Revelation

Fossilists and social reformers alike could claim 1824 as a banner year—the dawning of a new era of understanding. At the same time Buckland was busy exploring caves and fossiling at Stonesfield, those preoccupied with the harsh realities of their present world witnessed Parliament's repeal of the Combination Act.

The Combination Act, which had been in force for the previous twenty-five years, made it illegal for workers to join together to improve working conditions, effectively banishing trade unions. The decision to repeal the act, thereby legalizing unions, was not owing to any largesse of the Tory government. It merely reflected the government's acceptance of the recommendations of a Select Committee of Parliament. Indeed, the prime minister, Lord Liverpool, was a staunch opponent of reform, and had introduced some sternly repressive measures during his twelve years in office. These included the Six Acts, rushed into the statutes in 1819, following the Peterloo Massacre.

This massacre, which took place in St. Peter's Fields, Manchester, was named in ironic allusion to the Battle of Waterloo. At Waterloo British troops had been used against the French, but at Manchester they were used against the English. A crowd of some 50,000 people had gathered in a peaceful demonstration for parliamentary reform, but the local authorities overreacted and sent in the troops. The soldiers charged into the crowd with swords drawn, killing 11 and wounding 400.

The Six Acts were aimed at preventing further demonstrations for social reform. They included such Draconian measures as prohibiting public meetings of more than fifty people, except with prior consent. Publications judged to be blasphemous or seditious were banned, and certain radical publications that had previously avoided paying stamp duty now had to pay. Stamp duty required every publication to pay a levy of four pence per copy, putting the cost of newspapers out of reach of all but the rich.

Peterloo and its aftermath spurred Shelley to pen "Men of England," one of several poems he wrote to rally the spirit of the working classes. Had he been living in England at the time, rather than in Italy, he would undoubtedly have been arrested for sedition—if not for the first verse, then certainly for the sixth:

Men of England, wherefore plough For the lords who lay ye low? Wherefore weave with toil and care The rich robes your tyrants wear?

Sow seed—but let no tyrant reap: Find wealth—let no impostor heap: Weave robes—let not the idle wear: Forge arms—in your defence to bear.

Mantell had been so appalled by the Peterloo Massacre that he had drafted a set of resolutions to the Prince Regent. He was no doubt delighted when the Tories struck down the Combination Act. But Buckland, lacking Mantell's zeal for social reform, was probably too busy with other things to pay it much heed. The most pressing engagement on his calendar for 1824 was his presentation of a paper on the Stonesfield giant to the Geological Society.

In preparation for the February meeting Buckland had engravings made of the newly discovered jaw, together with parts of the skeletons of several other individuals. Although he had nothing that even approached an entire skeleton, he hoped to give his audience sufficient information to enable them to recognize the creature, should they encounter its remains in their own collections. That way they might contribute to a more complete picture of the beast.

With a little over three weeks to go before his presentation, Buckland received some unexpected news from the West Country. The news was sufficiently important to cause him to drop everything and dash off to Lyme Regis, calling on his good friend Conybeare on the way.

The Reverend William Conybeare's passion for geology was second only to his devotion to God and to his church. He was Rector of Sully at the time, in South Wales, but lived in Bristol. His ecclesiastical duties kept him busy, but he still had some time for geology. Living in Bristol helped him keep abreast of the marine reptile discoveries being made in the West Country by Mary Anning, and by local collectors nearby.

Conybeare was very astute. Even before Anning's discovery of the world's first plesiosaur skeleton, which he subsequently studied and named in 1821, he had anticipated its existence. Not that he knew what kind of creature it was. All he knew was that a second kind of marine reptile coexisted with ichthyosaurs. This surmise was based on his discovery of some unusual vertebrae in a collection of ichthyosaur vertebrae belonging to a local fossilist. The unidentified vertebrae had relatively thicker centra (spools), and each centrum was permanently fused to its neural arch. He was eventually able to match the mystery vertebrae to those of Anning's first plesiosaur skeleton, confirming his suspicion.

