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 • •
1ze 1
0
Estimation and
Biological Implications

�· •··.....
•,
.

•I ,
}
... .

Edited by
John Domuth and
Oruce J. MocFodden
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

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 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

Copyrighted material
 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

 . Introduction: body size and its estimation 7

the more fragmentary material representing the other members of the

group.
In summary, we are just beginning to develop the empirical base and
the analytical tools necessary for reliable reconstruction of the body
masses of fossil species. Further refinement of body mass estimation
techniques will make possible the reconstruction of a wide range of
biological aspects of fossil species and faunas, and an exciting new di­
mension of time can be added to ecological and evolutionary studies
that heretofore have had to rely primarily on patterns observed in the
extant biota.

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 Introduction: body size and its estimation 9

Jungers, W. L. (ed.) ..1985. Size and Scaling in Primate Biology. New York:
Plenum Press.
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J. Plrys. Antlrropol. 43: 195-216.
Kiltie, R. A. 1984. Seasonality, gestation time, and large ma.mmal extinctions.
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MacFadden, B. J. 1987. Fossil horses from "Eohippus" (Hyracotherium) to
Equus: scaling, Cope's law, and the evolution of body size. Paleobiology
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McDonald, J. N. 1984. The reordered North American selection regime and
Late Quaternary megafaunal extinctions. In Quaternary Extinctions: A Pre•
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University of Arizona Press.
McMahon, T. A., & Bonner, J. T. 1983. On Size and Life. Scientific American
Library. New York: W. H. Freeman.
McNab, B. K. 1978. The evolution of endothermy in the phylogeny of mammals.
Am. Nat. //2:1-21.
Marshall, L. 0., & Corruccini, R. S. 1979. Variability, evolutionary rates, and
allometry in dwarfing lineages. Paleobiology 4:101-119.
Martin, R. D., & Harvey, P. H. 1985. Brain size allometry: ontogeny and
phylogeny. In Size and Scaling in Primate Biology, ed. W. L. Jungers,
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Padian, K., & Clemens, W. A. 1985. Terrestrial vertebrate diversity: episodes
and insights. In Phanerozoic Diversity Patterns: Profiles in Macroevolution,
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Radinsky, L. 1978. Evolution of brain size in carnivores and ungulates. Am.
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Schmidt-Nielsen, K. 1984. Scaling: Why l s A,iimal Size So Important? Cam­
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Marep1-1an. 3aU11-1U1eHHb1 � asropcK1-1M npasoM

 2
The physiological significance of body. size
BRIA.N K•. MCNAB

All organisms are defined in terms of a distinctive combination of char­

acters, including those associated with anatomy, physiology, behavior,
and reproduction. Fundamental to the definition of a species is its body
size, food habits, and thermal biology, which interact to form the basis
for derived characters like its distribution. If such patterns can be spec­
ified in well-known living species, they can be applied to species about
which our knowledge is unavoidably fragmentary - namely, to rare and
extinct species. The view advocated here is that studies on the biology
of living mammals have generated a series of size-rel�ted relationships
for physiology and reproduction that can be applied (with care) to eco­
logically similar fossil species to suggest ecologically relevant aspects of
their physiology and reproduction. As an example, Lillegraven, Thomp­
son, McNab, and Patton (1987) use size-related aspects of physiology
in a reconstruction of the origin of eutherian mammals.
The principal impediment to the application of these relationships is
that they invariably are fitted to idiosyncratic sets of data derived from
living species, and therefore contain much residual variation. More often
than not, this variation is "swept under the rug" in the unfortunate
belief that it represents an "error" term. In fact, a detailed analysis
generally shows that much of this residual variation contains biologically
important variation (see, for example, McNab 1988a). This variation is
ignored at the peril of imprecision in predicting the characteristics of
living and extinct species. Consequently, the more that is.known about

In Body Size in Mam,nalian Paleobiology: Estimation and Biological Implica­

tions, John Damuth and Bruce J. MacFadden, eds.
o Cambridge University Press 1990.

11

Copyrighted material
12 Brian K. McNab

the biology of fossil mammals, the more that can be inferred about the
size-related aspects of their physiolagy, behavior, and reproduction.
Most size-related patterns seen in living species are complex; they are
most robust when derived from a phylogenetically, morphologically,
and ecologically uniform array of species. Some limitations in the ability
to analyze size relationships, then, are potentially present if the pattern
is derived from a biased sample of the living fauna, or if the fauna is a
biased sample of those capable of existing. This difficulty is at the heart
of the controversy as to whether dinosaurs were, or were not, endother­
mic (see Thomas & Olson 1980). Our judgments on this question are
directly or indirectly based on an analysis of the energetics of mammals,
which may not provide an adequate basis for judging the physiology of
dinosaurs: No mammals are known to be true inertial homoiotherms,
which may have been the case in many large dinosaurs (McNab &
Auffenberg 1976; Reid 1987).
The plan of this article is (1) to summarize the information derived
from extant species of mammals on size-related aspects of physiology
and reproduction, and (2) to caution against the blind use of these
relations, being mindful of the many factors other than size that con­
tribute to physiology and reproduction.

The influence of body size on the physiology and reproduction

of mammals

Many studies since the 1960s and 1970s have demonstrated the quan­
titative correlation of physiological functions with body size, which is
usually defined in terms of body mass, expressed in either grams or
kilograms. These studies have been summarized by Peters (1983), Calder
(1984), and Schmidt-Nielsen (1984).
Because the interest here is in the application of various physiological
functions to fossil mammals, I shall summarize those physiological func­
tions that are most likely to have a consequence for the ecology, dis­
tribution, and evolution of mammals, the subjects of greatest interest
to paleontologists. These functions will be grouped into the headings of
(1) metabolism, (2) temperature regulation, (3) locomotion, (4) inges­
tion, (5) respiration, (6) water balance, (7) reproduction and growth,
and (8) production. Some appropriate equations for these relations are
summarized in Table 2.1. All equations for these relations have a power
form, in part following the original suggestion of Huxley (1932) and in
part owing to its mathematical simplicity; other mathematical models,

Copyrighted material
Table 2.1. Selected scaling functions for physiological variables
in mammals

System and factor Equa1ions• Reference

Metabolism
Basal rate
All mammals M(J/h) = 86.9 m0· ,.. Hays.�en & Lacy 1985
All mammals M (J/h) = 69.3 m0·" McNab 1988a
Mar5upials M(J/h) =
50.0 mo. " McNab 1988a
Euthcrians M(J/h) = 70.9 m"·n McNab 1988a
Field expenditures
Marsupials M(J /h) = 491.8 m"s,. Nagy 1987
Euthcrians M(J/h) = 140.0 m"·"' Nagy 1987

Temperature regulation
Minimal basal rate for
continu.ous endothermy M (J/h) = 312.0 m 0
·" McNab 1983

Locomotion
All terrestrial animals v,.., (km/hr)= 2.70 m"·34 Bonner 1965
M(Jib) = 0.026 m""" ·
V + 0.048 m
0.7'1
Taylor et al. 1982

Ingestion
Carnivores M (Jib) =330.3 m":"' Farlow 1976
Herbivores M(J /h) =264.6 rn°·" Farlow 1976
Herbivores M(J/h) = 247.9 m0'' Nagy 1987

Respiration
Frequency /(I/min)= 322.1 m- 0,,. Stahl 1967

Water balance
Water intake Wi (ml/h) =· 0.010 m0 "" Adolph 1943
Evaporative water loss EWL (ml/h) = 0.0043
n,o.86 Altman & Dittmer 1968
Urinc:plasma concentration
ratio CIC
• p -
- 24.7 m-o.,o Calder & Braun 1983

Reproduction
Gestation period OP (d) =
12.0 m ••
o.
Millar 1981
Neonatal mass NM (g) = 0.045 m0 .. Millar 1981
Litter size I = 24.1 m-0-"' Millar 1981
Growth
Growth rate GR (gld) m 0.04 mo. "' Mil.lar 1977
Growth ra1.e GR (g/d) a 0.04 mo. 71 Case 1978
Growth constant K (I/d) = 0.126 m0·"' Zullinger et al. J 984

Production
Population PIBiomass (Jlglyr) =
24180 m-o.2, Farlow 1976
Homoiotherms rM.. (lid) = 0.040 m_,,_,, Fenchel 1974
Organisms r,.., (lid) = 0.025 m-11 ·"' Bluewciss cl al. 1978

'Body mass (m) is in grams; velocity (v) is in km/h.

13
Copyrighted material
14 Brian K. McNab

namely polynomials, are difficult to interpret biologically, even though

they may be more "accurate" in describing the data. These relations
are discussed here in relevant detail; some functions are included be­
cause they have an ob�ious application to paleontology, whereas others
are included because of their importance to physiological ecology.

Metabolism

The function having the broadest implications probably is the association

of rate of energy expenditure with body size. This relation was first
described for basal rate of metabolism by Kleiber (1932), and by many
others since then. The difficulty with any description of rate of metab­
olism as a function of body si.ze in endotherms is that it complexly
depends on ambient temperature, so that several scaling relationships
can be described, each with its own implications and mathematical char­
acteristics. In theory, the basal rate is the lowest rate of metabolism in
an endotherm that is compatible with temperature regulation (for an
exception, see McNab 1988b), and therefore represents a minimal ex­
penditure of energy. The advantage of using the basal rate is that it is
comparable in all species (McNab 1989), not that it directly estimates
field energy expenditures. The most recent estimates of the scaling re­
lation for basal rate in mammals have been made by Hayssen and Lacy
(1985) and McNab (1988a). The difficulty with these scaling relations,
and all others, is that they incorporate the influence of various factors
other than body mass on basal rate (see last section), even though they
are not explicitly acknowledged in the equations.
Most early descriptions of the scaling of basal rate dwelled on the
power of the metabolism function. The power for total basal rate was
usually thought to fall between 0.67 and 0.75. McMahon (1973) tried
to derive 0.75 from physical principles, and Heusner (1982) argued that
the "true" intraspecific power is 0.67, interspecific powers (presumably)
having no particular biological significance. Comparative biologists,
however, are particularly interested in interspecific comparisons. Hays­
sen and Lacy (1985) found that the total basal rate was proportional to
mass raised to the 0.69 power in a sample of 297 species, and McNab
(1988a) found that it was proportional to mass raised to the 0.71 power
in a sample of 321 species. A detailed analysis of this relation suggests
that its power does not simply reflect the influence of body mass (McNab
1988a). As is to be expected, these scaling relations reflect the sample
•

The physiological significance of body size 15

•

of species used to construct the relationship, so that caution should

prevail when one concludes that a particular relation represents the
"true" relation between two variables, here basal rate and body mass,
independent of sample composition.
Although the basis of the relationship between basal rate and body
mass is unclear, its consequences are obvious: Large mammals have
higher total rates of metabolism than small species. The increase in rate
associated with an increase in mass is not proportional to mass, but to
mass raised to a power less than 1.0. Using the power 0.71, a doubling
of mass leads to a 64% increase in basal rate, and a halving of mass
leads to a 39% decrease in total basal rate. So, contrary to a widely
held belief, even among some physiologists, an increase in body size
requires an increase in the rate of energy expenditure, and in· the amount
of food and energy available in the environment. In other words, an
increase in mass does not reduce energy expenditure. A mammal can
respond to a shortage in available energy by a reduction in mass or by
a reduction in rate of metabolism at a fixed mass.
More is probably known about the residual variation in the basal rate­
mass function than for any other scaled relationship. At a given mass,
basal rate varies by a factor of 3: 1 to 10: 1 (McNab 1988a). Much of this ,
variation is related to food habits. Mammals that specialize on grass and
on vertebrates tend to have basal rates that are two or more times those
of species of the same body mass that specialize on fruits, the leaves of
woody plants, and invertebrates (McNab 1986). The correlation of basal
rate with food habits is accentuated at masses greater than a kilogram,
presumably because energy and nutrient demand, as noted, increase
with mass. So, as mass increases, the differential liabilities associated
with the various foods, such as seasonal availability, digestibility, and
toxicity, are increasingly encountered by . the consuming mammal.
Additional factors that contribute to the residual variation
.
around the
basal metabolism-body mass curve are activity level (and muscle mass),
temperature regulation, and climate. For example, fruit and leaf eaters
have especially low basal rates if these mamm�ls have arboreal habits
(McNab 1986). This association occurs because arboreal species, at least
at masses greater than a kilogram, are often characterized by small
muscle masses. Basal rate is proportional to the fraction of total mass
that is muscle (McNab 1978b). SmaU mammals that go into daily or
seasonal torpor have lower basal rates than similarly sized species that
do not enter torpor (McNab 1983, 1988a). Species living-in cold climates

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16 Brian K. McNab

tend to have higher basal rates than those living in warm climates, but
this correlation may partially reflect the distribution of foods with respect
to climate.
The principal difficulty with using a basal-rate function to estimate
rates of energy expenditure in living or fossil mammals is that most
mammals spend little time in a standard state. Basal rates may be a
convenient index to mammalian energy expenditures, but they do not
represent those expenditures directly. Recent improvements in the abil­
ity to measure directly energy expenditures in the field, especially by
means of doubly labeled water, permitted the development of scaling
relations for field expenditures in mammals (see Nagy [1987] for a sum­
mary). These relations differ from basal-rate functions in both intercept
and power, indicating that eutherian field expenditures are approxi­
mately double those of basal rate. The difficulty with scaling field es­
timates is that they incorporate the multiplicity of factors operating at
the time the measurements were made, including season, activity level,
reproduction, etc. This makes field measurements very difficult to in­
terpret, as their scaling functions surely do not reflect the influence of
body mass alone. At present, the best analysis of energy expenditure
derives from combining field measurements with laboratory measure­
ments of rate of metabolism at ecologically relevant temperatures
(McNab 1989).

Temperature regulation
Although temperature regulation is often considered in qualitative terms
(i.e., a particular species regulates body temperature or it does not),
McNab (1983) demonstrated that a unique relationship exists between
the basal rate of metabolism and body mass in those species character­
ized by continuous endothermic temperature regulation. In this rela­
tion, total basal rate is proportional to mass raised (approximately) to
the 0.33 power. All mam.mals having basal rates that fall below this re­
lation enter daily torpor, at least upon occasion. One consequence
of this relation is that all small marsupials enter torpor in association
with their low rates of metabolism. From the viewpoint of paleontol­
ogy, this relation may be difficult to apply, unless one can establish
that some factor sets basal rate in a species so low that it falls below
this scaling relation and, therefore, that the species is likely to enter
daily torpor.

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 Tht physiological significance of body sizt 17

Locomotion
All mammals move from one place to another and search for food and
mates. Some have specialized behaviors, such as digging shelters and
long-distance migration. Each of these activities requires a n expenditure
of energy. Extensive measurements in mammals have demonstrated that
the cost of terrestrial locomotion increases with body mass and the
velocity of movement (see Taylor, Heglund, & Maloiy [1982] for a
summary). Body mass in this relation is raised (approximately) to the
0.69 power. Because large mammals can run faster than small mammals
(Bonner 1965), they may save time by moving a distance rapidly, but
they do not conserve energy by doing so.

Ingestion
The intake of food can be measured directly, or estimated from the
ene rgy and nutrient requirements of species, when the assimilation
efficiency is known. Farlow (1976), summarizing food intake for car­
nivorous and herbivorous mammals, showed that total intake is pro­
portional to mass raised to the 0. 70 and 0.73 powers, respectively. These
equations are similar to those proposed by Nagy (1987) for field ex­
penditures. They are approximately four times the basal rate; that is,
assimilation efficiency is about 50%.

Respiration

Relatively little use has been made in paleontology of the correlations

of respiration with body mass. One use was in explaining the evolution
of the secondary palate in the phylogenies of the mammals and the
bauriamorph therapsids. The development of the secondary palate was
associated with a marked decrease in body mass, a shift from ectothermy
to endothermy, and a consequent increase in respiratory frequency
(McNab 1978a). Respiratory frequency is proportional to mass raised
to the - 0.26 power (Stahl 1967).

