9 The Earliest Putative Homo Fossils

Handbook of Paleoanthropology

Prof. Dr. Dr. h. c. Winfried Henke

Institut fur Anthropologie (1050)

Fachbereich 10 ‐ BiologieJohannes Gutenberg‐Universitat Mainz

D‐55099 Mainz

Germany

email: [email protected]

Prof. Dr. Ian Tattersall

Division of Anthropology

American Museum of Natural History

New York, NY 10024‐5192USA


Dipl. Biol. Thorolf Hardt


Fachbereich 10 ‐ BiologieJohannes Gutenberg‐Universitat Mainz

D‐55099 Mainz

Germany


ISBN‐13: 978‐3‐540‐32474‐4

This publication is available also as:Electronic publication under ISBN 978‐3‐540‐33761‐4 andPrint and electronic bundle under ISBN 978‐3‐540‐33858‐1

Library of Congress Control Number: 2006936414

This work is subject to copyright. All rights are reserved, whether the whole or part of thematerial is concerned, specifically the rights of translation, reprinting, reuse of illustrations,recitation, broadcasting, reproduction on microfilms or in other ways, and storage in databanks. Duplication of this publication or parts thereof is only permitted under the provisions ofthe German Copyright Law of September 9, 1965, in its current version, and permission for usemust always be obtained from Springer‐Verlag. Violations are liable for prosecution under theGerman Copyright Law.

Springer is part of Springer Science+Business Media

springer.com

# Springer‐Verlag Berlin Heidelberg New York 2007

The use of registered names, trademarks, etc. in this publication does not imply, even in theabsence of a specific statement, that such names are exempt from the relevant protective lawsand regulations and therefore free for general use.

Product liability: The publishers cannot guarantee the accuracy of any information about theapplication of operative techniques and medications contained in this book. In every individualcase the user must check such information by consulting the relevant literature.

Editor: Dieter Czeschlik, Heidelberg/Sandra Fabiani, HeidelbergDevelopment Editor: Susanne Friedrichsen, HeidelbergProduction Editor: Frank Krabbes, HeidelbergCover Design: Frido Steinen‐Broo, Spain

Printed on acid‐free paper SPIN: 2109 ‐ 5 4 3 2 1 0

We dedicate these volumes to our long-time colleagues

Hartmut Rothe and Theodoros Pitsios

in appreciation of their friendship and unique contributions to

primatology and paleoanthropology.

Preface to the Series

Palaeoanthropology is perhaps the most multidisciplinary of all the sciences. Any

complete account of the evolution and of the cultural and biological contexts

of Homo sapiens must combine information from geology, paleoecology, prima-

tology, evolutionary biology and a host of other fields. Above all, historical

information garnered from the fossil record needs to be combined with, and

interpreted in the light of, what we know of the living world. In these volumes we

have brought together contributions by a variety of leading specialists that reflect

the broad spectrum of modern paleoanthropology, in an attempt to provide a

resource that we hope will be useful to professionals and students alike.

Volume I of this three-volume Handbook deals with principles, methods,

and approaches. In recent years enormous advances have been made in such areas

as phylogenetic analysis, evolutionary theory and philosophy, paleoecology, and

dating methods. The contributions aim to present the state of the art in these

and other relevant fields, as well as to furnish succinct introductions to them and

to reflect the many ways in which they interact. Human beings are primates,

and Volume II is devoted to primate origins, evolution, behavior, and adaptive

variety. In this compilation the emphasis is on the integration of fossil data with

the vast amount that is now known of the behavior and ecology of living primates

in natural environments. The third and final volume deals directly with the fossil

and molecular evidence for the evolution of Homo sapiens and its fossil relatives

(the family Hominidae or subfamily Homininae, according to taste, a matter that

we have left to each individual contributor). Paleoanthropology is a pluralistic

and actively developing field in which much remains to be settled, and we have

not tried to impose any uniformity of viewpoint on our authors. Instead, while

maintaining an emphasis on the data, we have encouraged them to express their

individual interpretations rather than to cover all possible points of view. This has

inevitably led to a certain degree of heterogeneity of opinion between the covers

of this Handbook; but we believe that this is the best way of reflecting the

excitement and momentum of the field and that it is best for the reader to be

left to reach his or her own conclusions. Science is, after all, a process rather than

a static product, and one of our primary aims here is to reflect the ongoing

dynamism of that process in paleoanthropology.

We thank all of the contributors to these volumes for their participation.

Some initially responded enthusiastically while others needed convincing about

the basic strategy of the Handbook, but all responded marvellously to the

viii Preface to the Series

particular needs of a corporate effort such as this one. We are particularly grateful

to those authors who responded at short notice to needs that became apparent

only as the project progressed. This series was conceived in collaboration with

Prof. Hartmut Rothe of the University of Gottingen, who was later forced to

withdraw for reasons beyond his control. We thank him most warmly for his

creativity in the conceptual stages and for his subsequent moral support. The

laborious process of putting together the volumes could not have been accom-

plished without the cheerful help of Thorolf Hardt, whose active involvement was

indispensable throughout.

This project could never have come to fruition without the enthusiastic

support of Dr. Dieter Czeschlik, editor life sciences at Springer Publishing, and

the efficient assistance of Mrs. Ursula Gramm. We express our deep gratitude to

Mrs. Susanne Friedrichsen and Mrs. Caroline Simpson, who showed both care

and commitment during the phases of copyediting and product development.

The continuous cooperation and dialogue with them and their professionalism

gave us the courage to see the project through. Further thanks go to Mrs. Sandra

Fabiani and her colleagues at Springer Publishing, who prepared the eReference.

Ms. Nitya Swaruba, compositor at SPi Technologies, deserves warm thanks for

her efficient help, and finally our gratitude goes in addition to Britta Hardt, Peter

Menke and Monika Sandfuhr, who also rendered much valuable assistance.

Winfried Henke and Ian Tattersall

Mainz and New York City

November, 2006

Table of Contents

Volume 1

1 Historical Overview of Paleoanthropological Research . . . . . . . . . . . . . . . 1Winfried Henke

2 Evolutionary Theory in Philosophical Focus . . . . . . . . . . . . . . . . . . . . . . . 57Philippe Huneman

3 The Ontogeny–Phylogeny Nexus in a Nutshell: Implications for

Primatology and Paleoanthropology . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103Peter R. Menke

4 Principles of Taxonomy and Classification: Current Procedures

for Naming and Classifying Organisms . . . . . . . . . . . . . . . . . . . . . . . . . . 141Michael Ohl

5 Quantitative Approaches to Phylogenetics . . . . . . . . . . . . . . . . . . . . . . . 167Kaila E. Folinsbee . David C. Evans . Jorg Frobisch . Linda A. Tsuji .

Daniel R. Brooks

6 Homology: A Philosophical and Biological Perspective . . . . . . . . . . . . . 217Olivier Rieppel

7 Taphonomic and Diagenetic Processes . . . . . . . . . . . . . . . . . . . . . . . . . . 241Gisela Grupe

8 Archeology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 261Miriam N. Haidle

9 Contribution of Stable Light Isotopes to Paleoenvironmental

Reconstruction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 289
Julia Lee‐Thorp . Matt Sponheimer
10 Chronometric Methods in Paleoanthropology . . . . . . . . . . . . . . . . . . . . 311Gunther A. Wagner

xxvi

11 Geological Background of Early Hominid Sites in Africa . . . . . . . . . . . . 339Ottmar Kullmer

12 Paleoclimate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 357Keith Alverson

13 Paleosols . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383Gregory Retallack

14 Quaternary Deposits and Paleosites . . . . . . . . . . . . . . . . . . . . . . . . . . . . 409Klaus‐Dieter Jager

15 Zoogeography: Primate and Early Hominin Distribution and

Migration Patterns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 421Alan Turner . Hannah O’Regan

16 Patterns of Diversification and Extinction . . . . . . . . . . . . . . . . . . . . . . . . 441Walter Etter

17 Paleoecology: An Adequate Window on the Past? . . . . . . . . . . . . . . . . . 503Thorolf Hardt . Britta Hardt . Peter R. Menke

18 Hominin Paleodiets: The Contribution of Stable Isotopes . . . . . . . . . . . 555Matt Sponheimer . Julia Lee ‐Thorp

19 Estimation of Basic Life History Data of Fossil Hominoids . . . . . . . . . . 587Helmut Hemmer

20 Population Genetics and Paleoanthropology . . . . . . . . . . . . . . . . . . . . . 621John H. Relethford

21 Ancient DNA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 643Susanne Hummel

22 Paleodemography of Extinct Hominin Populations . . . . . . . . . . . . . . . . 673Janet Monge . Alan Mann

23 Modeling the Past: The Primatological Approach . . . . . . . . . . . . . . . . . 701R. W. Sussman . Donna Hart

Table of Contents

24 Modeling the Past: The Paleoethnological Evidence . . . . . . . . . . . . . . . 723Paolo Biagi

xxvii

25 Modeling the Past: The Linguistic Approach . . . . . . . . . . . . . . . . . . . . . 747Bernard Comrie

26 General Principles of Evolutionary Morphology . . . . . . . . . . . . . . . . . . . 769Gabriele A. Macho

27 Computer‐Based Reconstruction: Technical Aspects

and Applications . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 787Lilian Ulhaas

