Note: These materials are intended as supplements for students in Ant. 301. These pages are in development and will contain errors.
The Discovery of the Fossil
is a Prehistory
The stage was set in 1860 for
a debate over whether evolution did occur and whether or not humanity
had a prehistory. Some anti-evolutionists held prominent scientific
positions. Richard Owen (1804-1892), director of the natural science
section of the British Museum, became champion of Cuvier's catastrophism
in England. Owen was a prestigious zoologist and paleontologist,
indeed it was through Richard Owen's efforts that the natural
history collections were moved to a facility dedicated to biology,
the British Museum of Natural History. Though Owen deduced the
distinction between analogies and homologies, he viewed them as
part of a divine plan, not an evolutionary process. Similarly,
one of the greatest American naturalists, the Swiss-born Louis
Agassiz (1807-1873), demonstrated the existence of European Ice
ages, but this Harvard professor of zoology remained a strong
opponent of Darwinism. Darwin himself did not publish on human
evolution until 1871. The most articulate and aggressive defender
of the idea of human evolution in England was Thomas Henry Huxley
(1825-1895), author of Man's Place in Nature (1863), a book that
applied Darwinism to human evolution. There were many talented
scientists on both sides of the debate. Even Lyell, whose work
gave Darwin the gift of geologic time, could never be entirely
convinced of the transmutation of species inherent in Darwin's
Industrial development and
associated population growth in Europe provided a continuous influx
of discoveries of archaeological materials, fossil animals, and
human bones, a tide that no church dogma could stem. Nevertheless,
religious arguments stormed down upon Darwin with a vengeance.
A retiring individual, and not motivated by any anti-church bias,
Darwin retreated from controversy and refused public debates and
Geologists and amateur archaeologists
alike began to visit such sites as Kent's Cavern in Devon.
Broken stone tools lay scattered among the bones of mammoth and
other animals of the Ice Age. It was becoming abundantly clear
to the lay world in England that human beings were contemporaries
at one time with mammoth, rhinoceros, reindeer, and other extinct
In France, Edouard Lartet (1801-1871),
geologist and archaeologist, excavated thousands of bones and
artifacts from various caves. The sites that he and his colleagues,
Henry Christi and Hugh Falkner described gave names to lithic
cultures that they contained: Magdalenian, Mousterian,
Aurignacian, etc. Lyell used these archaeological discoveries
to argue that humans had lived on Earth far longer than previously
imagined in The Geological Evidence for the Antiquity of Man (1863).
Sir John Lubbock (1834-1913), an English prehistorian, published
a best selling book, Prehistoric Times (1865), in which he divided
the Stone Age into two parts, the Neolithic and the Paleolithic.
Though the Neolithic was not controversial, a date for the start
of the Paleolithic was debated during the next 100 years of archeology.
(return to outline)
Humans Have Modern Anatomy and Abilities
Louis Lartet, son of Edward,
made a vital discovery in the Dordogne Valley of France in 1868.
Workmen laying a railway line along the river valley dug into
deposits of an ancient rock shelter, exposing archaeological strata.
Lartet found bones of three adult males, a woman, and an infant
buried in deposits rich in archaeological materials stylistically
similar to those found in a cave at Aurignac. These became the
famous type specimens of early humans known as Cro-Magnon. It
was the scientists' first look at the anatomy of Paleolithic people,
and what a surprise! They were extraordinarily like modern Europeans.
They had none of the pongid-like characteristics that might have
been expected from the Darwinian ideas of evolution, but there
could be no doubt about their antiquity.
Lartet's discoveries pointed
to a new element in the understanding of prehistoric humans. He
found numerous shell necklaces and reindeer antlers engraved with
drawings of other animals. Prehistoric peoples were artists! His
most famous discovery of the period, a thin piece of ivory depicting
the head and body of an elephant drawn with intricate, shallow
lines, was clear proof not only of contemporary association between
humans and elephant, but evidence that Paleolithic people were
both intelligent and artistic.
Acceptance by a growing public
in Europe was evidenced by two events in Paris (the bastion of
catastrophism) in 1867: (1) the innermost gallery of the exhibition
at the World's Fair was devoted to exhibits from prehistoric periods
and (2) the first International Congress of Prehistory was held.
There were still many opponents, but a mainstream body of scientific
writing argued that fossil humans did exist. That realization
meant fundamental changes in Cuvier's France.
Darwin published another controversial
book in 1871 (The Descent of Man and Selection in Relation to
Sex). In it, he introduced a new concept, the idea of sexual selection,
that is, individuals of the same sex may compete with each other
in propagation of the species. He also gave a more extensive discussion
of his thoughts on human evolution. Humans, he argued, are members
of the animal kingdom and similarities in embryonic development,
anatomy, and intellect suggest that humankind descended from some
less highly organized form, most likely a catarrhine (Old World)
primate. Since humans most closely resemble African apes, Darwin
concluded that Africa was the likely birthplace of the human species.
Archaeological data continued
to unfold. Dr. Paul Rivière demonstrated the presence of
deliberate burials and ornamented clothing in 1872 at the Grimaldi
caves. Similar discoveries came from other sites, including Lartet's
old site across the river from the Cro-Magnon rock shelter. Cro-Magnon
activities began to look human like indeed with art work, the
practice of burial, ornamented clothing, and funeral rites.
One of the most exciting finds
of the 19th century was made by a Spanish nobleman, Sr. Don Marcelino
de Sautoula, on his estate on the northern coast of Spain. An
amateur prehistorian, he partially excavated a cave on a hill
called Altamira. He found stag, horse, bison bones, oyster shells,
numerous implements and bone tools characteristic of the Magdalenian
culture. Exploring deeper into the cave, he encountered cave bear
bones and extensive ashes. On one of his excavation trips, he
took his 12 year old daughter, Maria, with him into the cave.
While he sat digging in damp darkness under the light of a lantern,
the girl suddenly cried out, "Papa, papa. Mira. Toros pintados."
