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Body mass in cercopithecidae (Primates Mammalia) Estimation and scaling in extinct and extant taxa.

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Book Reviews
Delson, Carl J. Terranova, William L. Jungers,
Eric J. Sargis, Nina G. Jablonski, and Paul C.
Dechow. New York: American Museum of Natural
History. 2000. 159 pp. ISSN 0065-9452. $16.50
This monograph, Body Mass in Cercopithecidae
(Primates, Mammalia); Estimation and Scaling in
Extinct and Extant Taxa, by Delson et al., is of
interest to scholars (especially graduate students
and professionals) whose research focuses on allometry, sexual dimorphism, estimation of body size in
fossil taxa, cercopithecoid evolution, evolutionary
changes in body size through time, and the relationship between ecological and morphological variables
and body size. The main purpose of this monograph
is to develop reliable equations for determining body
size in fossil cercopithecoid taxa, using ordinary bivariate least squares regression. Once this is accomplished, the second half of the monograph focuses on
using the equations to estimate body mass for nearly
all known fossil Old World monkeys.
The monograph is an extremely valuable resource
for anyone studying Old World monkeys. First, it provides body weights for male and female extant cercopithecoids by individual (Appendix Table 1) and
grouped data by sex within species (Appendix Table 2).
The authors took great pains to compile body weight
data for extant monkeys from individual museum
specimens and from highly reliable published primary
and secondary sources. Second, it provides what may
be the best current estimates for fossil cercopithecoids
(Tables 14 –17). The monograph is likely to be cited
frequently because of these tables.
The first half of the monograph describes the materials and methods used by the authors to develop their
equations, and compares the predictive reliability of
their equations to those of previous studies. Unlike
studies that use all primate or all anthropoid regressions to predict the body weight of extinct cercopithecoids, this study restricts the analysis to extant cercopithecoids, and even to subfamily and sexed subfamily
groups in an effort to make more accurate body mass
predictions when sex and subgroup are known for a
fossil taxon. The equations for predicting body mass
are generated based on 35 variables from the postcranium, dentition, and cranium for all cercopithecoids,
for each subfamily, and for sexed groups within each
subfamily (Table 7). The relative reliability of each
equation is judged by the mean prediction error (MPE,
discussed on p. 18), which has an inverse relationship
with reliability. A further test for reliability is carried
out by calculating mass for a subset of 20 extant spe©
cies (Table 9), including some (e.g., Mandrillus sphinx
and Macaca sylanus) for which reliable body weight
data became available after the study. Of 95 total
cases tested (by group and body region), 66 estimates
(about 70%) were within 20% of the actual compiled
body mass for a species. The authors argue that 20% is
the closest that one can expect to come to a species’
mean, based on the regression equations. Most errors
in estimating body mass were encountered when
mean mass was calculated for the largest and smallest
species, or when cranial variables were used. In general, postcranial and dental variables were found to be
more reliable indicators of body mass than cranial
variables. A series of reduced major-axis regression
equations was used to further investigate the relationship between skeletal or dental dimensions and body
mass for each subgroup (Table 11).
The application of the equations to estimating
body mass for fossil cercopithecoid taxa is fairly
straightforward. The most problematic estimate is
for the Miocene fossil colobine Microcolobus tugenesis. In fact, the only mistake I found in the tables
concerns this species, for which predictions of body
weight using the male regression model are listed as
postcranial body weight estimates rather than dental estimates (Table 14, p. 52). The only known specimen of M. tugenensis is a mandible identified by
myself as female, based on the minimal size of the P3
honing facet. One of the authors (Eric Delson) questions the sex determination for the specimen, because the body mass prediction based on the female
colobine regression seems too high at 8 kg (p. 50).
Alternative hypotheses are never considered by the
authors, e.g., that a cercopithecine regression may
be more appropriate for the species since it may not
yet have evolved a sacculated stomach (the specimen has a lower than expected shear quotient for a
colobine), or perhaps that the species does not fit any
of the extant cercopithecoid-based models. A tendency to use body mass predictions to determine the
sex and correct species designation of specimens
occurs at least two more times. The suggestion is
made that the unsexed mandible of Prohylobates
simonsi is “likely to be male,” based on the fact that
the male equations produce an estimate of 20 kg and
the female regression equations an estimate of
30 – 40 kg for the species (p. 82). P. simonsi is highly
unusual among cercopithecoids in terms of its tooth
proportions. I would be surprised if modern cercopithecoid regressions could reliably predict its body
mass. To suggest the sex of this specimen based on
its predicted mass seems absurd. Estimates of male
and female body mass for fossil Papio hamadryas
are similarly used to suggest that levels of sexual
dimorphism are not so low as to “question its inclusion in the living species” (p. 76).
