Arsuaga, Martinez - The Chosen Species. The Long March of Human Evolution (1998) PDF
Arsuaga, Martinez - The Chosen Species. The Long March of Human Evolution (1998) PDF
Arsuaga, Martinez - The Chosen Species. The Long March of Human Evolution (1998) PDF
Bernard Wood, Henry R. Luce Professor of Human Origins, George Washington University
THE CHOSEN
SPECIES
The Long March of Human Evolution
The Chosen Species
For Lourdes
For Chon
Spanish edition ß 1998 by Juan Luis Arsuaga Ferreras, Ignacio Martı́nez Mendizábal, and
Ediciones Temas de Hoy, Ltd. (T.H.)
Illustrations ß 1998 by Mauricio Antón
English translation ß 2006 by Blackwell Publishing, Ltd.
BLACKWELL PUBLISHING
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The right of Juan Luis Arsuaga Ferreras and Ignacio Martı́nez Mendizábal to be identified as
the Authors of this Work has been asserted in accordance with the UK Copyright, Designs,
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system, or transmitted, in any form or by any means, electronic, mechanical, photocopying,
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1988, without the prior permission of the publisher.
Originally published in 1998 by Temas de Hoy as La especie elegida
English edition published 2006 by Blackwell Publishing
Translated into English by Rachel Gomme
1 2006
Library of Congress Cataloging-in-Publication Data
Arsuaga, Juan Luis de.
[Especie elegida. English]
The chosen species : the long march of human evolution / Juan Luis Arsuaga, Ignacio
Martı́nez ; illustrations by Mauricio Antón ; [translated into English by Rachel Gomme].
p. cm.
‘‘Originally published in 1997 by Temas de Hoy as La especie elegida’’—T.p. verso.
Includes bibliographical references and index.
ISBN-13: 978-1-4051-1532-2 (hard cover : alk. paper)
ISBN-10: 1-4051-1532-7 (hard cover : alk. paper)
ISBN-13: 978-1-4051-1533-9 (pbk. : alk. paper)
ISBN-10: 1-4051-1533-5 (pbk. : alk. paper)
1. Fossil hominids. 2. Human evolution. 3. Primates—Evolution. 4. Anthropology,
Prehistoric. I. Martı́nez, Ignacio. II. Antón, Mauricio. III. Title.
GN282.A69813 2006
599.93’8—dc22
2004027275
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Contents
Acknowledgments xi
Introduction: Prehistory 1
Little Lucy 1
Intrepid Paleontologists 3
2 We the Primates 17
The Ecological Definition and Diversity of Primates 17
The Classification of Primates 20
Hominoids, Apes of Our Own Branch 22
History of the Primates 27
v
Contents
Milankovitch Cycles 40
A Climatic Model for Equatorial Africa 43
The Controversial Gas 45
The End of Paradise 46
vi
Contents
10 Development 145
The Rhythm of the Molars 145
Birth and the Newborn 146
Childhood and Adolescence 152
Summary 171
vii
Contents
Epilogue 264
The Never-ending Story 264
Bibliography 268
Index 275
viii
For now consider, like the zoologists and the anatomists,
that man has more of the ape than of the angel, and that
he has few grounds for vanity or conceit.
Santiago Ramón y Cajal
Acknowledgments
Little Lucy
Little Lucy trudged her way laboriously through the African savanna. As a
result of continuous effort over generations, she walked on two legs rather
than four like her ancestors. With her child weighing heavy in her arms,
she felt herself weakening as she drew nearer to the clump of thorny
acacias which could be glimpsed on the horizon, under the torrid tropical
sun. Small as she was – little more than a meter in height, and weighing
less than 30 kilograms – it was only by her wits that she had evaded
powerful predators. She had no stone tools. A million years had passed
since her ancestors, the first hominids, had decided to leave the protection
of the forest and move out into the savanna which continued to expand,
as a result of the great climate change then taking place. Her relatives,
the ancestors of chimpanzees and gorillas, had preferred the safety of the
forest and would remain there forever. For destiny belonged to the
bold, those who defied the danger of open environments. One day these
bolder beings would evolve, their brains and their intelligence would
develop, they would manufacture all kinds of tools, they would discover
fire and would banish forever the lion, the leopard, and the hyena.
1
Introduction: Prehistory
Eventually, they would conquer the world. All of this depended on Lucy
and her baby surviving and joining the little group of australopithecines
waiting in the stand of trees, who represented the future of humanity.
Lucy literally carried our future on her two legs.
But no, dear reader, this is not one of those books; we have not allowed
our imagination to run away with us. This is not a fanciful tale about
human evolution. Its aim is not to keep you in suspense, holding your
breath over the vicissitudes of our ancestors. After all, the ending of the
story of human evolution is well known. Lucy survived, if you like to think
of it those terms. Ultimately we are all descended from many Lucys.
Misia Landau has drawn attention to the narrative structure of histories
of human evolution, the rhetoric which surrounds them and its similarity
to mythological and religious literature. This is evident in the biblical
references in some of the names given to scientific hypotheses about our
origin, such as the Black Eve hypothesis, the Noah’s Ark hypothesis, or
even the title of this book. In fact in other cases the only aim of such
names is to catch the reader’s attention with familiar-sounding titles, like
the Out of Africa hypothesis, or the East Side Story hypothesis. But the
really important issue is not the name given to these hypotheses, but that
they can be judged against the facts, modified and even rejected if they are
not compatible with them: this is what makes them scientific hypotheses
rather than simply opinions or fantasies. Only dogma is immutable.
For however her story is told, Lucy is much more than a myth. Lucy is
real. The man who discovered her, Donald Johanson, gave this name in
1974 to an extremely well-preserved skeleton of a female hominid who
lived in what is now known as Ethiopia some three million years ago. And
she really was small. She no longer lived in the enclosed, humid forests,
but rather in more open, arid spaces: although this environment increased
the risk of falling victim to predators, it offered new plant resources for
food. Lucy could not speak as we do; her brain was not much bigger than
that of a chimpanzee, and she had no stone tools, but she was bipedal. All
the other elements of our tale – the split between the lines leading to
chimpanzees and to our species, when and where it took place, the climate
change which occurred and the reduction in the tropical forest which
resulted from it – all of this is based on scientific data. Our only artistic
license is in relating the life-story of a particular hominid individual.
Nevertheless, there are elements in the structure of this story which
remain hidden from view, but which have profound implications for
correct understanding not only of human evolution, but of evolution
in general. It is therefore worth pausing a moment to analyze these
narratives.
2
Introduction: Prehistory
Intrepid Paleontologists
3
Introduction: Prehistory
4
Introduction: Prehistory
the end of the novel only to find that the solution depends on evidence
which only the detective knew about and which has been concealed from
the reader until that point. But what is really unforgivable in a detective
novel is if the solution to the case is not explained in a satisfactory way,
since the most interesting thing is not ‘‘whodunit,’’ but how it was
worked out. This is because the detective novel appeals to the reader’s
intelligence. But if the reader is to have all the information at his disposal
and then appreciate the detective’s shrewd work, he needs to be present at
the interviews with all the witnesses, to observe the scene of the crime at
his leisure, to investigate suspects’ past history, study the results of labora-
tory analysis, and take time to reflect in order to try and fit all the pieces of
the puzzle together.
Well, this book also appeals to the intelligence of our readers, and we
have therefore tried to ensure that you will find in its pages all the facts and
arguments on which our conclusions are based. To this end, the book
follows the chronological axis of the evolution of hominids, and tackles
the key questions at different points. As in a detective novel, the reader can
skip straight to the end to find the solution, but this will mean missing
most of the plot; moreover, the file remains ‘‘open’’ on many of the
questions of human evolution.
Two quotations cited in Fernando Trueba’s Diccionario del cine [Dic-
tionary of Cinema] influenced the conception of this book, we hope to
the good. One of these will be discussed in the epilogue. The other is
apposite here, because it was a great solace when we were attempting to
make some of the really very complicated scientific problems easily com-
prehensible for a general audience. Under the keyword ‘‘simplicity’’ in
Trueba’s book, he quotes the following phrase from Albert Einstein:
‘‘Everything must be made as simple as possible, but no simpler.’’
5
I
Children of Africa
1
Basic Principles of
Evolutionary Theory
9
Basic Principles of Evolutionary Theory
Natural Selection
10
Basic Principles of Evolutionary Theory
that if they were not checked, human populations would tend to grow in
geometric progression, outstripping the increase in resources.
There is a kind of sentimental ecological thinking which has become
popular among city-dwellers, and which hampers correct understanding
of natural selection. Many people believe that animals, when they are not
persecuted by humans, lead a pleasant, risk-free life in nature. If this were
the case, why would living beings have to adapt to become more efficient?
On the contrary, simple arithmetic shows us that, in the natural world,
the life of animals is far from smooth. In demographically stable popula-
tions – in other words, populations which are not growing, which includes
all populations in normal circumstances over the long term, although
there may be short-term fluctuations – each reproducing pair is replaced
by another two individuals in the succeeding generation. Nevertheless, in
favorable conditions, a zebra living on the African plains gives birth to a
foal each year from the age of 4, over a period of 15 or more years; a
gazelle gives birth to one kid every 6 months from the age of 2. It is
obvious that the majority of those born will not reach adulthood or
reproduce. The predators fare no better: lions begin to reproduce at the
age of 4 (they can easily live up to 15 years in the wild), and have 2 or 3
cubs every 20 or 30 months; leopards living in the wild reproduce from
the age of 2 to the age of 12, producing from 1 to 3 cubs at intervals of
about 25 months.
The same reasoning applies to primates, and, over the period of their
evolution, to humans as well, although our situation has changed in
recent times, since the infant mortality rate has fallen so much that
without birth control we see an explosion of the population. The propor-
tion of fertilized eggs which do not become reproducing adults is almost
one hundred percent in the majority of aquatic vertebrates (amphibians,
fish), and in almost all invertebrates. The upshot of these simple figures is
that individuals in the different species are constantly under the threat of
death, and that consequently small genetic advantages may be crucial to
reaching adulthood and reproducing, or to continuing to reproduce. This
is what Darwin meant when he spoke of the struggle for existence, which
is not necessarily a bloody battle: plants and herbivores also compete with
one another.
Unlike artificial selection, performed by humans with animals and
plants and gradually enhancing specific characteristics in order to increase
productivity, natural selection pursues no particular goal. Moreover, no
genetic variant is better than another in an absolute sense; everything
depends on the environmental circumstances. What is favorable at one
moment may be unfavorable at another. In addition, through the
11
Basic Principles of Evolutionary Theory
12
Basic Principles of Evolutionary Theory
factor has been invoked to support the model of slow evolution, suggest-
ing that infinitesimal variations would give those who carried them the
tiniest of advantages which would only become dominant over many
generations. And there is no shortage of time in paleontology. After all,
life has existed on earth for more than 3,500 million years. It is around
this tension between two extremes, gradual evolution or evolution by
leaps, that current debate within evolutionary theory is focused.
13
Basic Principles of Evolutionary Theory
are ‘‘better’’ than reptiles, that primates are ‘‘better’’ than other mam-
mals, and finally, that humans are ‘‘better’’ than chimpanzees and gorillas?
According to Darwin, evolution has no goal, it follows no preconceived
design; it is simply opportunist, and is not directed toward any idea of
perfection. To put it another way, all species (including our own) are
equally perfect, each one marvelously adapted to its life habits through
natural selection. In other words, unlike artificial selection which the
farmer or stockbreeder carries out with a particular aim, natural selection
has no objective. Although in common parlance (as well as in politics and
business) the word ‘‘evolution’’ signifies change for the better, in Dar-
winian terms ‘‘evolution’’ means simply change, nothing more.
Among the mammals, humans are distinguished as tailless, bipedal
primates with a large brain, but otherwise we have few original features.
We still retain five digits on our hands and feet, while horses support
themselves on the third phalange of their single digit. We show nothing
like the transformations which bats or dolphins underwent, to evolve from
their quadruped ancestors. Are we more evolved, in the sense of being
more changed, than they are? Granted, a geranium cannot write a book –
that is one of our specialties – but with the aid of light it can synthesize
organic matter from mineral salts, water, and carbon dioxide; there is no
doubt that the geranium has a well-equipped laboratory, and it is difficult
to see it as an ‘‘inferior’’ being.
But those who prefer to see evolution as an arrow which has been
pointing toward us since the beginning will have to answer the question
of what obscure internal forces could be driving evolution in the right
direction, independent of what happens around it. Or are we really
dealing with forces outside of the natural world? In this case we move
outside of the sphere of science, which is the terrain of this book and its
authors. The object of science is to explain natural phenomena, like the
existence of our species (and all the others), in terms of natural causes.
So, to return to the terrain of science, Lamarck believed in the idea of
progress in evolution. However, the mechanism he proposed as the motor
driving evolution forward was adaptive (like that of Darwin), and did not
take any particular direction (even to ‘‘divert’’ organisms along ‘‘aber-
rant’’ blind alleys). Confronted with this paradox, Lamarck resolved it by
adding to his theory of use and lack of use the idea that all living forms
‘‘tended’’ gradually and inevitably toward ever higher (in other words,
more complex) levels of organization. Lamarck never explained the cause
of this tendency toward perfection, but later authors attributed it to ‘‘vital
impulses,’’ and therefore became known as vitalists, or finalists, because
they believed that evolution had directionality.
14
Basic Principles of Evolutionary Theory
For those who believe that the Story of Life reflects a program unfold-
ing over time, evolution is in some ways comparable to the process of
development which leads from the embryo to the adult, following pre-
established laws (natural, but which we do not as yet understand very
well). Clearly, those who wish to find significance, meaning, or an inten-
tion in the Story of Life will always have recourse to mysterious internal
forces, still to be discovered or unknowable.
But if it is the case that invisible threads have, from the beginning of
time, guided evolution in a linear and orderly manner to result in us, why
do we find ourselves among such a diversity of living forms? As we shall
see, we are not descended from chimpanzees, but we have a common
ancestor with them. Chimpanzees are our brothers, not our parents. Nor
are we descended from any organism like a sea urchin. Nevertheless, the
phylum of echinoderms, to which sea urchins belong, and that of the
chordates, which includes our species, share a very remote ancestor which
was neither a sea urchin nor a human being. Living species are not ordered
in sequence. What we have here is not a staircase to nowhere, but a tree
with many, many branches and with no trunk or central axis. Evolution is
not linear, but divergent.
Despite all of this some authors, although aware that, whether looking
at the past or the present, evolution does not appear to have moved in one
single direction, still express the view that life followed various tendencies,
and that ours is the tendency of increasing intelligence. However, they
never explain how these tendencies arise: they appear to obey mysterious
impulses which have nothing to do with the adaptation of organisms, but
rather act of their own accord. The definition of primates still frequently
includes the term ‘‘tendency toward cerebral expansion,’’ as if a tendency
could serve in and of itself to characterize an entire group including fossil
and living species, which would thus become a ‘‘unit of evolutionary
destiny.’’ Naturally, contemporary primates that do not exhibit such
cerebral expansion are held simply to represent relics of the past, or ‘‘living
fossils.’’
The authors remember a time when groups of primates or hominids
which were not in a direct evolutionary line with our species were de-
scribed as deviant forms; these ideas can still readily be found in serious
texts. If such forms became extinct, as a just punishment for their rebel-
lion, they were tarred as abortive or even aberrant forms, described as
trials or failed experiments (carried out by whom?), as blind alleys, or in
other terms which made it clear that it was not a good idea to diverge
from the path marked out by evolution.
15
Basic Principles of Evolutionary Theory
But let us leave, for now, the debate between vitalism and materialism,
or finalism and Darwinism. Over the chapters of this book we will review
what we know today of human evolution through its different stages and
circumstances. The last chapter will give us the opportunity to discuss, in
the light of past events, the nature of our history. But first, in the next
chapter, we will start to get to know ourselves a little better, locating
ourselves among the diversity of species of living and fossil primates.
16
2
We the Primates
17
We the Primates
18
We the Primates
grasp and climb trees, and jump from branch to branch. One such
adaptation is the first toe (the big toe), which is very large and mobile
and can be opposed to the other toes (except in our case). Primates also
have flat nails on all the digits of hands and feet, rather than the claws of
their ancestors. Some primates have claws, sometimes on all the digits
except for the first toe, but it appears that these were originally flat nails
which converted back into claws at a later date.
The teeth of mammals are very important in paleontology for two
reasons: one is that dentition reflects the type of diet, and the other is
that they are the most frequently found fossils – often the only fossils found
for certain groups. A first step toward the study of dentition is counting the
number of teeth an individual has. All living primates descend from an
ancestor who had 36 teeth. However, not all mammals have the same kinds
of teeth – they are specialized for different functions. Four groups of teeth
are distinguished, from front to back: incisors, canines, premolars, and
molars (Figure 2.1). The primate ancestor of all modern primates had, on
each side of the mouth and at both top and bottom, two incisors, one
canine, three premolars, and three molars. Some primates have modified
Molars
Premolars
Canine
Incisors
19
We the Primates
this formula over the course of their evolution, losing some elements, but
never increasing the number of teeth in any of the four categories.
20
We the Primates
needs to look at the nearest cat or dog. This arrangement of the nose
severely limits the facial expression of emotions. Haplorrhinae, including
ourselves, do not have this hairless skin around the nostrils, and the upper
lip is undivided and mobile. The continuous upper lip facilitates greater
facial expressivity, a well-known characteristic of haplorrhine primates.
The Strepsirrhinae include lemurs, indris, and the aye-aye (lemurs in the
broader sense), which evolved and diversified in isolation on the island of
Madagascar (off the east coast of Africa). Some species are nocturnal and
others diurnal. Unfortunately, the recent arrival of humans has resulted in
the degradation of their forest paradise and the disappearance of many
species.
The Strepsirrhinae also include the Asian lorises, and the African bush
babies and pottos (the Lorisidae in the broader sense), all of which are
nocturnal.
The Haplorrhinae are divided into three groups. One of these is the
tarsiers (Tarsiiformes), small nocturnal primates found in the Philippines,
Borneo, Sumatra, and other Southeast Asian islands, with enormous eyes,
very long tails, and very long back legs, well adapted to jumping. The
other groups of Haplorrhinae are the catarrhines (Catarrhinae), which
include our own species, and the American monkeys, known as platyrrhine
primates (Platyrrhinae).
The catarrhines and platyrrhines are usually grouped together in a
common category used more or less informally, that of anthropoid apes
(technically, Anthropoidea) – from here on we will use the term anthro-
poid. Both groups are diurnal, with the exception of the South American
owl monkey, Aotus trivirgatus, which appears to have become nocturnal,
although it is descended from diurnal ancestors.
Another feature of anthropoids is the completely frontal position of the
eyes, allowing for a wide field of stereoscopic or three-dimensional vision,
which requires that the fields of vision of the two eyes overlap. This type of
vision makes it possible to calculate the distance to objects, whether tree
branches or prey, very precisely. Anthropoids have a large brain, although
it appears that platyrrhines and catarrhines developed this (through evo-
lution) separately. The olfactory lobes of their brains are very much
reduced in size. As anthropoids, we perceive the world essentially in
terms of images, not in odors.
Platyrrhines have the same number of teeth as the first primates, except
for the marmosets and tamarins (Callitrichinae), a group which has lost the
last molar. The catarrhines, however, have lost a premolar (Figure 2.1),
although many of our readers will also find that the last molar (the wisdom
teeth) will never emerge. This absence of the third molar in the adult jaw is
21
We the Primates
22
We the Primates
Figure 2.3 Types of hominoid. Both sexes are shown. Top left : a pair of gibbons
(Hylobates lar); top right : orangutans (Pongo pygmaeus); below (left to right): gorillas
(Gorilla gorilla), common chimpanzees (Pan troglodytes), and humans
23
We the Primates
24
We the Primates
25
We the Primates
26
We the Primates
themselves on the soles of their feet and on the dorsal (back) side (not the
knuckles, as is often stated) of the intermediate phalanges of the second to
fifth fingers (the index to the small finger). Quadruped primates walk on
the soles of their feet and the palms of their hands. Gorillas’ and chim-
panzees’ particular mode of travel has also resulted in modifications to the
bones of the arms, to give them more stability when supporting the
weight of the body on either one.
The first known fossil thought to be that of a primate (though this is not
certain) is a molar from the late Cretaceous, the last period of the Meso-
zoic or Secondary era, when the dinosaurs were still alive. It is around
65 million years old and was found in Montana. It was named Purgatorius
ceratops, as it is contemporary with the well-known three-horned dinosaur
Triceratops.
Purgatorius is allocated to the group Plesiadapiformes, which continues
into the following era, the Cenozoic, made up of the Tertiary period
together with the Quaternary, or Pleistocene. Plesiadapiformes (Figure
2.6) are the only primate fossils known from the Paleocene, the first epoch
of the Tertiary (between 65 and 55 million years ago); during this epoch
they split into a number of lines. The last of these became extinct in the
subsequent, Eocene epoch (between 55 and 36 million years ago). The
fact that Plesiadapiformes have been found in both North America and
Europe indicates that the two continents were joined together before they
became definitively separated by the Atlantic Ocean. Throughout the
Cenozoic, Asia and North America were also joined from time to time,
in the region of what is now the Bering Strait. However, Africa was
isolated, like a large island, during most of the Cenozoic, although there
were sometimes land bridges between Eurasia and Africa which allowed
for interchanges of fauna.
There is ongoing debate as to whether the Plesiadapiformes should be
considered true primates or not (for example, they do not appear to have
had flat nails, nor an opposable big toe). Nevertheless, Plesiadapiformes
are the group closest, in evolutionary terms, to living primates, which
themselves form a natural group with an exclusive common ancestor.
Some authors suggest that primates can be divided into two large cat-
egories, Plesiadapiformes, or archaic primates, and others, the euprimates
or ‘‘true’’ primates (see Figure 2.7).
27
We the Primates
Euprimates appear in the fossil record in the Eocene (the epoch be-
tween 55 and 36 million years ago), and are well represented in Eurasia
and North America by two large groups, the Adapiformes and the Omo-
myidae, although both groups probably also lived in Africa. The Adapi-
formes show some general similarities with lemurs and lorises, but we
cannot say for certain that they are the direct ancestors of these animals
(Figure 2.8). Many authors consider the Omomyidae to be haplorrhines,
but this classification is also open to doubt. Finally, some middle Eocene
fossils from China and Algeria (about 45 million years old) have been
identified as anthropoids, though not all authors recognize them as such.
The transition from the Eocene to the subsequent epoch, the Oligo-
cene, is well represented in the El Fayum deposits in Egypt (Figure 2.9).
Numerous anthropoid fossils, between 30 and 37 million years old, have
been found here. The anthropoids, let us recall, are the group formed by
the platyrrhines, or New World monkeys, together with the catarrhines,
which in their turn are divided into the Old World monkeys and the
hominoids; the latter comprise apes and humans.
28
We the Primates
Strepsirrhinae Haplorrhinae
Anthropoids
Catarrhinae
Platyrrhinae
0 Pleistocene
Tarsiers
Cercopithecidae
Hominoids
Pliocene
Adapiformes
Miocene
−20
CENOZOIC
Oligocene
Omomyidae
Years (millions)
El
Fayum
Plesiadapiformes
−40
Eocene
fossils
First
fossil
anthropoids
Cretaceous Paleocene
−60
MESOZOIC
−80
29
We the Primates
Figure 2.8 Notarctus. A North American Eocene adapiform, with hind limbs
longer than its front limbs and a capacity for jumping similar to that of the modern
lemurs of Madagascar
these are the caviomorph rodents, which include guinea pigs, capybaras,
chinchillas, viscachas, and so on.
During the Miocene, the epoch which succeeded the Oligocene and
lasted from 24 million to 5 million years ago, the first hominoids – the
group to which we and the apes belong – emerged on the African
continent. The oldest fossils, found in East Africa, have been assigned to
the genus Proconsul (Figure 2.10), a type of hominoid which lived before
the line which led to the gibbons split off. The name Proconsul (pre-
Consul) was coined in 1933 in honor of a famous chimpanzee at London
Zoo called Consul, in the mistaken idea that the fossil species was the
direct ancestor of modern chimpanzees.
After Proconsul come Morotopithecus, Afropithecus, and Kenyapithecus.
Fossils of these four species dated between 23 and 14 million years ago
have been found. Proconsul does not appear to have developed the
locomotive adaptations characteristic of modern hominoids. However,
some details of the lumbar anatomy and of the joint between the shoulder
blade and humerus of Morotopithecus, 20.6 million years old, suggest that
30
We the Primates
this genus may offer the oldest fossil evidence of a body organization
similar to that of living apes.
Around 17 million years ago, Africa (at that time joined to the Arabian
peninsula), which had remained quite isolated since the Eocene, drew
31
We the Primates
Figure 2.10 Proconsul. The first fossil hominoid, which lived in East Africa
during the early Miocene
32
We the Primates
Dryopithecus laietanus. With its long arms and short legs, this fossil gives
us an idea of the way in which these hominoids, of about 9.5 million years
ago, moved around (Figure 2.11). They appear very similar to modern
orangutans, which hang from the branches of trees and move slowly
through them. A well-known but enigmatic long-armed primate known
Figure 2.11 Dryopithecus. Fossil hominoid which lived in Europe during the
Miocene
33
We the Primates
as Oreopithecus, which lived during the same period in the swampy forests
of central Italy and Sardinia (which at that time formed a single large
island), could be close, in evolutionary terms, to Dryopithecus. However,
in view of the very specific characteristics of its dentition (type and ar-
rangement of teeth), some authors do not even recognize Oreopithecus as
a hominoid.
The fossil trace of Eurasian hominoids disappears about 7 million years
ago, until forms already very close to the modern orangutan and gibbons
appear; the one notable exception is Gigantopithecus. A species of this
genus (Gigantopithecus blacki) survived in China and Vietnam until only a
few thousand years ago, and therefore must have lived at the same time as
humans. We have three jawbones and more than a thousand individual
teeth from this species. The first fossils of Gigantopithecus blacki to be
found were four molars which the paleontologist Ralph von Koenigswald
(1902–82) purchased in drugstores in Hong Kong and Canton between
1935 and 1939 (in traditional Chinese medicine, fossils are credited with
healing properties). The Gigantopithecus jawbones are larger than those of
gorillas – particularly one of them, believed to be that of a male. These
were probably the largest primates which ever existed.
The front teeth (incisors and canines) were relatively small, while the
premolars and molars were large and wide, with a thick layer of enamel.
The jawbones are also very robust. In these characteristics they are similar
to Paranthropus, a type of hominoid with very strong teeth and jaws,
which we shall describe later. This similarity has led some people to believe
that Gigantopithecus belonged to our evolutionary group, but in fact this
is a case of adaptive convergence – in other words, a resemblance due to
more or less similar chewing activity in two independent lines. The
evolutionary affinities of Gigantopithecus are more likely to be with Siva-
pithecus and other species in the orangutan line.
In view of their size, we can only assume that these enormous apes lived
on the ground, and that they fed on what must have been very abundant
plant resources. The strength of the jawbones and the type of dentition
indicate that they ate a hard, fibrous type of plant which required much
chewing. Some authors believe that this may have been bamboo, eaten by
modern pandas.
34
3
The hay appeareth, and the tender grass sheweth itself, and herbs of the
mountains are gathered.
Proverbs 27:25
The origin and the disappearance of species are two of the fundamental
problems of evolutionary biology. By selecting animals and plants, man
has managed to produce many different breeds and varieties but not, so
far, any new species. All breeds of a given species can interbreed, produ-
cing fertile offspring which can reproduce again. This is not the case with
different species of animals: although two different species can be bred
together, such as horses and donkeys, their offspring – in this case the
mule – is sterile. Obviously this may be a matter of time, given that we
have only been domesticating animals and plants on a large scale for
10,000 years.
During his famous voyage of five years and two days around the world
in the brig Beagle, Charles Darwin began to get an idea of a basic
mechanism for the development of new species. If a few individuals
remained isolated in a marginal region, they could adapt to the conditions
prevailing there and transform into new species. This would explain the
diversity of chaffinches found in the Galápagos Islands (Ecuador). Each
island had its own distinct species of chaffinch – in some cases more
than one – with different specializations (or, as an ecologist would say,
35
Climate and Evolution
occupying different niches); all developed from a single species, which had
come from the American continent.
The emergence of new species is not always the result of a seed or a pair
of a particular animal species being passively transported to a remote place
by winds or currents, as occurred with the Galápagos chaffinches or the
platyrrhine monkeys of South America. The area of distribution of a
species may be split by a new geographical barrier, giving rise to new
species on either side of the barrier. And there are also cases where species
are subjected to a change of landscape without having to move or be
isolated by changes in topography. This is the case of the hominoids,
which lost their environment because of a great, planet-wide ecological
change.
36
Climate and Evolution
Today
Average global
temperature
Warm
Cold
CARBONIFEROUS
PLEISTOCENE
CRETACEOUS
ORDOVICIAN
PALEOCENE
OLIGOLENE
CAMBRIAN
DEVONIAN
PLIOCENE
JURASSIC
MIOCENE
SILURIAN
TRIASSIC
PERMIAN
EOCENE
Absolute
Absolute age
age
(millions
(millions of
of years)
years)
4000 3000 2000 1000 570 225 65 1,7
37
Climate and Evolution
Paleotemperature Scales
The spectacular traces left by the ice during its advances and retreats in the
northern hemisphere have been used to establish a scale of changes in
temperature in Europe over time. Nevertheless the Ice Ages, although
they were large-scale climatic episodes, were also local phenomena which
affected different regions in different ways, even within the small Euro-
pean continent. Moreover, it is difficult to track them further back than
one million years.
The scales now used are based on marine paleotemperature curves,
derived from investigation of the sedimentary deposits accumulated in
the seabed. These marine records go back much further and are more
continuous than the continental records, and are a more reliable reflection
of global changes of temperature.
