Scientific American Special Edition - New Look at Human

WWW.SCIAM.
COM
Display until August 25, 2003
COPYRIGHT 2003 SCIENTIFIC AMERICAN, INC.

LETTER FROM THE EDITOR
Established 1845
New Look at Human Evolution

is published by the staff of
Scientific American, with
project management by:
The Original Human
EDITOR IN CHIEF: John Rennie
EXECUTIVE EDITOR: Mariette DiChristina
ISSUE EDITOR: Mark Fischetti
ISSUE CONSULTANT: Kate Wong
Interest Story
ART DIRECTOR: Edward Bell
ISSUE DESIGNER: Jessie Nathans
It’s quite a tale. Perhaps five million to 10 million years ago, a primate
PHOTOGRAPHY EDITOR: Bridget Gerety species diverged from the chimpanzee line. This was the forerunner of human-
PRODUCTION EDITOR: Richard Hunt ity— and a host of other beings who were almost but not quite human. For a
COPY DIRECTOR: Maria-Christina Keller time, a throng of hominid species shared the planet; at least four even coexist-
COPY CHIEF: Molly K. Frances ed in the same region. By around four million years ago, our progenitors and
COPY AND RESEARCH: Daniel C. Schlenoff,
Rina Bander, Shea Dean, others had mastered the art of walking upright. Some two million years later
Emily Harrison, David Labrador they strode out of Africa and colonized entirely new lands. Certain groups
EDITORIAL ADMINISTRATOR: Jacob Lasky learned to make sophisticated tools and, later, artwork and musical instru-
SENIOR SECRETARY: Maya Harty ments. The various species clashed, inevitably. Modern humans, who entered
ASSOCIATE PUBLISHER, PRODUCTION: Europe 40,000 years ago, may have slaugh-
William Sherman tered Neandertals (when they weren’t inter-
MANUFACTURING MANAGER: Janet Cermak
ADVERTISING PRODUCTION MANAGER: Carl Cherebin breeding with them). Eventually only one spe-
PREPRESS AND QUALITY MANAGER: Silvia Di Placido cies, Homo sapiens, was left. We thus find our-
PRINT PRODUCTION MANAGER: Georgina Franco
selves alone and yet the most numerous and
PRODUCTION MANAGER: Christina Hippeli
CUSTOM PUBLISHING MANAGER: successful primates in history.
Madelyn Keyes-Milch Reading the cracked brown fragments of
ASSOCIATE PUBLISHER/VICE PRESIDENT, CIRCULATION: fossils and sequences of DNA, however, scien-
Lorraine Leib Terlecki tists have found clues that the story of human
CIRCULATION DIRECTOR: Katherine Corvino
CIRCULATION PROMOTION MANAGER: origins has more convolutions. The account of
Joanne Guralnick our shared human heritage now includes more
FULFILLMENT AND DISTRIBUTION MANAGER:
controversial plot twists and mysteries. Was the
Rosa Davis MOSAIC of primitive and advanced remarkable seven-million-year-old skull found
VICE PRESIDENT AND PUBLISHER: Bruce Brandfon features marks Sahelanthropus
ASSOCIATE PUBLISHER: Gail Delott
in July 2002 in Chad really one of our first fore-
tchadensis, known from a seven-
SALES DEVELOPMENT MANAGER: David Tirpack million-year-old skull. bears, or a distant dead-end cousin with preco-
SALES REPRESENTATIVES: Stephen Dudley, ciously evolved features? Did modern humans
Hunter Millington, Stan Schmidt, Debra Silver
really originate in Africa alone, as is widely held, or in multiple locales? When
ASSOCIATE PUBLISHER, STRATEGIC PLANNING:
Laura Salant
(and how often) did we emigrate? Were Neandertals the crude, brutish cave-
PROMOTION MANAGER: Diane Schube men of comic strips or— as fresh evidence suggests— did they have a refined,
RESEARCH MANAGER: Aida Dadurian artistic culture? Did they copy and steal toolmaking technologies from the mod-
PROMOTION DESIGN MANAGER: Nancy Mongelli
ern humans, or did they invent them independently? Might they even have con-
GENERAL MANAGER: Michael Florek
ceived children with the moderns? And of course, why didn’t our kind perish
BUSINESS MANAGER: Marie Maher
MANAGER, ADVERTISING ACCOUNTING with the rest of the hominids? Were we luckier, more lingual or just more lethal
AND COORDINATION: Constance Holmes than the rest?
DIRECTOR, SPECIAL PROJECTS: Barth David Schwartz In this special edition from Scientific American, we have collected articles
MANAGING DIRECTOR, SCIENTIFICAMERICAN.COM: about the latest developments in the field of human evolution— written by the
Mina C. Lux experts who are leading the investigations. We invite you to explore the pages
DIRECTOR, ANCILLARY PRODUCTS: Diane McGarvey that follow, to learn more about that fascinating first chapter in everybody’s
PERMISSIONS MANAGER: Linda Hertz family history.
MANAGER OF CUSTOM PUBLISHING: Jeremy A. Abbate
CHAIRMAN EMERITUS: John J. Hanley

CHAIRMAN: Rolf Grisebach
John Rennie
Editor in Chief
KAZUHIKO SANO
PRESIDENT AND CHIEF EXECUTIVE OFFICER:

Gretchen G. Teichgraeber Scientific American
VICE PRESIDENT AND MANAGING DIRECTOR,
INTERNATIONAL: Charles McCullagh
editors@sciam.com
VICE PRESIDENT: Frances Newburg
www.sciam.com NEW LOOK AT HUMAN EVOLUTION 1

contents 2003
SCIENTIFIC AMERICAN Volume 13 Number 2
New Look at Human Evolution

1 Letter from the Editor
ORIGINS
38
4 An Ancestor to Call Our Own
By Kate Wong
Controversial new fossils could bring scientists closer than ever to the origin of humanity.
14 Early Hominid Fossils from Africa

By Meave Leakey and Alan Walker
A recently discovered species of Australopithecus, the ancestor of Homo, pushes back
the onset of bipedalism to some four million years ago.
EMERGENCE
20 Once We Were Not Alone
By Ian Tattersall
We take for granted that Homo sapiens is the only hominid on earth. Yet for at least
four million years, many hominid species shared the planet. What makes us different?
28 Who Were the Neandertals?

By Kate Wong
72 With contributions by Erik Trinkaus and Cidália Duarte;
by João Zilhão and Francesco d’Errico; and by Fred H. Smith
Contentious evidence indicates that these hominids interbred with anatomically
modern humans and sometimes behaved in surprisingly modern ways.
38 Out of Africa Again ... and Again?

By Ian Tattersall
Africa is the birthplace of humanity. But how many human species
evolved there? And when did they emigrate?
46 The Multiregional Evolution of Humans

By Alan G. Thorne and Milford H. Wolpoff
Both fossil and genetic clues argue that ancient ancestors of various
human groups lived where they are found today.
54 The Recent African Genesis of Humans

By Rebecca L. Cann and Allan C. Wilson
Genetic studies reveal that an African woman of 200,000 years ago
was our common ancestor. 28
2 SCIENTIFIC AMERICAN NEW LOOK AT HUMAN EVOLUTION
ADAPTATION
62 Food for Thought
By William R. Leonard
Dietary change was a driving force in human evolution.
72 Skin Deep
By Nina G. Jablonski and George Chaplin 4
Throughout the world, human skin color has developed to be dark
enough to prevent sunlight from destroying the nutrient folate
but light enough to foster the production of vitamin D.
80 The Evolution of Human Birth

By Karen R. Rosenberg and Wenda R. Trevathan
The difficulties of childbirth have probably challenged humans and their ancestors
for millions of years— which means that the modern custom of seeking assistance
during delivery may have a similarly ancient foundation.
86 Once Were Cannibals

By Tim D. White
Clear signs of cannibalism in the human fossil record have been rare,
but it is now becoming apparent that the practice is deeply rooted
in our history.
62
FAST-FORWARD
94 If Humans Were Built to Last

By S. Jay Olshansky, Bruce A. Carnes and Robert N. Butler
We would look a lot different—inside and out— if evolution
had designed the human body to function smoothly
not only in youth but for a century or more.
Cover painting by Kazuhiko Sano. This depiction of Sahelanthropus tchadensis—potentially the

oldest hominid yet found—is based on cranial and dental remains.
Scientific American Special (ISSN 1048-0943), Volume 13, Number 2, 2003, published by Scientific American, Inc.,
415 Madison Avenue, New York, NY 10017-1111. Copyright © 2003 by Scientific American, Inc. All rights reserved.
No part of this issue may be reproduced by any mechanical, photographic or electronic process, or in the form of
a phonographic recording, nor may it be stored in a retrieval system, transmitted or otherwise copied for public
or private use without written permission of the publisher. Canadian BN No. 127387652RT; QST No. Q1015332537.
To purchase additional quantities: U.S., $10.95 each; elsewhere, $13.95 each. Send payment to Scientific Amer-
ican, Dept. EVOL, 415 Madison Avenue, New York, NY 10017-1111. Inquiries: fax 212-355-0408 or telephone 212-
451-8890. Printed in U.S.A.
www.sciam.com SCIENTIFIC AMERICAN 3

ORIGINS
An
Ancestor
to Call Our Own
By Kate Wong
P
OITIERS, FRANCE —Michel Brunet removes the cracked,
brown skull from its padlocked, foam-lined metal car-

rying case and carefully places it on the desk in front of
Controversial
me. It is about the size of a coconut, with a slight snout
new fossils and a thick brow visoring its stony sockets. To my inexpert eye, the
could bring face is at once foreign and inscrutably familiar. To Brunet, a paleon-
tologist at the University of Poitiers, it is the visage of the lost relative
scientists closer he has sought for 26 years. “He is the oldest one,” the veteran fossil
than ever hunter murmurs, “the oldest hominid.”

Brunet and his team set the field of paleoanthropology abuzz when
to the origin they unveiled their find in July 2002. Unearthed from sandstorm-
of humanity scoured deposits in northern Chad’s Djurab Desert, the astonishingly
complete cranium— dubbed Sahelanthropus tchadensis (and nick-
named Toumaï, which means “hope of life” in the local Goran lan-
guage) — dates to nearly seven million years ago. It may thus represent
the earliest human forebear on record, one who Brunet says “could
KAZUHIKO SANO
touch with his finger” the point at which our lineage and the one lead-
ing to our closest living relative, the chimpanzee, diverged.
APE OR ANCESTOR? Sahelanthropus tchadensis, potentially the oldest hominid on

record, forages in a woodland bordering Lake Chad some seven million years ago.
Thus far the creature is known only from cranial and dental remains, so its body in
this artist’s depiction is entirely conjectural.
4 SCIENTIFIC AMERICAN Updated from the January 2003 issue

Less than a century ago simian human precursors from
Africa existed only in the minds of an enlightened few. Charles AFRICAN ROOTS
Darwin predicted in 1871 that the earliest ancestors of humans
would be found in Africa, where our chimpanzee and gorilla RECENT FINDS from Africa could extend in time and space the fossil
cousins live today. But evidence to support that idea didn’t record of early human ancestors. Just a few years ago remains more
come until more than 50 years later, when anatomist Raymond than 4.4 million years old were essentially unknown, and the oldest
Dart of the University of the Witwatersrand described a fossil specimens all came from East Africa. In 2001 paleontologists
skull from Taung, South Africa, as belonging to an extinct hu- working in Kenya’s Tugen Hills and Ethiopia’s Middle Awash region
man he called Australopithecus africanus, the “southern ape announced that they had discovered hominids dating back to nearly
from Africa.” His claim met variously with frosty skepticism six million years ago (Orrorin tugenensis and Ardipithecus ramidus
and outright rejection— the remains were those of a juvenile kadabba, respectively). Then, in July 2002, University of Poitiers
gorilla, critics countered. The discovery of another South
African specimen, now recognized as A. robustus, eventually
vindicated Dart, but it wasn’t until the 1950s that the notion
of ancient, apelike human ancestors from Africa gained wide-
spread acceptance.
In the decades that followed, pioneering efforts in East
Africa headed by members of the Leakey family, among oth-
ers, turned up additional fossils. By the late 1970s the austra-
lopithecine cast of characters had grown to include A. boisei,
A. aethiopicus and A. afarensis (Lucy and her kind, who lived
between 2.9 million and 3.6 million years ago during the
Pliocene epoch and gave rise to our own genus, Homo). Each
was adapted to its own environmental niche, but all were bi-
pedal creatures with thick jaws, large molars and small ca-
nines— radically different from the generalized, quadrupedal
Miocene apes known from farther back on the family tree. To
probe human origins beyond A. afarensis, however, was to fall
into a gaping hole in the fossil record between 3.6 million and
12 million years ago. Who, researchers wondered, were Lucy’s
forebears?
Despite widespread searching, diagnostic fossils of the right Sahelanthropus tchadensis
from Toros-Menalla, Chad
age to answer that question eluded workers for nearly two
decades. Their luck finally began to change around the mid-
1990s, when a team led by Meave Leakey of the National Mu-
seums of Kenya announced its discovery of A. anamensis, a
four-million-year-old species that, with its slightly more archaic
characteristics, made a reasonable ancestor for Lucy [see “Ear-
ly Hominid Fossils from Africa,” on page 14]. At around the
Overview/The Oldest Hominids

■ The typical textbook account of human evolution holds
that humans arose from a chimpanzeelike ancestor
between roughly five million and six million years ago in
East Africa and became bipedal on the savanna. But until
recently, hominid fossils more than 4.4 million years old
were virtually unknown.
■ Newly discovered fossils from Chad, Kenya and Ethiopia
may extend the human record back to seven million years

ago, revealing the earliest hominids yet.
■ These finds cast doubt on conventional paleoanthro-
pological wisdom. But experts disagree over how these

creatures are related to humans— if they are related at all.

paleontologist Michel Brunet and his Franco-Chadian
Paleoanthropological Mission reported having unearthed a nearly
PATRICK ROBERT Corbis Sygma (Sahelanthropus tchadensis skull); © 1999 TIM D. WHITE Brill Atlanta\National Museum of Ethiopia (A. r. kadabba fossils); GAMMA (O. tugenensis fossils); EDWARD BELL (map illustration)
seven-million-year-old hominid, called Sahelanthropus tchadensis, Ardipithecus ramidus kadabba
from Middle Awash, Ethiopia
at a site known as Toros-Menalla in northern Chad. The site lies some
2,500 kilometers west of the East African fossil localities. “I think
the most important thing we have done in terms of trying to
understand our story is to open this new window,” Brunet remarks.
“We are proud to be the pioneers of the West.”
HADAR
A. afarensis
MIDDLE AWASH
A. afarensis
A. garhi
CHAD Ardipithecus ramidus kadabba
A. r. ramidus
TOROS-MENALLA KONSO
Sahelanthropus tchadensis A. boisei
OMO Orrorin tugenensis

A. afarensis from Tugen Hills, Kenya
A. aethiopicus
ETHIOPIA A. boisei
WEST TURKANA
A. aethiopicus
A. boisei KOOBI FORA
A. boisei
A. afarensis
LOMEKWI ALLIA BAY
Kenyanthropus KENYA KANAPOI A. anamensis
platyops A. anamensis
TUGEN HILLS
Orrorin tugenensis
TANZANIA OLDUVAI GORGE
A. boisei
LAETOLI
A. afarensis
MAKAPANSGAT
A. africanus
KROMDRAAI
A. robustus
DRIMOLEN
A. robustus
SOUTH SWARTKRANS
AFRICA A. robustus
TAUNG STERKFONTEIN
Australopithecus A. africanus
africanus

It is the visage of the lost relative he has sought
for 26 years. “He is the oldest one,” the veteran
fossil hunter murmurs, “the oldest hominid.”
same time, Tim D. White of the University of California at ning discoveries— Brunet’s among them— that may go a long
Berkeley and his colleagues described a collection of 4.4-mil- way toward bridging the remaining gap between humans and
lion-year-old fossils recovered in Ethiopia that represent an their African ape ancestors. These fossils, which range from
even more primitive hominid, now known as Ardipithecus roughly five million to seven million years old, are upending
ramidus ramidus. Those findings gave scholars a tantalizing long-held ideas about when and where our lineage arose and
glimpse into Lucy’s past. But estimates from some molecular what the last common ancestor of humans and chimpanzees
biologists of when the split between chimps and humans oc- looked like.
curred suggested that even older hominids lay waiting some- Not surprisingly, they have also sparked vigorous debate.
where to be discovered. Indeed, experts are deeply divided over where on the family
Those intriguing predictions have recently been borne out. tree the new species belong and even what constitutes a hom-
Over the past few years, researchers have made a string of stun- inid in the first place.
ANATOMY OF AN ANCESTOR
KEY TRAITS link putative hominids Ardipithecus ramidus kadabba, Orrorin and Sahelanthropus to humans and distinguish
them from apes such as chimpanzees. The fossils exhibit primitive apelike characteristics, too, as would be expected of
creatures this ancient. For instance, the A. r. kadabba toe bone has a humanlike upward tilt to its joint surface, but the bone is
long and curves downward like a chimp’s does (which somewhat obscures the joint’s cant). Likewise, Sahelanthropus has a
number of apelike traits— its small braincase among them— but is more humanlike in the form of the canines and the
projection of the lower face. (Reconstruction
of the Sahelanthropus cranium, which is CRANIUM
distorted, will give researchers a better Modern human
understanding of its morphology.) The Orrorin
Sahelanthropus Chimpanzee
femur has a long neck and a groove carved
out by the obturator externus muscle— traits
typically associated with habitual bipedalism
and therefore with humans—but the distribution
of cortical bone in the femoral neck may be
more like that of a quadrupedal ape.
Large,
TOE BONE Small, more sharp
incisorlike canine canine
Modern
human A. r. kadabba Chimpanzee
Joint Joint surface
surface cants downward
cants
upward
Vertical
lower
face
Moderately projecting Strongly
lower face projecting
lower face
© C. OWEN LOVEJOY\Brill Atlanta (human, A. r. kadabba and chimpanzee toe bones); CHRISTIAN SIDOR New York College of Osteopathic Medicine (human skull and human femur);
MISSION PALÉOANTHROPOLOGIQUE FRANCO-TCHADIENNE (Sahelanthropus skull); © 1996 DAVID L. BRILL\DIVISION OF MAMMALS, NATIONAL MUSEUM OF NATURAL HISTORY, SMITHSONIAN
INSTITUTION (chimpanzee skull); GAMMA (Orrorin femur); C. OWEN LOVEJOY Kent State University (chimpanzee femur)

Standing Tall large and pointed relative to human canines, and its arm and
THE FIRST HOMINID CLUE to come from beyond the 4.4- finger bones retain adaptations for climbing. But the femur
million-year mark was announced in the spring of 2001. Pa- characteristics signify to Pickford and Senut that when it was
leontologists Martin Pickford and Brigitte Senut of the Na- on the ground, Orrorin walked like a man.
tional Museum of Natural History in Paris found in Kenya’s In fact, they argue, Orrorin appears to have had a more hu-
Tugen Hills the six-million-year-old remains of a creature they manlike gait than the much younger Lucy did. Breaking with
called Orrorin tugenensis. To date, the researchers have paleoanthropological dogma, the team posits that Orrorin gave
amassed 21 specimens, including bits of jaw, isolated teeth, fin- rise to Homo via the proposed genus Praeanthropus (which
ger and arm bones, and some partial upper leg bones, or fe- comprises a subset of the fossils currently assigned to A. afaren-
murs. According to Pickford and Senut, Orrorin exhibits sev- sis and A. anamensis), leaving Lucy and her kin on an evolu-
eral characteristics that clearly align it with the hominid fam- tionary sideline. Ardipithecus, they believe, was a chimpanzee
ily— notably those suggesting that, like all later members of our ancestor.
group, it walked on two legs. “The femur is remarkably hu- Not everyone is persuaded by the femur argument. C. Owen
manlike,” Pickford observes. It has a long femoral neck, which Lovejoy of Kent State University counters that published com-
would have placed the shaft at an angle relative to the lower puted tomography scans through Orrorin’s femoral neck—
leg (thereby stabilizing the hip), and a groove on the back of which Pickford and Senut say reveal humanlike bone struc-
that femoral neck, where a muscle known as the obturator ex- ture— actually show a chimplike distribution of cortical bone,
ternus pressed against the bone during upright walking. In oth- an important indicator of the strain placed on that part of the
er respects, Orrorin was a primitive animal: its canine teeth are femur during locomotion. Cross sections of A. afarensis’s fe-
moral neck, in contrast, look entirely human, he states. Love-
joy suspects that Orrorin was frequently— but not habitually—
bipedal and spent a significant amount of time in the trees. That
wouldn’t exclude it from hominid status, because full-blown
FEMUR bipedalism almost certainly didn’t emerge in one fell swoop.
Rather Orrorin may have simply not yet evolved the full com-
Modern human Orrorin Chimpanzee plement of traits required for habitual bipedalism. Viewed that
Long femoral neck Short femoral neck way, Orrorin could still be on the ancestral line, albeit further
removed from Homo than Pickford and Senut would have it.
Better evidence of early routine bipedalism, in Lovejoy’s
view, surfaced a few months after the Orrorin report, when
Berkeley graduate student Yohannes Haile-Selassie announced
the discovery of slightly younger fossils from Ethiopia’s Middle
Location of
obturator No Awash region. Those 5.2-million- to 5.8-million-year-old re-
Location of obturator
externus mains, which have been classified as a subspecies of Ardi-
obturator groove externus
externus groove pithecus ramidus, A. r. kadabba, include a complete foot pha-
groove lanx, or toe bone, bearing a telltale trait. The bone’s joint is an-
gled in precisely the way one would expect if A. r. kadabba
“toed off” as humans do when walking, reports Lovejoy, who
has studied the fossil.
Other workers are less impressed by the toe morphology.
“To me, it looks for all the world like a chimpanzee foot pha-
lanx,” comments David Begun of the University of Toronto,
noting from photographs that it is longer, slimmer and more
curved than a biped’s toe bone should be. Clarification may
come when White and his collaborators publish findings on an
as yet undescribed partial skeleton of Ardipithecus, which
White says they hope to do within the next year or two.
Differing anatomical interpretations notwithstanding, if ei-
ther Orrorin or A. r. kadabba were a biped, that would not
only push the origin of our strange mode of locomotion back
by nearly 1.5 million years, it would also lay to rest a popular
idea about the conditions under which our striding gait
evolved. Received wisdom holds that our ancestors became
bipedal on the African savanna, where upright walking may
have kept the blistering sun off their backs, given them access

Humanity may have arisen more than a million years
earlier than a number of molecular studies had estimated. More
important, it may have originated in a different locale.
to previously out-of-reach foods, or afforded them a better coveries when Brunet’s fossil find from Chad came to light.
view above the tall grass. But paleoecological analyses indicate With Sahelanthropus have come new answers— and new ques-
that Orrorin and Ardipithecus dwelled in forested habitats, tions. Unlike Orrorin and A. r. kadabba, the Sahelanthropus
alongside monkeys and other typically woodland creatures. In material does not include any postcranial bones, making it im-
fact, Giday WoldeGabriel of Los Alamos National Laborato- possible at this point to know whether the animal was bipedal,
ry and his colleagues, who studied the soil chemistry and ani- the traditional hallmark of humanness. But Brunet argues that
mal remains at the A. r. kadabba site, have noted that early a suite of features in the teeth and skull, which he believes be-
hominids may not have ventured beyond these relatively wet longs to a male, judging from the massive brow ridge, clearly
and wooded settings until after 4.4 million years ago. links this creature to all later hominids. Characteristics of Sa-
If so, climate change may not have played as important a helanthropus’s canines are especially important in his assess-
role in driving our ancestors from four legs to two as has been ment. In all modern and fossil apes, and therefore presumably
thought. For his part, Lovejoy observes that a number of the in the last common ancestor of chimps and humans, the large
savanna-based hypotheses focusing on posture were not espe- upper canines are honed against the first lower premolars, pro-
cially well conceived to begin with. “If your eyes were in your ducing a sharp edge along the back of the canines. This so-
toes, you could stand on your hands all day and look over tall called honing canine-premolar complex is pronounced in
grass, but you’d never evolve into a hand-walker,” he jokes. In males, who use their canines to compete with one another for
other words, selection for upright posture alone would not, in females. Humans lost these fighting teeth, evolving smaller,
his view, have led to bipedal locomotion. The most plausible more incisorlike canines that occlude tip to tip, an arrangement
explanation for the emergence of bipedalism, Lovejoy says, is that creates a distinctive wear pattern over time. In their size,
that it freed the hands and allowed males to collect extra food shape and wear, the Sahelanthropus canines are modified in
with which to woo mates. In this model, which he developed the human direction, Brunet asserts.
in the 1980s, females who chose good providers could devote At the same time, Sahelanthropus exhibits a number of
more energy to child rearing, thereby maximizing their repro- apelike traits, such as its small braincase and widely spaced eye
ductive success. sockets. This mosaic of primitive and advanced features,
WITNESS/GAMMA
Brunet says, suggests a close relationship to the last common
The Oldest Ancestor? ancestor. Thus, he proposes that Sahelanthropus is the earliest
THE PALEOANTHROPOLOGICAL community was still di- member of the human lineage and the ancestor of all later hom-
gesting the implications of the Orrorin and A. r. kadabba dis- inids, including Orrorin and Ardipithecus. If Brunet is correct,
HUNTING FOR HOMINIDS:

Michel Brunet (left),
whose team uncovered
Sahelanthropus, has
combed the sands of the
Djurab Desert in Chad for
nearly a decade. Martin
Pickford and Brigitte
Senut (center) discovered
Orrorin in Kenya’s Tugen
Hills. Tim White (top right)
and Yohannes Haile-
Selassie (bottom right)
found Ardipithecus in the
Middle Awash region
of Ethiopia.

humanity may have arisen more than a million years earlier manlike as Brunet claims. Along similar lines, in a letter pub-
than a number of molecular studies had estimated. More im- lished last October in the journal Nature, in which Brunet’s
portant, it may have originated in a different locale than has team initially reported its findings, University of Michigan pa-
been posited. According to one model of human origins, put leoanthropologist Milford H. Wolpoff, along with Orrorin dis-
forth in the 1980s by Yves Coppens of the College of France, coverers Pickford and Senut, countered that Sahelanthropus
East Africa was the birthplace of humankind. Coppens, not- was an ape rather than a hominid. The massive brow and cer-
ing that the oldest human fossils came from East Africa, pro- tain features on the base and rear of Sahelanthropus’s skull,
posed that the continent’s Rift Valley— a gash that runs from they observed, call to mind the anatomy of a quadrupedal ape
north to south— split a single ancestral ape species into two with a difficult-to-chew diet, whereas the small canine suggests
populations. The one in the east gave rise to humans; the one that it was a female of such a species, not a male human an-
© 1998 DAVID L. BRILL Brill Atlanta (top); GAMMA (bottom left); © 1998 DAVID L. BRILL Brill Atlanta (bottom left)
in the west spawned today’s apes [see “East Side Story: The cestor. Lacking proof that Sahelanthropus was bipedal, so their
Origin of Humankind,” by Yves Coppens; Scientific Amer- reasoning goes, Brunet doesn’t have a leg to stand on. (Pick-
ican, May 1994]. Scholars have recognized for some time that ford and Senut further argue that the animal was specifically
the apparent geographic separation might instead be an arti- a gorilla ancestor.) In a barbed response, Brunet likened his de-
fact of the scant fossil record. The discovery of a seven-million- tractors to those Dart encountered in 1925, retorting that
year-old hominid in Chad, some 2,500 kilometers west of the
Rift Valley, would deal the theory a fatal blow.
Most surprising of all may be what Sahelanthropus reveals
about the last common ancestor of humans and chimpanzees.
Paleoanthropologists have typically imagined that that crea-
ture resembled a chimp in having, among other things, a
strongly projecting lower face, thinly enameled molars and
large canines. Yet Sahelanthropus, for all its generally apelike
traits, has only a moderately prognathic face, relatively thick
enamel, small canines and a brow ridge larger than that of any
living ape. “If Sahelanthropus shows us anything, it shows us
that the last common ancestor was not a chimpanzee,” Berke-
ley’s White remarks. “But why should we have expected oth-
erwise?” Chimpanzees have had just as much time to evolve as
humans have had, he points out, and they have become high-
ly specialized, fruit-eating apes.
Brunet’s characterization of the Chadian remains as those
of a human ancestor has not gone unchallenged, however.
“Why Sahelanthropus is necessarily a hominid is not particu-
larly clear,” comments Carol V. Ward of the University of Mis-
souri. She and others are skeptical that the canines are as hu-

HOMINIDS IN TIME
FOSSIL RECORD OF HOMINIDS shows that multiple species existed alongside one another
during the later stages of human evolution. Whether the same can be said for the first
half of our family’s existence is a matter of great debate among paleoanthropologists,
however. Some believe that all the fossils from between seven million and three million
years ago fit comfortably into the same evolutionary lineage. Others view these
specimens not only as members of mostly different lineages but also as representatives
of a tremendous early hominid diversity yet to be discovered. (Adherents to the latter
scenario tend to parse the known hominid remains into more taxa than shown here.)
The branching diagrams (inset) illustrate two competing hypotheses of how the
recently discovered Sahelanthropus, Orrorin and Ardipithecus ramidus kadabba are
related to humans. In the tree on the left, all the new finds reside on the line leading to
humans, with Sahelanthropus being the oldest known hominid. In the tree on the right, in Kenyanthropus platyops A. garhi
contrast, only Orrorin is a human ancestor. Ardipithecus is a chimpanzee ancestor and
Sahelanthropus a gorilla forebear in this view.
A. africanus
A. r. ramidus
Orrorin
tugenensis
A. aethiopicus
Ardipithecus
ramidus kadabba
Sahelanthropus tchadensis Australopithecus anamensis A. afarensis

7 6 5 4 3
Millions of Years Ago
Sahelanthropus’s apelike traits are simply primitive holdovers Thus, cladistically “what a hominid is from the point of
from its own ape predecessor and therefore uninformative with view of skeletal morphology is summarized by those charac-
regard to its relationship to humans. ters preserved in the skeleton that are present in populations
The conflicting views partly reflect the fact that researchers that directly succeeded the genetic splitting event between
disagree over what makes the human lineage unique. “We have chimps and humans,” explains William H. Kimbel of Arizona
trouble defining hominids,” acknowledges Roberto Macchiar- State University. With only an impoverished fossil record to
elli, also at the University of Poitiers. Traditionally paleoanthro- work from, paleontologists can’t know for certain what those
pologists have regarded bipedalism as the characteristic that traits were. But the two leading candidates for the title of sem-
first set human ancestors apart from other apes. But subtler inal hominid characteristic, Kimbel says, are bipedalism and
changes— the metamorphosis of the canine, for instance— may the transformation of the canine. The problem researchers now ILLUSTRATIONS BY PATRICIA J. WYNNE AND CORNELIA BLIK
have preceded that shift. face in trying to suss out what the initial changes were and
To understand how animals are related to one another, evo- which, if any, of the new putative hominids sits at the base of
lutionary biologists employ a method called cladistics, in which the human clade is that so far Orrorin, A. r. kadabba and Sa-
organisms are grouped according to shared, newly evolved traits. helanthropus are represented by mostly different bony ele-
In short, creatures that have these derived characteristics in comments, making comparisons among them difficult.
mon are deemed more closely related to one another than they
are to those that exhibit only primitive traits inherited from a How Many Hominids?
more distant common ancestor. The first occurrence in the fos- M E A N W H I L E T H E A R R I V A L of three new taxa to the table
sil record of a shared, newly acquired trait serves as a baseline has intensified debate over just how diverse early hominids
indicator of the biological division of an ancestral species into were. Experts concur that between three million and 1.5 mil-
two daughter species— in this case, the point at which chimps lion years ago, multiple hominid species existed alongside one
and humans diverged from their common ancestor— and that another at least occasionally. Now some scholars argue that
trait is considered the defining characteristic of the group. this rash of discoveries demonstrates that human evolution was

years ago, only one hominid species is known to have existed
at any given time. “Where’s the bush?” Brunet demands. Even
at humanity’s peak diversity, two million years ago, White
GORILLA CHIMP HUMAN GORILLA CHIMP HUMAN says, there were only three taxa sharing the landscape. “That
ain’t the Cambrian explosion,” he remarks dryly. Rather,
White suggests, there is no evidence that the base of the fami-
ly tree is anything other than a trunk. He thinks that the new
finds might all represent snapshots of the Ardipithecus lineage
through time, with Sahelanthropus being the earliest hominid
and with Orrorin and A. r. kadabba representing its lineal de-
Homo habilis Sahelanthropus Orrorin A. r. kadabba scendants. (In this configuration, Sahelanthropus and Orror-
in would become species of Ardipithecus.)
Investigators agree that more fossils are needed to elucidate
how Orrorin, A. r. kadabba and Sahelanthropus are related to
one another and to ourselves, but obtaining a higher-resolu-
H. erectus tion picture of the roots of humankind won’t be easy. “We’re
going to have a lot of trouble diagnosing the very earliest mem-
bers of our clade the closer we get to that last common ances-
tor,” Missouri’s Ward predicts. Nevertheless, “it’s really im-
portant to sort out what the starting point was,” she observes.
H. sapiens
“Why the human lineage began is the question we’re trying to
answer, and these new finds in some ways may hold the key
A. robustus
to answering that question— or getting closer than we’ve ever
gotten before.”
It may be that future paleoanthropologists will reach a point
at which identifying an even earlier hominid will be well nigh
A. boisei impossible. But it’s unlikely that this will keep them from try-
ing. Indeed, it would seem that the search for the first hominids
is just heating up. “The Sahelanthropus cranium is a messenger
2 1 PRESENT [indicating] that in central Africa there is a desert full of fossils
of the right age to answer key questions about the genesis of our
clade,” White reflects. For his part, Brunet, who for more than
a complex affair from the outset. Toronto’s Begun— who be- a quarter of a century has doggedly pursued his vision through
lieves that the Miocene ape ancestors of modern African apes political unrest, sweltering heat and the blinding sting of an un-
and humans spent their evolutionarily formative years in Eu- relenting desert wind, says that ongoing work in Chad will keep
rope and western Asia before reentering Africa— observes that his team busy for years to come. “This is the beginning of the
Sahelanthropus bears exactly the kind of motley features that story,” he promises, “just the beginning.” As I sit in Brunet’s of-
one would expect to see in an animal that was part of an adap- fice contemplating the seven-million-year-old skull of Sahelan-
tive radiation of apes moving into a new milieu. “It would not thropus, the fossil hunter’s quest doesn’t seem quite so unimag-
surprise me if there were 10 or 15 genera of things that are inable. Many of us spend the better part of a lifetime searching
more closely related to Homo than to chimps,” he says. Like- for ourselves.
wise, in a commentary that accompanied the report by Brunet
and his team in Nature, Bernard Wood of George Washington Kate Wong is editorial director of ScientificAmerican.com
University wondered whether Sahelanthropus might hail from
the African ape equivalent of Canada’s famed Burgess Shale, MORE TO E XPLORE
which has yielded myriad invertebrate fossils from the Cam- Late Miocene Hominids from the Middle Awash, Ethiopia. Yohannes
Haile-Selassie in Nature, Vol. 412, pages 178–181; July 12, 2001.
brian period, when the major modern animal groups explod-
Extinct Humans. Ian Tattersall and Jeffrey H. Schwartz. Westview
ed into existence. Viewed that way, the human evolutionary Press, 2001.
tree would look more like an unkempt bush, with some, if not Bipedalism in Orrorin tugenensis Revealed by Its Femora. Martin
all, of the new discoveries occupying terminal twigs instead of Pickford, Brigitte Senut, Dominique Gommercy and Jacques Treil in
coveted spots on the meandering line that led to humans. Comptes Rendus: Palevol, Vol. 1, No. 1, pages 1–13; 2002.
Other workers caution against inferring the existence of A New Hominid from the Upper Miocene of Chad, Central Africa.
Michel Brunet, Franck Guy, David Pilbeam, Hassane Taisso Mackaye
multiple, coeval hominids on the basis of what has yet been et al. in Nature, Vol. 418, pages 145–151; July 11, 2002.
found. “That’s X-Files paleontology,” White quips. He and The Primate Fossil Record. Edited by Walter C. Hartwig. Cambridge
Brunet both note that between seven million and four million University Press, 2002.

