THESIS
CARNIVORE ATTRITION OF THE KAPLAN-HOOVER BISON BONEBED: LATE
HOLOCENE PREDATORY ECOLOGY OF THE CACHE LA POUDRE BASIN,
COLORADO PIEDMONT
Submitted by
Chrissina C. Burke
Department of Anthropology
In partial fulfillment of the requirements
For the degree of Master of Arts
Colorado State University
Fort Collins, Colorado
Summer 2008
COLORADO STATE UNIVERSITY
May 9th, 2008
WE HEREBY RECOMMEND THAT THE THESIS PREPARED UNDER
OUR SUPERVISON BY CHRISSINA COLEEN BURKE ENTITLED CARNIVORE
ATTRITION OF THE KAPLAN–HOOVER BISON BONEBED: LATE
HOLOCENE PREDATORY ECOLOGY OF THE CACHE LA POUDRE BASIN,
COLORADO PIEDMONT BE ACCEPTED AS FULFILLING IN PART
REQUIREMENTS FOR THE DEGREE OF MASTER OF ARTS.
Committee on Graduate Work
__________________________________________________
Kenneth J. Berry
__________________________________________________
Jason M. LaBelle
__________________________________________________
Advisor: Lawrence C. Todd
__________________________________________________
Anthropology Department Chair: Kathleen A. Galvin
ii
ABSTRACT OF THESIS
CARNIVORE ATTRITION OF THE KAPLAN–HOOVER BISON BONEBED:
LATE HOLOCENE PREDATORY ECOLOGY OF THE CACHE LA POUDRE
BASIN, COLORADO PIEDMONT
This thesis presents the results of zooarchaeological, taphonomic, and
ethological investigations of carnivore modification at the Kaplan–Hoover bison
bonebed (5LR3953) in Windsor, Colorado. Kaplan–Hoover is a Late Archaic
Yonkee bison bonebed dated to approximately 2724+/-35 RCYBP. Prehistoric
hunters used an arroyo to trap approximately 200+ bison. After the kill, limited
use of the carcasses by hunters left a surplus of bison meat available for nonhuman scavengers and predators. Carnivore attrition is present on over 40% of
the limb bones included in this study. Taphonomic analysis indicates that the
Kaplan–Hoover collection was modified and used by a range of non-human
scavengers. Using an interdisciplinary approach to methodology as well as
identifying key patterns relevant to a variety of fields of research, including
conservation biology is done. This thesis demonstrates how biogenic factors
influence the taphonomy of a faunal assemblage. In addition, this project is a
iii
part of the push to integrate zooarchaeological research and conservation
management decisions currently occurring in the field of archaeology. This
assessment suggests that in order to understand human interactions with present
and future environments, a researcher must first understand the prior behaviors
that assisted in the development of those events.
Chrissina Coleen Burke
Anthropology Department
Colorado State University
Fort Collins, CO 80523
Summer 2008
iv
ACKNOWLEDGEMENTS
This thesis is the culmination of a lot of lab time, tears, and excitement. I would have
never completed it, let alone even began it without the encouragement and help from a good
many people. First of all I could not have done this thesis without my advisor Dr. Larry Todd.
Not only has he been instrumental in my success here at CSU, but he is a major reason I was able
to preserver in all of the difficult situations I have encountered during my stay. I cannot say
enough wonderful things about Dr. Todd. He is an amazing mentor, teacher, friend, and
supporter. I have never felt respect from another faculty member as I have from Dr. Todd. His
ability to discuss every interest a student has in archaeology and help push them along to greater
and greater heights is one of amazement. I appreciate his willingness to take me on as a student
and I will always strive to be a mentor to my future students as he was with me.
Dr. Jason LaBelle also deserves a great deal of credit for helping me discover my path in
archaeology. His encouragement has always been of the utmost importance and his willingness
to sit and talk every research topic, obstacle, and success out thoroughly with me has been
extremely helpful. When I was frustrated and worried I couldn’t finish, Dr. LaBelle managed to
keep me going and help me realize that the whole process of graduate school is difficult for
everyone and that the ability to keep going was a trait that was developed and learned from
understanding set backs and problems. Dr. LaBelle is a great professor and mentor and has been
an important influence for the type of mentor I would like to be in the future.
Dr. Kenneth Berry was an important addition to my committee and helped ensure that my
statistics were correct and spent a number of hours discussing graduate school and life with me.
His encouragement and willingness to join my committee has been greatly appreciated and his
v
advice for future research and instruction on proper formatting was very helpful for the
completion of this thesis.
I could not have finished writing this thesis without the support of my friends. Bethany
Mizushima and Benjamin Jewell (nerd corp!) have been incredibly supportive in every facet of
the writing process. Without them I would not have completed let alone stayed sane the entire
time. When I thought I couldn’t keep writing, I knew I could always talk to either one of them
and they would help me get back on track, even if they were “culturals” and didn’t know what I
was talking about. Erik Otárola-Castillo has also been an important friend for and mentor in my
pursuits with archaeological research. His willingness to include me in research projects has not
only been very helpful for my future, but has also increased my understanding of working with
colleagues on presentations and papers. Lastly, Robin Roberts has been an important part of this
support network. Her ability to encourage and support my every decision and ability to always
push me to have fun has increased my sanity as well.
Finally, Abe Thompson deserves an award for living with me while I was writing my
thesis and not killing me during the many times I unloaded my stress and frustration on him. I
can not imagine living with me while I was panicking and he magically did so without moving
out or running away! My sister Michelle was and always has been my biggest supporter. No
matter how much doubt I have had about myself she has always told me that she knew I could do
it. She also saved my thesis by editing the 100 or so pages, while taking care of her own family.
To everyone above, you are greatly appreciated and without you this thesis would have
never been done. I appreciate your influence in my life every single day and hope that in the
future, I will be able to repay your thoughtfulness and I promise to always support your dreams
and goals.
vi
TABLE OF CONTENTS
ABSTRACT OF THESIS ................................................................................................. iii
ACKNOWLEDGEMENTS.............................................................................................. v
TABLE OF CONTENTS ................................................................................................ vii
LIST OF FIGURES........................................................................................................... ix
LIST OF TABLES............................................................................................................. xi
CHAPTER 1: INTRODUCTION .................................................................................... 1
Questions for Research................................................................................................ 1
Site Description and Information .............................................................................. 4
Kaplan–Hoover and Other Yonkee Bison Kill Sites................................................ 8
Summary of Chapters................................................................................................ 13
CHAPTER 2: FOUNDATIONS FOR RESEARCH .................................................... 15
Methodological Changes to Understanding Biogenic Factors ............................ 16
FAUNMAP: Choosing Non-Human Scavengers to Explore............................... 20
Ethological Research: Understanding Scavenging Behaviors ............................. 27
Conservation Research.............................................................................................. 41
Summary of Chapter ................................................................................................. 50
CHAPTER 3: AN INTERDISCIPLINARY APPROACH TO METHODS .............. 53
Data Collection Procedures ...................................................................................... 54
Extant Non-human Scavengers: Ethological Methods......................................... 75
Summary of Chapter ................................................................................................. 76
CHAPTER 4: RESULTS OF DATA ANALYSIS......................................................... 78
Herd Characteristics Analysis.................................................................................. 79
Differential Destruction Analysis ............................................................................ 90
Carnivore Modification Analysis .......................................................................... 104
CHAPTER 5: CONCLUSIONS AND FUTURE DIRECTIONS ............................. 124
Implications for Conservation Research............................................................... 133
Future Directions ..................................................................................................... 136
Literature Cited ............................................................................................................ 142
APPENDIX A: FAUNMAP DATA............................................................................ 155
APPENDIX B: CODING SYSTEM............................................................................. 157
vii
APPENDIX C: LANDMARK AND MEASUREMENT DESCRIPTIONS AND
CODES........................................................................................................................... 159
APPENDIX D: KAPLAN-HOOVER DATA............................................................. 165
APPENDIX E: ANALYSIS OF SEX FOR SPECIFIC SKELETAL ELEMENTS .... 188
viii
LIST OF FIGURES
Figure 1.1: Late Archaic projectile ................................................................................. 6
Figure 1.2: Plan map of the Kaplan–Hoover bison bonebed ..................................... 8
Figure 1.3: Map of Late Archaic Yonkee bison kill sites........................................... 10
Figure 2.1: Number of sites with specific non-human scavengers ......................... 24
Figure 2.2: Percentage of non-human scavengers present in sites.......................... 26
Figure 2.3: Canis lupus maxilla ..................................................................................... 29
Figure 2.4: Carnassial pair of Canis lupus ................................................................... 30
Figure 2.5: Canis latrans maxilla ................................................................................... 32
Figure 2.6: Ursus arctos maxilla .................................................................................... 35
Figure 2.7: Ursus americanus maxilla ........................................................................... 40
Figure 3.1: Chipping back ............................................................................................. 67
Figure 3.2: Crenellations ............................................................................................... 67
Figure 3.3: Furrowing .................................................................................................... 68
Figure 3.4: Pitting ........................................................................................................... 68
Figure 3.5: Punctures ..................................................................................................... 69
Figure 3.6: Scooping out................................................................................................ 69
Figure 3.7: Tooth scoring............................................................................................... 70
Figure 3.8: Light utilization .......................................................................................... 71
Figure 3.9: Light/moderate utilization ........................................................................ 71
Figure 3.10: Light/moderate utilization ...................................................................... 72
Figure 3.11: Moderate utilization................................................................................. 72
Figure 3.12: Moderate/heavy utilization..................................................................... 73
Figure 3.13: Moderate/heavy utilization..................................................................... 73
Figure 3.14: Heavy utilization ...................................................................................... 74
Figure 3.15: Extreme heavy utilization ....................................................................... 74
Figure 4.1: Scatter plot of %MAU and bone mineral densities ............................ 103
Figure 4.2: Presence or absence of modification...................................................... 105
Figure 4.3: Percentage of carnivore utilization for the humerus........................... 106
Figure 4.4: Carnivore utilization for the radius-ulna.............................................. 110
Figure 4.5: Carnivore utilization for the femur........................................................ 113
Figure 4.6: Carnivore utilization for the tibia .......................................................... 117
Figure 5.1: Bear modification Bos taurus tibia from Dr. Haynes collection ......... 132
ix
Figure 5.2: Probable bear modification on Bison bison............................................ 132
Figure E.1: Sex analysis scatter plot for the humerus ............................................. 188
Figure E.2: Sex analysis scatter plot for the radius-ulna ........................................ 189
Figure E.4: Sex analysis scatter plot for the metacarpal ......................................... 190
Figure E.5: Sex analysis scatter plot for the metacarpal ......................................... 190
Figure E.6: Sex analysis scatter plot for the metacarpal ......................................... 191
Figure E.7: Sex analysis scatter plot for the metacarpal ......................................... 191
Figure E.8: Sex analysis scatter plot for the femur .................................................. 192
Figure E.9: Sex analysis scatter plot for the femur .................................................. 192
Figure E.10: Sex analysis scatter plot for the femur ................................................ 193
Figure E.11: Sex analysis scatter plot for the tibia ................................................... 193
Figure E.12: Sex analysis scatter plot for the tibia ................................................... 194
Figure E.13: Sex analysis scatter plot for the tibia ................................................... 194
Figure E.14: Sex analysis scatter plot for tibia.......................................................... 195
Figure E.15: Sex analysis scatter plot for the tibia ................................................... 195
Figure E.16: Sex analysis scatter plot for the metatarsal ........................................ 196
Figure E.17: Sex analysis scatter plot for the metatarsal ........................................ 196
Figure E.18: Sex analysis scatter plot for the metatarsal ........................................ 197
Figure E.19: Sex analysis scatter plot for the metatarsal ........................................ 197
Figure E.20: Sex analysis scatter plot for the astragalus......................................... 198
Figure E.21: Sex analysis scatter plot for the calcaneus.......................................... 198
Figure E.22: Sex analysis scatter plot for the calcaneus.......................................... 199
x
LIST OF TABLES
Table 2.1: FAUNMAP age categories.......................................................................... 22
Table 2.2: FAUNMAP species codes ........................................................................... 22
Table 4.1: Cross tabulation of side and sex for MNI analysis of the humerus...... 86
Table 4.2: Cross tabulation of side and sex for MNI analysis of the radius-ulna . 86
Table 4.3: Cross tabulation of side and sex for MNI analysis of the metacarpal.. 87
Table 4.4: Cross tabulation of side and sex for MNI analysis of the femur........... 87
Table 4.5: Cross tabulation of side and sex for MNI analysis of the tibia.............. 88
Table 4.6: Cross tabulation of side and sex for MNI analysis of the metatarsal ... 88
Table 4.7: Cross tabulation of side and sex for MNI analysis of the astragalus.... 89
Table 4.8: Cross tabulation of side and sex for MNI analysis of the calcaneus..... 89
Table 4.9: Bone mineral densities for Bison bison ..................................................... 92
Table 4.10: MNE portions for the humerus................................................................ 93
Table 4.11: MNE portions for the radius-ulna ........................................................... 94
Table 4.12: MNE portions of the femur....................................................................... 95
Table 4.13: MNE portions of the tibia ......................................................................... 96
Table 4.14: Frequency table of landmarks on the humerus ..................................... 98
Table 4.15: Frequency table of landmarks on the radius-ulna ................................ 98
Table 4.16: Frequency table of landmarks on the femur .......................................... 99
Table 4.17: Frequency table of landmarks on the tibia ........................................... 100
Table 4.18: Minimum animal units and % MAU values for Kaplan–Hoover..... 101
Table 4.19: %MAU based on MNE portion codes for Kaplan–Hoover................ 102
Table 4.20: Spearman correlation of %MAU and bone mineral density crosses 104
Table 4.21: Chi-square analysis of sex and carnivore utilization .......................... 107
Table 4.22: Chi-square analysis of sex and modification ....................................... 107
Table 4.23: Chi-square analysis of carnivore utilization and side......................... 108
Table 4.24: Carnivore utilization and side cross-tabulation .................................. 108
Table 4.25: Chi-square analysis of carnivore utilization and portion................... 109
Table 4.26: Carnivore utilization and portion cross-tabulation ............................ 109
Table 4.27: Chi-square analysis of carnivore utilization and portion................... 111
Table 4.28: Cross-tabulation of carnivore utilization and portion ........................ 112
Table 4.29: Chi-square analysis of sex and carnivore utilization .......................... 114
Table 4.30: Chi-square analysis of sex and modification ....................................... 114
xi
Table 4.31: Chi-square analysis of carnivore utilization and side......................... 115
Table 4.32: Carnivore utilization and side cross-tabulation .................................. 115
Table 4.33: Chi-square analysis of carnivore utilization and portion................... 116
Table 4.34: Carnivore utilization and portion cross-tabulation ............................ 116
Table 4.35: Chi-square analysis of sex and carnivore utilization .......................... 118
Table 4.36: Chi-square analysis of sex and modification ....................................... 118
Table 4.37: Chi-square analysis of carnivore utilization and side......................... 119
Table 4.38: Chi-square analysis of carnivore utilization and portion................... 119
Table 4.39: Bone mineral densities from bison ........................................................ 120
Table 4.40: Bone mineral densities for bison ............................................................ 121
xii
CHAPTER 1: INTRODUCTION
Questions for Research
In any archaeological site there are specific factors that illustrate formation
processes essential to understanding the spatial and temporal aspects of
paleoecology. If the site is composed of a dense bonebed of skeletal material, it is
necessary to assess the taphonomic factors that are associated. Of particular
interest to this thesis is carnivore modification, a biogenic factor that influences
the overall destruction and formation of the bonebed as well as illuminates the
predatory and scavenging behaviors of both human and non-human scavengers.
The goals of this thesis are wide-ranging, with basal questions and
integrated questions. The basal questions are useful for setting the framework
from which the integrated questions can be asked. The integrated questions
incorporate an understanding of the basal questions so that the information
presented by this thesis is useful for other researchers at a variety of different
faunal assemblages. Finally, these questions are meant to establish a framework
for analyzing carnivore modification in faunal assemblages and how that
1
information is useful for understanding human and non-human scavenger
interactions.
Basal Questions:
1. What are the herd characteristics at Kaplan–Hoover?
2. What types of carnivore modification and intensity of carnivore
utilization is present on the Kaplan–Hoover collection?
3. Where is the modification located on the appendicular skeleton and
which specific elements exhibited more destruction than others?
4. What non-human scavenger behaviors are important to understand
when discussing the taphonomy of a faunal assemblage?
Integrated Questions:
1. What relationships between prehistoric human hunters and nonhuman scavengers can be observed from the faunal remains at
Kaplan–Hoover?
2. How can these same methods be used for other sites?
3. Why is the interdisciplinary approach used in this thesis beneficial for
answering all of the preceding questions?
The first question, that will build the basis for the remaining questions
asks, what are the herd characteristics? It is essential to know the demographics
of the herd to understand if carnivores were selectively scavenging the remains,
therefore analysis on size and sex of the skeletal material must be accomplished.
The second question: where is the carnivore modification located on the skeletal
elements, is important to understanding how the collection was damaged. A
2
record of the types of carnivore modification present as well as where it is
located anatomically and in what abundances is relevant for comprehending
overall destruction of the material, utilization of the material, and site formation
processes. In conjunction with the remaining basal questions, ethological
literature is useful for understanding non-human scavenger behaviors,
specifically species that could be associated with the region the site is located
within.
The integrated questions for contemplation and analysis seek to
understand the interactions between human hunters and non-human scavengers
that can be observed by comparing and contrasting the answers from the basal
questions. How do the herd demographics and carnivore modification data
support inferences that specific carnivores were present at the site? Included will
be a discussion on how these data sets can be used to understand the overall
paleoecology of a region, further discussing present and future conservation
issues that may have resulted from the interactions created in the past by
humans between themselves and their environments
These questions are important for a number of reasons. First of all,
archaeologists are in a prime position to influence decision making in the natural
resources (Lyman and Cannon 2004:xv). Research done by archaeologists is
useful for understanding interactions between species through time, therefore,
3
having the ability to assist in management decisions for eradication or
reintroduction of species into habitats and regions. Currently, the natural
resources and social sciences are working in parallel directions with a distinct
lack of integration and collaboration. Instead of a parallel line of thinking by
both fields, an integrated line would benefit both in their pursuits.
In general, the questions presented by this research are important to
zooarchaeology and a number of archaeological, paleontological, and ecological
sites. When answered, these questions will provide a basic description of the
Kaplan–Hoover bison bonebed as well as an understanding of four non-human
scavenger species: Canis lupus, Canis latrans, Ursus arctos, and Ursus americanus.
Finally, this research describes an interdisciplinary methodological framework to
zooarchaeological research and carnivore modification.
Site Description and Information
The Kaplan–Hoover bison bonebed located in Windsor, Colorado is a
highly non-human scavenger modified Bison bison bonebed. Therefore, it is an
important example of why it is imperative to research the behaviors of carnivore
and rodent species to assess the specific mammals that altered the skeletal
elements after prehistoric peoples had left. To test the usefulness of the
methodological questions presented in the preceding paragraphs, the Kaplan–
Hoover site was used. Excavations were undertaken by Colorado State
4
University (CSU) from 1997-2001 and overseen by Dr. Lawrence C. Todd from
the CSU Anthropology Department.
Kaplan–Hoover is a late archaic (middle Holocene) bison arroyo trap and
single catastrophic event kill (Todd, et al. 2001:137). Located in Larimer County
Colorado, the site sits at an elevation of 1475 m and is located approximately 800
m north of the Cache la Poudre River (Todd, et al. 2001:126). Radiocarbon dating
was accomplished using several charcoal samples that were collected and
mapped in the bonebed (Todd, et al. 2001:132-133). Using AMS dating, two
larger chunks of charcoal were dated to 2740+/-40 and 860+/-40 radiocarbon years
before present (RCYBP) (Todd, et al. 2001:132-133). In addition, an intact
metatarsal was dated to 2690+/-60 RCYBP and when averaged with the date of
one of the charcoal chunks of 2740+/-40 RCYBP, Kaplan–Hoover is given a date
of 2724+/-35 RCYBP (Todd, et al. 2001:132-133).
The seasonality of the site has been determined as being a SeptemberOctober kill based on the eruption and wear of the bison mandibular molars
(Todd, et al. 2001:137). The site is composed of a dense accumulation (Figure 1.2)
of Bison bison skeletal material with the excavated portion measuring 4-5 m wide
and at least 1 m thick (Todd, et al. 2001:127-128). The minimum number of
individual animals (MNI) at the time of the first report was 44 based on the
crania recovered (Todd, et al. 2001:135). Additionally, 4000+ identifiable skeletal
5
elements have been removed from the site and are currently housed in the
Anthropology Department at Colorado State University, with exception of a
number of elements that were left in situ and thus not available for analysis
(Todd, et al. 2001). Estimations from minimum number of skeletal elements
(MNE) and number of identified specimens (NISP) suggest that the deposit could
hold roughly 200+ bison, therefore, 150 have yet to be exhumed (Todd, et al.
2001:135). From these data, the herd composition is approximately 33-39% bulls
and 61-67% females and sub-adults (Todd, et al. 2001:137). Research presented in
this thesis, which includes analysis of materials recovered subsequent to the 2001
publication, will change these numbers slightly.
Figure 1.1: Late Archaic projectile points from the Kaplan–Hoover site (Todd et al. 2001).
The majority of the skeletal elements missing from the bonebed are ribs,
thoracic vertebrae (hump meat), and femora all of which have high food utility
6
values (Todd, et al. 2001:136-137). In general the preservation of the elements is
excellent, with very little weathering cracks, which according to Todd, et al.
(2001:134) indicate burial of the remains soon after the animal’s death. However,
the site preservation is compromised due to heavy modification of the
abandoned skeletal elements by carnivores and rodents and post-depositional
deterioration.
Heavy modification due to carnivores is what separates the Kaplan–
Hoover bonebed from other sites in the prehistory of North America. During
initial examination of the skeletal elements 45 humeri were studied for degree of
carnivore damage, of those 80% had carnivore damage consuming the entire
proximal end and Todd et al. (2001:140) remark that “overall, 98% of the humeri
from the site have some carnivore damage.” These estimates surpass the Casper
site which has 37%, the Jones–Miller site with 28%, and the Bugas–Holding site
with 17% carnivore damage to the humeri (Todd, et al. 1987; Todd, 1997; Todd, et
al. 2001:140). In Chapters 4 and 5 discussions on the types of carnivore
modification and intensity of modification will clarify these numbers and give
more details pertaining to the remaining skeletal elements used for analysis in
this thesis.
7
Figure 1.2: Plan map of the Kaplan–Hoover bison bonebed (Todd, et al. 2001:134, Figure 7).
Kaplan–Hoover and Other Yonkee Bison Kill Sites
The projectile points recovered from Kaplan–Hoover include Yonkee
points, described from the Powers–Yonkee site (24PR5) in southeastern Montana
(Bentzen 1961; Bentzen 1962b; Bump 1987; Frison 1978; Roll, et al. 1992). Yonkee
points (Figure 1.1) are typically side and or corner notched with a basal notch or
indention (Frison 1978:50). The type site of this technology is the Powers–
Yonkee (24PR5) site, located in southern Montana (Bentzen 1961, 1962b; Bump
1987; Frison 1978). Other sites that contain Late Archaic Yonkee points include
Kobold (24BH406) in southern Montana, Powder River (48SH312), and
Mavrakis–Bentzen–Roberts (48SH311) both of which are in northern Wyoming
(Bentzen 1962a; Bump 1987; Frison 1968; Frison 1970; Frison 1978). The last site
8
to be discussed is Ayers–Frazier (24PE30), another bison trap in Montana (Clark
1981) (Figure 1.3).
Powers–Yonkee (24PR5) was initially dated to 4450+/-125 years before
present (Bentzen 1961). Several years later Bump (1987) dated bison skeletal
material to 2290+/-50 years before present. The site was excavated by the
Sheridan Chapter of the Wyoming Archaeological Society in August of 1961
(Bentzen 1961). The Powers–Yonkee bison kill is an arroyo trap that sits upon a
high terrace at about 1097 meters above sea level; the site is located on the north
bank of a small arroyo upon this terrace (Bentzen 1961). Bentzen (1961) states
that the bison were driven into the north-south branch of the arroyo and then
were shunted or trapped into the east-west branch of the arroyo. A bison kill is
represented at Powers–Yonkee, however, there are other faunal remains,
including one canid (Bump 1987:31). The remains are very well preserved and
major concentrations lie at approximately 89–104 cm below the surface sediments
(Bump 1987:33). As will be seen in most of the sites designated as Yonkee,
projectile points are usually recovered from rib skeletal elements (Bentzen 1961:7;
Frison 1968:32-33, 1978:203). Muscle stripping is evident from butchery cut
marks on specific elements as well at Yonkee sites (Frison 1968:33-34, 1978:203).
9
Figure 1.3: Map of Late Archaic Yonkee bison kill sites
on the Great Plains in Montana, Wyoming, and Colorado.
The Powder River site (48SH312) located in the Powder River Basin has
not been dated. The site was excavated in 1966 and is an arroyo trap single kill
event of Bison bison, with an MNI of approximately 12 (Frison 1968:32). Of the 25
projectile points found at the site, 16 are located inside rib or vertebral column
skeletal elements (Frison 1968:32-33). Butchering marks indicate that the hunters
removed meat that was easiest to obtain (Frison 1968:33-34). The majority of the
meat removed at Powers-Yonkee was hind limbs, thus leaving most of the
forelimbs present at the site (Frison 1968:33). In addition, hunters did not
remove the brains, hides, or tongues, further reinforcing the idea that the meat
10
was chosen based on easiest to access in the arroyo (Frison 1968:33). Finally, the
author makes mention that other damage to the skeletal elements is most likely
indicators of non-human scavenging behaviors, although he does not go into
detail on this topic (Frison 1968:33).
The Mavrakis–Bentzen–Roberts bison trap (48SH311) is a single kill event
located in the Powder River Basin just as the Powder River site (Bentzen 1962a;
Frison 1968, 1978). Mavrakis–Bentzen–Roberts site was excavated in 1962 and is
an arroyo trap kill with an MNI of approximately 17-26 bison (Bentzen 1962a:32).
Dated to 2600+/-200, site 48SH311 shows little evidence of marrow removal
because of few stone tools imbedded directly into shaft of the elements (Bentzen
1962a:32). Finally, 48SH311 shows butchering cut mark evidence of muscle
stripping similar to the other Yonkee sites (Frison 1978:206).
The Kobold site (24BH406) has not been dated, however, the projectile
points at the site are Yonkee points (Frison 1970b). Kobold is a multiple
component bison jump from a 7.62 meter cliff in southern Montana (Frison
1970b:7). Two of the levels at the site contain faunal remains, level two contains
badly decomposed material with some long bones broken for marrow with an
MNE of approximately 65 (Frison 1970b:15). The second level to contain faunal
materials is level four, which is mostly scapulae, humeri, radii, and metacarpals
with possible removal of marrow (Frison 1970b:22-23).