The most commonly used scale is based on the oxygen found in the
minute fossil shells of microorganisms known as foraminifera. These
minuscule protozoa live in the sea, and when they die their shells gradually
accumulate at the bottom. Dozens of test wells have been drilled in deep
water all over the world in order to obtain a microfossil sequence covering
a long period of time. In nature, oxygen occurs in two different isotopes,1
both of which are stable: oxygen-16, light and very abundant, and oxy-
gen-18, heavier and very rare. The ratio of the two oxygen isotopes in
seawater, and in the carbon dioxide dissolved in it, depends on the
temperature. Because this oxygen passes into the shells of the foraminifera
while they are alive, the temperature is recorded in their bodies for all
time; when they die this information is gradually deposited at the bottom
of the sea.
In order to relate these marine scales, which tell us the temperature of
the seawater, with continental climatic cycles, the deposits of dust trans-
ported from the land by the wind have also been traced in the seabed, in
order to see how these deposits have changed; it is assumed that the drier
1
Isotopes are atoms of a specific element which are chemically distinct from one
another. Because the small difference occurs in the nucleus of the atoms (in the number
of neutrons, to be precise), it has little effect on the normal properties of the element,
except for a minute difference in weight. The fundamental importance of isotopes, and
the reason they are so well known, is their nuclear properties: some are stable, like
carbon-12, and others are radioactive (i.e., they decompose naturally by emitting
radiation), like carbon-14. Some, such as uranium-235, are used to make atomic
bombs, while others, like uranium-238 and others, are not.
38
Climate and Evolution
and more arid the adjacent continental regions, and the less vegetation
they have, the more dust will have blown toward the ocean. Fossil pollen
grains in the seabed have also been studied, in order to find out what kind
of plants covered the surface of the earth.
Marine records tell us what happened over the last few million years,
but they do not tell us why. What are the main factors contributing to the
changes in the earth’s climate?
The basic factors leading to climate change can be grouped into five main
categories: (a) catastrophic events; (b) geodynamic evolution of the
planet; (c) behavior of the hydrosphere-atmosphere system; (d) natural
fluctuations in the earth’s orbit around the sun; (e) the effect of the
biosphere, including human activity.
These factors, some of them very intricately enmeshed, produce very
different effects. Catastrophic events, which are sudden and unpredict-
able, such as the impact of giant meteorites or huge volcanic eruptions,
lead to marked changes of short duration; only if the change produced is
extremely drastic will it affect entire species.
At 10:00 a.m. on August 27, 1883, the largest recorded explosion in
history, much larger than any nuclear test, occurred in the volcanic crater
of Krakatoa (Indonesia). It was heard 3,500 km away; it destroyed an
entire island and spewed forth 21 sq m of rock. It shot enormous quan-
tities of gas and ash into the atmosphere, forming a vertical stream 80 km
high. The fine dust particles expelled into the stratosphere traveled
around the world several times and produced spectacular sunsets for
years, even in Europe. The resulting tsunami (‘‘tidal waves’’) traveled as
far as Hawaii and South America. More than 50,000 people died. It is
possible that the global temperature of the earth fell by half a degree, but
no long-term change occurred; no species disappeared. Currently there is
only one catastrophe theory which is taken seriously. This is the hypothesis
of a major meteorite impact, which is suggested as the cause of the
extinction of the dinosaurs – although this is still the subject of fierce
debate.
Geodynamic evolution includes a wide range of phenomena such as a
reduction in the flow of heat from the interior of the earth to the surface,
shifts in the geographic and magnetic poles, volcanic activity, and vertical
39
Climate and Evolution
Milankovitch Cycles
The earth’s orbit around the sun results in two extremely regular tem-
perature cycles which we know well. The first is the alternation of day
(warm period) with night (cool period), caused by the rotation of the
earth about an imaginary axis passing through the North and South Poles;
as we know, this occurs every 24 hours. The second is the annual succes-
sion of seasons, which in the northern hemisphere gives rise to the series
spring (temperate), summer (warm), fall (temperate), and winter (cold).
This second cycle is due to the inclination of the earth’s axis of rotation
with respect to the plane of its orbit, an inclination which is currently
about 23.58.
There are four events which mark the beginning and end of the seasons,
as we know them in the temperate zones of the earth: two equinoxes
(spring, or vernal, and autumnal), and two solstices (summer and winter).
At the equinoxes day and night are of exactly equal length. In the
northern hemisphere, the summer solstice sees the day with the most
40
Climate and Evolution
hours of daylight in the year, while the winter solstice sees the longest
night (see Figure 3.2). On the human scale, these are the only cycles we
perceive, and they are too short to leave geological traces.
The hypothesis that large-scale climate changes were due to natural
fluctuations in the earth’s orbit was first put forward in the 19th century.
However, the development of a quantitative theory relating the orbital
movements of the earth, the level of solar radiation, and the earth’s
climate was the impressive work of engineer Milutin Milankovic
(1879–1958), who dedicated over thirty years to the study of these
phenomena.
If the earth’s orbit around the sun was exactly circular (which it ‘‘vir-
tually’’ is), if the sun was exactly in the geometric center of the orbit
(which it ‘‘virtually’’ is), and the inclination of the earth’s axis of rotation
Daily rotation
Solar N N
radiation N
ator
Equ
E quato r Equ a t o r
Day Night
S
S S
Fall equinox
Vernal (spring)
equinox Trajectory of the
earth in its yearly orbit
Figure 3.2 The cause of the seasons. The inclination of the earth’s north–
south axis of rotation in relation to the plane of its orbit around the sun gives rise
to the seasons. In the northern hemisphere, the days are long and solar radiation is
strong in summer. In winter the days are short, and in addition solar radiation
meets the earth at a tangent (note that the opposite occurs in the southern
hemisphere). Note that the seasons are not directly related to the earth’s greater
or lesser distance from the sun
41
Climate and Evolution
was always 23.58 (as is ‘‘virtually’’ the case), there would be no large-scale
climate changes caused by fluctuations in the level of solar radiation: all
summers would have been the same for thousands of millions of years.
However, all those ‘‘virtually’’s added together, along with some others
we have not mentioned, result in very gradual changes in the amount of
solar radiation reaching the earth each year. According to Milankovic’s
theory, this is the slow but inexorable motor driving climate change.
In addition to equinoctial precession (Figure 3.3), which results in
periods of very hot summers alternating with periods of temperate
solstice,
INT
June
ER
21
Sun
152 million km 147 million km Perihelion
Aphelion
Empty focus
Center of Focus
ellipse Winter solstice, December 21
SU
In 11,000 years
M
ER
M
LL Fall equinox
Fall equinox, FAL
L FA S
September 22
UM
Winter
ME
solstice
R
Sun
Aphelion Perihelion
Summer
WI
solstice
NT
R
E
ING
Vernal equinox SPR
(spring)
Figure 3.3 Precession of the equinoxes. The position of the equinoxes and the
solstices which mark the beginning of the seasons changes slightly each year.
Currently, winter in the north begins when the sun is closest to the earth (the
perihelion); winters are therefore not particularly cold. In about 11,000 years’
time the winter will occur when the sun is farthest away (the aphelion), and
northern winters will be harsher. Thus equinoctial precession periodically tempers
or heightens the intensity of the seasons
42
Climate and Evolution
43
Climate and Evolution
44
Climate and Evolution
The natural presence of water vapor (H2 O), carbon dioxide, and other
atmospheric gases ensures that the average temperature of the earth’s
surface is 158C, rather than 158C: if it were not for the natural green-
house effect, the earth’s surface would be a layer of ice! For this reason,
when we talk of the greenhouse effect as something potentially danger-
ous, we should really use the term ‘‘heightened greenhouse effect’’ – in
other words, the additional overheating which can be caused by emissions
resulting from human activity.
CO2 is the slightly sour, usually harmless gas that gives us the bubbles in
our sodas. It makes up only a very small proportion of the atmosphere,
barely 0.03 percent, but it is nevertheless vitally important: not only is it
the main agent in the beneficial greenhouse effect (when this is not
magnified by human activity), it is also the basic source of organic
carbon – the carbon of which all living beings are made.
With the help of sunlight, plants convert water and CO2 into organic
matter through photosynthesis. Most plants belong to a group known as
the C3 plants, because they fix carbon dioxide through a mechanism
which uses a molecule with three carbon atoms. A minority of plants,
mostly grasses with tough, fibrous stems, belong to another group known
as the C4 plants, since their mechanism for fixing CO2 uses a molecule
with four carbon atoms. A few cultivated crops, such as maize and sugar
cane, belong to this second group.
With the right humidity and temperature conditions, and with abun-
dant CO2 , C3 plants develop much better than C4, but with the current
level of CO2 in the atmosphere, which is very low compared with other
geological epochs, C3 plants find it difficult to survive in hot, dry envir-
onments. Today, C3 plants still predominate in the temperate and cold
climates. In the hot climates of equatorial latitudes, C3 plants grow in
abundance in the humid forests and also in the so-called rain forests
(where an unlimited amount of water is available). The C4 plants, on
the other hand, are grasses and reeds typical of the open, dry, sunny
environments grazed in Africa by zebra and antelopes such as gazelles,
hartebeest, impala, and gnu, and by elephants, hippopotamus, and other
herbivores of the great savannas. As these C4 plants have fibrous, mineral-
rich stems they cause much wear on the teeth of the mammals which eat
them; for this reason their teeth have high crowns, so that they will last a
long time.
45
Climate and Evolution
46
Climate and Evolution
Figure 3.4 The giant gelada Theropithecus oswaldi (right), shown on the
same scale as a female hominid, Paranthropus boisei
forest which extended over much of the Old World becoming fragmented
and declining from the end of the Miocene, and particularly during the
Pliocene and Pleistocene. This loss of habitat probably resulted in the
disappearance of many species of hominoids, although this may not have
been the only cause of the reduction to the very narrow range which exists
today. In the Miocene there were various species of hominoids living
within the same region, whereas today no more than two species live
together. Another major cause of the decline may have been ecological
competition with the other Old World monkeys, the Cercopithecidae,
which are much more abundant and varied today.
However, this same climate change led to the appearance and spread
of more open ecosystems over much of Africa during the late Miocene
and Pliocene, with new species of plants and animals. Among these the
hominids (our ancestors and our closest relatives) soon appeared, as
we shall see in the next chapter. The ancestors of the patas monkey, and
of the baboons and the geladas, also took advantage of this change in
environment. One form of gelada, Theropithecus oswaldi, developed to
enormous size during the Pleistocene (weighing up to 100 kg in excep-
tional cases), and lived in the same zones as humans, who may even have
hunted them and contributed to their becoming extinct (Figure 3.4).
Numerous remains of this giant monkey, together with an enormous
47
Climate and Evolution
48
4
So for every man who has ever lived, in this universe, there shines a star.
Arthur C. Clarke, 2001: A Space Odyssey
Molecular Clocks
Studies in molecular biology indicate that our line separated from the
chimpanzee line between 4.5 and 7 million years ago – in other words, at
approximately the same time that the gradual decrease in atmospheric
CO2 levels was beginning to affect the African ecosystems, as we saw in
Chapter 3. This coincidence makes it tempting to speculate that hominids
originated as a direct result of ecological change and the spread of open
environments, to which they adapted from the beginning. However, as we
shall see later, it now seems that the oldest representatives of our group,
the first hominids, were as much forest-dwellers as are today’s chimpan-
zees, and that the gradual adaptation to drier, less densely forested envir-
onments occurred later.
However this may be, molecular biologists have calculated this
interval between 4.5 and 7 million years using their molecular clocks.
The basis of these molecular clocks is that the genetic difference between
species, such as our own and the chimpanzees, ought to be related to the
time which has passed since the two lines separated. In other words,
genetic divergence increases with time, as does the morphological differ-
ence between two lineages which diverge from one another to follow
different evolutionary paths.
49
The Origin of Humanity
But this assertion that genetic difference is related to the length of time
the lines have been separated is only valid if appropriate genes are chosen
for analysis. The genes suitable for use as molecular clocks are those
known as ‘‘neutral,’’ those which confer neither advantage nor disadvan-
tage, and on which natural selection therefore does not act. In neutral
genes, the naturally occurring spontaneous mutations accumulate at a
constant rate, without being either eliminated or favored, like snow falling
steadily.
On the other hand, the ‘‘non-neutral’’ genes that selection does target
may be modified at different, varying rates depending on the intensity
of the pressure for selection exerted upon them. In other words, if a
particular gene (the correct technical term is an allele) is very beneficial
for the individual that carries it, it is sure to spread rapidly throughout
the population. If, however, it confers a disadvantage, its frequency of
occurrence in the population will fall rapidly because it has natural
selection, a powerful enemy, against it. Furthermore, what is beneficial
today may not be so tomorrow, or may not be so in another species – so
these non-neutral genes are of no use for measuring time in evolution.
To take an example from everyday life, the power and capacity of
personal computers, which are subjected to market pressure for selection,
increase very rapidly, and not at a constant rate. This is a clock which
gains time.
But in order to calculate the rhythm of change of neutral genes, what
is known as the rate of mutation, we have to return once again to
fossils, measuring the genetic difference between two species for which
we know, from fossils, how long their lines have been separated. For
example, in order to work out how long it is since the human and
chimpanzee lines separated, we can use the human/orangutan pair.
Measuring the genetic distance between the two is the easy part, although
since not every gene can be used the calculations vary depending on
which ones are chosen. Establishing when the orangutan line separated
is a very different matter. Sometimes the figure of 13 million years is
used: this corresponds to the first fossils attributed to the species
Sivapithecus, which themselves are believed to mark the beginning of the
evolution of orangutans.
In other words, in order for the ‘‘molecular clock’’ to work, many
things are needed: genes which natural selection does not ‘‘see’’ but
which are known to us, constant rates of mutation, and a good paleonto-
logical reference point – too many things – but nevertheless, this interval
of between 4.5 and 7 million years for the human/chimpanzee separation
is acceptable to paleontologists, as we shall see shortly.
50
The Origin of Humanity
51
Bahr el Ghazal Australopithecus bahrelghazali
Figure 4.1 Location of the main deposits containing fossils of Australopithecus and Ardipithecus
The Origin of Humanity
Change of Habitat
53
The Origin of Humanity
Although hominids are not the only primates living in the savannas and
grasslands, the erect posture is a completely new innovation.
The next million years (roughly speaking) correspond to another East
African species, known as Australopithecus afarensis (which can be trans-
lated as ‘‘australopithecine of the Afar region’’). Most of the fossils of this
species have been found in the Hadar area, on the lower reaches of the
Awash River (in the Afar region, Ethiopia), and in Laetoli (Tanzania).
The Laetoli fossils, which include a mandible classified as the type
specimen (holotype) of the species (known as L.H. 4), are dated at 3.5
million years old; the Hadar fossils are between 3 and 3.4 million years
old. Donald Johanson’s team, working in Hadar since 1972, has discov-
ered numerous remains, so that Australopithecus afarensis has a reasonably
complete fossil record. This includes the skull of a male (A.L. 444-2),
discovered by Yoel Rak in 1992 (Figure 4.2), and a large part of the
skeleton of a female (A.L. 288-1), known worldwide as Lucy, as she was
named by Johanson when he found her in 1974.
An assemblage of fossils which includes the most complete known
mandible of this species was found in the Maka deposit, 3.4 million
years old, on the middle reaches of the Awash River. Some teeth a little
over 4 million years old found in Fejej, southern Ethiopia, and a fragment
of frontal bone from Belohdelie in the Middle Awash, approximately 3.9
million years old, have also been attributed to Australopithecus afarensis.
However, the assignation of these fossils to the species is subject to review
and they could, given their chronology, belong to Australopithecus
anamensis.
The associations of vertebrates in the Hadar deposits suggest changes in
the paleoecological conditions over the 400,000 years of geological his-
tory recorded there. Australopithecus afarensis appears to have lived both
in a fairly dry forest environment and in a cool savanna with corridors of
forest along the river valleys – in other words, neither a humid forest nor
an arid steppe, but rather an intermediate habitat.
One of the fundamental problems of paleontology is that of grouping
fossils into species, since unfortunately the remains do not appear with
labels in the deposits. It is the paleontologist’s job to find their place in
evolution.
Living species which are related to one another are often morphologic-
ally very similar, or may differ only in external features such as color, fur,
behavior, and other characteristics which, although they may be very
striking, are not reflected in the skeleton, the part that becomes fossilized.
Ian Tattersall has pointed out that many modern species of primates
would not be recognized as different species if we only considered their
54
The Origin of Humanity
Sagittal ridge
Nuchal ridge
55
The Origin of Humanity
56
The Origin of Humanity
If the sample from deposit A.L. 333 had included only large individuals,
or only small ones, or individuals with a particular morphology, then
we might conclude that fossils of different species had been artificially
included in Australopithecus afarensis. Conversely, if the sample included
all the sizes and morphotypes found at Hadar and Laetoli, we could be
sure that Australopithecus afarensis is a real species and not a hodgepodge
of odds and ends. In fact, there is wide variation within the A.L. 333
sample – as wide as has been suggested by the researchers who identified
Australopithecus afarensis.
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The Origin of Humanity
known up to this day have been found. However, at the end of 1995
Michel Brunet and his colleagues published the discovery north of N’Dja-
mena, in Chad, of the front portion of an australopithecine mandible and
an isolated premolar, which are dated to between 3 and 3.5 million years
old on the basis of the accompanying fauna. The authors suggested, in a
subsequent study, that this was a species different from Australopithecus
afarensis, which they named Australopithecus bahrelgazali (the term bah-
relgazali refers to the region of Chad in which the fossils were found –
Bahr el ghazal, which in Arabic means ‘‘river of the gazelles’’). This
discovery suggests that, early on in their history, hominids spread far to
the west of their East African birthplace – if this is in fact where they
originated.
The African deposits we have discussed thus far, which are lacustrine or
fluvial sediments (from lakes or rivers), usually contain aquatic species
such as turtles, crocodiles, fish, and hippopotamus. This does not help
us to determine in what kind of ecosystem the hominids lived: other
animals, not aquatic or amphibian, are more useful to us. This is no simple
matter, for a sedimentary basin may contain the remains of animals that
lived and died in very different environments, and have been transported
there by streams and rivers. Thus, everything accumulates at the bottom
of the basin, creating many problems for paleontologists, who try to
resolve them principally on the basis of common sense. The application
of common sense to the study of the formation of such deposits consti-
tutes a paleontological discipline in itself, a very important one, known as
taphonomy. Using taphonomy we can establish, for example, whether a
bone has been transported over a long distance or whether the animal
died close to where it is found. Fortunately, the study of the adaptations
found in fossil species, or functional paleomorphology, also contributes to
determining the place of these species in the ecosystem (their niche), and
what their environment was like.
We shall return to the question of the environment in a later chapter
when we look at the diet of the first hominids. For now let us take a
moment to consider the fossils associated with Australopithecus bahrelga-
zali. Michel Brunet imagines that these australopithecines lived in a
variety of environments, including forest corridors (characterized by the
presence of bushpigs), wooded savanna with elephants, and grasslands
where rhinoceros grazed.
The main differences between the Chad fossil and Australopithecus
afarensis are in the interior face of the symphysis, or anterior (front)
part of the jawbone, which has a fairly flat and vertical surface, without
the strong transverse reinforcements or tori (ridges – from the Latin torus)
58
The Origin of Humanity
Dating Fossils
The reader has perhaps already wondered how we know the age of these
fossils of our very distant ancestors. The paleontologist Yves Coppens,
who gives many lectures, is often asked this question, to which he recom-
mends the reply: ‘‘Trust us, we know the age of fossils, we have methods
for determining it, we are professionals.’’ At the risk of boring the reader,
we shall disregard his advice and attempt a brief discussion of this funda-
mental question.
Because of the intensity of the internal forces which come into play,
fracturing of the earth’s crust is often accompanied by volcanic activity,
which may cause ash to be thrown up into the air during the course of
eruptions. Winds and water transport the ash, which is finally deposited in
beds intercalated between the layers of sediment which contain the fossils.
These layers of volcanic ash, or tuffs, are very useful in correlating and
dating formations. Even two successive eruptions from the same volcano,
with very little time between them, have distinct characteristics, known as
their ‘‘chemical fingerprints.’’ Using chemical analysis, we can compare
two layers of volcanic tuff and find out whether they are the same, even if
they do not run continuously in the field because the rocks are fractured
into blocks, as is typical of the geology of rift valleys.
During the 1980s a method of dating these tuffs was invented, based
on melting a single small crystal of a mineral of the potassium feldspar
group using a laser beam. When the crystal melts, the laser releases a given
59
The Origin of Humanity
60
The Origin of Humanity
minerals such as flint or quartz, like teeth and bones, act as natural Geiger
counters, accumulating the radiation received over time.
Another method used in the measurement of geological time (geo-
chronology) is paleomagnetism. The earth acts as a magnet, creating a
geomagnetic field around it, with two poles (north and south). This field
orients the compass needle, indicating the position of the magnetic
north pole, which is today close to the geographic North Pole. But it
also orients ferrous clay minerals, like tiny compasses, provided that
they are deposited slowly in a calm environment, such as an undisturbed
pool or lake.
Over extremely long periods, the magnetic poles exchange position (so
that the magnetic north pole is located close to the geographic South
Pole). The location of the poles at any given moment is recorded in the
clay minerals making up the layers of sediment. These changes in the
earth’s magnetic polarity have been dated, enabling us to establish a
scale showing the sequence of alternating periods of one or other type
of polarity. Each of these long periods of time is called a chron. A subchron
is a shorter unit of time, with a polarity opposite to that of the chron
within which it occurs; even shorter periods are known as excursions.
Paleomagnetism cannot give us an absolute date for a formation, but it
can help with dating when combined with other methods.
In addition to these methods of dating, the fossils of the animals
associated with the hominids also help to establish their relative age,
since the evolution of species means that animals, including hominids,
change over time. Thus biochronological scales can be created, which can
be calibrated with the absolute dating obtained by physical methods.
61
The Origin of Humanity
Distinguishing Marks
62
The Origin of Humanity
63
The Origin of Humanity
64
The Origin of Humanity
stagnated among those who opted for the ‘‘comfort’’ of the forest and its
abundant fruits, or the warm African continent.
Although Dart’s interpretation of our origins was not universally
accepted, he was not alone. In a famous book entitled Man-apes or Ape-
men: The Story of Discoveries in Africa, published in 1967, Sir Wilfrid Le
Gros Clark (1895–1971) opined that the australopithecines were too
poorly defended, with their small canine teeth, to survive without
weapons, whether of stone, bone, horn, or teeth. According to Le Gros
Clark, these hominids were hunters and carrion-eaters cast into a hostile
environment; however, walking on two feet left their hands free to ma-
nipulate tools, and it was this that was the stimulus for the development of
intelligence. Le Gros Clark was a great authority on human evolution, and
it was he who, following a trip to South Africa in 1947 to see the original
fossils, helped to change the opinion of the majority of the scientific
community, allowing the australopithecines to be recognized as primitive
members of our line.
In short, the hunter hypothesis suggests a somewhat bloody beginning
to human evolution, though it is nevertheless a beginning. But was this
how things began? Have we been ‘‘murdering apes’’ and makers of tools
from the start, perhaps even before we became bipedal?
In order to answer this question, we shall address the question of
australopithecine locomotion in the next chapter. Later we shall look at
other forms of hominids, Paranthropus, and the first humans, together
with the first stone tools, and then we will discuss evolutionary changes in
intelligence, diet, growth, and sociability.
65
5
The human being is not the only mammal capable of walking on its hind
limbs. We have already noted that anthropoid apes are in the habit of
maintaining their trunk vertical as they move through the trees hanging
by their arms, or simply when they are sitting. But holding the trunk erect
is only half the battle in achieving erect posture and walking on two legs.
The other half involves aligning the legs with the trunk – in other words,
extending the entire body. The great apes sometimes walk on two legs,
but although they hold their trunk almost vertical, they keep the hip and
knee joints flexed, just as when they walk on four legs. Only we humans
are able to take stable steps without large movements of the trunk, and
long strides when we walk, extending our legs far behind the hip; other
mammals take only small, wobbly steps, with large shifts of the trunk.
Part of the reason for this major difference lies in the pelvis. When we
are standing still, more or less steady, the body is stable and the pelvis is
66
The Bipedal Chimpanzee
Figure 5.1 Position of the gluteus medius muscle in humans and chimpanzees.
In humans the muscle fibers are oriented laterally, and the muscle therefore acts
as an abductor, balancing the pelvis when the opposite foot moves forward. In
chimpanzees the fibers are oriented toward the back, so that the muscle acts to extend
the joint between the pelvis and femur
67
The Bipedal Chimpanzee
68
The Bipedal Chimpanzee
Gluteus maximus
Gluteus medius
Gluteus
maximus
Gluteus
minimus
push themselves out of the starting blocks, where they position themselves
in a quadruped posture to begin with, with the hip joint highly flexed,
before moving immediately into a bipedal posture.
In apes, as in other mammals, the function of the gluteus medius and
gluteus minimus is different from our own, because the muscles have a
different line of action. In chimpanzees and gorillas the three gluteal
muscles (maximus, medius, and minimus) act as hip extensors, never as
abductors; this means that habitual bipedal walking is not possible, but it
favors quadrupedal locomotion.
We have already noted that the function of a muscle is determined by its
line of action. Why is the line of action of the gluteus medius and minimus
different in humans? The answer, to put it simply, is the orientation of the
bony region in which both muscles originate, the iliac crest of the pelvis.
The pelvis is composed of the two coxal bones, one on either side,
and the sacrum at the back, which in its turn is formed by the sacral
vertebrae (the sacrum is thus part of the spinal column and continues into
the coccyx). The coxal bone is actually the result of the fusion of three
bones during adolescence – the ilium, which forms the upper part of
the coxal bone, the ischium, which forms the lower posterior part, and
69
The Bipedal Chimpanzee
the pubis, which represents the lower anterior part. These three bones
meet in the acetabulum, where the coxal bone articulates with the head of
the femur (see Figure 10.1).
Of all parts of the postcranial skeleton, i.e., the skeleton excluding the
skull and jawbone, the pelvis is probably that which most distinguishes
humans from apes. This is obviously the result of our particular mode of
walking, since the pelvis of apes is not substantially different from that of
other quadruped mammals. Let us look at the main innovations in the
revolutionary architecture of the human pelvis, and consider their biome-
chanical significance.
In quadrupeds the weight of the trunk is transmitted through the four
limbs, but in humans, because we are bipedal, it is transmitted through
the spinal column down to the sacrum, and from there through the coxal
bones to the heads of the two femurs, continuing down to the feet. In
walking, during the phases when we are supported on only one leg almost
all of the body’s weight is transmitted through one coxal bone (on the
supporting side). In order to reduce the stress on the bar of bone running
from the sacro-iliac joint to the coxo-femoral joint, the two joints have
become much closer than they are in chimpanzees and other quadrupeds.
This biomechanical improvement of the pelvis has, as we shall see later, an
undesirable side effect: it greatly complicates childbirth because it reduces
the size of the bony channel through which the full-term fetus has to pass
in order to be born.
The three gluteal muscles originate in the iliac crest and insert in the
femur. The iliac crest forms the larger part of the iliac bone or ilium, and
in apes consists of a high, narrow plate of bone. In humans it is propor-
tionally shorter, because, as we have already noted, the joints between the
coxal bone and the spine and femur are closer together, but most import-
antly, it is relatively much wider than in other apes.
The other major difference in the human iliac crest is its orientation (see
Figure 10.1). In quadrupeds the surface on which the gluteal muscles
originate faces directly backward. This orientation means that contraction
of the gluteal muscles can only produce extension of the hip joint, because
its line of action is posterior: in other words, if the legs are stable they
draw the pelvis backward, while if the pelvis is fixed it draws the legs
backward.
In humans, however, the orientation of the iliac crest has changed
radically, so that the surface where the gluteus medius and minimus
originate faces toward one side rather than backward. The biomechanical
result of this is that when the hip pulls sideways, the contraction of these
two muscles produces the effect of abduction, making bipedal walking
70
The Bipedal Chimpanzee
possible, as the trunk can be balanced at each stride. The gluteus max-
imus, on the other hand, which originates in the hindmost part of the
ilium and in the sacrum, continues to act as an extensor in humans, since
its line of action is posterior rather than lateral.
One of the biggest questions of evolutionary biology is how the great
anatomical transformations which create organisms radically different
from their predecessors come about. A bipedal primate represents a
revolutionary change, not simply a slight variation on other types of
hominoids. We have to rule out the idea that the entire skeleton altered
drastically all at once, but it is not easy to imagine how an organism can
move gradually from walking on four legs to walking on two. One inter-
esting hypothesis is that the initial modification which made it possible for
primates to begin walking bipedally was a change in the orientation of the
iliac crest. Merely altering in this orientation, causing the iliac crest to face
more to the side, would generate some capacity for abduction, one of the
basic features of bipedalism. If walking on two legs increased the possi-
bilities for surviving and reproducing, other modifications would subse-
quently continue to be selected until the entire skeleton was affected.
We will not exhaust the reader with any more biomechanical explan-
ations, although this is a fascinating discipline which allows the paleon-
tologist to play God for a moment, and say to a fossil skeleton: ‘‘Get up
and walk!’’ Just one final detail on the pelvis: in humans the capacity for
extension of the hip, which is very useful for climbing trees and for
walking on four legs on the ground, is also reduced in the hamstring
muscles, which originate in the lower part of the pelvis (Figure 5.3).