Aside from making taxonomic inferences from
body mass predictions, the monograph makes very
few inferences based on the large number of body mass
estimates made for all fossil primates. Plots of body
mass estimates for male and female fossil colobines
(Fig. 15) and cercopithecines (Figs. 16–19) are fascinating and should be useful to everyone studying the
evolution of Old World monkeys and of body size.
In summary, this monograph is an excellent one.
Given its low price, it is a bargain.
PRIMATE TAXONOMY. By Colin Groves. Washington,
DC: Smithsonian Institution Press. 2001. 350 pp.
ISBN 1-56098-872-X. $65.00 (cloth).
Readers of this Journal should need no convincing
that sound taxonomy is required if we are to be
effective at researching any aspect of primate biology. However, it is such a fundamental requirement
that we take it for granted that someone, somewhere, is working away curating the taxonomy of
the group we are interested in. Perhaps because of
my own background, the best analogy I can think of
for what we expect of taxonomy is related to medicine. When our specialists advise us about how to
treat a medical condition, we want them to be up to
date, but also to have a healthy skepticism about the
hype that so often accompanies announcements
about the latest advances in therapy. So it is with
taxonomy. We want it to reflect recent advances in
the relevant sciences, but we also want it to have
sufficient stability so that it does not change in response to every passing fad. Colin Groves is one of a
small and dwindling number of scientists who concentrate their energy and scholarship on matters
taxonomic. His interests extend beyond the primates, but he has sensed the need for scientists who
are consumers of primate taxonomy to have access
to an up-to-date statement about the best way to
reflect primate diversity in a taxonomy.
Perhaps it is appropriate that this book is a hybrid. Just over 50 of its 350 pages are devoted to
explaining what taxonomy is and how it should be
practiced. First, the good news. The 10-page history
of primate taxonomy is a delight. It is “vintage
Groves,” with interesting historical detail interlarded with engaging asides about the personalities
involved. The guide to nomenclatural rectitude is
also helpful. The bad news, for me at least, is that
the sections on taxonomic methods, and on what
species are, failed to provide what I was looking for.
The section on “relatedness” is shaky. There are
occasional confusions between a phylogenetic tree
and a cladogram and between paradigm and hypodigm, there is unnecessary ambiguity about the
role that cladistics plays in alpha taxonomy, and the
definition of polyphyly is unconventional. As for the
section on species definitions, without denying the
occasional difficulties encountered by neontologists,
the real problem with species concepts is whether
and how they can be applied to the paleontological
Department of Sociology and Anthropology
New Mexico State University
Las Cruces, New Mexico
DOI 10.1002/ajpa.20020
Published online in Wiley InterScience (www.interscience.wiley.
record. You only have to look at the range of taxonomic solutions being offered for the last two million
years of fossil evidence for human evolutionary history
to realize that paleontologists are in need of some
sound, unambiguous advice from an expert about how
to convert the variation we observe in the fossil record
into durable hypotheses about specific diversity.
The options for the section on the concept of species were to either comprehensively review all the
literature and come out with a distillation of the
strengths and weaknesses of all the various proposals, or to “cut to the chase” and provide some brisk,
didactic advice about how to do the taxonomy of the
primate fossil record. What we have is neither a
comprehensive survey, nor any practical advice
other than “I employ the phylogenetic species concept for living Primates because there is no alternative, and the composite species concept for fossils,
again because there is no alternative.” But exactly
how does the author convert Cracraft’s definition of
the phylogenetic species concept (PSC) into practical
advice? How do you recognize “a parental pattern of
ancestry and descent?” As for the composite species
concept, just how do you marry “the PSC to the internodal concept?” If this book is to be a practical manual,
then examples of how these different concepts would
be applied to some real living animals and some real
fossils would have been hugely helpful. There is only
one diagram, a rendition of the shortest platyrrhine
tree, in the whole of the introductory section. To tackle
thorny issues like relatedness and species in the absence of any diagrams is a tall order.
