Missing Links 1981
 Arthur Keith (1866-1955), anatomist, was one of a British scientific triumvirate whose beliefs and work profoundly affected investigations into the evolution of man for nearly fifty years. His associates were Arthur Smith Woodward (1864-1944), paleontologist and Keeper of Geology at the British Museum of Natural History, and Grafton Elliot Smith (1871-1937), an anatomist whose speciality was the study of the brain. All three gentlemen were knighted for their contributions to science, and their talents were memorably displayed in discussions concerned with the significance of alleged fossil human remains found at Piltdown, Sussex, between 1908 and 1915.
In an autobiography written late in life, Keith remarked that 'the ideas which a man devotes his life to exploring are, for the greater part, those which come to him in the first tide of his inquiries'. 1 The observation is certainly sustained by the facts of Arthur Keith's career. In the first year of his medical studies, Keith was awarded a copy of The Origin of Species for his distinctive work in the anatomy class. While working as medical officer to a mining company in Siam immediately after he qualified, Keith dissected monkeys in the hope of determining whether or not the animals shared the affliction of malaria that plagued the human population and discovered an absorbing interest in comparative anatomy as the means of elucidating the evolutionary development of mankind. Within a year he dissected thirty-two assorted primates, and on returning to England in 1892 arranged to receive and dissect primate carcasses from the London Zoo. He dissected human foetuses as well, and the comparative anatomy of the ligamentous structure of the feet and hands of monkeys and human babies became both the subject of his doctoral dissertation and the basis of his views on the evolution of man's erect posture. 2 At home in Scotland Keith performed cerebral dissections on farmyard cats of all ages to clarify his understanding of the development of the individual brain, and in London he studied primate  skulls at the Royal College of Surgeons and at the British Museum of Natural History. He carefully noted about 150 observations on each of over 200 skulls and it was here, in the winter of 1894, Keith writes 3 that he learned 'the alphabet by which we spell out the long-past history of man and ape' from the evidence of fragmentary fossils.
In 1895 Keith met Dubois; he made a reconstruction of the Java skull and wrote an article on the specimen for a popular journal (concluding the specimen was essentially human, but of a lowly kind). Thereafter he wrote frequently on the subject of human ancestry and, on his appointment as Hunterian Professor of Anatomy at the Royal College of Surgeons in 1908, vowed to uncover and write 'the anthropological history of the British'. 4
In The Descent of Man, Charles Darwin had implied that the early forerunners of man probably retained some characteristics of the ancestor they shared with the apes. Males were probably furnished with great canine teeth at one time, he said, but as they acquired the habit of using stones, clubs and other weapons for fighting they would have used their jaws less and less, with consequent decrease in the size of the canines and some restructuring of the jaw. 5 Scientists of the day drew several important conclusions from Darwin's observation.. Reshaping the jaw would have provided the space essential for the movement of the tongue in articulate speech; the ability to handle stones and clubs presumed an erect posture; both speech and erect posture require a considerable development of the mental abilities and thus the crucial developments on mankind's evolutionary path were clearly defined. But which came first? Development of the brain? The erect posture? Or the ability to speak? Darwin hardly commented upon the question but, as the slowly accumulating fossil remains inspired competing interpretations of their imprecise evidence, it was clear that some idea of the manner in which human evolution had proceeded would help by suggesting the features that fossils ought to possess. If man had walked before he could talk, then fossils could be expected to demonstrate the fact.
As a result of his work on the feet and hands of apes and human babies, Arthur Keith believed erect posture was an ancient attribute and the large brain mankind's most recent acquisition. 6 An opposing view was ardently championed by Grafton Elliot Smith, whose important and pioneering work on the function and evolution of the vertebrate brain had convinced him that 'the brain led the way'. At its most primitive, the brain had discerned little more than the sensation of smell, he said. Later, vision had been acquired and then 'an arboreal mode of life started man's ancestors on the way to pre-eminence' for, while they  avoided the fierce competition for size and supremacy waged among carnivores and ungulates on the earth below, 'the specialization of the higher parts of the brain gave them [the primates] the seeing eye, and in the course of time also the understanding ear; . . . all the rest followed in the train of this high development of vision working on a brain which controlled ever-increasingly agile limbs'. Thus `the Primates found in the branches the asylum and protection necessary for the cultivation of brain and limbs,' he said, and the erect posture developed when they had become 'powerful enough to hold their own and wax great'. It was 'not the real cause of man's emergence from the Simian stage, but ... one of the factors made use of by the expanding brain as a prop still further to extend its growing dominion'.7
Arthur Smith Woodward appears to have contributed little to the early stages of the debate. In 1885 he remarked upon the preponderance of 'Missing Links' in the fossil record; 8 and subsequently he stated that 'we have looked for a creature with an overgrown brain and ape-like
face', 9 missing from the chain connecting man and his primate ancestors, but by far the greater part of his career was devoted to the study of fossil fish rather than fossil man. In all Smith Woodward published more than six hundred papers on fossil fish and related paleontological subjectsmore than three hundred before he became Keeper of Geology at the British Museum in 1901and only thirty or o on fossil man. These figures, of course, accurately reflect the relative abundance and otherwise of the fossils in question.
By 1908 the fossil evidence of early man was still very slight. Specimens from France and Belgium had confirmed the existence of a Neanderthal race; there were the enigmatic Java discoveries, and a lower jaw found in a sandpit near Heidelberg in 1907 suggested that mankind's late Pleistocene ancestors had been able to talk and bore a receding chin, as evidence of their simian associations. These specimens added a measure of information to the story of mankind's evolution in general, but of the British anthropological history that Arthur Keith had vowed to write there was no indisputable evidence whatsoever. Fossils had been sought in the caves and gravels of England, Ireland, Scotland and Wales; some remains had been found, but their antiquity was questionable and their aspect distinctly modern. For the most part, therefore, the remains of fossil man in Britain were granted little credence. It was believed more likely that each was a case of recent remains interred in old deposits than a genuine example of early man in Britain.
The Galley Hill skeleton was a case in point. The bones were found to the east of London, in gravels deposited by the River Thames when it  flowed one hundred feet above its present level.