ORIGINS
early hominid fossils from
AFRICA
The year was 1965.
Bryan Patterson, a paleoanthropologist ern humans than the one other Australo- Yet Patterson’s fossil would eventu-
from Harvard University, unearthed a pithecus humerus known at the time. ally help establish the existence of a new
fragment of a fossil arm bone at a site And yet the age of the Kanapoi fossil species of Australopithecus— the oldest
called Kanapoi in northern Kenya. He proved somewhat surprising. Although yet to be identified— and push back the
and his colleagues knew it would be hard the techniques for dating the rocks where origins of upright walking to more than
to make a great deal of anatomical or the fossil was uncovered were still fairly four million years ago. But to see how
evolutionary sense out of a small piece of rudimentary, the group working in Ken- this happened, we need to trace the steps
elbow joint. Nevertheless, they did rec- ya was able to show that the bone was that paleoanthropologists have taken in
ognize some features reminiscent of a probably older than the various Austra- constructing an outline for the story of
species of early hominid (a hominid is lopithecus specimens that had previous- hominid evolution.
MATT MAHURIN (illustration); ROBERT CAMPBELL (left); ALAN WALKER; © NATIONAL MUSEUMS OF KENYA (center and right)
any upright-walking primate) known as ly been found. Despite this unusual result,
Australopithecus, first discovered 40 however, the significance of Patterson’s An Evolving Story
years earlier in South Africa by Raymond discovery was not to be confirmed for an- S C I E N T I S T S C L A S S I F Y the immediate
Dart of the University of the Witwater- other 30 years. In the interim, researchers ancestors of the genus Homo (which in-
srand. In most details, however, Patterson identified the remains of so many impor- cludes our own species, Homo sapiens)
and his team considered the fragment of tant early hominids that the humerus in the genus Australopithecus. For sev-
arm bone to be more like those of mod- from Kanapoi was rather forgotten. eral decades it was believed that these
ancient hominids first inhabited the
AUSTRALOPITHECUS earth at least three and a half million
ANAMENSIS (right) lived
roughly four million
years ago. The specimens found in South
years ago. Only a few Africa by Dart and others indicated that
anamensis fossils have there were at least two types of Austra-
been found— the ones lopithecus— A. africanus and A. robus-
shown at the left tus. The leg bones of both species sug-
include a jawbone and
part of the front of the
gested that they had the striding, bipedal
face (left), parts of an locomotion that is a hallmark of humans
arm bone (center) and among living mammals. (The upright
fragments of a lower leg posture of these creatures was vividly
bone (right)— and thus confirmed in 1978 at the Laetoli site in
researchers cannot
determine much about
Tanzania, where a team led by archae-
the species’ physical ologist Mary Leakey discovered a spec-
appearance. But tacular series of footprints made 3.6 mil-
scientists have lion years ago by three Australopithecus
established that individuals as they walked across wet
anamensis walked
upright, making it the
volcanic ash.) Both A. africanus and A.
earliest bipedal creature robustus were relatively small-brained
yet to be discovered. and had canine teeth that differed from
14 SCIENTIFIC AMERICAN Updated

NEW LOOK
from AT
theHUMAN
June 1997
EVOLUTION
issue
A new species of
Australopithecus,
the ancestor of Homo,
pushes back the origins
of bipedalism to some
four million years ago
By Meave Leakey and Alan Walker
those of modern apes in that they hard- of hominid bones and teeth discovered
ly projected past the rest of the tooth at Laetoli, as well as a large and very im-
row. The younger of the two species, A. portant collection of specimens from the
robustus, had bizarre adaptations for Hadar region of Ethiopia (including the
chewing— huge molar and premolar famous “Lucy” skeleton). The group
teeth combined with bony crests on the named the new species afarensis. Radio-
skull where powerful chewing muscles metric dating revealed that the species
would have been attached. had lived between 3.6 and 2.9 million
Paleoanthropologists identified more years ago, making it the oldest Aus-
species of Australopithecus over the next tralopithecus known at the time.
several decades. In 1959 Mary Leakey This early species is probably the best
unearthed a skull from yet another East studied of all the Australopithecus rec-
African species closely related to robus- ognized so far, and it is certainly the one
tus. Skulls of these species uncovered that has generated the most controversy
during the past 45 years in the north- over the past 30 years. The debates have
eastern part of Africa, in Ethiopia and ranged over many issues: whether the
Kenya, differed considerably from those afarensis fossils were truly distinct from
found in South Africa; as a result, re- the africanus fossils from South Africa;
searchers think that two separate robus- whether there was one or several species
tus-like species— a northern one and a at Hadar; whether the Tanzanian and
southern one— existed. Ethiopian fossils were of the same spe-
In 1978 Donald C. Johanson, now at cies; and whether the fossils had been
the Institute of Human Origins at Ari- dated correctly.
zona State University, along with his col- But the most divisive debate con-
leagues, identified still another species of cerns the issue of how extensively the
Australopithecus. Johanson and his bipedal afarensis climbed in trees. Fossils
team had been studying a small number of afarensis include various bone and

joint structures typical of tree climbers. BONOBO
Some scientists argue that such charac-
teristics indicate that these hominids CHIMPANZEE
must have spent at least some time in the HOMO
trees. But others view these features as Ardipithecus ?
simply evolutionary baggage, left over Orrorin ramidus
Au. afarensis Au. robustus
from arboreal ancestors. Underlying this
Au. africanus
discussion is the question of where Aus-
tralopithecus lived— in forests or on the Sahelanthropus Au. anamensis
open savanna.
Au. aethiopicus
By the beginning of the 1990s, re-
searchers knew a fair amount about the Au. boisei
various species of Australopithecus and
how each had adapted to its environ-
6 MYR 5 MYR 4 MYR 3 MYR 2 MYR 1 MYR
mental niche. A description of any one of AGO AGO AGO AGO AGO AGO
the species would mention that the crea-
FAMILY TREE of the hominid Australopithecus (red) includes a number of species that lived between
tures were bipedal and that they had ape-
roughly 4 million and 1.25 million years (Myr) ago. Just over 2 Myr ago a new genus, Homo (which
size brains and large, thickly enameled includes our own species, H. sapiens), evolved from one of the species of Australopithecus.
teeth in strong jaws, with nonprojecting
canines. Males were typically larger than 1982, expeditions run by the National River dominated the Turkana area for
females, and individuals grew and ma- Museums of Kenya to the Lake Turkana much of the Pliocene (roughly 5.3 to 1.8
tured rapidly. But the origins of Aus- basin in northern Kenya began finding million years ago) and the early Pleis-
tralopithecus were only hinted at, because hominid fossils nearly four million years tocene (1.8 to 0.7 million years ago). Only
the gap between the earliest well-known old. But because these fossils were main- infrequently was a lake present in the
species in the group (afarensis, from ly isolated teeth— no jawbones or skulls area at all. Instead, for most of the past
about 3.6 million years ago) and the pos- were preserved— very little could be said four million years, an extensive river sys-
tulated time of the last common ancestor about them except that they resembled tem flowed across the broad floodplain,
of chimpanzees and humans (about six the remains of afarensis from Laetoli. proceeding to the Indian Ocean without
million years ago, according to molecular But our excavations at an unusual site, dumping its sediments into a lake.
evidence) was still very great. Fossil just inland from Allia Bay on the east The Allia Bay fossils are located in
hunters had unearthed only a few older side of Lake Turkana [see maps on page one of the channels of this ancient river
fragments of bone, tooth and jaw from 18], yielded more complete fossils. system. Most of the fossils collected
the intervening 1.5 million years to indi- The site at Allia Bay is a bone bed, from Allia Bay are rolled and weathered
cate the anatomy and course of evolution where millions of fragments of weath- bones and teeth of aquatic animals—
of the earliest hominids. ered tooth and bone from a wide variety fish, crocodiles, hippopotamuses and the
of animals, including hominids, spill out like— that were damaged during trans-
Filling the Gap of the hillside. Exposed at the top of the port down the river from some distance
DISCOVERIES IN KENYA over the hill lies a layer of hardened volcanic ash away. But some of the fossils are much
past several years have filled in some of called the Moiti Tuff, which has been better preserved; these come from the
the missing interval between 3.5 million dated radiometrically to just over 3.9 animals that lived on or near the river-
and 5 million years ago. Beginning in million years old. The fossil fragments banks. Among these creatures are sever-
lie several meters below the tuff, indi- al different species of leaf-eating mon-
THE AUTHORS
MEAVE LEAKEY and ALAN WALKER, to- cating that the remains are older than keys, related to modern colobus mon-
gether with Leakey’s husband, Richard, the tuff. We do not yet understand fully keys, as well as antelopes whose living
have collaborated for many years on the why so many fossils are concentrated in relatives favor closely wooded areas.
discovery and analysis of early hominid this spot, but we can be certain that they Reasonably well preserved hominid fos-
fossils from Kenya. Meave Leakey is a were deposited by the precursor of the sils can also be found here, suggesting
researcher and former head of the divi- present-day Omo River. that, at least occasionally, early homi-
sion of paleontology at the National Mu- Today the Omo drains the Ethiopian nids inhabited a riparian habitat.
seums of Kenya in Nairobi. Walker is highlands located to the north, emptying Where do these Australopithecus
Evan Pugh Professor of Anthropology into Lake Turkana, which has no outlet. fossils fit in the evolutionary history of
and Biology at Pennsylvania State Uni- But this has not always been so. Our col- hominids? The jaws and teeth from Al-
versity. He is a MacArthur Fellow and a leagues Frank Brown of the University of lia Bay, as well as a nearly complete ra-
SLIM FILMS
member of the American Academy of Utah and Craig Feibel of Rutgers Uni- dius (the outside bone of the forearm)
Arts and Sciences. versity have shown that the ancient Omo from the nearby sediments of Sibilot just

CHIMPANZEE ANAMENSIS HUMAN
MANDIBLE
The human jaw

widens at the
back of the
mouth
The jawbones
in anamensis and
chimpanzees are
U-shaped
TIBIA
The top of the

tibia, near the In the tibias of anamensis
knee, is and humans, the top of the
somewhat bone is wider because of
T-shaped in the extra spongy bone
chimpanzee tissue present, which
serves as a shock absorber
Primates such as in bipedal creatures
chimpanzees that
walk on their
knuckles have a
HUMERUS deep, oval hollow at
the bottom of the
ALAN WALKER; © NATIONAL MUSEUMS OF KENYA (chimpanzee and anamensis); VIDEO SURGERY Photo Researchers, Inc. (human)
humerus where the Human and

humerus and the anamensis
ulna lock in place, bones lack this
making the elbow feature,
joint more stable suggesting
that, like
humans,
anamensis did
not walk on its
knuckles
FOSSILS from anamensis (center) share a number of features in common their interrelationships and thereby piece together the course of
with both humans (right) and modern chimpanzees (left). Scientists hominid evolution since the lineages of chimpanzees and humans
use the similarities and differences among these species to determine split some five or six million years ago.
to the north, show an interesting mix- story began. One of us (Leakey) has the layers of sediment, also carried out
ture of characteristics. Some of the traits mounted expeditions from the National by Feibel, reveal that the fossils here
are primitive ones— that is, they are an- Museums of Kenya to explore the sedi- have been preserved by deposits from a
cestral features thought to be present be- ments located southwest of Lake Turka- river ancestral to the present-day Kerio
fore the split occurred between the chim- na and to document the faunas present River, which once flowed into the Tur-
panzee and human lineages. Yet these during the earliest stages of the basin’s kana basin and emptied into an ancient
bones also share characteristics seen in history. Kanapoi, virtually unexplored lake that we call Lonyumun. This lake
later hominids and are therefore said to since Patterson’s day, has proved to be reached its maximum size about 4.1 mil-
have more advanced features. As our one of the most rewarding sites in the lion years ago and thereafter shrank as
team continues to unearth more bones Turkana region. it filled with sediments.
and teeth at Allia Bay, these new fossils A series of deep erosion gullies, known Excavations at Kanapoi have pri-
add to our knowledge of the wide range as badlands, has exposed the sediments at marily yielded the remains of carnivore
of traits present in early hominids. Kanapoi. Fossil hunting is difficult here, meals, so the fossils are rather fragmen-
Across Lake Turkana, some 145 kilo- though, because of a carapace of lava tary. But workers at the site have also re-
meters (about 90 miles) south of Allia pebbles and gravel that makes it hard to covered two nearly complete lower jaws,
Bay, lies the site of Kanapoi, where our spot small bones and teeth. Studies of one complete upper jaw and lower face,

the upper and lower thirds of a tibia, bits confident in both the age of the fossils to a changed diet— possibly much hard-
of skull and several sets of isolated teeth. and Brown’s and Feibel’s understanding er food— even though its jaws and some
After careful study of the fossils from of the history of the lake basin. skull features were still very apelike. We
both Allia Bay and Kanapoi— including A major question in paleoanthro- also know that anamensis had only a
Patterson’s fragment of an arm bone— pology today is how the anatomical mo- tiny external ear canal. In this regard, it
we felt that in details of anatomy, these saic of the early hominids evolved. By is more like chimpanzees and unlike all
specimens were different enough from comparing the nearly contemporaneous later hominids, including humans,
previously known hominids to warrant Allia Bay and Kanapoi collections of which have large external ear canals.
designating a new species. So in 1995, in anamensis, we can piece together a fair- (The size of the external canal is unre-
collaboration with both Feibel and Ian ly accurate picture of certain aspects of lated to the size of the fleshy ear.)
McDougall of the Australian National the species, even though we have not yet The most informative bone of all the
University, we named this new species uncovered a complete skull. ones we have uncovered from this new
Australopithecus anamensis, drawing on The jaws of anamensis are primi- hominid is the nearly complete tibia—the
the Turkana word for “lake” (anam) to tive— the sides sit close together and par- larger of the two bones in the lower leg.
refer to both the present and ancient lakes. allel to each other (as in modern apes), The tibia is revealing because of its im-
To establish the age of these fossils, rather than widening at the back of the portant role in weight bearing: the tibia
we relied on the extensive efforts of mouth (as in later hominids, including of a biped is distinctly different from the
Brown, Feibel and McDougall, who have humans). In its lower jaw, anamensis is tibia of an animal that walks on all four
been investigating the paleogeographic also chimplike in terms of the shape of legs. In size and practically all details of
history of the entire lake basin. If their the region where the left and right sides the knee and ankle joints, the tibia found
study of the basin’s development is cor- of the jaw meet (technically known as at Kanapoi closely resembles the one
rect, the anamensis fossils should be be- the mandibular symphysis). from the fully bipedal afarensis found at
tween 4.2 and 3.9 million years old. Mc- Teeth from anamensis, however, ap- Hadar, even though the latter specimen
Dougall has determined the age of the pear more advanced. The enamel is rel- is almost a million years younger.
so-called Kanapoi Tuff—the layer of vol- atively thick, as it is in all other species Fossils of other animals collected at
canic ash that covers most of the fossils of Australopithecus; in contrast, the Kanapoi point to a somewhat different
at this site— to be just over four million tooth enamel of African great apes is paleoecological scenario from the setting
years old. Now that he has successfully much thinner. The thickened enamel across the lake at Allia Bay. The chan-
ascertained the age of the tuff, we are suggests anamensis had already adapted nels of the river that laid down the sedi-
ments at Kanapoi were probably lined
3.9 MILLION 4.2 MILLION with narrow stretches of forest that grew
YEARS AGO YEARS AGO OMO RIVER close to the riverbanks in otherwise open
OMO RIVER
country. Researchers have recovered the
remains of the same spiral-horned ante-
lope found at Allia Bay that very likely
MODERN lived in dense thickets. But open-coun-
LAKE TURKANA
try antelopes and hartebeest appear to
have lived at Kanapoi as well, suggesting
that more open savanna prevailed away
ALLIA BAY from the rivers. These results offer equi-
MODERN vocal evidence regarding the preferred
SLIM FILMS; SOURCE: FRANK BROWN AND CRAIG FEIBEL (1991)
LAKE TURKANA
habitat of anamensis: we know that
bushland was present at both sites that
have yielded fossils of the species, but
LAKE there are clear signs of more diverse
LONYUMUN habitats at Kanapoi.
KANAPOI An Even Older Hominid?

A T A B O U T T H E S A M E T I M E that we
were finding new hominids at Allia Bay
KERIO RIVER
and Kanapoi, a team led by our colleague
Tim D. White of the University of Cali-
TURKANA BASIN was home to anamensis roughly four million years ago. Around 3.9 million years ago a
river sprawled across the basin (left). The fossil site Allia Bay sat within the strip of forest (green) fornia at Berkeley discovered fossil hom-
that lined this river. Some 4.2 million years ago a large lake filled the basin (right); a second site, inids in Ethiopia that are even older than
Kanapoi, was located on a river delta that fed into the lake. anamensis. In 1992 and 1993 White led

an expedition to the Middle Awash area en the human fossil record back close to
of Ethiopia, where his team uncovered the time of the chimp-human split. More
hominid fossils at a site known as Ara- recently, White and his group have found
mis. The group’s finds include isolated parts of a single Ardipithecus skeleton in
teeth, a piece of a baby’s mandible (the the Middle Awash region. As White and
lower jaw), fragments from an adult’s his team extract these exciting new fos-
skull and some arm bones, all of which sils from the enclosing stone, reconstruct
have been dated to around 4.4 million them and prepare them for study, the pa-
years ago. In 1994, together with his col- leoanthropological community eagerly
leagues Berhane Asfaw of the Paleoan- anticipates the publication of the group’s
thropology Laboratory in Addis Ababa analysis of these astonishing finds.
and Gen Suwa of the University of To- But even pending White’s results,
kyo, White gave these fossils a new name: new fossil discoveries are offering other
Australopithecus ramidus. In 1995 the surprises. A team led by Michel Brunet of
group renamed the fossils, moving them the University of Poitiers has found frag-
to a new genus, Ardipithecus. Earlier fos- ments of Australopithecus fossils in
sils of this genus have now been found Chad. Surprisingly, these fossils were re-
dating back to 5.8 million years ago. covered far from either eastern or south-
Other fossils buried near the hominids, ern Africa, the only areas where Aus- FOSSIL HUNTER Alan Walker ( foreground) and
such as seeds and the bones of forest tralopithecus had appeared. The Chad two colleagues excavate the bone bed at Allia
Bay, where several anamensis fossils have been
monkeys and antelopes, strongly imply sites lie 2,500 kilometers west of the
recovered. The bone bed appears as a dark band
that these hominids, too, lived in a western part of the Rift Valley, thus ex- about 18 inches thick at the top of the trench.
closed-canopy woodland. tending the range of Australopithecus
This new species represents the most well into the center of Africa. The significance of these exciting dis-
primitive hominid known— a link be- These fossils debunk a hypothesis coveries is now the center of an active
tween the African apes and Australo- about human evolution postulated by debate.
pithecus. Many of the Ardipithecus ram- Dutch primatologist Adriaan Kortlandt The fossils of anamensis that we have
idus fossils display similarities to the and expounded in Scientific American by identified should also provide some an-
anatomy of the modern African great Yves Coppens of the College of France swers in the long-standing debate over
apes, such as thin dental enamel and [see “East Side Story: The Origin of Hu- whether early Australopithecus species
strongly built arm bones. In other fea- mankind,” May 1994]. This idea was lived in wooded areas or on the open sa-
tures, though— such as the opening at that the formation of Africa’s Rift Valley vanna. The outcome of this discussion
the base of the skull, technically known subdivided a single ancient species, iso- has important implications: for many
as the foramen magnum, through which lating the ancestors of hominids on the years, paleoanthropologists have accept-
the spinal cord connects to the brain— east side from the ancestors of modern ed that upright-walking behavior origi-
the fossils resemble later hominids. apes on the west side. nated on the savanna, where it most like-
Describing early hominids as either Brunet’s latest discovery, an impor- ly provided benefits such as keeping the
primitive or more advanced is a complex tant cranium older than six million years, hot sun off the back or freeing hands for
issue. Scientists now have almost deci- is also from Chad and shows that early carrying food. Yet our evidence suggests
sive molecular evidence that humans hominids were probably present across that the earliest bipedal hominid known
and chimpanzees once had a common much of the continent. This cranium, to date lived at least part of the time in
ancestor and that this lineage had previ- which the team called Sahelanthropus wooded areas. The discoveries of the
KENNETH GARRETT National Geographic Image Collection
ously split from gorillas. This is why we tchadensis, together with fragmentary past several years represent a remarkable
often use the two living species of chim- jaws and limb bones from about six mil- spurt in the sometimes painfully slow
panzee (Pan troglodytes and P. panis- lion years ago in Kenya [see “An Ances- process of uncovering human evolution-
cus) to illustrate ancestral traits. But we tor to Call Our Own,” on page 4], are ary past. But clearly there is still much
must remember that since their last even older than the Ardipithecus fossils. more to learn.
common ancestor with humans, chim-
panzees have had exactly the same MORE TO E XPLORE
amount of time to evolve as humans Australopithecus ramidus, a New Species of Early Hominid from Aramis, Ethiopia. Tim D. White,
Gen Suwa and Berhane Asfaw in Nature, Vol. 371, pages 306–312; September 22, 1994.
have. Determining which features were
New Four-Million-Year-Old Hominid Species from Kanapoi and Allia Bay, Kenya. Meave G. Leakey,
present in the last common ancestor of Craig S. Feibel, Ian McDougall and Alan Walker in Nature, Vol. 376, pages 565–571; August 17, 1995.
humans and chimpanzees is not easy. From Lucy to Language. Donald C. Johanson and Blake Edgar. Simon & Schuster, 1996.
But Ardipithecus, with its numerous The Earliest Known Australopithecus, A. anamensis. C. V. Ward, M. G. Leakey and A. Walker in
chimplike features, appears to have tak- Journal of Human Evolution, Vol. 41, pages 255–368; 2001.

EMERGENCE
TODAY WE TAKE FOR GRANTED THAT HOMO SAPIENS

FOUR MILLION YEARS MANY HOMINID SPECIES
ONCE we
SHARING A SINGLE LANDSCAPE, four kinds of hominids lived about 1.8 million years ago in what is now part of northern Kenya.
Although paleoanthropologists have no idea how— or if— these different species interacted, they do know that Paranthropus boisei,
Homo rudolfensis, H. habilis and H. ergaster foraged in the same area around Lake Turkana.
20 SCIENTIFIC AMERICAN Updated from the January 2000 issue

IS THE ONLY HOMINID ON EARTH. YET FOR AT LEAST
SHARED THE PLANET. WHAT MAKES US DIFFERENT?
were not alone
By Ian Tattersall • Paintings by Jay H. Matternes

Homo sapiens has had the earth to itself
for the past 25,000 years or so, free and years ago in what is now northern Kenya, tinct geographical regions offer some in-
clear of competition from other mem- that the single-species hypothesis was triguing insights.
bers of the hominid family. This period abandoned. Yet even then, paleoanthro-
has evidently been long enough for us to pologists continued to cleave to a rather A Suite of Species
have developed a profound feeling that minimalist interpretation of the fossil FROM THE BEGINNING, almost from
being alone in the world is an entirely record. Their tendency was to downplay the very moment the earliest hominid
natural and appropriate state of affairs. the number of species and to group to- biped— the first “australopith”— made
So natural and appropriate, indeed, gether distinctively different fossils un- its initial hesitant steps away from the
that during the 1950s and 1960s a der single, uninformative epithets such forest depths, we have evidence for hom-
school of thought emerged that claimed, as “archaic Homo sapiens.” As a result, inid diversity. The oldest-known poten-
in essence, that only one species of hom- they tended to lose sight of the fact that tial hominid is Sahelanthropus tchaden-
inid could have existed at a time because many kinds of hominids had regularly sis, represented by a cranium from the
there was simply no ecological space on contrived to coexist. central-western country of Chad [see il-
the planet for more than one culture- Although the minimalist tendency lustration on page 26]. Better known is
bearing species. The “single-species hy- persists, recent discoveries and fossil Australopithecus anamensis, from sites
pothesis” was never very convincing— reappraisals make clear that the biolog- in northern Kenya that are about 4.2
even in terms of the rather sparse homi- ical history of hominids resembles that million years old.
nid fossil record of 40 years ago. But the of most other successful animal families. A. anamensis looks reassuringly simi-
implicit scenario of the slow, single- It is marked by diversity rather than by lar to the 3.8- to 3.0-million-year-old
minded transformation of the bent and linear progression. Despite this rich his- Australopithecus afarensis, a small-
benighted ancestral hominid into the tory— during which hominid species de- brained, big-faced bipedal species to
graceful and gifted modern H. sapiens veloped and lived together and compet- which the famous “Lucy” belonged.
proved powerfully seductive— as fables ed and rose and fell— H. sapiens ulti- Many remnants of A. afarensis have
of frogs becoming princes always are. mately emerged as the sole hominid. The been found in various eastern African
So seductive that it was only in the reasons for this are generally unknow- sites, but some researchers have suggest-
late 1970s, following the discovery of inable, but different interactions between ed that the mass of fossils described as A.
controvertible fossil evidence that hom- the last coexisting hominids— H. sapiens afarensis may contain more than one
inid species coexisted some 1.8 million and H. neanderthalensis— in two dis- species, and it is only a matter of time
PARANTHROPUS BOISEI HOMO RUDOLFENSIS

had massive jaws, was a relatively
equipped with huge large-brained
grinding teeth for a hominid, typified by
presumed vegetarian the famous KNM-ER
diet. Its skull is 1470 cranium. Its
accordingly strongly skull was distinct
built, but it is not from the apparently
known if in body size it smaller-brained H.
was significantly larger habilis, but its body
than the “gracile” proportions are
australopiths. effectively unknown.

until the subject is raised again. In any tinctive P. robustus and possibly a close- could wish for a better record of this
event, A. afarensis was not alone in ly related second species, P. crassidens. movement, and particularly of its dat-
Africa. A distinctive jaw, from an aus- I apologize for inflicting this long list ing, but there are indications that hom-
tralopith named A. bahrelghazali, was of names on readers, but in fact it actu- inids of some kind had reached China
found in 1995 in Chad. It is probably ally underestimates the number of aus- and Java by about 1.8 million years ago.
between 3.5 and 3.0 million years old tralopith species that existed. What is A lower jaw that may be about the same
and is thus roughly coeval with Lucy, as more, scientists don’t know how long age from Dmanisi in ex-Soviet Georgia
is the recently named new form Kenyan- each of these creatures lasted. Neverthe- is different from anything else yet found
thropus platyops. less, even if average species longevity [see “Out of Africa Again ... and Again?”
In southern Africa, scientists reported was only a few hundred thousand years, by Ian Tattersall, on page 38]. By the
evidence in 1999 of another primitive it is clear that from the very beginning million-year mark H. erectus was estab-
bipedal hominid species. As yet un- the continent of Africa was at least pe- lished in both Java and China, and it is
named and undescribed, this distinctive riodically— and most likely continual- possible that a more robust hominid spe-
form is 3.3 million years old. At about ly— host to multiple kinds of hominids. cies was present in Java as well. At the
three million years ago, the same region The appearance of the genus Homo other end of the Eurasian continent, the
begins to yield fossils of A. africanus, the did nothing to perturb this pattern. The oldest-known European hominid frag-
first australopith to be discovered (in 2.5- to 1.8-million-year-old fossils from ments—from about 800,000 years ago—
1924). This species may have persisted eastern and southern Africa that an- are highly distinctive and have been
until not much more than two million nounce the earliest appearance of Homo dubbed H. antecessor by their Spanish
years ago. A 2.5-million-year-old species are an oddly assorted lot and probably a discoverers.
from Ethiopia, named Australopithecus lot more diverse than their conventional About 600,000 years ago, in Africa,
garhi in 1999, is claimed to fall in an in- assignment to the two species H. habilis we begin to pick up evidence for H. hei-
termediate position between A. afaren- and H. rudolfensis indicates. Still, at delbergensis, a species also seen at sites
sis, on the one hand, and a larger group Kenya’s East Turkana, in the period be- in Europe— and possibly China— be-
that includes more recent australopiths tween 1.9 and 1.8 million years ago, tween 500,000 to 200,000 years ago. As
and Homo, on the other. Almost exact- these two species were joined not only we learn more about H. heidelbergensis,
ly the same age is the first representative by the ubiquitous P. boisei but by H. er- we are likely to find that more than one
of the “robust” group of australopiths, gaster, the first hominid of essentially species is actually represented in this
Paranthropus aethiopicus. This early modern body form. Here, then, is evi- group of fossils. In Europe, H. heidel-
form is best known from the 2.5-mil- dence for four hominid species sharing bergensis or a relative gave rise to an en-
lion-year-old “Black Skull” of northern not just the same continent but the same demic group of hominids whose best-
Kenya, and in the period between about landscape [see illustration on opposite known representative was H. nean-
2 and 1.4 million years ago the robusts page and below]. derthalensis, a European and western
were represented all over eastern Africa The first exodus of hominids from Asian species that flourished between
by the familiar P. boisei. In South Africa, Africa, presumably in the form of H. er- about 200,000 and 30,000 years ago.
during the period around 1.6 million gaster or a close relative, opened a vast The sparse record from Africa suggests
years ago, the robusts included the dis- prospect for further diversification. One that at this time independent develop-
HOMO HABILIS HOMO ERGASTER,

(“handy man”) was sometimes called “African
so named because it H. erectus,” had a high,
was thought to be the rounded cranium and a
maker of the 1.8- skeleton broadly similar
million-year-old to that of modern
stone tools humans. Although H.
discovered at Olduvai ergaster clearly ate meat,
Gorge in Tanzania. its chewing teeth are
This hominid relatively small. The best
fashioned sharp specimen of this hominid
flakes by banging is that of an adolescent
one rock cobble from about 1.6 million
against another. years ago known as
Turkana boy.

TUC D’AUDOUBERT CAVE in France was entered sometime between perhaps
11,000 and 13,000 years ago by H. sapiens, also called Cro Magnons, who
sculpted small clay bison in a recess almost a mile underground.
ments were taking place there, too— in- involving the cop-out of stuffing all vari- than gradual accretions. Over the past
cluding the emergence of H. sapiens. ants of Homo of the past half a million five million years, new hominid species
And in Java, possible H. erectus fossils or even two million years into the species have regularly emerged, competed, co-
from Ngandong were dated to around H. sapiens. existed, colonized new environments
40,000 years ago, implying that this area My own view, in contrast, is that the and succeeded— or failed. We have only
had its own indigenous hominid evolu- 20 or so hominid species invoked (if not the dimmest of perceptions of how this
tionary history for perhaps millions of named) above represent a minimum es- dramatic history of innovation and in-
years as well. timate. Not only is the human fossil teraction unfolded, but it is already evi-
The picture of hominid evolution just record as we know it full of largely un- dent that our species, far from being the
sketched is a far cry from the “Australo- acknowledged morphological indica- pinnacle of the hominid evolutionary
pithecus africanus begat Homo erectus tions of diversity, but it would be rash to tree, is simply one more of its many ter-
begat Homo sapiens” scenario that pre- claim that every hominid species that minal twigs.
vailed 40 years ago— and it is, of course, ever existed is represented in one fossil
based to a great extent on fossils that collection or another. And even if only The Roots of Our Solitude
have been discovered since that time. the latter is true, it is still clear that the A L T H O U G H T H I S is all true, H. sapi-
Yet the dead hand of linear thinking still story of human evolution has not been ens embodies something that is undeni-
lies heavily on paleoanthropology, and one of a lone hero’s linear struggle. ably unusual and is neatly captured by
even today quite a few of my colleagues Instead it has been the story of na- the fact that we are alone in the world
would argue that this scenario overesti- ture’s tinkering: of repeated evolution- today. Whatever that something is, it is
mates diversity. There are various ways ary experiments. Our biological history related to how we interact with the ex-
of simplifying the picture, most of them has been one of sporadic events rather ternal world: it is behavioral, which

HOMINIDS of modern body form most
likely emerged in Africa around 150,000
years ago and coexisted with other
hominids for a time before emerging as
the only species of our family. Until
about 30,000 years ago, they overlapped
with H. neanderthalensis (left) in Europe
and in the Levant, and they may have
been contemporaneous with the
H. erectus (right) then living in Java.
means that we have to look to our ar- who did the same, the Neandertals fur- lack the “grave goods” that would attest
chaeological record to find evidence of nish us with a particularly instructive to ritual and belief in an afterlife. The
it. This record begins some 2.5 million yardstick by which to judge our own Neandertals, in other words, though ad-
years ago with the production of the first uniqueness. The stoneworking skills of mirable in many ways and for a long
recognizable stone tools: simple sharp the Neandertals were impressive, if time successful in the difficult circum-
flakes chipped from parent “cores.” We somewhat stereotyped, but they rarely if stances of the late ice ages, lacked the
don’t know exactly who the inventor ever made tools from other preservable spark of creativity that, in the end, dis-
was, but chances are that he or she was materials. And many archaeologists tinguished H. sapiens.
something we might call an australopith. question the sophistication of their hunt- Although the source of H. sapiens as
This landmark innovation representing skills. a physical entity is obscure, most evi-
ed a major cognitive leap and had pro- Further, despite misleading early ac- dence points to an African origin perhaps
found long-term consequences for hom- counts of bizarre Neandertal “bear between 150,000 and 200,000 years
inids. It also inaugurated a pattern of cults” and other rituals, no substantial ago. Modern behavior patterns did not
highly intermittent technological change. evidence has been found for symbolic emerge until much later. The best evi-
It was a full million years before the next behaviors among these hominids or for dence comes from Israel and its sur-
significant technological innovation the production of symbolic objects— cer- rounding environs, where Neandertals
came along: the creation about 1.5 mil- tainly not before contact had been made lived about 200,000 years ago or per-
lion years ago, probably by H. ergaster, with modern humans. Even the occa- haps even earlier. By about 100,000
of the hand ax. These symmetrical im- sional Neandertal practice of burying years ago, they had been joined by
plements, shaped from large stone cores, the dead may have been simply a way of anatomically modern H. sapiens, and
were the first tools to conform to a “men- discouraging hyenas from making in- the remarkable thing is that the tools
tal template” that existed in the tool- cursions into their living spaces or have and sites the two hominid species left be-
maker’s mind. This template remained a similar mundane explanation. This hind are essentially identical. As far as
essentially unchanged for another mil- view arises because Neandertal burials can be told, these two hominids behaved
lion years or more, until the invention of
“prepared-core” tools by H. heidelber- IAN TATTERSALL and JAY H. MATTERNES have worked together since the early 1990s, no-
THE AUTHOR AND THE ARTIST
gensis or a relative. Here a stone core was tably on the Hall of Human Biology and Evolution at the American Museum of Natural His-
elaborately shaped in such a way that a tory in New York City and at the Gunma Museum of Natural History in Tomioka, Japan (where
single blow would detach what was an the Tuc d’Audoubert mural on the opposite page is installed). Tattersall was born in England
effectively finished implement. and raised in East Africa. He is a curator in the department of anthropology at the Ameri-
Among the most accomplished practi- can Museum of Natural History. His books include Becoming Human: Evolution and Human
tioners of prepared-core technology Uniqueness (Harvest Books, 1999) and The Last Neanderthal: The Rise, Success, and Mys-
were the large-brained, big-faced and terious Extinction of Our Closest Human Relatives (Westview Press, 1999, revised).
low-skulled Neandertals, who occupied Matternes is an artist and sculptor who has for more than 40 years specialized in fos-
Europe and western Asia until about sil primates and hominids. In addition to his museum murals, he is well known for his illus-
30,000 years ago. Because they left an trations for books, periodicals and television, including Time/Life Books and National Geo-
excellent record of themselves and were graphic. The research for his depictions has taken him to many parts of the U.S., Canada,
abruptly replaced by modern humans Mexico, France, Colombia and Africa.

