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the
multiregional
evolution of humans
By Alan G. Thorne and Milford H. Wolpoff
T
hree decades ago the pa- humanity the idea that humans origi- of modern humans across the globe.
leoanthropological com- nated in Africa and then developed their Nevertheless, mitochondrial DNA is
munity was locked in a modern forms in every area of the Old not the only source of information we
debate about the origin World. On the other side are researchers have on the subject. Fossil remains and
of the earliest humans. who claim that Africa alone gave birth to artifacts also represent a monumental
The disagreement centered on whether a new species of modern humans within body of evidence and, we maintain, a
the fossil Ramapithecus was an early hu- the past 200,000 years. Once again the considerably more reliable one. The sin-
man ancestor or ancestral to both human molecular geneticists have entered the gular importance of the DNA studies is
and ape lineages. Molecular biologists en- fray, attempting to resolve it in favor of that they show that one of the origin the-
tered that discussion and supported the the African hypothesis with a molecular ories discussed by paleontologists must
minority position held by one of us clock. Once again their help must be re- be incorrect.
(Wolpoff) and his students that Rama- jected because their reasoning is flawed. With Wu Xinzhi of the Institute of
pithecus was not a fossil human, as was Genetic research has undeniably pro- Vertebrate Paleontology and Paleoan-
then commonly believed. Their evidence, vided one of the great insights of 20th- thropology in Beijing, we developed an
however, depended on a date for the century biology: that all living people are explanation for the pattern of human
chimpanzee-human divergence that was extremely closely related. Our DNA evolution that we described as multire-
based on a flawed molecular clock. We similarities are far greater than the dis- gional evolution. We learned that some
therefore had to reject their support. parate anatomical variations of human- of the features that distinguish major hu-
Paleoanthropologists are again en- ity might suggest. Studies of the DNA man groups, such as Asians, Australian
gaged in a debate, this time about how, carried by the cell organelles called mito- Aborigines and Europeans, evolved over
when and where modern humans orig- chondria, which are inherited exclusive- a long period, roughly where these peo-
inated. On one side stand some re- ly from ones mother and are markers for ples are found today, whereas others
searchers, such as ourselves, who main- maternal lineages, now play a role in the spread throughout the human species be-
tain there is no single home for modern development of theories about the origin cause they were adaptive.
Multiregional evolution traces all
POINT-COUNTERPOINT: For an opposing view of how humankind arose around modern populations back to when hu-
the globe, see The Recent African Genesis of Humans, on page 54. mans first left Africa almost two million
WAY
PATH
SIAN
RALA
AUST Willandra Lakes 50
Ngandong 1 (Upper Pleistocene)
(Indonesia)
years ago, through an interconnected web Harvard University as the Noahs ark in Africa approximately 200,000 years
of ancient lineages in which the genetic model, posited that modern people arose ago. Only mitochondrial DNA that can
contributions to all living peoples varied recently in a single place and that they be traced to Eve, these theorists claim, is
regionally and temporally. Today dis- subsequently spread around the world, found among living people.
tinctive populations maintain their phys- replacing other human groups. That re-
ical differences despite interbreeding and placement, recent proponents of the the- Paddling in a Pool
population movements; this situation has ory believe, must have been complete. HOW COULD THIS BE? If Eves de-
existed ever since humans first colonized From their genetic analyses, Allan C. Wil- scendants mixed with other peoples as
Europe and Asia. Modern humanity orig- son and his colleagues at the University of their population expanded, we would
MILFORD H. WOLPOFF
inated within these widespread popula- California at Berkeley concluded that the expect to find other mitochondrial DNA
tions, and the modernization of our an- evolutionary record of mitochondrial lines present today, especially outside
cestors has been an ongoing process. DNA could be traced back to a single fe- Africa, where Eves descendants were in-
An alternative theory, developed by male, dubbed Eve in one of Wilsons vaders. The explanation offered for the
paleontologist William W. Howells of first publications on the subject, who lived current absence of other mitochondrial
The Eve theory makes five predictions can find none. For instance, whereas the the time when Eves descendants are sup-
that the fossil evidence should corrobo- hand ax was a very common artifact in posed to have left Africa suggests that
rate. The first and major premise is that Africa, the technologies of eastern Asia any new African technology emerged or
modern humans from Africa must have did not include that tool either before or spread to other continents. All in all, as
completely replaced all other human after the Eve period. There is no evi- we understand them, the Asian data re-
tinguishing features of the earliest of terial on the origins of Homo sapiens. Thorne is adjunct fellow in the department of ar-
these Javan remains, dated to more than chaelogy and natural history at the Australian National University. He graduated from the
one million years ago, show that they University of Sydney in 1963 and later taught human anatomy at the medical school there.
MILFORD H. WOLPOFF
had developed when the region was first Thornes excavations at Kow Swamp and Lake Mungo produced most of the Pleistocene hu-
inhabited. man remains in Australia. Wolpoff is professor of anthropology at the University of Michi-
Compared with human fossils from gan at Ann Arbor, where he directs the paleoanthropology laboratory. He received his Ph.D.
other areas, the Javan people have thick in 1969 from the University of Illinois at Urbana-Champaign. Wolpoff would like to thank
skull bones, with strong continuous lecturer Rachel Caspari of the University of Michigan for her help in drafting the epilogue.
MATERNAL LINEAGE RECONSTRUCTIONS based solely on the mitochondrial DNA types found today are
inherently flawed. A hypothetical tree inferred from only five surviving types (left) leaves out the the Eve theory predictions, the evidence
branches and mutational histories of extinct lines (right). Consequently, it sets the date for a common points indisputably toward the continu-
ancestor much too recently by presenting evidence of too few mutations. ity of various skeletal features between
Focus on Features
IF AFRICA REALLY WAS the Garden
of Eden from which all living people
emerged, one would expect to find evi-
dence for the transition from archaic to
modern forms there and only there.