Let us turn now to the fossils. So far we have been engaging in an
exercise of comparative anatomy, the best starting point for paleonto-
logical analysis. We have a limited sample of fossil pelvises from australo-
pithecines. The most complete is that of Lucy, which has retained the
sacrum and all of the left coxal bone, and those of two Sterkfontein
skeletons, Sts 14 and Stw 431 (the latter is fairly fragmented and has
not yet been analyzed). Although they show some distinctive character-
istics, their morphology is entirely human and does not resemble that of
apes in any way. The ilium is short and wide (favoring abduction), and the
ischium is short (reduced capacity for extension). Furthermore, the sac-
rum in these pelvises is proportionately wide, not narrow as in apes.
Various authors offer different interpretations of the exact orientation of
the iliac crest, which has had to be reconstructed in Sts 14 and Lucy
because both had been distorted by being crushed in the deposits. Some
authors suggest that the iliac crest in these fossils faces less to the side
and more toward the back than in modern humans, but all researchers
71
The Bipedal Chimpanzee
72
The Bipedal Chimpanzee
73
The Bipedal Chimpanzee
Cervical curve
Sagittal ridge (lordosis)
Nuchal ridge
Orientation of the
Orientation of the
foramen magnum
foramen magnum
Thoracic curve
(kyphosis)
Lumbar curve
(lordosis)
Figure 5.4 Curves of the spine and orientation of the foramen magnum in
gorillas and humans
while others believe that it was less ‘‘perfect.’’ The discussion might
have continued ad infinitum had not the team directed by Mary Leakey
(1913–96) made a completely unexpected discovery in 1978 and 1979:
several meters of tracks formed by footprints left by three hominids
who walked through an area of what is now Tanzania about 3.5 million
years ago.
The Laetoli Beds are located not far from the famous Serengeti National
Park. During one of its eruptions a nearby volcano, Sadiman, threw ash
out into the air, and rain converted this to mud in which the tracks of
many animals were recorded and fossilized. Among these animals were the
hominids we have mentioned. There are two parallel tracks, but that on
the right (looking in the direction of walking) appears to be that of two
74
The Bipedal Chimpanzee
individuals, one of them walking in the tracks of the other. The individual
on the left is very small, perhaps a female or a child.
The characteristics of these footprints are incredibly modern. They
show no sign of insecure or ‘‘imperfect’’ bipedalism; rather they indicate,
even in the smallest details, a way of walking identical to our own. The
foot of a chimpanzee, gorilla, or orangutan is very different from a human
foot. In fact it is more like our hand – flat, with a big toe which is shorter
than the others and can separate from them laterally, and extends away
from the other toes on each step (Figure 5.5).
In the Laetoli footprints we can study the mode of walking of the
hominids which produced them. If you are fortunate enough to have a
beach nearby, you can compare these fossil footprints with your own, and
appreciate their extraordinary similarity. In each step taken by the Laetoli
hominids the front foot was supported first on the heel, leaving a deep
impression in the soft ground. Part of the body weight was then trans-
ferred through the arch or instep. After this, the foot flexed over the toes,
giving a final impulse to lift the foot from the ground and extend the leg
forward. As in modern humans, the big toe was fundamental to this final
phase (being the last to leave the ground), and it therefore pointed
forward like the other toes and was aligned with them (Figure 5.5).
Fossils of Australopithecus afarensis of the same age as the footprints
have been found in the Laetoli Beds, and we can therefore only suppose
75
The Bipedal Chimpanzee
76
The Bipedal Chimpanzee
77
The Bipedal Chimpanzee
through the trees and over the ground. It appears that gorillas later largely
abandoned life in the trees when they became very heavy.
Peter Wheeler has identified a further advantage in the shift of the body
to the vertical, related to body temperature. An individual standing on
two feet receives less solar radiation, especially when the sun is at its
highest point, than a quadruped. Moreover, when the body leaves the
ground it moves away from a focus of heat and benefits from cooling
breezes. Combining this advantage with that we noted above, we might
conclude that bipedal locomotion is perhaps the best solution for a
hominid who has to travel long distances while exposed to the sun. The
first bipedal hominids did not live in the savanna, but they might never-
theless have had to travel between patches of vegetation separated by open
spaces.
One indisputable benefit of bipedalism is the ability to use the hands
and arms for carrying, whether food or perhaps children. In this, quad-
rupeds are at a clear disadvantage to us. We shall return to this capacity
for carrying in the chapter on the social biology of the first hominids,
because one proposed explanation for the origin of bipedalism is partly
based on this factor. Without jumping ahead, we can already anticipate
that such an explanation will have many problems, in terms of what we
intuit about the social biology of the first hominids. Thus we can still
today consider the significance of our acquisition of our special way of
traveling as one of, if not the biggest problem confronting human
paleontology.
Lucy was a very small individual. She was about 105 cm tall and weighed
no more than 30 kg. But she was not exceptional among her species. Even
the first fossils discovered by Donald Johanson in 1973, consisting of a
knee joint (the lower end of the femur and the upper end of the tibia),
correspond to an individual of the same size as Lucy; some of the hom-
inids from site 333 are of similar size (as we noted above, this assemblage
of fossils was christened the First Family). Other fossils are larger and
correspond to individuals around 135 cm tall, weighing about 45 kg. It is
thought that the small individuals, like Lucy, were females, and the larger
ones males. The male and female averages may have been somewhat closer
in Australopithecus africanus, but in any case australopithecines were small
hominids, similar in size to chimpanzees.
78
The Bipedal Chimpanzee
79
The Bipedal Chimpanzee
80
The Bipedal Chimpanzee
81
The Bipedal Chimpanzee
82
The Bipedal Chimpanzee
83
The Bipedal Chimpanzee
84
The Bipedal Chimpanzee
85
The Bipedal Chimpanzee
86
6
Paranthropus – Hominids of
the Open Plains
Gert was found, and drew from his trouser pocket four of the most
wonderful teeth ever seen in the world’s history.
Robert Broom, The South African Fossil Ape-Men
87
Konso Paranthropus boisei
fossils he had found in the South African Kromdraai deposit. The term is
very appropriate, since Homo and Paranthropus emerged during the same
era and lived alongside one another for about 1.5 million years, inhabiting
a vast region extending from present-day Ethiopia to the southern tip
of Africa.
The majority of paleontologists believe that there were three species
within the genus Paranthropus. Two of these are exclusively East African:
Paranthropus aethiopicus (aethiopicus refers to Ethiopia, the country
where the first remains of this species were found) and Paranthropus boisei
(the term boisei is in honor of an English sponsor named Boise); the third
has been found only in deposits in the south of the continent, and is
known as Paranthropus robustus (robustus meaning ‘‘robust’’ in Latin).
Although the first remains attributed to the genus Paranthropus –
specifically to Paranthropus robustus – appeared in South Africa, the oldest
known fossils were found thousands of kilometers away. In 1967, Camille
Arambourg (1885–1969) and Yves Coppens published details of a hom-
inid jawbone (Omo 18-1967-18) found in one of the deposits of the
lower reaches of the Omo River (close to where it flows into Lake
Turkana), in Ethiopia; the age of these fossils was established as 2.6
million years. This discovery passed almost unnoticed until, in 1986,
Richard Leakey and his collaborators published the find of an extraordin-
ary skull (WT 17000) belonging to the same species as the jawbone found
by Arambourg and Coppens two decades before, in one of the deposits on
the eastern shore of Lake Turkana. This fossil is one of the amazing
‘‘missing links’’ which paleontologists are occasionally lucky enough to
come upon, since it documents an evolutionary stage intermediate be-
tween Australopithecus afarensis and Paranthropus boisei.
Dated at around 2.5 million years old, WT 17000 is the most repre-
sentative fossil of the species Paranthropus aethiopicus (Figure 6.2). Many
of the features typical of the anatomy of Paranthropus are not yet present
in this species, while others are only in the incipient stages of develop-
ment. The most recent fossils attributed to Paranthropus aethiopicus are a
series of remains of skulls and mandibles found in the river Omo deposits,
and dated at around 2.3 million years old.
The first fossil of Paranthropus boisei was found in 1959, in Olduvai
Gorge, Tanzania, by Mary Leakey and Louis Leakey (1903–72), Richard
Leakey’s parents. It consists of an almost complete skull which its finders
dubbed ‘‘Dear Boy,’’ since it belonged to a young specimen. The age of
this fossil (known as OH5) is around 1.8 million years, and the Leakeys
attributed it to a new species and genus, Zinjanthropus boisei, although it
came to be included in the Paranthropus genus soon afterward.
89
Paranthropus – Hominids of the Open Plains
Sagittal ridge
Temporalis
muscle
Mandibular
ramus Mandibular
body
Masseter
muscle
90
Paranthropus – Hominids of the Open Plains
91
Paranthropus – Hominids of the Open Plains
92
Paranthropus – Hominids of the Open Plains
Temporalis
Zygomatic bone
Zygomatic arch
Maxilla
Masseter
Temporalis
Zygomatic bone
Zygomatic arch
Maxilla
Masseter
Figure 6.4 Bones and muscles involved in mastication. The temporalis muscle
is fan-shaped. Its wider end originates on the external surface of the side walls of the
cranium, while the narrower part passes through the temporal fossa to insert in the
mandible. Contraction of these muscles can easily be felt by palpating the temples
while clenching the teeth. The masseter muscles extend from the lower edge of
each zygomatic arch to the outer faces of the mandibular ramus and the posterior
parts of the mandibular body (see Figure 6.1). It is also easy to locate these
muscles by touching the cheekbones while clenching the molars
93
Paranthropus – Hominids of the Open Plains
The Specialist
94
Paranthropus – Hominids of the Open Plains
us, noting that vertical movements of the mandible are generated basically
by two groups of muscles located symmetrically at either side of the head:
the temporalis muscles and the masseter muscles (Figure 6.4). The hori-
zontal movements of the mandible, on the other hand, depend principally
on the action of the lateral pterygoid muscles.
The three species of Paranthropus show different degrees of specializa-
tion of the masticatory apparatus, which is very marked in Paranthropus
boisei, less pronounced in Paranthropus robustus, and only sketchily ap-
parent in Paranthropus aethiopicus.
Beginning with dentition: the molars of Paranthropus have a fairly thick
layer of enamel, thicker than that of any other hominid or ape. This
characteristic is clearly related to a diet which causes heavy wear on the
molars, either because the food contains hard particles or because it
requires prolonged chewing, or both. Moreover, the surface of the teeth
used to crush food is very large in relation to body weight. This is due to
the increased size of the molars and particularly to the fact that the
premolars are ‘‘molariform’’ – they are shaped like molars, substantially
increasing the total area of teeth dedicated to crushing food. Parallel with
the increase in the size of the molars and premolars, the anterior teeth of
Paranthropus, the canines and incisors, are much reduced in size.
In apes the premolars are not ‘‘molariform,’’ so the area of the teeth
dedicated to crushing food is restricted to the molars (and the reader will
recall that the incisors and canines are large). In Paranthropus, in contrast,
the mastication zone has extended to the premolars, which in these species
are ‘‘molariform.’’
The maxilla (upper jawbone) of Paranthropus is further back under the
cranium than is usually seen in apes and australopithecines, and
the premolar area is thus also further back. The zygomatic bones, on the
other hand, are further forward, bringing the zone in which the masseter
muscles originate further forward. These modifications increase the
strength of the bite in the premolar area (Figure 6.4).
Furthermore, the force exerted by a muscle is directly related to the
number of muscle fibers it contains – in other words, to its thickness.
There is a physical limit on the size of the temporalis muscle: it has to pass
through the temporal fossa, the diameter of which determines the max-
imum thickness of the muscle. In Paranthropus the zygomatic arch (which
forms the lateral end of each of the temporal fossae) is strongly curved
outward, markedly increasing the diameter of each fossa and indicating
extremely thick and powerful temporalis muscles. These muscles were so
big in Paranthropus that in males they result in the appearance of a sagittal
ridge along the mid-line of the upper surface of the cranium.
95
Paranthropus – Hominids of the Open Plains
96
7
The term homo (‘‘man,’’ in the generic sense of ‘‘human being’’) was used
by Linnaeus in 1758 to denote the genus to which our own species (Homo
sapiens) belongs. The terms ‘‘humanity’’ and ‘‘human’’ are usually
restricted to representatives of our genus, so when we explore the origin
and evolution of the genus Homo we are referring to the origin and
evolution of humans.
Until the early 1990s the oldest fossils assigned to our genus consisted
of an assemblage of isolated teeth (around 2.1 million years old) and
jawbone fragments (around 2.5 million years old) found in the deposits
of the Omo River valley. However, the allocation of these fossils to the
genus Homo was disputed by many authors, who called for more solid
evidence that Homo did in fact exist more than 2 million years ago. Those
who believed that the genus was in existence at this time based their
argument on the identification of stone tools in sediments about 2.3
million years old, also in the Omo deposits. But this was a debatable
argument based on a premise which had yet to be proved: that the only
hominids capable of manufacturing stone tools are members of the genus
Homo.
97
A New Kind of Hominid
Since the early 1990s, a series of fossils over 2 million years old, and
attributed to our genus, has been presented to the scientific community.
In 1992 it was reported that a fragment of temporal bone (known as
BCI, and found in 1965 in a deposit close to Lake Baringo in Kenya)
belonged to our own genus; this was dated at around 2.5 million
years old. However, as we do not know exactly where it was found,
its position in the stratigraphy of the deposit cannot be determined, and
its age cannot be reliably established. Moreover, not all researchers
accept that this fossil belongs to the genus Homo. In our view, the
Lake Baringo fossil does belong to our genus, although it cannot be
assigned to a particular species. A very complete mandible (UR 501),
found by Schrenk and Bromage’s team on the western shore of Lake
Malawi, is also attributed to Homo (specifically, to Homo rudolfensis).
This fossil is dated at between 2.3 and 2.5 million years old, on the basis
of paleontological data; this age is considered approximate and could be
reappraised. Again, the attribution of this mandible to Homo cannot
be considered definitive, given that, as we understand it, it could be that
of a Paranthropus.
In 1994 Johanson’s team found a very complete maxilla (A.L. 666-1) in
the Hadar region; the allocation of this find to the genus Homo seems
clear. The fossil was found immediately below a volcanic tuff dated at
around 2.3 million years old on the basis of radiometric techniques.
Around twenty stone tools were found together with the human fossil;
this represents the oldest known case of an association of stone tools with
hominid remains. This find has reinforced the hypothesis that it was
representatives of Homo who made the first tools. The oldest known
worked tools were found in another deposit in the Hadar region, Gona,
which is dated at around 2.5 million years old.
Systematic dressing of stone to obtain tools appears to have been one of
the keys to our genus’s ability to inhabit very varied environments and to
gain access to new resources. It is therefore worth leaving bones aside for a
moment to consider stones.
Humans are not the only animals to use tools. Chimpanzees also do so,
but their tools are natural objects suited to their function, such as the
sticks they prepare by stripping away the side shoots and then insert into
the openings of termites’ nests to ‘‘fish’’ for the insects. In these cases the
98
Hadar Homo
Figure 7.1 Location of deposits containing fossils of the first representatives of the genus Homo
A New Kind of Hominid
Figure 7.2 Some typical Mode 1 stone tools. Top: two views of a chopping
tool; below: two views of a polyhedron
100
A New Kind of Hominid
even making such simple tools is more difficult than it seems. Archeologist
Nick Toth and psychologist Sue Savage-Rumbaugh conducted an experi-
ment to teach a male bonobo (pygmy chimpanzee) named Kanzi, who has
shown remarkable capacity for manipulating symbols, to carve stone. (We
shall return to Kanzi later, in the chapter on language.)
In front of Kanzi, Nick Toth struck a pebble with another to obtain
flakes; with the edge of the flakes he then cut a string and opened a box
containing food. Kanzi immediately understood the usefulness of the
flakes and learned to select the best ones and to use them, but when it
came to producing them all his efforts were in vain. There is no doubt that
Kanzi has in his mind the idea of the flake he wants to obtain, but he is
unable to judge how to strike one stone against the other. The reader may
also try to do it: you will see that it is not so easy, particularly if you value
your fingers! In order to carve it, the stone needs to be held steady with
one hand and struck on its edge, slightly at a tangent, with another stone
which acts as a hammer or mallet. Flakes will come away from the
stone which is struck, leaving a carved core. Both the flakes and the core
can be used as tools, because they have edges. Nevertheless, Kanzi
discovered that by throwing a stone against the ground or against a rock
the stone would break apart and produce usable edges, although the result
did not resemble prehistoric tools.
As we have already noted, chimpanzees’ hands are not very well adapted
to manipulating small objects with precision, because of the length of the
palm and all the fingers except for the thumb, the tip of which is some
distance away from the tips of the other fingers. It is a hand made for
hanging from branches, not for carving. However, it is likely that the main
problem lies in the inability to coordinate the necessary movements of the
arms, wrists, and hands.
The australopithecines, at least since Australopithecus afarensis, have
hands which are already very similar to our own, and we may therefore
suppose that they had the mental capacity (just as chimpanzees do) and
the necessary coordination to produce flakes. If they did not do so, it is
perhaps because they did not need sharpened edges to cut with. It may be
that this need arose when the first humans began to eat meat, and
required edges both to cut the thick skin of large animals and to cut
tendons and slice up muscle. They also used stones to break open bones
and extract the marrow, although this activity is similar to that practiced
by chimpanzees when they open nuts using a stone as a hammer and
another, fixed on the ground, as an anvil. The animals that humans ate
would not necessarily have been hunted by them: often they would have
been carrion. They probably also used the sharp edges of stone tools to
101
A New Kind of Hominid
cut plants, and some believe that what Paranthropus did to prepare tough
plant material using their hyperdeveloped chewing apparatus, the earliest
Homo did with their stone tools.
The fossil record of Homo from deposits less than 2 million years old is
much richer and more varied. Until a few years ago, all the African fossils
dated between 2 and 1.4 million years old were attributed to two species.
The oldest and most primitive were assigned to the species Homo habilis,
while the more recent and more evolved were seen as African representa-
tives of Homo erectus, a species defined on the basis of fossils found in Java
and China (which we shall consider later). However, since the 1980s
the discovery of fossils with different morphology has led scientists to
rethink the number of species represented by African fossils over one
million years old.
Bernard Wood, who studied the cranial remains and fossil teeth from
Koobi Fora, has been the main agent of this revision. In his opinion, three
species can be distinguished among the oldest human fossils of Africa:
Homo habilis, Homo rudolfensis (the name rudolfensis refers to the old
name for Lake Turkana, Lake Rudolf), and Homo ergaster (ergaster mean-
ing ‘‘worker’’ in Greek). The fossils previously attributed to Homo habilis
are divided between the first two species, while the latter includes virtually
all the remains initially assigned to Homo erectus (with the exception of
skull OH 9, of which more later).
According to Wood, Homo rudolfensis is distinguished by the combin-
ation of a large brain with highly developed masticatory apparatus, similar
in some respects to that of Paranthropus. Homo habilis, on the other hand,
had a brain somewhat larger than that of australopithecines and Paran-
thropus, but smaller than that of Homo rudolfensis, combined with a
primitive overall architecture of the skull very similar to that of Australo-
pithecus africanus. It was distinguished from the latter by a less developed
and shorter masticatory apparatus, the smaller size of the molars and the
presence of a brow ridge separated from the rest of the frontal bone by a
shallow furrow (Figure 7.3).
Nevertheless, in our opinion the main fossils used to justify the exist-
ence of Homo rudolfensis could easily be included in Homo habilis if we
accept that the species showed a marked sexual dimorphism, like the
australopithecines and Paranthropus.
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A New Kind of Hominid
Brow ridge
103
A New Kind of Hominid
104
A New Kind of Hominid
105
A New Kind of Hominid
Brow ridge
Nasal bones
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A New Kind of Hominid
would have been more than 1.80 m tall when he finished growing. Another
very interesting feature which can be reliably established for WT 15000 is
the ratio between the lengths of the humerus and femur (Figure 7.5). This
ratio is 74 percent, entirely human and contrasting with that estimated for
the Homo habilis fossil OH 62 (95 percent).
Two very complete skulls known as ER 3733 and ER 3883 were found
at Koobi Fora. The first of these was found in 1975, in sediments around
1.8 million years old; the second was found a year later in another, more
recent stratum, around 1.6 million years old. Femurs ER 1472 and ER
1481, and coxal bone ER 3228, found at Koobi Fora, and coxal bone OH
28 from Olduvai, may also belong to Homo ergaster.
Outside of East Africa Homo ergaster is represented by an assemblage of
fossils recovered in the South African deposit of Swartkrans (in the same
sediments which contained fossils of Paranthropus robustus, about 1.8
million years old); these include remains of jawbones, teeth, and part of
a skull (SK 847).
Homo ergaster appears as the most human of the early species of Homo.
In addition to its large brain, this species exhibits a height and limb
proportions similar to those of later humans. Moreover, shortly after
Homo ergaster emerged, a new form of stone-dressing appeared, known
as the Acheulian industry.
The Acheulian, or Mode 2 industry, much more elaborate than the
Oldowan or Mode 1 culture, includes cores or large flakes worked on two
sides, known as bifaces and including handaxes, cleavers, and picks. These
tools show a marked degree of standardization in manufacture, and they
require a long sequence of gestures to make them, including turning the
core in the hand while continuing to strike it with the hammer to obtain
flakes. The result is a tool in which all or almost all of its edge forms a
blade. Handaxes are symmetrical bifaces with lateral cutting edges which
converge to a pointed tip; cleavers are bifaces with a straight edge on one
end (Figure 7.6). The handaxes were probably multipurpose tools, used
to cut meat, work wood, and perhaps also to prepare skins. The oldest
Acheulian industry known is around 1.6 million years old, and was found
in the Olduvai deposit in Tanzania.
Family Relationships
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A New Kind of Hominid
108
A New Kind of Hominid
Figure 7.6 Typical Mode 2 tools. Top: two views of a handax; below: two views
of a cleaver
make up the Story of Life. Until we have a time machine to take us back
into the past, no hypothesis of ancestor–descendant relationship can be
confirmed. Nevertheless, it is possible to continue reducing the number of
likely hypotheses, discarding others which are incompatible with three
types of data on the species represented by fossils: their age, their geo-
graphical distribution, and their degree of evolutionary relationship.
It is clear that more recent species cannot be proposed as ancestors of
older species, nor the latter as descendants of the former. Nor can we
assume an ancestor–descendant relationship between species which are
strictly contemporary with one another.
Moreover, in order for a species to be accepted as the ancestor of
another, it is not enough that it be older: it is also essential that it lived
in the same geographical region in which the assumed descendant species
originated.
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A New Kind of Hominid
110
A New Kind of Hominid
111
A New Kind of Hominid
This is not always an easy task, given the complexity of the evolutionary
process and above all the scarcity and state of preservation of the fossils
themselves. These difficulties mean that different researchers arrive at
different conclusions on the nature of the characters used in the analysis.
As a result, there is no consensus on the hominid evolutionary tree, and
various hypotheses have been put forward by different authors.
If we were to describe all the evolutionary trees put forward for hominids,
explaining how the appearance of new fossils has progressively eliminated
some and resulted in the presentation of other new theories, we would no
doubt try the patience of our readers. We therefore prefer to discuss the
evolutionary tree which in our view is the most compatible with the
evidence currently available (Figure 7.7).
In view of its antiquity and the primitive nature of the majority of its
traits, the species Ardipithecus ramidus appears the ideal candidate for the
0
Pleistocene
P. boisei P. robustus
−1
H.habilis/ H. ergaster
H. rudolfensis
A. africanus
Years (millions)
−2
First Homo
P. aethiopicus
−3 ?
Pliocene
A. afarensis
−4
A. anamensis
A. ramidus
−5
Figure 7.7 The authors’ evolutionary diagram for the first hominids
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A New Kind of Hominid
113
A New Kind of Hominid
Pleistocene
P. boisei P. robustus
−1
H.habilis/ H. ergaster
H. rudolfensis
A. africanus
Years (millions)
−2
First Homo
P. aethiopicus
−3 ?
Pliocene
A. afarensis
−4
A. anamensis
A. ramidus
−5
similar in the two species (compare Figure 4.4 and Figure 7.4). This seems
to us to indicate a very close relationship between Australopithecus afri-
canus and Homo. In our view, the common ancestor of the two lived
between 4 and 3 million years ago.
As far as evolutionary relationships between species in our own genus
are concerned, the question of whether Homo rudolfensis existed or not
makes very little difference. If we recognize only the existence of Homo
habilis the problem is somewhat simpler, since only two species remain to
be related, and on the basis of both chronology and cranial anatomy Homo
habilis could be the ancestor of Homo ergaster.
In any case, if we agree the existence of Homo rudolfensis, the species has
to be considered an evolutionary offshoot within the Homo genus, since
the anatomy of its masticatory apparatus is too specialized to propose it as
the ancestor of the other species of the genus. The position of Homo
habilis as the original species of Homo would thus remain unchallenged.
Our evolutionary tree for hominids implies that Paranthropus origin-
ated in East Africa. From there, representatives of this lineage reached the
ecosystems of the south of the continent, where they evolved separately
from their East African fellows and gave rise to a different species,
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A New Kind of Hominid
115
8
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The Evolution of the Brain
Cerebrum
Wernicke's area
Broca's area
Cerebellum
Brainstem
Figure 8.1 The human brain The areas of the cerebral cortex directly related
to language are shaded
When, in our lectures, we ask which is the animal species with the largest
brain, almost all students immediately reply: humans. The conviction that,
in line with our superior intelligence, our species is the most encephalized
in the animal kingdom, and that this is the basic characteristic that
distinguishes us from other animals, is embedded in our culture. How-
ever, this is not the case, or at least not in absolute terms.
The average size (mass) of the human brain is around 1,250 g,
and although this is greater than that of any other primate species
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The Evolution of the Brain
(chimpanzees and gorillas have average brain mass of around 400 and
500 g, respectively), and of the majority of animals, it is markedly less than
that of the large mammals, whose brains are much larger than ours: the
blue whale (the largest animal that has ever existed) has a brain mass of
around 6,800 g, while the African elephant (the largest living land mam-
mal) has a brain mass of about 5,700 g.
Does this mean that blue whales and elephants are more encephalized,
and therefore more intelligent than humans? Of course not. What hap-
pens is that since the work of the brain is to coordinate the functions of
the rest of the body, larger bodies predictably require larger brains to
ensure sufficient coordination. Thus when we compare the brain size of
two species we have to consider the influence of body size on the size
of the brain.
A simple way of relating body size to brain size is to divide brain mass by
body mass; in this way we can find the ratio between the two. The greater
the ratio, the more brain an animal would have per unit of body mass. On
first view, this might seem the most appropriate way of comparing the
brain size of animals of different sizes. However, this method also fails to
reveal humans as the most encephalized species in the animal kingdom.
Paradoxically, we are overtaken by the smallest mammals, which have
proportionately larger brains.
Should we then resign ourselves to accepting that the smallest mammals
are more encephalized than we are? We can rest easy: shrews do not have a
more developed brain than we do. The truth is that brain size increases
more slowly than body size: in other words, the size of the brain becomes
proportionately smaller in larger mammals. This phenomenon, the
change in the proportions of the organs as body size increases, is very
common in living beings, and was discovered in the 1920s; it was named
allometry, or the Law of Disharmony.
Allometry may occur during the growth of an organism: thus, for
example, a newborn has a proportionately bigger head than an adult,
and its limbs are relatively shorter. If a baby maintained these proportions
as it grew, the result would be an adult of very alarming appearance
(Figure 8.2).
Allometry may also occur between adults of the same species. A good
example of this is the variation in ‘‘ideal weight’’ in relation to height. As
height increases, the ideal weight becomes relatively greater: for a woman
150 cm tall the ‘‘ideal weight’’ is 47 kg (315 g per centimeter), while
another woman 180 cm tall has an ‘‘ideal weight’’ of 67 kg (372 g per
centimeter). But the kind of allometry which most interests us is the variation
between species in the same group. Thus in mammals, brain size varies in
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The Evolution of the Brain
relation to body size, and as we have already noted, brain size becomes
proportionately smaller as body weight increases. While in chimpanzees
the ratio between brain mass and body mass averages 0.01 (one gram of
brain per 100 g of body weight), in the mouse lemur (a small primate from
Madagascar with a body mass of only 60 g and brain mass of 1.8 g) the ratio
is around 0.03 (i.e. 3 g of brain for each 100 g of body weight).
Given that mammals’ brains grow allometrically in relation to body size,
the only way of comparing the brains of different species is to calculate the
brain mass that each should have in relation to its body mass, and to compare
these ‘‘ideal brain masses’’ with their actual brain mass. The species with the
greatest surplus brain mass would be the most encephalized.
Clearly, the key to comparing species of different size lies in calculating
the expected values of the brain mass of each. To this end the mathemat-
ical law which relates the body mass of an organism to its ‘‘ideal brain
mass’’ has been calculated, on the basis of analysis of extensive data on
brain mass and body mass in a large number of mammal species.
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The Evolution of the Brain
Weighing Ghosts
Although the brain does not become fossilized, the skull, which contains
the cavity which houses it, does. In skulls where the whole or a good part
of the neurocranium is preserved, it is possible to measure the volume of
the cavity which houses the brain. Since the brain occupies virtually the
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The Evolution of the Brain
whole of this space, the volume of the cranial cavity (cranial capacity) can
be used as a measure of the volume of the brain itself. Experimental
studies on primates and humans have shown that cranial capacity is almost
directly equivalent to brain mass: a capacity of 1,000 cc corresponds to a
mass of 971 g.
Estimating body mass from fossils is a different matter. Here the prob-
lem is more complicated because there is no measurement we can take
from the skeleton which would directly correspond to the individual’s
body mass. In tackling this problem, paleontologists start from an obvious
fact: the size of the bones is related to the size of the body itself, so large
bones should correspond to similarly large individuals.