Water balance
Total water intake and evaporative water loss are proportional to mass
raised to powers that range from 0.86 to 0.88, which means that large

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18 Brian K. McNab

species require more water than small species and that small species
need more water on a mass-specific basis. The capacity of mammals to
produce a concentrated urine relative to the plasma, in contrast, scales
proportionally to mass raised to the -0.10 power (Calder & Braun
1983). This function means that small species are more parsimonious
with water than are large species, which may be necessary because of
the greater rate of water turnover associated with a small mass.

Reproduction and growth

These processes are essential for survival of every species, and both
vary with body mass. Reproduction bears a complex relationship to
mass. Like other time functions (Lindstedt & Calder 1981). gestation
period and the time from birth to sexual maturity increase with mass
raised approximately to the 0.25 power, whereas neonate mass varies
with maternal mass raised to the 0.89 power, and litter size is propor­
tional to mass raised to the -0.30 power. Total growth rates (in grams
per day, g/day) are proportional to mass raised to the 0.69 power (Millar
1977), or to the 0.72 power (Case 1978). Growth, measured in terms
of a Gompertzian exponential-growth rate constant, is proportional to
mass raised to the -0.30 power (Zullinger, Ricklefs, Redford, & Mace,
1984), which if converted to growth in grams per day, bas a 0.70 power,
a value similar to those described by Millar and Case.

Production
Net population production is proportional to population metabolism
(Humphreys 1979; Lavigne 1982; McNeil & Lawton 1970), which means
that production should scale proportionally to mass raised to the 0. 75
power, which is the case (Farlow 1976; Lavigne 1982). As a consequence,
production per unit biomass should be proportional to mass raised to
the -0.25 ( = m075/m'00) power. In fact, Fenchel (1974) and Blueweiss
et al. (1978) showed that the maximal growth constant of populations,
'm••• which is the ultimate measure of effective production by a popu­
lation, is estimated to be proportional to -0.27 and -0.26, respectively.
That the concept of production can be applied to fossil faunas was
demonstrated by Farlow (1976), who examined living mammalian com­
munities to evaluate the possibility that large herbivorous dinosaurs had
levels of production representative of endotherms.

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 The physiological significance of body size 19

Scaling basal rate in mammals: a cautionary tale

All mass functions, as noted, are calculated from data collected on a
sample of living species. The powers and intercepts of these relations
therefore vary as the species composition of the sample varies (McNab
1988a), a condition that has led to differing interpretations. For example,
various authors (Glazier 1985; Hennemann 1983; McNab 1980; Schmitz
& Lavigne 1984) have argued that much of the residual variation in the
reproduction and growth of mammals is related to ecologically relevant
variations in (basal) rate of metabolism. Others (Hayssen 1984; Robin­
son & Redford 1986) have denied this correlation. The correctness or
error in this suggestion, however, is not as relevant here as is the re­
alization that all scaling relationships are approximations at best and
incorporate many factors other than body size. So these relations should
be viewed as approximations, not as physical or biological "laws." The
complexity of scaling relations can best be demonstrated in the basal
rate of metabolism in mammals.
The factors setting basal rate in mammals have be�n under dispute
recently, and this dispute has significance for the interpretation of scaling
functions generally. McNab (1980, 1986) argued that food habits in­
dependent of taxonomic affiliation are an important factor influencing
the basal rate of mammals, whereas Hayssen and Lacy (1985) and Elgar
and Harvey (1987) maintained that most of the residual variation is
related to taxonomic affiliation.
The comparative contributions of body mass, food habits, activity
level, climate, and taxonomic affiliation to basal rate are difficult to
separate because these factors are often correlated with one another.
Some of these complexities have been explored by McNab (1988a). For
example, if all 321 species of mammals for which reasonable estimates
of basal rate exist are pooled, a curve similar to the Kleiber curve is
obtained (Table 2.2). If, however, the curve is restricted to the 272
eutherians, a parallel, but slightly higher curve is obtained. These curves
are similar because 85% of the species belonging to the inclusive curv�
are eutherians. In contrast, a curve consisting only of the 46 marsupials
available is significantly lower than the eutherian curve. Many biologists,
including MacMillen and Nelson (1969) and Dawson and Hulbert (1970),
have concluded that marsupials have lower basal rates than eutherians.
But that conclusion is more complicated than appears at first glance.
All small marsupials go into torpor, but most small eutherians do not.
Marsupials that do not enter torpor have a scaling relation for basal

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20 Brian K. McNab

Table 2.2. Basal rate of metabolism of mammals

Condition Species (n) Equations•

All mammals 321 M(Jlh) - 69.3m0·713

Marsupials
All 46 M(J/h) = 50.0m0 73 7
No torpor 32 M(J/h) = 67.0m0 .,.
Euthtrians
All 272 M(J/h) = 70.9m0•716
No torpor 212 M(J/h) = 76.7m0•111
No invertebrate-eating specialists 219 M(J/h) = 65.1m ·"1
0

•Body mass (m) is in grams. All equations from McNab (1988a).

rate similar to that found in eutherians that do not enter torpor (Table
2.2). Thus, some of the difference in basal rate found between eutherians
and marsupials is related to the differential occurrence of torpor in these
two groups: The scaling relationship of basal rate to mass includes a
correlation of torpor with body mass and a correlation of torpor with
basal rate. Furthermore, if one eliminates all invertebrate-eating spe­
cialists from the eutherian curve, the remaining eutherians have a lower,
but steeper, curve than do all eutherians. This change occurs because
a disproportionate number of s.mall, invertebrate-eating mammals at
small size were excluded from the curve. That is, invertebrate-eating is
related to body size and to basal rate. I conclude that all factors that
are related to body mass and to basal rate are incorporated into the
basal-rate-body-mass function, unless steps are taken explicitly to pre­
vent the inclusion of these factors, which is unlikely to occur as long as
the data are grouped by taxa. This conclusion applies to all scaling
relations.

Summary
Paleontologists can greatly profit in the analysis of the biology of extinct
faunas by using the extensive set of correlations that have been shown
in living animals between various aspects of physiology and body mass.
These so-called "scaling" functions limit the range of possible physio­
logical configurations found in organisms. This range is further reduced

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 The physiological significance of body size 21

if additional knowledge about the organisms is available, including food

habits, activity level, climate, and phylogeny. The more information
that is known about an extinct species, the more one is able to specify
its physiology and behavior. Nevertheless, knowledge of body size alone
is sufficient to reduce significantly the range of possible combinations
of physiology and behavior found in a species.
Of course, the observations of paleontologists have potential value
for physiological ecologists. Thus, observations on the body size and
bone structure of dinosaurs have made major contributions to the debate
on the physiology of dinosaurs (Reid 1987; Thomas & Olson 1980).
Detailed descriptions of the reduction in body mass and of the concom­
itant evolution of the secondary palate in cynodont and bauriamorph
therapsids have contributed to our understanding of the evolution of
endothermy in the early phylogeny of mammals (McNab 1978a). These
observations demonstrate that some physiological combinations used in
the past are not used today, a conclusion that makes a significant con­
tribution to our understanding of the potentiality for physiological ad­
aptation to the environment. The morphological observations made by
paleontologists that will contribute further to our understanding of the
evolution of physiological function are difficult to predict. Without
doubt, however, they will include the concept of body size.

References
Adolf, E. F. 1943. Physiological Regulations. Lancaster, Penn.: Cattell Press.
Altman, P. L., & Dittmer, D. S. 1968. Metabolism. Bethesda, Md.: Federation
of American Societies of Experimental Biology.
Blueweiss, L., Fox, H., Kudzma, V., Nakashima, D., Peters, R., & Sams, S.
1978. Relationships between body size and some life history parameters.
Oecologia 37:257-272.
Bonner, J. T. 1965. Size and Cycle: An Essay on the Structure of Biology.
Princeton, N.J.: Princeton University Press.
Calder, W. A. 1984. Size, Function and Life History. Cambridge, Mass.: Har­
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and birds. Am. J. Physiol. 244:R60t-R606.
Case, T. J. 1978. On the evolution and adaptive significance of postnatal growth
rates in the terrestrial vertebrates. Q. Rev. Biol. 53:243-282.
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22 Brian K. McNab

Farlow, J. 0. 1976. A consideration of the trophic dynamics of a Late Cretaceous

large-dinosaur community (Oldman formation). Ecology 57:841-857.
Fenchel, T. 1974. Intrinsic rate of natural increase: the relationship with body
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Hennemann, W. W. 1983. Relationship among body mass, metabolic rate and
the intrinsic rate of natural increase in mammals. Oecologia 56:104-108.
Heusner, A. A. 1982. Energy metabolism and body size. I. Is the 0.75 mass
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 The physiological significance of body size 23
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 3
The behavioral/ecological significance of
body size in the Mammalia
JOHN F. EISENBERG

In 1984, Calder published the comprehensive review Size, Function, and

Life History, which attempts to document the overwhelming importance
of body size in determining the form and function of a species (individ­
ual) within an ecosystem. I recommend this book to anyone interested
in problems of biological scaling.
Given a mammal (endothermic and sometimes homoiothermic), Brian
McNab (this volume) has eloquently summarized the physiological
consequences of becoming large or ,remaining small. There is an
implied directionality in that statement which derives from the fact
that I am addressing paleontologists. Most researchers will concede
that in the evolutionary history of mammals, the primitive condi­
tion was a small body size; larger body size was often a derived char­
acter within any of the separate lineages. As McNab has pointed
out, when various physiological and behavioral values are plotted
against body size in a double common-log plot, it is possible to per­
form a regression analysis and specify the form of the curve in terms
of an allometric equation. He has also pointed out, and I wish to
emphasize, that the best-fit regression line often has a great many scat­
tered points around it, and an explanation of the residual values can
be extremely enlightening to a biologist. In fact, most of the scat­
ter can be accounted for by considering either phylogenetic affinity
or trophic adaptation. This will become evident as I proceed with my
discussion.

In Body Sizt in Mamm41ian Paltobiology: Estimation and Biological Implica­

tions, John Damulh and Bruce J. MacFadden, eds.
o Cambridge University Press 1990.

25

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26 John F. Eisenberg

Mammalian body size and niche

Some years ago, I began to analyze the size of mammals and study the
distribution of size classes within certain niche categories. I defined
"niche" according to the substrate used, such as aquatic, semiaquatic,
terrestrial, arboreal, etc., and the trophic strategy as folivore, grazer,
frugivore, etc. It was then possible to see the manner in which niches
were distributed over different geographic areas of the globe (Eisenberg
1981). Predictably, certain niche categories were absent from the tem­
perate zone and present in the tropics, and this further led me to con­
centrate on two tropical areas that were comparable in geographic
position and size: Panama and Malaya. When the frequency of size
classes were plotted for the various trophic categories, a remarkable
similarity could be demonstrated. Of course, there were some differ­
ences: The megafauna of Malaya is still intact, but Late Pleistocene
extinctions in Panama have eliminated some of the larger forms (Webb
1985). The distribution of size classes over trophic categories might even
be more similar if recently extinct forms were included in the tabulations.
This result suggests that specialjzation for a certain niche sets certain
constraints on body size, and that Darlington's principle of comple­
mentarity is a reality (Eisenberg 1981; see Figure 3.1).
Since I am addressing paleontologists, another way of looking at the
problem is to study communities from different parts of the globe
through time. Here paleontological data become important. For ex­
ample, when one looks at mammalian predatory guilds through time
and compares relative body proportions and size, one can discern some
remarkable convergences in community structure. This was demon­
strated by Van Valkenburgh (1984, 1985) when the carnivore guild of
the Serengeti was compared with a carnivore guild in North America
during the Pliocene. In a similar fashion, Larry Martin (1990) has been
able to demonstrate that even when carnivore assemblages become ex­
tinct, the replacement forms are remarkably similar to their predecessors
in size and body proportion. This again implies that there are some
general rules governing form and function as well as community structure
and dynamics, and that, given an appropriate range of mammalian
stocks, selection will inevitably tend to produce similar forms of sec­
ondary consumers within those communities that are comparable in
vegetative cover, primary productivity, and primary consumers.
A similar case could be made for the replacement of mammalian
primary consumers such as the extinctions at the end of the Eocene,

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 Significance of body size in mammalian ecology 21

leading to the loss of the Uintatheriidae and subsequent replacement

by the Brontotheriidae in Nort.h America (Osborn 1929).

Body size, trophic strategy, and population dynamics

In an attempt to unravel some rules or patterns of mammalian com­
munity structure, my colleagues and I have published a series of papers
concerning body size, diet, population biomass, and the rate of repro­
duction (Eisenberg 1980; Eisenberg, O'Connell, & August 1979; Kin­
naird & Eisenberg in press; Robinson & Redford 1986a,b). We chose
Neotropical communities for analysis because we had a great deal of
direct experience in the area; and, furthermore, there was a pressing
need to develop management plans for some species that required fun­
damental information concerning tropical community structure and
function. I will attempt to summarize briefly several general rules that
we uncovered. Much of the following information is included in Eisen­
berg et al. (1979) and Eisenberg (1980, 1981). In all communities, her­
bivores - in particular, browsers and grazers - collectively comprise the
largest proportion of the mammalian biomass within a community. In
short, primary consumers, or those feeding directly upon primary pro­
ductivity, are the standing-crop biomass dominants. Secondary and ter­
tiary consumers not only exist at much lower densities but contribute a
smaller proportion to the total mammalian biomass in a community.
These results are broadly consistent with the global. analysis presented
by Damuth (1987).
On the other hand, the smaller forms, although not comprising a
significant portion of the standing-crop biomass, are often the key forms
in terms of annual productivity and may account for the major con­
sumption of the primary producers. Eisenberg et al. (1979) demon­
strated that in the llanos of Venezuela the cane rat Zygodontomys
brevicauda (at 35 g) was not a biomass dominant, but in terms of annual
production had a very significant impact on the community.
A large pQrtion of the standing-crop biomass of a closed-canopy,
multistratal, tropical evergreen forest tends to be comprised of arboreal
species as opposed t o terrestrial, because of the reduced primary pro­
ductivity on the shaded f?rest floor of a mature rain forest. As one
proceeds through a series of communities showing reduced vegetational
cover, a more open canopy, or an earlier form of plant growth (second
growth forest), the proportion of community biomass contributed by
terrestrial forms increases until, in an almost treeless open savanna, the

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28 John F. Eisenberg

MALAYA

44 Fr/Gr

11 ••• • 15 Corn

• ■ I• 12 Fr/Br

II I ■ 14 Fr/Om

•■ I •• I 12 Br/Gr

,t • • ·J■ I • II In/Om

■ • 11 ■ 7 Plsc

• Myrm

2 3 4 5
Loo1o Body Wei91'1t (Gm)

Figure 3. l. Frequency of size classes according to trophic strategy: Malaya (A)

compared with Panama (B. opposite page). Log10 body weight (g) (abscissa) is
plotted against frequency (ordinate). The number of genera within each trophic
class is given to the right of each size series. Fr/Or = frugivore/granivore; Fri
Om = frugivore/omnivore; In/Om = insectivore/omnivore; Fr/Br = frugi­
vore/browser; Br/Gr = browser/grazer; Carn = carnivore; Pisc = piscivore;
Myrm = myrmecophage. (Data in part based on Handley 1966 and Medway
1978.)