The first plesiosaur skeleton, recall, had no skull, so Conybeare still did not know what sort of head plesiosaurs had. But sometime after its discovery, an isolated and incomplete lower jaw was found at Lyme Regis. The jaw was obviously not ichthyosaurian, and Conybeare suspected it probably belonged to a plesiosaur. Then a complete but badly crushed skull was discovered in the town of Street, partway between Bristol and Lyme Regis. Again, it was not ichthyosaurian, and Conybeare suspected that it too belonged to a plesiosaur, but he had no way of knowing. What was desperately needed was a complete skeleton, but Conybeare, in the meantime, had to make do with what was available.

Conybeare's studies of the available material convinced him that plesiosaurs formed a link between ichthyosaurs and crocodiles. Like Buckland, and most of the others in their geological circle, Conybeare thought that all organisms formed part of a "connected chain of organized beings" that God had created. Adherents to this belief were quick to deny any transmutational [evolutionary] connotation, and Conybeare made the point quite clear in his first paper on plesiosaurs: "When alluding to the regular gradation . . . the linked . . . series of animal forms, we would wish carefully to guard against the absurd and extravagant application which has sometimes been made of this notion [of transmutation]."

He went on to explain how every niche capable of supporting life had been filled, so there was a wide variety of different creatures, each providing:

Striking proof of the infinite riches of creative design, [and] of the infinite wisdom ... [of the Creator]. Some . . . however (and Lamarck is

Real Life Dragon Skeleton

The isolated lower jaw from Lyme Regis that Conybeare suspected was part of a plesiosaur (top), and the crushed skull from Street.

Mary Anning's second plesiosaur discovery—the first complete skeleton ever found. The specimen is on display at London's Natural History Museum.

more especially censurable on this account) have most ridiculously imagined that the links hence arising represent real transitions ... an idea so monstrous, and so completely at variance . . . with the evident permanency of all animal forms, that [it required] . . . nothing less than its bigotry to defend it.

Transmutation was clearly anathema to Conybeare.

It is unfortunate that Mary Anning had collected most of the first plesiosaur skeleton as separate bones, thereby losing their exact relationships. Conybeare's reconstruction of the fore paddle was accordingly conjectural, and the hind one was so incomplete that he did not even attempt a reconstruction. Some of the vertebrae appeared to have been lost during collection, so he could not be absolutely sure that the creature had quite such a long neck as he suspected. And, of course, the skull was missing. But this unsatisfactory situation was about to change.

Conybeare was busily working on a guest sermon he had to preach in Oxford when Buckland, en route to Lyme, came bursting in on him. Buckland, larger than life and doubtless in a great state of excitement, probably blurted his news: Mary Anning had just discovered a second plesiosaur! But this one, unlike the first, was a complete skeleton. Buckland told Conybeare that Anning had offered it to the Duke of Buckingham for £100. The duke had

Henry De la Beche, who collaborated with Conybeare on the study of marine reptiles, lived in Lyme Regis during his teenage years. He is often romantically connected with Anning, two years his junior, but there is no good evidence.

asked Buckland to examine the specimen and complete the transaction, if the skeleton was what it was purported to be.

Conybeare's peace and quiet was completely shattered by the visit, preventing him from getting on with his sermon, but he could not help being caught up in the excitement. He was anxious to know whether the new specimen confirmed his conjectures on the anatomy of plesiosaurs. "I begged him of course to send me immediate intelligence thence," Conybeare wrote his geological friend Henry Dc la Beche, who was currently in Jamaica, "and three days afterwards received a very fair drawing by Mary Anning of the most magnificent specimen. . . . You may imagine the fuss all this occasioned."

The Bristol Philosophical Society, which Conybeare had helped to establish, was meeting that very week, so he resolved to break the news there: "and to the Society I went, delighted ... of making this strange monster first known to the public. ..." Conybeare had only Anning's drawing to show his audience, but it was enough to demonstrate the singular features of the remarkable new discovery. "Such a communication could not fail to excite great interest," Conybeare told De la Beche. "Some of the folk ran off instantly (it was Friday evening) to Gutch's printing office [Bristol Press], whither I was obliged to follow to prevent some strange blunders . . ." being made by the journalists.

