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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

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
Sampling in Archaeology, CLIVE ORTON
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.
Zooarchaeology, 2nd edition, ELIZABETH J. REITZ and ELIZABETH S. WING
R. Lee Lyman University of Missouri-Columbia
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
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.

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

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

8. Final Thoughts 299
Counting as Exploration 302

Glossary 309
References 313
Index 345

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

list of ¬gures

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

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

7.8. Relationships between number of cut marks and the amount of
¬‚esh removed from six hindlimbs in each of three carcass sizes. 295
7.9. Relationship between number of cut marks and the amount of ¬‚esh
removed from eighteen hindlimbs. 295
8.1. Relationship between NISP and MNI in seven paleontological
assemblages of bird remains from North America. 304
8.2. Relationship between NISP and MNI in eleven paleontological
assemblages of mammal remains from North America. 304
8.3. Relationship between NISP and MNI in twenty-two archaeological
assemblages of bird remains from North America. 305
8.4. Relationship between NISP and MNI in thirty-¬ve archaeological
assemblages of mammal remains from North America. 306

1.1. An example of the Linnaean taxonomy. page 6
1.2. Fictional data on the absolute abundances of two taxa in six
chronologically sequent strata. 15
1.3. Description of the mammalian faunal record at Meier and at
Cathlapotle. 19
2.1. Fictional data on abundances of three taxa in ¬ve strata. 32
2.2. Data in Table 2.1 adjusted as if each individual of taxon A had ten
skeletal elements per individual, taxon B had one skeletal element per
individual, and taxon C had ¬ve skeletal elements per individual. 33
2.3. The differential exaggeration of sample sizes by NISP. 36
2.4. Some published de¬nitions of MNI. 40
2.5. A ¬ctional sample of seventy-one skeletal elements representing a
minimum of seven individuals. 45
2.6. Statistical summary of the relationship between NISP and MNI for
mammal assemblages from Meier and Cathlapotle. 53
2.7. Statistical summary of the relationship between NISP and MNI for
mammal assemblages from fourteen archaeological sites in eastern
Washington State. 55
2.8. Maximum distinction and minimum distinction MNI values for six
genera of mammals in a sample of eighty-four owl pellets. 55
2.9. Adams™s data for calculating MNI values based on Odocoileus sp.
remains. 57
2.10. Differences in site total MNI between the MNI minimum distinction
results and the MNI maximum distinction results. 59
2.11. The most abundant skeletal part representing thirteen mammalian
genera in two (sub)assemblages at Cathlapotle. 62

xviii list of tables

2.12. Fictional data showing how the distribution of most abundant
skeletal elements of one taxon can in¬‚uence MNI across different
aggregates. 63
2.13. Fictional data showing how the distribution of skeletal elements of
two taxa across different aggregates can in¬‚uence MNI. 64
2.14. Fictional data showing that identical distributions of most common
skeletal elements of two taxa across different aggregates will not
in¬‚uence MNI. 65
2.15. Ratios of abundances of pairs of taxa in eighty-four owl pellets. 72
3.1. Biomass of deer and wapiti at Cathlapotle. 86
3.2. Meat weight for deer and wapiti at Cathlapotle, postcontact
assemblage. 90
3.3. Comparison of White™s conversion values to derive usable meat with
Stewart and Stahl™s conversion values to derive usable meat. 91
3.4. Variation by age and sex of wapiti butchered weight as a percentage
of live weight. 92
3.5. Weight of a 350-kilogram male wapiti in various stages of butchering. 93
3.6. Descriptive data on animal age, bone weight per individual, and
soft-tissue weight per individual domestic pig. 97
3.7. Statistical summary of the relationship between bone weight and
weight of various categories of soft-tissue for domestic pig. 98
3.8. Descriptive data on dry bone weight per anatomical portion and
total weight per anatomical portion for domestic sheep. 100
3.9. Statistical summary of the relationship between bone weight and
gross weight or biomass of skeletal portions of two domestic sheep. 101
3.10. Relationship between NISP and bone weight of mammalian taxa in
seventeen assemblages. 104
3.11. Results of applying the bone-weight allometry equation to ¬ve
randomly generated collections of domestic sheep bone. 105
3.12. Deer astragalus length and live weight. 110
3.13. Ubiquity and sample size of twenty-eight mammalian taxa in
eighteen sites. 116
3.14. Ubiquity and sample size of mammalian taxa across analytical units
in two sites. 118
3.15. Fictional data illustrating in¬‚uences of NISP and the number of pairs
on the Lincoln“Petersen index. 126
3.16. Abundances of beaver and deer remains at Cathlapotle, and WAE
values and ratios of NISP and WAE values per taxon per assemblage. 135
list of tables xix

3.17. Estimates of individual body size of seventeen white-tailed deer
based on the maximum length of right and left astragali. 139
4.1. Volume excavated and NISP of mammals per annual ¬eld season at
the Meier site. 144
4.2. Annual NISP samples of mammalian genera at the Meier site. 145
4.3. Annual NISP samples of mammalian genera at Cathlapotle. 147
4.4. Mammalian NISP per screen-mesh size class and body-size class for
three sites. 155
4.5. Two sets of faunal samples showing a perfectly nested set of faunas
and a poorly nested set of faunas. 169
5.1. Sample size, taxonomic richness, taxonomic heterogeneity,
taxonomic evenness, and taxonomic dominance of mammalian
genera in eighteen assemblages from eastern Washington State. 181
5.2. NISP and NTAXA for small mammals at Homestead Cave, Utah. 183
5.3. NISP and NTAXA for ungulates at Le Flageolet I, France. 184
5.4. NISP per taxon in two chronologically distinct samples of
eighty-four owl pellets. 188
5.5. Expected values and interpretation of taxonomic abundances in two
temporally distinct assemblages of owl pellets. 188
5.6. Derivation of the Shannon’Wiener index of heterogeneity for the
Meier site. 193
5.7. Total NISP of mammals, NISP of deer, and relative abundance of
deer in eighteen assemblages from eastern Washington State. 199
5.8. Frequencies of bison and nonbison ungulates per time period in
ninety-one assemblages from eastern Washington State. 204
5.9. Frequencies of elk, deer, and medium artiodactyl remains per
stratum at Emeryville Shellmound. 207
5.10. Frequencies of two taxa of small mammal per stratum at Homestead
Cave. 209
6.1. MNE values for six major limb bones of ungulates from the FLK
Zinjanthropus site. 219
6.2. Fictional data showing how the distribution of specimens of two
skeletal elements across different aggregates can in¬‚uence MNE. 223
6.3. NISP and MNE per skeletal part of deer and wapiti at the Meier site. 224
6.4. Frequencies of major skeletal elements in a single mature skeleton of
several common mammalian taxa. 228
6.5. MNE frequencies of left and right skeletal parts of pronghorn from
site 39FA83. 230
list of tables