(Father, father. Look. Painted bulls.) Maria showed her astonished
father that the roof of the cavern was covered with a superb fresco
of red and black bison. Lantern light gave them three-dimensional
form since the artist had incorporated the natural projections
of the ceiling with great skill into his art. Sautuola was deeply
moved as he and his daughter explored the cavern roof and saw
marvelous polychrome animals, executed in three-dimensional skill,
all painted by an artist who had known the animals before they
became extinct from Spain of the distant past. Don Marcelino was
later to explore chamber after chamber of extraordinary drawings:
polychrome horses in flight, painted prints of human hands, outlines
of human hands. Here again was extraordinary evidence of the humanness
and artistic skills of Paleolithic artists. This was no childhood
art, but painting that merited attention itself as art. Don Marcelino
hired a French painter to make sketches of the art work so that
he could publish them. This well-written paper, An Account of
Certain Prehistoric Discoveries in the Province of Santander (1880)
included the paintings and a detailed description of the archeology
of the cave.
Don Marcelino was disappointed
with the skepticism and ridicule that the volume received. Everyone
knew that prehistoric humans were little more than apes, incapable
of finer art. Obviously the Don had perpetrated a fraud. Dispute
about the authenticity of Altamira continued for a decade, and
regardless of the damage it did Don Marcelino, it did turn attention
to deeper galleries of rock shelters and grottoes of France. In
1895, a few months after the death of Don Marcelino, Emile Riviera
found a series of paintings and engravings similar to those of
Altamira in the grotto of La Mouthe. Riviera by-passed the Prehistoric
Society of France that had resoundingly condemned cave art as
fraud, and took his discovery directly to the Academy of Science
in Paris. Altamira, validated by discovery of cave art in France,
was labeled the "Sistine Chapel of Prehistory." It attracted
thousands of visitors, including the King and Queen of Spain.
The Englishman's idea of a
Cro-Magnon life way at that point in time was hardly more sophisticated
than views of French or Spanish scientists. Cro-Magnons were commonly
portrayed in essays of that period as naked, living by strength
and wit, distinctly subhuman in character and abilities, and often
warring among themselves over possession of females (Darwin's
sexual selection). How did we know primitive peoples had no clothing?
No clothing meant no pockets. The absence of pockets accounted
for the vast quantities of stone tools cast about the countryside.
It seemed evident that Cro-Magnons had to make a new tool every
time they sat down to do something.
Lack of clothing posed another
problem. Without expensive clothing or badges, how is an important
person distinguished from a commoner? Tribal people were often
portrayed carrying a baton that could serve as a badge of status
among a society without clothing.
Perhaps the strangest element
in the attitude that developed toward prehistoric people at the
close of the 19th century was the idea that life was better then.
This myth, purported that prehistoric humans had no problems finding
food, they just plucked it from bush or tree. They had few worries;
were healthy; happy; and were free of the plagues and drudgery
of the modern world.
One prominent figure in German
biology at the close of the 19th century was Ernst Heinrich Haeckel
(1834-1919). Well known as a naturalist and evolutionist, he is
credited with coining the term "ecology." Darwinian
thought was applied to organic evolution in Haeckel's General
Morphology (1866) and The History of Creation (1867). Haeckel's
"biogenic law" emphasized the idea that ontogeny recapitulates
phylogeny, that is, that during embryonic development, an individual
fetus repeats the evolutionary stages through which its ancestors
passed. From the advantage of 20th century biology we know that
his idea is not literally accurate, but there are similarities
in embryonic development between embryos of closely related taxa.
Haeckel wrongly interpreted embryology as a literal reflection
of evolutionary history. He also turned the older idea of a "great
chain of being" and turned it upside down, arguing that it
was human vanity to presume our origins from gods or demigods.
"Just as most people much prefer to trace their family back to some degenerate baron of some famous prince rather than to an unknown peasant, so most men would rather have as parent of the race a sinful and fallen Adam than an advancing and vigorous ape." (The Evolution of Man, 1905, page 870)
Haeckel also argued that scientists must reject the idea of dualism, the idea that the body with its brain is separate from the soul. The soul, or "spark of divinity" (presumed in the late 19th century to be responsible for human reasoning abilities) was supposed to be an exclusive possession of humanity. He advocated monism, the idea that all mental and psychic activities are physiological functions of the brain and spinal chord. Further, there was no physiological or embryological basis for separating characters of a human brain from those of higher mammals. Both attributes, the unique human soul and human reason were, in Haeckel's view, untenable superstition. Indeed, he argued that intelligence evolved throughout the animal kingdom in the same fashion as other physiological characters. Huxley had pointed out that differences in organization between humans and apes are much less than corresponding differences between apes and "lower" primates. Thus the hypothetical Tertiary human ancestor was a catarrhine ape. Haeckel felt that the common ancestor at the branching between humans and contemporary apes was an early form of gibbon. He had been impressed by similarities between human and gibbon embryological features, and he thought the gibbon was nearer to this stem ancestor than the other apes. This hypothetical human ancestor, with general structure (especially in the limbs) as humans but lacking articulate speech and higher intelligence, he gave the name Pithecanthropus. (return to outline)
Another Human Species is Found
In 1886, Marcel de Puydt and Max Lohest found bones of three prehistoric skeletons and flints similar to those of Mousterian tool assemblages at Spy in Belgium. They called in other experts to verify that the human materials were in unquestionable geologic context and in clear association with mammoth and woolly rhinoceros bones. The importance of this discovery is that the anatomy of these remains was similar to the specimen from the Neander Valley discovery in 1857 (to which; the anatomist William King had given the name Homo neanderthalensis King, 1864). The Neanderthal skull was thus validated as being representative of Mousterian culture and was considered a type anatomically different from Cro-Magnon and modern humans. It could no longer be dismissed as a pathological idiot since it was unreasonable to assume that all known representatives of Mousterian people were pathological in the same way.