However, not all the bones of the body follow this rule. For example,
we have seen that Paranthropus had a highly developed jawbone,
owing to its particular adaptation to diet. So if we applied the rule
that large bones (and teeth) reflect large body size to the jawbones of
Paranthropus, we would have to conclude that these hominids had a
body size close to that of a female gorilla. This mistake was in fact made
in the past, and is the source of the adjective ‘‘robust’’ usually given to
Paranthropus.
In order to arrive at reliable estimates of the body mass of fossil
hominids, we need to use bones or parts of bones which are clearly
involved in supporting body weight. Modern humans have several
bones which satisfy this requirement. In our species the weight of the
trunk, the upper limbs and the head is transmitted through the lumbar
region of the spine to the sacrum, in the pelvis. From here it passes
alternately at each stride to each of the femurs (which also have to support
the weight of the leg which is in the air), and through them down to the
bones of the feet (which support virtually the entire body weight at each
step). The lumbar and sacral vertebrae, the coxal bones and the bones of
the lower limbs are thus good candidates for estimating an individual’s
body mass.
Nor are all the dimensions of a bone equally suitable for estimating
body mass. For example, the length of the femur has more to do with an
individual’s height than his body mass. Since the body weight is transmit-
ted from one bone to another through the joint surfaces of the two bones,
these are directly related to body mass. The pressure exerted on a joint is
directly proportional to body mass and inversely proportional to the
joint’s surface area; therefore, in order to avoid the pressure on a joint
increasing with an increase in weight (and consequent injury to cartilage
and bone tissue), a correlative increase in surface area is required. Thus,
large joint surfaces indicate high body mass.
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Having identified the bones and the anatomical regions which are
suitable for calculating body mass, there is still another problem to re-
solve. What is the mathematical relationship between the joint surfaces of
each bone and the body mass?
This question can be approached by analyzing data on joint surfaces and
body mass in living primates, in order to determine the mathematical
function relating them. But we have already seen that not all primates
walk in the same way, nor do they have the same body proportions, and
these factors affect how the body weight is transmitted. For example, in
apes, which support themselves on four limbs when they move over the
ground, the humerus is more involved in the transmission of weight,
while the femur supports less weight than in modern humans. Given
that the first hominids were fully bipedal, comparison with our own
species seems most appropriate. Nevertheless, as we have noted in previ-
ous chapters, the body proportions of australopithecines and Paranthro-
pus appear to have been intermediate between those of apes and our own.
Thus the data on these primates also needs to be taken into account when
estimating the body mass of fossil hominids.
Paleontologists meet a further obstacle when estimating the weight of
bygone hominids: the scarcity of the fossil record itself. Few even roughly
complete skeletons of the first hominids are known, and only in one or
two cases has it proved possible to deduce an individual’s mass on the
basis of a range of bones (allowing us to refine the calculation). We also
have very few bones suitable for use for each species, and therefore a
very limited number of estimates for any one species. One final added
difficulty is the fact that it is not always easy to assign a bone of the
postcranial skeleton, found in an isolated position, to a specific species
of hominid.
Despite all these problems, researchers such as Henry McHenry and
William Jungers have undertaken a series of investigations which, using
the available evidence and a variety of statistical techniques (together with
a good dose of common sense), have yielded comparable results.
Australopithecines and Paranthropus appear to have had very similar
body mass. Let us recall that the average body mass of male and female
Australopithecus afarensis, the species for which body mass has been most
accurately estimated, was 45 kg and 30 kg, respectively. Homo habilis does
not appear to have been any bigger than the australopithecines and
Paranthropus, whereas the values estimated for Homo ergaster are over
55 kg, indicating a marked increase in body mass toward values similar to
those of our species.
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1
Although we can calculate an average brain mass for each of the different species,
the ranges of variation may overlap, with the higher values in one species overlapping
with the lower values for another. This is the case, for example, with Paranthropus boisei
and Homo habilis: the former has an average brain mass of 508 g, but a range of
variation from 470 g (fossil ER 13750) to 590 g (Konso specimen). Homo habilis,
on the other hand, has an average of 619 g but includes specimens with a brain mass of
only 503 g (such as ER 1813) and others up to 661 g (OH 7), or 736 g, if fossil ER
1470 is included in this species (as it is for the purpose of our calculations).
2
Our calculation of indices of encephalization was based on a different formula from
that used in the section where we compared the encephalization of all mammals, which
gave a value of 7 for our species. For this calculation we used the formula which
estimates ideal brain mass on the basis of data from haplorrhine primates. This new
calculation gives a value of 2.9 for our species. This means that we have a brain mass
seven times the expected value for a mammal of our body mass, but only 2.9 times
greater than that which would correspond to a haplorrhine primate of our size.
Remember that primates are highly encephalized mammals.
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124
The Evolution of the Brain
Frontal lobe
Parietal lobe
Occipital lobe
Temporal lobe
activity, we can only track the evolution of a few of these in fossils because
our knowledge of the brain surface of fossil hominids is limited by various
circumstances.
First, we have no fossil brain, only casts (natural or artificial) of the
cranial cavity, and although the internal walls of the cranium reproduce
the general morphology of the brain surface they do not do so with
sufficient detail to identify the exact limits of most areas of the brain
with any precision. This is because the brain is covered by a triple layer
of membrane, the meninges.
Moreover, the larger the brain of an organism, the more faint and
indistinct the impressions of the brain on the inner surface of the cranium.
Some regions of the brain surface also leave more detailed impressions
than others. Thus, for example, the impressions of the occipital lobe
are lighter than those of the frontal lobe. In short, studies of the
paleoneurology of the first hominids are limited to a few specific aspects
of brain morphology.
The two world authorities in this area, Ralph Holloway and Dean Falk,
hold different opinions on the brain of australopithecines and Paranthro-
pus. Holloway believes that these hominids already show evidence of
reorganization of the brain compared with apes; these are reflected, for
example, in a degree of development of Broca’s area (an area related to the
production of speech, which we shall look at in the chapter dedicated to
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The Evolution of the Brain
language). However, Falk takes the view that the brain surface of Paran-
thropus and australopithecines is clearly similar to that of apes and that
there is no evidence of reorganization of the brain.
Despite their differences on this matter, Holloway and Falk agree that
there are clear indications of cerebral restructuring, similar to that of
modern humans, in the fossils of the first representatives of Homo.
These indications can be summed up in two major processes: the appear-
ance of marked asymmetry between the cerebral hemispheres, and a more
complex morphology of the frontal lobe.
The human brain is clearly asymmetrical, and although apes, and an-
thropoids in general, show cerebral asymmetries, these are not of the same
kind or as extensive as those typical in humans. The asymmetries charac-
teristic of the human brain are not directly related to the functional
specialization of the cerebral hemispheres observed in humans. This phe-
nomenon is known as brain lateralization, and seems to be confined to
our species. To generalize, the left hemisphere specializes in functions
related to language, precision movements of the right hand, the capacity
for analysis, and the perception of temporal sequence. The right hemi-
sphere is devoted more to tasks such as the skill of the left hand, the
capacity for overall understanding of processes, recognition of faces,
spatial vision, musical skills, the control of the tone of the voice, and the
expression of emotions and recognition of them in others.
In addition to other functions (such as control of primary motor
functions), the frontal lobe is responsible for a series of mental capacities
which are exclusive to or most fully developed in humans. These include
the capacity to determine the sequence of movements of the phonetic
apparatus which make up speech, control of the emotions, the ability to
concentrate on a task, planning, anticipation of events, maintaining an
idea in the mind for a long time, and control of the use of the memory to
integrate previous experiences and learning when taking decisions. Meta-
phorically speaking, we could say that the frontal lobe is the ‘‘conductor’’
of the orchestra of our brain.
Over the course of human evolution the frontal lobe has expanded a
great deal, both in absolute terms and in relation to the rest of the brain.
But recent investigations by a group of scientists led by Katerina Semen-
deferi indicate that the volume of our frontal lobe is what would be
expected in a primate with a brain the size of ours. Given that the frontal
lobe appears to be the most ‘‘human’’ region of our brain, perhaps the
increase in brain size is merely the result of the selective advantage that
having an increasingly more developed frontal lobe conferred on our
ancestors. In other words, it is reasonable to suppose that the large size
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The Evolution of the Brain
of the brain in our species is not the cause, but the result of our large
frontal lobes. If this is the case, development of the frontal lobe would
have been the driving force in our encephalization.
What we can at least say with certainty is that the surface of the frontal
lobe has become increasingly complicated, owing to an increase in the
number of sulci which cannot be explained purely in terms of the greater
size of the human brain.
These characteristics – the cerebral asymmetry linked to functional later-
alization, and greater structural complexity of the frontal lobe – are clearly
marked in the endocranial casts of Homo habilis/Homo rudolfensis and
Homo ergaster. So even in the first humans, in addition to an increase in
brain size, we find clear indications of a brain structure similar to our own.
To complete our examination of the brain of the first hominids, we
might speculate as to the nature of the vital advantage the new type of
brain gave our ancestors, favoring their selection. There are two hypoth-
eses on this subject.
The first, put forward by Robin Dunbar and Leslie Aiello, links the
increase in the neocortex (i.e., the cerebrum) to the improvement in social
skills within the group. According to these authors, the increase in size
and the reorganization of the human brain are linked to the development
of ‘‘social intelligence.’’ We shall return to this hypothesis when we look at
the social biology of the first hominids.
Falk, on the other hand, relates the modifications in the brain of the first
humans to a very specific capacity, that of language. She argues that given
that one of the crucial centers of human speech is located in the frontal lobe,
and that the production and decoding of language are functions which are
clearly lateralized in the human brain, both lateralization and the increased
structural complexity of the frontal lobe are related to the development of
the linguistic capacities of our first ancestors. It is immediately evident that
these two hypotheses are complementary, since the primary social skill is
perhaps the capacity to communicate efficiently with others.
However, the linguistic capacities of the first humans are the subject of
major controversy. We shall return to this subject in the chapter on
language.
During the last quarter of the 19th century it was fashionable in some
scientific circles to link human intelligence with brain size. During that
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The Evolution of the Brain
128
9
Types of Diet
129
Teeth, Guts, Hands, Brain
130
Teeth, Guts, Hands, Brain
131
Teeth, Guts, Hands, Brain
Carnivorous mammals have adapted their teeth to the functions they need
them to perform. Carnivores vary widely in form, but all of them have
certain teeth specialized in cutting meat, known as carnassial teeth (these
are the last upper premolar and the first lower molar). Carnassial teeth
have a very sharp edge which allows them literally to slice up the flesh of
dead animals (Figure 9.1).
Proteins and animal fats are rich in energy, having a high calorific value,
and they are also easy to digest. This means that this type of food does not
need to be prepared in the mouth before it is swallowed, so chewing is
brief, its purpose being only to reduce the flesh of the dead animal to
pieces of a size that can pass through the esophagus. The carnassial teeth
operate like shears, or scissors, the upper one passing to the outside of the
lower when the mouth is closed. This scissor-like movement causes the
carnassial teeth to sharpen themselves on one another, and as they wear
against one another they become razor-sharp. Unlike our kitchen knives,
this wear improves the cutting edge of the carnassial teeth and increases
their efficiency, rather than dulling or chipping them.
Carnivores also need tools to kill their prey, and the canines, often
known as eye-teeth, are highly developed in order to perform this func-
tion. The terrestrial carnivores with the most highly developed canines are
the Felidae, the family which includes cats, lions, tigers, and so on. In
these animals the incisors are small and the canines very large; the carnas-
sial teeth are highly developed and are the hindmost teeth in the jaw
(although the vestige of a molar remains behind them).
Hyenas have a similar dentition to that of the big cats: in fact their diet
is similar to that of the Felidae, although they do not always kill their prey.
(It has been established, however, that the larger spotted hyena is a
powerful group hunter.) In addition to the range of teeth seen in Felidae,
hyenas also have highly developed conical teeth (the penultimate pre-
molar, both upper and lower), which they use as hammers to crush the
bones of large herbivores. When the predators and the majority of carrion
eaters have finished eating, the tougher bones still remain: these enclose a
very fatty, and therefore energy-rich, material, the marrow. Hyenas can
gain access to this resource using their specialized premolars. At a certain
point, some other animals also began to compete for the marrow con-
tained in the strong-walled bones of the large herbivores; these were
primates – humans, who, lacking the appropriate teeth, used stone tools
to split the bones.
132
Molar
(carnassial tooth)
Premolars
Canine Incisors
Molars
Premolars
Canine
Incisors
But not all land-based carnivores have such a specialized diet, and
therefore dentition, as the cats. The Mustelidae (the family which includes
badgers, otters, martens, mink, and weasels), the Canidae (the group of
the foxes, wolves, coyotes, African wild dogs, etc.), the Ursidae (bears),
and the Viverridae (such as genet and mongoose) have molars behind the
carnassial teeth with chewing surfaces instead of cutting edges. While the
front part of the lower carnassial teeth has a cutting edge, the back does
not. This indicates that their diet also includes vegetable matter which is
more difficult to absorb, and requires prior chewing, insalivation, and
predigestion in the mouth. The famous cave bear, a gigantic animal bigger
than any modern bear, which lived at the same time as Neanderthal and
Cro-Magnon man, was basically vegetarian, as its large molars indicate.
It goes without saying that the large herbivores also modified their
teeth to the function they had to perform. The great grasslands – the
steppes, prairies, and pampas – provide enormous quantities of plant food
which is, however, of low nutritional value because the plants which make
it up are rich in fibers like cellulose. In order to absorb the food, a high
level of predigestion in the mouth is required, where the fibers are
reduced to a pulp which is then processed in the digestive tract. This
processing requires the assistance of symbiotic microorganisms (basically
bacteria) which live in the intestines of herbivores and can perform the
miracle of decomposing cellulose into carbohydrates which can be
absorbed by the animal.
The teeth of these mammals have a ridged crown which increases the
efficiency of their chewing surfaces. The work of maceration and reducing
the vegetable matter to a pulp is speeded up if the surfaces rubbing
against one another are ridged, like millstones in traditional grain mills
(Figure 9.1).
The premolars and molars of herbivores have to withstand intensive
wear, since the fibrous stalks of plants like grasses also contain mineral
particles which make them harder and more abrasive. They therefore have
very high crowns, so that they last longer, and very wide chewing surfaces;
they have no cutting edge for slicing flesh.
What kind of teeth do our closest relatives, the great apes, have? And are
ours similar to those of herbivores, or carnivores, or like those of the great
apes? The latter have very large incisors and canines, particularly the males.
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The front teeth (incisors and canines) work to slice up fruits of a certain
size so that they can then be chewed by the back teeth (premolars and
molars).
We modern humans have modified the dentition of our ancestors
considerably. To begin with, our incisors are not as large as those of the
apes. We have no diastema, and the tips of the canines project hardly at all
beyond the chewing surfaces of the other teeth; moreover, with wear the
point of the canines very quickly becomes level with the surface of
the adjacent teeth. In humans and apes the incisors are spatulate, or fairly
flat, widening from the base of the crown (or neck) to the cutting edge. In
apes the canines are conical, but in humans their shape is closer to that of
the incisors, spatulate, and becoming wider until just below the tip of the
tooth (which is in any case fairly rounded).
In humans the first lower premolar has two cusps like the others, and it
does not slide against the posterior edge of the upper canine. In apes the
incisors are far in front of the canines, which are aligned with the pre-
molars and molars, so that the dental arches have parallel sides and are U-
shaped. In humans the dental arches are parabolic or elliptical.
Of all the fossil hominids, the one which is most similar to living apes is
Ardipithecus ramidus. In this species the canines project considerably,
although not as much as in apes, and the first lower premolar is sectorial.
Other typical features of apes which have become lost over the course of
human evolution are found in this hominid, including the morphology of
the dental arch, which is U-shaped with parallel sides. No palates have
been preserved, but judging from the size of the canines we can imagine
that there was a well-developed upper diastema.
In Ardipithecus ramidus the first milk molar has a very prominent
principal cusp, as in apes; this characteristic had never before been seen
in a hominid (all the others have several cusps of similar size). This trait is
so significant that a small fragment of jawbone showing a conical first milk
molar appeared on the cover of the historic issue of Nature in which, on
September 22, 1994, the discovery of Ardipithecus ramidus was pub-
lished.
We have already noted that the enamel is also very thin in this species.
We have to conclude that the diet of Ardipithecus ramidus differed little
from that of modern chimpanzees. However, there is one trait which
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merits further attention. Although the canines project much further than
in modern humans, they do not project as much as in apes, nor are they
the same shape. As we have mentioned, in humans the canines are spatu-
late or incisiform (having the shape of the incisors), and this condition
appears already incipient in Ardipithecus ramidus. Moreover, the anterior
edge of the first lower premolar is not sharpened through wear against the
upper canine (as it is in apes). In fact, this change in the function of
the upper canine/first lower premolar pair, small though it is, is one
of the main arguments for classifying Ardipithecus ramidus as a hominid.
This combination of traits prompted Tim White and his colleagues to
attribute a crucial role in human evolution to this species, as ancestor of all
the subsequent hominids, and thus our first known ancestor. Given its
antiquity (4.4 million years), it is unlikely that much more primitive forms
of hominid will be found.
Although when the first fossils of Australopithecus afarensis were discov-
ered they seemed the height of primitivism (as had Australopithecus afri-
canus, long before), the archaic traits of Ardipithecus ramidus give an idea
of how many things had changed in Austalopithecus afarensis. Some of the
fossils of the latter still show first lower premolars with a very prominent
cusp (while others already have two cusps), the diastemas in the upper
dental arches are small or nonexistent, and the canines project less and are
more incisiform than in Ardipithecus ramidus. The most important feature
is that in Australopithecus afarensis the molars have become larger, and
particularly wider, than in Ardipithecus ramidus. This, combined with the
increase in thickness of the tooth enamel, indicates that a major ecological
change had taken place, resulting in a diet which is still plant-based, but
now contains a large proportion of tough, abrasive foods. There is no trait
in the dental morphology which indicates significant consumption of flesh.
Our first ancestors, both those who lived in the humid forest like Ardi-
pithecus ramidus and those who had begun to exploit the resources of dry
woods and clearings like Australopithecus afarensis, were not hunters but
vegetarians. To put it more dramatically, the first hominids did not come
down from the trees to become ‘‘murdering apes,’’ nor was it their taste for
meat that prompted them to abandon the forest.
The fossil record for Australopithecus anamensis is still scarce, but we
have some information on the masticatory apparatus of these fossils which
are intermediate between Ardipithecus ramidus and Australopithecus afar-
ensis. The Kanapoi maxilla and mandible, together with the teeth that
have been preserved, show very primitive characteristics, but it appears
that the enamel had already become thicker and the size of the molars
increased to a certain extent.
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Teeth, Guts, Hands, Brain
formed by the last premolar and the first two molars). The value of the
index for Australopithecus afarensis is almost double that for chimpanzees,
indicating that diet had already changed to include tougher plants. The
relative size of the chewing surface increases somewhat in Australopithecus
africanus and continues to increase in Paranthropus robustus and Paran-
thropus boisei (to three times that of chimpanzees). However, in Homo
habilis the value of the index returns to the level of Australopithecus
afarensis, and in Homo ergaster it approaches that of the chimpanzee,
very similar to our own.
Paranthropus are the hominids with by far the largest chewing surface,
both in absolute terms and in relation to body mass. And not only the
molars, but also the premolars are larger, particularly the posterior pre-
molar, which has become ‘‘molariform,’’ increasing the number of cusps
and acquiring the form of a molar. The incisors and canines, on the other
hand, have become very small, creating an imbalance between the very
reduced anterior teeth (incisors and canines) and the enlarged posterior
teeth (premolars and molars).
Large canines which project beyond the level of the adjacent teeth and
have to be slotted into diastemas make it difficult to move the front part of
the jaw from side to side. This type of lateral movement combines with the
vertical movements of opening and closing the mouth to produce rota-
tion. Rotational movements are necessary to grind tough plant foods. The
great apes do not have this problem because their diet does not require so
much grinding. In baboons the face is so large that some rotation of the
posterior part of the jaw is possible, although in males the anterior part is
blocked by the hugely developed canines. In Australopithecus, Paranthro-
pus, and Homo, the wear on the premolars and molars is very intensive; as
soon as these teeth appear their chewing surfaces characteristically become
flat and with virtually no relief, the enamel disappearing rapidly and
wearing closer and closer to the dentin, the tissue beneath the enamel.
What kind of diet did Paranthropus have? No doubt similar to that of
modern baboons, based on hard grass seeds, fresh or dry legumes, and nuts
(Figure 9.2). It would also have included fruits like blackberries, berries, and
stone fruit, which were easier to chew, and the underground storage organs
of plants like bulbs, tubers, fleshy rhizomes, and tuberous and swelling roots
(baboons eat practically everything, including small animals).
We have already noted that grass seeds and legumes are rich in carbo-
hydrates (which they store principally in the form of starch), and are thus
very nutritious, although low in protein (legume proteins, moreover, have
to be cooked in order to be absorbed). As processing seeds in the mouth is
very laborious, Paranthropus developed their own personal mill, through
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Figure 9.2 Paranthropus boisei and Theropithecus brumpti. These two primates developed somewhat similar specializations in their
chewing apparatus, owing to a similar diet, containing a high proportion of tough plant matter
Teeth, Guts, Hands, Brain
the expansion of the premolars and molars and of the muscles which
moved them and the bones in which the muscles were anchored. This
powerful chewing apparatus would also have been useful in cracking nuts,
rich in vegetable oils. The underground storage organs of some plants
provide carbohydrates and some proteins, although the soil which they
would inevitably have ingested with them would have contributed to wear
on the teeth. Baboons, and particularly their relatives the geladas which
live on the high, virtually treeless plains of Ethiopia, can digest grasses and
grass stems, but this is a unique specialization which cannot be assumed
without evidence to have existed in hominids.
Some authors believe that Paranthropus, and also the australopithec-
ines, used sticks to dig bulbs or tubers, for example, out of the earth. It is
also possible that both genera included animal products in their diet, as do
baboons and chimpanzees when they can. However, the theory main-
tained by Dart and Le Gros Clark, that the first hominids were predators
on a large scale, using tools to kill their prey, is now completely discre-
dited. In a magnificent example of taphonomic analysis, Bob Brain dem-
onstrated that the hominids whose bodies accumulated in the caves of
South Africa were not the fierce occupants of the caves, but rather prey
brought there by the real hunters, leopards. The baboons, gazelles, and
other herbivores whose remains are found together with the hominids
would have met the same fate. Lee Berger and Ron Clarke have recently
postulated that even the Taung Child was the victim of a predator,
although this time a winged one – an eagle.
One interesting aspect of baboons’ diet is that it requires a certain
capacity for manipulation of small objects, such as seeds, with the hand.
There is no published data on what Ardipithecus ramidus’ hands were like,
but it is possible that in the first hominids (this species or an earlier one)
the morphology of the hand was similar to that of apes, with a short
thumb and long palm and fingers. Remember that this is an adaptation for
hanging from trees by forming a hook with the hand.
In Australopithecus afarensis the hand is already almost the same as
ours, although the phalanges are more curved, somewhat like those of
chimpanzees; moreover, the thumb is proportionately shorter and the rest
of the hand slightly longer than in our own species. Hand bones of a
modern type have been found in the Swartkrans deposit. Most of the
Swartkrans fossils are from Paranthropus robustus, but as there are also
some from Homo, we cannot be sure to whom these hand bones belong.
However, we can venture the hypothesis that, at least since Australopith-
ecus afarensis, hominids had modified their hands so as to have full
capacity for manipulation of small objects, as a result of their new diet.
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Our vision of the first hominids has changed greatly with time. Origin-
ally they were envisaged as fierce predators of the savanna, and then as
inoffensive vegetarians, also living in the savanna. Ardipithecus ramidus is
now believed to have lived in the humid forests, and all species of Aus-
tralopithecus are thought to have been vegetarians living in a mixed
environment, whether this is called dry forest and clearing or densely
wooded savanna. Two types of hominids which were genuinely specialized
in open environments, Paranthropus and the first Homo, seem only to
have appeared 2.5 million years ago.
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meat is easy to digest. Herbivores, on the other hand, need long digestive
tracts in order to metabolize the plants they eat, particularly if these are
rich in cellulose.
We have already noted that about 2.5 million years ago two types of
hominid settled in open environments. This is a point in time which many
authors consider important from the point of view of climate change,
because a general cooling of the planet occurs – reflected in Africa by the
definitive expansion, at the cost of the enclosed forest environments, of
the great grasslands and the savannas (albeit still more or less wooded).
One of these two types of hominid is Paranthropus, which adapts its
chewing apparatus to eat the tough but nutritious plants of the savanna,
as baboons do today.
However, the brain of Paranthropus does not increase in size as much as
that of Homo. Bearing in mind that this increase in brain size implies an
increased energy consumption, only two solutions remain. One is increas-
ing the basal metabolic rate of the organism as a whole (the total energy
consumption). This is not what happened, since humans have the rate
which corresponds to a mammal of our size. The other solution is to
reduce the energy consumption of another organ in order to balance the
body’s energy economy. The question is, which organ will have to reduce
in size? Not the heart, nor the kidneys, nor the liver, which are vital parts.
However, the digestive tract can become smaller if diet is improved, by
increasing the proportion of high-quality nutrients, those which can be
easily digested and have high calorific value. What are these high-quality
products which do not form part of the diet of Paranthropus? The only
answer can be animal fats and proteins. The first humans must have shifted
to incorporating a larger proportion of meat than any other primate in
their diet; they would have got this first as carrion-eaters, and then
increasingly as hunters.
For the first time in the history of mammals, this change in diet was not
reflected in a change in dental morphology. We do not find humans with
teeth that work like hammers to crush bones, nor with teeth that act like
knives to cut meat, because the tools required for splitting bones and
cutting skin and meat are outside the body: they consist of stones and the
edges of stones worked by humans.
Thus, the enlargement of the brain in Homo could only have occurred
with a change in diet; this in its turn is reflected in the reduction in size of
the digestive tract and a correlative reduction in the chewing apparatus.
Aiello and Wheeler insist that this does not mean that change in diet
automatically leads to an increase in brain size; they maintain only that we
had to become carnivores in order to become intelligent (although this is
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144
10
Development
Yes. One goes on. And the time, too, goes on – till one perceives ahead a
shadow-line warning one that the region of early youth, too, must be
left behind.
Joseph Conrad, The Shadow Line
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Development
over about twenty years, almost double that of the great apes (chimpan-
zees, gorillas, and orangutans). On average chimpanzees get their first
adult molar when they are a little over three years old; in our species it
erupts at the age of six. In chimpanzees the second molar emerges at the
age of six and a half, while in humans it appears at around eleven years;
finally, the third molar appears in chimpanzees at around age eleven, and
in ourselves at around eighteen (although among modern humans the
four wisdom teeth sometimes do not appear at all, and when they do it can
be well into adulthood). In chimpanzees the first period of estrus, with its
characteristic swelling of the ano-genital region, occurs between the ages
of nine and almost fourteen, depending on the nutritional condition of
the female (in humans too the first menstrual period, or menarche, occurs
earlier, on average, in girls who are well fed and healthy). In chimpanzees
as in humans, this is usually followed by a period of infertility before the
first conception. In chimpanzees the first birth occurs on average at the
age of fourteen. A chimpanzee can live to the age of forty or more.
Despite this difference in length of the periods of development between
apes and ourselves, the relatively close correspondence between the stages of
life and the appearance of the molars helps us to establish the state of
development of a fossil hominid: although we do not know its ‘‘chrono-
logical age’’ (i.e., how old it was when it died), we do know its ‘‘physiological
age’’ (whether it was an infant, an older child, an adolescent, or an adult).
Thus the famous Taung Child was emerging from early childhood
when the eagle snatched him, because the first molar was erupting pre-
cisely at that moment. So how old would he have been? If he died between
the ages of three and four this would mean that the rhythm of develop-
ment of Australopithecus africanus was similar to that of chimpanzees,
while if he died at six, it would be slower, like our own. When Turkana
Boy died the second molars had already erupted. Was he around eleven
years old, like a modern child in terms of the development of his skeleton
and teeth, or was he around seven years old, like a chimpanzee with the
same ‘‘physiological age’’? How can we find out? In order to tackle this
question, we shall start at the beginning of existence – birth.
Shortly before birth the fetus shifts into an inverted position, with the
head in the upper part of the mother’s pelvis (formed by the iliac crests
and known as the greater pelvis or false pelvis). During birth, the full-term
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Development
fetus has to pass through the lower part of the pelvis (known as the lesser
or true pelvis) via a bony tube called the pelvic cavity or birth canal.
Among chimpanzees, gorillas, and orangutans birth is easy and rapid,
because the birth canal is large in relation to the head of the fetus. In
these apes the entry to the birth canal (known as the pelvic inlet) is oval,
with its major diameter running from front to back (a sagittal or ante-
roposterior orientation, in anatomical terms), and its minor diameter
oriented transversely (from side to side). As we have seen, the modifica-
tion in pelvic architecture required to make it possible to walk upright
meant that the joints between the coxal bone and the spine, and the coxal
bone and the femur, drew closer together. As a result, the sagittal diameter
of the birth canal reduced, and this is why complications can occur in the
human birth process.
In human females the entry to the birth canal is not oval but round
(Figure 10.1). The major diameter of the pelvic inlet is not sagittal: it is
sometimes transverse but more often neither sagittal nor transverse, but
on the two oblique or diagonal diameters. Moreover, the head of the full-
term fetus is elongated, so that its major diameter is anteroposterior (from
the forehead to the nape of the neck). The position of the fetus’ head
adapts to the greater diameter of the pelvic inlet, generally one of the
oblique diameters. As the greatest dimension of the outlet to the birth
canal is always sagittal (in humans as well as in other primates), the skull
and shoulders of the human fetus have to enter the birth canal with one
orientation (transverse or oblique) and leave with another (sagittal). Thus
there is a rotation within the birth canal both of the head and, afterwards,
of the shoulders (Figure 10.2).