6.6. Expected MNE frequencies of pronghorn skeletal parts at site 39FA83,
and adjusted residuals and probability values for each. 232
6.7. Frequencies of skeletal elements in a single generic artiodactyl
skeleton. 233
6.8. MNE and MAU frequencies of skeletal parts and portions. 236
6.9. MAU and%MAU frequencies of bison from two sites. 239
6.10. Skeletal-part frequencies for two taxa of artiodactyl. 246
6.11. Expected frequencies of deer and wapiti remains at Meier, adjusted
residuals, and probability values for each. 249
6.12. Ratios of NISP:MNE for four long bones of deer in two sites on the
coast of Oregon State. 252
6.13. African bovid size classes. 255
6.14. NISP and MNE frequencies of skeletal parts of bovid/cervid size class
II remains from Kobeh Cave, Iran. 256
6.15. NISP and MNE frequencies of skeletal parts of saiga antelope from
Prolom II Cave, Ukraine. 257
6.16. Relationship between NISP and MNE in twenty-nine assemblages. 259
6.17. NISP and MNI frequencies of skeletal parts of caprines from
Ngamuriak, Kenya. 260
6.18. Relationship between NISP and MNI per skeletal part or portion in
twenty-two assemblages. 261
7.1. Weathering stages as de¬ned by Behrensmeyer. 267
7.2. Weathering stage data for two collections of mammal remains from
Olduvai Gorge. 268
7.3. Expected frequencies of specimens per weathering stage in two
collections, adjusted residuals, and probability values for each. 269
7.4. Frequencies of cut marks per anatomical area on six experimentally
butchered goat hindlimbs. 288
7.5. Frequencies of arm strokes and cut marks on sixteen limbs of cows
and horses. 290
7.6. Number of cut marks generated and amount of meat removed from
eighteen mammal hindlimbs by butchering. 294
8.1. Statistical summary of relationship between NISP and MNI in
collections of paleontological birds, paleontological mammals,
archaeological birds, and archaeological mammals. 303

Several years ago I had the opportunity to have a relaxed discussion with my doctoral
advisor, Dr. Donald K. Grayson. In the course of that discussion, I asked him if
he would ever revise his then 20-year-old book titled Quantitative Zooarchaeology,
which had been out of print for at least a decade. He said “No” and explained that the
topic had been resolved to his satisfaction such that he could do the kinds of analyses
he wanted to do. A spur-of-the-moment thought prompted me to ask, “What if I
write a revision?” by which I meant not literally a revised edition but instead a new
book that covered some of the same ground but from a 20-years-later perspective.
Don said that he thought that was a ¬ne idea.
After the conversation with Grayson, I began to mentally outline what I would
do in the book. I realized that it would be a good thing for me to write such a
book because, although I thought I understood many of the arguments Grayson had
made regarding the counting of animal remains when I was a graduate student, there
were other arguments made by other investigators subsequent to the publication of
Grayson™s book that I didn™t know (or if I knew of those arguments, I wasn™t sure I
understood them very well). I also knew that the only way for me to learn a topic
well was to write about it because such a task forced me to learn its nuances, its
underpinning assumptions, the interrelations of various aspects of the argument,
and all those things that make an approach or analytical technique work the way that
it does (or not work as it is thought to, as the case may be).
As I mentally outlined the book over the next several months, it occurred to me
that at least one new quantitative unit similar to the traditional ones Grayson had
considered had become a focus of analytical attention over the two decades subse-
quent to the publication of Grayson™s book (MNE, and the related MAU). And an
increasing number of paleozoologists were measuring taxonomic diversity “ a term
that had several different meanings for several different variables as well as being
measured several different ways. What were those measurement techniques and
xxii preface

what were those measured variables? Finally, there were other kinds of phenomena
that zooarchaeologists and paleontologists had begun to regularly tally and analyze.
These phenomena “ butchering marks, carnivore gnawing marks, rodent gnawing
marks, burned bones “ had become analytically important as paleozoologists had
come to realize that to interpret the traditional quantitative measures of taxonomic
abundances, potential biases in those measures caused by differential butchery, car-
nivore attrition, and the like across taxa had to be accounted for. As I indicate in this
volume, there are several ways to tally up carnivore gnawing marks and the like, and
few analysts have explored the fact that each provides a unique result.
Finally, it had become clear to me during the 1990s that many paleozoologists were
unaware of what I took to be two critical things. First, zooarchaeologists seemed
to seldom notice what is published in paleontological journals; at least they sel-
dom referenced that literature. Thus, they were often ignorant of various sugges-
tions made by paleontologists regarding quantitative methods. Paleontologists were
equally unaware of what zooarchaeologists have determined regarding quanti¬ca-
tion of bones and shells and teeth. Therefore, it seemed best to title this volume
Quantitative Paleozoology for the simple reason that were it to be titled “Quantitative
Zooarchaeology,” it likely would not be read by paleontologists. A very interesting
book with the title Quantitative Zoology coauthored by a paleontologist (Simpson et
al. 1960) already exists, so that title could not be used, aside from it being misleading.
Quantitative Paleozoology is a good title for two reasons. The ¬rst reason is that the
subject materials, whether collected by a paleontologist or an archaeologist, do not
have a proximate zoological source (though their source is ultimately zoological) but
rather have a proximate geological source, whether paleontological (without associ-
ated human artifacts) or archaeological (with associated and often causally related
human artifacts). I conceive of all such remains as paleozoological. The second rea-
son Quantitative Paleozoology is a good title is that the volume concerns how to
count or tally, how to quantify zoological materials and their attributes, speci¬cally
those zoological remains recovered from geological contexts. Not all such topics are
discussed here, but many are; for an introduction to many of those that are not, see
Simpson et al. (1960), a still-useful book that was, fortunately, reprinted in 2003.
The second critical thing that many paleozoologists seem to be unaware of is basic
statistical concepts and methods. I was stunned in 2004 to learn that an anonymous
individual who had reviewed a manuscript I submitted for publication did not know
what a “closed array” was and therefore did not understand why my use of this par-
ticular analytical tool could have been in¬‚uencing (some might say biasing, but that
is a particular kind of in¬‚uencing) the statistical results. In the 1960s and early 1970s,
many archaeologists and paleontologists did not have very high levels of statistical
sophistication; I had thought that most of them did have such sophistication (or at
preface xxiii

least knowledge of the basics) in the twenty-¬rst century. The anonymous reviewer™s
comments indicate that at least some of them do not. Therefore, it seemed that any
book on quantitative paleozoology had to include brief discussions of various sta-
tistical and mathematical concepts. In order to not dilute the central focus of the
volume “ quantitative analysis of paleozoological remains “ I have kept discussion
of statistical methods to a minimum, assuming that the serious reader will either
already know what is necessary or will learn it as he or she reads the book. I have,
however, devoted the ¬rst chapter to several critical mathematical concepts as well
as some key paleozoological concepts.
Many of the faunal collections used to illustrate various points in the text were
provided over the years by friends and colleagues who entrusted me with the analysis
of those collections. Many of the things I have learned about quantitative paleozo-
ology are a direct result of their trust. To these individuals, I offer my sincere thanks:
Kenneth M. Ames, David R. Brauner, Jerry R. Galm, Stan Gough, Donald K. Grayson,
David Kirkpatrick, Lynn Larson, Frank C. Leonhardy, Dennis Lewarch, Michael J.
O™Brien, Richard Pettigrew, and Richard Ross. Perhaps more importantly, any clarity
this book brings to the issues covered herein is a result of the collective demand for
clarity by numerous students who sat through countless lectures about the counting
units and methods discussed in this book. A major source of inspiration for the ¬rst
several chapters was provided in 2004 by the Alaska Consortium of Zooarchaeol-
ogists (ACZ). That group invited me to give a daylong workshop on the topics of
quanti¬cation and taphonomy, and that forced me to think through several things
that had previously seemed less than important. I especially thank Diane Hansen
and Becky Saleeby of the ACZ for making that workshop experience memorable.
An early draft of the manuscript was reviewed by Jack Broughton, Corey Hudson,
Alex Miller, and an anonymous individual. Broughton and the anonymous reviewer
ensured that a minimum of both glaring errors in logic and stupid errors in mathe-
matics remain in this version. Broughton and the anonymous reviewer insisted that
I include several recently described analytical techniques, and they identi¬ed where
I overstepped and where I misstepped. These individuals deserve credit for many of
the good things here.
I wrote much of the ¬rst draft of this volume between July 2005 and August 2006.
During that time, I lost my younger brother and both parents. They all had an indirect
hand in this book. My parents taught me to hunt and ¬sh, and all of the things that
accompany those activities. My brother did not discourage me from collecting owl
pellets from his farm equipment shed, or laugh too hard when I collected them; he
even grew to appreciate what could be learned from the mouse bones they contained.
I miss them all, and I dedicate this book to the three of them.
June 2007
Tallying and Counting: Fundamentals

