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Book review Human Origins and Environmental Backgrounds.

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Book Reviews
PERSPECTIVES. Edited by Katerina Harvati
Terry Harrison. New York: Springer. 2006. $129.00
Paleoanthropologists, no matter which part of the
hominin fossil record they work on, all seek answers to
similar, if not the same, questions. How many taxa are
represented in the part of the fossil record they are
interested in? How are those taxa related to each other
and to the taxa that came before and after them? What
are the morphological characteristics of those taxa? How
did they behave? How did the behavioral niche they
occupied differ from that of any coeval hominin taxa?
And so on.
Those of us whose research focuses on earlier phases
of human evolutionary history, where some sample sizes
are pitifully small, now and then cast envious glances in
the direction of the Neanderthal fossil record. By the
standards of what is available for taxa such as Homo
habilis, the Neanderthal hypodigm is extraordinarily
rich. Because much of it was deliberately buried, taphonomic factors have not wrought the havoc they have on
the earlier record, so Neanderthal juveniles are preserved, and regions like the hand and the foot are relatively well represented. And to cap it all, there is even
ancient DNA available for some specimens, albeit in
small fragments. Thus, it is sometimes a puzzle to those
of us who look in from the outside, why the research
questions outlined above are still so much in play with
respect to the Neanderthals. Surely, with this excellent
record, all you have to do is to run the relevant information about the fossils through an appropriate computer
program, and in a few seconds you will have generated
the solution to your phenetic, phylogenetic, functional,
or adaptive conundrum?
At the beginning of 2005, Katerina Harvati and Terry
Harrison hosted a conference entitled ‘‘Neanderthals
Revisited: New Approaches and Perspectives,’’ which
was supported by the Center for the Study of Human
Origins at New York University and by the Department
of Human Evolution at the Max Planck Institute for Evolutionary Anthropology. They deliberately mixed senior
and junior researchers and asked the young and the not
so young alike to set out how fresh approaches might
reduce our ignorance of Neanderthal paleobiology. The
papers run the gamut of research themes and styles,
ranging from a study of the Neanderthal hand to one
that claims to have found evidence of selection acting on
mtDNA. Nonetheless, two sets of questions predominate
among the papers. The first concerns the nature of
Neanderthals. Are they distinctive? And if so, what does
their distinctiveness tell us about their relationship with
modern humans? The second set of questions assumes
that Neanderthals are distinctive at some biologically
meaningful level and goes on from that premise to try to
derive function from form. Just what were these morphologically distinctive creatures adapted (or maladapted) for?
Most of the contributions use one or other circumlocution (e.g., ‘‘an unambiguously demarcated morphoC 2008
species’’; ‘‘a historically individuated entity’’), which,
roughly translated, means they accept the working hypothesis that Neanderthals are a distinct species.
Three papers (Antonio Rosas et al., and both nameorder permutations of Maria Ponce de León and Christoph Zollikoffer) look to development for clues about
the origins of the Neanderthals. Rosas et al. set up the
‘‘accretion’’ model in opposition to what they term the
‘‘organismic’’ model of Neanderthal evolution. But to
me they seem complementary explanations; the accretion model is about mode, and the organismic model is
about mechanisms.
Ponce de León and Zollikofer track the cranial ontogenies of modern humans, Neanderthals, chimpanzees,
and bonobos in shape space in order to compare their
growth trajectories. It is implied that the differences
between the chimpanzee and bonobo trajectories are
likely to be the yardstick for a species-level difference.
Although all four of these creatures are part of the
hominid (sensu molecular-influenced taxonomy) twig of
the Tree of Life, it seems to me that while parsimony
would suggest that developmental constraints are
likely to have been similar within the Pan-Homo clade,
there is no logical reason to assume that all hominid
sister taxa became distinct via the exact same ontogenetic modifications. The ecological context for the emergence of the Pan and Homo clades was most likely not
the same as that which accompanied speciation events
within the later part of the hominin clade, so why
should development be modified in the same way?
Others have made the case that evolution is the effect
of ecology on development, and I, for one, am convinced
that if one can recover the developmental history of
morphological traits, then this will help sort homoplasies from homologies. However, for all their obvious
promise, these investigations of Neanderthal ontogeny
have not, as yet, taken us much further than AnneMarie Tillier did when she pointed out that the distinctive Neanderthal morphology was obvious even in
infants. You need a lot of data to compare ontogenies,
and, excellent though the Neanderthal fossil record is,
it is not yet quite good enough.
