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Book Reviews
AND CONSERVATION. Edited by Serge A. Wich, S. Suci
Utami Atmoko, Tatang Mitra Setia, and Carel P. van
Schaik Oxford: Oxford University Press. 2009. 464 pp.
ISBN 978-0-19-921327-6. $125.00 (cloth).
Orangutans, the sole great apes of Asia, are large, arboreal frugivores with a unique, semisolitary but complex
social organization. Fifty years ago orangutans in the
wild were the great unknown. As late as 1995, a book
entitled The Neglected Ape could be published, indicating
the relative lack of knowledge concerning orangutan
populations in the wild. To a great extent, Orangutans:
Geographic Variation in Behavioral Ecology and Conservation remedies this neglect. In the last 15 years, our
knowledge of wild orangutan populations has grown
exponentially, particularly in Sumatra where so little
was known before. The words ‘‘geographic variation’’ say
it all. This brilliant book details the extent of variation
among orangutan populations in different geographical
areas. Although captive data are aptly used, the emphasis is on applying recent field work to explicate differences and similarities among the four orangutan populations found in the wild: the Sumatran species Pongo abelii, which occupies the most western orangutan range,
and the three Bornean subspecies, Pongo pygmaeus pygmaeus, P. p. wurmbii, and P. p. morio. While highly technical, this book is also strangely fascinating. A friend,
seeing the volume on my desk, picked it up and started
reading. ‘‘Wow!’’ he said, looking genuinely surprised, ‘‘I
had no idea orangutans eat slow lorises!’’ His response
mirrored my own reaction to this book; I found it fascinating and hard to put down.
The main question, of course, is how different are Bornean from Sumatran orangutans and, secondarily, are
Bornean populations all that different from each other?
Although we are left with an overwhelming impression
of basic adaptations and behavior shared by all orangutans, the differences among orangutan populations are,
in some ways, quite profound. I have always thought
that, in this era of taxonomic inflation, if the Sumatran
and Bornean orangutan populations were granted species status, then perhaps we should take another look at
the three Bornean populations and evaluate whether
their differences warrant it as well. Although the islands
of Borneo and Sumatra were joined as late as twelve
thousand years ago, the mighty rivers that separate
Bornean orangutan populations have been flowing for
hundreds of thousands of years. The genetic picture,
as detailed here by Goossens et al., is complex but generally supports a Sumatra-Borneo bifurcation. Of course,
as we learn more about orangutans, our ideas about the
differences among the various populations change. For
instance, wild behaviors that were initially considered
possible cultural variants are now regarded as universals, among them the ‘‘kiss squeak with hands’’ and
‘‘snag crashing.’’
To answer the question of difference among orangutan
populations, the volume examines aspects of many
behaviors, sometimes in overwhelming detail, and the
particulars of orangutan ecology in the various regions
C 2010
where they range. For instance, there are no differences
in terms of postures exhibited by Bornean and Sumatran
orangutans, only in relative frequencies. Although there
is consistency in locomotor behavior between the species,
there are interspecific differences that may be related to
interstudy differences. Thorpe and Crompton do not discount the fact that the largest differences in positional
behavior may exist at the habitat level rather than the
species level.
An absorbing chapter by Taylor indicates that P. p.
morio and P. p. wurmbii are better adapted to resisting
large and/or frequent jaw loads than the Sumatran species, while P. p. pygmaeus is intermediate. Taylor points
out that while both Bornean and Sumatran orangutans
favor ripe fruit, they diverge in terms of their foraging
when fruit is limited. Sumatran orangutans spend more
time eating figs and insects, whereas Bornean orangutans depend on inner bark and fibrous vegetation. This
is probably a function of differential island ecology, with
soil fertility and habitat productivity both less in Borneo,
a theme echoed throughout this volume. This may
explain why Bornean orangutans forage on a wider
range of foods than Sumatran orangutans, which are
more selective feeders. Borneo also exhibits differential
effects from the El Niño weather phenomenon, with
eastern Borneo suffering from more severe droughts
than the western side of the island. Thus, it is not surprising that orangutans in east Borneo, P. p. morio, rely
most on alternative foods, as fruit is scarce during extensive droughts. Taylor concludes that orangutans ‘‘display
a relationship between variance in energy intake, feeding efficacy and relative brain size’’ (p. 15), with P. p.
morio exhibiting a relative reduction in brain size compared with other orangutan populations.