Table of Contents

28 Prospects and Pitfalls . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 815Jean-Jacques Hublin

Volume 2

1 Primate Origins and Supraordinal Relationships: Morphological

Evidence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 831Mary T. Silcox . Eric J. Sargis . Jonathan I. Bloch . Doug M. Boyer

2 Molecular Evidence on Primate Origins and Evolution . . . . . . . . . . . . . 861Hans Zischler

3 Fossil Record of the Primates from the Paleocene to the Oligocene . . . . . 889D. Tab Rasmussen

4 Fossil Record of Miocene Hominoids . . . . . . . . . . . . . . . . . . . . . . . . . . . . 921David R. Begun

5 The Biotic Environments of the Late Miocene Hominids . . . . . . . . . . . . 979Jordi Agustı

6 Postcranial and Locomotor Adaptations of Hominoids . . . . . . . . . . . . 1011Carol V. Ward

7 Hominoid Cranial Diversity and Adaptation . . . . . . . . . . . . . . . . . . . . . 1031Alan Bilsborough . Todd C. Rae

8 Dental Adaptations of African Apes . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1107.
Mark F. Teaford Peter S. Ungar
9 Evolution of the Primate Brain . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1133Dean Falk

xxviii

10 Primate Life Histories . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1163Elke Zimmermann . Ute Radespiel

11 The Biology and Evolution of Ape and Monkey Feeding . . . . . . . . . . . 1207Joanna E. Lambert

12 Great Ape Social Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1235Angela Meder

13 Primate Intelligence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1273Richard W. Byrne

14 Chimpanzee Hunting Behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1295Nicholas E. Newton‐Fisher

15 Cooperation, Coalition, and Alliances . . . . . . . . . . . . . . . . . . . . . . . . . . 1321Charlotte K. Hemelrijk . Jutta Steinhauser

Volume 3

1 Potential Hominoid Ancestors for Hominidae . . . . . . . . . . . . . . . . . . . . 1347George D. Koufos

2 Defining Hominidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1379Jeffrey H. Schwartz

3 Origins of Homininae and Putative Selection Pressures

Acting on the Early Hominins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1409Bogusław Pawłowski

4 Role of Environmental Stimuli in Hominid Origins . . . . . . . . . . . . . . . . 1441Elisabeth S. Vrba

5 The Origins of Bipedal Locomotion . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1483William E. H. Harcourt‐Smith

6 The Earliest Putative Hominids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1519Brigitte Senut

Table of Contents

7 The Species and Diversity of Australopiths . . . . . . . . . . . . . . . . . . . . . . 1539William H. Kimbel

xxix

8 Defining the Genus Homo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1575Mark Collard . Bernard Wood

9 The Earliest Putative Homo Fossils . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1611Friedemann Schrenk . Ottmar Kullmer . Timothy Bromage

10 Homo ergaster and Its Contemporaries . . . . . . . . . . . . . . . . . . . . . . . . . 1633Ian Tattersall

11 Defining Homo erectus: Size Considered . . . . . . . . . . . . . . . . . . . . . . . . 1655Susan C. Anton . Fred Spoor . Connie D. Fellmann . Carl C. Swisher III

12 Later Middle Pleistocene Homo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1695G. Philip Rightmire

13 Neanderthals and Their Contemporaries . . . . . . . . . . . . . . . . . . . . . . . 1717Katerina Harvati

14 Origin of Modern Humans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1749Gunter Brauer

15 Analyzing Hominid Phylogeny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1781David Strait . Frederick E. Grine . John G. Fleagle

16 Phylogenetic Relationships (Biomolecules) . . . . . . . . . . . . . . . . . . . . . . 1807Todd R. Disotell

17 Population Biology and Population Genetics of

Pleistocene Hominins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1825Alan R. Templeton

18 Species Concepts and Speciation: Facts and Fantasies . . . . . . . . . . . . 1861Colin Groves

19 Human Environmental Impact in the Paleolithic and Neolithic . . . . . 1881Wolfgang Nentwig

20 The Dentition of American Indians: Evolutionary Results

Table of Contents

and Demographic Implications Following Colonization

from Siberia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1901Christy G. Turner II . G. Richard Scott

xxx

21 Overview of Paleolithic Archeology . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1943Nicholas Toth . Kathy Schick

22 The Network of Brain, Body, Language, and Culture . . . . . . . . . . . . . . 1965Steven Mithen

23 Cultural Evolution in Africa and Eurasia During the Middle

and Late Pleistocene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2001Nicholas Conard

24 Paleoanthropology and the Foundation of Ethics: Methodological

Table of Contents

Remarks on the Problem of Criteriology . . . . . . . . . . . . . . . . . . . . . . . . 2039Mathias Gutmann . Michael Weingarten

About the Editors

Editors in Chief

Prof. Dr. Dr. h. c. Winfried Henke


Fachbereich 10 - Biologie

Johannes Gutenberg-Universitat Mainz

D‐55099 Mainz

Germany


Prof. Dr. Ian Tattersall

Division of Anthropology

American Museum of Natural History

New York, NY 10024–5192

USA


Editorial Assistant

Dipl. Biol. Thorolf Hardt


Fachbereich 10 - Biologie

Johannes Gutenberg-Universitat Mainz

D‐55099 Mainz

Germany


Winfried Henke is currently Professor of Anthropology at the Johannes

Gutenberg University of Mainz. He was born in 1944 in Ludwigshorst/

Pomerania, Germany, and studied biology, anthropology and geosciences in

Kiel and Braunschweig. He received his Ph.D. from the University of Kiel in

1971, his thesis focusing on a prehistoric anthropological topic. In 1990 he

habilitated with a monograph on the ‘‘Anthropology of Early Paleolithics and

Mesolithics’’ at the University of Mainz.

Research activities in various countries (Iceland, Israel, Jordan, US, Greece)

and extensive teaching assignments in the Erasmus exchange program at

numerous European Universities followed. From 1996 to 2004, he acted as

anthropology referee for the German Research Foundation (DFG). He served at

the advisory boards of many scientific journals, and was advisory consultant to

museums, e.g. the Neanderthal Museum (Mettmann, Germany). In 2006 he was

awarded with the honorary doctorate of the National and Kapodistrian

University of Athens and is an elected member of the German Academy of

2072 About the Editors

Sciences Leopoldina. Areas of research and teaching: paleoanthropology,

primatology, prehistoric anthropology, comparative morphology, systematics,

demography and sociobiology.

He has published approximately 180 original papers in scientific journals and

anthologies, and over 600 book reviews. He is author, co-author (together with

H. Rothe) and editor of several books, including such standard works as

‘‘Palaoanthropologie’’ and ‘‘Stammesgeschichte des Menschen’’ (published at

Springer-Verlag).

Ian Tattersall is currently Curator in the Division of Anthropology of the

American Museum of Natural History in New York City. Born in England and

raised in East Africa, he has carried out both primatological and paleontological

fieldwork in countries as diverse as Madagascar, Vietnam, Surinam, Yemen and

Mauritius. Trained in archeology and anthropology at Cambridge, and in

geology and vertebrate paleontology at Yale, Tattersall has concentrated his

research since the 1960s in two main areas: the analysis of the human fossil

record and its integration with evolutionary theory, and the study of the

ecology and systematics of the lemurs of Madagascar. Tattersall is also a

prominent interpreter of human paleontology to the public, with several trade

books to his credit, among them The Monkey in the Mirror (2002), Extinct

Humans (with Jeffrey Schwartz, 2000), Becoming Human: Evolution and Human

Uniqueness (1998) and The Last Neanderthal: The Rise, Success and Mysterious

Extinction of Our Closest Human Relatives (1995; rev. 1999) as well as several

articles in Scientific American and the co-editorship of the definitive Encyclopedia

of Human Evolution and Prehistory. He lectures widely, and, as curator, has also

been responsible for several major exhibits at the American Museum of Natural

History, including Ancestors: Four Million Years of Humanity (1984); Dark Caves,

Bright Visions: Life In Ice Age Europe (1986); Madagascar: Island of the Ancestors

(1989); The First Europeans: Treasures from the Hills of Atapuerca (2003); and the

highly acclaimed Hall of Human Biology and Evolution (1993).

Thorolf Hardt graduated in biology and is currently a PhD student at the

Institute of Anthropology, University of Mainz. He was born in 1973 in

Neustadt (RhP/Germany) and studied anthropology, paleontology and zoology

in Kiel and Mainz. At present his research activities are focused on Geometric

Morphometrics, functional morphology and evolutionary biology in Primates.