This new type of human had a robust brow ridge and a long, low skull, protruding occipital bun, and a weakly developed chin. In the intellectual climate of the end of the 19th century, Mousterians (or Neanderthals) were described as more primitive and more ape-like than they in fact were. They were pictured as stupid animals, incapable of fully erect posture, efficient locomotion, or higher intellectual activities. They did know how to use fire, make flint tools, and were clearly more human than ape. (return to outline)
The First Link: A Fossil Clearly Less Than Human
Among those students interested in prehistory that were excited by prehistoric discoveries and ideas of Darwin and Haeckel was a young Dutch student at the University of Amsterdam, Eugene Dubois (1858-1940). Dubois, greatly influenced by embryonic similarities between human, orangutan, and gibbon, saw Asia as the cradle of humankind. He attempted to obtain private funds for fieldwork in the Dutch East Indies. He was unsuccessful and, against the advice of his friends, eventually resigned a university lectureship in anatomy and accepted a post as a military surgeon in the East Indies in 1887. He was stationed first in Sumatra where he began to excavate caves at his own expense, searching for prehistoric humans.
A friend sent him a fossil from central Java which stimulated him to pull military strings to get a transfer. Dubois excavated intensely at small sites near the village of Trinil in Java between 1890-1892. Original discoveries included a mandible fragment, several teeth, an intact skull-cap, and a complete left femur from a deposit that was particularly rich in fossil bones. Dubois was elated and astounded. He published his discoveries in 1894, claiming that his Trinil fossil was the precursor and intermediate form between the anthropoid apes and humans. The isolated teeth were unquestionably human. He pointed out that, although it was human in shape, the skull had a volume larger than that of apes, but only about two-thirds that of humans. The femur, surprisingly, was absolutely modern in form. Dubois concluded that his ape-man walked upright as does modern humans (and Haeckel's hypothetical Asian precursor), but that its brain volume (estimated at 850 cc) was intermediate between ape (400 cc) and humans (1250 cc). He applied the species name Pithecanthropus erectus Dubois, 1894 to the material.
Dubois returned to Europe in 1895 with a large collection of undescribed fossils. He exhibited Pithecanthropus at the Third International Congress of Anthropology at Leiden and later toured Europe with fossils. Dubois proposed the name Prohylobates for Haeckel's hypothetical gibbon ancestor for human and ape, and expected his discoveries to draw support for Haeckel's theories. Generally, he was not noticed by those whose attention he sought. Most scientists, including the eminent German anthropologist Rudolf Virchow (1821-1902), regarded Pithecanthropus as some sort of fossil gibbon. Clergymen such as Dr. John Lightfoot (Chancellor of Cambridge) attacked the concept of Pithecanthropus as a human ancestor. Dubois' ideas (and Haeckel's) were opposed by Virchow, so Dubois' presentations drew mixed reactions from scientists. Dubois was appointed professor of geology at Amsterdam University where he found himself vulnerable to scientific skepticism and religious attack. He withdrew from the controversy, locked his materials (most unannounced or exhibited) in strongboxes, and lapsed into relative silence on the subject for 30 years.
Meanwhile evidence of human prehistory continued to accumulate. Prince Albert I of Monaco excavated spectacular burials from the Grotte des Enfants at Grimaldi (1895). More than a thousand mammoths were found associated with Aurignacian materials near Predmost in Czechoslovakia, and Professor Gorjanovic-Kramberger found more Neanderthals at Krapina in Yugoslavia. Site reports and discoveries are too numerous to list or describe as the twentieth century opened. Krapina is noteworthy because although its human fossils are fragmentary, more than 10 individuals are represented, and it allowed scientists to examine children and adolescents that belonged to the Neanderthal population. There could be no more talk of pathological idiots.
The first decade of the twentieth century is a continuation of the same kinds of discoveries (La Chapelle-aux-Saint, Le Moustier, Le Ferrassie, Combe-Capelle, La Quina,...). Pithecanthropus had generated enough excitement in Belgium that an expedition was mounted to find more fossils. Due to the untimely death of its leader, the expedition was led by his widow, Lenore Selenka. Unfortunately, no further remains of Pithecanthropus were located. The most bizarre chapter in the discovery of human prehistory begins in this context. (return to outline)
A Fake Ancestor to Fit Expectations
The years 1909-1915 mark the discovery of fossils at Piltdown, near Sussex, England. The first discoveries were announced in 1912 by Charles Dawson and Arthur Smith Woodward (1864-1944). Subsequently a series of discoveries were made until the death of Dawson in 1916. At least three skulls are represented. The first consisted of the right half of a lower jaw with two molar teeth, left temporal, parietal, nasal, turbinate, and parts of frontal and occipital bones. The second skull is known only from a molar tooth and parts of frontal and occipital. The third skull (known as the Barcombe Mills skull) included part of frontal, right parietal, both zygomatic bones, and a mandibular right second molar tooth. A separate discovery in 1915 consisted of an isolated canine tooth from gravel near the location of the original discovery. These fossils were assigned the name Eoanthropus dawsoni Woodward, 1912.
Piltdown presumably came from gravel deposits of Tertiary age, making them much older than any other human discovery, with the possible exception of the Mauer mandible from Heidelberg. Anatomically, they presented a mosaic of human and ape traits. The skull was human, almost modern, but the mandible was that of an ape. Molar wear was flat like a human, but the canine was large and pointed with a diastema like that of an ape. There was much argument about whether an ape-like mandible could belong to the human-like skull, but the general mosaic fit what early 20th century evolutionists expected a human/ape transition to have, namely a human brain and an ape's face. However, there was an element of prejudice in the acceptance this fossil. Even if evolutionists had turned the great chain of being upside down, Piltdown offered evidence of human evolution on English soil, not Africa or Asia. God might still be an Englishman after all - at least Englishmen did not have Africans or Australians as ancestors. With Piltdown in hand, even the status of the European Neanderthal was questionable as a British ancestor.