The gradient first mentioned by Taylor in orangutan
behavior and habitat from eastern Borneo to Sumatra
becomes a recurrent premise. Wich et al. document that
interbirth intervals decrease from Sumatra in the west
to eastern Borneo, whereas Husson et al. demonstrate
that orangutan population densities in Sumatra are
higher than in comparable Bornean habitat, indicating
once again that Sumatra’s forests, being more productive, provide better habitat. In Borneo itself, interestingly, there are no significant differences in density.
Marshall et al. present exhaustive, quantitative comparisons of orangutan habitat quality. These analyses indicate that Sumatran study sites have higher percentages
of trees in fruit, on average, and experience fewer and
shorter periods of fruit shortage than Bornean forests.
Noordjwik et al. conclude that there are no major differences between Bornean and Sumatran orangutans in
early development. Nonetheless, in Borneo immature
individuals are found alone occasionally at younger ages
than in Sumatra. Bornean orangutans shorten the association between mother and offspring rather than prolonging the average birth interval. There is also a gradient of increasing home-range size from east Borneo
west to Sumatra (Singleton et al.) that mirrors many
other aspects of behavior and ecology in orangutan populations. Finally, the absence of tigers in Borneo frees
orangutans, particularly males, to be more terrestrial
and thus, perhaps, more mobile. By the time the last
chapter is reached, the reader needs little convincing
that much of wild orangutan social organization,
behavior, and ecology may be explained by habitat differences, particularly the temporal and spatial variability of food.
There is a special irony to the intense and intimate
bond that orangutans share with the tropical forests
that constitute their only habitat. Perhaps the eastern
orangutans, having to handle longer and more frequent
periods of food deprivation in their normal lives are better preadapted than the western orangutans to a world
where Southeast Asian forests are both disappearing at
an ever-accelerating rate and becoming increasingly
fragmented. If this is so, then there should be little surprise that Sumatran orangutans, now fewer than 7,000
in the wild, are critically endangered. But, in the end,
an orangutan is still an orangutan. It is unthinkable
that creatures that have survived two million years in
their present form should go extinct because of the greed
Jean-Jacques Hublin and Michael P Richards. New York:
Springer. 2009. 264 pp. ISBN 978-1-4020-9698-3. $129.00
In 1825, Brillat-Savarin published his famous aphorism, ‘‘tell me what you eat and I will tell you what you
are’’ (Brillat-Savarin’s The Physiology of Taste, translated and annotated by M.F.K. Fischer [1971]. New York:
Alfred A. Knopf. p. 3). One-hundred years later,
pondering the place of a young australopithecine in our
ancestry, Raymond Dart also saw diet as a key to our
humanity, linking anatomical evolution to questions of
environment and technology, and the Taung skull to an
omnivorous, tool-using way of life. Now, nearly a century
after Dart’s arguments, this edited volume takes up
those same dietary themes, as discussed at a 2006 symposium at the Max-Planck Institute for Evolutionary Anthropology in Leipzig, Germany. The 20 articles collected
in this volume sample an interesting selection of topics,
but one that is narrower in scope than the ambitious
title of the book might suggest. The chapters are dedicated to the symposium themes of meat eating/hunting
and resource intensification, and, overall, the topical coverage of the book is skewed toward the Late Pleistocene
and Europe, in particular. Four introductory chapters
provide updated reviews of the ecological principles that
shape contemporary human and nonhuman primate
diet; these articles weave together themes of anatomy,
physiological processes, and ecological context and frame
interpretations made by other articles in the volume.
Within this introductory set of chapters, Snodgrass et al.
provide a particularly useful summary and expansion of
earlier work on energetics, brain size, and dietary quality in human evolution. Subsequent chapters focus on
interpreting hominin behavioral evidence, largely from
archaeological contexts, with a final set of chapters
returning to biological themes. Some present empirical
case studies, whereas others summarize or re-examine
larger-scale patterns.