To complicate matters further, in humans the vagina is angled forward,
forming a right angle with the uterus, so that as it passes through the birth
canal the fetus moves not in a straight line, but around a pronounced
curve which ends immediately below the pubic bone, where the baby’s
head emerges. In order to accommodate this curved trajectory, the fetus’
spinal column arches, the head flexes strongly toward the back and the
crown of the head faces forward as it is born, so the baby is facing
backward (in the opposite direction to the mother). In other primates
the vagina lies at the same angle as the uterus, with which it is aligned, and
the full-term fetus follows a straight trajectory toward the back as it is
born; furthermore the face is forward (in the same direction as the
mother). To sum up, birth in humans is ventral, while in other primates
it is dorsal.
Karen Rosenberg and Wenda Trevathan have pointed out that the
mother ape can help her baby to be born, guiding it through birth with
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Development
Ilium
Sacrum
Acetabulum
Ischium
Pubis
Iliac crest
Sacrum
Pelvic
Acetabulum inlet
Pubis
Ischium
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Development
Figure 10.2 Flexion and torsion of the fetus during birth. To begin with
(top), the head lies in a transverse or diagonal position and the shoulders in a
sagittal position (along the mid-plane of the body). Later (center) the head lies
sagittally and the shoulders transverse or diagonally. As it comes out the head
flexes strongly backwards. Finally (below) the head, once it has emerged, shifts
transversely, and the shoulders turn to a sagittal position to come out
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Development
her hands, clearing its nose and mouth so that it can breathe better and
freeing it from the umbilical cord if this is wound around its neck. Birth in
primates is a solitary event, with no outside help. However, in humans the
mother cannot see the newborn’s face because it faces in the opposite
direction, and any attempt to pull it could, given the extreme dorsal
flexion of the head, damage the spinal cord. Because of this women in
all cultures seek assistance at the moment of birth; human birth is a social
activity rather than a solitary behavior.
We have two australopithecine pelvises in a reasonably good state of
preservation: that of Lucy, and that of Sts 14 (an Australopithecus africa-
nus from Sterkfontein). Both belong to small individuals, and are there-
fore probably female. In Lucy’s pelvis the entrance to the birth canal is
very wide in the transverse direction. This could be a trait of the species or
simply an individual character, since in Sts 14 the entrance to the birth
canal is more rounded (in some women the birth canal is very wide from
side to side, a condition known as platypellic). In any case, in Lucy the
fetus’ head would have lain transversely, accommodating itself to the form
of the pelvic inlet. Robert Tague and Owen Lovejoy believe that the
position of the head would be the same on leaving the birth canal (there
would thus be no rotation), and that the fetus’ trajectory would be linear
and toward the back, rather than curved and toward the front (Figure
10.1). So this would be a transverse type of birth, a form without known
equivalent among primate species.
However, the morphology of the ischium and the pubis leads us, like
Christine Berge, to conclude that the vagina was angled forward, not
backward, in female australopithecines; this would mean that the birth
process in australopithecines would be similar to that of modern humans,
with rotation and a curved trajectory.
Before moving on in our discussion of birth in early hominids, let us
spend a few moments considering the implications if this theory were
proved true. In primates the vagina opens dorsally (toward the back) and
penetration during copulation is from behind. Gibbons and orangutans,
which spend much time in the trees, often mate hanging from their arms
and face to face, but chimpanzees and gorillas copulate from behind. The
bonobo has very varied positions, including face to face. In humans this is
the position favored by the ventral (forward) position of the vulva, and it is
characteristically human. We are also distinct in this, and if our conclusion
about the direction of the vagina in australopithecines is correct, face-to-
face copulation would also be the rule among these primitive hominids.
There has been much discussion of whether birth for Lucy would be as
difficult as it is for modern women, or as simple as it is for the great apes.
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Development
In fact the latter are the exception to the rule among primates: in gibbons
and other catarrhine and platyrrhine monkeys the diameters of the head of
the fetus are fairly close to those of the birth canal. This gives rise to a
certain number of pelvic–cephalic conflicts which result in the death of the
fetus, the mother, or both. This is an unfavorable consequence of the fact
that anthropoids have large brains in proportion to their generally small
body size.
In women the pubis is longer (in relation to the overall size of the
pelvis) than in men. As the pubic bone forms the anterior wall of the birth
canal, this lengthening of the pubic bone in women produces a wider
birth canal. Lucy and Sts 14 have an extraordinarily long pubis, and thus a
much larger birth canal, in relation to body size, than modern women.
The extremely elongated pubis in Lucy and Sts 14 gives rise to a pelvic
morphology known as hyperfeminine according to modern standards.
These and other characteristics lead us to conclude that the pelvis of
australopithecines was already designed to resolve the problem of birth
that bipedalism had created. Although this problem was perhaps not as
acute as in our species, the advantages enjoyed by female chimpanzees,
gorillas, and orangutans had been lost.
We can consider this question of the rhythms of development right
from the moment of birth, since among apes the newborn comes into the
world with a brain more than one-third of the volume of the adult brain,
while in our species it is less than one-quarter. This means that at birth the
newborn human is much less developed cerebrally, and therefore more
helpless, than any other primate. Primates in general are a group of
mammals which give birth to few offspring at a time (usually only one),
but these are well developed when they are born. This developmental
precocity is also found among apes, so the less developed state of our
newborns must be a condition our evolutionary line has acquired since the
time of any ancestors whose babies were more developed.
We have no direct information as to the size of the newborn australo-
pithecine’s brain, since no neonate skull has been found, but we can
approach the problem in another way. The pelvises of Lucy and Sts 14
tell us the dimensions of the birth canal through which the head and
shoulders of the fetus would have to pass during birth. The dimensions of
the birth canal establish the maximum size of the skull and the brain of the
newborn; in the case of Sts 14 and Lucy this would be comparable to that
of a newborn chimpanzee, or perhaps a little bigger. Furthermore, the
brain size of the average adult Australopithecus afarensis (Lucy’s species)
and Australopithecus africanus (the species of Sts 14) would also be
approximately the same size as that of a chimpanzee (about 400–410 g),
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Development
or perhaps bigger. Thus, the ratio between the brain size of the newborn
and that of the adult (which indicates the degree of cerebral development
of the newborn) would be comparable in the great apes and in the first
hominids. So the newborn babies of the first hominids would have come
into the world much more developed than modern human babies, al-
though the birth process was already like that of modern humans because
the australopithecines were bipedal like us.
We have already seen that in Homo ergaster body size had increased
substantially in relation to australopithecines, Paranthropus, and Homo
habilis. But at the same time the brain volume of the adult doubled.
The brain of the newborn may also have doubled in size, leaving the
proportion between the two brain sizes the same. But it is also possible
that the dimensions of the birth canal did not increase by the same
amount; in this case the brain of the fetus could not be as large (since
it had to pass through the birth canal), and it would represent a smaller
percentage of the adult brain than in australopithecines. This would
imply that the newborn came into the world less developed, needing
more care.
Although this supposition accords with the generally held view (which
we shall discuss later) that Homo ergaster already exhibited major changes
in the social sphere compared to australopithecines, and we imagine a
more protective social environment, we have no definitive proof that
young Homo ergaster were born less mature. Unfortunately, the only
pelvis of the species we have is that of the Turkana Boy, which in addition
to being that of a child is very incomplete and allows us to draw no firm
conclusions on this subject. Nevertheless, it is worth mentioning that
Christopher Ruff and Alan Walker have estimated, on the basis of the
Turkana Boy, that the newborn of the species would have a brain one-
quarter the size of the adult brain. Bearing in mind that any such conclu-
sion is as yet unproven, this would nevertheless imply that the newborn
Homo ergaster was already less mature.
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Development
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Development
stages. The greater the average brain of a species, the longer the life of the
individuals of that species and, for example, the later the second adult
molar appears. The brain volume of Homo ergaster is thought to have
averaged between 800 and 900 cc, midway between that of chimpanzees
(and australopithecines and Paranthropus) and our own. On this basis we
can, in principle, assume that the rhythm of the development of the
species would also be roughly midway between the two, and that Turkana
Boy died at the age of nine or ten. If this is the case, it would mean that a
significant change in the biology of development had occurred in this
species compared with australopithecines and Paranthropus.
There is one further aspect we need to consider in order to complete
this look at development. Turkana Boy is estimated to have been about
160 cm tall. However tall he might be when fully grown, this height
would be too tall for a modern child at his stage of development (as we
have noted, equivalent to about 11 years old). In other words, the height
of Turkana Boy would be more suited to a modern adolescent of 15 or
older (even among very tall populations such as the Maasai).
However, in our species an acceleration in growth occurs at an age older
than that of Turkana Boy – the so-called ‘‘growth spurt’’ of puberty
(around the age of twelve in girls and fourteen in boys). The conclusion
drawn by Holly Smith, who has studied the development of Turkana Boy,
is that in Homo ergaster this change in the rate of growth did not yet exist,
and that it only occurs in our species. In apes and other primates devel-
opment is more continuous, without any marked accelerations in adoles-
cence, and the percentage of growth remaining is lower in apes of an
equivalent ‘‘physiological age.’’ No doubt this was also true of Homo
ergaster (though even so, Turkana Boy would have grown to be over
180 cm tall).
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11
Social Intelligence
Five years after her last period of sexual activity, the female orangutan
comes into season once more. During this long break she has been
gestating, giving birth to and nursing her latest offspring. Now the
moment has come for weaning and beginning a new cycle, a new gesta-
tion. An ovum awaits fertilization, probably by the same male as the
previous time. After this brief receptive period, the sex life of the female
orangutan ceases for another five years or so (unless the baby she has
conceived miscarries or dies).
Female chimpanzees and gorillas also have seasons (known in zoology
as periods of estrus) separated by breaks of several years, generally more
than three years and less than six, but while female gorillas have sexual
relations with only one male, female chimpanzees will have several lovers.
Human women, on the other hand, do not have a period of estrus, and
thus the moment of ovulation cannot be detected; unlike female apes,
their sexuality is not governed by being in season and it includes the long
infertile periods of gestation, lactation, and menopause. In other words,
whereas female sexuality in apes is linked exclusively to reproduction, in
our species it also exists beyond this function.
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Behavior as Adaptation
With their research, Konrad Lorenz and Niko Tinbergen (who shared the
Nobel Prize for Medicine with Karl von Frisch in 1973) laid the founda-
tions for understanding animal behavior, creating a new scientific discip-
line – ethology, or the study of behavior. Ethology establishes that there is a
degree of genetic programming which codetermines behavior (in other
words, it determines it only in part). Just like morphological structure and
physiological function, behavior has to adapt to the way of life of the
individuals of different species, and therefore the genes which determine it
are subject to natural selection.
Ethologists have demonstrated that many of the behavior patterns of
animal species are innate, and moreover that they develop, like the organs,
over the life course of individuals. Thus young animals have certain infantile
behaviors which are only useful for surviving at this stage of life, when they
are strongly dependent on their parents, in a situation of direct competition
for food with other young. Complex behaviors of courtship, mating, and
care of offspring, however, mature at the same time as the reproductive
organs, and in many cases develop even if the animal has grown up in
extreme isolation, reflecting their innate rather than learned nature.
Konrad Lorenz (1903–89) discovered that the geese he studied at his
family home in Altenberg were programmed to recognize as mother the
first moving object the goslings saw at the moment when they hatched. In
normal conditions this would be the true biological mother, but they can
be made to adopt as mother a person or even an inanimate object which
appears in front of them at the decisive moment. Thus the different signals
which trigger behaviors in animals can be broken down and analyzed,
making ethology an experimental science.
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All anthropoids (Old and New World monkeys, apes, and humans) are
social, with one exception. The term ‘‘social species’’ is used to refer to
those in which lasting relationships between adults are established. The
only exception to this rule are the orangutans, which are solitary animals
in that the only stable bonds are those between mothers and their non-
adult offspring. Adult males and females only come together during the
brief and very widely spaced periods of estrus of the females. There are no
relationships or alliances between males or among females. Each female
lives in her small territory with her offspring, and the wider territories of
the males encompass those of several females, with whom, however, they
only come into contact to reproduce. Males compete with one another for
territory and for the females within it, and this competition results in them
becoming much heavier, twice the weight of females. It could be said that
male orangutans form harems, but these are harems in which the females
live dispersed rather than all together.
Gorillas, on the other hand, are highly social. As their food supply is
abundant and constantly available they do not need large territories and
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they do not travel great distances in one day. Each group of gorillas is
formed by an adult male, the silverback, and his harem, a group of females
with their offspring, all descendants of the silverback. When a female or a
male reaches puberty they leave the group. Males compete with one
another for females, and this is why they are so big, although the differ-
ence between the sexes (known as sexual dimorphism) is less marked than
in orangutans: the average female gorilla weighs about 60 percent of the
average male, or a little less than two-thirds.
Common chimpanzees do not form harems. When females reach adult-
hood they generally leave the group; however, when males reach
adulthood they stay in the group. This means that all the males in a
group of common chimpanzees are related, while the adult females are
not. Each community controls a territory, which the males defend against
other alliances of males. These struggles between groups of related males
are violent fights, sometimes to the death.
The trees which supply the fruit eaten by chimpanzees are dispersed,
and the fruit ripens at different times; for this reason within each territory
there are moments of fusion, when many individuals gather around a tree
with ripe fruit, and of fission, or dispersal in search of sources of less
abundant resources. When a female is in season, marked by a spectacular
anogenital tumescence (swelling of the zone around the sex organs), there
is no competition among males; rather, several of them mate with the
female at different times. Because of this promiscuity and the absence of
harems the males are not much bigger than the females, who on average
weigh at least 80 percent, or almost four-fifths, of the average male.
In discussing the behavior of apes we cannot emphasize too strongly that
we are talking about primates, not insects; thus their rules of behavior are
very varied and any attempt to generalize them is a blatant simplification.
For example, sometimes male chimpanzees at the top of the hierarchy have
been observed to push in front of those lower down the hierarchy when
mating with females in estrus; there are also cases where male–female
couples form, lasting several days, or even the full two weeks of estrus.
Genetic analyses of the relationship between individuals in chimpanzee
communities are currently underway; in the future these will throw a great
deal of light on how all of these behavioral strategies are reflected in
reproductive success. In order to make these ‘‘paternity tests,’’ hairs
which have fallen out in the ‘‘nests’’ the chimpanzees build in the trees
to sleep, and mouth cells left on bitten fruit, are used. On the basis of
this method, Paul Gagneux and his colleagues have observed that females
frequently ‘‘escape’’ from their community and are fertilized by a male
from another group.
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the sexual relations of each man with a woman, at least for a period of
time. In other words, something like monogamy.
But having set out what we know and what we do not know about the
social biology of our species and other species of primate, it is time to
investigate how the first hominids behaved in society.
Rob Foley observes that there are no examples among apes of groups
based on related females (whereas this is common among Cercopitheci-
dae, or Old World monkeys: only in the red colobus monkey are there
relationships between related males). He concludes that the social struc-
ture found among chimpanzees, involving alliances between related males
to defend a common territory, and dispersal of adult females outside their
birth territory, must also have been the rule among the first hominids,
whose social biology would initially have differed little from that of
chimpanzees.
However, australopithecines and especially Paranthropus lived in more
arid environments than modern chimpanzees, in sparser woods where the
sources of food would have been more dispersed and less abundant. This
leads Foley to believe that australopthecines and Paranthropus would have
had larger territories than chimpanzees. They would probably have main-
tained alliances between related males, both for defense of resources
against other coalitions of males, and against predators, which would
have become more dangerous as the tree cover decreased. However,
smaller social units would have formed within these large territories,
since the resources available in the environment would not have allowed
all members of a given community to be together at all times. Thus, the
social system would also have been one involving fusion, with much of the
group gathered around a very abundant source of food, to travel long
distances over open ground, or to sleep, and fission, with the group
separating into smaller units to find food during the day.
Determining what these smaller units might have consisted of is an-
other question. Jane Goodall and her colleagues have studied the spatial
distribution of a population of just under 150 chimpanzees in Gombe
National Park, in Tanzania, since 1960. The most closely studied com-
munity in the park was made up, over the years, of between 4 and 13 adult
males, 8 to 18 adult females and 18 to 31 immature individuals. The
group’s territory ranged from 6.75 to 15 sq km, occupying between
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3 and 6 valleys in the central region of the park. The adult females spend
more than 65 percent of their time alone with their young, feeding in
their own small areas of around 2 sq km, which partially overlapped with
those of others. The adult males are more social, and travel throughout
the community’s territory, patrolling and defending their borders to-
gether. This type of spatial distribution, with females spending time
alone with their young, is unlikely for australopithecines, even more so
for Paranthropus, because of the risk from predators in a more open
environment.
However, the family units might have been comparable to those of
hamadryas baboons (Papio hamadryas) and geladas, i.e., formed by one
male with a number of females (and their young); these may occasionally
have joined together with other small harems to form large groups to find
food, travel, or sleep. This is a social system which we know to function
successfully in ecosystems similar to those of our ancestors, like the arid
environments where the hamadryas baboons live. But one author, Owen
Lovejoy, believes that australopithecines were monogamous, and more-
over that monogamy is closely related to the development of the hom-
inids’ bipedal posture. Let us look at his argument.
Monogamy, or pairing off for reproduction, is not found only in
humans. It is also observed in many birds and primates, including gib-
bons. But as we have already noted, our species has the additional unusual
feature of a permanent sexual relationship, for most of the time without
reproductive function. To put it even more clearly, defining human sexu-
ality purely in terms of procreation is not natural (in the biological sense),
but the absolute opposite. Among humans sex also exists to keep the
couple together; in other words, it is at the service of love.
However, this romantic function of sex does not contradict Darwinist
principles, but in fact reinforces them. The long period of development of
human young makes it impossible for a mother to look after several
offspring at once (in the context of a hunter-gatherer economy). The
stable couple, in monogamy, means that the father is involved in the task
of supporting the family, which operates as an economic as well as a
reproductive unit.
There has been much debate as to what is understood by the contribu-
tion of fathers (males) to the raising of young, how this contribution can
be measured, and to what extent it occurs in the different primates and
human societies. Whatever the case, the situation in our species bears no
relation to the habits of chimpanzees, gorillas, and orangutans (our closest
relatives in other senses), where males have no connection with their
offspring; all we can say is that among chimpanzees and gorillas (not
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orangutans, which are not social), the males are tolerant of the young and
protect them against predators and being killed by other males (a serious
risk, as we shall see).
Lovejoy maintains that bipedalism has nothing to do with our ances-
tors’ adaptation to open environments, as has so often been suggested,
and that it probably arose when we still lived in the forest. Bipedalism,
according to Lovejoy, was not related to temperature regulation, to
efficiency of locomotion, or to freeing the hands to make tools. On the
contrary, bipedalism would have freed our hands and arms to carry food.
In Lovejoy’s view males transported food in this way to the base camp to
feed the females and their young, who thus avoided many dangers by not
having to travel with their mothers.
Lovejoy even believes that this made it possible to reduce the long
period between births characteristic of chimpanzees, gorillas, and orangu-
tans, and to increase the number of young over the fertile life of the
females. This supposed advantage of the first hominids is highly debat-
able, because even in modern human hunter-gatherer societies (i.e., pre-
agriculture and pre-animal husbandry) the interval between births is still
long, from three to four years.
We are used to seeing natural selection from the point of view of
individuals, but in order to understand what follows we need to look at
it from the point of view of genes. As Richard Dawkins noted in his
famous book The Selfish Gene, individuals die, but genes remain. Or
more correctly, they are preserved as copies in other bodies, those of our
children. Dawkins takes his argument to the extreme, affirming that genes
use us for their own benefit and even sacrifice our bodies if necessary, as in
the case of what is known as altruistic behavior, when parents put their
own lives at risk to save their children. In fact, the time and energy
expended in raising offspring, although less heroic, can also be considered
altruistic, since it is not aimed at satisfying the interests of the parents, but
those of the children. Everything has a point according to the logic of
natural selection, because the genes of parents who do not behave altru-
istically and abandon their children (thus dispatching them to their death)
will not be present in the succeeding generation.
These explanations enable us to understand behaviors which appear mon-
strous if we view them in moral terms. We noted earlier that when they reach
puberty, female gorillas and chimpanzees leave their native territory and their
community to enter another which is completely alien to them, and into
which they are nevertheless accepted. However, we are talking here of
females without young, because if they appear with a nursing infant in the
alien group it is virtually certain that the baby will be killed. The dominant
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male in the case of gorillas, or the related males in the case of chimpanzees,
have no interest in the genes carried by the newly arrived young individual,
but they are very interested in its mother being available as soon as possible to
be fertilized. Interrupting lactation by the rapid method of infanticide places
the mother in a situation where she can begin a new ovarian cycle. The reader
can imagine what happens to the young in a harem of gorillas when the
silverback is replaced by another male (as a result of natural death or defeat in
a fight). The logic of the genes is implacable.
If we follow this logic, a male of the first bipedal hominids providing
food for a female with young, as Lovejoy suggests, would have to be sure
that the young carried his genes. If the females of the species had periods
of estrus they would have to be watched closely during the whole of this
period. On the other hand, if the female had no estrus, making it impos-
sible to know when she was ovulating (and thus monopolize her during
this period), the only viable alternative to ensure paternity was monogamy
and sexual fidelity.
In fact menstruation may have evolved as an indicator of fertility,
because a few days after it has occurred there is a fertilizable egg. How-
ever, Beverly Strassmann points out that other primates do not appear to
make these calculations, since chimpanzees (like macaques, Old World
monkeys, and baboons), which menstruate copiously, indicate ovulation
to the males by means of swelling of the sexual organs.
As we pointed out above, social life is not preserved in the fossil record.
But we also noted that among gorillas and orangutans, males compete
with one another for females, and that as a result there are great differ-
ences in body size between the two sexes, whereas the two sexes are more
similar in chimpanzees and humans (although there are still differences; in
gibbons, which are strictly monogamous, there is no sexual dimorphism
of weight). If this is a universal rule linking anatomy to social biology, it
could offer a key to approaching the problem in extinct species. This type
of exercise, which consists of applying relationships observed in the mod-
ern biosphere to fossil species (with the premise that such relationships
have always existed), is known as actualism, and is one of the tools most
frequently used to study life in the past.
Australopithecus afarensis, the oldest hominid species of which there is
sufficient fossil record to study the difference between the sexes, shows
great variation. So much that some authors saw two or more distinct
species among these fossils; it was Tim White who grouped all the
fossils into a single species, as we have already noted. In fact this is a
species which shows great sexual dimorphism of size between males and
females (around 66 percent of body weight, very close to that of gorillas),
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170
Summary
Here ends the part of human evolution that took place exclusively in the
African continent – our birthplace, and the place where we literally took
our first steps. It is time to pause for a moment to look back, and sum up
what we have discussed and draw the main conclusions from it.
Darwinism, in the updated form of the modern synthesis, is the tool
which evolutionists of the end of the second millennium use in their
analyses. Although this is not a closed, dogmatic system and it is open to
criticism, improvement, and amendment, it forms the basis for our current
understanding of the phenomenon of evolution. One important aspect of
Darwinism is the adaptive nature of its basic mechanism, natural selection,
which does not appear to impose any specific direction on evolution. But in
this book we are describing, on the basis of the fossil record, how human
evolution arose; not until the final chapter will we discuss whether this has
any meaning, whether it corresponds to some Law of Evolution.
After setting out the terms of the problem (what happened, how it
happened, and why it happened), we have opted to follow the advice of
Chilo (usually attributed to Socrates): ‘‘Know thyself.’’ We are primates,
more specifically anthropoids, and as such we are basically visual, intelli-
gent mammals, diurnal, tropical, living in forests and trees. Many of our
morphological, physiological, and ethological characteristics match this
ecological definition of the group. The fact that humans, and to a lesser
extent other primates, now live in climates, regions, and ecosystems very
different from the backdrop to our evolution is still an anomaly, and a very
recent phenomenon in terms of the long history of the primates. In this
book we have briefly recounted this history, particularly that of the
hominoids, the class of anthropoids to which we belong.
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Summary
172
Summary
173
Australopithecus bahrelghazali Bahr el Ghazal Hadar Australopithecus afarensis
Homo
Australopithecus anamensis
Lake Turkana Paranthropus aethiopicus
Paranthopus boisei
Homo habilis/rudolfensis
Homo ergaster
Figure S.1 Location of the main deposits with fossils of Ardipithecus, Australopithecus, Paranthropus, and the first
representatives of Homo
Summary
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II
A New Home
12
Since the first humans were neither European nor Asian, but African, at
some point in the past the first human inhabitants of Eurasia must have
made their way there from Africa. When did this happen?
Since 1891, when Eugène Dubois (1858–1941) discovered a calotte (or
skullcap) and femur at Trinil, in Java, the island has supplied a large
number of human fossils. The oldest, over one million years old, are
some remains from the Sangiran region and the calvarium of a child
from Modjokerto (a calvarium is a neurocranium or braincase, a skull
without the skeleton of the face). Recently geochronologists Carl Swisher
and Garniss Curtiss and their colleagues, using the new laser fusion
method of argon-39/argon-40 dating, have dated the Modjokerto child
at 1.8 million years old, and two fragmented skulls from Sangiran (num-
bers 27 and 31) at 1.6 million years old. One partially preserved skull,
Sangiran 4, and some jawbone fragments are probably of the same age.
However, we need to treat these dates with caution, given that the relation
between the fossils and the dated volcanic sediments is not at all clear
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New Locations for Human Evolution
(these and many other fossils from Java were collected by amateurs, rather
than being professionally excavated).
Other Javanese fossils, such as the Trinil skullcap, Sangiran 2 (very
similar), and Sangiran 17 (the best preserved skull of any found in Java)
are between 500,000 and a little less than one million years old. Lastly, the
most recent of all the human fossils from Java are the Sambungmacan
calvarium and 14 more or less complete calvaria, two tibias, and pelvic
remains found in the terraces of the Solo river in Ngandong.
Eugène Dubois was a Dutch doctor who enlisted as an army doctor
with the aim of traveling to Java, where he hoped to find the missing link
between ‘‘ape’’ and man, following the theories of the famous evolution-
ary biologist Ernst Haeckel (1834–1919), who believed that we origin-
ated in the forests of Asia, rather than those of Africa. With the skullcap
and femur he found at Trinil, Dubois believed he had discovered the link
he was searching for, and he named it Pithecanthropus erectus (‘‘erect ape-
man’’), known today as Homo erectus. All the Javanese fossils are ascribed
to this species, although the Ngandong remains, the most recent, are
somewhat different, reflecting a degree of evolutionary change.
The Javanese fossils such as Sangiran 4, Sangiran 2, and the Trinil
skullcap itself differ little in their general architecture from the African
fossils of Homo ergaster. Their cranial capacity is estimated at between 800
and 950 cc. In fact the two species share many primitive traits, their
common inheritance from earlier hominids. For example, although the
brain is much larger than in australopithecines, the neurocranium is still
low (or flat), with a very broad forehead, and widest at the base, at the
level of the temporal bone; from here the side walls converge toward the
top, as can clearly be seen from the back.
However, the Javanese fossils show some characteristics not found in
Africans of the species Homo ergaster, and typical of the entire species of
Homo erectus. Essentially the neurocranium is more robust, with thicker
walls, a straight and highly developed brow ridge overhanging the eye
sockets, and another, also very conspicuous, transverse bony reinforce-
ment at the back of the cranium; this is known as the occipital ridge. There
are also some bony thickenings at the top of the cranial vault and in other
areas of the neurocranium. The side view shows that the occipital bone is
sharply angled.
Although there are fossils which do not belong to Homo erectus which
show some of these traits indicating a robust neurocranium (albeit gen-
erally in attenuated form), the combination of all these features is only
found in this species. Moreover, the base of the skull of Homo erectus
shows a series of specializations.
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The only fossil specimen from Java with a well-preserved facial skeleton
is Sangiran 17; this has a very broad, fairly flat face, which looks like a
robust version of Homo ergaster fossils such as ER 3733 and WT 15000.
Furthermore, a jawbone fragment and some cranial remains from Java
which have been very sketchily described are extremely robust, so much so
that there are still some authors who seek to create a new genus, Mega-
nthropus, on the basis of these fossils. Nevertheless, until we have more
complete fossil documentation it is too soon to accept the coexistence of
two human species in Java.
From the evolutionary point of view, it seems reasonable to accept that
Homo erectus derives from Homo ergaster, although there are important
changes which justify considering them as separate species. In 1961 Louis
Leakey found an incomplete braincase (OH 9), dated between 1.4 and
1.2 million years old, at Olduvai. From the morphology of what little
remains, this appears to represent an intermediate link between Homo
ergaster and Homo erectus. However, if it is confirmed that Java was first
populated 1.8 million years ago, it would mean that this specimen could
not occupy such a position in human evolution.
In China two presumed stone tools, a jawbone fragment with a molar
and premolar, and an isolated human incisor from the Longgupo deposit,
could be of a similar age to that currently attributed to the first Javanese
fossils. However, the age is debatable, and the jawbone may, according to
some experts, belong to a relative of the orangutans rather than a human
ancestor. The incisor is certainly human, although we do not yet have
enough information on the position of this fossil in the stratigraphy of the
deposit (the sequence of layers which make up the deposit).
Leaving aside the problematic finds of Longgupo, the oldest Chinese
fossil is a very poorly preserved skull from Gongwangling (Lantian), which
appears to be a little less than one million years old. The Chenjiawo
jawbone (also from Lantian) may be of the same age or a little more recent.
