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Human Evolution

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398 Human Cognition
Petitto LA, Zatorre RJ, Gauna K, Nikelski EJ, Dostie D
and Evans AC (2000) Speech-like cerebral activity in
profoundly deaf people processing signed language.
Proceedings of the National Academy of Sciences of the USA
97: 13961±13966.
Plotkin H (2002) The Imagined World Made Real: Towards a
Natural Science of Culture, London, UK: Allen Lane.
Human Evolution
Marta MirazoÂn Lahr, University of Cambridge, Cambridge, UK
CONTENTS
Introduction
Hominin origins and early hominin diversity
Human evolution in the Pleistocene
The origin of human species and their relationships
with the apes have been reassessed in the light of
genetic studies. Hominins are now believed to have
emerged in Africa about 7±5 million years ago.
INTRODUCTION
Human evolution is more than the evolution of
humans. More than 15 species have existed since
the time of our last common ancestor with the
apes. The features that make humans different
from apes did not evolve at once, or in the same
ancestor, and no single trait made the difference.
Humans are unique among the apes in the way
they move, their global distribution, the size of
their brains, their dependency on technology,
their linguistic abilities, their slow growth, and
their demography. These distinguishing features
evolved at different times, each giving an advan-
tage to some individuals at the time they were
competing with those lacking them, only to then
become part of the heritage of subsequent gener-
ations. Progress in understanding human evolution
has come through the amazing discoveries of new
fossils and archeological sites, the development of
sound chronological and paleoecological frame-
works, and the application of new techniques
such as molecular genetics to the subject. Paleoan-
thropology is an interdisciplinary subject by its
very definition.
Stuss DT, Gallup GG and Alexander MP (2001) The
frontal lobes are necessary for theory of mind. Brain 124:
279±286.
Tomasello M (1999) The Cultural Origins of Human
Cognition. Camridge, MA: Harvard University Press.
Introductory article
Human dispersals
Human technologies
Hominin evolution: the evidence
HOMININ ORIGINS AND EARLY
HOMININ DIVERSITY
When and Where Hominins Evolved
Fossil and genetic evidence indicates that hominins
emerged in Africa between 7 million and 5 million
years ago (Mya) in the late Miocene. The earliest
fossils that have been clearly identified as hominin
date to 4.4 Mya. Earlier, seemingly hominin remains
have been found dating from 6.0±5.8 Mya, although
their relationship to the hominin or chimpanzee
lines remains controversial given the lack of fossil
remains of the apes that would have given rise to
them. Between the earliest hominin and fossil apes
dated to about 9.0 Mya, the African hominoid fossil
record does not exist. However, genetic evidence
allows us to establish who, among living apes, are
our closest relatives, as well as to confirm the dates
established from fossils. In fact, these very early
hominin fossils have been discovered only in the
last few years, and it has been on the basis of
genetic evidence that the age of divergence be-
tween apes and hominins has been established.
The genetic comparison of apes and humans,
with a view towards establishing their pattern of
relationships, was pioneered by Sarich and Wilson
in the late 1970s. These comparisons, now based on
the study of a large number of different genes and
gene products, established three key facts: first, that