Across the long period of human biocultural evolution
covered in the volume, early hominin fossil evidence is
American Journal of Physical Anthropology
and obliviousness of close relatives, humans, who have
been on this planet for only a few hundred thousand. I
hope that this excellent volume, which broadens our
understanding of orangutan adaptations, brings us one
small step further toward saving orangutans as biologically viable, wild populations. They deserve no less.
Orangutan Foundation International
and Department of Archaeology
Simon Fraser University
Burnaby, British Columbia, Canada
DOI 10.1002/ajpa.21382
Published online 25 August 2010 in Wiley Online Library
the focus of a few articles, although the Early Stone Age/
Lower Paleolithic archaeological record receives relatively little attention, and dental microwear is neglected.
Reviews of living primate dietary ecology and biogeochemistry provide a broad comparative context for interpretations of early hominin tooth morphology and fossil
signatures of trace elements and stable isotopes. However, questions which arise about the habitat preferences, subsistence strategies, and dietary breadth of early
hominin species remain unresolved here, as elsewhere.
The majority of articles examine subsistence contrasts
between the Middle Paleolithic (MP)/Middle Stone Age
(MSA) and the Upper Paleolithic (UP)/Later Stone Age
(LSA), whether from selected archaeological data or from
behavioral evidence from the hominin fossils themselves.
All these articles seem to agree that meat eating was a
major component of the Paleolithic diet. Even in the one,
lonely chapter focused on archaeobotanical evidence,
Jones discusses possible Upper Paleolithic plant foods in
the context of ‘‘a human ecology largely concerned with
meat’’ (p. 171). Several articles evaluate faunal and lithic
evidence for temporal or ecological patterns in meat acquisition strategies in different regions. Together they
argue that MP/MSA humans had prey preferences that
varied with habitat; that they were skilled hunters; and
that they would have been very dependent on large
game, supplementing their diet with smaller, easily
acquired prey and other ‘‘slow’’ resources. For one case
study in the Caucasus, discontinuity in the material-culture record is associated with continuity in a variety of
faunal exploitation patterns; thus, Neanderthal hunters
may have been just as adept and seasonally specialized
as the later anatomically modern human (AMH) hunters
who replaced them. However, the transition from MP to
UP is marked by a significant shift in subsistence patterns in other areas. Increased dietary breadth and
larger home ranges are paced by the exploitation of
more expensive, labor-intensive resources using new
kinds of UP tools. These dietary patterns are consistent
with the stable isotope evidence presented in two chapters for Neanderthals and early anatomically modern
humans. In an updated review of MSA and LSA zooarchaeological assemblages in sub-Saharan Africa, Steele
and Klein renew arguments that a similar expansion of
early AMH dietary breadth occurs with the advent of
LSA technologies in Africa. They review evidence from
across the continent that more mobile, flexible subsistence strategies are associated with this cultural shift;
however, the role that biological identity (Neanderthal or
AMH) played in this apparent shift in cultural ecology
remains ambiguous.
Technological investment in subsistence is a theme in
several chapters. Two complementary articles focus on
the identification of projectile points. Shea provides evidence for the early use of MSA points on spears by anatomically modern humans in Africa, and notes that
data are still lacking that would settle the question
of whether spears were used in a comparable way by
Neanderthals. A lively and wide-ranging chapter by Villa
and Lenoir contributes new evidence that low-velocity
thrusting or throwing spears may indeed have been an
early addition to the hunting toolkit in Paleolithic
Europe, as well as in Africa and the Near East. Both
these articles demonstrate how careful attribute analysis
can add clarity to functional questions, which can sometimes be obscured by typological or technological
approaches to understanding artifacts.
Archaeological interpretations of Neanderthal subsistence are placed within a biological context by two
thoughtful articles. A new analysis of Neanderthal and
AMH shoulder morphology by Churchill and Rhodes
creates opportunities for both adaptive and developmental tests of the technological hypotheses of spear
use. Their work complements a chapter by MacDonald
et al., which applies energetic perspectives to interpretations of the subsistence shifts from Middle to Upper
Paleolithic. Both these articles provide an evolutionary
OUT AND WE SURVIVED. Edited by Clive Finlayson.