Early in the twentieth century, paleontologist Chester Stock (1929) was, as he put
it, faced with “recording a census” of large mammals from the late Pleistocene as
evidenced by their remains recovered “from the asphalt deposits of Rancho La Brea,”
in Los Angeles, California. Paleontologist Hildegarde Howard (1930) was faced with
a similar challenge with respect to the bird remains from Rancho La Brea. Stock and
Howard could have merely listed the species of mammals and the species of birds,
respectively, that were represented by the faunal remains they had “ they could have
constructed an inventory of taxa “ but they chose to do something more informative
and more analytically powerful. They tallied up how many individuals of each species
were represented by the remains “ they each produced a census. The quantitative
unit they chose became known as the minimum number of individuals, or MNI, a
unit that was quickly (within 25 years) adopted by many paleozoologists. We will
consider this unit in some detail in Chapter 2, but here it is more important to outline
how Stock and Howard de¬ned it and why they decided to provide a census rather
than an inventory of mammals and an inventory of birds.
Stock (1929:282) stated that the tally or “count” of each taxon was “determined
by the number of similar parts of the internal skeleton as for example the skull,
right ramus of mandible, left tibia, right scaphoid. In many cases the total number
of individuals for any single group [read taxon] is probably a minimum estimate.”
Howard (1930:81 “82) indicated that “for each species, the left or the right of the
[skeletal] element occurring in greatest abundance was used to make the count. . . . It
is probable that in many instances the totals present a minimum estimate of the
number of individuals [per taxon] actually represented in the collection.” We will
explore why the procedure Stock and Howard used provides a “minimum” estimate
of abundance in Chapter 2. Stock and Howard each produced a type of pie diagram
to illustrate their respective censuses of mammalian and of avian creatures based on
the bony remains of each (Figure 1.1).
quantitative paleozoology

figure 1.1. Chester Stock™s pie diagram of abundances of ¬ve mammalian orders repre-
sented in faunal remains from Rancho La Brea. Redrawn from Stock (1929).

An inventory of the mammalian taxonomic orders Stock identi¬ed among the
bones and teeth he studied would look like this:

Clearly, the pie diagram in Figure 1.1 reveals more about the structure of the Rancho La
Brea mammalian fauna because it contains not only the same set of taxonomic orders
as the inventory, but it also contains measures of the abundances of animals belonging
to each order. This example illustrates one of the major reasons why paleozoologists
count or tally the animal remains they study. Taxa present in a collection can, on
the one hand, be treated as attributes or as present or absent from a fauna, such
as is given in the inventory above (sometimes referred to as a “species list” if that
taxonomic level is used). On the other hand, abundances of each taxon provide a great
deal more information about the prehistoric fauna. There are times when knowing
only which taxa are present, or knowing only what the frequencies of different taxa
are is all that is wanted or needed analytically. (Two faunas may have the same, or
quite different, frequency distributions of individual organisms across taxa, and the
research question may only require knowing the frequency distributions and not
the taxa.) Knowing both, however, means we know more than when we know just
one or the other. And that is a good reason to count faunal remains and to determine
tallying and counting: fundamentals 3

a census. Counting faunal remains, particularly old or prehistoric remains, and the
variety of attributes they display, whether the remains are from archaeological or
paleontological contexts, is what this book is about.
There is already a book about counting animal remains recovered from archae-
ological and paleontological sites (Grayson 1984), and several other volumes cover
some of the same ground, if in less detail (e.g., Hesse and Wapnish 1985; Klein and
Cruz-Uribe 1984; Reitz and Wing 1999). Noting this, one could legitimately ask why
another book on this topic is necessary. There are several reasons to write a new
book. Much has happened in the ¬eld since Grayson (1984) published his book (and
his book has been out-of-print for several years). Some of what has happened has
been conceptually innovative, such as the de¬nition of new quantitative units meant
to measure newly conceived properties of the paleozoological record. Some of what
has happened has been technically innovative, such as designing new protocols for
tallying animal remains that are thought to provide more accurate re¬‚ections of what
is represented by a collection of remains than tallies based on less technologically
sophisticated methods. And, some of what has happened is misguided or archaic,
such as arguing that if certain biological variables are not mathematically controlled
for, then any count of taxonomic abundances is invalid. It is time (for these reasons)
for a new, up-to-date examination of the quantitative units and counting protocols
paleozoologists use in their studies.
There is yet another reason to produce a new book on quantitative paleozoology.
Today, early in the third millennium, there are more people studying paleozoological
collections than there were 20 years ago. These folks need to be able to communi-
cate clearly and concisely with one another regarding their data and their analyses
because the use of ambiguous terminology thwarts ef¬cient communication and
results in confusion. This point was made more than a decade ago with respect to the
plethora of terms, many unfamiliar to those in the ¬eld, used for quantitative units
in zooarchaeology (Lyman 1994a). Yet, the problem continues today. This problem
had originally been identi¬ed more than 15 years earlier still by Casteel and Grayson
(1977). For whatever reason, terminological ambiguity seems to plague paleozoology
and continues to do so despite it being explicitly identi¬ed twice in the past 30 years.
In my earlier discussion of terminological ambiguity (Lyman 1994a), I did not
advocate a particular terminology, nor am I doing so here. Clearly there are terms
I prefer “ the ones I use in this volume are the ones I learned as a student. What I
am arguing here is that whatever terms or acronyms one uses, these must be clearly
de¬ned at the start so as to avoid misunderstanding. In reading and rereading the
literature on quantitative paleozoology as I prepared this book, I was often dumb-
founded when people used terms such as “bone” and “relative abundance” when it
quantitative paleozoology

was quite clear that they were discussing teeth and absolute abundances, respectively.
Much of the remainder of this chapter is, therefore, devoted to terminology and def-
initions. For quick reference, I have included a glossary of key terms at the end of
this volume.
In this introductory chapter, several basic mathematical and statistical concepts are
de¬ned. This is necessary because these concepts will be used throughout subsequent
chapters and thus the concepts must be understood in order to follow the discussion
in later chapters. Several basic paleozoological concepts are introduced and de¬ned
for the same reason. I begin with these concepts before turning to the mathematical
and statistical concepts.


