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Quantitative Paleozoology

Quantitative Paleozoology describes and illustrates how the remains of long-dead
animals recovered from archaeological and paleontological excavations can be stud-
ied and analyzed. The methods range from determining how many animals of each
species are represented to determining whether one collection consists of more bro-
ken and more burned bones than another. All methods are described and illustrated
with data from real collections, while numerous graphs illustrate various quantitative
properties.

R. LEE LYMAN is professor of anthropology at the University of Missouri-Columbia.
A scholar of late Quaternary paleomammology and human prehistory of the Paci¬c
Northwest United States, he is the author of Vertebrate Taphonomy, and, most recently,
the coeditor of Zooarchaeology and Conservation Biology.
Cambridge Manuals in Archaeology
General Editor
Graeme Barker, University of Cambridge
Advisory Editors
Elizabeth Slater, University of Liverpool
Peter Bogucki, Princeton University

Cambridge Manuals in Archaeology is a series of reference handbooks designed for an inter-
national audience of upper-level undergraduate and graduate students and for professional
archaeologists and archaeological scientists in universities, museums, research laboratories,
and ¬eld units. Each book includes a survey of current archaeological practice alongside
essential reference material on contemporary techniques and methodology.



Books in the series
Pottery in Archaeology, CLIVE ORTON, PAUL TYERS, and ALAN VINCE
Vertebrate Taphonomy, R. LEE LYMAN
Photography in Archaeology and Conservation, 2nd edition, PETER G. DORRELL
Alluvial Geoarchaeology, A. G. BROWN
Shells, CHERYL CLAASEN
Sampling in Archaeology, CLIVE ORTON
Excavation, STEVE ROSKAMS
Teeth, 2nd edition, SIMON HILLSON
Lithics, 2nd edition, WILLIAM ANDREFSKY, JR.
Geographical Information Systems in Archaeology, JAMES CONOLLY and MARK LAKE
Demography in Archaeology, ANDREW CHAMBERLAIN
Analytical Chemistry in Archaeology, A. M. POLLARD, C. M. BATT, B. STERN, and S. M. M.
YOUNG
Zooarchaeology, 2nd edition, ELIZABETH J. REITZ and ELIZABETH S. WING
Quantitative
Paleozoology
R. Lee Lyman University of Missouri-Columbia
CAMBRIDGE UNIVERSITY PRESS
Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, São Paulo

Cambridge University Press
The Edinburgh Building, Cambridge CB2 8RU, UK
Published in the United States of America by Cambridge University Press, New York
www.cambridge.org
Information on this title: www.cambridge.org/9780521887496

© R. Lee Lyman 2008


This publication is in copyright. Subject to statutory exception and to the provision of
relevant collective licensing agreements, no reproduction of any part may take place
without the written permission of Cambridge University Press.
First published in print format 2008


ISBN-13 978-0-511-38646-6 eBook (EBL)

ISBN-13 978-0-521-88749-6 hardback

ISBN-13 978-0-521-71536-2 paperback



Cambridge University Press has no responsibility for the persistence or accuracy of urls
for external or third-party internet websites referred to in this publication, and does not
guarantee that any content on such websites is, or will remain, accurate or appropriate.
CONTENTS




List of ¬gures page xi
List of tables xvii
Preface xxi

1. Tallying and Counting: Fundamentals 1
Paleozoological Concepts 4
Mathematical and Statistical Concepts 8
Scales of Measurement 8
Measured and Target Variables: Reliability and Validity 11
Absolute and Relative Frequencies and Closed Arrays 13
Discussion 16
Background of Some Faunal Samples 17

2. Estimating Taxonomic Abundances: NISP and MNI 21
The Number of Identi¬ed Specimens (NISP) 27
Advantages of NISP 28
Problems with NISP 29
Problems, Schmoblems 30
A Problem We Should Worry About 36
The Minimum Number of Individuals (MNI) 38
Strengths(?) of MNI 43
Problems with MNI 45
Aggregation 57
De¬ning Aggregates 67
Discussion 69
Which Scale of Measurement? 71
Resolution 78
Conclusion 81
vii
contents
viii