Arthur Keith and the Galley Hill specimen
 The deposits were considered to be of early Pleistocene age; crude stone tools found near the skeleton were similar to others found elsewhere in association with extinct fauna of the same period, so the related geological and archaeological evidence seemed to suggest that Galley Hill Man was very old indeed. The anatomical evidence, however, clearly showed that the specimen represented Homo sapiens. The cranial capacity (about 1400 cubic centimetres) was well within the modern range, the chin did not recede, the jaw structure was compatible with the faculty of speech and the thighbone indicated an habitual erect posture. Furthermore, the skeleton was unusually complete and had been found with its parts in close proximity. All these factors combined to convince most authorities that the Galley Hill remains were of recent origin and had been entombed by the hand of men, not nature. 10
The Galley Hill skeleton was discovered in 1888 and soon thereafter consigned to the obscurity of a private collection in East London, where it remained until 1910, when Arthur Keith decided that the evidence merited reappraisal. Over a period of months he confirmed the skeleton's affinities with Homo sapiens and agreed that the remains had been buried in the ancient deposits but, whereas these factors had caused earlier investigators to reject the claims of great antiquity, Keith deduced from them that Galley Hill Man was as ancient as the deposits in which he had been buried, and the burial was not recent, but ancient. 'We hardly do justice to the men who shaped the [artefacts],' he wrote, 'if we hold them incapable of showing respect for their dead." 11 The implication of these deductions were considerable if large-brained, talking, walking Homo sapiens had existed at the beginning of the Pleistocene, as Keith claimed, then all the crucial evolutionary development of man must have taken place long before, true men had remained unchanged for a very long time, and the Neanderthal and Java fossils represented not the ancestors of man, but a "degenerate cousin'.
Geologists, in particular, did not agree that true man could be so ancient. But Keith claimed this was because they had grown up with a belief in the recent origin of man and therefore expected to see a sequence of anatomical change marking the evolution of man. 12 In fact the geologists' views simply conformed with the rationale of their paleontological training, which taught that the long and absolute lack of evolutionary change Keith proposed would be unique among the higher vertebrates, and therefore unlikely. But Keith, the anatomist, was untroubled by these niceties of paleontology and quite prepared to believe that mankind is unique. Thereafter his concept of true man's  exceptional antiquity became a point of faith. He called upon it to
explain the anomaly of apparently modern human remains found beneath ancient deposits at Ipswich 13 and, at a meeting of the British Association for the Advancement of Science held at Dundee in 1912, Arthur Keith offered perhaps the first full and authoritative exposition of the belief that mankind had acquired both large brain and erect stature a very long time ago and has remained relatively unchanged ever since.
Remarking upon the Neanderthal and Java fossils, Keith told his distinguished audience: 'thus we have a knowledge a very imperfect knowledge of only two human individuals near the beginning of the Pleistocene period. The one was brutal in aspect, the other certainly low in intellect.' If these are the ancestors of modern man, he said, then we have to accept 'that in the early part of the Pleistocene, within a comparatively short space of time, the human brain developed at an astounding and almost incredible rate'. To Keith it seemed more reasonable to reject Neanderthal and Java from man's ancestry and assume they were simply contemporaries and cousins of true man's large-brained ancestor, whose evolutionary development had occurred long before. 'Is it then possible,' he asked, 'that a human being, shaped and endowed as we are, may have existed so early as the Pliocene?'' Briefly reviewing the evidence he had presented, Keith concluded that it was. 'The picture I wish to leave in your minds,' he said, 'is that in the distant past there was not one kind but a number of very different kinds of men in existence, all of which have become extinct except that branch which has given origin to modern man. On the imperfect knowledge at present at our disposal it seems highly probable that man ' 14 At the same meetings, Grafton Elliot Smith confidently extended the pedigree of man's ancestors back to the Eocene, explaining how the 'steady and uniform development of the brain along a well-defined course throughout the Primates right up to Man ... gives us the fundamental reason for "Man's emergence and ascent"'. 15
Thus the leading anatomists of the day expounded the theories of human evolution that they hoped paleontologists might one day substantiate with the evidence of fossil human remains. just a few months later, in December 1912, Arthur Smith Woodward, perhaps the leading paleontologist of the day, unveiled Piltdown Man and thus presented the anatomists with a conundrum in which the expectation of theory and the logic of observed fact were wonderfully counterpoised.
Though he was responsible for the reconstruction and presentation of  Piltdown Man, Smith Woodward had not discovered the remains. The first pieces of fossil skullbones were brought to him by Charles Dawson, an amateur geologist who had already contributed important paleontological specimens to the British Museum collections. Dawson (1864-1916) was a solicitor practising at Uckfield in Sussex, a position which left time for a wide range of other activities (in 1909 he published A History of Hastings Castle in two volumes and at the onset of his terminal illness was investigating a case of incipient horns on the head of a carthorse) but geology and paleontology were the interests he pursued most energetically. In 1885 at the age of twenty-one Dawson was elected a fellow of the Geological Society for his contributions to the science, though little appeared in the literature under his own name. 'He preferred to hand over his specimens to experts who have made a special study of the groups to which they belonged,' an obituary recounts, 16 rather than describe them himself, and Smith Woodward was surely rewarding this respectful deference to greater knowledge when he named Piltdown Man Eoanthropus dawsoni (Dawson's Dawn Man), made Dawson principal author of their joint paper 17 and let him present the geological and archaeological evidence of the discovery while he, paleontologist Smith Woodward, spoke first on the anatomy of the specimen and then on the significance of the faunal remains found in the same deposit.
The details of Dawson's initial discovery are imprecise; not even the exact year is known. In the paper read at a meeting of the Geological Society on 18 December 1912, Dawson told how he had been attracted to the site 'several years' before when he traced some unusual brown flints found on a farm near Fletching to a small gravel pit on a farm adjacent to Piltdown Common, Sussex. He asked the labourers who occasionally worked there to keep any fossils they might encounter, and on a subsequent visit, they presented him with a small, concave, tabular object. It was part of a coconut shell, the men said, which they had found and broken in the course of their digging. They, had kept one piece for Dawson and discarded the rest. Dawson, however, realized that the object was part of a fossil human skull. Thereafter he visited the site frequently and 'some years later in the autumn of 1911' - he found another fragment of the same skull among the spoil heaps. Believing the pieces might match the proportions of the Heidelberg jaw, in May 1912 Dawson took his finds to the British Museum for more accurate assessment. Smith Woodward was impressed with the importance of the discovery and joined Dawson in the search for more remains that summer though only as a private, weekend holiday affair, and without the  involvement of any British Museum resources.
A DISCOVERY OF SUPREME IMPORTANCE TO ALL
INTERESTED IN THE HISTORY OF THE HUMAN RACE
Artist's portrayal of Piltdown Man with the original specimens and some of Dawson's many letters to Smith Woodward
. Apparently they wished to keep the discovery wholly to themselves. At first, just one labourer was employed to do the heavy digging in the small pit, but later they were joined by Father Teilhard de Chardin and a colleague, both French priests then studying at the Jesuit College near Hastings. Teilhard de Chardin shared Dawson's amateur interest in fossils and geology: they had met by chance in their rambles about the Sussex countryside. In later years Teilhard de Chardin became an authority on the fossil evidence of early man.