Oxford: Oxford University Press. 2009. 273 pp. ISBN
978-0-199-23918-4. $29.95 (hardcover).
This book might be more aptly titled How Climate
Change Shaped Human Evolution: Why We Survived
and Others Did Not. It is not a book about Neandertals,
but rather a book about human evolution, albeit with an
emphasis on the Late Pleistocene. Finlayson’s primary
theme is the role of climate change, chance, and luck in
influencing human evolution, especially the success of
our own species.
Finlayson’s 20-page prologue takes us from the mammalian radiation of the Cenozoic, through the spread of
early Eocene primates, and onward to Miocene apes and
early hominins. Not all readers will have the patience to
make their way through this background, but the concepts
presented in the prologue provide an important foundation that sets the proverbial stage for what is to come. And
the concept of a stage is exactly what Finlayson is striving
for as he likens human evolution to a play with actors
(species), a manager (climate), and an ever-changing set
(the environment).
There are many actors in the human evolution play.
Neandertals, as a widely recognized and fascinating
context for analyzing the costs and benefits of different
subsistence strategies, an ecological alternative to various sociocognitive hypotheses considered elsewhere in
the volume.
Many articles in the volume highlight methodological
issues that must temper interpretations. In particular,
there is welcome emphasis on the critical importance of
evaluating the taphonomic integrity and comparability
of samples, variation within assemblages, and incongruous scales of evidence. Thus the scientific challenge of
the volume title—integration––is difficult, and most
chapters juxtapose different types of evidence rather
than actually integrating analyses. Unfortunately,
because the editors provide no comparative discussion at
the end, readers are left to synthesize the results and
relationships of research in these overlapping fields for
themselves. Beyond the quality of individual articles, the
main value of this edited volume is that it illustrates
both the current promise and attendant frustrations of
some important approaches to exploring ancestral diets
as a guide to our evolutionary origins and identity.
Department of Anthropology
Indiana University
Bloomington, Indiana
DOI 10.1002/ajpa.21384
Published online 25 August 2010 in Wiley Online Library
group of humans, get to play a leading role. Why they
disappeared is a question that ranks right up there with
whether or not they interbred with Homo sapiens. Finlayson’s primary aim (and also the point of his previous
book) is to discredit the idea that Neandertals were
‘‘dumb brutes from the north’’ (p. 105) who went extinct
as the result of the encroachment of intellectually superior Homo sapiens. Instead, Finlayson argues, Neandertals were the victims of environmental conditions and
climate change, which stressed them beyond the point of
recovery. Homo sapiens simply came in and occupied the
areas left empty by Neandertal populations that died
out. To make this point, Finlayson provides myriad
examples, within and outside the hominin lineage,
where climate changes benefitted some species and led
to the extinction of others.
Finlayson contends that the most successful groups of
organisms are those living at the peripheries of environments. It is those groups who have learned to exploit
marginal habitats that are less prone to extinction when
times get tough. Throughout human evolution, Finlayson
argues, these peripheral, marginally living species survived best. For example, anatomically modern humans
‘‘proto-Ancestors’’ expanded onto the dry and harsh
steppes, where they learned to survive and thrive even in
the absence of trees for shelter or fire. There, they develAmerican Journal of Physical Anthropology
oped new hunting strategies and cultural innovations.
Most importantly, according to Finlayson, they learned
the concepts of surplus and storage. These concepts would
benefit their descendants, who would eventually become
Using Finlayson’s terminology, these steppe-living
Ancestors were the ‘‘innovators,’’ and Neandertals, who
kept to their old ways, were the ‘‘conservatives.’’ Projectile weapons were of no use in the open woodlands,
Finlayson argues, so Neandertals never made them.
That left them at a disadvantage when their hunting
grounds shrank in response to climate changes and
they faced the encroachment of the open steppes.
