyoming, where the students had the once-in-a-lifetime opportunity to dig dinosaurs with a celebrity scientist. Although at the time I had little prior experience, I was brought on as a teaching assistant, Paul’s right-hand man as we herded the students—a diverse lot, from pre­meds to philosophy majors—across the high desert.

Paul’s field sites were located near the tiny town of Shell, secluded between the Bighorn Moun­tains to the east, and Yellowstone National Park a hundred miles to the west. Only eighty-three people were counted during the last census. When we were there in 2005 and 2006, the road signs boasted of merely fifty residents. But that’s a good thing for paleontologists. The fewer people in the way of the fossils, the better. And although Shell is a forgettable dot on the map, it can rightly stake its claim as one of the world’s dinosaur capitals. It is built on the Morrison Formation, surrounded by beautiful hills carved out of muted green, red, and gray rocks bursting with dinosaurs. So many dinosaurs have been found here that it’s hard to keep track, but the count is probably well over a hundred skeletons by now.

As we drove west from Sheridan, on a surprisingly treacherous road across the rugged Bighorns, I felt I was on the trail of giants. Some of the biggest dinosaurs of all have been found in the Shell area: long-necked sauropods like Brontosaurus and Brachiosaurus, and the huge carnivores, like Al- losaurus, that ate them. But I also felt I was walking in the footsteps of another type of giant: the ex­plorers who found the first bones in this area in the late nineteenth century, the railwaymen and labor­ers who started a dinosaur rush and seized the moment to reinvent themselves as mercenary fossil collectors on the payrolls of gilded institutions like Yale University. They were a ragtag bunch, Wild West ruffians with cowboy hats, mustaches, and unkempt hair, who hacked giant bones out of the ground for months on end, and spent their free time raiding one another’s sites, constantly feuding and sabotaging and drinking and shooting. But these unlikely characters revealed a prehistoric world that nobody knew existed.

The first Morrison fossils were surely noticed by the many Native American tribes scattered across the West, but the first recorded bones were collected by a surveying expedition in 1859. In March 1877 the real fun started. A railroad worker named William Reed was returning home from a success­ful hunt, rifle and pronghorn antelope carcass in tow, when he noticed some huge bones protruding out of a long ridge called Como Bluff, not too far from the railroad tracks in an anonymous expanse of southeastern Wyoming. He didn’t know it, but at the same time a college student, Oramel Lucas, was finding similar bones a few hundred miles to the south, in Garden Park, Colorado. That same month, a schoolteacher named Arthur Lakes had just found a cache of fossils near Denver. By the end of that March, the fever of discovery was spreading throughout the American West, to even the most remote villages and railway outposts.

Like any prospecting rush, the dinosaur frenzy attracted a horde of questionable characters to the Wyoming and Colorado backcountry. Many of these men were grizzled opportunists on one mission: to convert dinosaur bones into cash. It didn’t take long for them to realize who was paying top dollar: two dapper East Coast academics, Edward Drinker Cope of Philadelphia and Othniel Charles Marsh of Yale University, the same men we briefly met two chapters ago, who studied some of the first Triassic di­nosaurs found in western North America. Once chummy, these two scientists had let ego and pride metastasize into a full-on feud, which was so radioactive that they would do anything to one-up each other in an insane battle to see who could name the most new dinosaurs. Cope and Marsh were op­portunists, too, and with each letter from a ranch hand or railway porter reporting more new dinosaur bones from the Morrison badlands, they saw the opportunity they had been craving but had been un­able to yet fulfill: a chance to beat the other guy once and for all. And they both went for it.

Cope and Marsh treated the West like a battlefield, employing rival teams that often acted more like armies, scooping up fossils wherever they went and sabotaging the other side whenever they could. Loyalties were fluid. Lucas worked for Cope, and Lakes teamed up with Marsh. Reed worked for Marsh, but members of his team defected to Cope. Pillaging, poaching, and bribing were the rules of the game. The madness continued for over a decade, and when it was over, it was hard to separate the winners from the losers. On the plus side, the so-called Bone Wars led to the discovery of some of the most celebrated dinosaurs, the ones that roll off the tongue of every schoolchild: Allosaurus, Ap­atosaurus, Brontosaurus, Ceratosaurus, Diplodocus, Stegosaurus, just to name a few. On the other hand, the mentality of constant warfare caused a lot of sloppiness: fossils haphazardly excavated and hastily studied, scraps of bone mistakenly christened as new species, different bits of the skeleton of the same dinosaur regarded as belonging to totally different animals.

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