Only a few people saw or handled the original Piltdown specimens. Casts were made for distribution or study and the originals were locked away in a safe at the British Museum of Natural History. They were available to scientists who had a justifiable need to examine originals, so the security of the museum is no excuse for the mistakes of judgment in those who described and reconstructed Eoanthropus. Although it was subject to frequent debate about details of its anatomy, Piltdown became established as a human ancestor and cast its shadow across human paleontology for the next 30 years.
Marcellin Boule (1861-1942), Director of Human Paleontology at the French National Museum of Natural History, who wrote extensively about Neanderthals between 1911-1913, based his analysis upon a skeleton from La Chapelle-aux-Saints. Neanderthals are portrayed as unable to extend the knee fully and thus postured hunched forward. The skull (according to Boule) indicated an intellect markedly inferior to modern humans. This analysis was endorsed by Grafton Elliott Smith, and thus the Neanderthal continued to be viewed in scientific literature as a subhuman, with less than human posture and wit. (return to outline)
Ape men: Humans with Ape Sized Brains
As the effects World War I (1914-1918) subsided, a new rush of reports of fossil discoveries began. A skull, pelvis, and long bones of a human were found in a lead mine near Broken Hill in Rhodesia (1921). The anatomy of the skull could only be described as extraordinary - although it has a modern sized brain volume, it exhibits a large face and enormous brow ridges. Though it clearly had some similarities to Neanderthal, it was given the scientific designation Homo rhodesiensis Woodward, 1921.
The most important fossil discovery of the twentieth century was recognition of a fossil endocranial cast as that of a hominid by Raymond Dart (1893-1989). Dart, a neuroanatomist trained at University College (London) in Grafton Elliot Smith's lab, was teaching anatomy at Witwatersrand University in South Africa. Industrialization of South Africa produced a growing demand for concrete and related building materials, so limestone quarrying to meet these demands had become an important industry. Quarries were also popular collecting localities for fossils. Caves and cracks in limestone would be filled with a fossil-bearing breccia. Miners piled this aside in spoil heaps since breccia contained impurities inappropriate for producing lime. Dart purchased a box of fossils from a limestone quarry at Taung. Among the specimens was a primate endocranial cast with matching face, which Dart recognized as being unlike that of a nonhuman primate. Dart's 1925 description of the discovery in Nature, assigned the name Australopithecus africanus Dart, 1925, to a juvenile hominid with a long and narrow brain of at least 520 cc volume. The form was more human than that any other anthropoid, but well below modern human size. Dart described Australopithecus as a hominid ancestor with a human face, bipedal locomotion, and a brain only slightly larger than that of an ape.
This presented a serious problem to those who tried to make sense out of the human fossil record. The prevailing thinking (influenced by the existence of Piltdown) was that the human precursor should have an ape-like face and body beneath an expanded human-like brain. To make matters worse, the specimen came from Africa! After some hesitation (Dart had excellent scientific credentials), all of the scientists that had been involved with Piltdown (the most important experts on human evolution in England) rejected Australopithecus, presuming that it was some type of fossil ape, most likely a chimpanzee. The British press had a field day with the controversy. "Taung" became a joke in the media.
A most notable exception to the nay-sayers was a retired physician, Robert Broom (1866-1951), who had an international reputation as a paleontologist for his collections of mammal-like reptiles from South Africa. Although trained in the biology of the 19th century, he was a scientist of great energy and genius. Broom examined Australopithecus and concluded that Dart's assessment was correct. Broom immediately sent short articles to Nature and Natural History confirming Dart's judgment that although the juvenile fossil was about the same size as a chimpanzee, it was a human ancestor, not a fossil ape. More importantly, the energetic Broom applied his fossil collecting resources to a search for more Australopithecines. (return to outline)
Exploration Between the World Wars
In 1923, Ales Hrdlická (1869-1943), from the Smithsonian Institution, traveled to Europe where he examined both Eoanthropus and Pithecanthropus. Hrdlická went on to visit India, Java, China, and South Africa, in a world wide tour of human paleontology. The result is an excellent and influential account of human paleontology, The Skeletal Remains of Early Man (1930). Hrdlicka's reaction to Eoanthropus was strongly negative. First he pointed out the complete absence of stratigraphic context. He concludes:
"No amount of trust and benevolence can quite fill these defects of the evidence (1930; page 89)." Further, the association between the Piltdown mandible and skull was not credible to Hrdlicka. "The more the lower jaw is studied and understood the less in harmony it appears with the skulls and is not unlikely that these latter belong to totally different, possible chronologically much younger human individuals" (1930; page 86).
Conversely, Hrdlicka was impressed with the intermediate anatomy of the Pithecanthropus skull cap. It clearly had ape-like features but contained a near-human brain volume (900 cc). The name Pithecanthropus seemed extremely appropriate. Dubois' discovery began to take its proper place in the pantheon of human ancestors. However, Dart's Australopithecus did not fare as well. Even though Hrdlicka examined the fossil and visited Taung, it is passed over as a "chimpansoid juvenile." Word of discoveries at Zhoukoudian in China arrived in Washington after the manuscript was typeset. A brief note and photographs were included as an addendum.