Contributors

J. Agustı Ballester

ICREA-Institut of Human

Paleoecology,

Universitat Rovira i Virgili,

Pl. Imperial Tarraco, 1,

43005-Tarragona,

Spain

K. Alverson

Ocean Observations and Services,

IOC/UNESCO,

Global Ocean Observing System,

1 rue Miollis,

75732 Paris Cedex 15,

France

S. C. Anton

Center for the Study of Human

Origins,

Department of Anthropology NYU,

25 Waverly Place,

New York, NY 10003,

USA

D. R. Begun

Department of Anthropology,

100 St George Street, Rm 1037,

University of Toronto,

Toronto, ONT M5S 3G3,

Canada

P. Biagi

Department of Science of Antiquities

and the Near East,

Ca’ Foscari University, Venice,

Palazzo Bernardo, S. Polo 1977,

30125 Venezia,

Italy

A. Bilsborough


University of Durham,

43 Old Elvet, Durham,

DH 1 3HN,

UK

J. I. Bloch

Vertebrate Paleontology,

Florida Museum of Natural History,

Dickenson Hall, University

of Florida,

Gainesville, Florida 32611‐7800,USA

D. M. Boyer

Department of Anatomical

Sciences,

Stony Brook University,

Stony Brook, NY 11794‐8081,USA

G. Brauer

Institut fur Humanbiologie,

Universitat Hamburg,

Allende-Platz 2,

D‐20146 Hamburg,

Germany

2074 Contributors

T. Bromage

Department of Biomaterials and

Biomimetics,

New York University College of

Dentistry,

345 East 24th Street, Room 804-S,

New York, NY 10010,

USA

D. R. Brooks

Department of Ecology &

Evolutionary Biology,

University of Toronto,

25 Harbord Street,

Toronto, Ontario M5S 3G5,

Canada

R. W. Byrne

School of Psychology,

University of St. Andrews,

St. Andrews, Fife KY16 9JU,

Scotland

M. Collard

Laboratory of Biological

Anthropology,


University of British Columbia,

6303 NW Marine Drive,

Vancouver, British Columbia V6T 1Z1,

Canada

B. Comrie

Department of Linguistics,

Max Planck Institute of Evolutionary

Anthropology,

Deutscher Platz 6,

D‐04103 Leipzig,

Germany

N. J. Conard

Institut fur Ur- und Fruhgeschichte,

Abteilung Altere Urgeschichte und

Quartarokologie,

Eberhard-Karls-Universitat Tubingen,

Schloss Hohentubingen,

D-72070 Tubingen,

Germany

T. R. Disotell


Origins,


New York University,

25 Waverly Place,

New York, NY 10003,

USA

W. Etter

Naturhistorisches Museum Basel,

Abteilung Geowissenschaften,

Augustinergasse 2,

CH-4001 Basel,

Switzerland

D. Evans

Department of Biology,

University of Toronto at Mississauga,

3359 Mississauga Road,

Mississauga, ON L5L 1C6,

Canada

D. Falk


Florida State University,

Tallahassee,

FL 32306‐7772, 1847 West,

Tennessee St.,

USA

Contributors 2075

C. D. Fellmann


Origins,


New York University,

25 Waverly Place,

New York City, NY 10003,

USA

J. G. Fleagle

Department of Anatomical Sciences,

Health Sciences Centre, Stony

Brook University,

Stony Brook, New York 11794‐8081,USA

K. E. Folinsbee



3359 Mississauga Road,


Canada

J. Frobisch



3359 Mississauga Road North,


Canada

F. E. Grine


State University of New York,

Stony Brook, NY 11794‐4364,USA

C. P. Groves

School of Archaeology and

Anthropology,

Australian National University

Canberra, ACT 0200,

Australia

G. Grupe

Dept. I fur Biologie,

Bereich Biodiversitatsforschung/

Anthropologie,

Universitat Munchen,

Grosshaderner Straße 2,

D-82152 Planegg-Martinsried,

Germany

M. Gutmann

Institut fur Philosophie,

Philipps-Universitat Marburg,

Wilhelm-Ropke-Strasse 6, Block B

D‐35032 Marburg,

Germany

M. N. Haidle

Institut fur Ur- und Fruhgeschichte

und Archaologie des Mittelalters,

Abt. Altere Urgeschichte und

Quartarokologie,

Universitat Tubingen,

Burgsteige 11,

D-72070 Tubingen,

Germany

W. E. H. Harcourt-Smith

Division of Paleontology,

American Museum of Natural History,

Central Park West and 79th Street,

New York, NY 10024,

USA

B. Hardt

Institut fur Anthropologie (1050),

Fachbereich 10 – Biologie,

Johannes Gutenberg-Universitat

2076 Contributors

Mainz,

D‐55099 Mainz,

Germany

T. Hardt




Mainz,

D‐55099 Mainz,

Germany

D. L. Hart


University of Missouri at St. Louis,

St. Louis, MO 63130, USA

K. Harvati

Department of Human Evolution,

Max Planck Institute for

Evolutionary Anthropology,

Deutscher Platz 6,

D‐04103 Leipzig,

Germany

C. K. Hemelrijk

Theoretical Biology,

Centre for Ecological and

Evolutionary Studies,

University of Groningen,

Biological Centre,

Kerklaan 30,

9751 NN Haren,

The Netherlands

H. Hemmer

Anemonenweg 18,

D‐55129 Mainz,

Germany

or

Institut fur Zoologie,



Mainz,

Johannes v. Muller-Weg 6,

D‐55128 Mainz,

Germany

W. Henke




Mainz,

D‐55099 Mainz,

Germany

J.-J. Hublin

Department of Human Evolution,

Max Planck Institute for Evolutionary

Anthropology,

Deutscher Platz 6,

D‐4103 Leipzig,

Germany

S. Hummel

Johann Friedrich Blumenbach-

Institut fur Zoologie und

Anthropologie, Historische

Anthropologie und Humanokologie,

Georg August‐Universitat Gottingen,Burgerstrasse 50,

D-37037 Gottingen,

Germany

P. Huneman

Institut d’Histoire et de Philosophie

des Sciences et des Techniques

CNRS/Universite Paris I

Sorbonne,

Contributors 2077

13 rue du Four,

75006 Paris,

France

K.-D. Jager

Institut fur Prahistorische

Archaologie,

Martin Luther-Universitat Halle,

Brandbergweg 23c,

D‐06099 Halle,

Germany

W. H. Kimbel

Institute of Human Origins,

Arizona State University,

P.O. Box 874101,

Tempe, AZ 85287‐4101,USA

G. D. Koufos

Department of Geology,

University of Thessaloniki,

GR‐54124 Thessaloniki,

Greece

O. Kullmer

Forschungsinstitut und

Naturmuseum,

Abt. Palaoanthropologie und

Quartarpalaontologie,

Senckenberganlage 25,

D-60325 Frankfurt am Main,

Germany

J. E. Lambert

Departments of Anthropology and

Zoology,

University of Wisconsin-Madison,

5317 Wm Sewell Social Science

Building,

Madison, WI 53706,

USA

J. Lee-Thorp

Department of Archaeological

Sciences,

University of Bradford,

Bradford, West Yorkshire,

BD7 1DP,

UK

G. A. Macho

Palaeoanthropology Research Group,

Centre for Research of Evolutionary

Anthropology,

Whitelands College,

Roehampton University,

London SW15 4JD,

England

A. E. Mann


Princeton University,

Princeton,

NJ 08544,

USA

A. Meder

Augustenstrasse 122,

D‐70197 Stuttgart,

Germany

P. Menke




Mainz,

D‐55099 Mainz,

Germany

2078 Contributors

S. Mithen

School of Human and

Environmental Sciences,

The University of Reading,

Whiteknights, P.O. Box 217,

Reading, Berkshire,

RG6 6AHm UK

J. Monge


University of Pennsylvania,

University Museum,

3260 South Street,

Philadelphia, PA 19104-6398,

USA

W. Nentwig

Zoologisches Institut,

Universitat Bern,

Baltzer Strasse 6,

CH‐3012 Bern,

Switzerland

N. E. Newton-Fisher


Marlowe Building,

University of Kent,

Canterbury, CT2 7NR,

UK

H. O’Regan

School of Biological and Earth

Sciences,

Liverpool John Moores University,

Liverpool L3 3AF,

UK

M. Ohl

Humboldt-Universitat zu Berlin,

Museum fur Naturkunde,

D‐10115 Berlin,

Invalidenstraße 43,

Germany

B. Pawłowski


University of Wrocław,

ul. Kuznica 35,

Wrocław 50‐138,Poland

U. Radespiel


Tierarztliche Hochschule Hannover,

Bunteweg 17,

D-30559 Hannover,

Germany

T. C. Rae


University of Durham,

43 Old Elvet, Durham,

DH1 3HN,

UK

D. T. Rasmussen


CB 1114, Washington University,

One Brookings Drive,

St. Louis, Missouri 63130,

USA

J. H. Relethford


SUNY College at Oneonta,

Oneonta, NY 13820,

USA

G. J. Retallack


University of Oregon,

Contributors 2079

Eugene, OR 97403‐1272,USA

O. Rieppel


The Field Museum,

1400 S. Lake Shore Drive,

Chicago, IL 60605‐2496,USA

G. P. Rightmire


Peabody Museum,

Harvard University,

Cambridge,

MA 02138,

USA

E. J. Sargis


Yale University,

P.O. Box 208277,

New Haven, CT 06520,

USA

K. Schick

Stone Age Institute,

1392 W. Dittemore Road,

Gosport, IN 47433,

USA

F. Schrenk


Naturmuseum,





Germany

J. H. Schwartz


3302 WWPH,

University of Pittsburgh,

Pittsburgh, PA 15260,

USA

G. R. Scott


University of Nevada,

Reno, NV 89557,

USA

B. Senut

Laboratoire de Paleontologie,

Museum National d’Histoire

Naturelle & UMR 8569,

CNRS,

8, rue Buffon,

75005 Paris,

France

M. T. Silcox

University of Winnipeg,


515 Portage Ave.,

Winnipeg, Manitoba R3B 2E9,

Canada

M. Sponheimer


University of Colorado at Boulder,

Boulder, CO 80309,

USA

F. Spoor

Department of Anatomy &

Developmental Biology,

2080 Contributors

University College London,

Gower St.,

London WC1E 6BT,

UK

J. Steinhauser

Theoretical Biology Group,

Centre for Ecological and

Evolutionary Studies,