During the summer of 1912 solicitor, paleontologist and priests scrutinized every spadeful dug from the pit and sifted through all the spoil heaps of previous years. In one heap they found three pieces of the right parietal bone from the skull one piece on each of three successive days and later Smith Woodward found another fragment which fitted the broken edge of the occipital and connected with the left parietal found by Dawson. 'Finally, on a warm summer evening after an afternoon's vain search,' Smith Woodward recounts,` 'Mr Dawson was exploring some untouched remnants of the original gravel at the bottom of the pit, when we both saw half of the human lower jaw fly out in front of the pick-shaped end of the hammer he was using. Thus was recovered the most remarkable portion of the fossil which we were collecting.' In addition the pit supplied three artefacts, fragments of an elephant tooth, some beaver teeth and one much-rolled fragment of a mastodon tooth, while on the surface of an adjacent field the party found a piece of red deer antler and a horse's tooth both fossilized and both presumed to have been thrown over the hedge by the workmen. In all, a remarkably comprehensive haul.
According to Dawson's determination, the Piltdown gravel bed lay about eighty feet above the level of the River Ouse, deposited there before the river began excising the valley through which it presently flows. The gravel bed could be divided into four distinct strata, he said, and the fossils had come from the third, which lay about three and a half feet below the land surface and was distinguished by a dark ferruginous appearance and the presence of ironstone. The fossil fauna from the pit were of early Pleistocene even Pliocene age, and the flint artefacts appeared to be similarly ancient all of which strongly implied that the human remains found in the same deposit represented the earliest known example of true man.
Arthur Smith Woodward completed an anatomical reconstruction of the remains sometime during the autumn of 1912. He worked alone, apparently without the advice, assistance or even knowledge of his  colleagues at the British Museum. 19 He made a cast of the interior of the reconstructed skull and invited Grafton Elliot Smith to comment upon the brain of Piltdown Man when the specimen was unveiled at the Geological Society In December. but Elliot Smith has made it clear that he did not help with reconstruction of the skull. 20
The audience that packed the Geological Society's lecture room on 18 December 1912 was larger than any before, and the Piltdown skull they had come to see perfectly fulfilled the expectations of many who attended. Smith Woodward's reconstruction had produced a creature with the jaw of an ape, the skull of a man and a cranial capacity (1070 cubic centimetres) appropriate to an intermediate stage between man and ape. A small minority thought the combination of ape and human characteristics was just a little too good to be true, and Smith Woodward himself accurately defined the point when he told the meeting: 'while the skull, indeed, is essentially human, only approaching a lower grade in certain characters of the brain [as described in Elliot Smith's contribution to the same meeting] . . . the mandible appears to be almost precisely that of the ape'. 21 Elliot Smith, however, did not find the combination beyond the bounds of reason. The Piltdown brain was the most primitive and the most simian so far recorded, he said, and its association with an ape-like jaw was not surprising to anyone familiar with recent research into the process by which the human brain had evolved from the brain of the ape.' 22
Arthur Keith, on the other hand, was among those who had misgivings. He was jealous that the remains had been given to a paleontologist at the British Museum and not him, an anatomist known to be specially interested in the anthropological history of the British. But then he knew that Smith Woodward regarded his belief in the antiquity of Homo sapiens as 'an amusing evolutionary heresy'. 23 At the Geological Society meeting, Keith hailed Piltdown as perhaps the most important discovery of fossil human remains ever made, but took exception to some aspects of the reconstruction. The chin region and the form of the front teeth were too much like the chimpanzee, he said. 24
The arguments concerning the association of jaw and skull, and the form of the reconstruction. would never have arisen if the remains had been more complete. But the problem was, of course, that the anatomical features capable of proving or confounding the association of jaw and skull were exactly those missing from the original specimens the chin region of the jaw which would have clearly demonstrated the form of the canines; and the articular knob which would have shown the form of the skull joint. In June 1913 Arthur Keith acquired casts of the remains and  made his own reconstruction of the Piltdown skull, one which assembled the available parts in accordance with anatomical principles and supplied the missing parts in accordance with his belief in the antiquity of Homo sapiens. The chin and front teeth were entirely human and the cranial capacity was 1500 cubic centimetres greater than the average for modern man. A few weeks later the conflicting interpretations of anatomist and paleontologist were reviewed by the participants at an International Congress of Medicine held in London during August. At the British Museum the distinguished gentlemen of medical science viewed Smith Woodward's anatomical reconstruction of the original remains, and at the Royal College of Surgeons Arthur Keith showed them where the paleontologist had gone wrong.
Because of a misconceived notion of the nature of the jaws and teeth in fossil man, Smith Woodward had found a chimpanzee palate and jaw on a skull that could not possibly carry them, Keith said, with the result that the upper joints of the backbone were so close to the palate that there was no room for windpipe or gullet and Piltdown man would have been unable to either eat or breathe. The Smith Woodward reconstruction was anatomically impossible, he said.` If skull and jaw truly belonged together then the skull must be large enough to accommodate the massive
jaw and the front teeth must be human enough to match the articular capacity.
In the course of the ensuing discussions Grafton Elliot Smith lent his support to the paleontologist's reconstruction, but Smith Woodward did not respond publicly to Keith's challenge until 16 September 1913, when he told a meeting of the British Association for the Advancement of Science assembled in Birmingham of further discoveries at Piltdown.
Fortunately, Mr Dawson has continued his diggings during the past summer,' he said, 'and, on August 30, Father P. Teilhard, who was working with him, picked up the canine tooth which obviously belong to the half of the mandible originally discovered. In shape it corresponds exactly with that of an ape, and its worn face shows that it worked upon the upper canine in the true ape-fashion. It only differs from the canine of my published restoration in being slightly smaller, more pointed, and a little more upright in the mouth. Hence, we have now definite proof that the front teeth of Eoanthropus resembled those of an ape, and my original determination is justified. 26
The Piltdown canine settled the argument about the chin so conclusively in Smith Woodward's favour that Arthur Keith was forced to concede an important point of his belief in the antiquity of Homo sapiens.
The Piltdown canine rests on an Illustrated London News photograph of Dawson (left) and Smith Woodward at the Piltdown excavation
. Despite the evidence of Galley Hill and Ipswich, it seemed that some of man's ancestors in the early Pleistocene had displayed distinctly ape-like characteristics after all. But the argument about the size of the Piltdown brain-case was still unresolved. Smith Woodward believed it must have been small because, in his view, the association of the primitive jaw with a large brain-case would be a most improbable combination of primitive and modern features. Keith, on the other hand, insisted that the brain must have been large because the association of a large jaw with a small brain-case was anatomically impossible. The 300 cubic centimetres separating the estimates of anatomist and paleontologist soon became a
subject of heated debate in the pages of Nature and elsewhere. But Smith Woodward was only slightly involved; in his stead, Grafton Elliot Smith promoted the smaller estimate and thus the paleontologist was able to remain a passive observer while the merits of his Piltdown reconstruction were argued by two anatomists. In Keith's view the anatomical errors responsible for the small brain-case were manifest and glaringly obvious; he was surprised and irritated by Elliot Smith's refusal to acknowledge them. Acrimony developed between the two men and, although Keith eventually came to terms with most of the colleagues with whom he argued during his career, he and Elliot Smith were never friends again.