the African apes form a clade within the great apes;
second, that within that clade, chimpanzees and
humans are more closely related to each other
than either is to the gorillas; and third, that on the
basis of a molecular clock, hominins and chimpan-
zees diverged from each other between 7 Mya and
5 Mya. These results have had a major impact on
hominoid systematics (Figure 1), as well as estab-
lishing a chronological and ancestral framework
for the study of hominin origins.
However, controversy exists as to who were the
ancestors of the living African apes and where they
lived. The fossil record of Miocene apes is a rela-
tively rich one, and yet two very contrasting hy-
potheses exist regarding the biogeography of
hominin origins. The Miocene is a long period
(25±5 Mya), which can be divided into two phases.
The first of these is characterized by climatic
warming and expansion of African forests, in
which early apes diversified. The second phase is
characterized by global cooling, contraction of
Hominidae
Hylobatidae
Hylobatidae
Hominidae
Pongidae
Pongidae
G O Go Ch H G O Go Ch
(a) Early classification and (b) Accepted correct classification
inferred phylogenetic and phylogenetic relationships
relationships of living of living hominoids
hominoids
Figure 1. Hominoid systematics. Traditionally the
superfamily Hominoidea has been seen as consisting of
three families, the Hylobatidae, the Pongidae, and the
Hominidae. These groupings emphasized the similarities
of the great apes and the distinctiveness of humans.
Molecular approaches have radically changed this, and
shown a close relationship between humans and African
apes, especially the chimpanzees. As one of the aims of
taxonomy is to reflect phylogeny, the traditional classifi-
cation has changed. Orang-utans are the only living rep-
resentatives of the family Pongidae, while gorillas,
chimpanzees and humans form the family Hominidae.
This reclassification also changes the vernacular of these
scientific names, so that instead of using the term `hom-
inid' to describe humans and their ancestors, the term
`hominin' (reflecting the subfamily status Homininae) is
used. Ch, chimpanzees, genus Pan; H, humans, genus
Homo; G, gibbons, genus Hylobates; Go, gorillas, genus
Gorilla; O, orang-utans, genus Pongo.
Human Evolution 399
forests, lowering of sea levels, and major geotec-
tonic changes. During this period the continents
and oceans acquired the general shape by which
we know them, the Rift Valley and the Himalayas
were formed, the Antarctic partially froze, and the
Arabian peninsula emerged forming a permanent
land bridge between Asia and Africa. These two
continents exchanged fauna through this land
bridge, and some apes dispersed to Eurasia for
the first time. Among those Eurasian apes we find
fossils that represent forms ancestral to orang-
utans.
For many years the ancestors of African apes
were sought among those apes who in the middle
Miocene remained in Africa. In this context, the
origin of hominins has been interpreted as part of
the process by which East and West African faunas
differentiated as the Rift Valley was formed. Homi-
nins would represent an African ape who adapted
to the drier and more open environments of East
Africa from an arboreal forest ancestor. However,
the genetic evidence implies that Asian and African
great apes share a common ancestor who lived after
apes had dispersed into Eurasia, suggesting that
the ancestors of the African apes (and hominins)
should be found among late Miocene Eurasian
forms. In this context, the differentiation of African
apes, including the origin of hominins, would
H result from the geographical separation of popula-
tions of an ancestral species as it dispersed into
Africa from the north-east. Therefore, although we
know hominins evolved in Africa from a common
ancestor with chimpanzees around 6±5 Mya, fas-
cinating issues relating to the origin of that
common ancestor will be resolved only by new
fossil finds from the crucial period between 8 Mya
and 5 Mya in Africa and possibly Arabia.
What Made Early Hominins Different?
Early anthropologists had two main expectations in
relation to the origins of humans and their ances-
tors: that they had diverged from apes a very long
time ago, and that this divergence was based upon
the development of a large brain. Both were proved
wrong. Hominins have a relatively short history,
and we now know that brain expansion began in
the last 2 million years. What set hominins apart
from other apes was their locomotion.
Bipedalism has long been recognized as the key
adaptation of the hominin lineage. In terms of
morphology, it requires the modification of most
of the skeletal system, although such modifications
were not all in place at once. The selective advan-
tages of bipedalism are much debated. As a means
400 Human Evolution
of terrestrial locomotion, it clearly provides an effi-
cient way of moving in open environments for long
periods, especially when compared with knuckle-
walking animals, such as gorillas and chimpanzees,
rather than true quadrupeds. It also results in more
efficient thermoregulation during the middle of the
day, as less body surface is exposed to the sun in
comparison with quadrupeds. Other advantages
may relate to predator vigilance and potential for
carrying. Most researchers would argue that a com-
bination of these factors gave bipedal apes a com-
petitive advantage in relation to their quadrupedal
relatives at the time.
However, when considering the selective pres-
sures that led to the evolution of bipedal apes, two
factors should be taken into account. First, early
hominin bipedalism was significantly different
from ours. The fragmentary fossil record of the
earliest hominins shows clear evidence that these
animals had already changed key aspects of their
anatomy towards a bipedal stance. However, the
earliest associated fossil skeleton ± that of the
famous Ethiopian Australopithecus afarensis fossil
nicknamed `Lucy' ± shows that several of the
changes that make bipedalism more efficient, such
as the shape of the ribcage, the curvature of fingers
and toes, and the elongation of the legs, had yet
to take place. Therefore, although it sets early
hominins apart from other apes, bipedalis itself
took millions of years to evolve. The second factor
is related to the first. It has become clear that early
hominins still spent significant amounts of time
in trees. Models of energy expenditure stress that
the advantages of bipedalism would become
meaningful only when hominin ancestors had to
spend a large percentage of their time on the
ground.
The Origin of the Lineage That Gave
Rise to Modern Humans
Early hominins were clearly successful in the niche
they colonized ± that of a terrestrial bipedal ape,
with a body and brain size similar to present-day
chimpanzees and a broadly similar diet based on
fruits, young leaves, and probably some meat. This
success led to demographic growth and a greater
distribution in East and South Africa, and eventu-
ally, competition among themselves for the re-
sources offered in this new niche. Competition
can lead to progressive specialization, in which
animals become better and better at processing
the resources on which they depend, and to differ-
entiation, in which animals try to exploit new re-
sources to avoid their competitors. Hominin
evolution in the Pliocene (the period between
5 Mya and 2 Mya) is characterized by both.
Among these bipedal apes, two adaptive trends
became established. One of these was towards
comparatively specialized animals, who adapted
their teeth, masticatory muscles and the shape of
the face to maximize their ability to crush large
quantities of rough food. They are known as the
robust australopithecines, and they consist of
several very successful species, usually grouped
under the genus Paranthropus. This group of homi-
nins was well established before 2 Mya, and sur-
vived until approximately 1 Mya in Africa. The
other adaptive trend was towards animals who
became encephalized, had larger body sizes, slower
growth, and appeared behaviorally more flexible,
as suggested by their association with stone tools,
their wider distribution, and the apparent con-
sumption of larger quantities of meat. In this suite
of characteristics, researchers recognize the appear-
ance of many features that we identify with
humans, and indeed, this group of animals repre-
sent the origins of the genus Homo to which we
belong.
Establishing the precise point of beginning of
new adaptive trajectories in evolution is always
difficult, and accordingly much controversy exists
as to which fossils mark the threshold between
Australopithecus and Homo. The hominin fossil
record in East Africa around 2 Mya shows, besides
the big-toothed Paranthropus, a very variable collec-
tion of animals, grouped under the species name
habilis, with some but not all of the Homo character-
istics listed above. Nevertheless, by about 1.8 Mya,
a new group of hominin can be identified ± Homo
erectus/ergaster, who shows the complete combin-
ation of incipient Homo traits, including the ability
to disperse widely. Homo erectus/ergaster is the first
hominin to leave Africa, and fossils of this group
from 2±1.5 Mya are found not only throughout East
and South Africa, but also in North Africa, the
Caucasus, and Java.
How can we interpret the combination of fea-
tures that differentiates the genus Homo from
earlier hominins and represents our own distinct
evolutionary heritage? Some of these traits can be
seen as fundamental biological changes in behav-
ior, anatomy and growth, while others represent
the consequences of these changes given energetic
and morphological constraints. Among those fun-
damental changes it is impossible to determine
`which came first', as their evolutionary success
depended on their concerted modification. How-
ever, it could be argued that new adaptive trajec-
tories become established when new behaviors