11
The final site is the Ayers–Frazier site (24PE30) dated to 2180+/-150 years
before present (Clark and Wilson 1981:23). The site was excavated in 1978 and is
interpreted as being a single kill event in an arroyo trap with 700 elements
recovered from excavations and approximately 300 more from a looter’s back
dirt pile (Clark and Wilson 1981:37). There is evidence of butchering in terms of
cut marks, chop marks, and skinning marks on the skeletal remains from Ayers–
Frazier (Clark and Wilson 1981:38). Of the 700 skeletal elements analyzed from
the test excavation area, approximately 15% have carnivore modification and the
authors go into great deal discussing the bone tools of the site, which appears to
be a description of the carnivore modification types of chipping back and
salivary polishing that will be defined later in this thesis (Clark and Wilson
1981:50–51).
Yonkee complex sites are similar in more ways than just the type of
projectile points. The bison kills from the Yonkee Late Plains Archaic are
typically arroyo traps with exception of the Kobold site, which is a jump
(Bentzen 1961, 1962; Bump 1987; Clark and Wilson 1981; Frison 1968, 1970b,
1978). Faunal material at Yonkee sites typically have points lodged in the ribs
and vertebral columns and butchering evidence suggests stripping of muscle
meat as well as some removal of marrow and long bones (Bentzen 1961, 1962b;
Bump 1987; Clark and Wilson 1981; Frison 1968, 1970b, 1978). With limited
12
carcass utilization, expecting non-human scavenger modification is reasonable.
The only authors discussing carnivore modification are Clark and Wilson with
the Ayers–Frazier site, which was published in 1981.
Focus on the Yonkee sites indicates that most of the sites with Yonkee
projectile points were arroyo traps and butchering patterns suggest that
prehistoric hunters did not use all of the bison meat available. At the Ayers–
Frazier site, there is a record of carnivore modification being present, suggesting
that a new analysis of the other sites may yield similar results considering that
the other publications were prior to intense taphonomic scrutiny at
archaeological sites.
Summary of Chapters
The remaining chapters will discuss a wide range of topics and end with a
synthesis of ideas. Chapter 2 discusses a number of background research
projects and methods. Chapter 2 begins by identifying the methodological
framework on carnivore attrition, then presents exploratory research using the
FAUNMAP database, and concludes with a discussion of non-human scavenger
behaviors and predator conservation issues in North America. Chapter 3
describes and explains the methods used to collect and analyze data from the
Kaplan–Hoover collection. Chapter 4 discusses the final results of analysis,
13
specifically discussing sex analysis, herd characteristics, carnivore modification,
and overall description of carnivore destruction on the collection. Finally
Chapter 5 allows discussion of the most important results and patterns as well as
correlations between the skeletal analysis and ethological literature review.
Chapter 5 concludes with future directions for research in biogenic factors in
taphonomy, carnivore management, and future interdisciplinary approaches to
current environmental problems.
14
CHAPTER 2: FOUNDATIONS FOR RESEARCH
This chapter presents information collected on carnivore modification in
zooarchaeology, presence of non-human scavengers in Great Plains
archaeological sites, and non-human scavenger ethological literature. First, it is
essential to discuss where and how the understanding of carnivore modification
on faunal remains emerged in the long history of archaeology and
zooarchaeology. Next a discussion on the use of the FAUNMAP database assists
in identifying non-human scavenger remains in Holocene archaeological and
paleontological sites. Finally, a review of the ethological literature on the species
of interest Canis lupus, Canis latrans, Ursus arctos, and Ursus americanus is
accomplished. This section serves as both part of the literature review and part of
the results section, as a means to illustrate animal behaviors that may have
contributed to the taphonomy of faunal assemblages. This chapter ends with a
discussion on conservation issues in regards to wolves, coyotes, and bears. In
view of the long relationship humans and these scavengers have had, it is
important to discuss how zooarchaeological research can contribute to long term
management decisions. Discussion of conservation can open interdisciplinary
15
communications that can influence an understanding of how humans have an
impact on their environment.
Methodological Changes to Understanding Biogenic Factors
In the earliest archaeological research, faunal remains were typically
thrown to the side and in some early cases recorded in lists (Reitz and Wing
1999:16-17). The reaction to exclude faunal remains from archaeological
interpretation is in part due to the lack of understanding that humans and their
environments are related and interacting (Reitz and Wing 1999:15-16). Initially,
archaeologists did not believe any data or significant information could be
gathered from faunal remains, and then gradually, archaeologists began
recording what species the faunal materials exhibited and counting how many
individuals were present (Reitz and Wing 1999:15-16). Over time, archaeologists
began interpreting damage to faunal remains as bone tools, and in several cases
the modification was interpreted as indicating tool construction (Binford 1981;
Brain 1981; Dart 1953; Dart 1956; Dart 1971). An example of the most famous
misinformed assumption that bones were altered to be tools or weapons was
established by Dart and his Osteodontokeratic Culture (Binford 1981; Brain 1981;
Dart 1953, 1956; Dart and Wolberg 1971). Dart believed that in the South African
caves, the bone accumulations were indications of a violent past in human
culture and he suggested that the breaks and crenellations exhibited in the
16
skeletal elements were created by early man (Binford 1981; Brain 1981; Dart 1953;
Dart 1956; Dart 1958; Dart 1971).
Brain (1981) took a different approach to the evidence proposed by Dart;
he studied the behaviors of African carnivores and the geology of caves and
discovered a drastically different scenario. After observing wild dogs, he found
that the types of modification Dart had been recording on bones was very similar
to the types of gnawing and crushing marks created by various carnivores
devouring a carcass (Brain 1981:22-24).
In the Great Plains, a similar situation occurred. Bison kill sites in various
settings were being excavated and skeletal elements were being analyzed. At the
Glenrock Buffalo Jump site, Frison (1970a:26-33) suggested that bone tools were
constructed as “expediency tools” and used to skin and remove meat from the
bison. At the Casper site, Frison (1974:28-31) again suggested the use of skeletal
elements as choppers and hide scrapers. Similar to Brain, Binford (1981) was
working on understanding the different processes possible to change the
dynamics of an archaeological site. In the seminal publication, “Bones: Ancient
Men and Modern Myths”, Binford illustrates the differences between bone tools
and carnivore modification of elements (Binford 1981). Moreover, he
demonstrates, with extensive actualistic research the differences between animal
and human modification to skeletal elements (Binford 1981). Finally, Binford
17
(1981:81) states that while humans may be the sole agent for change in stone by
making tools; a similar correlation does not exist for human impact to skeletal
material, especially considering that numerous mammals rely completely on
other animal resources for survival.
Complementary research by Haynes (1980a, 1980b, 1981, 1982, and 1983)
also emerged in the early 1980s. Instead of Binford’s system to understand the
types of specific marks produced by scavenging predators, Haynes (1981 and
1982) sought out to understand how entire carcasses were utilized. Instead of
coding the types of modification present on the skeletal elements, Haynes
(1982:275) recorded the amount of destruction to the elements and compared this
to actualistic research of scavenging behaviors. Combining the methods
presented by Binford and Haynes, such as: ethological literature, actualistic
research, and skeletal material properties, changes have been made to how
archaeologists understand the influence of biogenic factors in site formation
processes.
Researchers in current carnivore modification studies have begun to ask
how one could identify the specific predator that left the marks on the remains
found at hominid sites in Africa (Coard 2007; Dominguez-Rodrigo and Piqueras
2003; Pickering, et al. 2004; Selvaggio and Wilder 2001). Arguments could be
made that these researchers cannot discover the specific species consuming
18
carcass materials from faunal remains. It is difficult to assess specific carnivore
modifying assemblage by tooth marks because of the nature of carnivore
scavenging, which has been shown to be multiple marks overlapping one
another. Further it can be argued that they are not discussing the ethological
literature or doing ethological research as Binford (1981), Haynes (1980a, 1980b,
1982, and 1983), and Burgett (1990) had done. Trying to determine species from
tooth marks invariably led to the assessment (thus far) that the size class of
predators responsible could be discovered, however because of the processes of
gnawing and the plasticity and density of the skeletal material, accurate
measurements are not attained and comparison with the various African
carnivores can not be determined to any defining degree (Selvaggio and Wilder
2001). Selvaggio and Wilder (2001) specifically found that it is very difficult to
assess species of scavenger on skeletal elements based on size and shape of tooth
marks, specifically because skeletal element densities affect the resistance to force
of the skeletal material.
Appreciating how the methodological framework for studying faunal
remains developed over the course of archaeological history assists a researcher
in developing their own methods. Thus, illustrating what methods have worked
in the past and what needs to be changed to improve the field as a whole. Faunal
remains have not always been considered an important part of archaeological
19
assemblages, however, an increasing awareness of the dynamic nature of
archaeological sites has improved the questions asked and thus the methods
needed to resolve them. One possible method used in this research to help
decide which non-human scavenger species to focus on is the use of the
FAUNMAP database of faunal remains at archaeological and paleontological
sites across North America.
FAUNMAP: Choosing Non-Human Scavengers to Explore
This part of the project was used to achieve an understanding of the
specific species of predators represented in Bison bison archaeological
assemblages and to evaluate the overall changes, through time, space, and in
terms of population numbers between various species and bison. Collection of
data from FAUNMAP assists in the increase of information on how the Kaplan–
Hoover bonebed was used and why it is used at such a high intensity by nonhuman scavengers when compared to other Holocene Great Plains bison kills.
The FAUNMAP database was created in an effort to document the
mammalian species in paleontological and archaeological deposits in North
America during the Quaternary period (Graham, 1995). This database is not
exhaustive, but is fairly extensive on species within paleontological and
archaeological deposits. Initially the database was created to facilitate the
20
information on the evolution and movement of mammalian communities, but
can be used as an important resource for archaeologists attempting to
understand the biogenic ecology of a region, such as the locality of their specific
site (Graham and Lundelius 1995).
The FAUNMAP database was used as a method for understanding what
species could have been present during the late Holocene on the Great Plains.
From FAUNMAP information has been collected on the states of Montana,
Wyoming, and Colorado (coded as MT, WY, and CO respectively). The
information collected for each site includes: site name, number, state, county,
latitude and longitude, FAUNMAP age group (Table 2.1), species and family
(Table 2.2), minimum number of individuals (MNI), number of individual
specimens per taxon (NISP), and literature citation (to seek out any information
not provided by the FAUNMAP database). The species of interest for this
research study were Canis familiaris, Canis latrans, Canis lupus, Ursus arctos, Ursus
americanus, and Bison bison. To begin, all sites containing Bison bison in Montana,
Wyoming, and Colorado were documented, followed by information on above
species, and finally collection of the rest of the information listed above.
21
FAUNMAP Age Categories
Age Categories
FMAGE
Holocene 0-10,000 B.P.
HOLO
Early Holocene 7,500-10,500 B.P.
EHOL
Early/Middle Holocene 3,500-10,500 B.P.
EMHO
Middle Holocene 3,500-8,500 B.P.
MHOL
Middle/Late Holocene 0-8,500 B.P.
LMHO
Late Holocene 450-4,500 B.P.
LHOL
Late Holocene/Post-Columbian 0-4,500 B.P.
HIHO
Post-Columbian 0-550 B.P.
HIST
Table 2.1: FAUNMAP age categories used for background research.
FAUNMAP Species Codes
Species
FMSP
Canidae
CAN
Canis sp.
CA
Canis familiaris
CA fa
Canis lupus
CA lu
Canis latrans
CA la
Ursus arctos
UR ar
Ursus americanus
UR am
Bison bison
BI bi
Table 2.2: FAUNMAP species codes used for background research.
Analysis began by assessing the overall numbers of sites in each state and
within specific latitudes and longitudes. There are 30 sites located in Montana,
46 in Wyoming, and 13 in Colorado. This disparity could be accounted for by
bison populations during the middle Holocene. An increase in numbers in the
northern plains occurred because bison thrive on the abundance of C4 grasses
which are tolerant of drought and thus typical of the warm and dry
22
Hypsithermal period (8,000 – 4,000 B.P.), which began in the northern plains and
moved south through time (Kay 1998:25-27). This was then overlapped and
followed by the Neoglacial which began around 4,500 B.P. and lasted into
present times in the northern plains. The Neoglacial period contrastingly was
associated with higher precipitation and thus expansion of boreal forests, this
could have also increased bison population numbers and bison kills by
providing more precipitation to grasses, further this could have also expanded
Ursidae habitats (Kay 1998:27-28). Interestingly, Montana and Wyoming (Figure
2.1) have both grizzly bears and black bears present in faunal assemblages,
whereas Colorado (Figure 2.1) does not, and may have been too warm and dry to
accommodate the bears. Finally, these relationships correspond with the number
of sites containing non-human scavengers in the late Holocene.
The numbers of predators in archaeological assemblages increased in the
late Holocene (Figure 2.2), from 7% to 45%! From the time when the Neoglacial
began about 4,500 B.P. at the beginning of the FAUNMAP age group for the late
Holocene, it would be assumed that this drastic increase in predator
representation was related to moisture in the northern plains. Increasing
precipitation increases boreal forests, grasses, and other botanical species, thus
increasing amount of resources for a multitude of mammalian and ornithologic
species as well. It should be assumed that Canis familiaris, Canis latrans, Canis
23
lupus, Ursus arctos, and Ursus americanus were not solely sustaining themselves
on Bison bison. Therefore, the increase in numbers could have come in part on
the increase in the family Lagomorpha or the variety of berries whose population
numbers could have increased as well.
8
Number of Sites
7
6
Canis familiaris
5
Canis latrans
4
Canis lupus
3
Ursus arctos
2
Ursus americanus
1
0
Montana
Wyoming
Colorado
State
Figure 2.1: Number of sites with specific non-human scavengers in Montana, Wyoming, and
Colorado from the FAUNMAP database.
Transitioning from the late Holocene to the Post-Columbian is an
interesting shift as well. Again, 45% of late Holocene sites contain predator
remains, whereas the late Holocene/Post-Columbian switch has 23% and then
decreases again in the Post-Columbian period to 12% (Figure 2.2). EuroAmerican settlement of the Great Plains, decimation of bison populations, and
human death caused by disease, all of which limited bison kills, may have
impacted the number of bison kill sites on the Great Plains. Therefore, the
numbers of predators represented in bison faunal assemblages would have been
24
impacted. Furthermore, as bison began to disappear, so too would have the
predators that likely used them. This could have occurred by population
movements to the east, west, or south and by reduction in number of offspring
produced.
In relation to the paleoclimatic fluctuations, there are distinct differences
in which species live in which regions. Ursidae and Canidae can live in a wide
range of environments (Fitzgerald, et al. 1994). Ursidae, are typically scavengers,
and scavenge in the spring months after their winter lethargy period has ended
(Green, et al. 1997:1040; Mattson 1997:165). If these bears were scavenging
human produced carrion, then it could be assumed that human-predator
interactions would have occurred and possibly led to domestication.
Canids account for 97% of predator remains in the sites in Montana,
Wyoming, and Colorado. To what extent did wolves and coyotes follow humans
for sustenance? Later in this chapter a discussion on coyote behaviors indicates
that coyotes follow wolves to kills and wait to scavenge the remains. It has been
suggested that dog domestication and or hybrid dog remains are present as early
as 6,500 B.P. at the Hawken Site in Wyoming and 4,300 B.P. at the Dead Indian
Creek site in Wyoming (Walker and Frison 1982:127-128). Given that dog
domestication occurred at this time it is plausible that the increase in canid
remains in the late Holocene could be related to their domestication by humans.
25
Figure 2.2: Percentage of non-human scavengers present in sites within FAUNMAP age groups.
Use of the FAUNMAP database is important for recognizing patterns in
the archaeological and paleontological record of the Great Plains. It is evident
now from FAUNMAP research that canids are slightly more represented in
Holocene archaeological sites on the Great Plains. Additionally, it is evident that
more bison kill sites occur in Montana and Wyoming. Finally, there are more
non-human scavengers present in Montana and Wyoming. This information has
placed the Kaplan–Hoover bonebed in the context of other sites within the Great
Plains, within the Holocene, within bison sites, and within mammalian
population fluctuations. Besides placing Kaplan–Hoover in context, it is
important for questioning why this site is heavily modified and why other sites
may not be. Therefore, a discussion of non-human scavenger behaviors is
essential for understanding how scavengers used the bonebed.
26
Ethological Research: Understanding Scavenging Behaviors
There are thousands of books, articles, conference proceedings, and
reports published on the behaviors of carnivore and omnivore scavengers in the
Great Plains, North America, and the world. Because of this, the research
presented in this section is not conclusive or exhaustive. This research
background gives a brief description of Canis lupus, Canis latrans, Ursus arctos,
and Ursus americanus behaviors and feeding habits to inform further questions
about interactions with bison kill remains from prehistory. Research on wolves
and grizzly bears is very expansive; including numerous books and articles
whereas research on coyotes and black bears is slightly limited. For this project,
future research could include behaviors of other mammal scavengers that utilize
faunal remains.
Canis lupus: Wolves
Wolves were at one point common all over the United States, which
indicates that they could have easily been located in various places on the Great
Plains. With the introduction of domesticated cattle, however, the species has
been eradicated from most states for livestock slaughter (Fitzgerald, et al.
1994:307). Canis lupus can occupy a wide range of environments, including high
altitudes and the species typically lives in regions where there are high
populations of large bodied ungulate species (Fitzgerald, et al. 1994:306).
27
Documentation of kills demonstrates that the gray wolf mostly predates elk,
mule deer, bison, and mountain sheep and in some instances beavers (Fitzgerald,
et al. 1994:306; Mech 1970).
Anatomically, canids (Figure 2.3) are designed to make numerous shallow
bites while attacking to take down its prey, whereas a felid can take one deeper
bite and hold on to take their prey down (Peterson and Ciucci 2003:112-113).
Wolves having heterodont sharp high cusped teeth are able to break and chew
through a variety of gross materials, the molars are used to rip and shred meat,
while the canines and premolars are used to crush and crack bone material
(Peterson and Ciucci 2003). In addition to the dental specialization for meat
consumption, the mandibular structure of a wolf is robust with several massive
muscles that act in unison to close the jaw rapidly and efficiently (Peterson and
Ciucci 2003). Further adaptation for the lifestyle of a carnivore is the carnassial
pair (Figure 2.4), which are the upper fourth molar and the lower first molar that
act together as scissors for shearing and slicing through hide materials (Peterson
and Ciucci 2003).
28
Figure 2.3: Canis lupus maxilla (Myers, et al. 2008).
Wolves can swallow pieces of carcasses whole; in one study two whole
caribou tongues were found in a wolf stomach (Mech 1970:169-170). Wolf
stomach contents have been known to contain anywhere from 2.40 to 5.98
kilograms of meat, bone and hair, with the largest known amount being close to
8.62 kilograms (Mech 1970:170-171). Typically wolves begin feeding on the
internal organs after tearing open the carcass, then move to hind limbs and other
parts, while avoiding the entrails and stomach contents (Mech 1970; Peterson
and Ciucci 2003). Wolves will feed until their stomachs are full and their sides
distended and in the winter, after feeding, wolves will collapse and sleep up to 5
hours, which aids in digestion after gorging (Peterson and Ciucci 2003:124). In
addition, wolves rely heavily on skeletal materials from kills, often scavenging
on them to sustain their mineral nutrient intake (Peterson and Ciucci 2003:125).
29
Further, single wolves that have not had access to a fresh kill can sustain
themselves for long periods of time on only skeletal material and then may be
willing to eat any part of the gross tissue (Peterson and Ciucci 2003:125).
Figure 2.4: Carnassial pair of Canis lupus (Myers, et al. 2008).
Gray wolves have been recorded scavenging significantly more bison
carcasses as opposed to red deer and wild boars in Bialowieza Primeval Forest in
Poland (Selva, et al. 2005:1593). Interestingly, of all the predators monitored in
Bialowieza Primeval Forest (including: ravens, buzzards, eagles, red foxes,
raccoon dogs, pine martens, and domestic dogs), gray wolves were the only
species able to open dead bison carcasses (Selva, et al. 2005). In the winter
season, Huggard (1993) noted that wolves scavenged in shallower snow than
deep snow and would hunt more often if the snow was at great depths. Wolves
are even known to cache carcass remains for later use, specifically in the summer
to keep flies away and preserve the meat out of the summer heat (Mech 1970;
30
Peterson and Ciucci 2003). Further this caching can be done to save food for later
after the wolf is satiated, and typically wolves will take their food for caching
long distances (up to 5 km) from the kill to avoid theft by other scavengers
(Peterson and Ciucci 2003:117).
Canis latrans: Coyotes
Canis latrans (Figure 2.5) is still considered common all across Colorado
and is easily adapted to all elevations (Fitzgerald, et al. 1994:302). In addition,
the species is well adapted to living amongst human populations and may have
thrived after the eradication of the gray wolf (Fitzgerald, et al. 1994:302).
Typically, Canis latrans is an opportunistic feeder, often preferring animal meat,
but also consuming vegetation and insects in some instances (Fitzgerald, et al.
1994:303). In most situations coyotes will eat jackrabbits, cottontail rabbits, and
rodents, likewise; they are known to scavenge carrion of cattle and big game
which has been killed by other larger predators and do often themselves kill
livestock such as sheep and goats (Fitzgerald, et al. 1994; Hilton, 2001; Kleiman,
2001; Paquet, 1992). Coyotes can and will alter their natural diet to exploit foods
introduced by humans and other scavengers to lessen the amount of energy
needed to gain food (Kleiman and Brady 2001). Coyote stomachs typically
contain large quantities are scavenged foods (indicated by large quantities of
31
maggots) as well, suggesting that in the wild they are not necessary hunters, but
scavengers (Kleiman and Brady 2001:168-169).
Figure 2.5: Canis latrans maxilla (Myers, et al. 2008).
Canis latrans is known for defecating on their kills and carrion to mark
their property and deter other animals from consuming it (Acorn and Dorrance
1990; Acorn and Dorrance 1998; Wade and Bowns 1985). When hunting, coyotes
attack the neck/throat first of sheep and goats and attack the hind limbs in calves
of other ungulates (Wade and Bowns 1985). They primarily begin feeding in the
hind limbs or just below the ribs and choose viscera first when consuming
carcasses (Acorn and Dorrance 1990; Acorn and Dorrance 1998; Wade and Bowns
1985). In almost every case of carrion scavenging, however, the coyote will either
follow gray wolves or scavenge gray wolf kills (Paquet 1992). In some instances
it is difficult to assess the amount of material in a coyote stomach as carrion or
32
hunted; however, the presence of maggots and fly larvae have been used as a
means to determine if the coyote hunted its meal (Kleiman and Brady 2001:168).
Comparing Wolves and Coyotes
A research study at Riding Mountain National Park in Manitoba,
documented 194 ungulate wolf kills from July 1982 through March 1986 (Paquet
1992:338). Within this same time frame Canis latrans killed 59 ungulates and
were documented to visit every wolf kill (Paquet 1992:338). On average, large
gray wolf packs consume more of the killed carcasses than do coyotes; this is
likely due to size and energy expenditure needed by each non-human scavenger
(Hilton 2001; Paquet 1992). During this research study Paquet (1992:341) found
that all wolf-killed ungulates remains (n = 194) were scavenged by coyotes and
this was evident by coyote tracks to every wolf-kill and skeletal disarticulation
and hide removal of carcasses. Moreover, Paquet (1992:341) observed coyotes
waiting 100 m from a fresh wolf kill and noticed that the coyotes moved in
quickly to consume the remains immediately following wolf departure. This of
course led to the demise of some coyotes that were impatient; however, this
danger did not deter them (Paquet, 1992; Wilmers, 2004).
Anatomically, wolves and coyotes are similar; however, the coyote is
constrained by being significantly less powerful than the wolf and much smaller
in size (Hilton 2001). In terms of scat, wolf and coyote scats are similar in
33
appearance, contents, and can overlap in size; however, coyote scat rarely
exceeds three centimeters in diameter, while wolf scats can exceed three
centimeters and typically go beyond or up to 4 centimeters in diameter (Mech
1970). In appearance, wolf and coyote scat is arranged with the skeletal
fragments in the center while hide and hair are wrapped around the outside,
therefore protecting the intestines (Mech 1970). Unlike wolves, coyotes scavenge
and typically hunt alone, allowing more time to be dedicated to following other
hunters (Kleiman and Brady 2001).
Ursus arctos: Grizzly Bears
Ursus arctos (Figure 2.6) is known to live in a wide variety of environments
from plains grasslands to alpine tundra and is most content in a habitat of
seasonally changing food stuffs (Fitzgerald, et al. 1994; Servheen 1999). Use of
FAUNMAP to get a rough estimate of where grizzly bears were on the Great
Plains during the Holocene indicates that they were present at various
archaeological sites during all geological time periods, including Colorado
during the early and middle Holocene. Predominately, Ursus arctos consumes
vegetation; nevertheless, the species is known for scavenging carrion (especially
in the spring), and killing small mammals such as marmots, large mammals such
as elk and other ungulates, and livestock such as cattle and sheep (Craighead, et
al. 1995; Fitzgerald, et al. 1994; Wade and Bowns 1985).
34
Figure 2.6: Ursus arctos maxilla (Myers, et al. 2008).
These mammals are incredibly efficient as omnivores, and are fairly
inefficient as carnivores, so scavenging is the main way for them to get meat
protein (Craighead and Craighead 1972:304). Winter-killed ungulate species are
of importance to grizzly bears, specifically in the spring after they have
awakened from winter lethargy; however, carrion feeding is at its peak from
March through May (Craighead, et al. 1995; Green, et al. 1997; Mattson 1997). In
Yellowstone National Park, grizzly bears consume mostly elk, bison, and moose
meat (Craighead, et al. 1995; Mattson 1997). Carrion availability can drastically
affect these behaviors by increasing the amount of time bears use carrion
(Craighead, et al. 1995:258-260). Mattson (1997:171-172), states that the frequency
in which grizzly bears used ungulate carcasses, varied during months, years, and
regions of the park. In addition, use of ungulates was related to availability of
whitebark pine seeds (Mattson 1997:169). However, if large numbers of
carcasses exist in a grizzly bear’s territory, they will forego eating other foods
35
and just sustain themselves on carrion. Bears typically do not feed in groups or
at the site of the kill and typically remove pieces of carcasses and retreat to
concealed areas for feeding purposes (Craighead, et al. 1995). Bears are also
known for caching carcasses for future use and will return to feed numerous
times if the carcass is not located by other scavengers (Craighead, et al. 1995:259).
Moreover, Green, et al. (1997) discovered that date of death was less important in
determining if a bear would scavenger a carcass if the death occurred between
February and early March and more important between middle March to late
April. This could be due to increase in temperature and rate of decomposition of
the carcasses. Green, et al. (1997:1047) further reported that Ursus arctos typically
scavenges more Bison bison than Cervus elaphus in Yellowstone National Park.
Ursus arctos exploits carrion more frequently in higher altitudes in Yellowstone
National Park than lower altitudes during the late spring (Green, et al.
1997:1048).
Ursus arctos is an animal that, as argued by Craighead, et al. (1995) and
Craighead and Craighead (1971), is conditioned by humans for food resources.
There is a large body of literature on grizzly bears feeding in garbage dumps,
campgrounds, and boneyards (cattle carcass piles) in Montana and Yellowstone
National Park (Craighead and Craighead 1971; Craighead, et al. 1995; Rogers
1987; Wilson et al. 2005). After consumption of ungulate species in the early
36
spring, snow decreases and tourist activity increases allowing grizzlies to stock
up on more calorie rich foods from garbage dumps, which are particularly
important to the feeding habits of Yellowstone National Park bears (Craighead,
et al. 1995:44-47). Garbage dump feeding does not have a specific seasonal time
period however, as bears use this resource continually throughout their nonlethargy season as a consistent and stable food supply (Craighead, et al.