Harvati and Weaver make good use of geometric
morphometrics to investigate the relative utility of different regions of the hominin cranium for assessing
population affinities and for phylogeny reconstruction.
I coauthored a paper suggesting that conventional cranial morphometrics do not seem to be capable of reliably recovering the branching pattern of cladograms
based on molecular data. But Harvati and Weaver
show that when morphology is captured using more sophisticated 3D methods, the cranial base is better at
recovering deeper relationships than regions like the
face and vault. This is an important study that should
encourage others to undertake similar analyses using
other data sets.
To my mind the star of the show is Steve Churchill’s
report of his elegant attempt to provide a heuristically
useful model of Neanderthal bioenergetics. He focuses
on the large chest cavity seen in Neanderthals and
tries to discriminate between two functional explanations. Is its large size related to reducing heat loss or
is its large size a mechanism for heat production?
Churchill comes down in favor of the latter, but
because he writes so well and explains his complex
arguments so clearly, even if you do not agree with his
conclusions, you will be both informed and educated if
you read this paper.
Most of the chapters in this book are a rich source of
information and inspiration for students, researchers,
and teachers. My take is that despite the application of
the new methods featured in many of the book’s chapters, there is much still to do to understand the paleobiology of the Neanderthals. The depressing inference is
that if this is the case with a hypodigm as rich as that
for the Neanderthals, then what hope is there for those
by Hidemi Ishida, Russell Tuttle, Martin Pickford,
Naomichi Ogihara, and Masato Nakatsukasa. New
York: Springer. 2006. 281 pp. ISBN 0-387-29638-7.
$139.00 (hardcover).
The primate fossil record of the Miocene, considered
the golden age of ape-like primates, is characterized by
highly diversified groups of proconsulids, dryopithecids,
and the successive radiation of Old World monkeys, particularly the cercopithecoids. East Africa is blessed with
an abundance of Early to Middle Miocene localities bearing hominoids, which are rare in the Middle to Late Miocene epoch. The rarity of apes in the fossil record toward
the end of the Miocene has sparked some debates on the
effects of climate forcing, particularly on the evolution of
extant African apes. Human Origins and Environmental
Backgrounds, edited by Ishida and colleagues, is perhaps one of a few compilations of detailed studies of
East African Miocene hominoids and their depositional
The peer-reviewed volume is the tenth in Russell Tuttle’s successful Developments in Primatology series published by Springer. It exemplifies the range of research
topics and methodological issues addressed by primatologists and paleoanthropologists today and is devoted to
research and scholarship on the functional morphology,
positional behaviors, stratigraphy, and geochronology of
East African Miocene primates. Although the volume
originates from a conference dedicated to Dr. Hidemi
Ishida’s contribution to the study of primatology and
hominoid evolution, it is a great source of scholarly information related to current issues in human evolution.
Human Origins and Environmental Backgrounds is
divided into three major sections: ‘‘Fossil Hominoids and
Paleoenvironments,’’ ‘‘Functional Morphology,’’ and ‘‘Theoretical Approaches.’’ The first section provides some historical background on East African Miocene primate
research, especially work on primate positional behavior,
paleoenvironmental interpretations, geology, and the
depositional histories of East African sediments. The second section deals with issues related to functional morphology and inferred locomotor repertoires of Miocene
primates, while the third and last section deals with theoretical frameworks and research approaches currently
used in East African paleoanthropology.
In the first section, an introductory tribute by Nakatsukasa and coworkers to Ishida’s contributions to priAmerican Journal of Physical Anthropology
of us who focus on parts of the fossil record where the
evidence is, as yet, much sparser?
Center for the Advanced Study of
Hominid Paleobiology
Department of Anthropology
George Washington University
Washington, District of Columbia
DOI 10.1002/ajpa.20862
Published online 18 August 2008 in Wiley InterScience
matology and paleoanthropology is followed by five contributed articles that range in scope, covering issues
such as the history of East African Miocene primate
research, evolution of the vertebral column in Miocene
hominoids, and terrestriality of Old World monkeys. Tuttle’s paper on seven decades of East African Miocene
anthropoid studies provides a general overview on the
pace of fossil primate research as well as a basis for subsequent contributions. Gommery, for example, presents a
very compelling study on the evolution of the vertebral
column in Miocene hominoids and Plio-Pleistocene hominins, while Blue et al. discuss positional behaviors of
Middle Miocene monkeys, with an emphasis on Victoriapithecus from Maboko Island.