resulting in greater reproductive success lead to
selection of individuals whose genetic make-up
facilitates those behaviors. In this context, the fun-
damental behavioral shift associated with early
Homo was probably the shift to a more carnivorous
diet in conjunction with the manufacture of tools
that allowed its better exploitation. A more carniv-
orous diet, rich in protein and fats, would have
both released the energetic constraints to increasing
the size of the body and the brain (the most ener-
getically expensive organ), and favored more intel-
ligent individuals with greater memory, and
capable of more complex cooperation and plan-
ning. Furthermore, greater carnivory is associated
in mammals with larger geographical ranges and
less dietary exclusivity, and thus the underlying
condition for dispersal. Encephalization, in turn,
would have posed new pressures upon these
animals, both in terms of the need for greater ma-
ternal energy intake during pregnancy and lacta-
tion (most of the period during which human
brains grow), and in terms of maintaining meta-
bolic rate relative to body size.
These animals seem to have pursued two separ-
ate strategies for meeting these pressures, strategies
that have a feedback effect. On the one hand, re-
searchers have argued that the increasing energetic
Sociality
Tools/technology
Memory
Language?
Problem-solving
Range expansion
Figure 2. The complex adaptive system that underlies the evolution of the genus Homo. BMR, basal metabolic rate.
Human Evolution 401
costs to mothers imposed by encephalized infants
were met by `budgeting' these costs over a longer
period, thus leading to slower growth rates. On the
other hand, research has shown that while humans
have brains significantly larger than expected for
their body size, their guts show the reverse condi-
tion. On this basis, researchers have argued that
there was a trade-off between the energy channeled
towards brains and guts so as to maintain stable
metabolic rates, and that this was possible through
the smaller absorptive gut surface necessary in car-
nivores in relation to herbivorous animals. These
correlated changes and their feedback effects
(Figure 2) represent the underlying adaptive trend
of the genus Homo.
HUMAN EVOLUTION IN THE
PLEISTOCENE
Paleoenvironmental Context for the
Evolution of Neanderthals and Humans
One of the distinguishing features of the genus
Homo is its wide geographical distribution. For
more than 3 million years, hominins existed and
diversified within sub-Saharan Africa. In the sub-
sequent period, the Pleistocene epoch (1.8±0.01
Longer ontogeny
BRAINS
= BMR
MEAT Guts
EATING
Average
body
size
Longer legs Tall individuals
402 Human Evolution
Mya), hominins colonized first the northern and
southern (temperate) extremes of Africa (by 1.8
Mya), south-western Asia (by 1.7 Mya), tropical
south-east Asia (by 1.8 Mya), then east Asia (by
1.0 Mya) and Europe (by 0.8 Mya). Human evolu-
tion in the Pleistocene is no longer an African affair,
and allopatry becomes a key issue in interpreting
the patterns of differentiation among species and
populations. Under a simple allopatric model, Ne-
anderthals and modern humans represent, respect-
ively, the contemporaneous European and African
descendants of a Pleistocene hominin species
(Figure 3).
However, in order to understand why Neander-
thals and modern humans evolved, and why they
had such different fates, it is necessary to under-
stand the overall conditions that established their
selective environments. Evolution is the result of
changes in the competitive environment of species
that lead to new patterns of mortality. These estab-
lish new parameters of individual fitness, which
underlie the process of natural selection and conse-
quently adaptation. The competitive environment
of any species has both biotic and abiotic compon-
ents, which determine the conditions in which the
evolutionary process takes place. Understanding
climatic change, with its potential for promoting
expansion or contraction of ecological niches, is
essential for interpreting the diversification of
species. The Pleistocene climate was marked by
alternation between relatively cold glacial periods,
and warmer interglacial ones. The duration and
amplitude of the glacial periods both appear to
have increased around 800 000 years ago. These
climatic changes had effects on faunal distributions
and survivorship, some of which are predictable
and some of which are not.
A glacial cycle lasts approximately 100 000 years,
although each cycle varies in extent and severity.
Most of the cycle is characterized by a long period
of low, fluctuating temperatures, during which
continental ice accumulates at high latitudes. This
accumulation of water in the form of ice sheets
leads to a drop in sea levels, with the consequent
exposure of continental shelves, and to increasing
aridity in the tropics. As temperatures increase, the
ice sheets which had taken tens of thousands of
years to build start to melt, and interglacial phases
begin. That excess of circulating water leads to a
short period of extreme precipitation in the tropics,
creating extended freshwater networks and high
lake-level stands. When this water eventually
drains to the seas, conditions similar to those of
the present day set in. The overall effect for homi-
nins, depending on where they were, was one of
expanding and contracting ranges, of alternating
rich and scarce resources, and of highly variable
thermal conditions.
By 1 Mya hominin populations were established
throughout Africa, and in tropical and temperate
Asia as far north as northern China. Soon after-
wards, hominins occupied Europe for the first
time. This broad spatial distribution, in the context
of small population sizes, sets the conditions for
regional differentiation among the various groups
of encephalized hominins. That differentiation be-
comes very apparent in the Middle Pleistocene
(0.8±1.25 Mya). The main distinction in both
morphology and archeology is between east and
west, along what is known as the Movius line. In
the east, hominins have not changed from their
Homo erectus ancestry. Both in Java and China,
fossils in the time-span between 1.0 Mya and
300 000 years ago are Homo erectus, associated
with relatively simple tools. Homo erectus continued
to inhabit south-east Asia until about 50 000 years
ago, suggesting long-term isolation from other
regions of the world. The situation in China be-
comes increasingly more complex after 300 000
years ago, when fossils displaying a quite distinct
morphology make their appearance alongside
Homo erectus ones, and their affinities are still a
matter of debate. In the west, hominins change,
losing many of the erectus features, as well as show-
ing an overall increase in body size and brain size.
This morphology was for a long time referred to as
`archaic' Homo sapiens. Most researchers today at-
tribute it to a new species ± Homo heidelbergensis.
The main specimens of H. heidelbergensis are the
fossils from Bodo, Lake Ndutu, Kabwe, and
Elandsfontein in Africa, and from Arago, Box-
grove, and Petralona (among many others) in
Europe.
What makes H. heidelbergensis really interesting is
not only the strong similarities between the early
African and European forms (around 0.5±0.4 Mya),
but how they differentiate afterwards. In Europe, a
number of fossils of late Middle Pleistocene date
(0.3±0.125 Mya) show characteristics later to be
found only among Neanderthals. In Africa, differ-
entiation from H. heidelbergensis also takes place,
with the appearance of features that we identify
as modern human traits. Thus we have in H. heidel-
bergensis an even larger-brained hominin species
that in the middle of the Pleistocene epoch
extended from Africa to Europe. It is this species,
or its possible descendant, Homo helmei, which in
the opinion of many researchers represents the last
common ancestor of Neanderthals and modern
humans.
 Human Evolution 403

