Mark Maslin - The Cradle of Humanity - How The Changing Landscape of Africa Made Us So Smart-Oxford University Press (2017)

T he Cr adle of Hu m a ni t y
The Cradle
of Humanity
How the changing landscape
of Africa made us so smart
Mark Maslin
3
1
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Dedicated to Professor Martin Trauth without whom none
of this work on human evolution would have been possible.
For e wor d
The Cradle of Humanity is an important contribution to the exten-

sive literature on the subject of human evolution. Sadly there is no
absolute truth to be offered as to what and who became what and
who; any number of interpreters have published their views, and
sometimes even their arguments and reasons for holding such
views. The point that emerges is that data have yet to be found,
some may never be found, but each year, the aggregate of know-
ledge is more and more persuasive. We humans are indeed the
result of biological evolution and those facts exist in museums
and research collections. I stress this because even now in this
century, countless allegedly learned men and women, active in
science, speak of the ‘theory of human evolution’. Surely—and
this book fully attests to this—the evidence for evolution such as
the fossils, dates, archaeology, and countless re-evaluations lead-
ing to refinements in the data sets are now so comprehensive and
consistent that the idea of the ‘theory’ can be discarded entirely
and permanently.
The multidisciplinary fields of investigation are primarily con-
cerned with the analysis of the facts of evolution. For example,
how much can investigators learn regarding why, when, and how
our evolution has happened? We know for example that we had
small-brained early ancestors and later, larger brained ones. The
vii
for e wor d
relationship between body mass and brain size grew over several
million years. This is known, but why and in which populations
is yet to be fully understood. Today we are fully bipedal and we
use our hands (in conjunction with our brains) to manipulate
objects rather than using them to support our bodies. Upright
posture puts huge strains on the human frame (back problems,
hip joints, etc.) and yet it is clearly advantageous. What pressures
in the environment led to the shift from four legged to two legged
locomotion? Explanations abound, but the fact is it happened,
and this is important. Similarly, the question of the role that
speech played in our becoming us: when, why, and where? These
are questions which are important. Was the process gradual
or rapid?
Professor Mark Maslin does not dwell in depth on these and
many other evolutionary milestones concerning us, but what he
has done, and this makes the book so interesting, is to put our
evolution into a larger picture—a context that is increasingly
understood but seldom talked about in popular texts. For many,
the relationship between our place in the Earth’s history and the
history of our planet itself makes the story so much more accept-
able. The third, fourth, fifth, and sixth chapters deal with our
Earth and its history. The chapters discuss temperature gradients,
ocean currents, rainfall patterns, and much more on a global
scale. Many will find the account explaining mountain building,
lifting, plate tectonics, and their impact or influence on ecology,
changes in vegetation, and sedimentation useful when thinking
about why, over time, life forms have had to change. Most of this
discussion is written to explain Africa, and in particular East
Africa’s special circumstances for favouring, or one might say
driving, human evolution.
viii
for e wor d
I have found that many of the concerned questions that have

been put to me over forty years of public lecturing are now
answered in this one book. This is of great value to students and
the broader swathe of public interest. That Maslin’s account is in
some chapters different from other books should not be a worry
for the reader; it illustrates the complexities of science and the
inevitable debates arising from a variety of ideas to explain the
hard facts. There are countless fossil specimens now known that
simply make doubting our evolution an exercise in self delusion.
Read this excellent narrative and see for yourself.
Richard Leakey, FRS

4 March 2016
ix
Ack now l edgemen t s
I would like to thank Johanna, Alexandra, and Abbie Maslin for

being there for me. I would also like to thank my colleagues who
have put up with me continually asking questions who helped me
on my journey of discovery within the wonderfully contentious
field of human evolution. In random order they are: Beth
Christensen, Annett Junginger, Alice Roberts, Susanne Shultz,
Katie Wilson, Kit Opie, Leslie Aeillo, Craig S. Feibel, Robin
Dunbar, Brian Cox, Richard Leakey, Djuke Veldhuis, Rick Potts,
Adam Rutherford, Peter C. Kjaergaard, Alan Deino, Matt Pope,
Kaye Reed, Chris Stringer, Martin Trauth, Fred Grine, Meave
Leakey, Rob Foley, Sam Nicholson, David Price, Phil Hopley, Fiona
Scott, Victoria Ling, Marta Lahr, Peter de Menocal, Janet Hope,
Helen Douglas Dufresne, Pete Ilsley, John Kingston, Frank
Schäbitz, Simon Conway-Morris, Asfawossen Asrat, Chronis
Tzedakis, Matt Sponheimer, Fiona Jones, Verena Foerster, Julia
Lee-Thorp, Jim Rose, Chris Brierley, and the other two Marks
‘Thomas’ and ‘Collard’. Latha Menon and Jenny Nugee for the
excellent editing, nagging, and support. All the staff and friends at
UCL who make it the best place to work. Peter Kjaergaard and
Martin Trauth for reviewing the manuscript. And last, but not
least, Miles Irving for his excellent and distinctive illustrations.
xi
Con t en t s
List of Figures xv
1. Introduction 1
2. Early Human Evolution 13
3. Tectonics and Climate 41
4. Cradle of Humanity 61
5. Global Climate Change 79
6. Celestial Mechanics 100
7. African Climate Pulses 128
8. The Social Brain 149
9. The Future of Humanity 183
References and Further Reading 199

Index 225
xiii
L is t of F igur e s
1. Major biological events in the 4.6 billion years history of

the Earth. 5
Adapted from <https://en.wikipedia.org/wiki/History_of_Earth>.
Public Domain.
2. Key morphological features that define hominids. 8
3. Map of the location of the most important hominin fossil

finds in Africa.15
4. Location of the hole in the base of the skull through which

the spinal cord passes (foramen magnum) in humans and
chimpanzee, showing the central position essential for
bipedal movement.17
Adapted from figures 9.12, 9.13, and 9.14 in Lewin, R. and Foley,
R.A. © 2004. Principles of Human Evolution, John Wiley & Sons
© by Blackwell Science Ltd.
5. Evidence of bipedalism from the shape of the neck of

the femur (thighbone).19
Reprinted from Lovejoy, C. Owen. 2005. The Natural History of
Human Gait and Posture Part 1. Spine and Pelvis. Gait & Posture 21(1):
95–112 and Lovejoy, C. Owen. 2005. The Natural History of Human
Gait and Posture Part 2. Hip and Thigh. Gait & Posture 21(1): 113–24,
with permission from Elsevier. <http://www.sciencedirect.com/
science/journal/09666362>.
xv
l ist of figu r es
6. New divisions at the start of the Homo lineage suggested

by Susan Anton and colleagues based on skull morphology,
which challenges the original H. habilis, H. rudolfensis, early
Homo split. Figure includes the new stone tools (3.3 Ma)
and early Homo (2.8 Ma) finds.28
From Antón, Susan C., Potts, Richard, and Aiello, Leslie C. 2014.
Evolution of early Homo: An integrated biological perspective.
Science 345(6192). Reprinted with permission from AAAS. And
reprinted by permission from Macmillan Publishers Ltd:
Harmand, S., Lewis, J.E., Feibel, C.S., Lepre, C.J., Prat, S., and
Lenoble, A. 2015. 3.3-millionyear-old stone tools from
Lomekwi 3, West Turkana, Kenya. Nature 521: 310–15.
7. Records of hominin brain capacity and East African

hominin diversity over the last five million years. 33
Based on figure 6 in Maslin, M.A., Brierley, C.M., Milner, A.M.,
Shultz, S., Trauth, M.H., and Wilson, K.E. 2014. East African climate
pulses and early human evolution. Quaternary Science Reviews, 101: 1–17,
CC-BY-SA-3.0 <https://creativecommons.org/licenses/by/3.0/>.
8. Summary of the hominin species covering the last

five million years and major dispersal events. 39
CC-BY-SA-3.0 <https://creativecommons.org/licenses/by/3.0>.
9. Illustration of the huge effect the latitudinal location of

continents have on the Equator–Pole temperature gradient.
The present-day northern and southern hemisphere
temperature gradients are given as a comparison. 43
Based on figure 21 in Maslin, M. 2013. Climate: A Very Short
Introduction, Oxford University Press, Oxford, by permission of
Oxford University Press.
10. Illustration of the significant alteration of ocean currents with

different ocean gateways within longitudinal continents. 46
xvi
l ist of figu r es
Based on figure 23 in Maslin, M. 2013. Climate: A Very Short Introduction,

Oxford University Press, Oxford, by permission of Oxford University Press.
11. The effect of changing ocean gateways on the location

and intensity of deep ocean circulation. 49
Introduction, Oxford University Press, Oxford, by permission
of Oxford University Press.
12. Mountain rain-shadow. 52

13. The effects of mountains and plateaus on global rainfall. 53

14. The effects of plateaus and ice sheet on atmospheric

circulation. 55
Based on figure 27 in Maslin, M. 2013. Climate: A Very Short Introduction,
Oxford University Press, Oxford, by permission of Oxford University
Press.
15. Long-term links between tectonics, sea level, climate,

biodiversity, and extinctions. 60
Base on figure 29 in Maslin, M. 2013. Climate: A Very Short
16. Tectonic rifting and ocean formation. 62
17. Map of plate boundaries at the African Rift Valley triple

junction.64
Based on ‘Map of East Africa’, United States Geological
Survey <https://en.wikipedia.org/wiki/African_Plate#/media/
File:EAfrica.png>. Public Domain.
xvii
l ist of figu r es
18. Explanation of horst-graben and half-graben geological

formations.65
19. Tectonic development of the East African Rift Valley system. 66

Adapted from figure 3.11 from Lewin, R. and Foley, R.A. © 2004.
Principles of Human Evolution, John Wiley & Sons © by Blackwell
Science Ltd.
20. Climatic effects of the development of the East African

Rift Valley system. 68
Adapted from figure 3.11 from Lewin, R. and Foley, R.A. © 2004.
Principles of Human Evolution, John Wiley & Sons © by Blackwell
Science Ltd.
21. Influence of Tibetan uplift and the African Rift Valley

on regional atmospheric circulation during Northern
Hemisphere summer. 69
22. Vegetation effects of the development of the East

African Rift Valley system. 71
23. Carbon isotope differences and relative abundance from

different types of plants. The difference in carbon isotope
signature between C3 plants (mainly trees and shrubs)
and C4 plants (mainly tropical grasses) allows the
reconstruction of palaeo-diet and thus the dominate
vegetation on the landscape. 73
Based on figure 4 in Maslin, M.A. and Thomas, E. 2003.
Balancing the deglacial global carbon budget: The hydrate
factor. Quaternary Science Reviews 22(15–17): 1729–36, with
permission from Elsevier.
24. Global temperatures through the Cenozoic. 81

25. Ocean gateways both today and during the Cretaceous. 83
xviii
l ist of figu r es
Based on figure 22 in Mark Maslin, M. 2013. Climate: A Very Short

26. Drying out, salt deposit formation, and reflooding of the

Mediterranean Sea approximately 5 million years ago. 87
27. Oxygen isotopes showing the step-like changes in global ice

volume over the last 5 million years. 91
28. Pacific Ocean during El Niño and normal periods. 95

29. Orbital eccentricity, tilt, and precession. 102

30. Long-term changes in the orbital parameters. 103
31. Insolation at 65°N compared with global sea level or

global ice volume. 106
32. East Africa, modern and palaeo lake location map. 115
Adapted from original unpublished files for figure 1 in Junginger,
A. and Trauth, M.H. 2013. Hydrological constraints of paleo-Lake
Suguta in the Northern Kenya Rift during the African Humid
xix
l ist of figu r es
Period (15–5 ka BP). Global and Planetary Change 111: 174–88, with
permission from Elsevier and thanks to the authors.
33. East Africa north–south cross section with major

modern and palaeo lakes shown. 118
Adapted from original unpublished files for figure 1 in
Junginger, A. and Trauth, M.H. 2013. Hydrological constraints of
paleo-Lake Suguta in the Northern Kenya Rift during the African
Humid Period (15–5 ka BP). Global and Planetary Change 111: 174–88,
with permission from Elsevier and thanks to the authors.
34. Occurrence of shallow and deep lakes over 5 million years

compared with the precessional orbital forcing. 120
CC-BY-SA-3.0 <https://creativecommons.org/licenses/by/3.0>.
35. Age range of different dating methods using in the

study of human evolution. 123
Adapted from figure 2 in Veldhuis, D., Kjærgaard, P.C., and
Maslin, M. 2014. Human evolution: theories and progress, in
Smith, C. (ed.) Encyclopedia of Global Archaeology, Springer,
New York, pp. 3520–32. With permission of Springer.
36. Different modes of potential speciation with and without

geographical or niche separation. 134
© Ilmari Karonen/Wikimedia Commons/Permission is granted to
copy, distribute and/or modify this document under the terms of
the GNU Free Documentation License, Version 1.3 or any later version
published by the Free Software Foundation; with no Invariant
Sections, no Front-Cover Texts, and no Back-Cover Texts. A copy
of the license is included in the section entitled ‘GNU Free
Documentation License’ <https://en.wikipedia.org/wiki/Wikipedia:
Text_of_the_GNU_Free_Documentation_License>.
37. Variability selection hypothesis of early human

evolution.135
xx
l ist of figu r es
Adapted from figure 2 in Potts, R. 1999. Variability selection in

hominid evolution. Evolutionary Anthropology, 1999: 81–96.
Permission of John Wiley and Sons. Evolutionary Anthropology:
Issues, News, and Reviews © Wiley Periodicals, Inc.
38. Pulse climate variability framework. 139

Adapted from figure 3 in Maslin, M.A., Shultz, S., and Trauth,
M. 2015. A synthesis of the theories and concepts of early human
evolution. Philosophical Transactions of the Royal Society B 370
<http://dx.doi.org/10.1098/rstb.2014.0064>, CC-BY version 4.0
<https://creativecommons.org/licenses/by/4.0/>.
39. Relationship between growth and shrinkage of large lakes in the

African Rift Valley. The bifurcation is caused by the evaporation
from the lake making the surround region wetter and thus
resilient to initial reductions of overall precipitation. 140
Adapted from figure 4 in Maslin, M.A., Shultz, S., and Trauth, M.
2015. A synthesis of the theories and concepts of early human
40. An interpretation of how the turnover pulse hypothesis could

be placed within the pulsed climate variability framework. 143
41. An interpretation of how the aridity hypothesis could

be placed within the pulsed climate variability framework. 144
Adapted from figure 6 in Maslin, M.A., Shultz, S., and Trauth, M. 2015.
A synthesis of the theories and concepts of early human evolution.
Philosophical Transactions of the Royal Society B 370 <http://dx.doi.org/
10.1098/rstb.2014.0064>, CC-BY version 4.0 <https://creativecommons.
org/licenses/by/4.0/>.
xxi
l ist of figu r es
42. An interpretation of how the allopatric speciation

hypothesis could be placed within the pulsed
climate variability framework. 145
43. Estimated community size of different hominin species

over the last four million years. Community size is estimated
from converting cranial volumes to neocortex ratios which
have a strong relation to group size in primates. 155
Based on figure 3.3 in Dunbar, R. 2014. Human Evolution: A Pelican
Introduction, Penguin, London, p. 74. Reproduced by permission of
Penguin Books Ltd.
44. Robin Dunbar’s circles of human relationships. 156

Adapted from original image by Robin Dunbar, with permission.
45. A ‘Midwife’s’ view of the birth canal in a chimpanzee,

A. afarensis, and a modern human. 158
Based on figure 2a in Gruss, L.T. and Schmitt, D. 2015. The
evolution of the human pelvis: changing adaptations to
bipedalism, obstetrics and thermoregulation. Philosophical
Transactions of the Royal Society B 370: 1663, adapted from
Rosenberg, K. and Trevathan, W. 2002. Birth, obstetrics and human
evolution. BJOG 109: 1199–206 (doi:10.1046/j.1471-0528.2002.00010.x)
and Tague, R.G. and Lovejoy, O. 1986. The obstetric pelvis of
A.L. 288–1 (Lucy). Journal of Human Evolution 15: 237–55 (doi:10.1016/
S0047-2484(86)80052-5). With permission from The Royal Society.
46. Expensive brain hypothesis. 161

M. 2015. A synthesis of the theories and concepts of early
human evolution. Philosophical Transactions of the Royal Society B
xxii
l ist of figu r es
370 <http://dx.doi.org/10.1098/rstb.2014.0064>, CC-BY version 4.0

47. Comparison of the great apes and humans relative

body size and sexual organs. 165
Based on ‘Males, as females see them’, and ‘Females, as males see
them’, in Diamond, J. 1992. The Rise and Fall of the Third Chimpanzee,
Vintage, London, pp. 60–1. Created using data from table 3.3 in
Dixson, A.F. 2012. Primate Sexuality, 2nd edition, Oxford University
Press, Oxford (based on data from Harvey, P.H., Martin, R.D., and
Clutton-Brock, T.H. 1987. Life histories in comparative perspective,
in Smuts et al., (eds) Primate Societies, University of Chicago Press,
Chicago pp. 181–96) and table 9.3. With permission from Oxford
University Press.
48. Comparison of Homo sapiens skull morphology

showing feminization that has occurred since the
Middle Stone Age. 170
Based on unpublished diagrams by Robert L. Cieri, with
permission.
49. Dispersal of H. erectus, H. Neanderthalensis, and H. sapiens.174

Based on <https://en.wikipedia.org/wiki/Early_human
_migrations> CC BY-SA 3.0 with additional data from
Liu, W. et al. 2015. The earliest unequivocally modern humans
in southern China. Nature 526: 696–9.
50. Relationship between Homo sapiens and Homo erectus

reconstructed by Chris Stringer, based on the evidence we
currently have from fossils and genetics. 176
Based on diagrams by Chris Stringer, with permission.
51. A history of interbreeding between early modern humans,

Denisovans, and Neanderthals. 180
Figure 1 reprinted by permission from Macmillan Publishers
Ltd: Callaway, E. Evidence mounts for interbreeding bonanza in
ancient human species. Nature [website] 17 February 2016
xxiii
l ist of figu r es
<http://www.nature.com/news/evidence-mounts-for-interbreeding-
bonanza-in-ancient-human-species-1.19394>.
52. Mass extinctions, land-use changes, and rising atmospheric

greenhouse gas levels over the last 50,000 years. 184
Adapted from figure 1 in Lewis, S.L. and Maslin, M.A. 2015.
A transparent framework for defining the Anthropocene Epoch.
The Anthropocene Review 2(2): 128–46, doi: 10.1177/2053019615588792,
which in turn was adapted, expanded, and scaled appropriately
from Ruddiman, W.F., Ellis, E.C., Kaplan, J.O., and Fuller, D.Q.
2015. Defining the epoch we live in. Science 348: 38–9.
53. Shipping routes plotted from eighteenth- and

nineteenth-century logs. 188
© Benjamin M. Schmidt.
xxiv
1
Introduction
-
H umans are rather weak when compared with many other

animals. We are not particular fast and have no natural
weapons. But we have become the world’s apex predator and have
taken over the planet. We Homo sapiens currently number nearly
7.5 billion and are set to rise to nearly 10 billion by the middle of
this century. We have influenced almost every part of the Earth
system and as a consequence are changing the global environ-
mental and evolutionary trajectory of the Earth. We are also both
inadvertently and on purpose changing our own evolution.
Fundamental to our success is that we are very smart, both indi-
vidually but more importantly collectively. But why did evolution
favour the brainy ape? Because let face it there are lots of draw-
backs to having a large brain. First every child tries to kill their
mother, as it is difficult and dangerous to giving birth to offspring
with such a large head. Second you need a large amount of food
in order to keep the organ running, as big brains are energetically
expensive. Third it frequently goes wrong. So the large brain must
have given our ancestors a major advantage.
1
in t roduc t ion
This book pieces together the evidence for human evolution by

focusing on the major evolutionary changes that occurred in
Africa that led our ancestors to walk upright and then to become
progressively smarter. Without the unique combination of tec-
tonics, global and local environmental changes, and celestial
mechanics human evolution would never have occurred. In this
introductory chapter I briefly review the origin of our planet to
put our evolution in the context of all evolution on Earth. I then
set out the ten transitions that have occurred on the way to
humans becoming the new geological superpower. What I hope
this book will show is that this human ascendancy was not inev-
itable but caused but a unique combination of factors. As the
great evolutionary theorist Stephen J. Gould suggested, if we
reran the tape of evolution again and again the chances of humans
evolving is zero.
In the beginning
To comprehend human existence and how we got here I believe

we need to put our evolution into context. To do this we need to
understand the origins of our planet and how life has evolved. So
let us start right at the beginning of everything. The Universe,
according to cosmologists, is 13.8 billion years old. It all started
with the Big Bang, when all the matter in the Universe was created
and blasted outwards. About 100,000 years after the creation of
matter, its expansion allowed it to cool, so that negatively charged
electrons were trapped by positively charged protons forming
hydrogen gas. At this point there were no galaxies, no stars, no
planets, and no life—only an expanding cloud of gas. Over time,
this irregular cloud started to form clumps of matter. Gravity
2
in t roduc t ion
pulled on the gases within these clumps to form galaxies. Gravity

within each galaxy pulled matter together to form billions and
billions of stars.
Deep within each star the immense pressure resulting from
gravity forces the hydrogen atoms to collide. These collisions occa-
sionally lead to the creation of helium through the fusion of two
hydrogen atoms and results in a huge release of energy. Stars thus
burn hydrogen through nuclear fusion. As a star’s fuel is converted
from hydrogen to helium, the helium starts to collide and its fusion
creates carbon and oxygen. The carbon then burns to create oxy-
gen, neon, sodium, and magnesium. This nuclear fusion or ‘cook-
ing’ continues through the elements up to and including iron.
These lighter elements are the main ones found in rocky planets
and are often called the planet builders. Elements heavier than iron
can only be created by neutron capture that involves adding energy.
This only occurs in the ejecta of massive star, ten to twenty-five
times the size of the Sun, when they finally explode. Large stars that
have massive gravitational force have the most intense nuclear fires
and are extremely bright but relatively short-lived. These stars
produce all the elements and distribute them throughout galaxies
through supernova explosions at the end of their cosmically short
life. So we are all made of stardust. Though these massive stars cre-
ate the building blocks for planets and life they themselves are not
suitable for forming a habitable solar system. Smaller stars, like our
Sun, have a smaller gravitational pressure and thus cook their
hydrogen at a much slower rate and provide a more stable long-
term environment to allow solar systems to develop. Our own Sun
has already existed for 4.6 billion years, in 5 billion years time it will
expand forming a Red Giant, and will continue expanding until it
runs out of fuel in about 7.5 billion years time.
3
in t roduc t ion
Our Solar System formed 4.568 billion years ago from the
gravitational collapse of a giant interstellar gas cloud. The vast
majority of the system’s mass is in the Sun, with most of the
remaining mass contained in Jupiter. There are four smaller inner
or terrestrial planets—Mercury, Venus, Earth, and Mars. The four
outer planets are giant planets. Then there is the asteroid belt,
made up of numerous irregular shaped bodies and minor planets.
The total mass of the asteroid belt is about 4 per cent that of the
Moon, of which half is made up by the four largest asteroids. The
two largest planets, Jupiter and Saturn, are gas giants, and are
composed mainly of hydrogen and helium. It is the position and
mass of Jupiter that has helped to create our relatively stable Solar
System. It also influences the changing shape of the Earth’s orbit
around the Sun, and this has a profound effect on the Earth’s cli-
mate. The two outermost planets, Uranus and Neptune, are ice
giants, and are made of water, ammonia, and methane. Further
out are rocky fragments, one of which is Pluto, which to my
youngest daughter’s dismay has been downgraded to a ‘dwarf
planet’ or planetoid.
Central to the stability of the Earth is the Moon. The Moon
formed about 4.53 billion years ago, some 30–50 million years
after the origin of the Solar System. The prevailing hypothesis is
that the Earth–Moon system formed as a result of a giant impact.
A Mars-sized planet, which has been named Theia, hit the
proto-Earth, increasing the mass of the Earth while blasting
material into orbit around the proto-Earth, which accreted to
form the Moon. The Moon is thus made of the lighter elements
found in the surface of the proto-Earth. So the Moon does not
have a metal core and its density is lighter than Earth’s. The
Moon acts as the Earth’s gyroscope, so its axis of rotation in
4
in t roduc t ion
relation to the Sun has remained very similar for the last 4.5 bil-
lion years. As we will see later, the small wobbles of the axis of
the rotation are enough to push the Earth into and out of the
‘great ice ages’. About 100 million years after the formation of
the Solar System the Earth degassed, creating an early atmos-
phere of nitrogen and carbon dioxide as well as significant
amounts of water. The Earth overall has very low amounts of
the essential volatiles such as water, carbon dioxide, and meth-
ane, which are essential for life. Only one molecule in 3 million
on Earth is water, but through these early processes they have
been concentrated at the surface. At about 4 billion years ago we
have the first evidence of rock formation on Earth (Figure 1).
H. sapiens
230–65 Ma Dinosuars 4540±40 Ma Formation of the Earth
225 Ma Mammals
~380 Ma
first vertebrate land
animals
~530 Ma
Cambrian explosion 65 Ma 4.6 Ga
Ma 4100–3800 Ma first
251
Cenozoic
Mes
a signs of life
M Had
Pa
first animals 2 ean Prokaryotes

ozoic
54
leo
3.8
4 Ga
zo
Ga
ic
~3500 Ma
1 Ga Photosynthesis
an
c he
Pro
Ar
te
multicellular 3 Ga
ro
oi
c
z
life
2 Ga
a
2.5 G
Eukaryotes
~2300 Ma
Rise of oxygen in atmosphere
figure 1 Major biological events in the 4.6 billion years history of the Earth.
5
in t roduc t ion
Then, at about 3.5 billion years, we have the first evidence of life
on Earth—and that is where our story really begins.
Ten key steps in human evolution
The essential requirements to form a habitable planet seem to be

a small long-lived star, a stable Solar System, and a planet at the
right distance from the star so it is not too hot and not too cold
and which also has enough water and carbon at the surface to
support life. Once these elements were established then I think
there are ten key transitions that occurred that inadvertently
ended up with humans populating and dominating the whole
planet. I summarize these below, and we will examine a number
of them in more detail later.
1. Origin of life
Fossil evidence for life can be found as far back as 3.46 billion
years ago in the form of preserved cellular structures in the
Strelley Pool Formation in Western Australia. These are thought
to be created by ‘single-celled’ bacterial communities that formed
stromatolites and are still found today in the warm shallow seas
in Shark Bay, Australia. Little seems to have happened during the
following so-called ‘boring’ billion years until about 2.4 billion
years ago, when a significant rise in atmospheric oxygen known
as the Great Oxidation Event is recorded in the rocks. Until then,
only simple, ‘prokaryotic’ organisms existed on Earth. The more
complex ‘eukaryotic’ cells, with nuclei and other internal struc-
tures, emerged between 2.1 and 1.6 billion years ago from the
merger of two types of prokaryotes, and these are the basis of
almost all multicellular organisms.
6
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2. Origin of complex life

The origin of animals is currently uncertain, but simple animals
had probably evolved by about 600 million years ago. However,
most of what were to become the major animal phyla we know
today appear to have arisen during a period of rapid diversifica-
tion known as the Cambrian Explosion, which began about 541
million years ago, the start of what is known as the Phanerozoic
Eon, and lasted for over 40 million years. Some of the extraordin-
ary forms that existed at this time are captured in the remark-
ably preserved fossil assemblages of the Burgess Shale (Rocky
Mountains, Canada) and Chengjiang (Yunnan Province, China).
This melting pot of evolution included one branch—the verte-
brates—that evolved a backbone, and it is from these that we are
descended.
3. Evolution of mammals
The unique features of mammals are milk-producing mammary
glands in females, fur or hair, three bones in the inner ear which
evolved from the reptilian jaw, and a neocortex (Figure 2). The
neocortex is a region of the brain that controls higher functions
such as sensory perception, generation of motor commands, spa-
tial reasoning, and, in humans, conscious thought and language.
The earliest mammals are thought to have emerged from mam-
mal-like reptiles before the end of the Triassic Period, by about
225 million years ago.
4. Extinction of the dinosaurs

There have been several mass extinctions in Earth history, the
most extreme of which was at the end of the Permian some
250 million years ago, which destroyed most of life on land and
7
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figure 2 Key morphological features that define hominids.
8
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in the oceans. This has been referred to as the ‘mother of all extinc-
tions’, but somehow life managed to survive and then diversify
after this close call. The mass extinction most directly relevant to
our story however is the Cretaceous–Tertiary extinction also
called the ‘KT’ extinction for short. Around 66 million years ago, a
major period of volcanism resulting in massive outpourings of
lava, resulting in the Deccan Traps in India, compounded by the
Chicxulub meteorite impact event in Central America, caused a
mass extinction that killed off many species, including the non-
avian dinosaurs. This extinction event was essential in the story of
human evolution as it ended the 170 million-year domination of
dinosaurs and allowed the evolution and proliferation of mam-
mals, and the appearance of the first ancestors of primates.
5. Evolution of social primates

Ten million years after the extinction of the dinosaurs, during a
period of super-global warming, the first fossil evidence for true pri-
mates and social monkeys begins to appear. Anthropoids or simians,
a group that includes primates and monkeys (Table 1), began to live in
table 1. Common terminology used in human evolution and in this book
Anthropoids All primates (monkeys and great apes and their fossil
ancestors), hominins, and humans
Hominids All great apes (gorillas, orangutans, chimpanzees, bonobos,
gibbons), hominins, and humans
Hominins All our fossil ancestors (Ardipithecus, Australopithecus, Homo)
Anatomically Homo sapiens, but without substantial evidence for our
modern humans cultural accoutrements (art, burials, ornament, musical
instruments)
Humans Only modern humans, Homo sapiens, with clear evidence of
cumulative culture
9
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larger groups, which meant that each animal had to negotiate com-
plex webs of friendships, hierarchies, and rivalries. This is thought to
have been a major driver of brain expansion later, in hominins.
6. Evolution of hominins
The exact origins of hominins is disputed. However, the last com-
mon ancestor with chimpanzees is thought to have lived between
8 and 5 million years ago. Despite the relatively miniscule genetic
difference between modern humans and chimpanzees, our evo-
lutionary paths have been radically different since the split.
7. Bipedalism
Between 10 and 5 million years ago, hominins developed the abil-
ity to move efficiently on two legs. At the same time chimpanzees
and gorillas became better at tree-climbing and developed
knuckle-walking while on the ground. Bipedalism allowed our
ancestors to spread out from East Africa into Northern and
Southern Africa. Some of these bipedal hominins were already
using stone tools by at least 3.3 million years ago.
8. Brain expansion
About 2 million years ago new hominin species appeared with
brains up to 80 per cent larger than their ancestors, and for the
first time they dispersed out of Africa. This large brain was accom-
panied by other sweeping changes to life history (shortened inter-
vals between births, delayed child development), body size, the
shape of the pelvis, and a shoulder morphology which allowed
projectile use. These species show adaptations to long-distance
running, ecological flexibility, and social behaviour, including
food processing.
10
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9. Cumulative culture
Homo sapiens emerged about 200,000 years ago in East Africa and
dispersed into Eurasia. It is not until 100,000 years ago that there
is evidence of creative thinking. From 50,000 years ago this
becomes more consistent, with evidence of art, ornaments, and sym
bolic behaviour. These steadily increase in complexity and fre-
quency and demonstrate that knowledge was being generated
and passed on to the next generation. Culture was being accumu-
lated and grew with every generation.
10. Agriculture and industrial revolutions

At the end of the last ice age, about 11,000 years ago, agriculture
first appeared in South West Asia, South America, and northern
China. It then appeared 6,000–7,000 years ago in southern China
and Central America and 4,000–5,000 years ago in the savannah
regions of Africa, India, South East Asia, and North America. Agri
culture steadily replaced hunter-gathering as the major economic
mainstay and urban centres and city-states developed. Humanity
then went through the industrial, technological, and now the
information revolution. Along the way the impact of humanity
became so great that we emerged as a geological superpower and
entered our own geological period: the ‘Anthropocene’.
The rest of the book is focused on understanding why many of
the later key transitions seem to occur in East Africa. We will
consider how the environment in which our ancestors lived
changed through time and how that may have affected their evo-
lution. Plate tectonics, the movement of the Earth within the
Solar System, and changes in global climate all appear to have
played a part in our early evolution. Central to this whole story is
the formation and development of the East African Rift Valley
11
in t roduc t ion
system, one of the most extensive geological features on the

Earth’s surface. It runs north–south for around 4,500 km from
Syria through East Africa down to Mozambique. In Chapter 2,
I examine the fossil evidence of our evolution, much of which has
been found within the East African Rift Valley. This landscape is
tough and unforgiving—but it appears to have been the cradle of
humanity.
12
2
Early Human Evolution

-
T o understand our evolutionary history it is essential that

we understand the fossil record. Discoveries in the past few
decades have dramatically expanded the available hominin fos-
sil record, with thirteen new species suggested and four new
genera named since 1987. This richer variety of fossils has pro-
duced two major changes in our knowledge. First, it has led to
a much greater understanding of the range of variation in the
hominin ‘phenotype’—i.e. the observable characteristics or
traits. This includes insights into populations, with fossils of
multiple individuals found at Atapuerca (northern Spain),
Dmanisi (Georgia), Hadar (Ethiopia), and Rising Star (South
Africa). Second, the extensive use of new dating techniques has
provided chronological precision to link those phenotypes to
the environments in which they evolved. Even with all the recent
finds the hominin fossil record is still very limited, with many
gaps. There is also considerable discussion among researchers
about defining these new species and genera, which influences
our understanding of changes in overall hominin diversity.
13
e a r ly hu m a n e volu t ion
But conflating or expanding the defined species seems at present

to have little overall influence on the diversity pattern.
The other key debate concerns where all the new hominin spe-
cies evolved. The date of first appearance of each species depends
heavily on what happened after death, on the conditions in which
the fossil was preserved, and on sampling bias. Nevertheless, the
consistency of hominin first-appearance dates in East Africa cur-
rently suggests that the majority of the new hominin species
evolved in East Africa and then dispersed outwards. This is sup-
ported by the present evidence for brain capacity, which suggests
that brain expansion occurred first in East Africa and only appeared
later elsewhere, following a dispersal event. Some researchers
suggest South African, European, or Asian origins for hominin
speciation, but although these are valid possibilities there is cur-
rently no fossil evidence to support them. Genetic evidence also
highlights the fact that there was a large amount of interbreeding
between hominin species. This has lead to a growing realization
that the classical view of the branching hominin evolutionary tree
is incorrect. Instead we should consider human evolution as more
of a bush with a network of closely related species that frequently
interbred, blurring the lines between species.
Five stages of human evolution
The fossil record suggests five main stages in hominin evolution:

(1) the appearance of the earliest (proto) hominins attributed to
the genera Sahelanthropus, Orrorin, and Ardipithecus between 4 and
7 million years ago; (2) the appearance of the Australopithecus genus
around 4 million years ago; (3) the appearance of the Homo genus
and the Paranthropus genus around the Plio–Pleistocene boundary
14
figure 3 Map of the location of the most important hominin fossil finds in Africa.
between 2.8–2.5 million years ago; (4) the appearance of Homo

erectus 1.8 million years ago; (5) the appearance of H. heidelbergensis
800,000 years ago, Homo sapiens 200,000 years ago, and modern
humans around 50,000 years ago. The taxonomic classification
of many specimens, as well as their role in the evolution of mod-
ern humans, is continually discussed, which is a polite way of
saying continually fought over and argued about both in public
15
and in the literature. What is not disputed is that, apart from

Sahelanthropus remains from Chad, all the earliest specimens for
each of the main genera have been found in the East African Rift
System.
Stage 1: Earliest hominins
The earliest potential candidate for the first hominin is Sahelanthropus

tchadensis—though this is disputed. The fossils were discovered in
the Djurab Desert of Chad by a team of four led by a Frenchman,
Alain Beauvilain, and three Chadians, Adoum Mahamat, Djim
doumalbaye Ahounta, and Gongdibé Fanoné, and are dated to
approximately 7–6 million years ago. When the discovery was
announced in 2002, palaeoanthropologists were amazed by both
the location and date. The skull fragments were found in a desert in
Central Africa, far to the west from the other early hominin sites
(Figure 3). The remains are limited to cranial fragments that suggest
a mosaic of hominin and non-hominin features and a brain size
equivalent to modern chimpanzees. What is striking about the
skull, nicknamed ‘Toumai’ or ‘hope of life’, is how modern-looking
it is, but with a thick brow ridge. The lack of other bones from the
skeleton makes it extremely difficult to reconstruct its lifestyle,
whether it was bipedal, or whether it was truly a hominin. However,
the possible position and orientation of the ‘foramen magnum’—
the hole in the base of the skull where the spinal cord passes
through—strongly suggests a posture similar to other early bipeds
(see Figure 4). The fossil was recovered from what is now a very dry
desert area in Chad. The few teeth recovered suggest Sahelanthropus
ate mainly fruit, which means Chad has undergone a large change
in climate since Sahelanthropus was living there.
16
figure 4 Location of the hole in the base of the skull through which the spinal
cord passes (foramen magnum) in humans and chimpanzee, showing the central
position essential for bipedal movement.
17
The next putative hominin is Orrorin tugenensis, which means

‘original man’ in the local language of Tugen Hills in western
Kenya where it was found by a team led by Brigitte Senut and
Martin Pickford. The volcanic deposits above and below the fossil
have been accurately dated, and Orrorin lived about 5.8 million
years ago. The taxonomic position, lifestyle, and locomotion of
Orrorin are all disputed due to the fragmentary nature of the spe-
cimens. Unusually this species is known for other parts of its skel-
eton rather than its skull. There are fragments of jaws and teeth,
but the main evidence comes from two thighbones, which have
preserved the upper end that is connected to the hip. This is the
first direct evidence for a hominin walking upright on two legs.
Walking upright requires substantial changes to the hip joint,
including ligaments to stabilize the joint running between the
pelvis and the femur. The marks left by these ligaments can be
seen in modern humans and have been observed on the Orrorin
femurs. The structure of human femurs and those of other apes is
also different because when humans walk all the weight of the
body is alternatively held by the left and right hip joints. Nearly all
that stress is placed on the bottom of the femur neck, the narrow
section below the ball—so the bone is much denser in humans at
this section than in apes that use knuckle-walking to move
around (Figure 5). This is where the disputes about Orrorin bipedal-
ism arise, as this increased density of the femur bone is very clear
in later species such as Australopithecus afarensis, but less obvious in
the Orrorin femurs. Orrorin teeth are much more ape-like, though
the teeth have thick enamel like humans. Hence, both
Sahelanthropus and Orrorin have been assigned to hominins by
their discoverers based on very different evidence—they may or
may not be hominins, and even if they were hominins there is no
18
figure 5 Evidence of bipedalism from the shape of the neck of the femur
(thighbone).
way to tell if they were part of our ancestry. Both Sahelanthropus

and Orrorin have been suggested as members of a clade that
includes Ardipithecus—in other words, all three may have had a
common ancestor which we are yet to find.
The oldest member of the Ardipithecus genus is Ardipithecus kad-
abba, whose fossil evidence consists only of fragmentary teeth
and skeletal remains dated to approximately 5.6 million years ago.
A much more extensive fossil record exists for the second mem-
ber of the genus, Ardipithecus ramidus. The remains are from a
female, nicknamed ‘Ardi’, and includes most of the skull and teeth,
as well as the pelvis, hands, and feet. It was discovered by the pal-
aeoanthropologist Yohannes Haile-Selassie, who was part of a
team led by Tim White, in Ethiopia’s harsh Afar desert, at a site
called Aramis in the Middle Awash region. The name Ardipithecus
ramidus stems mostly from the Afar language in which Ardi means
‘ground or floor’ (borrowed from either Amharic or Arabic) and
ramid means ‘root’. The -pithecus portion of the name is from the
Greek word for ‘ape’. The bones and teeth of nine other A. ramidus
19
individuals have been discovered at As Duma in the Gona Western

Margin of Ethiopia’s Afar region, dated to between 4.35 and
4.45 million years old.
Ardipithecus had a small brain, measuring between 300 and 350 cm3.
This is slightly smaller than a modern bonobo or a female com-
mon chimpanzee brain, and much smaller than the brain of
Australopithecines like Lucy (~400 to 550 cm3). It is roughly 20
per cent the size of the modern human brain. Ardipithecus teeth
indicate a highly omnivorous diet and their post-crania suggest
a lifestyle of arboreality (tree dwelling) coupled with primitive
bipedality. The fauna and vegetation associated with the A. ramidus
specimens in the Awash Valley date to around 4.4 million years
ago, suggesting a woodland-forest matrix habitat associated with
significant rainfall and water availability. This appearance of
bipedality in closed woodland environments undermines theor-
ies of bipedality evolving exclusively as an adaptation to open
habitats. As we shall see, a number of researchers think that
bipedalism may have evolved many times in the past because it
proved useful for survival.
Stage 2: Appearance of Australopithecus
Australopithecus was first discovered in the 1920s and changed our

understanding of human evolution. Because the Australopithecus
species were small-brained, bipedal ancestors, this showed us that
the defining feature of hominins was walking upright, not larger
brains. Australopithecus means ‘southern ape’ and was coined by
Raymond Dart when he discovered A. africanus in South Africa in
1924. When other similar fossils were found in East Africa the
name was kept due to the close taxonomic similarity.
20
The first members of the Australopithecus genus, attributed to

A. anamensis, were found at two sites near Lake Turkana, Kanapo
and Allia Bay, and range in age from 3.8 to 4.2 million years ago.
These individuals show strong evidence of bipedality combined
with primitive cranial features. The species name anamensis comes
from the Turkana word ‘anam’, meaning ‘lake’. The sample of
A. anamensis fossils is small, but a well-preserved tibia (shin) bone
is very similar to a human tibia, suggesting bipedal locomotion.
A. anamensis is followed by A. afarensis, which is very well known
from the fossil record and includes the remarkably complete
‘Lucy’ specimen. Lucy was found by a team led by Donald Johan
son and Tom Grey on 24 November 1974 at Hadar in Ethiopia.
The story goes that they had taken a Land Rover out that day to
map another locality. After an unsuccessful morning survey-
ing for fossils, they decided to head back to the vehicle. Johanson
suggested taking an alternate route back to the Land Rover,
through a nearby gully. Within moments he spotted a right prox-
imal ulna (forearm bone) and identified it as a hominin bone. Over
the next two weeks they recovered several hundred fragments of
bone, representing 40 per cent of a single hominin skeleton—
‘Lucy’. How ‘Lucy’ got her name has gone down in folklore. On
the night of 24 November there was much celebration and excite-
ment over the discovery, with drinking, dancing, and singing,
mainly to the Beatles’ song Lucy in the Sky With Diamonds,
which was played over and over again. At some point during that
night, and no one has owned up to it, someone named the skel-
eton Lucy, and it has stuck. Our understanding of A. afarensis now
goes way beyond Lucy, with another two complete skulls and a
large collection of jaws and teeth, as well as a comprehensive col-
lection of limb bones.
21
The species name afarensis means ‘of the Afar region’, where the
fossil was initially found, and it has been dated to between 3.6 and
3 million years ago. A. afarensis still retains a small brain size, yet
the post-cranial morphology is more similar to modern humans
than to apes and suggests a lifestyle strongly adapted to long-
distance walking. It seems to have been a highly successful spe-
cies, and specimens have been found in Chad, Ethiopia, Kenya,
and Tanzania. The jawbone found in Chad has been assigned the
species name of Australopithecus bahrelghazali, partly to recognize a
unique location and some different characteristics, but mainly as
researchers desperately want to find new species and to name
them. The Australopithecus genus may be wider still because in
2001 on the western side of Lake Turkana, Meave Leakey found
what they named Kenyanthropus platyops. This skull was dated to
3.5 million years ago, and though similar to A. afarensis it has a
very flat face—platyops means ‘flat face’, and is derived from two
Greek words: platus, which means ‘flat’, and opsis, which means
‘face’. Usually a flat face is associated with strong muscles for
chewing, but there is no other evidence for this—for example, the
teeth are small. Much of the rest of the skull’s features are primi-
tive and very similar to A. afarensis, so other researchers now refer
to the species as Australopithecus platyops. Recent reanalysis of all
the data has, however, convinced Meave Leakey and her col-
leagues that this is a separate species, and I will use Kenyanthropus
platyops in this book.
The fossil evidence clearly indicates that A. afarensis was fully
bipedal, and not quadrupedal like most monkeys and chimpan-
zees, nor suspensory like orangutans or gibbons. It did not run
along the ground on all fours, or have long arms for climbing and
swinging. This terrestrial bipedalism is shown by the beautiful
22
and haunting fossilized footprint trail found in Laetoli in Tanzania,

dated at 3.5 million years ago. But it is also clear that A. afarensis
walked in a slightly different way to modern humans, as the
blades on their hip bones flare out further to the sides. In sum-
mary, A. afarensis was a similar size to a modern chimpanzee, and
was bipedal on the ground, but its relatively long arms and feet
and the curved bones of its feet and fingers would have been used
to climb trees for food, safety, and sleeping.
Recent finds in the Afar region of Ethiopia suggest that A. afarensis
may in fact be two species. In 2011, three fossilized jawbones were
found 35 km north of Hadar, where Lucy was found, in sediment
dated to between 3.5 and 3.3 million years ago. Though the new
fossils are similar to the Lucy A. afarensis specimen, the teeth have
different root structures and are smaller. The cheekbones seem to
be more forward-facing, the lower jaw is larger, and the teeth have
thicker outer enamel than A. afarensis, suggesting adaptation to
harder, tougher, and more abrasive food than A. afarensis. Yohannes
Haile-Selassie, the curator of physical anthropology at the Cleveland
Museum of Natural History, suggests these fossils may represent a
new species—Australopithecus deyiremeda. The name ‘deyiremeda’
means ‘close relative’ in the local Afar language. If these fossils
represent a new species, then some fossils identified as A. afarensis
may belong to A. deyiremeda. But the naming and identification of
A. deyiremeda as a new species is still controversial and requires a
lot more evidence.
A. afarensis may hold another huge surprise for us. In 2011 and
2012 the West Turkana Archaeological Project found the oldest
known stone tools, dated at 3.3 million years, on the western shore
of Lake Turkana in northern Kenya. These tools predate by a mil-
lion years the oldest fossil specimens attributed to Homo in West
23
Turkana at 2.35 million years ago, and by half a million years the
earliest Homo find from Ledi-Geraru, Afar, Ethiopia, at 2.8 million
years ago. This is revolutionary, as it has always been assumed
that the evolution of Homo was associated with the start of stone
tool-making. The only hominin species known to have been liv-
ing in the West Turkana region at a similar time is K. platyops at
about 3.5 million years ago, while in the Lower Awash Valley in
Ethiopia, A. afarensis is found at 3.39 million years ago. The signifi-
cance of these fossils is that they were found in association with
cut-marked bones indicating butchering. Bone from both impala
and buffalo-sized creatures were found to bear cut marks. Both
the stone tools and the cut-marked bones suggest that A. afarensis
or K. platyops were not only making and using tools, but also ven-
turing out of the safety of the forests and on to the plains in search
of meat.
Australopithecus africanus, which simply means ‘southern ape of
Africa’, is the oldest hominin found in South Africa (Figure 3)
with the first appearance dated to 3.3 million years ago. It is simi-
lar to A. afarensis but with more ape-like limb proportions yet less
primitive teeth. The longer femur in A. afarensis, as compared to
A. africanus, suggests a longer stride and more efficient walking
style. There are now hundreds of fossils associated with A. africanus,
largely from the cave sites of Sterkfontein and Makapansgat in
South Africa. Sterkfontein is close to Johannesburg, and is part
of a set of fossil sites named the Cradle of Humankind World
Heritage Site. A. africanus seems to have had a varied diet and the
proportions of carbon isotopes in their teeth has shown that their
diet may have contained either grass, grass seeds, or animals
whose primary food was grass. A. africanus teeth and jaws seem to
have no adaptations to eating grass or grass seeds. Given what we
24
have recently found out about A. afarensis and stone tool use, we
can speculate that A. africanus was also eating a significant amount
of meat. Though we know that chimpanzees hunt meat, it makes
up less than 5 per cent of their diet, but the carbon isotope evi-
dence suggests A. africanus was eating a lot more than 5 per cent
meat. But here we have a puzzle, as A. africanus teeth are also not
adapted to slice meat effectively.
The final australopithecine is A. garhi, associated with 2.5 million-
year-old deposits in the Awash Valley, Ethiopia (Figure 3). This is a
confusing set of fossils as it consists of a few skull fragments and
very large teeth; then, at a nearby site, a partial skeleton has been
found with no skull fragments; and, at another nearby site, there
are animal bones with evidence of cut marks. If these fossils do
represent one individual then it was a long legged creature with
long forearms that may have cut the meat off bones of animals.
Stage 3: Homo and Paranthropus
In 1964, Louis Leakey and his colleagues announced the discovery

of Homo habilis based on fossils found in the Olduvai Gorge in
Tanzania (Figure 3). They described it as bipedal, a tool-maker,
and with a larger brain than Australopithecus. The name Homo habilis
means ‘skilful man’ (sometimes incorrectly translated as ‘handy man’).
H. habilis has been found throughout Eastern and Southern Africa.
The age range for H. habilis is from 2.35 to 1.5 million years ago. In
2013, however, a fragment of a fossilized jawbone was found in the
Ledi-Geraru research area, Afar, Ethiopia, by an Ethiopian student,
Chalachew Seyoum. The fossil has been dated to 2.8 million years
ago and is considered the earliest evidence of the Homo genus
known to date. The new fossil seems to be intermediate between
25
Australopithecus and H. habilis. This discovery pushes back the ori-

gin of Homo by over 400,000 years.
H. habilis is a controversial species with much scholarly debate
regarding its placement in the genus Homo rather than the genus
Australopithecus. The small size and rather primitive attributes
have led some experts, such as palaeoanthropologists Bernard
Wood and Mark Collard, to propose excluding H. habilis from the
genus Homo and placing it instead in Australopithecus as Australopith
ecus habilis. This is because, in its appearance and morphology,
H. habilis is the least similar to modern humans of all species in
the genus Homo (except the equally controversial H. rudolfensis). H.
habilis was short and had disproportionately long arms compared
to modern humans; however, it had a less protruding face than
the Australopithecines from which it may have descended. H. habilis
had a cranial capacity of between 550 cm3 to 687 cm3, though a recent
study led by anatomist and palaeoanthropologist Fred Spoor
at University College London suggests one key specimen had
an endocranial volume of between 729 and 824 cm3, larger than
any previously published value. So, compared to Australopithecines,
H. habilis’ brain capacity was on average 50 per cent larger, but it
was still considerably smaller than the 1,350 to 1,450 cm3 range of
modern Homo sapiens. But what had until recently made H. habilis
special was that, despite the ape-like morphology of their bodies,
their remains were often accompanied by primitive stone tools at
locations such as Olduvai Gorge in Tanzania and Lake Turkana in
Kenya. The discovery of stone tools prior to the first appearance
of Homo raises the question again of whether Homo habilis should
be considered a member of the genus Homo.
The definition of H. habilis suffers from another problem: as
more and more specimens were classified as H. habilis there
26
seemed to be too much physical variation to occur in one species.

For example, many of the small specimens had more prominent
brow ridges—usually found in larger males in modern species of
apes. To get round this problem a new controversial species,
Homo rudolfensis, was proposed to encompass the larger individual
specimens with bigger brains and a slightly different facial struc-
ture and chewing mechanism. The species is named after Lake
Rudolf, the former name for Lake Turkana, in Kenya, and speci-
mens have been found that are between 2 and 2.5 million years
old. Some researchers warn, however, that we cannot call Homo
rudolfensis a bigger version of H. habilis as there are distinct differ-
ences. Homo rudolfensis has a much flatter face from side to side
and its cheekbones are further forward, which seems to echo fea-
tures of Australopithecines despite the more modern-looking face.
One theory is that these two species were eating different food
sources, which explains why they were able to live in the same
region without competing for resources.
The situation has become even more complicated with a new
synthesis of the early Homo fossils published by Susan Anton
(New York University), Richard Potts (Smithsonian Institution),
and Leslie Aiello (Wenner-Gren Foundation) in Science in 2014.
They contend that morphologically there are three distinct groups
instead of two (Figure 6). They define early Homo, which includes
the earliest H. habilis, starting at 2.35 million years ago; but now,
with the latest find, this should be extended to 2.8 million years
ago. Then they define two different groups, both appearing at
about 2.09 million years ago. One with a relatively tall, flat face; it
appears in the fossil record between 2.09 and 1.78 million years,
and the second with a more primitive face and appears in the fos-
sil record between 2.09 and 1.44 million years ago. What this new
27
figure 6 New divisions at the start of the Homo lineage suggested by Susan
Anton and colleagues based on skull morphology, which challenges the original
H. habilis, H. rudolfensis, early Homo split. Figure includes the new stone tools (3.3 Ma)
and early Homo (2.8 Ma) finds.
synthesis shows is that hominin evolution, rather than being a

simple linear transition between clearly defined species, is much
more of an evolutionary bush with lots of slightly different mor-
phologically defined species coexisting. We can speculate that
these very closely related species were interbreeding, hence why
their morphologies are very similar.
28
Our understanding of early Homo may be on the cusp of

another revolution. In September 2015, a new species, Homo naledi,
was announced. It is named after the Rising Star cave where the
fossils were first found; this is some thirty miles north-west of
Johannesburg in South Africa (Figure 3), and ‘naledi’ means ‘star’
in Sesotho, a local South African language. So far, the team has
recovered parts of at least fifteen individuals of the same species,
a small fraction of the thousands of fossils believed to remain in
the chamber. The team, led by palaeoanthropologist Lee Berger at
the University of the Witwatersrand in South Africa, suggests
that naledi looks like one of the most primitive members of the
Homo genus. H. naledi had a tiny brain, about 500 cm3, similar to
the Australopithecines and smaller than H. habilis, perched atop a
very slender body. H. naledi stood approximately 1.5 m (about 5 feet)
tall and weighed about 45 kg. H. naledi’s teeth and skull are similar
to Homo habilis. The shoulders, however, are more similar to those
of apes. In addition, H. naledi had extremely curved fingers—more
curved than almost any other species of early hominin—which
clearly demonstrates climbing capabilities. This contrasts with
the feet of H. naledi, which are virtually indistinguishable from those
of modern humans. Combined with its long legs, this suggests the
species was well-suited for long-distance walking. Like any new find,
H. naledi is very controversial. Some palaeoanthropologists have
suggested that it is in fact a small-brained Homo erectus, while oth-
ers dispute whether it is in the Homo genus. Regarding the later
dispute, the discoverers did not publish their results in the highest
impact journals, which is the normal route. Instead they pub-
lished in a less well-known open access journal which they sug-
gest was to enable everyone to have access the data; others have
suggested more cynically this was a way to get the genus name
29
Homo accepted in the literature. We currently do not have a date

for this find, and given its location it is unlikely we will ever have
a good date for the fossil. This means we cannot fit it into our jig-
saw of human evolution. This is a significant problem because if
the find is relatively old then it could be a new branch in the hom-
inin tree; however, if it is relatively young, it may just be a dwarf
species of an earlier lineage similar to H. floresiensis, which is
discussed later.
At the same time as early Homo was appearing in its various
forms, another completely different group of hominins was
evolving—the Paranthropus group. These hominins had robust
dentition and large jaw muscles, and appeared around 2.5 million
years ago. They are fascinating as they show us there was a radic-
ally different way of being a hominin. The Paranthropus genus
includes the East African P. aethiopicus (2.5 million years ago),
P. boisei (2.3–1.2 million years ago), and the Southern African
P. robustus (1.8–1.2 million years ago). All the Paranthropus species
are thought to have been bipedal, and they had robust skulls to
support the large jaws and strong chewing muscles. This adapta-
tion of the skull is thought to have enabled Paranthropus to eat
hard foods such as nuts, seeds, and tubers. This interpretation is
supported by evidence of extreme wear on the enamel of the
teeth that have been found. Intriguingly, these species are often
found at the same sites as our closer relatives, and it is very likely
that if you had visited South Africa between 2 and 1.5 million
years ago you could have seen two very different bipeds—one
looking vaguely familiar using tools to obtain meat and veget-
ables, and the other, P. robustus, moving across the landscape con-
suming mainly vegetables, particularly hard nuts and edible roots.
Another significant feature of the Paranthropus genus is the large
30
difference between the size of male and females—an example of

what is called sexual dimorphism. P. boisei and P. robustus males
were about twice the size of the females, which is the size diffe-
rence seen today in gorillas. This ratio is much larger than early
Homo or modern humans, where males tend to be only slightly
larger than females. The fossils of P. robustus found at Swartkrans
in South Africa also give us a stark reminder of the dangers faced
by hominins. Many of the bones show evidence that the individ-
uals were killed by predators. The best example is the skull of a
young hominin that has bite marks from a leopard, which had
jumped on its back and bitten down on the skull, puncturing
both the back of the skull and through the eye sockets.
Stage 4: Homo erectus
Arguably the most important episode in hominin evolution

occurred in East Africa around 1.9–1.8 million years ago, when
hominin diversity reached its highest level, with species of the
Australopithecus, Paranthropus, and Homo genera all coexisting along
side each other. At the same time, the most important leap for-
ward in human evolution appears to have occurred with the
appearance of Homo erectus (understood in the broadest sense),
which is associated with sweeping changes in brain size, life his-
tory, and body size and shape. It is also the first species of homi-
nin we know of that migrated out of the Rift Valley and into Eurasia.
I say Homo erectus as understood in the broadest sense because
African specimens, and sometimes those from Dmanisi in
Georgia, are described as Homo ergaster. The name ‘ergaster’ means
‘workman’ in Greek and was used as the African specimens have
all been found with stone tools. Moreover, H. ergaster does not
31
have all the features that are found in Asian populations of H. erectus.
To help the reader I will from now on refer to this group of spe-
cies simply as H. erectus and hope my palaeoanthropologist col-
leagues can forgive me. Post-cranially, H. erectus is very similar to
modern humans. Its brain size was about two-thirds the size of a
modern human which is much larger than earlier hominins. Early
representatives of Homo erectus in Africa had a brain that was over
80 per cent larger than Australopithecus afarensis and 40 per cent
larger than Homo habilis. In contrast, from the appearance of the
early Australopithecines until the appearance of the first member of
the genus Homo, there was remarkably little change in hominin
brain size (Figure 7). In fact, many of us would argue that the
Homo genus should really start with Homo erectus, given its simi-
larity to modern humans and the significant differences with
earlier Homo specimens.
For me, the emergence of H. erectus in East Africa represents a
fundamental turning point in hominin evolution. The dramatic
increase in brain size was also accompanied by major changes in
life history and body morphology. First the pelvis morphology
changed to allow the birth of a larger head associated with a big-
ger brain. There is also beautiful evidence from growth lines in
the fossilized teeth that Homo erectus was the first hominin to have
a delayed growth period during childhood. Modern human chil-
dren grow very differently to the offspring of other apes. A male
chimpanzee and male human both end up with the same body
weight but they have quite different growth patterns. At age one
the human child weighs twice that of the chimp, but at age eight
the chimpanzee is twice the weight of the human. The chimpan-
zee gains its adult weight at age twelve—a whole six years before
the human. A male gorilla is also a fast and consistent grower; a
32
figure 7 Records of hominin brain capacity and East African hominin diversity over the last five million years.
150 kg male gorilla weighs 50 kg by its fifth birthday and 120 kg

by its tenth birthday. This is because humans have a growth pause
or plateau, extending childhood and delaying full adult develop-
ment, and this pattern began with H. erectus. The body size of
H. erectus is larger than that of earlier hominins and comparable to
modern humans. This is well illustrated by the H. erectus skeleton
known as the Nariokotome Boy, found in 1984 by a team led by
Richard Leakey and Alan Walker at Lake Turkana. Hip analysis
shows that he was a boy, and teeth growth lines suggest he was
twelve years old when he died—but he was already 160 cm (5 foot
3 inches) tall. Dan Lieberman and colleagues at Harvard University
have shown that H. erectus had many key adaptations required for
long-distance running. More recently it has also been shown that
the shape of the shoulder in H. erectus would have allowed the
throwing of projectiles. H. erectus also produced a much more
sophisticated set of stone tools than previous hominins, referred to
as Acheulean tools. It has also been argued that H. erectus had learnt
to control fire because, as Richard Wrangham at Harvard University
eloquently argues, it is difficult to see how they could have main-
tained such a large, energy-intensive brain with such a small gut
without access to cooked meat. However, this runs contrary to
the prevailing view of most palaeoanthropologists that the con-
trol of fire and regular cooking came much later in human evolu-
tion. But if this is correct how did H. erectus get enough calories to
fuel its bigger brain? Katherine Zink, also at Harvard University,
and Daniel Lieberman may have found a solution. They experi-
mented with processing food with stone tools that would have
been available to H. erectus. By slicing meat and pounding root
vegetables and nuts they were able to improve the ability to chew
by at least 40 per cent. There was also a decrease in the masticatory
34
force needed, which corresponds to the observed reduction in

jaw size and strength in H. erectus. The control of fire and regular
cooking was, as Rick Potts at the Smithsonian suggests, essential
for the next significant increase in brain size between 600,000
and 700,000 years ago with the appearance of H. heidelbergensis
and then Neanderthals and H. sapiens.
So H. erectus was smart, with a large and flexible stone toolkit,
able to run long distances, throw projectile weapons, and process
food. If we are looking for the most successful hominin species in
history then the answer would have to be H. erectus, as it sur-
vived from 1.8 million years ago to 200,000 years ago and colon-
ized much of Africa and Eurasia. In fact, H. erectus may have
survived even longer. In 2003, partial skeletons of nine individual
hominins were recovered from a cave on the island of Flores in
Indonesia. These individuals stood about 1.1 m tall (3.5 feet). Not
only were they small but they also had a correspondingly small
brain, about 380 cm3, somewhere between chimpanzees and
Australopithecines. The new species was called Homo floresiensis or
Flores Man, and nicknamed ‘hobbit’ or ‘Flo’. The earliest evidence
for H. floresiensis are a few teeth and a mandible found at Mata
Menge in the Soa Basin and dated to 700,000 years ago. The par-
tial skeletons found at Liang Bua are much younger with strati-
graphic evidence from the cave and dates of associated stone
artefacts suggesting an age of between 100,000 and 50,000 years
ago. One suggestion is that, due to the similarity in morphology, H.
floresiensis is in fact a population of H. erectus that had undergone
island dwarfism: species that colonize isolated islands are often
found to decrease in size over time. Intriguingly, stories abound
that H. floresiensis may have survived after 50,000 years ago in
other parts of Flores, as the Nage people of the island tell stories
35
of the Ebu Gogo—small, hairy, language-poor cave dwellers, which

seem to be very similar to H. floresiensis. It has been suggested that
the Ebu Gogo were present at the time of the arrival of the first
Portuguese ships during the sixteenth century, and there are even
claims they existed as recently as the late nineteenth century. But
then again other cultures around the world have their own stories
of similar human-like monsters hiding in the forests or the moun-
tains such as trolls, dwarfs, Pukwudgie, Bigfoot, Yeren, and the
Yeti.
Stage 5: The journey towards Homo sapiens
With the appearance of H. erectus, brain size increased signifi-

cantly and continued to increase over the following half a million
years (Figure 7). Then, a million years after H. erectus first occurs in
the fossil record, a new species, Homo heidelbergensis, appears, with
the first-known fossil found in Ethiopia dated to 600,000–700,000
years ago. The name Homo heidelbergensis has been around a long
time, as it was first coined for a jawbone found near Heidelberg,
Germany, in 1907. The present concept of the species H. heidelber-
gensis is, however, new and it is understood to represent a des-
cendant of H. erectus in Africa which spread out into Europe.
H. heidelbergensis may even have been the common ancestor of
Neanderthals and Homo sapiens. H. heidelbergensis specimens are
recognizable from very human-like but massive robust skulls.
Brain capacity ranges from 1,100–1,400 cm3, which is three times
larger than an Australopithecus brain and, at the top end, overlaps
the range of modern humans. Early fossils of H. heidelbergensis
found in Greece, Spain, Ethiopia, and Zambia look very similar,
and this is why the species is thought of as both an African and
36
European species. Later European populations begin to look dif-

ferent, and this is why it is thought they may have evolved into
Neanderthals. H. heidelbergensis even made it to Britain—a popu-
lation which the palaeoanthropologist Chris Stringer lovingly
refers to as ‘Homo Britannicus’.
As you have probably become aware through reading this
chapter, nothing is simple when it comes to naming hominin fos-
sils, and there are two other species names you will come across.
The first is Homo rhodesiensis, the original species name given to
the Zambia specimen which most now consider a clear example
of H. heidelbergensis. The other is Homo antecessor, which is based on
the oldest known hominin fossils in Europe found in the moun-
tainous northern Spanish site of Atapuerca. This amazing site is
famous for its incredible collection of H. heidelbergensis fossils
dated to more than 400,000 years ago. But much older hominin
fossils have also been found there, dating from 800,000 to 1.2
million years ago. Eighty bones from six individuals have been
found, and because of the differences with H. heidelbergensis they
have been given the species name H. antecessor. H. antecessor was
about 1.6–1.8 m (5.5–6 feet) tall and their brain sizes were roughly
1,000–1,150 cm3, smaller than most H. heidelbergensis fossils.
Between 700,000 and 300,000 years ago there is evidence for
only one hominin species in Africa and Europe, H. heidelbergensis.
Fossilized wooden spears that are 400,000 years old have been
found in Germany, and 500,000-year-old hafted stone points
used for hunting have been discovered in South Africa, suggest-
ing that H. heidelbergensis was an active hunter. H. heidelbergensis,
like H. erectus before it, migrated out of Africa into both Europe
and Asia. Though the focus of this book is on our African ori-
gins it is worth noting that it is currently thought that European
37
populations of H. heidelbergensis gave rise to Homo neanderthalensis

around 400,000 years ago; Asian H. heidelbergensis were ances-
tral to Homo denisovan around 600,000 years ago; and African
H. heidelbergensis were the ancestors of H. sapiens around 200,000
years ago.
The last step on the road to modern humans is the appearance
of Homo sapiens. The earliest specimens found in the fossil record
are from Omo in south-west Ethiopia, dated to about 195,000
years ago. It is generally accepted that H. sapiens evolved in East
Africa and then dispersed out of Africa in a similar way to H. erectus
and H. heidelbergensis. But the intriguing point is that, for about
150,000 years, H. sapiens behaved in a very similar way to its
ancestors. It is only after about 100,000 years ago that there is
evidence for abstract thought and the use of symbolism to express
cultural creativity. From 50,000 years ago we find spectacular
cave paintings in France, Spain, and Indonesia and ‘Venus’ figur-
ines in Austria, France, and the Czech Republic. In Chapter 8,
I examine what kind of revolution may have occurred 50,000 years
ago to set in train the cumulative culture which continues today
and is one of the defining features of modern humans. Some
researchers have suggested that we should perhaps use subspe-
cies to define these changes. So Homo sapiens idaltu has been sug-
gested for the more robust primitive form of H. sapiens found at
sites including Herto in Ethiopia (dated at 160,000 years old) and
Skhul in Israel (dated at 90,000 years old), while the terms Homo
sapiens sapiens, or anatomically modern humans (AMH), or ana-
tomically modern Homo sapiens (AMHS), should be used for the
more gracile, less robust, and almost completely modern-looking
fossils found from the Upper Palaeolithic period (~50,000 years ago)
onwards. In this book I stick to the convention of H. sapiens and
38
figure 8 Summary of the hominin species covering the last five million years and major dispersal events. The dotted line shows the first
dispersal from East Africa into Southern Africa and the solid lines represents dispersals of hominins out of Africa.
use the term modern humans for those H. sapiens that appeared
after 50,000 years ago.
I hope this chapter has provided an adequate overview or
framework of human evolution over the past 7 million years.
With such a small and fragmented fossil record of hominins
available, there are, as I have tried to indicate, many different
opinions and controversies. Figure 8 summarizes what we cur-
rently know about the first and last appearance of the different
hominin species mentioned here.
For me, early human evolution can be simplified into three
stages. The first is the evolution and success of bipedalism, with
the spread of the Australopithecus species across the African con-
tinent. Second is the evolution of Homo erectus, with a significant
jump in brain size and body morphology to become close to that
of modern humans. Third is the appearance of Homo sapiens and
the acceleration of cumulative culture 50,000 years ago. Now,
with this framework in place, in the rest of the book we will
explore how plate tectonics, global and local climate change, and
celestial mechanics may have influenced the appearance and dis-
appearance of these different hominin species. What were the
wider environmental factors that drove the evolution of humans,
and why did they arise in East Africa?
40
3
Tectonics and Climate