The most complete human fossil record in China was found in the
Zhoukoudian cave, about 50 km from Beijing. Since 1921, when excav-
ations began, the remains of many different individuals, including two
braincases, have been found. Unfortunately, almost all the fossils were lost
in 1941 when they were sent, for ‘‘safety,’’ from Beijing to the USA,
under the guard of US marines. The fossils never arrived at their destin-
ation because the convoy was captured by the Japanese, although it does
not appear that they succeeded in seizing the fossils; whatever the case,
nothing more was ever heard of them. As a consolation we have the
magnificent study made by Franz Weidenreich (1873–1948), and casts
of the fossils.
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New Locations for Human Evolution
182
Swanscombe
Boxgrove Bilzingsleben
Ehringsdorf
Biache-Saint-Vaast
Steinheim
Vertesszöllös
Mauer
La Chaise
Arago
Atapuerca Dmanisi
Tighenif
Figure 12.1 Location of the main European deposits of the Lower and Middle Pleistocene
New Locations for Human Evolution
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New Locations for Human Evolution
When these human fossils were found their exact age was not known,
but it was clear that they were older than the Mauer mandible. This
conclusion was based on the fauna found in TD6, particularly the pres-
ence of a fossil rodent called Mimomys savini. About 500,000 years ago
this species of rodent gave way to a primitive form of the modern water
rat, known as Arvicola cantiana. The Mauer mandible and the Boxgrove
tibia were found associated with Arvicola cantiana, while TD6 contains
Mimomys savini.
The beginning of the epoch known as the Pleistocene is usually situated
around 1.7 million years ago (for the last 10,000 years we have been in an
interglacial period of the Pleistocene, sometimes considered as a separate
epoch known as the Holocene). The Pleistocene is divided into three
parts, Lower, Middle, and Upper. The boundary between the Lower
and Middle Pleistocene is marked by a change in the earth’s magnetic
field which occurred about 780,000 years ago, when the field changed
from having an ‘‘inverted’’ polarity to the current, ‘‘normal’’ polarity. The
Lower Pleistocene belongs to the chron of ‘‘inverted’’ polarity known as
the Matuyama chron; for the last 780,000 years we have been in the
Brunhes chron of ‘‘normal’’ polarity.
Geological and paleomagnetic studies carried out by Alfredo Pérez-
González and Josep Maria Parés situated the change in magnetic polarity
which marks the boundary between the Lower and Middle Pleistocene
above Gran Dolina level 6 (TD6). Thus, the human fossils and associated
stone tools are more than 780,000 years old. This evidence forces us to
accept that the first settlement of Europe occurred much earlier than we
had thought, although human presence in the continent may not have
been as extensive or as dense as in the last half-million years.
The human fossils in TD6 represent various parts of the skeleton of at
least six individuals who died at different ages. Bearing in mind the
restricted size of the initial survey, it is hoped that when this level is
excavated extensively in a few years’ time, it will provide a very rich sample
of human fossils and tools.
The association of stone tools in TD6, which includes neither handaxes
nor cleavers, is classified as Mode 1. This attribution poses an interesting
problem, because there is Acheulian culture (Mode 2) in East Africa 1.6
million years ago, and in ’Ubeidiya (Israel) shortly afterwards. So why are
the first European fossils not associated with this later culture?
The same problem is posed by the deposits in China and Java, which
similarly have no handaxes or cleavers. There are various possible solu-
tions. Perhaps the first humans to settle in Europe and Asia abandoned
the Acheulian style of carving of their ancestors; it has been suggested that
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New Locations for Human Evolution
the populations of the Far East may often have used another raw material
for their tools, such as bamboo.
Perhaps, as Eudald Carbonell and his team suggest, the people of East
Africa who developed the Acheulian industry had a more advanced culture
than those who manufactured Mode 1 tools. As a result, their population
increased and they forced groups who did not have the superior culture to
the marginal areas of the African continent, and finally to migrate out of
Africa.
Finally, the new dating we mentioned above for Java and China sug-
gests a new argument: perhaps the first Asians left Africa before the
Acheulian culture arose. The same explanation is also possible for Europe,
but here we have no reliable dates of such antiquity for the initial settle-
ment. Not far from Venta Micena, in Fuente Nueva-3, an assemblage of
stone tools (without handaxes or cleavers) somewhat older than those of
Gran Dolina has recently been found, and it is always possible that others,
even older, may appear somewhere in Europe at some time in the future.
In any case, the first European fossils of the Middle Pleistocene, such as
the Mauer and Boxgrove fossils, are associated with Acheulian technology.
We have to wonder how the Acheulian culture came to Europe – whether
it was with a new wave of settlers, or simply that the technology traveled
from Africa, passing from some populations to others without any migra-
tion of peoples or flow of genes. The problem of whether technological
changes observed in a particular place imply the arrival of a new type of
human is a recurrent one in prehistory, and we shall come across it again.
Prehistoric Cannibalism
In theory, the human fossils should not be in the Gran Dolina cave. In fact,
human remains ‘‘should’’ never be found in karst deposits hundreds of
thousands of years old. What ‘‘should’’ be found, however, and frequently
are found, are bones of carnivores and herbivores, and stone tools. The
reason for this is very simple. The carnivores (lions, panthers, lynx, hyenas,
wolves, wild dogs, foxes) settled in caves, making their shelters and lairs
there. They lived, reproduced, and died there. Bears also spent the winter
(hibernated) in caves. Herbivores do not live in caves, but carnivores and
carrion eaters transport their carcasses there to eat them. Human beings
occasionally used caves, spending some time there, usually not very long,
making and using stone tools, and then leaving them there. Humans also
brought animal prey to the deposit, and ate it there. Finally they left.
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New Locations for Human Evolution
Homo antecessor
When new fossils are found, the paleontologist compares them with those
found earlier in other deposits, in an attempt to determine to which
species they belong. Sometimes the comparison shows that the new fossils
are unlike any others, and at this point a new species is created for them.
This is the procedure which was followed with the human fossils of Gran
Dolina. Following many studies and comparisons, in 1997 José Marı́a
Bermúdez de Castro, Juan Luis Arsuaga, Eudald Carbonell, Antonio
Rosas, Ignacio Martı́nez, and Marina Mosquera created the species
Homo antecessor (antecessor – ‘‘pioneer,’’ the one who comes before
others).
Establishing the position of the new species in human evolution is
another matter. The Gran Dolina fossils show primitive traits in dentition
and other parts of the skeleton, which is to be expected given that they
are 800,000 years old. These archaic characters are no longer found in
later European fossils, and this is why the humans of Gran Dolina are
considered to be of a different species from fossils such as the Mauer
mandible, which is approximately 500,000 years old. On the other hand,
the species represented in Gran Dolina is not Homo erectus, since it lacks
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New Locations for Human Evolution
the specializations of the latter species. Finally, these first European hu-
mans could represent a late population of Homo ergaster. However, there
are a number of indications that this is not the case.
An adolescent jawbone fragment from Gran Dolina is less robust than
those of Homo ergaster. The canine and the third molar are also reduced.
Furthermore, we have the morphology of a child who died at the age of
11, which revealed something surprising. Part of the frontal bone of this
child is preserved, mainly the right-hand half, which has a well-developed
brow ridge. There is no doubt that in an adult the ridge would be very
robust. Some of the transverse diameters of the front part of the skull have
been estimated; these suggest that the Gran Dolina child had a brain
larger than that of Homo ergaster (the brain grows very little after the
age of 11). In the three best-preserved skulls of Homo ergaster (ER 3833,
ER 3733, and WT 15000) the cranial capacity is respectively 804 cc,
850 cc, and 900 cc, while that of the Gran Dolina Child is at least 1,000 cc.
The face of the Gran Dolina Child is astonishingly modern. In Homo
habilis, Homo ergaster and, as far as we know, in Homo erectus, the skeleton
of the face is still fairly flat. Our face, however, is more sculpted, because
the nasal opening is forward of the rest of the face, and the bones of the
cheeks (the maxilla and zygomatic bone) are hollowed below the cheek-
bones, which thus project markedly. It is this combination of a primitive
frontal bone with a modern face that makes the Gran Dolina Child not
just one more fossil, but a very important specimen in our search for
information about our origins.
It was always thought that the modern face was a recent development in
human evolution, appearing with our species; suddenly we find that it
already existed 800,000 years ago. Where do we find fossils with a modern
face, of an age in between these two? We have the answer in Gran Dolina
itself, where fragments of the facial skeleton of adult individuals have also
been found, these showing a less sculpted form. We now know that as the
individual grew up, the face grew to become very large and robust, and
also more rounded as the maxillary sinuses expanded, eventually hiding
the features of the child in the adult’s face.
Hundreds of thousands of years later, our direct ancestors underwent
an enlargement of the brain which altered the structure of the neurocra-
nium, and a reduction in the masticatory apparatus, affecting the face, the
mandible, and the teeth. These are the two cranial traits which distinguish
us. The enlargement of the brain involved a fairly complete reorganization
of the neurocranium together with a marked change in its shape, but
the reduction in the masticatory apparatus was achieved in the simplest
manner possible: the facial skeleton stops developing at an earlier point,
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New Locations for Human Evolution
retaining a childish appearance. In other words, our adult face is like those
of the children of our ancestors.
So we see that the Gran Dolina fossils occupy an evolutionary position
intermediate between Homo ergaster and ourselves, the only modern
humans. Homo antecessor precedes our species but, as we shall see, it
also precedes the Neanderthals (Figure 12.2), another human species
different from ourselves which became extinct a few thousand years ago
(practically yesterday if we think in the vast terms of geological time, and
even in terms of the short duration of human evolution).
In principle it is assumed that the first humans reached the Iberian
peninsula by exclusively terrestrial means, coming from Asia and crossing
the whole of Europe. There is no reason to believe that the Straits of
Gibraltar closed at any point during the last 3 million years, although this
may well have occurred during a brief interval at the end of the Miocene,
between 6.5 and 5 million years ago (i.e., too early for humans, who did
Paleomagnetism Epochs
? H. sapiens
0 H. neanderthalensis
Upper
Brunhes
Middle H. rodhesiensis
H. heidelbergensis
−0,5
H. erectus
Pleistocene
-0,78
Years (millions)
H. antecessor
−1 Jaramillo
Lower
Matuyama
−1,5
? H. ergaster
-1,7
Olduvai Pliocene
−2
Figure 12.2 Evolutionary diagram proposed by the authors for the genus
Homo, from the initial settlement of Eurasia. The recently created species
Homo antecessor is included
189
New Locations for Human Evolution
not yet exist, to cross over from Africa). As, moreover, the currents in the
Straits of Gibraltar are unfavorable for crossing, and the first humans are
not thought to have had seafaring skills, there are no solid arguments for a
western route, directly from Africa, for the colonization of Europe. We
shall see later that the oldest known human seafaring activity occurred a
few thousand years ago, and made possible the settlement of Australia and
Papua New Guinea. And those who achieved it were humans of our own
species (when sea level falls during the ice ages, Java and England can be
reached over land, but not Australia).
But if the European populations of Homo antecessor came from Asia,
and the Asian populations from Africa, where are the fossils of this species
outside of Europe? The answer is that they have not yet been found, partly
because there are no good African fossils of the same age, and the Asian
fossils which might be contemporary with Homo antecessor are those of
Homo erectus from the Far East. Three mandibles and a parietal bone were
found in North Africa, in the Tighenif (previously Ternifine) deposit in
Algeria; these are dated at 700,000–600,000 years old, later than the
Homo antecessor fossils. There are other mandibles of similar or slightly
more recent age in East Africa and Morocco. Unfortunately in Gran
Dolina we have only a fragment of an adolescent mandible to compare
with the African specimens.
We will therefore have to wait a little longer to get to know the African
relations of the Gran Dolina fossils. From that moment on, the European
branch of Homo antecessor, represented by the humans of Gran Dolina,
and the African branch, whose fossils have not yet been discovered,
followed different evolutionary histories.
Until recently, European fossil deposits were associated with one or other
of the cold or warm periods of the Alpine glaciations, identified on the
basis of the river terraces along the upper reaches of the Danube River.
However, as we have already noted in the chapter on climate, paleotem-
perature curves, established on the basis of the ratio of heavy and light
isotopes of oxygen in the calcareous shells of microfossils obtained from
deep-sea surveys, are now used. This record is much more consistent than
the continental record, and reflects changes of temperature at a global
level, since the ratio of oxygen isotopes is related to the size of the polar
190
New Locations for Human Evolution
ice caps and the corresponding rises and falls in sea level. Paleotempera-
ture curves are divided into a series of isotopic stages, numbered from the
present backward. The present day is isotopic stage number 1. All the
odd-numbered stages are warm, and the even-numbered stages are cold.
The Middle Pleistocene is a division of the Pleistocene which runs from
the Matuyama/Brunhes change in magnetic polarity 780,000 years ago to
the beginning of isotopic stage 5. This warm stage began around 127,000
years ago, at a point when the climate was particularly warm (perhaps even
warmer than now). The Middle Pleistocene is a very important period in
human evolution because it was this period that saw the development of
the two human species we know best – Neanderthals and ourselves.
The oldest human fossils of the European Middle Pleistocene, apart
from those of Mauer and Boxgrove, are those of Arago (France) and
Ceprano (Italy), all dated at more than 415,000 years old (before isotopic
stage 11). The fossils found in the Arago cave include two mandibles, a
facial skeleton and a right parietal bone which belong to the same indi-
vidual, a coxal bone, and some other human remains. The incomplete
braincase from Ceprano was discovered in a ditch dug during construc-
tion of a freeway. Morphologically it has been described as very archaic,
close to Homo erectus. This is a very interesting specimen, but there are
still many questions relating to both its chronology and its phylogenetic
(evolutionary) position.
The Middle Pleistocene human fossils of intermediate age, situated
chronologically between isotopic stages 11 and 8 (from 415,000 years
ago to 245,000 years ago), form another group. These include the cranial
remains from Bilzingsleben, Steinheim, and Reilingen (Germany), Swan-
scombe (England), Petralona (Greece), Vértesszöllös (Hungary), and
Sima de los Huesos (another deposit in the Atapuerca Mountains in
Spain, which we describe in detail below).
The most recent European Middle Pleistocene fossils are those of warm
stage 7 (from 245,000 to 190,000 years ago) and cold stage 6 (between
190,000 and 127,000 years ago); this group includes, among others, the
remains from Ehringsdorf (Germany) and Biache-Saint-Vaast and La
Chaise-Abri Suard (France). In 1993 a complete skeleton covered with
calcareous concretion was found in the bottom of a pothole in the
Lamalunga cave (Altamura, Italy). This may be a late Middle Pleistocene
fossil.
The teeth from Pinilla del Valle (Madrid, Spain) are also from the late
Middle Pleistocene. At other Spanish sites, the Bolomor molar (Valencia),
the Lezetxiki humerus (Guipúzcoa), the humerus and coxal bones from
Tossal de la Font de Vilafamés (Castellón), and the mandible and other
191
New Locations for Human Evolution
192
New Locations for Human Evolution
193
New Locations for Human Evolution
194
New Locations for Human Evolution
that everyone at that time died very young, before the age of 40? This
latter hypothesis is difficult to entertain, since we know that the period of
maturation in this fossil population was similar to our own, and we
therefore have to conclude that their potential life expectancy was roughly
the same as ours today. Even chimpanzees, whose period of maturation is
much shorter than that of humans, live beyond the age of 40 in the wild.
This is, in any case, just one of the problems currently being investigated;
it is also a matter of great interest in relation to finding out why so many
cadavers accumulated in La Sima de los Huesos.
The present authors, together with José Miguel Carretero, Ana Gracia,
and Carlos Lorenzo, have spent many years studying the human fossils of
La Sima; among their areas of interest is the difference between the sexes
in terms of body size. Many experts maintain that since sexual dimorph-
ism in body volume was much greater in the first hominids than it is now,
there must have been a more or less constant gradual reduction over the
course of human evolution. In this case, the Middle Pleistocene fossils,
including the Neanderthals of the Upper Pleistocene, should still show
greater sexual dimorphism than we do.
Two serious problems arise when we try to verify this hypothesis. One is
the proverbial scarcity of fossils, particularly of the postcranial skeleton,
which is used to estimate body size. Thus researchers are forced to
combine fossils from deposits in different places and eras in order to put
together a sample, which is in any case very small. The problem is not so
great in the case of the Neanderthals, for which there is a much larger
(though also dispersed) sample.
The second problem is methodological. In order to compare women
and men we first need to establish the sex of the fossils. The criterion of
size is generally the one used: the large bones are assumed to be those of
males, and the smaller bones to belong to females. But when the differ-
ence in size is not enormous, as it is in gorillas and orangutans, there is a
wide band of overlap of medium-sized and small male individuals and
large and medium-sized females, in which it is difficult to establish the sex
of the remains.
In La Sima de los Huesos we have a broad sample of individuals from a
single biological population. Moreover, our approach eliminates the
methodological problem, because what we are studying is the variation
in the sample, without assigning sex a priori to the fossils; we start from
the premise that the greater the difference between the sexes in the
population, the greater will be the variation observed in the sample.
In addition to measuring the bones of the postcranial skeleton, which
reflect differences in body weight, we also analyzed variation in brain size.
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New Locations for Human Evolution
In La Sima there are three skulls whose cranial capacity is known. One is
Skull 5, with a cranial capacity of 1,125 cc; the second is Skull 4, a
complete calvarium with a cranial capacity of 1,390 cc. The third is Skull
6, fairly complete, and corresponding to an adolescent aged about 14,
which has a cranial capacity of about 1,220 cc.
The result we have obtained is that the sexual dimorphism in body and
brain size in the population represented in La Sima (from about 300,000
years ago) was no greater than our own.
As La Sima de los Huesos is not a place in which humans would have
lived, we have no information on their behavior here. However, in one of
the Trinchera del Ferrocarril (Railroad Trench) deposits of the Atapuerca
Mountains, known as La Galerı́a (the Gallery), 13 archeological levels
have been excavated which give us some idea of human activity and
technology of the same period as La Sima and a little after. Two human
fossils, a jawbone fragment and a skull fragment, have also been found in
La Galerı́a.
196
13
The Neanderthals
‘‘The Red Flower?’’ said Mowgli. ‘‘That grows outside their huts in the
twilight. I will get some.’’
Rudyard Kipling, The Jungle Book
197
The Neanderthals
198
The Neanderthals
Figure 13.1 View of a Neanderthal with a wooden spear like those found in
Schöningen
199
The Neanderthals
200
The Neanderthals
201
The Neanderthals
“Bun”
Occipital ridge
Mastoid process
Retromolar space
Chin
Figure 13.2 Neanderthal skull (top) and modern human skull (below)
202
The Neanderthals
203
The Neanderthals
204
The Neanderthals
205
The Neanderthals
Figure 13.4 Some representative Mode 3 tools. Top: Levallois core and point
extracted from it; below left : retouched scraper of the la Quina type; below right :
bifacial point (Middle Stone Age)
relationship with the local populations, those responsible for the Acheu-
lian (Mode 2) industry? Or was it only the new technique that traveled,
without population movements? We shall return to this problem later.
Two aspects of Neanderthal behavior which bear striking similarity to
our own are the use of fire and the practice of burying the dead. There are
animals which, albeit in very simple ways, select and even modify natural
objects to use them as tools. However, no animal species other than ours
knows the technology of fire, nor does any bury its dead or conduct any
ceremony for them. The practice of burial is thus a ‘‘humanizing’’ trait. It
is difficult now to imagine human life in the wild without the use of fire.
In fact, what is surprising is that there have been humans who did not have
206
The Neanderthals
fire, at least in Europe and parts of Asia (in regions far from the equator).
In many senses fire makes us human, and the absence of fire places us on
the same level as animals.
It is not known for certain when the capacity to make and control fire
first appeared among humans. It is possible, although not certain, that it
was used in the Middle Pleistocene in places like Zhoukoudian in China,
Terra Amata in France, Vértesszöllös in Hungary, La Cotte de St. Brelade
(on the island of Jersey, Great Britain), and Bilzingsleben in Germany – all
deposits which are 200,000 years old or more. However, the generalized
and systematic use of the technology of fire, with all it implies in terms of
protection, heat, light, and so on, emerged somewhat less than 200,000
years ago. From this time on there are well-structured fireplaces in the
deposits, which leave no room for doubt that fire had been mastered. At
this point the humans living in Europe, those who made the fires, were
Neanderthals, now masters of the ‘‘Red Flower.’’
It has always seemed evident that the Neanderthals buried their dead,
and many of the Neanderthal skeletons excavated in cave deposits have
historically been considered the result of this burial practice (for example,
eight Neanderthals were found entombed in the French deposit of La
Ferassie). It is less certain whether the burials were accompanied by a
ritual, a ceremony with some symbolic significance, although in some
cases it has been suggested that there is evidence of this. For example,
the earth covering the skeleton of one of the Neanderthals from Shanidar
(Iraq) contained grains of pollen from flowers which were thought to have
been placed on the body; several pairs of mountain-goat horns found
around the skeleton of a nine-year-old child in Teshik Tash cave (Uzbeki-
stan) are thought to have been placed there deliberately; on the skeleton
of a two-year-old child in Dederiyeh (Syria) a triangular flint tool was
found at the level of the heart and a limestone ‘‘tombstone’’ next to
the head; a deer jawbone was found on the pelvis of a ten-month-old
child at Amud (Israel); the Le Moustier adolescent, in France, was
thought to have been sprinkled with ochre and buried in a flexed position,
accompanied by offerings.
However, in their book In Search of the Neanderthals, paleoanthropolo-
gist Chris Stringer and archeologist Clive Gamble very reasonably call into
question this evidence of ritual, which of course is open to other, more
prosaic interpretations. A casual association of human bones with animal
remains or tools, which produces the appearance of something inten-
tional, is always possible in a deposit where there is an abundance of
animals and stone tools (and pollen is brought in by the wind). Moreover,
some of the excavations, such as that of Le Moustier, are old, and there are
207
The Neanderthals
The Neanderthals properly speaking, i.e., with all or the majority of their
typical characteristics, existed in Europe some 230,000 years ago. The
fossils from Ehringsdorf, Biache-Saint-Vaast, la Chaise-Abri Suard, and
others from the end of the Middle Pleistocene can already be considered
Neanderthal.
The key to understanding the evolutionary position of the group of
Middle Pleistocene fossils of intermediate age (between 415,000 and
245,000 years old), such as those of Steinheim, Swanscombe, Reilingen,
Vértesszöllös, Petralona, and La Sima de los Huesos, is in the broad
sample from the Spanish deposit. Although in general their morphology
is primitive, the fossils from La Sima de los Huesos show incipient Nean-
derthal traits in the occipital bone and the face, and other more clearly
Neanderthal features in the mandible, for example the presence of a
retromolar space (see Figure 12.3). The postcranial skeleton also bears
traits in common with the Neanderthals, such as the morphology of the
pubis and humerus, among others. But there are so few remains of these
bones in the fossil record outside of La Sima de los Huesos that at present
it is difficult to know at what point in human evolution these traits
appeared (it may even have been before the colonization of Europe).
The Steinheim skull may possibly have been similar to those of La Sima
de los Huesos, though it is difficult to tell because it is incomplete and
badly deformed. More fragmented fossils, such as those from Swan-
scombe, show a more typically Neanderthal occipital morphology (with
a ridge that is depressed in the center and a broad suprainiac fossa). The
208
The Neanderthals
Petralona skull has a more Neanderthal face than those from La Sima de
los Huesos, but its occipital bone is less Neanderthal. Taken together, this
group of fossils indicate that the Neanderthals evolved in Europe over
hundreds of thousands of years in geographical and genetic isolation. In
this sense the Neanderthals are the original Europeans, a native, local
human species – what is known in biology as an endemic species.
The Neanderthals’ roots extend far back into the past, but how far back
do they go? Some authors, such as Chris Stringer, group the Mauer
mandible with the more modern fossils of Arago, Bilzingsleben, and
Petralona together to form a species known as Homo heidelbergensis,
with the Mauer mandible as the type specimen.
African specimens dated at between 600,000 and 250,000 years old,
such as the skulls of Bodo (Ethiopia), Ndutu and Eyasi (Tanzania), Salé
(Morocco), Elandsfontein (South Africa), and Broken Hill (Zambia), are
added to the European fossils of this species.
It is also suggested that Homo heidelbergensis is represented in Asia by
the Dali skull and the Jinnishuan skeleton, both from China, and dated at
between 200,000 and 300,000 years old. Precise dating of these fossils is
important, because they might be contemporaneous with the last Homo
erectus fossils in China, Skull 5 from Zhoukoudian, and an incomplete
skullcap from Hexian cave.
According to this theory, the place of Homo heidelbergensis in human
evolution would be that of the last common ancestor of Neanderthals and
modern humans (Figure 13.5). The cranial capacity of the species ranges
from 1,000 to 1,400 cc, the lower value in the range approximating to the
average for Homo erectus, and the upper to the average for Neanderthals
and modern humans.
In a recent article Robert Foley and Marta Lahr elaborate an evolution-
ary model which they call the Mode 3 hypothesis. According to them, the
species which invented Mode 3, between 300,000 and 250,000 years ago,
was a species which evolved in Africa from Homo heidelbergensis. They
suggest that this new species (whose Latin name we shall omit, so as not to
add further to the terminological confusion) had a larger brain and was
more intelligent than previous species, as demonstrated by their advanced
(Mode 3) technology. Some members of the Mode 3 Species migrated to
Europe around 250,000 years ago, in time replacing the European popu-
lation of Homo heidelbergensis and evolving into the Neanderthals. In
Africa the Mode 3 Species evolved to create our species, Homo sapiens.
However, in our view all these authors are mistaken. Our analysis of
morphological traits leads us to conclude that the European populations
such as that of Mauer, living around 500,000 years ago, were already on
209
The Neanderthals
Paleomagnetism Epochs
? H. sapiens
0 H. neanderthalensis
Upper
Brunhes
Middle
H. heidelbergensis
−0,5
Pleistocene
H. erectus
-0,78
Years (millions)
−1 Jaramillo
Lower
Matuyama
−1,5
? H. ergaster
-1,7
Olduvai Pliocene
−2
the evolutionary line of the Neanderthals. The rules for creation of species
state that the name of the species should be related to its type specimen;
as the type specimen for Homo heidelbergensis is the Mauer mandible,
this species should actually be exclusively European and the ancestor of
Neanderthals. The last common ancestor of Neanderthals and modern
humans is Homo antecessor, created on the basis of the Gran Dolina
fossils, which are more than 780,000 years old (see Figure 12.2). As we
shall see later, there are data from molecular biology which support
our position.
The species Homo heidelbergensis, as we understand it, would cover
fossils from the Mauer mandible to the fossils of La Sima de los Huesos
and all those in which primitive traits predominate, although they may
show some incipient characters indicating that they are ancestors of the
Neanderthals. The fossils of the end of the Middle Pleistocene (from
about 230,000 years ago onward), on the other hand, can to all intents
and purposes be considered true and complete Neanderthals.
210
The Neanderthals
The two skulls from Saccopastore (on the outskirts of Rome) and the
set of fossils from Krapina (Croatia) are from the beginning of the Upper
Pleistocene (the latter have been reliably radiometrically dated, but the
date for the Saccopastore fossils is less certain).
From this time on Neanderthals appear in abundance in European
deposits (relative, of course, to the general scarcity of the fossil record),
and they are also found in Central Asia and the Middle East, regions to
which they migrated from Europe. Fossils as emblematic as those of Le
Moustier (which gives its name to the Mousterian), Guattari 1 (Monte
Circeo), and La Chapelle-aux-Saints lived in Europe less than 60,000
years ago. These Neanderthals show some new characteristics with respect
to earlier specimens, and are often called ‘‘classic Neanderthals.’’ The
description of a Neanderthal that we gave earlier corresponds most par-
ticularly to these classic forms. As we shall see, there were still Neander-
thals in Europe 30,000 years ago, perhaps even more recently, before their
trail disappeared forever.
The Spanish record of the Upper Pleistocene is rich in Neanderthals
and includes fossils from the deposits of Agut (Barcelona), Axlor (Viz-
caya), Cova Negra (Valencia), Gibraltar (Devil’s Tower and Forbes
Quarry), Gabasa (Huesca), La Carihuela (Granada), Los Casares (Guada-
lajara), Mollet I (Gerona), and Zafarraya (Málaga). There is good reason
to hope that many more Neanderthals will be discovered in the Iberian
peninsula over the next few years.
211
14
212
The Fossil Evidence
Figure 14.1 Representative Mode 4 tools. Top: end-scraper; left : bone spear;
right : double-beveled burin
213
The Fossil Evidence
214
The Fossil Evidence
215
The Fossil Evidence
and bison on the walls of the Chauvet cave, and 10,000 years after the first
modern humans settled in the Iberian peninsula. However, this is not to
give the impression that modern humans advanced from east to west like a
steamroller, crushing all the Neanderthals in their path. The two species
must have shared the map of Europe for thousands of years, forming a
mosaic of intermingled populations. As we have seen, about 40,000 years
ago there were already modern humans in the regions of Catalonia and
Cantabria in Spain, and thousands of years later Neanderthals still sur-
vived well to the north of the Pyrenees. We may imagine that the pockets
of Neanderthal populations gradually diminished until the last of them
disappeared.