University of Groningen,

P.O. Box 14,

9750 AA Haren,

The Netherlands

D. Strait


University at Albany,

1400 Washington Avenue,

Albany, New York 12222,

USA

R. W. Sussman


Washington University at St. Louis,

St. Louis, MO 63130,

USA

C. C. Swisher III

Department of Geological Sciences,

Rutgers University,

610 Taylor Road,

Piscataway,

USA

I. Tattersall

Division of Anthropology,

American Museum of Natural

History,

New York, NY 10024‐5192,USA

M. F. Teaford

Center for Functional Anatomy and

Evolution,

Johns Hopkins University School of

Medicine,

1830 E. Monument St., Room 303,

Baltimore, MD 21205,

USA

A. R. Templeton


Washington University in

St. Louis,

1 Brookings, Campus Box 1137,

St. Louis, MO 63130,

USA

N. Toth

Stone Age Institute,

1392 W. Dittemore Road,

Gosport, IN 47433,

USA

L. A. Tsuji

Museum fur Naturkunde,

Humboldt-Universitat zu Berlin,

D‐10099 Berlin,

Germany

A. Turner

School of Biological and

Earth Sciences,

Liverpool John Moores University,

Liverpool L3 3AF,

UK

C. G. Turner II

School of Human Evolution and

Social Change,

Arizona State University,

Contributors 2081

Tempe, AZ 85287‐2402,USA

L. Ulhaas


Naturmuseum,





Germany

P. S. Ungar


University of Arkansas,

Old Main 330,

Fayetteville, AR 72701,

USA

E. S. Vrba

Department of Geology and

Geophysics,

Yale University,

New Haven, CT 06520,

USA

G. A. Wagner

Geographisches Institut,

Universitat Heidelberg,

Im Neuenheimer Feld 348,

D‐69120 Heidelberg,

Germany

C. V. Ward

Department of Pathology and

Anatomical Sciences,

M263 Medical Sciences Building,

University of Missouri,

One Hospital Drive,

Columbia, MO 65212,

USA

M. Weingarten

Institut fur Philosophie,

Philipps-Universitat Marburg,

Wilhelm-Ropke-Strasse 6, Block B,

D‐35032 Marburg,

Germany

B. Wood

Center for the Advanced Study of

Human Paleobiology, Department of

Anthropology,

The George Washington University,

2110 G St. NW,

Washington, DC 20052,

USA

E. Zimmermann


Tierarztliche Hochschule

Hannover,

Bunteweg 17,

D-30559 Hannover,

Germany

H. Zischler




Mainz,

D‐55099 Mainz,

Germany

9 The Earliest Putative HomoFossilsFriedemann Schrenk . Ottmar Kullmer . Timothy Bromage

Abstract

The earliest fossil remains of the genus Homo have been discovered in eastern,

southeastern, and southern Africa. The sample comprises about 200 skeletal

fragments attributable to about 40 individuals and assigned to two species:

Homo rudolfensis (2.5–1.8 Ma) showing a combination of primitive dentition

with Homo‐like locomotion andHomo habilis (2.1–1.5 Ma) exhibiting a progres-

sive reduction of tooth roots but resembling great apes rather than humans in

the postcranial skeleton. Another significant difference between early Homo and

the australopithecines is the brain size, which was larger in early Homo than

Australopithecus but smaller than in Homo erectus. Endocasts of H. habilis from

Olduvai Gorge and Koobi Fora reveal a number of distinctive features some of

which are recognized as Homo autapomorphies. Differences in tooth wear

betweenH. rudolfensis, with megadont teeth and more horizontal tooth abrasion,

and H. habilis, with more gracile molars and higher relief in worn teeth indicate

significant differences in diet and ecology of early Homo species. The origin of

the genusHomo coincided with the onset of material culture. Between ca. 2.8 and

2.5 Ma, extensive open habitats comprising more arid‐tolerant vegetation devel-

oped in Africa. The selective pressures of this habitat change resulted in the

increased survival of more megadont species varieties. Megadonty allowed these

species to feed on tougher fruit and open woodland–open savannah food items

resulting in the phyletic splitting of Australopithecus afarensis into Paranthropus

and Homo lineages by ca. 2.5 Ma. An evolutionary scenario that complies with

both the Habitat Theory and early hominid biogeography is provided. It deline-

ates the association between faunal turnover and climate change, and suggests a

single origin for the Paranthropus lineage but separate origins for H. rudolfensis

and H. habilis from A. afarensis and A. africanus ancestors, respectively.

# Springer-Verlag Berlin Heidelberg 2007

1612 9 The earliest putative Homo fossils

9.1 Introduction

The search for the roots of the genus Homo is of particular interest in the field of

paleoanthroplogy. The taxonomic determination of the earliest putative Homo

fossils provides the basis for the definition of the taxonHomo to which all modern

humans belong.

Carolus Linnaeus (1758) established the genus Homo in the tenth revision of

his Systema Naturae. In his opinion, Homo subsumed six groups: H. sylvestris

H. troglodytes—a mixture of orangutan and myths, H. sapiens, and four geo-

graphical variants from Africa, America, Asia, and Europe. In the two centuries

that followed H. neanderthalensis (King 1864), H. heidelbergensis (Schoetensack

1908), H. erectus (Dubois 1892; Mayr 1944), H. habilis (Leakey et al. 1964),

H. ergaster (Groves and Mazak 1975), H. rudolfensis (Alexeev 1986), H. antecessor

(Bermudez de Castro et al. 1997), and others were included in the genus Homo.

The history of research, the order of discoveries, and existing paradigms

heavily influence the formation and change in interpretations of human evolu-

tion. This holds true especially regarding ideas on the origin of the genus Homo

since the 1960s. Debates on the attribution of fossil specimens and the definition

of the genus Homo continue today (Wood 1992).

Opinions differ regarding the number of species and also the specimens

included in the genus. Some even assign all putative Homo specimens to living

humans (H. sapiens). Species names in paleoanthropology are labels rather than

natural species, and the taxonomic determination of fossils is more or less a

question of the philosophy followed by the authors. There are both theoretical

and practical reasons to erect taxa, as chronospecies for time equivalent appear-

ance or morphospecies for a complex of shared anatomical features, and in the

worst case there are political reasons for the allocation of species.

9.2 Fossil evidence

Early research on the origin of the genus Homo is closely related to the African

fieldwork of Louis Leakey (1903–1972). He strongly believed in Africa as

the cradle of humankind and in 1932 discovered the first evidence of early

Homo at Kanam, (Kenya) east of Lake Victoria—a specimen, which today is

attributed to H. erectus. He also undertook archeological surveys in Olduvai

Gorge, Tanzania (> Figure 9.1), where later he discovered early pebble tools in

Bed I (ca. 1.8 Ma)—remains of what he termed the ‘‘Oldowan’’ industry. The

search for the artifacts creator led to the discovery of robust australopithecine

remains(Zinjanthropus boisei) (Leakey 1959). However, due to its small brain size,

. Figure 9.1African early hominid sites. Homo rudolfensis and Homo habilis sites in bold

The earliest putative Homo fossils 9 1613

this specimen Olduvai Hominid 5 (OH5) was not a convincing candidate for the

first toolmakers.

A year later, Jonathan Leakey, the son of Louis, discovered two fragments of a

relatively gracile skull, a lower jaw (> Figure 9.2), and several hand bones of OH 7

(Leakey 1961), deriving from the same stratigraphic level (Bed 1) in Olduvai.

Brain volume was estimated at around 680 cm3, a significantly higher value than

in robust australopithecines. Consequently, this fossil was interpreted as repre-

senting a progressive hominid type of unknown species affiliation. Later Leakey

et al. (1964) decided on a new speciesH. habilis. Raymond A. Dart, the founder of

modern paleoanthropology in Africa, who in 1925 had introduced the genus

Australopithecus, had suggested this name to them. The Latin term ‘‘habilis’’ means

. Figure 9.2Type specimen of Homo habilis: mandible OH 7, Olduvai Gorge, Tansania


‘‘handy, skillful, able’’: finally, the producer of the Oldowan culture seemed to have

been identified.

Apart from OH 7, the species description of H. habilis included skull frag-

ments and teeth (OH 4 and OH 6), part of an adult foot (OH 8), and the

incomplete skull of an adolescent (OH 13). Further, Leakey et al. (1964) referred

a collection of juvenile cranial pieces (OH 14) and the fragmented cranial vault

and dentition (OH 16) of a young adult to the new species.