The vast majority of the text is a detailed speciesby-species treatment of the living primates, and this
is where the great value of this book lies. However,
even this invaluable resource would have benefited
from summary taxonomies for each of the major
groups. As it is, if you want to know Groves’ advice
about the taxonomy of the living higher primates,
you have to read through detailed sections to work
out that he advocates recognizing two species each
for Pan, Gorilla, and Pongo. He ducks fossil primates altogether, except for a summary classification of the Catarrhini.
The glossary is useful, but there are some puzzling omissions such as “hypodigm,” and inconsistencies between the text and the glossary. Also,
some statements, such as the assertion that evidence from morphology and molecules converges as
the number of characters increases, need more sub-
stantiation, and three paragraphs about the contribution of DNA evidence to taxonomy are probably
not enough.
This volume is useful, but future editions will be
even more useful if they focus on the taxonomy of
living primates and abandon much of the preamble.
I know it will add to the overall length, but having
the full citations of at least the recommended nomina, if not all the redundant synonyms, would
greatly add to this book’s value as a resource.
I hope Colin Groves will continue to accept the
daunting challenge of curating the taxonomy of the
living primates. No one is better qualified for the
Department of Anthropology
George Washington University
Washington, DC
By John H. Relethford. New York: Wiley-Liss.
2001. 252 pp. ISBN 0-471-38413-5. $69.95 (cloth).
tell us most about is demography: population size,
growth, change, subdivision, and migration. Historical reconstructions must be interpreted in this context. We are burdened by difficult nonidentifiability
problems, i.e., equivalently well-fitting explanations
based on different plausible values of the requisite
demographic parameters. The author presents these
carefully, showing (often by simple simulations)
some of the interpretive issues. A separate chapter
explains and evaluates the Neanderthal DNA data.
Nothing is perfect, and there are some limitations
in this book. To me, one of these is the absence of
historical depth regarding the competing models.
Long before genetics came on the scene, essentially
the same positions were held, for sociological and
other reasons that Loring Brace and others have
discussed at length, and the genetic arguments
themselves are not very different from what they
were 30 years ago. This should give readers pause in
the face of today’s sometimes-strident advocacy, because the combatants talk past each other in part
because they see through prefocused glasses.
Secondly, the genetics are not completely up to
date, especially in regard to the diversity of new
data, including haplotype analysis, detailed single
nucleotide polymorphism (SNP) surveys, and
genomic analysis of autosomal DNA. The story told
by these newer data is essentially the same, but
richer in detail. The author may be a bit too casual
about curious findings, such as Y and mtDNA reconstructions that suggest dramatically different male
and female origins for the same population. I think
some fanciful oversimplifications in the literature
could be avoided if the participants in the battle
introduced more anthropology, e.g., trivia such as an
understanding of cultural anthropology or huntergatherer demography. This book tries to put at least
some of that back into the picture.
More consideration may be due to the role of selection along with demic diffusion as a confounder of
drift-based reconstructions. For example, it has long
been known that “waves of advance” of advantageous genes could explain much of what we see
today, and could distort estimates of origin-times
made from drift-based assumptions (Weiss and Maruyama, 1976). Genomic analysis is revealing heter-
John Relethford has long weighed in on the problem of reconstructing human origins. He has written
balanced treatments in his numerous introductory
textbooks, and now has written another, extended
treatment of the subject, that should be on your
John presents the evidence on human origins,
ranging from population genetics to fossils to osteometrics. The contending models for human evolution
are presented in depth. The combating forces are the
“Out of Africa Replacement” and the “Regional Continuity” models. I think the pugilists have recoined
and hyped their theories without appropriate acknowledgment of historically venerable positions,
and I feel similarly about the extensive treatment
given to “mitochondrial Eve,” to me one of the most
damaging publicity stunts of our field. In a marketing age, I guess we’re stuck with pop-sci, but readers
can be confident that John’s book is groping for
understanding, not publicity.
The relevant population genetics theory and data
behind these contrasting models are explained with
enough mathematics to make the points clear, in a
balanced, well-written, smoothly digestible way.
Rather than burden the reader with formalism, the
author plays to his own strengths, which include
cogent simulations that illustrate major points without overstating them. After a brief introduction to
genes and population genetics, he examines most of
the major topics we need to consider, carefully evaluating what others have said, and presenting analyses of his own. He lays the debate out from the
point of view of the fossil evidence, and then the
genetic evidence and its interpretation are introduced. He does this initially in the modern sense of
sequence-coalescent reconstructions of our most recent common ancestor. The amount and nature of
regional genetic variation in our species, and the
way this affects historical interpretation and reconstruction, are given extended treatment.