Throughout this volume the focus is on vertebrates, especially mammals, because
that is the taxonomic group which much of the literature concerns and because it is
the group with which I am most familiar. However, virtually every thing that is said
about quantifying vertebrate remains and their attributes holds with equal force for
invertebrates (e.g., Claassen 1998:106“107).
In many discussions of how paleofaunal remains are tallied, and even in some dis-
cussions of how modern animal bones should be counted, the reader may encounter
the term “skeletal element.” Or, one might encounter the term “bone,” or “tooth,”
or “shell,” or any of many other similar, more or less synonymous general terms
for skeletal remains. But if one collection comprises ten “bones” of a skeleton and
another consists of eleven “bones” of another skeleton of the same species as the ¬rst,
is the latter more anatomically complete than the former? Is the taxon less abundant
in the ¬rst collection than in the second? If you think the answer is “Yes” to either
question, you might be correct. But you could be wrong if when the analyst tallied
specimens no distinction was made between anatomically complete bones and frag-
ments of bones. The lesson is simple. If we are going to tally up skeletal parts and want
to compare our tally with that of another analyst working with another collection,
we had best be sure that we counted skeletal parts the same way that the other person
did. What, then, exactly is a skeletal element?
Paleontologist Michael Voorhies (1969:18) distinguished between “fragments” and
“elements or bones,” but we need something more explicit and inclusive because
not all skeletal elements are, technically, bones. Some are teeth, some are horns, and
some are antlers, and so on. Following Arnold Shotwell (1955, 1958), Donald Grayson
(1984) and Catherine Badgley (1986) provide useful terminology and de¬nitions. A
tallying and counting: fundamentals 5

skeletal element is a complete discrete anatomical unit such as a bone, tooth, or shell.
The critical phrase is complete discrete anatomical unit. Each such item is a discrete
“anatomical organ” (Francillon-Vieillot et al. 1990:480) that does not lose its integrity
or completeness when it is removed from an organism. A humerus, a tibia, a carpal,
a ¬rst lower molar “ each is a skeletal element. One might correctly note that “dis-
creteness” depends on the age or ontogenetic stage of development of the organism,
but many paleozoologists would not tally the proximal epiphysis of a humerus and
the diaphysis of that humerus as two separate specimens if it was clear that the two
specimens went together (an issue we return to in Chapter 2). Those same paleozo-
ologists usually don™t tally up each individual tooth ¬rmly set in a mandible, along
with the dentary or mandible bone. These are potentially signi¬cant concerns but
may ultimately be of minimal analytical import once we get into tallying specimens.
Not all faunal remains recovered from paleozoological deposits are anatomically
complete; some are represented by only a part of the original skeletal element because
of fragmentation. Thus, another term is necessary. A specimen is a bone, tooth, or
shell, or fragment thereof. All skeletal elements are specimens, but not all specimens
are skeletal elements. A distal humerus, a proximal tibia, and a fragment of a premolar
are all specimens that derive from skeletal elements; phenomenologically they are
not, technically, anatomically complete skeletal elements. Specimen is an excellent
term for many counting operations because it is value-free in the sense that it does
not reveal whether specimen A is anatomically more complete, or less complete,
than specimen B. We can record whether specimen A is anatomically complete, and
if it isn™t, we can record the portion of a complete element that is represented by a
fragment, if our research questions demand such. Specimen is also a better generic
term than skeletal element for the individual skeletal remains we study because skeletal
element implies that a complete anatomical unit is represented. The problem with
the terms “bone” and “tooth” and the like are that sometimes when analysts use
them they mean both anatomically complete skeletal elements as de¬ned here and
incomplete skeletal elements. Failure to distinguish the two kinds of units “ skeletal
element and specimen “ can render separate tallies incomparable and make the
signi¬cance of various analyses obscure. Throughout this volume, I use the term
skeletal part as a synonym for specimen, but whereas the latter is a general category
that can include many and varied anatomical portions, skeletal part is restricted to
a particular category of anatomical portion, say, distal humerus. Skeletal portion is
sometimes used in the same category-speci¬c way that skeletal part is but will usually
mean a multiple skeletal element segment of a skeleton, such as a forelimb.
Henceforth, in this volume, specimen will be used to signify any individual skeletal
remain, whether anatomically complete or not. Unfortunately, the terms “skeletal
quantitative paleozoology

Table 1.1. An example of the Linnaean taxonomy

Taxonomic level Taxonomic name Common name
Kingdom Animalia Animals
Phylum Vertebrata Vertebrate
Class Mammalia Mammals
Infraclass Eutheria Placental mammal
Order Carnivora Carnivores
Family Canidae Canids
Genus Canis Dogs, coyotes, wolves, and allies
Species— latrans coyote

Technically, the species name is Canis latrans; latrans is the speci¬c epithet.

element” and “element” are still often used to denote anatomically incomplete items.
An effort is made throughout this book to make clear what exactly is being tallied
and how it is being tallied. In this respect, what are usually tallied are what are
termed “identi¬ed” or “identi¬able” specimens. Typically, this means identi¬ed as
to biological taxon, usually genus or species, represented by a bone, tooth, or shell
(Driver 1992; Lyman 2005a). To identify skeletal remains, one must know the structure
of the Linnaean taxonomy, an example of which is given in Table 1.1 . One must also
know the basics of skeletal anatomy, by which is meant that one must know the
difference between a scapula and a radius, a femur and a cervical vertebra, a clavicle
and a rib, and so on. Finally, the person doing the identi¬cations must be able to
distinguish intertaxonomic variation from intrataxonomic variation. Intrataxonomic
variation is also sometimes termed “individual variation” within the species level of
the taxonomy. I presume that readers of the book know these things, along with
anatomical location and direction terms used in later chapters.
The importance of the requirements for identi¬cation should be apparent when
one realizes that “identi¬cation” involves questions such as: Is one dealing with a
mammal or a bird? If it is a mammal, is it a rodent or a carnivore? If it is a car-
nivore, is it a canid, a felid, a mustelid, or any of several other taxa of carnivores?
The importance of the other knowledge requirement “ basic skeletal anatomy “ will
assist in answering the questions just posed. The importance of distinguishing inter-
taxonomic from intrataxonomic variation is usually (and best) met by consultation
of a comparative collection of skeletons of known taxonomic identity. The proce-
dure is simple. Compare the taxonomically unknown paleozoological specimen with
comparative specimens of known taxonomy until the best match is found. Often the
closest match will be obvious, and the unknown specimen is “identi¬ed” as belonging
to the same taxon as the known comparative specimen. Sometimes this means that
tallying and counting: fundamentals 7

one may be able to determine the species represented by the paleozoological speci-
men, but other times only the genus or perhaps only the taxonomic family or order
will be distinguishable.
Taxonomic identi¬cation is a complex matter that is discussed at length in other
contexts (e.g., Driver 1992; Lyman 2005a; and references therein). Blind tests of iden-
ti¬cation results (e.g., Gobalet 2001) highlight the practical and technical dif¬culties.
For one thing, what is “identi¬able” to one analyst may not be to another (e.g.,
Grayson 1979). Gobalet (2001) provides empirical evidence for such interanalyst
variation. It is precisely because of such interobserver differences and the interpre-
tive signi¬cance of whether, say, a bone is from a bobcat (Lynx rufus) or a North
American lynx (Lynx canadensis) that paleontologists developed a standardized for-
mat for reporting their results. Specimens (not necessarily anatomically incomplete
skeletal elements) are illustrated and are verbally described with taxonomically dis-
tinctive criteria highlighted so that other paleontologists can independently evaluate
the anatomical criteria used to make the taxonomic identi¬cation. Zooarchaeolo-
gists have been slow to understand the importance of this reporting form (see Driver
[1992] for a noteworthy exception). This is not the place to delve further into the
nuances of taxonomic identi¬cation and how to report and describe identi¬ed speci-
mens. What is important here is to note that skeletal remains “ faunal specimens “ are
usually tallied by taxon. “There are X remains of bobcats and Y remains of lynx.” So,
identi¬cation must precede tallying. To make taxonomic identi¬cations, one must
¬rst determine which skeletal element is represented by a specimen is order to know
whether the paleozoological unknown should be compared to femora, humeri, tib-
iae, and so on. And sometimes the frequencies of each skeletal element or each part
thereof are analytically important.
The ¬nal paleozoological concept that requires de¬nition is taphonomy. The term
was originally coined by Russian paleontologist I. A. Efremov (1940:85) who de¬ned
it as “the study of the transition (in all details) of animal remains from the bio-
sphere into the lithosphere.” Although not without precedent, Efremov™s term is the
one paleozoologists (and an increasing number of paleobotanists) use to refer to
the processes that in¬‚uence the creation and preservation (or lack thereof) of the
paleobiological record. We will have reason to return again and again to this basic
concept; here it suf¬ces to note that a taphonomic history concerns the formation
of an assemblage of faunal remains. Such a history begins with the accumulation
and deposition of the ¬rst specimen, continues through the deposition of the last
specimen, through the preservation, alteration, and destruction of remains, and up
to collection of a sample of the remains by the paleozoologist (see Lyman [1994c]
for more complete discussion). Along the way, faunal remains are modi¬ed, broken,
and even destroyed. The modi¬cation, fracture, and destruction processes create
quantitative paleozoology