Since then, numerous additional fossils of H. habilis have been discovered at

Olduvai Gorge: 9 fragmentary skulls, 4 mandible fragments, 19 teeth, and 8 post-

cranial fragments. Among these fragments was the squashed skull OH 24

(Twiggy), which was found in 1968. In 1986, a partial female skeleton (OH 62)

was assembled from a number of fragments (Johanson et al. 1987). This specimen

showed that H. habilis was fully bipedal and had a brain larger than all australo-

pithecines. For many years, the H. habilis remains from Olduvai Gorge were seen

as the most important early Homo specimens and consequently played the

leading role in most hypotheses regarding the origin of genus Homo.


Leakey et al. (1964) originally discussed cranial and mandibular traits, to

distinguish the Homo specimens of Olduvai from Australopithecines and

H. erectus. Maxillary and mandibular size is smaller than in Australopithecus

and tends in size to H. erectus and H. sapiens. The surface of the skull shows

slight to strong muscular markings and the parietal curvature in the sagittal plane

varies from slight to moderate. The frontal bone is more vertical and the torus

supraorbitalis is less developed than in australopithecines. In the occipital region,

the relatively open‐angled external sagittal curvature differs markedly from

Australopithecus.

In 1970 the picture of earliest Homo began to change significantly with the

success of the Koobi Fora Research Project in northern Kenya and led by Richard

Leakey, Louis’ son. In just a few years on the eastern shores of Lake Turkana, his

team recovered 9 skulls, 10 mandibles, 6 isolated teeth, and 5 postcranial frag-

ments (Leakey 1973a,b). Originally, all of the early Homo finds from East

Turkana, with an age similar to those from Olduvai (1.9–1.8 Ma), were inter-

preted as bearing similarities to H. habilis—then the only early species of Homo.

One cranial fragment from the Nachukui Formation on the western shores of

Lake Turkana was also assigned to H. habilis. However, two of the best preserved

skulls from Koobi Fora (KNM‐ER1470,> Figure 9.4; KNM‐ER 1813, > Figure 9.5)

later gave rise to an extended debate among researchers about the heterogeneity

of the H. habilis hypodigm, and finally led to the recognition of a new species,

H. rudolfensis.

In the 1970s a large number of isolatedHomo teeth were discovered north of

Koobi Fora, near the Omo‐River in southern Ethiopia, in Member G and H of the

Shungura Formation. From these it became clear that the origin of the genus

Homo extended well beyond 2 Ma. Already in 1965, a temporal bone had been

discovered by John Martyn at Chemero, Kenya, which nearly two decades later

was described by Hill et al. (1992) as a very early member of the genus Homo,

dated to around 2.4 Ma.

In 1976, an early Homo fossil was found at Sterkfontein, South Africa

(Stw 53), which belonged neither to H. erectus nor to Australopithecus (Hughes

and Tobias 1977). A partial facial skull (SK 847 from Swartkrans), assembled from

several fragments, originally attributed to a different species, is further evidence

for H. habilis, which probably migrated into southern Africa around 2 Ma (see

below).

The geographical gap between the southern and eastern African early homi-

nid sites was filled in the early 1990s through discoveries in the ‘‘Hominid

Corridor’’ of the northernMalawi Rift (Schrenk et al. 1993; Bromage and Schrenk

1995). In 1992, at Uraha, the Hominid Corridor Research Project (HCRP) has


recovered a mandibular corpus, UR 501 (> Figure 9.3), containing third and

fourth premolars and first and second molars in variable states of preservation

. Figure 9.3UR 501 from the Chiwondo Beds, northern Malawi (ca. 2.5–2.4 Ma), Homo rudolfensis(Drawing: Claudia Schnubel)

(Schrenk et al. 1993). Many absolute and relative measures defining molar and

premolar crown shape indices, relative cusp areas, fissure patterns and enamel

microanatomical features, as well as overall crown morphology, are within the

sample range of early Homo, although some may also be subsumed within the

limits of variation represented by Australopithecus (A. africanus and A. afarensis).

However, UR 501 has absolutely large molar crown areas, relative expansion of

the P3 talonid, plate like P3 and P4 roots, and some enamel microanatomical

features correspond more closely to the Paranthropus condition. UR 501 corre-

sponds closely to the subset of Late Pliocene fossils from east Turkana, Kenya,

which demonstrates relatively large brains and robust jaws and teeth and based

on the above have been assigned to H. rudolfensis by Alexeev (1986) and Wood

(1992), and to which UR 501 was also referred (Bromage et al. 1995). This Malawi

specimen has been dated by biostratigraphic correlation of suid material with

well‐dated sites in southern Ethiopia (Omo Shungura) and northern Kenya

(Koobi Fora), indicating an age of about 2.4–2.5 Ma whereas most early Homo

fossils are around 2 Myr old.


It is important to note that the appearance of earliest Homo is contempora-

neous with the origin of hyperrobust australopithecines (Paranthropus). The

earliest evidence for this co‐existence is based on further hominid discoveries in

the Chiwondo Beds of Northern Malawi. A maxillary fragment (RC 911) pre-

serves part of the left alveolar process, with badly damaged M1 crown and

fragmentary M2 crown. Size, morphology and abrasion occlusal pattern on the

surface suggest that RC 911 should be assigned to Paranthropus boisei (Kullmer

et al. 1999).

The biogeographic significance of these Malawi Rift hominids lay in their

association with the eastern African endemic faunal group. The associated bovid

and suid faunas show a small amount of overlap with southern African animals

and a greater overlap with eastern African faunal elements. Biogeographic varia-

tion, in the Malawi Rift may be linked to habitat change occurring due to climate

shifts, with maximum change occurring around 2.5 Ma.

9.3 Changing taxonomy

Hominid fossils are generally assigned to the genus Homo if they fulfill four main

criteria (Keith 1948; Tobias 1991; Wood and Collard 2001): a brain size above

600 cm3, ability for speech and tool‐making, as well as an opposable pollux. To

date, the hypodigm of earliest Homo attributed to H. habilis sensu stricto and

H. rudolfensis contains about 200 skeletal fragments attributable to about 40 indi-

viduals (> Tables 9.1 and > 9.2). Despite—or even due to—the large number of

specimens, the taxonomic interpretation of earliest Homo is highly controversial

(Wood 2000).

Originally, the interpretation of the early Homo hand as ‘‘modern’’ (Leakey

et al. 1964) supported the view of H. habilis as an early but ‘‘able’’ human as

opposed to the rather ‘‘clumsy’’ australopithecines. However, later skeletal finds

at Olduvai Gorge (OH 62) (Johanson et al. 1987) demonstrated that the postcra-

nial skeleton ofH. habilis indeed resembled Australopithecus africanus rather than

Homo. Yet the most distinctive character of H. habilis remains its relatively and

absolutely higher brain volume compared to that of Australopithecus. The fore-

head of H. habilis is more vertical and a weak supraorbital torus is present.

Whereas the morphological characters are quite uniform in the Olduvai sample,

the discussion started to heat up mainly over two very distinct skull fragments

from Koobi Fora: KNM‐ER 1470 (> Figure 9.4) (Leakey 1973a) and KNM‐ER1813 (> Figure 9.5) (Leakey 1973b).

In a comprehensive character analysis of all available putative H. habilis

fossils from Koobi Fora, Wood (1991) concluded that the variability exhibited

. Table 9.1

Significant morphological differences between H. habilis and H. rudolfensis

Homo habilis sensu stricto Homo rudolfensis

Skull and teethAbsolute brainsize (cm3)

An average volume of 610 An average volume of 751

Overall cranialvaultmorphology

Enlarged occipital contributionto the sagittal arc

Primitive condition

Endocranialmorphology

Primitive sulcal pattern Frontal lobe asymmetry

Suture pattern Complex SimpleFrontal Incipient supraorbital torus Torus absentParietal Coronal > sagittal chord Primitive conditionFace‐overall Upper face > midface breadth Midface > upperface breadth:

markedly orthognathicNose Margins sharp and everted;

evident nasal sillLess everted margins; no nasal sill

Malar surface Vertical or near vertical Anteriorly inclinedPalate Foreshortened LargeUpper teeth Probably two‐rooted

premolarsPremolars three‐rooted; absolutelyand relatively large anterior teeth

Mandibular fossa Relatively deep ShallowForamenmagnum

Orientation variable Anteriorly inclined

Mandibularcorpus

Moderate relief on externalsurface; rounded base

Marked relief on external surface;everted base

Lower teeth Buccolingually narrowed;postcanine crowns; reducedtalonid on P4; M3 reduction;mostly single‐rootedmandibular premolars

Broad postcanine crowns; relativelylarge P4 talonid; no M3 reduction;twin, platelike P4 roots, and bifid,or even twin, platelike P3 roots

PostcraniumLimb proportions Apelike ?Forelimbrobusticity

Apelike ?

Hand Mosaic of apelike and modernhumanlike features

?

Hindfoot Retains climbing adaptations Later Homo‐likeFemur Australopithecine‐like Later Homo‐like

After Wood (1992).


by the sample was not only the result of sexual dimorphism as was suspected

at the time, but that highly significant differences exist throughout the entire

skeleton. There is a mosaic of Australopithecus and Homo characters in both early

species: WhereasH. rudolfensis exhibits a combination of ancestral dentition with

Homo‐like locomotion, H. habilis shows a progressive reduction of tooth roots

and resembles great apes rather than humans postcranially.

