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Published by the Press Syndicate of the University of Cambridge
The Pitt Building, Trumpington Street, Cambridge CB2 1 RP
40 West 20th Street, New York, NY 10011, USA
10 Stamford Road, Oa.kleigh, Melbourne 3166, Australia

Cl Cambridge University Press 1990

First published 1990

Library of Congress Cataloging-in-Publication Da1a

Body size in mammalian paleobiology : estimation and biological
implications/ edited by John Damuth, Bruce J. MacFadden.
p. cm.
ISBN 0-521-36099-4
1. Mammals - Size - Congresses. 2. Adaptation (Biology) -
Congresses. I. Damuth, John Douglas, 1952- . II. MaeFadden,
Bruce J.
QL739.B63 1990
599 - dc20 87-71308
CIP

British Library Cataloguing in Publication Data

Body size in mammalian paleobiology.
I. Fossil mammals. Size
I. Damuth, John II. MacFadden, Bruce J.
569
ISBN 0-521-36099-4 hardback

Transferred to digital printing 2003

The excerpt on page 49, from The Time Machine, by H. G. Wells,

appears with the permission of A. P. Watt Ltd. on behalf of the
Literary Executors of the Estate of H. G. Wells.

Bahan dengan hak cipta

 Contents

Preface page vu
..
Contributors
IX
1 Introduction: body size and its estimation
JOHN DAMUTH AND BRUCEJ. MACFADDEN 1
2 The physiological significance of body size 11
BRIAN K. MCNAB

3 The behavioral/ ecological significance of body size in 25

the Mammalia
JOHN F. EISENBERG
4 The functional anato.my of body mass 39
THEODOREJ.GRAND
5 Estimating body mass and correlated variables in extinct 49
mammals: travels in the fourth dimension
ROBERT A. MARTIN
6 Evolutionary strategies and body size in a guild 69
of mammals
YJRGJNJA C , MAJQRANA

7 Problems and methods in reconstructing body size in 103

fossil primates
WIJ LIAM J,, JUNGERS

8 Body mass and hindlimb bone cross-sectional and 119

articular dime.nsions in anthropoid primates
CHRISTOPHER RUFF

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 . Contents
VI

9 Insular dwarf elephants: a case study in body mass 151

estimation and ecological inference
Y, LOUISE ROTH

10 Skeletal and dental predictors of body mass in carnivores 181

Bl AIRE YAN YAIKENRJJRQH

11 Problems with using fossil teeth t o estimate body sizes of 207

extinct mammals
MIKAEL FORTELJUS

12 Problems in estimating body masses of archaic ungulates 229

using dental measurements
JOHN DAMUTH

13 Correlation of cranial and dental variables with body size 255

in ungulates and macropodoids
CHRISTINEM. JANIS

14 Postcranial dimensions of ungulates as predictors of 301

body mass
KATHLEEN M. SCOTT

15 Body size estimates and size distribution of ungulate 337.

mammals from the Late Miocene Love Booe Bed
of Florida
BRUCE J. MACFADDEN AND RICHARD C. HULBERT, JR.

16 Appendix: Prediction equations 365

Index 391

Bahan dengan hak cipta

 Preface

Recent years have seen an increased interest among neontologists in

the importance of body size as a major influence on an animal's adap­
tation to its environment and its place in the community. Along with
this, there has been a surge of research by paleobiologists that uses
estimates of body size for fossil species, particularly Cenozoic mammals.
This book is the result of a workshop held in Gainesville, Florida,
at the University of Florida in April 1987. The purpose of the work­
shop was to bring together people with mutual interests in using body
size as a tool in studies of mammalian evolution and paleobiology. We
felt that much complementary research was being done by different in­
vestigators and that it was· time to try to assess the relative reliability
and comparability of body mass estimation techniques for fossil mam­
·mals ► and to do so in the context of some of the applications of body
mass estimates. We restricted our attention to the mammals for two
reasons: Mammalian body si.z e and related topics are curre11tly highly
active areas of research in both neontology and paleontology; and
body mass estimation in other important fossil vertebrate groups, which
have no (or only a few) extant members, necessarily requires differ­
ent kinds of approaches both to estimation and to assessment of
the outcome. In addition to review chapters, we sought primary re­
search results, because we wanted this book to serve both as a prac­
tical introduction to the subject and as an indicator of where the field
is heading.

John Damuth is grateful to the Smithsonian Institution for supporting

him as a Visiting Scientist under the auspices of the Evolution of Ter­
restrial Ecosystems Program, Department of Paleobiology,. 1986-8.
We thank the Florida Museum of Natural History at the University
..
Vil

Materi al chroniony prawem autorskim

 ...
VIJI Preface

of Florida for allowing us to host the Body Size Workshop there in

April 1987.
We also thank Peter-John Leone of Cambridge for his encourage,nent
during the development of this book.

