B l A C k w E l l
C o m PA n i o n s t o
A n t h R o P o l o g y
ISBN 978-1-4443-3116-5
“This collection of concise reviews of current research by an
All-Star team of authors will be a valuable resource for students
of paleoanthropology at all levels.”
John G. Fleagle, Stony Brook University
“A Companion to Paleoanthropology brings new ideas to light on
human evolution, brilliantly coordinated by David Begun. It
provides an unparalleled account of all that is new in
palaeoanthropology, and it is essential reading for students,
teachers, and researchers alike.”
Peter Andrews, Natural History Museum, London
“This is a wonderful addition to the Companions series: focused but
extensive and inclusive, with authoritative chapters by experts
combining just the right amount of history and cutting-edge
results. Suitable for advanced undergraduate and graduate-level
course readings.”
Eric Delson, City University of New York
The Editor
David R. Begun is Professor in the Department of Anthropology,
University of Toronto. A paleoanthropologist with 30 years of
experience in the analysis of fossil apes, Begun’s current research
focuses on the relationships between European and African fossil
and living great apes and the origin of the African apes and
humans. He has published numerous articles and book chapters on
that topic, and on the description and analysis of fossils from
Europe, Asia, and Africa. A Companion to Paleoanthropology presents
a
compendium of readings representing the state of the art in our
knowledge relating to the study of ancient humans as found in
fossil hominid evidence. Contributions from top scholars in
paleoanthropology and related fields offer accessible overviews of
paleontological methods and research topics for established
researchers, instructors, students, and non-professionals alike.
These original essays define new trends and current interpretations
of the paleoanthropological record. Chapters are organized into
sections that survey the history and techniques of
paleoanthropology, evaluate the fossil evidence and the current
classification of hominids, and review the trends throughout ape
and human evolution in cranial, postcranial, and brain
evolution.
By presenting the latest findings and developments in the field, A
Companion to Paleoanthropology represents an important contribution
to our understanding of the current social discourse on human
origins and evolution.
A Companion to Paleoanthropology
B l A C k w E l l C o m PA n i o n s t o A n t h R o P o l o g y
BlACkwEll
ComPAnions to
PG3628
A Companion to Paleoanthropology
The Blackwell Companions to Anthropology offers a series of
comprehensive syntheses of the traditional subdisciplines, primary
subjects, and geographic areas of inquiry for the field. Taken
together, the series represents both a contemporary survey of
anthro- pology and a cutting edge guide to the emerging research
and intellectual trends in the field as a whole.
1. A Companion to Linguistic Anthropology edited by Alessandro
Duranti 2. A Companion to the Anthropology of Politics edited by
David Nugent and Joan
Vincent 3. A Companion to the Anthropology of American Indians
edited by Thomas Biolsi 4. A Companion to Psychological
Anthropology edited by Conerly Casey and Robert
B. Edgerton 5. A Companion to the Anthropology of Japan edited by
Jennifer Robertson 6. A Companion to Latin American Anthropology
edited by Deborah Poole 7. A Companion to Biological Anthropology,
edited by Clark Larsen (hardback only) 8. A Companion to the
Anthropology of India, edited by Isabelle Clark-Decès 9. A
Companion to Medical Anthropology edited by Merrill Singer and
Pamela I.
Erickson 10. A Companion to Cognitive Anthropology edited by David
B, Kronenfeld, Giovanni
Bennardo, Victor de Munck, and Michael D. Fischer 11. A Companion
to Cultural Resource Management, Edited by Thomas King 12. A
Companion to the Anthropology of Education, Edited by Bradley A.U.
Levinson
and Mica Pollack 13. A Companion to the Anthropology of the Body
and Embodiment, Edited by Frances
E. Mascia-Lees 14. A Companion to Paleopathology, edited by Anne L.
Grauer 15. A Companion to Folklore, edited by Regina F. Bendix and
Galit Hasan-Rokem 16. A Companion to Forensic Anthropology, edited
by Dennis Dirkmaat 17. A Companion to the Anthropology of Europe,
edited by Ullrich Kockel, Máiréad
Nic Craith, and Jonas Frykman 18. A Companion to Border Studies,
edited by Thomas M. Wilson and Hastings
Donnan 19. A Companion to Rock Art, edited by Jo McDonald and Peter
Veth 20. A Companion to Moral Anthropology, edited by Didier Fassin
21. A Companion to Gender Prehistory, edited by Diane Bolger 22. A
Companion to Paleoanthropology, edited by David R. Begun
Forthcoming
A John Wiley & Sons, Ltd., Publication
This edition first published 2013 © 2013 Blackwell Publishing
Ltd
Blackwell Publishing was acquired by John Wiley & Sons in
February 2007. Blackwell’s publishing program has been merged with
Wiley’s global Scientific, Technical, and Medical business to form
Wiley-Blackwell.
Registered Office John Wiley & Sons, Ltd, The Atrium, Southern
Gate, Chichester, West Sussex, PO19 8SQ, UK
Editorial Offices 350 Main Street, Malden, MA 02148-5020, USA 9600
Garsington Road, Oxford, OX4 2DQ, UK The Atrium, Southern Gate,
Chichester, West Sussex, PO19 8SQ, UK
For details of our global editorial offices, for customer services,
and for information about how to apply for permission to reuse the
copyright material in this book please see our website at
www.wiley.com/wiley-blackwell.
The right of David R. Begun to be identified as the author of the
editorial material in this work has been asserted in accordance
with the UK Copyright, Designs and Patents Act 1988.
All rights reserved. No part of this publication may be reproduced,
stored in a retrieval system, or transmitted, in any form or by any
means, electronic, mechanical, photocopying, recording or
otherwise, except as permitted by the UK Copyright, Designs and
Patents Act 1988, without the prior permission of the
publisher.
Wiley also publishes its books in a variety of electronic formats.
Some content that appears in print may not be available in
electronic books.
Designations used by companies to distinguish their products are
often claimed as trademarks. All brand names and product names
used in this book are trade names, service marks, trademarks or
registered trademarks of their respective owners. The publisher is
not associated with any product or vendor mentioned in this book.
This publication is designed to provide accurate and authoritative
information in regard to the subject matter covered. It is sold on
the understanding that the publisher is not engaged in
rendering professional services. If professional advice or other
expert assistance is required, the services of a competent
professional should be sought.
Library of Congress Cataloging-in-Publication Data
A companion to paleoanthropology / edited by David R. Begun. pages
cm Includes bibliographical references and index. ISBN
978-1-4443-3116-5 (hardback) 1. Paleoanthropology. 2. Fossil
hominids. 3. Human evolution. I. Begun, David R., editor of
compilation. GN281.C5845 2013 569.9–dc23
2012036585
A catalogue record for this book is available from the British
Library.
Cover image: Top: © Natalia Lukiyanova / frenta / Shutterstock.
Centre: photo of archaeologist © topal / Shutterstock. Bottom:
The Mauer Mandible, found in 1907 near Heidelberg, Germany. Type
specimen of Homo heidelbergensis. Possibly MIS 15. Heidelberg
University Institute of Geology and Paleontology.
Cover design by Richard Boxall Design Associates.