-
T o understand long-term changes in global climate and their

effects on evolution we need to understand the underlying
driving force of tectonics. The Earth’s climate is simply a function of
how the Sun’s energy falling on the Earth is redistributed around
the globe. This heat redistribution is influenced by the position of
the continents and the oceans, which is controlled by plate tec-
tonics. This is why 100 million years ago the Earth was much
warmer and more humid than today and dinosaurs were happily
living in Antarctica.
Tectonics has two main effects on climate. First there are the dir-
ect effects, which include the creation of mountains and plateaux,
which block atmospheric circulation and change the hydrological
cycle. They also include the movement of continents that changes
the shape of the oceans, creating ocean gateways. Second there
are indirect effects, which regulate atmospheric greenhouse gases
through subduction, volcanism, and the chemical weathering of
rocks. Understanding how tectonics influences climate is essen-
tial if we are to understand human evolution, because without
41
t ec tonics a nd cl im at e
the uplift of the Tibetan Plateau and the formation of the East
African Rift Valley hominin evolution would not have occurred.
In this chapter I break down the influences into horizontal tec-
tonics, which looks at what happens if you simply move the con-
tinental plates around the globe; and vertical tectonics, which
looks at what happens if you create mountains or plateaux. Lastly
we will consider how these changes influenced hominin evolu-
tion in Africa.
Horizontal tectonics
Latitudinal continents
The north–south position of the continents has a huge effect
on the thermal gradient between the poles and the equator.
Geologists have run simple climate models to look at this effect
(Figure 9). If you put all the continents around the equator—
the so-called ‘tropical ring world’—the temperature gradient
between the poles and the equator is about 30°C (Figure 9). When
the poles are covered by ocean, they cannot go below freez-
ing, due to a trick of both the atmosphere and the oceans. A fun-
damental aspect of climate is that hot air rises and cold air sinks.
At the poles it is cold so the air sinks, and as it hits the ground
it spreads outwards away from the pole. When sea water at the
pole freezes it forms sea ice, and this ice is then blown away
from the pole towards warmer water where it melts. This main-
tains the balance and prevents the temperature at the poles going
below zero. However, as soon as you place land at the poles, or
even around the poles, ice can form permanently. If you have
a landmass like Antarctica over a pole with ice, then ground
temperatures drop below −35°C, which creates an equator–pole
42
(i) (ii)
+30 +30
Average surface temperature (°C)
tropical ‘ring world’
1
present northern
0 0 hemisphere
Polar ‘cap world’
2 (no ice)
Polar ‘cap world’ (with ice)

present southern
hemisphere
3
–30 –30
90° 60° 30° 0° 90° 60° 30° 0°

latitude latitude
(1) tropical ‘ring world’ (2) Polar ‘cap world’ (3) Polar ‘cap world’
with ice caps
continent
ice-cap
ocean
figure 9 Illustration of the huge effect the latitudinal location of continents have
on the Equator–Pole temperature gradient. The present-day northern and southern
hemisphere temperature gradients are given as a comparison.
temperature gradient of over 70°C (Figure 9)—exactly what we

have today.
In contrast, if you consider the present northern hemisphere,
the continents are not on the pole, but surround it. So instead of
one huge ice sheet, as we have in Antarctica, there is a smaller one
on Greenland, and the continents act like a fence and keep all the
sea ice in the Arctic Ocean. So the equator–pole temperature
43
g radient of the northern hemisphere is somewhere between the

extremes of Antarctica and an ice-free continent—about 50°C.
The size of the equator–pole temperature gradient is a fundamen-
tal driver of our climate, because ocean and atmospheric circula-
tion is in part driven by the need to move heat from the equator
to the poles. This temperature gradient defines what sort of cli-
mate the world will have. A cold Earth has an extreme equator–
pole temperature gradient and thus a very dynamic climate. This
is why we have strong hurricanes and winter storms as the cli-
mate system tries to pump heat away from the hot tropics towards
the cold poles.
Longitude continents
Ocean circulation is controlled by how the oceans are contained
by the continents around them. If there are no continents in the
way then oceans will just continue to circulate round and round
the globe. However, when an ocean current encounters a contin-
ent it is deflected both north and south. If we look at the modern
configuration of the continents, there are three main longitudinal
continents: (1) the Americas, (2) Europe down to Southern Africa,
and (3) North East Asia down to Australasia. A hundred million
years ago the continents would have been still recognizable, but
in slightly different positions. The two striking features are that,
first, there was an ocean across the whole of the tropics through
the Tethyan Sea and the Deep Central American passage, and sec-
ond, there was no ocean circulating around Antarctica. These
changes have had huge effects on the circulation of the surface
ocean and thus deep ocean circulation and global climate.
The circulation of the surface ocean is important in transport-
ing heat around the globe. The circulation of the surface oceans is
44
primarily controlled by the prevailing winds, because as the wind

blows on the surface water the friction allows the energy to be
transferred from winds to the surface water that leads to the
major currents. The wind energy is transferred to greater depths
in the water column turbulence, which allows wind-driven cur-
rents to be very deep. The rotation of the Earth adds an extra twist,
because of the Coriolis effect, which means in the northern hemi-
sphere the ocean currents will move to the right of the prevailing
wind direction while in the southern hemisphere they will move
to the left.
In the tropics the prevailing wind direction is from both the
north-east and south-east, and hence the surface ocean currents
move westward. In the subtropics the wind come from the west
and thus surface ocean currents are pushed eastward. In the high
latitudes the winds are again from the west and so the surface
ocean is pushed eastward. These currents will flow as far as they
can until they hit an obstacle such as a continent, when the cur-
rents are deflect both north and south along the continental mar-
gin. These giant currents join up within each ocean basin and
form gyres. These large systems of rotating ocean currents are
found in the present-day North and South Atlantic Oceans, North
and South Pacific Oceans, and the Indian Ocean.
By understanding the modern circulation of the oceans the
effects of ocean gateways on ocean circulation can be investi-
gated by modelling three very different scenarios. The first is a
simple double-slice world with longitudinal continents on either
side (Figure 10a). Because ocean currents are driven westward
by the surface winds in the tropics and high latitudes the ocean
currents are pushed to the west, while in the mid-latitudes they
are pushed to the east. This produces the classic two gyre s olution
45
figure 10 Illustration of the significant alteration of ocean currents with

different ocean gateways within longitudinal continents.
46
in both hemispheres. Today, both the North Pacific Ocean and

the North Atlantic Ocean have this type of circulation. The sec-
ond scenario is a double-sliced world with a low-latitude sea-
way. This produces a large tropical ocean circulating continually
westward around the world. There are then two smaller gyres in
each hemisphere (Figure 10b). This is the circulation seen during
the Cretaceous Period, with the two gyres in each hemisphere
occurring in the Pacific Ocean. The remnants of the Cretaceous
low-latitude seaway, the Tethys Sea, can still be seen today in the
Mediterranean Sea. The linkage between the Tethys Sea and the
Indian Ocean finally closed between 11 and 7 million years ago.
This coincides with the first sand dune deposits found in the
Sahara desert dated at 7 million years ago. It was originally
assumed that the Sahara desert had come into existence with the
start of the great ice ages at about 3 million years ago. Climate
modelling work now suggests that the shrinking of the Tethys Sea
weakened the African summer monsoon, allowing desert condi-
tions to expand across North Africa and created a split between
African and Asian flora and fauna. This work also suggests that
the final closing of the Tethys Sea enhanced the sensitivity of the
African monsoon to orbital forcing (see Chapter 6), which later
drove the expansions and contractions of the Sahara desert.
About 5.5 million years ago tectonics closed off the link between
the Mediterranean Sea and the North Atlantic Ocean, and over a
period of 200,000 years it repeatedly dried out and had a pro-
found effect on the climate of North Africa.
The third scenario is a double-sliced world with high-latitude
seaways. This produces strong circumpolar ocean currents and a
single tropical gyre in each hemisphere (Figure 10c). Today, the
southern hemisphere resembles this scenario, with a circumpolar
47
current around Antarctica. The Southern Ocean thus acts like a

giant heat extractor and was instrumental in the huge build-up of
ice on Antarctica.
Deep ocean circulation
Deep ocean circulation is also an important consideration, as it

influences the circulation of the surface ocean and the distribu-
tion of heat between the hemispheres. The presence or absence of
ocean gateways has a profound effect on deep ocean circulation.
For example, our modern-day North Atlantic Deep Water (NADW),
which helps pull the Gulf Stream northwards, maintaining a mild
European climate, may be only 4–5 million years old—about the
same time Australopithecines were spreading across Africa. If we
run computer simulations of ocean circulation with and without
the Drake Passage and the Panama ocean gateway, only the mod-
ern-day combination produces significant NADW. Hence, our
modern-day deep ocean circulation is due to an open Drake
Passage from about 25 million years ago and the closure of the
Panama ocean gateway from about 4 million years onwards
(Figure 11). It is all due to salt. Because of the greater effect of evap-
oration in the North Atlantic region, the North Atlantic Ocean is
saltier than the Pacific Ocean. NADW forms today when the
warm salty water from the Caribbean travels across the Atlantic
Ocean and cools down. The high salt load and colder temperature
both increase the density of the water, so it is able to sink north of
Iceland. When the Panama passage was open, fresher Pacific
Ocean water leaked in and reduced the overall salt content of the
North Atlantic Ocean. The surface water, even when cooled, was
thus not dense enough to sink, and so not as much NADW was
48
figure 11 The effect of changing ocean gateways on the location and intensity of
deep ocean circulation. NADW = North Atlantic Deep Water, AABW = Antarctic
Bottom Water, and Sv = Sverdrup that is a 1 million cubic metres per second.
49
formed compared to today. So one of the fundamental elements

of our modern climate system—the competition between the
Antarctic Bottom Water and the NADW—turns out to be a very
young feature.
Monsoons
The position of the continents in the tropics is also very import-

ant as they can create monsoonal systems. The name monsoon
comes from the Arabic word mausim, which means season, as
most of the rains that fall in South East Asia occur during the
northern hemisphere summer. In the tropics the Sun’s energy is
most intense when it is overhead. The seasons are caused by the
Earth’s axis of rotation being tilted at an angle of 23.5°, and so
each hemisphere receives different amounts of energy through-
out the year. Following the Sun through a year we can see how
this tilt affects the Earth through the seasons. If we start at 21
June, the Sun is overhead at midday at the Tropic of Cancer
(23.4°N)—the northern summer solstice. The angle of the Sun
moves southward until 21 September when it is overhead at mid-
day over the equator—the equinox or autumn equinox in the
northern hemisphere. The Sun appears to continue southward,
and on 21 December is overhead at midday at the Tropic of Capricorn
(23.4°S)—the southern summer solstice. The Sun then appears to
move northward until it is directly overhead at midday at the
equator on the 21 March—the equinox or spring equinox in the
northern hemisphere. And so the cycle c ontinues.
During the summer in each hemisphere the continents receive
the most solar energy and heat up. This causes the air above the
land to warm and rise, creating an area of low pressure beneath
50
it. This low pressure sucks air from the surrounding area and, if
this is from an adjacent ocean, very moist air is brought over the
continent. As this moist air is warmed it rises, and as it gets
higher it cools, the water condenses, and the extremely heavy
rains we refer to as monsoons are created. In Africa there are two
monsoonal systems. The first is in West Africa: during the north-
ern hemisphere summer, North Africa heats up and draws in
moist air from the Atlantic Ocean creating the Congo mon-
soonal system. This system is so strong that Atlantic Ocean-
derived moisture is pulled across the whole continent into East
Africa. The second monsoonal system is in East Africa. Again,
during northern hemisphere summer North Africa heats, up but
in East Africa the moist air is drawn from the Indian Ocean.
However, as we will see later, because of the East African Rift
Valley mountains and the Tibetan Plateau, much of this moist air
is deflected and joins the intense South East Asian monsoonal
system.
Vertical tectonics
As the tectonic plates bearing the continents move around the

surface of the Earth they frequently collide. When this happens
the land is pushed upwards. In some cases chains of mountains
are formed; or, when whole regions are uplifted, plateaux are
formed. These have a profound effect on the climate system. One
of these effects is a rain shadow, which is a dry area on the lee-
ward side of a mountain system. There is usually a correspond-
ing area of increased precipitation on the forward side. As a
weather system at ground level moves towards a mountain or
plateau it is usually relatively warm and moist (Figure 12). As the
51
figure 12 Mountain rain-shadow (RH = relative humidity).
air encounters the mountain it is forced to move up and over it.

Because atmospheric pressure decreases with increasing altitude
the air has to expand, and as it does it cools down. Cool air can
hold less moisture than warm air so the relative humidity rapidly
rises until it hits 100 per cent, and strong rainfall occurs. As the
air descends on the other side of the mountain, atmospheric pres-
sure increases, the air temperature rises, and the relative humidity
drops very low, because little or no moisture is left in the air.
Hence, on the descending side there is a rain-shadow area as there
is no moisture left to form rain, and this can lead to the creation
of a desert. This simple process can control the wetness or dry-
ness of whole continents. Figure 13 shows the effect of mountains
occurring on the western or eastern boundary of a continent.
There are three main rainfall belts in the world: one in the trop-
ics and one each in the mid-latitudes of the two hemispheres. Air
in the tropics moves from east to west, while in the mid-latitudes
it moves west to east. Having mountains on the western side
52
figure 13 The effects of mountains and plateaus on global rainfall.
roduces more rainfall on land and a wetter continent overall. By

p
coincidence, at the moment we have western mountain ranges
running down the west coast of North America—the Rockies—
and the west coast of South America—the Andes. These moun-
tains not only produce significant wet areas but also famous
deserts like the Atacama in Chile and Death Valley in the United
States—two of the driest deserts on Earth. In contrast, when the
East African Rift Valley was formed it created mountain ranges
up to 4 km high on the eastern boundary of the Africa conti-
nent—hence a lot of the moisture is prevented from falling over
East Africa and is lost to the Indian subcontinent.
Atmospheric barriers
When huge mountains or plateaux are thrust high up in the sky

they interfere with the circulation of the atmosphere. Not only do
they force air up and over them, but in many cases they deflect the
53
weather system around them. This effect is compounded as up

lifted areas also warm up in summer and cool down in winter
more than the surrounding lowlands. Figure 14 shows that if all
the continents in the northern hemisphere were flat then the
major circulation of the atmosphere would be nearly circular,
with maybe a slight deflection due to the difference between land
and the oceans. However, if you put the two modern plateaux in
place—i.e. the uplifted regions of the Tibetan–Himalayan and
Sierran–Coloradan complexes—then there are huge changes in
circulation. Both these plateaux are massive. The Tibetan Plateau
is the world’s highest and largest, with an area of 2.5 million km2,
which is about four times the size of France. The Colorado Plateau
covers an area of 337,000 km2 and is joined to numerous other
plateaux, which together make up the Sierran–Coloradan uplift
complex.
In the northern hemisphere winter these highlands are much
colder than the surrounding areas, creating a high-pressure sys-
tem and out-blowing anti-cyclonic circulation (see Figure 14a).
This deflects Arctic air northwards and keeps the middle of the
Asian and North American continents warmer than they would
otherwise be. In the northern hemisphere summer these two
major plateaux heat up more than the surrounding areas and thus
the air above them rises, creating an intense low-pressure zone
(Figure 14b). This sucks in surrounding air, producing a cyclonic
circulation-deflecting weather system much further north and
south. The summer cyclonic circulation around the Tibetan–
Himalayan plateau also greatly intensifies the South East
Monsoonal system. Because part of the air that is pulled towards
the Himalayas comes from the Indian Ocean it brings with it a lot
of moisture, part of which is taken away from East Africa. The
54
figure 14 The effects of plateaus and ice sheet on atmospheric circulation.
South East Monsoonal rainfall is essential for the well-being of

two-fifths of the world’s population.
Volcanic eruptions
Plate tectonics drives the formation of volcanoes, which have an

important influence on climate through the introduction of gases
and dust into the atmosphere. Normal-sized volcanoes inject sulphur
55
dioxide, carbon dioxide, and dust into the troposphere and can have
a considerable effect on our weather. The explosion of Krakatoa in
1883 killed over 36,000 people and is considered to be the loudest
sound ever heard in modern history, with reports of it being heard
nearly 3,000 miles away. The eruption was equivalent to 200 mega-
tons of TNT, which is about 13,000 times the nuclear yield of the
Little Boy bomb that devastated Hiroshima during World War II. The
sulphur dioxide and dust injected into the atmosphere increased
the amount of sunlight reflected back into space, and average glo-
bal temperatures fell by as much as 1.2°C in the year following the
eruption. Weather patterns continued to be chaotic for years and
temperatures did not return to normal until 1888.
The effect of Krakatoa on climate was short term and transient.
This is because the sulphur dioxide and dust were injected rela-
tively low in the atmosphere and the amount of water also injected
meant much of the material was washed out of the atmosphere
within a few years. Volcanic eruptions experienced during human
history have been very small compared to eruptions from super-
volcanoes. These are thousands of times larger than Krakatoa.
They can occur when magma in the Earth rises into the crust
from a hotspot but is unable to break through the crust. Pressure
builds in a large and growing magma pool until the crust is una-
ble to contain the pressure. Supervolcanoes can also form at con-
vergent plate boundaries—for example, Toba, which last erupted
about 74,000 years ago and ejected about 2,800 cubic kilometres
of material into the atmosphere. And they can form in continental
hotspot locations—for example, Yellowstone, which last erupted
2.1 million years ago and ejected 2,500 cubic kilometres of mater-
ial. Because of the scale of these events the sulphur dioxide and
dust are injected much higher in the atmosphere and therefore
56
the effects on global climate can last much longer. Modelling

work by the UK Meteorological Office suggests that a single trop-
ical supervolcano eruption would cause a drop of global temper-
atures of at least 6°C—up to 15°C in the tropics—for a minimum
of three years. Then, over a couple of decades, the climate would
slowly come back to within 1°C of normal. But imagine what
would happen if you had a succession of supervolcano eruptions.
This is exactly what happened about 66 million years ago in India;
making the global climate see-saw between warm and cold con-
ditions on a decadal timescale.
The Deccan Traps are massive lava flows which began forming
about 66.25 million years ago, at the end of the Cretaceous Period.
It is estimated that when formed over a 30,000-year period the
lavas covered 1.5 million km2, which is approximately half the size
of modern India. Due to erosion and plate tectonics the Deccan
Traps have been reduced to their current size of around 500,000 km2.
The release of volcanic gases, particularly sulphur dioxide, during
these frequent supervolcanic eruptions, is thought to have dropped
global temperatures by over 2°C. It has been speculated that these
frequent recurring cooling events may have played a role in the
Cretaceous–Paleogene extinction event, when non-avian dinosaurs
were made extinct. However, the current consensus among scien-
tists is that this extinction was triggered by the Chicxulub meteorite
impact event in Central America around 66 million years ago, as
this would have produced a sunlight-blocking dust cloud that killed
much of the plant life and reduced global temperature, during what
is now called an impact winter. However, recent work by Gerta
Keller at Princeton and others suggests the extinction may have
been caused by both the volcanism and the impact event. This
extinction event was essential in the story of human evolution as it
57
ended the 130 million-year domination of dinosaurs and allowed

the evolution and proliferation of mammals, in particular the
appearance of the first ancestors of primates.
Icehouse and greenhouse worlds
Plate tectonics drives the slow shift of the continents across the
globe, combining them into a supercontinent and then back into
fragmented continents again. The supercontinent Rodinia formed
about 1.1 billion years ago and broke up roughly 750 million years
ago. Another supercontinent, called Pannotia or the Vendian
Supercontinent formed about 600 million years ago, but only
lasted about 60 million years. One of the fragments included
large parts of the continents we now find in the southern hemi-
sphere. Plate tectonics brought the fragments of Rodinia back
together in a different configuration about 300 million years ago,
forming the best-known supercontinent, Pangaea. Pangaea subse-
quently broke up into the northern and southern supercontinents
of Laurasia and Gondwana, about 200 million years ago. Both of
these supercontinents have continued to fragment over the past
100 million years. Icehouse climates form when the continents are
moving together. The sea level is low due to lack of sea-floor pro-
duction. The climate becomes cooler and more arid because of
the reduction in rainfall due to the strong rain-shadow effect of
large superplateaux. On the other hand, greenhouse climates are
created as the continents are dispersed, with sea levels high due to
the high level of sea-floor spreading. There needs to be a relatively
large amount of carbon dioxide in the atmosphere, maybe over
three times current levels, due to production at oceanic rifting
zones. This produces a warm, humid climate.
58
The formation and break-up of these supercontinents had a

huge effect on evolution (Figure 15). Supercontinents are extremely
bad for life and are often associated with mass extinctions. First,
the interior of the continent is very dry and global climate is usu-
ally cold. Second, there is a massive reduction in the amount of
shelf seas. It is not surprising that the explosion of complex ani-
mals occurred during the Cambrian Period beginning 542 million
years ago, following the break-up of the Vendian supercontinent,
as the amount of shallow seas in which complex life first evolved
increased exponentially as the supercontinent fragmented. The
new continental fragments became less and less arid, and thus
increasingly hospitable for life. Mass extinctions are correlated
with the formation of supercontinents. For example, it is esti-
mated that up to 96 per cent of all marine species and 70 per cent
of terrestrial vertebrate species became extinct during the Permian–
Triassic extinction event 250 million years ago—nicknamed the
‘mother of all mass extinctions’ (Figure 15). This mass extinction
ended the dominance of the ‘proto-mammals’ and led to the evo-
lution of dinosaurs and mammals though, for the next 170 mil-
lion years, the dinosaurs reigned supreme.
Understanding how tectonics affects global climate suggests
the influence of the formation and break-up of supercontinents
on the development of complex animals 542 million years ago,
and the evolution of mammals at 225 million years ago. A com-
bination of supervolcanoes erupting in India and a massive
meteorite impact around 66 million years ago ended the domin-
ance of the dinosaurs and enabled the mammals to diversify,
including the appearance of the early ancestors of primates.
Tectonics also influences the location of the continents and thus
where monsoonal systems form. The uplift of mountain ranges
59
figure 15 Long-term links between tectonics, sea level, climate, biodiversity, and
extinctions.
and plateaux influences rainfall and temperature patterns. As we

will see in Chapter 4, the uplift of the Tibetan Plateau that started
20 million years ago intensified the South East Asian monsoon,
drawing moisture away from East Africa. This was further exacer-
bated by the formation of the East African Rift Valley mountains
which deflected much of the moisture-laden air towards the
Indian subcontinent. This aridification of East Africa and the cre-
ating of a mosaic of different vegetation types seems to be one of
the key driving forces of early human evolution.
60
4
Cradle of Humanity
-
T ectonics can cause significant changes in climate, hydrology,

and vegetation cover, both globally and regionally. In this
chapter we will look at the recent history of tectonics in East
Africa and how this has changed the local climate and vegetation.
The long-term climatic change in East Africa was controlled by
the progressive formation of the East African Rift Valley leading
to increased aridity, a fragmentation of the vegetation, and the
development of many lake basins. This made the region highly
responsive to climate change, and created a dynamic environ-
ment in which our ancestors lived and evolved.
Formation of the East Africa Rift System
East Africa is an example of an area that has undergone the pro-

cess of active rifting. This is initiated by significant uplift due to a
magmatic hot spot beneath the crust. Hot spots are caused by
plumes of hot magma, which rise from deep in the mantle. Where
a hotspot occurs beneath a continent, it heats the continental
61
cr a dl e of hu m a ni t y
figure 16 Tectonic rifting and ocean formation.
crust, which expands and stretches, and this bulging of the crust
leads to the formation of a high plateau (Figure 16). Finally, the
expansion becomes so great that the rocks fracture, creating
faults on either side of the plateau. These faults allow the central
part of the plateau to slip down, creating a giant hanging valley
62
and leaving large mountain ranges on either side. The final stage
of rifting occurs when the bottom of the hanging valley drops
so low the continental crust becomes very thin and upwelling
basaltic magma initiates the formation of oceanic crust. At this
stage, because of the subsidence of the relatively dense oceanic
crust, the Rift Valley becomes flooded with seawater, and this can
be the start of a new ocean. The Atlantic Ocean was formed this
way 130 million years ago when the Americas and Africa started
to move apart.
The East African Rift System is one of the most extensive geo-
logical features on the Earth’s surface. It runs north–south for
around 4,500 km from Syria through East Africa down to
Mozambique. We think it was formed by a hot spot that was cen-
tred on northern Ethiopia, the Horn of Africa, and the southern
Arabian region. As with most rift processes the hot spot created
a triple junction, with the fracturing of the rocks occurring in
three different directions (Figure 17). In this case there is fractur-
ing south along the East African Rift Valley; north-west, forming
the Red Sea; and north-east, creating the Gulf of Aden running
into the northern Indian Ocean. Volcanism in East Africa may
have started in Ethiopia as early as 45–33 million years ago, while
initial uplift may have occurred between 38–35 million years ago.
There is evidence for volcanism as early as 33 million years ago in
northern Kenya, but magmatic activity in the central and south-
ern rift segments in Kenya and Tanzania did not start until
between 15 and 8 million years ago. The high relief of the East
African Plateau developed between 18 and 13 million years ago,
while major faulting in Ethiopia occurred between 20 and 14 mil-
lion years ago and was followed by the development of east-dipping
faults in northern Kenya between 12 and 7 million years ago.
63
figure 17 Map of plate boundaries at the African Rift Valley triple junction.
Subsequent rift expansion has led to distinctive sets of faults, cre-

ating a landscape dominated by grabens and half grabens.
A graben is the geological name of the part of the crust that
drops when there are vertically orientated faults (Figure 18). A half
graben is when there is only one set of faults, creating very
distinctive highlands or horsts, with one side having a very steep
slope (the ‘footwall’) and the other side (the ‘hanging wall’) a
64
figure 18 Explanation of horst-graben and half-graben geological formations.
gently sloping gradient. In Kenya, half-graben morphologies devel-

oped between 9 and 6 million years ago, whereas the full-graben
morphology was established between 5.5 and 3.7 million years ago.
By 2.6 million years ago, the Kenya Rift was further fragmented by
new wet-dipping faults, which dropped part of the region down
to a 40-metre-wide inner rift, leaving a 30-kilometre-wide intrarift
plateau: the Kinangop Plateau.
This rifting process has created a unique landscape (Figure 19).
In the centre of the Rift Valley system there is a large hanging
valley about 1,000 m above sea level; to either side are asymmet-
ric mountain ranges with very steep sides facing the valley, and
then gentle slopes down the other side. It is amazing to fly
across this landscape when getting to and from fieldwork sites
in the Rift Valley. You get a real sense of the saw-tooth shape of
the Rift shoulder mountains which steadily get higher and
higher the further you get from the valley. Add to this a smaller
rifted valley within the central Rift Valley and you get a sense
65
figure 19 Tectonic development of the East African Rift Valley system.
of how complex and fragmented this landscape really is. In a

way, calling the East African Rift Valley the Cradle of Humanity
may be misleading because, although most of our ancestors
are found within the valley floor—‘the cradle’—the sides of
this cradle are not gentle and protective but rather harsh jag-
ged teeth rising thousands of metres on either side.
After 2 million years, though the rifting process slowed dra-
matically, there was still a lot of volcanism going on. Importantly
for our story, many of the Kenyan lake basins continued to frag-
ment due to this ongoing volcanic activity. This included the for-
mation of the Barrier Volcano separating Lake Turkana and the
Suguta Valley in the northern Kenyan Rift, which started to form
about 1.4 million years ago, and finally separated the two lake
basins about 0.7 million years ago (see Chapter 6 for more dis-
cussion about these lake basins). The Emuruangogolak Volcano
formed about ~1.3 million years ago, separating Lake Baringo and
66
the Suguta Valley. This means that before 1.4 million years ago an
interconnected lake system may have existed stretching from the
Omo National Park in the north to just north of Lake Baringo in
the south, a distance of over 500 km. Further south, in the Rift
Valley in Tanzania, initial rifting and basin formation started about
5 million years ago. But the major phase of rift faulting occurred
1.2 million years ago, resulting in the present-day rift escarpments
seen in Tanzania.
Rift systems develop from an initial triple junction, and the
activity along each of the three arms varies through time. In the
case of the Afar-centred triple junction, there is still a huge
amount of activity in Ethiopia, and the rifting process continues
in the Red Sea and the Gulf of Aden. The Somali Plate and the
Nubian Plate are currently moving away from the Arabian Plate at
a rate of 6–7 mm per year. However, in the lower East African Rift
System the rifting process became quiescent about 200,000 years
ago, and rifting has ceased. There is still tectonic activity within
East Africa, but the new faults do not run north–south—they run
NNW–SSE along the fractures of the very old basement rock
formed during the Pan-African Orogeny, a mountain-building
period over 600 million years ago.
Rifting influence on climate and vegetation
These tectonic changes are known to be associated with a variety of

biotic changes, which in turn may have affected hominin evolution.
Prior to the hot-spot-driven uplift, a swathe of rainforest stretched
from the Congo all the way across to East Africa. This was fed by
rains from the monsoon systems on either side of the African
continent, bringing moisture both from the Atlantic and Indian
67
figure 20 Climatic effects of the development of the East African Rift Valley
system.
Oceans (Figure 20). During the Oligocene and Miocene Epochs,

progressive uplift of East Africa split this pan-African rainforest.
This is shown by the appearance of new ‘endemic’ species of plants
and animals that evolved locally at about 33, 16, and 8 million years
ago.
This splitting of the east and west rainforests was only the start
of the huge influence rifting had on the climate of East Africa. As
the uplift progressed, rain shadows were created on both the
Congo and Indian Ocean side. This intensified the rainfall on the
outer sides of the mountains, but led to a decrease in moisture
within the Rift Valley (Figure 20). The uplift of the Tibetan Plateau
and the Eastern Rift Shoulder mountain range created the Findlater
Jet. This narrow, low-level, atmospheric jet stream blows diag-
onally across the Indian Ocean, parallel to the coasts of Somalia
and Oman. The Findlater Jet accelerates the transport of mois-
ture from the Southern Indian Ocean, past East Africa, and on to
68
the Indian subcontinent, contributing to the aridification of East