0 H. sapiens (Worldwide)
H. neanderthalensis
(Europe and Western Asia) in similar ways despite their anatomical
differences. And as long as they did so,
they somehow contrived to share the
H. heidelbergensis (throughout Old World)
Levantine environment.
The situation in Europe could hardly
H. erectus (Eastern Asia) be more different. The earliest H. sapi-
1 H. antecessor
(Spain) ens sites there date from only about
40,000 years ago, and just 10,000 or so
years later the formerly ubiquitous Ne-
andertals were gone. Significantly, the
H. sapiens who invaded Europe brought
H. habilis with them abundant evidence of a fully
(Sub-Saharan Africa) formed and unprecedented modern sen-
K. rudolfensis
(Eastern Africa) sibility. Not only did they possess a new
“Upper Paleolithic” stoneworking tech-
2
Homo ergaster P. robustus P. boisei nology based on the production of mul-
(Eastern Africa) (South Africa) (Eastern Africa) tiple long, thin blades from cylindrical
cores, but they made tools from bone
Au. africanus and antler, with an exquisite sensitivity
(South Africa) Paranthropus

aethiopicus to the properties of these materials.
Au. garhi (Eastern Africa) Even more significant, they brought
(Ethiopia) with them art, in the form of carvings,
engravings and spectacular cave paint-
3 ings; they kept records on bone and
stone plaques; they made music on wind
instruments; they crafted intricate per-
Au. bahrelghazali sonal adornments; they afforded some
(Chad) of their dead elaborate burials with
grave goods (hinting at social stratifica-
tion in addition to belief in an afterlife,
Kenyanthropus Au. afarensis for not all burials were equally fancy);
platyops (Ethiopia and Australopithecus
4 (Kenya) anamensis
and their living sites were highly orga-
Tanzania)
(Kenya) nized, with evidence of sophisticated
hunting and fishing. The pattern of in-
termittent technological innovation was
gone, replaced by constant refinement.
Clearly, these people were us.
Competing Scenarios
Ardipithecus ramidus IN ALL THESE WAYS, early Upper Pa-
5 (Ethiopia)
leolithic people contrasted dramatically
with the Neandertals. Some Neandertals
in Europe seem to have picked up new
ways of doing things from the arriving
H. sapiens, but we have no direct clues
as to the nature of the interaction be-
Orrorin tugenensis tween the two species. In light of the Ne-
(Kenya)
andertals’ rapid disappearance and of
6
PATRICIA J. WYNNE (drawings)
the appalling subsequent record of H.

sapiens, though, we can reasonably sur-
SPECULATIVE FAMILY TREE shows the variety of hominid mise that such interactions were rarely
species that have populated the planet— some identified by happy for the former. Certainly the re-
only a fragment, others known to exist for a specific time
period (solid lines). The emergence of H. sapiens has not peated pattern found at archaeological
been a single, linear transformation of one species into Sahelanthropus sites is one of short-term replacement,
another but rather a meandering, multifaceted evolution. tchadensis and there is no convincing biological ev-
(Chad)

The pattern of intermittent technological innovation
was gone, replaced by constant refinement.
Clearly, these people were us.
idence of any intermixing of peoples in see that, profound as the consequences sufficiently advantageous, this behav-
Europe. of achieving symbolic thought may have ioral novelty could then have spread
In the Levant, the coexistence ceased— been, the process whereby it came about rapidly by cultural contact among pop-
after about 60,000 years or so— at right was unexceptional. ulations that already had the potential to
about the time that Upper Paleolithic– We have no idea at present how the acquire it. No population replacement
like tools began to appear. About 40,000 modern human brain converts a mass of would have been necessary to spread the
years ago the Neandertals of the Levant electrical and chemical discharges into capability worldwide.
yielded to a presumably culturally rich what we experience as consciousness. It is impossible to be sure what this in-
H. sapiens, just as their European coun- We do know, however, that somehow novation might have been, but the best
terparts had. our lineage passed to symbolic thought current bet is that it was the invention of
The key to the difference between the from some nonsymbolic precursor state. language. For language is not simply the
European and the Levantine scenarios The only plausible possibility is that medium by which we express our ideas
lies, most probably, in the emergence of with the arrival of anatomically modern and experiences to one another. Rather
modern cognition— which, it is reason- H. sapiens, existing exaptations were it is fundamental to the thought process
able to assume, is equivalent to the ad- fortuitously linked by a relatively minor itself. It involves categorizing and nam-
vent of symbolic thought. Business had genetic innovation to create an unprece- ing objects and sensations in the outer
continued more or less as usual right dented potential. and inner worlds and making associa-
through the appearance of modern bone Yet even in principle this deduced sce- tions between resulting mental symbols.
structure, and only later, with the ac- nario cannot be the full story, because It is, in effect, impossible for us to con-
quisition of fully modern behavior pat- anatomically modern humans behaved ceive of thought (as we are familiar with
terns, did H. sapiens become complete- archaically for a long time before adopt- it) in the absence of language, and it is
ly intolerant of competition from its ing modern behaviors. That discrepan- the ability to form mental symbols that
nearest— and, evidently, not its dearest— cy may be the result of the late appear- is the fount of our creativity. Only when
co-inhabitors. ance of some key hardwired innovation we are able to create such symbols can
To understand how this change in sen- not reflected in the skeleton, which is all we recombine them and ask such ques-
sibility occurred, we have to recall cer- that fossilizes. But this seems unlikely, tions as “What if...?”
tain things about the evolutionary pro- because it would have necessitated a We do not know exactly how lan-
cess. First, as in this case, all innovations wholesale Old World–wide replacement guage might have emerged in one local
must necessarily arise within preexisting of hominid populations in a very short population of H. sapiens, although lin-
species— for where else can they do so? time, something for which there is no guists have speculated widely. But we do
Second, many novelties arise as “exap- evidence. know that a creature armed with sym-
tations,” features acquired in one con- It is much more likely that the modern bolic skills is a formidable competitor—
text before (often long before) being co- human capacity was born at— or close and not necessarily an entirely rational
opted in a different one. For example, to— the origin of H. sapiens, as an abili- one, as the rest of the living world, in-
hominids possessed essentially modern ty that lay fallow until it was activated cluding H. neanderthalensis, has discov-
vocal tracts for hundreds of thousands by a cultural stimulus of some kind. If ered to its cost.
of years before the behavioral record
gives us any reason to believe that they MORE TO E XPLORE
employed the articulate speech that the Dark Caves, Bright Visions: Life in Ice Age Europe. Randall White. W. W. Norton/American Museum
of Natural History, 1986.
peculiar form of this tract permits.
Language and Species. Reprint edition. Derek Bickerton. University of Chicago Press, 1992.
And finally, it is important to bear in
The Fossil Trail: How We Know What We Think We Know about Human Evolution. Ian Tattersall.
mind the phenomenon of emergence— Oxford University Press, 1995.
the notion that a chance coincidence Getting Here: The Story of Human Evolution. Updated edition. William Howells. Compass Press,
gives rise to something totally unexpect- 1997.
ed. The classic scientific example in this African Exodus: The Origins of Modern Humanity. Reprint edition. Christopher Stringer and
Robin McKie. Henry Holt, 1998.
regard is water, whose properties are
The Origin and Diversification of Language. Edited by Nina G. Jablonski and Leslie C. Aiello.
wholly unpredicted by those of hydro- University of California Press, 1998.
gen and oxygen atoms alone. If we com- The Last Neanderthal: The Rise, Success and Mysterious Extinction of Our Closest Human
bine these various observations, we can Relatives. Revised edition. Ian Tattersall. Westview Press, 1999.

EMERGENCE
Who Were the

NEANDERTALS?
28 SCIENTIFIC AMERICAN Updated from the April 2000 issue
Controversial evidence indicates that these
hominids interbred with anatomically modern humans
and sometimes behaved in surprisingly modern ways
By Kate Wong
It was such a neat and tidy story.

No match for the anatomically modern humans who swept in with a sophisticated cul-
ture and technology, the Neandertals— a separate species— were quickly driven to ex-
tinction by the invading moderns. But neat and tidy stories about the past have a way
of unraveling, and the saga of the Neandertals, it appears, is no exception. For more
than 200,000 years, these large-brained hominids occupied Europe and western Asia,
battling the bitter cold of glacial maximums and the daily perils of prehistoric life. To-
day they no longer exist. Beyond these two facts, however, researchers fiercely debate
who the Neandertals were, how they lived and exactly what happened to them.
REFLECTION OF THE PAST The steadfast effort to resolve these elusive issues stems from a larger dispute over how
reveals a face that is modern humans evolved. Some researchers posit that our species arose recently (around
at once familiar and 200,000 years ago) in Africa and subsequently replaced archaic hominids around the world,
foreign. The 130,000- whereas others propose that these ancient populations contributed to the early modern
year-old skull of an adult
human gene pool. As the best known of these archaic groups, Neandertals are critical to
female from the Krapina
rock-shelter in the origins controversy. Yet this is more than an academic argument over certain events
northwestern Croatia of our primeval past, for in probing Neandertal biology and behavior, researchers must
inspired this Neandertal wrestle with the very notion of what it means to be fully human and determine what, if
reconstruction. anything, makes us moderns unique. Indeed, spurred by recent discoveries, paleoan-
thropologists and archaeologists are increasingly asking, How much like us were they?
Comparisons of Neandertals and modern humans first captured the attention of re-
searchers when a partial Neandertal skeleton turned up in Germany’s Neander Valley
in 1856. Those remains— a heavily built skull with the signature arched browridge and
massive limb bones— were clearly different, and Neandertals were assigned to their own
species, Homo neanderthalensis (although even then there was disagreement: several Ger-
man scientists argued that these were the remains of a crippled Cossack horseman). But
it was the French discovery of the famous “Old Man” of La Chapelle-aux-Saints some
50 years later that led to the characterization of Neandertals as primitive protohumans.
Reconstructions showed them as stooped, lumbering, apelike brutes, in stark contrast to
upright, graceful Homo sapiens. The Neandertal, it seemed, represented the ultimate
“other,” a dim-witted ogre lurking behind the evolutionary threshold of humanity.
Decades later reevaluation of the La Chapelle individual revealed that certain anatom-
ical features had been misinterpreted. In fact, Neandertal posture and movement would
CROATIAN NATURAL HISTORY MUSEUM
have been the same as ours. Since then, paleoanthropologists have struggled to determine
whether the morphological features that do characterize Neandertals as a group— such
as the robustness of their skeletons, their short limbs and barrel chests, prominent
browridges and low, sloping foreheads, protruding midfaces and chinless jaws—warrant
designating them as a separate species. Researchers agree that some of these characteris-
tics represent environmental adaptations. The Neandertals’ stocky body proportions, for
example, would have allowed them to retain heat more effectively in the extremely cold
weather brought on by glacial cycles. But other traits, such as the form of the Neander-

NEANDERTAL
LARGER RECEDING a fossil from a site in southwestern Germany called
OCCIPITAL FOREHEAD
Vogelherd, which combines the skull shape of mod-
BUN
erns with features that are typically Neandertal,
STRONG
BROWRIDGE such as the distinct space between the last molar
and the ascending part of the lower jaw known as
a retromolar gap, and the form of the mandibular
foramen— a nerve canal in the lower jaw. Addi-
tional evidence, according to Frayer and Milford H.
PROJECTING
MIDFACE
Wolpoff of the University of Michigan at Ann Ar-
bor, comes from a group of early moderns discov-
ered in Moravia (Czech Republic) at a site called
Mladeč. The Mladeč people, they say, exhibit char-
acteristics on their skulls that other scientists have
described as uniquely Neandertal traits.
RETROMOLAR GAP
NO CHIN
Although such evidence was once used to argue
that Neandertals could have independently evolved
into modern Europeans, this view has shifted some-
EARLY MODERN EUROPEAN what. “It’s quite clear that people entered Europe as
well, so the people that are there later in time are a
SMALLER mix of Neandertals and those populations coming
OCCIPITAL STEEP into Europe,” says Wolpoff, who believes the two
BUN FOREHEAD groups differed only as much as living Europeans
DELICATE and aboriginal Australians do. Evidence for mixing
BROWRIDGE also appears in later Neandertal fossils, according to
Fred H. Smith, a paleoanthropologist at Loyola Uni-
versity of Chicago. Neandertal remains from Vin-
dija cave in northwestern Croatia reflect “the as-
similation of some early modern features,” he says,
VERTICAL
MIDFACE referring to their more modern-shaped browridges
and the slight presence of a chin on their mandibles.
Those who view Neandertals as a separate spe-
cies, however, maintain that the Vindija fossils are
NO RETROMOLAR GAP; too fragmentary to be diagnostic and that any sim-
LAST LOWER MOLAR
MISSING ON THIS CHIN ilarities that do exist can be attributed to conver-
SPECIMEN gent evolution. These researchers likewise dismiss
the mixing argument for the early moderns from
Mladeč. “When I look at the morphology of these
ˇ (bottom)
CHARACTERISTIC DIFFERENCES are shown between a Neandertal, represented by a French people, I see robustness, I don’t see Neandertal,”
specimen, La Ferrassie 1, and an early modern, Dolní Věstonice 16, from the Czech counters Christopher B. Stringer of the Natural
Republic. Each aspect can be found in both groups, varying in degree and frequency,
ERIK TRINKAUS/MUSÉE DE L’HOMME (top); ARCHEOLOGICK Ý ÚSTAV AV CR

but they tend to appear as suites of features.
History Museum in London.
Another reason to doubt these claims for in-
tal browridge, lack any clear functional significance and seem terbreeding, some scientists say, is that they contradict the con-
to reflect the genetic drift typical of isolated populations. clusions reached by Svante Pääbo, now at the Max Planck In-
For those scholars who subscribe to the replacement mod- stitute for Evolutionary Anthropology in Leipzig, Germany,
el of modern human origins, the distinctive Neandertal mor- and his colleagues, who in July 1997 announced that they had
phology resulted from following an evolutionary trajectory sep- retrieved and analyzed mitochondrial DNA (mtDNA) from a
arate from that of moderns. But for years, another faction of re- Neandertal fossil. The cover of the journal Cell, which con-
searchers has challenged this interpretation, arguing that many tained their report, was unequivocal: “Neandertals Were Not
of the features that characterize Neandertals are also seen in the Our Ancestors.” From the short stretch of mtDNA they se-
early modern Europeans that followed them. “They clearly have quenced, the researchers determined that the difference be-
a suite of features that are, overall, different, but it’s a frequency tween the Neandertal mtDNA and living moderns’ mtDNA
difference, not an absolute difference,” contends David W. Fray- was considerably greater than the differences found among
er, a paleoanthropologist at the University of Kansas. “Virtually living human populations. But though it seemed on the sur-
everything you can find in Neandertals you can find elsewhere.” face that the species question had been answered, undercur-
He points to one of the earliest-known modern Europeans, rents of doubt have persisted [see “Ancestral Quandary,” by

GUIDE TO TERMINOLOGY
Kate Wong, News and Analysis, January 1998]. Since then, Neandertal can also be spelled Neanderthal. Around 1900
mtDNA from three more specimens has been retrieved and an- German orthography changed, and the silent “h” in certain
alyzed, with similarly inconclusive results. words, such as “thal” (meaning “valley”), was dropped. The
Recent fossil evidence from western Europe has intensified designation Homo neanderthalensis remains the same, but
interest in whether Neandertals and moderns mixed. In Janu- the common name can be spelled either way.
ary 1999 researchers announced the discovery in central Por-
tugal’s Lapedo Valley of a largely complete skeleton from a Paleolithic, or Old Stone Age, is the period ranging from the
four-year-old child buried 24,500 years ago in the Gravettian beginning of culture to the end of the last glaciation. It is
style known from other early modern Europeans. According to subdivided into Lower, Middle and Upper stages.
Erik Trinkaus of Washington University, Cidália Duarte of the Mousterian is a Middle Paleolithic stone tool– based cultural
Portuguese Institute of Archaeology in Lisbon and their col- tradition associated with Neandertals and with early moderns
leagues, the specimen, known as Lagar Velho 1, bears a com- in the Near East.
bination of Neandertal and modern human traits that could
only have resulted from extensive interbreeding between the Aurignacian is an Upper Paleolithic cultural tradition associated
two populations [see “The Hybrid Child from Portugal,” on with moderns that includes advanced tools and art objects.
the next page]. Châtelperronian is an Upper Paleolithic cultural tradition
If the mixed-ancestry interpretation for Lagar Velho 1 associated with Neandertals. It resembles both the
holds up after further scrutiny, the notion of Neandertals as a Mousterian and the Aurignacian.
variant of our species will gain new strength. Advocates of the
DAY IN THE LIFE of Neandertals at the Grotte du Renne in France is along with evidence of huts and hearths, were once linked to modern
imagined here. The Châtelperronian stratigraphic levels have yielded humans alone, but the Grotte du Renne remains suggest that some
a trove of pendants and advanced bone and stone tools. Such items, Neandertals were similarly industrious.
MICHAEL ROTHMAN

THE HYBRID CHILD FROM PORTUGAL BY ERIK TRINKAUS AND CIDÁLIA DUARTE
ON A CHILLY AFTERNOON in late November does have a suite of features that align it had not been previously documented for
1998, while inspecting the Abrigo do Lagar predominantly with early modern western Europe. We therefore conclude
Velho rock-shelter in central Portugal’s Europeans. These include a prominent that Lagar Velho 1 resulted from
Lapedo Valley, two archaeology scouts chin and other details of the mandible interbreeding between indigenous Iberian
spotted loose sediment in a rodent hole (lower jaw), small front teeth, a short face, Neandertals and early modern humans
along the shelter’s back wall. Knowing that the nose shape, minimal brow dispersing throughout Iberia sometime
burrowing animals often bring deeper development, muscle markings on the after 30,000 years ago. Because the child
materials to the surface, one of the scouts thumb bone, the narrowness of the front of lived several millennia after Neandertals
reached in to see what might have been the pelvis, and several aspects of the are thought to have disappeared, its
unearthed. When he withdrew his hand, he shoulder blade and forearm bones. anatomy probably reflects a true mixing of
held in it something extraordinary: bones Yet intriguingly, a number of features these populations during the period when
of a human child buried nearly 25,000 also suggest certain Neandertal affinities. they coexisted and not a rare chance
years ago. Specifically, the front of the mandible mating between a Neandertal and an early
Subsequent excavation of the burial, slopes backward despite the chin, there is modern human.
led by one of us (Duarte), revealed that a porous depression above the neck Fieldwork conducted in 1999 yielded
the four-year-old had been ceremonially muscles, the pectoral muscles are major pieces of the skull and most of the
interred— covered with red ocher and laid strongly developed, and the lower legs are remaining teeth. An international team
on a bed of burnt vegetation, along with short and stout. Thus, the Lagar Velho then assembled to fully interpret this
pierced deer teeth and a marine shell— in child exhibits a complex mosaic of remarkable specimen. Aside from detailed
the Gravettian style known from modern Neandertal and early modern human comparative analyses of individual
humans of that time across Europe. Based features. portions of the skeleton, all the remains
on the abrupt cultural transition seen in This anatomical amalgam is not the were CT scanned and a virtual, computer-
archaeological remains from the Iberian result of any abnormalities. Taking normal assisted reconstruction of the skull was
Peninsula, it seemed likely that when human growth patterns into undertaken.
moderns moved into the area after 30,000 consideration, our analysis indicates that Such rigorous technological study is
years ago, they rapidly replaced the native except for a bruised forearm, a couple of
Neandertals. So it stood to reason that this lines on the bones indicating times when MORPHOLOGICAL MOSAIC found on this 24,500-
specimen, called Lagar Velho 1, growth was trivially arrested (by sickness year-old skeleton from Portugal indicates that
Neandertals and modern humans are members of the
represented an early modern child. In fact, or lack of food) and the fact that it died as
same species who interbred freely. The child—called
it didn’t occur to us at first that it could be a child, Lagar Velho 1 developed normally. Lagar Velho 1—is modern overall but bears some
anything else. The combination can only have resulted Neandertal traits, such as short lower-limb bones
This wonderfully complete skeleton from a mixed ancestry— something that and a backward-sloping mandible.
replacement model do allow for isolated instances of inter- tugal’s then cold climate. But this interpretation is problemat-
breeding between moderns and the archaic species, because ic, according to Jean-Jacques Hublin of France’s CNRS, who
some other closely related mammal species interbreed on oc- points out that although some cold-adapted moderns exhibit
casion. But unlike central and eastern European specimens that such proportions, none are known from that period in Europe.
are said to show a combination of features, the Portuguese For his part, Hublin is troubled that Lagar Velho 1 represents
child dates to a time when Neandertals are no longer thought a child, noting that “we do not know anything about the vari-
to have existed. For Neandertal features to have persisted thou- ation in children of a given age in this range of time.”
sands of years after those people disappeared, Trinkaus and
Duarte say, coexisting populations of Neandertals and mod- Survival Skills
erns must have mixed significantly. TAXONOMIC ISSUES ASIDE, much research has focused on
Their interpretation has not gone unchallenged. In a com- Neandertal behavior, which remained largely misunderstood
mentary accompanying the team’s report in the Proceedings of until relatively recently. Neandertals were often portrayed as in-
the National Academy of Sciences USA in June 1999, paleoan- capable of hunting or planning ahead, recalls archaeologist
thropologists Ian Tattersall of the American Museum of Nat- John J. Shea of the State University of New York at Stony
ural History in New York City and Jeffrey H. Schwartz of the Brook. “We’ve got reconstructions of Neandertals as people
University of Pittsburgh argued that Lagar Velho 1 is most like- who couldn’t survive a single winter, let alone a quarter of a mil-
ly “a chunky Gravettian child.” The robust body proportions lion years in the worst environments in which humans ever
that Trinkaus and his colleagues view as evidence for Nean- lived,” he observes. Analysis of animal remains from the Croa-
dertal ancestry, Stringer says, might reflect adaptation to Por- tian site of Krapina, however, indicates that Neandertals were

necessary because the discovery of an individual
with such a mosaic of features has profound
implications. First, it rejects the extreme Out of Africa
model of modern human emergence, which proposes
that early moderns originating in Africa
subsequently displaced all archaic humans in other
regions. Instead the Lagar Velho child’s anatomy
supports a scenario that combines a dispersal of
anatomically modern humans out of Africa with
mixing between that population and the archaic
populations it encountered. (For example, the
African ancestry of early modern Europeans is
reflected in their relatively long lower-leg bones, a
tropical adaptation. Lagar Velho 1, however, has the
short shins of the cold-adapted Neandertals.)
Lagar Velho 1 also provides insights into the
behavioral similarities of Neandertals and early
modern humans. Despite the paleontological
evidence indicating anatomical differences between
these two groups, their overall adaptive patterns,
social behaviors and means of communication
(including language) cannot have contrasted
greatly. To their contemporaries, the Neandertals
were just another group of Pleistocene hunter-
gatherers, fully as human as themselves.
ERIK TRINKAUS is a paleoanthropologist

at Washington University.
CIDÁLIA DUARTE is a researcher at the Portuguese

Institute of Archaeology in Lisbon.
Mary C. Stiner and Steven L. Kuhn have shown

that Neandertal subsistence strategies varied wide-
ly with the environment and the changing seasons.
Such demonstrations refute the notion that Ne-
andertals perished because they could not adapt.
But it may be that moderns were better at it. One
popular theory posits that modern humans held
some cognitive advantage over Neandertals, per-
haps a capacity for the most human trait of all:
symbolic thought, including language. Explana-
skilled hunters capable of killing even large animals such as rhi- tions such as this one arose from observations that after 40,000
noceroses, according to University of Cambridge archaeologist years ago, whereas Neandertal culture remained relatively stat-
JOSÉ PAULO B. RUAS/PORTUGUESE
Preston T. Miracle. And Shea’s studies suggest that some Ne- ic, that of modern Europeans boasted a bevy of new features,
INSTITUTE OF ARCHAEOLOGY
andertals employed sophisticated stone-tipped spears to con- many of them symbolic. It appeared that only moderns per-
quer their quarry— a finding supported in 1999, when re- formed elaborate burials, expressed themselves through body
searchers reported the discovery in Syria of a Neandertal-made ornaments, figurines and cave paintings, and crafted complex
stone point lodged in a neckbone of a prehistoric wild ass. bone and antler tools— an industry broadly referred to as Up-
Moreover, additional research conducted by Shea and investi- per Paleolithic. Neandertal assemblages, in contrast, contained
gations carried out by University of Arizona archaeologists only Middle Paleolithic stone tools made in the Mousterian style.

A CASE FOR NEANDERTAL CULTURE BY JOÃO ZILHÃO AND FRANCESCO D’ERRICO
EVER SINCE THE DISCOVERY nearly 150 this to mean that Neandertals picked up Similarly, the new knapping
years ago of the specimen that defined the these ideas from moderns, either techniques and tool types that appear
Neandertals, researchers have tended to collecting or trading for items among late Neandertals at other sites in
deny Neandertals the behavioral manufactured by moderns or imitating the France, Italy and Spain fail to show any
capabilities of modern humans, such as newcomers’ practices without really influence from the Aurignacian. Instead
the use of symbols or of complex grasping the underlying symbolic nature they maintain affinities with the preceding
techniques for tool manufacture. Instead of some of the objects. local traditions, of which they seem to
Neandertals were characterized as Our reassessment of the evidence represent an autonomous development.
subhuman, stuck in primitive technical from the Grotte du Renne shows that the If the Neandertals’ Châtelperronian
traditions impervious to innovation. And Neandertal-associated ornaments and culture was an outcome of contact with
when sophisticated cultural remains were tools found there did not result from a moderns, then the Aurignacian should
linked to late Neandertals at several sites mixing of the strata, as demonstrated by predate the Châtelperronian. Yet our
in western Europe, the evidence was the presence of finished objects and the reanalysis of the radiometric dates for the
explained away. The most spectacular of by-products of their manufacture in the archaeological sequences reveals that
these sites, a cave in north-central France same stratigraphic level. Moreover, the apart from a few debatable instances of
named Grotte du Renne (one in a string of Châtelperronian artifacts recovered at the mixture, wherever both cultures are
sites collectively known as the Arcy-sur- Grotte du Renne and other sites, such as
Cure caves), yielded a wealth of complex Quinçay, in the Poitou-Charentes region of
bone and stone tools, body ornaments and France, were created using techniques
decorated objects, found in association different from those favored by
with Neandertal remains. Other sites in Aurignacians. With regard, for example, to
France and along the Cantabrian and the pendants— modified bear, wolf and
Pyrenean mountain ranges bore similar deer teeth, among others—Neandertals
artifacts made in this tradition, called the carved a furrow around the tooth root so
Châtelperronian. that a string of some sort could be tied
Because early modern Europeans had around it for suspension, whereas
a comparable industry known as Aurignacians pierced their pendants. As
Aurignacian—which often appears at the archaeologist François Lévêque and a
same sites that contain Châtelperronian colleague have described, even when, as
materials—some researchers have they did on occasion, Neandertals put a
suggested that the archaeological layers hole through a tooth, they took an unusual
were disrupted, mixing Aurignacian approach, puncturing the tooth. Moderns
artifacts into the Neandertal-associated preferred to scrape the tooth thin and then
levels. Other scholars have interpreted pierce it.
Yet hints that Neandertals thought symbolically had of moderns. Then, in

popped up. Neandertal burials, for example, are well known 1996, Hublin and his
across Europe, and several, it has been argued, contain grave co-workers made a
goods. (Other researchers maintain that for Neandertals, in- startling announcement. Excavations that began in the 1940s
terment merely constituted a way of concealing the decompos- at the Grotte du Renne at Arcy-sur-Cure near Auxerre, France,
ing body, which might have attracted unwelcome predators. had yielded numerous blades, body ornaments and bone tools
They view the purported grave goods as miscellaneous objects and revealed evidence of huts and hearths— all hallmarks of the
that happened to be swept into the grave.) Evidence for art, in Upper Paleolithic. The scant human remains found amid the
the form of isolated pierced teeth and engraved bone fragments, artifacts were impossible to identify initially, but using com-
and red and yellow ocher, has been reported from a few sites, puted tomography to examine the hidden inner-ear region pre-
too, but given their relative rarity, researchers tend to assign served inside an otherwise uninformative skull fragment, Hub-
alternative explanations to these items. lin’s team identified the specimen as Neandertal.
The possibility that Neandertals might have engaged in mod- In response, a number of scientists suggested that Neander-
ern practices was taken more seriously in 1980, when researchers tals had acquired the modern-looking items by stealing them,
reported a Neandertal from the Saint-Césaire rock-shelter in collecting artifacts discarded by moderns or perhaps trading for
Charente-Maritime, France, found along with stone tools man- them. But this view has come under fire, most recently from ar-
ufactured according to a cultural tradition known as the Châ- chaeologists Francesco d’Errico of the University of Bordeaux
telperronian, which was assumed to have been the handiwork and João Zilhão of the University of Lisbon, who argue that the

represented at the same site, the Neandertals were already moving toward social roles within Neandertal cultures.
Châtelperronian always underlies the modernity on their own. In other words, the Thus, “modern” behavior seems to have
Aurignacian, suggesting its priority. Châtelperronian and other late Neandertal emerged in different regions and among
Furthermore, consideration of the cultures, such as the Uluzzian of Italy, different groups of humans, as would
hundreds of datings available from this emerged in Europe around 40,000 years happen later in history with the invention
period in Europe and the Near East shows ago, long before any moderns established of agriculture, writing and state society.
that wherever the context of the dated themselves in those areas. An alternative explanation, taking into
samples is well known, the earliest That this autonomous development account the broadly simultaneous
occurrences of the Aurignacian are included the manufacture and use of appearance of personal ornaments in
apparently from no earlier than around symbolic objects created for visual display many parts of the Old World, is that
36,500 years ago. The same radiometric on the body, as are often observed in contacts between modern and archaic
data, however, indicate that by then traditional societies, reflects various humans challenged each group’s personal,
social and biological identities, igniting an
PENDANTS, BONE TOOLS AND KNIVES from the Grotte du Renne site seem to be the handiwork of explosion of production of symbolic objects
Neandertals. That the advanced items underlie early modern human cultural remains from the
same site and are manufactured according to methods different from those favored by the moderns
by all those involved. On the strength of the
suggests that some Neandertals independently developed a modern culture. available data, however, we favor the
hypothesis of independent invention.
Regardless of which is eventually
proved correct, the behavioral barrier that
FROM “LES DERNIERS NÉANDERTALIENS,” LA MAISON DES ROCHES, 1999;

seemed to separate moderns from
Neandertals and gave us the impression of
being a unique and particularly gifted
COURTESY OF DOMINIQUE BAFFIER (left and right panels),
human type— the ability to produce

symbolic cultures— has definitively
collapsed.
JOÃO ZILHÃO is professor of prehistoric

FRANCESCO D’ERRICO (center panel)
archaeology at the University of Lisbon

in Portugal.
FRANCESCO D’ERRICO is a CNRS

researcher at the Institute of Prehistory
and Quaternary Geology, University of
Bordeaux, in France.
[Neandertals] only started to do these things after the

modern humans had arrived in western Europe or at
least in northern Spain,” he asserts. Unfortunately, be-
Châtelperronian artifacts at the Grotte du Renne and elsewhere, cause scientists have been unable to date these sites with suffi-
though superficially similar to those from the Aurignacian, re- cient precision, researchers can interpret the data differently.
flect an older, different method of manufacture [see “A Case for From his own work on the Grotte du Renne body ornaments,
Neandertal Culture,” above]. New York University archaeologist Randall White argues that
Most researchers are now convinced that Neandertals man- these artifacts reflect manufacturing methods known— albeit at
ufactured the Châtelperronian tools and ornaments, but what lower frequencies— from Aurignacian ornaments. Given the
prompted this change after hundreds of thousands of years is complicated stratigraphy of the Grotte du Renne site, the mod-
unclear. Cast in this light, “it’s more economical to see that as ern-looking items might have come from overlying Aurignacian
a result of imitation or acculturation from modern humans than levels. But more important, according to White, the Châtelper-
to assume that Neandertals invented it for themselves,” reasons ronian does not exist outside of France, Belgium, Italy and
Cambridge archaeologist Paul A. Mellars. “It would be an ex- northern Spain. Once you look at the Upper Paleolithic from a
traordinary coincidence if they invented all these things short- pan-European perspective, he says, “the Châtelperronian be-
ly before the modern humans doing the same things arrived.” comes post-Aurignacian by a long shot.”
Furthermore, Mellars disagrees with d’Errico and Zilhão’s pro- Still, post-Aurignacian does not necessarily mean after con-
posed order of events. “The dating evidence proves to me that tact with moderns. The earliest Aurignacian sites do not in-

THE FATE OF THE NEANDERTALS BY FRED H. SMITH
STRONG EVIDENCE has accumulated in It seemed that the Neandertals were sent the most desirable real estate in central
recent years that the emergence of into complete extinction by a superior Europe as late as 28,000 years ago. These
modern humans in Europe resulted human species— us. dates, the most recent known for
largely from the immigration of peoples Evidence from an important site in Neandertal fossils, show that these
into the continent, probably from the Near northwestern Croatia calls aspects of this humans were not quickly relegated to
East, starting sometime between 40,000 conventional wisdom into question. By the periphery; they competed quite well
and 30,000 years ago. Most researchers performing accelerator mass with intruding modern populations for
envisioned these early modern spectrometry dating directly on two a long time.
populations as having moved into Neandertal specimens from Vindija cave, This overlap of Neandertal and early
Anatolia and the Balkans, then up through my colleagues and I have demonstrated modern peoples for several millennia in
the plains and valleys of central Europe, that Neandertals were living in some of the heart of Europe allowed considerable
and finally into northern and western
Europe. Meanwhile the indigenous
Neandertals, it was thought, were
systematically pushed into more
peripheral and undesirable parts of the
landscape by these expanding
populations of moderns. The Neandertals’ MLADEČ
last bastion appeared to have been the (32,000 – 35,000 years ago)
Iberian Peninsula, where fossils from a
Spanish site called Zafarraya have been VOGELHERD
(32,000 years ago)
dated to 32,000 years ago and tools
attributed to Neandertals have been VINDIJA
LAGAR VELHO
dated to around 28,000 years ago. (24,500 years ago) (28,000 years ago)
A number of scholars argued that after
this time no traces of Neandertals
remained in Europe and that the
Neandertals did not make any biological
contributions to early modern humans. ZAFARRAYA
(28,000 – 32,000 years ago)
clude any human remains. Researchers have

assumed that they belonged to moderns be-
cause moderns are known from younger
Aurignacian sites. But “who the Aurigna-
cians were biologically between 40,000 and
35,000 years ago remains very much an Early Modern
unanswered question,” White notes. Neandertal
He adds that if you look at the Near East
around 90,000 years ago, anatomically mod-
ern humans and Neandertals were both making Mousterian lumbia University, all the brain asymmetries that characterize
stone tools, which, though arguably less elaborate than Auri- modern humans are found in Neandertals. “To be able to dis-
gnacian tools, actually require a considerable amount of know- criminate between the two,” he remarks, “is, at the moment, im-
how. “I cannot imagine that Neandertals were producing these possible.” As to whether Neandertal anatomy permitted speech,
kinds of technologically complex tools and passing that on from studies of the base of the skull conducted by Jeffrey T. Laitman
generation to generation without talking about it,” White de- of the Mount Sinai School of Medicine suggest that if they
clares. “I’ve seen a lot of people do this stuff, and I can’t stand talked, Neandertals had a limited vocal repertoire. The signifi-
over somebody’s shoulder and learn how to do it without a lot cance of such physical constraints, however, is unclear.
of verbal hints.” Thus, White and others do not buy the argu-
ment that moderns were somehow cognitively superior, espe- Fading Away
cially if Neandertals’ inferiority meant that they lacked lan- I F N E A N D E R T A L S P O S S E S S E D basically the same cognitive
guage. Instead it seems that moderns invented a culture that re- ability as moderns, it makes their disappearance additionally
SUSAN CARLSON
lied more heavily on material symbols. puzzling. But the recent redating of Neandertal remains from
Researchers have also looked to brain morphology for clues Vindija cave in Croatia emphasizes that this did not happen
to cognitive ability. According to Ralph L. Holloway of Co- overnight. Loyola’s Smith and his colleagues have demonstrated

opportunity for various interactions, and them. Morphologically, the Vindija same). Yet fossils from the Near Eastern
Vindija may reflect some of them. Work by Neandertals look more modern than do – and Qafzeh, which
sites of Skhul
my Croatian colleagues Ivor Karavanić of most other Neandertals, which suggests presumably represent the ancestors of
the University of Zagreb and Jakov that their ancestors interbred with early modern Europeans, do not have this
Radovčić of the Croatian Natural History early moderns. morphology. It is hard to explain how the
Museum has revealed a combination of The likelihood of gene flow between growth phenomenon responsible for this
Mousterian and Aurignacian tools in the the groups is also supported by evidence bunning could reappear independently and
same stratigraphic level as the dated that Neandertals left their mark on early ubiquitously in early modern Europeans.
Neandertal fossils, indicating that modern Europeans. Fossils representing Instead it is far more logical to recognize
Neandertals either made advanced early modern adults from central this morphology as a link to the
implements or traded with moderns for European sites such as Vogelherd in Neandertals. The Portuguese child
southwestern Germany and Mladeč in discovered late in 1998 in the Lapedo
Moravia (Czech Republic) have features Valley offers more intriguing clues [see “The
that are difficult to explain unless they Hybrid Child from Portugal,” on page 32].
have some Neandertal contribution to I believe the evidence shows that the
their ancestry. behavioral and biological interactions
For example, Neandertals and early between Neandertal and early modern
modern Europeans virtually all exhibit a human populations were very complex—too
projection of the back of the skull called an complex for the origins of modern humans
occipital bun (aspects of the shape and in Europe to have involved a simple,
position of the buns differ between them complete biological replacement of the
because the overall skull shapes are not the Neandertals. Neandertals as organisms no
longer exist, and Neandertal genes may not
MOVEMENT OF MODERNS (purple) into Europe have persisted to the present day, but
did not displace the Neandertals, who were still
living in central and western Europe 28,000 those genes were there in the beginnings of
years ago. A number of the early modern modern European biological history.
European specimens bear some Neandertal
features, which suggests that during the long FRED H. SMITH is a paleoanthropologist
period of overlap the two populations mixed. at Loyola University of Chicago.
that Neandertals still lived by the newcomers. Thousands of years of interbreeding be-
in central Europe 28,000 tween the small Neandertal population and the larger modern
years ago, thousands of human population, he surmises, diluted the distinctive Nean-
years after moderns had dertal features, which ultimately faded away.
QAFZEH moved in [see “The Fate of “If we look at Australians a thousand years from now, we
SKHUL
(around 90,000 the Neandertals,” above]. will see that the European features have predominated [over
years ago) Taking this into considera- those of native Australians] by virtue of many more Euro-
tion, Stringer imagines that peans,” Wolpoff asserts. “Not by virtue of better adaptation,
moderns, whom he views not by virtue of different culture, not by virtue of anything ex-
as a new species, replaced Neandertals in a long, slow process. cept many more Europeans. And I really think that’s what de-
“Gradually the Neandertals lost out because moderns were a scribes what we see in Europe— we see the predominance of
bit more innovative, a bit better able to cope with rapid envi- more people.”
ronmental change quickly, and they probably had bigger social From the morass of opinions in this contentious field, one
networks,” he supposes. consensus emerges: researchers have retired the vision of the
On the other hand, if Neandertals were an equally capable shuffling, cultureless Neandertal. But whether these ancient
variant of our own species, as Smith and Wolpoff believe, long- hominids were among the ancestors of living people or a close-
term overlap of Neandertals and the new population moving ly related species that competed with our own for the Eurasian
into Europe would have left plenty of time for mingling, hence territory and lost remains to be seen. In either case, the details
the mixed morphology that these scholars see in late Neander- will be extraordinarily complicated. “The more we learn, the
tals and early moderns in Europe. And if these groups were ex- more questions arise, the knottier it gets,” muses archaeologist
changing genes, they were probably exchanging cultural ideas, Lawrence G. Straus of the University of New Mexico. “That’s
which might account for some of the similarity between, say, why simple explanations just don’t cut it.”
the Châtelperronian and the Aurignacian. Neandertals as enti-
ties disappeared, Wolpoff says, because they were outnumbered Kate Wong is editorial director of ScientificAmerican.com