The difference in the estimates of the Piltdown brain size resulted solely from the manner in which the existing fragments were assembled and their curves projected to delineate the parts missing in between. A large area of the forehead, and the roof of the skull were among the 27 important missing pieces, as Keith pointed out in a letter to Nature, but he believed it was possible to reconstruct a complete skull from what remained of it by applying the principle of symmetry. 'The right and left halves of the mammalian head and skull are approximately alike,' he continued, but Dr Smith Woodward's reconstruction showed a 'great discrepancy between the right and left halves', 28 the extent of which was largely responsible for Smith Woodward's low estimate of brain size if the symmetry of the paleontologist's reconstruction was restored by adding to the right side of the skull, then the cranial capacity was substantially increased.
In reply, Elliot Smith stressed the position of the middle line of the skull as the most important factor in reconstructing a skull, and attributed Keith's alleged misplacement of this feature to his having worked from casts of the Piltdown fragments without reference to the original speci mens." He made little mention of the asymmetry of the skull, though it Is interesting to note that such a feature was fundamental to his subsequent theory that the leftor right-handedness of a person is shown in the relative sizes of certain parts of the left and right hemispheres of the brain. 30 A few weeks later Keith pointed out that Elliot Smith's remarks did less than justice to Mr F. 0. Barlow, who had made the casts, and even reflected badly upon the conduct of Dr Smith Woodward who, he wrote. had permitted 'the freest access to the specimens', even to those who like himself 'regarded the original reconstruction of the skull and brain cast as fundamentally erroneous'. 31
Keith championed his views in The Times and challenged Grafton Elliot Smith's conclusion at a memorable meeting of the Royal Society where he claims to have earned the reputation of a brawler, but behind the public controversy he was, by his own admission, 32 beginning to have private doubts about the validity of the argument. The problem he saw was quite simple: can a skull ever be reconstructed accurately from just a few fragments of the original? Such skills obviously were essential to anyone aspiring to study the progress of human evolution because most of the evidence was contained in fragmentary fossil remains. Several authorities claimed to possess the skills needed to reassemble such fragments, but in the case of Piltdown Man, their differing results begged the question: is there, or is there not, a science of fossil reconstruction?
To test his own skills scientifically, Keith arranged that some colleagues should cut fragments exactly duplicating the Piltdown remains from a modern skull in their possession, which he would then reconstruct according to his anatomical principles and afterwards compare with a cast of the original. The results were close enough to restore Keith's confidence in his skills and re-confirm his belief that Piltdown Man had possessed a large brain; the experiment was described at a meeting of the Royal Anthropological Institute and published in a scientific manner. 33 But nevertheless, the reconstruction was wrong in one important respect: Keith failed to reproduce the proper form of the forehead, of which there was almost no evidence at all among the Piltdown remains
While the deliberations concerning the form and size of the Piltdown skull were exercising the skills of the experts in London, Charles Dawson continued the search for more remains. The gravel pit seemed to have been worked out, and in wandering further afield, he examined newly ploughed land and the heaps of stones raked from the fields. His endeavours were attended by extraordinary good fortune. Sometime before 20 January 1915, in a field about two miles from the site of the original discovery,  Dawson found a fragment of fossil bone which, he was certain, had belonged to a second Piltdown Man. The fragment was a piece of the forehead, retaining a portion of the eyebrow ridge and the root of the nose. In July 1915 he found a molar tooth at the same site; and on another occasion a piece of the back of the skull.
Unhappily, Dawson became seriously ill with anaemia in the autumn of 1915. The condition worsened and turned to septicaemia, of which he died in August 1916. His last discoveries were presented before the Geological Society by Arthur Smith Woodward in February 1917, where it was concluded that the new evidence must support the contention that Eoanthropus dawsoni was a definite and distinct form of early man, as originally supposed, for, as Smith Woodward pointed out, the occurrence of the same type of bone with the same type of molars in two separate localities must add to the probability that they belonged to one and the same species. 34 The frontal bone revealed the form of only a small area of the interior surface, and one devoid of obtrusive features, but nonetheless Grafton Elliot Smith found that its evidence corroborated his opinion that the Piltdown skull presented features 'more distinctly primitive and ape-like than those of any other member of the human family at present available for examination '. 35 Arthur Keith attended the meeting too. and in the ensuing discussion effectively abandoned his opposition to the specimen. He never accepted the small brain Smith Woodward postulated, but he succumbed to the persuasive logic of the amazing discoveries and their presentation. The new Piltdown finds 'established beyond any doubt that Eoanthropus was a very clearly differentiated type of being', Keith said; adding that the frontal bone was particularly valuable because it cleared up any doubt as to the contour of the forehead. 36
The triumvirate of British paleoanthropological science was now united in its belief that the Piltdown remains represented the earliest known ancestor of Homo sapiens, a unique link between mankind and the ape-like creatures from which he had evolved. And did their knighthoods (Sir Arthur Keith 1921; Sir Arthur Smith Woodward 1924; Sir Grafton Elliot Smith 1934) reflect a shade of patriotic pride colouring this conviction that the ancestor of man was an Englishman?