Support from the Rockefeller Foundation resulted in establishment of a medical school in China, the Peking [Beijing] Union Medical College. Fossils from a nearby locality, Zhoukoudian (Chou Kou Tien in older texts), attracted the attention of Davidson Black (1884-1934), a Canadian anatomist at the college. Funded partly by the Rockefeller Foundation, an international team started to excavate Zhoukoudian in 1927. Black was a motivating force behind fieldwork, but the official director was Dr. V.K. Ting, a Chinese scientist who established the Geological Survey of China. Actual field work was overseen by a Chinese geologist, C. Li, and a Swedish paleontologist, Dr. Birger Bohlin. The first season produced a human tooth and the species name Sinanthropus pekinensis Black, 1927. In subsequent seasons a great variety of other scientists joined work at the site. Two Chinese scientists, W. C. Pei and C.C. Young, play an important role. Visiting scientists included Père Teilhard de Chardin (1881-1955) and the Abbé Henri Breuil (1877-1961). The 1929 season produced a long awaited skull and justified Black's enthusiasm about Zhoukoudian. As each season progressed, Black worked long hours with the material and with his other duties. As more discoveries came to light, the site and its materials demanded more time. His health failed and Black died late at night at his bench in the Cenozoic Research laboratory. One noteworthy tribute to Black came from Dr. Ting:
""...In politics, Black was a conservative, but in his dealings with his Chinese colleagues, he forgot altogether about their nationality or race, because he realized that science was above such artificial and accidental things" (Quoted in Shapiro. 1974; page 54)
Davidson Black was replaced temporarily by Teilhard de Chardin until Franz Weidenrich (1873-1948), a German anatomist, could arrive to take over permanently. Work at Zhoukoudian was stopped by military activity in 1937. Weidenrich prepared careful casts, photographs, drawings, detailed descriptions, and measurements of the human fossils. With the beginning of the Second World War in 1939, preparations were made to secure the laboratory and materials from war. Weidenrich fled to the U.S. safely in 1941 with his casts and records, leaving the original fossils carefully packed and stored in the laboratory. Fearing the Japanese would search for the materials, Dr. Wong, then the director of the Geological Survey, asked the American ambassador to arrange shipment of valuable specimens to America. The fossils were carefully packed in two boxes that were among footlockers sent by rail with a U.S. marine detachment at Peking to meet the U.S.S. President Harrison. Japanese soldiers captured the marines on the morning the ship was to sail, and the President Harrison was grounded while trying to evade a Japanese warship. The marines were sent to prison camps, and the fossils were lost in the confusion of war. Part of the booty taken by the Japanese at Ching-wang-tao where the marines were captured was loaded on a Japanese vessel that sank in the Straits of Chilli. Only Weidenreich's casts and records survived. There is no reliable evidence of the fate of Sinanthropus.
The 1930's were, until the start of World War II, a time of great discovery. A young East African raised British anthropologist, L.S.B. Leakey (1903-1972), organized his first expedition to East Africa in 1926. By 1931, his third expedition was on route to Olduvai Gorge. He was an energetic and imaginative archaeologist who was later to sponsor (and often find the funding for) an extraordinary number of scientific projects in East Africa. He established a routine in which his lecture and fund raising tours of Europe and U.S. would help support projects in East Africa. His first popular book, Adam's Ancestors, appeared in 1934.
Numerous European discoveries were still being made, but perhaps the two most historically important were a largely intact skull (Homo steinheimensis Berckhemer, 1936) from the Steinheim Quarry (near Stuttgart in Germany) and parts of a calvarium from the Barnfield gravel pit near Swanscombe, Kent, England. In the Middle East, a joint expedition of the British School of Archaeology in Jerusalem and the American School of Prehistoric Research excavated a series of small caves on Mount Carmel, south-east of Haifa, Israel. The expedition, under the direction of D.A. E. Garrod, found two skeletons in Tabun cave, ten individuals in Skuhl cave, and numerous isolated bone fragments. A young American physical anthropologist, Theodore D. McCown (1908-1969) spent weeks in Sir Arthur Keith's basement, extracting the better preserved skulls from breccia with dental tools. The Mt. Carmel specimens exhibited an exciting mosaic of Neanderthal and Cro-Magnon traits.
In South Africa, Robert Broom (1866-1951) retired from the practice of medicine and was appointed to a post in the Transvaal Museum. At the age of 68, he began to devote his full energy to finding another Australopithecus. By 1936, he was searching the cave deposits around Pretoria. He learned of breccia deposits at the caves at Sterkfontein, and in about two weeks, found an adult skull similar to Dart's specimen from Taung. Broom gave it the name Australopithecus transvaalensis Broom, 1936, later changing it to Plesianthropus transvaalensis Broom, 1937. Numerous fragments of Plesianthropus were found in 1937 and 1938. These discoveries were profoundly important because they demonstrated the anatomy an adult Australopithecus and validated Dart's predictions. In 1938, he found a skull of a different type of hominid at Kromdraai which he called Paranthropus robustus Broom 1938. These discoveries indicated two forms of early hominids: one gracile form, Australopithecus, and another much more robust form Paranthropus, both of which combined human and pongid characters in ways not previously seen.
Meanwhile, Gustav H. Ralph von Koenigswald (1902-1982) was appointed to a post as paleontologist to the Geological Survey Service in Java in 1930, and between 1931 and 1941 announced a number of important fossil discoveries.. Financed in part through the Carnegie Foundation, he began a systematic survey in Java. Meanwhile another member of the Geological Survey Service, C. ter Harr, found eleven fragmentary skulls of a more modern type near Ngandong along the Solo river about six miles from Trinil and the Pithecanthropus site. They had some of the large brow ridges of Pithecanthropus, but had endocranial volumes closer to those of modern humans. One of von Koenigswald's assistants brought him a piece of a Pithecanthropus skull in 1937. In his search for more material, the geologist offered to purchase additional fossil bones for one cent for each tooth and ten cents for each skull fragment. Unfortunately, this caused some of the specimens to be broken into splinters by the native helpers in order to maximize the number of pieces they could sell. One of the skull caps, the first Sangiran calvarium was an exact duplicate of Dubois' Pithecanthropus calvarium.
A very robust mandible fragment was designated Meganthropus palaeojavanicus Weidenreich, 1945. A maxilla fragment from Sangiran gave everyone their first look at a pithecanthropine face, and a juvenile calvarium from Modjokerto prompted this limerick (from Hooton's Up From the Ape, 19, page 297):
ODE TO HOMO SOME-JERKTENSIS
Young pithy from your Djetis bed
You raise a scarcely human head,
With all its soft spots ossified
And sutures closed that should gape wide.
Your marked postorbital constriction
Would clearly justify prediction
That had you lived to breed your kind,
They would have had the childish mind
That feeds upon the comic strips
And reads with movements of the lips.
