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Human Cognition
Human Evolution
Introductory article
Introduction
Hominin origins and early hominin diversity
Human evolution in the Pleistocene
Human dispersals
Human technologies
Hominin evolution: the evidence
INTRODUCTION
Human Evolution
Ch
Hominidae
Pongidae
O Go
Hylobatidae
Pongidae
Hominidae
Hylobatidae
G O Go Ch
399
forests, lowering of sea levels, and major geotectonic 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 orangutans.
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. Hominins 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
result from the geographical separation of populations 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 65 Mya, fascinating 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.
400
Human Evolution
of terrestrial locomotion, it clearly provides an efficient way of moving in open environments for long
periods, especially when compared with knucklewalking 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 combination of these factors gave bipedal apes a competitive advantage in relation to their quadrupedal
relatives at the time.
However, when considering the selective pressures 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.
Human Evolution
401
Longer ontogeny
Sociality
BRAINS
= BMR
Tools/technology
Memory
Language?
Problem-solving
MEAT
EATING
Guts
Average
body
size
Range expansion
Longer legs
Tall individuals
Figure 2. The complex adaptive system that underlies the evolution of the genus Homo. BMR, basal metabolic rate.
402
Human Evolution
Human Evolution
Paleoclimate
cold
hot
Technology
Evolutionary patterns
Europe
Mode 4
and
Mode 5
Blades/
microliths
Mode 3
Prepared
cores/
Levallois
technique
Middle
Paleolithic
(Mousterian) and
Middle Stone Age
Africa
Asia/Australia
Late
Pleistocene
modern
expansions
Modern human
extinctions,
dispersals and
regional isolation
Early
modern
human
dispersals
Dispersal of
H. helmei with
Mode 3
industries
from Africa to
Levant/Europe
?
Possible
dispersal
of Eurasian
H. heidelbergensis
and H. helmei
to north-east
Asia
Dispersal of
H. heidelbergensis with
Mode 2 industries
from Africa to the
Levant and Europe
Formation of the
Movius line
Persistence of
Mode 1
industries in Asia
and parts of
Europe
Dispersal of
H. erectus with
Mode 1 industries
from Africa to
Europe (H. antecessor)
Mode 2
Bifaces/
handaxes
Acheulean
Dispersal of
south-east Asian
H. erectus
to north-east
Asia
Dispersal of
H. erectus with
Mode 1 and 2
industries
from Africa to
the Levant
(separate strata,
Ubeidiya)
Dispersal of
H. erectus with
Mode 1 industries
from Africa to W. Asia
and the Caucasus
Mode 1
Choppers/
pebble tools
Oldowan
African H. erectus/
H. ergaster with
Mode 1 industries
403
404
Human Evolution
Figure 3. (cont.)
Evolutionary events
Evolution of modern
human diversity
Evolution of H. helmei in
the context of sub-Saharan
Africa, and establishment
of the shared level of
Neanderthal and
H. sapiens cognitive
abilities
Extinction of the
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
scarcity
Genetic history
Recent expansions and
extensive gene flow
Early human population
expansions
Ancestral bottleneck.
Coalescence of mtDNA
and Y chromosome
lineages
? Separation of
Neanderthal and
human populations
Coalescence of
Neanderthal and
human mtDNA
Evolution of
H. heidelbergensis
in Africa and
rapid expansion into
the Levant and Europe
Periodic extinction
of hominid
populations in
Europe?
Evolution of H. antecessor in Europe?
Stabilizing
selective forces
in Asia
Extinction of the last
australopithecines
Paleobiology
Relative gracilization,
encephalization and
language specialization
in African H. sapiens
Thermoregulatory
specializations and
body weight increase in
European
H. neanderthalensis
Body size reduction?
Encephalization
Modern life-history
parameters
Coalescence of
human nuclear
DNA genes
Specializations
towards cold
adaptations in
European Middle
Pleistocene hominins
Specialization
of Asian H. erectus
Extinction of early
Homo species
Period of greatest
hominid diversity
African
H. ergaster
H. erectus
Body shape:
increased size
linearity
Encephalization:
slower growth rates
Dietary change?
increased meat
consumption
faster growth
Increased geographical
range
Human Evolution
405
humans these differentiating features are more difficult 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 nonprotruding 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 contractions, 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 associated with the gracilization of the skeleton, are consistent with selective pressures on resources, by
which a `cheaper' skeleton would have been advantageous. Recent studies on the ontogeny of Neanderthal 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 lineages. Genetic studies have made a major contribution 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 similar way to the use of molecular clocks to understand 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 provide 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
Figure 4. Basic principles of coalescence theory, showing as an example the coalescence of a maternally inherited gene.
Human Evolution
HUMAN DISPERSALS
Dispersal is an essential element of the evolutionary 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 dispersals. However, that pattern of colonization is a
407
HUMAN TECHNOLOGIES
The archeological record represents the appearance
and development of hominin technological abilities, 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 possible association with late australopithecines have
been found. What is the importance of the archeological 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
Human Evolution
Aurignacian
Solutrean
Magdalenian
Wilton
Dabban
100
UPPER
PALEOLITHIC
MODE 3
50
Archeological
periods and
technological modes
MODE 4/5
Some
archeological
assemblages or
cultures
409
Possible
associated hominin
species
Homo sapiens
MIDDLE
PALEOLITHIC/
MIDDLE STONE
AGE
Mousterian
Still Bay
Szeletian
Aterian
H. heidelbergensis
(H. helmei )
H. neanderthalensis
H. sapiens
250
LOWER
PALEOLITHIC/
EARLY STONE
AGE
MODE 2
Acheulean
Australopithecus
garhi
? A. boisei
H. habilis
H. rudolfensis
H. ergaster
H. erectus
(H. antecessor )
MODE 1
500
1000
2000
H. ergaster
H. erectus
H. heidelbergensis
Oldowan
pebble
tool
410
Human Evolution
Intermediate article
Introduction
Cognitive science, HF/E and the psychology of action
Human performance capabilities and limitations
Human interaction with technology
INTRODUCTION
Most individuals reading an Encyclopedia of Cognitive Science will have more than a passing familiarity 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 cognitive 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 psychology and its historical antecedents.