Africa (Figure 21).
By between 5.5 and 4 million years ago the East Africa Rift
Valley had expanded to such a scale that the rain-shadow effect
led to much lower rainfall, so that rainforest was no longer able
to survive (Figure 22). This drying of East Africa seems to have
been progressive and has continued over the past 5 million years.
Evidence for this comes from pollen records and stable carbon
figure 21 Influence of Tibetan uplift and the African Rift Valley on regional
atmospheric circulation during Northern Hemisphere summer. H = high pressure
area and L = low pressure area.
69
isotopes measured in fossilized soil carbonates, organic matter

in marine sediment, and fossilized mammal teeth. The carbon
isotope records (see Box 1) show that there was a progressive
shift in vegetation from mainly trees and shrubs (C3 plants) to
mainly tropical grasses (C4 plants). This aridity trend is also sup-
ported by the results of climate model simulations. These studies
demonstrate that as uplift increased, wind patterns became less
east to west and more south to north, resulting in an overall
decrease in regional rainfall. Hence, as elevation increased, a
rain-shadow effect occurred that reduced moisture availability
on the Rift Valley mountainside, producing the strong aridifica-
tion trend evident in palaeoenvironmental records. Until recently
this aridity was interpreted as just a long-term trend to drier
conditions; but as we will see in Chapter 6, the African climate
swings cyclically from wetter to drier conditions approximately
every 20,000 years. We now think that what has happened over
the last 5 million years is that the dry periods have got much
drier and harsher, while there has been less change in the wet
periods.
In addition to contributing towards the aridification of East
Africa, the tectonic activity described earlier also produced
numerous basins suitable for lake formation. This is shown by
the increased frequency and locations of lake sediments found in
the East Africa over the last 10 million years. If we imagine a
relatively flat East Africa 20 million years ago, then the landscape
would have been dominated by large rivers, very much like the
Congo Basin is today. The southward propagation of rifting,
including the formation of faults and magmatic activity, is also
reflected in the appearance of the earliest lake basins in the
northern parts of the rift. For example, about 5 million years ago
70
Ardipithecus
ramidus
chimpanzee
figure 22 Vegetation effects of the development of the East African Rift Valley
system.
Carbon isotopes
Carbon isotopic records from carbonates and organic matter from

oceans, lakes, soils, and teeth are of interest to palaeoclimatologists
because they help us understand how the local to global carbon system
functions. There are two stable isotopes of carbon: 12C (98.89 per cent)
and 13C (1.11 per cent). There is no difference in the chemistry of these
two isotopes—only the physical difference of weight. But the lighter
isotope is used preferentially in photosynthesis, and this is an example
of an isotope separation or ‘fractionation’ process. As a result, carbon
isotopes can provide an insight into past changes in vegetation and
productivity. All carbon in organic compounds in the biosphere is
ultimately derived from photosynthetically produced material.
Photosynthesis is a very inefficient process—despite billions of years of
evolution, only about 1 per cent of sunlight falling on a leaf is converted
into glucose. Hence, plants use any trick they can to make this process
more efficient. One trick is to use carbon dioxide containing the lighter
continued >
71
12
C isotope as it take less energy to fix. Photosynthetic fixation of carbon
therefore involves a large fractionation of the 12C and 13C isotopes. The
organic matter produced by photosynthesis is very light or depleted in
13
C. Stable carbon isotope ratios are reported in comparison to a globally
accepted standard and given a delta notation, δ13C, which represents the
proportion of 13C to 12C in the sample compared to that in the standard,
given as parts per thousand, or ‰. δ13C values of organic carbon in
land plants (higher plants) vary according to the chemical pathway of
photosynthesis. Plants such as trees, shrubs, and temperate grasses use
the C3 (Calvin–Benson or non-Kranz) photosynthesis pathway and have
a fractionation of between −22‰ and −30‰ compared with the isotopic
composition of the atmosphere of −6‰ to −7‰. Sometime before 12
million years ago a new photosynthetic pathway evolved: the C4 (Hatch–
Slack or Kranz) pathway. This includes tropical and marsh grasses
with a δ13C range of between −9‰ and −15‰ (Figure 23). C4 plants evolved
under low atmospheric carbon dioxide levels as they outcompete C3
plants at low humidity and low CO2 conditions—for instance, during
glacial periods. This is because they have an internal ‘carbon pump’
system, and their primary carbon-fixing enzyme (PEP-carboxylase) does
not react with oxygen—hence, increased photorespiration under low
carbon dioxide conditions does not inhibit photosynthesis. This different
photosynthetic pathway also helps explain why C4 photosynthesis
does not fractionate carbon dioxide as much as C3 plants, as they do not
need this additional efficiency trick. There is also a third type of plant
metabolism—the CAM (Crassulacean acid metabolism)—which uses
either C3 or C4 depending on water availability; but it is minor compared
with the other two pathways.
Carbon isotopes can therefore discern whether the prevailing vegeta-
tion of a region was more C3 (tree and shrub) or more C4 (tropical grass
dominated). These records can be generated from marine, lake, or palaeo-
soil carbonates, or organic matter. Carbonate and bulk organic matter can
be used directly. A more discerning approach is the use of biomarkers—
continued >
72
figure 23 Carbon isotope differences and relative abundance from different

types of plants. The difference in carbon isotope signature between C3 plants
(mainly trees and shrubs) and C 4 plants (mainly tropical grasses) allows the
reconstruction of palaeo-diet and thus the dominate vegetation on the landscape.
particular organic compounds—which are known to originate only from

terrestrial plants. For example, long-chain n-alkyl compounds are major
components of epicuticular waxes from the leaves of vascular plants.
These compounds are relatively resistant to degradation, which makes
them suitable for use as higher plant biomarkers, and include n-alkanes,
n-alkanols, n-alkanoic acids, and wax esters. Carbon isotope records can
be generated from the teeth of animals, and in some cases hominins, and
provide a valuable insight into the diet of early humans. For example,
using δ13C from fossil teeth from the Turkana Basin, Thure Cerling and his
colleagues showed that A. anamensis was mainly forest dwelling; K. platy-
ops, P. aethiopicus, and early Homo were inhabiting grassy woodlands; while
P. boisei was living in much more open conditions in wooded grassland.
73
we find evidence for the beginning of lakes in Afar, Ethiopia,

Omo-Turkana, and the Baringo-Bogoria Basins in northern
Kenya, while the oldest lacustrine sequences in the central and
southern segments of the rift in Kenya and Tanzania occur about
2.5 million years ago, at the start of the Pliocene Epoch. Palaeo-
lakes in the northern part of the East African Rift Valley, thus
formed earlier than in the south. This aridification of East Africa
and the shift from a riverine to lacustrine hydrology must have
had a profound effect on the local environment and also early
human evolution.
Changing landscapes and the origins of bipedalism
For the first four decades of the twentieth century it was generally
assumed that bipedalism had an arboreal ‘tree dweller’ origin.
Hence our ancestors evolved to standing upright within forests as
a way of getting between trees both on the ground and in the air.
This was superseded in the past seventy years by the terrestrial
origin model or theory. Field studies of gorillas and chimpanzees
that began in the 1960s showed that their primary mode of loco-
motion was ‘knuckle-walking’ or quadrupedalism. When gorillas
and chimpanzees move on the ground, they move as if on four
legs by using the knuckles of their hands to support their weight.
Compared to real quadrupeds this is a very inefficient way to move
around. Because of the recognized genetic closeness of humans
and chimpanzees, it was assumed that our common ancestor was
also a knuckle-walking quadruped. So the savannah hypothesis
suggested that the formation of the Rift Valley meant that dense
tropical forest was slowly replaced by grassland. Forest food sources
would have become fragmented, and therefore the distances
74
between habitats increased. One evolutionary strategy to cope

with the new conditions would have been to evolve the ability to
walk upright, as a much more efficient way for a primate to cover
large distances.
This classical view of the origins of bipedalism has recently
been questioned, and we have returned to the original arboreal
theory, based on a better understanding of the lifestyle and evolu-
tion of the great apes and on new hominin fossil finds. All the
great apes walk bipedally, but most do so in an arboreal context.
For example, orangutans spend only 2 per cent of their time using
only their hind legs to move in trees, but a further 6 per cent of
their time where one of other of their arms is used to support
their bipedal movement. Though this is a small amount of time,
it is critical to orangutans as it allows them to access smaller
branches where fruit hangs. It also allows them to cross from one
tree to another, avoiding the effort of climbing down to the ground,
and the high predator risk involved in crossing the ground. The
second line of evidence for a forest origin of bipedalism comes
from the beautiful 4.4 million-year-old fossils of Ardipithecus rami-
dus found in Afar in Ethiopia. The extensive fossil remains of
A. ramidus show that it is a bipedal hominin, but with very long
arms. These long arms are clearly adapted to climbing and swing-
ing in trees, not knuckle-walking, suggesting that Ardipithecus
ramidus was able to travel fairly long distances using its bipedal
adaptation, while retaining the ability to climb trees—essential
for food gathering and escaping the large numbers of predators in
East Africa.
It seems very likely therefore that our ancestors learnt to stand
upright in the trees using their forelimbs for support. This also
implies that the origin of bipedalism is much older than the
75
hominin record suggests, and that it could have evolved many

times. There is fossil evidence of several species of Miocene ape
that show the ability to stand upright in an arboreal context. One
species, Oreopithecus bambolii, an extinct primate found in Italy,
may even have been regularly bipedal on the ground. Oreopithecus
bambolii existed 9 to 7 million years ago when Tuscany and Sardinia
were isolated islands, part of a chain stretching from Central
Europe to North Africa. Its foot has been described as chimp-like,
but its short pelvis was closer to those of hominins than those of
chimpanzees and gorillas. Although it has been suggested that
Oreopithecus was bipedal, it had a peculiar form of bipedalism, dif-
ferent from that of Australopithecines. This is because its big toe
could be placed at an angle of 100° to the other toes, enabling the
foot to act as a tripod in erect postures. So, although Oreopithecus
could stand erect and walk, it would not have been able to develop
particularly fast bipedal locomotion.
The possible repeated appearance of the bipedal adaptation
in different hominin lineages and its origin in a forest setting fits
with our current knowledge of the landscape changes over East
Africa between 10 and 5 million years ago. This is because, as the
uplift and rifting of East Africa proceeded, there was no simple
switch from forest to grassland: far from it. What would have
happened was that the tropical forest started to fragment and
a mosaic of different vegetation types appeared. Today in East
Africa the vegetation ranges from cloud rainforest to arid deserts,
and from open savannah to humid swampland. This fragmenta-
tion of the forest landscape would have driven adaptations within
primates. One adaptation was probably for chimpanzees and
gorillas to become even better at using forest resources as the
total area of forest shrank. This included better tree-climbing to
76
ensure continual access to canopy fruits while still enabling access

to fall-back foods on the ground. This increased commitment to
vertical climbing would have led to more flexible hips and knees,
which would have favoured knuckle-walking when on the ground.
The second adaptation was the likely hominin approach—
adopting terrestrial bipedalism to ensure access to distant food
sources but retaining the ability to climb both for food access and
safety.
The legacy we have inherited from our arboreal ancestors are
feet and legs that can cope with a large range of terrains, which
allowed us later on to develop the ability to run very efficiently.
For example, modern humans have been shown to outrun horses
in different trials, such as those over 22 miles of hilly mid-Wales
or 50 miles of sand dunes in the United Arab Emirates. We are
not, however, fast runners compared to African predators, so the
retention of long arms and powerful leg muscles to climb trees
would have been valuable for our ancestors.
Over the past 10 million years, then, East Africa has changed
from a rolling landscape with tropical forest covering much of
the region to a mountainous fragmented landscape with almost
all possible vegetation types present. The development of the Rift
Valley between 10 and 5 million years created two mountain
ranges, the Rift shoulders, up to 4 km high in places, running
roughly north to south. These mountains created rain shadows
and deflected atmospheric moisture westward back into the Congo
and eastward across to the Indian subcontinent. During this
period the region experienced a huge environmental change. As
we will see in Chapter 5, this change was compounded by the re-
evolution of tropical grasses, adapted to very dry conditions, which
in part drove the emergence of the savannah. The East African
77
primates responded to these massive environmental changes in

two ways: chimpanzees and gorillas became better at tree-climbing
while the hominins enhanced their arboreal bipedalism to enable
them to move around efficiently between the fragmented food
sources on the ground.
78
5
Global Climate Change

-
M assive changes have occurred in East Africa over the past

10 million years due to tectonic uplift and rifting. At the
same time revolutionary changes were occurring in the Earth’s
climate system as it switched from a ‘greenhouse’ to an ‘icehouse’
world. Fifty million years ago the Earth was a very different place.
The world was both warmer and wetter, with rainforest extend-
ing all the way up to Canada and all the way down to Patagonia.
In this chapter I will examine how it transformed from this lush
vibrant state to the ice-locked cool planet we have today. We will
explore what triggered the series of ice ages and the effects these
changes had on early human evolution in Africa.
Super-lush Earth
Before we investigate what caused the massive cooling of the

Earth, we should consider one key climatic event which occurred
during the period when Earth was hot, warm, and lush, that had
a profound effect on mammal evolution. The Palaeocene–Eocene
79
globa l cl im at e ch a nge
Thermal Maximum (PETM) is a climatic event that defines the

temporal boundary between the Palaeocene and Eocene Epochs
in the rock record (Figure 24). The event occurred about 55.8 mil-
lion years ago and lasted about 170,000 years. During this hot-
house event, scientists think that 1,500 gigatonnes of gas hydrates
may have been released. Gas hydrates, or clathrates, are mixtures
of water and methane which are solid at low temperatures and
high pressures. They are made up of cages of water molecules
which hold individual molecules of methane or other gases. The
methane comes from decaying organic matter found deep in
ocean sediments and in soils beneath permafrost. Gas hydrate
reservoirs can be unstable, as an increase in temperature or
decrease in pressure can cause them to destabilize, releasing the
trapped methane. It is thought that the warmth of the Palaeocene
period caused the breakdown of the global gas hydrate reserves,
releasing the trapped methane. Methane in the atmosphere is
twenty-one times more powerful as a greenhouse gas than car-
bon dioxide, and this catastrophic release may be the cause the
observed 5°C temperature rise in global temperatures. On land,
the temperature rises were even larger.
During the PETM the atmosphere became more humid, and
mangroves and rainforests spread as far north as England and
Belgium and as far south as Tasmania and New Zealand. Turtles,
hippopotami, alligators, and palm trees have been found at
Ellesmere Island in the Canadian Arctic. The Arctic Ocean was
very warm but stagnant, with sea temperatures rising from 18°C
to over 23°C. This was a period of rapid extinctions and evolu-
tionary change as local species struggled to adapt to changing
environments while facing fierce competition from migrating
species. It is during the PETM that we have the first fossil evidence
80
PETM
Antarctic Ice Sheet
12
Temperature (°C)
8 India Collides
with Eurasia Northern Hemisphere
Ice Sheets
0
Antarctic Glaciation
60 50 40 30 20 10 0
Time/Millions of years ago
figure 24 Global temperatures through the Cenozoic.
for true primates (Teilhardina brandti and Teilhardina belgica) and the
first social monkeys. It seems that one adaptation to the rapid cli-
mate changes during the PETM was for primates to band together
into large social groups—an essential development, as we think
sociality is a key driver in hominin brain expansion. In addition,
the ancestors of many modern mammals such as hoofed ani-
mals (pygmy horses and elk), tapirs, rodents, bats, owls, ele-
phants, and early whales also appeared during or shortly after
the PETM, heralding a new period of mammal diversification—
the Eocene Epoch.
Growth of ice sheets
If you compare a map of the world about 50 or 60 million years

ago to today, they seem identical—until you look in detail
81
(Figure 25). In Chapter 3 we saw that movements of the continents

around the face of the planet are very slow, but that minor changes
in location have had a profound effect on global climate. Over the
past 60 million years these small changes have moved the Earth’s
climate from a Greenhouse to an Icehouse World. Of particular
importance is having continents on or surrounding the poles and
a means of cooling them down. In the case of Antarctica the ice
did not start building up until about 35 million years ago (Figure 24).
Prior to that Antarctica was covered in lush temperate forest;
even bones of dinosaurs have been found there before they went
extinct 65 million years ago. What changed 35 million years ago
was a culmination of minor tectonic movements. South America
and Australia were slowly moving away from Antarctica, as the
large southern continent of Gondwana fragmented. About 35
million years ago the ocean opened up between Australia and
Antarctica. This was followed about 30 million years ago by the
opening of the Drake Passage between South America and
Antarctica, one of the most feared stretches of ocean. The open-
ing up of these seemingly small ocean gateways between the con-
tinents produced an ocean, the Southern Ocean, which could
circulate completely around Antarctica, continually drawing heat
from the continent and releasing it into the Atlantic, Indian, and
Pacific Oceans with which it mixes. Such has been the efficiency
of this process that there is enough ice on Antarctica today to
produce a global sea-level rise of over 65 metres if it were all to
melt—enough to cover the head of the Statute of Liberty. This
tectonic cause of the glaciation of Antarctica is also the reason
that scientists are confident that the Eastern Antarctic ice sheet,
with the potential of producing about 60 metres of sea-level rise,
will not melt due to future climate change.
82
figure 25 Ocean gateways both today and during the Cretaceous.
The ice-locked Antarctica of 30 million years ago did not, how-

ever, last long. Between 25 and 10 million years ago Antarctica
ceased to be completely covered with ice. This raises some ques-
tions. Why did the world start to cool all over again 10 million
years ago? And why did the ice start building up in the northern
hemisphere 2.5 million years ago? Palaeoclimatologists believe
83
that relatively low levels of atmospheric carbon dioxide are essen-

tial to maintaining a cold planet. Computer models have shown
that if you have high levels of atmospheric carbon dioxide you
cannot get ice to form on Antarctica, even with the ocean heat
extractor. So somehow atmospheric carbon dioxide levels
dropped and ice started building in Antarctica, and then much
later in the Arctic.
What caused the big freeze?
In 1988, palaeoceanographer Bill Ruddiman and his then graduate

student Maureen Raymo, at the Lamont-Doherty Earth Observatory,
wrote an influential paper. They suggested that the observed long-
term global cooling and build-up of ice sheets in the northern
hemisphere were caused by the uplift of the Tibetan–Himalayan
and Sierran–Coloradan regions. As we saw in Chapter 3, huge
plateaux can alter the circulation of the atmosphere, and they
argued that this cooled the northern hemisphere, allowing snow
and ice to build up. However, what they did not realize at the time
was that most of the Himalayan uplift occurred much earlier—
between 20 and 17 million years ago—and thus too early to have
been the direct cause of the ice in the north. But Maureen Raymo
then came up with the startling suggestion that this uplift may
have caused massively increased erosion, using up atmospheric
carbon dioxide in the process. When you make a mountain range
you also produce a rain shadow: one side of the mountain has a
lot more rain on it as air is forced up and over the mountain. This
is also why mountains erode much faster than gently rolling hills.
She argued that this extra rainwater and CO2 from the atmos-
phere formed a weak carbonic acid solution, which dissolves
84
rocks. But interestingly, only the weathering of silicate minerals

makes a difference to atmospheric CO2 levels, as weathering of
carbonate rocks by carbonic acid returns CO2 to the atmosphere.
As much of the Himalayas is made up of silicate rocks, their
chemical weathering could remove a lot of atmospheric carbon
dioxide. The resultant minerals dissolved in the rainwater are
then washed into the oceans and used by marine plankton to
make shells out of the calcium carbonate. The calcite skeletal
remains of the marine biota are ultimately deposited as deep-sea
sediments and hence lost from the global carbon cycle for the
duration of the life cycle of the oceanic crust on which they were
deposited. It’s a fast-track way of getting atmospheric carbon
dioxide out of the atmosphere and dumping it at the bottom of
the ocean. Geological evidence for long-term changes in atmos-
pheric carbon dioxide does support the theory that it has dropped
significantly over the past 20 million years. The only problem sci-
entists have with this theory is what stops the process. With the
amount of rock in the Tibetan Plateau that has been eroded over
the past 20 million years, all the carbon dioxide in the atmos-
phere should have been stripped out. So there must be other nat-
ural mechanisms which help to maintain the balance of carbon
dioxide in the atmosphere, as the long-term concentration of car-
bon dioxide is a balance between what is removed by weathering
and deposition in the deep ocean and what is added when it is
recycled at subduction zones and emitted by volcanoes.
With atmospheric carbon dioxide lowering between 10 and 5
million years ago the Greenland ice sheet started to build up. So,
by 5 million years ago, we had huge ice sheets on both Antarctica
and Greenland, very much like today. The great ice ages, when
huge ice sheets waxed and waned on North America, Northern
85
Europe, New Zealand, and Patagonia, did not start until 2.5 mil-
lion years ago.
The Great Salt Crisis
Between 11 and 7 million years ago, tectonic changes caused the

eastern end of the Tethys Sea gradually to close. This led to the
shifting the African summer monsoons, and we have evidence
for the first sand dunes in North Africa about 7 million years
ago. This is also when we have evidence for possibly the earliest
hominin, Sahelanthropus tchadensis, in Chad. About 6 million
years ago, the gradual tectonic changes in western parts of the
Mediterranean Sea resulted in the closure of the precursor of
the Strait of Gibraltar. This led to the transient isolation of the
Mediterranean Sea from the Atlantic Ocean. At 5.6 million
years ago, the strait closed for the last time and, because of the
generally dry climate conditions, within a millennium the
Mediterranean basin had completely dried out, leaving a deep
dry basin in some places between 3 to 5 km below the world
ocean level (Figure 26). There were a few hypersaline Dead Sea-
like lakes. Around 5.5 million years ago, less dry climatic condi-
tions allowed the basin to resume receiving more fresh water
from rivers, and brackish water lakes—like the Caspian Sea
today—were created, becoming progressively less hypersaline,
until the Strait of Gibraltar finally reopened at 5.33 million
years ago. During the isolation of the Mediterranean region,
desiccation led to the deposition of vast salt deposits. Imagine
a huge version of the Dead Sea where a few metres of seawater
covers a vast area. This event, called the Messinian Salinity
86
Crisis, was a global climate event because nearly 6 per cent of

all dissolved salts in the world’s oceans were removed. By five
and a half million years ago, the Mediterranean Sea was com-
pletely isolated and a salt desert (Figure 26). This was roughly at
the same time palaeoclimate records indicate that the northern
hemisphere was starting to glaciate. This desiccation of the
Mediterranean Sea would have had a profound effect on the cli-
mate of North Africa. Intriguingly, the first evidence for
Ardipithecus kadabba is also dated at 5.6 million years ago. It is
tempting to suggest that the drying out of the Mediterranean
region, and thus East Africa, led to the first known truly bipedal
hominin. But we have so few fossils from 10 and 4.4 million
figure 26 Drying out, salt deposit formation, and reflooding of the Mediterranean
Sea approximately 5 million years ago.
87
years ago that this would be pure speculation; fun, but not very
scientific.
At about 5.33 million years ago, the Strait of Gibraltar opened
up and caused the Terminal Messinian Flood, which is also called
The Zanclean Flood or Zanclean Deluge. Scientists have envis-
aged an immense waterfall higher than today’s Angel Falls in
Venezuela (979 m), and far more powerful than either the Iguazu
Falls on the boundary between Argentina and Brazil or the
Niagara Falls on the boundary between Canada and the USA.
More recent studies of the underground structures at the Gibraltar
Strait show that the flooding channel may have descended in a
more gradual way to the dry Mediterranean. The flood could have
occurred over months or a couple of years, but meant lots and
lots of dissolved salt was pumped back into the world’s oceans via
the Mediterranean–Atlantic gateway. This stopped the start of the
great ice ages at this time—all because of how oceans circulate.
The Gulf Stream in the Atlantic Ocean not only keeps Europe
warm, but also drives the deep ocean circulation and keeps the
whole planet relatively warm. Five million years ago this deep
ocean circulation was not as strong as it is today. This is because
fresher Pacific Ocean water was still able to leak through the
Panama ocean gateway, which is discussed later in this chapter.
The sudden massive increase in salt due to the Terminal Messinian
Flood increased the salinity and the density of the North Atlantic
Ocean, ensuring a very vigorous Gulf Stream and sinking water
in the Nordic seas. With all this tropical heat being efficiently
pumped northwards, the slide into the great ice ages was halted
about 5 million years ago. We had to wait another 2.5 million
years before the global climate system was again on the threshold
of a great ice age.
88
Appearance of tropical grasses
The evolution, emergence, and expansion of grasses using the C4

photosynthetic pathway (Box 1), which took place during the Mid
to Late Miocene, is thought to have been driven by a lowering of
atmospheric carbon dioxide levels. This was a global climate event,
as the evolution of tropical C4 grass-dominated biomes had a long-
lasting impact on continental biota, including major shifts in vege-
tation structure and, in Africa, accelerated forest shrinkage and
the emergence of more open landscapes accompanied by large-
scale evolutionary shifts in animal communities. Although the
theoretical threshold in atmospheric carbon dioxide concentra-
tions for the evolution of C4 photosynthetic pathway was breached
~30 million years ago, evidence for its existence starts 12 million
years ago in Asia. Carbon isotope evidence from fossil soils and
biogenic carbonate suggest that tropical grass-dominated biomes
emerged between 8 and 7 million years ago in Africa.
The Panama Paradox
Another important tectonic control, which geologists believe to

be a trigger for the great ice ages, is the closure of the Pacific–
Caribbean gateway. The Panama ocean gateway began to close 4.5
million years ago, and finally closed around 2 million years ago.
However, its closure creates a paradox, as it would have both
helped and hindered the start of the great ice ages. First, the
reduced inflow of Pacific surface water to the Caribbean would
have increased the salinity of the Caribbean because Pacific
Ocean water is fresher than its counterpart in the North Atlantic
Ocean. This would have increased the salinity of water carried
89
northward by the Gulf Stream and North Atlantic Current and, as

we noted earlier, this would have enhanced deep-water forma-
tion. The increased strength of the Gulf Stream and deep-water
formation would have worked against the start of the great ice
age as it enhances the oceanic heat transport to the high latitudes
and would have opposed ice sheet formation. So, after the aborted
attempt to start the great ice age about 5 million years ago, the
progressive closure of the Panama ocean gateway kept increasing
the heat transported northward, keeping the chill at bay. But here
is the paradox, because two things are need to build large ice
sheets: cold temperatures and lots of moisture. The enhanced
Gulf Stream also pumped a lot more moisture northward, ready
to stimulate the formation of ice sheets. And the building of large
ice sheets in the northern hemisphere could start at a warmer
temperature because of all the extra moisture being pumped
northward, ready to fall as snow and to build up ice sheets.
Why 2.5 million years ago?
Tectonic forcing alone cannot explain the amazingly fast intensi-

fication of northern hemisphere glaciation (Figure 27). My own
research using ocean sediments suggests that there were three
main steps in the transition to the great ice ages. The evidence is
based on when rock fragments ripped off the continent by ice
were deposited in the adjacent ocean basin by icebergs. First, ice
sheets started growing in the Eurasian Arctic and North East Asia
regions approximately 2.74 million years ago, with some evidence
of growth of the North East American ice sheet. Second, an ice
sheet started to build up on Alaska 2.7 million years ago. And
third, the biggest ice sheet of them all on the North East American
90
continent reached its maximum size 2.54 million years ago. So, in
less than 200,000 years, we go from the warm balmy conditions
of the early Pliocene to the great ice ages.
figure 27 Oxygen isotopes showing the step-like changes in global ice volume
over the last 5 million years.
91
The timing of the start of the intensification of northern hemi-

sphere glaciation must have had another cause. It has been sug-
gested that changes in orbital forcing—i.e. changes in the way the
Earth spins round the Sun—may have been an important mech-
anism contributing to the global cooling. The details of the Earth’s
numerous wobbles, and how they cause the waxing and waning
of individual ice ages, are discussed in Chapter 6. But though
these individual wobbles are on the scale of tens of thousands of
years, there are much longer variations as well. For example, one
of the most important is obliquity or angle of the tilt, which is the
wobble of the Earth’s axis of rotation up and down with respect
to the plane of its orbit. Over a period of 41,000 years, the Earth’s
axis of rotation will lean a little bit more towards the Sun, and
then a little less. It is not a large change, varying from between
21.8° and 24.4°. The tilt of the axis of rotation produces Earth’s
seasons, hence a larger tilt will result in greater difference between
summer and winter. Over a period of 1.25 million years the amp-
litude of this cycle of the tilt itself changes—that is, the maximum
variation in angle of the axis during the cycle varies on this longer
period. Both times the Earth tried to enter a glaciated state in the
northern hemisphere, at 5 million years and 2.5 millions year ago,
the variation of tilt had been at its largest value. This made the
changes in each season exceptionally large. Of particular import-
ance were the cold summers in the north, which allowed ice to
survive through the summer and develop into ice sheets.
The tropics react to the ice ages
The onset of the intensification of northern hemisphere glaci-

ation between 2.75 and 2.55 million years ago has, for the past
92
thirty-five years, been cited as the main cause for the appearance
of Homo in East Africa. With the recent discovery of a possible
species of Homo at 2.8 million years, and Homo habilis appearing at
2.35 million years, the two events do seem to overlap. However,
there is little evidence for major environmental change in Africa
at this time. It has also been argued that Homo habilis was not that
different to those members of the Australopithecus genus. With the
discovery of stone tools at 3.3 million years the special place of
Homo habilis has gone. This should help discount the obsession
with the view that intensification of northern hemisphere glaci-
ation was the great cause of human evolution. It would appear
that it was half a million years after the start of the great ice ages
that things started to change in the tropics.
Before 1.9–1.8 million years ago it seems there was very little
east–west sea surface temperature gradient in the Pacific Ocean,
but there was afterwards. This indicates a switch in the tropics
and subtropics to a modern mode of circulation, with relatively
strong Walker circulation and cool subtropical temperatures. The
Walker circulation is the atmospheric east–west component of
the Hadley cell and is instrumental in controlling rainfall in the
topics. It is also a key element in the El Niño-Southern Oscillation
or ENSO (see Box 2). So before 2 million years ago, ENSO may not
have existed in its modern form because there was a relatively
weak Walker circulation. The development of the Walker circula-
tion after 1.9 million years ago also seems to have been linked to
major changes in the African environment. Carbon isotopes
from fossil mammals suggest that, although there was a general
trend towards more open environments after 3 million years ago,
the most significant environmental change to open, grassy land-
scapes occurred after 2 million years ago, rather than 2.6–2.4
93
million years ago, as earlier research had suggested. The reanaly-

sis of terrestrial dust records found in marine sediment from the
Arabian Sea, the eastern Mediterranean Sea, and off subtropical
West Africa, all suggest an increase in aridity and great climatic
variability on the continent after 1.9–1.5 million years ago. As we
will see in Chapter 7, there is also evidence for large, deep, but
fluctuating lakes occurring throughout East Africa. And from 1.8
million years ago H. erectus, with an 80 per cent increase in brain
size from previous hominins, evolved in East Africa, and the first
hominin migrations out of Africa began.
El Niño-Southern Oscillation
One of the most important and mysterious elements in global climate

is the periodic switching of direction and intensity of ocean currents
and winds in the Pacific region. Originally known as El Niño (‘Christ
child’ in Spanish) because it usually appears at Christmas, it is now
generally known as ENSO (El Niño-Southern Oscillation), and it
typically occurs every three to seven years. It may last from several
months to more than a year. ENSO is an oscillation between three
climate states: the ‘normal’ conditions, La Niña, and ‘El Niño’. ENSO
has been linked to changes in monsoons, storm patterns, and the
occurrence of droughts throughout the world. For example, the
prolonged ENSO event in 1997–8 caused severe weather events all over
the Earth, including droughts in East Africa, northern India, north-east
Brazil, Australia, Indonesia, and Southern US, and heavy rains in
California, parts of South America, the Pacific, Sri Lanka, and east-
central Africa.
In an El Niño event, the warm surface water in the Western Pacific
moves eastward to the centre of the Pacific Ocean (Figure 28). Hence,
the strong convection cell or column of warm rising air is much closer
continued >
94
to South America. Consequently the trade winds are much weaker and
the ocean currents crossing the Pacific Ocean are weakened. This
reduces the amount of cold nutrient-rich water upwelling off the coast
of South America, and without those nutrients the abundance of life in
the ocean declines and fish catches are dramatically reduced. This mas-
sive shift in ocean currents and the position of the rising warm air
changes the direction of the jet streams and upsets the weather in North
America, Africa, and the rest of the world. However, if you ask what
causes El Niño the answer is a sort of chicken and egg one. Does the
figure 28 Pacific Ocean during El Niño and normal periods.
95
westward ocean current across the Pacific reduce in strength, allowing

the warm pool to spread eastward moving with the wind system? Or
does the wind system relax in strength, reducing the ocean currents,
which then allow the warm pool to move eastwards? Many scientists
believe that long-period waves in the Pacific Ocean that move between
South America and Australia over time help shift the ocean currents
which produce either an El Niño or La Niña event.
A La Niña event is a more extreme version of the ‘normal’ condi-
tions. Under normal conditions, the Pacific warm pool is in the Western
Pacific and there are strong westerly winds and ocean currents keeping
it there. This results in upwelling off South America and lots of nutri-
ents, making for excellent fishing. During a La Niña period the tem-
perature difference between the Western and Eastern Pacific becomes
extreme, and the westerly winds and ocean currents are enhanced. The
impacts of La Niña on the world’s weather are less predictable than
those of El Niño. This is because during an El Niño period the Pacific jet
stream and storm tracks get stronger and straighter and it is therefore
easier to predict its effects. La Niña, on the other hand, weakens the jet
stream and storm tracks, making them more looped and irregular. So
the behaviour of the atmosphere, and in particular of storms, becomes
more difficult to predict. In general, where El Niño is warm, La Niña is
cool; where El Niño is wet, La Niña is dry.
Longer, deeper, more intense ice ages
The Early to Middle Pleistocene transition (EMPT) is the name given

to the marked prolongation and intensification of glacial–intergla-
cial climate cycles that began about 1 million years ago and finished
about 800,000 years ago (Figure 27). Prior to the EMPT, the glacial–
interglacial cycles seem to have occurred every 41,000 years, which
corresponds to the slow changes in the tilt of the Earth. After about
800,000 years ago the glacial–interglacial cycles seem to have
96
become much longer, averaging over 100,000 years. The shape of

these cycles also changes. Before the EMPT the transition between
glacial and interglacial periods was smooth and the world seems to
have spent a roughly equal amount of time in each climate. After
the EMPT the cycles become saw-toothed, with ice building up over
a period of 80,000 years to produce deep intense ice ages, followed
by rapid deglaciation or loss of all that ice within 4,000 years. The cli-
mate then stays in an interglacial period like our current climate for
about 10,000 years before descending back into an ice age. One
suggestion for this saw-toothed pattern is that the much larger ice
sheets are very unstable and therefore with a slight change in cli-
mate they collapse rapidly and the whole climate system rebounds
back into an interglacial period. It is also interesting to note that
prior to the EMPT the ice age cycles appear to have had very little
effect on the tropics, which seemed to have a completely different
climate cycle lasting only 21,000 years, driven by different parts of
the orbital forcing system. After the EMPT the glacial–interglacial
cycles seem to have had an increasing influence on the tropics, and
in particular Africa. I am sure you will also have noticed that the
end of the EMPT, when the much longer glacial–interglacial cycles
were established, occurred around the time when we find the first
evidence for H. heidelbergensis.
Unstable ice ages
In many ways, ice ages or glacial periods should really be called