Paleoclimate Technology Evolutionary patterns

cold hot Europe Africa Asia/Australia
Late Modern human
Mode 4 Pleistocene extinctions,
and modern dispersals and
Mode 5 expansions regional isolation
Blades/
Early
microliths
modern
human
dispersals
Dispersal of
H. helmei with Possible
Mode 3 dispersal
Mode 3 of Eurasian
industries
Prepared H. heidelbergensis
from Africa to
cores/ and H. helmei
Levant/Europe
Levallois to north-east
?
technique Asia
− Middle
Paleolithic
(Mousterian) and
Middle Stone Age
Dispersal of
H. heidelbergensis with
Persistence of
Mode 2 industries
Mode 1
from Africa to the
industries in Asia
Levant and Europe
and parts of
Formation of the
Europe
Movius line

Dispersal of
H. erectus with Dispersal of
Mode 1 industries south-east Asian
from Africa to H. erectus
Europe (H. antecessor) to north-east
Asia

Dispersal of
H. erectus with
Mode 1 and 2
industries
from Africa to
the Levant
(separate strata,
Mode 2 Ubeidiya)
Bifaces/
Dispersal of
handaxes
H. erectus with
− Acheulean
Mode 1 industries
from Africa to W. Asia
and the Caucasus

Dispersal of H. erectus with

Mode 1 Mode 1 industries from
Choppers/ Africa to south-east Asia
pebble tools African H. erectus/
− Oldowan H. ergaster with
Mode 1 industries

Figure 3. The record of human evolution in the Pleistocene.

404 Human Evolution

Figure 3. (cont.)

Evolutionary events Genetic history Paleobiology

Evolution of modern
Extinction of the
human diversity
last archaic populations
African and European
speciation events:
origin of Neanderthals in
the context of directional
selection and origin of modern
humans in the context of
African fragmentation
and resource
Evolution of H. helmei in scarcity
the context of sub-Saharan
Africa, and establishment
of the shared level of
Neanderthal and
H. sapiens cognitive
abilities
Recent expansions and
extensive gene flow
Relative gracilization,
Early human population
encephalization and
expansions
language specialization
Ancestral bottleneck.
in African H. sapiens
Coalescence of mtDNA
and Y chromosome
lineages Thermoregulatory
specializations and
body weight increase in
European
H. neanderthalensis
? Separation of
Neanderthal and
Body size reduction?
human populations
Encephalization
Modern life-history
Coalescence of parameters
Neanderthal and
human mtDNA

Evolution of
H. heidelbergensis Increase in body size
in Africa and and robusticity
rapid expansion into Language?
the Levant and Europe