Buckland arranged for the specimen to be shipped to London, to be placed on temporary exhibition at the Geological Society. He charged Conybeare to be there to meet it, to make sure it did not fall into the hands of Sir Everard Home (1756-1832).

Sir Everard Home, first president of the Royal College of Surgeons, studied under John Hunter, the celebrated surgeon, who later married Home's sister. When Hunter died in 1793, Home became the executor of his will, and was entrusted with Hunter's unpublished manuscripts. Home was not noted for his productivity or brilliance, but he began to publish more papers than he had before. The suspicion naturally arose that he was plagiarizing Hunter's work. The suspicion was founded in fact, but the guileful Home destroyed the evidence by burning Hunter's manuscripts. Some of the geological circle knew of these nefarious deeds, which helps explain why he was not widely trusted or respected.

Buckland was to be elected president of the Geological Society at the forthcoming annual meeting, and hoped the plesiosaur would arrive in time for this auspicious occasion. So too did Conybeare, who was scheduled to deliver an address on its anatomy at the same meeting. Conybeare arrived in town as arranged, only to learn that the ship carrying the precious cargo had been delayed in the English Channel. Undaunted, he was determined to make the most of his unexpected trip to London. He stayed at the Salopian Coffee House, a hotel in Charing Cross Road that was Buckland's favorite haunt.

As it happens, the plesiosaur skeleton failed to arrive in time for the meeting, but Conybeare was not unduly concerned. He told his friend De la Beche that the annual dinner was:

one of the pleasantest public meetings I have ever attended. Buckland as the new Pres.[iden]t was put to his oratory, & some dozen of us talked in our turn, but in place of the usual trash on such occasions every one had some interesting facts ... to communicate. We adjourned to the Society's rooms at 112 past eight, & there I lectured on my Mstr [monster]

Conybeare's presentation was an enormous success. He did not have the same flair for storytelling as Buckland, but he nevertheless painted a graphic picture of the bizarre anatomy of plesiosaurs. They were as alien to his audience as if they had hailed from another planet, invoking ethereal images of primordial seas ruled by wondrous reptiles. Using Anning's detailed drawing of the skeleton, he showed how the new discovery confirmed several of his conjectural points on plesiosaurian anatomy. The neck was as unbelievably long as he had predicted—longer than in any other animal. Initially he had counted twelve neck vertebrae, but the new specimen had about thirty-five. No other animal had so many neck vertebrae, the closest approach being birds, though he commented that they too fell short, with no more than twenty-three (some swans have upward of twenty-five). He went on to suggest:

It swam upon or near the surface, arching back its long neck like the swan, and occasionally darting it down at ... fish. ... It may perhaps have lurked in shoal water along the coast, concealed among the seaweed, and raising its nostrils to a level with the surface from a considerable depth ... a secure retreat from the assaults of dangerous enemies . . . the length and flexibility of its neck may have compensated for the want of strength in its jaws and its incapacity for swift motion through the water

Conybeare's image of a plesiosaur cruising along near the surface, with its long neck arched above the waves, became very popular, inspiring artists for

An early nineteenth-century vision of life in the sea during the Age of Reptiles, showing a swan-necked plesiosaur.

generations to come. But it was not very plausible. This is because plesiosauri-an vertebrae, unlike those of birds, have restricted dorso-ventral (up and down) mobility, most of their flexibility being lateral (side to side). The notion of their lurking in the shallows with their nostrils raised above the surface is also implausible, not only because of the limited dorso-ventral neck flexibility, but also because of the problems of water pressure. When an animal is below the surface, water pressure tends to collapse its lungs, and this pressure increases with depth, by one atmosphere of pressure for every thirty-two feet (10 meters). The lungs therefore have to work against water pressure during inspiration, and this becomes increasingly difficult with increasing depth, which is why scuba divers must breathe compressed air.