‘climate rollercoasters’, because ice sheets are naturally unstable
and cause the climate to veer violently from one state to another as
the ice sheets dramatically collapse and then regrow. Most of the
variations occur on the millennial timescale, but the start of these
97
extreme events can occur in as little as three years. The most

impressive of these are the Heinrich events, named by Professor
Wally Broecker, a palaeoceanographer at Lamont-Doherty Earth
Observatory, after a paper by Hartmut Heinrich describing them in
1988. Heinrich events are massive collapses of the North American
Laurentide ice sheet that resulted in millions of tonnes of ice being
poured into the North Atlantic Ocean. Wally Broecker in his usual
flamboyant way described them as armadas of icebergs floating
from North America across the Atlantic Ocean to Europe. Huge
gouges have been found on the north French coast where these
enormous icebergs ran aground. These Heinrich events occurred
against the general background of unstable glacial climate, and
represent the brief expression of the most extreme glacial condi-
tions around the North Atlantic region. The Heinrich events are
evident in Greenland ice core records as a further 2–3°C drop in
temperature from the already cold glacial climate. The Heinrich
events have been found to have had a global impact, with evidence
for major climate changes described from as far afield as the Americas,
China, the Arabian Sea, and East Africa. During these events
around the North Atlantic region much colder conditions are
found both in North America and Europe. In the North Atlantic
Ocean the huge number of melting icebergs added so much cold
fresh water that sea surface temperatures and salinity were reduced
to the extent that the surface water could not sink. This stopped all
deep-water formation in the North Atlantic Ocean, switching off
the global ocean conveyor belt. Heinrich events seem to occur
every 13,000 to 7,000 years during an ice age. We now know that
between these massive Heinrich events there are smaller events
occurring at about every 1,500 years, which are referred to as
Dansgaard–Oescheger events or cycles. There is evidence from
98
African lake sediments that these millennial climate cycles may

have been influencing African climate as far back as 2.7 million
years, further complicating our picture of the climate in which our
ancestors lived.
During the period of early human evolution in Africa, then,
there were five major transitions or climate events that had a sig-
nificant influence on African climate. The first was the emergence
and expansion of C4 grass-dominated biomes from 8 million
years ago onwards, which would have helped dry out the region
and led to the creation of the iconic savannahs of Africa. Second
was the Messinian Salinity Crisis between 5.95 and 5.33 million
years ago, and the desiccation of the whole Mediterranean region.
The third transition was the intensification of northern hemi-
sphere glaciation and the start of the great ice ages between 2.75
and 2.55 million years ago. The fourth was the development of the
Walker circulation between 1.9–1.7 million years ago and the
development of a modern ENSO system, both of which seem to
have caused significant changes in the climate of East Africa. And
the fifth was the Early–Middle Pleistocene transition, between 1
and 0.8 million years ago, when Africa first became influenced by
the northern hemisphere-driven glacial–interglacial cycles and
the millennial-scale climate cycles associated with the collapse of
giant North American ice sheets. Many of the climate events
coincide with major changes in human evolution, but how and
why these changes occurred will be discussed in later chapters.
In Chapter 6 we will consider how changes to the Earth’s orbit
have influenced climate—in particular, the climate of Africa.
99
6
Celestial Mechanics
-
T ectonics is the main driving force for long-term global and

regional changes in climate. In the shorter term there is
another major control—celestial mechanics. The Earth’s orbit
around the Sun and the angle of its axis of rotation vary or ‘wob-
ble’. We now know that this ‘orbital forcing’ drove the glacial–
interglacial cycles that are fundamental characteristics of the
Quaternary Period, the past 2.5 million years. More recently we
have found that orbital forcing also has a profound effect on the
seasons in the tropics, changing the occurrence of rainfall, and in
East Africa has led to the expansion and shrinking of large, deep-
water lakes within the Rift Valley.
Orbital forcing
Orbital forcing is one of the hardest subjects I teach, and despite

my repeated efforts we have never been able to show it properly
on TV. So please stick with me through the next section and I will
take you though it step by step. There are three main orbital
100
ce l est i a l mech a nics
parameters or wobbles: eccentricity, obliquity (tilt), and preces-

sion, which have a significant effect on the long-term climate of
the Earth.
Eccentricity describes the shape of the Earth’s orbit around the
Sun. It varies from nearly a circle to an ellipse, due to the gravita-
tional force exerted by Jupiter. The Earth’s orbit varies from nearly
circular, with an eccentricity of 0.005, to quite elliptical, with an
eccentricity of 0.06 (Figure 29). Currently we have an eccentricity
of 0.0174—nearly its minimum. This variation occurs over a
period of about 96,000 years with an additional long cycle of
about 413,000 years (Figure 30). Described another way, the long
axis of the ellipse varies in length over time. Imagine the Earth’s
orbit around the Sun as a perfectly circular rubber band. You put
two fingers into the rubber band and slowly expand it so it makes
an ellipse shape. When you release your fingers the rubber band
goes back to a circle. Today, the Earth’s orbit is a small ellipse and
the Earth is closest to the Sun on 3 January, at about 146 million
km; this position is known as the perihelion. On 4 July the Earth
is most distant from the Sun—about 156 million km at the aphe-
lion. Changes in eccentricity cause only minor variations in total
insolation (solar energy that falls on the Earth’s surface), but can
have a significant effect on the insolation received throughout the
year and thus a big seasonal effect. If the orbit of the Earth were
perfectly circular there would be no seasonal variation in solar
insolation. Today, the average amount of radiation received by
the Earth at perihelion is ~351 Watts per square metre (W/m2), and
329 W/m2 at aphelion. This represents a difference of ~6 per cent,
but at times of maximum eccentricity (ellipse length) over the
past 5 million years the difference could have been as large as
30 per cent.
101
figure 29 Orbital eccentricity, tilt, and precession.
Obliquity, or the tilt of the Earth’s axis of rotation with respect to the
plane of its orbit, varies between 21.8° and 24.4° over a period of
41,000 years (Figure 29). It is the tilt of the axis of rotation that causes
the seasons. Currently the Earth is tilted at 23.44° from its orbital
plane, roughly halfway between its extreme values. The larger the
obliquity, the greater the difference between the insolation received
in summer and winter. As we saw earlier, before about 1 million years
ago, global climate was dominated by this 41,000 years cycle. Even
before the great ice ages 2.5 million years ago the climate swung
gently between warmer and cooler conditions, driven by obliquity.
Obliquity also has a long-term cycle, with the variations in tilt slowly
increasing and then decreasing over 1.5 million years (Figure 30).
Precession is the most complicated forcing of the three main

orbital parameters, but has the most effect on the tropics.
102
figure 30 Long-term changes in the orbital parameters.
103
Precession has one component relating to the elliptical orbit of

the Earth (eccentricity) and its axis of rotation (Figure 29). The
Earth’s rotational axis moves around a full circle, or precesses,
every 26,000 years. This gyroscopic motion is due to the tidal
forces exerted by the Sun and the Moon on the solid Earth,
which are amplified as the Earth is shaped as an oblate spheroid
rather than a sphere. This is similar to the gyrations of the rota-
tional axis or ‘plunger’ of a toy spinning-top. Precession of the
axis of rotation causes a change in the Earth–Sun distance for
any particular date—for example, the beginning of the north-
ern hemisphere summer. However, this will only influence cli-
mate if the orbit of the Earth is an ellipse, because when the
Earth’s orbit around the Sun is a perfect circle there would be no
difference in the Earth–Sun distance. So the effect of precession
of the axis of rotation is modulated by eccentricity, and combining
these two orbital parameters results in a periodicity in the effect
of 19,000 years.
The second component relates to the precession of the whole
of the Earth’s orbit around the Sun (Figure 29). Every 105,000
years the Earth’s whole orbital rotation around the Sun swings
around the Sun. Imagine the Earth’s orbital path around the Sun
as hula hoop, and that you are one of those gifted people who can
spin a hoop around on one ankle without falling down. That spin
of the hula hoop is similar to what happens to the Earth’s orbit
around the Sun. This effect also modulates the influence of the
precession of the axis of rotation, creating a periodicity of 23,000
years. These two periodicities combine so that perihelion coin-
cides with the summer season in each hemisphere—on average
every 21,700 years. Precession has the most significant impact in
the tropics, because obliquity has no direct impact at the equator.
104
So although obliquity clearly influences high-latitude climate

change, which may ultimately influence the tropics, the direct
effects of insolation in the tropics are due to eccentricity-modu-
lated precession alone. By modelling the positions of the planets
in the Solar System, astronomers can combine the effects
of eccentricity, obliquity, and precession to provide the means of
calculating insolation for any latitude back through time. Of
course, small variations in the different gravitational influences
disrupt the calculations and so they become less accurate the fur-
ther back we try to calculate insolation on Earth.
Waxing and waning of the great ice ages
We know about the orbital parameters because of nearly five dec-

ades of scientific endeavour to understand the driving forces
behind the great glacial–interglacial cycles. The waxing and wan-
ing of the huge continental ice sheets were initiated by changes in
the Earth’s orbit around the Sun. In 1949, Milutin Milankovitch, a
brilliant Serbian mathematician and climatologist, was the first
scientist to combine the effects of all three orbital parameters to
calculate the solar energy received by the Earth. He suggested that
summer insolation at 65°N, which is just south of the Arctic
Circle, was critical in controlling glacial–interglacial cycles
(Figure 31). He argued that, if the summer insolation was reduced
enough then ice could survive through the summer, start to build
up, and eventually produce an ice sheet.
Orbital forcing does have a large influence on this summer
insolation. The maximum change in solar radiation in the past
0.5 million years is equivalent to reducing the amount of sum-
mer radiation received today at 65°N to that received now over
105
figure 31 Insolation at 65°N compared with global sea level or global ice volume.
550 km to the north at 77°N. In simplistic terms, it brings the

current ice limit in mid-Norway down to the latitude of mid-
Scotland. These lows in 65°N insolation are caused by eccentricity
elongating the summer Earth–Sun distance, obliquity being shal-
low and precession placing the summer season at the longest
Earth–Sun distance produced by eccentricity. The reason why it
is 65°N and not 65°S, which controlled global climate in the
Cenozoic, is because of the current configuration of the contin-
ents: thus any ice that builds up in the northern hemisphere has
lots of continents to grow upon. In contrast, in the southern
hemisphere, the ice growth is limited by the Southern Ocean,
because any extra ice produced on Antarctica falls into the ocean
and is swept away to warmer seas. It is almost as though Antarctica
is already full up with ice and unable to accept any more. So the
106
conventional view of glaciation is that low summer solar energy

in the temperate northern hemisphere allows ice to survive sum-
mers, and thus ice sheets start to build up on the northern contin-
ents. But this simple clockwork view of the world is really much
more complicated as the effects of orbital changes on the seasons
is very small, and it is feedbacks in the climate system that amp-
lify these changes.
Glacial–interglacial cycles
I always tell students off if they say that orbital forcing causes gla-
cial–interglacial cycles. First, there isn’t a one-to-one relationship
between cause and effect. For example, the position of the Earth’s
orbital parameters is very similar today to that of 21,000 years
ago during the last ice age. So it is not the exact orbital position
that controls climate, but rather the changes in the orbital posi-
tions. Second, orbital forcing in itself is insufficient to drive the
observed glacial–interglacial variability in climate. Instead, the
Earth system amplifies and transforms the changes in solar
energy received at the Earth’s surface through various feedback
mechanisms. For example, let us start with building an ice age.
The first thing to happen is a slight reduction in summer temper-
atures. As snow and ice accumulate due to initial changes in sum-
mer temperature, the albedo—the reflection of sunlight back into
space—increases. Reflecting more sunlight back into space sup-
presses local temperatures, this promotes the accumulation of
more snow and ice increasing the albedo of the region, producing
the so-called ‘ice–albedo’ feedback. So once you have a small ice
sheet, it changes the environment around it to make more snow
and ice, and will get bigger and bigger.
107
Another feedback cycle is triggered when the ice sheets, par-

ticularly the Laurentide ice sheet on North America, become
big enough to deflect the atmospheric planetary waves (see
Figure 14b). This changes the storm path across the North
Atlantic Ocean and prevents the Gulf Stream and North
Atlantic Drift penetrating as far north as it does today. This
surface ocean change, combined with the general increase in
melt-water in the Nordic Seas and Atlantic Ocean due to the
presence of large continental ice sheets, ultimately leads to
reduction in the production of deep water. Deep-water produc-
tion in the Greenland and Labrador Seas is the heartbeat of the
modern climate. By reducing the formation of deep water it
reduces the amount of warm water pulled northwards, all of
which leads to increased cooling in the northern hemisphere
and expansion of the ice sheets.
There is currently debate among palaeoclimatologists about
the relative roles of the ‘physical climate’ feedbacks described
here and of greenhouse gases in the atmosphere. Air bubbles
trapped in polar ice have shown us that carbon dioxide
dropped by a third, and methane by a half, during each glacial
period. These changes would have compounded the cooling
that occurred during each glacial period, helping to build
more ice. The argument continues: do changes in the Earth’s
orbit affect the production of greenhouse gases, cooling down
the Earth to make the northern hemisphere continents sus-
ceptible to the build-up of large ice sheets? Or do changes in
the Earth’s orbit start to build up large ice sheets in the north-
ern hemisphere that then change global climate and reduce
the production of greenhouse gases, prolonging and deepen-
ing the ice age? The jury is still out on this one. However, what
108
we do know is that greenhouse gases played a critical role in

glacial–interglacial cycles. We also know that changes in green-
house gas concentration always comes before changes in global
temperatures.
An important question is why don’t these feedbacks end up
becoming runaway and freezing the whole Earth. They are pre-
vented from doing so by ‘moisture limitation’. To build an ice
sheet you need the climate to be cold and wet. As the warm sur-
face water is forced further and further south, the supply of the
moisture required to build ice sheets decreases. So the ice sheets,
by changing the atmospheric and ocean circulation, end up starv-
ing themselves of moisture—a negative feedback loop that limits
the effects of the positive feedbacks.
In the past million years it took up to 80,000 years to build up
ice sheets to reach the maximum extent of ice. The last time this
occurred was about 21,000 years ago. However, getting rid of the
ice is much quicker. This is called deglaciation and it usually took
a maximum of only 4,000 years. Now this deglaciation is trig-
gered by an increase in solar energy received in the summer at
about 65°N, encouraging the northern hemisphere ice sheets to
melt slightly. The rise of atmospheric carbon dioxide and meth-
ane promotes warming globally and encourages the melting of
the large continental ice sheets. But these processes have to work
against the ice sheets’ albedo effect, which produces a microcli-
mate that tries to keep them intact. What causes the rapid removal
of ice is the rise in sea level due to the melting ice sheets. Large ice
sheets adjacent to the oceans are undercut by rising sea levels
because the coldest seawater can be is about −1.8°C while the base
of the ice sheet is usually colder than −30°C, so it is like putting
hot water under a tub of ice cream. Undercutting of the ice sheet
109
leads to more melting and ice calving into the ocean. This in turns
increases sea levels, which causes even more undercutting. This
sea-level feedback mechanism can be extremely rapid. Once the
ice sheets are in full retreat the other feedback mechanisms dis-
cussed here are thrown into reverse.
Orbital forcing and the African climate
Orbital forcing has an obvious impact on high-latitude climates,

drives glacial–interglacial cycles, and influences global climate
transitions such as the intensification of northern hemisphere gla-
ciation. It also has a huge influence in the tropics, particularly
through precession and its effect on seasonality and, thus, rainfall.
There is a growing body of evidence for precession-forcing of
moisture availability in the tropics, in Africa, Amazonia, and Asia.
The precessional control on tropical moisture has also been illus-
trated by climate modelling. Amy Clement, a climate modeller at
the University of Miami, used a complex climate model, usually
used to model future climate changes, to switch between extremes
of precession to see what effect it had on climate. She and her col-
leagues found that a 180° shift in precession could change annual
precipitation in the tropics by at least 200 mm/year, and cause a
significant shift in seasonality. This is on the same order of magni-
tude as switching between a full glacial and interglacial period. In
contrast, precession has almost no influence on global or regional
temperatures.
The mechanism by which precession changes the hydrological
cycle in the tropics is very similar to that which causes the mon-
soons. In the tropics, the Sun’s energy is most intense when it is
overhead. This heats up the land or sea and thus warms the air
110
above. This warm moist air rises, leaving an area of low pressure
beneath it, which sucks in air from the surrounding area. This
suction results in the trade winds, which can travel from high lati-
tudes to this area of rising air. As the winds come from both the
northern and southern hemisphere, this area is known as the
Intertropical Convergence Zone (ITCZ). As the air at the ITCZ
rises it forms huge towering clouds and produces large amounts
of rain. The ITCZ moves north and south with the seasons as the
position of the most intense sunlight shifts up and down across
the equator. It is also strongly influenced by the position of the
continents. This is because the land heats up faster and to a greater
extent than the ocean, and thus it can pull the ITCZ even further
north or south during the respective season.
Eccentricity affects the Sun–Earth distance as the Earth circles
the Sun. Precession affects which time of year coincides with the
closest Sun–Earth distance, and thus the amount of insolation
received during each season. For example, in the northern hemi-
sphere summer the tropics and subtropics heat up as the Sun
steadily moves from overhead at the equator to overhead at the
Tropic of Cancer. At the maximum positive precession then, the
Sun–Earth distance will be closest when the Sun is overhead at
the Tropic of Cancer. This significantly increases the amount of
solar energy hitting the subtropics, and thus the amount of con-
vection, strengthening trade winds and rainfall within the ITCZ.
Hence the amount of rainfall in the northern hemisphere tropics
is greatly increased.
At the same time, on the flip side, the southern hemisphere
summer will coincide with the longest Sun–Earth distance; thus
the opposite will occur, and rainfall will be greatly reduced in
the southern hemisphere tropics. About 21,000 years later the
111
opposite will occur, and the southern hemisphere tropics will

have the most intense insolation and thus greatly increased rain-
fall. This inverse relationship between the northern and southern
hemisphere hydrological cycles can be seen in palaeorecords over
the past 10,000 years. North African lakes have steadily been dry-
ing out, while the Amazon River discharge has steadily increased
over this period of time. We should also remember that this influ-
ence of precession increases at the peak of eccentricity every
96,000 years, and is greatest once every 413,000 years when the
Earth’s orbit around the Sun is at its most elliptical.
In North and East Africa there are excellent records of this pre-
cessional forcing of the hydrological cycle over the past 5 million
years. First there are the marine dust records for the East
Mediterranean which reflect the periodic increase in aridity over the
eastern Algerian, Libyan, and western Egyptian lowlands north of
the central Saharan watershed. Second, there are complementary
dust records from ocean sediment cores in the North Atlantic
Ocean adjacent to the West African coast and from the Arabian
Sea. Third are the sapropel formations found in marine sediment
recovered by deep-sea drilling in the Mediterranean Sea. A sapropel
is a dark organic-rich layer found in marine sediments, resulting
from a reduction in the oxygen content of the bottom waters.
Sapropels are common within Mediterranean marine sediments
and are caused by increased Nile River discharge, which changes the
circulation of the Mediterranean Sea. The dust records that show
aridity, and the sapropel records which show increased rainfall, are
inversely related and have a dominant periodicity of 21,000 years—
the beat of precession. There is also a growing body of evidence for
precessional forcing of East African lakes, which is discussed in
detail later in the next section.
112
We have gone far into the complex world of orbital forcing

and seen how precession dominates the climate variability of
the tropics so maybe I should stop here. But although the dir-
ect influence of orbital forcing on the African climate seems
straightforward, there are additional complexities of preces-
sion that need to be considered, as these increased the variabil-
ity of the climate in East Africa experienced by our ancestors.
During any year in the tropics, there are two insolation max-
ima when the Sun is overhead at the equator—the spring and
autumn equinoxes—which, at the moment, occur on 21 March
and 21 September. There also two insolation minima when
the Sun is overhead at the Tropics of Cancer and Capricorn—
summer and winter solstice—which, at the moment, occur on
21 June and 21 December. The magnitude of the maxima and
minima, and thus of insolation at equinox and solstice, is, as
we have seen, controlled by precession via the changing Earth–
Sun distance. In East Africa today, these changes create two
rainy seasons, roughly coinciding with the spring and autumn
equinoxes—one in March–April, and another in late September–
early October. Hence, when precession is increasing insola-
tion of the spring equinox, it is also decreasing insolation
during the autumn equinox; therefore one rainy season is get-
ting strong and longer, while the other gets weaker and shorter.
So there are two peaks in rainfall for the tropics every half pre-
cessional cycle, approximately every 11,500 years. There are
also four maximum peaks in seasonality—i.e. the difference
between the maximum and minimum solar insolation occur-
ring in any one year in the tropics. This is because the strength
and duration of the dry periods between the rainy seasons is
determined by insolation at summer and winter solstice. Hence,
113
maximum variation in seasonality in the tropics occurs every

5,500 years.
As I have often argued, it is unsurprising that the tropics are a
hotbed of evolution, because even if there were no major climate
changes, the whole region undergoes massive hydrological shifts
every 5,500, 11,000, and 21,000 years. The key to precessional for-
cing, however, is how sensitive the region is to either overall changes
in yearly rainfall or changes in seasonality, and this determines
which periodicity is the dominant control on the environment.
East African Rift System lakes
The sedimentary record of East Africa is rich in lake deposits.

This is because the southward propagation of rifting created lake
basins along the entire length of the Rift Valley. The map in
Figure 32 shows where these lake basins occur, and whether they
currently contain a lake. The tectonic movements within the Rift
Valley have led to many past lake sediments being exposed by
faulting or uplift. They range from huge vertical deposits that,
from a distance, look like the white cliffs of Dover, to large white
bands sandwiched between fossilized soil horizons.
Because of the dispersed nature of these deposits, the geologist
and palaeoclimatologist Martin Trauth at Potsdam University has
developed a new twenty-first-century style of fieldwork and very
kindly drags me along. He sets up a base camp on the Rift shoul-
der where the daily temperature range is much smaller than in
the Rift Valley. The mobile camps also take advantage of local for-
est and water sources. This is no safari lodge—it consists of tents
and holes in the ground for the toilets—but the food is always
wonderful. To get to the sampling sites in the Rift Valley we get
11 4
figure 32 East Africa, modern and palaeo lake location map.
up at 6.30 a.m. for breakfast, and as the Sun comes up we board

the ‘Rift Valley taxi’—the helicopter. Thirty to forty five minutes
later we will be at the first site we have identified. Throughout the
day, as soon as we have finished with a site, we can then use the
helicopter to move to the next nearest site of interest. When col-
lecting samples and mapping lake and tectonic features, this is an
extremely efficient way to run a field campaign in a study area of
over 12,000 km2 that includes sand dunes, rivers, jagged lava
flows, and swamps. Otherwise it involves hours and hours in a
4×4 just trying to get to a site, and if there is nothing of interest at
that site the whole day is wasted. By around 2.30 p.m., after seven
hours’ work, and as temperatures in the Rift Valley start to climb
over 40°C, we can escape back to the relative cool of the camp to
collate and analyse our samples and data.
Using this field method, Martin Trauth and his team have iden-
tified and mapped large, deep freshwater lakes that filled the Rift
Valley in the past. The key indicator is the presence of pure white,
and frequently laminated, or finely layered, sediments made up of
diatoms. Diatoms are a major group of photosynthetic algae,
found in both freshwater and seawater. They are unicellular,
although they can form colonies in the shape of filaments or rib-
bons, fans, and even stars. The diatom cell is enclosed within a
cell wall made of silica (hydrated silicon dioxide) called a frustule.
These frustules are unique to each species and show a wide diver-
sity in form, but they are usually bilaterally symmetrical—hence
the group name. Diatom species and communities are a useful
tool for monitoring environmental conditions, and have been
used in seminal studies of water quality and the effects of acid
rain. In African lake deposits, the diatom species found can tell us
how fresh the water in the lake was, and the approximate depth of
116
the lake. These ‘diatomite’ sediments are also of commercial

value, and in parts of Kenya they are mined, heated in a furnace,
and the resultant fine-grained material is shipped all over the
world to soft drink manufacturers to filter out any impurities in
their drinks. Shallow and more alkaline lakes can also be identi-
fied from analysis of lake deposits. These deposits have more
mud and sand in the sediment, which would have been washed
into the shallower lake, and diatom species that prefer higher
alkaline conditions. From our fieldwork we found that deep fresh-
water lakes were characterized by an area of several hundred
square kilometres, water depths in excess of 150 metres, and a
neutral pH, and are documented by these pure white and fre-
quently layered diatomites. In contrast, the shallow and more
alkaline lakes were typically less than 150 km2 in size, had water
depths much less than 100 metres (often only a few metres), and
dried out episodically. The pH of these lakes is often around 8,
but may reach significantly higher values. The corresponding
sediments are diatomites with significant amounts of clays and
silts. In extreme alkaline lake sediments, we have found silicates
such as zeolites, that were generated in situ and document chem-
ical weathering of silica volcanic glass due to the extreme pH.
Martin Trauth, myself, and colleagues have compiled a record
of these lake occurrences within the Rift Valley. The collation is
based on our colleagues’ detailed published geological evidence,
and our own fieldwork, identifying the appearance of either deep
or shallow alkaline lakes within East Africa. To make things easier,
we looked at the lakes within seven major basins. These basins,
shown in Figure 33, are: Olduvai (Tanzania), Magadi-Natron-
Olorgesailie (north Tanzania and south Kenya), Central Kenya Rift
(Kenya), Baringo-Bogoria (Kenya), Omo-Turkana-Suguta (north
117
figure 33 East Africa north–south cross section with major modern and palaeo lakes shown.
Kenya), Ethiopian Rift (south and central Ethiopia), and Afar

(north Ethiopia). Dating of the sediments in East Africa is rela-
tively straightforward because of all the volcanic material, which
lends itself well to radioactive Argon-Argon dating (see Box 3).
The first of these lake compilations was published in the jour-
nal Science in 2005, showing that there were key periods of time
when many of these basins contained deep lakes. Our most recent
compilation (Figure 34) shows that there were lake periods 4.6 to
4.4 million years ago, 4 to 3.9 million years ago, 3.5 to 3.3 million
years ago, 3.1 to 3 million years ago, 2.7 to 2.5 million years ago, 2
to 1.7 million years ago, 1 to 0.8 million years ago, and 0.2 to 0
million years ago. These occurrences of deep lakes correlates
with the 400,000-year component of the eccentricity cycle, and
was our first hint of the role of precession in lake formation in
East Africa. Moreover, the last four lake periods also correlate
with major hominin evolutionary and dispersal events.
Over very long timescales of hundreds of thousands of years,
changes in lakes are primarily determined by tectonics, initially
creating but also destroying lake basins. However, tectonics also
affects conditions in a lake over shorter timescales, through
changes to the shape and size of catchment areas and drainage
networks for example. Furthermore, tectonics shapes the morph-
ology of lake basins and hence contributes to the sensitivity of
these lakes to changes in the precipitation and evaporation bal-
ance. The rifting process in East Africa not only created many
lake basins, but has contributed to their sensitivity to small
changes in rainfall, and they have become referred to as amplifier
lakes. These amplifier lakes are very sensitive to moderate climate
change. For example, the reconstructed water level of palaeo-
Lake Suguta changed from zero to over 300 metres deep due to a
119
figure 34 Occurrence of shallow and deep lakes over 5 million years compared with the precessional orbital forcing.
The dating challenge
To understand the causes of human evolution we need to be able to

date the fossils, the hominin artefacts, and the environmental records.
This is a considerable challenge, as our ancestors’ fossilized bones and
artefacts are not directly datable, hence we have to try to date the
sediments in which they were found. Key to this is the stratigraphy of
the fossil site or location, because very often it is layers above or below
the fossil bed that contain material that can be dated. This gives a
minimum or maximum age of the fossils. Working out the stratigraphy
of a site or location is not easy especially in East Africa, which has been
tectonically very active and thus moved and shifted the sediments
around. Geologists spend many years in the field just piecing together
the sequence which the sediments were laid down to ensure that we can
constrain the fossil and artefact ages as much as possible. The hominin
fossils found in South Africa are even harder to date as most of them
have been found in cave deposits bereft of datable material. For
example, despite the excitement of the finding H. naledi, it does not help
us understand human evolution as we have no idea when they were
roaming around South Africa, and given the site we are unlikely to ever
know. Moreover our knowledge of each fossil site continually improves
and so does our ability to date material. An example of this is H.
floresiensis—the Indonesian cave deposits were initially thought to be
11,000 years old, but after another ten years of detailed painstaking
work this has been re-evaluated and the deposits are now thought to be
between 50,000 and 100,000 years old.
There are two primary dating technologies that apply to the study of
hominin evolution; palaeomagnetism and radiogenic isotopes
(Figure 35). Palaeomagnetism uses the direction of magnetic particles
that are found in layers of sediment and compares these to the known
shifts in the Earth’s magnetic field over time. These shifts include full
reversals when the north magnetic pole suddenly becomes the south
continued >
121
magnetic pole. These seem to occur completely randomly through

Earth history and can occur as close as 100,000 years or as far as
part 1.5 million years. You can also use smaller changes in direction of
the magnetic field due to magnetic pole wander and variations in mag-
netic field strength. At the moment the magnetic North Pole is not at
the same position as the Earth axis of rotation North Pole, and we know
that in the past it has wandered as far south as the US-Canadian boarder.
The drawback of palaeomagnetism is that it is not a direct dating
method, as you need to know roughly where you are in time before you
can interpret the results. This is because what you get is a sequence
of normal (north is north) and reversed (north is south) sediments and
hence you could be anywhere in the geological record. So some way
of tying down roughly where you are in time is essential before pal-
aeomagnetics can help you refine the date. For example, in a known
Miocene sequence of rocks palaeomagnetics was used to date Sivapithecus,
a genus of extinct primate and probable ancestor to orangutans, to
12.5 million years ago.
Another very useful relative dating method is orbital forcing stratig-
raphy. This method can be used on long continuous oceans and lake
sediments records. As discussed in Chapter 5, there are regular changes
in global climate between glacial and interglacial periods. For the last
1 million years these have occurred about every 100,000 years, and
before that they occurred every 41,000 years. So by measuring climate
proxies in either lake or ocean cores, which respond to these large
changes in global climate, you can count back successive glacial–
interglacial cycles and get a chronology for the whole of your record
(see Figure 27). This approach can be refined in the tropics, as we
know that the tropical climate responds more to precession than to
the glacial–interglacial cycles, and so long records that contain the
21,000 year precessional cycle can be used to produce and even higher
resolution chronology.
continued >
122
What we all dream of, however, is sediments that can be dated dir-
ectly using radiogenic isotopes to give a real date. We also dream that
we can date the sediment in which the fossils or artefacts are found, but
that rarely happens. The French physicist, Henry Becquerel, discovered
the natural radioactive decay of uranium at the turn of the twentieth
century. Further work by physicists Ernest Rutherford and Bertram
Borden Boltwood indicated that the predictable decay of radioactive
elements could be used to keep track of time. Not only did this approach
confirm the Earth to be several billion years old, it also provided the
earliest empirical timescale for the fossil record. Common chemical
elements used in dating include carbon, uranium, potassium, and
argon (Figure 35). Which elements researchers use depends on how
quickly they decay. For example, radiocarbon (carbon-14 or 14C) dating
is extensively used to date organic and carbonate finds less than 60,000
years old, as it has a relatively small half-life (the time it takes for one-
half of the atoms in 14C to disintegrate) of about 5,730 years. This has
allowed us to document accurately the origins of agriculture, to around
11,000 years ago, and the later occurrence of H. sapiens and the extinc-
tion of Neanderthals. Radiocarbon dating does have its drawbacks.
First, you can only get a date if you have enough uncontaminated car-
bon in your sediment. Second, the amount of 14C in the environment
figure 35 Age range of different dating methods using in the study of

human evolution.
continued >
123
varies due to its changing production rates in the upper atmosphere

and the release of old ‘dead’ carbon with no 14C from the oceans or from
old sedimentary rocks like limestone and chalk. So huge efforts have
been made to calibrate radiocarbon years to calendar or actual years so
that we can covert between the two.
Earlier on I complained that trying to piece together the geology of
East Africa is difficult because of the tectonics, which has resulted in
massive changes in the landscape. But this also means that there is a
large amount of volcanic ash or lava interleaved with the lake and river
deposits in which fossils and artefacts are found. These ashes and lavas
contain potassium-rich minerals such as potassium feldspars, which
are essential for potassium-argon and the newer argon-argon dating.
Of the naturally occurring isotopes of potassium, 40K is radioactive and
decays into argon (40Ar). This decay is at a known rate, as 40K has a half-
life of 1.25 billion years, so that the ratio of 40K to 40Ar in minerals is
always proportional to the time elapsed since the mineral was formed.
The problem is that at room temperature potassium is a solid while
argon is a gas, making the isotopic comparison difficult. To get round
this, samples are placed in a nuclear reactor to convert the stable form
of potassium (39K) into the radioactive 39Ar. It is then possible to meas-
ure the ratio of 40Ar-39Ar and thus to calculate the age of the sample.
Argon-argon dating has been used successfully to date the majority of
the fossil sites in East Africa due to the ubiquitous nature of volcanic
sediments in the region. Argon-argon was used to date hominin
remains at Dmanisi in Georgia, giving a date of 1.8 million years ago. It
has also been used to date the fluvial sandstone layer from which the
oldest known fossils of H. floresiensis were excavated in 2014, suggesting
that this hominin existed around 700,000 years ago.
There other less common dating methods such as thermolumines-
cence and electron spin resonance (ESR), which are used when other
datable material is not present. These measure the quantity of electrons
that get trapped inside a rock or an artefact over time. First developed in
continued >
12 4
the 1950s, thermoluminescence works on the basis that crystals, such

as quartz, trap electrical charges (electrons) at a known rate over time.
If a crystal, for example a flint blade, is heated, these electrons are liber-
ated, emitting a measurable amount of light. This allows researchers to
determine when an object was last heated. What is actually determined
is the amount of elapsed time since the sample had previously been
exposed to high temperatures. It is used to date volcanic materials and
meteorite impact craters, but can also directly date past human activity
such as when ceramics or flint materials were heated to improve
strength and durability. ESR also measures electrons trapped in the lat-
tice, but unlike thermoluminescence it does not destroy the sample
being tested, allowing samples to be dated multiple times. ESR mostly
tests calcium carbonate in limestone, coral, fossil teeth, molluscs, egg
shells, quartz, and flint. Both methods are typically used to date sam-
ples from the last 300,000 years, though theoretically they could be
used on much older samples.
25 per cent increase in precipitation during the African Humid