Periodic extinction
of hominid
populations in Coalescence of
Europe? human nuclear
Evolution of H. antecessor in Europe? DNA genes
Specializations
Stabilizing towards cold
selective forces adaptations in
in Asia European Middle
Extinction of the last Pleistocene hominins
australopithecines

Specialization
of Asian H. erectus

Greater habitat and

climatic tolerance

Extinction of early Body shape:

Homo species African increased size
H. ergaster linearity
H. erectus Encephalization:
slower growth rates
Dietary change?
increased meat
Period of greatest consumption
hominid diversity faster growth
Increased geographical
range

The Origins of Modern Humans and
Neanderthals
The climatic cycles of the last half-million years
promoted periods of recurrent contact between Af-
rican and European hominins (as shown by the
strong similarities between Homo heidelbergensis in
both continents), while the timing of such cycles
suggests that the direction of contact was from
Africa to Europe. It is in these contacts that the
origin of the lineages leading to modern humans
on the one hand, and Neanderthals on the other,
are to be found. As mentioned above, different
opinions exist as to when the African and European
lineages exchanged significant genes for the last
time, whether it was around half a million years
ago in the form of H. heidelbergensis, or whether it
was more recently, around 300 000 years ago, in the
form of H. helmei. Independent of this, from around
130 000 years ago African and European fossils
show modern human and Neanderthal features
respectively.
The period preceding the appearance of Nean-
derthals and modern humans, 200 000±130 000
years ago, was one of extensive glaciation. These
extreme climatic conditions would have generated
different pressures on the populations of both con-
tinents. In Europe, thermoregulation would have
been the main factor affecting survivorship, while
in Africa the immediate effect would have been
scarcity of resources because of aridity and eco-
logical fragmentation. It is in response to these
different selective pressures that researchers inter-
pret the adaptive differences between Neander-
thals and modern humans.
Many of the features of H. sapiens and H. nean-
derthalensis were inherited from their large-brained
and large-bodied ancestor, who had a relatively
long ontogeny, who manufactured complex tools,
used a degree of planning and social organization
in its strategy to exploit the environment, and most
importantly, probably had some linguistic abilities.
The differences between the two species thus re-
flect their modification of this ancestral heritage in
order to meet the pressures posed by their respect-
ive evolutionary environments. In Neanderthals,
these adaptations are represented by changes to
the face, in particular how much it protruded and
how large the nasal aperture was, together with a
broad chest, and an overall dense and robust skel-
eton, with relatively short distal limb proportions.
These changes are thought to reflect thermoregula-
tory adaptations to the warming of inhaled air and
overall heat retention, as well as to a tough lifestyle
that required extensive nomadism. In modern
Human Evolution 405
humans these differentiating features are more dif-
ficult to specify, largely because the history of the
species is one of early geographical differentiation,
thus making it difficult to identify the universal
traits. However, a few can be described ± these
are represented in the small size of a nonprotrud-
ing face with a chin in the jaw, in a differentiated
cranial shape that resulted in a rounded head, and
an overall gracile body. The increased aridity
throughout Africa that led to increasingly scarce
resources would also have led to demographic con-
tractions, which might in turn have increased the
role of genetic drift, as well as set new selective
pressures. The changes observed in the evolution
of a modern morphology, to a large extent associ-
ated with the gracilization of the skeleton, are con-
sistent with selective pressures on resources, by
which a `cheaper' skeleton would have been ad-
vantageous. Recent studies on the ontogeny of Ne-
anderthal and modern human cranial growth
suggest that the differences characteristic of both
species were present at birth.
Genetic evidence has significantly increased our
understanding of the evolution of these two lin-
eages. Genetic studies have made a major contribu-
tion towards establishing a recent African origin for
modern humans, and revealing their relationship
to Neanderthals through the retrieval of ancient
DNA. The genetics of living populations can be
used to draw inferences about their past, in a simi-
lar way to the use of molecular clocks to under-
stand the phylogenetic relationship between living
apes and humans, and to infer, through genetic
distances, the time lapsed since their divergence.
The study of the genetic origins of modern humans
is based on several genes and a range of different
techniques.
For a long time geneticists used classical markers
(polymorphic gene products) to study the variation
among human populations. These are measured as
percentages of different alleles in different groups,
and their study has been championed by the Italian
geneticist Luca Cavalli-Sforza. Classical markers
are interesting, particularly because a large number
of gene systems can be analyzed together to pro-
vide a tree of relationships of human populations.
These are population trees, in the sense that they
track the multivariate history of populations
through the genetic expression of many systems.
The results of studies of classical markers point
clearly to an African origin of all modern humans,
with the first branches of a human genetic tree
taking place among African populations.
Since the 1980s molecular markers have been
used to track the history of human populations.
406 Human Evolution
These studies were first developed using the
mitochondrial deoxyribonucleic acid (mtDNA)
genome, but are today based on a number of
genes, including mutations on the Y chromosome.
The advantages of the mtDNA and Y chromosome
genomes are that they are inherited through only
the maternal or the paternal line, and therefore do
not undergo recombination during meiosis. Fur-
thermore, most of these markers are believed to
be selectively neutral, which means that not only
does their distribution reflect history rather than
selection, but also they accumulate mutations faster
than functional genes. Together, they provide the
most powerful tool for the investigation of the evo-
lutionary history of living human populations (of
course, they cannot throw light on any group that
has since become extinct, with the rare exception of
the extraction of DNA from actual ancient
remains). Their disadvantage is that they provide
`gene trees' rather than `population trees', and the
history of any population is made up by the sum of
each gene's history. This is an important caution,
as, for example, mtDNA and Y chromosome histor-
ies are not the same because men and women have
had different histories determined by their patterns
of mating and dispersal behavior. These single
gene systems are analyzed in a variety of ways,
the most important of which are based on coales-
cence theory and network analysis (Figure 4).
The results of these studies all point to an African
origin of modern humans. Among the thousands
of individuals sampled for either mtDNA or Y
chromosome markers, Africans are the most di-
verse in having the greatest number of mutations.
Figure 4. Basic principles of coalescence theory, showing as an example the coalescence of a maternally inherited gene.
Furthermore, all the earliest branching events occur
among African individuals. In other words, these
data show that the last common ancestor of all
living humans was part of a population from
which Africans derive, and that only much later,
other populations branched off this group. The ap-
plication of a molecular clock to these various
systems provides a range of dates for the last
common ancestor of all humanity ± between
200 000 and 130 000 years ago, thus in full agree-
ment with the paleoanthropological record.
Molecular data can also provide information on
the demographic parameters of the ancestral popu-
lation of any given species. The basis for these is the
estimation of the effective population size of a
given system, derived from the equation
FST ˆ 2Ne …1†
Both FST (a measure of diversity in a gene) and m
(the mutation rate of a gene) are empirical observa-
tions, allowing the calculation of Ne, the effective
ancestral population size. From such calculations,
geneticists have suggested that the ancestral
population of Homo sapiens underwent a severe
demographic bottleneck before its diversification.
Estimates of the magnitude of the ancestral bottle-
neck vary, ranging from 1000 to 10 000 individuals.
Transforming these values into census values (i.e.
the whole population), it is possible to estimate that
the ancestral population of modern humans was at
one point composed of 3000 to 30 000 people. The
scar of this bottleneck is one of the main features of
Homo sapiens. Although our species contains more
than 6 billion individuals, we have a tenth of the