They would have had no need for braces
To warp their teeth into their places-
Equipped for general mastication
And not progressive education.
In place of brows a bony torus,
And no ideals with which to bore us.
Gustav H. Ralph von Koenigswald also had problems during the war years. He hid his fossils, and although he was interned in a prison camp, only one fossil skull was confiscated by the Japanese soldiers. It was presented to Emperor Hirohito and recovered after the war.
There were a few important publications during the war years, the most notable being Weidenreich's description of Sinanthropus. In a borrowed office at the American Museum of Natural History, Weidenreich reviewed the fossil record of human evolution, merging Sinanthropus and Pithecanthropus into a new taxon, Homo erectus. He published descriptions and assigned scientific names to some of von Koenigswald's discoveries - von Koenigswald was incorrectly presumed dead in a Japanese prisoner of war camp.
G.H.R. von Koenigswald's comments about Piltdown were prophetic, pointing out that all discoveries since 1917 had contradicted the idea that humans had an ancestor with a modern brain and ape jaw. Indeed, the trend of the discoveries suggested the opposite relationship. Weidenreich concluded:
"...I am only wondering why, if a human vault, a simian mandible and an anonymous 'canine' were combined into a new form, the other animal bones and teeth found in the same spot were not added to the 'Eoanthropus' combination; I do not believe in those miracles whether offered by anti-Darwinian or Darwinians. The sooner the chimaera 'Eoanthropus' is erased from the list of human fossils, the better for science. (Weidenreich, 1943; page 220)"
After World War II, scientists approached the question of human origins with new vigor and renewed resources. Broom was vigorously excavating (sometimes using explosives). Concern over his excavation techniques led the Historical Monuments Commission to forbid him to continue without the presence of "a competent field geologist." He ignored the ruling and at Sterkfontein in 1947, found an intact skull of Plesianthropus and a nearly intact pelvis. Numerous other Australopithecine fragments were coming out of the lime works. The small ape-like brain volume predicted by Dart for the adult Australopithecine continues to be validated by the almost complete specimens. The following year, 1948, Dart, assisted by personnel from the Department of Anatomy at Witwatersrand, recovered additional Australopithecus specimens from Makapansgat while Broom recovered Paranthropus material from Swartkrans. With numerous specimens of Australopithecus, Paranthropus, Pithecanthropus, Sinanthropus, and Neanderthals available, a modern synthesis of human paleontology was possible.
Two opposing traditions in anatomy conflicted in postwar England. One, championed by Sir Wilfred Edward Le Gros Clark (1895-1971), followed traditional functional anatomy, and is associated with the phrase "total morphological pattern". The other, championed by Lord Solly Zuckerman, uses mathematics and statistics to make phylogenetic decisions. Le Gros Clark argued that length and breadth measures can not distinguish between a square and a circle, but the Zuckerman metrical approach was more popular in a scientific community that was discovering the power and convenience of statistics. Le Gros Clark and the aging Sir Arthur Keith sided with Broom in placing Australopithecus among our human ancestors. Zuckerman, on the basis of tooth measurements, decided Australopithecus was an ape. (return to outline)
The Demise of Eoanthropus
Piltdown was finally debunked by Kenneth Oakley (1911-1981) and M. F. Ashley Montagu (1905-). They devised a technique for measuring fluorine content in bone which was practical to apply to fossils. They demonstrated (following the lead of Adolf Carnot, a 19th century French mineralogist) that fossils acquire fluorine from ground water. With certain limits, the procedure could determine the relative age of fossils. In 1948 Oakley determined that Piltdown specimens exhibited a great variety of ages, and did not represent a set of associated materials. Eoanthropus was removed as a valid taxon, since the skull and mandible were no longer in stratigraphic association. Joseph Weiner, Oakley, and Le Gros Clark exposed Piltdown as a fake in 1953.
The lessons of Piltdown are numerous, but the affair highlighted the importance of stratigraphic context and anatomical completeness. More troubling, it is a case of scientific predisposition toward interpretations that validate contemporary ideas about evolutionary events. Once such ideas gain wide acceptance, they are sometimes judged by the strength of opinion, not strength of evidence.
Mary Douglas Nicol Leakey (1913- ) made the first important discovery of an Australopithecine-like creature in East Africa at Olduvai Gorge in 1959. This relatively complete and robust skull (minus mandible) was given the name Zinjanthropus boisei Leakey, 1959. The specimen was fragmented but it was associated with stone artifacts and in a geologic context that could be associated with a rich fauna. The importance of these discoveries at Olduvai helped the Leakeys secure funding to conduct larger projects in East Africa in addition to the small budget expeditions they had sustained over the previous 25 years. The next few years saw an avalanche of discoveries. Notable was recognition of another species (Homo habilis Leakey, Tobias and Napier, 1964) that was near Australopithecines in time but more human in form, especially in brain volume. A most surprising discovery was a stone or rubble circle and living floor that appeared to represent a hut of some sort ( DK hut circle site).
Radiometric Dating Provides a Time Scale
Attempts to date the Zinjanthopus strata included a revolutionary new set of techniques. Jack Evernden and Garniss Curtis used radiometric dating techniques, especially the potassium argon method, to place a date of 1.75 million years ago for a hard tuff that overlay the DK hut circle site and the Zinjanthropus site. The basalt beneath produced an age of 1.9 million years. In just a few years availability of absolute dates from radiometric techniques overhauled everyone's perspective for the time depth of the human fossil record.
Shortly after new radiometric techniques began to improve geochronology, molecular biologists proposed time depths of separation between primate lineages based on estimated rates of molecular evolution. Molecular biologists like Morris Goodman at Wayne State University were demonstrating that phylogenetic relationships could be reliably demonstrated using serum proteins. Vincent Sarich, with the encouragement of Allan Wilson and Sherwood Washburn at Berkeley, proposed a molecular clock model that used proteins both to demonstrate phylogenetic relationships and to provide estimates of their time depth. The first and immediate controversy was the relationships between humans and African apes. Molecular biologists, especially Sarich, pushed for a young ape-human split - contesting the consensus among paleontologists for a much older division.