There, their body mass, which was so well adapted to
open woodland ambush hunting, would also have put
them at a disadvantage. Those who were already used
to exploiting more open environments (Homo sapiens)
had the advantage, as they merely had to tweak their
technology to make it work in this new setting. They
survived, even flourished, on the steppes, while Neandertals either died out or retreated to refugia (e.g., Gorham Cave in Gibraltar), where they were able to eke
out another couple-thousand years before finally
But the big question is why did those last Neandertals in various refugia eventually die out? Finlayson
suggests that in some cases disease or inbreeding could
have been the cause, but it was certainly not the superiority of incoming Homo sapiens. While I respect (and
agree with) Finlayson’s desire to paint Neandertals as
intelligent humans, I remain unconvinced by his argument that it was solely climatic changes that led to the
Neandertals’ demise. Neandertals survived many cold
snaps and climatic fluctuations over their 200,000-year
tenure on this earth. Their numbers may have
dwindled, but they did not go extinct. The one thing
that coincides very well with their extinction (and is a
change from the earlier circumstances) is a new actor
(the antagonist) on the scene: Homo sapiens. Finlayson
would have us believe that Neandertals were simply
dealt a bad hand, literally ‘‘doomed to extinction’’ (p.
116). But if, as Finlayson himself argues, the ‘‘innovative’’ Homo sapiens exploiting the plains developed
cultural advantages over the ‘‘conservative’’ Neander-
Gluckman, Alan Beedle, and Mark Hanson. Oxford:
Oxford University Press. 2009. 296 pp. ISBN: 978-019-923639-8. $65.00 (paperback).
Although the authors state in their introduction that
this textbook is ‘‘intended for the clinician’’ (p. 15), it
would be useful for any of the various fields of study
that examine health from an evolutionary perspective.
It comprises three parts: an overview of the evolutionary
process with a focus on human evolution, including cultural evolution; four chapters that illustrate how principles of evolution enhance understanding and treatment
of health issues; and a synthesis of the previous sections
to offer a new way of viewing human health and illness.
There is a thorough and up-to-date review of the fundaAmerican Journal of Physical Anthropology
tals, why should we not refer to this as intelligence? In
other cases presented in the book, Finlayson implicates
the need to innovate as a major force behind the evolution of intelligence. In the end, the insistence that
Neandertals went extinct without the help of Homo
sapiens feels more like an effort to be politically correct
than a reflection of reality.
Finlayson may not have succeeded in convincing me
that Neandertal extinction was simply the result of
being in the wrong place at the wrong time, but The
Humans Who Went Extinct does succeed as a general
book on human evolution. It does so in three ways: it
provides information that is probably new to many lay
readers; the information is generally accurate; and it
tells a memorable story. Finlayson takes the reader
from our humble beginnings through the eventual
spread of Homo sapiens all over the world, relying on
clear examples to illustrate his concepts. One of the
book’s strengths is its emphasis on the bushiness of
the human family tree. Another is its emphasis on the
roles of chance and luck in the evolution (including
the success or extinction) of hominin species. The importance of these concepts is often overlooked. The
book would make a useful accompaniment to introductory texts, which provide the much-needed details but
often offer a less readable story. Specialists may take
issue with some of the details or the glossing over of
certain ideas—oversimplification of the Out of Africa
and Multiregional theories, for example. But those of
us who teach introductory human evolution classes
will benefit from Finlayson’s multitude of examples
from the natural world, which help make explaining
the complex human evolutionary story just a little bit
Department of Anthropology
New York University, New York, NY
DOI 10.1002/ajpa.21395
Published online 24 September 2010 in Wiley Online Library
mentals of evolutionary theory followed by chapters on
genetics, epigenetics, and life history theory that draw
on human examples. The first section closes with an
overview of human evolutionary history that includes
evolution not only of physical characteristics but also of
behavioral (e.g., tool making and clothing wearing) and
psychological (e.g., language, theory of mind, and orders
of intentionality) qualities.
In Part 2, the authors get into the subject matter
of the title: the medicine part of evolutionary medicine. They approach this topic from what they call
‘‘four illustrative axes’’ (p. 15): reproduction, nutrition,
biological defenses, and behavior. There are familiar
concepts from sociobiology, including male and female
reproductive strategies, mating systems, and sexuality.