Set in 10/12.5pt Galliard by SPi Publisher Services, Pondicherry,
India
1 2013
Acknowledgments xxviii
1 The Past, Present and Future of Paleoanthropology 1 David R.
Begun
2 History 17 Matthew R. Goodrum
Part I Background to Paleoanthropology 35
Section 1 Method and Theory 35
3 Human Systematics 37 David S. Strait
4 Experimental Approaches to Musculoskeletal Function in Primates
55 Matthew J. Ravosa, Kimberly A. Congdon, and Rachel A.
Menegaz
5 Multivariate Quantitative Methods in Paleoanthropology 75 Michael
A. Schillaci and Philipp Gunz
6 Growth, Development, and Life History in Hominin Evolution 97 Jay
Kelley and Debra Bolter
Section 2 Anatomical Regions 118
7 Cranial Evolution in the Apes 119 Brian T. Shea
Contents
9 Hominin Diets 165 Peter S. Ungar and Matt Sponheimer
10 Origin and Evolution of Human Postcranial Anatomy 183 Brian G.
Richmond and Kevin G. Hatala
Section 3 Environment and Behavior 203
11 Multiproxy Paleoecology: Reconstructing Evolutionary Context in
Paleoanthropology 204 Kaye E. Reed
12 Reconstructing Social Behavior from Fossil Evidence 226 J.
Michael Plavcan
13 Geochronology 244 Alan L. Deino
14 The Origins and Evolution of Technology 265 Kathy Schick and
Nicholas Toth
Section 4 Genetics and Race 290
15 Genetic Perspectives on Ape and Human Evolution 291 Todd R.
Disotell
16 The Genetics of Morphology 306 Richard J. Sherwood and Dana L.
Duren
17 Paleoanthropology and Race 321 Milford H. Wolpoff and Rachel
Caspari
Part II The Fossil Record 339
Section 5 Paleogene Primates 339
18 Primate Origins 341 Mary T. Silcox
19 Anthropoid Origins 358 K. Christopher Beard
20 Catarrhine Origins 376 Terry Harrison
Section 6 Neogene/Quaternary Hominoids 397
21 The Miocene Hominoid Radiations 398 David R. Begun
contents vii
22 Before Australopithecus: The Earliest Hominins 417 Scott W.
Simpson
23 Australopithecus and Kenyanthropus 434 Ashley S. Hammond and
Carol V. Ward
24 Paranthropus 457 Bernard Wood and Kes Schroer
Section 7 The Age of Homo 479
25 Earliest Homo 480 Friedemann Schrenk
26 Homo erectus and Related Taxa 497 Susan C. Antón
27 The Middle Pleistocene Record: On the Ancestry of Neandertals,
Modern Humans and Others … 517 Jean-Jacques Hublin
28 Neanderthals 538 Katerina Harvati-Papatheodorou
29 Modern Human Origins 557 Mark Collard and Mana Dembo
30 Homo floresiensis 582 William L. Jungers
Index 599
Figure 3.1 Early hominin phylogenetic relationships. (a) A
possible cladogram describing hominin cladistic relationships; (b)
a simplified cladogram suggesting that robust australopiths and
Homo are closely related to each other; (c) a simplified cladogram
suggesting that robust and gracile australopiths have complex and
unresolved relationships, but that they are all more closely
related to Homo than they are to any of the pre- australopiths; (d)
a phyletic tree based on cladistic relationships depicted in (a).
The pre-australopiths give rise to the gracile australopiths, which
in turn give rise to both the robust australopiths (Paranthropus)
and Homo. The approximate time ranges of hominin species are shown
as black bars. Solid lines represent likely ancestor–descendant
relationships. Dashed lines represent possible ancestor–descendant
relationships. Adapted from Figures 6 and 7 in Strait,
2010.
Figure 5.1 Bivariate plot of scores for the first two
principal components (a) and canonical discriminant functions (b).
Filled boxes, H. heidelbergensis; cross, Neanderthals; boxes early
modern humans; filled circles, Upper Paleolithic Europeans; gray
diamonds, recent human populations.
Figure 5.2 Results from the multidimensional scaling (a),
UPGMA (b) and neighbor- joining (c) cluster analyses of Manhattan
distances derived from the principal components scores. The
neighbor-joining tree is rooted with H. heidelbergensis as the
primitive outgroup. Bootstrap support values greater than 60
percent are shown.
Figure 5.3 Principal components (PC) analysis of neurocranial
landmarks and semi- landmarks (illustrated on the Mlade 1 cranium)
in Procrustes shape space. These first two PC axes explain
approximately 65 percent of the total sample variation. There is no
overlap between modern humans and Neanderthals, including specimens
from the Upper Paleolithic (labeled
List of Illustrations
list of illustrations ix
as “fossil modern humans”). For fossil specimens, the nearest
neighbors in shape space are plotted as a black connecting line.
This illustrates that specimens that appear closest to each other
in the projection of the first two PCs need not be closest in all
dimensions of shape space.
Figure 5.4 Virtual reconstruction of the Taung child
(Australopithecus africanus). CT scans of the three parts of the
original fossil (a) are assembled virtu- ally (b). After
mirror-imaging, the missing parts are estimated using a thin-plate
spline interpolation (c–f). Here, landmarks and semiland- marks
measured on a modern human child (c) are used to complete the
missing neurocranial morphology of the partially preserved
fossil.
Figure 7.1 Illustrations of cranial anatomy (frontal,
superior, lateral views), and internal mid-sagittal views. Top to
bottom, the rows depict a male siamang, a male orang utan, a male
chimpanzee and a male gorilla. The images are shown at the same
approximate skull length, and are there- fore obviously not to
scale. See text for discussion.
Source: The line drawings were done by Nicolas Amorosi, an artist
in the anthropology department at the American Museum of Natural
History, in 1982–3. They were explicitly for use in publication.
Two of them (orangutan, chimpanzee) were published in Shea
(1985).
Figure 8.1 Evolution of cranial capacity. (a) Cranial capacity
vs. body size in modern and fossil primates. Data from Table
8.1. Average mammal: cranial capacity (cc) = 0 059(body mass g)0 76
(based on Martin 1981); average primate: cranial capacity (cc) = 0
087(body mass g)0 77 (data from Stephan et al. 1981). (b) Evolution
of primate cranial capacity. Data from Table 8.2. Best-fit
third-order polynomial of anthropoids through Homo sapiens sapiens,
excluding robust australopithecines (Paranthropus aethiopicus, P.
boisei, and P. robustus) and immature specimens: cranial capacity
(cc) = 76 814(log Ma)3 − 53 694(log Ma)2 − 681 44(log Ma) + 863 31
(r2 = 0 90, N =183).
Figure 10.1 Characteristic postcranial skeletal features of
(a) modern apes (from Fleagle 1999), and hypothesized
characteristic features of (b) the hominid (great ape and human)
last common ancestor (LCA) based primarily on evidence from
Eurasian hominid fossils, and (c) the homi- noid LCA (adapted from
Fleagle 1999). Derived features of the LCAs are denoted with
asterisks.
Figure 10.2 Phylogram showing known potential hominoid and
true hominoid (= clade uniting modern hominoids) taxa from
Africa, Europe, and Asia. Taxa that have associated postcranial
evidence are denoted in grey.
Figure 10.3 Characteristic postcranial skeletal features of
Australopithecus (left), Homo erectus (center), and H. sapiens
(right). The features noted on Australopithecus include the
combination of primitive and derived traits characteristic of this
genus; those noted on H. erectus are derived relative to
Australopithecus; and those indicated for H. sapiens are
autapomorphic.
x list of illustrations
Figure 11.1 Discriminant function plot of bovid species from
the Hadar hominin site. Measurements were taken on bovid mandibles
of extant species and com- pared with fossils recovered at Hadar.
Mean ratios were computed for each species and a discriminant
function was performed. The fossil taxa are positioned in quadrants
with living taxa, thus reconstructing their diets.
Figure 11.2 A cluster analysis of taxa from 23 African
localities. These data are derived from Dice similarity indices of
the species at each locality, and show that the South African sites
are quite different from East African ones. In addition, the site
of Makapansgat is not very similar to the later sites in South
Africa (Sterkfontein and Swartkrans). These differences between
South and East Africa are not all due to time, as many are roughly
the same age.
Figure 13.1 The dating range of each of the methods discussed
in this chapter.
Figure 14.1 Oldowan artifact forms from Koobi Fora, Kenya.
(From Schick and Toth 1993; copyright Kathy Schick and Nicholas
Toth.)
Figure 14.2 (a) Early Acheulean handaxes and a cleaver (bottom
right) from Bed II of Olduvai Gorge, about 1.5 million years ago.