EMERGENCE
out of
Africa is the birthplace of

afr
species evolved there? And when
I
t all used to seem so simple. The recent evidence does seem to indicate
human lineage evolved in Africa. that it was not necessarily H. erectus
Only at a relatively late date did who migrated from Africa— and that
early humans finally migrate these peregrinations began earlier than
from the continent of their birth, we had thought.
in the guise of the long-known
species Homo erectus, whose first A Confused Early History
representatives had arrived in eastern RECENT DISCOVERIES in Kenya of
Asia by around one million years ago. fossils attributed to the new species Aus-
All later kinds of humans were the de- tralopithecus anamensis have pushed
scendants of this species, and almost back the undoubted record of upright-
everyone agreed that all should be clas- walking hominids to about 4.2 to 3.9
sified in our own species, H. sapiens. To million years ago. The most recent finds
acknowledge that some of these de- in Kenya and Chad may push this back
scendants were strikingly different from to six million years ago or more. The A.
ourselves, they were referred to as “ar- anamensis fossils bear a strong resem-
chaic H. sapiens,” but members of our blance to the later and far better known
own species they were nonetheless con- species Australopithecus afarensis,
sidered to be. found at sites in Ethiopia and Tanzania
Such beguiling simplicity was, alas, in the 3.9- to 3.0-million-year range and
too good to last, and over the past few most famously represented by the “Lucy”
years it has become evident that the lat- skeleton from Hadar, Ethiopia.
er stages of human evolution have been Lucy and her kind were upright walk-
a great deal more eventful than conven- ers, as the structures of their pelvises and
tional wisdom for so long had it. This is knee joints particularly attest, but they
true for the earlier stages, too, although retained many ancestral features, no-
there is still no reason to believe that hu- tably in their limb proportions and in
mankind’s birthplace was elsewhere their hands and feet, that would have
“LUCY” SKELETON represents the best-known than in Africa. Indeed, for well over the made them fairly adept tree climbers.
species of early hominid, or human precursor, first half of the documented existence of Together with ape-size brains and large,
Australopithecus afarensis, often
characterized as a “bipedal chimpanzee.” the hominid family (which includes all protruding faces, these characteristics
The 3.18-million-year-old skeleton is from upright-walking primates), there is no have led many to call such creatures
the Hadar region of Ethiopia. record at all outside that continent. But “bipedal chimpanzees.” This is proba-

ica
By Ian Tattersall
again . . . and again?

humanity. But how many human
did they emigrate?
bly a fairly accurate characterization, es- more modern body structures. Exactly
pecially given the increasing evidence how many species of early hominids
that early hominids favored quite heav- there were, which of them made the
PHOTOGRAPH BY D. FINNIN AND J. BECKETT, FROM CAST ON DISPLAY AT THE AMERICAN MUSEUM OF NATURAL HISTORY (these two pages)
ily wooded habitats. Their preferred tools, and how they walked remain
way of life was evidently a successful among the major conundrums of human
one, for although these primates were evolution.
less adept arborealists than the living Physically, at least, the picture be-
apes and less efficient bipeds than later comes clearer after about 1.9 million
hominids, their basic “eat your cake and years ago, when the first good evidence
have it” adaptation endured for well occurs in northern Kenya of a species that
over two million years, even as species of is recognizably like ourselves. Best exem-
this general kind came and went in the plified by the astonishingly complete 1.6-
fossil record. million-year-old skeleton known as the
It is not even clear to what extent Turkana Boy, discovered in 1984, these
lifestyles changed with the invention of humans possessed an essentially modern
stone tools, which inaugurate our ar- body structure, indicative of modern
chaeological record at about 2.5 million gait, combined with moderately large-
years ago. No human fossils are associ- faced skulls that contained brains double
ated with the first stone tools known, the size of those of apes (though not
from sites in Kenya and Ethiopia. In- much above half the modern human av-
stead there is a motley assortment of erage). The Boy himself had died as an
hominid fossils from the period follow- adolescent, but it is estimated that had he
ing about two million years ago, mostly lived to maturity he would have attained
associated with the stone tools and a height of six feet, and his limbs were
butchered mammal bones found at Tan- long and slender, like those of people
zania’s Olduvai Gorge and in Kenya’s who live today in hot, arid African cli-
East Turkana region. By one reckoning, mates, although this common adaptation
at least some of the first stone toolmak- does not, of course, indicate any special
ers in these areas were hardly bigger or relationship. Here at last we have early
more advanced in their body skeletons hominids who were clearly at home on
“TURKANA BOY,” an adolescent Homo ergaster
than the tiny Lucy; by another, the first the open savanna. dated to about 1.6 million years ago, is
tools may have been made by taller, A long-standing paleoanthropologi- representative of the first hominids with an
somewhat larger-brained hominids with cal tradition seeks to minimize the num- effectively modern body skeleton.

PHOTOGRAPH BY ROBERT CAMPBELL; COURTESY OF MEAVE LEAKEY
NEWLY DISCOVERED
SPECIES: Australopithecus
anamensis is the earliest
well-documented
ber of species in the human fossil record hominid. This lower jaw
and to trace a linear, progressive pattern from Kanapoi, Kenya,
of descent among those few that are rec- seen as it was found in
ognized. In keeping with this practice, the field, has been dated
to around four million
the Boy and his relatives were originally years ago. A. anamensis
assigned to the species H. erectus. This closely resembles
species was first described from a skull- A. afarensis in dental
cap and thighbone found in Java a cen- details, and a partial tibia
tury ago. Fossils later found in China— (shinbone) indicates that
it walked upright.
notably the now lost 500,000-year-old
“Peking Man”— and elsewhere in Java
were soon added to the species, and
eventually H. erectus came to embrace a
wide variety of hominid fossils, includ- one million years old, the conclusion ap- standard-issue hominid of the 1- to 0.5-
ing a massive braincase from Olduvai peared clear: H. erectus (as exemplified million-year period, was in fact a local
Gorge known as OH9. The latter has by OH9 and also by the earlier Turkana (and, as I shall explain below, ultimate-
been redated to about 1.4 million years, Boy and associated fossils) had evolved ly terminal) eastern Asian development.
although it was originally thought to in Africa and had exited that continent
have been a lot younger. All these fossil not much more than one million years An Eastern Asian Cul-de-Sac
forms possessed brains of moderate size ago, rapidly spreading to eastern Asia T H E P L O T T H I C K E N E D in early 1994,
(about 900 to 1,200 milliliters in vol- and spawning all subsequent develop- when Carl C. Swisher of the Berkeley
ume, compared with an average of ments in human evolution, including Geochronology Center and his col-
around 1,400 milliliters for modern hu- those in Europe. leagues applied the newish argon/argon
mans and about 400 milliliters for apes), Yet on closer examination the speci- dating method to volcanic rock samples
housed in long, low skull vaults with mens from Kenya turned out to be dis- taken from two hominid sites in Java.
sharp angles at the back and heavy brow tinctively different in braincase con- The results were 1.81 and 1.66 million
ridges in front. The few limb bones struction from those of classic eastern years: far older than anyone had really
known were robust but essentially like Asian H. erectus. In particular, certain expected, although the earlier date did
our own. anatomical features that appear special- confirm one made many years before.
Whether H. erectus had ever occu- ized in the eastern Asian H. erectus look Unfortunately, the fossils from these two
pied Europe was vigorously debated, the ancestral in the African fossils of compa- sites are rather undiagnostic as to species:
alternative being to view all early human rable age. Many researchers began to re- the first is a braincase of an infant (juve-
fossils from that region (the earliest of alize that we are dealing with two kinds niles never show all the adult character-
them being no more than about 500,000 of early human here, and the earlier istics on which species are defined), and
years old) as representatives of archaic Kenyan form is now increasingly placed the second is a horrendously crushed and
H. sapiens. Given that the Javan fossils in its own species, H. ergaster. This spe- distorted cranium that has never been
were conventionally dated in the range cies makes a plausible ancestor for all satisfactorily reconstructed. Both speci-
of one million to 700,000 years and subsequent humans, whereas the cranial mens have been regarded by most as H.
younger and that the earliest Chinese specializations of H. erectus suggest that erectus, more for reasons of convenience
fossils were reckoned to be no more than this species, for so long regarded as the than anything else. Over the decades,
“PEKING MAN” is the name

PHOTOGRAPH BY CRAIG CHESEK, FROM A CAST AT
THE AMERICAN MUSEUM OF NATURAL HISTORY
given to this skull of a male

H. erectus from Zhoukoudian,
near Beijing. The skull was
reconstructed from
fragments of various
individuals, all probably
around 500,000 years old.
40 SCIENTIFIC AMERICAN
REPLICA OF OLDOWAN BASALT CORE illustrates how
sharp flakes were struck from the core to provide
cutting implements. Tools of this kind were first
made around 2.5 million years ago.
PHOTOGRAPHS BY WILLARD WHITSON
TWO ACHEULEAN TOOLS, from St. Acheul, France,

are probably around 300,000 years old, but
implements of this kind began to be made in
Africa as many as 1.5 million years ago. On the
left is a pointed hand ax and on the right a
blunt-ended cleaver.
sporadic debate has continued regarding A very early hominid departure from eastern Asia lacked such utensils, which
whether the Javan record contains one or Africa has the advantage of explaining led many to wonder why the first human
more species of early hominid. Further, an apparent anomaly in the archaeolog- immigrants to the region had not brought
major doubt has been cast on whether ical record. The stone tools found in sed- this technology with them, if their ances-
the samples that yielded the older date iments coeval with the earliest H. ergas- tors had already wielded it for half a mil-
were actually obtained from the same ter (just under two million years ago) are lion years. The new dates suggest, how-
spot as the infant specimen. Still, these essentially identical with those made by ever, that the first emigrants had left
dates do fit with other evidence pointing the first stone toolmakers many hun- Africa before the invention of the Ach-
to the probability that hominids of some dreds of thousands of years before. These eulean technology, in which case there is
kind were around in eastern Asia much crude tools consisted principally of sharp no reason why we should expect to find
earlier than anyone had thought. flakes struck with a stone “hammer” this technology in eastern Asia. Interest-
Independent corroboration of this from small cobbles. Effective cutting tools ingly, in 1989 Robin W. Dennell of the
scenario comes, for instance, from the though these may have been (experimen- University of Sheffield in England caused
Dmanisi site in the former Soviet repub- tal archaeologists have shown that even quite a stir by reporting very crude stone
lic of Georgia, where in 1991 a hominid elephants can be quite efficiently butch- tools from Riwat in Pakistan that are old-
lower jaw that its describers allocated to ered using them), they were not made to er than 1.6 million years. Their great age
H. erectus was found. Three different a standard form but were apparently pro- is now looking decreasingly anomalous.
methods indicated that this jaw was as duced simply to obtain a sharp cutting Of course, every discovery raises new
old as 1.8 million years, and with four edge. Following about 1.5 million years questions, and in this case the problem is
crania from the site now in hand, there ago, however, standardized stone tools to explain what it was that enabled hu-
is ample evidence of an unexpectedly began to be made in Africa, typified by man populations to expand beyond
early hominid exodus from Africa. Even the hand axes and cleavers of the Africa for the first time. Most scholars
the most parsimonious reading of the Acheulean industry (first identified in the had felt that it was technological ad-
admittedly imperfect record suggests mid-19th century from St. Acheul in vances that allowed the penetration of
that these pioneering emigrants must France). These were larger implements, the cooler continental areas to the north.
have been H. ergaster or something very carefully shaped on both sides to a tear- If, however, the first emigrants left Africa
much like it. drop form. Oddly, stone tool industries in equipped with only the crudest of stone-
DRAWINGS BY DON M C GRANAGHAN
SKULLCAP known as Olduvai

Hominid 9 (OH9) was dated to
1.4 million years old; it was
originally believed to have
been much younger. Its
affinities are still being debated.
SCIENTIFIC AMERICAN 41
SUCCESSIVE WAVES of early humans exited from
Africa to all parts of the Old World. The record of
these emigrations is incomplete, but it is evident
that this history is much longer and more
complex than has traditionally been believed.
Atapuerca Dmanisi
working technologies, we have to look
to something other than technological
prowess for the magic ingredient. And
because the first human diaspora appar-
ently followed hard on the heels of the
‘Ubeidiya
acquisition of more or less modern body
form, it seems reasonable to conclude
that the typically human wanderlust
emerged in concert with the emancipa-
tion of hominids from the forest edges
that had been their preferred habitat. Of
course, the fact that the Turkana Boy Hadar
and his kin were adapted in their body
proportions to hot, dry environments Turkana
does nothing to explain why H. ergaster Kanapoi
was able to spread rapidly into the cool-
er temperate zones beyond the Mediter- Olduvai
ranean; evidently the new body form that
made possible remarkable endurance in
open habitats was in itself enough to
make the difference.
The failure of the Acheulean ever to
diffuse as far as eastern Asia reinforces
the notion, consistent with the cranial the world. Further datings tend to con- ticism; but, if accurate, they have con-
specializations of H. erectus, that this firm this view. Swisher and his col- siderable implications for the overall
part of the world was a kind of paleo- leagues reported in 1996 dates for the pattern of human evolution. For they are
anthropological cul-de-sac. In this re- Ngandong H. erectus site in Java that so recent as to suggest that the long-lived
gion, ancient human populations large- center on only about 40,000 years ago. H. erectus might even have suffered a fate
ly followed their own course, indepen- These dates, though very carefully ob- similar to that experienced by the Nean-
dent of what was going on elsewhere in tained, have aroused considerable skep- dertals in Europe: extinction at the hands
PHOTOGRAPHS COURTESY OF RUSSELL CIOCHON University of Iowa
FOSSILS FROM LONGGUPO, such as the lower jaw fragment (side and top views at left), may
indicate the presence of hominids in China as many as 1.9 million years ago.
it that aligns it with any particular human
species. Future fossil finds from Long-
Beijing gupo will, with luck, clarify the situation;
meanwhile the incisor and stone tools are
clear evidence of the presence of humans
in China at what may be a very early date
Riwat indeed. These ancient eastern Asians were
Longgupo the descendants of the first emigrants
from Africa, and, whatever the hominids
of Longgupo eventually turn out to have
been, it is a good bet that Huang and his
colleagues are right in guessing that they
Indian represent a precursor form to H. erectus
Ocean
rather than that species itself.
All this makes sense, but one anom-
aly remains. If H. erectus was an indige-
nous eastern Asian development, then
LAURIE GRACE AND JANA BRENNING

Less than 0.1
we have to consider whether we have
0.1 to 0.5
correctly identified the Olduvai OH9
0.5 to 1.0 Java braincase as belonging to this species. If
1.0 to 1.5 we have, then H. erectus evolved in east-
1.5 to 2.0 ern Asia at quite an early date (remem-
More than 2.0 ber, OH9 is now thought to be almost
1.4 million years old), and one branch of
the species migrated back to Olduvai in
of late-arriving H. sapiens. Here we find inid lineages that had evolved elsewhere. Africa. But if these new Asian dates are
reinforcement of the gradually emerging At the other end of the scale, in 1996 accurate, it seems more probable that as
picture of human evolution as one of re- an international group led by Huang we come to know more about OH9 and
peated experimentation, with regional- Wanpo of Academia Sinica in Beijing re- its kind we will find that they belonged to
ly differentiated species, in this case on ported a remarkably ancient date for a different species of hominid altogether.
opposite sides of the Eurasian continent, Longgupo Cave in China’s Sichuan The opposite end of the Eurasian con-
being ultimately replaced by other hom- Province. This site had previously yield- tinent was, as I have hinted, also isolated
ed an incisor tooth and a tiny lower jaw from the human evolutionary main-
fragment with two teeth that were ini- stream. As we saw, humans seem to have
tially attributed to H. erectus, plus a few arrived in Europe fairly late. In this re-
very crude stone artifacts. Huang and gion, the first convincing archaeological
his colleagues concluded that the fossils sites, with rather crude tools, show up at
and tools might be as many as 1.9 mil- about 800,000 years ago or thereabouts
lion years old, and their reanalysis of the (although in the Levant, within hailing
fossils suggested to them a closer resem- distance of Africa, the site of ’Ubeidiya
blance to the earliest African Homo spe- has yielded Acheulean tools dated to
cies than to H. erectus.
This latter claim has not gone unex- IAN TATTERSALL was born in England
THE AUTHOR
amined. As my colleague Jeffrey H. and raised in East Africa. He is chair of

Schwartz of the University of Pittsburgh the department of anthropology at the
COURTESY OF ERIC DELSON
and I pointed out, for instance, the teeth American Museum of Natural History in
in the jaw fragment resemble African New York City. His latest books include
Homo in primitive features rather than in The Monkey in the Mirror: Essays on the
the specialized ones that indicate a special Science of What Makes Us Human (Har-
relationship. What is more, they bear a vard Books, 2003), Becoming Human:
striking resemblance to the teeth of an Evolution and Human Uniqueness (Har-
orangutan-related hominoid known from court, 1998) and The Fossil Trail: How
PARTIAL MANDIBLE (top and side views) from a much later site in Vietnam. And al- We Know What We Think We Know about
Dmanisi, in former Soviet Georgia, may be as though the incisor appears hominid, it is Human Evolution (Oxford University
old as 1.8 million years. Although it was initially fairly generic, and there is nothing about Press, 1995).
assigned to H. erectus, its species is still uncertain.
around 1.4 million years ago, just about In 1994 excavations at that site pro- noted various primitive traits in the fos-
as early as any found to the south). The duced numerous simple stone tools, plus sils, which they provisionally attributed
problem has been the lack of a sign of the quite a few human fossil fragments, the to H. heidelbergensis. This is the species
toolmakers themselves. most complete of which is a partial up- into which specimens formerly classified
This gap began to be filled by finds per face of an immature individual. All as archaic H. sapiens are increasingly be-
made by Eudald Carbonell of the Uni- came from a level that was dated to ing placed. Carbonell and his colleagues
versity of Tarragona in Spain and his co- more than 780,000 years ago. No traces see their fossils as the starting point of an
workers at the Gran Dolina Cave site in of Acheulean technology were found indigenous European lineage that grad-
the Atapuerca Hills of northern Spain. among the tools, and the investigators ually evolved into the Neandertals.
These latter, large-brained hominids
are known only from Europe and west-
ern Asia, where they flourished in the pe-
riod between about 200,000 years and
30,000 years ago, when they were ex-
tinguished in some way by invading H.
sapiens.
This is not the only possibility, how-
ever. With only a preliminary description
of the very fragmentary Gran Dolina
fossils available, it is hard to be sure, but
it seems at least equally possible that
they are the remains of hominids who
made an initial foray out of Africa into
Europe but failed to establish themselves
there over the long term. Representa-
tives of H. heidelbergensis are known in
Africa as well, as long ago as 600,000
years ago, and this species quite likely re-
GRAN DOLINA CAVE in the Atapuerca Hills of northern Spain has produced the earliest colonized Europe later on. There it
human fossils yet found in Europe. These fossils, dated to about 780,000 years ago and would have given rise to the Neander-
initially attributed to H. heidelbergensis, may in fact represent a distinct form. The mature tals, whereas a less specialized African
cranium (below) is from Sima de los Huesos, about one kilometer from Gran Dolina, where a population founded the lineage that ul-
huge trove of mostly fragmentary but exquisitely preserved human fossils is dated to timately produced H. sapiens.
about 300,000 years ago.
At another site, just a kilometer from
Gran Dolina, Juan-Luis Arsuaga of
Complutense University in Madrid and
his colleagues have discovered a huge
cache of exquisitely preserved human
fossils, about 400,000 years old. These
are said to anticipate the Neandertals in
certain respects, but they are not fully
Neandertal by any means. And although
they emphasize that the Neandertals
(and possibly other related species) were
an indigenous European development,
these fossils from Sima de los Huesos
JAVIER TRUEBA Madrid Scientific Films
(“Pit of the Bones”) do not establish an

unequivocal backward connection to
their Gran Dolina neighbors.
Born in Africa
E V E R Y L O N G T I M E R E A D E R of Sci-
entific American will be familiar with the
competing models of “regional continu-
ity” and “single African origin” for the

Homo sapiens H. sapiens
emergence of our own species,

H. neanderthalensis
H. sapiens [see “The Multire-
gional Evolution of Hu-
mans,” on page 46; and “The
Recent African Genesis of LEADING THEORIES of the origins of
Humans,” on page 54]. The modern humans are contrasted in
first of these models holds that these diagrams. According to the
the highly archaic H. erectus notion of “regional continuity,” all
H. heidelbergensis
modern human populations trace
(including H. ergaster) is their beginnings to H. erectus, but
nothing more than an ancient each regional population evolved H. erectus
variant of H. sapiens and that along its own distinctive lines,
for the past two million years exchanging enough genes with its
the history of our lineage has neighbors (arrows represent gene
exchange) to remain part of the
been one of a braided stream same species; all eventually
of evolving populations of this became H. sapiens. The “single
species in all areas of the Old origin” theory holds that H. ergaster
World, each adapting to local H. sapiens descended from a single
conditions, yet all consistent- ancestral population that emerged
H. erectus
in one place, probably Africa.
ly linked by gene exchange. REGIONAL CONTINUITY SINGLE ORIGIN
The variation we see today
among the major geographi-
cal populations of humans is, by this fossil record shows that from the earli- Most important, the new dates from
reckoning, simply the latest permutation est times, Africa was consistently the eastern Asia show that human-popula-
of this lengthy process. center from which new lineages of hom- tion mobility dates right back to the ori-
The other notion, which happens to inids sprang. Clearly, interesting evolu- gins of effectively modern bodily form.
coincide much better with what we tionary developments occurred in both Finds from Europe demonstrate that al-
know of evolutionary processes in gen- Europe and eastern Asia, but they in- though distinctive regional variants
eral, proposes that all modern human volved populations that were not only evolved there, the history of occupation
populations are descended from a single derived from but also eventually sup- of that region may itself not have been at
ancestral population that emerged in one planted by emigrants from Africa. In Af- all a simple one. As ever, though, new
place at some time between about rica our lineage was born, and ever since evidence of the remote human past has
150,000 and 100,000 years ago. The fos- its hominids were first emancipated served principally to underline the com-
sil evidence, thin as it is, suggests that this from the forest edges, that continent has plexity of events in our evolution. We
place of origin was somewhere in Africa pumped out successive waves of emi- can only hope that an improving fossil
(although the neighboring Levant is an grants to all parts of the Old World. record will flesh out the details of what
LAURIE GRACE; SOURCE: THE LAST NEANDERTHAL, BY IAN TATTERSALL (MACMILLAN, 1995)
alternative possibility); proponents of What we see in the human fossil record was evidently a richly intricate process
this scenario point to the support afford- as it stands today is without doubt a of hominid speciation and population
ed by comparative molecular studies for shadowy reflection at best of what must movement over the past two million
the notion that all living humans are de- have been a complex sequence of events. years.
scended from an African population.
In view of what I have already said MORE TO E XPLORE
about the peripheral roles played in hu- Three New Human Skulls from the Sima de los Huesos Middle Pleistocene Site in Sierra de
man evolution by early populations both Atapuerca, Spain. J.-L. Arsuaga et al. in Nature, Vol. 362, No. 6420, pages 534–537; April 8, 1993.
in eastern Asia and Europe, it should Age of the Earliest Known Hominids in Java, Indonesia. C. C. Swisher III et al. in Science, Vol. 263,
No. 5150, pages 1118–1121; February 25, 1994.
come as no surprise that between these
Early Homo and Associated Artefacts from Asia. W. Huang et al. in Nature, Vol. 378, No. 6554,
two possibilities my strong preference is pages 275–278; November 16, 1995.
for a single and comparatively recent Whose Teeth? J. H. Schwartz and I. Tattersall in Nature, Vol. 381, No. 6579, pages 201–202;
origin for H. sapiens, very likely in Af- May 16, 1996.
rica— the continent that, from the be- Latest Homo erectus of Java: Potential Contemporaneity with Homo sapiens in Southeast Asia.
ginning, has been the engine of main- C. C. Swisher III et al. in Science, Vol. 274, No. 5294, pages 1870–1874; December 13, 1996.
stream innovation in human evolution. A Hominid from the Lower Pleistocene of Atapuerca, Spain: Possible Ancestor to Neandertals
The rise of modern humans is a recent and Modern Humans. J. M. Bermúdez de Castro et al. in Science, Vol. 276, pages 1392–1395;
May 30, 1997.
drama that played out against a long
Earliest Pleistocene Hominid Cranial Remains from Dmanisi, Republic of Georgia: Taxonomy,
and complex backdrop of evolutionary Geological Setting, and Age. Leo Gabunia et al. in Science, Vol. 288, pages 1019–1025;
diversification among hominids, but the May 12, 2000.

EMERGENCE
the
multiregional
evolution of humans
By Alan G. Thorne and Milford H. Wolpoff
Both fossil and genetic evidence argues

that ancient ancestors of various human
groups lived where they are found today
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 rely from one’s 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

AFRICAN PATHWAY
Qafzeh 9 N Kow Swamp 1

(Upper Pleistocene) LU TIO
(recent Australian)
L EVO
GI ONA
LT IRE
MU
WAY
PATH
SIAN
RALA
AUST Willandra Lakes 50
Ngandong 1 (Upper Pleistocene)
(Indonesia)
ALTERNATIVE ANCESTRIES for a modern individual are

illustrated by various skulls. The progressive changes
in the skulls from Australasian sites (Kow Swamp,
Ngandong, Willandra Lakes and Sangiran) suggest that
the local modern people developed in Australasia over
hundreds of thousands of years. The Eve theory
(African pathway) claims that an early African was the
ancestor of all modern people, but significant features
of the skull from Qafzeh in Israel differ considerably
from those of the modern Australian skull. Multiregional
Sangiran 17
evolution combines these two pathways.
(Middle Pleistocene)
years ago, through an interconnected web Harvard University as the “Noah’s 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 Eve’s 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 Eve’s descendants were in-
An alternative theory, developed by male, dubbed “Eve” in one of Wilson’s vaders. The explanation offered for the
paleontologist William W. Howells of first publications on the subject, who lived current absence of other mitochondrial

SERIES OF CHINESE SKULLS shows continuity in form without evidence of a
replacement by African characteristics. From left to right, the male skulls are from
the Zhoukoudian Lower Cave (Middle Pleistocene period), Dali site (early Upper
Pleistocene period) and Zhoukoudian Upper Cave (late Upper Pleistocene).
DNA lineages is that none of the local groups. Second, implicit

women mixed with the invading modern within this idea is that the
men from Africa—which means that Eve earliest modern humans
founded a new species. Wilson’s recon- appeared in Africa. Third,
struction of the past demands that over a it also follows that the
period of no more than 150,000 years earliest modern humans
there was a complete replacement of all in other areas should have
the preexisting hunter-gatherers in Africa African features. Fourth,
and the rest of the then inhabited world; modern humans and the
later, the original African features of the people that they replaced
invading human species presumably gave should never have mixed
way to the modern populational features or interbred. Fifth, outside
we see in other regions. of Africa an anatomical
An analogy can highlight the differ- discontinuity should be
ence between our multiregional evolu- evident between the human fossils before dence for the introduction of a novel
tion theory and Wilson’s Eve theory. Ac- and after the replacement. technology.
cording to multiregional evolution, the Geoffrey G. Pope of William Paterson
pattern of modern human origins is like No Trace of Invasion University has pointed out that six
several individuals paddling in separate WE ARE SURPRISED by the allegation decades of research on the Asian Paleo-
corners of a pool; over time, they influ- that beginning about 200,000 years ago lithic record have failed to unearth any in-
ence one another with the spreading rip- one group of hunter-gatherers totally re- dication of intrusive cultures or tech-
ples they raise (which are the equivalent placed all others worldwide. Although it nologies. Types of artifacts found in the
of genes flowing between populations). is not uncommon for one animal species earliest Asian Paleolithic assemblages
In contrast, the total replacement re- to replace another locally in a fairly short continue to appear into the very late Pleis-
quirement of the Eve theory dictates that time, the claim that a replacement could tocene. If invading Africans replaced the
a new swimmer must jump into the pool occur rapidly in every climate and envi- local Asian populations, they must have
with such a splash that it drowns all the ronment is unprecedented. adopted the cultures and technologies of
other swimmers. One of these two views We would expect native populations the people they replaced and allowed
of our origin must be incorrect. to have an adaptive and demographic their own to vanish without a trace.
Mitochondrial DNA is useful for advantage over newcomers. Yet accord- Archaeological evidence for an inva-
guiding the development of theories, but ing to the Eve theory, it was the new- sion is also lacking in western Asia,
only fossils provide the basis for refuting comers who had the upper hand. To use where Christopher B. Stringer of the
one idea or the other. At best, the genet- a modern analogy, however, despite the Natural History Museum in London and
ic information explains how modern hu- overwhelming forces of destructive tech- a few other researchers believe the earli-
mans might have originated if the as- nologies and infectious diseases, most est modern humans outside of Africa can
sumptions used in interpreting the genes American and Australian indigenous be found at the Skhul- and Qafzeh sites in
are correct, but those conditions are only populations and their genes have con- Israel. The superb record at Qafzeh
hypothetical, and one theory cannot be tinued to persist through adaptation and shows, however, that these “modern”
used to test another. The fossil record is interbreeding. people had a culture identical to that of
the real evidence for human evolution, If a worldwide invasion and com- their local Neandertal contemporaries:
and it is rich in both human remains and plete replacement of all native peoples they made the same types of stone tools
archaeological sites stretching back for by Eve’s descendants actually took with the same technologies and at the
two million years. Unlike the genetic place, we would expect to find at least same frequencies; they had the same styl-
data, fossils can be matched to the pre- some archaeological traces of the be- ized burial customs, hunted the same
dictions of theories about the past with- haviors that made them successful. Yet game and even used the same butchering
out relying on a long list of assumptions. examining the archaeology of Asia, we procedures. Moreover, no evidence from
MILFORD H. WOLPOFF
The Eve theory makes five predictions can find none. For instance, whereas the the time when Eve’s 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-

fute the archaeological predictions im- browridges forming an almost straight lations but were otherwise remarkably
plied by the Eve theory. bar of bone across their eye sockets and similar to much earlier individuals in the
Perhaps that refutation explains why a second well-developed shelf of bone at region.
Wilson turned to a different advantage, the back of the skull for the neck muscles.
asserting that the invasion was successful Above and behind the brows, the fore- Australians and Eve
because Eve’s descendants carried a mi- head is flat and retreating. These early In- T H E F I R S T I N H A B I T A N T S of Aus-
tochondrial gene that conferred language donesians also have large projecting faces tralia arrived more than 60,000 years
ability. This proposal is yet to be widely with massive rounded cheekbones. Their ago, and their behavior and anatomy
accepted. Not only does it conflict with teeth are the largest known in archaic hu- were clearly those of modern human be-
paleoneurology about the language abil- mans from that time. ings. Some of their skeletons show the Ja-
ities of archaic humans, but if it were A series of small but important fea- van complex of features, along with fur-
true, it would violate the assumption re- tures can be found on the most complete ther braincase expansions and other
quired of Wilson’s clock that mitochon- face and on other facial fragments that modernizations. Several dozen well-pre-
drial mutations are neutral. are preserved. These include such things served fossils from the late Pleistocene
The remaining predictions of the Eve as a rolled ridge on the lower edge of the and early Holocene demonstrate that the
theory relate to abrupt anatomical eye sockets, a distinctive ridge on the same combination of features that dis-
changes and whether the earliest recog- cheekbone and a nasal floor that blends tinguished those Indonesian people from
nizably modern humans resembled ear- smoothly into the face. their contemporaries distinguishes some
lier regional populations or Africans. Most of this unique morphology was ancestors of indigenous Australians from
With the fossil evidence known at this retained for at least 700,000 years while other living peoples.
time, these questions can be resolved in other modern characteristics continued If the earliest Australians were all de-
at least two and possibly three regions of to evolve in the Javan people. For exam- scendants of Africans, as the Eve theory
the world. The most convincing data are ple, the large fossil series from Ngan- requires, the continuity of fossil features
from southern and northern Asia. dong, which evidence suggests is as old would have to be no more than apparent.
The hominid fossils from Australasia as 200,000 years, offers striking proof All the features of the early Javans would
(Indonesia, New Guinea and Australia) that the Javans of that time had brain need to have evolved a second time in the
show an anatomical sequence during the sizes similar to modern Australian popu- population of invaders. The repeated evo-
Pleistocene that is uninterrupted by a
new African species at any time. The dis- ALAN G. THORNE and MILFORD H. WOLPOFF have extensively studied the original fossil ma-
THE AUTHORS
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 Thorne’s 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.