But while the three experts were gathering around Piltdown Man, the creature was still under attack from other quarters. And the attacks were concerned not simply with the size of the skull, but with the more fundamental question of whether or not the jaw and skull belonged together. Arthur Smith Woodward, it will be remembered, had drawn attention to this problem when the remains were first presented to  science; and, at that same meeting, Professor Waterston had remarked that it was very difficult to believe that the two specimens could have come from the same individual. 37 Later Waterston strengthened his opinion; in a letter to Nature he wrote: 'it seems to me to be as inconsequent to refer the mandible and the cranium to the same individual as it would be to articulate a chimpanzee foot with the bones of an essentially human thigh and leg.` 38
Similar views were voiced elsewhere. In America Gerrit Miller compared casts of the Piltdown fossils (he never saw the originals) with the corresponding parts of twenty-two chimpanzees, twenty-three gorillas, seventy-five orang utans and a series of human skulls, concluding that 'a single individual cannot be supposed to have carried this jaw and skull' without assuming 'the existence of a primate combining brain case and nasal bones possessing the exact characters of a genus belonging to one family, with a mandible, two lower molars and an upper canine possessing the exact characters of another' without any blending of their distinctive characteristics. Miller concluded that the remains must represent two individuals despite the amazing coincidence of their
discovery in such close proximity, and created a completely new species of chimpanzee (Pan vetus) to accommodate the peculiarities of the Piltdown jaw. 39
In 1921 The French authority on fossil man, Marcellin Boule, approached 'the paradoxical association of an essentially human skull with an essentially simian jaw' with the question: 'Is Eoanthropus an Artificial and Composite Creature?' Boule considered the evidence and concluded that Piltdown Man was at least composite (if not artificial). The jaw had come from a chimpanzee, he said, and the skull was human but had belonged to a race of men quite distinct from the Neanderthals and closely related to the ancestry of modern man. 40
Many authorities in America and France supported the views put forward by Miller and Boule; objections to the paradoxical association of ape jaw and human skull were also raised in Italy and Germany. In all the anatomical evidence seemed so strong as to be incontrovertible; indeed, if the jaw and skull had been discovered in separate excavations no expert would have dreamt of suggesting they belong to the same species, but at Piltdown the scientific evidence of anatomy collapsed against the circumstantial evidence of paleontology (or so it was said) and therefore must belong together; and the discovery of matching remains some distance away supported the conclusion. As Grafton Elliot Smith remarked: there was no reason to assume 'that  Nature had played the amazing trick of depositing in the same bed of gravel the brain-case (without the jaw) of a hitherto unknown type of early Pleistocene Man displaying unique simian traits, alongside the jaw (without the brain-case) of an equally unknown Pleistocene Ape displaying the human traits unknown to any Ape'. 41 Rational minds found it much more likely that the jaw and skull belonged together especially since the creature thus formed so closely resembled the form of man many authorities believed had existed at that stage of human evolution.
In 1922 Elliot Smith, in conjunction with a colleague, made another reconstruction of the skull, supplementing those by Keith and Smith Woodward. The cranial capacity was 1200 cubic centimetres, its form was more in keeping with the structure of the jaw and Marcellin Boule, for example, found Elliot Smith's contribution persuasive. The new facts should eliminate or at least lessen the 'anatomical paradox', he wrote; expressing the view that the balance of the argument now inclined more towards Smith Woodward's theory. Boule was glad of this, he said, for he esteemed 'both the knowledge and the personal attributes of this scientist'. 42
There can be no doubt that he prestige and status of Arthur Smith Woodward were very largely responsible for the degree of acceptance that Piltdown Man achieved. His pronouncements on the fossils constituted a very small part of his work, as we have seen, but they carried the ring of
conviction. Sir Arthur Keith has remarked that Smith Woodward liked to set a puzzling specimen on a table where the light from a window caught it at all hours of the day, so that as he passed and re-passed it in the course of his work, a chance glance might reveal aspects he had not seen before and the significance of the fossil would gradually become clear. The Piltdown specimens were afforded this treatment. 43 so we must assume the paleontologist was satisfied with what he saw.
In the early stages of the debate Elliot Smith had said examination of the originals was essential to correct interpretation. Indeed opposition stemmed mainly from those who had dealt with casts only, and several sceptics who subsequently handled the originals are known to have changed their minds. The American-based anthropologist, Ales Hrdlicka, was one of these. Having examined the originals extensively he remarked on the great difference that exists between the study of a cast and its original. 'It is very probable,' he reported, 'that . . . some of the conclusions arrived at by some authors would not have been made had they been able to study the jaw itself.' Hrdlicka accepted Smith Woodward's designation of the remains as 'a being from the dawn of the human period' .44
 One Sunday morning in July 1921 another sceptic, Henry Fairfield Osborn, President of the American Museum of Natural History, spent two hours after church examining the Piltdown remains. He concluded that 'paradoxical as it had appeared to the sceptical comparative anatomists, the chinless Piltdown jaw, shaped exactly like that of a chimpanzee . . . does belong with the Piltdown skull, with its relatively high. well-formed forehead and relatively capacious brain case.' 45
But behind the controversy, it must be noted, there was unanimous agreement among the experts on a point which had a most pervasive effect, particularly in respect of the beliefs and predispositions that were presented to the anthropology students of the day. The experts may have disputed the association of the Piltdown jaw and skull, they may have argued about the absolute size of the brain, but of one thing they were all certain. the Piltdown remains proved beyond doubt that mankind had already developed a remarkably large brain by the beginning of the Pleistocene. And the implications of this were very important firstly, a brain so large at that time must have begun its development long before, which implied that true man was very ancient indeed; and secondly, since the Piltdown remains of this 'true man' were older (as it was believed) than the Java and Neanderthal fossils. they firmly dismissed those small-brained, 'brutish' creatures from the human line to the status of 'aberrant offshoots' evolutionary experiments that led to extinction cousins of mankind perhaps, but not ancestors.
Thus Piltdown Man contributed to the intellectual climate of the 1920 and 1930s when some significant discoveries were scorned because they did not conform with accepted beliefs, while others, less accurately founded, were welcomed because they conformed only too well. Regardless of the continuing controversy over their details, the Piltdown remains became a standard requiring mention in related literature, and a measure against which subsequent discoveries had to be compared. This Piltdown effect, as it might be called, is well demonstrated in the work of Louis Leakey (1903-1972), who studied anthropology at Cambridge during the 1920s and was an admiring disciple of Sir Arthur Keith.
In 1934, Leakey published a popular book on fossil man called Adam's Ancestors. In it he supported Keith's appraisal of the Galley Hill remains and described Piltdown Man as a good candidate for the ancestry of man (he took care to explain why the specimen had not been presented in the preceding chapter on 'Our Stone Age Cousins'). 'The Piltdown skull is probably very much more nearly related to Homo sapiens than to any other yet known type,' Leakey wrote, and would have granted the specimen full ancestral status if it had been 'vastly more ancient' than the  Kanam mandible he had recently found in East Africa and which, he believed, must represent the oldest ancestor of true man. 46
We have concentrated so far on the anatomists' and paleontologists' views of Piltdown Man, but a third approach was available to science geology and it was from this direction that the riddle eventually was solved. Charles Dawson had said that the remains were found in a gravel bed lying about eighty feet above the level of the River Ouse. This implied an antiquity not much less than that of other river terraces in Britain and Europe, and indeed, on the evidence of the extinct fauna they were said to contain the Piltdown gravels could hardly have been younger. Thereafter Dawson's estimate was repeated as fact by other authorities and, in particular, gained considerable respect from the support of W. J. Sollas, Professor of Geology at Oxford who even improved upon the Dawson estimate.
In his book, Ancient Hunters (1924), Sollas converted eighty feet to twenty-five metres, bracketed twenty-five with thirty and thus correlated the Piltdown gravels with those lying on terraces thirty metres above other rivers (the Thames for instance), concluding that the Piltdown remains must, therefore, date from the early Pleistocene. Thereafter, Sollas's assessment became the most authoritative reference on the geology and age of the Piltdown deposits. 'Thirty metres'
was frequently converted to 'one hundred feet' and in turn offered to support a contention that the fossils might be even older than originally thought, perhaps even of Pliocene age though the only Pliocene deposits known from that part of England were of marine origin and above the five hundred-foot contour. All of which undoubtedly helped to obscure the fact that Dawson's original estimate was based on an erroneous assumption.