Santa Barbara, California John Da,nuth

Gainesville, Florida Bruce J. MacFadden

Materi al chroniony prawem autorskim

 Contributors

John Damuth Christine M. Janis

Department of Biological Division of Biology and Medicine
Sciences Brown University
University of California Providence, Rhode Island
Santa Barbara, California William L. Jungers
John F. Eisenberg Department of Anatomical
Florida Museum of Natural Sciences
History School of Medicine
University of Florida SUNY at Stony Brook
Gainesville, Florida Stony Brook, New York
Mikael Fortelius Bruce J. Mac Fadden
Department of Geology Florida Museum of Natural
University of Helsinki History
Finland University of Florida
Gainesville, Florida
Theodore I. Grand
Department of Zoological Brian K. McNab
Research Department of Zoology
National Zoological Park University of Florida
Washington, DC Gainesville, Florida
Richard C. Hulbert, Jr. Virginia C. Maiorana
Florida Museum of Natural Department of Ecology and
History Evolution
University of Florida University of Chicago
Gainesville. Florida Chicago, Illinois

IX

Materi al chroniony prawem autorskim

x Contributors

Robert A. Martin Kathleen M. Scott

Department of Biology Department of Biological
Berry College Sciences
Mt. Berry, Georgia Rutgers University
Piscataway, New Jersey
V. Louise Roth
Blaire Van Valkenburgh
Zoology Department
Department of Biology
Duke University
University of California
Durham, North Carolina
Los Angeles, California
Christopher Ruff
Department of Cell Biology and
Anatomy
Johns Hopkins University School
of Medicine
Baltimore, Maryland

Materi al chroniony prawem autorskim

Body size in mammalian paleobiology

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 1
Introduction: body size and its estimation
JOHN DAMUTH AND BRUCE J. MACFADDEN

Body size In biology and paleobiology

In recent years there has been growing interest in the biological impli­
cations of body size in animals. Body mass is correlated with a host of
metabolic and physiological variables; with' ecologically relevant char­
acteristics such as life history traits, diet, population density, population
growth rate, home range size, and behavioral adaptations; and with
larger-scale patterns in community structure and biogeography (Brown
& Maurer 1989; Calder 1984; Damuth 1981, 1987; Eisenberg 1981, this
volume; Emmons, Gautier-Hion, & Dubost 1983; Fleming 1973; Jarman
1974; McMahon & Bonner 1983; McNab, this volume; Peters 1983;
Schmidt-Nielsen 1984).
At the same time, there has been increasing use of body size in a
wide range of applications in vertebrate paleobiology. Studies of func­
tional morphology often employ measurements that must be related to
body size before functional interpretations can be applied to fossil spe­
cies (e.g., Emerson 1985; Gantt 1986; Kay 1975; Thomason 1985). In­
ferences concerning the metabolism and energetics of fossil vertebrates
often depend critically on body mass estimates of fossil species. and
body size has figured prominently in evolutionary explanations of the
evolution of vertebrate metabolic physiology (McNab 1987; Thomas &
Olson 1980; Tracy, Turner, & Huey 1986; Turner & Tracy 1986). The
evolution of body proportions and scaling relationships themselves. and
the resulting functional implications, form an important area of paleo­
biological research (Fortelius 1985, this volume; Gingerich & Smith
In Body Size in M11,,,n,alian Paleobiology: E.stitnalion and Biological ln1plico-
1io11.s, John Damuth and Bruce J. Macfadden, eds.
"'Cambridge University Press 1990.