Mammalian biostratigraphic correlative studies rarely
receive attention in paleoanthropology beyond their importance in relative dating of hominoid-bearing deposits.
A detailed and generalized study of mammalian biostratigraphy and faunal turnover events in relation to
paleoenvironments and hominoid evolution during the
Cenozoic is well presented by Nakaya and Tsujikawa. It
is complemented by Sawada et al.’s study on the geology
and depositional history of Miocene fossiliferous sediments bearing Nacholapithecus, Samburupithecus, and
Orrorin from Kenya. This particular biostratigraphic correlative study provides a rich, informative, time-averaged stratigraphy illustrated by columnar sections
describing various facies, which provide readers with a
better understanding of temporal relationships of fossiliferous beds that have been associated with these important hominoids.
The second section, which consists of seven contributions, provides detailed discussions of primate comparative functional morphology in relation to locomotion.
Nakano’s study of patterns and modes of arboreal locomotion, using three-dimensional techniques, offers some
insightful information on the varying degree of functionality of the forelimb and hindlimb muscles in verticalclinging primates. This short paper is complemented by
Richmond’s detailed comparative study of primate hand
anatomy and its relevance to human bipedality. In this
section, primate positional behaviors and locomotor preferences continue to dominate where topics on energetics
and biomechanics are used to develop models for the evolution of upright stance in our ancestors. For example,
Matsumura et al. present probably one of the few mammalian examples of hindlimb morphology and mechanical loading that can be associated with varying postural
adaptation. This article goes together with four other
contributions: Hirasaki et al. on experimental biomechanical studies using trained primates; Nakatsukasa
et al. on locomotor energetics in nonhuman primates;
and Jouffroy and Medina on the form, action, and function
of gluteus maximus as it relates to different postural
behaviors across an array of primates. Finally, Ogihara
and Yamazaki conclude this section with their studyemploying computer simulations of bipedal locomotion
using musculoskeletal system constraints during a given
locomotor repertoire-on issues of primate postural positional behavior relevant to the evolution of bipedality.
The third and final section sums up key issues associated with East African Miocene primate studies. Tuttle’s
article on neontological perspectives on East African
Middle and Late Miocene anthropoids complements most
of the articles in the first section, thus providing a
reader with a better summary on the evolution of East
African Miocene hominoids. Pickford provides a detailed
discussion on paleoenvironments, paleoecology, adaptation, and the origins of bipedalism and posits some
questions that challenge our understanding of Miocene
primate adaptation. Senut’s work on arboreal origins of
bipedalism describes some important morphological
features of Miocene hominoids that link arboreal bipedality with terrestrial upright walking. Okada’s article
presents more evidence that not only supports Senut’s
by Pauline Asingh and Niels Lynnerup. Højbjerg,
Denmark: Aarhus University Press. 2007. 351 pp.
ISBN 8-788-41529-5. $46.00 (hardcover).
The Grauballe Man was found by peat cutters in 1952
in a bog in Jutland, Denmark. The preservation of the
body, one of many such bog bodies across Northern
Europe, is attributed to the tanning action of the bog
water. The body was removed to a local museum, where
studies including radiocarbon dating, radiologic and forensic examination, and examination of intestinal contents were initiated by Professor P. V. Glob. The body
was then conserved for museum exhibition. Fifty years
later, another group of scientists gathered around the
Grauballe Man, this time at a hospital, to apply the latest in medical and scientific technology to this visitor
from the past. Their efforts have resulted in a superbly
illustrated volume, providing much more information
and correcting some earlier misconceptions regarding his
life and death.
My first step in reviewing this volume was to reread
the 25-page chapter on Grauballe from the 1969 English
translation of Glob’s The Bog People: Iron Age Man Preserved. He concluded that the body was a 30-year-old
man with fractures of the skull and left femur and tibia,
with death due to a deep cut across the throat that severed the trachea, esophagus, and great vessels. The lack
of any artifacts associated with this naked body made archeological dating impossible. Repeated radiocarbon dating has now moved the date of his death from about 210–
410 AD back to about 400–200 BC, i.e., from the Late
proposal but also elucidates the importance of biomechanics and functional morphological studies as locomotor indicators in primates. Although the article by
Furuichi on the evolution of the social structure of hominoids digresses from the main themes of this volume,
still it offers some theoretical frameworks equally important in construction of social and positional behaviors of
Miocene apes. A thought-provoking article by Tuttle on
the question of whether human beings are apes or apes
are people too is a reminder of the many challenges the
field of paleoanthropology faces, particularly from popular opinion. Ishida sums up in this section with some
thoughts on terrestrial locomotion and the origin of
human bipedalism. In general, I highly recommend this
volume to students of East African hominoid paleoanthropology and human evolution.