1995:270).
A significant research study from 1977 to 1987 suggested that grizzly
bears on the east front of the continental divide of Montana were frequenting
rancher boneyards as a secondary source of all protein (Craighead, et al. 1995;
Wilson, et al. 2005; Wilson et al. 2006). These sites are typically frequented
during the spring when bears need to gain calories and protein quickly after
winter lethargy (Mace et al. 1987; Wilson et al. 2006).
According to Craighead and Craighead (1972), grizzlies who feed at
garbage dumps, campgrounds, and boneyards exhibit less fear of humans and
human smells; however, in other areas of national parks and human landscapes
they are fearful and tend to avoid contact with humans. Within Yellowstone
National Parks campgrounds, grizzlies return for garbage foraging during spring
and fall migratory movements to gain access to another food source with limited
energy expenditures and the animals that frequent campgrounds on a regular
37
basis become conditioned to human presence, a difference from grizzlies that
frequent isolated garbage dumps where humans move into and out of the area
on a more predictable regular basis (Craighead and Craighead 1972:309-311).
The conditioned behaviors exhibited by grizzly bears could be attributed
interactions with humans (Craighead and Craighead 1972). Given that these
interactions are occurring more frequently, due to human encroachment on
natural habitats, grizzly bears will become acclimated to human scents and
environments (Craighead and Craighead 1972). Therefore, bears become less
fearful, further causing more conflicts in homes, cars, and camp sites (Craighead
and Craighead 1972).
Understanding feeding ecocenters as defined by Craighead, et al. (1995)
assists in the general knowledge of how bears feed and the patterns that may be
observed from their feeding behaviors. Craighead, et al. (1995:321), define bears
as an ecocentered population “that congregates at a high-quality food source in a
relatively confined, predictable portion of an entire ecosystem during an
extensive, annually predictable time period.” In view of the fact that bears have
a regimented feeding pattern based on availability of food stuffs, it would be
expected that bears could have evolved behaviors that rely heavily on this
feeding style to ensure existence.
38
Finally, boneyards as feeding ecocenters are important in terms of where
bear behaviors have been constant over time. Across the Great Plains and
Western United States, cattle ranchers have sometimes used boneyards or cattle
carcass dumps when their livestock have died (Craighead, et al. 1995; Mace et al.
1987; Wilson et al. 2005; Wilson; et al. 2006). Craighead et al. (1995:324–326)
compare boneyards to bison kill sites, suggesting that these were predictable and
stable events across the Great Plains, allowing bears to be conditioned further by
humans by expecting large quantities of carrion to be available throughout the
year. As stated previously, Craighead, et al. (1995:322-326) indicate that this
phenomenon of using feeding ecocenters is a biological phenomenon that has
existed in bears not necessary because of a reliance on humans, but possibly
because of the environment bears have been a part of since human hunters began
hunting large numbers of bison in the Great Plains. In the present time grizzly
bears have been shut out of garbage dumps thus increasing human-bear
conflicts.
Ursus americanus: Black Bears
Ursus americanus (Figure 2.7) is arboreal by nature and is therefore
commonly found in montane shrublands and forests and subalpine forests at
moderate elevations (Fitzgerald et al. 1994:318). Primarily, black bears are
vegetarians; however, they have been known to consume carrion and will kill elk
39
calves and other wild ungulate calves, sheep, goats, and pigs (Fitzgerald et al.
1994; Wade and Bowns 1985).
Ursus americanus commonly predates in the spring and summer (Wade
and Bowns 1985). Green et al. (1997), discuss Ursus americanus use of lower
altitude carrion as opposed to Ursus arctos. Additionally, the black bear is more
likely to use carcasses during the late spring than the early spring (Green, et al.
1997:1052). In the attack, black bears typically use their paws and break the back
or neck with strong blows, eventually killing the prey by biting the neck and
shoulders (Wade and Bowns 1985). Black bears are inclined to drag their food to
a secluded area for feeding; however they will defecate on the carrion as Canis
latrans to discourage other non-human scavengers from consuming it (Wade and
Bowns 1985:10). Finally, both bears do not scatter, chew, and break up carcasses,
which is typical canid behavior (Wade and Bowns 1985).
Figure 2.7: Ursus americanus maxilla (Myer et al. 2008).
40
Non-human scavenger behaviors vary between species. When discussing
canids and ursids it is important to note that they are very distinct in terms of
type of dentition, scavenging styles and patterns, and various other behaviors.
Differences in dentition are due in large part to the type of carnivore or omnivore
it is. Canids are generally hunters therefore sharp cusped teeth are necessary to
rip flesh and muscle, whereas ursids are typically omnivores and need rounded
low cusped teeth useful for eating a larger variety of food stuffs.
There are great differences in terms of how canids and ursids scavenge.
First of all canids typically scavenge and hunt in groups, therefore when there
are a number of individuals together at one carcass, fighting for food increases
the tearing, dragging, and movement of a carcass. Bears on the other hand are
solitary or with cubs, therefore individuals generally take a limb or piece of the
carcass and drag it away to a concealed location for feeding.
Conservation Research
Zooarchaeological research in the past and present is addressing the use
of faunal remains associated with human behaviors by non-human scavengers
(Reitz and Wing 1999:135). Given that research on carnivore modification lends
information to how a faunal assemblage was altered through time, this research
can be useful when applied to management decisions in animal conservation.
41
Managers deciding whether to reintroduce or eradicate a particular species can
benefit from understanding the expansive time period in which a species has coexisted with others. As stated previously, archaeologists are in a prime position
to influence management decisions by sharing their data with the natural
sciences to assist in understanding the impacts of species management changes.
The following discussion highlights the history of eradication and specific
conservation issues currently for wolves, coyotes, grizzly bears, and black bears.
Canis lupus and Canis latrans: Conservation Issues
Wolves and coyotes have always been considered a nuisance by ranchers
and hunters (Clark and Rutherford 2005; Fritts et al. 2003; Smith, Brewster, and
Bangs 1999; Wilmot and Clark 2005). Public interest is varied, with some people
believing that wolves and coyotes destroy their products (livestock) and profits
while others believe that they should be left alone, to allow them to live naturally
in the wilderness (Clark and Rutherford 2005). On another side, hunters argue
that introduction or reintroduction of non-human predators significantly reduces
population numbers for wild game hunting and eliminates the traditional
heritage of American settlers (Wilmot and Clark 2005).
While settling the west, ranchers would kill wolves and coyotes since they
were seen as decimating their livestock and adding competition to hunting wild
game (Smith, et al. 1999:108-109). After prey species populations of the wolf,
42
such as bison, deer, elk, and pronghorn declined in the west, wolves began to
feed on livestock provided by the ranchers (Young 1946). The rough lifestyle of
western settlers and the depredation of their livestock caused animosity towards
the wolves, which in a short period of time led to the eradication of wolves
(Young 1946). Wolves were almost completely eradicated, with few singles and
pairs from the lower United States, except for northern parts of Minnesota by the
1930s (Smith, et al. 1999:108). In 1974, wolves were listed on the Endangered
Species Act and mandated to be reintroduced to Yellowstone National Park and
not until 1995 and 1996 were 31 individuals reintroduced (Smith, et al. 1999;
Smith, et al. 2003). After a year, 44 adult wolves and an unknown number of
litters were present in and around the park (Smith, et al. 1999). Prior to the
count, 26 had been killed from human caused deaths such as illegally or legally
being shot and hit by cars (Smith, et al. 1999). To this day, humans are the largest
cause in wolf mortality, and influence the behaviors and predatory ecology of the
species (Fritts et al. 2003). Finally, this reintroduction has changed the numbers
of coyotes drastically and changed their breeding and hunting behaviors as well
(Smith, et al. 1999).
The importance of wolves in ecosystems cannot be overstated. The
species is responsible for much of the available carcasses for scavenging species
(Mech 1970). In addition, wolves are imperative to some ungulate species
43
population control; without predator-prey relationships, some species such as
elk become overpopulated, which will lead to starving and more accidents with
cars. A research project on the influence of reintroduction of wolves to Cervus
elaphus (elk) and Bison bison suggests that the elk population suffered a shift in
diet quality to low, while bison remained stable with females and calves
increasing vigilance (Hernandez and Laundre 2005; Laundre, et al. 2001).
Coyotes have been seen in a similar light throughout western settlement
and are still seen as a problematic nuisance with ranchers today (Beckoff and
Gese 2003). Livestock predation amongst coyotes is a contentious issue, with a
disparity between human belief that coyotes kill livestock or leave livestock
alone (Beckoff and Gese 2003:475). As the wolf’s habitat has decreased and
human eradication of them has occurred, coyotes have been able expand their
habitat ranges (Nowak 2001). In addition, coyotes are very efficient at adapting
to human environments such as neighborhoods, towns, and larger cities by
exploiting garbage, livestock, and pets (Beckoff and Gese 2003; Nowak 2001).
For the sheep industry, stockmen have stated that coyote depredation is the most
problematic cause of profit loss (Beckoff and Gese 2003; Sterner and Shumake
2001). Coyote population control and determent from sheep and livestock
depredation has been done using a number of non-lethal methods, including:
exclusion fences, aversive agents, and chemosterilants (Acorn and Dorrance
44
1998; Acorn and Dorrance 1990; Beckoff and Gese 2003; Sterner and Shumake
2001). Of the methods to deter coyote depredation, aversive agents such as
olfactory and gustatory products have done little to decrease predation (Beckoff
and Gese 2003). In recent years, other methods, such as sheep collars that release
toxic chemicals and trapping have been somewhat productive (Beckoff and Gese
2003). Interestingly, with all of the control methods and killing of coyotes by
humans, population numbers of coyotes have remained stable (Beckoff and Gese
2003).
Ursus arctos and Ursus americanus: Conservation Issues
Human relationships with bears are a highly contentious issue in wildlife
conservation (Craighead, et al. 1995; Gilbert 1989). Bears are typically
omnivores, and easily adapt to change. They are able to adapt so well to human
surroundings, that people have been accidentally conditioning bears for a long
time (Gilbert 1989). From the opening of Yellowstone National Park, humans in
the west have been feeding and taking care of bears, to which bears have
responded by being less fearful of humans (Craighead, et al. 1995; Gilbert 1989).
Grizzly bear populations today in the lower 48 states are located in
Greater Yellowstone Ecosystem, with another smaller population located in the
northern Cascades, with approximately 1000 bears between the two (Schwartz, et
al. 2003:558). From the 1930s through the 1960s, visitors of the Park would feed
45
grizzlies and did not anticipate that building a reliance and acceptance of human
presence could influence future problems, such as personal injuries to humans
and property damage (Craighead, et al. 1995:24). Early attempts to educate the
public on wildlife included lecture series, where park rangers would place food
wastes in a central location and then visitors would sit on bleachers watching the
bears feed and hear about their behavior and ecology (Craighead, et al. 1995:4447). The last of the lecture series food waste sites were closed by the mid-1940s
in a hope to decrease human-bear interactions; however, bears did not stop
feeding in these locations or other established tourist dumps and may have
become accustomed to including garbage into their seasonal feeding habits at
this point (Craighead, et al. 1995:270). These human induced bear interactions
led researchers to believe that the major causes of bear mortality had less to do
with habitat and more to do with their relationships with humans (Gilbert 1989).
Mortality rates in grizzly bear populations are caused mostly by human impacts
to populations through hunting, poaching, and habitat loss (Servheen 1999;
Schwartz, et al. 2003). Cause for eradication of grizzly bears was due to fear of
attack from them or destruction of campsites and livestock (Craighead, et al.
1995; Primm and Murray 2005).
Human-bear interactions are among the most important to consider when
discussing conservation, specifically for the grizzly bear, which has been
46
conditioned to trust and in some instances rely on humans to provide food
(Gilbert 1989). Researchers believe that in parks, like Yellowstone National Park,
when humans began feeding bears or when lectures were centered on bear
feeding, that bears became conditioned or accepting of human smells and
therefore less fearful (Craighead, et al. 1995; Gilbert 1989). Beginning in 1968,
Yellowstone National Park personnel began reducing the amount of garbage
held in the remaining Park tourist dumps and by 1971 closed the rest of the
dumps (Primm and Murray 2005). Other researchers believed that the dumps
should not have been closed instantaneously and that the bears should have been
weaned off of human garbage as sustenance (Craighead, et al. 1995; Primm and
Murray 2005).
After the closure of the dumps, there was a dramatic decline in the
number of grizzly bears in the park. With the constant source of garbage
unavailable, the bears began going into camps, livestock areas, and surrounding
community garbage dumps outside of Yellowstone National Park, leading to
more human-bear conflicts, and more deaths caused by human shootings
(Primm and Murray 2005). After this, researchers and public relationships
declined dramatically. Finally, in 1975 the grizzly bear was declared threatened
under the Endangered Species Act (Primm and Murray 2005). For the next
decade, grizzly bear populations declined because of the force to learn to find
47
natural foods in the environment, thus causing fecundity issues, given that they
were malnourished (Primm and Murray 2005). Today, grizzly bears have made
a comeback and have become accustomed to feeding in natural environments;
however, because of the two-way movement of humans into bear habitats and
bears into human habitats, more interactions have led to increased animosity
between ranchers, surrounding communities, and the national park systems
(Primm and Murray 2005).
Black bears are a wide ranging species capable of adapting to human
presence; therefore, causes of their mortality are human hunting, poaching, and
roads kills (Pelton et al. 1999; Pelton 2003; Rogers 1989). Similar to grizzly bears,
black bears have a tendency to feed on human garbage in and around parks
(Fitzgerald et al. 1994; Green, Mattson, and Peek 1997; Wade and Bowns 1985).
Typically, black bears will use garbage dumps when there is a reduction in their
habitat and, therefore, natural food range (Rogers 1989). Human and black bear
interactions are more frequent than grizzly bear interactions at dumps and this is
primarily due to the fact that black bears simply ignore human presence at the
dumps and continue on with what they are doing (Rogers 1989). In the event
that people begin to throw food or rocks at the bears feeding, they will either
ignore or move away, rarely if ever defending their food (Rogers 1989). Most of
the injuries incurred by black bears have been a reaction to a person handing a
48
bear some food (Rogers 1989). In Wisconsin, complaints of bear-related
destruction were the main reason for bear control initiatives (Rogers 1989). To
curb bear damage, trapping, shooting, and similar to coyotes, chemical aversion
techniques, have been employed (Pelton 2003; Rogers 1989).
When it comes to carnivores, both canids and ursids, it is important to
realize that humans influence their behaviors (Beckoff 2001:xix). This idea is
controversial and conservation of carnivores is emotionally intense for a large
number people, leading to great debates (Beckoff 2001:xix). To better the
dialogue on carnivore management, understanding the whole story including
the length of time humans and carnivores have been interacting, the role that
these non-human scavengers play in the biodiversity, stability, and integrity of a
multitude of other mammal species and community constructions is extremely
relevant (Beckoff 2001:xvii-xix). To encourage this dialogue, it is becoming
increasingly necessary for conservationists and government agencies to gather all
of the information they can, instigating an interdisciplinary approach to
understanding carnivore management. Zooarchaeological studies have a
potentially central role to play in elucidating these long-term human/carnivore
interactions. Beginning with the earliest settlers of the world, humans have been
interacting with their environments, for better or for worse; changes have been
made (Lyman and Cannon 2004).
49
As presented in the preceding paragraphs, humans impact other
mammals in a variety of ways. Since these impacts began in North America over
13,000 years ago, it is important to review archaeological sites for an improved
understanding of our previous interactions with non-human scavengers to
become better prepared to make conservation decisions (Lyman and Cannon
2004). Finally, if managers and government agencies have the information
archaeologists can provide of the past and our understandings of taphonomy,
species distributions in archaeological assemblages, and manipulations humans
have made to their environments, a more informed decision can be made about
how to sustain non-human scavenger populations. Further discussion of this
topic will follow in Chapter 5 to illustrate how the data provided by the Kaplan–
Hoover bison bonebed could be relevant to conservation issues on the Great
Plains.
Summary of Chapter
This chapter presented multiple areas of literature from various
disciplines. First, a methodological background for how questions were asked
and what prior researchers influenced the methods chosen to analyze the
Kaplan–Hoover collection was undertaken. From the methodological
framework it is shown that carnivore modification research has undergone a
50
number of changes and will continue to change as the archaeologist’s integration
of literature from the natural sciences is incorporated into archaeological
research. Further, the integration of natural science research and methods can
open up a dialogue between the disciplines to enhance management decisions in
the future. Second, a sample of the literature on archaeological sites from
FAUNMAP in the states of Montana, Wyoming, and Colorado discussing
presence of bison and one of the four main non-human scavengers was
presented. After reviewing FAUNMAP, an understanding of the relationships
between bison and these animals in the archaeological literature was
accomplished. This illustrated important patterns in the archaeological record on
the Great Plains. Some of those important patterns include: increase in bison kill
sites as one goes north from Colorado to Montana, increase in diversity of nonhuman scavengers in bison kill sites as one goes north, and the presence of more
canids in sites containing bison faunal remains than bears. Finally, this chapter
ends with a review of Canis lupus, Canis latrans, Ursus arctos, and Ursus
americanus behavior in feeding events and conservation issues relevant to these
species and the role of zooarchaeological analyses in contributing to these
conservation related topics. This is useful in identifying the differences between
canids and ursids and the minor differences between wolves and coyotes as well
as grizzlies and black bears. After assessing the literature background
51
information it is important to identify specific methods used to collect data from
the Kaplan–Hoover bonebed.
52
CHAPTER 3: AN INTERDISCIPLINARY
APPROACH TO METHODS
Zooarchaeological research is the result of the cultural ecology movement
of the mid-twentieth century established by Julian Steward (Krause 1998; Reitz
and Wing 1999). Prior to this paradigm shift in anthropology, archaeologists
dealt with faunal remains associated with archaeological deposits differently.
Instead of thorough research and data collection, faunal remains were reported
in lists and tables in appendices and rarely discussed in any further detail (Reitz
and Wing 1999). The late twentieth century was met with the onset of functional
and processual approaches to faunal remains to which the majority of
zooarchaeological research is conducted today are still maintained (Reitz and
Wing 1999). Site formation studies in zooarchaeological research changed
temporally as well. While taphonomy was defined by Efremov (1940) in the field
of paleontology, the use of taphonomic research in archaeology was not clearly
established until the 1980s as a reaction to Binford’s concept of middle-range
research (Binford 1981). Since that time, taphonomy has become one of the most
53
important factors for understanding faunal assemblages and archaeological site
formation processes (Lyman 1994; Reitz and Wing 1999).
An interdisciplinary approach to site formational processes and
paleoecological reconstructions has become increasingly important to
archaeological research. For this project, research was undertaken using three
methodological frames of inquiry: zooarchaeological, taphonomic, and
ethological. By collecting more data from various fields of the natural and social
sciences, a researcher may become familiar with the interacting dynamic
processes that produced the bonebed. For instance, use of ethological literature
will increase the understanding of non-human scavenger behaviors that can vary
across species and would have influenced the taphonomy of the bonebed
dramatically. This chapter outlines those three approaches by illustrating
specific methods used for research. In addition, each section outlines the
importance of the method used and how it has improved the holistic nature of a
interdisciplinary approach to site formational processes and paleoecological
reconstructions.
Data Collection Procedures
For this research, the majority of the appendicular skeleton of the species
Bison bison for the Kaplan–Hoover site was analyzed: humerus, radius-ulna,
metacarpal, femur, tibia, metatarsal, astragalus, calcaneus, and third phalanx.
54
These elements were used for both practical and methodological reasons. In
terms of practicality, each of these elements has been reliably used to assess sex
in bison herds and is consistently well preserved in archaeological sites (Bedord
1974; Morlan 1991; Todd 1983, 1987; von den Driesch 1976). Second, these
particular skeletal elements have been standardized in terms of measurements
and have been discussed in non-human scavenger modification studies more
often than axial skeletal remains. Therefore alleviating the need to create
methods for determining herd demographics, instead focus was directed on
carnivore modification methodologies that have not been standardized (Bedord
1974; Haynes 1980a, 1980b, 1981, 1982, 1983, 1991; Morlan 1991; Todd 1983, 1987;
Todd, Hill, Rapson, Frison 1997; Todd and Rapson 1988, 1999; von den Driesch
1976). For statistical analysis, basic descriptive statistics, scatter-dot graphs, chisquare goodness-of-fit, and Pearson and Spearman correlations were prepared
using the SPSS program version 15.0.
Multiple-Component Coding System
The coding system (Appendix B) used for the data collection was
established by Todd (1983:324-327 and 1987:121-123) and included: element,
portion, side, sex, proximal fusion, distal fusion, and bone breakage. This system
has been used for the majority of zooarchaeological research in the Great Plains.
Systematic coding after Todd (1983 and 1987) was used as a standard means to
55
collect data used in analysis of herd demographics. These methods allow for
organization of the data in a clear manner to illustrate patterns and give a
necessary basis for statistical analysis.
The landmark coding system (Appendix C) was used to assess overall
destruction of the assemblage (Hill 1994:169). Skeletal elements with landmark
codes included: humerus, radius-ulna, femur, and tibia. This is another back-up
to the coding system that recorded portion and segment remaining in the skeletal
elements from the collection. Additionally, the landmarks were used to
understand overall non-human scavenger destruction in terms of specific skeletal
landmarks that in most cases indicate muscular attachments that could increase
understanding of non-human scavenger behaviors including meat selection
choices. Specific landmarks discussed for the humeri include: deltoid tuberosity,
tubercle for attachment of medial collateral ligament, major tuberosity, proximal
olecranon fossa, posterolateral nutrient foramen, and teres major tubercle. For
the radius-ulna, landmarks are posterolateral nutrient foramen and radial
tuberosity. On the femora the anterior nutrient foramen, supracondyloid fossa,
and the major and minor trochanter were recorded. Finally, on the tibia the
following landmarks were recorded: anterior crest, posterolateral nutrient
foramen, and anterior nutrient foramen.
56
Modifications to this coding system, however, were designed to better
represent the collection for non-human scavenger modification. Carnivore
modification (Appendix B) was coded based on multiple authors’ typological
descriptions and relevant images will be discussed later on in this chapter
(Binford 1981; Fisher 1995; Haynes 1980, 1982, 1983, 1991; Lyman 1994; Stiner
1994). These data collection methods are useful for a holistic approach to
understanding bonebed formation dynamics by increasing the available data set,
which allows a more diverse and comprehensive set of analysis from which to
work.
Todd’s coding system increases speed and efficiency of data collection
while allowing a researcher to obtain a greater breadth of data, hence increasing
the ability to ask further questions of the data if needed at a later time. The
approach of collecting more data rather than less data is not always feasible due
to time restraints and funding constrictions; however, in this instance the
collection of more data allowed for an increased learning environment as well as
a variety of empirical methods that can be used in future projects and for future
directions with this particular collection by other researchers.
Herd Characteristics: Zooarchaeological Methods
To begin it is important to define specific terms that will be used
throughout this thesis, as these terms can be defined differently by different
57
archaeologists (Lyman 1994). First of all, for the entirety of this project “skeletal
element”, “element”, and “specimen” refers to any identified or unidentified
bone or fragment of bone. This nomenclature is necessary as the Kaplan–Hoover
material used for this thesis represents both complete and fragmented bones to
varying degrees and interchanging the terminology would weigh down the
understanding of the information being presented. In instances where
understanding the intensity of the destruction is necessary, there will be
reference back to the coding system and all tables, graphs, or figures will
appropriately explain where on the skeletal element the information is being
derived.
Metric quantification for the collection was undertaken using
measurements (Appendix C) previously defined by Bedord (1974:201-203),
Morlan (1991:222-223), Todd (1987:371-388), and von den Driesch (1976:92-93).
These measurements were used to approximate sex in the collection to ascertain
if any difference was present between males and female/sub-adult usage by nonhuman scavengers. Sex determination for the assemblage was difficult to assess
very quickly and accurately due to the high degree of destruction by non-human
scavengers. The elements that were easily sexed (Appendix E) include: humerus
(M11 × M7; Todd 1987:162-165), radius-ulna (M9 × M4, M11 × M7; Todd 1987:162167), astragalus (M3 × M5; Morlan 1991:223-224), calcaneus (M4 × M4, M6 × M7;
58
Morlan 1991:223-224) and metapodials. Sex for the metapodials was
accomplished using measurements 1–4 crossed against M5 (Bedord 1974:208).
Bedord (1974:208) suggested that greatest breadth of distal end is the most
accurate measure to use for sexing the metapodials; therefore this measure was
crossed against the remaining four measures collected for the assemblage. The
reason these were relatively easy includes the frequency distribution of data
measurements available and bivariate methods used are well defined in the
literature. The remaining skeletal elements, however, were not as easy to record
sex.
Sexing the femur and tibia was complicated by the overall destruction by
non-human scavengers, making it difficult to use the measurements taken by
Todd (1983:142), and von den Driesch (1976:84-85); therefore, these elements
were sexed differently. In order to overcome the destruction obstacle,
measurements with the greatest value distributions were crossed using bivariate
scatter plots to assess sex for these elements (Appendix E). This included a
number of crosses; therefore, columns were created in the database to account for
all the crosses and following completion these were assessed to determine sex
(Appendix Disc). Furthermore, in the event that the sex estimates were even,
meaning female and male identifications were the same an assignment of “not
sexed” was used to avoid biased results. For the femur, measurement
59
distributions suggest that measurements 8, 10, and 17 are usable for sex
determination. To use these measurements they were all crossed against each
other using bivariate scatter plots (Appendix E) and the following crosses
worked out for sex determination (M17 × M10, M8 × M10, and M8 × M17). For
the sex of the tibia there were a number of measurements that were usable
including: M1, M2, M6, and M7. Measurements 9 and 10 also had large
frequency distributions in terms of measurements; however, bivariate scatter
plots were too difficult to read and therefore sex based on those measurements
was not reliable. Even given the use of M1, M2, M6, and M7, sexing of the tibia
was difficult as there were not many measurements to cross (Appendix E),
however, crosses did include: M1 × M2, M1 × M6, M1 × M7, M2 × M6, and M2 ×
M7. Finally, sex for the third phalanx was not undertaken due to the difficulty in
assessing fore from hind limb, which could have drastically affected results and
there are no confirmed means to accurately assess sex of the elements.
Quantification of the skeletal remains present is the first task of any faunal
analysis (Grayson 1984; Lyman 1994). Under the coding system described above,
Bison bison skeletal elements are quantified using a set of previously defined
general indices. First of which is number of identified specimens (NISP), where
“identified” indicates to skeletal element. NISP will be used to indicate the entire
quantity of skeletal elements used for this project, thus allowing for an
60
understanding of why results may vary due to increases or decreases to the
quantity of lines of data represented. Similarly, NISP is used to identify the total
number of skeletal elements within each specific identified element; thus
identifying sample size (Lyman 1994:100).