(b) Later Acheulean handaxes (top) and cleavers (bottom) from
Kalambo Falls, Zambia, about 400,000 years ago. (From Schick and
Toth 1993; copyright Kathy Schick and Nicholas Toth.)
Figure 14.3 (a) Middle Paleolithic prepared core technologies.
Levallois “tortoise” core and flake (top) and Levallois point core
and flake (bottom). (b) A range of Middle Paleolithic/Middle Stone
Age tool forms. (From Schick and Toth 1993; copyright Kathy Schick
and Nicholas Toth.)
Figure 14.4 (a) Upper Paleolithic blade production by indirect
percussion or punch technique. (b) A range of Upper
Paleolithic/later Stone Age tool forms. (From Schick and Toth 1993;
copyright Kathy Schick and Nicholas Toth.)
Figure 15.1 Schematic representation of the different
populations’ contributions to modern human diversity. Neandertals
are proposed to have contributed approximately 2·5 percent of the
alleles to the modern Eurasian gene pool but not to that of
Africans. Denisovans, while being most closely related to
Neandertals, appear to have contributed to a limited number of
Asian populations.
Figure 16.1 Linear (a) and angular (b) measures taken from
human radiographs. Angle A = Ba–S–N; Angle B = N–S–PNS; Angle C =
S–N–A (Sherwood et al. 2008a).
Figure 16.2 Example of a string plot for the trait Facial
Taper. Chromosomes are identified as numbered straight lines,
curves indicate strength of link- age signal. Statistically
significant linkage (LOD score >3·0) is found on chromosomes 6.
Linkage between a LOD of 1·9 and 3·0 are described as suggestive
and can be seen on chromosomes 1 and 5.
list of illustrations xi
Figure 16.3 Example of a LOD plot showing the detailed linkage
results for Posterior Facial Height on chromosome 6. Maximal LOD =
4·52 (Sherwood et al. 2011).
Figure 18.1 Different approaches to defining taxonomic groups.
Redrawn based on de Queiroz and Gauthier 1990: fig. 1. (a)
Node-based definition: the name refers to all of the descendents of
the common ancestors of Y and Z, but no stem taxa. (b) Stem-based
definition: the name refers not only to the products of the most
recent common ancestor of Y and Z, but also all taxa more closely
related to this common ancestor than to X. (c) Apomorphy-based
definition: the first appearance of a shared, derived feature (=
apomorphy; indicated by the solid black bar) is used to delineate a
common ancestor, and the name applies to all descend- ents of that
ancestor.
Figure 18.2 Possible basal euprimates. Modified from Rose,
1995: box 2. Illustrations by E. Kasmer.Altiatlasius
koulchii: (a) LM1–3 in occlusal view; (b) LP3–M3 in buccal and (c)
occlusal views. P4 and M3 are not currently known (reconstructed).
Altanius orlovi: (d) LP3–M3 in occlusal view; (e) LP3–M3 with
alveoli for mesial teeth in buccal and (f ) occlusal views.
Figure 18.3 Hypothesis of relationships based on Bloch et al.
(2007), with signifi- cant character complexes mapped on. Named
nodes: 1 = euarchonta; 2 = primates; 3 = euprimateformes; 4 =
euprimates.
Figure 19.1 Partial lower dentition of the amphipithecid
primate Ganlea mega- canina from the late Middle Eocene of Myanmar,
based on mirror- image restoration of the holotype right dentary
(Beard et al. 2009). This image was produced by X-ray synchrotron
microtomography on beamline ID19 at the European Synchrotron
Radiation Facility, Grenoble, France. Image provided courtesy of
Dr. Paul Tafforeau. Scale bar equals 1 cm.
Figure 19.2 Alternative phylogenies of early anthropoids. Tree
topology shown on the left follows Beard et al. (2009), while the
tree topology shown on the right derives from the work of Seiffert
et al. (2005). Certain taxa have been eliminated to make both
studies comparable. Living catar- rhines should be positioned as a
sister taxon of Propliopithecidae.
Figure 20.1 Cladogram showing the inferred phylogenetic
relationships between the major groups of catarrhines.
Figure 20.2 Comparison of crania of stem catarrhines. (a–c)
Cranium of Propliopithecus zeuxis (CGM 40237) from the Early
Oligocene of the Fayum, Egypt. (a) right lateral view (with
unassociated mandible); (b) anterior view; (c) ventral view.
Courtesy of and © Eric Delson. (d–f) Partial skull of Pliopithecus
vindobonensis from the Middle Miocene locality of Neudorf-Spalte,
Devínská Nová Ves, Slovakia. (d) right lateral view; (e) frontal
view; (f) palatal view. Courtesy of and © Eric
xii list of illustrations
Delson. (g–h) Partial cranium of Saadanius hijazensis (SGS-UM
2009-002) from the mid-Oligocene locality of Harrat Al Ujayfa,
Saudi Arabia. (g) left lateral view (image reversed); (h) oblique
antero-dorsal view; (i) ventral view. Courtesy of Iyad Zalmout and
William J. Sanders.
Figure 21.1 Selected Miocene apes. (a) Afropithecus; (b)
Proconsul cranium; (c) Proconsul mandible; (d) Çandr mandible
(Griphopithecus); (e) Equatorius; (f) Kenyapithecus; (g)
Nacholapithecus; (h) Samburupithecus. Figure 21.1d previously
published as figure 2b, p. 13 in Begun et al. 2012.
Figure 21.2 (a) Rudapithecus cranium anterior; (b)
Rudapithecus cranium lateral; (c) Oreopithecus; (d) Ouraopithecus;
(e) “Ouranopithecus” from Turkey. Figures 21.2a and b
previously published as part of Figure 6C, p. 15, in Begun et
al. 2012. Figure 21.2c is adapted from Figure 20.10, p.
358, in Begun 2002. Figure 21.2d is adapted from
Figure 20.8, p. 355, in Begun 2002.
Figure 22.1 Shaded relief map of Africa with the locations of
the major Late Miocene–Early Pliocene hominin sites.
Source: US Department of Commerce, National Oceanic and Atmospheric
Administration (NOAA), National Geophysical Data Center (NGDC),
http://www.ngdc.noaa.gov/mgg/topo/img/af.gif.
Figure 23.1 Type specimen of Australopithecus africanus Taung
1. Image is a CT scan reconstruction courtesy of Dr. Philipp Gunz.
Endocast is colored dark gray; most of braincase is missing and
reconstruction indicated by transparent outline.
Figure 23.2 Some well-preserved crania of Australopithecus and
Kenyanthropus illustrating the overall similarities. Top row: Four
Australopithecus afri- canus specimens illustrating diversity
within a single species, from left to right Taung 1, STS 5, STW 71,
STW 505 (reconstructed CT scan images courtesy of Dr. Philipp
Gunz). Bottom row, left to right: pho- tographs of Australopithecus
afarensis A.L. 444-2 (photo courtesy of Dr. William Kimbel),
Australopithecus sediba MH-1 (photo courtesy of Dr. Lee Berger),
and Kenyanthropus platyops KNM-WT 40000 (photo copyright National
Museums of Kenya, courtesy of Dr. Meave Leakey, with photo credit
to Bob Campbell). Note in all Australopithecus spec- imens the
relatively small brain compared with modern humans and moderate
degree of facial prognathism, and the lack of enlarged canine
teeth. Kenyanthropus has deeper zygomatics (cheekbones) placed fur-
ther forward in the skull than the other specimens.
Figure 24.1 Locations of sites where evidence of Paranthropus
has been confirmed.
Figure 24.2 Significant events in the discovery and analysis
of the fossil record of Paranthropus.
Figure 24.3 Comparison of Paranthropus crania. Left lateral
views of the well- preserved cranium of Paranthropus aethiopicus,
the holotype of
list of illustrations xiii
Paranthropus boisei, and a representative cranium of Paranthropus
robustus. Images of KNM-WT 17000 and SK 48 are taken from the
NMNH-SI Human Origins website. Not to scale.