lution of an individual feature would be prehistoric and living Asian incisors is many have been described as mixtures.
conceivable but rare; the duplication of unique to the region. Clearly, the European Neandertals were
an entire set of unrelated features would This combination of traits is also ex- not completely replaced by Africans or by
be unprecedentedly improbable. hibited at the Zhoukoudian Cave area in people from any other region.
Northern Asia also harbors evidence northern China, where fully a quarter of Instead the evidence suggests that Ne-
linking its modern and ancient inhabi- all known human remains from the Mid- andertals either evolved into later humans
tants. Moreover, because the similarities dle Pleistocene have been found. As Wu or interbred with them, or both. David
involve features that are different from Rukang and Zhang Yinyun of the Chi- W. Frayer of the University of Kansas and
those significant in Australasia, they nese Academy of Sciences have pointed Fred H. Smith, now at Loyola University
compound the improbability of the Eve out, even within the 150,000 or more of Chicago, have discovered that many
theory by requiring that a second com- years spanned by the Zhoukoudian indi- allegedly unique Neandertal features are
plete set of features was duplicated in an- viduals, evolutionary changes in the found in the Europeans who followed the
other population. modern direction, including increases in Neandertals— the Upper Paleolithic,
The very earliest Chinese fossils, brain size and decreases in tooth size, can Mesolithic and later peoples. In fact, only
about one million years old, differ from be seen. Our examinations of the Chi- a few Neandertal features completely
their Javan counterparts in many ways nese specimens found no anatomical ev- disappear from the later European skele-
that parallel the differences between idence that typically African features ever tal record.
north Asians and Australians today. replaced those of the ancient Chinese in Features that persist range from high-
Our research with Wu Xinzhi and inde- these regions. Instead there is a smooth ly visible structures, such as the promi-
pendent research by Pope demonstrated transformation of the ancient popula- nent shape and size of the nose of Nean-
that the Chinese fossils are less robust, tions into the living peoples of east Asia dertals and later Europeans, to much
have smaller and more delicately built and the Americas. more minute traits, such as the form of
flat faces, smaller teeth and rounder Paleontologists have long thought Eu- the back of the skull and the details of its
foreheads separated from their arched rope would be the best source of evidence surface. A good example is the shape of
browridges. Their noses are less promi- for the replacement of one group, Nean- the opening in the mandibular nerve
nent and more flattened at the top. Per- dertals, by more modern humans. Even canal, a spot on the inside of the lower
haps the most telling indication of mor- there, however, the fossil record shows jaw where dentists often give a pain-
phological continuity concerns a pecu- that any influx of new people was neither blocking injection. The upper part of the
liarity of tooth shapes. Prominently complete nor without mixture. The most opening is covered by a broad bony
“shoveled” maxillary incisors, which recent known Neandertal skull, from bridge in many Neandertals, but in oth-
curl inward along their internal edges, Saint-Césaire in France, apparently had ers the bridge is absent. In European fos-
are found with unusually high frequen- the behavioral characteristics of the peo- sils, 53 percent of the known Neander-
cy in living east Asians and in all the ear- ple who succeeded the Neandertals in Eu- tals have the bridged form; 44 percent of
lier human remains from that area. Stud- rope. The earliest post-Neandertal Euro- their earliest Upper Paleolithic successors
ies by Tracey L. Crummett of San José peans did not have a pattern of either do, too, but in later Upper Paleolithic,
State University show that the form of modern or archaic African features, and Mesolithic and recent groups, the inci-
dence drops to less than 6 percent.
Inferred History of Actual History of In contrast, the bridged form is seen
Mitochondrial DNA Branching Mitochondrial DNA Branching
rarely in fossil or modern people from
Common ancestor
Asia and Australia. In Africa the few
jaws that date from the suggested Eve pe-
Hypothesized
common ancestor riod do not have it. This mandibular trait
and others like it on the skull and the
skeleton must have evolved twice in Eu-
rope for the Eve theory to be correct.
In sum, the evolutionary patterns of
three different regions—Australasia, Chi-
Only three mutations Five mutations na and Europe— show that their earliest
back to common ancestor back to common ancestor modern inhabitants do not have the com-
plex of features that characterize Africans.
Surviving types Extinct types
There is no evidence that Africans com-
pletely replaced local groups. Contrary to
LAURIE GRACE
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

the earliest human populations and liv-
ing peoples in different regions. Like ge-
Nerve
netic variation, human anatomical vari-
ation reflects significant differences in
occurrence for characteristics found in
all populations.
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 drift—the accidental loss of lines—is 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 there—and 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 1’s, which generally appears more substantiated. We must wonder why the guished some lines of mitochondrial

DNA from Eve’s descendants and her Hemisphere, some human populations could not— those lost were gone forever.
contemporaries. shrank because of climate fluctuations Human populations with dissimilar
Any interpretation of the surviving during the ice ages. Archaeological evi- demographic histories can therefore be
mitochondrial DNA mutations in pop- dence from both Africa and Australia expected to preserve different numbers
ulations consequently depends on a suggests that similar population reduc- of mutations since their last common mi-
knowledge of how the size of the popu- tions may have taken place there as well. tochondrial DNA ancestor. They cannot
lations has changed over time and how These reductions could have exacerbat- be used together in a model that assumes
many maternal lines may have vanished. ed genetic drift and the loss of mito- the lengths of mitochondrial lineages re-
Random losses from genetic drift alter a chondrial DNA types. flect the age of their divergence. One can-
reconstruction of the tree of human mi- At the end of the ice ages, along with not assume that all the variation in a pop-
tochondrial DNA branching by pruning the first domestication of animals and ulation’s mitochondrial DNA stems sole-
off signs of past divergences. Each un- plants, some populations expanded ex- ly from mutations: the history of the
counted branch is a mutation never tak- plosively throughout a wide band of ter- population is also important.
en into account when determining how ritory from the Mediterranean to the
long ago Eve lived. Pacific coast of Asia. Although the num- No Molecular Clock
Changes in population sizes have ber of people expanded, the number of A M A J O R P R O B L E M with the Eve the-
been dramatic. In parts of the Northern surviving mitochondrial DNA lines ory, therefore, is that it depends on an ac-
curate molecular clock. Its accuracy must
be based on mutation rates at many dif-
ferent loci, or gene positions. Yet genes
in the mitochondrial DNA cannot re-
combine as genes in the nucleus do. All
the mitochondrial DNA genes are the
EUROPE equivalent of a single locus. The molec-
AFRICA EAST ASIA AUSTRALASIA
AND LEVANT ular clock based on mitochondrial DNA
Lagar Velho Shandingdong
is consequently unreliable.
Kow Swamp
LATE
^
Afalou Mitochondrial DNA may not be neu-
Predmostí 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
Qafzeh Klasies Dingcun

Krapina Omo Kibish Xujiayao of mitochondrial DNA evolution that
are dramatically slower than Wilson and
Ehringsdorf Sambungmachan his colleagues have claimed for humans.
Ngaloba Dali
LATE
Biache 1, 3 A number of molecular geneticists dis-

Florisbad Jinniushan
MIDDLE PLEISTOCENE
Zuttiyeh Ngandong
agree with Wilson’s interpretation of the
MIDDLE
Sima de los Huesos

Kabwe Zhoukoudian H mitochondrial genetic data.
Petralona Hexian The molecular clock has, we believe,
Ndutu
Arago Nanjing
Steinheim major problems: its rate of ticking has
probably been overestimated in some cas-
EARLY
Bodo Zhoukoudian Sangiran 2, 10,

Gran Dolina Ternifine D, E, L 12, 17 es and underestimated in others. Rebecca
Olduvai 12 Chenjiawo Trinil
Yunxian L. Cann of the University of Hawaii at
Manoa and Mark Stoneking of Pennsyl-
LATE
Buia, Bouri Gongwangling

Olduvai 9 Yuanmou vania State University, two of Wilson’s
LOWER PLEISTOCENE
students, have acknowledged that their

Konso Gardula
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

most recent common ancestor of all ex- similar to Asian Homo erectus remains, mtDNA is most like that of living hu-
isting human mitochondria, but she is and is anatomically intermediate between mans, whereas the oldest is least alike—
not the most recent common ancestor of earlier and later Africans, suggests that the the opposite of what we would expect
all living people. Mitochondrial history evolving Homo lineage in the early and from unaltered Neandertal mtDNA
is not population history, just as the his- middle Pleistocene was a single species, evolving in a separate genetic line.
tory of names mentioned earlier is not not a mix of different species evolving in More recently, researchers have ob-
the same as the history of populations. different places. Early specimens of “mod- tained sequences of nuclear DNA, and
Such an interpretation can fully reconcile erns” are also instructive. In the Australian they provide a different picture. Most
the fossil record with the genetic data. We case, significant ancestry in the Ngandong fundamentally, nuclear genes prove to be
propose that future research might more fossils from Indonesia could not be ex- older than the mitochondrial gene, in
productively focus on attempts to dis- cluded. In the European case, a 50 percent some cases by millions of years. If the ori-
prove this hypothesis than on attempts to contribution by Neandertals for the earli- gin of today’s mtDNA was also the ori-
recalibrate a clock that does not work. est moderns could not be excluded. These gin of a new species, all the older nuclear
The dramatic genetic similarities anatomical studies support the idea of variations should have been eliminated,
across the entire human race show the multiregional evolution. and most genes should be the approxi-
consequences of linkages between people Meanwhile genetic research has be- mate age of the species or younger. This
that extend to when our ancestors first come more definitive. The rate of change is the most significant disproof of the Eve
populated the Old World. They are the of mitochondrial DNA (mtDNA) was theory. Nuclear genes are much older
results of an ancient history of popula- first estimated over millions of years from than Eve and preserve evidence of past
tion connections and mate exchanges comparisons with chimpanzees, but with migrations, mostly out of Africa but also
that has characterized the human race modern intergenerational studies the rates from some other regions, followed by
since its inception. Human evolution have been found to be many times as fast. population mixtures that preserve past
happened everywhere because every area The effects of accidental loss of mtDNA variation. This genetic evidence signifi-
was always part of the whole. variation were greatly underestimated. cantly supports multiregional evolution.
Neither anatomical nor genetic analy- Then came the realization that because A greater focus on epistemology also
ses provide a basis for the Eve theory. In- mtDNA is a single molecule, it cannot re- has made it clear that the original debate
stead the fossil record and the interpreta- combine or have crossover, so selection over modern human origin was indeed a
tion of mitochondrial DNA variation can on any part of it is selection on the whole. debate about the pattern of human evo-
be synthesized to form a view of human Natural selection has repeatedly reduced lution. The multiregional model is an in-
origins that does fit all the currently its variation; the same has been found in traspecific, network model, fundamen-
known data. This synthetic view com- the nonrecombining parts of the nuclear tally different from the tree-based Eve
bines the best sources of evidence about chromosomes. If selection and not pop- theory. This was important because an
human evolution by making sense of the ulation history accounts for mtDNA vari- assumption that tree (branching) attri-
archaeological and fossil record and the ation, it does not address the Eve theory. butes describe population histories un-
information locked up in the genetic vari- MtDNA has also been recovered derlies the acceptance of gene trees as
ation of living people all over the world. from Neandertals and from ancient population trees. The increasing molec-
The richness of human diversity, which Australians, and some of it is unlike the ular and anatomical evidence against re-
contrasts with the closeness of human ge- modern form. This evidence addresses cent speciation underscores the appro-
netic relationships, is a direct consequence the issues of how, and how quickly, priateness of such a network model. Mo-
of evolution. We are literally most alike mtDNA changes, but it does not help re- lecular and anatomical variation reflect
where it matters— under the skin. solve the pattern of evolution. Also less something different than the time since
than helpful is the possibility that all the the separation of populations. They in-
Epilogue Neandertal mtDNA recovered so far clude the complexities of gene flow be-
I N T H E D E C A D E since this article orig- may have been altered by contamination tween groups, different histories of se-
inally appeared in Scientific American, sig- or DNA breakdown. This is suspected lection, and different population struc-
nificant discoveries and analyses have because the most recent Neandertal tures across space and over time.
changed the nature of the debate about
the pattern of human evolution. The find- MORE TO E XPLORE
ing of a 25,000-year-old Portuguese child Race and Human Evolution. Milford Wolpoff and Rachel Caspari. Simon & Schuster, 1997.
from Lagar Velho who has a combination Modern Human Ancestry at the Peripheries: A Test of the Replacement Theory. Milford H. Wolpoff,
of Neandertal and “modern European” John Hawks, David W. Frayer and Keith Huntley in Science, Vol. 291, pages 293–297; 2001.
characteristics suggests that Neandertals Mitochondrial DNA Sequences in Ancient Australians: Implications for Modern Human Origins.
Gregory J. Adcock et al. in Proceedings of the National Academy of Sciences USA, Vol. 98, No. 2,
mixed with other populations and there- pages 537–542; January 16, 2001.
fore were the same species. A million-year- Number of Ancestral Human Species: A Molecular Perspective. D. Curnoe and A. Thorne in
old Ethiopian skull found in Bouri that is Homo: Journal of Comparative Human Biology, Vol. 53, Issue 3, pages 201–224; March 2003.

EMERGENCE
African
the recent
Genesis
of humans
Genetic studies
reveal that an
African woman
from less than
200,000 years ago
was our common
ancestor
By Rebecca L. Cann
and Allan C. Wilson
POINT-COUNTERPOINT: For an opposing view of how humankind arose

around the globe, see “The Multiregional Evolution of Humans,” on page 46.

I
n the quest for the facts about had been right and they had been wrong. lived about 200,000 years ago, probably
human evolution, we molecular Once again we are engaged in a de- in Africa. Modern humans arose in one
geneticists have engaged in two bate, this time over the latest phase of place and spread elsewhere.
major debates with the paleon- human evolution. The paleontologists Neither the genetic information of
tologists. Arguing from their fos- say modern humans evolved from their living subjects nor the fossilized remains
sils, most paleontologists had archaic forebears around the world over of dead ones can explain in isolation
claimed the evolutionary split between the past million years. Conversely, our how, when and where populations orig-
humans and the great apes occurred as genetic comparisons convince us that all inated. But the former evidence has a cru-
long as 25 million years ago. We main- humans today can be traced along ma- cial advantage in determining the struc-
tained human and ape genes were too ternal lines of descent to a woman who ture of family trees: living genes must have
similar for the schism to be more than a
AFRICAN ORIGIN for all modern humans is indicated by the genetic evidence. A genealogy based on
few million years old. After 15 years of 182 current mitochondrial DNA types (outer edges) points to the existence of a common female
disagreement, we won that argument ancestor from Africa. The arrows on the map (center) indicate the route and the minimum number of
when the paleontologists admitted we unrelated females (red circles) who colonized various areas, as inferred from the branching pattern.
110 100 90 80 70
120 60
130
50
140
40
36 31
150
30
160
20
18
170
Ancestor
15 10
JOE L E MONNIER (map); LAURIE GRACE
African Asian Australian New Guinean Caucasian

180
0 0.2 0.4 0.6 0.6 0.4 0.2 0

Divergence in DNA Divergence in DNA
Sequence (percent) Sequence (percent)

ancestors, whereas dead fossils may not The fossil record, on the other hand, tory at the University of California at
have descendants. Molecular biologists is infamously spotty because a handful Berkeley, challenged a fossil primate
know the genes they are examining must of surviving bones may not represent the called Ramapithecus. Paleontologists
have been passed through lineages that majority of organisms that left no trace had dated its fossils to about 25 million
survived to the present; paleontologists of themselves. Fossils cannot, in princi- years ago. On the basis of the enamel
cannot be sure that the fossils they ex- ple, be interpreted objectively: the phys- thickness of the molars and other skele-
amine do not lead down an evolutionary ical characteristics by which they are tal characteristics, they believed that
blind alley. classified necessarily reflect the models Ramapithecus appeared after the diver-
The molecular approach is free from the paleontologists wish to test. If one gence of the human and ape lineages and
several other limitations of paleontol- classifies, say, a pelvis as human because that it was directly ancestral to humans.
ogy. It does not require well-dated fos- it supported an upright posture, then Sarich measured the evolutionary
sils or tools from each part of the fami- one is presupposing that bipedalism dis- distance between humans and chim-
ly tree it hopes to describe. It is not viti- tinguished early hominids from apes. panzees by studying their blood pro-
ated by doubts about whether tools Such reasoning tends to circularity. The teins, knowing the differences reflected
found near fossil remains were in fact paleontologist’s perspective therefore mutations that have accumulated since
made and used by the population those contains a built-in bias that limits its the species diverged. (At the time, it was
remains represent. And it concerns itself power of observation. much easier to compare proteins for
with a set of characteristics that is com- As such, biologists trained in modern subtle differences than to compare the
plete and objective. evolutionary theory must reject the no- genetic sequences that encode the pro-
A genome, or full set of genes, is tion that the fossils provide the most di- teins.) To check that mutations had oc-
complete because it holds all the inherit- rect evidence of how human evolution curred equally fast in both lineages, he
ed biological information of an individ- actually proceeded. Fossils help to fill in compared humans and chimpanzees
ual. Moreover, all the variants on it that the knowledge of how biological pro- against a reference species and found
appear within a population— a group of cesses worked in the past, but they that all the genetic distances tallied.
individuals who breed only with one an- should not blind us to new lines of evi- Sarich now had a molecular clock;
other— can be studied, so specific pecu- dence or new interpretations of poorly the next step was to calibrate it. He did
liarities need not distort the interpreta- understood and provisionally dated ar- so by calculating the mutation rate in
tion of the data. Genomes are objective chaeological materials. other species whose divergences could
because they present evidence that has be reliably dated from fossils. Finally, he
not been defined, at the outset, by any Molecular Clock applied the clock to the chimpanzee-hu-
particular evolutionary model. Gene se- ALL THE ADVANTAGES of our field man split, dating it to between five mil-
quences are empirically verifiable and stood revealed in 1967, when Vincent lion and seven million years ago—far lat-
not shaped by theoretical prejudices. M. Sarich, working in Wilson’s labora- er than anyone had imagined.
The Inheritance of Mitochondrial DNA

Egg Fertilized egg
37 genes
Mitochondrial DNA
Mitochondrion Nuclear DNA MOST OF AN INDIVIDUAL’S GENES are located on DNA molecules
in the cell nucleus. Mitochondria, the specialized structures that
provide cells with energy, also carry some genes for their own
manufacture on a ring of DNA. When a sperm and an egg cell unite,
they contribute equally to the DNA in the nucleus of the resulting
Sperm cell. All the mitochondria and the DNA they contain, however,
LAURIE GRACE
derive from the egg. Studies of mitochondrial DNA can reveal

an individual’s maternal ancestry.

Mitochondrial DNA is inherited from the
mother alone, so all of it today had one female ancestor.

At first, most paleontologists clung convert food into a form of energy the chondrial DNA is inherited from the
to the much earlier date. But new fossil rest of the cell can use. Unlike the DNA mother alone, unchanged except for
finds undermined the human status of of the nucleus, which forms bundles of chance mutations. The father’s contribu-
Ramapithecus: it is now clear that Ra- long fibers, each consisting of a protein- tion ends up on the cutting-room floor,
mapithecus is actually Sivapithecus, a coated double helix, the mitochondrial as it were. The nuclear genes, to which
creature ancestral to orangutans and not DNA comes in small, two-stranded the father does contribute, descend in
to any of the African apes at all. More- rings. Whereas nuclear DNA encodes an what we may call ordinary lineages,
over, the age of some sivapithecine fos- estimated 100,000 genes— most of the which are of course important to the
sils was downgraded to only about six information needed to make a human transmission of physical characteristics.
million years. By the early 1980s almost being— mitochondrial DNA encodes For our studies of modern human ori-
all paleontologists came to accept Sarich’s only 37. In this handful of genes, every gins, however, we focus on the mito-
more recent date for the separation of one is essential: a single adverse muta- chondrial, maternal lineages.
the human and ape lines. Those who con- tion in any of them is known to cause Maternal lineages are closest among
tinue to reject his methods have been re- some severe neurological diseases. siblings because their mitochondrial
duced to arguing that Sarich arrived at For the purpose of scientists studying DNA has had only one generation in
the right answer purely by chance. when lineages diverged, mitochondrial which to accumulate mutations. The de-
Two novel concepts emerged from DNA has two advantages over nuclear gree of relatedness declines step by step
the early comparisons of proteins from DNA. First, the sequences in mitochon- as one moves along the pedigree, from
different species. One was the concept of drial DNA that interest us accumulate first cousins descended from the mater-
inconsequential, or neutral, mutations. mutations rapidly and steadily, accord- nal grandmother, to second cousins de-
Molecular evolution appears to be dom- ing to empirical observations. Because scended from a common maternal great-
inated by such mutations, and they ac- many mutations do not alter the mito- grandmother and so on. The farther
cumulate at surprisingly steady rates in chondrion’s function, they are effective- back the genealogy goes, the larger the
surviving lineages. In other words, evo- ly neutral, and natural selection does not circle of maternal relatives becomes, un-
lution at the gene level results mainly eliminate them. til at last it embraces everyone alive.
from the relentless accumulation of mu- This mitochondrial DNA therefore Logically, then, all human mito-
tations that seem to be neither harmful behaves like a fast-ticking clock, which chondrial DNA must have had an ulti-
nor beneficial. The second concept, mo- is essential for identifying recent genetic mate common female ancestor. But it is
lecular clocks, stemmed from the obser- changes. Any two humans chosen ran- easy to show she did not necessarily live
vation that rates of genetic change from domly from anywhere on the planet are in a small population or constitute the
point mutations (changes in individual so alike in most of their DNA sequences only woman of her generation. Imagine
DNA base pairs) were so steady over that we can measure evolution in our a static population that always contains
long periods that one could use them to species only by concentrating on the 15 mothers. Every new generation must
time divergences from a common stock. genes that mutate fastest. Genes con- contain 15 daughters, but some mothers
trolling skeletal characters do not fall will not produce a daughter, whereas
Mitochondrial Clue within this group. others will produce two or more. Be-
W E C O U L D B E G I N to apply these Second, unlike nuclear DNA, mito- cause maternal lineages die out whenev-
methods to the reconstruction of later
stages in human evolution only after REBECCA L. CANN and ALLAN C. WILSON applied the tools of genetics to paleontology dur-
THE AUTHORS
1980, when DNA restriction analysis ing many of their collaborations. Cann is professor of genetics and molecular biology at the
made it possible to explore genetic dif- John A. Burns School of Medicine of the University of Hawaii at Manoa. She received both
ferences with high resolution. Workers her bachelor’s degree in genetics and her Ph.D. in anthropology from the University of Cal-
at Berkeley, including Wes Brown, Mark ifornia, Berkeley. As a postdoctoral fellow, she worked at Berkeley with Wilson and at the
Stoneking and us, applied the technique University of California, San Francisco. Cann is using mitochondrial DNA to assay the ge-
to trace the maternal lineages of people netic diversity of birds in the Hawaiian Islands. Until his death in 1991, Wilson was profes-
sampled from around the world. sor of biochemistry at Berkeley. A native of New Zealand, he received his doctorate from
The DNA we studied resides in the Berkeley. Wilson also worked at the Weizmann Institute of Science in Rehovot, Israel, at the
mitochondria, cellular organelles that University of Nairobi and at Harvard University.

er there is no daughter to carry it on, it homeland: the global distribution of mi- Working at Berkeley with Stoneking,
is only a matter of time before all but one tochondrial DNA types he saw could we expanded on Kocher’s work by ex-
lineage disappears. In a stable popula- then be explained most easily as the re- amining a larger genealogical tree made
tion, the time for this fixation of the ma- sult of no more than three migration up of 182 distinct types of mitochondri-
ternal lineage to occur is the length of a events to other continents. al DNA from 241 individuals. The mul-
generation multiplied by twice the popu- A crucial assumption in this analysis tiple occurrences of mitochondrial DNA
lation size. is that all the mitochondrial lineages types were always found among people
evolve at the same rate. So when Kocher from the same continent and usually in
Eve in Africa conducted his comparison of the human persons who lived within 100 miles of
ONE MIGHT REFER to the lucky wo- mitochondrial DNAs, he also included one another. Because the tree we con-
man whose lineage survives as Eve. Bear analogous sequences from four chim- structed had two main branches, both of
in mind, however, that other women panzees. If the human lineages had dif- which led back to Africa, it, too, sup-
were living in Eve’s generation and that fered in the rate at which they accumu- ported the hypothesis that Africa was
Eve did not occupy a specially favored lated mutations, then some of the 14 hu- the place of origin for modern humans.
place in the breeding pattern. She is man sequences would be significantly One noteworthy point that jumps
purely the beneficiary of chance. More- closer or farther away from the chim- out of our study is that although geo-
over, if we were to reconstruct the ordi- panzee sequences than others. In fact, all graphic barriers do influence a popula-
nary lineages for the population, they 14 human sequences are nearly equidis- tion’s mitochondrial DNA, people from
would trace back to many of the men tant from the chimpanzee sequences, a given continent do not generally all be-
and women who lived at the same time which implies that the rates of change long to the same maternal lineage. The
as Eve. Population geneticists Daniel L. among humans are fairly uniform. New Guineans are typical in this respect.
Hartl, now at Harvard University, and The chimpanzee data also illustrated Their genetic diversity had been suspect-
Andrew G. Clark, now at Cornell Uni- how remarkably homogeneous humans ed from linguistic analyses of the re-
versity, estimate that as many as 10,000 are at the genetic level: chimpanzees markable variety of language families—
people could have lived then. The name commonly show as much as 10 times the usually classified as Papuan— spoken on
“Eve” can therefore be misleading— she genetic variation of humans. That fact this one island [see “The Austronesian
is not the ultimate source of all the ordi- alone suggests that all of modern hu- Dispersal and the Origin of Languages,”
nary lineages, as the biblical Eve was. manity sprang from a relatively small by Peter Bellwood; Scientific Ameri-
From mitochondrial DNA data, it is stock of common ancestors. can, July 1991]. On our genealogical tree,
possible to define the maternal lineages
of living individuals all the way back to 1
a common ancestor. In theory, a great 2
number of different genealogical trees
could give rise to any set of genetic data. 3
To recognize the one that is most prob- 4
ably correct, one must apply the parsi- 5
mony principle, which requires that sub-
6
jects be connected in the simplest possi-
ble way. The most efficient hypothetical 7
Generation
tree must be tested by comparison with 8

other data to see whether it is consistent
9
with them. If the tree holds up, it is ana-
lyzed for evidence of the geographic his- 10
tory inherent in elements. 11
In 1988 Thomas D. Kocher of Berke-
12
ley (now at the University of New Hamp-
shire) applied just such a parsimonious 13
interpretation to the interrelatedness of 14
the mitochondrial DNA of 14 humans 15
from around the world. He determined
that 13 branching points were the fewest 16
that could account for the differences he
LAURIE GRACE
UNIVERSAL MATERNAL ANCESTOR can be found for all the members of any population. The example
found. Taking the geographic consider- shown here traces the lineages of 15 females in a stable population. In each generation, some
ations into account, he then concluded maternal lineages proliferate and others become extinct. Eventually, by chance, one maternal
that Africa was the ultimate human lineage (dark blue) replaces all the others.

Huge levels of gene flow between early continents—very
unlikely—would have been needed for multiregionalism.
New Guineans showed up on several dif- tochondrially speaking, races are not ed that constraint. The reaction makes it
ferent branches, which proved that the like biological species. We propose that possible to duplicate DNA sequences
common female ancestor of all New the anatomical characteristics uniting easily, ad infinitum; a small starting sam-
Guineans was not someone in New New Guineans were not inherited from ple of DNA can expand into an endless
Guinea. The population of New Guinea the first settlers. They evolved after peo- supply.
must have been founded by many moth- ple colonized the island, chiefly as the re- The polymerase chain reaction en-
ers whose maternal lineages were most sult of mutations in nuclear genes spread abled Linda Vigilant of Pennsylvania
closely related to those in Asia. by sex and recombination throughout State University to redo our study using
That finding is what one would ex- New Guinea. Similarly, the light skin mitochondrial DNA data from 120 Af-
pect if the African origin hypothesis color of many whites is probably a late ricans, representing six diverse parts of
were true: as people walked east out of development that occurred in Europe af- the sub-Saharan region. Vigilant traced
Africa, they would have passed through ter that continent was colonized by a genealogical tree whose 14 deepest
Asia. Travel was probably slow, and Africans. branches lead exclusively to Africans
during the time it took to reach New During the early 1980s, when we and whose 15th branch leads to both Af-
Guinea, mutations accumulated both in were constructing our genealogical tree, ricans and non-Africans. The non-Afri-
the lineages that stayed in Asia and in we had to rely on black Americans as cans lie on shallow secondary branches
those that moved on. substitutes for Africans, whose mito- stemming from the 15th branch. Con-
Thus, people who are apparently re- chondrial DNA was difficult to obtain in sidering the number of African and non-
lated by membership in a common geo- the required quantities. Fortunately, the African mitochondrial DNAs surveyed,
graphic race need not be very closely re- development of a technique called the the probability that the 14 deepest
lated in their mitochondrial DNA. Mi- polymerase chain reaction has eliminat- branches would be exclusively African is
one in 10,000 for a tree with this branch-
ing order.
Satoshi Horai and Kenji Hayasaka of
the National Institute of Genetics in
Mishima, Japan, analogously surveyed
population samples that included many
more Asians and individuals from few-
er parts of Africa; they, too, found that
the mitochondrial lineages led back to
Africa. We estimate the odds of their ar-
riving at that conclusion accidentally
were only four in 100. Although these
statistical evaluations are not strong or
rigorous tests, they do make it seem like-
African human ly that the theory of an African origin for
Non-African person human mitochondrial DNA is now fair-
Chimpanzee ly secure.
Pygmy chimpanzee
200,000 Years or Less
BECAUSE OUR COMPARISONS with
the chimpanzee data showed that the
human mitochondrial DNA clock has
ticked steadily for millions of years, we
knew it should be possible to calculate
LAURIE GRACE
INTERRELATEDNESS of 14 humans and four chimpanzees was inferred

from similarities discovered in their mitochondrial DNA sequences. The
when the common mother of humanity
chimpanzee data help researchers to measure when various evolutionary lived. We assumed that the human and
divergences in the human lineages occurred. chimpanzee lineages diverged five mil-

lion years ago, as Sarich’s work had might run by fits and starts, we ran a test occupied by archaic people who had mi-
shown. We then calculated how much to measure how much mitochondrial grated from Africa to Asia nearly a mil-
humans had diverged from one another DNA has evolved in populations found- lion years ago. Such famous fossils as
relative to how much they had diverged ed at a known time. Java Man and Beijing Man are of this
from chimpanzees— that is, we found The aboriginal populations of New type. This finding and the hypothesis
the ratio of mitochondrial DNA diver- Guinea and Australia are estimated to that the archaic Eurasian population un-
gence among humans to that between have been founded less than 50,000 to derwent anatomical changes that made
humans and chimpanzees. 60,000 years ago. The amount of evolu- them resemble more modern people led
Using two different sets of data, we tion that has since occurred in each of to the multiregional evolution model:
determined that the ratio was less than those places seems about one third of similar evolutionary changes in separate
1:25. Human maternal lineages there- that shown by the whole human species. geographic regions converted the inhab-
fore grew apart in a period less than 1⁄ 25 Accordingly, we can infer that Eve lived itants from archaic small-brained types
as long as five million years, or less than three times 50,000 to 60,000 years ago, to modern big-brained types.
200,000 years. With a third set of data or roughly 150,000 to 180,000 years Huge levels of gene flow between
on changes in a section of the mito- ago. All our estimates thus agree that the continents, however, would be neces-
chondrial DNA called the control re- split happened not far from 200,000 sary to maintain human populations as
gion, we arrived at a more ancient date years ago. one biological species. The multiregion-
for the common mother. That date is Those estimates fit with at least one al evolution model also predicts that at
less certain, however, because questions line of fossil evidence. The remains of least some genes in the modern east
remain about how to correct for multi- anatomically modern people appear first Asian population would be linked more
ple mutations that occur within the con- in Africa, then in the Middle East, and closely to those of their archaic Asian
trol region. later in Europe and east Asia. Anthropol- predecessors than to those of modern
One might object that a molecular ogists have speculated that in east Africa Africans. We would expect to find deep
clock known to be accurate over five the transition from anatomically archaic lineages in Eurasia, especially in the Far
million years could still be unreliable for to modern people took place as recently East. Yet surveys in our laboratories and
shorter periods. It is conceivable, for ex- as 130,000 years ago [see “The Emer- in others, involving more than 1,000
ample, that intervals of genetic stagna- gence of Modern Humans,” by Christo- people from Eurasia and its mitochon-
tion might be interrupted by short bursts pher B. Stringer; Scientific American, drial DNA satellites (Australia, Oceania
of change when, say, a new mutagen en- December 1990]. and the Americas), have given no hint of
ters the environment, or a virus infects On the other hand, a second line of that result.
the germ-line cells, or intense natural se- evidence appears to conflict with this It therefore seems very unlikely that
lection affects all segments of the DNA. view. The fossil record shows clearly any truly ancient lineages survive unde-
To rule out the possibility that the clock that the southern parts of Eurasia were tected in Eurasia. We simply do not see
the result predicted by the regional mod-
el. Moreover, geneticists such as Masa-
PRESENT AFRICAN EUROPEAN EAST ASIAN AUSTRALIAN
toshi Nei of Pennsylvania State Univer-
sity, Kenneth K. Kidd of Yale Universi-
ty, James Wainscoat of the University of
100,000 Klasies Neandertal Ngandong Oxford and Luigi L. Cavalli-Sforza of
Dali Stanford University have found support
for an African origin model in their stud-
Age (years)
ies of nuclear genes.

Zhoukoudian
300,000 Saldanha Petralona (“Beijing”) Sambungmachan Multiregional Mystery
P R O P O N E N T S O F the multiregional
evolution model typically emphasize
that they have documented a continuity
of anatomical morphologies between
700,000 Olduvai European Lantian Java
the archaic and modern residents of dif-
ferent regions; they insist that these mor-
phologies would be unlikely to evolve
Homo erectus
independently in any invading people.
LAURIE GRACE
ARCHAIC HUMAN GROUPS were gradually replaced throughout the Old World by modern humans who For that argument to hold true, howev-
arose in Africa. Archaic females do not seem to have contributed mitochondrial genes to the modern er, it must also be shown that the cranial
people of Europe, east Asia and Australia. features in question are truly indepen-

dent of one another— that is, that natur-
Son
al selection would not tend to favor cer-
tain constellations of functionally relat- Father Mother
ed features anyway. Yet we know that
powerful jaw muscles may impose
changes on the mandible, the browridge
and other points on the skull; circum-
stances that promoted the evolution of
these features in one population might
do so again in a related population.
Other paleontologists also dispute
the evidence for continuity. They argue
that modern populations are not linked
to past ones by morphological charac-
teristics that evolved uniquely in the fos- Male
sil record. Instead fossils and modern Female
populations are united by their shared
retention of still older ancestral charac- Mitochondrial DNA source
teristics. The continuity seen by believ- PEDIGREE of one individual illustrates the difference between the patterns of nuclear and
ers in multiregional evolution may be an mitochondrial inheritance. All 32 ancestors from five generations ago contributed equally to his
illusion. nuclear DNA. His mitochondrial lineage (blue line) leads back to only one person in every generation.
The idea that modern humans could
cohabit a region with archaic ones and Lucotte, while at the College of France, proved methods of extracting DNA
replace them completely without any and his colleagues have indirectly com- from still older fossilized bone now ap-
mixture may sound unlikely. Neverthe- pared such sequences in an effort to pear close at hand. With them, we may
less, some fossil finds do support the trace paternal lineages to a single pro- begin building the family tree from a root
idea. Discoveries in the caves at Qafzeh genitor —“Adam,” if you will. Those that was alive when the human family
in Israel suggest that Neandertals and preliminary results also point to an was young.
modern humans lived side by side for African homeland, and with further re-
40,000 years, yet they left little evidence finements this work on paternal lineages Epilogue
of interbreeding. may be able to provide an invaluable S I N C E T H I S A R T I C L E was first pub-
How one human population might check on our results for maternal lin- lished, further genetic work on the mi-
have replaced archaic humans without eages. Unfortunately, base changes ac- tochondrial DNA sequences of three
any detectable genetic mixing is still a cumulate slowly on useful regions of the Neandertal specimens upholds our con-
compelling mystery. One of us (Cann) Y chromosome, making it technically clusions about the lack of a mixture be-
suspects that infectious diseases could difficult to conduct a detailed genealog- tween ancient and modern Homo sapi-
have contributed to the process by help- ical analysis. ens. Furthermore, whole mitochondrial
ing to eliminate one group. Cavalli- More progress can be expected soon, genome sequencing— all 16,569 base
Sforza has speculated that the ancestors as molecular biologists learn to apply pairs from more than 50 donors— gives
of modern humans may have developed their techniques to materials uncovered more precise resolution to the timescale
some modern trait, such as advanced by our friendly rivals, the paleontolo- of our emergence. It now seems that the
language skills, that effectively cut them gists. Preliminary molecular studies have earliest migration out of Africa is closer
off from breeding with other hominids. already been conducted on DNA from to 120,000 years ago than 200,000 years
This and related questions may yield as mummified tissues found in a Florida ago— more recent, yet still within the
molecular biologists learn how to link bog and dated to 7,500 years ago. Im- range we had originally estimated.
specific genetic sequences to the physical
and behavioral traits that those se- MORE TO E XPLORE
quences ultimately influence. Mitochondrial DNA and Human Evolution. Rebecca L. Cann, Mark Stoneking and Allan C. Wilson in
Even before then, further studies of Nature, Vol. 325, No. 6099, pages 31–36; January 1–7, 1987.
both nuclear and mitochondrial DNA Mitochondrial DNA. M. Stoneking and A. C. Wilson in The Colonization of the Pacific: A Genetic Trail.
Edited by Adrian V. S. Hill and Susan W. Serjeantson. Oxford University Press, 1989.
will render more informative genetic
Mitochondrial DNA Sequences in Single Hairs from a Southern African Population. Linda Vigilant,
trees. Particularly enticing are the se-
LAURIE GRACE
Renee Pennington, Henry Harpending, Thomas D. Kocher and Allan C. Wilson in Proceedings of the
quences on the Y chromosome that de- National Academy of Sciences USA, Vol. 86, No. 23, pages 9350–9354; December 1989.
termine maleness and that are therefore Sequence Evolution of Mitochondrial DNA in Humans and Chimpanzees. T. D. Kocher and A. C.
inherited from the father alone. Gerard Wilson in Evolution of Life. Edited by S. Osawa and T. Honjo. Springer-Verlag, Tokyo, 1991.