The combined effect during the late 1960s and early 1970s was to push estimates for human divergence from ape lineages from the Oligocene or Early Miocene to late Miocene. Relationships between fossil and extant taxa were intensely discussed and reviewed to accommodate new sources of information and revised views of earlier finds.
Late Twentieth Century Fossil
In 1968, Richard Erskine Frere Leakey (1944- ), son of Mary and Louis Leakey, mounted an expedition to East Turkana in northern Kenya. Assisted by his wife Mauve Leakey, Richard and a field team explored the eroded hills and waddis of the Turkana drainage basin. Although the area eventually produced more than a hundred hominids, a fragmented skull (KNM-ER-1470), was a watershed discovery. Seasonal surveys of the Koobi Fora area have since produced a series of marvelously preserved Australopithecus and Homo fossils from datable strata. After the death of his father in 1972, Richard Leakey took a progressively larger role in fossil hunting. In 1984 the area yielded an almost complete skeleton of Homo erectus at Nariokotome in West Turkana which more than doubled the H. erectus postcranial material known, and contradicted the view that H. erectus was generally shorter than modern people.
In 1974, Mary Leakey turned her attention to Laetoli, a site that the Leakey family had occasionally visited since the 1930's. Between 1974 and 1978, Laetoli yielded not only a diverse fauna, it produced a 3.5 million year old trail of hominid footprints. The prints of three individuals were preserved in a thin layer of ash that was excavated carefully to preserve its features and expose over 24 m of trail surface. The footprint and stride characteristics are human.
The Afar Depression, a remote lowland in southeastern Ethiopia lies at an intersection of three rifts--the East African, Arabian, and Gulf of Aden rifts-- formed by the separation of the African, Arabian, and East African plates. It is the only such "triple junction" of active rifts on dry land. World wide attention began to focus on this area through studies in plate tectonics in the 1960's. In 1970, a French geologist, Maurice Taieb, discovered the fossil site of Hadar in the Afar's lower Awash river valley. The following year, Taieb was joined by an American geologist, Jon Kalb, then with the Ethiopian Geological Survey. Kalb conducted independent surveys of fossil-bearing areas immediately south and north of Hadar. In late 1971, Taieb and Kalb formed the nucleus of the International Afar Research Expedition (IARE), under the patronage of Louis and Mary Leakey, to conduct detailed studies of Hadar and other sites. The IARE was joined in mid-1972 by anthropologist, Don Johanson, who had been working with a team from the University of California at Berkeley in Ethiopia's Omo Valley. In late 1974, Kalb broke from the IARE in a dispute with Johanson precipitated by documentation procedures, and in early 1975 formed the Rift Valley Research Mission in Ethiopia (RVRME), an Addis Ababa-based organization made up of Ethiopians and Americans. The RVRME conducted extensive explorations of a 5000 km2 area in the Middle Awash Valley, extending well south of Hadar. During this time, they discovered extensive fossil deposits ranging from late Miocene, several million years older than Hadar, to late Pleistocene, several million years younger.
There are numerous hominid finds from Hadar. One noteworthy discovery in 1974 is approximately 40% of a single individual, assigned the name Australopithecus afarensis (Johanson, White and Coppens, 1978); but affectionately called "Lucy" (AL-288) and dated to about 3 million years before present. A second phenomenal discovery in 1975 included dozens of fragments representing at least 13 individuals from the same locality (AL-333).
The first hominid discovered in the Middle Awash was a largely complete, early to middle Pleistocene skull of an archaic Homo called the Bodo skull. It was found associated with a rich fossil fauna and abundant Acheulian handaxes, cleavers, and other artifacts, including enormous basalt core boulders (c. 60 Kg) that had apparently been rolled to the site from a volcanic flow nearly a half a mile away.
After 3 1/2 years of rigorous surveys, however, the work of the RVRME collapsed following Kalb's expulsion from Ethiopia by security officials amid unfounded rumors that he was in the CIA. The RVRME research permit was then turned over to archaeologist J. Desmond Clark (1916- ), for work on the sites discovered by RVRME. In 1981 a team led by Desmond Clark and Tim White found fragmentary hominid remains of approximately 4 m.y., which White identified as A. afarensis. The ill-will generated by Kalb's expulsion, however, spread, involving government officials and Ethiopians formerly with the RVRME. In 1982 the Ethiopian government temporarily banned all prehistory work in the country by foreign expeditions pending a revision of Ethiopia's national policies concerning antiquities.
A number of fragmentary hominid fossils had remained unnamed from East Africa. Mary Leakey (and some of her colleagues) had refrained from applying taxons in some cases bcause of the uncertianty of relevant morphologies. Mary Leakey did not agree with the composition of the proposed A. afarensis taxon and friction between the different camps of paleoanthropologists ran high at this time. Johanson, White and Coppens chose a type specimen for A. afarensis that limited Mary's options. Each fossil species is defined by reference to a type specimen (called a holotype) against which all other proposed members of that species is compared. The "type specimen" is strongly tided to its scientific name. The scientific name remains valid if it is the first appropriately applied to the "type." If she was right and the various specimens attributed to A. afarensis included several taxa, the name was tied to the type specimen in her laboratory from Laetoli. The Ethiopian team could apply another taxonmic name to "Lucy" and their other Hadar material, but Mary was stuck with A. afarensis for some of her important specimens.
Subsequent to 1983, archaeological and fossil materials continued to be recovered world-wide. Subsaharan Africa was particularly productive. Notable discoveries include the discovery of a complete skull of a robust Australopithecine at Lomekwi I in 1985 (KNM-WT-1700) and more hominids from Olduvai, especially postcranial material (OH 62). The South African breccia cave sites, especially Swartkrans, continued to be productive.