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 Significance of body size in mammalian ecology 29

PANAMA

-■ 28 Fr/Gr

•• •• 20 Fr/Om

II a• I. • 12 In/Om

I I • 10 Fr/Br

• • •••• ■ 7 Br/Gr

•• •• 7 Corn

• • 2 Plsc

• ■■ ■ 4 Myrm

3 4 5 6
Loo10 Body W.iOht CGml

terrestrial components dominate completely over alt other locomotor

categories (scansorial or arboreal). This is intuitively obvious, but is
nevertheless an important factor to bear in mind when trying to recon­
struct extinct communities (Eisenberg and Seidensticker 1976).
The proportion of total standing-crop biomass contributed by differ­
ent size classes bears the following relationship: If one plots average

Copyrighted material
30 John F. Eisenberg

standing-crop biomass for each species against its average weight class,
the relationship is positive. That is to say, larger forms contribute more
to total standing-crop biomass than do smaller species (Kinnaird and
Eisenberg in press). However, there is considerable scatter around the
regression line, and in an attempt to account for the scatter, multiple
regression analyses were performed. More than 70% of the variation
can be accounted for by trophic strategy. For example, within any given
size class, herbivores contribute more to standing-crop biomass than do
secondary and tertiary consumers.
Population densities of a species are predictably related to their size
and trophic specialization. Again, a plot of population densities against
mean body size demonstrates a negative relationship. Smal1er forms can
exist at higher densities than larger ones. Again there is a great deal of
scatter around the regression line. Almost all of this scatter can be
accounted for when the trophic specializations are plotted separately.
Herbivores within a given size class exist at higher densities than sec­
ondary or tertiary consumers (Robinson and Redford 1986a).
Although a species may be a biomass dominant, it does not necessarily
follow that this species shows a high productivity value. Indeed, the
intrinsic rate of population increase covaries negatively with body mass.
This trend does not necessarily correlate with dietary specialization, and
in fact, seems to be more closely tied to the phylogenetic history of the
organism than any special adaptation for a particular diet (Robinson
and Redford 1986b).

Body size, behavior, and life history

Body size has profound implications for how a mammal can locomote,
whether it be on land, sea, or air. For example, there is a maximum
limit on size if an animal becomes specialized for steady, powered flight
rather than hovering flight. Such relationships have been studied not
only for bats but of course for birds and insects (Pirlot 1977).
If we consider nonvolant mammals and terrestrial locomotion, then
the following generalizations can be made: Maximum speed covaries
positively with body size, even though step frequency covaries negatively
with body size. That a large mammal can achieve a higher speed derives
from the fact that the stride length is relatively greater as size increases
(Calder 1984). Larger-bodied mammals have the capacity to use a larger
area of space more efficiently because, although metabolic rate increases
in all species as speed increases, the per-gram cost of rapid locomotion

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 Significanct of body size in mammalian tcoldgy 31

is higher for smaller than for larger forms (Taylor, Heglund, & Maloiy
1982). (This becomes apparent when metabolic rate during locomotion
is plotted against speed, with large-bodied forms plotted separately from
small-bodied ones (Calder 1984): The slope is less for larger forms than
· for smaller forms.) It follows that l.arger forms can more efficiently
exploit a larger home range; and, although the total energy consumed
· by a larger form is greater than that of a smaller one (and thus, ulti­
mately, larger-bodied forms, whatever trophic specialization, need
larger areas), it is a fact that home-range size increases with body size
at a faster rate than do individual metabolic require.ments (Harestad &
Bunnell 1979; McNab 1963). Once again there is considerable scatter
around the regression line which can in part be accounted for by trophic
specializations. For any given size class, given comparable primary pro­
duction values for the habitat, a herbivore has a smaller home range
than a carnivore.
What a mammal can perceive often scales with body size. For ex­
ample, in regard to sound production with vocal chords, larger-bodied
forms produce a range of sounds having lower frequencies than those
produced by smaller-bodied forms; and with respect to audition in ter­
restrial forms, the highest frequency perceived covaries negatively with
body size (Calder 1984; Dooling 1980).
As indicated in the previous section, trophic strategy and body size
are inextricably linked. For example, mammals feeding on small insects
scattered widely in the forest floor litter are often of small body size.
Those taxa specialized for feeding on social insects, such as ants or
termites, are a special case. These "anteaters" may be of modest, to
even large, body size. Granivores, mammals that harvest small seeds,
are often of modest body size. Of course, man, who cultivates and
processes cereal grains on a large scale, is an exception. Ruminants, or
foregut fermenters, have a lower limit on the body size they can attain
because of the necessity for ceasing feeding while fermentation takes
place in the foregut, thus reducing the rate of passage for ingested
herbaceous material (Janis 1976; Parra 1978). Constraints on niche oc­
cupancy as a function of body size have been extensively reviewed in
Eisenberg (1981).
Body size profoundly affects life history strategies, a phenomenon
expansively dealt with by Bonner in his 1965 book. For example, gen­
eration interval scales positively with body size, longevity scales posi­
tively with body size, and the reproductive strategy of a species is very
much influenced by adult body size. Mammals that opt for a near se-

,
•
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32 John F. Eise,1berg

melparous mode of reproduction are almost always of modest body size

and attain their adult growth within a year. On the other hand, those
species showing extreme iteroparity, with a litter size of one and a long
interbirth interval, are often of large body size (Eisenberg 1981). Ad­
aptation for various foraging strategies has profound implications with
respect to the form of social organization that a species exhibits (Ei­
senberg 1966; Jarman 1974).
In a previous publication (Eisenberg 1981), I attempted to arrive at
some general correlations between trophic strategy and social structure.
The argument is difficult to condense, but the rest of this section sum­
marizes the major hypotheses. Throughout the evolution of the Mam­
malia there has been pressure from competitors (both intraspecific and
interspecific), predators, pathogens, and parasites coupled with a fluc­
tuating global climate, resulting in fluctuating levels of primary produc­
tivity. The role of pathogens and parasites in shaping mammalian
communities has often been overlooked (Barbehenn 1978; Muul 1978).
Yet I concur with Jerison (1973) that the history of the Mammalia on
the "contiguous" continents (North America, Europe, Asia, Africa) has
resulted in an overall increase in relative brain size. The corollary to
this statement is that the neocortex has increased in size; thus the stage
was set for a feedback loop between individually acquired experience
and learning by descendents, enabling them to operate in a more and
more unpredictable world as new niches were invaded. The evolution
of complex, interdependent social groupings is not the only outcome o f
such processes; rather it is only one of four possible contemporary out­
comes of the total, historical selective forces (as illustrated in Figure
156 of Eisenberg [1981]). Within the extant species of the class Mam­
malia, a relatively large brain tends to be associated with selective forces
that have shaped a species that is K-selected; that is, the individual within
the species has a relatively long reproductive life and a small litter size.
Often, but not always, within a historical lineage such a species has a
relatively large body size (Eisenberg 1981 ).

Using current data to project t.o the past

The above represents the argument for the important influence of trophic
strategy upon the density a mammalian species can achieve within a
community. Likewise the standing-crop biomass exhibited by a species
{or a population of a species) within a community is also determined
by trophic strategy and life history strategy. Life history strategies are

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 Significance of body size in ,nammalian ecology 33

constrained by trophic specializations, but demographic processes are

ultimately the outcome of age at first reproduction, mean annual litter
si.ze, and age at last reproduction (Cole 1954). For a mammalian pa­
leontologist, the genuine problem in reconstructing extinct communities
is to decide on which trophic category to assign a species that one has
never seen. Moreover, one confronts the problem that one will never
see the species in nature. In the early years of community studies, the
trophic categories allocated to secondary consumers were broad: her­
bivore, omnivore, carnivore.
Let us take the category of herbivore for closer scrutiny. In the original
sense of the tenn, it implied feeding on plants or parts of plants. We
now know that there are profound consequences for the life history of
a species if it is selected to become a frugivore rather than a folivore,
or a browser rather than a grazer. In short, our science is moving toward
a finer definition of trophic specialization for primary and secondary
consumers. Consider the work of Hoffmann and Stewart (1972) in es­
tablishing the correlations between the stomach structure and feeding
habits of East African ungulates, correlating with the behavioral ob­
servations of Jarman (1974). The feeding habits of browsers and grazers
in the East African ungulate community arc a continuum; yet mor­
phological and behavioral data also form a concordant continuum. Un­
fortunately, behavioral data and the anatomy of the "soft parts" are
unavailable to the paleontologist. Yet, I feel we can make progress in
paJeoecology even with only skulls, teeth, and. hopefully, postcranial
elements at our disposal.
The work of Kay and Cartmill (1977), Kay and Hylander (1978), and
Cartmill (1972) justifies the previous assertion. In his article, Cartmill
asked the question, What morphological specializations can one discern
in the anatomy of extant arboreal species (both diurnal and nocturnal)
that may have relevance for the interpretation of fragmented fossils
from the Paleocene-Eocene boundary? Great insight was demonstrated
in selecting the extant Sciurus carolinensis and Didelphis virginiana as
two of several possible examples for comparison and contrast. Both
species were readily observable and available in North America. Kay
and Cartmill undertook a more ambitious project when they attempted
to infer the habits of the Paleocene Palaechthon naci"1ie11ti. In this case,
the task was to determine diet, and probable hunting strategy (i.e.,
major sensory inputs), from a set of pitiful fragments. The authors took
many measurements from extant forms to establish such facts as relative
size of orbit and activity cycle, relative size of the foramina, and vibrissal

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34 John F. Eisenberg

development to create a picture of the sensory world of Pa/aechthon.

Kay and Hylander addressed, and answered, the question, Can the
degree of feeding on leaves in extant primates be correlated with molar
tooth measurements? The three articles cited are not indicative of a new
trend in paleontological research. Undoubtedly, from the time the first
fossil skull was discovered by a human possessing an inquiring mind,
the habitus of the fossil was postulated based on the discoverer's own
experience with extant forms. Indeed, all museum exhibitions of fossils
represent "educated guesses" at natural history. I propose nothing new
in one sense, but I do propose that as neontologists become more so­
phisticated, so should paleontologists. The dialogue established at the
symposium should - no, must - continue.

Concluding remarks
As this brief review indicates, there are many rules of biological scaling
that we can discern in extant forms that are probably excellent predictors
for extinct species. Once the radiation of mammals had commenced in
the Paleocene, and the major lines of descent had diverged, recognizable
lineages developed which through time have become specialized for
various niches. And even with extinction events, the remaining stocks
of mammals have seemingly been able to adaptively radiate into similar
niches. This is especially true for the period following the extinctions at
the end of the Oligocene, when the Late Miocene and Pliocene com­
munities emerged. These communities have more or less persisted
through time, although shifting geographically depending on global clima­
tic changes. Some cautionary words must be uttered, however, in that
when any of these allometric relationships have been established, there
usually is a great deal of residual scatter around the regression line. As
I have indicated in this paper, and as McNab has indicated in his, much
of this scatter can be accounted for if one knows something about the
trophic strategy of the species. One should never discard or discount
the residual variation; it often is of the greatest biological interest.
Let us consider the problem of scaling and regression analysis. Many
biologists would hope to develop mathematical expressions of their mea­
surements that would allow a predictive power emulating those equations
derived by physicists (Lotka 1956). Yet, biology is a young science, as
compared to astronomy, let alone physics. Furthermore, even though we
revere physics for its precision and foundation in the experimental
method, the discipline is as subject to the influence of political forces as

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 Significance of body size in mammalian ecology 35

any other science dependent on government subsidies (McCormmach

1982). Most biologists, in plotting a supposedly dependent variable
against an independent variable, consider the scatter about a regression
line as "noise." This is unfortunate, since the scatter may reflect our ina­
bility to appropriately categorize our data set int.o biologically meaningful
subsets. qiven the inertia of institutionally sanctioned jargon, I say, be­
ware of "hardening of the categories."
For example, once again we may consider the regression analysis p e r ­
formed b y Kinnaird and Eisenberg (in press) for Neotropical mammals.
Standing-crop biomass for species was regressed against mean body
weight. There wa� enormous scatter, but greater than 70% of the vari­
ance could be accounted for by trophic specialization. Similar results
were established by Robinson and Redford (1986b) for species densities
of Neotropical mammals. Scatter about. a regression line should not be
considered an annoyance, but rather a challenge.

Rererences
Barbehenn, K. R. 1978. Discussion: concluding comments, from the worm's
view "eco-pharmacodynamics" and 2000 A.O. In Populations ofSmall Mam­
mals under Natural Conditions, ed. D. P. Snyder, pp. 231-236. Special
Publication Series, Vol. 5. Linesville, Penna.: Pymatuning Laboratory of
Ecology. University of Pittsburgh.
Bonner, J. T. 1965. Size and Cyde. Princeton, N.J.: Princeton University Press.
Calder, W. A., Ill. 1984. Size, Function, and Life Hi.story. Cambridge, Mass.:
Harvard University Press.
Cartmill, M. 1972. Arboreal adaptations and the origin of the order Primates.
In The Function and Evolutionary Biology of Primates, ed. R. Tuttle,
pp. 97-122. Chicago/New York: Aldine-Atherton.
Cole, L. C. 1954. The population consequences o f life history phenomena.
Q. Rev. Biol. 29:103-137.
Darnuth, J. 1987. Interspecific allometry of population density in mammals and
other animals: the independence of body mass and population energy-use.
Biol. J. Linn. Soc. 31: 193-246.
Dooling, R. J. 1980. Behavior and psychophysics of hearing in birds. In Com­
paraJive Studies of Hearing in Vertebrates, ed. A. N. Popper & R. R. Fay,
pp. 261-285. New York: Springer.
Eisenberg, J. F. 1966. The social organizations of mammals. Handbuch der
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· & B. A. Wilcox, pp. 35-55. Sunderland, Mass.: Sinauer Press.
Eisenberg, J. F. 1981. The Mammalian Radiations. Chicago: University of Chi­
cago Press.

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Copyrighted material
 4
The functional anatomy of body �as.,
.
THEODORE I.GRAND

One central challenge for the paleontologist is to reconstruct body mass

from fragments of bone and teeth. Estimates of volumes, whether of
specific soft tissues or the entire body mass, are derived from linear
measures. Moreover, this becomes a closed system: One makes suc­
cessive approximations, but can never verify volumetric/mass statements
about e.xtinct animals. The paleontologist, unable to make direct anal­
_
yses, looks to living species for guidance and the luxuries of choice and
abundance. The anatomist, on the other hand, works relatively uncon­
strained: There are infinite opportunities to dissect; t.o make direct mea­
surements of skin, adipose stores, muscle, bone, and body mass; to
observe directly the biomechanics of motion, of physiological processes,
of social behavior. How can the "loop" of linear measures of volume
and body mass open up? What can the anatomy of living animals say
to the paleontologist?
I have studied more than 150 genera of metatherian and eutherian
mammals (from 10-g bats to 1000-kg whales). But a survey is not ap­
propriate: The student of fossil carnivores has no direct interest in mod­
em bats; the student of ungulate origins need not know about the
anatomy of modem whales. One conclusion about mammalian body
composition, however, is important: For the largest tissues of the body
(skin, muscle, bone, adipose stores), the dominating components of
weight vary with function and, to a degree, independently of one an­
other. How does evolutionary process "tinker" with the component
tissues, while weight itself remains constant?

In Body Sizt in Mammalian Paleobiology: Estimation and Biological Implica­

tions, John Damuth and Bruce J. MacFadden, eds.
o Cambridge University Press 1990.

39

Copyrighted material
40 Theodore I. Grand

To demonstrate that locomotpr adaptation has a substantive effe.ct on

body composition, I will compare pairs of species of equal body mass:
the potto, Perodicticus potto, with the greater galago, Ga/ago crassi­
caudatus, at 1000 g; the red panda, Ailurus fulgens, with the raccoon,
Procyon lotor, at 4000 g. Differences in the proportion of skin and
muscle correlate directly with locomotor and antipredator behavior.
Since this is true across my wide study sample, I suggest that an analysis
of body mass must begin by interpreting functionally the largest tissues­
for example, cutaneous adaptations such as quill or carapace, scaling
effects on skin proportion, differences between arboreal and terrestrial
locomotor behaviors, and patterns of adipose storage. Then, we have
a basis for understanding body mass and the potential for better esti­
mates among extinct species.