ADAPTATION

SALAD DAYS: Australopithecus
afarensis, a human ancestor,
forages for plant foods in
an African woodland some
3.5 million years ago.

SKELETAL REMAINS indicate that our ancient forebears the australopithecines were
bipedal by four million years ago. In the case of A. afarensis (right), one of the earliest
hominids, telltale features include the arch in the foot, the nonopposable big toe, and
certain characteristics of the knee and pelvis. But these hominids retained some apelike
traits— short legs, long arms and curved toes, among others— suggesting both that they
probably did not walk exactly like we do and that they spent some time in the trees. It
wasn’t until the emergence of our own genus, Homo (a contemporary representative of
which appears on the left), that the hind limb features required for upright walking evolved.
These include the fully modern limb and foot proportions and pelvis morphology.
We humans are strange primates.

We walk on two legs, carry around natural selection acting to maximize di- from those of other primates? Further, to
enormous brains and have colonized etary quality and foraging efficiency. what extent have modern humans de-
every corner of the globe. Anthropolo- Changes in food availability over time, parted from the ancestral dietary pattern?
JOHN GURCHE (preceding pages and above)
gists and biologists have long sought to it seems, strongly influenced our homi- Scientific interest in the evolution of
understand how our lineage came to dif- nid ancestors. Thus, in an evolutionary human nutritional requirements has a
fer so profoundly from the primate sense, we are very much what we ate. long history. But relevant investigations
norm in these ways, and over the years Accordingly, what we eat is yet an- started gaining momentum after 1985,
all manner of hypotheses aimed at ex- other way in which we differ from our when S. Boyd Eaton and Melvin J. Kon-
plaining each of these oddities have been primate kin. Contemporary human pop- ner of Emory University published a sem-
put forth. But a growing body of evi- ulations the world over have diets richer inal paper in the New England Journal of
dence indicates that these miscellaneous in calories and nutrients than those of our Medicine entitled “Paleolithic Nutrition.”
quirks of humanity in fact have a com- cousins, the great apes. So when and how They argued that the prevalence in mod-
mon thread: they are largely the result of did our ancestors’ eating habits diverge ern societies of many chronic diseases—
64 SCIENTIFIC AMERICAN Updated from the December 2002 issue

obesity, hypertension, coronary heart dis- The type of environment a creature of Liverpool John Moores University
ease and diabetes, among them— is the inhabits will influence the distribution of submits that moving upright allowed
consequence of a mismatch between energy between these components, with early humans to better regulate their
modern dietary patterns and the type of harsher conditions creating higher main- body temperature by exposing less sur-
diet that our species evolved to eat as pre- tenance demands. Nevertheless, the goal face area to the blazing African sun.
historic hunter-gatherers. Since then, of all organisms is the same: to devote The list goes on. In reality, a number
however, understanding of the evolution sufficient funds to reproduction, which of factors probably selected for this type
of human nutritional needs has advanced ensures the long-term success of the spe- of locomotion. My own research, con-
considerably—thanks in large part to new cies. Thus, by looking at the way animals ducted in collaboration with my wife,
comparative analyses of traditionally liv- go about obtaining and then allocating Marcia L. Robertson, suggests that bi-
ing human populations and other pri- food energy, we can better discern how pedalism evolved in our ancestors at least
mates—and a more nuanced picture has natural selection produces evolutionary in part because it is less energetically ex-
emerged. We now know that humans change. pensive than quadrupedalism. Our analy-
have evolved not to subsist on a single, ses of the energy costs of movement in
Paleolithic diet but to be flexible eaters, an Becoming Bipeds living animals of all sizes have shown
insight that has important implications W H E N T H E Y A R E on the ground, liv- that, in general, the strongest predictors
for the current debate over what people ing nonhuman primates typically move of cost are the weight of the animal and
today should eat in order to be healthy. around on all fours, or quadrupedally. the speed at which it travels. What is
To appreciate the role of diet in hu- Scientists generally assume therefore that striking about human bipedal movement
man evolution, we must remember that the last common ancestor of humans and is that it is notably more economical than
the search for food, its consumption and, chimpanzees (our closest living relative) quadrupedal locomotion at walking rates.
ultimately, how it is used for biological was also a quadruped. Exactly when the Apes, in contrast, are not economical
processes are all critical aspects of an or- last common ancestor lived is unknown, when moving on the ground. For in-
ganism’s ecology. The energy dynamic but clear indications of bipedalism— the stance, chimpanzees, which employ a pe-
between organisms and their environ- trait that distinguished ancient humans culiar form of quadrupedalism known as
ments— that is, energy expended in rela- from other apes— are evident in the old- knuckle walking, spend some 35 percent
tion to energy acquired— has important est known species of Australopithecus, more calories during locomotion than
adaptive consequences for survival and which lived in Africa roughly four mil- does a typical mammalian quadruped of
reproduction. These two components of lion years ago. Ideas about why bipedal- the same size— a large dog, for example.
Darwinian fitness are reflected in the way ism evolved abound in the paleoanthro- Differences in the settings in which hu-
we divide up an animal’s energy budget. pological literature. C. Owen Lovejoy of mans and apes evolved may help explain
Maintenance energy is what keeps an an- Kent State University proposed in 1981 the variation in costs of movement.
imal alive on a day-to-day basis. Produc- that two-legged locomotion freed the Chimps, gorillas and orangutans evolved
tive energy, on the other hand, is associ- arms to carry children and foraged in and continue to occupy dense forests
ated with producing and raising off- goods. More recently, Kevin D. Hunt of where only a mile or so of trekking over
spring for the next generation. For Indiana University has posited that the course of the day is all that is needed
mammals, this must cover the increased bipedalism emerged as a feeding posture to find enough to eat. Much of early
costs that mothers incur during preg- that enabled access to foods that had pre- hominid evolution, on the other hand,
nancy and lactation. viously been out of reach. Peter Wheeler took place in more open woodland and
grassland, where sustenance is harder to
Overview/Diet and Human Evolution come by. Indeed, modern human hunter-
gatherers living in these environments,
■ The characteristics that most distinguish humans from other primates are who provide us with the best available
largely the results of natural selection acting to improve the quality of the model of early human subsistence pat-
human diet and the efficiency with which our ancestors obtained food. Some terns, often travel six to eight miles daily
scientists have proposed that many of the health problems modern societies in search of food.
face are consequences of a discrepancy between what we eat and what These differences in day range have
our Paleolithic forebears ate. important locomotor implications. Be-
■ Yet studies of traditionally living populations show that modern humans are cause apes travel only short distances
able to meet their nutritional needs using a wide variety of dietary strategies. each day, the potential energetic benefits
We have evolved to be flexible eaters. The health concerns of the industrial of moving more efficiently are very small.
world, where calorie-packed foods are readily available, stem not from For far-ranging foragers, however, cost-
deviations from a specific diet but from an imbalance between the energy effective walking saves many calories in
we consume and the energy we expend. maintenance energy needs—calories that
can instead go toward reproduction. Se-

0.0
Early H. sapiens
1,150 cc
0.5
Modern chimpanzee
400 cc
1.0 H. erectus
900 cc Modern H. sapiens
1,350 cc
Time (millions of years ago)
1.5 A. boisei
500 cc Homo habilis
600 cc
2.0
2.5 A. africanus
415 cc
3.0 BRAINS GREW BIGGER— and hence more

energetically demanding— over time.
The modern human brain accounts for
10 to 12 percent more of the body’s
3.5 Australopithecus afarensis resting energy requirements than the
385 cubic centimeters average australopithecine brain did.
4.0
5 7 9 11 13 15 17 19 21 23 25
Percent of Resting Energy Allocated to Brain
lection for energetically efficient loco- living apes, showing only a modest in- brains. In fact, at-rest brain metabolism
motion is therefore likely to be more increase in brain size, from around 400 cu- accounts for a whopping 20 to 25 per-
tense among far-ranging animals because bic centimeters four million years ago to cent of an adult human’s energy needs—
they have the most to gain. 500 cubic centimeters two million years far more than the 8 to 10 percent ob-
For hominids living between five mil- later. Homo brain sizes, in contrast, bal- served in nonhuman primates, and more
lion and 1.8 million years ago, during the looned from 600 cubic centimeters in H. still than the 3 to 5 percent allotted to the
Pliocene epoch, climate change spurred habilis some two million years ago up to brain by other mammals.
this morphological revolution. As the 900 cubic centimetersin early H. erectus By using estimates of hominid body
African continent grew drier, forests gave just 300,000 years later. The H. erectus size compiled by Henry M. McHenry of
way to grasslands, leaving food resources brain did not attain modern human pro- the University of California at Davis,
patchily distributed. In this context, bi- portions (1,350 cubic centimeters on av- Robertson and I have reconstructed the
pedalism can be viewed as one of the first erage), but it exceeded that of living non- proportion of resting energy needs that
strategies in human nutritional evolu- human primates. would have been required to support the
tion, a pattern of movement that would From a nutritional perspective, what brains of our ancient ancestors. Our cal-
have substantially reduced the number of is extraordinary about our large brain is culations suggest that a typical, 80- to
calories spent in collecting increasingly how much energy it consumes— roughly 85-pound australopithecine with a brain
dispersed food resources. 16 times as much as muscle tissue per size of 450 cubic centimeterswould have
unit weight. Yet although humans have devoted about 11 percent of its resting
Big Brains and much bigger brains relative to body energy to the brain. For its part, H. erec-
Hungry Hominids weight than do other primates (three tus, which weighed in at 125 to 130
N O S O O N E R H A D humans perfected times larger than expected), the total rest- pounds and had a brain size of some 900
their stride than the next pivotal event in ing energy requirements of the human cubic centimeters, would have earmarked
human evolution— the dramatic en- body are no greater than those of any about 17 percent of its resting energy—
CORNELIA BLIK
largement of the brain— began. Accord- other mammal of the same size. We that is, about 260 out of 1,500 kilocalo-
ing to the fossil record, the australopith- therefore use a much greater share of our ries a day— for the organ.
ecines never became much brainier than daily energy budget to feed our voracious How did such an energetically costly

SAGITTAL CREST NO SAGITTAL CREST
(to anchor
chewing muscles)
MASSIVE
CHEEKBONES MORE DELICATE
(to anchor CHEEKBONES
chewing muscles)
SMALLER, MORE
VERY LARGE, THICKLY THINLY ENAMELED
ENAMELED MOLARS MOLARS
LARGER
INCISORS
SMALL INCISORS AND CANINES
ROBUST AUSTRALOPITHECINES like A. boisei (left) had pronounced adaptations

to eating tough, fibrous plant foods. H. erectus (right), in contrast, evolved to eat
a softer, higher-quality diet—one that most likely featured meat regularly.
brain evolve? One theory, developed by er products). Modern chimps, in com- man family tree that lived alongside
Dean Falk of Florida State University, parison, obtain only 5 to 7 percent of members of our own genus— had espe-
holds that bipedalism enabled hominids their calories from these comestibles. An- cially pronounced adaptations for grind-
to cool their cranial blood, thereby free- imal foods are far denser in calories and ing up fibrous plant foods, including mas-
ing the heat-sensitive brain of the tem- nutrients than most plant foods. For ex- sive, dish-shaped faces; heavily built man-
perature constraints that had kept its size ample, 3.5 ounces of meat provides up- dibles; ridges, or sagittal crests, atop the
in check. I suspect that, as with bipedal- ward of 200 kilocalories. But the same skull for the attachment of powerful
ism, a number of selective factors were amount of fruit provides only 50 to 100 chewing muscles; and huge, thickly
probably at work. But brain expansion kilocalories. And a comparable serving enameled molar teeth. (This is not to say
almost certainly could not have occurred of foliage yields just 10 to 20 kilocalories. that australopithecines never ate meat.
until hominids adopted a diet sufficient- It stands to reason, then, that for early They almost certainly did on occasion,
ly rich in calories and nutrients to meet Homo, acquiring more gray matter meant just as chimps do today.) In contrast, ear-
the associated costs. seeking out more of the energy-dense fare. ly members of the genus Homo, which
Comparative studies of living ani- Fossils, too, indicate that improve- descended from the gracile australopith-
mals support that assertion. Across all ments to dietary quality accompanied ecines, had much smaller faces, more del-
primates, species with bigger brains dine evolutionary brain growth. All australo- icate jaws, smaller molars and no sagit-
on richer foods, and humans are the ex- pithecines had cranial and dental features tal crests— despite being far larger in
treme example of this correlation, boast- built for processing tough, low-quality terms of overall body size than their pre-
ing the largest relative brain size and the plant foods. The later, robust australo- decessors. Together these features suggest
choicest diet [see “Diet and Primate Evo- pithecines—a dead-end branch of the hu- that early Homo was consuming less
lution,” by Katharine Milton; SCIENTIF-
THE AUTHOR
IC AMERICAN, August 1993]. According WILLIAM R. LEONARD is professor of anthropology and co-director of the Laboratory for Hu-
to recent analyses by Loren Cordain of man Biology Research at Northwestern University. He received his Ph.D. in biological an-
Colorado State University, contempo- thropology at the University of Michigan at Ann Arbor in 1987. The author of more than 80
DAVID BRILL
rary hunter-gatherers derive, on average, research articles on nutrition and energetics among contemporary and prehistoric popu-
40 to 60 percent of their dietary energy lations, Leonard has studied indigenous agricultural groups in Ecuador, Bolivia and Peru
from animal foods (meat, milk and oth- and traditional herding populations in central and southern Siberia.

INTO THE FIRE
EATING MORE ANIMAL FOODS is one way of boosting the caloric and nutrient
density of the diet, a shift that appears to have been critical in the evolution
of the human lineage. But might our ancient forebears have improved dietary
quality another way? Richard Wrangham of Harvard University and his
colleagues recently examined the importance of cooking in human evolution.
They showed that cooking not only makes plant foods softer and easier to
chew, it substantially increases their available energy content, particularly
for starchy tubers such as potatoes and manioc. In their raw form, starches
are not readily broken down by the enzymes in the human body. When
heated, however, these complex carbohydrates become more digestible,
thereby yielding more calories.
The researchers propose that Homo erectus was probably the first
hominid to apply fire to food, starting perhaps 1.8 million years ago. They
argue that early cooking of plant foods (especially tubers) enabled this
species to evolve smaller teeth and bigger brains than those of their
predecessors. Additionally, the extra calories allowed H. erectus to start
hunting— an energetically costly activity— more frequently.
From an energetics perspective, this is a logical enough line of reasoning.
What makes the hypothesis difficult to swallow is the archaeological evidence
Wrangham’s team uses to make its case. The authors cite the East African
sites of Koobi Fora and Chesowanja, which date to around 1.6 million and 1.4
million years ago, respectively, to indicate control of fire by H. erectus. These
localities do indeed exhibit evidence of fires, but whether hominids were
responsible for creating or harnessing the flames is a matter of some debate.
The earliest unequivocal manifestations of fire use—stone hearths and
burned animal bones from sites in Europe—are only some 200,000 years old.
Cooking was clearly an innovation that considerably improved the EARLY COOKING of plant foods, especially tubers,
quality of the human diet. But it remains unclear when in our past this enabled brain expansion, argue Richard Wrangham
practice arose. — W.R.L. of Harvard University and his colleagues.
plant material and more animal foods. developing the first hunting-and-gather- played a critical role in enabling that
As to what prompted Homo’s initial ing economy in which game animals be- change. After the initial spurt in brain
shift toward the higher-quality diet nec- came a significant part of the diet and re- growth, diet and brain expansion prob-
essary for brain growth, environmental sources were shared among members of ably interacted synergistically: bigger
change appears to have once more set the the foraging groups. Signs of this behav- brains produced more complex social
stage for evolutionary change. The con- ioral revolution are visible in the archae- behavior, which led to further shifts in
tinued desiccation of the African land- ological record, which shows an increase foraging tactics and improved diet,
scape limited the amount and variety of in animal bones at hominid sites during which in turn fostered additional brain
edible plant foods available to hominids. this period, along with evidence that the evolution.
Those on the line leading to the robust beasts were butchered using stone tools.
australopithecines coped with this prob- These changes in diet and foraging A Movable Feast
lem morphologically, evolving anatomi- behavior did not turn our ancestors into T H E E V O L U T I O N of H. erectus in
cal specializations that enabled them to strict carnivores; however, the addition Africa 1.8 million years ago also marked
subsist on more widely available, diffi- of modest amounts of animal foods to a third turning point in human evolution:
cult-to-chew foods. Homo took a differ- the menu, combined with the sharing of the initial movement of hominids out of
I. DEVORE Anthro-Photo File
ent path. As it turns out, the spread of resources that is typical of hunter-gath- Africa. Until recently, the locations and
grasslands also led to an increase in the erer groups, would have significantly in- ages of known fossil sites suggested that
relative abundance of grazing mammals creased the quality and stability of hom- early Homo stayed put for a few hun-
such as antelope and gazelle, creating op- inid diets. Improved dietary quality dred thousand years before venturing out
portunities for hominids capable of ex- alone cannot explain why hominid of the motherland and slowly fanning
ploiting them. H. erectus did just that, brains grew, but it appears to have out into the rest of the Old World. Ear-

lier work hinted that improvements in
tool technology around 1.4 million years NEANDERTAL HUNTERS
ago— namely, the advent of the Acheu-
lean hand ax—allowed hominids to leave TO RECONSTRUCT what early humans ate, researchers have traditionally studied
Africa. But new discoveries indicate that features on their fossilized teeth and skulls, archaeological remains of food-related
H. erectus hit the ground running, so to activities, and the diets of living humans and apes. Increasingly, however,
speak. Rutgers University geochronolo- investigators have been tapping another source of data: the chemical composition
gist Carl Swisher III and his colleagues of fossil bones. This approach has yielded some especially intriguing findings with
have shown that the earliest H. erectus regard to the Neandertals.
sites outside of Africa, which are in In- Michael Richards, now at the University of Bradford in England, and his colleagues
donesia and the Republic of Georgia, date recently examined isotopes of carbon (13 C) and nitrogen (15 N) in 29,000-year-old
to between 1.8 million and 1.7 million Neandertal bones from Vindija cave in Croatia. The relative proportions of these
years ago. It seems that the first appear- isotopes in the protein part of human bone, known as collagen, directly reflect their
ance of H. erectus and its initial spread proportions in the protein of the individual’s diet. Thus, by comparing the isotopic
from Africa were almost simultaneous. “signatures” of the Neandertal bones to those of other animals living in the same
The impetus behind this newfound environments, the authors were able to determine whether the Neandertals were
wanderlust again appears to be food. deriving the bulk of their protein from plants or from animals.
What an animal eats dictates to a large The analyses show that the Vindija Neandertals had 15N levels comparable to
extent how much territory it needs to those seen in northern carnivores such as foxes and wolves, indicating that they
survive. Carnivorous animals generally obtained almost all their dietary protein from animal foods. Earlier work hinted that
require far bigger home ranges than do inefficient foraging might have been a factor in the subsequent demise of the
herbivores of comparable size because Neandertals. But Richards and his collaborators argue that in order to consume as
they have fewer total calories available to much animal food as they apparently did, the Neandertals had to have been skilled
them per unit area. hunters. These findings are part of a growing body of literature that suggests
Large-bodied and increasingly de- Neandertal subsistence behavior was more complex than previously thought [see
pendent on animal foods, H. erectus “Who Were the Neandertals?” on page 28]. — W.R.L.
most likely needed much more turf than
the smaller, more vegetarian australo-
pithecines did. Using data on contempo-
rary primates and human hunter-gather-
ers as a guide, Robertson, Susan C. An-
tón of Rutgers University and I have
estimated that the larger body size of H.
erectus, combined with a moderate in-
crease in meat consumption, would have
necessitated an eightfold to 10-fold in-
crease in home range size compared with
that of the late australopithecines—
enough, in fact, to account for the abrupt
expansion of the species out of Africa. NEANDERTAL MEALS consisted mostly of meat (from, for example, reindeer), according
to analyses of carbon and nitrogen isotopes in fossilized bone.
Exactly how far beyond the continent that
shift would have taken H. erectus remains
unclear, but migrating animal herds may northern settings today. The Siberian cost of living up further still. Indeed,
have helped lead it to these distant lands. reindeer-herding populations known as whereas a 160-pound American male
As humans moved into more north- the Evenki, which I have studied with Pe- with a typical urban way of life requires
HELLIO & VAN INGEN Photo Researchers, Inc.
ern latitudes, they encountered new di- ter Katzmarzyk of Queen’s University in about 2,600 kilocalories a day, a diminu-
etary challenges. The Neandertals, who Ontario and Victoria A. Galloway of the tive, 125-pound Evenki man needs more
lived during the last ice ages of Europe, University of Toronto, and the Inuit (Es- than 3,000 kilocalories a day to sustain
were among the first humans to inhabit kimo) populations of the Canadian Arc- himself. Using these modern northern
arctic environments, and they almost cer- tic have resting metabolic rates that are populations as benchmarks, Mark
tainly would have needed ample calories about 15 percent higher than those of Sorensen of Northwestern University
to endure under those circumstances. people of similar size living in temperate and I have estimated that Neandertals
Hints at what their energy requirements environments. The energetically expen- most likely would have required as many
might have been come from data on tra- sive activities associated with living in a as 4,000 kilocalories a day to survive.
ditional human populations that live in northern climate ratchet their caloric That they were able to meet these de-

and grains. Similarly, the development of
liquid nutritional supplements and meal
Dmanisi,
Georgia replacement bars is a continuation of the
Longgupo, trend that our ancient ancestors started:
China?
gaining as much nutritional return from
our food in as little volume and with as
little physical effort as possible.
Bahr el Ghazal, Overall, that strategy has evidently
Chad
Hadar, Ethiopia worked: humans are here today and in
record numbers to boot. But perhaps the
Turkana,
Kenya strongest testament to the importance of
energy- and nutrient-rich foods in human
Olduvai Gorge, evolution lies in the observation that so
Tanzania
Laetoli, Tanzania Java, Indonesia many health concerns facing societies
Homo erectus around the globe stem from deviations
Homo habilis
from the energy dynamic that our ances-
Australopithecines
Sterkfontein, Swartkrans, tors established. For children in rural
South Africa South Africa populations of the developing world,
low-quality diets lead to poor physical
AFRICAN EXODUS began as soon as H. erectus evolved, around 1.8 million years ago, probably in part growth and high rates of mortality during
because it needed a larger home range than that of its smaller-bodied predecessors.
early life. In these cases, the foods fed to
mands for as long as they did speaks to be considered tactics for boosting the youngsters during and after weaning are
their skills as foragers [see box on pre- quality of the human diet. Cooking, for often not sufficiently dense in energy and
ceding page]. one, augmented the energy available in nutrients to meet the high nutritional
wild plant foods [see box on page 68]. needs associated with this period of rapid
Modern Quandaries With the advent of agriculture, humans growth and development. Although
JUST AS PRESSURES to improve di- began to manipulate marginal plant spe- these children are typically similar in
etary quality influenced early human cies to increase their productivity, di- length and weight to their U.S. counter-
evolution, so, too, have these factors gestibility and nutritional content— es- parts at birth, they are much shorter and
played a crucial role in the more recent sentially making plants more like animal lighter by the age of three, often resem-
increases in population size. Innovations foods. This kind of tinkering continues bling the smallest 2 to 3 percent of Amer-
such as cooking, agriculture and even as- today, with genetic modification of crop ican children of the same age and sex.
pects of modern food technology can all species to make “better” fruits, vegetables In the industrial world, we are facing
the opposite problem: rates of childhood
VARIOUS DIETS can satisfy human nutritional requirements. Some populations subsist almost entirely and adult obesity are rising because the
on plant foods; others eat mostly animal foods. Although Americans consume less meat than do a
number of the traditionally living people described here, they have on average higher cholesterol
energy-rich foods we crave— notably
levels and higher levels of obesity (as indicated by body mass index) because they consume more those packed with fat and sugar— have
energy than they expend and eat meat that is higher in fat. become widely available and relatively in-
Energy Intake Energy from Energy from Total Blood Body Mass Index
Population (kilocalories/day) Animal Foods Plant Foods Cholesterol (weight/height
(percent) (percent) (milligrams/deciliter) squared)
HUNTER-GATHERERS
!Kung (Botswana) 2,100 33 67 121 19
Inuit (North America) 2,350 96 4 141 24
PASTORALISTS
Turkana (Kenya) 1,411 80 20 186 18
Evenki (Russia) 2,820 41 59 142 22
AGRICULTURALISTS
Quechua (Highland Peru) 2,002 5 95 150 21
LAURIE GRACE (map)
INDUSTRIAL SOCIETIES
U.S. 2,250 23 77 204 26
Note: Energy intake figures reflect the adult average (males and females); blood cholesterol and body mass index (BMI) figures are given for males.
Healthy BMI = 18.5–24.9; overweight = 25.0–29.9; obese = 30 and higher. BMI is weight (kilograms)/height (meters) squared.

expensive. According to recent estimates,
more than half of adult Americans are A DIVERSITY OF DIETS
overweight or obese. Obesity has also ap-
peared in parts of the developing world THE VARIETY OF SUCCESSFUL dietary strategies employed by traditionally living
where it was virtually unknown less than populations provides an important perspective on the ongoing debate about how
a generation ago. This seeming paradox high-protein, low-carbohydrate regimens such as the Atkins diet compare with those
has emerged as people who grew up mal- that underscore complex carbohydrates and fat restriction. The fact that both these
nourished move from rural areas to ur- schemes produce weight loss is not surprising, because both help people shed
ban settings where food is more readily pounds through the same basic mechanism: limiting major sources of calories. When
available. In some sense, obesity and oth- you create an energy deficit— that is, when you consume fewer calories than you
er common diseases of the modern world expend— your body begins burning its fat stores and you lose weight.
are continuations of a tenor that started The larger question about healthy weight-loss or weight-maintenance diets is
millions of years ago. We are victims of whether they create eating patterns that are sustainable over time. On this point it
our own evolutionary success, having de- appears that diets that severely limit large categories of foods (carbohydrates, for
veloped a calorie-packed diet while min- example) are much more difficult to sustain than are moderately restrictive diets. In
imizing the amount of maintenance en- the case of the Atkins-type regimen, there are also concerns about the potential
ergy expended on physical activity. long-term consequences of eating foods derived largely from feedlot animals, which
The magnitude of this imbalance be- tend to contain more fat in general and considerably more saturated fats than do
comes clear when we look at traditional- their free-ranging counterparts.
ly living human populations. Studies of In September 2002 the National Academy of Sciences’s Institute of Medicine put
the Evenki reindeer herders that I have forth new diet and exercise guidelines that mesh well with the ideas presented in
conducted in collaboration with Michael this article. Not only did the institute set broader target ranges for the amounts of
Crawford of the University of Kansas carbohydrates, fat and protein that belong in a healthy diet— in essence,
and Ludmila Osipova of the Russian acknowledging that there are various ways to meet our nutritional needs— the
Academy of Sciences in Novosibirsk in- organization also doubled the recommended amount of moderately intense physical
dicate that the Evenki derive almost half activity to an hour a day. By following these guidelines and balancing what we eat with
their daily calories from meat, more than exercise, we can live more like the Evenki of Siberia and other traditional societies—
2.5 times the amount consumed by the and more like our hominid ancestors. — W.R.L.
average American. Yet when we com-
pare Evenki men with their U.S. peers, diet—an oversimplification embodied by most all animal foods among popula-
they are 20 percent leaner and have cho- the current debate over the relative mer- tions of the Arctic to primarily tubers and
lesterol levels that are 30 percent lower. its of a high-protein, high-fat Atkins-type cereal grains among populations in the
These differences partly reflect the diet or a low-fat one that emphasizes high Andes. Indeed, the hallmarks of hu-
compositions of the diets. Although the complex carbohydrates. This is a funda- man evolution have been the diversity of
Evenki diet is high in meat, it is relative- mentally flawed approach to assessing strategies that we have developed to cre-
ly low in fat (about 20 percent of their di- human nutritional needs. Our species ate diets that meet our distinctive meta-
etary energy comes from fat, compared was not designed to subsist on a single, bolic requirements and the ever increas-
with 35 percent in the average U.S. diet), optimal diet. What is remarkable about ing efficiency with which we extract en-
because free-ranging animals such as rein- human beings is the extraordinary vari- ergy and nutrients from the environment.
deer have less body fat than cattle and ety of what we eat. We have been able to The challenge our modern societies now
other feedlot animals do. The composi- thrive in almost every ecosystem on the face is balancing the calories we consume
tion of the fat is also different in free-rang- earth, consuming diets ranging from al- with the calories we burn.
ing animals, tending to be lower in satu-
rated fats and higher in the polyunsat- MORE TO E XPLORE
urated fatty acids that protect against Evolutionary Perspectives on Human Nutrition: The Influence of Brain and Body Size on Diet
heart disease. More important, however, and Metabolism. William R. Leonard and Marcia L. Robertson in American Journal of Human Biology,
Vol. 6, No. 1, pages 77–88; January 1994.
the Evenki way of life necessitates a much
Rethinking the Energetics of Bipedality. William R. Leonard and Marcia L. Robertson in Current
higher level of energy expenditure. Anthropology, Vol. 38, No.2, pages 304–309; April 1997.
Thus, it is not just changes in diet that Human Biology: An Evolutionary and Biocultural Approach. Edited by Sara Stinson, Barry Bogin,
have created many of our pervasive health Rebecca Huss-Ashmore and Dennis O’Rourke. Wiley-Liss, 2000.
problems but the interaction of shifting Ecology, Health and Lifestyle Change among the Evenki Herders of Siberia. William R. Leonard
diets and changing lifestyles. Too often et al. in Human Biology of Pastoral Populations. Edited by William R. Leonard and Michael H. Crawford.
Cambridge University Press, 2002.
modern health problems are portrayed
An Ecomorphological Model of the Initial Hominid Dispersal from Africa. Susan C. Antón,
as the result of eating “bad” foods that William R. Leonard and Marcia L. Robertson in Journal of Human Evolution, Vol. 43, No. 6, pages
are departures from the natural human 773–785; December 2002.

ADAPTATION
Throughout the world, human skin color has evolved to be
dark enough to prevent sunlight from destroying the nutrient
folate but light enough to foster the production of vitamin D
By Nina G. Jablonski and George Chaplin
Among primates, only humans have

a mostly naked skin that comes in different colors. Geographers and anthro-
pologists have long recognized that the distribution of skin colors among in-
digenous populations is not random: darker peoples tend to be found nearer
the equator, lighter ones closer to the poles. For years, the prevailing theory has
been that darker skins evolved to protect against skin cancer. But a series of dis-
coveries has led us to construct a new framework for understanding the evolu-
tionary basis of variations in human skin color. Recent epidemiological and
physiological evidence suggests to us that the worldwide pattern of human skin
color is the product of natural selection acting to regulate the effects of the sun’s
ultraviolet (UV) radiation on key nutrients crucial to reproductive success.
The evolution of skin pigmentation is linked with that of hairlessness, and
to comprehend both these stories, we need to page back in human history.
Human beings have been evolving as an independent lineage of apes since at
least seven million years ago, when our immediate ancestors diverged from
those of our closest relatives, chimpanzees. Because chimpanzees have
changed less over time than humans have, they can provide an idea of what
human anatomy and physiology must have been like. Chimpanzees’ skin is
light in color and is covered by hair over most of their bodies. Young animals
have pink faces, hands, and feet and become freckled or dark in these areas

only as they are exposed to sun with age. The earliest humans serves the dual purpose of physically and chemically filtering the
almost certainly had a light skin covered with hair. Presumably harmful effects of UV radiation; it absorbs UV rays, causing
hair loss occurred first, then skin color changed. But that leads them to lose energy, and it neutralizes harmful chemicals called
to the question, When did we lose our hair? free radicals that form in the skin after damage by UV radiation.
The skeletons of ancient humans— such as the well-known Anthropologists and biologists have generally reasoned
skeleton of Lucy, which dates to about 3.2 million years ago— that high concentrations of melanin arose in the skin of peo-
give us a good idea of the build and the way of life of our an- ples in tropical areas because it protected them against skin
cestors. The daily activities of Lucy and other hominids that cancer. James E. Cleaver of the University of California at San
lived before about three million years ago appear to have been Francisco, for instance, has shown that people with the disease
similar to those of primates living on the open savannas of xeroderma pigmentosum, in which melanocytes are destroyed
Africa today. They probably spent much of their day foraging by exposure to the sun, suffer from significantly higher than
for food over three to four miles before retiring to the safety normal rates of squamous and basal cell carcinomas, which are
of trees to sleep. usually easily treated. Malignant melanomas are more fre-
By 1.6 million years ago, however, we see evidence that this quently fatal, but they are rare (representing 4 percent of skin
pattern had begun to change dramatically. The famous skele- cancer diagnoses) and tend to strike only light-skinned people.
ton of Turkana Boy— which belonged to the species Homo er- But all skin cancers typically arise later in life, in most cases af-
gaster— is that of a long-legged, striding biped that probably ter the first reproductive years, so they could not have exerted
walked long distances. These more active early humans faced enough evolutionary pressure for skin protection alone to ac-
the problem of staying cool and protecting their brains from count for darker skin colors. Accordingly, we began to ask
overheating. Peter Wheeler of Liverpool John Moores Uni- what role melanin might play in human evolution.
versity has shown that this was accomplished through an in-
crease in the number of sweat glands on the surface of the body The Folate Connection
and a reduction in the covering of body hair. Once rid of most IN 1991 O N E O F U S (Jablonski) ran across what turned out
of their hair, early members of the genus Homo then encoun- to be a critical paper published in 1978 by Richard F. Branda
tered the challenge of protecting their skin from the damaging and John W. Eaton, now at the University of Vermont and the
effects of sunlight, especially UV rays. University of Louisville, respectively. These investigators
showed that light-skinned people who had been exposed to sim-
Built-in Sunscreen ulated strong sunlight had abnormally low levels of the essen-
IN CHIMPANZEES, the skin on the hairless parts of the body tial B vitamin folate in their blood. The scientists also observed
contains cells called melanocytes that are capable of synthesiz- that subjecting human blood serum to the same conditions re-
ing the dark-brown pigment melanin in response to exposure sulted in a 50 percent loss of folate content within one hour.
to UV radiation. When humans became mostly hairless, the The significance of these findings to reproduction— and
ability of the skin to produce melanin assumed new importance. hence evolution— became clear when we learned of research
Melanin is nature’s sunscreen: it is a large organic molecule that being conducted on a major class of birth defects by our col-
leagues at the University of Western Australia. There Fiona J.
Overview/Skin Color Evolution Stanley and Carol Bower had established by the late 1980s that
folate deficiency in pregnant women is related to an increased
■ After losing their hair as an adaptation for keeping cool, risk of neural tube defects such as spina bifida, in which the
early hominids gained pigmented skins. Scientists arches of the spinal vertebrae fail to close around the spinal
initially thought that such pigmentation arose to protect cord. Many research groups throughout the world have since
against skin-cancer-causing ultraviolet (UV) radiation. confirmed this correlation, and efforts to supplement foods
■ Skin cancers tend to arise after reproductive age, with folate (folic acid) and to educate women about the im-
however. An alternative theory suggests that dark skin portance of the nutrient have become widespread.
might have evolved primarily to protect against the We discovered soon afterward that folate is important not
breakdown of folate (folic acid), a nutrient essential for only in preventing neural tube defects but also in a host of oth-
fertility and for fetal development. er processes. Because folate is essential for the synthesis of
■ Skin that is too dark blocks the sunlight necessary for DNA in dividing cells, anything that involves rapid cell prolif-
catalyzing the production of vitamin D, which is crucial eration, such as spermatogenesis (the production of sperm
IRAIDA ICAZA (preceding pages)
for maternal and fetal bones. Accordingly, humans have cells) , requires folate. Male rats and mice with chemically in-
evolved to be light enough to make sufficient vitamin D duced folate deficiency have impaired spermatogenesis and are
yet dark enough to protect their stores of folate. infertile. Although no comparable studies of humans have been
■ As a result of recent human migrations, many people conducted, Wai Yee Wong and his colleagues at the Universi-
now live in areas that receive more (or less) UV radiation ty Medical Center of Nijmegen in the Netherlands have re-
than is appropriate for their skin color. cently reported that folic acid treatment can boost the sperm
counts of men with fertility problems.
74 SCIENTIFIC AMERICAN Updated from the October 2002 issue