The second measure of importance is minimum number of individuals per
specific skeletal element (MNI), where number of individuals will be crosstabulated with sex (male or female/sub-adult) and side (left, right, not sided) to
gain the most accurate demographic description of the sample (Lyman 1994:100).
The following quantitative measures will be used for descriptive purposes
as well; however, these measures are more useful for understanding the overall
destruction of the humerus, radius-ulna, femur, and tibia by non-human
scavengers. Minimum number of elements (MNE) is used to assess the number
of specimens for each specific element examined. Accordingly, this
measurement is used to assess specific patterns in completeness of the skeletal
remains. These quantitative measures will be MNEpr, minimum number of
elements proximal portion (including any portion code for proximal or proximal
epiphysis); MNEds, “ds” for distal portion (including any code for distal or distal
epiphysis); MNEsh, “sh” for shaft portion (including any code for shaft or
diaphysis); and finally, MNEco, where “co” denotes complete portion. Further,
several ratios will be assessed using the MNE portions to illustrate differential
61
destruction between the specific elements and between specific portions (Lyman
1994:102-104). Descriptive statistics of the landmarks will be assessed at this
point as well and will be compared with the MNE indices created above for
comparison. This minor empirical test will evaluate the use of collecting
landmark data to understand overall destruction. In addition to collecting data
on portions, data are collected on mineral density values of skeletal elements as
put forth by Kreutzer (1992:276-281) and Lyman (1994:240-248).
The last quantitative measure used for this collection is minimum number
of animal units necessary to account for the elements in a collection (MAU)
(Lyman 1994:104-105). The formula for MAU is MNE₁ / number of times ₁ occurs
in one skeleton (Binford 1984:51; Lyman 1994:104-105). Finally, %MAU will be
assessed for the collection using the entire collection and then MNE portion
values. %MAU is found by taking MAU x 100 and dividing it by the maximum
MAU value (Binford 1984:51; Lyman 1994:104-105).
The MNE portions are used to assess independence from sex, side, and
utilization. The chi-square goodness-of-fit will assess if these variables are
independent from each other. Related to the MNE portions, the %MAU values
will be crossed in a scatter plot and analyzed using the chi-square goodness-of-fit
analysis with the mineral density values for Bison bison skeletal elements put
forward by Kreutzer (1992:276-281) and Lyman (1994:240-248).
62
Finally, bone mineral densities have often been used as a proxy for
understanding nutritional utility of specific skeletal elements (Brink 1997:264;
Kreutzer 1992:272; Kreutzer 1996:115). Kreutzer (1996:116) indicates that
“nutritionally valuable parts tend to be low-density parts.” It has further been
stated that carnivore modification is in most regards density mediated (Faith, et
al. 2007:2025-2026; Marean and Spencer 1991:651-652). Faith, et al (2007:2026)
also indicate that in scavenging events of less competition, animals may be
selective in their feeding by differentially choosing low density elements to
consume. However, in events of high competition animals will not differentiate
what elements and both low and high density portions of elements will be eaten
similarly (Faith, et al. 2007:2026). Given the information provided and for the
scope of this thesis, density values will be used as indicator of nutritional utility.
In the future directions section of Chapter 5 there will be a discussion on the use
of nutritional utility as a stepping stone for future projects on carnivore
modification.
The methods put forth in this section pertaining to indices and measures
will give an overall description of the herd demographics as well as illustrate the
scavenging patterns statistically allowing for a statistical representation to back
up the photographic representation of non-human scavenger destruction at
Kaplan–Hoover.
63
Carnivore Modification: Taphonomic Methods
Non-human scavenger modification has not always been addressed in the
literature. Typically, articles mention carnivore modification as present or not
and in most instances it is given in a percentage of presence in the collection.
Literature in the past decade (Binford 1981; Fisher 1995; Haynes 1980, 1982, 1983,
1991; Lyman 1994; Stiner 1994) have discussed the amount of modification in
terms of where it is on the skeleton, intensity, types of marks, and percentage on
the entire collection. It is important to consider non-human scavenger
modification in all aspects to accurately question the paleoecology and site
formation processes.
Two methods of data collection will be used to assess non-human
scavenger modification. Use of these methods, with alterations to each, will
achieve a holistic examination of non-human scavenger impacts on the collection.
The first method, as described previously, is based on a literature review of
carnivore modification typologies and images (Binford 1981; Fisher 1995; Lyman
1994; Stiner 1994). After an evaluation of previous and current typologies of
carnivore modification, it was necessary to survey the Kaplan–Hoover collection
for carnivore modification. As a result, the following modification types were
assessed: chipping back, crenellation, furrowing, pitting, puncture, salivary
polishing, scooping out, and tooth scoring (Figures 3.1-3.7). The second method
64
codes non-human scavenger modification based on intensity of skeletal element
utilization (Haynes 1982). This method allows for an overall view of the
destruction of the skeletal material and therefore gives a better representation of
the processes leading to burial of the carcass (Haynes 1982). Furthermore,
analysis of skeletal element utilization will allow for a more holistic view of
destruction by encouraging researchers to review the entire process instead of
cataloging specific marks, therefore, collecting data necessary to discover the
paleoecology and complete record of site formation processes (Haynes 1982).
In order to appreciate how these elements were coded for this project,
descriptions need to be defined in images as well as text. Chipping back (Figure
3.1) is defined by Binford (1981:51) as being caused by “strong carnassial teeth”
applying pressure to the compact bone and hence leaving a “mashed off”
appearance. Salivary polishing, which is caused by the salivary fluid of a tongue
moving over the bone, therefore wearing it smooth and chipping back are coded
in this project (when they appear together) as crenellations (Figure 3.2).
Crenellations are defined in the Webster’s New World Dictionary and Thesaurus
as “to furnish with battlements or with squared notches” as would be seen in a
fortress or castle. On skeletal elements, crenellations appear as sharp notches cut
from the proximal or distal ends. This code was only used on skeletal elements
that exhibited both chipping back and salivary polishing. Furrowing (Figure 3.3)
65
is defined for research in this project as the “gouging out” of
cancellous/trabecular bone. Pitting (Figure 3.4) is typically a product of an
animal gnawing at bone without intention of removing meat, thus leaving a
number of tiny pit marks (Binford 1981:46). Punctures (Figure 3.5) are easily
defined and visible by any observer. These marks are defined by Binford
(1981:44) as an area “where the bone has collapsed under the tooth,” leaving as
Lyman (1994:206) states “a clear, more or less oval depression in the bone, often
with flakes of the outer wall of the bone pressed into the puncture.” Scooping
out is defined in this research differently than it has been previously defined by
Lyman (1994:210), who defines it more as furrowing. In order to account for a
phenomenon seen on the collection, scooping out is similar to chipping back;
however, it is a larger chunk pulled back (Figure 3.6) off of the medullary cavity.
This is similar to Binford’s mark of channeling, although there is less of a crushed
edge and the piece removed was removed in one act (Binford 1981:51). Tooth
scoring (Figure 3.7) has been described by Binford (1981:46) as the “result of
either turning the bone against the teeth or dragging across” the bone. Binford
(1981:46-47) notes that scoring can resemble cut marks in that it looks like linear
dragging marks. In the Kaplan–Hoover collection these particular marks ranged
in breadth and therefore in many cases did not look like cut marks, but more like
deep grooves in the bone surface.
66
Figure 3.1: Chipping back on proximal end of femur, cranial view from
Kaplan–Hoover (Element # F28-8-440).
Figure 3.2: Crenellations on proximal end of humerus, cranial view from
Kaplan–Hoover (Element # F28-4-248).
67
Figure 3.3: Furrowing on proximal end of tibia, cranial view from
Kaplan–Hoover (Element # F28-3-150).
Figure 3.4: Pitting on proximal end of humerus, cranial view from
Kaplan–Hoover (Element # F27-24-711).
68
Figure 3.5: Punctures on the distal end of a femur, caudal view from
Kaplan–Hoover (Element # F28-3-226).
Figure 3.6: Scooping out on the proximal end of a humerus, caudal view from
Kaplan–Hoover (Element # F28-9-417).
69
Figure 3.7: Tooth scoring on the proximal end, head of the humerus from
Kaplan–Hoover (Element # F28-7-14).
Skeletal element utilization is coded as light, light-moderate, moderate,
moderate-heavy, and heavy (Figures 3.8-3.15). This is a slight variation on
Haynes (1982), who used full as a measure for moderate-heavy. This technique
is argued by Haynes (1982:274) to present a more holistic view of the non-human
scavenger damage at a site as well as illustrate that not all modification results in
specific marks and therefore would be difficult to code. Furthermore, Haynes
(1982:279-280) argues that without this holistic perspective, a researcher may
associate particular marks with specific animals instead of approaching the
modification from an un-biased stance. In light of taphonomic research, it is
70
therefore important to incorporate both methods to illuminate the entire range of
processes leading up to site formation.
Figure 3.8: Light utilization of a tibia, cranial view from Kaplan–Hoover (Element # F28-8-106).
Figure 3.9: Light/moderate utilization of a tibia, cranial view from
Kaplan–Hoover (Element # F28-4-250).
71
Figure 3.10: Light/moderate utilization of a femur, cranial view from
Kaplan–Hoover (Element # F27-15-116).
Figure 3.11: Moderate utilization of distal femur, cranial view from
Kaplan–Hoover (Element # F28-12-199).
72
Figure 3.12: Moderate/heavy utilization of a femur, cranial view from
Kaplan–Hoover (Element # F27-25-39).
Figure 3.13: Moderate/heavy utilization of a humerus, medial view from
Kaplan–Hoover (Element # F27-25-39).
73
Figure 3.14: Heavy utilization of two humeri, lateral view from
Kaplan–Hoover (Right element # F28-4-248 and left element # F27-18-103) .
Figure 3.15: Extreme heavy utilization of two humeri from Kaplan–Hoover
(Top element # F27-25-54 and bottom element # F27-23-227).
74
Finally, specific non-human modification types are assessed with
descriptive statistics. Percentages of non-human scavenger modification versus
no modification, compared to complete versus incomplete elements is assessed
to distinguish the overall amount of destruction. Chi-square goodness-of-fit is
used to determine any significance or independence of carnivore utilization from
sex, side, and element. Assessment of types of carnivore modification marks
compared with degree of utilization and bone mineral densities is discussed as
well.
Extant Non-human Scavengers: Ethological Methods
Scavenging, predatory, and nutrient attaining behaviors affect the
movement, and destruction of kill sites. Recognizing which predators prefer
which carrion and the general behavioral ecology of mammals in the Great
Plains will greatly increase the knowledge base on prehistoric hunting in North
America and increase awareness of outside mammalian components that affect
zooarchaeological site formation. In order to get at who contributed to the site
formation, it is necessary to research the specific non-human scavengers in the
Great Plains today and during the middle Holocene and discuss their
scavenging, hunting, and predating patterns.
75
Mammals of interest for this research project are Canis lupus and Canis
latrans (gray wolf and coyote) and Ursus arctos and Ursus americanus
(brown/grizzly and black bears). Foxes, wild cats, mustelids, and various birds
like vultures and ravens would also have been informative, it is assumed they
would not have ignored the carcasses; however, literature on them is not as
frequent, especially pertaining to carrion scavenging behaviors. A discussion of
the specific behaviors of non-human scavengers preceded this chapter in the
literature review and will continue further in the discussion where results will be
compared to assess similarities and differences.
Summary of Chapter
This chapter presented the methodologies used to assess the faunal
material from the Kaplan–Hoover bison bonebed. A holistic methods approach
using zooarchaeological, taphonomic and ethological data collection procedures
greatly increases the ability to infer the processes that altered the bonebed after
the hunt and before, during, and possibly after deposition. Zooarchaeological
methods will improve the understanding of herd demographics and descriptive
statistics on biogenic factors. Taphonomic methods will assist in the
understanding of bonebed formational processes; specifically those biogenic
influences and ethological research will increase the understanding of non-
76
human carnivore behaviors. Finally, these methods are not only applicable for
Kaplan–Hoover, but if adapted accordingly could be used for other faunal
assemblages, including single death events.
77
CHAPTER 4: RESULTS OF DATA ANALYSIS
This chapter presents the results of data analysis on the Kaplan–Hoover
Bison bison fauna collection. To begin, basic descriptive statistics concerning herd
characteristics are addressed. Basic frequencies for number of individual
specimens (NISP), minimum number of individuals (MNI), minimum number of
elements (MNE), and minimum animal units (MAU) are described. Included in
the descriptive statistics are analyses of sex from bivariate scatter plots. MNI is
based on sex and side cross-tabulations, and MNE will be described by specific
portions of elements, proximal, distal, shafts, and completes. These results are
useful in answering the first question in this thesis: what are the herd
characteristics?
Differential destruction analysis is performed using MNE, landmarks,
MAU, and the index %MAU. These statistics are used to discover specific
patterns in the destruction of the skeletal elements by non-human scavengers,
further answering the second question of inquiry: what are the patterns of
carnivore modification?
78
Finally, this chapter concludes with a discussion and analysis of carnivore
modification in terms of descriptive results pertaining to percentages and
degrees of specific modification marks as well as results of intensity and degree
of utilization. These final results are useful for illustrating the interactions
between the first two questions to make inferences of human and non-human
scavenger interactions and illustrate the relationships initiated by the institution
of large bison kills on the Great Plains.
Herd Characteristics Analysis
Analysis of herd characteristics is always of importance when doing
research with faunal assemblages. The importance lies in the ability to ask
questions of the collection that may illustrate information for taphonomy.
Important questions akin to whether sex and size of skeletal elements are related
to overall destruction or if scavengers preferentially selected specific elements.
Besides data collection of portion, segment, side, carnivore modification, and
measurements, sex was assessed to understand whether the collection was a
nursery herd or a male dominated herd.
Sex and determination of herd demographics are useful to results later in
this chapter. By comparing sex with specific types of carnivore modification
marks and degrees of utilization, patterns will emerge illustrating if there was
79
feeding preference by non-human scavengers. In addition, sex can be compared
with bone mineral densities to understand further if preferential selection exists.
By using chi-square goodness-of-fit, sex is compared to various measures to
determine significance of sex in relation to side, portions, utilization and
presence or absence of modification.
Sex Analysis
Assessing sex on the Kaplan–Hoover collection is complicated.
Destruction of the skeletal elements by non-human scavengers has made it
difficult to assess sex. Approximately 512 of the 1204 skeletal elements had some
carnivore modification. This damage made it difficult in a number of skeletal
elements to identify sex, because key measurements could not be accurately
taken due to lack of complete material. In other elements, breakage resulting
from interactions of post-depositional deterioration coupled with archaeological
collection damage added to the lack of ability to take accurate measurements. In
total 482 elements could be sexed. Of those 482, 66.2% (n = 319) elements are
female/sub-adult while 33.8% (n = 163) are male. The results for sex were
accomplished using bivariate scatter plots designating the particular
measurements that were in high enough abundance that accurate crosses could
be accomplished. For all of the sex scatter plot graphs, red indicates female or
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sub-adult and blue represents male. These graphs are located in Appendix E;
however, a brief discussion of the results for each element is given below.
Humerus
The humerus (Appendix E – Figure E.1) is the easiest element to sex (in
terms of number of scatter plots needed) and of the 93 humeri measured; only 57
could be sexed. Of those 70.2% (n = 40) are female and/or sub-adult and 29.8% (n
= 17) are male. Measurements 11, greatest depth of the medial distal end and 7,
breadth of distal articular surface are the most reliable measures used to sex
incomplete humeri (Todd 1987:162-163).
Radius-Ulna
Sex of the radius-ulna (Appendix E – Figures E.2 and E.3) is based on 63 of
the 127 elements that were possible to sex. It is estimated that 66.7% (n = 42) are
female and/or sub-adults and 33.3% (n = 21) are male. The radius-ulna was sexed
using measurements 9, greatest depth of proximal end, crossed with 4, greatest
breadth of proximal articular surface, and 11, greatest depth of distal end,
crossed with, 7 greatest breadth of distal end.
Metacarpal
The metacarpal and metatarsal (results discussed later) were sexed by
crossing measurement 5, greatest breadth of the distal end, by the remaining
measurements: greatest length (M1), greatest breadth of the proximal end (M2),
81
smallest breadth of the diaphysis (M3), and smallest depth of the diaphysis (M4).
According to Bedord (1974:208), greatest breadth of the distal end is good for
assessing sex of the metapodials. Thus, all of the remaining measurements were
crossed with this measure. Of the 107 metacarpals only 59 could be sexed
(Appendix E – Figures E.4-E.7). Of those 59, 61% (n = 36) are females and/or subadults and 39% (n = 23) are males.
Femur
The measurements used for the femora are greatest length from the head
(M3) and greatest depth of the distal epiphysis (M18) according to Todd
(1983:142). Unfortunately, the femur is damaged enough that those
measurements were not possible, specifically greatest length from the head. The
proximal ends of the femora were highly damaged by non-human scavengers
causing greatest length to be, in most instances, impossible. Therefore, it was
important to assess frequency distributions of measurements with the most
recorded data and then cross those with other measurements of similar amounts
of data. For this assessment, least depth of diaphysis (M17), least breadth of
diaphysis (M10), and greatest depth of head (M8) had the highest frequency
distributions of recorded data; therefore, they were crossed against each other to
gain sex information (Appendix E – Figures E.8-E.10). In total, of the 112 femora
82
only 40 could be sexed. Therefore, 23.2% (n = 26) are female/sub-adult and 12.5%
(n = 14) are male.
Tibia
The measurements used to assess sex on the tibia according to Todd
(1987:169) are depth of the proximal end (M15) and greatest breadth of the
proximal end (M4). However, similar to the femur, these measurements were
rare due to the non-human scavenger damage on the proximal ends. Therefore,
frequency distributions of the remaining measurements were used to determine
which would be useful for gaining sex of the tibia. The measurements with the
highest frequencies of data are: greatest length (M1), medial length (M2), least
breadth of the diaphysis (M6), and greatest breadth of the distal end (M7). These
were crossed against each other using bivariate scatter plots. The total number of
female/sub-adult and males is 50% (n = 14) each a total of 101 elements not sexed
out of 129 tibias (Appendix E – Figures E.11-E.15). This is due in large part to the
furrowing destruction on the proximal end of the tibia, which was caused by
non-human scavenging.
Metatarsal
Similar to the metacarpal, the metatarsal is sexed by crossing
measurement 5, greatest breadth of the distal end, by the remaining
measurements: greatest length (M1), greatest breadth of the proximal end (M2),
83
smallest breadth of the diaphysis (M3), and smallest depth of the diaphysis (M4).
For the metatarsal (Appendix E – Figures E.16-E.19) there are 59.3% (n = 32)
female/sub-adults and 40.7% (n = 22) males, with 70 elements not sexed out of
124 elements. Several of the problems with sexing the metapodials are due to the
breakage of the bones. Carnivore scavenging is limited on both the metacarpal
and metatarsal; however, many were broken completely in half, therefore
making greatest length difficult to measure. Other measurements were difficult
as well due to breakages on the medial and lateral surfaces of both the proximal
and distal ends.
Astragalus
For the astragalus, Morlan (1991) suggests use of measurements of the
distal width (M3) and medial length (M5) as accurate for determining sex
(Appendix E – Figure E.20). Crossing these measurements using a scatter-plot
identifies the sex distribution out of the 102 elements sexed: 56.9% (n = 58) are
female/sub-adults and 43.1% (n = 44) are males.
Calcaneus
The calcaneus was sexed using measurements from Morlan (1991). For
the calcaneus, the measurements most useful for determining sex include: distal
width (M4), distal depth (M5), length of talus (astragalus) facet (M6), and length
of tarsal c + 4 facet (M7). M5 and M4 are crosses as well as M6 and M7. After
84
analysis of the 114 calcanei only 70 could be sexed. Therefore, sex was
determined to be 89.9% (n = 71) are female/sub-adults and 10.1% (n = 8) are males
(Appendix E – Figures E.21-E.22). The difficulty in sexing the male individuals
in the calcaneus is likely due to the lack of reliable measurements and methods to
assess difference in size. Therefore, the assessment of sex on the calcanei could
be doubtful in both male and female cases, given that sex is difficult to ascertain.
Future empirical testing may illustrate a better means of determining sex for the
calcaneus.
Number of Individual Specimens
The total number of individual specimens (NISP) was 1204 based on the
humerus, radius-ulna, metacarpal, femur, tibia, metatarsal, astragalus, calcaneus,
third phalanx. Specific skeletal element NISP frequencies include: humerus 93,
radius-ulna 127, metacarpal 107, femur 112, tibia 129, metatarsal 124, astragalus
120, calcaneus 114, and third phalanx 278.
Minimum Number of Individuals
In order to assess the MNI (minimum number of individuals) for the
collection, cross tabulation was used in SPSS to illustrate the distribution of sex
and side, where side (SD) is coded as left (L), right (R), and (N) is not sided, and
where female/sub-adult is designated by (F), male is (M), and (N) is not sexed.
These codes have been defined in Appendix B.
85
Forelimb
Based on the cross tabulation for the humerus (Table 4.1), the MNI is 33,
indicated by the total number of sexed right humeri. Based on the side by sex
cross tabulation (Table 4.2), the radius-ulna has a MNI of 33, illustrated by the
total number of female/sub-adult left side elements and male right side elements
present. The MNI of the metacarpal (Table 4.3) is based on the total number of
female/sub-adult right side elements and number of male left side elements at 22
individuals.
Humerus SD * SEX Crosstabulation
Count
SEX
M
F
SD
N
Total
L
20
4
12
36
N
0
0
10
10
R
20
13
14
47
40
17
36
93
Total
Table 4.1: Cross tabulation of side and sex for MNI analysis of the humerus.
Radius-Ulna SD * SEX Crosstabulation
Count
SEX
M
F
L
SD
Total
22
N
10
Total
35
67
N
0
0
2
2
R
20
11
27
58
42
21
64
127
Table 4.2: Cross tabulation of side and sex for MNI analysis of the radius-ulna.
86
Metacarpal SD * SEX Crosstabulation
Count
SEX
M
F
L
SD
16
N
12
Total
22
50
N
0
0
2
2
R
20
11
24
55
36
23
48
107
Total
Table 4.3: Cross tabulation of side and sex for MNI analysis of the metacarpal.
Hind Limb
Cross tabulation of the femur (Table 4.4) assigns the MNI at 24, based on
the total number of sexed right side elements. Cross tabulation for the tibia
(Table 4.5) indicates that the MNI is 16, based on the total number of left
elements for the female/sub-adults and right side elements for the males. For the
metatarsal (Table 4.6) cross tabulation, the MNI is 29 based on the total number
of female/sub-adult and male right side elements.
Femur SD * SEX Crosstabulation
Count
SEX
M
F
SD
Total
N
Total
L
10
6
28
44
N
0
0
19
19
R
16
8
25
49
26
14
72
112
Table 4.4: Cross tabulation of side and sex for MNI analysis of the femur.
87
Tibia SD * SEX Crosstabulation
Count
SEX
M
F
SD
N
Total
L
8
6
44
58
N
0
0
7
7
R
6
8
50
64
14
14
101
129
Total
Table 4.5: Cross tabulation of side and sex for MNI analysis of the tibia.
Metatarsal SD * SEX Crosstabulation
Count
SEX
M
F
SD
Total
N
Total
L
14
11
41
66
N
0
0
7
7
R
18
11
22
51
32
22
70
124
Table 4.6: Cross tabulation of side and sex for MNI analysis of the metatarsal.
Astragalus, Calcaneus, and Third Phalanx
Finally, the remaining elements to assess MNI are the astragalus (Table
4.7) and calcaneus (Table 4.8). MNI was not determined for the third phalanx
because of the lack of methods to assign sex, and besides lateral or medial side, it
is difficult to assign right or left side. Cross tabulations for the astragalus and the
calcaneus assign MNI as 55, based on the left and 37, based on the total number
of female/sub-adult right elements and left side male elements respectively.
88
Astragulus SD * SEX Crosstabulation
Count
SEX
F
SD
M
N
Total
L
32
R
26
21
9
56
58
44
18
120
Total
23
9
64
Table 4.7: Cross tabulation of side and sex for MNI analysis of the astragalus.
Calcaneus SD * SEX Crosstabulation
Count
SEX
M
F
SD
Total
N
Total
L
33
4
18
55
N
0
0
3
3
R
38
4
14
56
71
8
35
114
Table 4.8: Cross tabulation of side and sex for MNI analysis of the calcaneus.
Approximately, 55 individuals are present at Kaplan–Hoover based on the
total number of left astragali in the sample assemblage. This number would be
different if the damage to the skeletal elements was not so great and if
excavations could have continued to the bottom of the bonebed. It has been
estimated by Todd, et al. 2001:135) that the bonebed likely held as many as 200
individual bison carcasses. To continue discussing the damage sustained by the
skeletal elements because of non-human scavenging the following section will
address the amount of destruction on the skeletal elements at Kaplan–Hoover.
89
Differential Destruction Analysis
This section discusses the results of zooarchaeological analysis to
understand the specific skeletal remains by portion. To fully appreciate the nonhuman scavenger damage, it is important to understand how much of a
collection is present or absent based on proximal, shaft, distal, and complete
portions of the skeletal elements. Further analysis of those portions and
landmarks in relation to bone mineral densities on the humerus, radius-ulna,
femur, and tibia using correlation coefficients will increase the comprehension of
the final section on carnivore modification.
Minimum Number of Elements by Coded Location
Minimum number of elements (MNE) was given additional codes to
assess the overall destruction of the collection. Accordingly, MNE was divided
up into four separate skeletal locations: proximal (PR), distal (DS), shaft (SH),
and complete elements (CO). Later on in this chapter the complete code will be
broken up differently. Multiple portion codes fell in to each of these particular
location codes. Proximal skeletal elements were any with the following codes:
proximal (PR), proximal plus less than half the shaft (PRS), proximal plus more
than half the shaft (PSH – coded as one PR and one SH), proximal epiphysis
(PRE), proximal diaphysis (DPR), diaphysis plus fused proximal epiphysis
(DFP), olecranon portion of the ulna (OLC), trochlear notch of the ulna (ANC),
90
and head (HE) (after Todd 1983, 1987). For distal skeletal elements the following
codes applied: distal (DS), distal plus less than half the shaft (DSS), distal plus
more than half the shaft (DSH – coded as one DS and one SH), distal epiphysis
(DSE), distal diaphysis (DDS), and diaphysis plus fused distal epiphysis (DFD).
To use these measurements as comparison with the landmarks the only skeletal
elements that MNE was assessed for are the humerus, radius-ulna, femur, and
tibia. For skeletal elements with a portion of shaft the following codes were
relevant: shaft (SH), diaphysis (DF), blade of the ulna (BL) and the PSH and DSH
codes were coded with one PR/DS respectively and one SH each. Finally,
specific codes that were not useful for the MNE coding system were assigned a
value of not applicable (N); those include: flake (FK), impacted flake (IMK),
condyle (CDL), epiphysis (EP), and unspecified (US). In addition to the coding
system developed, the MNE results will be compared to mineral densities (Table
4.9) from Kreutzer (1992) and Lyman (1994) for Bison bison skeletal remains.