and destroy different kinds of phenomena the observation of which can generate
quantitative data.
A ¬nal note about how paleozoological data are presented in the book. Capital
letters are used to denote upper teeth, lower case letters to denote lower teeth, and
a lowercase d to denote deciduous premolars. Thus, a permanent upper second
premolar is P2, a deciduous lower third premolar is dp3, and a lower ¬rst molar is
m1. The capital letter L is used to signify the left element of bilaterally paired bones,
and the capital letter R is used to signify the right element. In general, D stands for
distal, and P stands for proximal. The critical thing to remember is the difference
between a specimen and a skeletal element; both terms will reappear often in what
follows, and both kinds of units can be identi¬ed and tallied.


This book is about quanti¬cation, but the topics covered include different sorts of
quanti¬cation, particularly counting or tallying units, methods of counting, and
analyzing counts. A term that might have been used in the title of the book, were
it not for its generality, is measurement. Typically this term is de¬ned as assigning a
numerical value to an observation based on a rule governing the assignment. The
rule might be that length is measured in linear units of uniform size, such that we
can say something like “Pencil A is 5 cm long and pencil B, at 10 cm of length,
is twice as long as pencil A.” Measurement more generally de¬ned concerns writing
descriptions of phenomena according to rules. An estimate is a measurement assigned
to a phenomenon (making a measurement) based on incomplete data. The process
of estimation can involve judging how tall someone is in centimeters without the
bene¬t of a tape measure, or studying a ¬‚ock of birds and suggesting how many
individuals there are without systematically tallying each one. Making estimates,
like taking measurements, is a way to describe phenomena. Descriptions involve
attributes of phenomena that may or may not have numerical symbols or values
associated with them. Whether they do or not concerns what is often referred to as
the scale of measurement of the attribute that is under scrutiny.

Scales of Measurement

Stock™s census of Rancho La Brea mammals (Figure 1.1 ) illustrates that quanti-
tative data describing taxonomic abundances are more revealing than taxonomic
tallying and counting: fundamentals 9

presence“absence data. Quantitative data often are subjected to a variety of math-
ematical manipulations and statistical analyses. Those manipulations and analyses
are only valid if the data are of a certain kind. Four distinct scales of measurement
are often distinguished (Blalock 1960; Shennan 1988; Stevens 1946; Zar 1996), and it
is important that these be explicitly de¬ned at the start because they will be referred
to throughout the book.
Nominal scales of measurement are those that measure differences in kind. Of
the several scales they contain the least amount of information. Numbers may be
assigned to label nominal scale phenomena, such as 0 = male, 1 = female; or 11 =
quarterback, 32 = fullback, and 88 = wide receiver on a football team. Or, numbers
need not be assigned, but rather labels used such as Italian citizen, French citizen,
and German citizen; or coyote (Canis latrans), wolf (Canis lupus), and domestic
dog (Canis familiaris). Nominal scales of measurement do not include an indication
of magnitude, ordering, or distance between categories, and are sometimes labeled
qualitative attributes or discontinuous variables. They are qualitative because they
record a phenomenon in terms of a quality, not a magnitude or an amount. They are
discontinuous (or discrete) because it is possible to ¬nd two values between which
no other intermediate value exists; there is (normally) no organism that is halfway
between a male and a female within a bisexual species. Other scales of measurement
tend to be quantitative because they specify variation more continuously. Continuous
variables are those that can take any value in a series, and there is always yet another
value intermediate between any two values. A tally of skeletal specimens of coyote
in an archaeological collection may be 127 or 128, but there won™t be a collection
in which there are 127.5 or 127.3 or 127.924 specimens of coyote. But the lengths of
coyote humeri are continuous; think about the numbers just noted as millimeters of
Ordinal scales of measurement are those that record greater than, less than relation-
ships, but not the magnitude of difference in phenomena. They allow phenomena
to be arranged in an order, say, from lesser to greater. “I am older than my children”
is a statement of ordinal scale difference, as is “The stratum on the bottom of the
stratigraphic column was deposited before the stratum on the top of the column”
and “A year is longer than a month.” There is no indication of the magnitude of
difference in my age and the ages of my children, or in the length of time between
the deposition of the bottom and top strata, nor in the duration of a year relative
to the duration of a month. Instead, we only specify which phenomenon is older
(or younger), or which was deposited ¬rst (or last), or which is longer in duration
(or shorter). Sometimes when one uses an ordinal scale, measurements are said to
be relative measurements because a measure of phenomenon A is made relative to
quantitative paleozoology

phenomenon B; A is older/shorter/heavier than B. Ordinal scale measurements may
be (and often are said to be) rank ordered from greatest to least, or least to greatest,
but the magnitude of distance between any two measurements in the ordering is
unknown. Ordinal scale measurements are discrete insofar as there is no rank of
“¬rst and a half” between the rank of ¬rst and second (ignoring tied ranks).
Interval scales of measurement are those that record greater than or less than
relationships and the magnitude of difference between phenomena. Both the order
of measurements and the distance between them are known. My children are 23 and
25 years old; I am 56 years old, so I am 33 and 31 years older than my two children,
respectively. The stratum on the bottom of the stratigraphic column has an associated
radiocarbon date of 3000 BP and the stratum on top has an associated date of 500 BP,
so the stratum on the bottom was deposited about 2,500 (14 C) years before the stratum
on top (assuming the dated materials in each stratum were formed and deposited at
the same time as the strata were deposited). On average, a year is 365.25 days long
whereas an average month is about 30.4 days in duration; the difference in duration
of an average year and an average month is thus 334.85 days. The distance between 10
and 20 units (days, years, centimeters) is the same as the distance between 244 and
254 of those units, the same as the distance between 5337 and 5347 of those units, and
so on. Interval scales are typically used to measure what are referred to as quantitative
variables. Interval scale measurements, like ordinal scale ones, can be rank ordered
from greatest to least, or least to greatest, but unlike with ordinal scale measures, the
distance between any two interval scale measurements is known. Indeed it must be
known else the variable is not interval scale. Interval scale measurements are generally
continuous but may be discrete. If age is recorded only in whole years, then age is
continuous but it is also discrete (ignoring for the sake of discussion that one might
be 53.7 years old). Importantly, interval scale measures have an arbitrary zero point.
It can be 0—¦ Celsius outside, but there is still heat (if seemingly only a little) caused
by the movement of molecules. The zero point on the Celsius scale is placed at a
different location along the continuum of amount of molecular movement than is
the zero point of the Fahrenheit temperature scale. Both zero points are arbitrary
with respect to the amount of heat (molecular movement), thus both measures of
temperature are interval scale.
Ratio scales of measurement are identical to interval scales but have a natural zero.
Thus, the theoretical natural zero of temperature is “273 —¦ Celsius (or 0 Kelvin, or “
459—¦ Fahrenheit). There is no molecular movement at that temperature. Similarly,
a mammal in a cage comprises 1 individual consisting of more than 100 bones and
teeth, but if the cage is empty there are 0 (zero) individuals, 0 bones, and 0 teeth in
the cage. Thus, if a taxon is represented by 0 skeletal specimens in an assemblage, it is
tallying and counting: fundamentals 11