. Table 9.2

Fossil remains of Homo habilis sensu stricto

Homo habilis (better‐preserved specimens in bold)

Sites Skulls and crania Mandibles Isolated teeth Postcranial

Olduvai(OH)

6, 7, 13, 14, 16, 24,52, 62

7, 13, 37, 62 4, 6, 15, 16, 17, 21,27, 31, 32, 39, 40,41, 42, 44, 45, 46,47, 55, 56

7, 8, 10, 35,43, 48, 49,50, 62

Koobi Fora(KNM‐ER)

807, 1478, 1805,1813, 3735

1501, 1502,1506, 1805, 3734

808, 809, 1462,1480, 1508, 1814

813, 1472,1481, 3228,3735

Omo L894‐1 Omo 222‐2744 L28‐31; L398‐573,1699; Omo 33‐3282,Omo 47‐47; Omo74‐18; Omo123‐5495; Omo166‐781;OmoK7‐19;OmoSH1‐17; P933‐1

WestTurkana

Kangaki I site(w/o number)

– – –

Sterkfontein Stw 53, SE 255,1508, 1579, 1937,2396; Sts 19

– – –

Swartkrans SK 847 (?) – – –


Based on the work of Wood (1991) and subsequent re‐evaluation of fossils it

is evident that two distinct types can be separated. Whereas one group, repre-

sented by KNM‐ER 1813, follows the original description of H. habilis from

Olduvai Gorge, a new group is represented by KNM‐ER 1470, for which no

comparison existed in Olduvai at the time. Although the subsequent find of

OH 65 at Olduvai (Blumenschine et al. 2003) showed a mixture ofH. habilis sensu

stricto and KNM‐ER 1470 features, it is still a valid conclusion that about half of the

early Homo material from Koobi Fora belongs to H. habilis sensu stricto, with an

age of 2.1–1.5 Ma, which includes also specimens from Koobi Fora,

West‐Turkana, Omo, Olduvai Gorge, and southern Africa (> Table 9.2). Hominid

remains from Ubeidiyah in Israel (Leakey et al. 1964) and Meganthropus palaeo-

javanicus from Java, Indonesia, which at one stage were tentatively assigned to

H. habilis (Tobias and von Koenigswald 1964), are not considered as such today.

The Koobi Fora early Homo material not assigned to H. habilis is defined as

the more recently named species H. rudolfensis, with an age of 2.5–1.8 Ma, which

also includes specimens from Chemeron, West‐Turkana (Prat et al. 2005), and

Omo (Suwa et al. 1996; Ramirez Rozzi 1997) as well as northern Malawi (Schrenk

et al. 1993) (> Table 9.3).

. Figure 9.4KNM‐ER 1470 from Koobi Fora, Kenya (ca. 1.9 Ma), Homo rudolfensis (Drawing: ClaudiaSchnubel)


The species name rudolfensiswas coined by Russian paleontologist V.A. Alexeev,

who in 1986 described KNM‐ER 1470 as ‘‘Pithecanthropus rudolfensis,’’ after Lake

Rudolf, the name of Lake Turkana prior to Kenyan independence in 1963.

9.4 Postcranial skeleton

The postcranial skeleton of H. habilis was characterized by Leakey et al. (1964)

using a number of characteristics: the clavicle resembles that of H. sapiens, the

hand shows broad terminal phalanges, and capitate and MCP articulations also

resemble H. sapiens, but differ in respect of the scaphoid and trapezium, attach-

ments of the superficial flexor tendons, and the robusticity and curvature of the

phalanges. The foot bones resemble H. sapiens in the stout and adducted hallux

and well‐defined foot arches, but differ in shape of the talan trochlea surface and

the relatively robust third metatarsal.

. Figure 9.5KNM‐ER 1813 from Koobi Fora, Kenya (ca. 1.9 Ma), Homo habilis (Drawing: ClaudiaSchnubel)


All these features indicate an affinity towards H. sapiens and underline that

the early postcranial material from Olduvai seems to be different from that of the

australopithecines. Functional implications based on more detailed descriptions

of the foot (OH 8, OH 10) and leg (OH 35), both of which are probably from the

same individual (Stern and Susman 1983), were generally more cautious (Wood

1992). According to Stern and Susman (1983), H. habilis had not reached the

characteristic bipedal gait of H. sapiens, since the functional morphology of the

knee joint was not well adapted for striding. Further analysis of the OH 8 foot

demonstrated that some features, common in nonhuman hominoids, are also

present in the foot bones (Lewis 1989). As more postcranial material was uncov-

ered at Koobi Fora, interpretation of hind limb function of early Homo became

more complex. However, some specimens, such as the femur KNM‐ER‐1472Aand the talus KNM‐ER‐813, may belong toH. erectus orH. ergaster rather than to

H. habilis (Tobias 1991). Although the partial skeleton OH 62 from Olduvai

. Table 9.3

Fossil remains of Homo rudolfensis

Homo rudolfensis (better‐preserved specimens in bold)

SitesSkulls andcrania Mandibles Isolated Teeth Postcranial

Koobi Fora(KNM‐ER)

1470, 1590,3732, 3891

819, 1482, 1483,1801, 1802

– –

Chemeron KNM‐BC 1 – – –Omo Omo 75‐14 ? L7‐279; L26‐1g; L28‐30,

L628‐10; Omo 29‐43;Omo 33‐740, 5495, 5496;Omo 75i‐1255; Omo 75s‐15, 16; Omo 177‐4525;Omo 195‐1630

WestTurkana

Lokalalei? WT42718

– – –

Uraha (UR) – 501 – –


Gorge was interpreted as evidence for fully upright human bipedal locomotion,

Haeusler and McHenry (2004) demonstrated that there is little evidence in

support of ancestral body proportions with short legs and long arms inH. habilis.

Their results suggest that it is more likely that earliest Homo possessed an

elongation of the legs relative to A. africanus and A. afarensis, whereas long

forearms were still retained (Johanson et al. 1987). With its upper‐to‐lowerlimb shaft length proportions, OH 62 falls within the upper range of modern

humans and the lower range of chimpanzees due to the partial overlap between

these taxa at small body sizes. KNM‐ER 3735, the larger‐bodied early Homo from

Koobi Fora, falls well outside chimpanzees and reflects the average proportions of

modern humans. Comparison of the Koobi Fora and Hadar postcranial remains

leads to the interpretation that H. habilis did probably possess a modern pattern

of limb shaft proportions and the body proportions of OH 62 are in agreement

with other available evidence of H. habilis postcranial material (Johanson et al.

1987). The change in the proportions of limb length toward the development of

long legs may be indicative of long‐distance terrestrial walking in early Homo and

probably implies a shift in hominid ecology.

9.5 Brain and language

A significant difference between early Homo and Australopithecines exists in

brain size, which was larger in early Homo than in Australopithecus but smaller


than H. erectus. Endocasts of H. habilis from Olduvai and Koobi Fora reveal a

number of distinctive features, some of which are recognized as autapomorphies

of the genus Homo. Tobias (1987) defined the principal morphological trait to

distinguishH. habilis from Australopithecus as a larger mean endocranial capacity

in the former (640 cm3) than in A. africanus (441 cm3), A. boisei (513 cm3), and

A. robustus (530 cm3). This suggests that the evolutionary trend toward brain

expansion was already well under way more than 2 Ma. The H. habilis mean

(640 cm3) is close to the lower limit (647 cm3) of the 95% population range of

H. erectus but well above the upper limit of the A. africanus range (492 cm3). The

brain capacity of the H. rudolfensis type specimen (750 cm3) is larger than the

known range for H. habilis from Olduvai Gorge and Koobi Fora and falls within

the lower range of H. erectus.

A prominent feature of the H. habilis brain is the bilateral transverse expan-

sion of the cerebrum, especially in the frontal and parieto‐occipital areas, and a

posterior heightening. The increased bulk of cerebral frontal and parietal lobes

and the sulcal and gyral pattern of the lateral frontal lobe have been interpreted as

derived features for the genusHomo (Tobias 1987). TheH. habilis brain showed a

well‐developed left superior parietal lobule, and a prominent development of the

inferior parietal lobule.

The endocast of KNM‐ER 1470 shows a left frontal lobe sulcal pattern that is

associated with Broca’s area in living people (Falk 1987), a finding that has led to

the conclusion that H. rudolfensismay have been capable speech. This conclusion

is in accordance with Holloway’s observation of a pronounced left‐occipital‐right‐frontal petalia pattern in the KNM‐ER‐1470 endocast that may indicate

functional cortical asymmetry (Holloway 1983). Surprising corroborative evi-

dence has been provided by Toth’s analyses of stone flakes, which indicate that

hominids were predominantly right‐handed by 2 Ma (Toth 1985). Adjacent areas

in the left frontal lobe control the speech organs and the right hand. Tobias (1991)

stated that H. habilis is the earliest hominid to show prominent enlargement of

Broca’s and Wernicke’s areas. If so, the same should be seen in H. rudolfensis.