91
Bison Mineral Densities (adapted from
Kreutzer 1992 and Lyman 1994)
Element
Scan
Site
Portion
Density
Value
Humerus
HU1
PR
.24
Humerus
HU2
PR
.25
Humerus
HU3
SH
.45
Humerus
HU4
DS
.48
Humerus
HU5
DS
.38
Ulna
UL1
PR
.34
Ulna
UL2
PR
.69
Radius-ulna
RA1
PR
.48
Radius-ulna
RA2
PR
.56
Radius-ulna
RA3
SH
.62
Radius-ulna
RA4
DS
.42
Radius-ulna
RA5
DS
.35
Femur
FE1
PR
.56
Femur
FE2
PR
.73
Femur
FE3
PR
.66
Femur
FE4
SH
.70
Femur
FE5
DS
.39
Femur
FE6
DS
.48
Tibia
TI1
PR
.45
Tibia
TI2
PR
.53
Tibia
TI3
SH
.87
Tibia
TI4
DS
.74
Tibia
TI5
DS
.56
Table 4.9: Bone mineral densities for Bison bison, adapted from Kreutzer (1992:276-281 – Figure 2
and Table 2) and Lyman (1994:240-248 - Figures 7.4-7.6 and Tables 7.6-7.7).
Humerus
Humerus MNE portions (Table 4.10) indicate that there are approximately
42.4% of the distal ends remaining. When this is compared to the mineral
densities for two sites on the proximal end of the humerus by (Kreutzer 1992;
Lyman 1994), the density values for HU1 and HU2 are 0.24 and 0.25,
92
respectively. Compared to the distal end whose mineral density values are 0.48
(HU4) and 0.38 (HU5), it is clear that mineral densities in the humerus account
for the higher frequency of MNEds and MNEsh results, suggesting more distal
ends and shafts remaining.
Humerus MNE Portions
Portion
CO
Frequency
11
Percent
7.3
DS
64
42.4
N
1
.7
PR
10
6.6
SH
65
43.0
151
100.0
Total
Table 4.10: MNE portions for the humerus.
Radius-Ulna
Radius-ulna MNE portions (Table 4.11) are mixed and vary from the
humerus results. The mineral densities for the proximal sites UL1, UL2, RA1,
and RA2 are 0.34, 0.69, 0.48, and 0.56 (Kreutzer 1992, Lyman 1994). Ulna damage
is typically to the very proximal portion of the olecranon process, which has the
density value of 0.34, and the density for the proximal end of the radius
compares with the MNEpr frequencies at approximately 29.7%. Approximately
38.4% of the elements were complete, which is reasonable given that the density
values for the rest of the radius-ulna are 0.62 (RA3), 0.42 (RA4), and 0.35 (RA5).
93
Radius-Ulna MNE Portions
CO
Frequency
53
Percent
38.4
DS
30
21.7
Portion PR
41
29.7
SH
14
10.1
138
100.0
Total
Table 4.11: MNE portions for the radius-ulna.
Femur
Femora MNE portions (Table 4.12) indicates that more of the proximal
end and shafts of the elements are remaining, at approximately 35.3% and 32.4%
respectively. Where as the distal end and completes are the least represented.
When these results are compared to the mineral density values from Kreutzer
(1992) and Lyman (1994), the results begin to make sense. The proximal scan
sites for mineral densities FE1, FE2, and FE3 are all higher densities at 0.56, 0.73,
and 0.66 respectively (Kreutzer 1992, Lyman 1994). For the shaft and distal end
the remaining scan sites FE4, FE5, and FE6 show the following densities: 0.70,
0.39, and 0.48, all of which are denser than the proximal end of the humerus,
suggesting difficulty in breaking through the elements (Kreutzer 1992:276-281,
Lyman 1994:246-248).
94
Femur MNE Portions
Portion
CO
Frequency
19
Percent
13.7
DS
25
18.0
N
1
.7
PR
49
35.3
SH
45
32.4
139
100.0
Total
Table 4.12: MNE portions of the femur.
Tibia
MNE portions of the tibia (Table 4.13) are similar to that of the humerus;
however, the mineral densities are very different. Approximately 30.2% of the
distal tibias are present in the collection and 36.3% of the 129 total tibias are shaft
remains. Compared to the mineral densities for the proximal end of 0.45 (TI1)
and 0.53 (TI2) there is question as to why there are similar results to the humerus
MNEds, but a drastically higher density measure by almost 50% higher density
(Kreutzer 1992, Lyman 1994). For the shaft and distal ends the densities are as
follows: 0.87 (TI3), 0.74 (TI4), and 0.56 (TI5), all of which are very dense
correlating with the lower frequencies of MNEsh and MNEpr (Kreutzer 1992,
Lyman 1994).
95
Tibia MNE Portions
Portion
CO
Frequency
47
Percent
26.3
DS
54
30.2
N
1
.6
PR
12
6.7
SH
65
36.3
179
100.0
Total
Table 4.13: MNE portions of the tibia.
Use of MNE portions is more descriptive than MNE alone. Lyman
(1994:103-104) discusses the use of MNEends, MNEshafts, and MNEcomp
(indicating completes) as an important tool for deriving an understanding of
what a faunal assemblage represents in terms of portion (Lyman 1994:103-104).
Given that the faunal assemblage used for this thesis is damaged by non-human
scavengers any means of deriving the amount of skeletal material remaining will
be useful for understanding the overall destruction of the material.
Minimum Number of Individuals by Skeletal Landmarks
Skeletal landmarks are unique anatomical marks on specific skeletal
elements that indicate either muscle attachments or areas where vessels may pass
through the bone itself. For this thesis, landmarks were recorded as another
means of indicating portion remaining of skeletal elements in the sample used
from Kaplan–Hoover. Hill (1994:169) used skeletal landmarks in a similar way
96
to increase the identification of what portions were remaining. To identify the
more useful tool for this thesis, the landmarks were coded based on portion
(proximal, distal, shaft) and frequency tables were produced for the humerus,
radius-ulna, femur, and tibia to assess the usefulness between MNE portion
codes as listed in the previous section and landmarks.
Humerus
For the humerus there were six landmarks (Table 4.14) recorded: deltoid
tuberosity (LM1), tubercle for attachment of medial collateral ligament (LM2),
major tuberosity (LM3), proximal olecranon fossa (LM4), posterolateral nutrient
foramen (LM5), and teres major tubercle (LM6) (Hill 1994:169). In order to
understand how these measurements could be used as comparative with MNE
portion codes, it is important to assign them portion locations for where they
reside on the element. LM1, LM3, and LM6 are all proximally located therefore
they are receiving the code PR. LM2 and LM4 are both distally located, therefore
coded DS and LM5 is the only shaft location, SH. Landmarks were coded as
either being present (P), absent (A), or half-present (H). These data are useful for
understanding overall remains of particular portions, but to what extent that will
be useful for overall data analysis is difficult to ascertain given that there are
more proximal portion landmark codes than distal and shaft. When frequencies
97
of the landmarks are considered, there are significantly more distal ends as well
as shafts, which does corroborate with the MNEds, 10.8% and MNEpr, 68.8%.
Humerus Landmark Frequencies
HalfPresent
Absent
Landmark
Portion Present
LM1
PR
30
24
39
LM2
DS
68
0
25
LM3
PR
2
1
90
LM4
DS
67
3
23
LM5
SH
65
0
28
LM6
PR
70
2
21
Table 4.14: Frequency table of landmarks on the humerus.
Radius-Ulna
The radius-ulna has two landmarks (Table 4.15) recorded: posterolateral
nutrient foramen (LM1) and radial tuberosity (LM2) (Hill 1994:169). For these
landmarks, LM1 is in the shaft (SH) location and LM2 is on the proximal end. A
comparison of these data with the MNEpr results of 32.3%, indicating a slight
similarity.
Radius-Ulna Landmark Frequencies
Present
HalfPresent
Absent
Landmark
Portion
LM1
SH
56
32
39
LM2
PR
62
32
33
Table 4.15: Frequency table of landmarks on the radius-ulna.
98
Femur
Four landmarks were recorded for the femur (Table 4.16): anterior
nutrient foramen (LM1), supracondyloid fossa (LM2), major trochanter (LM3),
and minor trochanter (LM4) (Hill 1994:169). LM1 is a shaft (SH) portion location,
while LM2 is a distal portion location. The last two landmarks 3 and 4 are
proximal portion locations. In view of the fact that there is an overrepresentation of the proximal end, this could bias results. However, when
looking at Table 4.16, it is evident that more proximal portions and shafts are
absent when compared to the distal portions.
Femur Landmark Frequencies
Half_
Present
Absent
76
Landmark
Portion Present
LM1
SH
36
0
LM2
DS
58
3
51
LM3
PR
4
1
107
LM4
PR
42
4
66
Table 4.16: Frequency table of landmarks on the femur.
Tibia
Three landmarks (Table 4.17) were recorded for the tibia: anterior crest
(LM1), posterolateral nutrient foramen (LM2), and anterior nutrient foramen
(LM3) (Hill 1994:169). LM1 is a proximal portion location, while landmarks 2
and 3 are shaft portion locations. Similarly to the radius-ulna these data on
landmarks for the tibia are useless for understanding destruction since there is
99
not a representation of distal portion locations. This may be changed if more
landmarks were recorded, or if the landmarks chosen were more comparable to
actual gross anatomical regions of muscle attachments.
Tibia Landmark Frequencies
HalfPresent
Absent
Landmark
Portion Present
LM1
PR
37
39
53
LM2
SH
81
0
48
LM3
SH
21
0
108
Table 4.17: Frequency table of landmarks on the tibia.
Finally, when comparing the MNE portions (MNEpr, MNEsh, MNEds) to
the landmarks portions for the humerus, radius-ulna, femur, and tibia, it is
apparent that collecting information for the MNE portions is more encompassing
than the landmarks used. Because of the lack of obvious distinctions as with the
MNE portion codes, there are limited reasons, besides as a back up to the already
well established and defined coding system used, to collect landmark codes for
this particular type of project. Given this information, increasing the number of
landmarks collected could alter this assumption. In addition, choosing
landmarks related to specific muscles that may be of importance to hunters or
non-human scavengers could elucidate a different pattern and therefore provide
evidence that landmarks are highly useful.
100
Minimum Animal Units and %MAU
Minimum number of animal units equals the MNE of one specific element
divided by the number of times that element occurs in one individual
skeleton/animal (Lyman 1994:105). For the Kaplan–Hoover collection, the
unmodified, just skeletal element MAU values are located in Table 4.18.
However, to gain a more holistic view of the assemblage in terms of damage, it is
important to derive MAU and %MAU based on the MNE portion values for the
humerus, radius-ulna, femur, and tibia (Table 4.19) and use these results for a
comparison with bone mineral densities to assess the destruction to the
assemblage. MNEpr, MNEds, and MNEsh will be used and any elements with
MNEco data will be coded as one MNEpr, MNEds, and MNEsh. This will allow
for a more complete view of the overall destruction and can then be crossed with
mineral densities.
Kaplan-Hoover MAU and %MAU
Element
Humerus
MNE
# IN
SKELETON
MAU
%
MAU
93
2
46.5
72.093
Radius-ulna
127
2
63.5
98.450
Metacarpal
107
2
53.5
82.946
Femur
112
2
56.0
86.822
Tibia
129
2
64.5
100.000
Metatarsal
124
2
62.0
96.124
Astragalus
120
2
60.0
93.023
Calcaneus
114
2
57.0
88.372
Third Phalanx
278
8
34.8
53.876
Table 4.18: Minimum animal units and % MAU values for Kaplan–Hoover collection.
101
Analysis of the %MAU for the entire collection illustrates that the most
abundant skeletal element in the collection is the tibia (Table 4.19). When MNE
portions are considered in the %MAU assessment, the most abundant element is
the shaft of the tibia (Table 4.19). When this is compared to the mineral density
of the shaft of the tibia, 0.87, it appears to be a reasonable assessment. To help
discover if %MAU and bone mineral densities are related, a Spearman
correlation analysis is undertaken below.
Skeletal Element Portions MAU and %MAU
MNE
Portion
Totals
Element
MNE
Portion
# in
Skeleton
MAU
%
MAU
Mineral
Densities
Humerus
MNEpr
21
2
10.5
18.910
.245
Humerus
MNEsh
75
2
37.5
67.560
.450
Humerus
MNEds
75
2
37.5
67.560
.430
Radius-ulna
MNEpr
99
2
49.5
89.180
.518
Radius-ulna
MNEsh
67
2
33.5
60.360
.620
Radius-ulna
MNEds
78
2
39.0
70.270
.385
Femur
MNEpr
67
2
33.5
60.360
.650
Femur
MNEsh
64
2
32.0
57.560
.700
Femur
MNEds
45
2
22.5
40.540
.435
Tibia
MNEpr
58
2
29.0
52.250
.490
Tibia
MNEsh
111
2
55.5
100.000
.870
Tibia
MNEds
102
2
51.0
91.820
.650
Table 4.19: %MAU based on MNE portion codes for Kaplan–Hoover collection.
For Kaplan–Hoover, there is slight correlation between the two variables
(Figure 4.1) and the Spearman correlation test is 0.399 (Table 4.20), which is not
significant at the alpha 0.05 level, suggesting that there are no specific selection
biases during the non-human scavenging based on bone mineral densities.
102
Further, this indicates that there were likely other taphonomic factors such as
fluvial transport and sedimentation that also influenced the movement and
destruction of the bonebed.
Figure 4.1: Scatter plot of %MAU and bone mineral densities for the humerus, radius-ulna,
femur, and tibia.
103
Spearman Correlations
%MAU
%MAU
Correlation
Coefficient
Mineral Density
1.000
.399
Sig. (2-tailed)
N
Spearman's
rho
Mineral
Density
.199
12
12
Correlation
Coefficient
.399
1.000
Sig. (2-tailed)
.199
N
12
12
Table 4.20: Spearman correlation of %MAU and bone mineral density crosses.
Carnivore Modification Analysis
This section presents the results of carnivore modification analysis;
specifically, the descriptions of carnivore modification and utilization as well as
results pertaining to overall destruction and intensity of destruction.
General Descriptive Statistics
Entire Collection
The skeletal elements from the Kaplan–Hoover sample are more
destroyed than other recorded sites. For the entire collection, approximately
42.5% (n = 1204) of elements measured for this thesis exhibit carnivore
modification (Figure 4.2). When applying sex to this, female/sub-adult
specimens were modified by non-human scavengers 2:1, with 115 (n = 482 sexed)
being female and 62 male. The most prominent types of carnivore modification
include: furrowing, punctures, and pitting; however, in most elements there are
104
countless marks overlapping each other, therefore, making it difficult to fully
understand which are present and which could have been had they not been
covered up by others. For utilization, the entire collection fell mainly into heavy
or light utilization; approximately 13% are heavily utilized and 17% light.
Figure 4.2: Presence or absence of modification on the entire collection used for this project.
Modification and Intensity: Specific Skeletal Elements
Humerus
For the humerus, 93.55% of the elements have some carnivore
modification. The three most prominent types of carnivore modification on the
humerus were crenellations, pitting, and furrowing. The amount of carnivore
utilization (Figure 4.3) is mostly heavy with 65.59% (n = 45).
105
Carnivore Utilization - Humerus
Number of Elements
70
60
50
40
30
20
10
0
H
MH
M
LM
L
I
N
Carnivore Utilization
Figure 4.3: Percentage of carnivore utilization for the humerus.
In order to understand the damage to the humerus, chi-square goodnessof-fit tests were performed on a number of the variables. First, sex of the
elements is crossed against carnivore utilization (Table 4.21). For sex and
carnivore utilization, chi-square test for independence is 30.242, with df = 12. The
asymptotic significance value is 0.003, which is significant at the alpha 0.05 level.
Therefore, the sex of the skeletal element is not independent of carnivore
utilization. This indicates that scavengers were not preferentially selecting
skeletal material for feeding. Second, analysis of whether sex and presence or
absence of modification is significant was analyzed using the chi-square test of
independence as well (Table 4.22). Accordingly, the chi-square test for
independence is 0.130, with df = 2. The asymptotic significance value is 0.937
106
which is not significant at the alpha 0.05 level. Therefore, the sex of the skeletal
element is independent of the presence or absence of modification. This
indicates that sex is not a factor is selection of elements for consumption by
scavengers. This makes sense taking into consideration almost all of the humeri
are modified.
Sex and Carnivore Utilization Chi-Square
Analysis
Value
Pearson Chi-Square
N of Valid Cases
Asymp. Sig.
(2-sided)
df
30.242
12
.003
93
Table 4.21: Chi-square analysis of sex and carnivore utilization for the humerus.
Sex and Modification Chi-Square Analysis
Value
Pearson Chi-Square
N of Valid Cases
.130
Asymp. Sig.
(2-sided)
df
2
.937
93
Table 4.22: Chi-square analysis of sex and modification for the humerus.
Carnivore utilization is an important variable to consider in relation to
side and portion as well. For carnivore utilization and side, the chi-square test of
independence is used (Table 4.23). For this test, the chi-square test for
independence is 59.246, with df = 12. The asymptotic significance value is 0.000,
which is significant at the alpha 0.01 level. For carnivore utilization and portion
107
(Table 4.24), the chi-square test for independence is 157.595, with df = 24. The
asymptotic significance value is 0.000, which is significant at the alpha 0.01 level.
Carnivore Utilization and Side Chi-Square
Analysis
Value
Pearson Chi-Square
df
59.246
N of Valid Cases
Asymp. Sig.
(2-sided)
12
.000
93
Table 4.23: Chi-square analysis of carnivore utilization and side for the humerus.
CU * SD Crosstabulation
Count
SD
L
CU
Total
N
R
Total
H
26
0
35
61
I
1
10
5
16
L
3
0
3
6
LM
2
0
0
2
M
1
0
0
1
MH
1
0
1
2
N
2
0
3
5
36
10
47
93
Table 4.24: Carnivore utilization and side cross-tabulation for the humerus.
Both of these tests indicate that carnivore utilization is not independent of
side or portion, suggesting that specific sides may have more utilization than
others. This could be due to the order or manner in that the elements were
processed by human hunters. For portion these results suggest that particular
portions have specific types of carnivore modification. In fact, when a simple
108
cross-tabulation of carnivore utilization with side and portion is accomplished it
is apparent that there are relationships between which sides are modified more
often (Table 4.25). Of specific interest is heavy utilization with side and portion.
In Table 4.24, there are 35 right and 26 left side humeri that are heavily modified,
while in Table 4.26 there are 53 distal elements that had heavy utilization and 56
of the shaft elements, meaning that the entire proximal portion of the humerus
was removed by non-human scavenging.
Carnivore Utilization and Portion
Chi-Square Analysis
Value
Pearson Chi-Square
157.595
N of Valid Cases
Asymp. Sig.
(2-sided)
df
24
.000
151
Table 4.25: Chi-square analysis of carnivore utilization and portion for the humerus.
CU * POR Crosstabulation
Count
POR
CO
CU
Total
DS
N
PR
SH
Total
H
3
53
0
0
56
112
I
0
4
1
9
3
17
L
3
2
0
1
2
8
LM
2
0
0
0
0
2
M
1
0
0
0
0
1
MH
2
0
0
0
0
2
N
0
5
0
0
4
9
11
64
1
10
65
151
Table 4.26: Carnivore utilization and portion cross-tabulation for the humerus.
109
Radius-Ulna
Given that the radius-ulnae are fairly dense elements with bone density
values at scan sites ranging from 0.34–0.69 (Table 4.9) and less meat in general,
this element exhibits less destruction than the humerus. Accordingly,
approximately 48.82% exhibit carnivore modification. The most prominent type
of carnivore modification is furrowing, which accounts for over 50% of the type
of modification on the radius-ulna. In terms of carnivore utilization, 33.07% of
the elements illustrate light modification and over 50% have no modification at
all (Figure 4.4).
Carnivore Utilization - Radius-Ulna
Number of Elements
70
60
50
40
30
20
10
0
H
MH
M
LM
L
I
N
Carnivore Utilization
Figure 4.4: Carnivore utilization for the radius-ulna.
Similar to the humeri, chi-square test of independence were performed for
sex against carnivore utilization, sex against presence or absence of modification,
110
and carnivore utilization against side. For these three tests, the chi-square tests
are insignificant. For each test the asymptotic significance was greater than the
alpha 0.05 level, suggesting that these variables are independent of each other.
This could be due to the density of the elements, since non-human scavenging
would have been difficult. However, when carnivore utilization is crossed with
portion (Table 4.27), the chi-square test for independence is 28.098, with df = 12.
The asymptotic significance value is 0.005 which is significant at the 0.05 level.
This test suggests there is no independence between carnivore utilization and
portion. When a cross-tabulation of carnivore utilization and portion is
displayed it is apparent (Table 4.28), that there are slightly more complete
elements with light modification than distal and proximal elements. However
this is a problem with the data because most of the portion coded elements fall
into “no” carnivore utilization, which is where the real correlation is.
Carnivore Utilization and Portion
Chi-Square Analysis
Value
Pearson Chi-Square
N of Valid Cases
28.098
Asymp. Sig.
(2-sided)
df
12
.005
138
Table 4.27: Chi-square analysis of carnivore utilization and portion for the radius-ulna.
111
CU * POR Crosstabulation
Count
POR
CO
CU
DS
PR
SH
Total
H
1
1
2
3
7
L
22
8
12
2
44
LM
3
0
6
0
9
M
0
1
4
0
5
N
27
20
17
9
73
53
30
41
14
138
Total
Table 4.28: Cross-tabulation of carnivore utilization and portion for the radius-ulna.
Femur
Similar to the humeri, the femora were heavily modified by non-human
scavengers. Approximately, 93.75% of the femora have evidence of carnivore
modification. The most prevalent types of carnivore modification are furrowing
and punctures, with the rest of the types occurring between 1–15% of the time.
Carnivore utilization (Figure 4.5) is broken up disproportionately with
approximately 41% of the elements being heavily utilized, light to moderate
utilization occurring on 23.2% of the elements, and light and moderate occurring
on 11.6% equally.
112
Number of Elements
Carnivore Utilization - Femur
50
40
30
20
10
0
H
MH
M
LM
L
I
N
Carnivore Utilization
Figure 4.5: Carnivore utilization for the femur.
For the femur, sex, modification, side, carnivore utilization, and portion
variables were used in chi-square analyses to understand any relationships. First
carnivore utilization and sex were analyzed (Table 4.29), the chi-square test for
independence is 32.323, with df = 10. The asymptotic significance value is 0.000
which is significant at the 0.01 level. This suggests that carnivore utilization and
sex for the femur are not independent of each other. However, when sex and
modification are analyzed (Table 4.30) using the chi-square test for
independence, the value is 4.148 with df = 2 and the asymptotic significance is
0.126, which is not significant at the 0.05 level. These results suggest, as they did
for the humerus and the radius-ulna, that there is no relationship between sex
and modification.
113
Sex and Carnivore Utilization Chi-Square
Analysis
Value
Pearson Chi-Square
N of Valid Cases
Asymp. Sig.
(2-sided)
df
32.323
10
.000
112
Table 4.29: Chi-square analysis of sex and carnivore utilization for the femur.
Sex and Modification Chi-Square Analysis
Value
Pearson Chi-Square
N of Valid Cases
4.148
Asymp. Sig.
(2-sided)
df
2
.126
112
Table 4.30: Chi-square analysis of sex and modification for the femur.
As in the humerus and radius-ulna, carnivore utilization and side are
analyzed using the chi-square test of independence. For the femur, the chisquare test for independence value is 39.442, with df = 10 (Table 4.31). The
asymptotic significance value is 0.000 which is significant at the 0.01 level. This
chi-square analysis indicates that carnivore utilization and side are not
independent of each other. When a cross-tabulation (Table 4.32) of the two
variables is presented there is an obvious trend towards heavy utilization on
both the left and right sides of the elements.
114
Carnivore Utilization and Side Chi-Square
Analysis
Value
Pearson Chi-Square
39.442
N of Valid Cases
Asymp. Sig.
(2-sided)
df
10
.000
112
Table 4.31: Chi-square analysis of carnivore utilization and side for the femur.
CU * SD Crosstabulation
Count
SD
L
CU
Total
N
R
Total
H
22
0
24
46
L
4
2
7
13
LM
4
13
9
26
M
8
1
4
13
MH
3
0
4
7
N
3
3
1
7
44
19
49
112
Table 4.32: Carnivore utilization and side cross-tabulation for the femur.
Portion is important to consider when discussing carnivore utilization,
since the scavengers are the agents that affected the amounts of elements
remaining. When carnivore utilization and portion are used in a chi-square
analysis the independence value is 69.343, with df = 20 (Table 4.33). The
asymptotic significance value is 0.000 which is significant at the 0.01 level. These
results indicate that carnivore utilization is not independent of portion, similar to
the humeri and radius-ulnae. When a cross-tabulation of the two variables is
presented there are mostly shafts with heavy utilization (Table 4.34).
115
Carnivore Utilization and Portion
Chi-Square Analysis
Value
Pearson Chi-Square
69.343
N of Valid Cases
Asymp. Sig.
(2-sided)
df
20
.000
139
Table 4.33: Chi-square analysis of carnivore utilization and portion for the femur.
CU * POR Crosstabulation
Count
POR
CO
CU
DS
N
PR
SH
Total
H
1
10
0
18
36
65
L
6
0
0
7
0
13
LM
6
6
0
13
2
27
M
4
3
0
6
3
16
MH
2
3
0
2
3
10
N
0
3
1
3
1
8
19
25
1
49
45
139
Total
Table 4.34: Carnivore utilization and portion cross-tabulation for the femur.
Tibia
The tibia falls somewhere between the humerus or femur and radius-ulna
in terms of amount of modification and destruction. First of all 71.32% of the
tibias exhibit some form of modification. Most prominent carnivore modification
types include: furrowing, chipping back, and pitting. Approximately, 34.88% of
the elements were heavily utilized, with 24.81% lightly and 28.68% not utilized at
all (Figure 4.6).
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Number of Elements
Carnivore Utilization - Tibia
50
40
30
20
10
0
H
MH
M
LM
L
I
N
Carnivore Utilization
Figure 4.6: Carnivore utilization for the tibia.
In the same way as the previous skeletal elements, destruction of the tibia
is analyzed using the chi-square test of independence. The first variables to be
analyzed are carnivore utilization and sex (Table 4.35). The chi-square test for
independence value is 50.426, with a df = 6. The asymptotic significance value is
0.000 which is significant at the alpha 0.01 level. Illustrating again, that carnivore
utilization and sex on the tibia is not independent of each other, as was the case
in the humerus and femur. A chi-square test for independence of sex and
modification (Table 4.36) has a value of 3.623, with df = 2. The asymptotic
significance value is 0.163 which is not significant at the alpha 0.05 level. These
results indicate that as with the humerus, radius-ulna, and femur, sex and
modification are independent of each other.
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Sex and Carnivore Utilization Chi-Square
Analysis
Value
Pearson Chi-Square
N of Valid Cases
Asymp. Sig.
(2-sided)
df
50.426
6
.000
129
Table 4.35: Chi-square analysis of sex and carnivore utilization for the tibia.
Sex and Modification Chi-Square Analysis
Value
Pearson Chi-Square
N of Valid Cases
3.623
Asymp. Sig.
(2-sided)
df
2
.163
129
Table 4.36: Chi-square analysis of sex and modification for the tibia.