diversity of any of our close primate relatives. From
their African origin, modern humans underwent
various periods of demographic expansion that
led to multiple dispersals into Europe and Asia. A
large proportion of the diversity among human
populations can be related to this early fragmenta-
tion as small groups of hunter-gatherers dispersed
and colonized the world. One such dispersal took
modern humans into Europe around 40 000 years
ago, into the Neanderthals' homeland.
The nature of the relationship between modern
humans and Neanderthals has been much debated.
After modern humans dispersed into Europe,
Neanderthal fossil remains and their associated
Mousterian technology disappeared from most of
the continent. However, in certain areas these two
hominins appear to have coexisted for some 10 000
years. Studies of Neanderthal mtDNA, obtained
through the extraction of ancient DNA from three
fossils, show that 600 000±500 000 years ago there
lived a population from which both Neanderthals
and modern humans descend, although not when
this population separated into its European and
African descendants. Furthermore, the absence of
the particular mutations observed in the Neander-
thal samples among the thousands of recent people
studied indicate that Neanderthals did not contrib-
ute to the living mtDNA gene pool, offering further
evidence for the extinction of Neanderthals, al-
though whether small amounts of interbreeding
took place 40 000 years ago cannot be ruled out.
The last 30 000 years have been probably the first
time in the last 5 million years in which only one
species of hominin exists in the world.
HUMAN DISPERSALS
Dispersal is an essential element of the evolution-
ary process for all organisms. Most species start as
small localized populations, and if successful
spread more widely to fill up the available habitat.
The rate and direction of any such dispersal are
functions of the ability of the species to survive
and thrive in the environments encountered, and
of its own characteristics. Looking broadly across
the animal kingdom, it is clear that not all species
disperse to the same extent: some are much more
widely distributed than others. Lions, for example,
existed across the whole of Africa and Eurasia,
whereas most monkeys are very localized.
Human evolution fits into this biogeographic
pattern. From their localized African origins,
humans have come to live all over the world, and
that can only have happened as a result of disper-
sals. However, that pattern of colonization is a
Human Evolution 407
complex one, occurring irregularly over long
periods. The first part of human evolution, that of
the period 5±2 Mya, was dominated by australo-
pithecines, and they never managed to disperse
beyond Africa. In this sense they were much like
the other anthropoid apes. With the emergence of
Homo more extensive dispersals occurred. From
2±1.5 Mya Homo ergaster/erectus spread into the
southern parts of Asia, becoming the first non-
African hominin. After 1 Mya they gradually dis-
persed into the colder, more northerly latitudes of
northern Eurasia. However, rather than this being a
single continuous dispersal, it is probably best seen
as a series of repeated events, probably inter-
spersed with extinction of populations. The pri-
mary driving force of this process would have
been the climate, particularly after the onset of gla-
cial conditions.
The dispersal of modern humans is another of
these events, beginning during the warmer phase
of the last interglacial period, around 120 000 years
ago. Again, the dispersal of modern humans
was not a continuous single event, but is best de-
scribed as a series of multiple dispersals. The major
events in this process were the first dispersals out
of Africa and along the southern rim of Asia,
through to Australia, around 60 000 years ago; the
dispersals into northern Eurasia from around
40 000 years ago; recolonization following the ex-
tremes of the last glaciation around 15 000 years
ago, resulting in the first peoples in the New
World; and, perhaps most important in terms of
the current human population, the spread of the
first farming peoples in the last 10 000 years. While
archeologists and anthropologists have argued
about the relative importance of migration com-
pared with local development, it is becoming in-
creasingly evident that the ability to migrate and
colonize has been an important element of human
evolution.
HUMAN TECHNOLOGIES
The archeological record represents the appearance
and development of hominin technological abil-
ities, and it extends to the last 2.5 million years.
Stone tools are thus associated with the second
half of hominin evolution, most markedly with
species of the genus Homo, although tools in pos-
sible association with late australopithecines have
been found. What is the importance of the archeo-
logical record?
. It provides very strong evidence of behavior in the
past, ranging from use of the landscape, to subsistence
strategies, to defense organization.
408 Human Evolution
. It provides unique evidence of cognitive processes in
extinct taxa, through the inferred thought processes
required to produce either the tools themselves or