Period, between 15,000 and 5,000 years ago. On the other hand,
as hydrological modelling suggests, large water bodies buffer
rapid shifts in climate due to their delayed response to changes in
the precipitation–evaporation balance. Thus, theoretically, lakes
can be very quick to form, but their influence on the local climate
creates an inertia that resists their removal or disappearance.
Subsequently other groups have supported our work, find-
ing more evidence for the role of precession. For example, the
geochronologist and Chelsea fan Alan Deino (Berkeley Geochron
ology Centre) and palaeoanthropologist John Kingston (University
of Michigan) have found that the major lake period found in the
Baringo Basin in the Central Kenyan Rift, between 2.7 and 2.55
million years ago, actually consisted of five tick diatomite deposits
125
representing distinct palaeo-lake phases separated by a the preces-

sional cyclicity of ~21,000 years. Clayton Magill and colleagues at
Pennsylvania State University have found biomarker stable carbon
isotope evidence (see Box 1) in Olduvai lake sediment of preces-
sional forced variations between open C4 grasslands and C3 forest
between 1.8 and 1.9 million years ago. There is also evidence for
precessional forcing of the lake phase 1.9–1.7 million years ago
identified in the KBS Member of the Koobi Fora Formation in the
North East Turkana Basin in Kenya. During the same period, an
oxygen isotope record from the Buffalo Cave flowstone in the
Makapansgat Valley, Limpopo Province, South Africa, shows clear
evidence of precessionally forced changes in rainfall. The occur-
rences of these environmental changes are in phase with increased
freshwater discharge into the Mediterranean Sea, shown by the
sapropel records, and out of phase with the dust transport records
in sediments from the Arabian Sea, tropical North Atlantic Ocean,
and Mediterranean Sea.
In summary, then, the palaeoclimate records from East Africa
and surrounding regions show that there were distinct periods
when the local climate became highly variable. These periods
coincide with the approximately 400,000 years maxima in eccen-
tricity. During each of the periods, the climate of East Africa
cycled between extremely wet periods, when the whole of the
Rift Valley was full of deep freshwater lakes, to hyper-arid condi-
tions when huge amounts of dust were blown into the surrounding
oceans. This extreme climate variability varied on a 20,000 year
frequency, suggesting that precession forces the local climate to
switch between extremely wet and dry phases. These periods of
highly variable climate seem to only last 100,000 years. In
between there are long periods of time when there seems to have
126
been very little change in the East African climate, and no lakes,
apart from large permanent ones such as Lake Turkana.
These short periods of extreme climate variability seem to cor-
relate with key periods of hominin evolution, such as the first
appearance of A. ramidus (4.4 million years ago); of the genus
Homo (2.8 million years ago) and the genus Paranthropus (2.5 mil-
lion years ago); of P. robustus, H. erectus, and H. ergaster (1.8 million
years ago); of Homo heidelbergensis (700,000 years ago); and of
H. sapiens (200,000 years ago).
In Chapter 7, we will look at how this new understanding of
past environmental changes in Africa are shaping theories of
how and why hominins evolved.
127
7
African Climate Pulses

-
I n previous chapters we have seen the huge changes to the land-

scape of East Africa wrought by tectonics, celestial mechanics,
and climate change. The hominin fossil record gives us brief
glimpses of our different ancestors and when they first and last
appeared in Africa. In this chapter we will look briefly at the dif-
ferent theories that have been put forward in which a changing
environment is the key control on hominin evolution. My col-
leagues and I have developed a new synthesis of the palaeoenvi-
ronmental data from East Africa that allows us to bring together
all the different theories of human evolution into one framework.
It seems as if no single theory may be right, but I instead will
argue that all of them may be right.
Theories of evolution
As we have seen, environmental pressures have long been assumed

to play a key role in hominin speciation and adaptation. A num-
ber of iconic theories have been put forward to frame and develop
128
a fr ic a n cl im at e pu l ses
the discussion of hominin evolution. In Table 2 I have tried to put

these theories into the context of overarching theories of evolution.
There are two main schools of thought when it comes to over-
all evolutionary theory. The first is phylogenetic gradualism, based
on the idea of evolution presented in Darwin’s Origin of Species,
which suggests that most speciation is due to one species slowly,
uniformly, and gradually evolving into a new and distinct species.
However, most scientists would agree that there is little or no evi-
dence for this uniform rate of evolution, and evolutionary biolo-
gist Richard Dawkins argues that Charles Darwin himself was not
a constant-rate gradualist. For example, in the first edition of On
the Origin of Species, Darwin stated that ‘Species of different genera
and classes have not changed at the same rate, or in the same
degree’. Hence, we must differentiate between constant and vari-
able phylogenetic gradualism.
The second school of thought that developed from Darwin’s
original ideas was first proposed in 1972 by palaeontologists Niles
Eldredge and Stephen Jay Gould—they called their new theory
punctuated equilibrium. They propose that once species appear in
the fossil record they will become stable, showing little evolution-
ary change for most of their geological history—a state they
called stasis. When a rare and geologically rapid event occurs then
significant evolutionary change occurs through branching speci-
ation called cladogenesis. Cladogenesis is the process by which a
species splits into two distinct species, rather than one species
gradually transforming into another. When we look at the whole
geological record, life is dominated by extinctions and speciations
due to the five large mass extinctions, the last of which ended the
rule of the dinosaurs and ushered in the age of mammals. So,
with a very large timescale, punctuated equilibrium seems a good
129
table 2. Theories of evolution and how they correspond to the many postulated
theories of early human evolution
CLIMATE STRESS
Long-term state Threshold

None Variability
Constant
change event
Allopatic
Phyletic gradualism
Red Queen
speciation
Variability
EVOLUTIONARY FORM
Turnover pulse selection

Variable
hypothesis hypothesis
Red Queen Allopatic
speciation
Savannah Pulsed climate
hypothesis variability
hypothesis
equilibrium
Punctuated
Court Jester Court Jester Pulsed climate

Aridity
(e.g. impact Allopatic variability
hypothesis
event) speciation hypothesis
fit with our knowledge of the fossil record. But with smaller time-
scales, such as those we are investigating for hominin evolution,
punctuated equilibrium may not be so applicable. Of course, we
must remember that the split between phylogenetic gradualism
and punctuated equilibrium is artificial, but it does provide a
starting point for discussing theories of early human evolution.
Theories of early human evolution
Various evolutionary theories have sought to explain the evolu-

tion of bipedalism, encephalization (brain expansion), and our
ancestors’ dispersals out of Africa in terms of environmental
pressures. The first key environmental theory to explain bipedal-
ism was the savannah hypothesis, which suggests that as grasslands
expanded, hominins were forced to descend from the trees and
130
adapt to life on the savannah, which was facilitated by walking

erect on two feet. The savannah hypothesis has a long history and
has influenced the debate about early human evolution for over
200 years. It was first suggested in 1809 by the French naturalist,
Jean Baptiste Lamarck, in his book Philosophie zoologique:
If the individuals of which I speak were impelled by the desire to

command a large and distant view, and hence endeavoured to stand
upright, and continually adopted that habit from generation to gen-
eration, there is again no doubt that their feet would gradually
acquire a shape suitable for supporting them in an erect attitude.
(Lamarck 1809, 170)
This view was echoed sixty-two years later by Charles Darwin,

who also envisioned primeval humanity leaving the forest and
adapting to open plains. In his book Descent of Man he wrote:
As soon as some ancient member in the great series of the Primates

came, owing to a change in its manner of procuring subsistence, or to
a change in the conditions of its native country, to live somewhat less
on trees and more on the ground, its manner of progression would
have been modified; and in this case it would have had to become
either more strictly quadrupedal or bipedal. (Darwin 1871, 140–1)
The savannah hypothesis led to the knuckle-walking hypothesis,

which states that human ancestors used quadrupedal locomotion
on the savannah, as a precursor to bipedal locomotion. But, as
discussed at the end of Chapter 4, the fossil record suggests that
our ancestors did not knuckle-walk—instead, they retained long
arms needed for tree-climbing at the same time as they developed
bipedalism. For example, Ardipithecus ramidus, when it existed 4.4
million years ago in Afar in Ethiopia, was clearly able to travel
fairly long distances using its bipedal adaptation, but retained the
ability to climb trees that would have been essential for food gath-
ering and escaping the large numbers of predators in East Africa.
131
We also now know much more about the development of the

East African palaeoenvironment and the slow fragmentation of
the vegetation, as opposed to a simple binary switch between for-
est and savannah.
I can easily understand why scientists working in East Africa
could have seen the modern landscape and assumed it was a bin-
ary system. This is because as you fly up the Rift Valley there are
vast areas that look just like savannah, with mainly grass and a
few large acacia trees scattered around the landscape, with some
isolated areas of forest. But this landscape is 1,000 metres above
sea level, with an annual rainfall that could easily support open
forest. The reason it is grassland is because of deforestation and
constant grazing by cattle, goats, and camels. The worst offenders
for this type of deforestation are the goats, which can climb small
trees and strip plants instead of nibbling them, preventing the
plants from regrowing. Therefore much of the present-day land-
scape of East Africa is the result of human activity, making it
much harder for us to envisage the past. The current predomin-
ant view of the origins of bipedalism is that that it had an arboreal
origin. Our ancestors learnt to stand upright in the trees using
their forelimbs for support. Bipedalism was a response to the
need to move greater distances as efficiently as possible between
different food sources as the forest cover started to fragment and
change.
The savannah hypothesis was refined as the aridity hypothesis,
which suggested that the long-term trend towards increased aridity
and the expansion of the savannah was a major driver of hominin
evolution, particularly the evolution of Homo and encephalization.
This idea was encapsulated in a paper published in Science in 1995 by
the palaeoceanographer Peter deMenocal of Columbia University.
132
He used dust records in marine cores off the coasts of the Sahara
and Ethiopia to look at changes in aridity on the African continent,
and suggested that there were three distinct periods when aridifica-
tion of East Africa accelerated, related to the major thresholds in
the global climate system discussed in Chapter 5:
Major steps in the evolution of African hominids and other vertebrates

are coincident with shifts to more arid, open conditions near 2.8 Ma,
1.7 Ma, and 1.0 Ma, suggesting that some Pliocene (Plio)- Pleistocene
speciation events may have been climatically mediated. (deMenocal
1995, 59)
The aridity hypothesis was probably the first of the early human
evolution theories that embraced the evolutionary pattern of
punctuated equilibrium. For the next decade, this neat and straight-
forward theory became the textbook explanation for human evo-
lution, until it was challenged.
Another set of theories focused on how different types of spe-
cies react differently to environmental stress. The turnover pulse
hypothesis, suggested by Elisabeth Vrba at Yale University, was ori-
ginally developed in 1988 to explain discrete patterns in ungulate
(hoofed animal) speciation, and suggests that acute climate shifts
drove adaptation and speciation. She recognized that environ-
mentally induced extinctions hurt specialist species more than
generalist species. Hence, when there is environmental disrup-
tion the generalists will tend to thrive by utilizing new environ-
mental opportunities and by moving elsewhere to take advantage
of other areas that have lost specialist species. The specialists will
experience more extinctions, and an increased speciation rate
within their group. This would lead to more rapid evolution in
isolated areas—referred to as allopatric speciation (Figure 36)—
whereas the generalists will become more spread out.
133
figure 36 Different modes of potential speciation with and without geographical

or niche separation.
Rick Potts, the director of the Smithsonian Institution Museum

of Natural History’s Human Origins Program, later refined
Elisabeth Vrba’s theory and coined the variability selection hypoth-
esis. This theory advocates the role of environmental unpredict-
ability in selecting for behavioural or ecological flexibility. It
develops the original turnover pulse hypothesis, but instead splits
species in terms of their differing ability to adapt and evolve in a
more variable and unpredictable environment (Figure 37). The
variability selection hypothesis originally emphasized the long-
term trends towards a drier and more variable climate. However,
it struggled to explain the recent palaeoanthropological evidence
13 4
figure 37 Variability selection hypothesis of early human evolution.
suggesting a pulsed/threshold nature to hominin speciation and

migration events. So, over the past few years, Potts has revised the
variability selection hypothesis so that the selection for more
flexible adaptations occurs during the climatically variable peri-
ods of time.
More recently, it has been suggested that periods of climate sta-
bility may be equally important in driving human evolution, dis-
persal, and technological innovation. Relatively long periods of
climate stability could produce the scenario predicted by the Red
135
Queen hypothesis, or sympatric evolution due to sexual selection, dis-

cussed in Chapter 8.
Leigh van Valen, an American evolutionary biologist, proposed
the Red Queen hypothesis in 1973, which suggests that continued
adaptation is needed for a species to maintain its relative fitness
among co-evolving systems, and that biotic interactions, rather
than climate, are driving evolutionary forces. It is based on the
Red Queen’s race in Lewis Carroll’s Through the Looking-Glass, in
which the Queen says ‘It takes all the running you can do, to
keep in the same place’. The idea is that evolution is an arms race;
for example, as antelopes evolve to run faster so cheetahs have to
get faster too. However, for this to occur it is reasonable to
assume a relatively highly productive environment, so that com-
petition, rather than resources, forms the dominant control. At
Koobi Fora (northern Kenya) there is evidence for the presence
of multiple hominin species, including Paranthropus boisei, Homo
erectus, H. habilis, and H. rudolfensis, attributed to the period of
maximal lake coverage (~1.8–1.9 million years ago), and hence
the highest availability of resources. It might be postulated that
these hominins evolved as a result of competition with each
other and other animals. It is interesting to note that, at the end
of this period, only one species survives—Homo erectus. The
antithesis to the Red Queen hypothesis is the Court Jester hypoth-
esis, coined by the palaeontologist Anthony Barnosky at Berkeley
University. He suggested that abiotic forces such as tectonics and
meteorite impacts, rather than biotic competition between spe-
cies, are the major drivers behind the process of evolution.
Finally, a direct development of the variability selection hypoth
esis, which incorporates the role of both stability and instability
and challenges the aridity hypothesis, is the pulsed climate variability
136
framework. This theory came out of the original palaeo-lake work

of Martin Trauth, myself, and our colleagues, and highlights the
role of short periods of extreme climate variability in East Africa
in driving hominin evolution. This framework is discussed in the
next section, along with how the other evolutionary theories may
be applied, given the new environmental context. Instead of any
one of these theories being the right one, they could all play an
important part in the story of early human evolution.
Pulsed climate variability
Our pulsed climate variability framework suggests that there

were periods of extreme climate variability every 400,000 or
800,000 years, driven by the eccentricity maxima, when lakes
grew and filled much of the Rift Valley, and then disappeared on
an approximately 20,000-year precessional timescale. These
periods of highly variable climate in East Africa seem to coincide
with the majority of the first and last appearances of hominin
species over the past 5 million years. A period of highly variable
climate between 4.6 and 4.4 million years ago coincides with the
ages of A. ramidus discovered at As Duma in the Afar region of
Ethiopia. Another highly variable climate period between 3.4
and 3.2 million years ago coincides with the record of A. afarensis
at Hadar, Ethiopia, covering a period from about 3.4 to 3 million
years ago. Between 2.7 to 2.5 million years ago there is another
period to which the five diatomite deposits of the Baringo Basin
have been identified. These correspond to the peak in precession
and the age, about 2.8 million years ago, to which the first Homo
species have been identified, as well as to the appearance of the
Paranthropus genus, 2.5 million years ago. The largest and most
137
prolonged period with variable climate is between 2.1 and 1.7

million years ago. This coincides with distinct variants of H.
rudolfensis and H. habilis identified by Susan Anton of New York
University and her colleagues, at 2.09 million years ago, the first
appearance of H. erectus 1.9 million years ago, and the first disper-
sal of hominins outside Africa about 1.8 million years ago. Then,
1.1 to 0.9 million years ago, another highly variable period coincides
with the last appearance of H. erectus in East Africa and the second
dispersal of hominins out of Africa. Recent modelling work by Rick
Potts and Tyler Faith at the University of Queensland suggests that
these correlations are not down to sampling bias or coincidence.
Fundamental to understanding what evolutionary mechan-
isms could have applied to hominins in East Africa are the speed
and form of the transitions between lakes appearing and
disappearing from the landscape. At first, it may appear that
orbitally forced climatic oscillations may be too long-term to
have created rapid changes in lake occurrence. However, this does
not take account of the sinusoidal nature of orbital forcing or the
threshold nature of the African lake systems. All orbital param-
eters are sinusoidal, which means that there are periods of little or
no change followed by periods of large change. For example, the
sinusoidal precessional forcing at the equator consists of periods of
less than 2,000 years, during which 60 per cent of total variation in
daily insolation and seasonality occurs. These are followed by
~8,000 years when relatively little change in daily insolation occurs.
Hence, precession does not result in smooth forcing, but rather
produces a combination of brief periods of strong forcing and long
periods of relatively weak forcing. If this is combined with the idea
that many of the East African lakes are amplifier lakes that respond
very quickly to a small increase or decrease in the precipitation–
138
evaporation balance, then it is relatively easy to envisage threshold

responses of the landscape to precessional forcing.
Katie Wilson, my former doctoral student at University College
London, showed that the diatomite deposits in the Baringo
Basin do indeed suggest that the lakes appear rapidly, remain
part of the landscape for thousands of years, then disappear in a
highly variable and erratic way. In fact, the absence of shallow-
water diatom species at the start of many palaeo-lake deposits
suggests that the lakes could have appeared in only a few hun-
dred years. Figure 38 shows a compilation of what a generic
extreme wet–dry cycle may have looked like, with a threshold at
the beginning of the wet phase and a prolonged highly variable
period at the end of the wet phase. There would be four or five
of these cycles during each of the periods of extreme climate
variability.
figure 38 Pulse climate variability framework.
139
The different speeds at which the lakes appeared and disap-

peared is also consistent with the idea of a bifurcated relationship
between climate and lake presence. Figure 39 shows that precipi-
tation needs to increase significantly before lake growth can initi-
ate, but once it has, there are some key feedbacks which accelerate
the expansion of the lakes. The most important is the change in
the local climate due to increased moisture in the atmosphere.
Hence, increased local relative humidity reduces the amount of
evaporation occurring, thus increasing the moisture in the atmos-
phere. When lakes become more established, the general increase
in moisture changes the vegetation and more bushes and trees
appear, subsequently increasing the evapo-transpiration and fur-
ther increasing the moisture in the atmosphere. These same feed-
backs also resist the drying out of the lake when precession starts
to reduce overall rainfall. This leads to a period of up to 2,000
years when the lake expands and contracts, until finally there is
not enough moisture in the region to sustain any sort of lake.
figure 39 Relationship between growth and shrinkage of large lakes in the

African Rift Valley. The bifurcation is caused by the evaporation from the lake
making the surround region wetter and thus resilient to initial reductions of overall
precipitation.
1 40
More recent evidence of this sort of lake behaviour has been

found by palaeoliminologist Annett Junginger, now at the
University of Tübingen in Germany, and Martin Trauth. Between
them they compiled a large number of radiocarbon dates (see Box 3)
for palaeo-Lake Suguta in northern Kenya (see Figures 32 and 33).
These show that the lake appeared rapidly 14,700 years ago,
achieving a depth of 300 metres in less than a few hundred years.
The lake survived until about 7,000 years ago, and then took
about 2,000 years finally to dry up completely.
It should be stressed, however, that this pulsed climate variabil-
ity framework, whereby eccentricity-modulated precession is the
main influence, only applies up to 800,000 years ago. After this
time, as mentioned in Chapter 5, the Early–Middle Pleistocene
transition occurred, intensifying and prolonging the glacial–inter-
glacial climate cycles, which thus have an increasing influence on
tropical climates. Hence, over the past 800,000 years the climate
of the tropics became more complicated and fragmented as it was
influenced both by localized influences of orbital forcing, and the
ever stronger global influence of the glacial–interglacial cycles.
Human evolution within the new framework
The big problem with most previous theories of human evolution is

that they have no timescale over which they operate. This is import-
ant because a slow environmental change over one hundred gener-
ations is going to have little or no effect on the evolution of a species.
However, the drying up of a lake over a few generations will have an
effect on the survival of individuals, and thus potentially drive evo-
lution. It seems harsh, but evolutionary change is equally about
those individuals and their traits which do not survive as it is about
1 41
those that make it through. So the pulsed climate variability frame-

work provides a temporal framework for key periods of time when
we know that new species of hominin evolved, and so it can helps us
to conceptualize the different theories of early human evolution.
Figure 40 shows how the turnover pulse hypothesis would operate
through one of these extreme climate cycles.
The palaeontologist and zoologist, Elisabeth Vrba, suggested
that environmental changes would affect specialist and generalist
species differently. During dry periods, the extinction rates of
specialists would go up as they would struggle to find food, having
lost their environmental niche and their competitive advantage.
In contrast, the generalists would have a lower extinction rate, as
they would be better adapted to finding resources during stressful
periods (Figure 40). Speciation would be much higher among the
specialists during dry periods, as they adapt to new habitats. In
contrast, during the wet periods, and to a lesser extent the highly
variable periods, generalist species would suffer as specialists
would have a lot more niches to fill and thus would outcompete
the generalists. Figure 41 illustrates possible changes that could
have occurred due to the aridity hypothesis, which suggests that
speciation mainly occurs during periods of dryness with low
resources. Extinctions would have occurred during both wet and
dry periods, but for different reasons. They would have occurred
during dry periods as some species would not have been able to
adapt to the harsh conditions, which conversely drives speci-
ation. There would have also been extinctions during the wet
period as some of the species that adapted to the harsh dry condi-
tions may not have retained the ability to cope with the wet plen-
tiful periods, or may have been outcompeted by those more able
to take advantage of a high-resource environment.
1 42
figure 40 An interpretation of how the turnover pulse hypothesis could be

placed within the pulsed climate variability framework.
The variability selection hypothesis developed from the ori-

ginal turnover pulse hypothesis, but instead splits species into
their varying ability to adapt and evolve in a more variable and
unpredictable environment. Hence, generalists undergo more
extinction, and specialists more speciation, during the highly
1 43
figure 41 An interpretation of how the aridity hypothesis could be placed

within the pulsed climate variability framework.
variable climate period in between the long wet and arid phases.
In contrast, the Red Queen hypothesis suggests that continued
adaptation is needed in order to keep up with other species which
are also evolving. It is therefore reasonable to assume that a rela-
tively high-energy environment would provide more resources
and therefore more energy in the system to enable interspecies
competition.
Finally, the visual example in Figure 42 illustrating allopatric
evolution suggests that by geographically isolating populations,
144
figure 42 An interpretation of how the allopatric speciation hypothesis could

be placed within the pulsed climate variability framework.
each can then evolve independently with slightly different environ-

mental stresses. In the Rift Valley, during the extreme dry periods,
north–south and east–west migration is very difficult, so it would
have created isolated populations. The same is true of extreme wet peri-
ods, because when lakes completely fill the rift basins migration
north–south and east–west would again be difficult, also creating
isolated populations. Only during the high-variability period and
the threshold change would it be possible to move up and down
and across the Rift Valley easily. Recent evidence from Katie
1 45
Wilson’s work suggests that there were millennial-scale fluctu-

ations in lake levels during the extreme wet periods, probably
driven by the northern ice sheet collapse events. This may have
made movement between populations possible during the wet
phases, limiting the isolation.
The illustrations shown in this chapter are just my interpret-
ations of how the major theories of human evolution could be
placed within the pulsed climate variability framework. This is
my attempt to place a temporal framework on evolutionary the-
ories. I would encourage you to use this visual approach to play
with your own interpretation of how changing environments
would interact with different theories of early human evolution.
We should remember, however, that there are long periods of sta-
bility ranging from 200,000 to 400,000 years between the high-
variability climate periods in which evolutionary events could
have occurred. Moreover, as we will examine in Chapter 8, the
evolution of our brain may have also helped drive our evolution
independently of environmental change.
Hominin dispersals
The periodic hominin dispersal events also seem to correlate

with these periods of high climate variability. Dispersal events
seem to occur at about 3 million years ago within Africa, and at
1.8, 0.9, 0.6, and after 0.1 million years ago out of Africa (Figure 8).
It has been suggested that both lake presence and absence could
be associated with these dispersal events. For example, when the
lake basins are dry they become ‘hyper-arid’; thus the Rift Valley
becomes uninhabitable, and hominin populations would have
been forced to migrate to the north and south. However, severe
1 46
lack of resources would mean there was only a small and pos-
sibly shrinking population that could migrate. The absence of
lakes may have facilitated allopatric speciation in key refugia
such as Turkana, which may have remained wet. Dispersal is
thus more likely to have occurred when the basins were com-
pletely filled with water, as there would have been limited space
for the hominin populations on the tree-covered Rift shoulders
and river flood plains. The wet conditions could have been more
conducive to dispersal because hominin populations could
expand due to the availability of water and food, and could fol-
low the Nile tributaries northward and through a green Levant
region. There is some clear evidence that when it is wet in East
Africa it is also wet in the Levant and the Middle East, opening
up multiple corridors out of Africa and into the East. So the
occurrence of deep freshwater lakes would have forced expand-
ing hominin populations both northwards and southwards, gen-
erating a pumping effect, pushing them out of East Africa
towards the Ethiopian highlands and the Sinai Peninsula, or into
Southern Africa, with each successive precessional cycle. The
recent finding that the wet periods were also variable suggests
that there could have been small expansions and contractions of
the Rift Valley lakes on a millennial scale, enhancing this pump-
ing effect of hominins out of Africa.
The pulsed climate variability framework therefore takes the
latest palaeoclimate understanding of East Africa and provides a
framework within which to understand the causes of early human
evolution. There seem to have been periods of highly variable cli-
mate in East Africa, occurring every 400,000 or 800,000 years.
Within these periods the climate of East Africa and the Levant
seems to vary from extremely dry, with large quantities of dust
1 47
blowing into the adjacent oceans, to extremely wet, when the Rift
Valley basins were filled with deep freshwater lakes. These vari-
ations seem to have been driven by precession. The dry periods
may have lasted up to 8,000 years, and then lakes appeared on
the landscape very rapidly, maybe within a few hundred years.
The wet periods also lasted up to 8,000 years, and there is evi-
dence that the lake levels fluctuated up and down, driven by
millennial-scale events probably originating in the North Atlantic
Ocean. At the end of the wet period, the lakes took nearly 2,000
years finally to dry out, during which time lake levels were highly
variable. This means that there were environmental fluctuations
on the correct timescale to influence evolution. Different species
or, at the very least, different emerging traits within a species,
could have evolved through various mechanisms, including the
turnover pulse hypothesis, aridity hypothesis, variability selec-
tion hypothesis, or allopatric speciation. I argue that this is exem-
plified by the case of Homo erectus, which exhibited changes in life
history (shortened interbirth intervals, delayed development),
pelvic morphology, body size and dimorphism, a shoulder morph-
ology that enabled projectile use, adaptation to long-distance
running, ecological flexibility, social behaviour, and a new stone
tool kit allowing the processing of food. Each one of these traits
could have been forced by a different evolutionary mechanism
operating at a different part of the environmental cycle. So what
is emerging from our new understanding of the hominin fossil
record and the palaeoenvironment of East Africa is a much more
complicated and rich picture of how our species evolved.
In Chapter 8, we will consider how a large brain could have
helped our ancestors cope with these rapid environmental
changes.
1 48
8
The Social Brain

-
T he previous chapters give us some insight into the rapid envi-

ronmental changes that occurred in East Africa coincident
with the emergence and disappearance of hominin species. It is
easy to imagine that many of these environmental changes would
have created stress due to reduced access to food and water, and
would have created positive selective advantages for those ances-
tors whose particular traits enabled them to maintain access to
essential resources. The fossil record seems to suggest that there
were many different hominins roaming throughout Africa at the
same time, providing fertile ground for evolution. At critical peri-
ods, when populations of different hominin species were geo-
graphically together—such as on the shores of Lake Turkana 1.8
million years ago—we can imagine species to species competi-
tion and possibly interbreeding. This species to species competi-
tion may simply have been about winning or losing access to
food and water. Or it may have led to direct conflict. The seminal
work of the renowned primatologist Jane Goodall on chimpan-
zees shows that they can sustain long-term intraspecies warfare
1 49
t he soci a l br a in
until one group of male chimpanzees is wiped out. There may

have also been interbreeding between hominin species—for
example, recent genetic studies show that there was some inter-
breeding between H. sapiens and H. neanderthalensis. There is, how-
ever, another evolutionary driving force that needs to be
examined—sexual selection. In this chapter I examine how this
may have driven the evolution of a larger brain through the need
to control large social groups.
What good is a large brain?
It may be possible that a large brain became a key characteristic in

sexual selection, when both males and females actively selected
partners with traits associated with a bigger brain. This would
have lead to sympatric evolution (Figure 36), when new species
emerge from within the original population and then disperse.
The social brain hypothesis suggests that enhanced cognitive ability
would provide the ability strongly to influence groups or tribes of
hominins, and hence control the distribution of resources, and
thus it would be strongly selected for, driving brain expansion in
hominins.
Originally, large brains were thought to be essential to enable
the making of stone tools, and this is why H. habilis was thought
to be the start of the Homo genus. We now know that many ani-
mals make and use tools, and that stone tools originated much
earlier than Homo. So what was the large brain used for if it was
not essential for tool-making? Anthropologist Robin Dunbar and
colleagues at Oxford University argue that it takes a huge amount
of cognitive ability to exist in large social groups. Our position in
the group, our access and ability to get mates and resources, rely
150
on how good we are at playing the social game. As the groups get
larger, so the computational power needed to keep up with the
interconnections grows exponentially. For example, if you are
talking to your best friend about the weather, this is a simple lin-
ear conversation with little brainpower needed. If you now talk to
your best friend about their mother you need to start to think.
First, what is their relationship to their mother? Has it been good
or bad in the past, and what is like now? What is your relationship
like with their mother, and how does this relate to your relation-
ship with your best friend? Let’s take it up a notch—how about a
simple question to your best friend about how their mother is
getting on with her husband, who is in fact your best friend’s
stepfather. This is a simple example with someone you know
well. Imagine what happens when it is someone you know less
well and who you want to impress.
I have to say I am very mean when I illustrate this to my first-
year undergraduates—usually by asking one who has been repeat-
edly late to my lectures a set of simple questions. Remember, this
person is answering a professor in control of a class of 130 stu-
dents. The first question, ‘How are you?’, usually gets a simple
answer and sometimes a bit of humour such as, ‘I was better five
minutes ago’. The second question, ‘Do you like my class?’, is a dif-
ficult one because they want to appease me, but also do not want
to be seen as weak or cowering before me. The third question is,
‘Do you like this person sitting in the second row?’ (I point to that
person). They now have to decide what their relationship is to me,
to the person in the second row, and what they wish to tell me and
the other 128 students. Because in complex social situations the
truth is extremely fluid, the answer they give is usually one to
make them look good or simply extricate themselves from the
151
nightmare situation I have put them in. The fourth question is, ‘Do
you think they would go out with your friend sitting next to you?’
This one completely throws them as they have to work out their
answer based on their relationship with me, their friend, the per-
son sitting on the second row, and the other 127 students. Nine
times out of ten the student will use humour to get out of the situ-
ation. But remember, all these stressful social replies have to be
worked out in milliseconds so that the conversation continues.
So social groups are complex, with high stress levels, because
the rewards are high. Hence, our huge brain is developed to keep
track of rapidly changing relationships. My undergraduates come
to university thinking they are extremely smart as they can do dif-
ferential equations and understand the use of split infinitives. But I
point out to them that almost anyone walking down the street has
the capacity to hold the moral and ethical dilemmas of at least five
soap operas in their head at any one time, and that is why we have
a huge complex brain. And our brain is truly complex because to
just understand how the brain maintains our human intellect, we
would need to know about the state of all 86 billion neurons and
their 100 trillion interconnections, as well as the varying strengths
with which they are connected, and the state of more than 1,000
proteins that exist at each connection point. But neurobiologist
Steven Rose suggests that even this is not enough as we would
need to know how the connections have evolved over a person’s
lifetime and even the social context in which they had occurred. In
The New York Times, the neuroscientist Kenneth Miller suggested it
will take ‘centuries’ just to figure out basic neuronal connectivity.
This new understanding of the human brain and its capacity and
complexity has also changed our views about computers. In the
1970s and 1980s it was thought that artificial intelligence (AI) was
152
catching up with humans. With all our new knowledge of the

human brain and how complex it is, AI is now even further behind
than we thought. So, for all the recent hype, the takeover of the
world by sentient robots is scientifically even further away than it
was when initially envisaged.
However, the invention of computers has caused another prob-
lem, which is that many scientists think the human brain oper-
ates like a computer. However, Robert Epstein, a psychologist at
the American Institute for Behavioral Research and Technology,
says this is just shoddy thinking and is holding back our under-
standing of the human brain. He points out that humans start
with senses, reflexes, and learning mechanisms. What we do not
start with and never have are: information, data, rules, software,
knowledge, lexicons, representations, algorithms, programmes,
models, memories, images, processors, subroutines, encoders,
decoders, symbols, or buffers—which are key design elements that
allow digital computers to behave somewhat intelligently. Robert
Epstein argues that the ‘information processing’ metaphor of
human intelligence is completely wrong and is holding back the
whole of neuroscience. As he bluntly puts it, ‘We are organisms,
not computers. Get over it.’ The final difference is that computers
store exact copies of data that persist for long periods of time,
even when the power is switched off, while our brain only main-
tains our intellect as long as it remains alive.
I would argue that the incredible flexible and complex hominin
brain evolved to deal with complex social groups so that an indi-
vidual could maximize their access to the best resources and the
opportunity to mate with the best partners. I also do a small
experiment each year with my first-year undergraduates to show
how sexual selection for bigger brains may have operated. So
153
every year, for the past twenty years, in my human evolution lec-
tures, I asked my undergraduates questions about how they go
about choosing a mate. As you can imagine there is a lot of snig-
gering. I randomly ask female undergraduates what traits they
look for if they are interested in a male partner. The answers are
usually humour, intelligence, charm, etc. Now these are great, but
I point out that I cannot see how those qualities would have
helped our early ancestors on the plains of Africa when faced by
a hungry lion. Would they not have preferred a ‘man who throws
spear furthest in a pub’, which sets off more laughter. This line of
questioning has a serious point because, though human societies
value good looks and youth, they also highly value humour, intel-
ligence, and individuals with access to large resources. This is
because those traits show us who are alphas in society with the
ability to control large groups of humans and thus co-opt a
greater share of resources for themselves and their family.
Extreme examples of this power are top comedians who can fill
stadiums and control and entertain 70,000 people, or politicians
who, through their rhetoric and charm, convince millions of us
to vote for them so they can run our countries.
Group size: the magic number 150
Robin Dunbar has created a magic number: 150. He and his col-
leagues found that modern humans like communities of about
150 people. This number is found in the population of Neolithic
villages (6,500–5,500 bc), villages in the Domesday Book (1085
ad), eighteenth-century English villages, modern hunter-gatherer
societies, Anglican church congregations, and Christmas card
distribution lists—to name just a few examples. He has also
154
shown that community size in primates is linked to the size of the