Figure 24.4 Phylogenetic hypotheses concerning Paranthropus
taxa. Hypotheses about the relationships among Paranthropus
considered in this review; (a) Paranthropus monophyly, (b)
Paranthropus polyphyly. Redrawn after Grine (1997).
Figure 25.1 African early hominin sites. Homo rudolfensis and
Homo habilis sites in bold.
Figure 25.2 UR 501 from the Chiwondo Beds, northern Malawi
(about 2·5–2·4 Ma), Homo rudolfensis (Drawing: Claudia
Schnubel).
Figure 25.3 KNM-ER 1470 from Koobi Fora, Kenya (about 1·9
Ma.), Homo rudolfensis. KNM-ER 1813 from Koobi Fora, Kenya (about
1·9 Ma.), Homo habilis (Drawing: Claudia Schnubel).
Figure 26.1 The temporal and geographic distribution of H.
erectus localities and some important specimens discussed in the
text. On the far left is the geomagnetic polarity timescale, with
normal periods in black and reversed in white. Radiometric time is
indicated in millions of years on the far right. Within regional
columns: solid lines on either side of site names indicate time
spans suggested by multiple H. erectus individuals from a site;
dashed lines indicate possible time range around a single/ few
specimens. In the Africa column: sites are grouped from left to
right as South Africa, Kenya, Tanzania, Ethiopia; OG is
Olorgesailie, OH is Olduvai Hominid, numbers on the Koobi Fora line
are KNM-ER numbers, WT15k refers to KNM-WT 15000. In the Java
column: the Sangiran box delineates the timing of the hominin
bearing Sangiran and Bapang formations; however, hominins are
present only from about 1·6 Ma, as indicated by S4, 27, 31 (which
refer to Sangiran specimen numbers), younger overlying sediments
are not (currently) hominin-bearing and are not shown. Gray boxes
indicate a period of temporal and geographic overlap with other
species. The overlapping species are: In Africa, H. habilis; In
China, archaic H. sapiens or H. heidelbergensis; In Java, H.
sapiens.
Figure 26.2 Homo erectus and alternative related taxa that
have been named to accommodate these fossil specimens. The type
specimen for each taxon is listed in parentheses next to the first
use of the species name in the table.
*Only rarely have Indonesian and Chinese remains been separated at
the species level (or into multiple species within regions) since
their synonimization into H.erectus in the 1940s (Mayr 1944).
**H. ergaster is the most common designation for this group of
African and Georgian remains; however, all the other African
“species” were named before H. ergaster and by the laws of
zoological nomenclature the oldest should be the name-bearer.
However, the type for H. ergaster
xiv list of illustrations
is always included in this grouping, whereas OH 9 (the type for H.
leakeyi; Heberer 1963) is included by some in H. erectus s.s., and
the Ternifine mandibles (At. mauritanicus; Arambourg 1954) are
often excluded from the grouping and placed in archaic H. sapiens,
and the Swartkrans material assigned to T. capensis (Robinson
1953a, b) is also often excluded from this group entirely.
*** See previous footnote for different taxonomic placement of
these type specimens.
Figure 26.3 Cranial features of H. erectus illustrated on the
KNM-ER 3733 face, the posterior vault of Sangiran 4, the superior
supraorbital region of OH 9, and a close-up of the mastoid region
of Sangiran 4.
Figure 26.4 Cranial comparisons of presumed male crania from
different regions. From left to right bottom row: Africa (Olduvai
hominid 9; cranial capacity 1067 cc), Indonesia (Sangiran 4 cast;
904 cc) and the Republic of Georgia (Dmanisi 2280; 775 cc). Top
row: superior view of OH 9 and D2280 supraorbital regions. Note the
similarity in posterior vault form and position and development of
occipital superstructures. Note the similarity in form but size
difference in the supraorbital regions. Sangiran 4 and D2280 were
photographed together. The OH 9 image was matched to the scale (not
shown) in the Sangiran 4 and D2280 image. Note that the three
posterior views are registered on the approx- imate position of the
mastoid crest. Copyright Susan Antón 2012.
Figure 27.1 The Mauer Mandible, found in 1907 near Heidelberg,
Germany. Type specimen of Homo heidelbergensis. Possibly MIS
15.
Source: Heidelberg University Institute of Geology and
Paleontology.
Figure 27.2 Lateral view of the rear skull from Swanscombe
(UK). An example of early Neandertal morphology from MIS 11 in
Acheulian context. © 2012 Natural History Museum,
London.
Figure 27.3 Lateral view of the Kabwe Skull (Zambia), type
specimen of Homo rho- desiensis. © 2012 Natural History Museum,
London.
Figure 27.4 Hypothetical phylogenetic relationships between
the Middle and Late Pleistocene hominins. The Marine 18O isotope
record is indicated on the right of the tree. Horizontal dashes
indicate the limits of the Middle and Late Pleistocene. Image
created in the Department of Human Evolution, MPI-EVA. © 2012
Jean-Jacques Hublin.
Figure 27.5 Map of Europe during the last glacial maximum.
South of the ice cap, the permafrost area is dotted. The extension
of two major Middle Pleistocene glaciations (MIS 16 and MIS 6) are
indicated by dashed lines. Image created in the Department of Human
Evolution, MPI- EVA. © 2012 Jean-Jacques Hublin.
Figure 28.1 Map of the geographic distribution of
Neanderthals, showing important Neanderthal and pre-Neanderthal
sites. Adapted from Harvati 2007.
list of illustrations xv
Figure 28.2 Complete Neanderthal skeleton (left) reconstructed
using elements from five partial skeletons (principally La
Ferrassie 1 and Kebara 2) compared with a modern human skeleton
(right). Courtesy of and copyright Ian Tattersall, American Museum
of Natural History.
Figure 29.1 Lateral view of Omo Kibish 1 skull. Drawing © Matt
Cartmill, used with permission from The Human Lineage, by Matt
Cartmill and Fred H. Smith (2009).
Figure 29.2 (a) Lateral and (b) occipital views of Herto 1
cranium. Drawing © Matt Cartmill, used with permission from The
Human Lineage, by Matt Cartmill and Fred H. Smith (2009).
Figure 29.3 Lateral view of Skhul 5 skull. Drawing © Matt
Cartmill, used with permission from The Human Lineage, by Matt
Cartmill and Fred H. Smith (2009).
Figure 29.4 Lateral view of Qafzeh 9 skull. Drawing © Matt
Cartmill, used with permission from The Human Lineage, by Matt
Cartmill and Fred H. Smith (2009).
Figure 29.5 Lateral view of Mlade 5 cranium. Drawing © Matt
Cartmill, used with permission from The Human Lineage, by Matt
Cartmill and Fred H. Smith (2009).
Figure 30.1 (a) Scatterplot of stature versus body mass in
Asian and African human pygmies and Homo floresiensis. (b)
Scatterplot of the ponderal index versus the body mass index in
same. Ellipses represent 95 percent of the data points in each
human sample. Although the body mass of LB1 can be matched among
human pygmies, adult stature cannot, and this implies a very stocky
build in Homo floresiensis.
Figure 30.2 (a) Lateral views of casts of fossil crania, from
left to right – Homo habilis from East Africa, Homo erectus from
Dmanisi, and the cranium of LB1, the type specimen of Homo
floresiensis. Photo courtesy of Chris Stringer. (b) The mandibles
of LB6 (left) and LB1(right) resting on the hands of WLJ. LB6 was
damaged at Gadjah Mada University in 2005 and is permanently
distorted. Photo courtesy of Djuna Ivereigh. (c) Lateral view of
the brain endocasts of LB1 (left) and Homo erectus (right),
courtesy of the Mallinckrodt Institute of Radiology, Washington
University School of Medicine. Despite the obvious size difference,
their overall shapes are quite similar and neither resemble
microcephalic humans.
Figure 30.3 The assembled skeleton of LB1, the type specimen
of Homo floresiensis (photo by W. L Jungers).