Carnivore utilization is used to illustrate other comparisons, such as its
relationship to side and portion. For carnivore utilization and side, the chisquare test for independence is used (Table 4.37). For this test, the chi-square test
for independence is 7.019, with df = 6. The asymptotic significance value is 0.319,
which is not significant at the alpha 0.05 level. This indicates that carnivore
utilization is independent from side for the tibia. This could be related to the fact
that there are far more elements that are not sexed than are sexed. If those
results were changed, the results for this would change and could change to
reflect the results identified by the carnivore utilization and side analysis for the
humerus and femur. For carnivore utilization and portion (Table 4.38), the chisquare test for independence is 118.355, with df = 12. The asymptotic significance
118
value is 0.000, which is significant at the alpha 0.01 level. This test indicates that
carnivore utilization is related to portion and is not independent of portion.
Carnivore Utilization and Side Chi-Square
Analysis
Value
Pearson Chi-Square
N of Valid Cases
Asymp. Sig.
(2-sided)
df
7.019
6
.319
129
Table 4.37: Chi-square analysis of carnivore utilization and side for the tibia.
Carnivore Utilization and Portion
Chi-Square Analysis
Value
Pearson Chi-Square
N of Valid Cases
118.355
Asymp. Sig.
(2-sided)
df
12
.000
179
Table 4.38: Chi-square analysis of carnivore utilization and portion for the tibia.
Metapodials
Carnivore modification and utilization of the metapodials is difficult
because of the structural density (Table 4.39) of these elements. For the
metacarpal 12.15% of the elements exhibit some modification. Approximately
87.85% of those metacarpals have no utilization. The only types of carnivore
modification that are present on the metacarpal are furrowing, punctures,
pitting, and tooth scoring, all having less than five elements exhibiting the type
of mark. On the metatarsal, the results are almost identical. Only 12.10% of the
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elements exhibit some modification, with 87.90% having no utilization.
Chipping back, furrowing, punctures, pitting, and tooth scoring are all present
on the metatarsal; however, they are present in very low numbers, less than five.
Bone Mineral Densities (adapted from Kreutzer
1992 and Lyman 1994)
Element
Scan
Site
Density
Portion Value
Metacarpal
MC1
PR
.59
Metacarpal
MC2
PR
.63
Metacarpal
MC3
SH
.69
Metacarpal
MC4
DS
.60
Metacarpal
MC5
DS
.46
Metacarpal
MC6
DS
.53
Metatarsal
MR1
PR
.52
Metatarsal
MR2
PR
.59
Metatarsal
MR3
SH
.67
Metatarsal
MR4
DS
.51
Metatarsal
MR5
DS
.40
Metatarsal
MR6
DS
.48
Table 4.39: Bone mineral densities from bison, adapted from Kreutzer (1992) and Lyman (1994).
Astragalus, Calcaneus, and Third Phalanx
The remaining elements used for this thesis project are the astragalus,
calcaneus, and the third phalanx. The overall carnivore modification for these
elements is low because of their structural densities and sizes (Table 4.40).
Wolves and bears have been known to completely swallow a phalanx, therefore
it may be difficult to assess whether they were chewed on or swallowed whole,
except to look for pitting from stomach acids (Binford 1981). On the Kaplan–
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Hoover collection, there is no evidence in the elements used for this thesis that
these elements were swallowed whole. However, there are a number of scat pile
skeletal remains that have been used in another project that suggest some piles
were differentially digested and approximately 13 could have been from coyotes
and 21 could have been from wolves at Kaplan–Hoover (Gensler, Burnett, and
Todd 2002).
Bone Mineral Densities (adpated from
Kreutzer 1992 and Lyman 1994)
Element
Scan
Site
Density
Value
Astragalus
AS1
.72
Astragalus
AS2
.62
Astragalus
AS3
.60
Calcaneus
CA1
.46
Calcaneus
CA2
.80
Calcaneus
CA3
.49
Calcaneus
CA4
.66
Third Phalanx
P31
.32
Table 4.40: Bone mineral densities for bison, adapted from Kreutzer (1992) and Lyman (1994).
For the astragalus, 20% of the elements (n = 24) exhibit some evidence of
modification. Approximately 80% of the astragali have no modification and
approximately 15% have light utilization. The most common type of carnivore
modification on the elements is furrowing and punctures. The structural
densities of the astragalus (Table 4.40) would make it difficult to break through
the material; however, since the size of the element is much smaller than the
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other elements used for this project, possibly pulling or dragging of hind limbs
are reasons for punctures to show up on the astragalus.
The calcaneus is a very irregular shaped skeletal element therefore
modification is slightly less common than on the astragalus with 18.4% of the
calcanei (n = 21) exhibiting modification. Of the 18.4%, approximately 13.2% of
those elements have light utilization. The most prominent type of modification
found on the calcaneus is punctures, again, possibly related to the pulling of the
hind limbs by non-human scavengers.
Finally, the last element to discuss is the third phalanx. This element has
far more modification than the previous two elements with 32.5% of the elements
(n = 93) exhibiting modification. Approximately 22% of the phalanges have light
modification with punctures being the most prominent type of modification.
Again, this may be related to a pulling and dragging event with the bonebed in
which the non-human scavengers could have been pulling on the hind limbs to
move carcasses away from piles of bison to consume or in the case of some
behaviors exhibited by bears, possibly pulling carcasses away for concealed
feeding.
Generally, more carnivore modification and damage is present on low
density portions of skeletal elements. The most common types of modification
throughout the entire sample are furrowing and puncturing. Utilization on the
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skeletal elements was most prominent on the humerus, femur, and tibia. The
most modified element, the humerus has 93% of the elements exhibiting some
carnivore modification and approximately 65% of the humeri are heavily
modified. For the femur, approximately 41% of the elements are heavily
modified. Finally, for the tibia approximately 34% of the elements are heavily
modified.
The astragalus is an interesting element in terms of modification because
of the great amount of puncturing on these elements. Approximately 20% of the
astragali exhibit carnivore modification and the most common type are
punctures. The punctures on these elements are likely due to non-human
scavenger behaviors of dragging and pulling at carcasses. Considering the point
that canids fight over food at a kill and bears drag food for feeding concealment,
the behaviors causing punctures on the astragali could be related to either
scavenger.
The final chapter of this project discusses the results of data analysis in
general and the patterns that have emerged in terms of carnivore modification on
the skeletal elements and how those relate to animal behaviors. In addition,
future directions for this research are discussed.
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CHAPTER 5: CONCLUSIONS AND FUTURE
DIRECTIONS
The Kaplan–Hoover bison bonebed exemplifies human and non-human
interactions on the Great Plains during the Holocene. Predatory ecology of both
humans and other mammals is important on a variety of levels to understand
when human conditioning of other predators began and how those interactions
instituted the relationships that are present today. When coming to an end, it is
important to discuss where the research began. In order to accomplish this, the
questions that started the research will be evaluated given the results for data
collection and analysis.
Question: What are the herd characteristics at Kaplan–Hoover?
The Kaplan–Hoover assemblage is a nursery herd, dominated by female
and sub-adult bison with few males. Due to the fragmentary nature of the
assemblage, only 512 of the 1204 specimens (NISP) were able to be used in sex
analysis and of those, only 482 could be identified to a specific sex. Of the 482,
approximately 319 are female or sub-adults and 163 are male. From the first
publication on Kaplan–Hoover (Todd et al. 2001:137), it was documented that the
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kill occurred in September to October, therefore late summer to possibly early or
middle fall. This estimate was based on the mandibles of the collection, with
approximate age at death of 0.4–0.5 years (Todd et al. 2001:137). The minimum
number of individuals (MNI) for the collection is based on the cross tabulation of
sex and side for each element. Based on that analysis, there are approximately 55
individuals represented by the skeletal material incorporated into the sample
used in this thesis, based on the number of astragali skeletal elements present.
Questions: What types of intensity and carnivore modification is present on the
Kaplan–Hoover collection? Further, where is it located on the appendicular
skeleton and which specific elements exhibited more destruction than others?
It is apparent that the most modified skeletal elements in the collection are
the humerus and femur with approximately 94% of the elements exhibiting
carnivore modification. Following the humerus and femur, 73% of the tibia
exhibit carnivore modification. In terms of utilization, the heaviest utilized is the
humerus with 65.69% of these elements having almost the entire proximal end
being removed and in few instances having both the proximal and the distal end
completely removed (Figure 3.15). Other heavily utilized elements include the
femur with 41.07% and the tibia with 34.88% heavy destruction of either the
entire proximal or distal ends being removed.
To decipher any correlations between data sets, chi-square analyses were
completed. For the humerus there were a number of positive tests of
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significance. Sex and carnivore utilization, carnivore utilization and side, and
portion were all significant. Of particular interest is carnivore utilization and
portion, these results indicate that they are not independent of each other,
illustrating that the heavily utilized elements are coded as distal portions,
meaning the entire proximal end is removed. The same results were true of the
femur, where all three tests were significant. For the femur, when carnivore
utilization and portion were analyzed, there was significance and 49 of the
elements had portion codes of proximal and 45 had portion codes of shaft,
indicating that the elements were heavily destroyed on the proximal end.
The bone mineral densities of the humerus and femur are important to
consider when discussing these results (Table 4.9). The proximal portion of the
humerus is the least dense with values of 0.24 and 0.25. The femur values (Table
4.9) for the proximal end are 0.56 and 0.73, while the distal end values are 0.48
and 0.39. It is obvious why there is heavy destruction of the proximal end of the
humerus relative to the distal end; however, given the distal end of the femur
density values, why is there almost no destruction on the distal end of the
humerus where there are mineral density values of 0.38 and 0.48? In
comparison, it could be assumed from the femur density values that there is less
destruction on the proximal ends, in view of the high density values. In fact,
there is more destruction on the distal end than on the proximal end. This could
126
be explained by the shape of the distal end of both elements, possibly the shape
is too difficult to get a good grip on or put enough force on to cause major
destruction by a non-human scavenger. In addition, these results could be
related to the tightness of the ligaments between the humerus and radius-ulna or
the femur and tibia. Future research could help alleviate these questions about
the differences between the two skeletal elements.
The remaining elements not discussed previously, radius-ulna,
metapodials, astragalus, calcaneus, and third phalanx, are fairly dense; therefore,
a lot of force would be required to break into the bone. Further discussion of the
attritional behaviors of scavengers will help in the evaluation of these elements,
specifically the incidence of punctures on these elements.
Correlations: Skeletal Analysis and Ethological Information
The last question is important for a number of reasons. The previously
discussed questions and results are a good basis for understanding
paleoecological relationships in the Holocene between humans and non-human
predators. It is now imperative to determine to what degree the results from
those questions interact and how much can they provide in terms of interplays
between the multiple species, to do this, use of ethological literature is of
particular significance.
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Question: What relationships between prehistoric human hunters and nonhuman scavengers can be observed from the material remains at Kaplan–
Hoover?
It is obvious that canids were players in the taphonomy at Kaplan–
Hoover. The existence of a domesticated dog in the bonebed indicates that,
however, the modification types and degree of utilization influence the decision
that canids were present (Kinneer 2002). Given that the kill occurred in the late
summer to early fall, wolves may have been interested in scavenging these
remains. Bearing in mind denning season for wolves is in the late winter and
pups are born in the early spring when most birth-pulse ungulate species like
bison are born, wolves may be more active for scavenging to provide for
pregnant females (Packard 2003). In view of the fact that they do scavenge if the
opportunity exists (Mech 1970; Selva et al. 2005), wolves in the area of the kill
would have likely passed by, smelled the carcasses, or re-visited the kill later in
the winter or possibly early spring. In addition, coyotes likely visited the site for
foraging, specifically if any followed wolves that were feeding on the site given
that coyotes have been documented in Manitoba to visit every wolf kill and have
been recorded waiting approximately 100 m away from wolves feeding before
moving in (Paquet 1992). Another important point is that coyotes are known to
defecate on their kills to protect it from other predators (Wade and Bowns 1985).
As discussed earlier, at Kaplan–Hoover a number of scat bone piles were
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discovered and recorded (Gensler, et al. 2002). These piles were differentially
digested and approximately 13 could have been from coyotes and 21 could have
been from wolves based on size of element fragments indicating that both
wolves and coyotes likely used the site for feeding (Gensler, et al. 2002).
Wolves cause great destruction to the proximal ends of long bones,
specifically the humerus because of its grease content and low density and the
femora and tibia (Haynes 1980a:345-346). Correlating to that observation, over
93% of the Kaplan–Hoover humeri exhibit carnivore modification and over 65%
of the elements exhibit heavy utilization, where almost the entire proximal ends
are removed. Scavenging behaviors of canids suggest that they may have been
pulling and grappling with the limbs of the carcasses in order to remove some
meat from the area where the other wolves or coyotes were feeding (Mech
1970:185-186). In addition to pulling carcass parts away for feeding, wolves and
coyotes have been known to cache remains for later feeding, thus dragging parts
and likely inflicting puncture marks (Mech 1970; Paquet 1992; Peterson and
Ciucci 2003). In order to do this, it is assumed that the animals would grip the
end of a limb or the side of a limb to drag it away, therefore, leaving punctures
and possibly pitting marks on the skeletal elements. There are numerous
instances of punctures documented at Kaplan–Hoover (Figure 3.5), the skeletal
elements with the most punctures included: third phalanx, femur, astragalus,
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and calcaneus. Of these elements, the third phalanx and the femur have the
greatest incidences of puncturing. For the third phalanx, this may be associated
with the fact that the scavengers could have been dragging elements and for the
femur this could also have been possible. At Kaplan–Hoover, the butchering
patterns suggest that the limb bones were not highly utilized by humans, with
exception of some marrow extraction on the femora (Todd et al. 2001:138-139).
This could be essential as to why there are larger frequencies of punctures
present on the femora. After removal of marrow occurred on these elements, the
surrounding meat would have been opened up and available for scavengers to
have access without having to tear through hides and sinew. Further research on
whether scavengers would begin feeding in already opened areas of a carcass
could prove useful in comprehending this pattern. As discussed in Chapter 2,
wolves are the only species capable of opening up a bison carcass (Selva et al.
2005:1599); therefore, opened carcasses may have been more attractive to coyotes
and other smaller carnivores and scavengers.
Scavenging behaviors of bears are very different from the canids. First of
all, when canids feed on a carcass they scatter the remains in a wider area
directly around the carcasses than bears who can drag pieces away to far more
distant concealed areas (Wade and Bowns 1985). Given that bears do not scatter
remains and tend to drag food stuffs away for concealed feeding, several of the
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elements that show the most damage by bears may not be available for analysis
(Wade and Bowns 1985). Keeping in mind that these scavengers could have
made little damage to the site if they arrived first and removed anatomical pieces
and carried them away to concealed feeding locations, canid scavengers may
have covered up, through their gnawing any definitive marks made by bears
(Wade and Bowns 1985). Similar to coyotes, black bears are known to defecate
on carrion to protect it from other predators; therefore, scat bone may be a useful
indicator for illustrating the possible species present at bison kills (Wade and
Bowns 1985).
Bears have bunodont or low cusped molar dentition (Figure 2.8); because
of this, the marks they will leave on skeletal elements while scavenging will be
different than marks left by canids (Schwartz, Miller, and Haroldson 2003).
Canids, having the sharp high cusped carnassial cheek teeth leave sharp marks
(Figure 2.6). Documented bear marks (Figure 5.1) from Gary Haynes’s collection
at the University of Nevada Reno compared with almost identical marks at
Kaplan–Hoover (Figure 5.2) suggest that it is probable that bears were using the
bison pile for sustenance. Unfortunately, any overlaps in distinct types of
modification would make it impossible to distinguish whether specific marks are
from high cusp molars or from bunodont molars (Haynes 1980). In addition, it
may be incredibly difficult to identify specific scavengers using a carcass because
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of the nature of modification and number of predators that use carrion
throughout the year for sustenance (Haynes 1980 and 1982).
Figure 5.1: Bear modification Bos taurus tibia from Dr. Haynes collection.
Figure 5.2: Probable bear modification on Bison bison from
Kaplan–Hoover (Element # F27-24-230).
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Implications for Conservation Research
First, when discussing conservation it is applicable to appreciate how
animals become conditioned by human behaviors. Inherently, there are multiple
important behavioral patterns associated with canids and ursids. When thinking
on the subject of wolf behaviors, it is essential to include information on their
knowledge of prey species behaviors. For example, modern bison in constrained
settings like parks utilize specific winter ranges consistently and rarely modify
those ranges; therefore, if this occurred prehistorically wolves have become
adept at overexploiting those niches by staying close to these ranges in the
winter months, in turn relying on a dependable prey resource with little energy
expenditure (Haynes 1982). If wolves are capable of remembering the ranges of
bison in the winter it is completely plausible that they could have remembered
the hunting patterns of human populations during the Holocene. Coyotes, as
previously stated, follow wolves to feeding sites and tend to rely on wolves for
carrion of larger prey animals (Paquet 1992:341). Further, both brown and black
bears have been conditioned to rely on humans for garbage as a continuous
source of nutrients throughout the year and during some time relied heavily on
ranchers for cattle carcass piles as well (Craighead, et al. 1995:334-338). With that
said, it is evident that humans had influence in the sustainability of wolves,
133
coyotes, and bears from the early to middle to late Holocene and still do today.
The evidence lies in the adaptations seen by these predators to increasing
encroachment of human habitats on their habitats. Therefore, the relationship
between humans hunting and killing large numbers of bison and the ability of
non-human scavengers to sustain themselves is influential to their sustainability.
Taking into consideration conservation decisions in national parks, natural areas,
and towns or cities it is necessary to reflect on the impacts that humans have had
on these scavengers while humans have been in North America.
The relationship between archaeology and the natural sciences has been
recognized for a long period (Wintemberg 1919). As far back as 1919,
Wintemberg discussed the applicability of archaeology to paleontology and the
understanding of what species were present in which regions. Further,
Wintemberg (1919:64), lists 4 important reasons why zoologists and
archaeologists should work together: (1) discovery of which animals are now
extinct, (2) differentiation between animals that became extinct before and after
the arrival of Europeans in North America, (3) location of animal populations in
the past and now in the present, and finally (4) whether animals that are found in
archaeological sites still remain in the same region today. Recently, Lyman and
Cannon (2004) edited “Zooarchaeology and Conservation Biology” a collection
of articles discussing the importance of zooarchaeological analysis toward
134
conservation issues in North America. Of particular importance for this thesis is
the eradication and reintroduction debate pertaining to wolves, coyotes, and
bears. Wintemberg (1919) discusses the usefulness of archaeology to know
where animal populations once occupied and how those populations have
shifted within regions through history. Given that archaeologists are able to
identify that these non-human scavengers have been on the landscape for tens of
thousands of years, policy makers could use this knowledge to understand the
demise to other species in regions with the eradication or reintroduction of large
predators. It has been noted the significant role wolves play in assisting in the
sustainability of rodents and other smaller predators through their kills and
skeletal remains that provide nutrients. While conservationists do consider the
impacts their decisions make on landscapes, wouldn’t they be more informed
and therefore highly respected if they took into consideration the past and the
knowledge that archaeologists can provide to their cause?
In that effort to inform those management decisions, it is also important to
consider the position that archaeologists are currently in to make information
available and push coalescence between the fields of archaeology, ecology,
biology, zoology, natural resource managers, and finally government policy
makers. Finally, understanding there has never been a “pristine" environment
and taking into consideration that humans have been a major player in the
135
conditioning of species is important. Researchers like Lyman and Cannon (2004)
are trying to bridge that gap between archaeology and natural sciences. This
publication is a collection of articles by zooarchaeologists who were informed by
the editors to address a conservation issue, describe and analyze the
zooarchaeological data related to that concern, and off recommendations for
possible solutions to the conservation issue (Lyman and Cannon 2004:3). With
this publication, the bridge has been formed lending credence to integration and
collaboration in the natural sciences and archaeology.
Future Directions
This thesis project, presents data collection methods, data analysis, and
results that illustrate how taphonomy and ethological studies used in
conjunction can improve the understanding of archaeological faunal
assemblages. This thesis has also created a starting point for further research
using interdisciplinary methods.
Several more questions were exposed from this research. First of all
actualistic studies in North America have been done on a number of species,
mostly, wolves, coyotes, and bears (Binford 1981; Burgett 1990; Haynes 1981). In
Africa, far more species have been studied using actualistic research, such as
hyenas, wild dogs, and a number of the large felids (Blumenschine 1986, 1989;
136
Egeland et al. 2004; Marean and Spencer 1991). However, in North America
specifically, more research is needed on the smaller mammals such as rodents,
mustelids, foxes, felids, and birds. The incorporation of ethological methods by
archaeologists would be of great importance as well. As the literature review
suggests, little research by ethologists is dedicated to understanding how nonhuman scavengers destroy carcasses, where they feed first, and how they
destroy, use, and move the skeletal remains. If ethological research began
collecting data on these characteristics of carcass and skeletal material feeding
patterns, then collaboration between archaeologists and ethologists would
advance the understanding of human and non-human scavenger interactions in
the past and present.
Domestication of canids is of interest at this point as well (Morey 1986,
1992, 1994; Walker and Frison 1982). At what point in the domestication process
does conditioning of the animal build reliance for survival from the
domesticator? Various examples in the ethological literature, specifically the
behavior of bears towards humans in North America, suggests that the
conditioning of bears to believe that humans can and will provide food likely
began when humans first reached this continent. This indicates it is possible that
instead of a human based decision to domesticate, hunters may have discovered
that leaving food sources available for scavenging would cause non-human
137
scavengers to rely on them for sustenance (Morey 1994). Morey (1994) suggests
that it may not have been a conscious decision by humans to domesticate
animals; that instead, humans may have noticed a need and if the animals were
already in some events following them for sustenance they may have stumbled
on the idea. Morey (1994) indicates that it is difficult to focus on humans as the
only factor involved in domestication and any discussion of the wolves side of
the interaction should be included in the understanding of the relationships
necessary to create dependence. This is especially important when taking into
consideration the demise of numerous canid species that were not domesticated,
specifically the eradication of wolves and the desire to eradicate coyotes.
Furthermore, research of domestication practices and the consequences of not
completely domesticating an animal are relevant to understanding why bears
have been conditioned by humans and how that conditioning influences the
conservation of the species today.
Of equal importance is the management of non-human scavengers and
predators on the landscape. As discussed in this thesis, the disparity between
the public, policy managers, and scientists is so broad helpful management
decisions are not being made. An interdisciplinary approach to management
through the use archaeology, biology, zoology, ethology, and wildlife research, is
138
important to any progress on conservation issues in the Great Plains and North
America.
Lyman and Cannon (2004) illustrate the usefulness of understanding
faunal assemblages and how human interactions with their environments to the
understanding of where humans have been in the past and what can be done in
the future. A continuation and focus in zooarchaeology to understanding
human-animal interactions is a necessary goal for the future of archaeological
research and this thesis project assisted in pushing that goal into the forefront of
current research.
Finally, as discussed in Chapter 4, density values of skeletal elements
were used as a proxy for understanding differential selection by non-human
scavengers. It would be more relevant and useful in future research to address
whether a difference exists between feeding selection based on density and
nutritional utility. To better understand how carnivores use carcass material it is
necessary to not limit the indices and measures used to analyze the faunal
sample discussed.
Nutritional utility is understood based on bone grease content (Brink
1997:259). Bone grease is one of the most dependable fat sources in mammalian
carcasses and may be useful for indicating element selection of specific element
portions by non-human scavengers (Brink 1997:259). Using bone density as a
139
proxy for nutritional utility is an accurate assessment based on the analysis
completed by Brink (1997:262, Table 1). Brink (1997) found that the proximal
humerus, proximal femur, and proximal tibia all have the highest percentages of
fat content, which Kreutzer (1992:278-281, Table 2) illustrated are among the least
dense portions of those elements. Brink further states, however, that there is
great variability in amount of grease within individual skeletal elements (Brink
1997:263). This illustrates that the high variability cannot provide a solid
foundation for understanding differential selection of skeletal elements based on
grease content. Therefore density, although it is correlated with bone grease, is
less variable within individual skeletal elements, which could possibly be a more
reliable indicator of preferential selection (Brink 1997:264-265 and 272).
In future research, discussing all aspects of skeletal element morphology
would be useful for ascertaining the suite of characteristics that may indicate if a
selection bias occurred. Faith and Behrensmeyer (2006:1726-1727) and Faith, et
al. (2007:2033) discuss how the intensity of non-human scavenging can be
indicated by the percentage of low density element portions versus high density
element portions. They state that if there is high competition for skeletal
elements the scavengers will not preferentially select portions to eat and if there
is low competition, then scavengers will choose low density high nutritional
utility elements for feeding (Faith and Behrensmeyer 2006:1726-1727; Faith, et al.
140
2007:2033). Understanding the differences between density and nutritional
utility could therefore assist in assessing the numbers of non-human scavengers
present at a site and identify more taphonomic information. This future research
can accompany discussion on scavenging behaviors of canids and ursids. For
example, canids feed in groups, which results in a higher competition setting,
and ursids feed in solitude, which is a low competition setting. It could then be
argued that canids would not be preferentially selecting elements for
consumption like ursids because of the level of competition as indicated by Faith
and Behrensmeyer (2006:1726-1727) and Faith, et al. (2007:2033). If an ethologist
collected data on how non-human scavengers destroy carcasses and skeletal
remains then archaeologists could better reconstruct the taphonomic signatures
in a faunal assemblage.