neocortex region of the brain, where all cognitive processing occurs.
If one extrapolates the relation based on the other extant pri-
mates then humans with their very large neocortex extend the
graph to a community size of 150. This leads Robin jokingly to say
that anyone declaring that they have more than 150 friends on
Facebook is lying. Through detailed analysis of Twitter communi-
ties (those who exchange posts with each other, based on a sample
figure 43 Estimated community size of different hominin species over the last
four million years. Community size is estimated from converting cranial volumes
to neocortex ratios which have a strong relation to group size in primates. Median
shown with 50 per cent and 95 per cent range. Solid line is the major 150 community
size found within modern human societies.
155
of 1.7 million people) and email communities (those that exchange

regular emails based on a sample of 10 million people) his team
showed that these virtual communities also usually consisted of
between 100 and 200 individuals.
This relationship between community size and the size of the
neocortex is extremely useful, as it means we have a way of esti-
mating group size in extinct hominins. Hominins such as the
Australopithecines and early Homo have an estimated community
or group size of fifty. The big change occurs with Homo erectus,
when group size jumps up to nearly one hundred; then Homo hei-
delbergensis at 130 and, of course, modern Homo sapiens at 150
(Figure 43). Finally, Dunbar has shown that the size of the group
in primates directly correlates with the amount of grooming. He
figure 44 Robin Dunbar’s circles of human relationships.
156
argues that living in large groups is extremely stressful, so grooming

is a way of lowering individual and group stress. This grooming, or
‘paying attention’ to each other in a group, also translates to humans
who spend most time communicating and interacting with peo-
ple we call friends. Or, to flip it around, our friendship group con-
sists of the people with whom we currently spend most time
interacting and socializing. Human acquaintances can also be
mapped in both directions from the magic 150—with inner circles
of closer and closer relationships, a core circle of five in the centre,
and then outer circles of 500 acquaintances, and 1,500 people for
whom we can put a name to a face (see Figure 44).
Our expensive brain
Being able to control, manipulate, and/or influence groups of

individuals must have provided high rewards because there are
also high costs associated with having a large brain. The first is
the problem and danger of giving birth to a baby with a large
head. This is called the obstetric dilemma. Chimpanzees give
birth relatively easily as the size of the head is significantly
smaller than the birth canal (Figure 45). Moreover, the baby
comes out face forward so that it is relatively easy for the mother
to pick up the baby and lift it directly on to the breast for suck-
ling. With A. afarensis, the changes in the hips to allow bipedal
locomotion, and the slightly larger head means the baby’s head
comes sideways through the birth canal. Again, it is relatively
easy for the mother to reach down, pick up the baby, and place
it on her breast (Figure 45). In H. sapiens, the head of the baby
relative to the birth canal is much larger, and so the baby head
has to twist twice to get through the hips. Like A. afarensis, a
157
figure 45 A ‘Midwife’s’ view of the birth canal in a chimpanzee, A. afarensis, and

a modern human. Note that human babies undergo two 90° twists and usually are
born facing their mother’s bottom.
modern human baby twists sideways to enter the birth canal,

but halfway through it has to twist again to get past the hip
bones. This means that the majority of H. sapiens babies are born
facing the mother’s bottom. This makes it very awkward for the
mother to reach down to pick up the baby and twist it round so
it faces her breast for suckling without getting it tangled with
the umbilical cord. Palaeo-anatomists think that H. erectus was
probably the first hominin that had the double twist birth,
which must have evolved due to a significant increase in brain
size. This does mean that childbirth in later Homo species was
158
more dangerous to the mother and infant than for apes and ear-
lier hominins. This is due to the tightness of fit of the baby’s
head and the double twist, which adds complications such as
the umbilical cord getting wrapped around the infant’s neck.
Even today with modern hospitals, childbirth can occasionally
be fatal for mother and/or child.
So there are two fundamental questions we have to ask to
understand why the obstetric dilemma occurred. First, why can-
not the heads of babies be smaller? At the moment about 30 per
cent of adult brain size is achieved, on average, at birth. It seems
that, in all primates, 30 per cent is the minimum required to pro-
duce a viable primate infant. This is supported by modern medi-
cal data, which suggests the prevalence of special educational
needs in children rises sharply in babies born prematurely. Hence,
producing small-brained babies seems a non-viable option. Also,
it is not just the head that can cause problems in childbirth but
also the shoulders and relatively large body weight of the baby,
both of which can lead to maternal tearing and haemorrhaging
and muscular or spinal damage of the offspring. However, birth
weight is the single biggest predictor of survival in early life; so
any reduction in non-brain body weight will adversely affect the
survival chances of the infant.
The second question is: why can’t the mother’s pelvis be larger?
The pelvis is limited by the need to allow bipedal movement, but
there is variation in human pelvis size both front to back and side
to side; so the mother’s pelvis in some cases could be larger. These
modern variations seem to be tied to local environmental condi-
tions and mothers’ access to nutrition as they are growing up. For
example, Jonathan Wells at University College London points out
that African and Asian women have a narrower pelvis than
159
European women in response to hotter, more humid environ-

ments, but correspondingly have slightly shorter gestation peri-
ods. Within the African population, short mothers have the
highest risk of obstructed labour, suggesting their pelvis could
have been larger if they had had the required nutrition for full
growth during their own development. There is now evidence
that the balancing act between ensuring that the baby is large
enough to survive early life and avoiding its becoming too big to
fit through the pelvis are negotiated through metabolic signals
between the mother and the foetus. Interestingly, Jonathan Wells
notes that the current large burden of maternal mortality around
the world is due partly to the underdevelopment of many moth-
ers’ pelvises because of undernutrition early in life, and partly to
increased foetal birth weight related to the emerging obesity epi-
demic resulting from rapidly changing diets—both of which are
disturbing the balance achieved by the obstetric dilemma.
During the period of rapid evolution, when brain size was
expanding significantly and the pelvis had to change shape to
accommodate it, the obstetric dilemma would have been a con-
tinual process and must have resulted in a large increase in mater-
nal deaths. So there must have been huge rewards for being smart
and having a bigger head for the selection pressure to continue.
This has been called the expensive brain framework (Figure 46), which
tries to understand the advantages of having enhanced cognitive
ability. One of the most interesting things about the complicated
childbirth in later Homo is that mothers would have required help,
called allomaternal care. So individual females who were more
socially adept would get more help, and they and their infants
were more likely to survive—another positive feedback loop driv-
ing the evolution of larger brains as a means of having greater
160
figure 46 Expensive brain hypothesis.
social influence. The expensive brain framework tries to under-

stand the positive effects, such as food production and sharing,
predation reduction, and allomaternal care, compared to the
negative impacts such as increased food requirement and mother
mortality. Underlying all of this is the social brain hypothesis,
which can be seen as an internal arms race to develop the higher
cognitive skills to enable greater social control. Clearly, with the
emergence of Homo erectus, H. heidelbergensis, and H. sapiens, the
positives outweighed the negatives. We can only speculate as to
whether this was driven by stressful environmental change in
East Africa, increased competition for resources within the species
as populations expanded, or competition with other species—or
all of these.
Language
One of the missing pieces in the story of human evolution is the

appearance and development of language, as the social brain
hypothesis implicitly requires complex social interaction—or, as
Robin Dunbar would say, the ability to gossip. I have not men-
tioned language before simply because we have no idea when it
appeared. Scientists have strong views on this, based on their dif-
ferent theoretical backgrounds and their views on how modern
human babies develop language skills. The views can be split into
two main camps. The first are continuity theories, which suggest
that language now exhibits so much complexity it could not sim-
ply have appeared from nothing. It must therefore have evolved
from earlier pre-linguistic systems among our hominin ances-
tors. At the moment, most linguistic scholars accept the under-
lying idea behind continuity theories, but they vary in how they
162
envisage language development. Steven Pinker, experimental

psychologist and cognitive scientist at Harvard University, is
among those who see language as mostly innate, having evolved
in a gradual way.
Other scientists support discontinuity theories, according to
which language is a unique trait that cannot compare with any-
thing found among non-humans, and must therefore have
appeared suddenly with Homo sapiens. Noam Chomsky, linguist
and cognitive scientist, formerly at MIT, is a prominent propon-
ent of discontinuity theory. He argues that a single chance muta-
tion occurred in one individual around 100,000 years ago,
instantaneously installing the faculty of language in a ‘perfect’ or
‘near-perfect’ form. This view is supported by Ian Tattersall of the
American Museum of Natural History, who argues that evidence
of symbolic reasoning does not appear in the archaeological
record until after 100,000 years ago, and these early cultural arte-
facts are evidence for the spontaneous invention of language.
I would, however, counter this by pointing out that there are
examples of symbolic reasoning prior to H. sapiens. For example,
Josephine Joordens and colleagues at Leiden University published,
in Nature, an engraved freshwater mollusc pseudodon shell found
at a site known as Trinil on the island of Java. It was almost cer-
tainly etched by Homo erectus around 500,000 years ago, showing
the ability for abstract design and thus symbolic activity. While in
Germany, beautifully preserved wooden spears have been found
that were crafted and used by H. heidelbergensis 400,000 years ago.
It is difficult to see how these weapons could have been crafted
without teaching, and that requires sophisticated communication
techniques. My own view is that it is difficult to see how expansion
of the brain and our ancestors’ cognitive abilities could have
163
occurred without some form of language. One lesson we have

learnt in studying human evolution is that the archaeological
record is incomplete, and many stages of development in fact
occured much earlier than our records show. Stone tools are now
associated with A. afarensis, so it would not surprise palaeoanthro-
pologists if evidence were found for symbolic behaviour when H.
erectus first appeared, which would then mean that language was
central to the development of the genus Homo.
Sexual behaviour and human evolution
Another facet of human behaviour, which may have co-evolved

with our large brain, is our mating strategies. Primates exhibit all
sorts of mating behaviour, including polygynous, multimale-
multifemale, and monogamous. One control on which behaviour
is selected is the degree of size difference between males and
females. The greater this sexual dimorphism, the more likely the
mating strategy is either polygynous or multimale-multifemale.
This can be shown by observing our closest living relatives—
chimpanzees and gorillas. According to calculations using the
rate of DNA mutations, it has been calculated that our common
ancestor with chimpanzees lived about 8 to 5 million years ago
(see Box 4). There is a large difference in size between male and
female chimpanzees, and they have a multimale-multifemale
mating system. Essentially, male chimpanzees have sex with any
female, all the time, and with any excuse. Hence, a female may
contain sperm from multiple partners at any one time. So, des-
pite close genetic ties to humans, chimpanzees have adaptations
for direct sperm competition. For example, their testicles are
huge, to facilitate massive production of sperm multiple times a
164
figure 47 Comparison of the great apes and humans relative body size and
sexual organs.
day. A chimpanzee’s testes weigh more than a third of its brain

(Figure 47). In contrast, human testes are only about 3 per cent of
the weight of an adult’s brain.
There is also a large difference in size between male and female
gorillas, but they have a polygynous or harem-style mating sys-
tem—that is, many females to a single male. Gorillas, like humans,
therefore have relatively small testes. This suggests that sperm
competition is not the primary evolutionary driving force and
thus there have been no special adaptations of the male repro-
ductive system to facilitate this. This is similar to what is found in
modern humans, whose reproductive glands or testes are of a
very modest size (Figure 47); their sperm count reduces by more
165
than 80 per cent if men ejaculate more than two or three times a
day. Some experts have discussed whether the human penis is
exceptional in its size, and it does seems so when compared with
those of our closest primate relatives—chimpanzees, gorillas,
and orangutans. However, primatologist Alan Dixson (Victoria
University, New Zealand) in his wonderfully detailed book,
Primate Sexuality, suggests that if we look at all primates, includ-
ing monkeys, this is just wishful thinking. Comparative meas-
urements across all primates show that the human penis is not
exceptionally long, and its only unusual feature is its relatively
large circumference—though even this trait is not unique. It is
only large in comparison with the great apes (Figure 47).
If we compare human sexual dimorphism with that of all other
primates it suggests an evolutionary background involving a sig-
nificant degree of polygynous, rather than exclusively monogam-
ous, mating. This is supported by anthropological data studying
recent human populations, which shows that most populations
engage in polygynous marriage. Anthropologists Clellan Ford
and Frank Beach, in their book Patterns of Sexual Behavior first pub-
lished in 1951, suggested that 84 per cent of the 185 human cul-
tures they had data on engaged in polygyny. However, it is important
to note that it is usually only high-status or wealthy men who are
likely to have polygynous marriages; the majority of people
engage in monogamy, even though they are in a polygynous soci-
ety. In comparison, monogamy and serial monogamy is the main
system used by hunter-gather bands around the world, which
might provide a better analogy to our ancestors.
The problem is that the fossil record leaves no trace of our
ancestors’ sexual behaviour. However, comparing hominin sexual
dimorphism the fossil evidence is clear that the Australopithecus
166
and Parathropus genera had a much larger dimorphism than the

Homo genus. This suggests that early hominins may have had a
more polygynous mating system than later hominins. It is also
apparent that the sexual dimorphism of H. erectus was very similar
to that of modern humans, implying that human mating systems
may have evolved early on, at the same time as there was a signifi-
cant increase in brain size. Kit Opie and colleagues at University
College London analysed the behaviour of 230 primate species to
understand why a large proportion of human mating is monog-
amous or, at the very least, serial-monogamous. They argue that
social monogamy or pair-living is very uncommon in mammals—
less than 3 per cent—compared to birds, for which it is over 90 per
cent, though in primates a quarter of the known species do engage
in some form of social monogamy. Three main theories have been
put forward for social monogamy: parental care, mate guarding,
and infanticide risk. For males, it is advantageous to seek as many
opportunities to reproduce as possible, so something has to keep
males hanging around. Kit Opie’s analysis suggests that because
human infants are so vulnerable for so long, infanticide by other
males is a big risk. For example, it is well known that when a
younger male lion finally displaces the alpha male lion in charge of
a pride, they will systematically kill all cubs of the previous male
to ensure that all new offspring are their own. Hence, to ensure
that children are able to reach maturity the male is likely to stay to
protect them, both socially and physically, if needed. In contrast,
zoologists Dieter Lukas and Tim Clutton-Brock at Cambridge
University argue that it is less to do with infanticide but rather
linked to males being unable to defend access to multiple females.
If we view the evolution of monogamy or serial monogamy
mating systems in humans through the lens of the social brain
167
hypothesis, it is clear that it takes a huge amount of social effort

to maintain and protect more than one mate at a time. It is only in
more advanced societies, where males have access to wealth and
power, that they can protect multiple females, usually by paying
other men to protect them. So monogamy seems to be an adapta-
tion to protect one’s mate and children from other males. This is
re-enforced by the high social cost and stress of attempting to do
this for multiple partners, and it has been supported in complex
human societies by cultural norms.
The Homo sapiens problem
There are two problems with Homo sapiens. The first is that we
have very little idea why H. sapiens emerged about 200,000 years
ago. There is evidence that the climate of East Africa, after being
very dry and stable for 400,000 years, became more variable, and
this may have driven the small evolutionary changes from H. hei-
delbergensis to H. sapiens. But, in short, we really do not know. The
second problem is, for me, the more intriguing: that after H. sapi-
ens evolved in Africa and spread out into Europe and Asia they
seemed to do nothing special for the first 150,000 years. Over
these 150,000 years there are increasing records of symbolic
behaviour, starting with microliths, shell engravings, ochre, and
shell beads. But it is not until about 50,000 years ago that consist-
ent signs of creative thinking emerge—beautiful cave paintings in
Spain, France, and Indonesia, beautifully carved Venus figurines
in Germany, the Czech Republic, Austria, France, and Siberia,
and shell beads in North Africa and Europe. Around the same time,
modern humans appear that were more slender than their earlier
ancestors, and had less hair and smaller, less robust skulls—they
168
looked basically like us. But these changes weren’t just cosmetic
as the appearance of these smaller, more fine-boned humans was
accompanied by a revolutionary development of cumulative cul-
ture that led to the birth of agriculture and eventually to human
dominance of the planet.
There are, however, some tantalizing hints of what may have
happened from several very recent studies. The first is an analy-
sis of the fossilized skulls of our ancestors during this transi-
tional period, carried out by a team led by Robert Cieri at the
University of Utah and published in the journal Current
Anthropology. Cieri and colleagues found that the brow ridge (the
bony bit above the eye sockets) became significantly less promi-
nent and male facial shape became more similar to that of
females. They referred to this as craniofacial feminization, mean-
ing that, as H. sapiens slimmed down, their skulls became flatter
and more ‘feminine’ in shape. They think this must have been
due to lower levels of testosterone, as there is a strong relation-
ship between levels of this hormone and long faces with extended
brow ridges, which we may perceive today as very ‘masculine’
features (Figure 48). There is a second line of evidence that comes
from studying the relative finger lengths of our ancestors. There
seems to be a strong correlation between the ratio of the length
of the second (index) and fourth (ring) fingers to aggression,
promiscuity, and competitiveness in humans. The primary rea-
son for this relationship is because the digit ratio seems to reflect
prenatal testosterone levels. A hand with a shorter index finger
than the ring finger suggests higher testosterone levels. Emma
Nelson from Liverpool University and Susanne Shultz, now at
Manchester University, have found this relationship also holds
across the great apes and New World monkeys. Human digit
169
figure 48 Comparison of Homo sapiens skull morphology showing feminization

that has occurred since the Middle Stone Age.
ratios are intermediate between pair-bonded and the more pro-

miscuous great apes, supporting the idea that monogamy is
partly socially constructed to protect mates and children.
Moreover, there is a clear drop in digit ratio at the same time as
craniofacial feminization occurs, suggesting that there is a sig-
nificant drop in prenatal testosterone levels during this transi-
tional period.
People with lower levels of testosterone are less likely to be
reactively or spontaneously violent, and therefore this would
have greatly enhanced social tolerance. This has a huge knock-on
effect. As seen among humans today, we live in extremely high-
density populations with an incredible amount of social toler-
ance. So a reduction in reactive violence must have been an
essential prerequisite for our living in larger groups and develop-
ing a more cooperative culture. The idea that humans became
more feminine, less aggressive, and thus could cooperate in large
groups is certainly very intriguing as it would have allowed indi-
viduals with different skills to be valued and be reproductively
successful due to the reduction of, particularly, male–male vio-
lence. In most primates the physically strongest male tends to
170
dominate, but in early humans the smartest or the most creative

people may have come to the forefront.
Self-domestication of humans
The question remains: how did we become more feminine, less

violent, and more creative? A paper in the journal Animal Behaviour
by Brian Hare at Duke University and his colleagues may throw
some light on this. They compared chimpanzees (Pan troglodytes)
and bonobos (Pan paniscus) in West Africa—two closely related spe-
cies living in very similar environmental conditions either side of
the Congo River. One key distinction between the two species is
their sexual dimorphism. Male chimps are significant larger than
females, whereas the difference in bonobos is much smaller. This
difference is driven by different levels of testosterone. Size is just
one manifestation of deeper differences that show up in how the
animals interact with one another. Chimpanzees, particularly
males, are very aggressive; but violence within or between groups
is almost non-existent among bonobos. As both these species
have a common ancestor there must have been strong selection
going on to feminize the bonobos. Hare and his colleagues sug-
gest a process of self-domestication, whereby violent individuals
are punished and prevented from reproducing. The traits exhib-
ited by bonobos are very similar to the changes observed in spe-
cies that humans have domesticated, such as dogs, cows, guinea
pigs, and foxes. They postulate that the reason bonobos were able
to feminize and chimpanzees were not is because on the eastern
side of the Congo, where the chimps live, they are in direct com-
petition with gorillas, whereas the bonobos on the western side
have no competition. Harvard professor Richard Wrangham, a
17 1
co-author of the Hare paper, suggested the same process may

have happened to early humans.
Equality improves networking
This feminization through self-domestication may not only have

made groups of humans more peaceful, but may have also pro-
duced a more sexually equal society. A recent study in Science by
colleagues of mine at University College London showed that in
hunter-gatherer groups in the Congo and the Philippines deci-
sions about where to live and with whom were made equally by
both genders. Despite living in small communities, this resulted
in hunter-gatherers living with a large number of individuals with
whom they had no kinship ties. The authors argue this may have
proved an evolutionary advantage for early human societies, as
it would have fostered wider-ranging social networks, closer
cooperation between unrelated individuals, a wider choice of
mates, and reduced chances of inbreeding. The frequent move-
ment and interaction between groups also fostered the sharing of
innovations, which may have helped the spread of culture.
As Andrea Migliano, the leader of the study, points out, ‘sex
equality suggests a scenario where unique human traits, such as
cooperation with unrelated individuals, could have emerged in
our evolutionary past’. It may have only been with the rise of agri-
culture that an imbalance between the sexes re-emerged, as indi-
vidual men were suddenly able to concentrate enough resources
to maintain several wives and many children. Indeed, the Robert
Cieri-led study does show slightly more masculine facial shapes
emerging in recent agriculturalists relative to early humans and
recent human foragers.
172
So at the moment we have some tentative hints of what may

have happened between 50,000 and 10,000 years ago. Humans
may have undergone self-domestication and, over many gener-
ations, weeded out those individuals who were unable to control
their reactive violence. This is not as far-fetched as it sounds—
studies of the Gebusi tribe in Papua New Guinea by Bruce Knauft
showed significant levels of male mortality due to the tribe decid-
ing that if an individual’s behaviour was intolerable, then for
the good of the tribe they must be killed. So human proactive
violence—that is, thought-out, discussed, and planned violence—
is used to curb, control, and cull reactively violent individuals.
This process, combined with female mating choices which,
instead of selecting just for larger cognitive ability selected males
which dominated the group socially and not violently, over thou-
sands of years, could have resulted in males with lower testoster-
one and more feminine features, leading to a much more
gender-equal society and the start of our cumulative culture.
Dispersal of modern humans
The next stage in the H. sapiens’ takeover of the world was their
dispersal out of Africa into all the major continents, except
Antarctica. The evidence suggests that H. sapiens first evolved in
Africa about 200,000 years ago; they then dispersed out of Africa
into the Middle East about 120,000 years ago. New evidence from
fossilized teeth in China suggests that modern humans may have
made it all the way to China from between 100,000 and 80,000
years ago. Between 60,000 to 40,000 years ago, humans made it
into Australia. Then, about 40,000 years ago, humans made it
into Europe and started to compete for resources with Homo
173
figure 49 Dispersal of H. erectus, H. Neanderthalensis, and H. sapiens. Dates are in years before present and refer to the global dispersal of
H. sapiens.
neanderthalensis. Genetic studies have also shown that this

European expansion went east through what is now Russia, and
formed the base population which migrated into Mongolia and
Korea, and spread across the Bering Sea into North America,
Central America, and then South America (Figure 49). When
H. sapiens encounter other hominin species such as Neanderthals
and Denisovan there seems to have been multiple periods of
interbreeding which has lead to a genetic legacy of those species
within our own genome (see Box 4).
Neanderthals
Current evidence suggests that Neanderthals and humans shared

a last common ancestor sometime between 765,000 and 550,000
years ago. We speculate that this ancestor was H. heidelbergensis.
The story gets a little bit more complicated as there are also
Eastern Neanderthals, called Denisovans, whose genetics seem to
indicate a close affinity to European Neanderthals (Figure 50).
Evidence for H. denisovans has been found in Siberia, Asia, and
Melanesia. One major difference between early African humans
and Neanderthals was their diet. This is because the cold sparse
winters in Eurasia would have restricted plant availability, forcing
Neanderthals to rely more heavily on meat—indeed, chemical
analysis of their bones shows that it dominated their diet. It seems
Neanderthals were specialized seasonal hunters, and there is evi-
dence for them killing reindeer in winter and red deer in summer.
Neanderthal archaeological sites contain sharp wooden spears
and large numbers of bones of big game animals, showing evi-
dence of hunting and butchery. Neanderthals were, however,
adaptable, and at the Gibraltar coastal site there is evidence they
175
figure 50 Relationship between Homo sapiens and Homo erectus reconstructed by

Chris Stringer, based on the evidence we currently have from fossils and genetics.
exploited marine resources such as fish, molluscs, seals, and even

dolphins. They did also eat plants, and remains of starch grains
have been found in excavated Neanderthal molar teeth.
Neanderthals also had a sophisticated stone tool industry. This
differs from earlier ‘core tool’ traditions, such as the Acheulean
tradition of H. erectus. Acheulean tools are created by removing
flakes from the surface to ‘reveal’ a tool. In contrast, the
Neanderthal Mousterian stone tool industry is characterized by
flake tools that were detached from a prepared stone core. This
innovative technique allowed multiple tools to be fashioned from
a single suitable stone. Neanderthals used tools for activities such
as hunting and sewing. There is evidence for left–right arm asym-
metry, which was originally assumed to have developed from
the hunting technique of thrusting spears instead of throwing
them. This is in contrast to other hominins, such as Homo erectus,
176
H. heidelbergensis, and H. sapiens, that had shoulder adaptations to

allow them to throw weapons, and in the case of the later two
species there is archaeological evidence for thrown spears. The
Neanderthals’ style of hunting at close quarters has been used to
explain why Neanderthal bones have such a high frequency of
fractures, as these injuries are similar to those among profes-
sional rodeo riders who regularly interact with large, dangerous
animals. Colin Shaw from Cambridge University, however, has
suggested a less glamorous reason for asymmetry between the
arms. He suggests that Neanderthals may have spent hours scrap-
ing animal hides for clothing production, using stone tools. This
would have been an arduous and repetitive task, but essential in
the cold conditions in which they lived. The asymmetry in the
arms of Neanderthals has no directly analogy in modern humans
but some sports people, such as cricketers and tennis players, do
have very pronounced one-sided development.
One of the reasons Neanderthals are so interesting to palaeoan-
thropologists is because they are the first species to show evidence
of wearing clothes and of consistent symbolic behaviour. Scrapers
and stone and bone awls have been found, along with animal
bones, at Neanderthal sites. It is suggested the Neanderthals would
have used a scraper to first clean the animal hide, then an awl to
poke holes in it, and strips of animal tissue to lace together a
loose-fitting garment. It is only with modern humans that we find
evidence of the bone sewing needles needed to make tighter fit-
ting clothing. Neanderthals also controlled fire and lived in shel-
ters. There is evidence that Neanderthals deliberately buried their
dead and occasionally even marked their graves with offerings,
such as flowers. This may also explain why we have such a rich
history of Neanderthals, as the burials greatly enhance the
177
chances of preservation. They also made symbolic and ornamen-

tal objects such as grooved and perforated animal teeth, mam-
moth-ivory rings, decorated bone tools, and used colourants. No
other species had ever practised this sophisticated and symbolic
behaviour.
Neanderthals and modern Homo sapiens may have inhabited sim-
ilar geographic areas in Eurasia for nearly 50,000 years. They may
have had little direct interaction for most of this time as recent
genetic evidence suggests that the Neanderthal population was
very small compared with H. sapiens over the 400,000 years of their
existence. There is, however, clear genetic evidence that the two
species did interact at some stage. Scientists have recently sequenced
Neanderthal mitochondrial and nuclear genomes. When com-
pared with modern humans, it seems that many non-African peo-
ple have between 2–4 per cent Neanderthal ancestry, meaning that
Neanderthals and early humans must have interbred. It is also clear
that Europeans and Asians have inherited Neanderthal genetic
material, while some Asians have also inherited Denisovan genetic
material (Box 4). This admixture, though small, may have provided
adaptive advantages to non-African humans. Just a few thousand
years after modern humans moved into Europe, Neanderthal num-
bers dwindled to the point of extinction. All traces of Neanderthals
disappeared about 40,000 years ago. There is an endless debate as
to whether H. sapiens simply outcompeted Neanderthals, or
whether there was proactive culling, or both.
By 40,000 years ago we think that H. sapiens was the last homi-
nin species having outcompeted, interbred, and even killed off
the other hominin species. We humans are the only ultrasocial
creature on the planet. We co-habit in cities of tens of millions of
people and violence between individuals is extremely rare—even
178
Genetic revolution
Since the 1960s, the field of evolutionary genetics has arguably made
the greatest impact of any discipline upon human evolutionary studies.
Advances in technology, such as the development of the polymerase
chain reaction, which can amplify small quantities of DNA, have been
instrumental in this revolution, allowing, for example, the analysis of
ancient DNA taken from Neanderthal fossils. Allan Wilson, Emile
Zuckerkandl, and Linus Pauling pioneered the use of molecular
approaches, which examined evolution at the scale of DNA and
proteins, showing that relationships among living and extinct primates
can be inferred from genetics as well as fossils. DNA can also be used as
a ‘molecular clock’, involving comparison of the amount of genetic
difference (mutations) between living organisms. Since mutations
accumulate at approximately predictable rates over time, they can be
used to estimate how long ago two living species shared a common
ancestor. The molecular clock cannot assign concrete dates and must
be calibrated against independent evidence, such as the fossil record.
Nevertheless, bringing together the transdisciplinary evidence, we now
have a robust understanding of the relationships between humans and
apes. For example, we know that humans and chimpanzees split from
their common ancestor approximately 8 to 4 million years ago, and that
the genetic difference between humans and chimpanzees is miniscule,
at about 1.2 per cent.
Genetics has also confirmed that living humans have a limited
genetic diversity, indicating that there may have been a series of popu-
lation bottlenecks. These temporary, drastic reductions in population
size and therefore genetic variability may have been caused by rapid
climate changes, earthquakes, or even disease. Studies of genetic varia-
tion reveal that the greatest diversity can be found in African popula-
tions. This, combined with evidence from histories following the
female (mitochondrial DNA) and male (Y chromosome) lines, confirms
continued >
179
an African origin for our species and suggests that our direct ancestors
migrated out of Africa between 70,000 to 40,000 years ago.
Genetics has also contributed to our understanding of the complex
relationship and interbreeding between H. neanderthalensis, Denisovans,
and H. sapiens, pushing the boundary of DNA analysis to more than
400,000 years. It seems that there may have been at least five inbreed-
ing events in the last 0.5 million years (Figure 51). Genetics is demon-
strating that human evolution, rather than being a simple linear
evolution between species, is much more a network of slightly different
species interbreeding and exchanging DNA. We need to see human
evolution not as an evolution tree but more of a tangled bush with
many interconnections. We can speculate that the interbreeding that
figure 51 A history of interbreeding between early modern humans,

Denisovans, and Neanderthals.
180
occurred in the most recent hominins may have also occurred earlier in
our history, explaining why there were so many very morphologically
similar hominins coexisting in East Africa in the past. Genetics is also
making contributions towards revealing the evolution of human cog-
nition, for example language capacity (e.g. the FOXP2 gene) and identi-
fying human-specific neurological structures (e.g. the HARs, SRGAP2,
GPRIN2, GTF2I, and HYDIN2 genes).
though this is not the picture painted by the media. So when we

study human evolution we need to understand the cause of the
development of our extremely large flexible and complex ‘social
brain’. Of course, we can see many advantages in having a large
brain. First, it allows humans to exist in a group size of about 150,
which builds resilience to environmental changes by increasing
and diversifying food production and sharing. Humans have no
natural weapons, but living in large groups and having tools
allowed humans to become the apex predator, hunting animals as
large as mammoths. Larger groups also offer more protection
from other predators. Second, it allows specialization of skills
such as support for childbirth, tool-making, and hunting.
There is new genetic evidence by Aida Gomez-Robles and col-
leagues at The George Washington University that suggests the
modern human brain is indeed extremely flexible. They showed
that the human brain is more plastic and is modelled more by the
surrounding environment than that of chimpanzees. The anatomy
of the chimpanzee brain is strongly controlled by their genes,
whereas the human brain is extensively shaped by the environ-
ment, no matter what the genetics. This means the human brain
is pre-programmed to be extremely flexible; its cerebral organiza-
tion is adjusted by the environment into which it has been born
181
and raised. So each new generation’s brain structure could adapt

to new environmental challenges without the need to evolve
physically. This may also explain why we all complain that we do
not understand the next generation, as they have different brain
structures from us because they have grown up within a different
physical and social environment.
Having a large brain does not mean that culture and society
were inevitable—far from it, as there seem to be other require-
ments before human culture could start to build up. These may
have involved a reduction in reactive violence, and populations of
humans simply reaching a size at which inventions and new ideas
were not lost, and real accumulation of knowledge could occur.
In Chapter 9, I discuss what happened once this rate of accu-
mulation of culture started to accelerate.
182
9
The Future of Humanity

-
H uman evolution has clearly been heavily influenced by

tectonics and celestial mechanics through local environ-
mental change. Through the middle to the end of the last ice age,
H. sapiens dispersed into Australasia, Europe, and the Americas.
Once H. sapiens acquired cumulative culture, in the middle of the
last ice age, they started to have a larger influence on their local
environment. At the end of the last ice age, 11,000 years ago, agri-
culture first appeared in South West Asia, South America, and
north China. Independently, it then appeared 6,000–7,000 years
ago in south China and Central America, and 4,000–5,000 years
ago in the savannah regions of Africa, India, South East Asia, and
North America. There is evidence that this early agriculture influ-
enced atmospheric greenhouse gases, with the release of carbon
dioxide from deforestation and the release of methane through
wet rice cultivation (Figure 52). H. sapiens’ influence on the local
and global environment has accelerated through the industrial
revolution to the present day. Our influence is now so great that
scientists suggest we are a major force of geology, as powerful as
183
figure 52 Mass extinctions, land-use changes, and rising atmospheric greenhouse gas levels over the last
50,000 years.
t he f u t u r e of hu m a ni t y
tectonics or a meteorite impact. To acknowledge our dominant

role in the global environment, scientists have suggested we have
entered a new geological epoch called the Anthropocene. In this
chapter I will examine the evidence for the Anthropocene, and
what it means for human evolution.
Evidence for the Anthropocene
Are we really that important on planet Earth? Has, as Desmond