List of Tables
Table 3.1 The classification of humans. Humans and their close
extinct ancestors and relatives are found within the subtribe
Hominina, and are marked in bold text.
Table 4.1 Jaw-loading regimes in anthropoids.
Table 4.2 Mandibular and limb peak shear strains (γmax) in
primates and other tetra- pods during powerful masticatory and
locomotor behaviors.
Table 4.3 EMG analyses of primate appendicular muscles.
Table 4.4 Kinematic and kinetic analyses of primate limb
elements.
Table 5.1 Pooled-sex sample size information for populations
and fossil hominid groupings.
Table 5.2 Eigenvector loadings and eigenvalues from the
principal components analysis.
Table 5.3 Eigenvector loadings and eigenvalues from the
canonical discriminant analysis.
Table 5.4 Matrix describing phenotypic distances among groups
calculated from the means of the first three principal
components.
Table 6.1 Life-history stages of wild chimpanzees.
Table 6.2 Life-history stages of humans.
Table 6.3 Ages or durations of key life-history attributes in
great apes and humans (in years and as a percentage of ages or
durations for Homo).
Table 6.4 Select immature specimens of Plio-Pleistocene
hominins (note: not intended as a comprehensive inventory).
list of tables xvii
Table 6.6 Hypothesized life-history stages of Homo
ergaster/early Homo erectus.
Table 7.1 A list of selected features that have been proposed
as derived cranial nov- elties in the great ape skull.
Table 8.1 Estimated cranial capacities and body weights for
primate fossils.
Table 8.2 Estimated cranial capacities and body weights for
extant primate species.
Table 11.1 Definitions of terms in paleoecology.
Table 11.2 Habitat reconstructions for some Plio-Pleistocene
hominin sites.
Table 13.1 A summary of each of the geochronological methods
discussed in Chapter 13.
Table 14.1 Major stages of the Paleolithic.
Table 16.1 Heritability estimates (h2) and standard errors for
craniofacial traits. Significant covariates are indicated (Sherwood
et al. 2008a).
Table 20.1 Family-group classification of the Catarrhini
(after Harrison and Gu 1999; Harrison 2002, 2005, 2010; Andrews and
Harrison 2005).
Table 20.2 Classification of the Propliopithecoidea from the
Oligocene of Afro- Arabia.
Table 20.3 Classification of the Pliopithecoidea from the
Miocene of Eurasia (updated from Harrison and Gu 1999).
Table 20.4 Taxonomy of Saadanioidea, Dendropithecoidea and
early catarrhines of uncertain affinity from the Oligocene and
Miocene of Afro-Arabia (after Harrison 2002, 2010; Pickford et al.
2010; Zalmout et al. 2010).
Table 21.1 A list of the taxa included in Chapter 21.
Table 21.2 A classification of hominoids described in Chapter
21.
Table 23.1 Key modern human fossil-bearing sites discussed in
Chapter 23.
Table 24.1 Site, age and nature of evidence of
Paranthropus.
Table 25.1 Sgnificant morphological differences between H.
habilis and H. rudolfensis (after Wood 1992).
Table 25.2 Fossil remains of Homo habilis sensu stricto.
Table 25.3 Fossil remains of Homo rudolfensis.
Table 26.1 Selected dimensions by geographic region in H.
erectus individuals and isolated elements. (Following Ruff and
Walker (1993) or Graves et al. (2010).)
Table 28.1 Some proposed derived Neanderthal features.
Table 29.1 Key modern human fossil-bearing sites discussed in
Chapter 29.
Notes on Contributors
Susan C. Antón is a Professor in the Center for the Study of Human
Origins, Department of Anthropology, New York University. Her
research concerns the origin and evolution of genus Homo in
Indonesia and Africa and human impact on island ecosystems in the
South Pacific. She recently co-edited with Leslie Aiello Human
Biology and the Origin of Homo (2011), a special issue of Current
Anthropology. She is co-author with Craig Stanford and John Allen
of Introduction to Biological Anthropology: A Natural History of
Humankind 2012). She is co-founder of the Bones and Behavior
Working group (bonesandbehavior.org) and member of the Koobi Fora
Research Project (http://www.kfrp.com/).
K. Christopher Beard is curator of vertebrate paleontology at the
Carnegie Museum of Natural History (Pittsburgh, PA). A specialist
on early primates and the origin of anthropoids, Beard is the
author of the award-winning book The Hunt for the Dawn Monkey:
Unearthing the Origins of Monkeys, Apes and Humans (University of
California Press, 2004). Among his most notable scientific papers
are: K. C. Beard, et al. (1996) “Earliest complete dentition of an
anthropoid primate from the late middle Eocene of Shanxi Province,
China” Science, 272:82–85; K. C. Beard. (2008) “The oldest North
American primate and mammalian biogeography during the
Paleocene-Eocene Thermal Maximum” Proceedings of the National
Academy of Sciences of the USA 105:3815–3818; K. C. Beard, et al.
(2009) “A new primate from the Eocene Pondaung Formation of Myanmar
and the monophyly of Burmese amphipithecids” Proceedings of the
Royal Society B, 276:3285–3294; and J.-J. Jaeger, K. C. Beard, et
al. (2010) “Late middle Eocene epoch of Libya yields earliest known
radiation of African anthropoids” Nature, 467:1095–1098.
David R. Begun is a professor in the Department of Anthropology,
University of Toronto. He has directed or co-directed field
research at Miocene fossil ape sites in Spain and Hungary and is
mainly interested in the origins and evolution of the great ape and
human clade. Begun works on issues of hominoid phylogeny,
functional
notes on contributors xix
anatomy and paleobiogeography. His most recent scientific papers
include: D. R. Begun et al. (2012) “European Miocene Hominids and
the Origin of the African Ape and Human Clade” Evolutionary
Anthropology 21:10–23; D. R. Begun and T. L. Kivell (2011)
“Knuckle-walking in Sivapithecus: the combined effects of homology
and homoplasy and implications for the origin of human bipedalism.”
Journal of Human Evolution 60:158-170; D. R. Begun (2010) “Miocene
hominids” Annual Review of Anthropology 39:67–84; and D. R. Begun
(2010) “Catarrhine cousins: the origin and evolution of monkeys and
apes of the Old World” in C. S. Clark (ed.), A Companion to
Biological Anthropology. pp. 295–313. Wiley-Blackwell.
Debra Bolter is an anthropology professor at Modesto College in
Northern California. Research on primate life history, growth and
development focuses on the integration of information from multiple
systems: cranial, dental, skeletal, and from soft tissue, with
implications for hominin evolution. She has published growth and
development data on vervet monkeys (Journal of Zoology, 2003),
colobines (Anatomy Research International, 2011), chimpanzees
(Proceedings of the National Academy of Sciences of the USA 2004;
Journal of Zoology, 2007; American Journal of Physical
Anthropology, 2011 and 2012), and an overview article (“Primate
growth and development” in Primates in Perspective (2011), Oxford
University Press) that lays out her approach to primate growth and
development in an evolutionary context.
Rachel Caspari, Professor of Anthropology at Central Michigan
University, is a paleoanthropologist with a long-standing interest
in the relationship between sci- ence and the race concept. Recent
publications include: “The evolution of grandpar- ents” Scientific
American 305(2):44–49 (2011); “Older age becomes common late in
human evolution”(with S. H. Lee) Proceedings of the National
Academy of Sciences of the USA 101:10895–10900 (2004);
“Deconstructing race: race, racial thinking and geographic
variation” in C. Larsen (ed.) Companion to Biological Anthropology
pp.104–122 (2010) Wiley-Liss; and “1918: Three perspectives on race
and human variation” American Journal of Physical Anthropology
139:5–15 (2009).
Mark Collard is a Professor and Canada Research Chair in the
Department of Archaeology at Simon Fraser University in British
Columbia, Canada. He is also the director of SFU’s Human
Evolutionary Studies Program (http://hesp.irmacs.sfu. ca/).