This long coexistence of Neanderthals and modern humans (popularly
known as Cro-Magnon Man) is one of the chapters of prehistory which has
most excited popular attention, giving rise to tales such as La Guerre du Feu
(Quest for Fire, 1911) by J. H. Rosny-Aı̂né, made into a film by Jean-
Jacques Annaud in 1981, and Jean M. Auel’s saga The Clan of the Cave
Bear. One particularly intriguing aspect of this coexistence between Nean-
derthals and modern humans is the origin of the Chatelperronian and other
similar technologies. Had the Neanderthals developed the use of bone,
horn, and ivory to make tools and decorations, and the production of long,
thin stone blades, of their own accord? Did they do it independently in a
number of regions of Europe? Or did they copy it from modern humans by
watching them work, or examining abandoned objects? Might some of the
elements found in the Grotte de Renne at Arcy-sur-Cure, such as the ivory
pendants, come from an exchange between the two types of humans? As yet
we have no solution to these puzzles.
216
The Fossil Evidence
from the population of the father or the mother, or with one another. The
new species must be genetically isolated from others. In the case of the
Neanderthals, we know of no fossil hybrids of them and our ancestors;
moreover, modern Europeans do not carry genes inherited from the
Neanderthals. However, the fact that there was little exchange of genes
does not mean that it was impossible, and this is the condition for defining
a species on the basis of genetic criteria.
In paleontology, the evolutionary concept of the species proposed by
George Gaylord Simpson (1902–84) makes more sense. According to this
concept, a species is a continuum of populations which succeed one
another in time and follow their own evolutionary trajectory, independent
of other species, and continue over a considerable period: what counts is
that there is genetic continuity between generations and that isolation is
maintained. According to this criterion, Neanderthals would be an ‘‘evo-
lutionary species.’’ Something similar occurred with the two species of
chimpanzee, which are separated by the Congo River: common chimpan-
zees live to the north, and pygmy chimpanzees or bonobos to the south.
The different species of gibbon also developed through geographical
separation.
However, according to Simpson’s criterion the species Homo heidelber-
gensis, as we understand it, would not exist, since fossils like those of La
Sima de los Huesos are the ancestors of the Neanderthals and therefore
belong to the same ‘‘evolutionary species’’: they are simply very primitive
Neanderthals. While we recognize the validity of this argument, we be-
lieve that given the morphological difference between the two types of
fossils, the species Homo heidelbergensis should be retained for practical
reasons, as is normally the case in paleontology (and particularly if we look
at things from the common-sense point of view, which dictates that we
should give different names to things which are distinct from one an-
other).
On the other hand, if the Neanderthals were not human, no one would
dispute that they deserve their own species. However, many researchers
find it difficult to accept that human beings who buried their dead and
used fire were of a different species. What is more, European Neanderthals
even came to manufacture tools similar to those of modern humans (the
Chatelperronian industry). Lastly, we shall see below that the industry of
the first modern humans (in the Levant) was the same as that of the
Neanderthals (Mousterian).
However, all of these coincidences only mean that Neanderthals were
intelligent. The fact that we are the only intelligent human species cur-
rently in existence does not mean that it has always been so. We are also
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The Fossil Evidence
now the only bipedal primate species, and we have seen that in the past
there were several species of bipedal hominids living at the same time. The
Neanderthals represent another intelligent human species, among other
reasons because the common ancestors of Neanderthals and modern
humans (Homo antecessor) were also intelligent. There is nothing to rule
out the possibility that the different intelligent human species exchanged
information, produced the same type of tools, and shared the technology
of fire; two intelligent species can exchange information without exchan-
ging genes.
To put it another way, if an alien spaceship landed on our planet the day
after tomorrow, the beings which came out of it would no doubt be
intelligent, and we would find a way to communicate with them. Never-
theless, because they evolved in a different place, the aliens would be of a
different biological species from ours. Something similar happened, albeit
with fatal consequences for the Neanderthals, when our ancestors arrived
on the European continent for the first time.
In any case, assuming that technical change came with biological
change, as some researchers advocate, would imply four separate succes-
sive colonizations of Europe: first the manufacturers of Mode 1 (repre-
sented by the fossils of Gran Dolina) would arrive, then those who made
Mode 2 (Homo heidelbergensis), then those of Mode 3 (the ancestors of
the Neanderthals), and finally those of Mode 4 (modern humans). We
believe, on the contrary, that cultural diffusion between human popula-
tions or species was more frequent than replacement of one by another,
and therefore that Europe was settled only twice: first by Homo antecessor
about 800,000 years ago, and then by our ancestors around 45,000 years
ago.
There are researchers who believe they see a substantial difference, a
chasm, between the Neanderthal mind and that of modern humans. Thus
our superior cognitive capacity is manifested by, among other things, the
fact that we are the only ones capable of devising aesthetic and symbolic
concepts. We have already seen that there is debate as to whether the
Neanderthals buried their dead with ceremony. For us the mere fact that
they buried them already implies a ritual, and therefore a capacity for
symbolism.
On the other hand, we have to acknowledge that the explosion of art
occurred in the Upper Paleolithic. Before this there is some very doubtful
evidence, such as the series of cuts on bones found in the Bilzingsleben
deposit in Germany, over 350,000 years old, or the supposed sculpture of
a woman from Berekhat Ram in Israel, which is over 230,000 years old.
However, although the ancestors of the Neanderthals did not have art,
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The Fossil Evidence
nor did our ancestors at that time. The great examples of artistic expres-
sion like the paintings and engravings on walls and stone panels, and the
sculptures of animals and people, are found not in the early part of the
Upper Paleolithic, between 50,000 and 45,000 years ago, but only from
just before 30,000 years ago. It is possible that the Neanderthals did not
come to produce art simply because they became extinct before artistic
activity became widespread. However, we have already noted that 34,000
years ago the Neanderthals of the Grotte de Renne had items of decor-
ation similar to those of their modern human contemporaries, indicating
that they had an aesthetic sense.
But . . . if modern humans did not evolve in Europe from the Neander-
thals, where did they come from? In order to answer this question we
need to look at the fossil record of the Middle East.
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The Fossil Evidence
Figure 14.2 Reconstruction of the head of a woman from the Jebel Qafzeh
deposit
220
The Fossil Evidence
A skull fragment (basically the frontal part of the skull) of uncertain date
found in 1925 in the Zuttiyeh cave, also in Israel, corresponds to an even
older population in this region. But it is not clear whether this is an
ancestor of the Neanderthals or of modern humans, although the latter
seems more probable.
The Neanderthals and the modern humans described here share a
common class of stone technology, the Mousterian, within the great
category of Mode 3 or Middle Paleolithic industries.
We might imagine that the Neanderthals were in the Levant first, and
that modern humans came later and replaced them, as happened in
Europe. There is even the possibility that modern humans evolved
from Neanderthals in the Levant and then migrated to Europe to replace
the Neanderthals there. It is more difficult to imagine that the two human
types were contemporaneous in Israel, and that the Neanderthals lived in
Tabun cave a few minutes away from the modern humans living in the
Skhul rock shelter. This is why it is so important to know the age of both.
The human fossils from Tabun, Skhul, and Jebel Qafzeh have been
dated by the TL, ESR, and uranium series methods, giving an age of
about 100,000 years. However, we need to be more specific in the case of
the Tabun deposit. This cave was excavated between 1929 and 1934 by
the English archeologist Dorothy Garrod, who found a female skeleton
(Tabun 1) and an isolated mandible (Tabun 2). The skeleton is clearly
Neanderthal; the species identification of the mandible (Tabun 2), on the
other hand, is more doubtful, and it might equally belong to a modern
human. Both remains come, in principle, from the level dated at about
100,000 years old. However, Garrod herself admitted the possibility that
the female skeleton was of more recent date and corresponded to the level
above; the people of this upper level might have dug a hole to bury her
and placed her in the lower level.
The Neanderthals of Kebara, Amud, and Dederiyeh are more recent,
between 85,000 and 50,000 years old. These fossils are contemporaneous
with the ‘‘classic’’ European Neanderthals. If the Neanderthal Tabun 1
skull also belonged to this period (which is not certain), and the Tabun 2
mandible was that of a modern human, then all the data would indicate
that the ancestors of modern humans were the first to occupy the region,
as an extension of the settling of Africa; they were then replaced by the
Neanderthals. Archeologist Ofer Bar-Yosef believes that the Neanderthals
colonized the Levant during a period of intense cold in Europe, which saw
a displacement of the populations of central Europe toward the Mediter-
ranean.
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The Fossil Evidence
Finally, between 45,000 and 50,000 years ago the Mode 4 (or Upper
Paleolithic) industries appear in the Middle East, and we may assume that
those who made these tools were the modern humans who, having arrived
from Africa, had replaced the Neanderthals, then spread through the rest
of the world and replaced all other populations of other kinds of humans
they found on their way.
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The Fossil Evidence
spread of modern humans. The robust Australian fossils are in fact rem-
iniscent not of Homo erectus but of the first modern humans of Israel and
Europe.
As in Europe, this spread of modern humans into Asia had fatal conse-
quences for the native populations. In the easternmost area of the world
inhabited by humans, the island of Java, lived the last Homo erectus,
descendants of the populations represented by the Javanese Lower and
Middle Pleistocene fossils. We have a magnificent collection of braincases
of these last Homo erectus, found in the terraces of the Solo River in
Ngandong. As far as is known, in Java as in Europe a local process of
evolution, with a degree of geographic and genetic isolation, took place
here. Morphologically the Ngandong braincases show clear evolutionary
continuity with the older Javanese fossils, although brain size has in-
creased (with cranial capacities ranging from 1,035 to 1,225 cc), modify-
ing the structure of the neurocranium somewhat. The Sambungmacan
braincase is generally considered a good example of a fossil intermediate
between archaic specimens like those of Trinil and Sangiran and the
Ngandong skulls.
It has traditionally been thought, on the basis of the fauna and geology
of the region, that these human fossils from Ngandong belong to the
Upper Pleistocene (less than 127,000 years old), while the Sambungma-
can braincase is usually thought to be older, over 200,000 years old. The
Ngandong Homo erectus would thus be contemporary with the Neander-
thals and modern humans. If this chronology is correct, it would mean
that three different human species coexisted until very recently. Carl
Swisher and his colleagues have used uranium series and ESR dating to
date bovid teeth associated with the human fossils from Ngandong and
Sambungmacan. The result is very surprising, giving a range of ages
between 54,000 and 27,000 years. But once more, we need to wait for
this new dating to be confirmed, given that some eminent geochronolo-
gists, such as Christophe Falguères, disagree and believe the Ngandong
and particularly the Sambungmacan fossils to be considerably older, all
more than 200,000 years old.
We have already seen that the first humans who were basically like our-
selves, albeit with some archaic traits, were to be found in the Middle East
(Skhul and Qafzeh) between 90,000 and 120,000 years ago. In South
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The Fossil Evidence
224
The Fossil Evidence
a large brain volume (over 1,350 cc), and braincases which have become
less robust and are rounded in shape (with, for example, less angled
occipital bones than in previous forms). However, as Giorgio Manzi
points out, they have yet to undergo the radical transformation which,
undoubtedly in just one time and place, converted a more or less low
neurocranium with a flat forehead to a rounded, almost spherical neuro-
cranium, widest in the middle and with a high cranial vault and vertical
forehead – the shape we recognize as our own.
225
15
The value and utility of any experiment are determined by the fitness of
the material to the purpose for which it is used.
Gregor Mendel, Experiments in Hybridization of Plants
A Brilliant Idea
The origin of modern humanity has always been one of the most contro-
versial questions in human paleontology. Many paleontologists believe,
like Gunter Bräuer and Christopher Stringer, that modern humans ori-
ginated in Africa between 300,000 and 100,000 years ago. From this
African birthplace, our species spread through the rest of the Old World
and replaced the other human species (Neanderthals and Homo erectus)
which had appeared as a result of local evolution, in conditions of repro-
ductive isolation, in Europe and Asia. This has been dubbed the ‘‘Out of
Africa’’ hypothesis, in reference to Isak Dinesen’s wonderful book, on
which the film of the same name was based.
Moreover, the idea that Neanderthals and modern humans do not form
a sequence of the ancestor-descendant type, but belong to two independ-
ent evolutionary lines which separated very early on, has also been pro-
mulgated by various anthropologists since it was first put forward in 1912
by Pierre Marcelline Boule (1861–1942).
In the preceding chapters we have followed these approaches to
the origin of modern humanity and its evolutionary relationship with
226
The Genetic Evidence
the Neanderthals since, in our opinion, the fossil evidence fully confirms
this relationship.
However, it must also be admitted that other paleoanthropologists
find, in the same fossils, evidence of a multiple and very ancient origin
for modern humanity. These researchers believe that each of the different
human populations which occupied the Old World after the first depart-
ure from Africa, over one million years ago, evolved in their separate
geographic regions to give rise to the human populations (the different
‘‘races,’’ to use an older term) which today inhabit the globe.
In its original formulation this hypothesis, championed by Franz Wei-
denreich and Carleton Coon (1904–81) among others, suggested that
each human line had evolved independently and in parallel with the
others. This is not a Darwinist view, since it postulates that different
populations which evolve separately in different environments will ultim-
ately converge into the same species. In order to resolve this problem the
modern version of this theory, whose principal proponents are Milford
Wolpoff and Alan Thorne, suggests that there was a gene flow between all
the Pleistocene populations distributed throughout Africa, Asia, and
Europe. This gene flow, it is suggested, was of sufficient magnitude to
maintain the homogeneity of the human species dispersed through three
continents, but not so intense as to eliminate certain specific traits which
characterize humans in each region. This hypothesis is now known as the
hypothesis of multiregional origin.
The main reason that contradictory theories of the origin of modern
humans continue to exist is the nature of the fossil record. Paleontologists
are trying to unravel a process which took place over hundreds of millions
of years in three continents and involved thousands of individuals. In
order to carry out this task they have no more than a handful of fossils,
often fragmented, isolated, and dispersed in time and space. There is no
doubt that the missing pieces of the jigsaw are more numerous than the
ones we have.
The discovery of new fossils, the precise dating of these, and the ever-
deepening knowledge of the biology of species are the tools paleontolo-
gists use to support their hypotheses. But this procedure is slow and
tortuous, and depends in large measure on the pure chance of paleonto-
logical discoveries. Ideally, we need to have recourse to data from a field
independent of paleontology in order to put the hypotheses based on the
fossils to the test. But where can we turn to find such data?
The knowledge gained over the 20th century about the mechanisms of
genetic inheritance allows us to consider a new way of approaching the
problem of the evolutionary history of species. The idea is as simple as it is
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The Genetic Evidence
brilliant: it does not matter that there are few fossils of past species, since
the genetic material in living species contains the key to their evolutionary
history. We just need to know where to look.
We have already seen an example of this approach in the chapter on the
origin of hominids and their relation to gorillas and chimpanzees. But the
problem of the origin of modern humanity is different because only one of
the species involved in the process survives, and we therefore only have
genetic material from one of them (with one exception, which we shall
look at later in this chapter). Genetic studies dedicated to clarifying the
origin of modern humanity attempt to determine the genetic structure of
humans today; on this basis inferences can be drawn about the how, when,
and where of our origin.
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The Genetic Evidence
Black Eve
Our cells obtain their energy from a series of very complex chemical
reactions, most of which, particularly those which involve oxygen, take
place within a series of small organelles known as mitochondria. In add-
ition to performing this important role in cell life, the mitochondria are
exceptional in another way: they are the only organelles in the animal cell
which have their own genetic material.
The DNA of a mitochondrion is contained in a circular chromosome
smaller than the chromosomes of the cell nucleus, and very similar to that
of bacteria. Mitochondrial DNA is ideal for evolutionary studies for two
reasons: first because all variation is due exclusively to mutation, since it is
not subject to recombination, and secondly because the organelles in the
egg cell come only from the maternal ovum and are transmitted through
the mother’s line (in the process of fertilization the sperm only contrib-
utes its nuclear chromosomes, so the egg cell consists of the ovum plus the
nuclear chromosomes of the sperm).
Thus, we can trace the ancestry of a mitochondrial chromosome, from
woman to woman, over the generations. The mitochondrial DNA (for
which we shall use the standard abbreviation, mtDNA) in any of our cells
can be identified with a single ancestor in each generation: our mother,
our maternal grandmother, only one of our four great-grandmothers (the
mother of our grandmother), and so on.
One might naı̈vely suppose that geneticists analyze the entire mtDNA
of each individual to identify its peculiarities in order to compare them
with those of others, but such a task would require a great deal of time,
resources, and effort. Studies on variation in mtDNA are actually confined
229
The Genetic Evidence
230
The Genetic Evidence
result of chance (for example, the mtDNA of women who only give birth
to males will cease to be represented in the population); this alone would
be enough to explain the limited variation in non-African populations.
Another criticism is based on the argument that mtDNA studies may
offer a biased vision of the evolutionary history of humans, since they
consider only the history of women, which might not be the same as that
of the overall population.
These objections have met responses and counter-arguments, drawing
in a large number of scientists. Since all the arguments put forward are
reasonable, the only way to resolve the problem is to look for new
evidence by studying nuclear DNA.
We will return to the issue of calculating the time elapsed since the
origin of modern humans later in this chapter.
The best way of comparing results and interpretations based on the study
of mtDNA is by studying the variation of a part of the nuclear DNA which
is transmitted paternally and which, like the mitochondrial chromosome,
is not subject to recombination. The only nuclear chromosome fulfilling
these criteria is the Y-chromosome.
The variation in some of the polymorphisms identified in the
Y-chromosome can be categorized as one of a few types (or haplotypes)
among which we can determine which is the most primitive (by compar-
ing them with the condition found in modern apes). This is a valuable
innovation with respect to earlier studies of the mitochondrial chromo-
some, in which the primitive type was identified on the basis of
the distribution of different types of mtDNA among the populations
analyzed.
The results of various analyses of different polymorphisms of the
Y-chromosome all point in the same direction: modern humans had a
male ancestor who lived in Africa between 100,000 and 200,000 years
ago. The two most recent studies, carried out by teams led by Michael
Hammer and Peter Underhill, go even further, indicating that the
Khoisan people (the Bushmen) are the human population with the
highest frequency of primitive haplotypes.
But in addition to these results relating to the origin of our own species,
the Y-chromosome studies have given us very valuable information on
other aspects of our evolutionary history. First, that there was no one
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The Genetic Evidence
232
The Genetic Evidence
Pleistocene Park
Michael Crichton’s novel Jurassic Park, the starting point for Steven
Spielberg’s film series, is based on the possibility of recovering intact
DNA from fossils. When Crichton wrote his book, a series of scientific
articles reporting on the discovery of dinosaur DNA through insects
fossilized in amber around 100 million years ago had just been published.
However, these studies have now been completely discredited and it has
been recognized that the DNA found was due to modern-day contamin-
ation. Moreover, it has been demonstrated that it is impossible to find
fossil DNA of such an age, for the simple reason that the DNA molecule
cannot survive unaltered for this length of time; not even amber can
prevent the oxidation of DNA and its consequent deterioration.
The oldest DNA which has been recovered from fossils is much more
recent than that of the dinosaurs in Jurassic Park; it is mtDNA from
mammoths preserved in the ice of Siberia between 50,000 and 100,000
years ago. The low temperatures seem to have favored the preservation of
the mammoths’ mtDNA, and there is no expectation that anything similar
will be found outside of very specific environments such as the Siberian
permafrost.
One of the main problems for the ‘‘DNA paleontologist’’ (apart from
that of whether fossil DNA actually exists) is avoiding contamination with
modern DNA through the manipulation inherent in the excavation,
restoration, study, and other work carried out on fossils. The efforts
made to isolate mtDNA from the famous Ice Man of the Tirol (5,000
years old) offer a good example. The first attempts were thwarted by the
presence of contaminating modern DNA, and the techniques had to be
refined in order to find mtDNA whose authenticity was not in doubt.
Although the results obtained from the study of the mtDNA from the
Ice Man were modest (his mtDNA was shown to belong to a type
characteristic of central European populations), the study did offer some
very valuable lessons on the techniques and controls necessary in order to
get around the problem of contamination with modern human DNA.
Using these new procedures, an international team (comprising Mat-
thias Krings, Anne Stone, Ralph Schmitz, Heike Krainitzki, Mark Stone-
king, and Svante Pääbo) embarked on the search for mtDNA in
Neanderthal fossils. They took samples from the type specimen of this
species, the Neanderthaler skeleton. The study was very carefully planned
and surrounded with all possible precautions, both to avoid contamin-
ation as far as possible and to prevent pointless destruction of a fragment
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The Genetic Evidence
of this valuable fossil. The 3.5-g sample was taken from a place theoret-
ically inaccessible to contamination, the inner part of one of the bones –
specifically, the right humerus.
The next step in the study was to analyze the structure of the fragments
of what was presumed to be Neanderthal mtDNA. The DNA molecule is
made up of thousands of smaller units known as nucleotides, of which
there are only four different types in DNA. The structure of a fragment of
DNA is simply the sequence of nucleotides which make it up, so this part
of the work consisted of determining the nucleotide sequence in the
different fragments of DNA found in the sample taken from the fossil.1
Many of these fragments showed sequences which were identical over
some part of their length: in other words, they overlapped. This made it
possible to reconstruct the sequence of the original segment from which
they came. Thus, after three months of intensive work, the sequence of a
segment of 379 nucleotides was reconstructed, on the basis of 123
different fragments. This sequence corresponds to region I of the control
segment of mtDNA.
The researchers compared the mtDNA sequence from the fossil with 16
types of mtDNA from chimpanzees and 1,986 types from different mod-
ern human populations, and obtained some extremely valuable results.
First, they determined that on average humans and chimpanzees differ in
55 bases of the base sequence, while the mtDNA analyzed showed an
average difference of 27 bases from modern humans. Moreover, they
established that within the modern human sample the average difference
was only 8 bases. Thus, the fossil mtDNA is sufficiently similar to our own
to recognize that it comes from a human being, but different enough to
reject the suggestion that it belongs to a modern human, thus ruling out
the possibility of contamination and confirming it as an authentic frag-
ment of Neanderthal mtDNA.
The fact that the average difference between Neanderthal mtDNA and
that of modern humans is more than three times greater than the average
difference among modern humans (27 bases compared to 8) led the
1
A series of fragments of mtDNA, representing two types of sequence, was obtained
from the Neanderthal fossil. One of these sequence types was attributed to modern
mtDNA, present through contamination, while it was thought that the other might
correspond to the authentic mtDNA of the fossil. It is striking that despite all the
precautions taken, which theoretically rendered the presence of contaminating DNA
impossible, it did appear in the analyses. This fact indicates how difficult it is to avoid
contamination in this type of work, and throws into question the results of other, less
rigorous studies.
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The Genetic Evidence
researchers to conclude that the two lineages separated a very long time
ago.2 In order to establish when this separation occurred, the researchers
used a mutation rate determined on the basis of the average number of
changes observed between modern humans and chimpanzees (55) and
the time which is assumed to have passed since the two lines separated.
Their calculations gave an age of between 550,000 and 690,000 years for
the Neanderthal/modern human separation, and between 120,000 and
150,000 years for the origin of present-day human diversity.
The conclusions about the age of modern humans concur with those
derived from studies of mtDNA, the Y-chromosome, and other nuclear
chromosomes. One secondary result of the analysis is that the mtDNA of
modern humans of African origin again appears ancestral to that of the
other modern human populations.
In addition to reinforcing the theory of a single, African origin for
modern humanity, this study has another noteworthy feature of particular
relevance. The separation between Neanderthals and modern humans is
traced much further back than the majority of authors working on fossil
studies had suggested up to this point. However, this separation date is
compatible with the results of our investigations of the human fossils of
Atapuerca, both from La Sima de los Huesos and from Gran Dolina level
6. As we have already noted, the Gran Dolina fossils, which are close to
800,000 years old, represent the ancestor species of both the Neanderthal
and the modern human lines, Homo antecessor.
Perhaps the discovery of Neanderthal fossil mtDNA will give rise to a
new novel, a tale of cloning Neanderthal individuals. However, one of the
conclusions which can be drawn from our better knowledge of fossil DNA
is that we will never find sufficient nuclear DNA to allow us to dream of
the Pleistocene version of Crichton’s book.
It has frequently been suggested in the popular media that genetic studies
just by themselves have revealed the ‘‘how,’’ ‘‘when,’’ and ‘‘where’’ of our
2
Nevertheless, some comparisons of mtDNA within modern humans gave a greater
average difference (up to 24 bases) than that obtained from comparison of Neanderthal
mtDNA with that of certain modern humans (20 bases). Thus the results of the study
are not as convincing as has been suggested, but should rather be considered statistic-
ally reliable.
235
The Genetic Evidence
236
The Genetic Evidence
Standards of Beauty
Up to now we have been considering our species’ past, but what of its
future? Many believe that, now we have machines enabling us to depend
less and less on our physical strength to survive, and more and more on
our intelligence, the corresponding organs will be affected in the future.
Thus the human being of tomorrow is frequently represented as having an
atrophied body and a very big head, or rather, with an overdeveloped
brain, since the face and teeth are also depicted as reduced – in short, a
very unattractive individual, in terms of the ancient Greek canons of
237
The Genetic Evidence
238
16
Thus it was necessary for the said hole in the Adam’s apple to close when
meat is swallowed that it does not enter the throat, which would be a
vexing and harmful thing, of which we have experience every day when
we hurry in swallowing and something enters our throat, because there
follows from this a most troublesome and grievous coughing.
Bernardino Montaña de Monserrate, Libro de la anathomia del
hombre [Book of the Anatomy of Man]
Human beings are the only organisms which speak. That is to say, we
transmit to our fellows, and receive from them, all kinds of new informa-
tion, deliberately encoding our messages in combinations (words) of
pre-established sounds (syllables). Other animals are only capable of
exchanging very specific information on certain aspects of their life,
using a limited system of sounds and gestures which are not intentionally
encoded.
Konrad Lorenz, in a delightful book entitled King Solomon’s Ring,
makes reference to the legend that King Solomon possessed a ring
which enabled him to talk with beasts. Lorenz boasted that he too was
able to understand the simple vocabulary of animals, without needing a
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The Origins of Human Language
ring; but he added that animals do not have a true language; rather, each
individual innately possesses a code of signals formed by expressive sounds
and movements of which another individual of the same species has a
similarly innate understanding. However – and this is where they differ
fundamentally from humans – animals emit these signals automatically
when they find themselves in a particular state of being, even if there is no
one to witness them. Lorenz expressed this idea by saying that with their
sounds animals are emitting not ‘‘words’’ but ‘‘interjections.’’ The
Merriam-Webster Dictionary defines an interjection as ‘‘a cry or inarticu-
late utterance expressing an emotion.’’ And this is exactly what animals are
emitting, according to Lorenz.
These ideas appear to be true for the majority of animals, but they are
perhaps not quite so accurate in the case of primates. Dorothy Cheney and
Robert Seyfarth have studied vervet monkeys (Cercopithecus aethiops) in
the wild in Africa, observing that in addition to emitting signals, in the
form of sounds and gestures, which communicate their motivation or
state of mind, these animals also inform one another about certain aspects
of the environment. For example, they have different sounds to warn of
the presence of different predators – snake, eagle, or leopard. The reac-
tions that these calls stimulate in the hearers are different in each case: if
the warning is of a leopard they climb trees; if the predator is an eagle they
hide in the bush or look up; and if a snake is signaled they stand up and
scan the grass. Thus, each call has a distinct meaning which triggers a
different response; these are not simply cries of fear in reaction to the
presence of a predator.
Moreover, Cheney and Seyfarth’s research on other calls related to the
social life of the vervet monkey has revealed that these monkeys associate
sounds which have similar meanings, even if they are acoustically very
different (just as we would with the words ‘‘car’’ and ‘‘vehicle’’).
In the 1960s and 1970s the idea of direct communication with the
animals most like us, chimpanzees and gorillas, was taken very seriously in
some research programs. Since chimpanzees and gorillas are physically
unable to pronounce words, the task of communicating with us was
facilitated by teaching them sign language, which they were able to
reproduce. The chimpanzees and gorillas proved to be outstanding pupils,
and revealed their capacity for associating ideas which we express in words
with gestures, or with tokens of different shapes, with drawings and colors
(icons).
The bonobo Kanzi (whose efforts at carving stone tools we described in
a previous chapter) understands more than 150 words of spoken English.
And Kanzi was not the only chimpanzee to show a degree of linguistic
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The Origins of Human Language
241
The Origins of Human Language
upper temporal gyrus and the parietal lobe (a little behind and above the
ear), is responsible for encoding and decoding messages. A lesion in
Wernicke’s area renders the individual incapable of correctly understand-
ing and producing language, whether spoken or written.
According to Philip Tobias, the lower region of the parietal lobe, the
part related to Wernicke’s area, is more developed in the Homo habilis
fossils from Olduvai than in australopithecines, Paranthropus, and apes.
Moreover, Broca’s area is markedly enlarged both in Homo habilis/Homo
rudolfensis and in Homo ergaster. This area is much more developed in the
first humans than in australopithecines and Paranthropus, in which it is
only sketchily present.
Thus, the regions of the cerebral cortex most directly related to the
production of human language were already well developed in the first
representatives of our genus. Does this mean that these humans already
possessed the capacity of speech? Although this is the conclusion of the
majority of specialists who have studied brain casts from primitive hom-
inids, there are those who oppose this view.
In his book The Wisdom of the Bones, Alan Walker, who led the study of
the Turkana Boy, rejects the possibility that this individual, and by exten-
sion all those of his species, was able to speak. Walker bases his conclusion
on the fact that within the thoracic vertebrae of Turkana Boy the vertebral
canal is very narrow. This configuration is common among apes, but not
in modern humans, who have a much wider vertebral canal. In view of the
reduced diameter of the vertebral canal, Walker argues that the spinal cord
of the fossil specimen contained fewer neurons than that of modern
humans, so Turkana Boy’s thoracic region would have been less innerv-
ated than our own. The only plausible explanation of this fact, according
to Walker, is that the thoracic musculature involved in breathing was not
capable of performing the precise inhalations and exhalations that human
speech requires. So how do we explain the high level of development of
Broca’s area reflected in the inner surface of the braincase of Turkana Boy?