Methods
My published data (Grand 1977) on three P. potto and two G. crassi­
caudatus have been increased to five adult individuals of each species.
Results of the studies of five adult A. fulgens and six P. lotor have not
yet been published. The mass-related measurements have been de­
scribed in a series of papers (Grand 1977, 1978, 1983).
Body composition: Skin, muscle, skeleton, and adipose stores
were dissected and weighed separately, and each quantity
was divided by total weight to determine its percentage of
total body weight (%TBW).
Functional muscle groups: The weight of each muscle group
(forelimb, hindlimb, back extensors, masticatory, tail) was
divided by the total weight of all muscle.
The ratios of limb skeleton: The weight of the forelimb (scapula,
humerus, radius and ulna, band/paw) skeleton was divided
by that of the hindlimb (femur, tibia and fibula, foot).
The mean values for each tissue and muscle group are given in the
figures. Error bars show the 95% confidence intervals for these means.

Results

Pair contrasts - /000 g: the potto (Perodicticus potto) and the

greater galago (Galago crassicaudatus)
These animals represent the structural and behavioral extremes of the
lorisoid radiation of African prosimians (Charles-Dominique 1977;

Copyrighted material
 The functional anatomy of body mass 41

Grand 1977). Both are arboreal and nocturnal, and at 800-1200 g (Na­
pier & Napier 1967) are equal in body mass. There the similarity ends.
The potto is a slow-moving climber with a stumpy, almost ftaplike tail.
The extreme mobility of each limb joint provides the basis for the enor­
mous reach of hands and feet within three-dimensional space; fingers
and toes have a viselike grip; the vertebral column exhibits snakelike,
sinusoidal flexibility. Pulling with the forelimb and pushing with the
hindlimb enable the animal to glide across branch supports like a
"slinky" toy. Because one or more limbs is always in contact with a
branch, the potto is supported during the entire gait cycle. As a con­
sequence, branches deform slowly, and rustling noises are reduced. This
acoustic "crypsis" is a common antipredator strategy, and part of the
potto's own stalking technique with regard to insects, nestling birds, and
other live food.
By contrast, the galago is a long-legged, long-tailed vertical clinger
and leaper (Napier & Walker 1967). As in the case of Tarsius (Grand
& Lorenz 1968) hindlimb dominance is unequivocal: The great mass
and length of each hindlimb segment, the elongate and elaborate fulcrum
within the foot (which nevertheless retains prehensile digits) are ap­
parent. The ability to leap and change direction at landing resembles
the ricochetal hop and antipredator evasion of some small rodents,
insectivores, and the macropods.

Body composition. The potto has 15% more skin, but only 75% of
the muscle of the galago (Figure 4.1). Although the percentage of skel­
etal mass is similar, the ratios of forelimb to hindlimb differ significantly
- 1: 1 in the potto and 1 :2 in the galago.

Functional muscle groups. In the potto almost 30% of all muscle is

concentrated in the hindlimb; 40% is found in the forelimb (Figure 4.2).
This muscular balance reflects the push-pull nature of potto climbing
(Charles-Dominique 1977). Less than 10% of all muscle is concentrated
in the back extensors; with the tail reduced, there is almost no sacral
muscle.
In the galago a higher proportion of muscle is concentrated in the
hindlimb (over 40% of all muscle) than in the forelimb (under 30%).
Moreover, the back extensors constitute a higher proportion of muscle
than in the potto. Tail muscle adds another 3%.
There are "local" muscle-group differences as well. The quadriceps
femoris are larger (jumping is based upon powerful knee-joint exten­
sion) than those of the potto. However, the adductors, powerful climbing

Copyrighted material
42 Theodore I. Grand

...
"'
.r. 30
'ii
,
..
.,,,,' ,..-
t•
..

� 25
"8
CD

...., 20

-0
I-

...
0 15
.,
C

.,
�
Q.
10

Potto Galago

E:ZJ Skin 1SS1 Bone � Muscle

Figure 4.1. Body composition in (A) P . potto and (B) G . crassicaudatus: skin,
bone, and muse.le as percentages of total body weight.

and stabilizing muscles, are much smaller in the galago than they are in
the potto.
The potto has reduced the bulk of the lumbar extensor muscles as
well as the lengths of transverse and spinous processes. This complex
of elements enhances flexibility of the lumbar region, a primary adap­
tation to slow climbing. The galago has much larger lumbar extensors,
and the transverse and spinous processes are much more pronounced.
The loss of side-to-side mobility, which makes hindlimb propulsion more
effective, is like that of scansorial, hindlimb-dominant species such as
the kangaroo rat (Dipodomys), the brown four-eyed opossum (Meta­
chirus) (Grand 1983), and the elephant shrew (Elephantulus).

Pair contrasts - 4 kg: the red panda (Ailurus fulgens) and the
raccoon (Procyon lotor)
The panda is an arboreal climber that can walk on the ground; the
raccoon is a terrestrial quadruped that can climb (Ewer 1973; Lotze &

Copyri ghted material

 ,

The f11nctional anatomy of body mass 43

45
A 42.6
40 38.5

35

.!!
:::, 30
:::,
� 25

-...
0
20
C: 15

10

Potto Galago

� Tail � Hindlimb � Back extens. � Forelimb

Figure 4.2. Functional muscle groups in (A) P . polto and (B) G. crassicaudatus:
the mass of muscle groups in tail, hindlimb, back, and forelimb as percentages
of total body musculature.

Anderson 1979; MacClintock 1988; Roberts & Gittleman 1984). Limb

proportions are simi!.ar (Davis 1964), but the segments differ in relative
mass (Grand, unpublished data): The thigh of the panda is two-thirds
that of the raccoon; the tail is almost 50% heavier.

Body composition. The skin (16% TBW), adi_pose tissue (6% TBW),
and bone (15% TBW) in both species are equal (Figure 4.3). Data on
the raccoon (Grand, unpublished) show enormous seasonal fluctuations
in adipose stores. In terms of muscularity, the two animals differ sig­
nificantly. The red panda is 30% muscle; the raccoon, 35% muscle.

Proportions of the limb bones. The ratio of forelimb to hindlimb bone

is 1:1 in the red panda; 0.75:1 in the raccoon.

Functional muscle groups. Less than 30% of all muscle in the red panda
is concentrated in the hindlimb, slightly more than 30% in the forelimb

Bahan dengan hak cipta

Another random document with
no related content on Scribd:
 Che ne fait mies à demander se li prinches de Galles et
d’Acquittainne fu grandement tourblés en couraige et fort
courouchiés, quant soi se vei semonre et adjourner ou nom de ses
subgés en estraingne court, ilz qui se tenoit et presumoit ungs des
grans dou monde et à qui le plus droite gens d’armes et touttes
mannierres de gueriieurs obeyssoient. Toutteffois, il respondi adonc
enssi comme chy dessus est dit. Lors fissent ses gens partir et yssir
de sa cambre et de sa presence le chevalier et le clerc dessus
noummet, qui n’estoient mies à leur aise, car nulx ne les
accompagna à leur hostel, et sentirent bien qu’il avoient bien
durement courouchiet le prinche. Si se fuissent vollentiers tantost
parti, mès il leur fu dit que point ne partesissent jusquez à tant que
il aroient responsce du prince. Si obeirent et sejournèrent en
Angouloime quatre jours, sans point yssir de leur hostel. Au cinqime
jour, il leur fu dit qu’il pooient bien partir, se il volloient, car il avoient
bien fait leur messaige, et que li prinches n’y responderoit autrement
qu’il avoit respondu. Sur chou il se partirent, et prissent leur chemin
pour venir viers Toulouse et pour yssir de la princhipauté au plus tost
qu’il pewissent, car il n’y estoient mies trop bien asseur. Enssi qu’il
chevauchoient tout leur chemin à grant esploit, il furent rencontré et
aresté dou senescal d’Aghinois, monsigneur Guillaumme le Monne,
et pris et menet em prison à Pennes en Aginois. Ces nouvelles
s’espandirent tantost et vinrent en Franche, car li varlet des dessus
dit n’eurent nul empeschement, mès cheminèrent tant qu’il se
trouvèrent à Paris, et recordèrent à tous ceux qui leur demandèrent
de leurs mestres, coumment il leur estoit. Quant li roys de Franche
en seut le verité et la responsce dou prinche, il n’en fu mies mains
penssieux. Toutteffois, il s’en porta et passa adonc au plus bellement
qu’il peut, et n’en fist nul grant samblant... Fos 151 vo et 152.

§ 607. De ceste response.—Ms. d’Amiens: En ce meysme tamps

et en celle meysme sepmainne, estoit revenus à Paris messire Guis
de Blois qui adonc estoit encorres escuiers, mès il fu fais celle année
chevaliers en Prusse et y leva bannierre. Si estoit revenus, si comme
dessus est dit, d’Engleterre quittes de se foy, car il avoit esté
ostagiiers pour le roy de Franche et sejourné pour ceste cause en
Engleterre six ans ou environ. Si regardèrent adonc chil qui dallés lui
estoient et qui le gouvrenoient, doy escuier, Hues de Villers et
Jehans de Leglisuelle, qu’il perdoit son tamps; si traitièrent par
deviers le consseil dou roy englès, sus se delivranche, avoecques le
consseil et l’avis d’un sage baron de Franche, monsigneur Carle de
Montmorensi, qui se boutta sagement ou traitiet et fist tant qu’il fu
delivrez et quittes ossi de sa foy et de sa prison enviers le dit roy
englès, parmy deux mil frans qu’il paiia. Et li dis messires Guis de
Blois rendi, quitta et ahireta le roy englès de le comté de Soissons
qui estoit adonc ses hiretages. En ces traitiés et pourkas à faire, aida
grandement et mist pluiseurs painnes et conssaux li sires de Couchi
qui tendoit et tiroit à avoir la ditte comté de Soissons, ensi qu’il eut,
en escange d’autre terre et revenue dont il estoit assignés sour les
coffres dou dit roy englès, de par madamme Ysabiel sa femme, fille
au dit roy englès que li jonnes sires de Couchy avoit nouvellement
pris par mariaige: ceste est la cause pourquoy il parvint
premierement à le comté de Soissons. Or revenons à le matère dou
prinche qui estoit adonc en Espaingne. Fo 149.
P. 99, l. 27: selonch che.—Le ms. A 8 ajoute: grandement. Fo 299
vo.
P. 100, l. 5: Guillaumes.—Ms. A 8: Jehan. Fo 300.
P. 100, l. 29: de ce.—Ms. A 8: parmy ce.
P. 100, l. 30 et 31: pareçons.—Ms. A 8: pactions et couvenances.
P. 101, l. 3: l’ostagerie.—Ms. A 8: l’ostage.
P. 101, l. 14: Guillaummes Wikam.—Ms. A 8: messire Guillaume.
P. 101, l. 16: offices.—Le ms. A 8 ajoute: de chancellerie.
P. 101, l. 17: vaghièrent.—Ms. A 8: furent vacans.
P. 101, l. 23: messages.—Ms. A 8: messagiers.
P. 102, l. 3: que amiable à se composition.—Ms. A 8: qu’il lui feust
courtois et aimable à sa composicion.
P. 102, l. 4: l’euist.—Ms. A 8: eust le dit eveschié.
 P. 102, l. 14: en Engleterre.—Ms. B 6: Se fu le conte daufin
d’Auvergne mis à finanche, et paia trente mille frans, et le comte de
Poursien dix mille, et tout li aultres hostaigiers des chités et bonnes
villes de Franche furent espars parmi Engleterre et tenus en diverses
prisons. Fo 727.

§ 608. Vous devés savoir.—Ms. d’Amiens: Vous avés chy dessus

bien oy recorder coumment li prinches de Galles fu ajournés à venir
en le cambre de parlement à Paris à l’encontre dou comte
d’Ermignach, dou seigneur de Labreth, dou comte de Pierregorth,
dou comte de Comminges, dou viscomte de Quarmaing et de
pluisseurs grans signeurs de Gascoingne, à oïr droit et le declaration
de l’apel qu’il avoient fourmé contre lui, sus les griefs qu’il volloit
faire en leurs terres. Si sachiés que de cel adjour li prinches fu
durement courouchiés et le prist en grant despit, et dist bien que la
cose ne demourroit mies enssi. Non obstant ce, toudis procedoit il
dou fouaige que il volloit eslever, et li avoient chil de Poito, de
Saintonge, de le Rocelle, de Roherge, de Querzin et de Limosin
acordet; car il n’en pooient ne n’osoient ad present autre cose faire,
tant estoit leurs pays raempli d’Englès, offisciers au prinche, et ossi
touttes ces terres dessus noummées sont moult enclinnes et
obeissans à celui qu’il tiennent pour leur naturel seigneur, et adonc il
y tenoient le prinche et nul autre... Fo 152 vo, col. 1.
Si vous di que adonc avoit ung senescal en Roherge, qui s’appeloit
messire Thummas de Welkefare, chevalier englès, et se tenoit li di
senescaux à Villenove d’Aghinois. D’autre part, sus le frontière dou
pays estoient li viscomtes de Quarmaing, ungs mout appers
chevaliers, li sires de la Barde, li sires de Taride et li sires de
Picornet. Chil dit signeur et chevalier, qui s’estoient mis et bouteit en
l’apiel avoecq les autres, avoient pris en grant despit le prise de
monsigneur Caponnet de Caponval et de son compaignon. Si
s’avisèrent qu’il feroient embusce sus les gens dou prinche et en
atraperoient ossi aucuns. Si seurent par espies que messires Thumas
de Welkefare devoit chevauchier deviers Rodais, ensi qu’il fist, à
gens d’armes, pour entendre à le forterèche et rafreschir de tout
chou que il y besongnoit. Si tost qu’il seurent ces nouvelles, il se
queillièrent, et furent bien trois cens lanches, et se missent sus leur
embusce par où li chevaliers englès devoit passer. Enssi que
monsigneur Thummas de Welkefare et se routte chevauchoient, et
pooient estre environ soissante lanches et deux cens archiers, que
brighans, ceste embusche leur sailli au devant, les lanches abaissies,
en escriant et disant: «Vous n’en yrés mies ensi.» Lors se ferirent
ens de plains eslais, et en y eut de premier encontre pluisseurs
abatus, d’un lés et d’autre. Là se deffendirent Englès de leur costé
au mieux qu’il peurent, et se combatirent vassaument; mès
finablement, il ne peurent durer, car li Franchois estoient grant
fuisson et tout pourvueu de leur fait. Si furent li Englès desconfis, et
à grant meschief se sauva li senescaus de Roherge messires
Thummas de Welkefare, et s’en vint par force de bon courssier à
Montalben et se bouta ens ou fort, et ses gens furent tous espars, et
se sauvèrent chil qui sauver se peurent.
Ces nouvelles vinrent au prinche, qui se tenoit en le cité de
Angouloime, et n’estoit mies trop bien hetiés, coumment li sires de
Pincornet, li viscomtes de Quarmaing et li sires de la Barde avoient
rencontré son senescal de Roherge et desconfit et cachiet jusques à
Montalben. Si en fu li prinches durement courouchiés, mès amender
ne le peut, tant qu’à ceste fois.
Adonc estoit li dus d’Angho à Thoulouse, mès il se tenoit encorres
tous quois, fors tant qu’il traitoit et faisoit traitier toudis as
cappittainnes des Compaingnes, que il les pewist avoir à son acort,
enssi qu’il en eult pluisseurs, enssi comme vous orés chi apriès. Ossi
d’autre part, deviers le prinche estoient revenu messires Thumas de
Felleton, messires d’Agohrises, messires Hues de Hastingues,
messires Richars Tanton, messires Gaillars Vighier et chil qui avoient
estet pris en Espaingne en l’avant garde dou duc de Lancastre, si
comme il est chi dessus dit et contenu en l’istoire, et s’estoient
ranchounnet et delivret, li ung par mise de deniers, li autre par
escange. Ossi, dou costé des Franchois, se delivrèrent de prison et
finèrent au mieux qu’ils peurent, messires Ernouls d’Audrehen,
messires Jehans de Noefville, li Bèghes de Vellainnes, li Alemans de
Saint Venant et li chevalier et escuier de Franche qui avoient estet
pris à le bataille de Nasares. Et fu ranchounnés messires Bertrans de
Claiequin deviers monsigneur Jehan Camdos, qui estoit ses mestres,
à cent mil frans. Bien les paiia li di messires Bertrans en biaux florins
tous appareilliez... Fos 152 vo et 153.
Quant li prinches de Galles vit et entendi que c’estoit à certes, que
on le guerioit enssi de tous costés et que li Franchois se mettoient
en painne de li tollir son pays, si s’avisa quil se deffenderoit, mais il
n’estoit mies em point de chevaucher. Si envoia tantost deviers
monsigneur Jehan Camdos, qui se tenoit à Saint Sauveur le
Viscomte, en lui segnefiant et mandant que il retournast tantost.
Quant messires Jehans Camdos oy ces nouvelles, si ne li plaisurent
pas trop bien, car trop le deshetoit et anuioit la guerre renouvellée,
et sorti et dist tantost que grans maux en venroient. Nonpourquant,
il se hasta au plus tost qu’il peult, et s’en vint en Anghouloime
deviers le prinche, qui le vit mout vollentiers. Assés tost apriès le
revenue de monsigneur Jehan Camdos, fist li prinches ung grant
mandement de chevaliers et d’escuiers d’Acquittainne, de chiaux quil
tenoit à avoir le comfort. A son mandement vinrent li captaux de
Beus, li doy frère de Pumiers, messires Jehans et messires Elies, car
messires Aimmenons de Pummiers s’estoit partis et disoit que il s’en
yroit outre mer aventurer en estraingnes terres, et que point de celle
guerre ne se volloit ensonniier, ne franchois, ne englès. Si y vinrent
encorres li sires de Partenay, messires Aimmeris de Tarse, li sires de
le Ware, englès, messires James d’Audelée, senescaux de Poitou; et
les envoiea li prinches en le marche de Toulouse yaux tenir à
Montalben, pour deffendre le pays contre les Franchois qui là se
tenoient.
Quant messires Jehans Camdos et li captaux, qui estoient chief et
souverain de ceste cevaucie, furent venu à Montalben, et li chevalier
dessus noummez, assés tost apriès leur revinrent messires Loeis de
Halcourt, messires Rammons de Moroel, messires Loeis de Melval,
troy grant baron et de grant affaire; si fissent à Montalben une
bonne garnison, et coummenchièrent à chevaucher ou pays
thoulouzain et à faire mout de dammaiges. Adonc estoient les terres
en grant variement, car un jour estoient franchois et l’autre, englès;
ne point de estableté n’y avoit, fors li plus fors tenoit le plache:
quant plus fors revenoit, il reconcqueroit chou qui avoit estet
concquis.
A ce donc avoit ung senescal en Roherge, bon chevalier durement,
qui se noummoit messire Thummas de Wettevale, et tenoit une
fortrèche à quatorze lieuwes de Montpellier, que on appelloit la
Millau, sus les mettes de Roherge et de Limozin. Si sachiés qu’il se
tint en le ditte fortrèche moult vaillamment, si comme vous orés chy
apriès, et tout chil qui avoecques lui estoient. Fo 154 vo.
P. 102, l. 18: l’ajour.—Ms. A 8: l’ajournement. Fo 300 vo.
P. 102, l. 23: Compagnes.—Le ms. B 6 ajoute: telz que Naudons
de Bagherant, le bourch de Bretuel, le bourch Camus, le bourch de
Lespare, Lami, Espiote, Hanequin Franchois, messire Robert Brickés,
Cressuelle, messire Robiers Ceni, messire Perducas de Labreth,
messire Garsis de Castiel, messire Gaillart Vighier, Bernart de le
Salle, Bernart de Wesc, Hortigo et pluisseur aultre. Fo 728.
P. 102, l. 26: temprement.—Ms. A 8: brief.
P. 102, l. 27: faire.—Les mss. B 4 et A 7 ajoutent: et les
ensonnieroit. Fo 298.—Ms. A 8: et les embesongneroit. Fo 300 vo.
P. 102, l. 28: joiant.—Ms. A 8: joieux.
P. 102, l. 29: d’enfle.—Ms. A 8: d’enfleure.
P. 103, l. 4: ydropisse.—Ms. A 8: ydropisie.
P. 105, l. 7: Montpesier.—Ms. A 8: Montpellier. Fo 301.