Bahan dengan hak cipta

2 John Damuth and Bruce J. MacFadden

1985; Gould 1974, 1975; Jerison 1973; Kay 1975; Martin & Harvey 1985;
Radinsky 1978; Scott 1985).
Body size plays a major role in studies of mammalian paleoecology.
In fact, based upon modern knowledge, a mammal's body size may be
the most useful single predictor of that species' adaptations. In addition
to allowing reconstruction of trophic role and habitat preference, re­
lationships between body size and ecological factors help us to make
inferences about many more complex ecological characters of fossil
mammals. For example, interpretations of life history characters and
social behavior have played a part in explanations of the evolution of
horns in ungulates (Janis 1982) and explanations of megafaunal suscep­
tibility to extinction (Kiltie 1984; McDonald 1984). Fauna! and com­
munity structure can be characterized in part by body size range and
distribution, and has been used in studies of community evolution and
energetics, and in the interpretation of climate and ancient vegetation
(Andrews & Nesbit Evans 1979; Andrews, Lord, & Nesbit-Evans 1979;
Collinson & Hooker 1987; Farlow 1976, 1987; Heagle 1978; Janis 1982,
1984; Legendre 1986; Van Couvering 1980; Van Valkenburgh 1985,
1988).
The mammalian fossil record shows many examples of body size
change within lineages, including the sometimes spectacular cases of
gigantism and dwarfism on islands. Explanation of the island trends is
still a challenge (Geist 1987; Hooijer 1967; Marshall & Corruccini 1979;
Maiorana, this volume; Martin, this volume; Roth, this volume; Sondaar
1977, 1987; Thaler 1973).
Because body size is a character exhibited by every species, it poten­
tially bas an important role to play in studies of the tempo and mode
of evolution (e.g., Gingerich 1976; Gould &Eldredge 1977; MacFadden
1987). Cope's rule - that size tends to increase within lineages or taxa
- is one of the most widely discussed general macroevolutionary patterns
(though until recently there were few studies that used actual body-mass
estimates in quantitative studies of the phenomenon) (Gould 1988;
MacFadden 1987; Newell, 1949; Rensch, 1959; Stanley 1973). At least
the Pleistocene and Cretaceous mass extinction events have been notably
size-selective among terrestrial vertebrates, affecting the larger species
more strongly (Martin & Klein 1984; Padian & Clemens 1985). It has
been suggested that frequent "megafaunal" extinctions or periods of
size reduction may be characteristic of fossil vertebrate faunas, and may
happen more locally and on a different time scale than the extinctions
of the proposed long-term (i.e., 26 million years [myr]) global cycle

Materi al com direitos autorais

 Introduction: body size and its estimation 3

(Bakker 1977; Prothero 1985; Raup & Sepkoski 1984; Webb 1984). The
presumed different extinction probabilities (and other ecological differ­
ences) exhibited by large and small vertebrates have implications for
explaining the sorting of species in mammalian clades, but as yet this
has not received much attention (but see Martin, this volume; Stanley
1973).
Finally, it is clear that many taphonomic processes operate upon ver­
tebrate remains in a size-selective or size-dependent way (Behrensmeyer
� Hill 1980; Behrensmeyer, Western & Dechant Boaz 1979; Voorhies
1969). Knowledge of body size can thus be a useful tool in the inter­
pretation of mammalian fossil assemblages (Badgley 1986; Damuth
1982; Wolfe 1975).

Rationale and organization of this book

It may seem from the preceding sketch that body mass, having already
been so widely used, must not be a problematic character to infer for
fossil species. Certainly, size would appear to be one of the most straight­
forward characters that a fossil could exhibit. But the picture of satisfying
accuracy and general compatibility that this widespread use suggests is
misleading. We cannot measure body mass directly for fossil species,
and must derive estimates from skeletal remains that are usually frag­
mentary and incomplete. Some taxa are represented by better fossil
material than are others, and some have no living representatives. Some
groups, such as the primates, have received far more detailed attention
and have been analyzed by more sophisticated techniques than have
· others (e.g., Jungers 1985, this volume; Ruff, this volume). Estimates
for species of different taxa typically derive from different regressions
or other estimator techniques, and are based on different body parts.
Studies vary considerably in the degree of precision they demand of
body mass estimates. Some.require only that relative masses be correct,
some require only species averages, and some require accurate estimates
for particular fossil specimens.
Workers have tended to develop and employ estimation techniques
suited primarily to the study at hand, and intended to yield the level of
accuracy that a particular study requires. Often, a skeletal element well
represented among the fossil species is measured in a group of extant
relatives and regressed upon body mass; a high correlation coefficient
is taken as a sign of success and the regression equation is used to
estimate fossil body masses. Too little attention has been paid in the

Materi al com direitos autorais

4 John Damu1h and Bruce J. MacFadden

past to the choice of the reference sample of modern species, and to

the range of probable error in the resulting estimates. As is shown by
some of the contributions in this volume, regressions involving char­
acters commonly used to estimate body masses of fossil species do not
always produce what would be satisfactory results, even when reapplied
to the extant species upon which the regression is based.
We sought several different kinds of chapters for this book. Some
contributors work primarily with the modern fauna, investigating the
significance of size. Others are here primarily concerned with the tech­
niques of estimating mass in fossil mammals accurately. We decided to
include this range of topics, so that there would be in one volume a
compendium of techniques and basic practical information, and a source
of ideas (including caveats) and signposts to the literature about the
various applications of body size in paleobiological studies. The tables
of Chapter 16 (Appendix) combine in one place, for the reader's con­
venience, the regression equations discussed in the preceding chapters.
We have also included here additional regression equations provided
by some authors, which are based on characters that may not be as
reliable as those that the authors have judged to yield the "best" esti­
mates of body mass for a particular group. Nevertheless, they may be
of some value in cases where the "best" characters are not preserved
for the fossils in question. We warn the reader not to use the equations
of Chapter 16 uncritically, and also not to bypass the reading of the
relevant chapter.