Conybeare's studies enabled him to confirm that the isolated skull, found in Street a couple of years before, was indeed plesiosaurian, because the new skeleton had a similar type of skull:

We now also learn for the first time, that the head of this animal was remarkably small, forming less than the thirteenth part of the total length of the skeleton; while in the Ichthyosaurus its proportion is one-

fourth. This proportional smallness of the head, and therefore of the teeth, must have rendered it a very unequal combatant against the latter animal. . . .

The image of animals actively pitching into one another in mortal combat was a popular notion during Conybeare's time. De la Beche, for example, painted a watercolor depicting such a scene, sometime around 1831, and called it Duria antiquior or Ancient Dorset. The central figure, a large toothy ichthyosaur, bites through the long neck of a plesiosaur, and another plesiosaur launches a sneak attack from the shallows on a crocodile basking on the shore. The sea is a throng of biting, snapping jaws, and, in the distance, the long, disembodied neck of a plesiosaur launches itself into the air to snatch a low-flying pterosaur.

Such sanguine imagery of the ancient globe was based on a distorted view of the fossilists' nineteenth-century world. Today, thanks mainly to television, we know that animals in their natural habitat arc not locked in eternal battles to the death. We know, for example, that combat between meat eaters, or between any other groups of animals, rarely becomes an overt conflict of tooth and claw. Rather, it is a subtle competition for resources and for living space. People in Regency England rarely had the opportunity to witness interactions between animals in the wild. Some individuals had seen lions and tigers in menageries, and could see how fierce they were. It was therefore natural for them to suppose that such animals spent most of their time in open conflict in the wild. It is not surprising, then, that the early fossilists were so obsessed with depicting combat among the denizens of the prehistoric world.

Conybeare told his audience that the limbs of plesiosaurs were as unusual as their necks. Their digits were exceptionally long, with upward of ten phalanges (finger bones), compared with only three in our own fingers. Such long, slender paddles were unique among the quadrupeds. Conybeare freely admitted that he got the humerus the wrong way around in the first skeleton, and confused the bones of the forearm (the radius and ulna). He also acknowledged that he had wrongly depicted the edges of the paddles as being formed of rounded bones, but this was because "when the specimen . . . was found, the bones in question were loose, and had been subsequently glued into their present situation, in consequence of a conjecture of the proprietor."

The "proprietor" referred to here, of course, was Mary Anning. He could have used the word "collector," but may have chosen the alternate word to convey the pecuniary, rather than collegial, nature of her relationship with the geological circle. The fact that he did not mention her name once during his entire address, even though she discovered both of the plesiosaur skeletons, substantiates his intent to exclude her.

Conybeare was criticized by some geologists at the time he proposed the name plesiosaur (meaning "approximate to the saurians") because the name did not refer to their unusually long necks. He responded to this in his presentation to the Geological Society by saying, "I think it very probable, from specimens which I have examined, that species of Plesiosaurus with shorter necks exist. ..." He based this on differences in the relative proportions of their neck vertebrae. Conybeare was absolutely correct in this surmise because we now know that there are two distinctly different kinds of plesiosaurs. One group, today referred to as elasmosaurs, have long necks and small heads, like the first specimens he described. The other group, the pliosaurs, have short necks and large heads.That Conybeare could have made such a prophetic statement back in 1 824, based on the very limited material available to him, underscores his outstanding abilities.

The evening of February 20, 1824, was a singular night in the annals of paleontology because the second speaker to take the floor was Buckland, who had an important discovery to announce. Buckland's presentation was riveting. Some of the audience had already heard of his earlier findings of gigantic bones at Stonesfield. But now they learned he had discovered more material of this remarkable reptile, including part of the lower jaw with teeth still in place. This was the vital missing part of his osseous jigsaw puzzle. Now that he had part of the skull, albeit only a small part, he was able to determine what sort of a creature it was.

Buckland, the irrepressible storyteller, beguiled his audience with his account of the terrifying beast. It had dagger-shaped teeth, six inches long, each rooted in its own socket.They were curved and pointed, with serrations along their cutting edges, like a steak knife. These were the teeth of a ferocious flesh-eater, and he dwelt upon the way the old ones were replaced by the new ones.The new teeth were formed in distinct pits beside the old, and he speculated how these may have been expelled by pressure exerted by the new ones. He emphasized the remarkable nature of this rapid tooth replacement.