141
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APPENDIX A: FAUNMAP DATA
- adapted from: http://www.museum.state.il.us/research/faunmap/aboutfaunmap.html
Site Name
Badger House
Buick Campsite
Cedar Point
Dutch Creek
Fort Davey Crockett
Hall-Woodland Cave
Merino
Mesa Verde
Mesa Verde
Recon John Shelter
Roberts Buffalo Jump
Texas Creek Overlook
Wetherill Mesa
24CA287
Bootlegger Trail
County Line
24GF250
Antonsen
Antonsen
Ash Coulee
Big Lip
Birdtail Butte
Drake
Drake
Ellison's Rock
Ellison's Rock
False Cougar Cave
Hagen
Hoffer
Hoffer
Holmes Terrace
Kobold
Mangus
Montana Ice Cave
Morse Creek #1
Pictograph Cave
Pictograph Cave
Pictograph Cave
Red Rock Springs
Risley Bison Kill
Blacktail Cave
Site Number
1453
5EL1
5EL8
5JF463
5MF605
5JF9
5LG122
866
875
ST
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
5LR100
CO
5RB2435
CO
1644 CO
24CA287
MT
24TL1237
MT
24MO197
MT
24GF250
MT
24GA660
MT
24GA660
MT
MT
24CB75
MT
24BL1152
MT
24YL51
MT
24YL51
MT
24RB1020
MT
24RB1020
MT
24CB84
MT
24DW2
MT
24CH669
MT
24CH669
MT
24FR52
MT
24BH406
MT
24CB221
MT
KU-MT-41
MT
MT
24YL1
MT
24YL1
MT
24YL1
MT
24BE1230
MT
24LC1003
MT
24CL151
MT
COUNTY
Montezuma
Elbert
Elbert
Jefferson
Moffett
Jefferson
Logran
Montezuma
Montezuma
Pueblo
Larimer
Rio Blanco
Montezuma
Cascade
Toole
Missoula
Garfield
Gallatin
Gallatin
Prarie
Carbon
Blaine
Yellowstone
Yellowstone
Rosebud
Rosebud
Carbon
Dawson
Choteau
Choteau
Fergus
Big Horn
Carbon
Fergus
Granite
Yellowstone
Yellowstone
Yellowstone
Beaverhead
Lewis and Clark
Lewis and Clark
LAT
370700
392200
392200
393000
404500
394500
402200
370700
370700
383000
403700
393700
370700
473000
481500
465200
473000
453925
453925
464500
450000
480700
454500
454500
455200
455200
450700
470000
474500
474500
473700
451500
451500
464500
463700
453700
453700
453700
445200
472200
470500
LONG
1083000
1035200
1035200
1050700
1084500
1050700
1031500
1083000
1083000
1044500
1051500
1075200
1083000
1111500
1111500
1134500
1064500
1110950
1110950
1051500
1081500
1090000
1083700
1083700
1063700
1063700
1081500
1043700
1095200
1095200
1093700
1070000
1075200
1090000
1130700
1082200
1082200
1082200
1124500
1122200
1121700
FMAGE
LHOL
LHOL
LHOL
LHOL
HIHO
LHOL
LHOL
LHOL
LHOL
LHOL
HIHO
HIST
LHOL
HOLO
LHOL
LHOL
LHOL
HIHO
LHOL
HIHO
LHOL
LHOL
LHOL
HIHO
HIST
LHOL
LMHO
HOLO
HIHO
LHOL
HIHO
LHOL
LHOL
HOLO
HOLO
LMHO
LHOL
HIHO
LHOL
HIHO
MHOL
FMSP
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
FMSP1 FMSP2 FMSP3 FMSP4
CA fa
CA fa
CA la
CA fa
CA
CA
CAN
CA fa
CAN
UR am
CA la
UR ar
CAN
CA
UR am
CA la
CA la
CA lu
CA lu
CA
CA
CA fa
CA la
CA fa
CA lu
CA lu
CA la
CAN
CAN
CA lu
UR
CA lu
CA la
UR ar
UR ar
CA lu
155
Shield Trap
Sorenson
48AB301
48CA1391
48CA1729
48CA1751
48CA2227
48CA403
48CR4897
48LN74
48SH312
48SW998
48TE1090
48TE1102
48TE111
48TE114
48UT199
Austin Wash
Beehive
Bessie Bottom
Bottleneck Cave
Bottleneck Cave
Bottleneck Cave
Buffalo Creek
Bugas-Holding
Cache Hill
Castle Garden Access Road
Daughtery Cave
Dead Indian Creek
Deer Creek
Eagle Shelter
Espy-Cornwell
Gull Island
Horse Creek
John Gale
Lamar
Lamar
Maxon Ranch
McCleary
McKean
Piney Creek
River Bend
Rock Ranch Trading Post
Scoggin
Skull Point
Spring Creek Cave
Vore
Wardell Buffalo Trap
24CB91
24CB202
48AB301
48CA1391
48CA1729
48CA1751
48CA2227
48CA403
48CR4897
48LN74
48SH312
48SW998
48TE1090
48TE1102
48TE111
48TE114
48UT199
48UT390
48BH346
48UT1186
48BH206
48BH206
48BH206
48SH311
48PA563
48CR61
48FR1398
48WA302
48PA551
48BH18
48CR4001
48TE1067
48LA549
48CR303
48SW2590
48NA1152
48CK7
48JO312
48NA202
48GO123
48CR304
48LN317
48WA1
48CK302
48SU301
MT
MT
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
WY
Carbon
Carbon
Albany
Campbell
Campbell
Campbell
Campbell
Campbell
Carbon
Lincoln
Sheridan
Sweetwater
Teton
Teton
Teton
Teton
Uinta
Uinta
Big Horn
Uinta
Big Horn
Big Horn
Big Horn
Sheridan
Park
Carbon
Fremont
Washakie
Park
Big Horn
Big Horn
Carbon
Teton
Laramie
Carbon
Park
Park
Sweetwater
Natrona
Crook
Johnson
Natrona
Goshen
Carbon
Lincoln
Washakie
Crook
Sublette
450700
451500
421500
440000
433700
433700
433700
433700
420000
414500
444500
420700
435200
435200
435200
435200
413000
412200
441500
410100
445200
445200
445200
444500
443700
441500
425200
440700
443700
445200
445200
413000
435200
412200
414500
445200
445200
410700
425200
441500
443100
424500
420000
414500
413700
435200
443000
423000
1081500
1075200
1060000
1051500
1051500
1051500
1051500
1051500
1064500
1100000
1060700
1073000
1103700
1103700
1103700
1103700
1103700
1101500
1073000
1105200
1082400
1082400
1082400
1061500
1092200
1053000
1073000
1071700
1092200
1080000
1080700
1072200
1103700
1050000
1070700
1101500
1101500
1090700
1064500
1044500
1064700
1062200
1041500
1063000
1103000
1073000
1040700
1100000
MHOL
LHOL
HIST
LHOL
MHOL
HOLO
HOLO
LHOL
LMHO
HOLO
LHOL
LMHO
HIHO
LHOL
HIST
LHOL
MHOL
LHOL
LMHO
LHOL
LHOL
EHOL
HIHO
LHOL
HIHO
HIST
LHOL
HIHO
MHOL
LHOL
EHOL
LHOL
LHOL
MHOL
HIHO
HIHO
LHOL
LHOL
HIST
LMHO
HIHO
HIST
HIST
MHOL
HIHO
LHOL
HIST
LHOL
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
BI bi
CA
CA
UR ar
CAN
CAN
CA la
CA
CA
CA
CA
CA lu
CA lu
UR ar
CA la
UR am
CAN
CAN
CA
CA
CA
CA
CA fa
CA
UR am
CA la
CA la
CA lu
UR ar
156
APPENDIX B: CODING SYSTEM
Skeletal Element
ELE
Segment Codes
SEG
Humerus
HM
Complete
CO
Radius/Ulna
RDU
Posteromedial
PM
Radius
RD
Proximal
PR
Ulna
UL
Anteromedial
AM
Femur
FM
Distal
DS
Tibia
TA
Anterolateral
AL
Astragalus
AS
Lateral
LT
Metapodial
MP
Posterolateral
PL
Metacarpal
MC
Medial
ME
Metatarsal
MT
Cranial
CR
Calcaneous
CL
Caudal
CD
Third phalanx
PHT
Interior
IN
Dorsal
DR
Portion Codes
POR
Exterior
EX
Complete
CO
Ventral
VN
Proximal diaphysis
DPR
Edge
EG
Proximal
PR
Left
L
Diaphysis + fused distal epiphysis
DFD
Right
R
Proximal + < half the shaft
PRS
End
END
Diaphysis + fused proximal epiphysis
DFP
Condyle
CDL
Proximal + > half the shaft
PSH
Unidentified fragment
FR
Shaft
SH
Distal
DS
Side Codes
SD
Flake (< half circum of SH)
FK
Left
L
Distal + < half shaft
DSS
Right
R
Impact flake
IMK
Axial
A
Distal + > half shaft
DSH
Not sided
N
Condyle
CDL
Proximal epiphysis
PRE
PFUS and DFUS
Distal epiphysis
DSE
Unfused
0
Unidentified epiphysis
EP
partially fused
1
Diaphysis
DF
fused, but line visible
2
Distal diaphysis
DDS
fully fused
3
157
Head
HE
broken, can't tell
4
Trochlear notch
ANC
not applicable, no epiphyseal
5
Olecranon portion
OLC
Olecranon tuber
PRE
Breakage type
BR
Carnivore Utilization
CU
No breakage
N
Light
L
Dry break
D
Light/moderate
LM
Intermediate break
I
Moderate
M
Carnivore modification
C
Moderate/heavy
MH
Green
G
Heavy
H
Rodeal gnawing
R
None
N
Crushed
S
Indeterminate
I
Sex
SEX
F
Female/Subadult
Multiple breaks
M
Excavator breakage
E
Modification
MOD
M
Male
Rodent gnawing
R
N
Not Known
Carny mod present
P
I
Indeterminate
Carny mod absent
A
Landmarks
LM
Carnivore modification
C
Present
P
Puncture
PC
Absent
A
Furrow
FW
Half
H
Tooth scoring
TS
Chipping back
CB
Salivary polishing
SP
Crenellation
CT
Scooping out
SO
Pitting
PT
- adapted from Todd (1983 and 1987)
158
APPENDIX C: LANDMARK AND MEASUREMENT
DESCRIPTIONS AND CODES
Landmarks
Landmark Code Hill
HM
DT
Landmark Code Burke
HM
LM1
MEH
MT
OLC
PLF
TM
RD
PLF
RT
FM
ANF
SF
MTO
MO
TA
ACR
PLF
ANF
LM2
LM3
LM4
LM5
LM6
RD
LM1
LM2
FM
LM1
LM2
LM3
LM4
TA
LM1
LM2
LM3
Description
deltoid tuberosity
tubercle for attachment of medial
collateral ligament
major tuberosity
proximal olecranon fossa
posterolateral nutrient foramen
teres major tubercle
posterolateral nutrient foramen
radial tuberosity
anterior nutrient foramen
supracondyloid fossa
major trochanter
minor trochanter
anterior crest
posterolateral nutrient foramen
anterior nutrient foramen
*Adapted from Hill 1994 Master's Thesis pp. 169-170
159
Measurements
Measurement Code Todd
HM
Measurement Code Burke
HM
Description
HM1
M1
HM2
M2
greatest lateral length/osteometric board
greatest length from the head/osteometric
board
HM3
M3
HM4
M4
HM5
M5
HM6
M6
HM7
M7
HM8
M8
HM9
HM10
M9
M10
HM11
M11
HM12
M12
HM13
M13
least breadth of diaphysis/sliding calipers
greatest breadth of distal end/osteometric
board
breadth of the distal articular
surface/sliding calipers
least breadth of olecranon fossa/sliding
calipers
greatest depth of proximal
end/osteometric board
least depth of diaphysis/sliding calipers
greatest depth of distal end/sliding
calipers
greatest sagittal depth of head/spreading
calipers
greatest breadth of articular surface of the
head/spreading calipers
HM14
M14
least depth of distal end/sliding calipers
HM15
M15
HM16
M16
HM17
M17
depth of olecranon fossa/sliding calipers
least circumference of
diaphysis/measuring tape
greatest length of fragmentary
humerus/osteometric board
greatest medial length/osteometric board
greatest proximal breadth/osteometric
board
160
RD
RD1
RD2
RD
M1
M2
RD3
M3
RD4
M4
RD5
RD6
M5
M6
RD7
M7
RD8
M8
RD9
M9
RD10
M10
RD11
M11
RD12
M12
RD13
M13
RD14
M14
RD15
M15
RD16
M16
RDU1
UL
M17
UL
UL1
M1
UL2
M2
UL3
M3
UL4
M4
UL5
M5
UL6
UL7
M6
M7
UL8
M8
UL9
M9
UL10
M10
physiological length/spreading calipers
greatest length/osteometric board
greatest breadth of proximal
end/osteometric board
greatest breadth of proximal articular
surface/sliding calipers
least breadth of diaphysis/sliding calipers
least depth of diaphysis/sliding calipers
greatest breadth of distal end/osteometric
board
greatest breadth of distal articular
surface/sliding calipers
greatest depth of proximal end/sliding
calipers
greatest depth of proximal end lateral
margin/sliding calipers
greatest depth of distal end/osteometric
board
greatest breadth of articular surface for
radial carpal/sliding calipers
greatest length of diaphysis (only if
unfused)/osteometric board
greatest breadth of distal diaphysis (only if
unfused)/osteometric board
greatest depth of distal diaphysis (only if
unfused)/osteometric board
greatest length of fragmentary
radius/osteometric board
greatest length of radius-ulna/osteometric
board
greatest length of ulna/osteometric board
greatest height of "cavitas sigmoides
majors"/osteometric board
greatest length of olecranon/osteometric
board
greatest breadth of olecranon
tuberosity/osteometric board
greatest breadth of coronoid process of
ulna/sliding calipers
greatest depth of olecranon
tuberosity/osteometric board
least depth of olecranon/sliding calipers
least depth at anconeal process/sliding
calipers
depth of "cavitas sigmoides
majors"/sliding calipers
greatest depth of olecranon/osteometric
board
161
FM
FM
FM1
M1
FM2
M2
FM3
M3
FM4
M4
FM5
M5
FM6
M6
FM7
FM8
M7
M8
FM9
M9
FM10
M10
FM11
M11
FM12
M12
FM13
M13
FM14
M14
FM15
M15
FM16
M16
FM17
M17
FM18
M18
FM19
M19
FM20
M20
greatest length of femur/osteometric board
length of major trochanter to lateral
condyle/osteometric board
greatest length from the head/osteometric
board
length of diaphysis (only if
unfused)/osteometric board
greatest length of medial condyle/sliding
calipers
greatest length of lateral condyle/sliding
calipers
greatest breadth of proximal
end/osteometric board
greatest depth of head/sliding calipers
greatest breadth of proximal diaphysis
(only if unfused)/osteometric board
least breadth of diaphysis/sliding calipers
greatest breadth of distal diaphysis (only if
unfused)/osteometric board
greatest breadth of distal end/osteometric
board
greatest breadth of trochlea/sliding
calipers
least breadth of trochlea/sliding calipers
least breadth of intercondyloid
fossa/sliding calipers
greatest depth of proximal
epiphysis/osteometric board
least depth of diaphysis/sliding calipers
greatest depth of distal
epiphysis/osteometric board
greatest depth of medial trochlea/sliding
calipers
greatest length of fragmentary
femur/osteometric board
162
TA
TA1
TA2
TA
M1
M2
TA3
M3
TA4
M4
greatest length/osteometric board
medial length/osteometric board
greatest length of diaphysis/osteometric
board
greatest breadth of proximal
end/osteometric board
TA5
M5
greatest breadth of proximal diaphysis
(only if unfused)/osteometric board
TA6
M6
TA7
M7
TA8
M8
TA9
M9
TA10
M10
TA11
M11
TA12
M12
TA13
M13
TA14
M14
TA15
M15
TA16
M16
least breadth of diaphysis/sliding calipers
greatest breadth of distal end/osteometric
board
greatest breadth of distal diaphysis (only if
unfused)/osteometric board
least depth of diaphysis/sliding calipers
greatest depth of distal end/osteometric
board
least distance between intercondylar
tubercles/sliding calipers
greatest breadth of extensor sulcus/sliding
calipers
depth of extensor sulcus/sliding calipers
breadth of distal articular surface/sliding
calipers
depth of proximal end/osteometric board
greatest length of framentary
tibia/osteometric board
*Adapted from Todd 1987 Taphonomy of the Horner II Bone Bed pp. 121-124
163
Measurement Code Morlan Measurement Code Burke Description
AS
AS
DM
M1
medial depth/sliding calipers
Wp
M2
proximal width/sliding calipers
Wd
M3
distal width/sliding calipers
Ll
M4
lateral length/sliding calipers
Lm
M5
medial length/sliding calipers
Dl
M6
lateral depth/sliding calipers
*Adapted from Morlan 1991 Bison Carpal and Tarsal Measurements: Bulls versus Cows and Calves pp.223
Measurement Code Hill
CL
Measurement Code Burke
CL
Description
CL1
M1
greatest length medial view/osteometric
board
CL2
CL3
CL4
CL5
CL6
M2
M3
M4
M5
M6
greatest proximal width/sliding calipers
greatest proximal depth/sliding calipers
distal width/sliding calipers
distal depth/sliding calipers
length of talus facet/sliding calipers
CL7
M7
length of tarsal c+4 facet/sliding calipers
*Adapted from Morlan 1991 Bison Carpal and Tarsal Measurements: Bulls versus Cows and Calves pp. 223
Measurement Code Driesch
MP
Measurement Code Burke
MP
Gl
M1
Bp
M2
SD
M3
DD
M4
Bd
PHT
DLS
M5
PHT
M1
Ld
MBS
M2
M3
Description
greatest length/sliding calipers (Bedord
No.1)
greatest breadth proximal end/sliding
calipers (Bedord No.2)
smallest breadth diaphysis/sliding calipers
(Bedord No.10)
smallest depth diaphysis/sliding calipers
(Bedord No.9)
greatest breadth distal end/sliding calipers
(Bedord No.4)
diagonal length of sole/sliding calipers
length of dorsal surface/sliding calipers
middle breadth of sole/sliding calipers
*Adapted from Driesch 1976 pp. 92-93 and 101 and Bedord 1974
164
APPENDIX D: KAPLAN-HOOVER DATA
Humerus
BN #
F24-4-1609
F27-12-279
F27-12-301
F27-13-224
F27-13-314
F27-13-339
F27-13-386
F27-13-453
F27-17-330
F27-17-774
F27-17-898
F27-18-103
F27-18-125
F27-18-314
F27-18-344
F27-18-591
F27-19-277
F27-22-185
F27-22-268
F27-22-300
F27-23-181
F27-23-226
F27-23-227
F27-23-448
F27-23-547
F27-23-567
F27-23-837
F27-23-908
F27-23-969
F27-24-173
F27-24-241
F27-24-242
F27-24-272
F27-24-274
F27-24-334
F27-24-387
F27-24-543
F27-24-711
F27-24-828
SEX
N
N
M
M
N
N
N
N
N
N
N
M
N
M
F
F
M
F
N
F
M
F
N
N
N
F
F
N
N
F
F
M
F
F
M
M
N
F
N
ELE
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
POR
DSE
DSH
DSH
CO
DSH
DSH
DSH
PSH
PRE
PRE
FR
DSH
DSH
DSH
CO
DSH
CO
CO
DSH
DSH
DSH
DSH
SH
SH
SH
DSS
DSH
DF
DSE
DSH
DSH
DSH
DSH
DSH
DSH
CO
DSE
DSH
SH
SEG
CO
CO
CO
CO
CO
CO
CO
CD
CO
CO
END
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
MD
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
FR
SD PFUS DFUS BR1
N
4
3D
L
4
1D
L
4
3D
R
3
3G
L
4
3G
R
4
3D
L
4
3D
L
3
3D
N
0
4G
N
0
4G
R
3
3D
R
4
3G
L
4
3D
R
4
3G
R
3
3D
L
4
3D
L
3
3G
L
4
3D
R
4
3D
R
4
3D
R
4
3G
L
4
3G
R
4
4G
R
4
3D
L
4
3D
L
4
3D
R
4
3D
R
4
4D
R
4
3D
L
4
3G
R
4
3D
R
4
3G
R
4
3G
R
4
3G
R
4
3G
R
4
3D
R
4
3G
L
4
3D
N
4
4D
BR2
G
N
G
N
N
G
G
G
N
N
G
N
G
N
N
G
N
G
G
G
N
N
N
G
G
G
G
G
G
N
G
N
N
N
N
G
N
G
G
MOD
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
A
C1
CB
CT
CT
FW
CT
CB
CB
FW
TS
TS
TS
CT
CT
CT
FW
CB
FW
PT
CT
SP
CB
PC
CT
CT
CB
TS
SO
CB
TS
PT
PT
PT
CT
CT
SO
FW
TS
CT
N
C2
PT
PT
PT
TS
PT
TS
PT
N
FW
FW
FW
PT
PT
PT
PC
PT
TS
TS
N
N
FW
CT
N
FW
PT
N
CT
TS
FW
CB
CT
CT
TS
PT
CT
PC
FW
PT
N
C3
TS
TS
N
N
N
FW
N
N
PC
N
N
N
N
N
TS
SP
CB
FW
N
N
PT
PT
N
PT
SP
N
N
N
N
SP
N
N
PT
PC
PT
TS
PT
FW
N
CU
I
H
H
L
H
H
H
L
I
I
I
H
H
H
L
H
LM
H
H
I
H
H
H
H
H
L
H
H
I
H
H
H
H
H
H
H
H
H
I
LM1
A
H
H
P
H
H
A
A
A
A
A
P
H
P
P
H
P
H
A
H
P
H
A
H
A
A
A
A
A
H
P
P
P
H
H
A
A
H
A
LM2
A
P
P
P
P
P
P
A
A
A
A
P
P
P
P
A
P
P
P
P
P
P
A
A
A
P
P
A
A
P
P
P
P
P
P
P
A
P
A
LM3
A
A
A
P
A
A
A
H
A
A
A
A
A
A
P
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
LM4
A
P
P
P
P
P
A
A
A
A
A
P
P
P
P
P
P
P
P
P
P
P
A
A
A
H
H
A
A
P
P
P
P
P
P
P
A
P
A
LM5
A
P
P
P
P
P
A
P
A
A
A
P
P
P
P
P
P
P
P
A
P
P
P
A
A
P
A
P
A
P
P
P
P
P
P
P
A
P
A
LM6
A
A
P
P
P
P
P
P
A
A
P
P
P
P
P
P
P
P
P
P
P
P
A
P
P
A
P
H
A
P
P
P
P
P
P
P
A
P
A
165
F27-24-870
F27-25-149
F27-25-151
F27-25-154
F27-25-54
F27-25-8
F27-9-207
F28-12-170
F28-12-203
F28-12-270
F28-12-298
F28-12-299
F2812-332
F28-13-210
F28-13-275
F28-13-316
F28-13-378
F28-13-423
F28-13-432
F28-2-168
F28-2-339
F28-2-345
F28-3-212
F28-3-213
F28-3-388
F28-3-765
F28-3-77
F28-4-248
F28-4-428
F28-4-571
F28-4-641
F28-4-643
F28-4-743
F28-5-37
F28-7-109
F28-7-14
F28-7-424
F28-7-564
F28-8-247
F28-8-291
F28-8-330
F28-8-408
F28-8-474
F28-8-504
F28-8-932
F28-8-995
F28-9-120
F28-9-170
F28-9-215
F28-9-276
F28-9-320
F28-9-417
F28-9-420
F28-9-454
M
N
N
M
N
F
N
N
M
F
F
F
N
F
N
N
F
N
F
F
F
F
F
M
F
F
M
F
N
F
N
N
F
F
F
N
N
F
M
N
M
F
M
F
N
F
F
F
F
F
N
F
F
N
