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Nevraumont Book PTEROSAURS C O N T E N TS 1 DRAGONS OF THE AIR, 2 2 PTEROSAUR PLANET, 16 3 CONSIDERING MEDUSA, 30 4 A TREE FOR PTEROSAURS, 56 5 THE HEAD INSIDE OUT, 88 6 THE BODY INSIDE OUT, no 7 BABES ON THE WING, 140 8 HIGH FLIERS, 164 9 GROUNDED, 196 10 THE PTEROSAUR STORY, 224 11 POSTSCRIPT, 266 APPENDIX: LIST OF VALID PTEROSAUR SPECIES, 271 NOTES AND SOURCES, 275 BIBLIOGRAPHY, 311 ACKNOWLEDGMENTS, 329 INDEX, 331 DRAGONS OF THE AIR The dragon blinked in the fierce light of the sun as it emerged from the clouds and banked hard, its tremendous wings arching under the load. Reflected in a massive dark eye, the world below slowly tilted into view. Vast herds of dinosaurs were strung out across a dusty yellow-orange plain, occasionally gathered in knots where they had stopped to feed on patches of stunted vegetation. Then came marshes and— the dragon focused— a long, still, clear-blue lake. In its glassy depths, rainbow-hued fish hung almost motionless in the warm water, fins undulating, gills slowly pulsing. Suddenly, a huge dark shadow swept across the water's surface and, before it could dive into the safety of the weeds, a fish was gripped by long, sharp-pointed jaws and jerked violently into the air. As its consciousness faded, it gazed up into a limpid eye, set in a reptilian skull framed by fine, straggly hair. Slender, powerful wings beat smoothly up and down, membranes tensed and relaxed as the animal rose swiftly. At the top of its climb the Quetzalcoatlus paused, swallowed, then dived down again toward the shimmering lake far below.' FIGURE 1.1 A beautiful new fossil of the Upper Jurassic (approximately 148 million years old) Solnhofen Limestone pterosaur Anurognathus, only the second example yet known for this species. This pterosaur had a wingspan of about 16 inches (40 centimeters). (Photograph courtesy of Helmut Tischlinger.) 4 THE PTEROSAURS Around 215 million years ago, at about the same time as dinosaurs first spread across the continents, an altogether different group of reptiles took to the air. Reptiles had tried this before, on several occasions, but this time was different. This time they got beyond gliding, where all previous groups had given up, and, as bats and birds were to do millions of years later, they evolved a rare and complex adaptation: true flapping flight. Going boldly where no reptile had gone before, these intrepid aeronauts entered a new realm— the open sky— and developed into a riotous multitude of species. Some, such as the frog-headed beetle-cruncher Anurognathus, shown in Fig ure 1.1, were as small as a starling, but others, like Quetzalcoatlus, became as large as an airplane. They thrived for 150 million years, only to disappear forever in the events that also killed off many of their contemporaries, in cluding, most famously, the dinosaurs, and brought evolution's finest hour, the Mesozoic, to a close. This extraordinary group of animals— nature's real dragons— was the pterosaurs.2 The Trouble with Pterosaurs Pterosaurs, "winged reptiles," as their Greek name puts it, are familiar to most people as rather fierce-looking, leather-winged monsters featured in classic novels, such as Arthur Conan- Doyle's Lost World, or flit across the screen in TV documentaries3 and in movies from King Kong to Jurassic Park.4 No dinosaur scene is complete with out them— usually a Pteranodon, the most famous pterosaur of them all, with large teeth to give it added fierceness, even though this species was actually completely toothless.5 Trying to see beyond these superficial images and get a glimpse of the real thing triggers all sorts of questions: What were ptero saurs really like? How big did they get? Could they fly as well as birds? And one of the first questions that is always asked and hardest to answer: Why did they become extinct? Driven more than most by curiosity, scientists find these strange crea tures absolutely fascinating (pterosaur talks always fill the hall at scientific meetings) and some of the sharpest paleontological minds ever to ponder a fossil were so beguiled by these extraordinary animals that they made them the centerpiece of their researches. The first thing they discovered was that pterosaurs are really hard to understand. Even the term embodied in their name— winged reptile— seems contradictory. The word reptile, from the French "repere," meaning to creep, is not especially flattering, but describes living reptiles rather well. Wings, on the other hand, are about the last thing DRAGONS OF THE AIR 5 one might expect such a creature to have, and yet for pterosaurs, this defined them and their very existence. As soon as pterosaurs were discovered, at the end of the 18th century, naturalists started quarrelling about them. Disagreements came thick and fast: Pterosaur origins, for instance, were hotly debated on several occa sions, and, in one particular case