Without giving reasons, Buckland told the assembled gentlemen that the vertebral column and legs resembled those ol mammals, but that the teeth showed the animal to be reptilian. One of the mammal-like features that impressed him was probably the way the sacrum—the part of the backbone to which the hips are attached—was formed of five fused vertebrae. No other

The lower jaw of Buckland's Stonesfield giant. The teeth, which are still in place, have serrated edges, like a steak knife.

reptile known at that time had such a sacrum. The limb feature that reminded him of mammals was probably the way the head of the femur—the ball that fits into the socket of the hip joint—was set off from the shaft, almost at right angles. In modern reptiles the head of the femur is in line with the shaft, causing the bone to be splayed at the sides of the body. The same is also true for the forelimbs. This gives reptiles their sprawling posture. Mammals and birds, in contrast, have an upright posture, where the femur (and humerus) is held vertically beneath the body. Despite its mammalian leg structure, the Stonesfield giant was clearly reptilian in the way its daggerlike teeth were continuously replaced, a feature not found in any mammal.

Buckland would have been in his element with such an attentive audience, and such a rattling good story to tell. His estimate of the size of the leviathan, based on the length of its limb bones, would certainly have captured their attention. The largest femur found at Stonesfield was two feet nine inches (84 centimeters) long, which was huge when compared to that of a modern lizard. Scaling up a lizard until it had an equally large femur gave a size estimate for his giant of over forty feet (twelve meters). A reptile of this size was hard to imagine, and Buckland may have paused here to let the information sink in. He then told his audience that Gideon Mantell had discovered an even larger femur in the Tilgate Forest deposits of Sussex. Buckland inferred, erroneous ly, that this gigantic femur belonged to the same species, and comparisons to a modern lizard gave a length estimate of sixty to seventy feet (eighteen to twenty-one meters). This information, delivered with the correct timing and cadence, might have evoked a collective gasp of incredulity from his attentive listeners.To appreciate his audience's reaction we should remember the sensation that large exotic animals, like giraffes and elephants, had caused at zoological gardens. Here was a considerably larger animal, it was also a reptile, and a carnivorous one at that.

The large size inspired the name he coined for the beast. This name, meaning giant lizard, was announced without fanfare: "I have ventured, in concurrence with my friend and fellow-labourer, the Rev. W. Conybeare, to assign to it the name Megalosaurus." Neither Buckland, nor any of the other gentlemen present realized it at the time, but he was announcing the name of the world's first dinosaur.

Megalosaurus was a most remarkable discovery—far more significant than the most exciting of today's paleontological finds. This is because it showed, very graphically, that giant reptiles roamed the land during the remote antediluvian past, just as they had inhabited the seas and the air. Some of the gentlemen may have been acquainted with the idea of the Age of Reptiles through their readings of Cuvier. But here was solid proof, in the skeletal remains he set out before them, of one of the leviathans from that distant age—an age when reptiles ruled the Earth, as mammals dominated the modern world.

None of his audience would have doubted that these monstrous reptiles were extinct, like the mammoths that came long after them. Nor would any theological toes have been trodden on by the revelation, because the Book of Genesis told of a former time when "There were giants in the Earth."

If any of the gentlemen were having difficulties coming to terms with the Age of Reptiles, it was probably because of their lack of imagination, and the absence of any points of reference in their own world. There were, for example, no modern representatives of marine reptiles because neither plesiosaurs nor ichthyosaurs left any living descendants. But Buckland painted a more tangible picture of Megalosaurus. He saw it as a scaled-up version of a modern lizard, referring to it as the "great fossil lizard of Stonesfield." He also thought it was capable of venturing into the water, probably because of its association with crocodile and turtle fossils.

Having introduced Megalosaurus to his audience, Buckland spent some time discussing the associated fossils found at Stonesfield, comparing them to Mantell's Tilgate fossils. The two faunas were remarkably similar. Both includ ed terrestrial and aquatic animals, as well as terrestrial plants. And the two assemblages appeared to be formed under similar environmental conditions. However, Buckland pointed out that the thickness of the strata that intervened between the two formations ruled out the possibility that they were identical.