Australopithecus endocasts show Broca’s area, but not Wernicke’s region,

while anthropoid apes display neither of the two. The prominent develop-

ment of the two speech areas may thus be seen as an important autapomor-

phy of the genus Homo (Tobias 1991). Even if H. habilis and H. rudolfensis

possessed the neurological basis of speech, there is no evidence that either of them

used spoken language. The areas of the brain that control spoken communication

probably manifested themselves only when brain enlargement occurred and

marked encephalization started.


9.6 Material culture and food processing

Around 2.5 Ma, simultaneously with an increase in drier and harder food stuffs

due to increasing aridity in eastern Africa (de Menocal 2004), there occurred

the first hyperrobust australopithecines (Paranthropus) and the genus Homo in

the fossil record (Bromage et al. 1995). This demonstrates an evolutionary

alternative to the massive masticationary system of Paranthropus, which was

capable of dealing with very tough foods. The evolutionary alternative to mega-

donty was the manufacturing and use of stone tools. The oldest chopper tools

are known from Ethiopia (Hata, Bouri Formation) and from Tanzania, approxi-

mately 2.5 Ma (Kaiser et al. 1995). From Gona, east of Hadar in the Afar Triangle,

primitive pebble tools are dated to 2.6 Ma (Harris 1986). New discoveries from

west of Lake Turkana confirm the existence of an early tool culture around 2.5 Ma

(Roche et al. 2003). In the Hadar area of Ethiopia, stone tools were found

associated with early Homo remains (Kimbel et al. 1996). At many sites the

presence of more than one hominid species, occurring in the same horizon as

early Oldowan pebbles does not give clear evidence of who were the first tool-

makers. However, the distinct specialization of the skull and dental morphology

in robust australopithecines and brain expansion in earlyHomo point to the latter

as the most likely tool manufacturer.

Implements are widely used by higher primates. Yet during marked habitat

shifts, which led to pronounced changes in food resources, it probably was the

invention of stone tools, which supported the origin of the genus Homo around

2.5 Ma. Increasing independence from the environment led to an increase in the

dependence on culture.

If early Homo utilized stone tools to prepare food, the dentition might

actually indicate these behavioral changes in food acquisition. However, the

morphology of early Homo teeth does not seem to suggest extensive food

preparation before ingestion. The incisors are large compared to those of Aus-

tralopithecus and H. erectus, and the canines are large relative to the premolar

crown surfaces. The premolars are narrower than in Australopithecus and fall

within the range of H. erectus. Molar size overlaps the ranges for Australopithecus

and H. erectus. The cheek teeth of H. rudolfensis are enlarged and show affinities

to Paranthropus molars. All teeth are relatively narrow buccolingually and elon-

gated mesiodistally, especially the mandibular molars and premolars. InH. habilis

we see well‐developed third molars, while in H. rudolfensis the third molar forms

a smaller crown than the second molar. The occlusal surface of the cheek teeth is

not as broad as in australopithecine molars and indicates differences in chewing.

Tobias (1987) states that the crown’s cusps relief is still present even when the

teeth are in advanced wear and dentine is visible. This means that the attrition of


enamel is less pronounced than in earlier hominids. Differences in tooth wear

between H. rudolfensis, with megadont teeth and a more horizontal tooth abra-

sion, and H. habilis, with its more gracile molars and higher relief in worn teeth

are clearly visible. This indicates significant differences in diet and ecology of early

H. species. H. rudolfensis and the robust australopithecines share some cranial

and dental features regarding the morphology of the masticatory apparatus

(Wood 1992), which indicates that these hominids were able to cope with

tough fruits and plants. Since those features are judged as an adaptation to

drier climatic conditions, they also show that H. rudolfensis was relatively conser-

vative nutritionally and probably followed a more herbivorous strategy in food

acquisition.

9.7 Biogeographic scenario

The scenario presented here is derived mainly from hypotheses about early

hominid evolution in the context of environmental change and faunal biogeog-

raphy. It is thus a biogeographic perspective against which we understand the

relevance to studies of early Homo systematics in general, and morphology and

character transformation more specifically.

The behavioral inclination of the earliest hominids distributed along the

margin of the tropical rain forest, was to maintain a connection to and remain

near the borders between broad riparian habitats and open woodlands during the

ascendancy of more warm and humid times. Over short geological timescales, this

was typically a local, nondispersing tendency, but by approximately 4 Ma, several

species of Australopithecus had successfully dispersed throughout the reaches of

the African Rift Valley and into western Africa (> Figure 9.7). Over longer time

frames, this included dispersal through the riparian ‘‘corridor’’ connecting

eastern and southern Africa, permitting population dispersal into southern Africa

by 3 Ma. This dispersing population maintained habitat specificities to forested

environments (Rayner et al. 1993), though in more environmentally temperate

climes and in relative geographical isolation at the extreme distal edge of its

distribution. The dispersion along changing latitudinal circumstances covar-

ied with its transformation into, first, a geographic variant and, subsequently,

into A. africanus, joining ranks with other southern African endemic faunas.

Thus, A. afarensis was essentially an eastern African endemic and it follows that

no typical representatives are likely to be recovered from southern African

deposits older than 3.5 Ma. (> Figures 9.6 and > 9.7).

By approximately 2.8 Ma, the initiation of cooler and drier conditions

prevailed upon the African landscape, its vegetation, and its faunas, until

. Figure 9.6Hominid chronology


climaxing ca. 2.5 Ma (Bonnefille 1980; Vrba 1985, 1988; Prentice and Denton

1988; de Menocal 2004). During this time, A. afarensis in eastern Africa and

A. africanus in southern Africa were each subject to unique paleobiogeographic

. Figure 9.7Early hominid biogeography, dispersal and migration in Africa


consequences of this global aridification, in accordance with the ‘‘Habitat

Theory’’ of Vrba (1992).

For A. afarensis, then, the changing climate meant vicariance of its habitat

and its distribution into more removed ecotonal riparian and closed lake margin

environs. During the interim between ca. 2.8 and 2.5 Ma, these changing condi-

tions engendered more extensive open habitats comprising more resistant arid‐tolerant vegetation around the remaining relatively lush but narrowed ‘‘ribbons’’

of tree‐lined riverine forest. The selective pressures of this habitat change resulted

in the increased survival of more megadont varieties capable of feeding on

tougher fruit and open woodland–open savannah food items. This was so for

early hominid as well as numerous eastern and southern African large terrestrial

vertebrate lineages ca. 2.5 Ma (Turner and Wood 1993). These pressures were


likewise sufficient to result in the phyletic splitting of A. afarensis into Paranthro-

pus and Homo lineages by ca. 2.5 Ma (Vrba 1988) (> Figures 9.6 and > 9.7).

Ensuing cooler and drier conditions favored a tougher savannah vegetation

composed of plant species better able to retain their moisture under such condi-

tions. Selection favored more facially robust and large molar‐toothed mammals,

including early hominids, capable of efficiently processing the tougher, more

durable vegetation of the savannah. Our evidence suggests that the tropical

equatorial animals, including the hominids, of eastern Africa stayed in the

tropical African ecological domain, while during the drying and cooling of global

climates ca. 2.5 Ma, the southern and more temperate African faunas followed

their northward‐drifting vegetation zones. Thus, Homo and Paranthropus may

have emerged in tropical Africa as a result of the ca. 2.5‐Ma climatic cooling event

and remained there.

The beginnings of the Paranthropus lineage maintained a reliance on fruiting

resources on the riverine side of its ecotone, particularly during the dry season,

but it was equally adept at grinding on the postcanine dentition those food items

it required from more open habitats during more hospitable times of the year.

The beginnings of the Homo lineage, also ca. 2.5 Ma, and represented by

H. rudolfensis, was an endorsement of its recency of common ancestry with

A. afarensis, a distinction it shared with Paranthropus (Bromage et al. 1995).

However, while Paranthropus was principally adapted by means of a robust

masticatory system to its tough and abrasive diet, H. rudolfensis exhibited an

increased behavioral flexibility as its adaptation to climatic circumstances includ-

ed a larger and more provoking, inquiring, and capable brain. This included a

shift to proportionately less abrasive foodstuffs and more omnivorous habits.

Material culture ameliorated the effects of climate change to the degree that it

enabled H. rudolfensis to take advantage of other resources more efficiently than

was ever possible before.

P. boisei and H. rudolfensis remained endemic to tropical latitudes during

this time (Bromage et al. 1995). The eastern African tropical faunas, having

habitable alternatives, remained within their biogeographic domain rather than

brave the relative deterioration and paucity of habitats south of the African Rift

Valley.

The faunas of southern Africa were subject to a different set of environmental

sequelae during the ca. 2.5‐Ma cooling event. Waning of the forests and woodlands

in deference to more open arid grasslands invigorated not only evolutionary

adaptations to savannah life in tropical eastern Africa but also resulted in

the distribution drift northward of faunas tracking the equator ward shift of

grassland and woodland biomes into eastern Africa from the south ca. 2.5 Ma

(Bromage et al. 1995). The temperate zone ca. 2.5 Ma experienced more seasonal


extremes and many organisms unwittingly maintained their inherited preference

for moderate seasonal climes and temperate vegetation physiognomy by moving

northward with the shrinking of this biome toward the equator, effectively

transgressing the Zambezi Ecozone. Among these migrants was A. africanus

who, having been adapted to a modest temperate ecology, now found its suitable

habitats shifted to the north toward the African Rift Valley. While dispersing

toward the eastern African tropical domain, selection for increased behavioral

flexibility was related to the habitat diversity of the tropics and the presence of

other nonvegetative food resources available in their new region. This emerging

taxon, H. habilis, rapidly established itself as a categorical omnivore and found

that it could buffer itself more resolutely from environmental changes. This

enabled it to cross habitat boundaries more easily and also to advantage itself

of more resources with its material culture.