absent; that is a natural zero. Essentially all quantitative measures in paleozoology “
taxonomic abundances, frequency of gnawing damage, and so on “ are potentially
ratio scale. Whether they are in fact ratio scale or not is another matter.
Measurements of different scales allow (or demand, depending on your perspec-
tive) different statistical tests of different scales or power. Thus, ordinal scale mea-
surements require ordinal scale statistical tests; interval/ratio scale measurements
can be analyzed with either interval/ratio scale statistics or ordinal scale ones, but the
reverse “ applying interval/ratio scale statistics (or parametric statistics) to ordinal
scale data “ will likely violate various statistical assumptions.
Most paleozoologists working in the twentieth century sought ratio scale measures
of the attributes of the ancient faunal remains they studied, just as Stock and Howard
did. Although the optimism that such measures would eventually be designed has
waned somewhat, there are still many who hope for such, whether working with
human remains (e.g., Adams and Konigsberg 2004), paleontological materials (e.g.,
Vermeij and Herbert 2004), or zooarchaeological collections (e.g., Marean et al.
2001; Rogers 2000a). We now know a lot more about taphonomy than we did even
20 years ago when Grayson (1984), Klein and Cruz-Uribe (1984), and Hesse and
Wapnish (1985) noted that many problems with quantitative zooarchaeology orig-
inated in taphonomic histories. And we also know that many taphonomic analyses
and interpretations of taphonomic histories require quantitative data and analy-
ses of various sorts. Where taphonomy can in¬‚uence quantitative paleozoology is
noted throughout this volume, and it is occasionally suggested what we might do
about those in¬‚uences. The point here is that ratio scale measurements of faunal
remains and many of their attributes may be precluded because of taphonomic

Measured and Target Variables: Reliability and Validity

Other important statistical concepts concern the difference between a measured vari-
able and a target variable. A measured variable is what we actually measure, say, how
many gray hairs I have on my head. A target variable is the variable that we are
interested in, say, my age. The critical question is this: Are the measured variable and
the target variable the same variable, or are they different? If the latter, the question
becomes: Are the two variables suf¬ciently strongly correlated that measuring one
reveals something about the other? It is likely that the number of gray hairs on my
head will be correlated with my age, assuming I do not arti¬cially color my hair
(either not gray, or gray). But although the color of the shirt I am wearing today
quantitative paleozoology

can be measured rather precisely, it is unlikely to indicate or correlate with my age
(although the style of my shirt might).
The concepts of measured variable and target variable can be stated another way.
When we measure something, are we measuring what we think we are measuring?
Does the attribute we are measuring re¬‚ect the concept (e.g., length, age, color) we
wish to describe (Carmines and Zeller 1979)? These questions serve to de¬ne the
concept of validity. Is a radiocarbon age on a piece of burned wood a valid measure
of the age of deposition of a fossil bone with which the wood is stratigraphically
associated? Assuming no contamination of the sample of wood, and that the wood
was deposited more or less simultaneously with the bone, it will be a valid measure if
it derives from a plant that was alive at about the same time as the animal represented
by the bone. Validity is a different property of a measurement than reliability, which
simply de¬ned means replicability, or, if we measure something twice, do we get the
same answer? If, on the one hand, we measure the length of a femur today and get
12.5 cm, tomorrow we measure it and get 12.4 cm, and the next day we measure it and
get 12.5 cm, then we are producing rather consistent and thus reliable measures of
that femur™s length. On the other hand, femur length is unlikely to be a valid measure
of the time period when the represented animal was alive, regardless of the reliability
of our measurements of length.
Another set of measurements will help underscore the signi¬cance of the preceding
paragraph, and help highlight the differences between a target variable and a mea-
sured variable. A fundamental measurement (sometimes referred to as primary data
[Clason 1972; Reitz and Wing 1999]) is one that describes an easily observed property
of a phenomenon. Length of a bone, stage of tooth eruption in a mandible, and
taxon represented by a shell are all fundamental measurements. A derived measure-
ment (sometimes referred to as secondary data) is more complex than a fundamental
one because it is based on multiple fundamental measurements. Derived measure-
ments are de¬ned by a speci¬ed mathematical (or other) relation between two or
more fundamental measurements. A ratio of length to width exempli¬es a derived
measure. Derived measurements require analytical decisions above and beyond a
choice of scale; do we calculate the ratio of length to width, or width to length,
or width to thickness? As a result, derived measurements are sometimes dif¬cult
to relate clearly to theoretical or interpretive concepts. Derived measurements may
nevertheless reveal otherwise obscure patterns in data even though relating those
patterns to a target variable may be dif¬cult.
The MNI measure mentioned above is the most widely known derived measure-
ment in paleozoology. It depends on (i) tallies of (ii) each kind of skeletal element
of (iii) each taxon in a collection, and often (iv) (but not always) other information,
such as size of bones of a taxon. Each of the lower case Roman numerals denotes
tallying and counting: fundamentals 13

a distinct fundamental measurement; each plays a role in deriving an MNI, as can
several other fundamental measurements (considered in more detail in Chapter 2).
A ¬at or proxy measurement will likely be more complex than either a fundamental
or derived measurement because a ¬at measurement is more conceptual or abstract
and less easily observed. The distinction of fundamental, derived, and proxy mea-
surements is relevant to a measurement™s accuracy. “Accuracy” refers to “the nearness
of a measurement to the actual value of the variable being measured” (Zar 1996:5).
Throughout this volume, major concerns are the accuracy and validity of derived
measures or secondary data, and fundamental measures or primary data with respect
to a target variable of interest. Does a particular derived measure, such as MNI, accu-
rately re¬‚ect the abundance of individual organisms in a collection of bones and
teeth (or shells)? Of organisms in a deposit? Of organisms on the landscape?
Stock™s census of Rancho La Brea mammals was, he hoped, an accurate proxy
measure of the structure and composition of the mammalian fauna on the landscape
at the time of the deposition of the remains. That long-dead fauna is not directly
visible or measurable, so how well the remains from the tar pits actually re¬‚ect or
measure that fauna in terms of which taxon was most abundant and which was
least abundant and a host of other properties (how accurately MNI measures the
landscape fauna) cannot be determined. The validity of a ¬at or proxy measurement,
or a measured variable, for re¬‚ecting a target variable of some sort is the key issue
underpinning much of the discussion in this volume. This is so for the simple reason
that many target variables in paleozoology cannot be directly measured reliably or
validly with broken bones, isolated teeth, and fragments of mollusk shell. What this
book is in part about is how well the measured variables and proxy measurements
commonly used by paleozoologists measure or estimate the target variable(s) of
interest. Two key questions to keep in mind throughout this book are: What is the
target variable? How is the measured variable related to the target variable of interest?
As a prelude to how important these questions are, think about this. Was Stock wise
to use MNI (the derived and measured variable) to estimate the abundances of
mammals on the landscape (the target variable) given that he only had animals that
became mired in the pits of sticky tar at Rancho La Brea? Would he have been
better off using, say, the tally of skulls (a different measured variable) to estimate the
abundances of mammals trapped in the tar pits (a different target variable)?