their accumulation pattern.
. Owing to its durability, it provides the best evidence
for the spatial distribution of populations in the past,
allowing, in certain circumstances, paleodemographic
inferences to be made.
The range of tools possible is unlimited to our
minds, with new inventions appearing daily, but
this was not so in the past. Several factors acted to
constrain the range of tools earlier hominins made:
manual dexterity, cognitive ability, and availability
of raw materials. This has allowed archeologists to
categorize stone tool traditions according to the
shape of the tools produced or the methods by
which they were made. Several such categoriza-
tions with different levels of detail exist (Figure 5).
One of the most useful classifications of the archeo-
logical record is that developed by Graham Clark.
He organized stone tools according to what he
called `modes', and the timing and distribution of
these modes, as well as trends within the tech-
nocomplexes they define, have important con-
sequences for our interpretation of hominin
cognition through time (see Figure 3).
For those working within the framework of a
recent African origin of modern humans, Homo
heidelbergensis has been considered as the last
common ancestor of Neanderthals and modern
humans. This view argues that, some time between
600 000 and 500 000 years ago, during an intergla-
cial phase, hominins dispersed from Africa to
Europe, taking Mode 2 or Acheulean technologies
to Eurasia for the first time. However, around
300 000 years ago, a new technology appears in
Africa and soon afterwards in Europe. This tech-
nology, known as Mode 3, or Middle Stone Age
(Africa) or Middle Paleolithic (Europe), has some
striking features. Hominins were no longer turning
stone cores into tools, but rather flaking them to
obtain thinner, preshaped, and more abundant
objects. In order to do this, hominins needed to
prepare the cores prior to flaking, a step that re-
quired a level of mental abstraction. This technique
of preparing the core before flaking, first observed
within late Mode 2/Acheulean assemblages, is the
technological background of both Neanderthals
and modern humans.
Neanderthals and modern humans may have
inherited this ability from a more recent common
ancestor, known as Homo helmei. This would be
consistent not only with the archeological record,
but also with the little evidence there is for the
evolution of language. Language is considered
one of the key traits that define and to some extent
structure human cognition and human behavior.
However, the ability to speak does not affect the
skeleton, and thus its origins cannot be traced in the
fossil record. However, in recent years two indirect
lines of evidence for linguistic ability among homi-
nins have been explored ± one related to the ex-
pansion of the vertebral canal at the point of
innervation of the diaphragm, and the other related
to the size and shape of the hyoid bone. Both sug-
gest that some of the anatomy that allows human
speech evolved within the last half-million years,
and that Neanderthals probably had some lan-
guage ability. This would further reinforce the
view that the two lineages share a recent common
ancestor in the late Middle Pleistocene who had
increased cognitive abilities.
HOMININ EVOLUTION: THE EVIDENCE
Paleoanthropology has recovered a rich record of
the evolutionary history of humans. This record is
made up of fossils and of the material culture left
behind by various hominin species, integrated with
the information derived from geology, paleon-
tology, and paleoclimatology. It is further enriched
by the evolutionary inferences possible from our
own genetic diversity and that of our closest living
relatives, the apes. Together, this information
allows the reconstruction of a complex process, in
which we recognize some broad patterns:
. Hominin origins, the divergence of our lineage from
that leading to chimpanzees, date to 7±5 million years
ago, and are based on changes in locomotive strategy.
. The first 3 million years of this history, during the
Pliocene, are characterized by the diversification of `bi-
pedal apes' within eastern and southern Africa, leading
to the evolution of two separate lineages: one in which
megadontic adaptations are associated with the con-
sumption of large quantities of low-quality foods, the
other in which encephalization, increased carnivory,
tool use, and a longer ontogeny are associated with a
more opportunistic exploitation of resources.
. The last 2 million years, the Pleistocene, are character-
ized by dramatic geographical expansions in the con-
text of deteriorating climatic conditions, and the
establishment of distinct regional trajectories ex-
pressed in both morphology and material culture.
. Neanderthals and modern humans represent the
contemporaneous evolution, in Europe and Africa re-
spectively, of large-brained and behaviorally complex
hominins from a common ancestor who lived 500 000±
300 000 years ago, and thus uniquely share a number of
traits.
. From their African ancestral homeland, modern
humans dispersed throughout the world by means of
multiple events. These, together with the different
 Human Evolution 409