Professor Collard works on a number of topics in evolutionary
anthropology, including the identification of species in the
hominin fossil record, the reconstruction of fossil hominin and
nonhuman primate phylogenetic relationships, and the estima- tion
of body mass, stature and age from skeletal material. In addition,
he is using methods and theory from evolutionary biology to
investigate archeologically- and ethnographically-documented
patterns of material culture variation.
Kimberly A. Congdon is a PhD student in the Integrative Anatomy
program at the University of Missouri. Her research interests
include the plasticity, development and functional morphology of
hands and feet in arboreal primates, particularly as such topics
relate to climbing and grasping. She is also interested in applying
in-vivo exper- imental methods to questions of locomotor evolution
in primates. Her recent work has examined the correlation of pedal
phalangeal curvature with frequency of arboreal
xx notes on contributors
activity in extant apes, the role of mechanical loading on joint
development, and a novel 3D analysis of hominid metatarsals. Recent
publications include “Interspecific and ontogenetic variation in
great ape pedal phalangeal curvature” International Journal of
Primatology, 33:418–427 (2012); K. A. Congdon et al. (2012)
“Differential limb loading in miniature pigs (Susscrofa
domesticus): A test of chondral modeling theory” Journal of
Experimental Biology, 215:1472–1483; and K. A. Congdon et al.
(2011) “3D analysis of the first complete fourth metatarsal of
Australopithecus afarensis from Hadar, Ethiopia” American Journal
of Physical Anthropology Supplement (abstract) 144:111.
Alan L. Deino graduated from U.C. Berkeley in 1985, where he became
a founding member of the Berkeley Geochronology Center, a nonprofit
institution dedicated to the calibration of the timing of events in
Earth’s history. While most of his research has focused on dating
important milestones in human and faunal evolution in East Africa,
recent work has concentrated on tracking variations in
paleoclimatic induced by oscillations in the Earth’s orbit, as
revealed in cyclical deposits in the rock record.
Mana Dembo is a PhD candidate in the Department of Archeology at
Simon Fraser University in British Columbia, Canada. She is also a
member of SFU’s Human Evolutionary Studies Program
(http://hesp.irmacs.sfu.ca/). Her PhD research focuses on hominin
systematics. She is attempting to improve the reliability of the
methods paleoanthropologists use to reconstruct the evolutionary
relationships of the fossil hominins.
Todd R. Disotell is based at the Center for the Study of Human
Origins, Department of Anthropology, New York University. His
research interests are centered upon the theme of primate and human
evolution, at all levels from the population to the supra- ordinal.
His most recent scientific papers include: J. A. Hodgson and T. R.
Disotell “Anthropological genetics: inferring the history of our
species through the analysis of DNA” Evolution: Education and
Outreach 3:387–398 (2010).
Dana L. Duren is an Associate Professor in the Division of
Morphological Sciences and Biostatistics, Departments of Community
Health and Orthopaedic Surgery, Wright State University Boonshoft
School of Medicine, Dayton, Ohio. Her primary research focuses on
the genetic and environmental influences on bone and joint anat-
omy and function across the life span. Recent publications include
quantitative genet- ics and linkage analysis of childhood bone
mass, and heritability of joint cartilage thickness, a precursor to
osteoarthritis.
Matthew R. Goodrum is a professor of history of science in the
Department of Science and Technology in Society at Virginia Tech.
His research focuses on the history of paleoanthropology and the
history of prehistoric archeology and their rela- tionship with
geology, paleontology, and biology. In addition to many published
arti- cles he was the Subject Area Editor responsible for
paleoanthropology and physical anthropology for the New Dictionary
of Scientific Biography (2008). More recently he was an associate
editor and a major contributing author responsible for historical
and biographical entries for the Wiley-Blackwell Encyclopedia of
Human Evolution (2011).
notes on contributors xxi
Philipp Gunz is a physical anthropologist with a primary research
interest in paleoan- thropology. He obtained his PhD in
anthropology from the University of Vienna in 2005 and currently
works as a Research Fellow at the Max Planck Institute for
Evolutionary Anthropology in Leipzig, Germany. He studies
developmental and evolutionary changes in the growth patterns and
morphology of fossil hominins, extant humans and non-human
primates. Philipp works both in the area of “virtual
paleoanthropology,” where he applies computer modeling techniques
to reconstruct partial and damaged fossils for further analysis,
and in the application of statistical methods to analyze shapes of
fossil and living primates (geometric morphometrics). His most
recent scientific papers include: P. Gunz et al. (2012) “The
mammalian bony labyrinth reconsidered, introducing a new geometric
morphometric approach” in the Journal of Anatomy; Gunz P. Gunz et
al. (2010) “Brain development after birth differs between
Neanderthals and modern humans” Current Biology 20 (21):R921– 922;
and P. Gunz et al. (2009) “Principles for the virtual
reconstruction of hominin crania” Journal of Human Evolution 7
(1):48–62.
Ashley S. Hammond is a Life Sciences Fellow and PhD Candidate in
Integrative Anatomy (Department of Pathology and Anatomical
Sciences) as part of the Pathobiology Area Program at the
University of Missouri. She has published research evaluating the
effects of exercise on joint biology in mammals and finite element
evaluations of cranial morphology during growth in apes and humans.
In 2012, she was the recipient of the Ernest A. Hooten Prize from
the American Association of Physical Anthropologists for her
research using novel 3D methods to quantify hip joint size and
shape in fossil hominins.
Terry Harrison is Professor of Anthropology and Director of the
Center for the Study of Human Origins at New York University. He
obtained his PhD in Biological Anthropology from the University of
London. His research focuses on the phyloge- netic relationships
and paleobiology of fossil catarrhines, including the earliest
homi- nins. He has conducted paleontological fieldwork in Europe,
East Africa and Asia, and he is currently co-director of
paleoanthropological research at the early hominin locality of
Laetoli, Tanzania. He is Editor of the recently published
two-volume series on Paleontology and Geology of Laetoli: Human
Evolution in Context (Springer, Dordrecht).
Katerina Harvati-Papatheodorou is the Director of Paleoanthropology
at the Eberhard-Karls-Universität Tübingen and the Senckenberg
Center for Human Evolution and Paleoecology. She is also adjunct
Professor of Anthropology at the City University of New York
Graduate Center. After obtaining her PhD at the University of New
York in 2001, Katerina Harvati worked as an Assistant Professor at
New York University. From 2004 she was Senior Researcher at the
Max-Planck-Institute for Evolutionary Anthropology in Leipzig.
Professor Harvati’s research specializes in Neanderthal evolution,
modern human origins and the application of 3-D geometric
morphometric methods to paleoanthropology. Her general research
interests include primate and human evolution; evolutionary theory;
evolution of primate and human life-history; the relationship of
morphological variability to population history and the
environment; and Paleolithic archeology. She has conducted
fieldwork in Europe
xxii notes on contributors
and Africa, and recently directed paleoanthropological fieldwork in
Greece and Tanzania. Her research was named one of the top 10
scientific discoveries of the year 2007 by TIME magazine for
demonstrating the African origin of all modern humans. In 2010 she
was elected Fellow of the American Association for the Advancement
of Science for her contributions to Paleoanthropology. Among her
recent publications are: K. Harvati, and T. Harrison, (2006)
Neanderthals Revisited: New Approaches and Perspectives. Springer;
with S. Benazzi et al. (2011) “Early dispersal of modern humans in
Europe and implications for Neanderthal behavior” Nature
479:525–528; K. Harvati et al. (2011) “Morphologie und
Chronologie der Schädelkalotte aus Iwo Eleru (Nigeria, Later Stone
Age)”. PLoS ONE 6(9):e24024; K. Harvati et al. (2010) “Evolution of
middle-late Pleistocene human cranio-facial form: A 3-D approach”
Journal of Human Evolution 59:445–464.
Kevin G. Hatala is a doctoral candidate in the Hominid Paleobiology
Doctoral Program at the Center for the Advanced Study of Hominid
Paleobiology at the George Washington University. A 2009 graduate
of Duke University, his research interests are in postcranial
functional morphology and the evolution of human gait.