Absolute and Relative Frequencies and Closed Arrays

An absolute frequency is a raw tally of some set of entities, usually all of a particular
kind. To note that there are ten rabbit bones and ¬ve turkey bones in a collection is
quantitative paleozoology

to note the absolute frequencies of specimens of each species. If one were to note that
in that collection of ¬fteen specimens, 66.7 percent of the specimens were of rabbits
and 33.3 percent were of turkey, then one would be noting the relative frequency
of each species. Relative frequencies are termed such because they are relative to
one another. A relative frequency is a quantity or estimate that is stated in terms of
another quantity or estimate. The analyst could have different absolute abundances,
say thirty rabbit bones and ¬fteen turkey bones, but rabbit bones would comprise the
relative abundance of 66.7 percent of the collection and turkey bones would comprise
33.3 percent of that collection, the same as when there are ten rabbit bones and ¬ve
turkey bones. Percentages and proportions of a total are relative frequencies. The
term “relative frequencies” is sometimes used in the paleozoological literature to
signify estimates in which a quantity is not stated but rather only that A is greater (or
smaller, or less) than B. In such cases relative frequencies are equivalent to ordinal
scales of measurement. In this volume, the term “relative frequencies” is used in the
more typical sense of percentage or proportional abundances.
Relative frequencies are typically given as percentages of some total set of things,
and the summed relative frequency is always 100 percent (proportions are fractions).
When relative frequencies of kinds of things in a set of things are given as percent-
ages, all of those frequencies must sum to 100 percent rather than 90 percent or 110
percent. Such percentage relative frequencies comprise what is called a closed array
(proportions also form a closed array as they must sum to 1.0).
Another way to think about the difference between absolute and relative frequen-
cies involves comparison of measurements. Let™s say we have two collections of faunal
remains. In collection 1, taxon A is represented by 5 specimens and taxon B is rep-
resented by 10 specimens. In collection 2, taxon A is represented by 50 specimens
and taxon B is represented by 55 specimens. The absolute difference in abundances
of the two taxa in each collection is 5 specimens, but in collection 1, taxon A is only
50 percent as abundant as taxon B whereas in collection 2 taxon A is 90.9 percent as
abundant as taxon B. Or, one could say that in collection 1 the relative abundances
of taxa A and B are 33.3 percent and 66.7 percent, respectively, whereas the relative
abundances of those taxa in collection 2 are 47.6 percent and 52.4 percent, respec-
tively. The difference between absolute and relative frequencies is not a matter of
which is correct and which is not, but rather they are simply two different ways to
measure (describe) the frequencies of things.
Importantly, the absolute frequency of things of kind A in a collection will not
change value if the absolute frequency of kind B in that collection changes, but the
relative frequency of both A and B will change if the absolute frequency of either A
or B changes. This last property is a characteristic “ one could say diagnostic “ of
tallying and counting: fundamentals 15

Table 1.2. Fictional data on the absolute abundances
of two taxa in six chronologically sequent strata

Taxon A Taxon B
Stratum VI 50 (71.4) 20 (28.6)
Stratum V 50 (62.5) 30 (37.5)
Stratum IV 50 (55.6) 40 (44.4)
Stratum III 50 (50.0) 50 (50.0)
Stratum II 50 (45.4) 60 (54.6)
Stratum I 50 (41.7) 70 (58.3)

Relative (percentage) abundances in parentheses.

closed arrays; they must sum to 100 percent. Consider the set of ¬ctional data in
Table 1.2. If we examine these data, we see that Taxon A does not change in abso-
lute abundance over the stratigraphic sequence, but Taxon B does change in abso-
lute abundance. However, relative abundance data suggest that both taxa change in
abundance. This example reveals a ¬nal and critically important aspect of abundance
In paleozoology, absolute abundance data or raw tallies are often given, but when it
comes to interpreting abundance data, it is in terms of relative abundances. Using the
¬ctional data in Table 1.2 as an example, one might read something like the following:

Throughout the stratigraphic sequence (from stratum I as earliest or oldest to stratum
VI as youngest) Taxon A increased in abundance relative to Taxon B, which decreased in
relative abundance. Given that Taxon A prefers habitats that support vegetation adapted
to cool-moist climatic conditions, and Taxon B prefers habitats indicative of warm-dry
climatic conditions, then it seems that over the time span represented by strata I“VI,
the local climate became progressively cooler and moister.

Notice that in the interpretation no mention is made of the absolute abundances
of taxa A and B. Rather, their abundances relative to each other are the focus. The
abundances are not even taken as measures of how many of either taxon was present
on the landscape at the time the strata and faunal remains were deposited. Rather,
the interpretation involves postulating a cause for the shift in relative abundances of
two taxa. One could also postulate that hunting practices or procurement technology
shifted, resulting in the shift in which taxon was taken more frequently. Resolving
these sorts of issues is beyond the scope of this volume, but suf¬ce it to say that
regardless of the interpretive model one calls upon, relative abundances, in the case
of this example relative taxonomic abundances, are interpreted.
quantitative paleozoology


Quantitative data often comprise tallies of different kinds of phenomena. They might
also include a set of measurements of, say, the length of individual specimens. This
book concerns only the former kind of quantitative data “ tallies. It is about how
a paleozoologist might count phenomena (faunal specimens, or attributes thereof)
when one seeks a measure of the magnitude of a particular variable that demands
counts of phenomena (bones, teeth, shells, and fragments thereof, or burned bones,
gnawed bones, or broken bones). How one chooses to tally those phenomena, and
how those tallies are summarized and analyzed statistically, depend in large part on
the research question asked and the target variable that one hopes to measure in
order to answer that question. The choices likely will also depend on the presumed
relationship of the target variable and the chosen measured variable. Discussion of
how one determines the nature of that relationship in particular cases is beyond
the scope of this volume. When necessary to assist discussions in later chapters, a
particular relationship is assumed or identi¬ed.
The terms assemblage or collection denote an aggregate of faunal remains whose
setness has been de¬ned archaeologically (e.g., remains from an excavation unit),
geologically (e.g., remains from a trash pit or a stratum), or analytically (e.g., all
remains of a taxon). Graphs are used whenever possible to exemplify and illustrate
concepts and analytical results, and to display relationships between variables. Statis-
tical analyses are kept relatively simple and are used to evaluate particular properties
of collections. In a few cases, statistical complexities are described in a clearly delin-
eated box of text and may be skipped when reading the main content of a chapter.
Data are often presented in table form so that the reader may replicate the statistical
analyses (and graphs) to ensure understanding. This volume is, however, not meant
to be exhaustive with respect to all of the myriad ways that faunal remains might
be counted, or with respect to how the varied features faunal remains might display
can be counted. Rather, most of the commonly used quantitative units (measured
variables) and their attendant analyses serve as the background against which the
discussion is framed. Target variables are identi¬ed and de¬ned as necessary when
discussing particular quantitative units.
This is not a book about taphonomic, zooarchaeological, or paleontological analy-
ses. There are several excellent titles on each of these topics that are presently available
(e.g., Lyman 1994c; Reitz and Wing 1999; Simpson et al. 1960, respectively). Quantita-
tive Paleozoology is meant as a supplement to those other volumes because it covers in
detail a limited range of topics relevant to various analytical methods and techniques
described in each of those other volumes.
tallying and counting: fundamentals 17