Some Archeological Possible

archeological periods and associated hominin
assemblages or technological modes species
cultures

0
Aurignacian UPPER

MODE 4/5
Solutrean PALEOLITHIC
Magdalenian
Wilton Homo sapiens
Dabban

50

MODE 3
MIDDLE
PALEOLITHIC/
MIDDLE STONE
AGE
Mousterian
Still Bay H. heidelbergensis
100 Szeletian (H. helmei )
Aterian H. neanderthalensis
H. sapiens
Time (103 years ago)

250

LOWER
PALEOLITHIC/
EARLY STONE H. ergaster
AGE H. erectus
H. heidelbergensis
Acheulean
MODE 2

500

Australopithecus
garhi
? A. boisei
MODE 1

1000 H. habilis
H. rudolfensis
H. ergaster
H. erectus
(H. antecessor )
Oldowan
pebble
2000 tool

Figure 5. Temporal distribution of archeological assemblages and modes.

selective pressures in the areas colonized and the
effects of drift on small hunter-gatherer groups,
shaped the diversity of the species.
. The extinction of the Neanderthals was probably asso-
ciated with the migration of modern humans into
Europe and the changes in the competitive environ-
ment caused by this influx of people.
. In recent millennia, and in association with the shift to
an agricultural and sedentary lifestyle, humans have
undergone a demographic revolution that took the
species from a few thousand to over 6 billion individ-
uals.
. When compared with other species, Homo sapiens
shows a distinct lack of genetic diversity in spite of its
current population size (the heritage from a recent
ancestry as a very small population in Africa), and
most of that diversity is to be found within, rather
than between, populations.
410 Human Evolution
In summary, humans are the result of the same
evolutionary mechanisms that acted on other
animal lineages. The history of their lineage is
made up of the evolution, differentiation, and ex-
tinction of a large number of species during the last
7±5 million years. The history of the species is rela-
tively recent and characterized by unprecedented
geographic and demographic success, one in which
our unique cultural complexity and diversity
played a major part.
Further Reading
Boyd R and Silk JB (2000) How Humans Evolved, 2nd edn.
Conroy G (1997) Reconstructing Human Origins: A Modern
Synthesis. New York, NY: WW Norton.
Human Factors and Ergonomics
Peter A Hancock, University of Central Florida, Orlando, Florida, USA
Raja Parasuraman, Catholic University of America, Washington, DC, USA
CONTENTS
Introduction
Cognitive science, HF/E and the psychology of action
Human performance capabilities and limitations
Human interaction with technology
Human factors and ergonomics is the study and
practice of designing technology around the cap-
abilities, both physical and mental of the user. In-
cluded in the chapter are brief descriptions of
neuroergonomics, the combining of neuroscience
and ergonomics, and human factors and ergonom-
ics in the twenty-first century.
INTRODUCTION
Most individuals reading an Encyclopedia of Cogni-
tive Science will have more than a passing familiar-
ity with the research areas and issues that they
encounter in the text. Yet this may not be true for
human factors and ergonomics (HF/E), since these
terms appear to lie outside the mainstream of cog-
nitive science. However, we hope to persuade the
reader that HF/E represents a central concern of
any cognitive science that aspires to be relevant to
the real world. We support this claim by beginning
with a reference to the origins of cognitive psych-
ology and its historical antecedents.
View publication stats
Donnelly P and Foley RA (eds) (2001) Genes, Fossils and
Behaviour: An Integrated Approach to Human Evolution.
Brussels, Belgium: NATO.
Foley RA (1995) Humans Before Humanity: An Evolutionary
Perspective. Oxford, UK: Blackwell.
Gamble C (1994) Timewalkers: The Prehistory of Global
Colonization. Cambridge, MA: Harvard University
Press.
Johanson D and Edgar B (1996) From Lucy to Language.
New York, NY: Simon & Schuster.
Klein RG (1999) The Human Career: Human Biological and
Cultural Origins. Chicago, IL: Chicago University Press.
Stringer C and Gamble C (1993) In Search of the
Neanderthals. New York, NY: Thames & Hudson.
Tattersall I (1995) The Fossil Trail: How We Know What We
Think We Know About Human Evolution. Oxford, UK:
Oxford University Press.
Intermediate article
Cognition and the physical environment
Neuroergonomics
HF/E in the twenty-first century
Summary
Cognitive Psychology and Human
Factors and Ergonomics
Most modern cognitive scientists are unconcerned
with, or not fully cognizant of, the goals, issues and
findings of HF/E. This relative neglect is under-
standable, but paradoxical, since the post-Second
World War origin of modern cognitive psychology
owes much to HF/E. The `cognitive revolution' of
the late 1950s supplanted the behaviorist tradition
that dominated American (and, to a lesser extent,
European) psychology. This paradigm shift is
rightly seen as the beginnings of the cognitive sci-
ence movement, accompanied by developments in
artificial intelligence (Newell and Simon, 1956) and
linguistics (Chomsky, 1957). Two of the leading
figures of this cognitive revolution were George
Miller in the USA and Donald Broadbent in the
UK. The latter can be seen in action at the prime
of his career in Figure 1.
Broadbent's classic information-processing
model of the human cognitive system (Broadbent,