Outside Africa, excavation at Kebara Cave, Mt. Carmel, Israel in 1983 recovered a relatively complete adult male Neanderthal postcranial skeleton that provided our most intact Neanderthal pelvis and hyoid. Additional Neanderthal sites such as the Molare Shelter near Salerno, Italy, have been excavated in Mediterranean Europe.
Anatomical analyses of A. afarensis from Hadar in the mid 1980's established that species' mosaic of hominid and pongid characteristics. Estimates of the cranial volume of A. afarensis from Hadar (Falk, 1985) fell below 400 cm3. Although bipedal, A. afarensis exhibits somewhat pongid limb proportions, extensive body dimorphism, and hand and foot adaptations for climbing. The anatomical transition from an ape life-way to a human life-way appears to be effectively bridged in the fossil record, but there were only fragmentary hints of possible precursors to the Australopithecines in the late Miocene of Africa.
Perhaps the most controversial discussions of the 1980s were stimulated by molecular biologists who argued that mitochondrial DNA analyses indicated an African origin for the modern human species and estimated the first modern humans to have existed about 200,000 years ago. Recognition that modern anatomy might be 100,000 years or older (Omo I, Klasses River, Border Cave, Xujiango, Qafzeh) set the current stage for ongoing debate about the immediate origins of humans with a modern anatomy.
Scientists working in the Lake Turkana area of Kenya after 1992 were also finding hominid fossils that are close to 4 million years old. Mave Leakey led a team to Kanapoi, where, in deposits dating between 3.9 and 4.2 MYBP, they recovered fragments of a bipedal hominid, Australopithecus anamensis.
Aramis, one of several sites in the Awash River drainage basin in Ethiopia, was the focus of field work between 1992 and 1994 by a team lead by Tim white, Gen Suwa, and Berhane Asfaw. They collected numerous fragments of a fossil hominid that date approximately 4.4 MYBP. Most of the material is still unpublished but they proposed that it represents a previously unknown hominid and christened it Ardipithecus ramidus. Additional fragmentary materials dating between 5.2 and 5.8 mybp were later proposed as
A fossil Australopithecine skull had been reported from Chad in 1960 and assigned the name Tchadanthropus uxoris (Coppens, 1965). Uncertainties about the date, provenance, and morphology resulted in the material being generally ignored. It has been regarded very fragmentary and eroded modern skull fragments. The absence of Australopithecines from Central and West Africa led some scientists to view the East African rift valley as the cradle board of the hominids. Australopithecus was geographically extended to Central Africa with the description of Australopithecus bahrelghazali (Brunet et al. 1995) from Koro Toro in Chad. Sahelanthropus tchadensis, represented by an apelike skull dated between 6 and 7 mybp was found in Chad in 2002.
About this time (mid 1990's, with excavations continuing for a decade) parts of another Australopithecine skeleton began to be reported from Sterkfontein. At first it was a few foot bones that were thought to date to approximately 3.5 mybp, but further work by Ronald Clarke and his colleagues continued to uncover more of what turned out to be much of an australopithine skeleton that was nicknamed "Little Foot." Better dating techniques revised the age of "Little foot" to over 4 mybp.
A number of laboratories attempted to extract and sequence Neanderthal mtDNA. The first reports in 1997 suggested that Neandertal mtDNA contrasted from modern humans and supported that Neanderthals were a separate human species from modern Homo sapiens. Since then various laboratories have sequenced at least eight samples of Neanderthal mtDNA with similar conclusions.
Also in the late 1990's, archeological surveys at Gona, Ethiopia recovered stone tools (Mode I) in sediments between 2.5 and 2.6 mybp. A presumed archaic Homo sapiens cranium was found in Ertrea that dates to about 1 mybp. Australopithecine material from Bouri in Ethiopia was given the species name Australopithecus garhi (). Other Australopithecine fragments from deposits ranging from 5.2 mybp to 5.8 mybp were assigned first the name Ardipithecus ramidus kadbba (Haile-Selassie 2001) and then was renamed Ardipithecus kadbba in 2004.
Twenty-first Century Fossil Discoveries
Two Homo erectus skulls and a
mandible from Dmanisi in the Republic of Georgia were dated to
about 1.7 mybp. These materials again confirm that Homo
erectus was well established in Euroasia by that date.
More material was later recovered from Dmanis that have been proposed
as intermediates between H. habilis and H.
erectus. The species name Homo gerogicus
(Ghabunia et al. 2002) was proposed for these discoveries.
Probably the most important fossil identification of the turn of the Twentieth century was the naming of Kenyanthroupus platyops (Leakey et al 2001), a 3.6 million year old fossil from Kenya. The fossil record now included a variety of early hominids. The Australopithecines were not an ideal ancestor for Homo because of their specialized megadon postcanine dentition. There were a variety of bipedal hominids prior to Australopithecus, so the focus shifts to a search for a bipedal preaustralopithecine candidate who might be a logical precursor of Homo. Possibly it would be bipedal, and would lack a sectorial mandibular premolar. The dentition of Homo is primitive except for the canine/premolar complex, so the search was for a bipedal hominid with similar primitive teeth.
A French-Kenyan team described some hominid fragments as older (6 mybp) and as more advanced than the Australopithecines. Their material, nicknamed the "Millennium Man" was assigned the taxon Orrorin tugenensis. The significance of this material is currently uncertain because of its fragmentary nature.
Some very important discoveries came from Herto, Ethiopia in 2003. They are three skulls that are similar in some respects to archaic humans and modern human anatomy. They are important because of their estimated age, 160,000 ybp. A new subspecies name, Homo sapiens idaltu was proposed for them. If they are modern in form, they are the oldest dated material with modern anatomy.
In 2004, researchers found a H. erectus skull (OL 45500) at the famous Acheulean site of Olorgesailie in Kenya. It is dated about 0.95 mybp.
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21 Aug 2004
Department of Department of Anthropology, College of Liberal Arts , UT Austin
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