By approximately 2 Ma, Africa was rebounding from its relatively cool and dry

climate to return to slightly more warm and humid conditions (Bromage et al.

1995). A phase of biome expansion ensued that facilitated dispersions away from

the equator, ending nearly 1 Myr of relative endemism dominated by tropical

equatorial speciations. P. boisei dispersed southward along reestablished ecotonal

habitats into southern Africa, varied there as a geographic variant under more

temperate conditions, and evolved into Paranthropus robustus (> Figure 9.6).

H. habilis expanded southward into the southern African temperate domain,

but it maintained a very much broader niche and increased its distributional area

as a single species.H. rudolfensis remained endemic to the eastern African tropical

domain due partly to its preference for more open habitats around the rain

shadows of the African Rift Valley and partly, perhaps, to some small measure

of competitive exclusion from geographic realms occupied by H. habilis.

References

Alexeev VP (1986) The origin of the human

race. Progress Publishers, Moscow

Bermudez de Castro JM, Arsuaga JL, Carbonell

E, Rosas A, Martinez I, Mosqueria M (1997)

A hominid from the Lower Pleistocene of

Atapuerca, Spain possible ancestor to Nean-

derthals and modern humans. Science 276:

1392–1395

Blumenschine RJ, Peters CR, Masao FT, Clarke

RJ, Deino A, Hay RL, et al. (2003) Late Plio-

cene Homo and hominid land use from

western Olduvai Gorge, Tanzania. Science

299: 1217–1221

Bonnefille R (1980) Vegetation history of sa-

vanna in East Africa during the Pleistocene.

Proc IV Int Palyn Conf Lucknow 3: 75–89

Bromage TG, Schrenk F (eds) (1995) Evolu-

tionary history of the Malawi Rift. J Hum

Evol 28: 1–120

Bromage TG, Schrenk F, Zonneveld FW (1995)

Paleoanthropology of the Malawi Rift: an

early hominid mandible from the Chiwondo


Beds, northern Malawi. J Hum Evol 28:

71–108

de Menocal PB (2004) African climate change

and faunal evolution during the Pliocene‐Pleistocene, Earth Planet Sci Lett 220:

3–24

Dubois E (1892) Palaeontologische anderzoe-

kingen op Java. Verslag van het Mijnwezen.

Batavia 3: 10–14

Falk D (1987) Hominid paleoneurology. Annu

Rev Anthropol 16: 13–30

Groves CP, Mazak V (1975) An approach to the

taxonomy of the Hominidae: Gracile Villa-

franchian hominids of Africa. Casopis pro

Mineralogii A Geologii 20: 225–247

Haeusler M, McHenry HM (2004) Body pro-

portions of Homo habilis reviewed. J Hum

Evol 46: 433–465

Harris JWK (1986) Decouverte de materiel

archeologique oldowayen dans le rift de

l’Afar. L’Anthropologie 90: 339–357

Hill A, Ward S, Deino A, Curtis G, Drake R

(1992) The earliest Homo. Nature 355:

719–722

Holloway RL (1983) Human brain evolution: a

search for units, models and synthesis. Can J

Anthropol 3: 215–230

Hughes AR, Tobias PV (1977) A fossil skull

probably of the genus Homo from Sterkfon-

tein, Transvaal. Nature 265: 310–312

Johanson DC, Masao FT, Eck GG, White TD,

Walter RC, Kimbel WH, Asfaw B, Manega P,

Ndessokia P, Suwa G (1987) New partial

skeleton of Homo habilis from Olduvai

Gorge, Tanzania. Nature 327: 205–209

Kaiser T, Bromage TG, Schrenk F (1995) Hom-

inid corridor research project update: new

Pliocene fossil localities at Lake Manyara

and putative oldest Early Stone Age occur-

rences at Laetoli (Upper Ndolanya Beds),

northern Tanzania. J Hum Evol 28: 117–120

Keith A (1948) A new theory of human evolu-

tion. CAWatts & Company, London

Kimbel WH, Walter RC, Johanson DC, Red KE,

Aronson JL, Assefa Z, Marean CW, Eck GG,

Bobe R, Hovers E, Rak Y, Vondra C, Yemane

T, York D, Chen Y, Evensen NM, Smith PEB

(1996) Late Pliocene Homo and Oldowan

tools from the Hadar Formation (Kada

Hadar Member), Ethiopia. J Hum Evol 31:

549–562

King W (1864) The reputed fossil man of the

Neanderthal. Q J Sci 1: 88–97

Kullmer O, Sandrock O, Abel R, Schrenk F,

Bromage TG, Juwayeyi Y (1999) The first

Paranthropus from the Malawi Rift. J Hum

Evol 37: 121–127

Leakey RE (1973a) Evidence for an advanced

plio‐pleistocene hominid from East Rudolf,

Kenya. Nature 242: 447–450

Leakey RE (1973b) Further evidence of Lower

Pleistocene hominids from East Rudolf,

north Kenya. Nature 248: 653–656

Leakey LSB, Tobias PV, Napier JR (1964) A new

species of the genus Homo from Olduvai

Gorge. Nature 202: 7–10

Lewis OJ (1989) Functional morphology of the

evolving hand and foot. Oxford University

Press, Oxford

Linnaeus C (1758) Systema naturae. Laurentii

Salvii, Stockholm

Mayr E (1944) On the concepts and terminolo-

gy of vertical subspecies and species. Natl Res

Council Bull 262: 11–16

Prat S, Brugal J‐P, Tiercelin J‐J, Barrat J‐A,Boh M, Delagnes A, Harmand S, Kimeu K,

Kibunjia M, Texier P‐J, Roche H (2005) First

occurrence of early Homo in the Nachukui

Formation (West Turkana, Kenya) at 2.3–2.4

Myr. J Hum Evol 49: 230–240

Prentice ML, Denton GH (1988) In: Grine FE

(ed) Evolutionary history of the ‘‘Robust’’

Australopithecines. Aldine de Gruyter, New

York, pp 383–403

Ramirez Rozzi F (1997) Les hominides du Plio‐Pleistocene de la vallee de l’Omo. Cahiers de

paleoanthropologie CNRS editions, Paris

Rayner RJ, Moon BP, Masters JC (1993) The

Makapansgat australopithecine environment.

J Hum Evol 24: 219–231

Roche H, et al. (2003) Les sites arche ologi-

ques pliopleeistocenes de la formation de

Nachukui, Ouest Turkana: bilan synthetique

1997–2001. CR Palevol 2: 663–673

Schoetensack O (1908) Der Unterkiefer des

Homo heidelbergensis aus den Sanden

von Mauer bei Heidelberg. W Engelmann,

Leipzig


Schrenk F, Bromage TG, Betzler C, Ring U,

Juwayeyi Y (1993) Oldest Homo and Plio-

cene biogeography of the Malawi Rift. Nature

365: 833–836

Stern JT, Susman RL (1983) Locomotor anato-

my of Australopithecus afarensis. Am J Phys

Anthropol 60: 279–317

Suwa G, White TD, Howell FC (1996) Mandib-

ular postcanine dentition from the Shungura

Formation, Ethiopia: crown morphology,

taxonomic allocations, and Plio‐Pleistocenehominid evolution. Am J Phys Anthropol

101: 247–282

Tobias PV (1987) The brain of Homo habilis: a

new level of organization in cerebral evolu-

tion. J Hum Evol 16: 741–761

Tobias PV (1991) The skulls, endocasts and

teeth of Homo habilis. In: Olduvai Gorge,

vol 4, 2 vols. Cambridge University Press,

Cambridge

Tobias PV, von Koenigswald GHR (1964) A

comparison between the Olduvai hominines

and those of Java and some implications for

hominid phylogeny. Nature 204: 515–518

Toth N (1985) Archeological evidence for pref-

erential right‐handedness in the lower and

middle Pleistocene, and its possible implica-

tions. J Hum Evol 14: 607–614

Vrba ES (1985) Ecological and adaptive changes

associated with early hominid evolution. In:

Delson E (ed) Ancestors: the hard evidence.

Alan R. Liss, New York, pp 63–71

Vrba ES (1988) Late Pliocene climatic events

and human evolution. In: Grine FE (ed)

Evolutionary history of the ‘‘Robust’’ Austra-

lopithecines. Aldine de Gruyter, New York,

pp 405–426

Vrba ES (1992) Mammals as a key to evolution-

ary theory. J Mamm 73: 1–28

Wood BA (1991) Koobi Fora research project,

vol 4: Hominid Cranial remains. Clarendon

Press, Oxford

Wood BA (1992) Origin and evolution of the

genus Homo. Nature 355: 783–790

Wood BA, Collard M (2001) The meaning of

Homo. Ludus Vitalis vol. IX. 15: 63–74

9 The Earliest Putative Homo Fossils

Documents

Transcript of 9 The Earliest Putative Homo Fossils