3. Estimating Taxonomic Abundances: Other Methods 83
Biomass and Meat Weight 84
Measuring Biomass 85
Problems with Measuring Biomass (based on MNI) 86
Solving Some Problems in Biomass Measurement 88
Measuring Meat Weight 89
The Weight Method (Skeletal Mass Allometry) 93
Bone Weight 102
Bone Size and Animal Size Allometry 108
Ubiquity 114
Matching and Pairing 119
More Pairs Means Fewer Individuals 121
The Lincoln“Petersen Index 123
Identifying Bilateral Pairs 129
Correcting for Various Things 134
Size 137
Discussion 139

4. Sampling, Recovery, and Sample Size 141
Sampling to Redundancy 143
Excavation Amount 144
NISP as a Measure of Sample Redundancy 146
Volume Excavated or NISP 149
The In¬‚uences of Recovery Techniques 152
Hand Picking Specimens by Eye 152
Screen Mesh Size 154
To Correct or Not to Correct for Differential Loss 156
Summary 158
The Species’Area Relationship 159
Species’Area Curves Are Not All the Same 164
Nestedness 167
Conclusion 171

5. Measuring the Taxonomic Structure and Composition (“Diversity”)
of Faunas 172
Basic Variables of Structure and Composition 174
Indices of Structure and Similarity 178
Taxonomic Richness 179
Taxonomic Composition 185
contents ix


Taxonomic Heterogeneity 192
Taxonomic Evenness 194
Discussion 198
Trends in Taxonomic Abundances 203
Conclusion 209

6. Skeletal Completeness, Frequencies of Skeletal Parts, and
Fragmentation 214
History of the MNE Quantitative Unit 215
Determination of MNE Values 218
MNE Is Ordinal Scale at Best 222
A Digression on Frequencies of Left and Right Elements 229
Using MNE Values to Measure Skeletal-Part Frequencies 232
Modeling and Adjusting Skeletal-Part Frequencies 233
Measuring Skeletal Completeness 241
A Suggestion 244
Measuring Fragmentation 250
Fragmentation Intensity and Extent 250
The NISP:MNE Ratio 251
Discussion 254
Conclusion 261

7. Tallying for Taphonomy: Weathering, Burning, Corrosion,
and Butchering 264
Yet Another Quantitative Unit 266
Weathering 267
Chemical Corrosion and Mechanical Abrasion 273
Burning and Charring 274
A Digression 276
Gnawing Damage 277
Butchering Marks 279
Types of Butchering Damage 280
Tallying Butchering Evidence: General Comments 281
Tallying Percussion Damage 283
Tallying Cut Marks and Cut Marked Specimens 284
The Surface Area Solution 286
Discussion 291
Conclusion 296
contents
x


8. Final Thoughts 299
Counting as Exploration 302

Glossary 309
References 313
Index 345
LIST OF FIGURES




1.1. Chester Stock™s pie diagram of abundances of ¬ve mammalian
orders represented in faunal remains from Rancho La Brea. page 2
2.1. Schematic illustration of loss and addition to a set of faunal remains
studied by a paleozoologist. 23
2.2. Taxonomic relative abundances across ¬ve strata. 33
2.3. The theoretical limits of the relationship between NISP and MNI. 49
2.4. The theoretically expected relationship between NISP and MNI. 50
2.5. Relationship between NISP and MNI data pairs for mammal
remains from the Meier site. 52
2.6. Relationship between NISP and MNI data pairs for the precontact
mammal remains from the Cathlapotle site. 53
2.7. Relationship between NISP and MNI data pairs for the postcontact
mammal remains from the Cathlapotle site. 54
2.8. Relationship between NISP and MNI data pairs for remains of six
mammalian genera in eighty-four owl pellets. 56
2.9. Amount by which a taxon™s MNI increases if the minimum
distinction MNI is changed to the maximum distinction MNI in
eleven assemblages. 60
2.10. Change in the ratio of deer to gopher abundances in eleven
assemblages when MNImax is used instead of MNImin. 61
2.11. Relationships between NISP and MNImin, and NISP and MNImax
at site 45DO214. 66
2.12. Ratios of abundances of four least common taxa in a collection of
eighty-four owl pellets based on NISP, MNImax, and MNImin. 72
2.13. Frequency distributions of NISP and MNI taxonomic abundances
in the Cathlapotle fauna. 73


