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Cambridge Studies in Biological and Evolutionary Anthropology 38

Neanderthals and Modern Humans

Neanderthals and Modern Humans develops the theme of the close
relationship between climate change, ecological change and biogeo-
graphical patterns in humans during the Pleistocene. In particular, it
challenges the view that Modern Human ˜superiority™ caused the ex-
tinction of the Neanderthals between 40 000 and 30 000 years ago.
Clive Finlayson shows that to understand human evolution, the spread
of humankind across the world and the extinction of archaic popula-
tions we must start off from a theoretical evolutionary ecology base
and incorporate the important wider biogeographic patterns, including
the role of tropical and temperate refugia. His proposal is that Nean-
derthals became extinct because their world changed faster than they
could cope with, and that their relationship with the arriving Modern
Humans, where they met, was subtle.

Clive Finlayson is Director, Museums and Heritage in the Govern-
ment of Gibraltar, based at the Gibraltar Museum. He is also Professor
in the Department of Anthropology at the University of Toronto. His
research interests include Quaternary human“environmental patterns,
the biogeography of hominids, and changing environments and faunal
patterns in the Quaternary of southern Europe.
Cambridge Studies in Biological and Evolutionary Anthropology

Series Editors
human ecology
C. G. Nicholas Mascie-Taylor, University of Cambridge
Michael A. Little, State University of New York, Binghamton
Kenneth M. Weiss, Pennsylvania State University
human evolution
Robert A. Foley, University of Cambridge
Nina G. Jablonski, California Academy of Science
Karen B. Strier, University of Wisconsin, Madison

Selected titles also in the series
21 Bioarchaeology Clark S. Larsen 0 521 49641 (hardback), 0 521 65834 9 (paperback)
22 Comparative Primate Socioecology P. C. Lee (ed.) 0 521 59336 0 (hardback)
0 521 00424 1 (paperback)
23 Patterns of Human Growth, second edition Barry Bogin 0 521 56438 7 (paperback)
24 Migration and Colonisation in Human Microevolution Alan Fix 0 521 59206 2
25 Human Growth in the Past Robert D. Hoppa & Charles M. FitzGerald (eds)
0 521 63153 X
26 Human Paleobiology Robert B. Eckhardt 0 521 45160 4
27 Mountain Gorillas Martha M. Robbins, Pascale Sicotte & Kelly J. Stewart (eds)
0 521 76004 7
28 Evolution and Genetics of Latin American Populations Francisco M. Salzano &
Maria C. Bortolini 0 521 65275 8
29 Primates Face to Face Agust´n Fuentes & Linda D. Wolfe (eds) 0 521 79109 X
30 Human Biology of Pastoral Populations William Leonard & Michael Crawford
(eds) 0 521 78016 0
31 Paleodemography Robert D. Hoppa & James W. Vanpel (eds) 0 521 80063 31
32 Primate Dentition Davis Swindler 0 521 65289 8
33 The Primate Fossil Record Walter C. Hartwig (ed.) 0 521 66315 6
34 Gorilla Biology Andrea B. Taylor & Michele L. Goldsmith (eds) 0 521 79281 9
35 Human Biologists in the Archives D. Ann Hening & Alan C. Swedlund (eds)
0 521 80104 4
36 Human Senescence Douglas Crews 0 521 57173 1
37 Patterns of Growth and Development in the Genus Homo Jennifer L. Thompson,
Gail E. Krovitz & Andrew J. Nelson (eds) 0 521 57173 1
Neanderthals and
Modern Humans
An Ecological and Evolutionary Perspective

The Gibraltar Museum
The University of Toronto
cambridge university press
Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, São Paulo

Cambridge University Press
The Edinburgh Building, Cambridge cb2 2ru, UK
Published in the United States of America by Cambridge University Press, New York
Information on this title: www.cambridge.org/9780521820875

© Clive Finlayson 2004

This publication is in copyright. Subject to statutory exception and to the provision of
relevant collective licensing agreements, no reproduction of any part may take place
without the written permission of Cambridge University Press.
First published in print format 2004

isbn-13 978-0-511-18634-9 eBook (EBL)
isbn-10 0-511-18634-7 eBook (EBL)

isbn-13 978-0-521-82087-5 hardback
isbn-10 0-521-82087-1 hardback

Cambridge University Press has no responsibility for the persistence or accuracy of urls
for external or third-party internet websites referred to in this publication, and does not
guarantee that any content on such websites is, or will remain, accurate or appropriate.
To Geraldine and Stewart

page ix
Acknowledgements x

1 Human evolution in the Pleistocene 1
2 Biogeographical patterns 9
3 Human range expansions, contractions and extinctions 39
4 The Modern Human“Neanderthal problem 71
5 Comparative behaviour and ecology of Neanderthals and
Modern Humans 94
6 The conditions in Africa and Eurasia during the last glacial cycle 135
7 The Modern Human colonisation and the Neanderthal extinction 148
8 The survival of the weakest 195

References 209
Index 249


In 1848 a strange skull was discovered in Forbes™ Quarry, Gibraltar, close to
where I live. A second skull found eight years later in the Neander Valley, near
Dusseldorf in Germany, gave a new hominid its name “ the Neanderthal. This
name, and its relation to an individual that lived close to the edge of its range,
led to over a century of perception of the Neanderthals as a brutish people
of northern Europe who survived, through thick and thin, the cold of the ˜ice
ages™ until they were supplanted by the newly arrived and intelligent Modern
The image is still one that many regard as close to reality. Yet, paradoxically,
the Neanderthals were intelligent people of mild climates. They evolved across
the northern shores of the Mediterranean Sea and eastwards towards the Black
and Caspian Seas. They ventured north only during mild climatic episodes and
the unstable, cold and arid climate of late Pleistocene Europe eventually gave
them the blow that sent them on the road to extinction. The Modern Humans
hovered in the periphery and took advantage of the situations left vacant by the
Neanderthals. This book is an attempt to redress the balance of over a century
of misunderstanding.