Following the lead of German researcher
Reiner Protsch von Zieten of Goethe JAW MORPHOLOGY distinguishes many Neandertal skeletons. In most living people and in fossils,
University in Frankfurt, Germany, some the rim around the mandibular nerve canal opening is grooved (left), but in a number of
paleontologists did argue that modern Neandertals, it was surrounded by a bony bridge (right). Some later Europeans also had this
Neandertal feature, although it was less common.
Homo sapiens originated in Africa be-
cause they believed the earliest modern- modern in its higher skull and more analysis of mitochondrial DNA suggest-
looking humans were found there and rounded cranial rear. An associated man- ed a theory so contrary to the facts. Per-
that modern African features can be seen dible has a definite chin. Like the Levant haps the mitochondrial DNA has been
in these fossils. But the African evidence remains of similar age from Qafzeh and misinterpreted.
is similar to other regions in that modern - even this small Omo sample com-
Skhul, The basic difficulty with using mito-
features and not modern populations ap- bines a mix of archaic- and modern-ap- chondrial DNA to interpret recent evolu-
pear gradually and at about the same pearing individuals. tionary history stems from the very source
time as they appear elsewhere. The best excavated remains are from of its other advantages: in reproduction,
The African record differs from oth- Klasies River and are securely dated to the mitochondrial DNA clones itself in-
er regions in that the earlier, archaic pop- between 80,000 and 100,000 years ago. stead of recombining. Because mitochon-
ulations are more variable and have no Some of the skull fragments are small drial DNA is transmitted only through the
specifically African features. Modern-ap- and delicate and are said to prove that maternal line, the potential for genetic
pearing humans and technologies first modern humans were present. Yet a driftthe accidental loss of linesis great:
arise during the time between the last comparative analysis of the entire sample some mitochondrial DNA disappears
two glaciations. The technologies seem by Rachel Caspari of the University of every time a generation has no daughters.
regional and impermanent, not conti- Michigan at Ann Arbor showed that oth- The problem is analogous to the way
nent-wide, but anatomical features are ers are not modern-looking at all. Two of in which family surnames are lost when-
more widespread. We believe the main the four lower jaws do not have chins, so ever there is a generation without sons.
reason that Africa differs from the rest of thorough proof of a modern jaw is lack- Imagine an immigrant neighborhood in
the world at this time is that it is much ing. The single cheekbone from the site is a large city where all the families share a
more heavily populated many, if not not only larger than those of living surname. An observer might assume that
most, people lived thereand more pop- Africans but also larger and more robust all these families were descended from a
ulation movement is outward than in- than those of both the earlier transition- single successful immigrant family that
PATRICIA J. WYNNE, BASED ON WORK BY MARIA OSTENDORF SMITH
ward. The key specimens addressing mod- al humans and the archaic humans found completely replaced its neighbors. An al-
ernity span the continent, from Omo in Africa. The claim that this sample con- ternative explanation is that many fami-
Kibish in Ethiopia to Klasies River Mouth tains modern Africans is highly dubious lies immigrated to the neighborhood and
Cave in South Africa. The three Omo and does not justify the proposal that the intermarried; over time, all the surnames
Kibish crania date roughly to between earliest modern humans arose in Africa. but one were randomly eliminated
100,000 and 200,000 years ago and are through the occasional appearance of
similar to other African remains from DNA Reanalyzed families that had no sons to carry on their
this time in combining ancient and mod- W I T H T H E D I S P R O O F of the unique names. The surviving family name would
ern features. Omo 2 is the more archaic, African ancestry theory for the living have come from a single immigrant, but
with a lower skull and a much broader people of most areas and the lack of evi- all the immigrants would have con-
and more angled cranial rear, resembling dence showing that modern people first tributed to the genes of the modern pop-
those of Laetoli 18 from Tanzania. Its appeared in Africa, we conclude that the ulation. In the same way, generations
browridge, however, is smaller than predictions of the Eve theory cannot be without daughters could have extin-
Omo 1s, which generally appears more substantiated. We must wonder why the guished some lines of mitochondrial
^
Afalou Mitochondrial DNA may not be neu-
Predmost Ziyang Keilor
UPPER PLEISTOCENE
^
Lukenya
Mladec Liujiang Willandra tral enough to serve as the basis for a mo-
Vindija Lakes 50 lecular clock, because some data suggest
MIDDLE
Kebara Dar es Soltan Maba Lake Mungo 1, 3 that it plays a role in several diseases. Be-
La Ferrassie
La Chapelle cause of random loss and natural selec-
tion, some vertebrate groups have rates
EARLY
Zuttiyeh Ngandong
agree with Wilsons interpretation of the
MIDDLE
Lake Turkana Sangiran clock was able to date Eve to only be-
(east) 992 4, 27, 31 tween 50,000 and 500,000 years ago. Be-
cause of the uncertainty, we believe that
Lake Turkana
EARLY
Dmanisi (east) 730, 3883, 3733 Mojokerto for the past half a million years or more
(west) 15000 of human evolution, for all intents and
purposes, there is no molecular clock.
LAURIE GRACE
WELL-DATED FOSSILS point to the continuous, linked evolution of modern humans at sites around
the world. Modern human groups in different regions developed distinct anatomical identities.
Putting aside the idea of a clock, one
Nevertheless, gene flow between the groups through interbreeding spread important changes can interpret the genetic data in a much
throughout and was sufficient to maintain humans as a single species. more reasonable way: Eve carried the