Dawson had claimed the Piltdown gravels were part of a plateau lying above the one hundred-foot contour line, and he had calculated their height in relation to these features. But the gravels are actually part of a larger, well-defined terrace which maintains a constant height of fifty feet above the River Ouse throughout its extent. The portion in which the fossils were found is no exception, as was clearly revealed on the six inch to one mile Ordnance Survey map of the district published in 1911. If this had been noted in 1913, and the stratigraphy of the area accurately ascertained, it may have seemed more correct to correlate the Piltdown gravels with the fifty-foot terraces of the River Thames rather than with anything older, in which case the Piltdown fossils could only have been of Late, not Early Pleistocene. In such a scenario interest in the remains probably would have evaporated very quickly the fossils might have seemed odd and anomalous, but no one could have claimed any great  antiquity for them, especially since Smith Woodward had always said that the skull was the same age as the deposit in which it had been found.
But the error was not noted in 1913; and it drew no comment in 1926 when a map giving the correct elevation appeared in a Geological Survey publication with text repeating Dawson's estimate. 47 In fact, the error and its significance was first mentioned only in 1935, when the attributes of the Piltdown skull seemed difficult to reconcile with those of another skull found at Swanscombe. The Swanscombe specimen came from gravels of the one hundred-foot terrace of the River Thames itself, and when the problem of its correlation with the Piltdown remains arose at a meeting of the British Association at Norwich, Kenneth Oakley, a geologist at the British Museum, challenged Sollas's assertion that the Piltdown gravels were part of the thirty-metre terrace. He draw attention to the 1926 map of the area and its author's observation that the deposits more satisfactorily corresponded with those of the fifty-foot terrace.
The Swanscombe remains were found by Alvan T. Marston, a dentist with an interest in fossils, in a gravel pit not far from the site of the Galley Hill discovery. They comprise the rear half of a skull; there is no clue whatsoever to the form of the face, the law or the forehead. The cranial capacity was estimated to be 1325 cubic centimetres and comparative anatomists could find little to distinguish the specimen from Homo sapiens. Yet the Swanscombe skull had come from deposits no younger than those at Piltdown, and the Piltdown skull was held to be so old and so distinct from Homo sapiens as to merit the creation of a new genus, Eoanthropus. Clearly something was amiss. The geology at Swanscombe was well documented, and the Swanscombe skull's affinities were well defined, so Marston concluded that the fault must lie with the Piltdown specimen. The jaw must have belonged to an ape, he said, and the skull must have belonged to a man more recent
than even the Swanscombe remains whatever the circumstances of the discovery. Marston summarized the problem in 1937: 'that the Swanscombe skull had to be considered in its relations to the Piltdown was inevitable, and once this was embarked upon the gross inconsistencies of the large-brained, Pliocene, ape-jawed, eolithic medley became apparent. The relegation of the Piltdown skull to a later date will remove the disharmony which has occasioned so much difficulty for those who have tried to describe it as an early Pleistocene type'. 48
By the 1930s, fossil evidence accumulating from other parts of the world seemed to suggest that the brain had not led the way in the evolution of mankind. Fossils of no less antiquity than was proposed for Piltdown Man revealed distinctly man-like jaws and teeth, while the  brain remained relatively small, so that it became increasingly difficult to reconcile the large brain and ape-like jaw of Piltdown Man with a reasonable interpretation of mankind's evolution as suggested by the new evidence. Scientific papers repeatedly drew attention to the differences rather than the similarities between Piltdown and the new fossils. The triumvirate of British anthropology (they became two with the death of Sir Grafton Elliot Smith in 1937) remained convinced of the specimen's validity, and regarded the new evidence as proof of the theory that two lines (at least) of hominid evolution had once co-existed, the surviving line represented by modern man, Piltdown and little else, while all the new discoveries represented lines that had led to extinction. To other authorities, however, Piltdown simply did not fit; the specimen was a chimaera, a once-intriguing riddle about which there seemed little more to be said.
But the truth if discernible at all must be contained in the evidence. The question was: could it ever be extracted? Oakley actually had begun to answer this question in 1935 when he had referred to the observation that Dawson had estimated the height of the Piltdown gravels incorrectly. The implications of Dawson's error were clear: the age of Piltdown Man was derived solely from its association with extinct fauna of the early Pleistocene found in the same pit; but if the deposits were younger than had been claimed, then the older fossils must have come from somewhere else and there could be no compelling reason to believe that all the Piltdown fossils were of equal antiquity. The extinct fauna was indisputably older than the deposit, but Piltdown Man could be the same age or even younger. The next question was: is there some way of determining whether or not bones found close together in a single deposit are actually the same age? This question became a subject of Kenneth Oakley's research programme after the war, and although the solution of the Piltdown riddle was not his specific interest, that was an important result of his endeavours.
Fossils absorb fluorine from the soil in which they are buried, and Oakley's research explored the observation that the amount of fluorine in a fossil steadily increases with time and therefore might give some indication of its geological age. The phenomenon had been noted by J. Middleton in 1844, who remarked that the 'accumulation of fluorine seems to involve the element of time, so interesting to geological investigations' 49 and attempted to establish a timescale based on fluorine content by which the absolute age of fossils could be determined. Taking the quantity of fluorine in a bone of an ancient Greek known to be 2000 years old as his standard, Middleton dated fossils from the Siwaliks at  7700 years, and an extinct pachyderm at 24,200 years age estimates which then must have seemed a trifle ungenerous. Furthermore, Middleton had overlooked that fact that, because the soil's fluorine content varies considerably from place to place, fossils found in different deposits are likely to have absorbed quite different amounts and therefore cannot be dated one against the other reliably on this basis.
Middleton's observations were not pursued, and the significance of fluorine in fossil bones slipped into obscurity until it was discovered anew by Adolf Carnot, a French mineralogist. In 1892 Carnot published tables showing the increasing amounts of fluorine in fossil bones from progressively more ancient deposits; the following year he reported on the fluorine contents of a fossil mammoth bone and a human bone from the same deposit they were different, he said, and therefore the bones must be of different ages. 50 This observation was fundamental to the potential of fluorine content as a means of dating fossil bones. It could never provide an absolute timescale, such as Middleton had sought, but it could provide a useful relative scale and, furthermore, a means of assessing whether or not bones found together in a deposit had all been there for the same length of time. But even Carnot's work passed unnoticed; the principle of fluorine dating again slipped into obscurity, and there it remained for fifty years until it was discovered anew by Kenneth Oakley.