The author goes to appropriately great lengths to
support his major assertion, that what genetic data
DOI 10.1002/ajpa.20021
Published online in Wiley InterScience (www.interscience.wiley.
ogeneity in estimated evolutionary parameters, e.g.,
mutation and recombination, and selection across
the genome. And there is evidence from sequence
trees of possible long-standing population subdivision within Africa that was represented in the emigrants. These things could be important.
However, these are the reviewer’s quibbles, because John ends by presenting a “mostly out of Africa” scenario that does not rest delicately on any
one fact, and appropriately stresses demographic
considerations (Fig. 1). His model is based on the
changing roles of gene flow and drift occasioned by
periods of differential growth, perhaps due to human cultural or biological advance. One might wish
to view this compromise between “Killer Apes” and
“Good Neighbors” as a gentlemanly cop-out by someone finding himself in no-man’s land. Only time will
tell whether the one real truth is such a Golden
Proportion. But Relethford’s scenario is an appealing one, based on well-considered reasoning, and literally as this review was being written, Templeton
(2002) suggested a similar view. Analyses may need to
be developed to test specifically whether partial replacement is more plausible than that today’s pattern
is a kind of global projection of African patterns that
pre-existed a true replacement diaspora.
This is the kind of book anthropology students and
professionals should have handy, heavily marked
up, for reminders, lectures, and lessons in tempered
argument. It sets a refreshing standard of measured
dialogue that students should see before being recruited into their advisor’s army. Relethford will
leave you thinking, not swearing. The book is welldocumented and clearly explained, the bibliography
is good, and a “Notes” section gives clear and helpful
explanations of technical, mathematical, or other
fine points from which the reader wanting a smooth
flow of the main text is spared. In a rapidly moving
field the details will age, but the concepts will have
some staying power. Because the cost is outrageous,
Abrahams PH (2001) McMinn’s Interactive Clinical
Anatomy, Version 2.0. St. Louis, MO: Mosby. $99.95
Donovan TM, and Welden CW (2002) Spreadsheet Excercises in Conservation Biology and Landscape
Ecology. Sunderland, MA: Sinauer Associates, Inc.
464 pp. $24.95 (paper).
Donovan TM, and Welden CW (2002) Spreadsheet
Excercises in Ecology and Evolution. Sunderland,
MA: Sinauer Associates, Inc. 556 pp. $24.95 (paper).
Groves C (2001) Primate Taxonomy. Washington, DC:
Smithsonian Institution Press. 350 pp. $65.00 (cloth).
Henneberg M (ed.) (2001) Causes and Effects of Human
Variation. Adelaide, Australia: Australasian Society
for Human Biology. 164 pp. (paper).
Fig. 1. Relethford’s “Mostly Out of Africa” scenario for modern human origins (his Fig. 9.3).
especially without color or slick paper, I suggest that
all you professors buy a copy for your graduate students.
Department of Anthropology
Pennsylvania State University
University Park, Pennsylvania
Templeton A. 2002. Out of Africa again and again. Nature 416:
Weiss K, Maruyama T. 1976. Archeology, population genetics,
and studies of human racial ancestry. Am J Phys Anthropol
44:31– 49.
DOI 10.1002/ajpa.10115
Published online in Wiley InterScience (www.interscience.wiley.
Hoffecker JF (2002) Desolate Landscapes: Ice-Age Settlement in Eastern Europe. New Brunswick, NJ: Rutgers University Press. 298 pp. $32.00 (paper).
Hoppa RD, and Vaupel JW (eds.) (2002) Paleodemography: Age Distributions from Skeletal Samples.
New York: Cambridge University Press. 259 pp.
$80.00 (cloth).
Pietrusewsky M, and Douglas MT (2002) Ban Chiang,
a Prehistoric Village Site in Northeast Thailand, I:
The Human Skeletal Remains. Philadelphia: University of Pennsylvania. 493 pp. $100.00 (cloth).
Prufer OH, Pedde SE, and Meindl RS (eds.) (2001)
Archaic Transitions in Ohio and Kentucky Prehistory. Kent, OH: Kent State University Press. 359 pp.
$38.00 (paper).
Reynolds PA, and Abrahams PH (2001) McMinn’s
Interactive Clinical Anatomy: Head and Neck, Version 2.0. St. Louis, MO: Mosby. $99.95 (CD-ROM).
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mass, extinct, mammalia, taxa, primate, cercopithecid, body, scaling, estimating, extant
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