xi
list of ¬gures
xii


2.14. Frequency distributions of NISP and MNI taxonomic abundances
in the 45OK258 fauna in eastern Washington State. 74
2.15. Frequency distributions of NISP and MNI taxonomic abundances
in two lumped late-prehistoric mammal assemblages from the
western Canadian Arctic. 75
2.16. Frequency distributions of NISP and MNI taxonomic abundances
in four lumped historic era mammalian faunas. 76
3.1. Ontogenetic, seasonal, and sexual variation in live weight of one
male and one female Columbian black-tailed deer. 88
3.2. Relationship between bone weight per individual and soft-tissue
weight in domestic pig. 98
3.3. Relationship between bone weight per skeletal portion and gross
weight per skeletal portion in 6-month-old domestic sheep and
90-month-old domestic sheep. 101
3.4. Frequency distributions of biomass per taxon in two sites in Florida
State. 106
3.5. Frequency distributions of biomass per mammalian taxon in a site
in Georgia State. 107
3.6. Relationship between lateral length of white-tailed deer astragali
and body weight. 112
3.7. Relationship between NISP and ubiquity of six genera in a
collection of eighty-four owl pellets. 115
3.8. Relationship between NISP and ubiquity of twenty-eight taxa in
eighteen sites. 117
3.9. Relationship between NISP and ubiquity of ¬fteen taxa in seven
analytical units in site 45DO189. 119
3.10. Relationship between NISP and ubiquity of eighteen taxa in four
analytical units in site 45OK2. 120
3.11. A model of how the Lincoln“Petersen index is calculated. 125
3.12. Latero-medial width of the distal condyle and minimum
antero-posterior diameter of the middle groove of the distal condyle
of forty-eight pairs of left and right distal humeri of Odocoileus
virginianus and Odocoileus hemionus. 131
3.13. A model of how two dimensions of a bone can be used to determine
degree of (a)symmetry between bilaterally paired left and right
elements. 132
4.1. Cumulative richness of mammalian genera across cumulative
volume of sediment excavated annually at the Meier site. 146
list of ¬gures xiii


4.2. Cumulative richness of mammalian genera across cumulative
annual samples from the Meier site. 148
4.3. Cumulative richness of mammalian genera across cumulative
annual samples from Cathlapotle. 149
4.4. Relationship of mammalian genera richness and sample size per
annual sample at the Meier site. 150
4.5. Relative abundances of ¬fteen size classes of mollusk shells
recovered during hand picking from the excavation, and recovered
from ¬ne-mesh sieves. 153
4.6. The effect of passing sediment through screens or sieves on recovery
of mammal remains relative to hand picking specimens from an
excavation unit. 153
4.7. Cumulative percentage recovery of remains of different size classes
of mammals. 156
4.8. Model of the relationship between area sampled and number of taxa
identi¬ed. 160
4.9. Two models of the results of rarefaction. 161
4.10. Rarefaction curve and 95 percent con¬dence intervals of richness of
mammalian genera based on six annual samples from the Meier site. 166
4.11. Examples of perfectly nested faunas and poorly nested faunas. 168
4.12. Nestedness diagram of eighteen assemblages of mammalian genera
from eastern Washington State. 170
5.1. Two ¬ctional faunas with identical taxonomic richness values but
different taxonomic evenness. 176
5.2. Three ¬ctional faunas with varying richness values and varying
evenness values. 177
5.3. Relationship between genera richness and sample size in eighteen
mammalian faunas from eastern Washington State. 182
5.4. Relationships between NISP and NTAXA of small mammals per
stratum at Homestead Cave, Utah. 182
5.5. Relationship between NISP and NTAXA per stratum at Le Flageolet
I, France. 184
5.6. Two Venn diagrams based on the Meier site and Cathlapotle site
collections. 187
5.7. Bivariate scatterplot of relative abundances of mammalian genera at
the Meier site and Cathlapotle. 190
5.8. Rarefaction analysis of eighteen assemblages of mammal remains
from eastern Washington State. 191
list of ¬gures
xiv