There is a refreshing critique of many of the usual
sociobiology examples (e.g., why men prefer younger
women and women prefer older men as mates), with
the suggestion that cultural factors are often more
useful in explaining human behaviors than evolutionary biology.
As is common in evolutionary medicine, this book
invokes the extraordinary difference between the foods
consumed by our ancestors and those we eat today, with
clear impacts on our health and well-being. One of the
clearest differences has to do with patterns of metabolic
disease and obesity that result from this mismatch of
evolved biology and modern diets. This is the area of evolutionary medicine in which the authors of this text
have made major contributions, so, not surprisingly, it
receives much attention. Their critique and conclusions
bring together genes, prenatal biology, and postnatal
environments including diet, activity level, socioeconomic status, and lifestyle. The recent research and
arguments behind this explanation for rising rates of
obesity and metabolic disease reinforce the conviction
that factors from both biology and culture underlie most
health challenges and also buttress understanding of
the plasticity of human biology that has been the major
target of selection. Although they discuss clinical ways
of reversing some of these trends (e.g., with leptin injections soon after birth), the arguments presented in this
chapter on nutrition and metabolism also point to ways
of dealing with the rising epidemics through ‘‘evolutionary public health’’ measures. One measure that I was
pleased to see emphasized by the authors is increasing
rates and duration of breastfeeding. The baby bottle is
clearly an evolutionary novelty that has profound lifelong health consequences.
Some of the recent writing in evolutionary medicine
can be likened to the just-so stories that characterized
evolutionary psychology in its early years. This sort of
conjecture based on slim and sometimes flimsy evidence
can be off putting to medical researchers and scholars.
Thankfully, the authors of Principles of Evolutionary
Medicine refrain from presenting concepts that are not
firmly grounded in research with some level of factual
basis. They are also much more cautious in their use of
metaphors for describing evolutionary processes. For
example, rather than the common metaphor of maternal–fetal conflict in pregnancy, they suggest viewing
the relationship as ‘‘part of an evolved equilibrium
which mediates the . . . interaction between mother and
her present and future offspring’’ (p. 172). This academic caution will likely make the book more attractive
to medical students and graduate students in biology
and anthropology, but I have to admit that it is not as
engaging or thought provoking as some of the recent
work in evolutionary medicine. On the other hand, this
approach may be what is needed to bring the evolution-
ary perspective back into medical training, research,
and practice.
Although the book is very useful as an introduction
and overview of the field of evolutionary medicine, I
found myself frustrated by the lack of internal referencing and a bibliography. The only evidence that published material was consulted by the authors in writing
the book is the list of suggestions for further reading
found at the end of each chapter. Most of these are
books or chapters in books, and very few journal
articles are referenced. Not using parenthetical citations certainly makes reading easier (and more attractive to students), but it is problematic for a reader who
wants to know the sources of the information being presented. As an example, the authors discuss the concept
of evolutionary ‘‘spandrels’’ (exaptations) introduced by
Gould and Lewontin, but their paper is not cited. I was
intrigued by a box entitled ‘‘Why are pygmies short?’’
(p. 119), but there were no citations for follow-up.
Another box summarizes a study that used body lice to
assess when humans first started wearing clothing.
They quote the authors of the study as saying that the
estimated date of 100,000 years ago is ‘‘surprisingly
recent’’ but provide no further reference.
This book would be useful for an upper division or
graduate course in evolutionary medicine taught in anthropology, psychology, or biology programs, or for the
(unfortunately) rare course taught in medical schools.
It would also be useful for instructors who are seeking
a way of teaching about evolution that addresses
student concerns about relevance. Most of the evolutionary principles discussed are illustrated with medical
examples, ones that likely seem more ‘‘real world’’ to
students than some of the more esoteric examples
found in standard introductory texts in evolutionary
theory. The drawings and figures used to illustrate evolutionary concepts are particularly clear and useful.
Although supplementary readings would probably be
necessary to expand a course beyond the principles so
thoroughly described in this book, a successful course
could easily be built using Principles of Evolutionary
Medicine as the core textbook.
Department of Anthropology
New Mexico State University
Las Cruces, NM
DOI 10.1002/ajpa.21396
Published online 24 September 2010 in Wiley Online Library
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