In 1943, while Oakley was assessing Britain's phosphate resources, a colleague working on fluorosis (another of the country's wartime problems) showed him Carnot's 1892 tables giving the fluorine content of fossil bones. Oakley realized that Carnot's work suggested a means of comparing the age of fossils within a single deposit, and for several years thereafter believed this primary observation on fluorine dating was his alone. Only after he had refined and tested his methods did he learn of Carnot's 1893 paper, and later still of Middleton's work.
The quantity of fluorine absorbed by a fossil is never large, even in the oldest bone, and measurement involves complicated chemical analysis. At the instigation of Oakley and the British Museum, preliminary trials were conducted by The Government Chemist during 1948 to establish the most satisfactory procedure, and the perfected fluorine dating method was used to assess the
relative age of a fossil assemblage that same year. For this first ever test, Oakley and his colleague M. F. A. Montagu selected the Galley Hill skeleton. The results profoundly contradicted Sir Arthur Keith's belief in the great antiquity of the specimen.
Briefly, Oakley and Montagu showed that the fossil fauna from the Middle Pleistocene gravels contained about two per cent fluorine, those  from Upper Pleistocene deposits in the same sequence about one per cent, and the post-Pleistocene bones not more than 0.3 per cent. The Galley Hill skeleton which. it will be recalled, had been found in the oldest gravels contained only about 0.3 per cent fluorine. Therefore it matched the post-Pleistocene bones and cannot have been older; the skeleton must have been of recent origin. entombed in the ancient deposits by man, not nature. despite Keith's assertions to the contrary. The antiquity of the Swanscombe skull, on the other hand, was confirmed by the fluorine test; the bones contained two per cent fluorine, perfectly matching the Middle Pleistocene fauna with which it was associated.
So the antiquity of Galley Hill Man was dismissed. and of Swanscombe Man confirmed; where did that leave Piltdown Man, with his combination of 'ancient' ape-like jaw and 'recent' large brain? In October 1948 Kenneth Oakley was authorized to apply his fluorine dating method to the Piltdown material, the Keeper of Geology at the British Museum having deemed it likely that the results might help resolve the riddle of its age and association. Every available bone and tooth from the assemblage was analyzed, thirty-six specimens in all, including ten pieces of the Eoanthropus material, and six of the extinct fauna from which the Lower Pleistocene age of Eoanthropus had been derived. The fluorine content of the entire assemblage ranged from a minimum of less than 0.1 per cent to a maximum 0f 3.1 per cent. The higher levels all were found in the extinct fauna, confirming their antiquity; while the remains of Piltdown Man contained an average of only 0.2 per cent, clearly showing that he was not as old as the Lower Pleistocene fauna with which he was supposed to have been associated. Eoanthropus dawsoni - Dawson's Dawn Man was probably no older than Homo sapiens from Galley Hill, it seemed.
Oakley's results were published in March 1950. 52 'That the figures scarcely provide any differentiation between Eoanthropus and the recent bones requires some explanation,' he remarked, but the deeper implication of his observation was not fully realized until three years later.
Meanwhile, Oakley had simply added another twist to the riddle. The combination of jaw and skull was puzzling enough when both were believed to be of great antiquity, as we have seen. but if both stemmed from the recent past further problems arose. The skull was reasonable enough, but what about the law? No man could have possessed such an ape-like form at so late a stage in human-evolution. This may seem to have vindicated the contention that jaw and skull represented different individuals, but this in itself presented another problem: if the jaw did not belong to the skull, obviously it came from somewhere else. But where? The great apes are totally absent from the fossil record of Britain  and Europe, and highly unlikely to have inhabited the region during the upheavals of the Ice Age. So Oakley provided no comfort for either side of the Piltdown controversy.
Reconstructed skull of Piltdown Man. The brown parts represent the fragments found at Piltdown; the remainder is plaster reconstruction
 With the aging and passing of its protagonists the controversy had lost much of its impetus by 1950. Although Piltdown Man so completely contradicted the evidence of the small-brained hominid fossils with man-like laws subsequently found in Africa and the Far East, most anthropologists were content to consign the riddle to a 'suspense account' and await clarification, rather than actively search for it.
But one evening towards the end of July 1953 a chance remark by Kenneth Oakley prompted Joseph Weiner, an anatomist working with Professor le Gros Clark at Oxford, to ponder the problem again as he drove home. In the early hours of the morning he found the key to the riddle. If Piltdown Man was not as old as had been claimed, if the strange jaw with man-like features matched no apes living or extinct, if it belonged neither to the skull nor to the deposit in which the specimens were found, then all 'natural' explanations of the Piltdown phenomena were eliminated. Which left only an 'unnatural' alternative, reasoned Weiner. Could it be that the man-like features of the jaw were artificial, and the whole assemblage was deposited in the Piltdown gravels with the express intention of suggesting to its discoverers that man in the early Pleistocene had possessed an ape-like law and a large brain?
The proposition seemed outrageous, but as Weiner weighed the evidence, the case for a deliberate hoax gained strength on several counts. The discovery of the second Piltdown remains so precisely echoing the first some years before considerably lessened the likelihood of either or both being an accident. The fact that the chin region and the articular knob were missing strongly suggested that these critical diagnostic features had been deliberately removed. The pieces of the puzzle began to fall into place.
Next day, Weiner examined the casts of the Piltdown remains in the collection of the Department of Anatomy at Oxford and discussed his theory with Professor le Gros Clark. Even on the casts, the wear of the molar teeth seemed more compatible with artificial abrasion than natural; and similarly on the canine, where artificial abrasion would also explain the apparent paradox first noted by a dentist in 1916 54 of such excessive wear on such an immature tooth.
Weiner and le Gros Clark took their case to the British Museum, and during the autumn of 1953 the riddle of Piltdown Man was finally resolved forty-one years after it had first arisen. The teeth confirmed Weiner and le Gros Clark's preliminary observations. Further fluorine  testing revealed that the jaw was not just recent, but not long dead; the skull was slightly more ancient. The remains were all stained to match the Piltdown deposit, so too were the mammalian fossils with which they were associated. The hoax had been ingeniously planned, carefully carried out and completely unsuspected.
When the news was released in November 1953 it excited comment from many quarters. In the House of Commons a motion was put forward proposing a lack of confidence in the trustees of the British Museum 'because of the tardiness of their discovery that the skull of the Piltdown Man is partially a fake'. The proposers were angry at the 'sycophantic servility' of the museum tradition which had itself been playing a hoax on the public with this 'so-called Missing Link', they said, but the motion aroused more laughter than serious debate: Speaker 'not sure how serious the motion is (laughter), but sure [we] have many other things to do besides examining the authenticity of a lot of old bones' (loud laughter). Lord Privy Seal -'the government had found so many skeletons to examine when they came into office that there had not yet been time to extend the researches into skulls' (laughter)."