5.9. The relationship between taxonomic heterogeneity and sample size
in eighteen assemblages of mammal remains from eastern
Washington State. 194
5.10. Frequency distribution of NISP values across six mammalian genera
in a collection of owl pellets. 195
5.11. Relationship between taxonomic evenness and sample size in
eighteen assemblages of mammal remains from eastern Washington
State. 197
5.12. Relationship between sample size and the reciprocal of Simpson™s
dominance index in eighteen assemblages of mammal remains from
eastern Washington State. 198
5.13. Percentage abundance of deer in eighteen assemblages from eastern
Washington State. 202
5.14. Abundance of bison remains relative to abundance of all ungulate
remains over the past 10,500 years in eastern Washington State. 203
5.15. Bivariate scatterplot of elk abundances relative to the sum of all
ungulate remains in eighty-six assemblages from eastern
Washington State. 206
5.16. Bivariate scatterplot of elk’deer index against stratum at
Emeryville Shellmound. 208
5.17. Relative abundances of Neotoma cinerea and Dipodomys spp. at
Homestead Cave. 210
5.18. Bivariate scatterplot of elk abundances relative to the sum of all
ungulate remains in eighty-six assemblages from eastern
Washington State summed by age for consecutive 500-year bins. 212
6.1. Relationship of NISP and MNE values for deer remains from the
Meier site. 225
6.2. Relationship of NISP and MNE values for wapiti remains from the
Meier site. 225
6.3. Frequency distributions of NISP and MNE abundances per skeletal
part for deer remains from the Meier site. 226
6.4. Frequency distributions of NISP and MNE abundances per skeletal
part for wapiti remains from the Meier site. 227
6.5. Frequency distribution of skeletal parts in single skeletons of four
taxa. 229
6.6. Comparison of MNE of left skeletal parts and MNE of right skeletal
parts in a collection of pronghorn bones. 231
list of ¬gures xv


6.7. Frequencies of skeletal elements per category of skeletal element in a
single artiodactyl carcass. 234
6.8. MNE and MAU frequencies for a ¬ctional data set. 235
6.9. MNE values plotted against the MNE skeletal model. 236
6.10. MAU values plotted against the MAU skeletal model. 237
6.11. MAU values for two collections with different MNI values. 240
6.12. %MAU values for two collections with different MNI values. 241
6.13. Relationship between Shotwell™s CSI per taxon and NISP per taxon
for the Hemphill paleontological mammal assemblage. 243
6.14. Relationship between Thomas™s CSI per taxon and NISP per taxon
for the Smoky Creek zooarchaeological mammal collection. 245
6.15. Bar graph of frequencies of skeletal parts for two taxa. 247
6.16. Model of the relationship between fragmentation intensity and
NISP. 253
6.17. Model of the relationship between NISP and MNE. 254
6.18. Relationship between NISP and MNE values for size class II cervids
and bovids at Kobeh Cave, Iran. 257
6.19. Relationship between NISP and MNE values for saiga antelope at
Prolom II Cave, Ukraine. 258
6.20. Relationship between NISP and MNE values for caprine remains
from Neolithic pastoral site of Ngamuriak, Kenya. 260
6.21. Relationship between (lefts + rights) and MNI per skeletal part. 262
7.1. Weathering pro¬les for two collections of ungulate remains from
Olduvai Gorge. 269
7.2. Relationship between years since death and the maximum
weathering stage displayed by bones of a carcass. 271
7.3. Weathering pro¬les based on ¬ctional data for a collection of bones
with skyward surfaces representing one pro¬le and groundward
surfaces representing another pro¬le. 272
7.4. Frequency distribution of seven classes of burned bones in two
kinds of archaeological contexts. 275
7.5. Relationship between number of arm strokes and number of cut
marks on thirty-one skeletal elements. 291
7.6. Relationship between number of arm strokes necessary to de¬‚esh a
bone and the amount of ¬‚esh removed. 293
7.7. Relationship between number of cut marks and the amount of ¬‚esh
removed from thirty-one limb bones. 293
list of ¬gures
xvi


7.8. Relationships between number of cut marks and the amount of

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