On the basis of results obtained from modern techniques used to explore
brain activity (specifically positron emission tomography, or PET), which
show that the area of the cerebral cortex surrounding Broca’s area is also
related to manipulation of the right hand, Walker suggests that the
development of this area in the first humans was an adaptation related
not to speech, but to dressing stone.
To sum up, although different studies of the cerebral cortex of the
first hominids concur that the areas connected with language (particularly
Broca’s area) are more fully developed in the first humans than in austra-
lopithecines, Paranthropus, and apes, they do not agree on the
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243
The Origins of Human Language
But if our upper respiratory tract has lost efficacy in this respect (and also
for breathing and sense of smell), what is the compensation? The answer is
that our species has a pharynx larger than that of any other mammal, giving
us the capacity to modulate a wide range of different sounds.
Contrary to popular belief, the majority of the basic sounds which make
up human speech do not originate directly in the vocal cords. In the
production of vowel sounds (and also of voiced consonants, but for the
sake of simplicity we will refer only to vowels in what follows), the vocal
cords open and close rapidly with periodic breaths of air to produce a
‘‘base’’ sound, or laryngeal tone. This is always the same, regardless of the
vowel we are pronouncing.
The laryngeal tone is made up of one main frequency and a series of
‘‘accompanying’’ frequencies or harmonics. If the set of cavities above
the larynx (the pharynx, nasal cavity, and oral cavity) were not involved
Pharynx
Epiglottis
Nasal cavity
Oral cavity
Location of
vocal cords and
glottis
Alimentary canal
Hyoid bone
(esophagus)
Thyroid cartilage
Respiratory tract
(trachea)
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The Origins of Human Language
Pharynx
Epiglottis
Location of vocal
Hyoid bone cords and glottis
Thyroid cartilage
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The Origins of Human Language
246
The Origins of Human Language
of the tongue (Figure 16.1). The longer the tongue, the longer these
muscles are, and the slower and less precise their movements.
It is easy to estimate the length of the tongue from a fossil skull, since it
is proportional to the length of the hard palate. However, deducing the
position of the larynx is a different matter. As we have already noted, the
larynx is formed of cartilage and supported by muscles and ligaments,
none of which fossilize.
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The Origins of Human Language
In today’s world we are used to the idea that communication and infor-
mation are the key to progress, and the basis of our current technological
development. Thus it appears to us that possessing an articulated language
gives us an indisputable superiority over other living beings in the struggle
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The Origins of Human Language
Larynx
Piriform sinuses
To the lungs
To the stomach
Figure 16.2 Section of the head of a nursing infant. The larynx is connected
to the nasal cavity and liquid passes into the alimentary canal through the piriform
sinuses. It is thus possible to drink and breathe at the same time. In adult humans,
the low position the larynx means that it has no a direct connection to the nasal
cavity, and breathing therefore has to be interrupted during the ingestion of
liquids.
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The Origins of Human Language
for existence. But on reflection, it is easy to see that linguistic ability has no
advantage for a lone human confronting nature with only his own re-
sources: this characteristic only has meaning within a group to which the
individual necessarily belongs. Language is a property not of the individ-
ual, but of the collective. It is not the individual who communicates well;
it would be more appropriate to say that the group has good communi-
cation. The capacity to share and transmit information between individ-
uals and between generations by means of language confers a great
adaptive advantage on the group as a whole, but not on the isolated
individual.
This fact, which appears obvious to us, poses a serious problem for
evolutionary biology, because the theory of evolution through natural
selection – the Darwinian theory that is universally accepted, with minor
variations, by all scientists – is based on competition among individuals.
We have already noted that some authors have proposed a reductionist
version of natural selection, occurring at a level below that of the individ-
ual, such as the level of the genes. We have seen in this book that
competition can sometimes occur at the level of sperm – again, below
the level of the individual. Darwin himself put forward the mechanism of
natural selection in order to explain certain characteristics of individuals
which cannot be understood from the point of view of simple competition
for environmental resources and the struggle for survival; after all, what
matters is surviving in order to reproduce.
However, the appearance of properties which characterize groups, such
as communication, poses the problem of natural selection at the suprain-
dividual level, the level of the group. Thus, language was selected because
groups with a higher level of internal communication were more com-
petitive, more efficient in exploiting the resources of their environment,
and displaced other groups. In other words, an individual with the cap-
acity to produce and understand articulated language may be no more
competitive than another individual who does not have this ability, but a
group with language is more competitive than a group without.
It has been known since ancient times that competition between groups
of the same species occurs among social mammals, including primates.
Chimpanzees, to take a case in point, defend their territory and sometimes
invade that of other groups, or are invaded by neighboring groups. Not
only does communication serve to increase the cohesion and efficiency of
groups (‘‘information is power,’’ as we say in our society), but it appears
that behaviors of social cooperation (‘‘social altruism’’) within the group
are extremely important in selection between groups. Social cooperation
extends to many areas, such as defense of territory or resources, defense
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The Origins of Human Language
against predators, group hunting, caring for other adults’ offspring, shar-
ing food, and so on.
However, while competition between groups is easy to understand,
formulating a realistic model of group selection is a more complex matter.
If we start from the premise that cooperative behavior and the mental and
phonetic capacity for language have a genetic basis, we can approach the
problem by considering the existence of a gene for cooperative behavior
and a gene for language (this is, of course, an oversimplification, which we
hope the discriminating reader will forgive).
In order for selection between groups to occur, there must be a high level
of genetic homogeneity within each group – in other words, a high level of
consanguinity. To put it in more technical terms, the variability within each
group must be much lower than the variability between the different
groups. Groups in which the language and cooperation genes occur fre-
quently will be more competitive than others; since cooperation and com-
munication are not governed by the law of all or nothing, it would in fact be
the greater or lesser capacity for language and cooperation that made the
difference. In the case of social cooperation, there is the additional problem
that ‘‘selfish’’ individuals profit from the efforts of others without expend-
ing their own energies, and could thus benefit from natural selection and
leave more genes for the succeeding generation, with the result that ‘‘al-
truistic’’ behavior would never become dominant. One solution is that
there might be mechanisms of social rejection which prevent ‘‘selfish’’
individuals from reproducing, but then we are faced with the question of
how these mechanisms were selected, and we are back to square one.
Chimpanzees form social groups in which the males are related but not
the females, who move from one group to another. As we have seen, we
can picture something similar happening among the first hominids. Thus,
the social group is not a closed reproductive group, and hence genetic
uniformity is lost when females (genes, in short) are imported from other
groups. Even so, we can formulate mathematical models which make
selection at group level viable under certain conditions. But it is also
possible to imagine that our ancestors came to form larger social units,
which functioned as reproductive units and also competed with one
another. Whether or not this is the case, at present we have no better
mechanism than group selection to explain one of our most important
characteristics: articulated language.
Another question for which selection at group level appears the only
practical explanation is the replacement of the Neanderthals by modern
humans. It does not seem reasonable to suggest that this substitution took
place through selection at individual level. As we have already noted, the
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The Origins of Human Language
Neanderthals were physically stronger and their anatomy was better adapted
to the European climate (see Figure 13.1), and we therefore have to believe
that individually they were as well suited as modern humans to their envir-
onment (if not more so). It seems virtually certain that the success of our
ancestors was based on some group characteristic, but which one?
Many authors believe that the absence of a true language among
Neanderthals was one of the reasons, if not the main reason, for their
replacement by modern humans. To put it simply, the rudimentary lan-
guage of the Neanderthals limited their social complexity, restricting their
capacity to transmit information essential for exploiting the resources of
their environment. And when modern humans appeared on the horizon,
well equipped with their sophisticated language, the Neanderthals were
doomed to extinction.
Although the hypothesis of the linguistic superiority of modern humans
is an attractive one, we have already seen that it does not fit with the
fossils. Apart from the fact that it is not clear what kind of language
Neanderthals had, we have already noted that they were not ‘‘swept
away’’ by modern humans in a rapid, universal process: the replacement
of one by the other took place over about 10,000 years. If modern
humans had such an overwhelming advantage in terms of social complex-
ity and exploitation of the environment, why did it take them so long to
replace the Neanderthals?
We believe that what enabled our ancestors to displace the Neander-
thals was not the possession of a qualitative advantage, the type of lan-
guage, but rather the higher level of development of their capacities to
exploit resources: to put it simply, they had more of the same.
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The Origins of Human Language
254
17
Until tonight, you thought that life was absurd. From now on you will
know that it is mysterious.
Eric-Emmanuel Schmitt, The Visitor
In his book Wonderful Life, Stephen Jay Gould explains that we are not
the inevitable result of evolution, but rather a mere circumstance of it –
that if the videotape of life was rewound and played again from the
beginning, planet Earth would now be populated by a completely differ-
ent range of life forms, among which humans would be nowhere to be
found.
It is, of course, impossible to perform this experiment by returning to
the beginning of life. However, to a certain extent it has been conducted
by natural means. For example, the platyrrhine monkeys in America did
not evolve into forms of intelligence comparable to our own. Clearly, they
were not subject to any ‘‘impulse’’ driving them toward ‘‘progress’’ or
‘‘perfection’’ (the same could be said of marsupials in Australia, and other
similar cases of evolution in conditions of geographical isolation).
In any case, there is another way we can play Gould’s game. If we
assume that evolution is directed or tends spontaneously toward increas-
ingly ‘‘higher’’ or more complex forms of life, we would expect the fossil
record to reflect a history in which progressively more complex forms of
life replace others, owing to their clear superiority, until humans appear.
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The Meaning of Evolution
But this analysis runs up against the problem of how to measure com-
plexity. We need to be able to establish a scale which could be applied to
living species or fossils, so that we can say: this species is at complexity level
3 or level 7 (the reader perhaps imagines that our species would be the only
one at level 10). We have to admit that this is a difficult problem to solve.
In 1949 George Gaylord Simpson, one of the greatest paleontologists
of the 20th century, published a very influential book entitled The Mean-
ing of Evolution, in which he concluded that evolution had no purpose.
Among other themes, in a chapter headed ‘‘Progress in Evolution’’
Simpson analyzed the question of whether there has been an increase in
complexity over the course of the Story of Life. It was clear to him that
when multicellular organisms (those with many cells) appeared, they
represented an increase in complexity over the first, unicellular (single-
celled) forms of life. According to Simpson, a second step toward greater
complexity occurred when the major different types of multicellular or-
ganisms emerged (the technical term for these types is phylum – plural
phyla); however, this progress took many directions, rather than following
a single, privileged line. From this point on, it is impossible to compare
the level of complexity within each of the lines. Simpson, who specialized
in vertebrate paleontology, wrote that it would be a brave man who would
attempt to prove that a human being is more complex than an ostraco-
derm (a kind of fishlike aquatic vertebrate which appeared more than
4 million years ago).
Meanwhile, we still have not defined what complexity is – no easy task
(you might also like to try). One modern way of doing it would be to use
the concept of complexity as it is applied to systems. A system is a com-
bination of elements which interact with one another, giving rise to the
properties of the system, and the more separate elements it has, the more
different possibilities for interaction exist, rendering the system richer in
functions, or more complex – in the sense of less predictable, less rigid,
more variable, and also more adaptable.
Multicellular organisms are self-regulated systems composed of differen-
tiated cells which form tissues and organs; these in their turn are organ-
ized into systems – respiratory, digestive, reproductive, excretory,
circulatory, or nervous. The concept of the complexity of systems can be
used to compare organisms in very different groups, for example to
compare mammals with sponges or jellyfish: the latter are clearly much
simpler forms of organization than mammals, with fewer differentiated
elements, and we can consider them biological systems with a relatively
low level of complexity. Who, however, would venture to assess the
relative complexity of a bat and a lion?
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The Meaning of Evolution
257
The Meaning of Evolution
200 million years ago, Therapsida began to decline, and were gradually
replaced in terrestrial ecosystems by dinosaurs (birds later evolved from one
particular group of dinosaurs, making them the true ‘‘living dinosaurs’’).
If the term ‘‘superior’’ can be applied in evolutionary biology to any
particular group, in this case it would have to be applied to the dinosaurs.
The Therapsida eventually became extinct, and although some evolved into
the first mammals, these latter not only failed to win out over reptiles, but
led a very discreet life throughout the remainder of the Mesozoic (without
exception, all mammals were small in size). In fact it was the impact of a
meteorite, not the superiority of mammals, which finished off the dinosaurs
65 million years ago. If it had not been for this ‘‘providential’’ meteorite,
the evolution of terrestrial vertebrates would no doubt have been very
different. (Some authors believe that the extinction of the dinosaurs may
have been due to the atmospheric effects of a series of great volcanic
eruptions; the important thing for our argument is that the disappearance
of the dinosaurs had a nonbiological cause, and it is irrelevant whether this
was a meteorite, a volcanic phenomenon, or any other geological disaster.)
Let us now turn to another case from the pages of the fossil record, this
time even closer to ourselves. As we have noted in this book, within the
order of primates we belong to a group, the hominoids, which includes a
series of species, apes, with which we share many traits and, in truth, many
genes. In fact, apes are also the primates with the most developed brains.
Since they are the mammals most resembling us, it might be expected that
their superiority led them to win out at least over the other monkeys
existing at the time when they appeared. As we have noted, hominoids
originated in Africa at least 23 million years ago and, from the moment
when these primates made their way out of Africa to populate Europe and
Asia (about 17 million years ago), they became the group of primates with
the greatest evolutionary success, diversifying into a large number of
species inhabiting the wide belt of forest which extended throughout
much of the Old World. This might seem logical, since these were the
most intelligent primates and their success prefigured the glorious future
awaiting human beings.
Once more the fossil record tells us the opposite of what appears
‘‘logical.’’ Between about 8 and 7 million years ago, hominoids ceased
being the ‘‘kings of creation.’’ A great ecological change caused their
paradise to disappear. Astronomical factors, movements of continental
landmasses, and the emergence of mountain ranges changed the climate
and the composition of the atmosphere, resulting in deterioration of their
habitat over much of its once enormous expanse. The forest gave way to
more open ecosystems.
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The Meaning of Evolution
But it was not only that the change in vegetation reduced their living
space; other primates, the Cercopithecidae, became more numerous and
more varied than them. In military terms, the apes beat a retreat. Today
they remain reduced to two species of chimpanzee and the gorilla in
Africa, the orangutan in Sumatra and Borneo, and the gibbons of contin-
ental and island Southeast Asia. It is indeed significant that the little
gibbon, the hominoid least like humans, is the most successful in terms
of diversity and numbers. Despite pressure from humans, several million
gibbons, of nine different species, still survive.
But about 5 or 6 million years ago, in some place in Africa, possibly the
easternmost part, a specific type of hominoid began to emerge: the first
hominid, our oldest ancestor. To begin with it differed very little from the
ancestors of modern chimpanzees and gorillas. It could be considered the
East African version of the same group. Later, at least 4 million years ago,
this type of hominoid had developed a singular characteristic, never before
seen. This was not a hominoid more intelligent than the others. No – it
was a bipedal hominoid.
As time went on, bipedal hominoids adapted to the increasingly arid
ecosystems covering much of Africa. They developed specialized dentition
to cope with this. We have already seen that it is difficult to measure the
intelligence of fossil species (even of living species), but the index of
encephalization suggests that the most encephalized species 3 million
years ago were not hominids but dolphins.
By about 2.5 million years ago the hominids had split into two different
evolutionary lines. One of these, Paranthropus, developed a hypertrophied
(overdeveloped) masticatory apparatus. The other line was that of the first
representatives of the genus Homo, the first humans, who had a somewhat
larger brain. Only from this moment did hominids become unique among
living beings for their greater cerebral complexity. Paranthropus later became
extinct, and subsequent humans modified their body structure, increased
their brain size, and improved their technology. But even since the point
when intelligence appeared in the biosphere, human evolution has not
followed one single path, a straight line leading to ourselves. On the contrary,
until a few thousand years ago a number of intelligent human species existed
on earth. The fact that ours is the only one now in existence gives us the false
perspective that it has always been so, that our ancestors succeeded one
another in an ordered sequence, on a staircase we climbed step by step.
In short, neither the evolutionary history of mammals nor that of hom-
inoids reflects a pattern whereby these groups appeared and progressively
came to dominate other creatures thanks to their superior characteristics,
particularly their intelligence. On the contrary, the fossil record in both
259
The Meaning of Evolution
260
The Meaning of Evolution
Today we fear that the agricultural and industrial activities of the human
race could destroy the ecological balance, causing the widespread
261
The Meaning of Evolution
extinction of species, however well adapted they may be. The introduction
of animals, plants, or other organisms from one region into another (for
example, bringing rabbits to Australia) can also have catastrophic effects
on the ecological balance. We are likewise concerned about the effects on
the biosphere of climate change caused by the emission of greenhouse
gases such as carbon dioxide, and by the destruction of the ozone layer.
We are all increasingly aware that ecosystems, which are always in fragile
equilibrium, are composed of many species with a long history.
But all of these things about which we are (with reason) concerned have
occurred many times in the past, through natural causes. The continents
and oceans were not always distributed over the earth’s surface as they are
now, and the different regions where life is found have separated and come
together in many different ways, giving rise to various exchanges between
species in which some were favored while others lost out. Climate change
is a frequent occurrence in the Story of Life, and has a powerful impact on
ecosystems (the alternation between ice ages and interglacial periods such
as the one in which we are now living – which will not last forever – is but
one example). The composition of the atmosphere has also varied.
Finally, even if the physical environment remained unchanged, the
appearance, through evolution, of new species introduces a fundamental
instability factor into ecosystems, and means that they are always chan-
ging. Organisms show adaptations, developed over the period of their
evolution, which only make sense in relation to the ecological niches
occupied by their species. In a community, all populations are related to
one another in intricate networks, through which energy and matter flow.
To take the example of a primate, any change in the plants on which it
feeds or the animals it eats, the predators which prey on it, its competitors,
or its parasites, will have unpredictable consequences for the survival of
the species, which will have to evolve in its turn, adapting to its new
circumstances.
In other words, the biosphere is an enormously complex macrosystem,
made up of many different elements which organize themselves according
to a hierarchy of levels (cell, tissue, organ, system, individual, family
group, social group, population, community, ecosystem), and interact in
many different ways at all levels. This kind of system makes it extremely
difficult to identify the basic laws according to which it functions, even
when the constituent elements remain the same. However, to complicate
things further, the evolution of species means that the biosphere is an
unstable system, far from equilibrium, which has never remained static
and whose composition (species, and therefore their various interactions)
has changed over time.
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The Meaning of Evolution
It follows from all of this that the evolution of species is subject to the
influence of so many factors that in practice it is impossible to predict its
future. This does not mean that evolution depends on pure chance (in the
popular sense of incomprehensible chaos). On the contrary, it can be
understood, even if only with hindsight, like the weather. To a certain
extent, the well-known example of chaos theory called the ‘‘butterfly
effect’’ can be applied here: the flutter of a butterfly’s wings in Beijing
can cause rain to fall in New York (to say nothing of the case where the
delicate movement of an insect’s wings is replaced by a meteorite many
miles in diameter traveling at full speed toward earth).
But is this a merely technical question? Is our uncertainty due only to
the complexity of the problem? Chaos theory goes further, asserting that
even if we knew all these factors and interactions in full detail, the future
cannot be known, simply because it is not predetermined. This is the end
of certainty, which is replaced by probabilities.
In his book On Aggression, Konrad Lorenz recounts how Alfred Kühn
ended a lecture with Goethe’s words: ‘‘It is the greatest joy of the man of
thought to have explored the explorable and then calmly to revere the
unexplorable.’’ As the audience broke into applause, Kühn raised his voice
and cried: ‘‘No, not calmly, gentlemen; never calmly !’’ Should we assume
that at this point investigation into the nature of evolution is closed? Has
everything already been said?
We believe, on the contrary, that there is still much work to be done.
Newtonian physics speaks of trajectories which can be expressed in terms
of equations. If we know the initial conditions, these trajectories are
predictable and reversible, like a pendulum – first here, then there. In
these equations time does not exist: it is merely an illusion in which the
future and the past meet. Quantum physics simply replaces trajectories
with wave functions, but the symmetry with regard to time does not
change. Biological evolution, in contrast, is an irreversible process which
unfolds over time, surprising us at each moment, and follows no trajectory
(or trend). How can we reconcile physics and biology? If chaos theory is
correct, there is, as Ilya Prigogine (winner of the 1977 Nobel Prize for
Chemistry) says in his book The End of Certainty, a narrow path between
two equally alienating conceptions of the world. Either we live in a
deterministic world governed by immutable laws which leave no place
for novelty (and where the greatest novelty of all, evolution, would be
impossible), or we are in ‘‘a world . . . where everything is absurd, acausal
and incomprehensible,’’ subject to pure chance. It is the task of the
Darwins of the present and the future to travel this narrow path.
263
Epilogue
Living organisms had existed on earth, without ever knowing why, for
over three thousand million years before the truth finally dawned on
one of them. His name was Charles Darwin.
Richard Dawkins, The Selfish Gene
In his Dictionary of the Cinema, Fernando Trueba tells how the famous
French film director François Truffaut declared that when he went to the
cinema and saw a group of characters digging a tunnel for an hour and a
half and at the end of the film the tunnel was no use, he believed he should
get his money back. Although we cannot give our readers their money
back, we would not like to leave them with the unpleasant impression that
more than 3,000 million years of evolution have been completely point-
less, and that we are just a species like any other. Because in fact this is not
true.
In his well-known book The Meaning of Evolution, which we referred to
in a previous chapter, George Gaylord Simpson explains that evolution has
no aim, but then rejects the suggestion that this means that man is ‘‘just
an animal.’’ To begin with, Simpson does not see any animal species as
‘‘just an animal,’’ because each one has its own peculiarities, shared with
no other species. But ours is exceptional in many very important aspects.
We are the most intelligent species, and this intelligence (however de-
fined) has enabled us to occupy all the different landscapes of the planet
264
Epilogue
265
Epilogue
266
Epilogue
Only since Darwin has it been understood that we are not the chosen
species but, as Robert Foley suggests, a unique species among many other
unique species – albeit an amazingly intelligent one.
And it remains a paradox that so many centuries of science have led us
to know what any Kalahari Bushman, any Australian Aborigine, or any of
our ancestors who painted bison in the caves of Altamira knew full well:
that it is not the earth that belongs to man, but man who belongs to
the earth.
267
Bibliography
In addition to the books cited in the text, we felt it would be useful to provide a
brief list of some of the most important recent publications which might be of
interest to the reader who wants to study human evolution in more depth.
Inevitably some important works will have been omitted, although we have
attempted to ensure that, though not all the references available are listed here,
all those that are listed are available.
Books
268
Bibliography
Conroy, G.
1990 Primate Evolution. New York: W. W. Norton.
Day, M.
1986 Guide to Fossil Man. London: Cassell.
Fleagle, J.
1988 Primate Adaptation and Evolution. San Diego: Academic Press.
Foley, Robert
1987 Another Unique Species. Harlow: Longman.
Fossey, Dian
1983 Gorillas in the Mist. Boston: Houghton Mifflin.
Goodall, Jane
1971 In the Shadow of Man. Boston: Houghton Mifflin.
1990 Through a Window. Boston: Houghton Mifflin.
Johansen, D., and M. Edey
1981 Lucy: The Beginnings of Humankind. New York: Simon and Schuster.
Johansen, D., and B. Edgar
1996 From Lucy to Language. New York: Simon and Schuster.
Jones, S., R. Martin, and D. Pilbeam, eds.
1992 The Cambridge Encyclopedia of Human Evolution. Cambridge:
Cambridge University Press.
Klein, R.
1989 The Human Career. Chicago: University of Chicago Press.
Landau, M.
1991 Narratives of Human Evolution. New Haven: Yale University Press.
Leakey, Richard
1981 The Making of Mankind. New York: E. P. Dutton.
Leakey, Richard, and R. Lewin
1992 Origins Reconsidered. New York: Doubleday.
Le Gros Clark, Wilfrid
1969 The Antecedents of Man. Chicago: Quadrangle.
Lewin, R.
1987 Bones of Contention: Controversies in the Search for Human Origins.
New York: Simon and Schuster.
Martin, R.
1990 Primate Origins and Evolution. London: Chapman and Hall.
Napier, J., and P. Napier
1985 The Natural History of the Primates. London: British Museum (Natural
History).
Reader, J.
1981 Missing Links. Harmondsworth: Penguin.
Rightmire, G.
1990 The Evolution of Homo erectus. Cambridge: Cambridge University Press.
269
Bibliography
Articles
270
Bibliography
271
Bibliography
Falk, D.
1993 Evolution of the Brain and Cognition in Hominids. James Arthur Lec-
tures on the Evolution of the Human Brain. New York: American Museum of
Natural History.
Fernández-Jalvo, Y., et al.
1996 Evidence of Early Cannibalism. Science 271:269–270.
Foley, R., and M. Lahr
1998 Mode 3 Technologies and the Evolution of Modern Humans. Cambridge
Archaeological Journal 7:3–36.
Gagneux, P., D. Woodruff, and C. Boesch
1997 Furtive Mating in Female Chimpanzees. Nature 387:358–359.
Hammer, M., and S. Zegura
1996 The Role of the Y chromosome in Human Evolutionary Studies. Evolu-
tionary Anthropology 5:116–134.
Harcourt, A.
1994 Sexual Selection and Sperm Competition in Primates: What are Male
Genitalia Good For? Evolutionary Anthropology 4:121–129.
Hublin, J. J., et al.
1995 The Mousterian Site of Zafarraya (Andalucia, Spain): Dating and Impli-
cations on the Palaeolithic Peopling Processes of Western Europe. Comptes
Rendus de l’Académie des Sciences de Paris, série II 321(10):931–937.
1996 A Late Neanderthal Associated with Upper Palaeolithic Artefacts. Nature
381:224–226.
Johanson, D., et al.
1986 New partial skeleton of Homo habilis from Olduvai Gorge, Tanzania.
Nature 327:205–209.
Kappelman, J.
1997 They Might be Giants. Nature 387:126–127.
Kimbel, W., D. Johanson, and Y. Rak
1996 Systematic Assessment of a Maxilla of Homo from Hadar, Ethiopia.
American Journal of Physical Anthropology 103:235–262.
Klein, J., N. Takahata, and F. J. Ayala
1993 Mhc Polymorphism and Human Origins. Scientific American 269:78–83.
Krings, M., et al.
1997 Neanderthal DNA Sequences and the Origin of Modern Humans. Cell
90:1–20.
Leakey, M. G., et al.
1995 New Four-million-year-old Hominid Species from Kanapoi and Allia Bay,
Kenya. Nature 376:565–571.
Lieberman, P., et al.
1992 The Anatomy, Physiology, Acoustics and Perception of Speech: Essential
Elements in the Analysis of the Evolution of Human Speech. Journal of Human
Evolution 23:447–467.
272
Bibliography
Lovejoy, C.
1981 The Origin of Man. Science 211:341–350.
Manzi, G., A. Vienna, and G. Hauser
1996 Developmental Stress and Cranial Hypostasis by Epigenetic Trait Occur-
rence and Distribution: An Exploratory Study on the Italian Neanderthals.
Journal of Human Evolution 3:511–527.
McHenry, H.
1992 How Big were Early Hominids? Evolutionary Anthropology 1:15–20.
Moyà-Solà, S., and M. Köhler
1997 A Dryopithecus Skeleton and the Origins of Great-ape Locomotion.
Nature 379:156–159.
Parés, J. M., and A. Pérez-González
1995 Paleomagnetic Age for Hominid Fossils at Atapuerca Archaeological Site,
Spain. Science 269:830–832.
Plavcan, J., and C. Van Schaik
1996 Intrasexual Competition and Body Weight Dimorphism in Anthropoid
Primates. American Journal of Physical Anthropology 103:37–68.
Rosenberg, K., and Trevathan, W.
1997 Bipedalism and Human Birth: The Obstetrical Dilemma Revisited. Evo-
lutionary Anthropology 4:161–168.
Ruff, C., E. Trinkaus, and T. Holliday
1997 Body Mass and Encephalization in Pleistocene Homo. Nature 387:
173–176.
Schrenk, F., et al.
1993 Oldest Homo and Pliocene Biogeography of the Malawi Rift. Nature
365:833–836.
Semendeferi, K., et al.
1998 The Evolution of the Frontal Lobes: A Volumetric Analysis Based on
Three-dimensional Reconstructions of Magnetic Resonance Scans of Human
and Ape Brains. Journal of Human Evolution 32:375–388.
Seyfarth, R., and D. Cheney
1992 Meaning and Mind in Monkeys. Scientific American 267:122–129.
Suwa, G., et al.
1993 The First Skull of Australopithecus boisei. Nature 389:489–492.
Swisher III, C., et al.
1994 Age of the Earliest Known Hominids in Java, Indonesia. Science
263:1118–1121.
1997 Latest Homo erectus of Java: Potential Contemporaneity with Homo
sapiens in Southeast Asia. Science 274:1870–1874.
Szathmáry, E., and J. Smith
1995 The Major Evolutionary Transitions. Nature 374:227–232.
Tague, R., and C. Lovejoy
1986 The Obstetric Pelvis of A.L. 288–1 (Lucy). Journal of Human Evolution
15:237–255.
273
Bibliography
Vrba, E.
1987 Late Pliocene Climatic Events and Hominid Evolution. In Evolutionary
History of the ‘‘Robust’’ Australopithecines. F. Grine, ed. New York: Aldine de
Gruyter.
Wheeler, P.
1993 Human Ancestors Walked Tall, Stayed Cool. Natural History 102:65–67.
White, T., G. Suwa, and B. Asfaw
1993 Australopithecus ramidus, a New Species of Early Hominid from Aramis,
Ethiopia. Nature 371:306–312.
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