SKIN IN THE SUN
THE ULTRAVIOLET (UV) RAYS of the sun are a mixed blessing: by melanocytes protects against DNA damage and folate
they spur the production of vitamin D but destroy folate and breakdown. But keratinocytes must get enough UV rays to
can cause cancer by damaging DNA. Melanin pigment produced make vitamin D. — N.G.J. and G.C.
UVC rays are blocked

from reaching skin by ozone
layer in atmosphere
Hair follicle
Sweat gland UVB rays penetrate
epidermis, prompting
melanocytes to make
EPIDERMIS melanin pigment, which
is packaged in
structures called
melanosomes (detail at
bottom). Melanosomes
are taken up by
DERMIS keratinocytes to shield
their DNA by forming
a nuclear cap
Blood Oil gland

vessels
Folate
(folic acid)
MELANOCYTE
Breakdown products Melanosome

Nuclear cap
UVA rays permeate blood DNA
vessels in the dermis, where they Melanocyte
destroy folate (folic acid)
Keratinocyte HO 7-dehydro-
cholesterol
LIVER
KERATINOCYTE
Blood vessels
KIDNEY
UVB rays that reach keratinocytes convert 1,25-
cholesterol into basic vitamin D, which the liver dihydroxy-
and then the kidneys progressively convert into vitamin D3
KEITH KASNOT
the active form of vitamin D

IRAIDA ICAZA

Such observations led us to hypothesize that dark skin bution of UV radiation on the earth and relate the satellite data
evolved to protect the body’s folate stores from destruction. to the amount of UVB necessary to produce vitamin D.
Our idea was supported by a report published in 1996 by Ar- We found that the earth’s surface could be divided into
gentine pediatrician Pablo Lapunzina, who found that three three vitamin D zones: one comprising the tropics, one the sub-
young and otherwise healthy women whom he had attended tropics and temperate regions, and the last the circumpolar re-
gave birth to infants with neural tube defects after using sun gions north and south of about 45 degrees latitude. In the first,
beds to tan themselves in the early weeks of pregnancy. Our the dosage of UVB throughout the year is high enough that hu-
evidence about the breakdown of folate by UV radiation thus mans have ample opportunity to synthesize vitamin D all year.
supplements what is already known about the harmful (skin- In the second, at least one month during the year has insuffi-
cancer-causing) effects of UV radiation on DNA. cient UVB radiation, and in the third area not enough UVB ar-
rives on average during the entire year to prompt vitamin D
Human Skin on the Move synthesis. This distribution could explain why indigenous peo-
T H E E A R L I E S T M E M B E R S of Homo sapiens, or modern hu- ples in the tropics generally have dark skin, whereas people in
mans, evolved in Africa between 120,000 and 100,000 years the subtropics and temperate regions are lighter-skinned but
ago and had darkly pigmented skin adapted to the conditions have the ability to tan, and those who live in regions near the
of UV radiation and heat that existed near the equator. As mod- poles tend to be very light skinned and burn easily.
ern humans began to venture out of the tropics, however, they One of the most interesting aspects of this investigation was
encountered environments in which they received significantly the examination of groups that did not precisely fit the pre-
less UV radiation during the year. Under these conditions their dicted skin color pattern. An example is the Inuit people of
high concentrations of natural sunscreen probably proved detri- Alaska and northern Canada. The Inuit exhibit skin color that
mental. Dark skin contains so much melanin that very little UV is somewhat darker than would be predicted given the UV lev-
radiation, and specifically very little of the shorter-wavelength els at their latitude. This is probably caused by two factors. The
UVB radiation, can penetrate the skin. Although most of the ef- first is that they are relatively recent inhabitants of these climes,
fects of UVB are harmful, the rays perform one indispensable having migrated to North America only roughly 5,000 years
function: initiating the formation of vitamin D in the skin. Dark- ago. The second is that the traditional diet of the Inuit is ex-
skinned people living in the tropics generally receive sufficient tremely high in foods containing vitamin D, especially fish and
UV radiation during the year for UVB to penetrate the skin and marine mammals. This vitamin D–rich diet offsets the prob-
allow them to make vitamin D. Outside the tropics this is not lem that they would otherwise have with vitamin D synthesis
the case. The solution, across evolutionary time, has been for in their skin at northern latitudes and permits them to remain
migrants to northern latitudes to lose skin pigmentation. more darkly pigmented.
The connection between the evolution of lightly pigmented Our analysis of the potential to synthesize vitamin D allowed
skin and vitamin D synthesis was elaborated in 1967 by W. us to understand another trait related to human skin color:
Farnsworth Loomis of Brandeis University. He established the women in all populations are generally lighter-skinned than men.
importance of vitamin D to reproductive success because of its (Our data show that women tend to be between 3 and 4 percent
role in enabling calcium absorption by the intestines, which lighter than men.) Scientists have often speculated on the rea-
in turn makes possible the normal development of the skeleton sons, and most have argued that the phenomenon stems from
and the maintenance of a healthy immune system. Research led sexual selection—the preference of men for women of lighter col-
by Michael Holick of the Boston University School of Medi- or. We contend that although this is probably part of the story,
cine has, over the past 20 years, further cemented the signifi- it is not the original reason for the sexual difference. Females
cance of vitamin D in development and immunity. His team have significantly greater needs for calcium throughout their re-
also showed that not all sunlight contains enough UVB to stim- productive lives, especially during pregnancy and lactation, and
ulate vitamin D production. In Boston, for instance, which is must be able to make the most of the calcium contained in food.
located at about 42 degrees north latitude, human skin cells be-
gin to produce vitamin D only after mid-March. In the winter- NINA G. JABLONSKI and GEORGE CHAPLIN work at the California
THE AUTHORS
time there isn’t enough UVB to do the job. We realized that this Academy of Sciences in San Francisco, where Jablonski is Irvine
was another piece of evidence essential to the skin color story. Chair and curator of anthropology and Chaplin is a research asso-
During the course of our research in the early 1990s, we ciate in the department of anthropology. Jablonski’s research
sought in vain to find sources of data on actual UV radiation centers on the evolutionary adaptations of monkeys, apes and
levels at the earth’s surface. We were rewarded in 1996, when humans. She is particularly interested in how primates have re-
we contacted Elizabeth Weatherhead of the Cooperative In- sponded to changes over time in the global environment. Chaplin
stitute for Research in Environmental Sciences at the Univer- is a private geographic information systems consultant who spe-
sity of Colorado at Boulder. She shared with us a database of cializes in describing and analyzing geographic trends in biodi-
measurements of UV radiation at the earth’s surface taken by versity. In 2001 he was awarded the Student of the Year prize by
NASA’s Total Ozone Mapping Spectrophotometer satellite be- the Association of Geographic Information in London for his mas-
tween 1978 and 1993. We were then able to model the distri- ter’s thesis on the environmental correlates of skin color.

We propose, therefore, that women tend to be lighter-skinned
SKIN AND MIGRATION than men to allow slightly more UVB rays to penetrate their skin
and thereby increase their ability to produce vitamin D. In ar-
THE SKIN OF PEOPLES who have inhabited particular areas eas of the world that receive a large amount of UV radiation,
for millennia has adapted to allow vitamin D production while women are indeed at the knife’s edge of natural selection, need-
protecting folate stores. The skin tones of more recent ing to maximize the photoprotective function of their skin on the
immigrants will take thousands of years to catch up, putting one hand and the ability to synthesize vitamin D on the other.
light-skinned individuals at risk for skin cancer and dark-skinned
people in danger of vitamin D deficiency. — N.G.J. and G.C. Where Culture and Biology Meet
A S M O D E R N H U M A N S M O V E D throughout the Old World
LONG-TERM RESIDENT RECENT IMMIGRANT
about 100,000 years ago, their skin adapted to the environ-
SOUTHERN AFRICA: LATITUDE ~20–30 O S mental conditions that prevailed in different regions. The
skin color of the indigenous people of Africa has had the
longest time to adapt because anatomically modern humans
first evolved there. The skin color changes that modern humans
underwent as they moved from one continent to another— first
Asia, then Austro-Melanesia, then Europe and, finally, the
Americas— can be reconstructed to some extent. It is important
to remember, however, that those humans had clothing and
shelter to help protect them from the elements. In some places,
Khoisan Zulu: arrived about they also had the ability to harvest foods that were extraordi-
(Hottentot) 1,000 years ago narily rich in vitamin D, as in the case of the Inuit. These two
factors had profound effects on the tempo and degree of skin
AUSTRALIA: LATITUDE ~10–35 O S color evolution in human populations.
Africa is an environmentally heterogeneous continent. A
number of the earliest movements of contemporary humans
outside equatorial Africa were into southern Africa. The de-
scendants of some of these early colonizers, the Khoisan (pre-
PENNY TWEEDIE Corbis (Aborigine); DAVID M C LAIN Aurora (European); ERIC WHEATER Lonely Planet Images (Sudanese);
viously known as Hottentots), are still found in southern Africa
and have significantly lighter skin than indigenous equatorial
Africans do— a clear adaptation to the lower levels of UV ra-
diation that prevail at the southern extremity of the continent.
Aborigine European: ~300 years ago Interestingly, however, human skin color in southern
WAYNE EASTEP Getty Images (Arab); ROGER WOOD Corbis (Bengali); JEREMY HORNER Corbis (Tamil)
Africa is not uniform. Populations of Bantu-language speakers
BANKS OF RED SEA: LATITUDE ~15–30 O N
who live in southern Africa today are far darker than the
Khoisan. We know from the history of this region that Bantu
PETER JOHNSON Corbis (Khoisan); BARBARA BANNISTER Gallo Images/Corbis (Zulu);

speakers migrated into this region recently— probably within
the past 1,000 years— from parts of West Africa near the equa-
tor. The skin color difference between the Khoisan and Bantu
speakers such as the Zulu indicates that the length of time that
a group has inhabited a particular region is important in un-
derstanding why they have the color they do.
Cultural behaviors have probably also strongly influenced
Sudanese Arab: ~2,000 years ago the evolution of skin color in recent human history. This effect
INDIA: LATITUDE ~10–30 O N can be seen in the indigenous peoples who live on the eastern
and western banks of the Red Sea. The tribes on the western
side, which speak so-called Nilo-Hamitic languages, are
thought to have inhabited this region for as long as 6,000 years.
These individuals are distinguished by very darkly pigmented
skin and long, thin bodies with long limbs, which are excellent
biological adaptations for dissipating heat and intense UV ra-
diation. In contrast, modern agricultural and pastoral groups
on the eastern bank of the Red Sea, on the Arabian Peninsula,
have lived there for only about 2,000 years. These earliest Arab
Bengali Tamil: ~100 years ago
people, of European origin, have adapted to very similar envi-

WHO MAKES ENOUGH VITAMIN D?
POPULATIONS THAT LIVE in the
tropics receive enough ultraviolet
light from the sun (top map, brown
and orange) to synthesize vitamin
D all year long. But those that live
at northern or southern latitudes
do not. In the temperate zones
(light-shaded band), people lack
sufficient UV light to make vitamin
D one month of the year; those
nearer the poles (dark-shaded
band) do not get enough UV light
most months for vitamin D
synthesis. The bottom map shows
predicted skin colors for humans
based on UV light levels. In the Old
World, the skin color of indigenous
peoples closely matches
predictions. In the New World,
however, the skin color of long-
term residents is generally lighter
than expected— probably because
of their recent migration and factors
such as diet. — N.G.J. and G.C.
ronmental conditions by almost exclusively cultural means— min D, an insidious problem that manifests itself in high rates
wearing heavy protective clothing and devising portable shade of rickets and other diseases related to vitamin D deficiency.
in the form of tents. (Without such clothing, one would have The ability of skin color to adapt over long periods to the
expected their skin to have begun to darken.) Generally speak- various environments to which humans have moved reflects the
ing, the more recently a group has migrated into an area, the importance of skin color to our survival. But its unstable nature
more extensive its cultural, as opposed to biological, adapta- also makes it one of the least useful characteristics in determin-
tions to the area will be. ing the evolutionary relations between human groups. Early
Western scientists used skin color improperly to delineate hu-
Perils of Recent Migrations man races, but the beauty of science is that it can and does cor-
DESPITE GREAT IMPROVEMENTS in overall human health rect itself. Our current knowledge of the evolution of human
in the past century, some diseases have appeared or reemerged skin indicates that variations in skin color, like most of our
in populations that had previously been little affected by them. physical attributes, can be explained by adaptation to the en-
One of these is skin cancer, especially basal and squamous cell vironment through natural selection. We look ahead to the day
carcinomas, among light-skinned peoples. Another is rickets, when the vestiges of old scientific mistakes will be erased and
brought about by severe vitamin D deficiency, in dark-skinned replaced by a better understanding of human origins and diver-
peoples. Why are we seeing these conditions? sity. Our variation in skin color should be celebrated as one of
As people move from an area with one pattern of UV ra- the most visible manifestations of our evolution as a species.
diation to another region, biological and cultural adaptations
have not been able to keep pace. The light-skinned people of MORE TO E XPLORE
northern European origin who bask in the sun of Florida or The Evolution of Human Skin Coloration. Nina G. Jablonski and George
Chaplin in Journal of Human Evolution, Vol. 39, No. 1, pages 57–106;
northern Australia increasingly pay the price in the form of pre- July 1, 2000.
mature aging of the skin and skin cancers, not to mention the Why Skin Comes in Colors. Blake Edgar in California Wild, Vol. 53, No. 1,
unknown cost in human life of folate depletion. Conversely, a pages 6–7; Winter 2000. The article is also available at
www.calacademy.org/calwild/winter2000/html/horizons.html
number of dark-skinned people of southern Asian and African
SARA CHEN
The Biology of Skin Color: Black and White. Gina Kirchweger in Discover,
origin now living in the northern U.K., northern Europe or the Vol. 22, No. 2, pages 32–33; February 2001. The article is also available
northeastern U.S. suffer from a lack of UV radiation and vita- at www.discover.com/feb__01/featbiology.html

ADAPTATION
the evolution
CREDIT

Human
of
irth The difficulties of

By Karen R. Rosenberg and Wenda R. Trevathan
GIVING BIRTH IN THE TREETOPS is not the nor-

childbirth have mal human way of doing things, but that is exact-
probably challenged ly what Sophia Pedro was forced to do during the
humans and height of the floods that ravaged southern Mozam-
their ancestors for bique in March 2000. Pedro had survived for four
millions of years— days perched high above the raging floodwaters
which means that killed more than 700 people in the region. The
that the modern day after her delivery, television broadcasts and
custom of seeking newspapers all over the world featured images of
assistance during Pedro and her newborn child being plucked from
delivery may the tree during a dramatic helicopter rescue.
have similarly Treetop delivery rooms are unusual for humans
ancient roots but not for other primate species. For millions of
years, primates have secluded themselves in tree-
tops or bushes to give birth. Human beings are the
only primate species that regularly seeks assistance
during labor and delivery. So when and why did
our female ancestors abandon their unassisted and
CREDIT
solitary habit? The answers lie in the dif- head is close to the size of that opening.
ficult and risky nature of human birth. For humans, this tight squeeze is
Many women know from experience complicated by the birth canal’s not be-
that pushing a baby through the birth ing a constant shape in cross section. The
canal is no easy task. It’s the price we pay entrance of the birth canal, where the
for our large brains and intelligence: hu- baby begins its journey, is widest from
mans have exceptionally big heads rela- side to side relative to the mother’s body.
tive to the size of their bodies. Those who Midway through, however, this orienta-
have delved deeper into the subject know tion shifts 90 degrees, and the long axis
that the opening in the human pelvis of the oval extends from the front of the
Pubic bones
through which the baby must pass is lim- mother’s body to her back. This means
ited in size by our upright posture. But that the human infant must negotiate a
only recently have anthropologists begun series of turns as it works its way through Forehead
to realize that the complex twists and the birth canal so that the two parts of its
turns that human babies make as they body with the largest dimensions— the
travel through the birth canal have trou- head and the shoulders— are always Tailbone
bled humans and their ancestors for at aligned with the largest dimension of the
least 100,000 years. Fossil clues also in- birth canal [see illustration at right]. BABY BORN FACING BACKWARD, with the back of
dicate that anatomy, not just our social To understand the birth process from its head against the mother’s pubic bones,
nature, has led human mothers—in con- the mother’s point of view, imagine you makes it difficult for a human female to guide the
trast to our closest primate relatives and are about to give birth. The baby is most infant from the birth canal— the opening in the
almost all other mammals—to ask for likely upside down, facing your side, mother’s pelvis (insets)— without assistance.
help during childbirth. Indeed, this prac- when its head enters the birth canal.
tice of seeking assistance may have been Midway through the canal, however, it the family tree of human ancestors, we
in place when the earliest members of our must turn to face your back, and the would eventually reach a point where
genus, Homo, emerged and may possibly back of its head is pressed against your birth was not so difficult. Although hu-
date back to five million years ago, when pubic bones. At that time, its shoulders mans are more closely related to apes ge-
our ancestors first began to walk upright are oriented side to side. When the baby netically, monkeys may present a better
on a regular basis. exits your body, it is still facing back- model for birth in prehuman primates.
ward, but it will turn its head slightly to One line of reasoning to support this as-
Tight Squeeze the side. This rotation helps to turn the sertion is as follows: Of the primate fos-
T O T E S T O U R T H E O R Y that the prac- baby’s shoulders so that they can also fit sils discovered from the time before the
tice of assisted birth may have been between your pubic bones and tailbone. first known hominids, one possible re-
around for millennia, we considered first To appreciate the close correspondence mote ancestor is Proconsul, a primate fos-
what scientists know about the way a of the maternal and fetal dimensions, sil dated to about 25 million years ago.
primate baby fits through the mother’s consider that the average pelvic opening This tailless creature probably looked like
birth canal. Viewed from above, the in- in human females is 13 centimeters at its an ape, but its skeleton suggests that it
fant’s head is basically an oval, longest largest diameter and 10 centimeters at its moved more like a monkey. Its pelvis,
from the forehead to the back of the smallest. The average infant head is 10 too, was more monkeylike. The heads of
head and narrowest from ear to ear. centimeters from front to back, and the modern monkey infants are typically
Conveniently, the birth canal— the bony shoulders are 12 centimeters across. This about 98 percent the diameter of the
opening in the pelvis through which the journey through a passageway of chang- mother’s birth canal— a situation more
baby must travel to get from the uterus ing cross-sectional shape makes human comparable with that of humans than
to the outside world— is also an oval birth difficult and risky for the vast ma- that of chimps, whose birth canals are
shape. The challenge of birth for many jority of mothers and babies. relatively spacious.
primates is that the size of the infant’s If we retreat far enough back along Despite the monkey infant’s tight
squeeze, its entrance into the world is
KAREN R. ROSENBERG and WENDA R. TREVATHAN bring different perspectives to the study less challenging than that of a human
THE AUTHORS
of human birth. Rosenberg, a paleoanthropologist at the University of Delaware, specializes baby. In contrast to the twisted birth
in pelvic morphology and has studied hominid fossils from Europe, Israel, China and South canal of modern humans, monkeys’
Africa. About 15 years ago she began studying the pelvis as a way to reconstruct the evo- birth canals maintain the same cross-sec-
lution of the birth process. That’s when she met Trevathan, a biological anthropologist at tional shape from entrance to exit. The
NINA FINKEL
New Mexico State University, whose particular interests include childbirth, maternal be- longest diameter of this oval shape is ori-
havior, sexuality, menopause and evolutionary medicine. Both authors have experienced ented front to back, and the broadest
birth firsthand: Rosenberg has two daughters, and Trevathan is trained as a midwife. part of the oval is against the mother’s
82 SCIENTIFIC AMERICAN Updated from the November 2001 issue

back. A monkey infant enters the birth breathing passage for the infant, to re- that human
canal headfirst, with the broad back of move the umbilical cord from around its birth is seldom easy and
its skull against the roomy back of the neck or even to lift the baby up to her rarely unattended. Today virtually
mother’s pelvis and tailbone. That breast. If she tries to accelerate the de- all women in all societies seek assistance
means the baby monkey emerges from livery by grabbing the baby and guiding at delivery. Even among the !Kung of
the birth canal face forward— in other it from the birth canal, she risks bending southern Africa’s Kalahari Desert— who
words, facing the same direction as the its back awkwardly against the natural are well known for viewing solitary birth
mother. curve of its spine. Pulling on a newborn as a cultural ideal— women do not usu-
Firsthand observations of monkey at this angle risks injury to its spinal ally manage to give birth alone until they
deliveries have revealed a great advan- cord, nerves and muscles. have delivered several babies at which
tage in babies’ being born facing for- For contemporary humans, the re- mothers, sisters or other women are pres-
ward. Monkeys give birth squatting on sponse to these challenges is to seek assis- ent. So, though rare exceptions do exist,
their hind legs or crouching on all fours. tance during labor and delivery. Whether assisted birth comes close to being a uni-
As the infant is born, the mother reach- a technology-oriented professional, a lay versal custom in human cultures [see box
es down to guide it out of the birth canal midwife or a family member who is fa- on next page].
and toward her nipples. In many cases, miliar with the birth process, the assis- Knowing this— and believing that
she also wipes mucus from the baby’s tant can help the human mother do all this practice is driven by the difficulty
mouth and nose to aid its breathing. In- the things the monkey mother does by and risk that accompany human birth—
fants are strong enough at birth to take herself. The assistant can also compen- we began to think that midwifery is not
part in their own deliveries. Once their sate for the limited motor abilities of the unique to contemporary humans but in-
hands are free, they can grab their moth- relatively helpless human infant. The ad- stead has its roots deep in our ancestry.
er’s body and pull themselves out. vantages of even simple forms of assis- Our analysis of the birth process through-
If human babies were also born face tance have reduced maternal and infant out human evolution has led us to sug-
forward, their mothers would have a mortality throughout history. gest that the practice of midwifery might
much easier time. Instead the evolution- have appeared as early as five million
ary modifications of the human pelvis Assisted Birth years ago, when bipedalism constricted
that enabled hominids to walk upright O F C O U R S E , O U R A N C E S T O R S and the size and shape of the pelvis and birth
necessitate that most infants exit the even women today can and do give birth canal.
birth canal with the back of their heads alone successfully. Many fictional ac- A behavior pattern as complex as
against the pubic bones, facing in the op- counts portray stalwart peasant women midwifery obviously does not fossilize,
posite direction as the mother (in a posi- giving birth alone in the fields, perhaps but pelvic bones do. The tight fit between
tion obstetricians call “occiput anteri- most famously in the novel The Good the infant’s head and the mother’s birth
or”). For this reason, it is difficult for the Earth, by Pearl S. Buck. Such images canal in humans means that the mecha-
laboring human mother—whether squat- give the impression that delivering ba- nism of birth can be reconstructed if we
ting, sitting, or lying on her back— to bies is easy. But anthropologists who know the relative sizes of each. Pelvic
reach down and guide the baby as it have studied childbirth in cultures anatomy is now fairly well known from
emerges. This configuration also great- around the world report that these per- most time periods in the human fossil
ly inhibits the mother’s ability to clear a ceptions are highly romanticized and record, and we can estimate infant brain

and skull size based on our extensive walking on two legs would have con- line from the mother’s belly to her back.
knowledge of adult skull sizes. (The del- stricted the maximum size of the pelvis This starting position would have meant
icate skulls of infants are not commonly and birth canal in similar ways among that the shoulders probably also had to
found preserved until the point when hu- related species. turn sideways to squeeze through the
mans began to bury their dead about The anatomy of the female pelvis birth canal.
100,000 years ago.) Knowing the size from this time period is well known This simple rotation could have in-
and shape of the skulls and pelvises has from two complete fossils. Anthropolo- troduced a kind of difficulty in australo-
also helped us and other researchers to gists unearthed the first (known as Sts 14 pithecine deliveries that no other known
understand whether infants were born and presumed to be 2.5 million years primate species had ever experienced.
facing forward or backward relative to old) in Sterkfontein, a site in the Trans- Depending on which way the baby’s
their mothers— in turn revealing how vaal region of South Africa. The second shoulders turned, its head could have ex-
challenging the birth might have been. is best known as Lucy, a fossil discov- ited the birth canal facing either forward
ered in the Hadar region of Ethiopia and or backward relative to the mother. Be-
Walking on Two Legs dated at just over three million years old. cause the australopithecine birth canal is
I N M O D E R N H U M A N S , both bipedal- Based on these specimens and on esti- a symmetrical opening of unchanging
ism and enlarged brains constrain birth mates of newborns’ head size, C. Owen shape, the baby could have just as easi-
in important ways, but the first funda- Lovejoy of Kent State University and ly turned its shoulders toward the front
mental shift away from a nonhuman pri- Robert G. Tague of Louisiana State Uni- or back of its body, giving it about a
mate way of birth came about because versity concluded in the mid-1980s that 50–50 chance of emerging in the easier,
of bipedalism alone. This unique way of birth in early hominids was unlike that face-forward position. If the infant were
walking appeared in early human an- known for any living species of primate. born facing backward, the australopith-
cestors of the genus Australopithecus at The shape of the australopithecine ecine mother— like modern human
least four million years ago [see “Evolu- birth canal is a flattened oval with the mothers— may well have benefited from
tion of Human Walking,” by C. Owen greatest dimension from side to side at some kind of assistance.
Lovejoy; Scientific American, No- both the entrance and exit. This shape ap-
vember 1988]. Despite their upright pos- pears to require a birth pattern different Growing Bigger Brains
ture, australopithecines typically stood from that of monkeys, apes or modern I F B I P E D A L I S M A L O N E did not intro-
no more than four feet tall, and their humans. The head would not have rotat- duce into the process of childbirth
brains were not much bigger than those ed within the birth canal, but we think enough difficulty for mothers to benefit
of living chimpanzees. Recent evidence that in order for the shoulders to fit from assistance, then the expanding size
has called into question which of the sev- through, the baby might have had to turn of the hominid brain certainly did. The
eral australopithecine species were part its head once it emerged. In other words, most significant expansion in adult and
of the lineage that led to Homo. Under- if the baby’s head entered the birth canal infant brain size evolved subsequent to
standing the way any of them gave birth facing the side of the mother’s body, its the australopithecines, particularly in
is still important, however, because shoulders would have been oriented in a the genus Homo. Fossil remains of the
Childbirth across Cultures

THE COMPLICATED CONFIGURATION of the human birth canal is such that laboring
women and their babies benefit— by lower rates of mortality, injury and anxiety—
from the assistance of others. This evolutionary reality helps to explain why
attended birth is a near universal feature of human cultures. Individual women
throughout history have given birth alone in certain circumstances, of course.
But much more common is the attendance of familiar friends and relatives, most
of whom are women. (Men may be variously forbidden, tolerated, welcomed or
even required at birth.) In Western societies, where women usually give birth in
the presence of strangers, recent research on birth practices has also shown that
a doula— a person who provides social and emotional support to a woman in
SQUATTING is one of the most typical positions for
labor— reduces the rate of complications. women to give birth in non-Western cultures.
DANNY LEHMAN Corbis
In many societies, a woman may not be recognized as an adult until she has
had a baby. The preferred location of the delivery is often specified, as are the positions that the laboring women assume. The
typical expectation in Western culture is that women should give birth lying flat on their backs on a bed, but in the rest of the world
the most prevalent position for the delivery is upright— sitting, squatting or, in some cases, standing. — K.R.R. and W.R.T.

pelvis of early Homo are quite rare, and that took place from two million to BABY BORN FACING FORWARD makes it possible
the best-preserved specimen, the 1.6- 100,000 years ago. for a monkey mother to reach down and
carefully guide the infant out of the birth canal.
million-year-old Nariokotome fossil Fossils that span the past 300,000
She can also wipe mucus from the baby’s face
from Kenya, is an adolescent often re- years of human evolution support the to assist its breathing.
ferred to as Turkana Boy. Researchers connection between the expansion of
have estimated that the boy’s adult rel- brain size and changes in pelvic anatomy. desire for companionship and security.
atives probably had brains about twice In the past 20 years, scientists have un- Psychiatrists have argued that natur-
as large as those of australopithecines covered three pelvic fossils of archaic al selection might have favored such
but still only two thirds the size of mod- Homo sapiens: a male from Sima de los emotions— also common during illness
ern human brains. Huesos in Sierra Atapuerca, Spain (more and injury— because they led individuals
By reconstructing the shape of the than 200,000 years old); a female from who experienced them to seek the pro-
boy’s pelvis from fragments, Christo- Jinniushan, China (280,000 years old); tection of companions, which would
pher B. Ruff of Johns Hopkins Universi- and the male Kebara Neandertal—which have given them a better chance of sur-
ty and Alan Walker of Pennsylvania is also an archaic H. sapiens—from Israel viving [see “Evolution and the Origins
State University have estimated what he (about 60,000 years old). These speci- of Disease,” by Randolph M. Nesse and
would have looked like had he reached mens all have the twisted pelvic openings George C. Williams; Scientific Amer-
adulthood. Using predictable differences characteristic of modern humans, which ican, November 1998]. The offspring of
between male and female pelvises in suggests that their large-brained babies the survivors would then also have an
more recent hominid species, they could would most likely have had to rotate the enhanced tendency to experience such
also infer what a female of that species head and shoulders within the birth emotions during times of pain or dis-
would have looked like and could esti- canal and would thus have emerged fac- ease. Taking into consideration the evo-
mate the shape of the birth canal. That ing away from the mother— a major lutionary advantage that fear and anxi-
shape turns out to be a flattened oval challenge that human mothers face in de- ety impart, it is no surprise that women
similar to that of the australopithecines. livering their babies safely. commonly experience these emotions
Based on these reconstructions, the re- The triple challenge of big-brained during labor and delivery.
searchers determined that Turkana infants, a pelvis designed for walking up- Modern women giving birth have a
Boy’s kin probably had a birth mecha- right, and a rotational delivery in which dual evolutionary legacy: the need for
nism like that seen in australopithecines. the baby emerges facing backward is not physical as well as emotional support.
In recent years, scientists have been merely a contemporary circumstance. For When Sophia Pedro gave birth in a tree
testing an important hypothesis that fol- this reason, we suggest that natural selec- surrounded by raging floodwaters, she
lows from Ruff and Walker’s assertion: tion long ago favored the behavior of may have had both kinds of assistance.
the pelvic anatomy of early Homo may seeking assistance during birth because In an interview several months after her
have limited the growth of the human such help compensated for these difficul- helicopter rescue, she told reporters that
brain until the evolutionary point at ties. Mothers probably did not seek as- her mother-in-law, who was also in the
which the birth canal expanded enough sistance solely because they predicted the tree, helped her during delivery. Desire
to allow a larger infant head to pass. This risk that childbirth poses, however. Pain, for this kind of support, it appears, may
assertion implies that bigger brains and fear and anxiety more likely drove their well be as ancient as humanity itself.
roomier pelvises were linked from an
evolutionary perspective. Individuals who MORE TO E XPLORE
displayed both characteristics were more Human Birth: An Evolutionary Perspective. Wenda R. Trevathan. Aldine de Gruyter, 1987.
successful at giving birth to offspring Birth as an American Rite of Passage. Robbie Davis-Floyd. University of California Press, 1993.
who survived to pass on the traits. These Bipedalism and Human Birth: The Obstetrical Dilemma Revisited. Karen R. Rosenberg and
NINA FINKEL
changes in pelvic anatomy, accompanied Wenda R. Trevathan in Evolutionary Anthropology, Vol. 4, No. 5, pages 161–168; 1996.
by assisted birth, may have allowed the On Fertile Ground: A Natural History of Human Reproduction. Peter T. Ellison. Harvard University
dramatic increase in human brain size Press, 2001.