A letter to The Times asked: 'Sir, May we now regard the Piltdown Man as the first human being to have false teeth?"'
Humour may cover embarrassment satisfactorily, but it could never dispel the question: Who dunnit? Dawson, Smith Woodward, Elliot Smith, de Chardin, have all been accused by some and excused by others. Several books and many articles have been published on the subject; from time to time 'new' evidence is produced to throw new light, but so far there is no definite, incontrovertible answer. The evidence simply is not conclusive enough. A case could be made against each of the characters involved (Including some not mentioned in this chapter), but none would stand serious cross-examination and judgement would depend heavily upon the predispositions of the judge.
The inconclusive nature of the Piltdown affair reflects a fundamental problem of the science as a whole, for the fossil evidence of human evolution rarely offers just one clear interpretation. At the same time, however, the Piltdown affair makes two pertinent points: firstly. accurate geological and stratigraphical determinations are essential. And secondly, when preconception is so clearly defined, so easily reproduced, so enthusiastically welcomed and so long accommodated as in the case of Piltdown Man, science reveals a disturbing predisposition towards belief before investigation as perhaps the hoaxer was anxious to demonstrate.
1 KEITH, A. 1950 London: An Autobiography : 122,
2 KEITH, A. 1894 London: Journal of Anatomy, 28: 149-335.
3 as 1: 170.
4 as 1: 317.
5 DARWIN, C. 1871 London: The Descent of Man; 1922 printing: 80.
6 KEITH, A. 1912 London: The Human Body: 78.
7 SMITH, C. E. 1912 Dundee: Presidential Address, Anthropology Section,
Report of the British Association, 1912: 575-98.
8 WOODWARD, A. S. 1885 Macclesfield: Modern Ideas of the Creation, Macclesfield Courier and Herald, 28 March 1885.
9 WOODWARD, A. S. 1913 Birmingham: Missing Links among extinct animals, Report of
the British Association, 1913: 783.
10 KEITH, A. 1925 London: The Antiquity of Man; second edition: 15 8.
11 ibid 258..
12- ibid: 265.
13 a) MOIR, J. REID. 1912: The Occurrence of a Human Skeleton in a Glacial Deposit at
Ipswich, Proceedings of the Prehistory Society of East Anglia, 1: 194.
b) KEITH, A. 1915 London: The Antiquity of Man.
c) ANON. 1912 London: The Earliest Known Englishman, Illustrated London News, 140:
14 KEITH, A. 1912 Dundee: Modern problems relating to the antiquity of Man, Report of the
British Association, 1912: 758.
15 as 7: 577.
16 WOODWARD, A. S. W. 1916 London: Charles Dawson - An Obituary, Geological
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17 DAWSON, C. & WOODWARD, A. S. 1913 London: On the discovery of a Palaeolithic
human skull and mandible in a flint-bearing gravel overlying the Wealden (Hastings Beds) at Piltdown, Fletching, Sussex, Quarterly Journal of the Geological Society, 69: 117-44.
18 WOODWARD, A. S. 1948 London: The Earliest Englishman :11.
19 WEINER, J. S. 1955 London: The Piltdown Forgery: 120
20 SMITH, G. E. 1913 London: Preliminary report on the cranial cast [Piltdown skull],
Quarterly Journal of the Geological Society, 69: 145-7.
21 as 17: 134-5.
22 as 20: 147.
23 as 1 : 32-4.
24 KEITH, A. 1913 London: The Human skull etc. from Piltdown, discussion, Quarterly
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25 KEITH, A. 1913 London: Reported in The Times, 11 August 1913.
26 as 9: 786.
27 KEITH, A. 1913 London: The Piltdown Skull and Brain Cast, Nature, 92:197.
28 ibid: 198-9.
29 ibid. :267.
30 SMITH, G. E. 1914 London: The Evolution of Man: 184.
31 as 27:292.
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34 WOODWARD, A. S. 1917 London: Fourth note on the Piltdown gravel, with evidence of a
second skull of Eoanthropus dawsoni, Quarterly Journal of the Geological Society, 73: 1-10.
35 ibid: 8.
36 KEITH, A. 1917 London: Second Skull from Piltdown Gravel, discussion, Quarterly Journal of the Geological Society, 73: 10.
37 WATERSTON, D. 1913 London: The Human skull etc. from Piltdown;; discussion, Quarterly Journal of the Geological Society, 69:150.
38 WATERSTON, D.1913 London: The Piltdown Mandible, Nature, 92:319.
39 MILLER, G. S.1915 Washington: The jaw of Piltdown man. Smithsonian Miscellaneous Collections, 65: 1-31.
40 BOULE, M. 1923 Edinburgh: Fossil Men: 171 & 471.
41 SMITH, G. E. 1924 London: Essays on the Evolution of Man: 73.
42 as 40: 472.
43 KEITH, A. 1948 London: In a foreword to Woodward A. S., The Earliest Englishman.
44 HRDLICKA, A. 1930 Washington: The skeletal remains of early man, Smithsonian Miscellaneous Collections, 83: 65-90.
45 OSBORN, H. F. 1927 Princeton N.J.: Man Rises to Parnassus: 53.
46 LEAKEY, L. S. B. 1934 London: Adam's Ancestors: 221.
47 WHITE, H. J. 0. 1916 London: The Geology, of the Country, near Lewes, with map by Edmunds, F. Fl., Memoir Geological Survey of England and Wales. Expl. sheet 319.
48 MARSTON, A. 1937 London: The Swanscombe Skull, Journal of the Royal Anthropological Institute, 67: 394.
49 MIDDLETON, J. 1844 London: On fluorine in bones, its source, and its application to the determination of the geological age of fossil bones, Proceedings of the Geological Society, 4: 431-3.
50 CARNOT, A. 1893 Paris: Recherches sur la cotiiposition générale et la tenure en fluor des
os modernes et des os fossiles de different âges, Ann. Min. (9, Mem.) 3:155--95.
51 OAKLEY, K. P. & MONTAGU, M. F. A.. 1949 London: A re-consideration of the Galley Hill skeleton, Bulletin of the British Museum (Natural History), Geology, I, 2: 27-46.
52 OAKLEY, K. P. & HOSKINS, C. R. . 1950 London: New evidence on the antiquity of Piltdown man, Nature, 165: 379-2.
53 as 19:26-35.
54 LYNE, C. W. 1916 London: The significance of the radiographs of the Piltdown teeth. Proceedings of the Royal Society of Medicine, 9 (3 Odont. Sect.) : 33-62.
55 The Times, 1953 London: Parliamentary report 27 November 1953.
56 KRAMER, L. M. J. 1953 London: Letter to The Times, 28 November 1953.