Throughout this volume, extensive use is made of data derived from actual zooarchae-
ological and paleontological collections of vertebrate, usually mammalian, remains.
In several cases, the mammalian remains from a set of modern owl pellets collected
in the 1990s are used to illustrate an analytical procedure or a concept (see Lyman
and Lyman [2003] and Lyman et al. [2001, 2003] for more details on this collection).
In the chapters that follow, many points are illustrated by analyzing faunas from var-
ious places and dating to various time periods. This helps to emphasize that many
properties of the paleozoological record are in at least one sense universal, by which
is meant that those properties are typically found in an average paleo-faunal assem-
blage. By “average” is meant typical and having multiple taxa (usually more than a
half-dozen) and multiple identi¬ed specimens for the total collection (usually more
than, say, 50 specimens). What are rather atypical if not rare or unusual are those
collections that have hundreds if not thousands of specimens, all representing the
same species. The well-known bison (Bison spp.) kill sites of North America do not
seem particularly rare because publications on them are numerous, but in terms of
the faunal record they are rather atypical. An even more unusual paleofauna would
be one consisting of only a couple identi¬ed specimens (say, < 10), each represent-
ing a unique taxon. When necessary, these sorts of relatively unusual collections are
mentioned, but otherwise typical faunas are used to illustrate quantitative concepts
and analyses.
Using the same faunal samples throughout, it will be easy to track different kinds
of interdependence, and how one analytical result in¬‚uences whether or not another
analysis is reasonable or even feasible. And, by the same token, if two faunal samples
from basically the same geographic area and dating to the same time period are
available, then other sorts of analytical insights can be gained. Thus two collections of
mammal remains are used to illustrate signi¬cant points in later chapters. Analyses
of the artifacts and features at each site are ongoing, so some of the background
information is terse and incomplete. The lack of information on particular aspects
of the collections will not make a difference to the points made in this volume.
The collections are zooarchaeological “ they originate in an archaeological context.
These are faunal remains that had associated artifacts; such assemblages are some-
times referred to as archaeofaunas. Both collections consist of mammal remains
recovered from two late-prehistoric sites within 10 km of one another. Both sites are
found in what is locally known as the Portland Basin or the Wapato Valley of north-
western Oregon state and southwestern Washington state. All mammalian remains
from both sites were recovered from one-quarter-inch mesh screens in the ¬eld.
quantitative paleozoology

The Meier site (35CO5) is located downstream (north) of modern Portland, Oregon,
on the ¬‚oodplain of the Columbia River, on the Oregon side. The Meier site com-
prises a single large cedar-plank house that was occupied more or less continuously
between approximately ad 1400 and ad 1800, and associated midden deposits (Ames
1996; Ames et al. 1992). The site was tested in 1973 and 1984. It underwent extensive
excavations every year between 1987 and 1991, inclusively. The 1973 collection was
made by Pettigrew (1981) and studied by Saleeby (1983). The 1984 test was directed
by Ellis (n.d.); recovered faunal remains have not been analyzed. Kenneth Ames of
Portland State University directed the excavations that took place in the late 1980s
and early 1990s. I identi¬ed all mammalian remains collected by Ames during a 1993
research-sponsored leave when I worked with him in Portland.
The other site, Cathlapotle (45CL1), is located northeast of Meier, on the Wash-
ington side of the Columbia River, on a series of levees next to the river. The site
was visited by Meriwether Lewis and William Clark in March of 1806 as they lead
the Corps of Discovery eastward. At the time of their visit the site comprised several
large cedar-plank houses and associated midden deposits (Ames et al. 1999; Ames and
Maschner 1999:110). Radiocarbon dates indicate the main occupation began about
ad 1450. Ceramic trade goods indicate that abandonment of the site occurred about
ad 1834. Auger sampling of the Cathlapotle sediments took place in 1992“1993, and
excavations took place each year from 1993 through 1996. Both the auguring and the
excavations were under the direction of Ames. I identi¬ed all mammalian remains
recovered from this site at the University of Missouri-Columbia campus.
The assemblages of mammalian remains from Meier and Cathlapotle were recov-
ered from similar depositional contexts. At both sites, the deposits variously comprise
exterior (midden and “yard”) deposits and interior deposits (inside of a house). Exte-
rior deposits had very high organic content, lenses of fresh-water mussel shells, and
other indications that they formed as primary or secondary dumps (Ames et al. 1999).
Yard deposits are generally broad, sheet-like deposits that contain intact hearths,
activity areas, pits, evidence of small structures, and so forth. They usually lack the
very high organic content of middens though they can have organic content. Inte-
rior deposits were assigned to walls, benches (deposits below the 2 m-wide sleeping
benches or platforms that ran along the interior side of the house walls), storage pits,
and hearth areas. Faunal remains have not been sorted into these distinct depositional
contexts as yet. Were they to be so assigned, it is likely that assemblages would be
quite small. In later chapters, the in¬‚uences of small sample sizes receive considerable
The houses at Meier and Cathlapotle had extensive sub¬‚oor storage features that,
at Meier at least, formed a cellar almost 2 m deep that extended under the house
tallying and counting: fundamentals 19

Table 1.3. Description of the mammalian faunal record at Meier and at


Meier Precontact Postcontact

Didelphis— “ “ “ “ 10 1
Scapanus 14 4 “ “ 3 2
Sorex “ “ 3 1 1 1
Sylvilagus 16 2 “ “ 1
Lepus “ “ 3 2 40 4
Aplodontia 5 1 61 8 57 8
Tamias 1 1 “ “ “ “
Tamiasciurus 2 1 “ “ “ “
Thomomys 9 5 “ “ “ “
Castor 329 9 111 5 238 7
Peromyscus 35 21 2 1 3 2
Rattus— 1 1 “ “ “ “
Neotoma 1 1 “ “ “ “
Microtus 100 41 10 4 55 22
Ondatra 337 13 56 7 36 4
Erethizon 1 1 “ “ “ “
Canis 90 7 18 3 17 2
Vulpes 2 1 4 1 “ “
Ursus 82 5 45 3 53 4
Procyon 272 20 109 6 84 8
Martes 19 4 1 1 1 1
Mustela 130 17 12 3 9 2
Mephitis 4 2 3 1 “ “
Lutra 45 3 28 3 26 4
Puma 9 1 5 1 5 2
Lynx 22 3 8 1 15 2
Phoca 40 3 26 2 34 2
Ovis “ “ 1 1 1 1
Cervus 832 10 1,091 12 1,793 24
Odocoileus 3,504 56 775 14 1,347 27
Equus— “ “ “ “ 4 1
TOTAL 5,939 “ 2,372 “ 3,834 “

Taxa are historically introduced and not native to the area, hence they are intrusive
to site sediments.
quantitative paleozoology

¬‚oor between the sleeping platforms and the row of hearths in the house™s center. The
Cathlapotle features are less extensive, but are about 2 m wide by 2 m deep. They are
below the sleeping platforms rather than next to them as at Meier. The mammalian
remains from both sites were derived primarily from these storage pits and exterior
Because, with few exceptions, the mammalian genera identi¬ed are monotypic
(include only one species), the basic faunal identi¬cation and quantitative data are
presented by genus (Table 1.3). Stratigraphy at Meier is extremely complex as a
result of multiple episodes of house remodeling and rebuilding; temporally distinct

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