ADAPTATION
Clear evidence
of cannibalism in
the human fossil
Once
record has been rare,
but it is now becoming Were
CAN
apparent that the
practice is deeply
rooted in our history
BY TIM D. WHITE
NEANDERTAL CRANIUM from the Krapina
rock-shelter in Croatia. Physical anthropologists
and archaeologists have recently determined
that this specimen and hundreds of other
skeletal remains at this site attest to
cannibalism. This cranium was smashed so
the brain could be removed and consumed.
NIBALS
It can shock, disgust and fascinate in equal measure,
whether through tales of starved pio- methods. In the past several years, the to inherit their qualities or honor their
neers or airplane crash survivors eating results of their studies have finally pro- memory. And pathological cannibalism
the deceased among them or accounts of vided convincing evidence of prehistoric is generally reserved for criminals who
rituals in Papua New Guinea. It is the cannibalism. consume their victims or, more often,
stuff of headlines and horror films, Human cannibalism has long in- for fictional characters such as Hannibal
drawing people in and mesmerizing trigued anthropologists, and they have Lecter in The Silence of the Lambs.
them despite their aversion. Cannibal- worked for decades to classify the phe- Despite these distinctions, however,
ism represents the ultimate taboo for nomenon. Some divide the behavior ac- most anthropologists simply equate the
many in Western societies— something cording to the affiliation of the con- term “cannibalism” with the regular,
to relegate to other cultures, other times, sumed. Thus, endocannibalism refers to culturally encouraged consumption of
other places. Yet the understanding of the consumption of individuals within a human flesh. In the age of ethnographic
cannibalism derived from the past few group, exocannibalism indicates the exploration—which lasted from the time
centuries of anthropological investiga- consumption of outsiders, and autocan- of Greek historian Herodotus in about
tion has been too unclear and incom- nibalism covers everything from nail bit- 400 B.C. to the early 20th century— the
plete to allow either a categorical rejec- ing to torture-induced self-consumption. non-Western world and its inhabitants
tion of the practice or a fuller apprecia- In addition, anthropologists have come were scrutinized by travelers, mission-
tion of when, where and why it might up with classifications to describe per- aries, military personnel and anthropol-
have taken place. ceived or known motivations. Survival ogists. These observers told tales of hu-
New scientific evidence is now bring- cannibalism is driven by starvation. His- man cannibalism in different places, from
ing to light the truth about cannibalism. torically documented cases include the Mesoamerica to the Pacific islands to
It has become obvious that long before Donner Party— whose members were central Africa.
the invention of metals, before Egypt’s trapped during the harsh winter of Controversy has often accompanied DAVID BRILL (preceding pages); TIM D. WHITE (opposite page)
pyramids were built, before the origins 1846–47 in the Sierra Nevada— and these claims. Anthropologists partici-
of agriculture, before the explosion of people marooned in the Andes or the pated in only the last few waves of these
Upper Paleolithic cave art, cannibalism Arctic with no other food. In contrast, cultural contacts— those that began in
could be found among many different ritual cannibalism occurs when mem- the late 1800s. As a result, many of the
peoples— as well as among many of our bers of a family or community consume historical accounts of cannibalism have
ancestors. Broken and scattered human their dead during funerary rites in order come to be viewed skeptically.
bones, in some cases thousands of them,
THE AUTHOR
have been discovered from the prehis- TIM D. WHITE is co-director of the Laboratory for Human Evolutionary Studies of the Muse-
toric pueblos of the American Southwest um of Vertebrate Zoology at the University of California, Berkeley. He is also a professor in
to the islands of the Pacific. The osteol- Berkeley’s department of integrative biology and a member of the National Academy of Sci-
ogists and archaeologists studying these ences. White co-directs the Middle Awash research project in Ethiopia. His research interests
ancient occurrences are using increas- are human paleontology, Paleolithic archaeology, and the interpretation of bone modifica-
ingly sophisticated analytical tools and tion in contexts ranging from prehistoric archaeology to contemporary forensic situations.
88 SCIENTIFIC AMERICAN Updated from the August 2001 issue

In 1979 anthropologist William Arens critical assessment of these conclusions
of the State University of New York at appeared. Archaeologist Lewis Binford’s
Stony Brook extended this theme by re- book Bones: Ancient Men and Modern
viewing the ethnographic record of can- Myths argued that claims for early hom-
nibalism in his book The Man-Eating inid cannibalism were unsound. He built
Myth. Arens concluded that accounts of on the work of other prehistorians con-
cannibalism among people from the cerned with the composition, context
Aztec to the Maori to the Zulu were ei- and modifications of Paleolithic bone as-
ther false or inadequately documented. semblages. Binford emphasized the need
His skeptical assertion has subsequently to draw accurate inferences about past
been seriously questioned, yet he none- behaviors by grounding knowledge of
theless succeeded in identifying a signif- the past on experiment and observation
icant gulf between these stories and evi- in the present. His influential work cou-
dence of cannibalism: “Anthropology pled skepticism with a plea for meth-
has not maintained the usual standards odological rigor in studies of prehistoric
of documentation and intellectual rigor cannibalism.
expected when other topics are being
considered. Instead, it has chosen un- Standards of Evidence
critically to lend its support to the col- IT WOULD BE HELPFUL if we could
lective representations and thinly dis- turn to modern-day cannibals with our
guised prejudices of western culture questions, but such opportunities have
about others.” largely disappeared. So today’s study of
The anthropologists whom Arens this intriguing behavior must be accom-
was criticizing had not limited them- plished through a historical science. Ar-
selves to contemporary peoples. Some chaeology has therefore become the pri-
had projected their prejudices even mary means of investigating the exis-
more deeply— into the archaeological tence and extent of human cannibalism.
record. Interpretations of cannibalism One of the challenges facing archae-
inevitably followed many discoveries of ologists, however, is the amazing variety
prehistoric remains. In 1871 American of ways in which people dispose of their
author Mark Twain weighed in on the dead. Bodies may be buried, burned,
subject in an essay later published in placed on scaffolding, set adrift, put in
Life as I Find It: “Here is a pile of bones tree trunks or fed to scavengers. Bones
of primeval man and beast all mixed to- may be disinterred, washed, painted,
gether, with no more damning evidence buried in bundles or scattered on stones.
that the man ate the bears than that the In parts of Tibet, future archaeologists
bears ate the man— yet paleontology will have difficulty recognizing any mor-
holds a coroner’s inquest in the fifth ge- tuary practice at all. There most corpses
ologic period on an ‘unpleasantness’ are dismembered and fed to vultures and
which transpired in the quaternary, and other carnivores. The bones are then col-
calmly lays it on the MAN, and then lected, ground into powder, mixed with
CRUSHING adds to it what purports to be evidence barley and flour and again fed to vul-
Many different types of damage can be of CANNIBALISM. I ask the candid read- tures. Given the various fates of bones
seen on bones left by human cannibals. er, Does not this look like taking ad- and bodies, distinguishing cannibalism
When this damage is identical to that vantage of a gentleman who has been from other mortuary practices can be
seen on animal bones at the same sites, dead two million years. . . . ” quite tricky.
archaeologists infer that the human In the century after Twain’s remarks, Scientists have thus set the standard
remains were processed in the same archaeologists and physical anthropolo- for recognizing ancient cannibalism
manner and for the same reason: for gists described the hominids Australo- very high. They confirm the activity
consumption. In these metatarsal (foot) pithecus africanus, Homo erectus and H. when the processing patterns seen on
bones from Mancos Canyon in Colorado, neanderthalensis as cannibalistic. Ac- human remains match those seen on the
the spongy tissues at the ends were cording to some views, human prehisto- bones of other animals consumed for
crushed so that fat could be removed. ry from about three million years ago un- food. Archaeologists have long argued
(All the bones on the following pages are til very recently was rife with cannibalism. for such a comparison between human
from the same Anasazi site in Mancos.) But in the early 1980s an important and faunal remains at a site. They rea-
son that damage to animal bones and the same culture— and checked against dry, mostly intact skulls were then han-
their arrangement can clearly show that predictions embedded in ethnohistorical dled extensively, often creating a polish
the animals had been slaughtered and accounts. on their projecting parts. They were
eaten for food. And when human re- This comparative system of deter- sometimes painted and even mounted
mains are unearthed in similar cultural mining cannibalism emphasizes multiple on poles for display and worship. Soft
contexts, with similar patterns of dam- lines of osteological damage and con- tissue, including brain matter, was eaten
age, discard and preservation, they may textual evidence. And, as noted earlier, at the beginning of this process; thus, the
reasonably be interpreted as evidence of it sets the standard for recognizing can- practice would be identified as ritual
cannibalism. nibalism very high. With this approach, cannibalism. If such skulls were en-
When one mammal eats another, it for instance, the presence of cut marks countered in an archaeological context
usually leaves a record of its activities in on bones would not by themselves be without modern informants describing
the form of modifications to the con- considered evidence of cannibalism. For the cannibalism, they would not consti-
sumed animal’s skeleton. During life, example, an American Civil War ceme- tute direct evidence for cannibalism un-
varying amounts of soft tissue, much of tery would contain skeletal remains with der the stringent criteria that my col-
it with nutritive value, cover mammali- cut marks made by swords and bayo- leagues and I advocate.
an bones. When the tissue is removed nets. Medical school cadavers are dis- Nevertheless, adoption of these stan-
and prepared, the bones often retain a sected and their bones cut-marked. dards of evidence has led us to some
record of this processing in the form of With the threshold set so conserva- clear determinations in other, older sit-
gnawing marks and fractures. When hu- tively, most instances of past cannibal- uations. The best indication of prehis-
mans eat other animals, however, they ism will necessarily go unrecognized. toric cannibalism now comes from the
mark bones with more than just their A practice from Papua New Guinea, archaeological record of the American
teeth. They process carcasses with tools where cannibalism was recorded ethno- Southwest, where archaeologists have
of stone or metal. In so doing, they leave graphically, illustrates this point. There interpreted dozens of assemblages of hu-
imprints of their presence and actions in skulls of the deceased were carefully man remains. Compelling evidence has
the form of scars on the bones. These cleaned and the brains removed. The also been found in Neolithic and Bronze
One of the challenges facing archaeologists is the amazing

variety of ways in which people dispose of their dead.
same imprints can be seen on butchered
human skeletal remains.
The key to recognizing human can-
nibalism is to identify the patterns of CHOPPING
processing— that is, the cut marks, ham- Hack marks visible on the left side
mering damage, fractures or burns seen of this fragment of a human tibia
on the remains—as well as the survival of are testament to the removal of
different bones and parts of bones. Nu- muscle and tendon. Tools were also
tritionally valuable tissues, such as brains used to make finer slices, to remove
and marrow, reside within the bones and tissue or to sever heads from
can be removed only with forceful ham- bodies. Archaeologists have to be
mering— and such forced entry leaves re- careful in their interpretations,
vealing patterns of bone damage. When however, because humans process
human bones from archaeological sites their dead in many ways; not all
show patterns of damage uniquely slice or hack marks indicate
linked to butchery by other humans, the cannibalism.
inference of cannibalism is strengthened.
Judging which patterns are consistent
with dietary butchery can be based on
the associated archaeological record—
particularly the nonhuman food-animal
remains discovered in sites formed by
Age Europe. Even Europe’s earliest hom- In the past few years, yet another site
inid site has yielded convincing evidence has offered evidence. On the banks of
of cannibalism. the Rhône River in southeastern France,
Alban Defleur of the University of the
Early European Cannibals Mediterranean at Marseilles has been
T H E M O S T I M P O R T A N T paleoan- excavating the cave of Moula-Guercy
thropological site in Europe lies in for more than a decade. Neandertals oc-
northern Spain, in the foothills of the cupied this small cave 100,000 years
Sierra de Atapuerca. The oldest known ago. In one layer the team unearthed the
section so far is the Gran Dolina, cur- remains of at least six Neandertals, rang-
rently under excavation. The team ing in age from six years to adult. De-
working there has recovered evidence of fleur’s meticulous excavation and recov-
occupation some 800,000 years ago by ery standards have yielded data every bit
what may prove to be a new species of the equivalent of a modern forensic
human ancestor, H. antecessor. The crime scene investigation. Each fragment
hominid bones were discovered in one of fauna and Neandertal bone, each
horizon of the cave’s sediment, inter- macrobotanical clue, each stone tool has
mingled with stone tools and the re- been precisely plotted three-dimension-
mains of prehistoric game animals such ally. This care has allowed an under-
as deer, bison and rhinoceros. The hom- standing of how the bones were spread
inid remains consist of 92 fragments around a hearth that has been cold for
from six individuals. They bear unmis- 1,000 centuries.
takable traces of butchery with stone Microscopic analysis of the Nean-
tools, including the skinning and re- dertal bone fragments and the faunal re-
moval of flesh and the processing of the mains has led to the same conclusion
braincase and the long bones for mar- that Spanish workers at the Gran Dolina
row. This pattern of butchery matches site have drawn: cannibalism was prac-
that seen on the nearby animal bones, ticed by some Paleolithic Europeans. De-
providing the earliest evidence of homi- termining how often it was practiced
nid cannibalism. and under what conditions represents a
Cannibalism among Europe’s much far more difficult challenge. Neverthe-
younger Neandertals— who lived be- less, the frequency is striking. We know
tween 35,000 and 150,000 years ago— of just one very early European site with
has been debated since the late 1800s, hominid remains, and those were canni-
when the great Croatian paleoanthropol- balized. The two Croatian Neandertal
ogist Dragutin Gorjanovi č-Kramberger sites are separated by hundreds of gen-
found the broken, cut-marked and scat- erations, yet analyses suggest that can-
tered remains of more than 20 Neander- nibalism was practiced at both. And re-
tals entombed in the sands of the Krapina cently a Neandertal site in France was
rock-shelter. Unfortunately, these soft shown to support the same interpreta-
fossil bones were roughly extracted (by tion. These findings are built on exacting
today’s standards) and then covered with standards of evidence. Because of this,
thick layers of preservative, which ob- most paleoanthropologists these days
scured evidence of processing and made are asking, “Why cannibalism?” rather
interpretation exceedingly difficult. Some than “Was this cannibalism?” BURNING
workers believe that the Krapina bones Similarly, discoveries at much The dark and damaged areas on these
TIM D. WHITE (this and opposite page)
show clear signs of cannibalism; others younger sites in the American Southwest four mastoid regions— that is, the hard
have attributed the patterns of damage to have altered the way anthropologists bump behind each ear— indicate that
rocks falling from the cave’s ceiling, to think of Anasazi culture in this area. these human skulls were roasted.
carnivore chewing or to some form of Corn agriculturists have inhabited the Because the mastoid region is not
burial. But recent analysis of the bones Four Corners region for centuries, build- covered by much muscle or other tissue,
from Krapina and from another Croatian ing their pueblos and spectacular cliff damage from burning was often more
cave, Vindija—which has younger Nean- dwellings and leaving one of the richest intense in this area than on other parts of
dertal and animal remains—indicates that and most fine-grained archaeological cranial bone. Burning patterns therefore
cannibalism was practiced at both sites. records on earth. Christy G. Turner II of provide clues about culinary practices.
www.sciam.com
Arizona State University conducted pio-
Historical Accounts neering work on unusual sets of broken
and burned human skeletal remains
ETHNOHISTORICAL REPORTS from Anasazi sites in Arizona, New
of cannibalism have been Mexico and Colorado in the 1960s and
recorded for centuries in 1970s. He saw a pattern suggestive of
many corners of the globe. cannibalism: site after site containing
Although some involve human remains with the telltale signs.
well-documented accounts Yet little in the history of the area’s more
by eyewitnesses— such as recent Puebloan peoples suggested that
the Donner Party cannibalism was a widespread practice,
expedition— other and some modern tribes who claim de-
accounts by explorers, scent from the Anasazi have found the
missionaries, travelers and idea disturbing.
soldiers often lack The vast majority of Anasazi burials
credibility. For example, involve whole, articulated skeletons fre-
these two artists’ portraits quently accompanied by decorated ce-
depict cannibalism ramic vessels that have become a fa-
catalyzed by starvation in vorite target of pot hunters in this area.
China in the late 1800s But, as Turner recorded, several dozen
and a European view of sites had fragmented, often burned hu-
cannibalism in the New man remains, and a larger pattern began
World (based on a woodcut to emerge. Over the past three decades
from 1497). Such ethno- the total number of human bone speci-
historical accounts do not mens from these sites has grown to tens
carry the weight of of thousands, representing dozens of in-
archaeological and dividuals spread across 800 years of pre-
forensic evidence. history and tens of thousands of square
They may, however, serve kilometers of the American Southwest.
as rich sources of testable The assemblage that I analyzed in 1992
hypotheses, guiding future from an Anasazi site in the Mancos
archaeological excavations. Canyon of southwestern Colorado, for
instance, contained 2,106 pieces of bone
from at least 29 Native American men,
women and children.
These assemblages have been found
in settlements ranging from small pueb-
HULTON-DEUTSCH COLLECTION Corbis (top); LEONARD DE SELVA Corbis (bottom)

los to large towns and were often con-
temporaneous with the abandonment of
the dwellings. The bones frequently
show evidence of roasting before the
flesh was removed. They invariably in-
dicate that people extracted the brain
and cracked the limb bones for marrow
after removing the muscle tissue. And
some of the long bone splinters even
show end polishing, a phenomenon as-
sociated with cooking in ceramic vessels.
The bone fragments from Mancos re-
vealed modifications that matched the
marks left by Anasazi processing of
game animals such as deer and bighorn
sheep. The osteological evidence clearly
demonstrated that humans were skinned
and roasted, their muscles cut away,

their joints severed, their long bones bro-
ken on anvils with hammerstones, their HAMMERING
spongy bones crushed and the fragments It is clear from the
circulated in ceramic vessels. But articles archaeological record
outlining the results have proved con- that meat— fat or muscle
troversial. Opposition has sometimes or other tissue— on the
seemed motivated more by politics than bone was not the only
by science. Many practicing anthropol- part of the body that was
ogists believe that scientific findings consumed. Braincases
should defer to social sensitivities. For were broken open, and
such anthropologists, cannibalism is so marrow was often
culturally delicate, so politically incor- removed from long bones.
rect, that they find any evidence for it im- In these two examples,
possible to swallow. stone hammers split the
The most compelling evidence in upper arm bones
support of human cannibalism at the lengthwise, exposing
various Anasazi sites was published in the marrow.
2000 by Richard A. Marlar of the Uni-
versity of Colorado School of Medicine
and his colleagues. The workers exca-
vated three Anasazi pit dwellings dating
to approximately A.D. 1150 at a site
called Cowboy Wash near Mesa Verde
It remains much more difficult to establish why

cannibalism took place than to establish that it did.
in southwestern Colorado. The same strong additional support for numerous nibalism presented a way to get through
pattern of findings that had been docu- osteological and archaeological findings the lean times or a satisfying way to get
mented at other sites, such as Mancos, across the Southwest. rid of outsiders—requires knowledge not
was present: disarticulated, broken, scat- yet available to archaeologists. Even in
tered human bones in nonburial con- Understanding Cannibalism the case of the Anasazi, who have been
texts. Excellent preservation, careful ex- I T R E M A I N S M U C H more challenging well studied, it is impossible to determine
cavation and thoughtful sampling pro- to establish why cannibalism took place whether cannibalism resulted from star-
vided a chemical dimension to the than to establish that it did. People usual- vation or was rooted in religious beliefs,
analysis and, finally, direct evidence of ly eat because they are hungry, and most or was some combination of these and
human cannibalism. prehistoric cannibals were therefore other things. What is becoming clear
Marlar and his colleagues discovered probably hungry. But discerning more through the refinement of the science of
residues of human myoglobin— a pro- than that— such as whether the taste of archaeology, however, is that cannibal-
tein present in heart and skeletal mus- human flesh was pleasing or whether can- ism is part of our collective past.
cle— on a ceramic vessel, suggesting that
human flesh had been cooked in the pot. MORE TO E XPLORE
An unburned human coprolite, or an- Prehistoric Cannibalism at Mancos 5MTUMR-2346. T. D. White. Princeton University Press, 1992.
cient feces, found in the fireplace of one Does Man Eat Man? Inside the Great Cannibalism Controversy. L. Osborne in
of the abandoned dwellings also tested Lingua Franca, Vol. 7, No. 4, pages 28–38; April/May 1997.
positive for human myoglobin. Thus, Fijian Cannibalism: Osteological Evidence from Navatu. D. DeGusta in
American Journal of Physical Anthropology, Vol. 110, pages 215–241; October 1999.
osteological, archaeological and bio-
chemical data indicate that prehistoric Neanderthal Cannibalism at Moula-Guercy, Ardèche, France. A. Defleur, T. D. White,
P. Valensi, L. Slimak and E. Crégut-Bonnoure in Science, Vol. 286, pages 128–131; October 1, 1999.
TIM D. WHITE
cannibalism occurred at Cowboy Wash.

Biochemical Evidence of Cannibalism at a Prehistoric Puebloan Site in
The biochemical data for processing and Southwestern Colorado. R. A. Marlar, B. L. Leonard, B. R. Billman, P. M. Lambert and J. E. Marler
consumption of human tissue offer in Nature, Vol. 407, pages 74–78; September 7, 2000.

FAST-FORWARD
if humans were
BUILT TO LAST By S. Jay Olshansky, Bruce A. Carnes and Robert N. Butler
Illustrations by Patricia J. Wynne
Rewired eyes
Bigger ears
Curved neck
PERSON DESIGNED FOR A HEALTHY OLD AGE might possess

the features highlighted here, along with countless other
external and internal adjustments.
Forward-tilting
upper torso
Shorter limbs
and stature
Extra padding
around joints
Reversed
knee joint
94
B
ulging disks, fragile bones, fractured along despite the harmful consequences late in life.
hips, torn ligaments, varicose veins, Had we been crafted for extended operation, we
cataracts, hearing loss, hernias and would have fewer flaws capable of making us miserable
hemorrhoids: the list of bodily mal- in our later days. Evolution does not work that way,
functions that plague us as we age is however. Instead it cobbles together new features by tin-
long and all too familiar. Why do we kering with existing ones in a way that would have
fall apart just as we reach what should made Rube Goldberg proud.
be the prime of life? The upright posture of humans is a case in point. It
The living machines we call our bodies deteriorate was adapted from a body plan that had mammals walk-
because they were not designed for extended operation ing on all fours. This tinkering undoubtedly aided our
and because we now push them to function long past early hominid ancestors: standing on our own two feet
their warranty period. The is thought to have promoted
human body is artistically everything from food gather-
beautiful and worthy of all ing and tool use to enhanced
the wonder and amazement it
evokes. But from an engi-
We would look intelligence. Our backbone
has since adapted somewhat
neer’s perspective, it is a com-
plex network of bones, mus-
a lot different to the awkward change: the
lower vertebrae have grown
cles, tendons, valves and joints bigger to cope with the in-
that are directly analogous to if evolution creased vertical pressure,
the fallible pulleys, pumps, and our spine has curved a
levers and hinges in machines.
As we plunge further into our
had designed bit to keep us from toppling
over. Yet these fixes do not
postreproductive years, our
joints and other anatomical the human body ward off an array of prob-
lems that arise from our bi-
features that serve us well or pedal stance.
cause no problems at younger
ages reveal their imperfec-
to function What If?
tions. They wear out or oth-
erwise contribute to the health
smoothly for a R E C E N T L Y the three of us
began pondering what the
problems that become com- human body would look
mon in the later years. century or more like had it been constructed
In evolutionary terms, we specifically for a healthy long
harbor flaws because natur- life. The anatomical revi-
al selection, the force that molds our genetically con- sions depicted on the following pages are fanciful and
trolled traits, does not aim for perfection or endless incomplete. Nevertheless, we present them to draw at-
good health. If a body plan allows individuals to sur- tention to a serious point. Aging is frequently described
vive long enough to reproduce (and, in humans and as a disease that can be reversed or eliminated. Indeed,
various other organisms, to raise their young), then many purveyors of youth-in-a-bottle would have us be-
that plan will be selected. That is, individuals robust lieve that the medical problems associated with aging
enough to reproduce will pass their genes— and there- are our own fault, arising primarily from our decadent
fore their body design— to the next generation. Designs lifestyles. Certainly any fool can shorten his or her life.
that seriously hamper survival in youth will be weed- But it is grossly unfair to blame people for the health
ed out (selected against) because most affected indi- consequences of inheriting a body that lacks perfect
viduals will die before having a chance to produce off- maintenance and repair systems and was not built for
spring. More important, anatomical and physiological extended use or perpetual health. Our bodies would still
quirks that become disabling only after someone has re- wear out over time even if some mythical, ideal lifestyle
produced will spread. For example, if a body plan leads could be identified and adopted.
to total collapse at age 50 but does not interfere with This reality means that aging and many of its accom-
earlier reproduction, the arrangement will get passed Continued on page 99

WALK THIS WAY
A NUMBER OF the debilitating and even species to flourish. Every step we take single day, disks in the lower back are
some of the fatal disorders of aging stem in places extraordinary pressure on our feet, subjected to pressures equivalent to
part from bipedal locomotion and an ankles, knees and back— structures that several tons per square inch. Over a
upright posture— ironically, the same support the weight of the whole body lifetime, all this pressure takes its toll, as
features that have enabled the human above them. Over the course of just a does repetitive use of our joints and the
FLAWS
BONES THAT LOSE MINERALS AFTER AGE 30

Demineralization makes bones susceptible to
fractures and, in extreme cases, can cause
osteoporosis (severe bone degeneration),
curvature of the spine and “dowager’s hump”
RELATIVELY SHORT
FALLIBLE SPINAL DISKS RIB CAGE
Years of pressure on the spongy disks that Current cage
separate the vertebrae can cause them to slip, does not fully
rupture or bulge; then they, or the enclose and protect
vertebrae themselves, can press most internal organs
painfully on nerves
MUSCLES THAT LOSE MASS AND TONE

Such atrophy can impede all activities,
including walking. In the abdomen, hernias
can arise as the intestines (always pulled by
gravity) protrude through weak spots
in the abdominal wall. Flaccid abdominal
muscles also contribute to lower-back pain
LEG VEINS PRONE TO VARICOSITY

Veins in the legs become enlarged
and twisted when small valves that should
snap shut between heartbeats (to keep blood
moving up toward the heart)
malfunction, causing blood to pool.
Severe varicosities can lead
Normal direction to swelling and pain
of blood flow and, on rare occasions,
to life-threatening blood clots
JOINTS THAT WEAR
Malfunctioning As joints are used repetitively
check valve through the years, their lubricants
can grow thin, causing the bones
Pooled blood to grind against each other.
The resulting pain may be
exacerbated by osteoarthritis
and other inflammatory disorders
96 SCIENTIFIC AMERICAN Updated from the March 2001 issue

constant tugging of gravity on our tissues. tendons helps to tether our organs to the forever. Had longevity and persistent good
Although gravity tends to bring us spine, keeping them from slumping down health been the overarching aim of
down in the end, we do possess some and crushing one another. evolution, arrangements such as those
features that combat its ever present pull. But these anatomical fixes— like the depicted below might have become
For instance, an intricate network of body in general— were never meant to work commonplace.
FIXES
SHORTER STATURE
Would provide a lower FORWARD-TILTING UPPER TORSO
center of gravity, perhaps Would relieve pressure on vertebrae, thereby lessening the risk
preventing the falls of ruptured or slipped disks, which contribute, along with
that often fracture weakening abdominal muscles, to lower-back pain
demineralized bones
CURVED NECK WITH ENLARGED VERTEBRAE

CAGE WITH Would counterbalance the tilted torso and enable the
ADDED RIBS head to stay up and face forward
Could help prevent hernias
and other problems by
holding organs in
place more effectively
THICKER DISKS
Would resist destructive pressures
EXTRA MUSCLES AND FAT

Would add weight on the bones, which would help
counter the effects of demineralization; they
would also cushion bones against
breakage during falls
THICKER BONES
Would protect LEG VEINS WITH MORE
against breakage CHECK VALVES
during falls Would combat
the development
of varicose veins
Extra
valves
KNEE ABLE TO BEND BACKWARD Smooth-
Would make the bones less likely to grind flowing
and deteriorate, especially if the knee never blood
locked in place. But the absence of a locking
mechanism would make it hard to stand for
very long, so further modifications
would be needed LARGER HAMSTRINGS
AND TENDONS
Would help support
the leg and hip

PLAN A HEAD
EAR WITH FRAGILE FLAWS Detached retina
TRANSMITTERS
Hair cells of the inner Optic
ear, which relay nerve
VARIOUS PARTS of the head and sound information to
neck become problematic with the brain, become
damaged by exposure
disturbing regularity as people to loud noises
age. Consider the eye: the human
version is an evolutionary marvel,
but its complexity provides many
opportunities for things to go WEAK LINK BETWEEN RETINA
wrong over a long lifetime. AND BACK OF EYE
This frail connection exists in part
Our vision diminishes as the because the optic nerve, which carries
protective fluid of the cornea visual signals from the retina to the
becomes less transparent over brain, connects to the retina only from
the inside of the eye, not from the back
time. The muscles that control the
opening of the iris and the
focusing of the lens atrophy and Unwanted flow of food
lose responsiveness, and the lens
thickens and yellows, impairing
visual acuity and color COMMON UPPER PASSAGEWAY FOR FOOD AND AIR
When food travels toward the esophagus, a
perception. Further, the retina— flaplike tab of cartilage (the epiglottis) closes off
responsible for transmitting the trachea, or windpipe. With age, a progressive
images to the brain— can detach loss of muscle tone decreases the tightness of
Esophagus Trachea Epiglottis the seal, raising the risk of inhaling food or drink
fairly easily from the back of the
eye, leading to blindness.
Many of those problems would
be difficult to design away, but FIXES Retina
the squid eye suggests an

arrangement that could have
reduced the likelihood of retinal
detachment. A few anatomical
tweaks could also have preserved
hearing in the elderly.
Suboptimal design of the
upper respiratory and digestive
systems makes choking another OPTIC NERVE ATTACHED
risk for older people. A simple TO BACK OF RETINA
rearrangement would have fixed Might stabilize the retina’s
connection to the back of
that problem, albeit at the cost the eye, helping to prevent
of severe trade-offs. retinal detachment
Safer flow of food
ENLARGED, MOBILE OUTER EAR

Would collect sound with greater
efficiency, to compensate for
internal breakdowns RAISED TRACHEA
Would help food and drink to bypass the windpipe more
MORE PLENTIFUL AND effectively. This design would need refining, though,
DURABLE HAIR CELLS because it would disrupt breathing through the mouth
Would preserve hearing longer Esophagus and the ability to speak
Trachea

Continued from page 95 Health and Longevity
panying disorders are neither unnatural nor avoidable. No sim- O U R R E S E A R C H interest in redesigning the Homo sapiens
ple interventions can make up for the countless imperfections body is a reaction to the health and mortality consequences of
that permeate our anatomy and are revealed by the passage of growing old. We focus on anatomical “oddities” and “design
time. We are confident, however, that researchers in the vari- flaws” not only because they would be familiar to most read-
ous biomedical sciences will be able to ease certain of the mal- ers, but because they represent a small sample of lethal and dis-
adies that result from our extended life spans. Investigators are abling conditions that threaten the length and quality of life. It
rapidly identifying (and discerning the function of) our myriad is important to recognize that we live in a world in which hu-
genes, developing pharmaceuticals to control them, and learn- man ingenuity has made it possible for an unprecedented num-
ing how to harness and enhance the extraordinary repair capa- ber of people to grow old. Our redesign goal is thus to draw
bilities that already exist inside our bodies. These profound ad- attention to the health consequences associated with the aging
vances will eventually help compensate for many of the design of individuals and populations.
flaws contained within us all. One critical message we wish to convey is that people were
CALL A PLUMBER
AN EXPERIENCED PLUMBER MALE PROSTATE FEMALE BLADDER
side view front view
looking at the anatomy of
a man’s prostate might Ligament
suspect the work of
a young apprentice,
because the urethra, the Bladder
Ureter
tube leading from the
Ureter
bladder, passes straight from Bladder Wall muscle
through the inside of the kidney
gland. This configuration
may have as yet unknown Direction of
urine flow Enlarged
benefits, but it eventually prostate Sphincter
causes urinary problems in Direction of
Urethra urine flow
many men, including weak
flow and a frequent need
to urinate.
FLAW FLAW
URETHRA PRONE TO CONSTRICTION MUSCLES AND LIGAMENTS THAT WEAKEN WITH TIME
Women also cope with The prostate becomes enlarged in one of every two Particularly after multiple pregnancies,
plumbing problems as they males at some point in life. As it grows, it squeezes the muscles of the pelvic floor and the bladder, and the
the urethra, potentially obstructing the flow of ligaments that support the bladder, can sag,
age, particularly urine. Total obstruction can be fatal leading to incontinence
incontinence. Both sexes
could have been spared
much discomfort if
FIX STRONGER SPHINCTER MUSCLES IN BLADDER
FIX
URETHRA HUGGING OUTSIDE OF PROSTATE
evolution had made some Would not be squeezed if the AND MORE DURABLE LIGAMENTS
simple modifications prostate became enlarged Would increase control over bladder function
in anatomical design. Larger
ligament
Stronger
wall
Repositioned muscle
urethra
Larger
sphincter

We need to EXPLOIT OUR KNOWLEDGE of
evolution to enhance our quality of life as we grow older.
not designed by evolution for extended survival, so it is not their modify organisms (microbes, plants and animals) to suit their
fault that they ultimately suffer age-related ailments. Most of purposes. The most worrisome trade-off for genetic manipula-
what goes wrong with us as we grow older is a product of op- tion directed toward living longer would be an extension of
erating our living machines beyond their biological warranty frailty and disability rather than an extension of youthful health
period. Although we have considerable control over the quali- and vitality.
ty of our lives at any age, there is little we can do about the Though cobbled together by the blind eye of evolution, hu-
length of our lives other than shorten them. mans have proved to be a remarkably successful species. We
Even the term “flaw” requires clarification. Living things, have outcompeted almost every organism that we have en-
and everything they make, eventually fail. The cause of failure countered, with the notable exception of microbes. We have
is a flaw only when the failure is premature. A race car that fails blanketed the earth and even walked on the moon. We are also
beyond the end of the race has no engineering flaws. In the same one of the only species that has figured out how to escape pre-
way, bodies that fail in the postreproductive span of life may mature death and survive to old age.
contain numerous design oddities, but they have no design flaws At this point in history, we need to exploit our expanding
as far as evolution goes. Aging, disease and death are natural knowledge of evolution to enhance the quality of our lives as
by-products of bodies that were optimized for reproduction. we grow older, because the single-minded pursuit of life exten-
There are countless other aspects of human biology that sion without considering health extension could be disastrous.
would merit modification if health and longevity were nature’s Our fanciful designs of anatomically “fixed” humans are
primary objective. For example, gerontologists theorize that not intended as a realistic exercise in biomechanical engineer-
aging is caused, in part, by a combination of the molecular ing. Given what is known today about human aging, if the task
damage that inevitably arises from operating the machinery of of designing a healthy long-lived human from scratch were giv-
life within cells and the imperfect mechanisms for molecular en to a team comprising the father of evolution, Charles Dar-
surveillance, maintenance and repair that permit damage to win, the great painter Michelangelo, and the master engineer
accumulate over time. If this view of the aging process is cor- and scientist Leonardo da Vinci, they most certainly would have
rect, then modifying these molecular processes to lessen the fashioned a living machine that differs from the one we now oc-
severity or accumulation of damage, or to enhance the main- cupy. Indeed, anyone who tries his hand at redesign would
tenance and repair processes, should have a beneficial impact probably construct a human body that would look unlike the
on health and longevity. These wondrous modifications, how- ones we’ve created on these pages. Yet we invoke this approach
ever, would have little effect unless the common sense that is as an instructive way of communicating the important message
needed to avoid destructive lifestyles becomes more wide- from evolutionary theory that, to a significant degree, the po-
spread among people. tential length of our lives and, to a lesser degree, the duration of
Living things are exceedingly complex, and experience health and vitality are genetic legacies from our ancient ances-
teaches us that undesirable consequences invariably arise when- tors, who needed to mature quickly to produce children before
ever humans have taken over the reins of evolution in order to they were killed by the hostile forces of nature.
S. JAY OLSHANSKY, BRUCE A. CARNES and ROBERT N. BUTLER all MORE TO E XPLORE
THE AUTHORS
have an enduring interest in the processes that underlie human On Growth and Form. D’Arcy Wentworth Thompson. Expanded edition,
aging. Olshansky is professor in the School of Public Health at the 1942. (Reprinted by Dover Publications, 1992.)
University of Illinois at Chicago. He and Carnes, both senior re- The Panda’s Thumb: More Reflections in Natural History. Reissue
edition. Stephen Jay Gould. W. W. Norton, 1992.
search scientists at the National Opinion Research Center/Cen-
ter on Aging at the University of Chicago, collaborate on studies— The Scars of Evolution: What Our Bodies Tell Us about Human Origins.
Reprint edition. Elaine Morgan. Oxford University Press, 1994.
funded by the National Institute on Aging (NIA) and NASA—of the
The Blind Watchmaker: Why the Evidence of Evolution Reveals
biodemography of aging (examining the biological reasons for a Universe without Design. Reissue edition. Richard Dawkins.
age-related patterns of disease and death in populations). They W. W. Norton, 1996.
are co-authors of The Quest for Immortality: Science at the Fron- Why We Get Sick: The New Science of Darwinian Medicine. Randolph M.
tiers of Aging (W. W. Norton, 2001). Butler is president of the In- Nesse and George C. Williams. Vintage Books, 1996.
ternational Longevity Center in New York City and was founding The Olshansky and Carnes Web site is www.thequestforimmortality.com
director of the NIA. The International Longevity Center Web site is www.ilcusa.org


Scientific American Special Edition - New Look at Human

Încărcat de

Informații document

Drepturi de autor

Formate disponibile

Partajați acest document

Partajați sau inserați document

Opțiuni de partajare

Vi se pare util acest document?

Este necorespunzător acest conținut?

Drepturi de autor:

Formate disponibile

Scientific American Special Edition - New Look at Human

Încărcat de

Drepturi de autor:

Formate disponibile

WWW.SCIAM.

COPYRIGHT 2003 SCIENTIFIC AMERICAN, INC.

New Look at Human Evolution

CHAIRMAN EMERITUS: John J. Hanley

PRESIDENT AND CHIEF EXECUTIVE OFFICER:

www.sciam.com NEW LOOK AT HUMAN EVOLUTION 1

New Look at Human Evolution

14 Early Hominid Fossils from Africa

28 Who Were the Neandertals?

38 Out of Africa Again ... and Again?

46 The Multiregional Evolution of Humans

54 The Recent African Genesis of Humans

80 The Evolution of Human Birth

86 Once Were Cannibals

94 If Humans Were Built to Last

Cover painting by Kazuhiko Sano. This depiction of Sahelanthropus tchadensis—potentially the

www.sciam.com SCIENTIFIC AMERICAN 3

brown skull from its padlocked, foam-lined metal car-

than ever hunter murmurs, “the oldest hominid.”

APE OR ANCESTOR? Sahelanthropus tchadensis, potentially the oldest hominid on

4 SCIENTIFIC AMERICAN Updated from the January 2003 issue

Overview/The Oldest Hominids

may extend the human record back to seven million years

pological wisdom. But experts disagree over how these

6 SCIENTIFIC AMERICAN NEW LOOK AT HUMAN EVOLUTION

OMO Orrorin tugenensis

www.sciam.com SCIENTIFIC AMERICAN 7

COPYRIGHT 2003 SCIENTIFIC AMERICAN, INC.

www.sciam.com SCIENTIFIC AMERICAN 9

HUNTING FOR HOMINIDS:

10 SCIENTIFIC AMERICAN NEW LOOK AT HUMAN EVOLUTION

www.sciam.com SCIENTIFIC AMERICAN 11

Sahelanthropus tchadensis Australopithecus anamensis A. afarensis

12 SCIENTIFIC AMERICAN NEW LOOK AT HUMAN EVOLUTION

www.sciam.com SCIENTIFIC AMERICAN 13

early hominid fossils from

14 SCIENTIFIC AMERICAN Updated

www.sciam.com SCIENTIFIC AMERICAN 15

16 SCIENTIFIC AMERICAN NEW LOOK AT HUMAN EVOLUTION

The human jaw

The top of the

humerus where the Human and

www.sciam.com SCIENTIFIC AMERICAN 17

KANAPOI An Even Older Hominid?

18 SCIENTIFIC AMERICAN NEW LOOK AT HUMAN EVOLUTION

www.sciam.com SCIENTIFIC AMERICAN 19

TODAY WE TAKE FOR GRANTED THAT HOMO SAPIENS

20 SCIENTIFIC AMERICAN Updated from the January 2000 issue

were not alone

By Ian Tattersall • Paintings by Jay H. Matternes

www.sciam.com SCIENTIFIC AMERICAN 21

PARANTHROPUS BOISEI HOMO RUDOLFENSIS

22 SCIENTIFIC AMERICAN NEW LOOK AT HUMAN EVOLUTION

HOMO HABILIS HOMO ERGASTER,

www.sciam.com SCIENTIFIC AMERICAN 23

24 SCIENTIFIC AMERICAN NEW LOOK AT HUMAN EVOLUTION

www.sciam.com SCIENTIFIC AMERICAN 25

(South Africa) Paranthropus

the appalling subsequent record of H.

26 SCIENTIFIC AMERICAN NEW LOOK AT HUMAN EVOLUTION

www.sciam.com SCIENTIFIC AMERICAN 27

Who Were the

It was such a neat and tidy story.

www.sciam.com SCIENTIFIC AMERICAN 29