§ 609. Li rois de France.—Ms. d’Amiens: En ce tamps, estoit li

roys de Franche en grant branle pour gueriier le roy d’Engleterre, car
li roys englès li requeroit fortement qu’il se delivrast de parpaiier le
redemption dou roy son père, et que trop y metoit au paiier, ou
autrement il li feroit guerre; car, seloncq le teneur de le pais, on
devoit avoir tout paiiet dedens trois ans, et encorres y avoit seize
cens mil frans de Franche à paiier. Enssi estoient les grongnes de
l’un à l’autre: li roys englès courouchiés et dur emfourmez sour le
roy de Franche, pour tant qu’il ne se delivroit point de paiier la
somme des deniers où tenus il estoit, et qu’il avoit consenti que sez
filx li prinches de Galles estoit adjournés de ses soubjès em
parlement à Paris, et consenti encorres qu’il li faisoient guerre, et si
avoient cil leur retour en Franche et dalés le roy; et li roys Franchois,
d’autre part, ossi courouciez et dur enfourmés sour le roy englès,
pour tant qu’il soustenoit et comfortoit lez Compaignes, si comme on
disoit, et les envoieoit en Franche, et qu’il volloit tenir la duché
d’Acquittainne sans resort, qui estoit grandement ou prejudisce dou
royaumme et hors de le vollenté de touttes gens residans en celui
pays. Si n’en pooit longement estre ne demourer, que guerre ne se
remeuist entre ces deux rois, mès li rois de Franche ne le volloit mies
recoummenchier, se il ne savoit bien coumment: si s’estoit
pourvueus de loing temps, tout bellement et tout sagement, d’avoir
atrais et acquis à lui tous ses voisins, le duc Aubert, bail de
Haynnau, et les signeurs de celui pays, ossi le duc de Braibant et les
signeurs de celle terre, le comte de Clèves, l’evesque de Liège,
l’evesque de Miés, l’evesque de Verdun, le duc de Loerainne, le duc
de Bar, le comte de Montbliar, le comte de Genève, le comte de
Savoie et touttes ses gens, le signeur de Roussellon et les gentils
hommes de la duché de Prouvenche et de la comté de Venisin, ossi
le comte de Fois et touttes ses gens, le roy d’Arragon et touttes ses
gens, et tous les marcissans environneement autour de son
royaumme. Et sentoit encorres bien que pluisseurs signeurs, comte,
baron, chevalier et escuier de la duché d’Acquittaine, se retouroient
deviers lui, se la guerre estoit renouvellée, et mout de villes, de
chités et de castiaux de la ditte duché, qui point n’amoient les
Englez. Et par especial, il avoit tous les coers des gentils hommes de
Bretaingne, qui moult li pooient valloir, car il savoient bien que li dus
de Bretaigne estoit plus englès que franchois; mès, là où ses pays
vorroit estre pour lui, dou corps ne de le haynne dou dit ducq ne
faisoit il mies grant compte. Avoecq touttes ces coses, li rois Carles
s’estoit trop fort arestés à savoir se cil de Pontieu vorroient estre de
son acort; il trouva que oil moult vollentiers, car il ne pooient ainmer
les Englès. Che plaisy grandement au roy de Franche et traita
enviers chiaux de Abbeville tout secretement et bellement, et leur
proummist et jura, là où il se vorroient rendre deviers lui, que
jammais ne les metteroit hors dou demainne dou royaumme, et les
tenroit en plus grant francise et liberté que chiaux de Paris. Enssi se
composèrent ces besoingnes... Fo 153.
Or avint que li princes, pour mieux venir à sen entension, par le
consseil de l’evesque de Bade, envoiea grans messages deviers le
pappe Urbain qui adonc tenoit son siège à Romme, telx que son
marescal d’Acquittainne monsigneur Guichart d’Angle, monsigneur
Jehan Isoret son fil, monsigneur Guillaumme de Seris et maistre
Jehan Briffaut, un advocat de le Rocelle, pour impetrer cel fouage
sus le clergiet et pour pluisseurs autres besoingnes. Si lairons à
parler des dis messagiers qui fissent leur voiaige et esploitièrent
d’aucunes coses, et non pas de touttes, deviers le pappe, si comme
je leur oy recorder, car je me parti de Romme avoecq yaux et
rappassay les mons en leur compaignie, et parlerons de cel apiel des
barons de Gascoingne, et coumment il s’eslevèrent et parseverèrent
contre le prinche... Fo 152 vo.
En ce tamps, revenoit de Romme li marescaux d’Acquittainne
messires Guichars d’Angle, qui y estoit allés pour les besoingnes dou
prinche. Si entendi, entroes qu’il estoit en Savoie sur son retour, que
la guerre estoit renouvellée entre les deux roys. Si se doubta que il
ne fust pris ou espiiés, et se parti secretement de ses gens et se
mist en abit et estat d’un povre cappelain et laissa tout son arroy. Si
rapassa parmi Auvergne messires Guichars, enssi que je vous di, et
parmi Limozin, et entra en Poito. Et messires Guillaummes de Seris,
uns chevaliers de Poito, qui estoit en se compaignie, ne s’osa
aventurer enssi que li dis messires Guichars fist, mès s’en vint à
l’abbeie de Clugny et se mist en sainte terre, et se tint là plus de
cinq ans. Fo 154.
P. 107, l. 21: breton.—Le ms. B 6 ajoute: et en eult cent frans.
F 726.
o

§ 610. Tant esploita.—Ms. d’Amiens: Et avint que li comtes de

Tamcarville et messires Guillaumes de Dormans estoient allé en
Engleterre, de par le roy de Franche, pour parler au roy englès sus
l’estat dou ressort que li roys franchois volloit callengier et maintenir
que point ne l’avoit quitté, mès par le teneur de la cartre de le pais
tenu et reservé. Entroes que li dessus dit estoient en Engleterre, li
rois de Franche fist escripre unes lettres de deffianches au roy
englès, et les fist baillier à ung varlet breton de son hostel, et li fist
dire que il portast en Engleterre ces lettres au roy englès, mès il
fesist retourner, ainschois qu’il les mesist avant, le comte de
Tamkarville et monsigneur Guillaumme de Dormans, et que, au
retour, il aroit cent frans tous appareilliés. Li varlez, pour le
convoitise dou gaegnier, emprist volentiers le voiaige à faire, et dist
qu’il le feroit bien et sagement. Si se parti de Paris et se mist à voie,
et esploita tant qu’il vint à Bouloingne, et passa là le mer et ariva à
Douvres, et là trouva il le comte de Tamkarville et monsigneur
Guillaumme de Dormans, asquelx il dist une partie de sen entente,
et sur quel estat il alloit au roy englès. Si trestost que li dessus dit
l’entendirent, il hastèrent leur passaige et vinrent à Bouloingne, et li
varlés chevaucha vers Londres et fist tant que il y vint.
Adonc estoit li roys englès à Wesmoustier dehors Londres, et là
avoit ung moult grant parlement de chiaux de son pays pour avoir
consseil sour aucunnes requestes que li rois de Franche avoit faittes
par les dessus dis, le comte de Tancarville et monsigneur
Guillaumme des Dormans, et ossi sus l’estat de la duché
d’Acquittainne; car moult desplaisoit au roy englès que ses filz li
prinches s’esmouvoit ne herioit ses gens en Acquittainne; et estoit
bien sen entente que il y pourveroit temprement de remède, car il
ne volloit mies estre en le malevolense de ses subgès, et n’estoit pas
cose bien appertenans.
Enssi que cils parlemens estoit assamblés et que li roys et si doy
fil, li dus de Lancastre et li comtes de Cantbruge, et tout prelat,
baron et chevalier estoient mis enssamble pour parlementer et
conssillier pluisseurs coses, li varlés, qui les deffiances portoit, vint à
l’uis de le cambre et appella l’uissier, et dist qu’il estoit messagiers
au roy de France. Li wuissiers, pour le reverenche de celui dont il se
noumma, li dist: «Entrés ens, et vous tenés dallés moy, et je
regarderay coumment vous parlerés au roy.» Il entra ens et se tint
dallez l’uissier. Assés tost apriès, entra en le cambre li sires de Perssi,
à qui li Englès dist: «Monsigneur, se il vous plaist et il vous viegne à
point, si avanchés ce varlet, qui est, si comme il dist, au roy de
Franche et aporte lettrez au roy.» Li sires de Perssi respondi: «Mout
vollentiers.» Il passa avant et enclina le roy et ses enfans, et puis
tous les signeurs. Assés tost apriès, il dist que là estoit ung
messaiges de par le roy de Franche, qui apportoit lettres. Si tost que
li rois englès oy ce, pour le grant desir qu’il eut de savoir de quoy les
lettres parloient, il dist: «Faittes le avant venir.» On le fist venir. Il
s’agenouilla et bailla ses lettres. Li roys les prist et les ouvri, et puis
les fist lire. Si faisoient mention plainnement coumment li roys de
Franche le deffioit. De ces nouvelles furent li signeur d’Engleterre
tout esmervilliet, et regardèrent li ung l’autre sans point parler. Si fu
li varlés enquis de rechief, et examinés à savoir qui ces lettres li avoit
baillies. Il respondi: «li roys de France.» Adonc le fist on partir de la
cambre, et qu’il se tenist au dehors tant qu’il aroit responsce. Il fist
ce que on li dist. Lors demanda li roys englès consseil sour ce que
vous avez oy. On li consseilla que tantost et sans delay il envoiiast à
Callais, à Ghinnes, à Ardre, et par especial à Abbeville et en le comté
de Pontieu, car elle estoit en grant peril d’estre perdue; et ossi tous
les ostagiers de Franche, qui estoient pour le tamps en Engleterre,
tant baron et chevalier que bourgois des bonnes villes, on les
envoiiast en divers lieus en Engleterre, et fuissent là tenu tout court
em prison. Que vous feroie je loing compte? Chils parlemens finna
ensi, et fu respondut à celui qui les lettres de deffianches avoit
apportées, que il pooit bien partir quant il volloit, et que à ses lettres
ne couvenoit nulle responsce. Il se parti et s’en revint en Franche,
sans avoir nul griefs dou corps. Or vous diray de l’ordounnanche dou
roy de Franche.... Fo 153 vo.
Sitost que on peut savoir ne presummer certainnement que li roys
englès fu deffiiés, et que li comtes de Tamcarville et messires
Guillaummes des Dormans furent revenu à Bouloingne et eurent dit
les nouvelles dou varlet qui les deffianches portoit, li comtes Guis de
Saint Pol, qui estoit tous pourvueus de grans gens d’armes,
chevaucha tout couvertement à grant esploit deviers Abbeville. Si
trouva le porte toutte ouverte et les gens de le ville tout appareilliés
pour lui recepvoir. Si entra li dis comtes de Saint Pol dedens
Abbeville baudement à plus de cinq cens armures de fier, et se saisi
de le ville et dou marchiet, et prist ad ce jour messire Nichole de
Louvaing, qui estoit senescaus de Pontieu de par le roy englès, et le
tresorier dou pays, qui estoit englès, et tous les Englès qui y estoient
à ce jour, et les mist em prison. Et puis chevauchièrent à Saint
Wallery, et le saisirent et du castiel ossi, et puis de Noyelle et dou
Crotoi, et puis de Lonch en Pontieu. Apriès, chevaucha li comtes de
Saint Pol deviers le Pont de Remy; car il entendi que là avoit bien
deux cens Englès qui s’y estoient retret et s’i tenoient sus le comfort
de le fortrèche. Si vint là li dis comtes de Saint Pol et monsigneur
Moriaux de Fiennes, connestables adonc de Franche, avoecq lui, et
messires Hues de Castellon, li sires de Saintpi, li sires de Bremeu, li
sires de Loncvillers, li sires de Bassentin, li sires d’Aveluis, messires
Oudars de Renti, li sires de Reli, messires Engherans du Edins et
pluisseurs autres chevaliers et escuiers, et estoient bien six cens
combatans. Si vinrent droit au Pont de Remy. Là trouvèrent il les
Englès tous aprestés pour yaux attendre et deffendre le passage.
Adonc coummencha là li hustins mout durs et mout fors, et fist là li
comtes de Saint Pol son fil chevalier, monsigneur Wallerant, qui
estoit en l’eage adonc de quinze ans ou environ. Là eut une
escarmuche grande et forte, et maint homme blechié d’un lés et de
l’autre. Finablement, li Englès furent desconfi, et li Pons Remy, sus le
rivière de Somme, pris et gaegniés, et tout li Englès mort ou pris,
petit s’en sauvèrent. Que vous feroie je loinc recort? Tous li pays de
Pontieu fu delivrés des Englès, et les villes et les fortrèches mises et
rendues au comte de Saint Pol, qui y estoit establi de par le roy de
Franche. Si trouvèrent les Franchois le ville d’Abbeville en bon estat,
et bien rappareillie et fortefiiée, car li rois englès y avoit toudis fait
ouvrer; et li avoit li comtés de Pontieu cousté cent mil florins, dessus
touttes revenues, à remparer les villes et les castiaux qui y sont, car
il le tenoit pour son bon hiretaige.
Avoecques les deffiances et les nouvelles dessus dittes qui vinrent
au roy englès et à ses gens, cestes de le perte de Pontieu leur furent
mout diverses. Et se coummenchièrent moult à doubter, et estoient li
pluisseur parmy Engleterre enssi que tout foursené sus les
Franchois, qui pour ostaiges demouroient entre yaux; mès li roys fist
faire un ban, et sus le hart, que nuls ne fesist mal as Franchois, qui
là estoient: autrement il n’ewissent nient duret. Si se
ranchounnèrent li baron de Franche au plus tost qu’il peurent et
trouvèrent le roy englez assés courtois, et rappassèrent le mer; et
ossi les bonnes villes et les chités de Franche rachatèrent leurs
bourgois. Fo 154.
P. 109, l. 31: Evous.—Ms. A 8: Et vont. Fo 302.
P. 110, l. 1: donna.—Ms. A 8: donnèrent.
P. 111, l. 4: nequedent.—Ms. A 8: neanmoins.
P. 111, l. 12: quatre cens.—Mss. A 7, 8: trois cens. Fo 297.
P. 111, l. 26: six cens.—Mss. A 7, 8: six vingt.
P. 112, l. 14: Pont de Remi.—Ms. A 8: Pont de Saint Remy.
P. 112, l. 17: Gallerans.—Ms. B 6: Wallerant de Ligny, aysné filz
audit conte. Fo 730.