Morris called us, a naked ape really become the dominant geo-
logical power on the planet? Let’s look at the evidence. If we start
with the atmosphere there is clear evidence that human actions
have released 555 petagrams of carbon (where 1 Pg = 1015 g = 1
billion metric tonnes) into the atmosphere since 1750, increasing
atmospheric CO2 to a level not seen for at least 800,000 years,
and possibly several million years (Figure 52). There is even evi-
dence that these extra greenhouse gases have delayed the Earth’s
next ice age. The released carbon dioxide has also increased ocean
water acidity at a rate probably not exceeded in the past 300 mil-
lion years. Despite the focus on greenhouse gases due to concerns
about climate change, humans have also profoundly affected other
parts of the Earth system—for example, the nitrogen cycle. In the
early twentieth century, the invention of the Haber–Bosch pro-
cess allowed us to convert atmospheric nitrogen to ammonia for
use as fertilizer, revolutionizing agriculture and helping to feed
billions of people. But it has also altered the global nitrogen cycle
so fundamentally that the nearest suggested geological compari-
sons are events about 2.5 billion years ago.
There is clear evidence that anthropogenic greenhouse gases
are changing our climate. These changes include a 0.85°C increase
185
in average global temperatures and sea-level rise of over 20 cm in

the past 100 years. There is also evidence for significant shifts in
the seasonality and intensities of precipitation, changing weather
patterns, and the significant retreat of Arctic sea ice and nearly all
continental glaciers. It is estimated that Greenland is losing over
200 gigatonnes of ice per year, a six-fold increase since the early
1990s; Antarctica is losing about 150 gigatonnes of ice per year, a
five-fold increase since the early 1990s—and most of this loss is
from the northern Antarctic Peninsula and the Amundsen Sea
sector of West Antarctica.
Human drivers of evolution
Human actions are also affecting non-human life. Over recent

decades, global net primary productivity—the amount of new
plant growth each year—appears to have been relatively con-
stant. However, humans appropriate between 25 per cent and 38
per cent of this net primary productivity, which of course reduces
the amount available for millions of other species on Earth. In a
paper in Nature, Thomas Crowther and colleagues at Yale University
estimated that before the industrial revolution there were 6 tril-
lion trees on Earth, but now there are only 3 trillion trees left. This
deforestation and land-use conversion to produce food, fuel,
fibre, and fodder, combined with targeted hunting and harvest-
ing, has resulted in species extinctions some 100 to 1,000 times
higher than background rates, and probably constitutes the
beginning of the sixth mass extinction in Earth’s history. Species
removals are non-random, with a disproportionate removal of
animals with larger body sizes from both the land and the oceans.
This Megafauna Extinction began between 50,000 and 10,000
186
years ago, depending on when modern humans arrived on the

continent. Overall, during the Megafauna Extinction about half
of all large-bodied mammals worldwide, equivalent to 4 per cent
of all mammal species, were lost. The losses were not evenly dis-
tributed as Africa lost 18 per cent, Eurasia lost 36 per cent, North
America lost 72 per cent, South America lost 83 per cent, and
Australia lost 88 per cent of their large-bodied mammals. What
may surprise you is the massive shift from wild mammals to
humans and domestic mammals since this extinction began.
According to Vaclav Smil of the University of Manitoba, the bio-
mass of land mammals on Earth today is made up of 30 per cent
humans and 67 per cent domesticated animals; wild terrestrial
mammals make up just 3 per cent.
Organisms have also been transported around the world,
including crops, domesticated animals, and pathogens on land.
Similarly, boats have transferred organisms among once-discon-
nected oceans (Figure 53). Such movement has led to a small
number of extraordinarily common species, new hybrid species,
and a global homogenization of Earth’s biota. Just one example
of this is that nearly all the earthworms in North America are
European in origin because when they were inadvertently brought
over by the early colonists in the sixteenth century they outcom-
peted the local worms, and took over.
By far the biggest change was started in the sixteenth century,
when species from the Old World (Europe and Asia) were exchanged
with those in the New World (the Americas). This is known as the
Colombian Exchange. One result of the exchange was the globaliza-
tion of human foodstuffs. The New World crops maize/corn and
potatoes, and the tropical staple manioc/cassava, were subsequently
grown across Europe, Asia, and Africa. Meanwhile, Old World
187
figure 53 Shipping routes plotted from eighteenth- and nineteenth-century logs.
crops, such as sugarcane and wheat, were planted in the New World.
There was also movement of domesticated animals such as horses,
cows, goats, and pigs, all to the Americas. Human commensals, such
as the black rat, also made it to the Americas. There were other acci-
dental transfers from Europe too, such as smallpox, measles, and
typhus, which ended up killing over 50 million people in South and
Central America in the sixteenth century. These exchanges of plants
and animals continue today, and invasive species have become a
major concern on all continents. A study in Nature by ecologist Mark
van Kleunen and colleagues at the University of Konstanz suggests
that 4 per cent of plant species have been relocated around the globe,
equivalent to all the native plant species in Europe. These changes are
unique since the supercontinent Pangaea separated about 200 mil-
lion years ago, but Simon Lewis and myself at University College
London would argue that such trans-oceanic exchanges probably
have no geological analogue.
Human actions may well constitute the Earth’s most import-
ant evolutionary pressure. The intense harvesting of crops, the
movement of species to new habitats, and the selective pressure
of higher air temperatures resulting from greenhouse gas emis-
sions are all likely to alter evolutionary outcomes. Add to this
the development of diverse products, including antibiotics, pes-
ticides, and novel genetically engineered organisms, and humans
are now the major influence on evolution. Furthermore, given
that the average species survives between 1 and 10 million years,
the rates of anthropogenic environmental change in the near
future may exceed the rates of change encountered by many
species in the whole of their evolutionary history. So I would
suggest that human activity has clearly altered the lithosphere,
hydrosphere, cryosphere, atmosphere, and biosphere, and thus
189
we are changing the environmental and evolutionary trajectory

of the Earth.
Anthropocene paradigm shift
The physicist and philosopher of science Thomas Kuhn defined a

paradigm shift as a change in the basic assumptions within the
ruling theory of science. I would contend that within science the
evidence for humans being a major geological power has been
accepted, and the paradigm shift has already occurred. That
human activity has altered Earth fundamentally is rarely, if ever,
questioned in scientific publications. But I do want to be clear
that this shift in the scientific paradigm, though it is through
complex social processes, is a better, not just a different, under-
standing of the world. This is because a common misinterpret-
ation of paradigms is the belief that the discovery of paradigm
shifts and the dynamic nature of science is a case for relativism—
i.e. that science only has subjective value according to differences
in perception, consideration, or beliefs. Thomas Kuhn, and most
scientists, would vehemently deny this as the rational assessment
of the weight of scientific evidence means the new paradigm, if
evidence-based, is always superior to the previous theory.
What is now being argued about is exactly when the Anthropocene
Epoch began, so it can be ratified and added to the official geo-
logical timescale. Now, as I am sure you would expect, the argu-
ments about when humans became the dominant geological
power on Earth are robust, to say the least. The three main con-
tenders are the influence of early agriculturists from 5,000 years
onwards, the Colombian Exchange, with a date suggested by
Simon Lewis and myself of 1610, and the Great Acceleration,
190
when our environmental impacts grew exponentially in the 1950s

and 1960s. In many ways, these arguments detract from the pro-
found change created by recognition of the Anthropocene. This
is because it reverses 500 years of scientific discoveries, which
have continually moved humans to ever-increasing insignificance.
The Copernican revolution of the sixteenth century put the Sun
at the centre of the Solar System, downgrading the Earth. Modern
cosmology suggests that our Sun is one of 1024 stars in the
Universe, each one with the potential to have planets. Darwin’s
nineteenth-century discoveries, and the development of evolu-
tionary studies, established humans as a mere twig on the tree of
life, with no special place or origin. In the twenty-first century,
adopting the Anthropocene reverses this insignificance: humans
are not passive observers of Earth. The role of Homo sapiens is cen-
tral, because the future of the only place where life is known to
exist is being determined by our actions. In fact, I would argue
that not only has humanity become a geological superpower, it
has taken control of evolution itself.
The future of human evolution
Human evolution did not stop with the invention of agriculture;

instead, a different set of selection pressures arose. For example,
all humans are born with the ability to digest lactose in milk so
they can suckle from birth. However, in most humans the enzyme
lactase switches off in later life. In the early Holocene Epoch,
starting about 9,000 years ago, for small populations in Central
Europe, West Africa, and south-west India lactase persisted into
adulthood, allowing the digestion of non-human milk and dairy
products. Lactase persistence in adults seems to be due to the
191
presence of the key lactase gene. Today, this gene can be found in
80 per cent of Europeans and Americans of European ancestry,
while in sub-Saharan Africa and South East Asia its presence is
very low. It is absent in the Bantu of South Africa and most
Chinese populations. These geographical distributions strongly
correlate with the spread of domesticated cattle. Through the
early Holocene, dairy farming began and expanded from Central
Europe into Western and Northern Europe. Lactose tolerance
must have given a significant evolutionary advantage in early
agricultural communities. They were probably using milk prod-
ucts as a staple source of energy and as a fall-back food during
winter or drought periods. Non-human milk also provides an
additional source of Vitamin D, which is essential for normal
healthy bones, skin, and blood development. The global spread of
both dairy farming and European-derived populations has spread
lactase persistence around the globe—though it is interesting to
note that the majority of people around the world remain lactose
intolerant.
Lactase persistence is just one example of recent changes in
human evolution. Another is the appearance of blond(e), fair-
skinned peoples. However, the great Stephen Jay Gould declared
that he could see no biological change in modern humans over the
past 40,000 years. This view is echoed by the geneticist and col-
league of mine at University College London, Steve Jones. He
argues that modern medicine has halted natural selection, as
almost everyone can survive to the age of reproduction, and thus
pass on their genetic information. This is, however, a very Western
view of our world. First, at this time about 2 billion people do not
have access to a modern healthcare system; 1 billion people do not
have access to safe, clean drinking water; and 700 million people
192
do not have access to enough food, and go to bed hungry each

night. Second, even within a relatively rich developed country
there are clear differentiated costs involved in reproduction and
health, including access to education, health, contraception, junk
food, alcohol, and drugs. Moreover, religion can play a role in the
reproductive strategy of particular communities. Third, as Chris
Stringer from the Natural History Museum in London points out,
there are large numbers of mutations arising in our genome all the
time. One estimate suggests each one of us has fifty mutations
that we did not inherit from our parents. With the human popula-
tion at 7 billion, and set to rise to nearly 10 billion by the middle of
the century, that’s a lot of mutations—a lot of variation—for evo-
lution to play with. And moreover, increasing globalization means
that even previously isolated populations of humans are now
interbreeding, producing new genetic combinations.
Many geneticists and anthropologists now argue that culture, far
from holding evolution back, is accelerating it. Over the past 10,000
years we have gone through profound changes, from hunter-
gatherer to pastoralism to agriculture to urbanization, all of which
would have driven evolutionary processes. An example is the
human evolutionary reaction to disease. Unsanitary, densely packed
urban areas led to the rise of major diseases and parasites—for
example, smallpox, cholera, the plague, and yellow fever. Human
populations that co-evolved with these diseases built up some gen-
etic resistance. This is shown most dramatically by the devastation
of the native population of South America in the sixteenth century,
when these diseases were inadvertently introduced. The isolation
of this human population from Old World humans for over 10,000
years meant they hadn’t evolved immunity to these diseases, lead-
ing to over 50 million deaths. Meanwhile about 10 per cent of
193
Europeans had mutations, which provided resistance to smallpox

and may also confer resistance to HIV as well, as a fortuitous by-
product. In the tropics, the cutting down of forest and widespread
use of irrigation led to the spread of malaria across much of the
tropical and subtropical area, and the evolution of immunity or
resilience in some human populations.
It seems that not only our immune system is constantly evolv-
ing, but also our inner ear and eyesight. Recent mutations seem
to have changed the genes that produce proteins for the cilia and
their membranes in our inner ear, and the bones in our ears,
both of which help transmit sonic frequencies. Different muta-
tions with similar results are found in African, European, Chinese,
and Japanese populations, suggesting these changes have evolved
paralleling the development of different languages and their dis-
tinct sounds. Sight may also be undergoing evolutionary pres-
sure, with mutations found in key genes in East Asian populations,
changing the photoreceptors in the retina as well as the con-
struction of the inner ear. I also want to speculate as to whether
there is a genetic element to the clearly documented reduction in
human violence throughout history. Steven Pinker has shown
that reactive violence—i.e. individual on individual—can be
shown to have reduced in all cultures through time, but now
argues that this is due to the refinement and development of bet-
ter legal and cultural systems in some societies. But what if there
is a genetic component, and modern societies naturally select
less violent individuals? Chris Stringer speculates that perhaps
some of the one hundred or so recent mutations in brain neuro-
transmitters concerned with mood and demeanour may have
been selected to allow us to deal with large population sizes and
increased social tensions. However, others have argued that this
194
could be due to environmental causes. In the USA individual on

individual violence peaked in 1985, ten years after lead was
banned from petrol, and has steadily decreased every year since.
In the UK individual on individual violence peaked in 1990, ten
years after lead was banned from petrol, and has steadily
decreased every year since. Lead is a well-documented neuro-
toxin and it could be that it increased violence in the middle of
the twentieth century. Others have suggested the rise of violent
computer games distracts young males so they do not congre-
gate in large groups outside, and also provides an outlet for
young male aggression. Of course both these affects would also
lead to changes in brain pattern and structure, which would then
be passed on.
Humans are also taking charge of the evolutionary process,
both inadvertently and on purpose. Factors such as: the trans-
location of species, including human populations, to new habi-
tats; the selective removal of particular predators and large
mammals; environmental pressure from land-use changes,
deforestation, and climate change, are all altering evolutionary
outcomes. Add to this the pressures of new pollutants such as
antibiotics and other pharmaceuticals, pesticides, heavy metals,
microplastics, and novel genetically engineered organisms, and
the result is that humans are inadvertently influencing the evo-
lution of many organisms, including ourselves. We are also on
the cusp of a human genetic engineering revolution. Already we
can counsel parents about harmful DNA mutations that could
occur in their children, altering their decisions and changing the
future gene pool. Gene therapy will allow DNA to be altered in
the germline of an unborn child, changing their evolutionary
destiny forever.
195
If we stray into science fiction we can image a world where

humans are fully in control of their own evolution. The late Iain
M. Banks, the renowned and brilliant science fiction writer, imag-
ined a human society which was aptly named ‘The Culture’ that
had spread throughout the Universe. Once humans and their AI
partners, called ‘Minds’, had access to the stellar medium, then
resources were unlimited and humans controlled their own evo-
lution. Humans within ‘The Culture’ still have the basic form of
Homo sapiens, but they also have the ability to change gender
whenever they wish, can control their reproductive system, and
select particular gene combinations for their children. We are a
long way from this, but it does beautifully illustrate the point that
we really could be in charge of our evolutionary destiny, unlike
the situation today, in which our evolution is only partly inten-
tional and partly continues unplanned, driven by the blind force
of natural selection, with unpredictable effects and consequences
for the future.
Homo dominatus
Human evolution is the story of how we came to walk upright

and then became progressively smarter. All these major evolu-
tionary changes occurred in Africa. It seems that without the unique
impact of tectonics, celestial mechanics, and global climate vari-
ations on the landscape of East Africa, human evolution would
never have occurred. With the extinction of the Neanderthals
about 40,000 years ago, H. sapiens became the last surviving
hominin. With no natural weapons or defences, we dispersed out
of Africa as the apex predator and populated all the continents
except Antarctica. The complex flexible social human brain facili-
196
tated the development of cumulative culture. This led to the first

appearance of agriculture at the end of the last ice age. Throughout
the next 10,000 years, agriculture spread to almost every region
of the world, steadily replacing hunter-gathering as the major
economic mainstay, allowing urban centres and city-states to
develop. The intense competition between small nation states in
Europe, from the eleventh century onwards, created the perfect
centre for technological innovation, particularly of weapons. By
the fifteenth century, European powers had outstripped the
Chinese and Ottoman Empires and begun the great age of explor-
ation. Annexation and depopulation of the Americas in the six-
teenth century provided wealth and produce, allowing the
European population to grow beyond its own agricultural prod-
uctivity limits. Continued competition between the European
powers, and capital from their empires, led to the industrial revo-
lution of the eighteenth century. Post-World War II prosperity
created the technological revolution and the Great Acceleration
of environmental impacts. We have now entered what many see
as the fourth revolution—the information revolution. All these
revolutions have been driven by our cumulative culture and our
ultrasociality. It has also allowed the human population to expand
to nearly 7.5 billion people and is set to rise to nearly 10 billion
people by the middle of the century. We have influenced almost
every part of the Earth system. We are changing the global envir-
onmental and evolutionary trajectory of the Earth. We are also
both inadvertently and on purpose changing our own evolution.
Humanity has become in effect a geological superpower.
What I hope this book has demonstrated is that we were in many
ways created by rapid environmental changes that occurred in East
Africa, evolving a large social brain with intellectual flexibility and
197
group dynamics to deal with rapid and surprising changes. The

questions are: can we as a species cope with the global environ-
mental changes we ourselves have inadvertently caused? Can
humans, the only ultrasocial creature, knowingly self-organize
nearly 10 billion individuals in the future to mitigate and/or adapt
to the destruction of our environment? Or are we limited by our
evolutionary history? Because we are the only known species that
can predict the results of our own actions and truly understand
how we could positively manage the Earth’s environment for the
benefit of ourselves and all other species. We have named ourselves
Homo sapiens—‘wise person’. It is clear that as a species we have
taken control over the environmental and biological future of
Earth. Maybe Homo dominatus—which translates as ‘dominant per-
son’ or ‘tyrannical person’—would be more apt.
For me, the flexibility of the human brain and the fact that
many of its structures are determined not by genetics, but by the
environment and society into which we are born, means that we
can learn new ways of living very quickly. Each generation of
humanity can therefore adapt to the new environmental, social,
and political challenges facing us. Maybe in this century, if we
take our impacts on the Earth system seriously, we will earn our
species name ‘wise’. Whatever the future holds, we have come a
long way from our cradle in East Africa.
198
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22 4
Inde x
Afar 19, 20, 22–4, 67, 73, 75, 119, 131, 137 brain expansion 10, 14, 81, 130, 150
African lakes 112, 116, 138 Broecker (Wally) 98
Agriculture 11, 123, 169, 172, 183, 185,
191, 193 Cambrian Explosion 5, 7, 59
Aiello (Leslie) 27 carbon isotopes 24–5, 70, 71–3, 89, 93, 126
allopatric speciation 133, 145, 147–8 Chad 16, 22, 86
amplifier lakes 119, 138 Chicxulub 9, 57
anatomically modern humans 9, 38 chimpanzee 9, 22–3, 32, 35, 74, 76–7, 164,
Antarctica 41–4, 48, 82–5, 106, 186 171, 179, 181
Anthropocene 11, 185, 190, 191 birth 157–8
Anthropoids 9 brain size 16–7, 35, 164
Anton (Susan) 27–8, 138 common ancestor 10, 179
Arabian Plate 67 meat eating 25
Arctic Ocean 43, 80 penis 166
Ardi 19 quadrupedal 20, 74
Ardipithecus kadabba 19, 87 testes 165
Ardipithecus ramidus 19, 20, 71, 75, 127, tree-climbing 78
131, 137 war 149
Argon-Argon dating 119, 124 Chomsky (Noam) 163
aridity hypothesis 130, 132, 133, 136, Cieri (Robert) 169, 172
142, 144, 148 Clutton-Brock (Tim) 167
Atapuerca 13, 37 Collard (Mark) 26
Australopithecus afarensis 18, 21–5, 32, Colombian Exchange 187, 190
137, 157–8, 164 Congo 51, 67–8, 70, 77, 171–2
Australopithecus africanus 20, 24, 25 Court Jester hypothesis 136
Australopithecus anamensis 21, 73 Cradle of Humanity 12, 61, 66, 198
Australopithecus bahrelghazali 22 cumulative culture 9, 11, 38, 40, 169,
Australopithecus deyiremeda 23 173, 183, 197
Australopithecus garhi 25
Australopithecus platyops 22, 24, 73 Dart (Raymond) 20
Awash Valley 20, 24–5 Darwin (Charles) 129, 131, 191
Dawkins (Richard) 129
Barnosky (Anthony) 136 deep ocean circulation 44, 48, 88
Beauvilain (Alain) 16 Deino (Alan) 125
Berger (Lee) 29 Deccan Traps 9, 57
Big Bang 2 deMenocal (Peter) 132–3
bipedalism 10, 18–20, 22, 40, 73, 75–7, 130–2 Denisovans 175, 180
bonobo 9, 20, 171 diatom 116, 117, 139
225
inde x
diatomite 117, 125, 137, 139 hominin diversity 13, 31, 33
dinosaurs 7, 9, 41, 57–9, 82, 129 Homo antecessor 37
dispersal events 14, 39, 119, 124, 146 Homo Britannicus 37
Dixson (Alan) 166 Homo denisovan 38, 178, 180
Dmanisi 13, 31, 124 Homo dominatus 196, 198
DNA 179, 180, 195 Homo ergaster 31, 127
Drake Passage 48, 82, Homo erectus 15, 31, 32–40, 94, 127, 136, 138,
Dunbar (Robin) 150, 154, 156, 162 148, 162, 176
birth 158
Early to Middle Pleistocene transition brain size 94
(EMPT) 96, 97 food processing 34
Earth 1, 2, 4–7, 11–12, 41, 44, 51, 79, 104, group size 156
107, 123, 190 long-distance running 34
East African Rift Valley 11, 42, 51, 53, migration 37–8, 174
60–1, 63, 66, 68, 71, 73 naledi 29
Eccentricity 101–2, 104–5, 111–12, 119, sexual dimorphism 167
126, 137, 141 symbolic activity 163–4
Eldredge (Niles) 129 Homo floresiensis 30, 35, 36, 121, 124
El Niño-Southern Oscillation (ENSO) 93, Homo habilis 25, 26, 27–9, 32, 93, 136,
94, 99 138, 150
Epstein (Robert) 153 Homo heidelbergensis 15, 35, 36, 37–8, 97, 127,
equator–pole temperature gradient 43, 44 156, 162–3, 168, 175, 177
expensive brain framework 160, 164 Homo naledi 29, 121
Homo neanderthalensis 38, 150, 174,
Findlater Jet 68 175, 180
foramen magnum 16–17 Homo rhodesiensis 37
Homo rudolfensis 26, 27, 28, 136, 138
genetics 175–6, 179, 180–1, 198 Homo sapiens 1, 5, 9, 15, 36, 38, 40, 123,
gibbons 9, 22 163, 168, 176–8, 180, 183
Gomez-Robles (Aida) 181 birth 157, 158
Goodall (Jane) 149 brain size 26
Great Acceleration 190, 197 culture 11
Great Oxidation Event 6 dispersal 173, 174
Greenhouse World 58 drivers of evolution 191
Greenland 43, 85, 98, 108, 186, feminization 169, 170
Grey (Tom) 21 group size 156
glacial–interglacial cycles 96–7, 99, 100, interbreeding 150
105, 107, 109–10, 122, 141 self-domestication 170
glacial periods 72, 97, 122 ultrasocial 196
gorillas 9, 10, 31, 74, 76, 164–6, 171 Homo sapiens idaltu 38
Gould (Stephen J.) 2, 129, 192 Homo sapiens sapiens 38
graben 64, 65 hot spot 61, 63, 67
half-graben 64, 65 hunter-gathering 11, 197
Hadar 13, 21, 23, 137, Icehouse World 79, 82

Haile-Selassie (Yohannes) 19, 23 Indian Ocean 45, 47, 51, 54, 63, 68
Heinrich events 98 information revolution 1, 197
hominins 9, 10, 14, 16, 18, 20, 30–1, 34, 40, intensification of northern
73, 76, 94, 127, 130, 136, 138, 147, 149, hemisphere glaciation 90, 91,
150, 181 92, 93, 99, 110
group size 156 Intertropical Convergence Zone
mating system 167 (ITCZ) 111
226
inde x
Johanson (Donald) 21 North Atlantic Deep Water (NADW) 48, 49
Jones (Steve) 192 Nubian Plate 67
Joordens (Josephine) 163
Junginger (Annett) 141 obliquity 92, 101, 102, 104–6
Olduvai Gorge 25–6
Keller (Gerta) 57 Opie (Kit) 167
Kenyanthropus platyops 22, 24, 73 orangutans 9, 22, 75, 122, 166
Kingston (John) 125 orbital forcing 47, 92, 97, 100, 105, 107
Knauft (Bruce) 173 African climate 110, 113, 120
knuckle-walking 10, 18, 74, 75, 76, 131 African lakes 138, 141
Koobi Fora 126, 136 stratigraphy 122
Krakatoa 56 Oreopithecus bambolii 76
origin of life 6
Lake Turkana 21–3, 26–7, 34, 66, 127, 149 Orrorin tugenensis 18
Lamarck (Jean Baptiste) 131
language 7, 162–4, 181, 194 Palaeocene–Eocene Thermal Maximum
Leakey (Louis) 25 (PETM) 80–1
Leakey (Meave) 22 palaeomagnetism 121–2
Leakey (Richard) 34 Pan-African Orogeny 67
Lewis (Simon) 189 Panama Paradox 89
Lieberman (Dan) 34 Paranthropus aethiopicus 30, 73
Lucy 20, 21, 24, Paranthropus boisei 30, 31, 73, 136
Lucy in the Sky With Diamonds 21 Paranthropus robustus 30, 31, 73, 127
phylogenetic gradualism 129
magic number 150 154 Pickford (Martin) 18
Makapansgat 24, 126 Pinker (Steven) 163, 194
mass extinctions 7, 59, 129, 184 plate tectonics 40, 41, 55, 57–8
mating behaviour 164 Potts (Richard) 27, 35, 134–5, 138
Mediterranean Sea 47, 86–7, 94, 112, 126 precession 101, 102, 104–7, 120,
Megafauna Extinction 186–7 African lakes 112, 119–20, 125–6, 137–41
Messinian Salinity Crisis 99 Ice ages 106
Milankovitch (Milutin) 105 pulsed climate variability 147–8
Miller (Kenneth) 152 seasonality 110–14
modern humans 9, 10, 15, 38, 40, 168 primate 9, 58–9, 75–6, 78, 81, 131, 155–6
clothing 177 birth 159
dispersal 173, 178, 187 genetics 179
feet 29 sexuality 164, 166
group size 154 pulsed climate variability framework 137,
hip joint 18, 23 141–7
Neanderthal ancestry 178, 180 punctuated equilibrium 129–30, 133
running 77
sexual dimorphism 31, 167 quadrupedalism 74
social monogamy 167
testes 165 rain-shadow 52, 58, 69–70
monogamy 166–8, 170 Raymo (Maureen) 84
monsoon 47, 50, 51, 54–5, 59, 60, 67, 86, Red Queen hypothesis 136, 144
94, 110 Rising Star cave 13, 29
Moon 4, 104 Rose (Steven) 152
Ruddiman (Bill) 84
Neanderthals 35–7, 123, 175–8, 180, 196
neocortex 7, 155–6 Sahara desert 47, 112, 133, 192
North Africa 47, 51, 76, 86, 87, 186 Sahelanthropus tchadensis 16, 86
227
inde x
savannah hypothesis 75, 130–2 Tibetan Plateau 42, 51, 54, 60, 68, 85
Senut (Brigitte) 18 tilt see obliquity
sexual dimorphism 31, 164, 166–7, 171 Trauth (Martin) 114, 116–17, 137, 141
Seyoum (Chalachew) 25 Triassic extinction event 59
Shultz (Susanne) 169 Toumai 16
social brain hypothesis 150, 162 tree-climbing 10, 77–8, 131
Solar System 3–6, 11, 105, 191 turnover pulse hypothesis 130, 133–4,
Somali Plate 67 142–3, 148
South Africa 14, 20, 24, 29, 30–31, 37,
121, 126, 191 van Valen (Leigh) 136
South East Asian monsoonal system 51 variability selection hypothesis 130, 134–6,
Spoor (Fred) 26 143, 148
Sterkfontein 24 Vrba (Elisabeth) 133–4, 142
stone tools 24, 26, 28, 31, 34, 150, 164
Neanderthals 177 Walker (Alan) 34
oldest 10, 23, 93 Wells (Jonathan) 159–60
Stringer (Chris) 37, 176, 193–4 West Turkana 24
Suguta Valley 66–7 West Turkana Archaeological Project 23
Supercontinents 58–9 White (Tim) 19
supervolcano 56–7, 59 Wilson (Katie) 139, 146
Wrangham (Richard) 34, 171
Teilhardina belgica 81 Wood (Bernard) 26
Teilhardina brandti 81
Tethyan Sea 44 Zink (Katherine) 34
228
Moonstruck
How lunar cycles affect life
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ples the remarkable real effects that we are now
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shortly after the formation of the Moon, animals
evolved adaptations to the lunar cycle. He also
considers whether, if Moon-clock genes occur
in other animals, might they also exist in us?
Origins
The Scientific Story of Creation
-
Jim Baggott
“The collective mind of humanity has made

extraordinary progress in its quest to under-
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ation story. This book tells the version accord-
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Baggott helps us fulfil this desire, which is driven in part by simple curiosity
but also by a deeper emotional need to connect ourselves meaningfully with
the world which we call home.
The Improbable Primate
How Water Shaped Human Evolution
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Clive Finlayson
“Finlayson writes in a dry, clear, scholarly style

which somehow accentuates the sheer improb-
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Taking an ecological approach to our evolu-

tion, Clive Finlayson considers the origins of
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argues that environmental change, particularly
availability of water, played a critical role in
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shaping the direction of human evolution,
contributing to our spread and success.

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T he Cr adle of Hu m a ni t y

The Cradle of Humanity is an important contribution to the exten-

I have found that many of the concerned questions that have

Richard Leakey, FRS

I would like to thank Johanna, Alexandra, and Abbie Maslin for

References and Further Reading  199

1. Major biological events in the 4.6 billion years history of

2. Key morphological features that define hominids. 8

3. Map of the location of the most important hominin fossil

4. Location of the hole in the base of the skull through which

5. Evidence of bipedalism from the shape of the neck of

6. New divisions at the start of the Homo lineage suggested

7. Records of hominin brain capacity and East African

8. Summary of the hominin species covering the last

9. Illustration of the huge effect the latitudinal location of

10. Illustration of the significant alteration of ocean currents with

Based on figure 23 in Maslin, M. 2013. Climate: A Very Short Introduction,

11. The effect of changing ocean gateways on the location

12. Mountain rain-shadow. 52

13. The effects of mountains and plateaus on global rainfall. 53

14. The effects of plateaus and ice sheet on atmospheric

15. Long-term links between tectonics, sea level, climate,

16. Tectonic rifting and ocean formation. 62

17. Map of plate boundaries at the African Rift Valley triple

18. Explanation of horst-graben and half-graben geological

19. Tectonic development of the East African Rift Valley system. 66

20. Climatic effects of the development of the East African

21. Influence of Tibetan uplift and the African Rift Valley

22. Vegetation effects of the development of the East

23. Carbon isotope differences and relative abundance from

24. Global temperatures through the Cenozoic. 81

25. Ocean gateways both today and during the Cretaceous. 83

Based on figure 22 in Mark Maslin, M. 2013. Climate: A Very Short

26. Drying out, salt deposit formation, and reflooding of the

27. Oxygen isotopes showing the step-like changes in global ice

28. Pacific Ocean during El Niño and normal periods. 95

29. Orbital eccentricity, tilt, and precession. 102

30. Long-term changes in the orbital parameters. 103

31. Insolation at 65°N compared with global sea level or

33. East Africa north–south cross section with major

34. Occurrence of shallow and deep lakes over 5 million years

35. Age range of different dating methods using in the

36. Different modes of potential speciation with and without

37. Variability selection hypothesis of early human

Adapted from figure 2 in Potts, R. 1999. Variability selection in

38. Pulse climate variability framework. 139

39. Relationship between growth and shrinkage of large lakes in the

40. An interpretation of how the turnover pulse hypothesis could

41. An interpretation of how the aridity hypothesis could

42. An interpretation of how the allopatric speciation

43. Estimated community size of different hominin species

44. Robin Dunbar’s circles of human relationships. 156

45. A ‘Midwife’s’ view of the birth canal in a chimpanzee,

46. Expensive brain hypothesis. 161

370 <http://dx.doi.org/10.1098/rstb.2014.0064>, CC-BY version 4.0

47. Comparison of the great apes and humans relative

48. Comparison of Homo sapiens skull morphology

49. Dispersal of H. erectus, H. Neanderthalensis, and H. sapiens.174

50. Relationship between Homo sapiens and Homo erectus

References and Further Reading 199

2. Key morphological features that define hominids. 8

12. Mountain rain-shadow. 52

13. The effects of mountains and plateaus on global rainfall. 53

16. Tectonic rifting and ocean formation. 62

19. Tectonic development of the East African Rift Valley system. 66

24. Global temperatures through the Cenozoic. 81

25. Ocean gateways both today and during the Cretaceous. 83

28. Pacific Ocean during El Niño and normal periods. 95

29. Orbital eccentricity, tilt, and precession. 102

30. Long-term changes in the orbital parameters. 103

38. Pulse climate variability framework. 139

44. Robin Dunbar’s circles of human relationships. 156

46. Expensive brain hypothesis. 161

49. Dispersal of H. erectus, H. Neanderthalensis, and H. sapiens.174

But conflating or expanding the defined species seems at present

g radient of the northern hemisphere is somewhere between the

formed compared to today. So one of the fundamental elements

roduces more rainfall on land and a wetter continent overall. By