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
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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.
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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