Jean-Jacques Hublin started his career at the French CNRS, before
being hired as Professor of Anthropology at the University of
Bordeaux. He is now a Professor and Director at the Max Planck
Institute for Evolutionary Anthropology in Leipzig (Germany), where
he created the Department of Human Evolution in 2004. His research
mainly focuses on virtual paleoanthopology, the processes
associated with the emergence of Neandertals and modern humans, and
on the interactions between the two groups in Europe. His latest
book, co-edited with S. P. McPherron, is entitled Modern Origins: A
North African Perspective, Springer 2012.
William L. Jungers is Distinguished Teaching Professor and Chairman
in the Department of Anatomical Sciences at the School of Medicine,
Stony Brook University. His research interests include human
evolution, paleontology, and quanti- tative methods. His
publications include the following: B. G. Richmond and W. L.
Jungers (2008) “Orrorin tugenensis femoral morphology and the
evolution of homi- nin bipedalism” Science 319:1662–1665; W. L.
Jungers et al. (2009) “The foot of Homo floresiensis” Nature
459:81–84; and (2009) Paleoanthropological Research at Liang Bua,
Indonesia (M. Morwood and W. L. Jungers, eds.) Journal of Human
Evolution Special Issue 57:437–650.
Jay Kelley is a research affiliate at Arizona State University’s
Institute of Human Origins and associate professor at the
University of Illinois, Chicago. His most recent publication is:
Jay Kelley and Feng Gao (2012) “Juvenile hominoid cranium from the
late Miocene of southern China and hominoid diversity in Asia”
Proceedings of the National Academy of Sciences of the USA 109(18)
6882–6885.
Rachel A. Menegaz is a doctoral candidate in the Integrative
Anatomy graduate pro- gram at the University of Missouri. Her
research investigates the evolutionary and developmental
interactions between muscle and skeletal tissues in the mammalian
craniofacial complex, incorporating micro- and macro-anatomical
techniques to
notes on contributors xxiii
elucidate the relationship between morphology and environment. She
is a National Science Foundation Graduate Research fellow and
University of Missouri Life Sciences fellow, and has been
recognized by the American Association of Physical Anthropologists
(2008) and the American Society for Bone and Mineral Research
(2012) for her work in bone biology. Recent publications include:
R. A. Menegaz et al. (2010) “Evidence for the influence of diet on
cranial form and robusticity” Anatomical Record 293A:630–641; R. A.
Menegaz and E. C. Kirk (2009) “Septa and processes: convergent
evolution of the orbit in haplorhine primates and strigiform birds”
Journal of Human Evolution 59:672–687; R. A. Menegaz et al. (2009)
“Phenotypic plasticity and function of the hard palate in growing
rabbits” Anatomical Record 292A:277–284.
J. Michael Plavcan is Professor of Anthropology at the University
of Arkansas, Fayetteville, and a AAAS fellow. He has published
extensively on dental, cranial, and postcranial sexual dimorphism
in primate and human evolution, and has carried out field work in
North and South America and Africa. Recent publications include: J.
M. Plavcan (2012) “Sexual size dimorphism, canine dimorphism, and
male–male compe- tition in primates: where do humans fit in?” Human
Nature 23:45–67; (2011) “Understanding dimorphism as a function of
changes in male and female traits” Evolutionary Anthropology
20:143–155.
Matthew J. Ravosa is Professor of Biological Sciences, Aerospace
and Mechanical Engineering, and Anthropology at the University of
Notre Dame, and a Research Associate in Zoology at the Field
Museum. His research investigates major trans- formations in the
mammalian musculoskeletal system during development and across
higher-level clades, integrating diverse approaches to increase our
under- standing of the evolutionary and pathobiological
significance of anatomical, func- tional, behavioral and ecological
patterns. In addition to being a lifetime member of the Society for
Integrative and Comparative Biology and American Association of
Physical Anthropologists, he is a fellow of the American
Association for the Advancement of Science. Recent publications
include: J. E. Scott, J. B. Lack, and M. J. Ravosa (2012) “On the
irreversibility of mandibular symphyseal fusion” Evolution 66, in
press; M. J. Ravosa et al. (2010) “Allometry of masticatory loading
parameters in mammals” Anatomical Record 293A:557–571; E. Jašarevi
et al. and M. J. Ravosa (2010) “Masticatory loading, function and
plasticity: A microana- tomical analysis of mammalian circumorbital
soft-tissue structures” Anatomical Record 293A:642–650.
Kaye E. Reed is Professor in the Institute of Human Origins, School
of Human Evolution and Social Change, Arizona State University,
Tempe. Among her recent publications are: D. Geraads, R. Bobe,and
K. E. Reed (2012) “Pliocene Bovidae (Mammalia) from the Hadar
Formation of Hadar and Ledi-Geraru, Lower Awash, Ethiopia” Journal
of Vertebrate Paleontology 32:180–197; K. E. Reed, and F. Bibi
(2011) “Fossil Tragelaphini (Artiodactyla: Bovidae) from the late
Pliocene Hadar Formation, Afar Regional State, Ethiopia”
Journal of Mammalian Evolution 18:57–69; and K. E. Reed, and S. M.
Russak (2009) “Tracking ecological change in relation to the
emergence of Homo at the Plio-Pleistocene boundary,” in F.
Grine,
xxiv notes on contributors
and R. E. Leakey (eds.) The First Humans –Origin and Early
Evolution of the Genus Homo. SpringerLink Series in Vertebrate
Paleobiology and Anthropology.
Brian G. Richmond is an Associate Professor in the Department of
Anthropology at The George Washington University. He received his
PhD from Stony Brook University in 1998. His research focuses on
the origin and evolution of human gait and the functional anatomy
of the hand. At The George Washington University he teaches
undergraduate and graduate courses in human evolution, functional
anatomy, human anatomy, and analytical member. He is a faculty
member at the Center for the Advanced Study of Hominid Paleobiology
and the Graduate Advisor for the Hominid Paleobiology Program.
Recent publications include: D. J. Green, M. W. Hamrick, and B. G.
Richmond (2011) “The effects of hypermuscularity on shoulder
morphol- ogy in myostatin-deficient mice” Journal of Anatomy
218:544–557; and J. Chalk, R. G. Richmond et al. (2011) “A
finite element analysis of masticatory stress hypoth- eses”
American Journal of Physical Anthropology 145:1–10.
Kathy Schick is Professor of Anthropology at Indiana University
Bloomington and co-director at The Stone Age Institute. She
received her PhD in Anthropology from the University of California,
Berkeley. Her topical interests include paleoanthropol- ogy and
African prehistory, and her geographical areas of specialization
are Africa and China. She was elected as a Fellow of the American
Association for the Advancement of Science in 2004 and received the
Distinguished Faculty Research Award from Indiana University in
1997.
Michael A. Schillaci received his doctorate in anthropology from
the University of New Mexico in 2002. He is currently an Associate
Professor in the Department of Anthropology at the University of
Toronto Scarborough. Dr. Schillaci’s diverse research interests
include human and nonhuman primate evolution. His recent publi-
cations include “Latitudinal variation in cranial dimorphism in
Macaca fascicularis” American Journal of Primatology 72(2):152–160,
and “Estimating the probability that the sample mean is within a
desired fraction of the standard deviation of the true mean”
Journal of Human Evolution 56(2):134–138.
P. Thomas Schoenemann is an Associate Professor in Anthropology at
Indiana University, a Research Scientist at the Stone Age
Institute, and a member of the Cognitive Science Program at Indiana
University. His research interests are on the evolution of brain
and behavior, with a special focus on language. Previous publica-
tions include: “Evolution of the size and functional areas of the
human brain” (2006) Annual Review of Anthropology, v. 35:379–406,
and “Evolution of brain and language” (2012) in M. A. Hofman and D.
Falk (eds.), Progress in Brain Research, Vol. 195, pp.