Department of Anthropology
University of Colorado Denver
Denver, Colorado
DOI 10.1002/ajpa.20874
Published online 9 July 2008 in Wiley InterScience
Iron Age to the Early Iron Age and has certainly laid to
rest the early contention by some that Grauballe Man
was a contemporary Jutlander who had died in the bog.
Conservation of the body was initially undertaken by
the newly appointed Conservator of the Prehistoric Museum in Aarhus, Gunnar Lange-Kornbak. Quite a challenge on his first day at work! Reexamination of the
body has revealed Lange-Kornbak’s techniques, secrets
that he took ‘‘to the grave’’ (p 50). The process involved
creation of an artificial bog, tanning in an oak-bark
bath, and subsequent treatment with oils and resins. A
surprise was that numerous body parts, including the internal organs and several vertebrae, had been replaced
by what appear to be sponges, wax, and blackboard erasers. The vertebrae followed a tortuous path, from Denmark to Professor William Laughlin’s labs, first at the
University of Wisconsin and then at the University of
Connecticut, and finally to the Museum of the Aleutians
in Unalaska, Alaska. A photograph in the book even
shows Laughlin and Lange-Kornbak removing the vertebrae in 1956.
Based on interpretation of the intestinal contents as a
special sacrificial meal, Glob considered the bog bodies to
be sacrifices to the divine of the period. The detailed
modern analysis indicates that the last meal was simply
the porridge or gruel eaten by the common people of the
time rather than any special ritual meal. CT scans, not
available at the time of the initial studies, now reveal
the skull and femoral fractures to be postmortem and
due to the pressure of the peat. The tibial fracture is
perimortem, indicating the Grauballe man was knocked
to his knees and then seized by the head, with the fatal
American Journal of Physical Anthropology
wound inflicted by a right-handed person standing
behind him. Based on three-dimensional visualization, a
full size epoxy model of the skull was made, which
allowed a facial reconstruction, putting us face to face
with an Early Iron Age man.
Histologic study of the skeleton revealed that the
decalcification noted radiologically was not due to osteoporosis but rather to the chelating of bone calcium by
sphagnum moss. A similar effect was seen on the teeth,
with the enamel totally lost. Osteon analysis provided an
age of 34 years, consistent with changes in non-synovial
joint morphology, dental analysis, growth plate fusion,
and degenerative joint changes. Grauballe Man is a redhead now but probably was not during life, as the brown
eumelanin pigment in hair fades with time, leaving the
red phaeomelanin pigment. His hair also provided evidence of a terrestrial diet with primarily animal-based
protein. Microscopic examination of his preserved intestinal tract was not rewarding, with only collagen of the
intestinal wall remaining. In general, there was no evidence of pathology in the body other than the traumatic
wounds, dental wear, periodontitis, and tooth loss.
The question raised by this publication is whether the
application of techniques not available 50 years ago has
provided us with significant new information. This goal
has certainly been met. Perhaps disappointing in one
respect is that Grauballe Man was not a special person,
simply a typical man of the Early Iron Age. In a broader
American Journal of Physical Anthropology
context, he is very special indeed, a ‘‘man on the street’’
preserved for us to see as a representative of his time
over 2,000 years ago. The primary aims of the new studies, evaluating Grauballe’s state of preservation and
ensuring his future survival, have been achieved, and
we know much more about his life and death. As an
added bonus, a penultimate chapter on the bog people
offers a photographic review of many of the bodies found
in bogs in northern Europe, England, and Ireland. The
book is extensively referenced and, although lacking an
index, will be a valuable asset for paleopathologists and
Iron Age specialists, especially at this attractive price.
Department of Biology
Villanova University
Villanova, Pennsylvania
Department of Anthropology
University of Pennsylvania
Philadelphia, Pennsylvania
DOI 10.1002/ajpa.20904
Published online 18 August 2008 in Wiley InterScience
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