He was perfectly correct to assert that the Stonesfield and Tilgate assemblages were not the same. As noted earlier, they belong to the Middle Jurassic and Early Cretaceous, respectively. However, he was wrong to say that Megalosaurus occurred in the Tilgate fauna. Large carnivores similar to Megalosaurus were certainly known from Cuckfield, but the material was too incomplete to make a positive identification. Indeed, much of this material is still unidentified to the present day. He should therefore not have used the femur from Tilgate to estimate the size of Megalosaurus. His estimate of sixty to seventy feet was a gross exaggeration, by a factor of at least two, as we now know.

Although Megalosaurus was the largest and most impressive terrestrial fossil from the Mesozoic era, a tiny jaw had been discovered at Stonesfield that Buckland realized was "not less extraordinary." Buckland thought it was an opossum, because of its similarities with the living pouched mammals (marsupials). However, we now know that this primitive mammal, called Phascolotherium, was not a marsupial. What made it so special was that it was the first fossil mammal from the Mesozoic. Buckland, fully aware of the enormity of the find, told his audience that he identified the inch-long jaw as mammalian only on the authority of Cuvier, who had examined the specimen. He admitted that he would have hesitated to announce the unprecedented discovery of a mammal below the level of the chalk, had he not received the highest approbation of Cuvier.

The discovery was a major anomaly because it did not fit in with the accepted scheme of things. God had created different creatures at different times, beginning with the lowliest of animals, like fish, which first appeared in the older rocks. Life progressed through to the amphibians and reptiles, which first appeared at a higher level, to the mammals, which did not appear until later in the sedimentary series. The last mammal to appear in this continuous chain of beings was the human species, the last and greatest of God's creative works. So how could the presence of a mammal be explained in such ancient rocks?

Buckland did not offer any explanation at the time, and was quite content just to report the find, allowing Cuvier to take responsibility for authenticating the discovery. Others would sidestep the issue by arguing that the materi-

The Stonesfield "opossum" was an isolated lower jaw, only an inch (2.5 centimeters) long. This primitive mammal, called Phascolotherium ("pouched beast"), was the first mammal discovered from the Age of

Reptiles.

al must have been reworked from the overlying, younger, Tertiary deposits. This was not a satisfactory solution, however, and the problem was discussed and argued for years to come. Buckland discussed the issue at some length in his second book, Geology and Mineralogy, published in 1836. This was one of a series of books in the Bridgewater Treatises, commissioned by the late Earl of Bridgewater, to show "the Power, Wisdom, and Goodness of God, as manifested in the Creation . . ." Buckland made the point that the molar teeth of the Stonesfield "opossum" had a double root, so there was no question it was from a mammal, but he reasoned it was a marsupial. Marsupials bore their young in an immature state, and shared a number of other "inferior conditions." They clearly occupied "an intermediate place between viviparous [live-bearing] and oviparous [egg-laying] animals . . ." forming "a link between Mammalia and Reptiles . . ." leading "us to expect . . . that the first forms of Mammalia would have been Marsupials." Buckland thereby removed the anomaly of discovering a mammal so early in the geological strata by interpreting it as a new link in the "grand continuous chain which connects all past and present forms of organic life, as parts of one great system of Creation." This, of course, was not an evolutionary chain, but one ordained by God.

Buckland's announcement of the Stonesfield mammal would have caused a great deal of interest on that dark February evening. So too would Conybeare's disclosure of the bizarre anatomy of plesiosaurs. But Megalosaurus had taken center stage. Some of the gentlemen in the room may have wondered whether Megalosaurus was the only giant reptile to stalk the land in those remote ante diluvian times. There was, after all, more than one kind of sea dragon, so why not more than one kind of giant lizard? And although some may have speculated upon the matter, there were those present in the audience who knew, for sure, that other kinds of cold-blooded giants did exist: They had seen their fossilized remains with their own eyes.

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