§ 611. Quant li rois.—Ms. d’Amiens: Sitos que li dus d’Ango peut

connoistre et sentir que li rois englès estoit deffiiés, si en fu moult
joieaus, et dist que il feroit au prinche et à ses gens une très forte
guerre; car point ne lez amoit. Si avoit ja de son acord pluisseurs
chevaliers et escuiers de Gascoingne, et ossi bons cappitainnes des
Compaignes. D’autre part, se tenoit à Rion en Auvergne li dus de
Berri, qui metoit sus grant gens d’armes, telx que le signeur de
Biaugeu, le signeur de Perreus, son nepveult, monsigneur Joffroi de
Bouloingne et monsigneur Griffon de Montagut, monsigneur Robert
Daufin, monsigneur Huge Daufin, le signeur de Calenchon, le
signeur de la Tour, monsigneur Jehan de Bouloingne, le comte de
Ventadour et de Montpensé, le signeur de Sulli, le signeur d’Achier, le
signeur d’Achon, le signeur de Gonsaut, Ambaut dou Plachier et
tamaint chevalier et escuier d’Auviergne et des marches voisinnes. Si
entrèrent ces gens d’armes en le duché d’Acquittainne, et
coummenchièrent à prendre, à pillier et à rober, et à chevaucher
sour le pays dou prinche et à faire mout de maux. D’autre part,
couroient ossi et chevauchoient en Roherge messires Jehans
d’Ermignach, messires Jehans de Villemur, Rogiers de Biaufort, li
sires de Rocefort, li sires de Seregnach, et mettaient le pays en grant
misère de Roherge et de Limozin. Fo 154 vo.
P. 113, l. 14: remparer.—Mss. A 7, 8: reparer. Fo 303.
P. 113, l. 15: deseure.—Ms. A 8: pardessus.
P. 114, l. 7: l’archeveskié.—Ms. A 8: l’eveschié. Fo 303 vo.
P. 114, l. 23: Keranloet.—Mss. B 1 et B 4: Karuel.—Ms. A 8:
Carnet.—Ms. A 7: Kaerenloet. Fo 197 vo.

§ 612. Li dus de Lancastre.—Ms. d’Amiens: Vous avés bien chy

dessus oy recorder coumment les Compaignes avoient estet en
Franche, et ivernet et ostoiiet, siques, sitost que li Gascon
entendirent que li roys de Franche vorroit gueriier, il se partirent des
Englès. Si estoit li plus grans chiés de ces Compaignes gascons
messires Perducas de Labreth, et des Compaignes englesses,
messires Robers Bricqués et Jehans Carsuelle. Si se traissent li
Gascon deviers le ducq d’Ango, et estoient de leur routte li Petis
Meschins, li bours de Bretoeil, Ammenion de Lortige, Perrot de
Savoie, Jakes de Bray, Ernauton de Paus et pluisseur aultre, et
estoient bien troi mil combatans. Li Englès qui s’estoient
descompaigniet d’iaus, pooient estre environ quinze cens
combatans, et s’estoient retret deviers Normendie et par deviers les
Mans, et avoient pris Castiel Ghontier et une bonne ville que on
appelle le Vire, et pluisseurs autres fortrèches, dont il guerioient et
destruisoient tout le pays et y faisoient moult de maux. Dont il avint
que li prinches les remanda et leur fist asavoir que il vendesissent
leur forterèce et se retraissent deviers lui. Si trestost que il oïrent ces
nouvelles, il furent tout joieant et eurent grant desir de obeir au
coummandement du prinche. Si se delivrèrent de tout ce qu’il
tenoient, au plus bellement qu’il peurent, et se requeillièrent
enssamble pour passer plus efforchiement. En ce tamps que ces
routtes yssoient hors de Normendie, il avoient à ciaux du pays vendu
Castiel Ghontier et le Vire et tout chou que il y tenoient.
Et arivèrent au port de Saint Malo de l’Ille messires Aimmons,
comtes de Cantbruge, et li comtes de Pennebrucq et li sires de
Carbestonne, ungs banerés englès, à quatre cens hommes d’armes
et quatre cens archiers; et les envoieoit li roys englès en la duché
d’Acquittainne deviers le prinche, pour lui aidier à faire sa gerre. Si
oïrent nouvelles assés tost li uns de l’autre, dont il furent moult
joieant; car il dissent qu’il passeroient tout enssamble, si comme il
fissent. Et envoiièrent tantost li doy comte dessus noummet deviers
le ducq de Bretaingne, qui se tenoit à Nantes, et deviers le consseil
dou pays, em priant que on les volsist laissier passer paisiulement
parmy Bretaigne, bien paiiant tout ce que il y prenderoient. Li
messagier que li dit seigneur y envoiièrent, esploitièrent si bien, que
li passaiges leur fu ouvers et acordés par l’acord dou duch, qui ne le
fist mies envis, et de tous les barons dou pays, et passèrent
paisiulement parmy Bretaigne et au pont à Nantes. Et rechupt les
seigneurs li dis dus moult grandement, et les festia par deux jours
en le chité de Nantes; et puis s’em partirent et chevauchièrent et
esploitièrent tant que il entrèrent en Poito et vinrent en Anghouloime
deviers le prinche, qui les rechupt à grant joie, et ossi fist madamme
la princesse.
En ce tamps, estoit messires Hues de Cavrelée sus le marche
d’Arragon, à une routte de gens d’armes, de quoy, sitost que il
entendi que li Franchois guerioient le prinche, il se parti et s’en vint
en Angouloime deviers lui, et amena ce qu’il avoit de gens. Se le
rechupt li prinches à grant joie, et le fist tantost cappittainne et
souverain de touttes les Compaingnes, qui estoient nouvellement
venues de Normendie. Si le envoiea li dis prinches, et toutes ces
gens d’armes, en le terre le comte d’Ermignach et le seigneur de
Labreth, pour ardoir et destruire leur pays et faire y guerre, car chil li
estoient grant ennemit. Fo 155.
P. 115, l. 21: d’Ortige.—Ms. A 8: d’Ortinge. Fo 303 vo.
P. 115, l. 21: Perros.—Ms. A 8: Pierre.
 P. 115, l. 21: Jakes.—Ms. A 8: Raoul.
P. 115, l. 28: Rochewart.—Ms. A 8: Rochechouart.
P. 116, l. 6: d’arciers.—Ms. A 8: d’arbalestriers.—Ms. B 6: et plus
de trois mille combatans. Fo 730.
P. 116, l. 15: li dus de Bretagne.—Les mss. B 4 et A 7, 8 ajoutent:
messires Jehans de Montfort. Fo 301 vo.
P. 116, l. 19: Lagnigay.—Ms. A 8: Lagingay.
P. 117, l. 1: rihote.—Ms. A 8: riote.
P. 117, l. 31: deux mil.—Ms. B 6: bien mille. Fo 731.

§ 613. Li contes de Cantbruge.—Ms. d’Amiens: Encorres envoiea li

prinches son frère monsigneur Aimenon et le jone comte de
Pennebrucq, à tout grant fuisson de gens d’armes, en le comté de
Pieregorth. Si chevauchièrent chil seigneur en grant routte, et
estoient bien troi mil combatans, uns c’autres, et entrèrent en grant
effort en le comté dessus dite et vinrent mettre le siège devant ung
très fort castel que on claimme Bourdille. Si l’environnèrent tout
autour. Par dedens estoient en garnison li doy enfant de Batefol,
hardi homme durement et bons guerieurs, frères bastars à
monsigneur Seghin de Batefol, dont j’ay parlé chy en avant en
l’istoire.
Si estoient li doy escuier pourvueu mout bien dedens Bourdille de
toutte artillerie, d’espringalles, de kanons et d’ars à tours, et de bons
compaignons hardis et sceurs, pour le deffendre et tenir, et si
avoient assés par raison de quoy vivre. Si vous di que devant
Bourdille eut tamaint assaut, mainte eskarmuce et maint puignies, et
priesque tous les jours. Fo 155.
P. 118, l. 25 et 26: environ quatre jours.—Ms. A 8: trois jours.
F 304 vo.
o

P. 118, l. 27: d’aler en le Gascongne.—Ms. A 8: de partir

d’Angoulesme.
P. 119, l. 8: exillier.—Ms. A 8: assaillir.
 § 614. En le garnison.—Ms. d’Amiens: Enssi estoient les guerres
efforchies de tous costés ens ou royaumme de Franche, car messires
Jehans de Buel, uns très bons chevaliers bourghignons, et messires
Guillaummes des Bordes et Caruels estoient sour les marches de
Poito, à plus de quinze cens combatans, et faisoient là une moult
forte guerre environ Chastieleraut, qui se tenoit de monsigneur Loeis
de Halcourt, et coururent mout du plain pays de Poito à l’encontre
d’yaux. Ossi de par le prinche estoit uns bons chevaliers englès et
grans cappitainnes de gens d’armes, messires Simons de Burlé, qui
deffendoit et gardoit le pays ce qu’il pooit, et chevauchoit à le fois
sus les Franchois, et li Franchois sour lui, et avoient souvent des
durs rencontres: à le fois gaegnoient li ung, et puis li autre. Dont il
avint que messires Jehans de Buel et messires Guillaumes des
Bordes et Caruels, Bretons, et leurs routtes chevauchoient un jour; si
trouvèrent, entre Mirabel et Luzegnan, monsigneur Simon de Burlé
et ses compaignons. Là eut dur hustin et fort et bien combatu, et
pluisseurs reverssés d’un lés et de l’autre. Toutteffois, li Franchois
s’efforchièrent si, et si vaillamment se combatirent, que par force il
reculèrent les Englez et missent en cache. Et couvint messire Simon
fuir, et fu si dur et si roit encauchiet que, au destroit d’un passage
d’une desroute cauchie qui là estoit, il fu ratains, et trebuça ses
courssiers, et chei. Si fu tantost environnés de touttes pars, assaillis
fierement et requis que il se volsist rendre, ou autrement il estoit
mors. Quant messires Simons se vi à terre et en ce parti à tel
meschief, et que deffensce n’i valloit riens, si se rendi et fiancha
prison à monsigneur Jehan de Buel. Si retournèrent li Franchois à
grant joie, qui eurent le journée devant yaux, et ramenèrent leurs
prisonniers à sauveté. De le prise monsigneur Simon de Burlé fust li
prinches courouchiés, mès amender ne le peut tant c’à ceste fois.
Fo 155.
P. 119, l. 27: hardi.—Le ms. A 8 ajoute: entreprenans. Fo 304 vo.
P. 119, l. 28: amiroient.—Ms. B 4: cremoient. Fo 302.—Ms. A 8:
amoient.
P. 120, l. 8: Carenloet.—Mss. B 4 et A 8: Charnet, Carnet.—Ms. A
7: Jehan Kaeranloet. Fo 299.
 P. 121, l. 6: sept cens combatans.—Ms. B 6: quinze cens lanches.
Fo 732.
P. 121, l. 13: li aucun.—Ms. A 8: les Anglois.
P. 121, l. 17: puignie.—Ms. A 8: poingniée.
P. 121, l. 21: sievois.—Ms. A 8: poursuis.
P. 121, l. 30: moult.—Le ms. A 8 ajoute: il prisoit et. Fo 305.