detailed in Chapter 4, led to a protracted and acrimonious feud, while several other disagreements, for example, over wing shape and walking posture, continue even now. It was a long time be fore scientists could even agree as to what kind of animal pterosaurs might be— reptiles, as it turned out, but others argued that they were birds, and several eminent naturalists became convinced that they were bats. Debates also raged over the lifestyle of pterosaurs. Were they some kind of water creature? Or could they fly? And what did they do on the ground— strut around on their hind limbs like birds, or clump around on all fours? Even their likely metabolism became a scientific battleground: cool-blooded like crocodiles and lizards, insisted several authorities, warm-blooded like birds and mammals, countered others. Pterosaurs became a cause celebre, one of the great paleontological myster ies. Each generation of scientists "had a go" at them, and everyone had an opinion that, almost without exception, differed from that held by everyone else. The arguments, the confusion and the misunderstandings continued right up until a decade ago, and a few persist even today, but before we get into that, we should look a little more closely at why these ancient fliers caused so much controversy in the first place. A Fossil Problem Pterosaurs have successfully defied more than two centuries' worth of scientific probing for several reasons. The most obvious is the problem of trying to understand animals that are known only from fossils. Just a tiny proportion of all the pterosaurs that ever existed, prob ably less than one individual in a million, has actually made it into the fossil record. The processes by which their cadavers became fossils, normally sur vived only by the hardest parts of the body— bones and teeth6— mean that most of the important information about anatomy, movement and behavior, how pterosaurs were colored, what noises they made, was lost forever. Compounding the problem, even pterosaurs' hard parts were not well- suited for the rigors of fossilization. Pterosaurs were creatures of the air, with a relatively light and delicate skeleton constructed from slender, hollow 6 THE PTEROSAURS bones whose walls were often little thicker than a credit card. This is not a good design if you want to become a fossil. To begin with, it meant that even pterosaurs' skeletons were relatively easily destroyed, so, compared with other backboned animals, their fossil remains are rare. Worse still, if you pull open a museum drawer, you find that most of their fossilized remains consist of isolated, often broken, bones— dumb witnesses that tell us little more than "here be pterosaurs." To cap it all, most of the decent pterosaur fossils that we do have, whole skeletons and, very occasionally, fragments of fossilized soft tissues, come from just a few locations scattered across the world and are separated by vast, barren, pterosaur-less gaps of thousands of miles and millions of years. Thin-walled bones also mean that the complete skeleton of Anurogna- thus, shown in Figure 1.1, buried at the bottom of a Bavarian lagoon 148 mil lion years ago, is rather less helpful than one might expect. Like many other beautiful-looking pterosaurs, several of which are featured in this book, it is absolutely flat— a "picture" fossil— its hollow-tube bones unable to resist the inexorable crushing weight applied over countless millennia by the overlying rock. Without the three-dimensional, sticking-out-here, dimpled-in-there form of the skull, the pelvis, or any of the 300 or so bones that made up a pterosaur skeleton, and unable to measure the exact shape, size and position of the joints, paleontology is robbed of critical data. Confronted with a row of these "road-kills," it is often hard for an observer to establish even basic facts, such as: How many species are there? Two? Or more? Or just one, its representatives flattened in different ways? Trying to go further and find out, for example, how these pterosaurs might have stood, walked, or flown, is even harder. Strictly speaking, these are relatively simple questions (some of the really tough ones, for example, about physiology and breathing, will pop up later), but, enigmatic as the Mona Lisa, and sometimes just as smiley, the "picture" pterosaurs rarely give an answer. More Problems: Analogy and Chauvinism As if fossils themselves were not difficult enough to interpret, the ways in which scientists have gone about studying them also have their pitfalls. One trap that pterosaur researchers seem to have queued up to throw themselves into is a method much used, and no less often abused, by paleontologists: analogy. Confront ed with the fossilized remains of an organism from deep time— be it a tiddly little ammonite, a 10-ton dinosaur or a toothless Pteranodon, all extinct for half an eternity and tragically bereft of any living descendants— it is terri-