443–459.
Kes Schroer is a doctoral candidate, Center for the Advanced Study
of Hominid Paleobiology, the George Washington University. Her
research interest is premolar variation, particularly in
Paranthropus. She is using an extant primate model to research the
morphological integration and developmental processes at work in
the molariza- tion of the fourth premolar. Recent publications
include: M. M. Skinner, K. E. Schroer
notes on contributors xxv
et al. (2011) “Mandibular P4 morphology among Plio-Pleistocene
hominins: taxonomic implications and morphological trends” American
Journal of Physical Anthropology Suppl. 52 276; K. E. Schroer et
al. (2010) “How long were australopith- ecine toes?” American
Journal of Physical Anthropology Suppl. 50 209; and K. E. Schroer
(2009) “God and the Stegosaurus: presentations of creationism and
evolution in American museums” American Journal of Physical
Anthropology Suppl. 48 330.
Friedemann Schrenk is Section Head of the Department of
Paleoanthropology at the Senckenberg Museum in Frankfurt, Germany.
He is Professor of Paleoanthropology at the Johann Wolfgang Goethe
University.
Brian T. Shea is a Professor in Cell and Molecular Biology at
Northwestern University Feinberg School of Medicine. He received
his PhD in bioanthropology from Duke University and completed a
postdoctoral fellowship at the American Museum of Natural History.
He recently co-authored “Growth hormone binding protein, insu-
lin-like growth factor-I and short stature in two pygmy populations
from the Philippines” with N. Dávila et al. Journal of Pediatric
Endocrinology & Metabolism 2002 Mar; 15(3):269–76.
Richard J. Sherwood is a Professor in the Departments of Community
Health and Pediatrics, and the Director of the Division of
Morphological Sciences and Biostatistics, Boonshoft School of
Medicine, Wright State University, Dayton, Ohio. His training is in
comparative anatomy and he has spent most of his career
investigating the myriad forces influencing variation in the
craniofacial complex of modern humans as well as extinct and extant
nonhuman primates. His most recent work has focused on the genetic
underpinnings of craniofacial morphology. His publications range
from descriptions of fossil primates to genome-wide linkage scans
for QTL influencing craniofacial morphology.
http://www.wright.edu/~richard.sherwood/.
Mary T. Silcox is Associate Professor of Anthropology at the
University of Winnipeg. Her research focuses on understanding the
earliest events in the evolution of the Order Primates, using the
fossil record. She focuses especially on the evolution, anat- omy,
and ecology of plesiadapiforms. Recent publications include M.T.
Silcox, C.K. Dalmyn, and J.I. Bloch. “Virtual endocast of Ignacius
graybullianus (Paromomyidae, Primates) and brain evolution in early
primates” Proceedings of the National Academy of Sciences of the
USA 106:10987–92; and M.T. Silcox, J.I. Bloch, D.M. Boyer, M.
Godinot, T.M. Ryan, F. Spoor, and A. Walker “Semicircular canal
system in early primates” Journal of Human Evolution
56:315–327.
Scott W. Simpson is Professor of Anatomy at Case Western Reserve
University School of Medicine, Cleveland, Ohio, USA and at the
Laboratory of Physical Anthropology, Cleveland Museum of Natural
History. Recent publications include: C. O. Lovejoy,S. W. Simpson
et al. (2009) “Careful climbing in the Miocene: The forelimbs of
Ardipithecus ramidus and humans are primitive.” Science 326:73,
100–106; with G. Suwa et al. (2009) “Paleobiological implications
of the Ardipithecus ramidus dentition” Science 326:69, 94–99; S. W.
Simpson et al. (2008) “A female Homo erectus pelvis from Gona,
Ethiopia” Science 322:1089–1092.
xxvi notes on contributors
Matt Sponheimer is a Professor at the University of Colorado at
Boulder. He does research on the ecology of early hominins and
associated fauna in Africa. He has also directed and co-directed
several multi-disciplinary projects on the ecology of living
mammals, both large and small, in South Africa.
David S. Strait is an Associate Professor of Anthropology at the
University at Albany (SUNY). He is a paleoanthropologist with
interests in hominin phylogeny, the evolu- tion of feeding
biomechanics in primates and fossil humans, hominin biogeography,
and paleoanthropological fieldwork. Among his recent publications
are: Strait, D.S. et al. (2009) “The feeding biomechanics and
dietary ecology of Australopithecus afri- canus” Proceedings of the
National Academy of Sciences of the USA 106:2124–2129; and D. S.
Strait, and F. E. Grine (2004) “Inferring hominoid and early
hominid phylogeny using craniodental data: the role of fossil taxa”
Journal of Human Evolution 47:399–452.
Nicholas Toth is a Professor of Anthropology at Indiana University
Bloomington and co-director of The Stone Age Institute. He earned
his PhD in anthropology at the University of California, Berkeley.
His topical interests include paleoanthropol- ogy, lithic
technology, and African prehistory, and his geographical areas of
specializa- tion are Africa and China. He was elected as a Fellow
of the American Association for the Advancement of Science in
2004.
Peter S. Ungar is a Professor at the University of Arkansas. His
publications include The Diets of Early Hominins, co-authored with
Matt Sponheimer, Mammal Teeth: Origin, Evolution, and Diversity,
Johns Hopkins University Press 2010, and Evolution of the Human
Diet: The Known, the Unknown, and the Unknowable, Oxford University
Press 2007; his most recent publications include P. S. Ungar, and
M. Sponheimer (2011) “The diets of early hominins” Science
334:190–193.
Carol V. Ward is Professor and Director of Anatomical Sciences in
Integrative Anatomy in the University of Missouri School of
Medicine Department of Pathology and Anatomical Sciences. She has
published papers presenting new fossils of fossil monkeys, as well
as Proconsul nyanzae, P. heseloni, Afropithecus turkanaensis,
Australopithecus anamensis, A. afarensis, A. boisei, and Homo
habilis. She is a William T. Kemper Teaching Fellow and a Fellow of
the American Association for the Advancement of Science.
Milford H. Wolpoff is a paleoanthropologist, and since 1977, a
Professor of Anthro- pology and Adjunct Associate Research
Scientist, in the Museum of Anthropology at the University of
Michigan. He is the leading proponent of the multiregional evolu-
tion hypothesis that attempts to explain the evolution of Homo
sapiens as a conse- quence of evolutionary processes within a
single species. He is the author of Paleoanthropology, 1980 and
1999 editions with McGraw-Hill, New York, and the co-author (with
Rachel Caspari) of Race and Human Evolution: A Fatal Attraction,
which reviews the scientific evidence and conflicting theories
about how human evo- lution has been interpreted, and how its
interpretation is related to views about race.
notes on contributors xxvii
Bernard Wood is University Professor of Human Origins at the George
Washington University Center for the Advanced Study of Hominid
Paleobiology. His publications include: Koobi Fora Research
Project. Volume 4: Hominid cranial remains (1991) Clarendon Press;
B. A. Wood (1992) “Origin and early evolution of genus Homo”
Nature, 355:783–790; editor of Wiley-Blackwell Encyclopedia of
Human Evolution (2011); and Rui Diogo and Bernard Wood (2012)
Comparative Anatomy and Phylogeny of Primate Muscles and Human
Evolution. CRC Press–Taylor and Francis.
I am extremely grateful to all the friends and colleagues who
contributed to this book. All were eager to do so, and I am
delighted with the results. I would also like to thank Rosalie
Robertson for asking me to edit this volume and for her recognition
of the need for this kind of book. I am also grateful to Julie Kirk
and Helen Gray for their skillful managing activities during the
unfolding of this project. I thank Alec McAulay for his
copy-editing efforts that made that phase of the project easy for
me. Thank you as well to Mariam Nargolwalla for her help with the
proofing and index. Finally, I would like to thank my wife,
Dana Bovee, for her support and especially for our ongoing
collaboration at our excavations in Rudabánya.
Acknowledgments