§ 615. Apriès ceste avenue.—Ms. d’Amiens: Vous avés bien chy

dessus oy compter coumment messires Jehans Camdos se tenoit à
Montalben, et messires Loeys de Halcourt, messire Loeys de Melval,
monsigneur Rainmon de Maruel, li sires de Pierebufière, li sires de le
Ware, li captaux de Beus, li sires de Lespare, li soudis de Lestrade,
messires Thummas de Felleton et li doy frère de Pummiers et
pluisseur bon chevalier et escuier, et gardoient le frontière contre les
Franchois. Si faisoient souvent des yssues et des chevauchies d’un
lés et de l’autre, et ne demandoient autre cose que il pewissent
trouver leurs ennemis. Si partirent un jour de Montalben en grant
arroy et chevauchièrent deviers Toulouse, et vinrent mettre le siège
devant un fort castiel, que on appelle Terrières. Si l’environnèrent de
tous costés, et puis ordounnèrent et coummandèrent as mineurs qui
là estoient, qu’il s’aprestassent et se missent em painne et en
pourcach de l’avoir par mine. Li mineur, qui sont coustummiers et
usés de chou faire, eurent tantost adviset là où il coummencheroient
leur minne. Si abillièrent leurs instrumens et minèrent vistement et
fortement, et fissent grans petruis par desoubs lez murs. Avoec tout
ce, quant li chevalier sentirent que leur ouvrier estoient au dessus de
leur ouvraige, il se missent à assaillir chiaux de dedens. Là eut grant
assaut fort et bien ordounné, mès finablement chil qui estoient en le
mine, entrèrent par desoubz terre en le villine; et ensommèrent
tellement les deffendans, qu’il les reboutèrent arrière des murs, et
perdirent tout arroy et ordounnanche de deffendre, et entrèrent ens
li assallans par force. Si fu la ville de Terrières prise et gaegnie,
toutte pillée et robée, et y eut moult de gens ochis. Quant li Englès
en eurent fait leur vollenté, il s’en partirent et s’en revinrent arrière à
Montalben.
Assés tost apriès, fissent il une autre chevauchie, et avoient espiiet
et adviset le bonne ville c’on dist Laval, à trois lieuwes près de
Toulouse. Si avoient laissiet une grosse embusque en un bois,
environ demy lieuwe enssus de le ville, et yaux six vingt armés
couvertement et en cotes de vilains, et en venoient tout devant, et
fuissent sans faute entré en le ville; mais il furent descheu par l’un
de leurs compaignons, de qui la coute de fier passoit et appairoit
dessous sa coute de villain; et le perchupt un varlez dou pais, qui
venoit piet à piet avoecq yaux, siques, quant il durent approchier la
porte, il se mist au cours tout devant, et dist as gardes de le porte:
«Cloés, cloés, seigneurs! Traï! traï! Veci les Englès.» Si cloirent chil
tantost le porte et sounnèrent leur cloche et se missent as murs et
as deffensces de la ville. Par enssi fallirent li Englès à leur emprise,
dont il furent moult courouchiet, et retournèrent arrierre à
Montalben, dont il s’estoient parti.
En ce tamps, chevauchoient adonc, dou costé des Franchois, li
viscomtes de Quarmaing, li comtes de Comignes, li comtes de Laille,
li comtes de Pieregorth, li comtes de Murendon, li comtes de Talar, li
viscomtes de Brunikiel, les gens le seigneur de Labreth, et le comte
d’Ermignach, li sires de la Barde, li sires de Tharide, li sires de
Picornet, et les Compaingnes: messires Perducas de Labreth,
messires Berardet de Labret, messires Garcis dou Castiel, le Petit
Mescin, Janikot, d’Orteine, Lamit, le bourch de Tarse, le bourch de
Bretuel, et vous di que à ce donc il tenoient les camps. Si entrèrent
en Quersin, gastant et essillant le pays, et s’en vinrent devant
Roiauville en Quersin; si l’asegièrent. Par dedens avoit aucuns bons
escuiers englès et archiers, que li senescaux de Quersin y avoit
estaublis, qui jammès ne se fuissent rendu, quoyque les Englès de le
ville en fuissent bien en vollenté, mais dissent qu’il se tenroient bien
et vaillamment. Quant li Franchois furent venu devant Royauville, si
l’asegièrent de tous lés et dissent bien qu’il ne se partiroient mies
enssi. Si assaillirent chiaux de dedens fortement, et fissent drechier
grans engiens devant le ville, qu’il faisoient acariier avoecq yaux, qui
estoient de la chité de Thoulouse. Là eut, je vous di, par pluisseurs
jours, moult de grans assaus et de belles appertisses d’armes faittes.
Touttefois finablement, à un grant assaut qui fu entre les autres, li
Franchois assaillirent si ouniement et si bien continuèrent que de
forche ils prissent Royauville. Et furent tous li Englès qui dedens
estoient mort et ochis, sans nul prendre à merci. Et fissent jurer as
hommes de le ville que, de ce jour en avant, il seroient bons
franchois et loyal, et le jurèrent et eurent en couvent; et furent tout
joiant, quant par enssi il peurent escaper. Fo 155.
P. 122, l. 7: Montalben.—Le ms. B 6 ajoute: et estoient bien douze
cens lanches et trois mille combatans, parmy les archiers. Fo 732.
P. 122, l. 29: desous.—Ms. A 8: couvertement. Fo 305 vo.
P. 122, l. 31 et 32: aultrement.... Montalben.—Ms. A 8: lequel
descouvri la besoingne, et par ce ilz fallirent à avoir la ville et à leur
entente, et s’en retornèrent arrière à Montalben.
P. 123, l. 8: d’Ortige.—Ms. A 8: d’Ortinge.
P. 123, l. 9: Paus.—Ms. A 8: Pans.
P. 123, l. 10: dix mil.—Ms. B 6: plus de douze mille. Fo 734.
P. 123, l. 22: peuissent.—Ms. A 8: eussent peu.

§ 616. Endementrues.—Ms. d’Amiens: Entroes que ces gens

d’armes se tenoient sour le pays et chevauchoient tant d’un lés
comme de l’autre, se parti de Thoulouse li arcevesques de la ditte
chité, par le promovement dou duc d’Ango qui là se tenoit. Et s’en
vint en le chité de Chaours, dont ses frerres estoit evesques, qui le
rechupt liement. Chils arcevesques de Thoulouse preecha la querelle
dou roy de Franche si bellement et si sagement, et si volentiers
l’oïrent chil de Chaours, que briefment il se tournèrent et
relenquirent le prinche et les Englès, et jurèrent solempnelment à
estre bons françois et loyal. Apriès, chevaucha li dis arcevesques
vers Villefranche de Quersin, et preecha, en la ditte ville, la querelle
dou dit roy de Franche: si se tourna ossi la ditte ville et devint
franchoise. Et puis chevaucha vers Rodès, et le fist tourner, et
Figach, Gramach, Rocemadour et Capdonach. Et fist li arcevesques
de Toulouse retourner franchoises plus de soissante villes, cités et
castiaux, parmy le comfort de monsigneur Jehan d’Ermignach, de
monsigneur Jehan de Villemur, de Rogier de Biaufort, dou signeur de
Seregnac, qui chevauchoient et tenoient sour le pays grant routte.
Enssi estoient ces terres que je vous nomme, en grans variemens;
ne meysmement les chités, les villes ne li castiel ne savoient que
faire pour le milleur, ne li plus des gens dou pays ne savoient liquel
avoient droit ou tort: si estoient tout en grant branlle et vivoient en
grant tribulation.... Fo 155.
Or revenrons au roy d’Engleterre, qui fu durement courouchiés des
nouvelles qu’il avoit oyes et de celles encorres que il ooit tous les
jours, de che que on li tolloit et prendoit enssi le pays qui se tenoit
pour sien. Si envoiia tantost grans gens d’armes à Callais, à Ghinnes
et à Ardre, pour deffendre et garder les frontierres contre les
Franchois; car li comtes de Saint Pol, qui li estoit mout grans
ennemis, se tenoit à Saint Omer à plus de mil combatans, et
couroient li Franchois tous les jours jusques à Ardre. Et si envoiea
encorres li dis roys englès grant fuisson de gens d’armes sus les
mettes d’Escoche, à Bervich et à Rosebourch; car il se doubtoit que li
Escos ne se revellassent contre li et sus son pays, et que li rois
d’Escoce n’ewist fait nouvelles alianches au roy de Franche.
Encorres envoiea grans messaiges li roys englès deviers son
nepveut monsigneur Edouwart de Guerlle, en lui remoustrant, et
complaindoit des tors que li rois de Franche li faisoit, et que, par
linaige et pour droit aider à soustenir, il ne li vosist mies fallir.
Messires Edouwars de Guerlles et li dus de Jullers, ses serourges,
eurent en couvent au roy englès que il le serviroient et aideroient à
mil lanches contre le roy de Franche et les Franchois, et deffiièrent
cil doy seigneur tost et appertement le roy de Franche. Quant li rois
de Franche s’en vit deffiiés, il trouva voies et pourcach pour yaux
guerriier et ensonniier dou duc de Braibant, son oncle, et dou comte
de Clèves et de aucuns seigneurs d’Alemaigne qu’il atraist à son
acord.
Encorres envoiea grans messaiges li roys englès deviers sa
cousinne madamme Jehanne, duçoise de Braibant, en lui
complaindant des tors et des injures que li roys de Franche li faisoit.
Et prioit li roys à sa cousinne, à tout le mains, se elle ne ses pays ne
volloient estre de son acord ne tenir lez allianches de jadis, que li
dus de Braibant, ses cousins et pères à le dite damme, et li pays de
Braibant avoient juret et saiellet, elle ne fust mies ennemie ne
contraire à lui. Li chevalier qui envoiiet y estoient, messires Richars
Sturi et messires Thummas Rock, de par le roy englès, esploitièrent
si bien que la duçoise de Braibant eut en convent que elle ne ses
pays ne se mouveroit de ceste guerre. Et enssi eut li dus Aubers, car
grans messaiges li furent envoiiet.
Encorres escripsi fiablement li roys englès et manda à ce jentil
chevalier monsigneur Robert de Namur, que il fuist tous appareilliés
de venir deviers lui, quant il seroit mandés, et qu’il retenist
chevaliers et escuiers de tous lés, car il les paieroit et deliveroit
thous. Ces nouvelles pleurent moult bien à monsigneur Robert de
Namur, et festia et requeilli liement lez messaiges dou roy englès, et
leur dist que il estoit tous prês, quant li rois le vorroit mander, et
deux cens ou trois cens armures de fer en se compaignie. Ensi se
pourveoient li doy roy de gens d’armes, li uns d’un costé et li autre
d’autre, et prioient et requeroient leurs amis partout où il les
penssoient à avoir... Fo 154.
Se li roys Carles de Franche faisoit grant appareil par terre et par
mer pour gueriier le roy d’Engleterre, li roys englès otant bien se
pourveoit à l’autre lés, et fist tant que ses nepveux messires
Edouwars de Guerle, qui avoit à espeuse l’aisnée fille à monsigneur
le duc Aubert, deffia le roy de Franche et proummist à son oncle le
dit roy et ses cousins, ses enfans, que il feroit une très grande
guerre et forte en Franche; et avoit de son acort son serourge le duc
de Jullers, et devoient y estre tout doy de une alianche et d’une
yssue, et devoient mettre sus mil lanches de droite gens d’armes,
bien montés et bien armés.
Encorres escripsi li roys englès et envoiea grans messaiges deviers
ce gentil chevalier monsigneur Robert de Namur, en lui priant et
amonestant que il se pourveyst grandement, seloncq son estat, de
chevaliers et d’escuiers et de gens d’armes. Messires Robers
respondi qu’il estoit tous prês, quant il plairoit au roy ou à son fil le
duc de Lancastre qu’il trayst avant.
Entroes que ces besoingnes s’ordounnoient et que cil roy se
pourveoient enssi par terre et par mer, et que chacuns acqueroit
amis là où il les pooit ne penssoit à avoir, se avisa encorres li dis roys
de Franche que il envoieroit as barons de l’Empire, especialement en
la duché de Braibant et en le comté de Flandres et de Haynnau, de
Hollandes et de Zellandes, la certainne teneur de le chartre de le
pais qui jadis fu faitte et acordée à Bretegny, priès de Cartres, pour
mieus emfourmer les seigneurs en quoy li roys d’Engleterre, par son
seellé, et si emfant, estoient obligiet et aloiiet. Si le fist coppiier em
pluisseurs coppies, et en envoiea au ducq de Braibant, son oncle, au
ducq Aubert, son cousin, au comte de Clèves, à messire Jehans de
Blois, et enssi as barons et as conssaux des seigneurs, laquelle
chartre de le paix parloit enssi: «Edouwars, etc.[342]»
Encorres estoit escript en ceste grosse lettre une autre lettre
obligatoire tretians sus fourme de commission, envoiie dou dit roy
Edouwart d’Engleterre par ses marescaux, de fortrèce en fortrèce,
apriès le pais faite traitie à Bretegny dallés Chartrez, et confremée à
Calais; et en faisoit faire li dis roys de France mention, pour tant que
il volloit que on sewist clerement se li roys d’Engleterre et li prinches
de Galles, ses fils, avoient bien tenu et acompli ce que il avoient juré
et promis par leur seellé, laquelle teneur de la ditte coppie s’ensuist
ensi: «Edouwars, etc.[343]»
Si prioit li roys Carles de Franche humblement à tous les seigneurs
dessus dis et à leurs conssaux que ilz volsissent, à grant loisir, lire ou
faire lire ces presentes lettres, et regarder et ymaginer sus et bien
examiner de poinct em poinct, car il juroit et prommetoit en se
loyauté que oncque li roys d’Engleterre ne ses fils li prinches de
Galles, par especial, n’en avoient mies tenu ne acompli le dizime
partie: pour quoy il disoit que ilz avoient allé et erré contre leur
sierement, et enfraint et brisié le pais sans nul title de raison, et tenu
et envoiiés gens d’armes et Compaignes sus le royaumme de
Franche, et avoient retenu pluisseurs cappittainnes et autres des
Compaingnes, tant Englès comme Gascon, qui avoient estet pris ou
royaumme de Franche. Quant on les faisoit morir pour leurs villains
fais, tels que le bourch Camus, le bourch de Bretuel, Espiotte,
Batillier et Jehan le Nègre, sus leur mort, il confessoient que li
prinches de Galles les avoit envoiiés, et envoieoit ains le guerre
ouverte encorres tous les jours. Si devés sçavoir que telx parolles et
moustranches que li roys de Franche moustroit et declaroit,
coulouroient mout ses besoingnes. Si les fist il preechier, publiier et
remoustrer notoirement et generaument parmy son royaumme par
monsigneur Guillaume de Dormans, qui bien le savoit faire, et
pluisseurs autres prelas et ses offisciers, ydosnes et propisses à ce
faire; et en signe d’umelité et en cremeur de Dieu, il en faisoit faire
pourcessions publicques, et ils meysmes et la roynne de Franche y
aloient en grant devotion, tout à nus piés. D’autre part, li roys
Edouwart d’Engleterre, en son pays et par dechà le mer, as
seigneurs de son acord et à li alyé se complaindoit trop grandement
dou roy de Franche, et moustroit voies de droit et de raison, qui ooit
et entendoit ses lettres et ses proches. Et pour ce, se il envoieoient
enssi des uns as autres, ne se laissoient il mies à pourveir et à
gueriier l’un l’autre par mer et par terre. Fos 162 à 164.
P. 124, l. 8 à 13: telz que.... gens d’armes.—Ms. B 6: messire
Jehan d’Erminach, messire Jehan de Villemur, le sire de Bieaugieu, le
sire d’Alençon, messire Jehan de Boullongne, messire de Sailly,
messire Robert de Sansoire, marisal de Franche, ou lieu de messire
Ernoul d’Audrehem qui estoit nouvellement mort à Paris sus son lit.
Fo 734.
P. 124, l. 11: Serignach.—Ms. A 8: Sergnac. Fo 306.
P. 124, l. 15: apovrissoient.—Le ms. A 8 ajoute: dommagoient.
P. 124, l. 19: leurs gens.—Ms. A 8: le duc de Berry et ses gens.
P. 124, l. 25 et 26: si bellement.—Ms. A 8: par si bonne manière.
P. 124, l. 30: la querelle.—Ms. A 8: le bon droit.
P. 125, l. 4: Gramach.—Ms. A 8: Gramat.
P. 126, l. 18: envoiiet.—Ms. A 8: enjoint.
P. 126, l. 24: lés.—Ms. A 8: pais.
 P. 127, l. 30 et 31: soutieus.—Mss. A 7, 8: soubtilz. Fo 300 vo.
P. 128, l. 10: eu.—Ms. A 8: sur ce.
P. 128, l. 12: cambres.—Le ms. A 8 ajoute: et compaingnies.
P. 129, l. 4: trau.—Ms. A 7: dommage.—Ms. A 8: cam.
P. 129, l. 4: vint.—Ms. A 7: neuf.