I am grateful to the publishers, and in particular Tracey Sanderson, for the
opportunity to publish this book and for their support throughout. The ideas
put forward in this book were conceived after many discussions with friends
and colleagues over a number of years. I am particularly indebted to my wife,
Geraldine Finlayson, for her insightful discussions, ideas and support through-
out. The ecological approach followed in this book stems from many years
working in bird ecology. The ecological discussions have been particularly
intense and fruitful with my friend and colleague Darren Fa.
I ¬rst ventured into the ¬eld of human evolution in 1990 when I became
involved in the Gibraltar Caves Project. Two of its co-directors, Chris Stringer
and Andy Currant of the Natural History Museum in London, have had a lot
to do with my involvement and participation in this exciting ¬eld. I have been
especially welcomed into the archaeological side of this subject, and have learnt
vast amounts in the ¬eld, from the friendship and knowledge of Paco Giles of
the Museo de El Puerto Santa Mar´a. I have spent many good times discussing
and learning about the Palaeolithic from him and his team, especially Antonio
Santiago P©rez, Jos© Mar´a Gutierrez L´ pez and Esperanza Mata Almonte. I am
± o
also deeply indebted to my good friend and colleague Joaquin Rodriguez-Vidal
for the brilliant way in which he has made me understand the geomorphology
of the karstic landscapes that the Neanderthals lived in.
During the last ¬ve years in particular I have bene¬ted from discussions
with many colleagues, particularly during the two Calpe conferences organ-
ised in Gibraltar in 1998 and 2001: Emiliano Aguirre, Juan Luis Arsuaga,
Javier Baena Preysler, Nick Barton, Ofer Bar-Yosef, Jacques Blondel, Eudald
Carbonell, Miguel Cort©s, Francesco d™Errico, Yolanda Fern´ ndez Jalvo, Rob
Foley, Clive Gamble, Paul Goldberg, Marta Lahr, Richard MacPhail, Paul Mel-
lars, Marina Mosquera, Paul Pettitt, Marcia Ponce de Le´ n, Robert Sala, Larry
Sawchuk, Olga Soffer, Gerardo Vega Toscano, Erik Trinkaus, Manuel Vaquero,
Joao Zilhao, Christoph Zollikofer.

1 Human evolution in the Pleistocene

The origins of humanity may be traced to the tropical African Pliocene, around 6
million years ago (Myr). Genetic evidence has for some time predicted the ex-
istence of a common ancestor to chimpanzees and humans around 5“6 Myr
(Takahata & Satta, 1997; Gagneux & Varki, 2001). Recent discoveries of
African fossils that are claimed to be close to this common ancestor have been
dated to between 6 and 7 Myr (Brunet et al., 2002).
From this point until the emergence of Homo erectus 1.9 Myr ago and its
rapid subsequent range expansion (Aguirre & Carbonell, 2001), hominids were
con¬ned to sub-Saharan Africa. The estimated number of species that lived
during this long period in the Pliocene varies among authors. If we follow a
conservative approach (Klein, 1999) we observe a pattern of increasing hominid
species richness from about 4.6 Myr with a peak between 1.9 and 1.6 Myr and
a sharp decline thereafter (Fig. 1.1). The decline after 2 Myr ago is correlated
with increasing climate instability.
The peak in diversity coincides with the ¬rst appearance in the fossil record
of H. erectus. Recently this early African member of the genus Homo has been
separated from contemporary Asian forms. The name H. erectus has been re-
tained for the Asian forms and the name H. ergaster for the African (Klein,
1999). Recent evidence suggests, however, that the two signi¬cantly overlap in
morphology and that they should form part of a geographically diverse species
H. erectus (Asfaw et al., 2002). I follow this latter classi¬cation here. Subse-
quent forms have been given speci¬c status by different authorities although
there is considerable uncertainty regarding the precise boundaries of each. The
classi¬cation of fossils is fraught with dif¬culties as we shall see in Chapter 4.
In this book I consider H. erectus“H. sapiens to be a single chronospecies
(Cain, 1971) that has repeatedly produced divergent lineages through geo-
graphical isolation during the last 1.9 Myr. Some of the described forms are
clearly temporal entities within the H. erectus“H. sapiens continuum. I include
H. heidelbergensis and H. helmei in this category. Others are divergent lin-
eages that have subsequently become extinct. The Neanderthals are the clearest
example of such a divergent lineage and their relationship with mainstream
H. sapiens will occupy much of this book. Until equivalent fossils are found
in Africa it is probably best to regard the form H. antecessor from the Spanish
site of Atapuerca (Carbonell et al., 1995), and possibly also those of Ceprano

2 Neanderthals and Modern Humans

in Italy (Manzi et al., 2001) in this latter category, i.e. a divergent lineage that
became extinct.
The question of interbreeding between mainstream H. sapiens and diver-
gent lineages when geographical or ecological barriers broke down will be ad-
dressed, with speci¬c reference to Neanderthals and contemporary mainstream
H. sapiens, in Chapter 7. The degree of genetic isolation of the constituent pop-
ulations would be dependent on a range of factors at any point. These would
include distance effects and physical, climatic and ecological barriers. Popu-
lations would become isolated at some points and a process of genetic diver-
gence would ensue. Most often such a process would end with renewed contact
among populations. At other scales, metapopulations in different regions would
become isolated from each other. Gene ¬‚ow would continue within but not be-
tween regions. At even larger spatial scales entire regions would occasionally



Number of Species


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