General conclusions

The general theme that emerged at the Gainesville workshop, and that
runs through book, is that body mass estimation and functional mor­
phological interpretation are not separable. The reason is that, in using
data from modern species to derive estimation equations or other means
of estimation, one must choose a set of modem species that exhibit a
similar relationship between body parts and body mass. This requires
identification of broad functional/morphological groupings, which may
or (at least as often) may not faJI within traditional taxonomic lines
(Damuth; Fortelius; Janis; Van Valkenburgh, this volume). As Grand
(this volume) reminds us, "body mass" is a composite character, whose
components differ in animals pursuing different modes of life. The dif­
ferent ways the same anatomical element may be related to overall mass
in different species can to some extent be predicted from functional
considerations.

Copyrighted material
 Introduction: body size and its estin,ation 5

An analogous statement can be made for interpretations: Dividing

the modern fauna into functional groups improves predictive power of
body mass for both physiological and ecological variables (Eisenberg;
McNab, this volume). In particular, diet is related to much of the varia­
tion found in both kinds of variables.

Practical conclusions and caveats

A number of general practical conclusions, recommendations, and cau­

tions concerning body mass estimation in fossil mammals can be ab­
stracted from the conference proceedings and the papers published here:
1. Estimates based upon certain limb measurements, if available,
appear to be substantially more reliable than those based on
cranial or dental measurements. However, much more com­
parative work needs to be done on the scaling of postcranial
elements in modern forms before we can use limb measurements
for most groups. Proximal limb elements are more reliable cor­
relates of body mass than are distal elements in ungulates and
primates (Ruff; Scott, this volume). The problem with using
fossil limb elements, however, is that they first must be tax­
onomically assignable.
2. Because of their identifiability and preservability, teeth will con­
tinue to be of importance in body mass estimation. However,
dental measures alone, even when restricted to functionally sim­
ilar groups, may not be accurate enough for all purposes. Per­
cent standard errors below 30 are rare in the dental regressions
reported here.
3. In dealing with taxa with high ontogenetic dental variation (e.g.,
• high-crowned horses), the worker must carefully consider which
wear stage(s) should be used to predict body mass. Such vari­
ation differs among taxa and among different dental measures
and among different teeth within the same species.
4. For ungulates, tooth length measurements are generally better
predictors than widths or areas, as the latter vary more with
diet.
5. Techniques using more than one morphological variable (e.g.,
multiple regression) can increase accuracy of predictions when
feasible and appropriate. Particularly, combinations of dental
and postcranial measures, including body length, can result in
relatively accurate estimates for well-represented species (per­
cent standard error < 30).

Materyales na naka copyright

6 John Damuth and Bruce J. MacFadden

6. As in any other statistical prediction procedure, it is unwise to

extrapolate regression predictions beyond the range of the avail­
able data for modern forms. There is no guarantee that rela­
tionships holding within this range will be applicable to smaller
or larger forms (e.g., MacFadden & Hulbert, this volume). In
such cases use of a more general regression (such as "all un­
gulate" or "all mammal" regressions), with the resulting loss of
precision, may be preferable. This consideration adds particular
uncertainty to our estimates of the very largest fossil species,
which may lie beyond the size range of any living mammal; our
sample of living megafauna is also highly restricted and com­
posed of species exhibiting only a few body forms.
7. The use of general regression equations, even those with small
standard errors, does not guarantee accuracy for every species.
In particular, care should be taken to recognize fossil species
that may be aberrant in one or more characters that for most
species might yield good estimates (e.g., the enlarged third mo­
lars of suids, or the enlarged second molars of amynodonts).
8. Estimates remain only estimates. There are certainly numerous
unrecognized sources of error for fossil species. The statistical
errors reported here for regressions on living forms are under­
estimates of the actual inaccuracy in estimates for extinct spe­
cies. To the extent that fossil species deviate from the average
of the modem population used, inaccuracy will increase.

Future research directions

Some directions for future research in mammalian body-mass estimation
are also evident. The need for broader investigation of the scaling of
postcrania has already been mentioned. Craniodental and postcranial
scaling relationships analogous to those presented here are unknown
for many extant mammalian groups. These include most rodent taxa,
insectivores, small marsupials, and various "oddball" groups such as the
edentates. We need further refinement of functional groups among all
of the mammals, as this could dramatically increase the accuracy of our
predictions. Finally, there are extinct groups, such as the North Amer­
ican Merycoidodontidae (oreodonts), that exhibit what appear to be
unique body proportions and craniodental relationships, but for which
we have complete material for some members. Using the well­
represented species, reliable body-mass estimates may be obtainable,
and from these species correction factors could be derived for use on

Materyales na naka copyright

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