§ 617. Vous avés.—Ms. d’Amiens: En ce tamps, envoiea li comtes

Loeis de Flandres grans messaiges en Engleterre, pour requerre au
roy que il le volsist quitter d’aucuns couvens que il avoient
enssamble, pour le cause dou mariaige dont j’ay parlé chy dessus en
ceste histoire, de la fille dou dit comte et dou fil dou dit roy, le comte
de Cantbruge: lequel mariaige pappes Urbains ne vot oncques
dispensser, et cousta au roi d’Engleterre li pourcach très grant avoir,
mais li pappes avoit dit et juret que, pour lui detraire as chevaux, il
ne le dispensseroit ja. Et qant li roys englès vit chou que il n’en aroit
aultre cose, enssi comme tout tannés il quitta le comte de Flandres
et la damme ossi. Si vous di que, sitost que les quittanches furent
faittes, li mariaiges fu fais, car il estoit ja tout tretiés de celle
damme, fille au comte Loeis de Flandres, et de monsigneur Phelippe,
ducq de Bourgoingne, maisné frère dou roy Carle de Franche, parmy
tant que ly roys de Franche rendi et quitta tout liegement au comte
de Flandres et à le comtet, et à tousjours mès, Lille et Douay et
touttes lez appendanches; et encorres eut li comtes de Flandres,
pour lez frès de lui et de ses gens, six vingt mil frans franchois. Si
espousa li dis dus de Bourgoingne la fille au comte de Flandres en
l’abbeie de Saint Piere de Gand, et là eut grans festez et nobles et
mout de signeurs, et y jousta on par trois jours. Ce fu environ le
Saint Jehan Baptiste, l’an de grasce mil trois cens soissante et neuf.
Or revenrons as besoingnes d’Acquittainne. Fo 154 vo.
P. 129, l. 25: paraus.—Mss. A 7, 8: pareil. Fo 301.
P. 129, l. 28: Cil.—Le ms. A 8 ajoute: qui envoiez y furent.
P. 130, l. 5: li contes.—Le ms. A 8 ajoute: de Flandres.
P. 130, l. 12: wage.—Ms. A 8: gage.
 P. 130, l. 19: affreoit.—Ms. A 8: afferoit.
P. 130, l. 31 et 32: un petit plus frans et plus fors.—Ms. A 7: un
petit plus frans et plus durs.—Ms. A 8: plus dur et plus fel. Fo 307 vo.
P. 130, l. 32: grignes.—Ms. A 8: griefz.

§ 618. Li rois Edowars.—Ms. d’Amiens: En ce tamps, passa li roys

Carles de Navarre outre en Engleterre, et trouva le roy englès et le
duc de Lancastre, son fil, et une partie du consseil d’Engleterre en
l’ille de Cepée, en ung mout biel castiel que li roys englès avoit là fait
faire et fonder nouvellement sus le rivière de Tamise, assés priès de
Cantorbie. Si eurent chil doy roy parlemens, tretiés et aliances adonc
enssamble, et devoit li roys de Navarre faire guerre au roy de
Franche, lui revenut en Normendie. Enssi le proummist il au roy
englèz et l’eut en couvent; si s’en retourna arrière en son pays à
Chierebourch, où il se tenoit. Et le raconvoiièrent les gens dou roi
englès, tant qu’il fu là où il volloit y estre, asquelx il n’ezchei mies
trop bien à leur retour; car, ensi qu’il s’en ralloient en Engleterre, il
encontrèrent quatre nefs de Normans à qui il leur couvint parler. Si
furent assailli li Englès qui là estoient, mort et desconfit une nef tant
seullement; mais petit y avoit de gentilz hommez, fors que gens
d’offisse. Si desplaisoit il mout au roy englès, quant il oy recorder ces
nouvelles que ses gens avoient eu ung si dur rencontre. Fo 162.
P. 131, l. 12: lés.—Ms. A 8: costez. Fo 307 vo.
P. 131, l. 26: se tenoit.—Ms. A 8: se tenoient.
P. 131, l. 27: partout.—Ms. A 8: par toutes.
P. 132, l. 20: moult mal.—Les mss. A 15 à 17 ajoutent: car
endementres que le roi de Navarre, qui nouvellement estoit venu
d’Engleterre de parlementer avec le roi, si comme j’ai dit ci devant,
festioit ces chevaliers d’Engleterre qui raconduit et ramené l’avoient,
seurent aulcuns Normans et Bretons et aultres escumeurs de mer,
ceste avenue du roi de Navarre et des Englès, et comment il s’en
devoient tantost retourner en Engleterre. Si s’ordonnèrent et misent
en aguet sus mer, et assés tost rencontrèrent ces chevaliers
d’Engleterre qui parti estoient de Chierebourch et du roi de Navarre
et s’en retournoient en leur pays ne point ne se donnoient de garde.
Si rencontrèrent.... Fo 338 vo.
P. 132, l. 32: querre.—Ms. A 8: querir.
P. 133, l. 2: donna.—Le ms. A 8 ajoute: congiet.
P. 133, l. 14: le captal.—Le ms. A 8 ajoute: de Beuch.

§ 619. En ce temps.—Ms. d’Amiens: Si comme nous avons parlé

ung grant tamps des besoingnes et des avenues qui avinrent adonc
en Ango, em Poito, en Tourainne, en Saintonge, en le Rocelle et en
le Langhe d’Ok, et des rebellions des villes, des chités et des
castiaux, et ossi des gentilx hommes qui s’esmurent et se levèrent
contre le prinche d’Acquittainne et se tournèrent franchois, nous
couvient parler des avenues qui avinrent en celle saison en Picardie,
environ Saint Omer, Arde, Ghines et Calais. Si vous di que li roys
Carles de Franche avoit si grossement et si grandement pourvueu
ces frontières de bonnes gens d’armes, que on ne pooit entrer ne
chevauchier ou pays de Pikardie, fors à grant routte; et en estoit
cappittainnes et mestres souverains li comtes de Saint Pol, qui tenoit
plus de mil combatans ens ès fortrèches et sus les frontières par les
garnisons. Et estoit à ce donc cappittainnes de Boulongne li sires de
Saintpi, qui de garder son frontière faisoit mout bien son devoir. Si
s’esmurent en cel estet tout li chevalierz et escuier d’Artois, et de
Haynnau aucuns, et par especial messires Jehans de Werchin,
senescaux de Haynnau, li sires de Floion, et li connestables de
France pour le temps, messires Moriaux de Fiennes, et s’en vinrent
devant le bastide d’Arde, et s’y ordounnèrent et appareillièrent pour
l’assaillir. Et là eut à ceste assamblée plus de cinq cens chevaliers et
escuiers franchois et leurs gens; mès noient n’y fissent. Si furent il
par devant je croy cinq jours, et y fissent aucunes escarmuches, mès
riens n’y conquestèrent. Quant il virent que il ne pooient riens
gaegnier à assaillir Arde et que elle estoit trop bien gardée et
pourvueue, si s’avisèrent que il se retrairoient en leur fortrèce et
guerriroient par garnisons. Si se departirent, et s’en ralla chacuns là
où il estoit ordounnés pour li tenir, et li senescaux de Haynnau s’en
alla en Franche deviers le roy. Fo 161 vo.
 P. 133, l. 20: d’Arde.—Ms. A 8: d’Ardre. Fo 308.
P. 133, l. 23: seneschaus.—Ms. A 8: connestable.
P. 134, l. 5: frice.—Ms. A 8: frique.

§ 620. Nous revenrons.—Ms. d’Amiens: Là (devant Royauville)

eut, je vous di, par pluisseurs jours moult de grans assaus et de
belles appertisses d’armes faittes. Toutteffois finablement, à un grant
assaut qui fu entre les autres, li Franchois assaillirent si ouniement et
si bien continuèrent que de forche il prissent Royauville, et furent
tout li Englès qui dedens estoient mort et ochis, sans nul prendre à
merci. Et fissent jurer as hommes de le ville que, de ce jour en
avant, il seroient bons Franchois et loyal, et le jurèrent et eurent en
couvent, et furent tout joyant, quant par enssi il peurent escaper.
Et toudis se tenoit en la Millau li senescaux de Roherge, messires
Thummas de Wettevale, quoique li pays d’environ lui se tournast
franchois, et faisoit souvent des yssues et des chevauchies moult
honnerables pour lui; et tenoit encorres une fortrèche, que on
appelloit le Roche Vauclère, où souvent chevauchoit de l’un à l’autre,
pour rafreschir ses gens et yaux renouveller de comfort et de corage
et regarder coumment il leur estoit. Or retourrons nous au siège de
Bourdille, en le comté de Pieregorth, qui ne fait mies à oubliier.
Fo 155 vo.
P. 134, l. 19: gens.—Le ms. A 8 ajoute: d’armes.
P. 134, l. 25: besongnast.—Ms. A 8: besoing eust esté.
P. 135, l. 1: Pennebruch.—Ms. A 8: Pennebroch.
P. 135, l. 1: seoient.—Ms. A 8: estoient à siège.
P. 135, l. 22: Paus.—Ms. A 8: Pans.
P. 135, l. 24: Laille.—Mss. A 7, 8: Lisle.
P. 136, l. 3: Wettevale.—Ms. A 8: Witevale.
P. 136, l. 8: Roerge.—Ms. A 8: Bretaingne.
P. 136, l. 11: jusques adonc.—Ms. A 8: jusques à ce.
 § 621. Sur les marces.—Ms. d’Amiens: En ce tamps, si comme je
vous ay ja dit chy dessus, estoient sour les marces de Poito et de
Tourainne messires Guillaummes des Bordes, messires Jehans de
Buel, messires Loeis de Saint Juliien et Caruel le Breton et leurs
routtes. Si penssoient et estudioient nuit et jour coumment il
pewissent grever les Englès et leurs ennemis. Si avisèrent tant qu’il
prissent de nuit et emblèrent par esciellement le fort castiel qui
s’appelle le Roche de Ponsoy en Poito, sus le rivierre de Cruese, à
deux liewes de le Haie en Tourainne et assés priès de Casteleraut:
de quoy tous li pays d’environ fu durement effraés, et chil dou lés
des Franchois grandement recomfortés, car tantost il missent une
bonne grosse garnison dedens, sus le comfort de le fortrèce.
En ce meysme tamps, tenoient les camps en Poito messires
Guichars d’Angle, marescaux d’Acquittainne, et messires James
d’Audelée, senescaux de Poito, et messires Bauduins de Fraville,
senescaux de Saintonge; et avoient bien douze cens combatans, et
cevauchoient sus les marches de Berri et de Tourainne et mettoient
le pays où il converssoient, qui estoit contraire à yaux, en grant
tribulation. Et vinrent li dessus noummet chevalier et leur routtes
devant le ville de Breuse, qui est au seigneur de Cauvegny, ung
grant baron de Poito, liquelx viscomtez de Breuse et sires de
Cauvegny estoit tournés franchois: dont li prinches avoit grant
mautalent et avoit coummandé à monseigneur Guichart d’Angle que
la terre dou dessus dist chevalier fust toutte courue et gastée sans
deport, pour tant qu’il s’estoit tournés franchois et l’avoit relenqui,
qui estoit ses naturelx sires.
Quant messires Guichars d’Angle et li chevalier dessus noummet
et leurs gens furent venut devant le ville de Breuse, il s’arestèrent et
l’avisèrent, et ymaginèrent mout bien coumment il le poroient
assaillir à leur plus grant prouffit. Toutteffois, quant il l’eurent bien
consideré, il s’ordounnèrent à l’asaut, les archiers par devant, qui
coummencièrent à traire moult fort et mout roit, et chil de le
fortrèche à traire contre yaux et bien deffendre. Là eut grant assaut
et dur et bien continué, mès che premier jour il n’y fissent riens.
Dont se retraissent sus le soir li Englès et li Poitevin à leurs logeis, et
dissent que, à l’endemain, il feroient leur pooir dou concquerre, quoi
qu’il dewist couster. Si passèrent enssi celle nuit. L’endemain au
matin, il s’armèrent et ordounnèrent, et aprocièrent le ville de grant
vollenté et le coummencièrent à assaillir fortement et vistement, et
se prendre chacun priès de bien faire le besoingne. Si eut aucuns
compaignons appers et aventureux, qui vinrent jusquez as murs de
le ville, car li fosset n’estoit mies trop larges ne trop perfons, et
avoient piks et haviaux en leurs mains, dont il picquetoient les dis
murs; et pour le get des pierres qui leur pooient venir d’amont, il
estoient très bien paveschiet. Ossi y avoit archiers d’Engleterre qui
estoient aroutté sus les fossés: chil traioient si roit et si ouniement
que à painnes osoit nuls aparoir as gharittes pour deffendre le ville.
Et tant picquetèrent et assaillirent chil d’aval des fossés, qu’il
pertrusièrent le mur d’un grant trau et tant que doy homme y
pooient de froncq bien entrer. En peu d’eure, li murs fu si efforchiés
que li assallant en rompirent ung grant pan, par quoy touttes gens
sans dangier pooient bien par là entrer en le ville. Enssi Bruse prise,
et les gens contourné à grant meschief, car on les ochioit sans
merchy, dont c’estoit pités, et furent pris seize cens hommes d’armes
dou viscomte de Breuse, et tantost et sans delay pendu en leurs
armures meysmes. Ceste contrevenganche prissent les gens dou
prinche des hommes et de le ville de Breuse, affin que li autre
ewissent cause et matère de yaux castiier; et quant il en eurent fait
leur vollenté, il se retraissent deviers la chité de Poitiers. Ces
nouvelles vinrent au signeur de Cauvegny, qui se tenoit à Paris,
coumment sa ville de Breuse estoit prise et gastée et ses gens mors
et mis à destruction. Si en fu durement courouchiés, che fu bien
raisons, et dist qu’il l’amenderoit, quant il poroit, sus messire
Guichars d’Angle, qui estoit ses voisins et qui li avoit pourcachiet che
dammaige.
En ce tamps, manda li prinches de Galles en Angouloime, là où il
se tenoit tous malades, le viscomte de Rochuwart, que il venist
parler à lui, car li prinches estoit enfourmés que il se volloit tourner
franchois. Se vint li viscomtes deviers le prinche. Si trestost qu’il fu
venus, li prinches le fist prendre et mettre en prison courtoise et
bien garder, et jura qu’il ne partiroit de là jusques à tant que il aroit
bonne caution dou dit viscomte, qu’il seroit bons Englès et
demourroit dalés lui et ses gens en touttes besoingnes, ensi qu’uns
feables doit demourer dalés son seigneur. De le prise dou dit
viscomte et de le souppechon que li princes avoit sour lui, furent
tout si proïsme mout courouchiet, et si n’en peurent avoir autre
cose, tant qu’à ceste fois. Enssi estoient adonc li seigneur et les
terres en le duché d’Acquittainne en grant variement et guerriiet de
leurs voisins. Si ne savoient li pluisseur bonnement que faire. Fo 156
ro et vo.
P. 136, l. 17 et 18: Carenloet.—Mss. B: Charuel.—Ms. A 7:
Kaeranloet. Fo 302 vo.—Ms. A 8: Carnet. Fo 309.
P. 136, l. 18: douze cens.—Mss. B 4 et A 8: douze mille. Fo 306.
P. 136, l. 23: le Roce de Ponsoy.—Ms. A 8: la Roche de Posoy.
P. 137, l. 18: Rocewart.—Ms. A 8: Rochechouart.
P. 138, l. 3: Fraiville.—Ms. A 8: Franville.
P. 138, l. 10: contournés.—Ms. B 4: tournés. Fo 306 vo.—Ms. A 8:
tourmentez.
P. 138, l. 29: tamainte.—Le ms. A 8: ajoute: belle.
P. 139, l. 10: ou.—Ms. A 8: en.

§ 622. Messires Robers.—Ms. d’Amiens: Or entendi messires

Robers Canolles les nouvelles coumment li Franchois faisoient très
forte guerre au prinche, et li tolloient tous les jours ses villes et ses
castiaux, et ardoient et essilloient en la ducé d’Acquittainne mout
avant. Si s’avisa li dis messires Robers que il venroit deviers le
prinche et le serviroit et conforteroit à ce besoing de son corps et de
ses gens, car moult y estoit tenus pour tant que li roys ses pères
l’avoit mout amet et avanchiet en tous cas. Si fist ses pourveanches
pour lui et pour ses gens en quatre vaissiaux, sus ung havene de
mer en Bretaigne, que on appelle Konke; et quant il eut tout
apresté, il vint celle part et entra en son vaissiel à soixante hommes
d’armes, et naga et singla tant qu’il arriva ou havene au kay de le