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
HM
DSH
DSS
DF
CO
SH
DSH
PRE
DSH
DSH
CO
DSH
CO
CO
DSH
PRE
DSH
DSH
PRE
DSH
DSH
DSH
DSH
DSH
DSH
DSH
DSH
DSH
DSH
DSH
DSH
PRE
DSH
DSH
DSS
DSH
PRS
DSH
DSH
DSH
DSH
DSH
CO
DSH
DSH
DPR
DSS
DSH
DSH
DSH
DSS
PRE
DSH
CO
DSH
CO
CO
CD
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
AL
CO
CO
CO
PR
CO
CO
CO
CO
CO
CO
CO
CO
CD
CO
CO
CO
CO
CO
CO
CO
CO
CO
R
L
N
R
R
R
N
L
L
R
R
L
L
R
N
R
R
N
L
R
L
L
L
L
R
L
R
R
L
L
N
L
L
L
R
R
R
R
R
L
R
L
R
R
R
R
L
L
R
R
N
L
L
R
4
4
3
2
4
4
0
4
4
4
4
2
3
4
4
4
4
0
4
4
4
4
4
4
4
4
4
4
4
4
0
4
4
4
4
3
4
4
4
4
4
3
4
4
4
4
4
4
4
4
0
4
3
4
3
3
3
3
4
3
3
3
3
3
3
3
3
3
4
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
4
3
3
3
3
4
3
3
3
3
3
3
3
3
4
3
3
3
3
3
4
3
3
2
D
D
D
D
G
G
G
D
D
D
N
D
D
D
G
D
D
G
D
G
D
D
D
G
D
D
G
D
G
D
G
D
D
G
D
D
D
D
D
G
D
D
D
D
D
D
G
D
D
G
G
G
D
D
G
G
G
G
N
N
N
G
G
G
N
G
G
G
N
G
G
N
G
N
G
G
G
N
G
G
N
G
N
G
N
N
G
N
G
G
G
G
G
N
G
G
G
G
G
N
N
G
G
N
N
N
G
G
P
P
P
P
P
P
P
P
P
P
A
P
P
P
P
P
P
P
A
P
P
P
P
P
P
P
A
P
P
P
P
A
P
P
P
P
P
P
P
P
P
P
P
P
P
A
P
P
P
P
P
P
P
P
CT
TS
PT
FW
CT
CT
PC
CB
CB
TS
N
FW
CT
CB
FW
CT
TS
PC
N
PT
SO
CT
CT
CT
SO
CB
N
PT
CT
CT
PC
N
PT
SO
SP
TS
PT
CT
CT
PT
CB
TS
PT
CB
PT
N
CB
CT
CB
CT
TS
SO
FW
TS
PT
FW
FW
PC
PT
PT
FW
PT
PT
PC
N
TS
PC
PT
TS
PT
PC
FW
N
CT
CT
PT
PT
PT
CT
SP
N
CT
PT
N
TS
N
SP
CT
PT
CB
CB
PT
PT
CT
SP
FW
FW
SP
SP
N
TS
PT
SP
PT
FW
CT
CB
PT
N
N
N
TS
N
FW
TS
N
FW
FW
N
PC
FW
FW
N
N
N
N
N
N
N
TS
N
N
PT
PT
N
PC
N
N
FW
N
N
PT
N
FW
N
N
FW
N
PT
PC
CB
PT
N
N
FW
FW
PT
N
PC
PT
N
CT
H
I
I
MH
H
H
I
H
H
H
N
L
LM
H
I
H
L
I
N
H
H
H
H
H
H
H
N
H
H
H
I
N
H
H
H
I
H
H
H
H
H
MH
H
H
I
N
H
H
H
H
I
H
M
H
P
A
P
P
A
P
A
A
P
P
P
P
P
P
A
A
A
A
A
H
H
P
H
H
P
P
A
H
H
P
A
A
A
H
A
A
P
P
P
H
H
P
A
P
A
A
P
A
H
A
A
A
P
A
P
A
A
P
A
P
A
A
P
P
P
P
P
P
A
P
P
A
P
P
P
P
P
P
P
P
P
P
P
P
A
A
P
P
P
A
P
P
P
P
P
P
P
P
A
P
P
P
P
P
A
P
P
P
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
P
P
A
P
A
P
A
P
P
P
P
P
P
P
A
P
H
A
P
P
P
P
P
P
P
P
P
P
P
P
A
A
P
P
P
A
P
P
P
P
P
P
P
P
A
P
P
P
P
P
A
P
P
P
A
A
P
P
P
P
A
P
A
P
P
P
P
P
A
A
A
A
P
P
P
P
P
P
P
A
P
P
P
P
A
P
P
P
P
A
A
P
P
P
P
P
P
P
A
A
A
P
P
P
A
P
P
P
P
A
P
P
A
P
A
P
P
P
P
P
P
P
A
P
A
A
P
P
P
P
P
P
P
P
P
P
P
P
A
A
P
P
P
A
P
A
P
P
P
P
P
P
H
A
P
P
P
P
A
P
P
P
166
Radius-Ulna
BN #
NONUM3
NONUM2
NONUM
G28-3-22
G27-24-3
F28-9-416
F28-9-344
F28-9-253
F28-9-237
F28-9-237
F28-9-209
F28-9-209
F28-9-179
F28-8-967
F28-8-887
F28-8-739
F28-8-661
F28-8-512
F28-8-472
F28-8-333
F28-8-333
F28-8-214
F28-7-486
F28-7-372
F28-6-123
F28-4-858
F28-4-747
F28-4-575
F28-4-573
F28-4-572
F28-4-561
F28-4-301
F28-4-232
F28-4-230
F28-4-1617
F28-4-148
F28-3-669
F28-3-669
F28-3-416
F28-3-387
F28-3-386
F28-3-376
F28-3-369
F28-3-258
F28-3-242
F28-3-237
F28-3-186
F28-3-129
F28-3-128
SEX
N
M
F
N
N
F
M
F
F
F
F
F
N
N
F
M
N
F
F
M
M
M
F
N
F
F
N
M
F
F
F
N
F
M
N
N
F
F
N
N
F
N
N
N
F
N
N
N
F
ELE
UL
RD
RD
RD
RD
RDU
RDU
RDU
RD
UL
RD
UL
RD
RD
RD
RDU
RDU
RD
RD
RD
UL
RDU
RDU
UL
RDU
RDU
RD
RDU
RDU
RDU
RD
UL
RD
RD
UL
UL
RD
UL
UL
UL
RD
UL
RD
UL
RD
UL
RD
UL
RD
POR
OLC
PRS
CO
DSE
DSS
CO
CO
CO
CO
ANC
CO
CO
SH
DSE
CO
CO
DSS
PSH
CO
CO
CO
CO
CO
OLC
CO
PSH
CO
PSH
CO
CO
CO
OLC
DFP
DSE
OLC
OLC
PSH
OLC
OLC
OLC
CO
ANC
CO
OLC
CO
CO
CO
OLC
CO
SEG
CO
CO
CO
CO
DS
CO
CO
CO
CO
CO
CO
CO
CO
ME
CO
CO
CO
CO
CO
CO
CD
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
DS
CO
CO
CO
CR
CO
CO
CO
CO
CO
SD PFUS DFUS BR1
L
3
4D
L
3
4D
L
3
3D
L
4
0D
R
4
2D
L
3
3D
L
3
3D
R
3
3D
R
3
3D
R
4
4D
R
3
3N
R
4
4G
N
4
4D
R
4
0D
R
3
3D
L
4
3D
L
4
2D
R
3
4D
L
3
3D
R
3
3N
R
3
3D
R
3
3N
R
3
3D
R
4
4D
R
3
3N
L
3
4D
R
3
2D
L
3
4D
R
3
3D
L
3
3D
R
3
3D
R
0
4G
L
3
0N
L
4
0D
L
3
4D
R
0
4D
L
3
4D
L
4
4G
L
0
4D
R
4
4D
R
3
3D
L
4
4G
L
3
2D
L
4
4G
R
3
3D
L
4
3D
R
3
2D
L
4
4G
L
3
3D
BR2
N
N
N
N
N
G
G
G
N
G
N
N
G
N
N
N
G
N
N
N
N
N
G
G
N
G
N
G
G
G
N
D
N
G
N
G
N
D
G
N
N
D
G
N
G
G
G
D
N
MOD
A
A
A
A
A
P
P
P
A
P
A
P
P
A
A
A
P
A
A
A
A
A
P
P
A
P
A
P
P
P
A
P
A
P
A
P
A
P
P
A
A
P
P
P
P
P
P
P
A
C1
N
N
N
N
N
FW
FW
PT
N
CB
N
PT
PT
N
N
N
CB
N
N
N
N
N
FW
PT
N
PC
N
FW
FW
FW
N
PC
N
FW
N
FW
N
PT
FW
N
N
PC
TS
PC
FW
FW
FW
FW
N
C2
N
N
N
N
N
N
N
CB
N
FW
N
FW
CB
N
N
N
SP
N
N
N
N
N
N
FW
N
N
N
N
CB
PT
N
FW
N
N
N
N
N
FW
PC
N
N
FW
FW
PT
N
N
TS
CB
N
C3
N
N
N
N
N
N
N
SP
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
SP
N
N
N
N
N
TS
N
N
N
N
N
N
N
CB
N
N
N
N
PC
CB
N
N
CT
N
N
CU
N
N
N
N
N
L
L
L
N
LM
N
L
H
N
N
N
L
N
N
N
N
N
L
L
N
L
N
L
L
L
N
L
N
L
N
L
N
LM
L
N
N
M
L
M
L
LM
H
L
N
LM1
NA
A
P
A
A
P
P
P
A
NA
P
NA
A
A
P
P
A
P
P
P
NA
P
P
NA
P
P
A
P
P
P
P
NA
P
A
NA
NA
P
NA
NA
NA
P
NA
P
NA
P
NA
P
NA
P
LM2
NA
P
P
A
A
P
P
P
P
NA
P
NA
A
A
P
P
A
P
P
P
NA
P
P
NA
P
P
A
P
P
P
P
NA
P
A
NA
NA
P
NA
NA
NA
A
NA
P
NA
P
NA
P
NA
P
LM3
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM4
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM5
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM6
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
167
F28-2-349
F28-2-311
F28-2-304
F28-13-320
F28-13-295
F28-13-165
F28-13-105
F28-12-451
F28-12-355
F28-12-342
F28-12-300
F28-12-300
F28-12-227
F28-12-163
F27-9-208
F27-9-165
F27-9-152
F27-9-147
F27-25-47
F27-24-882
F27-24-833
F27-24-699
F27-24-544
F27-24-473
F27-24-329
F27-24-312
F27-24-312
F27-24-275
F27-24-275
F27-24-273
F27-24-273
F27-24-240
F27-24-203
F27-23-866
F27-23-860
F27-23-812
F27-23-762
F27-23-680
F27-23-579
F27-23-449
F27-23-419
F27-23-367
F27-23-358
F27-23-329
F27-23-260
F27-23-217
F27-23-196
F27-23-196
F27-23-1000
F27-22-375
F
F
M
N
F
M
M
N
F
N
N
N
M
N
N
N
N
F
M
N
N
N
N
N
M
N
N
N
N
F
F
N
M
N
N
F
F
F
N
N
F
N
N
M
F
N
F
F
N
N
RDU
RD
RD
RD
RD
RD
RD
RDU
RD
RD
RD
UL
RDU
RD
UL
UL
RD
RD
RDU
RD
RD
RD
RD
RD
RDU
RD
UL
UL
UL
RD
UL
UL
RDU
RD
RD
RD
RD
RD
RD
UL
RD
RDU
RDU
RD
RD
RD
RD
UL
RD
UL
CO
PSH
CO
DSE
CO
DFP
DSE
CO
DSH
PSH
CO
CO
CO
DSS
CO
CO
DF
DFP
CO
CO
DSS
DSE
PRS
DSS
CO
CO
ANC
BL
OLC
CO
OLC
OLC
CO
PRS
DSE
PRS
DSE
DSS
DSE
OLC
CO
DSS
DSS
PSH
CO
PSH
CO
ANC
PSH
OLC
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
ME
DS
CO
DS
CO
CO
CO
CO
CO
CO
DS
CO
ME
CO
CO
CO
CO
FR
CO
CO
CR
CO
CO
CO
DS
CO
CO
DS
CO
CO
CO
CO
DS
CO
CO
CO
CO
CO
CO
DS
L
R
L
R
L
R
R
R
L
R
L
L
R
R
L
L
L
R
R
L
L
R
R
R
R
L
L
N
L
L
R
R
R
R
L
L
L
R
R
R
L
L
L
R
R
R
L
L
L
L
3
3
3
4
3
3
4
3
4
3
3
4
4
4
4
0
0
3
3
3
4
4
3
4
3
3
4
4
4
3
4
4
3
3
4
3
4
4
4
0
3
4
4
3
3
2
3
4
3
4
3
4
3
0
3
0
0
3
3
4
3
3
4
3
3
4
0
0
3
3
3
0
4
3
3
3
4
4
4
3
4
4
2
4
0
4
0
3
3
4
3
3
3
4
3
4
3
4
4
4
D
D
D
D
D
D
N
G
D
D
D
D
G
D
G
G
D
D
D
D
D
D
D
D
D
D
D
D
G
D
D
D
G
D
D
D
N
D
G
D
D
D
D
D
D
D
D
G
D
D
G
N
N
G
N
N
N
D
N
N
N
N
D
G
N
N
G
N
G
G
N
N
N
N
G
N
G
N
D
N
G
G
N
G
N
G
N
N
N
G
N
G
N
N
G
G
N
D
N
G
P
A
A
P
A
A
A
P
A
A
A
A
P
P
P
A
P
A
P
P
A
A
A
A
P
A
P
A
P
A
P
P
P
P
A
P
A
A
P
P
A
P
A
A
P
P
A
P
A
P
FW
N
N
FW
N
N
N
FW
N
N
N
N
FW
TS
FW
N
FW
N
FW
FW
N
N
N
N
FW
N
PT
N
FW
N
CB
PT
FW
FW
N
PC
N
N
PC
PC
N
FW
N
N
PT
FW
N
PT
N
FW
N
N
N
TS
N
N
N
PT
N
N
N
N
TS
FW
PC
N
PC
N
N
N
N
N
N
N
PT
N
CB
N
PC
N
PT
CB
PC
N
N
N
N
N
TS
FW
N
TS
N
N
SP
PC
N
CB
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
PT
N
N
N
N
N
N
N
N
N
SP
N
FW
N
PT
N
FW
FW
N
N
N
N
N
N
N
N
N
SP
N
N
CB
CB
N
N
N
N
L
N
N
M
N
N
N
L
N
N
N
N
L
L
LM
N
H
N
L
L
N
N
N
N
L
N
M
N
LM
N
LM
LM
L
L
N
L
N
N
L
L
N
L
N
N
L
H
N
M
N
LM
P
P
A
A
P
P
A
P
A
P
A
NA
P
A
NA
NA
P
P
P
A
A
A
A
A
P
A
NA
NA
NA
P
NA
NA
P
P
A
A
A
A
A
NA
P
A
A
P
P
P
P
NA
A
NA
P
P
A
A
P
P
A
P
A
P
P
NA
P
A
NA
NA
A
P
P
P
A
A
P
A
P
P
NA
NA
NA
P
NA
NA
P
P
A
P
A
A
A
NA
P
A
A
P
P
P
P
NA
A
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
168
F27-22-273
F27-19-368
F27-19-319
F27-19-263
F27-18-971
F27-18-267
F27-18-126
F27-18-126
F27-17-949
F27-17-879
F27-17-831
F27-17-780
F27-17-685
F27-17-585
F27-17-469
F27-17-427
F27-17-1513
F27-13-449
F27-13-412
F27-13-175
F27-12-319
F27-12-175
E28-16-11
E27-6-16
E27-5-37
E27-16-51
E27-15-98
E27-15-79
N
M
F
N
N
N
M
M
N
N
N
N
F
N
M
M
N
N
N
N
F
N
N
N
N
F
F
F
RD
RDU
RD
RD
RD
UL
RD
UL
RD
RD
RD
UL
RD
UL
RD
RDU
RDU
RD
RD
RD
RDU
UL
RD
RD
UL
RD
RD
RD
CO
DSS
CO
PRS
DSS
OLC
CO
CO
PRS
DDS
DSE
OLC
CO
OLC
DSE
CO
PRS
DS
DSE
PSH
CO
CO
PRS
CO
OLC
DFP
CO
DFP
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CR
CO
CO
CO
CO
LT
AM
CO
CO
CO
DS
LT
CO
CO
CO
CO
CO
R
L
L
L
R
R
L
L
R
L
L
R
L
L
R
L
R
L
L
L
L
L
L
L
R
R
L
R
3
4
3
3
4
3
3
4
3
4
4
4
3
3
4
3
3
4
4
3
3
4
3
3
0
3
3
3
3
3
3
4
3
4
2
4
4
0
0
4
3
4
0
3
4
3
0
4
0
4
4
3
4
0
3
0
D
D
N
D
D
D
D
D
D
G
G
D
N
D
D
G
D
D
D
D
D
G
D
D
D
D
D
N
N
N
N
G
N
N
N
G
G
D
N
G
N
N
G
D
G
N
G
N
G
D
N
G
N
N
G
N
A
A
A
P
A
A
A
P
P
P
P
P
A
A
P
P
A
A
P
A
P
P
A
P
A
A
P
A
N
N
N
FW
N
N
N
FW
FW
FW
FW
CB
N
N
FW
FW
N
N
FW
N
FW
FW
N
FW
N
N
FW
N
N
N
N
PT
N
N
N
PT
TS
N
N
PT
N
N
N
N
N
N
N
N
SP
CB
N
PC
N
N
PC
N
N
N
N
SP
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
TS
SP
N
N
N
N
N
N
N
N
N
H
N
N
N
L
L
H
L
L
N
N
L
L
N
N
L
N
L
LM
N
L
N
N
L
N
P
A
P
A
A
NA
P
NA
P
A
A
NA
P
NA
A
P
P
A
A
P
P
NA
A
P
NA
A
P
P
P
A
P
P
A
NA
P
NA
P
A
A
NA
P
NA
A
P
A
A
A
P
P
NA
A
P
NA
P
P
P
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
C3
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
FW
FW
N
N
N
N
N
CU
N
N
N
N
N
N
N
L
N
N
N
N
N
N
N
LM
LM
N
N
N
N
N
LM1
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM2
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM3
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM4
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM5
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM6
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
Metacarpal
BN #
F28-9-415
F28-9-393
F28-9-357
F28-9-269
F28-9-233
F28-9-1001
F28-8-872
F28-8-844
F28-8-684
F28-8-614
F28-8-547
F28-8-521
F28-8-464
F28-8-379
F28-8-375
F28-8-174
F28-7-453
F28-7-177
F28-5-87
F28-5-36
F28-5-23
F28-4-832
SEX
F
N
F
F
N
N
F
N
M
M
F
F
N
M
F
N
N
F
F
N
M
N
ELE
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
POR
CO
CO
CO
CO
PSH
DSS
CO
DFP
CO
CO
CO
CO
DFP
CO
CO
DFP
DFP
CO
CO
PSH
CO
DSS
SEG
CO
CO
CO
CO
AL
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
SD
L
L
R
R
R
L
R
R
L
L
R
R
L
R
R
R
R
R
L
L
R
N
PFUS DFUS BR1
NA
3D
NA
3D
NA
3N
NA
3N
NA
4D
NA
3D
NA
3N
NA
0C
NA
3D
NA
3D
NA
3N
NA
3D
NA
0D
NA
3N
NA
3D
NA
0C
NA
0C
NA
3N
NA
3N
NA
4D
NA
3D
NA
3D
BR2
N
N
N
N
N
N
N
D
N
N
N
N
N
N
N
D
D
N
N
N
N
N
MOD
A
A
A
A
A
A
A
P
A
A
A
A
A
A
A
P
P
A
A
A
A
A
C1
N
N
N
N
N
N
N
PC
N
N
N
N
N
N
N
PC
PC
N
N
N
N
N
C2
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
TS
TS
N
N
N
N
N
169
F28-4-533
F28-4-470
F28-4-273
F28-4-226
F28-4-1665
F28-4-1593
F28-3-563
F28-3-540
F28-3-515
F28-3-438
F28-3-172
F28-3-154
F28-2-298
F28-13-289
F28-13-285
F28-13-276
F28-13-252
F28-12-66
F28-12-316
F28-12-180
F27-9-225
F27-9-119
F27-25-37
F27-25-118
F27-24-811
F27-24-701
F27-24-601
F27-24-570
F27-24-563
F27-24-430
F27-24-423
F27-24-330
F27-24-315
F27-24-281
F27-24-276
F27-24-245
F27-23-549
F27-23-479
F27-23-433
F27-23-282
F27-23-280
F27-23-262
F27-23-200
F27-23-197
F27-23-151
F27-22-253
F27-19-380
F27-19-332
F27-18-792
F27-18-742
F27-18-667
F27-18-597
F27-18-555
F27-18-44
F27-18-295
F27-18-140
F27-18-127
F
M
M
F
N
N
N
M
N
N
F
F
N
F
N
N
N
M
M
N
N
M
F
F
N
N
M
N
N
N
M
M
N
N
M
M
N
N
N
N
M
F
F
F
M
M
N
F
N
N
N
N
F
F
F
F
M
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
CO
CO
CO
CO
PSH
DSH
CO
CO
DSS
PSH
CO
CO
CO
CO
DFP
DFP
PSH
CO
CO
CO
DSS
CO
CO
CO
CO
PSH
CO
DFP
DSS
CO
CO
CO
PSH
CO
CO
CO
CO
DSS
CO
CO
CO
CO
CO
CO
CO
CO
PSH
CO
DFP
DFP
CO
PSH
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
R
L
R
R
R
L
R
R
N
L
L
R
L
L
R
L
R
L
L
L
R
R
L
R
L
R
R
R
R
R
R
R
L
L
R
L
R
L
R
R
L
L
L
R
L
R
R
L
L
R
R
R
L
R
L
R
L
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
3
3
3
3
4
3
3
3
2
4
3
3
3
3
0
0
4
3
3
3
3
3
3
3
3
4
3
0
3
3
3
3
4
3
2
3
3
3
3
0
3
3
2
3
3
3
4
3
0
0
3
4
3
2
3
3
3
N
N
D
D
D
D
C
D
D
D
D
D
D
D
D
D
D
D
N
N
G
N
N
D
D
D
D
D
D
D
D
D
C
D
N
D
C
G
D
C
N
D
D
D
N
N
D
D
C
D
D
D
D
D
D
D
N
N
N
N
N
N
N
D
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
D
N
N
N
D
N
N
D
N
N
N
N
N
N
N
N
D
N
N
N
N
N
N
N
N
A
A
A
A
A
A
P
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
P
A
A
A
P
A
A
P
A
A
A
A
A
A
A
A
P
A
A
A
A
A
A
A
A
N
N
N
N
N
N
PC
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
TS
N
N
N
PC
N
N
PC
N
N
N
N
N
N
N
N
FW
N
N
N
N
N
N
N
N
N
N
N
N
N
N
FW
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
FW
N
N
N
TS
N
N
FW
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
LM
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
LM
N
N
N
L
N
N
L
N
N
N
N
N
N
N
N
L
N
N
N
N
N
N
N
N
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
170
F27-17-964
F27-17-886
F27-17-835
F27-17-817
F27-17-694
F27-17-541
F27-17-433
F27-17-404
F27-17-362
F27-17-329
F27-17-328
F27-17-1505
F27-17-1501
F27-14-114
F27-13-433
F27-13-358
F27-13-115
F27-12-321
F27-12-298
E28-15-58
E27-5-67
E27-5-44
E27-5-10
E27-15-189
E27-15-182
E27-15-178
E27-15-129
E27-15-123
M
N
F
N
F
F
M
F
F
N
N
N
N
F
N
N
N
F
M
N
N
F
M
F
F
F
F
N
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
MC
CO
PRS
CO
DFP
CO
CO
CO
CO
CO
DFP
CO
CO
CO
CO
CO
DFP
CO
CO
CO
CO
DFP
CO
CO
CO
CO
CO
CO
DFP
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
DS
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
L
L
L
L
R
R
R
R
L
R
L
L
R
L
L
R
L
L
L
R
L
R
L
L
R
R
L
L
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
3
4
3
0
3
3
3
3
3
0
3
2
3
3
3
0
2
3
3
3
0
0
3
3
3
3
3
0
N
D
N
C
N
N
C
N
N
D
D
C
D
N
D
D
D
N
N
D
C
N
D
N
D
N
N
C
N
N
N
N
N
N
N
N
N
N
N
D
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
D
A
A
A
P
A
A
P
A
A
A
A
P
A
A
A
A
A
A
A
A
P
A
A
A
A
A
A
P
N
N
N
FW
N
N
PT
N
N
N
N
TS
N
N
N
N
N
N
N
N
PC
N
N
N
N
N
N
PC
N
N
N
N
N
N
N
N
N
N
N
PC
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
LM
N
N
L
N
N
N
N
LM
N
N
N
N
N
N
N
N
L
N
N
N
N
N
N
L
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
C2
PC
FW
N
FW
FW
N
PC
TS
N
SP
PC
N
SP
PC
N
FW
PC
FW
N
FW
PT
SP
FW
PC
C3
N
N
N
PT
N
N
N
FW
N
PT
FW
N
N
TS
N
TS
N
N
N
N
SP
PT
PC
PT
CU
LM
M
L
H
H
L
LM
M
LM
H
H
N
H
L
L
M
LM
L
LM
MH
H
H
M
LM
LM1
A
A
A
P
A
P
A
A
A
P
A
A
A
P
A
A
P
A
P
A
A
P
P
P
LM2
A
A
A
P
A
A
A
P
H
P
P
A
P
P
A
P
A
A
P
A
P
P
P
P
LM3
A
A
A
A
A
P
A
A
A
A
A
A
A
P
A
A
A
A
A
A
A
A
A
A
LM4
A
A
A
A
A
A
A
P
A
P
P
A
A
A
A
P
P
P
H
A
A
P
P
A
LM5
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM6
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
Femur
BN #
F28-9-406
F28-9-405
F28-9-305
F28-9-304
F28-8-983
F28-8-941
F28-8-816
F28-8-721
F28-8-624
F28-8-440
F28-8-383
F28-8-351
F28-8-329
F28-8-317
F28-8-142
F28-7-580
F28-7-548
F28-7-545
F28-7-470
F28-7-344
F28-7-31
F28-5-58
F28-5-15
F28-4-692
SEX
N
N
N
F
N
N
N
N
N
N
M
N
M
N
N
F
N
N
N
N
F
F
F
M
ELE
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
POR
PR
PRE
PRE
DPR
DSE
PRS
DS
CO
CO
SH
PSH
PRE
SH
CO
PRE
CO
DPR
PRS
CO
PSH
SH
PSH
CO
CO
SEG
ME
ME
ME
CO
ME
CO
ME
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
PR
CO
CO
CO
CO
CO
CO
SD PFUS DFUS BR1
R
3
4G
N
0
4G
L
0
4G
L
0
4D
L
4
0G
R
3
4G
R
4
3G
L
3
3G
R
3
3G
L
4
4G
L
3
4G
N
0
4D
R
4
4D
L
2
2G
N
0
4G
L
2
3G
R
0
4G
L
2
4G
R
2
4G
L
2
4G
L
4
4G
R
3
4G
L
3
3D
R
3
3G
BR2
N
N
N
G
N
D
N
D
D
N
D
N
G
D
N
D
D
D
D
D
D
N
G
D
MOD
P
P
P
P
P
P
P
P
P
P
P
A
P
P
P
P
P
P
P
P
P
P
P
P
C1
FW
PC
FW
CT
PC
PC
FW
PC
FW
CB
CB
N
CT
FW
FW
PC
FW
PC
FW
PC
CB
CB
PT
FW
171
F28-4-639
F28-4-589
F28-4-452
F28-4-421
F28-4-377
F28-4-245
F28-4-1578
F28-3-78
F28-3-76
F28-3-746
F28-3-737
F28-3-689
F28-3-57
F28-3-549
F28-3-533
F28-3-415
F28-3-397
F28-3-330
F28-3-286
F28-3-246
F28-3-226
F28-3-162
F28-3-143
F28-2-391
F28-2-375
F28-2-313
F28-2-258
F28-13-476
F28-13-462
F28-13-211
F28-12-460
F28-12-426
F28-12-324
F28-12-311
F28-12-248
F28-12-199
F28-12-185
F27-25-39
F27-25-38
F27-25-36
F27-25-34
F27-25-195
F27-25-144
F27-24-739
F27-24-696
F27-24-602
F27-24-569
F27-24-537
F27-24-530
F27-24-278
F27-23-849
F27-23-565
F27-23-427
F27-23-361
N
N
N
F
N
N
N
N
N
F
N
N
N
N
M
M
N
M
M
F
M
F
F
N
N
N
N
N
N
N
N
N
N
F
M
F
F
F
N
N
N
N
N
F
N
F
M
N
N
M
N
N
N
N
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
PR
PRE
PRS
SH
CO
PSH
SH
PRE
PSH
SH
SH
SH
DSS
DS
DPR
PSH
DSS
DSH
PSH
PSH
PSH
SH
PSH
PRE
PSH
DPR
PRS
CO
PRE
DSH
CO
DS
CO
SH
SH
DSH
PSH
CO
PRS
DS
DSS
PRE
CDL
PSH
SH
DF
PSH
PR
PRE
CO
DSE
PR
SH
PR
ME
ME
CO
CO
CO
CO
CO
ME
CO
CO
CD
CO
CO
ME
CO
CO
CO
CO
CO
CO
CO
CO
CO
ME
CO
CO
CO
CO
ME
CO
CO
ME
CO
CO
CO
CO
CO
CO
PR
ME
DS
CO
ME
CO
CO
CO
CO
ME
HE
CO
LT
ME
CO
ME
N
N
L
R
L
L
R
N
L
R
N
R
R
L
L
L
L
R
L
L
R
R
R
N
R
R
R
R
N
R
R
N
L
R
L
L
R
L
R
N
R
N
R
R
R
L
R
N
R
R
L
N
L
N
3
0
0
4
3
2
0
0
3
4
4
4
4
4
0
3
4
4
3
3
2
4
3
0
3
0
3
3
0
4
3
4
0
4
4
4
3
3
4
4
4
0
4
3
4
0
3
3
2
3
4
4
4
2
4
4
4
4
3
4
4
4
4
4
4
4
3
3
4
4
3
3
4
4
4
4
4
4
3
4
4
3
4
3
4
3
0
4
4
3
4
3
4
3
4
4
4
4
4
4
4
4
4
3
0
4
4
4
G
G
D
G
G
D
G
G
G
G
G
G
G
G
D
G
G
G
G
D
G
G
G
N
G
G
D
G
G
D
G
G
G
G
G
D
G
G
G
G
G
G
D
G
G
G
G
G
G
D
D
G
G
G
N
N
G
D
D
G
D
N
D
D
D
D
N
D
G
D
D
D
N
G
D
D
D
N
D
D
G
D
N
G
D
N
D
N
D
G
D
N
N
N
N
D
N
D
N
N
D
D
D
G
G
N
D
D
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
A
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
A
P
P
P
P
P
P
P
P
P
P
P
FW
FW
FW
CT
TS
FW
PT
PC
PT
CB
CB
CB
TS
FW
PC
CT
FW
PC
FW
PC
PC
CB
CB
N
PC
FW
FW
PC
FW
CB
FW
FW
PC
CB
CB
TS
FW
PC
SP
FW
FW
PC
N
PC
CB
CB
PC
FW
PC
FW
PC
FW
CT
FW
PC
PC
N
SO
FW
CB
CB
FW
FW
SP
SP
SP
FW
PC
PT
FW
PC
FW
CT
FW
FW
PT
PC
N
FW
PC
N
N
N
SP
TS
PC
FW
SP
SP
FW
PC
CB
FW
N
SP
FW
N
CB
SP
PC
CB
PC
FW
N
FW
PC
TS
PC
N
N
N
N
N
SP
SP
N
SP
N
N
PT
PC
N
CB
PC
SP
CT
PC
SP
CB
SP
FW
N
TS
N
N
N
N
PC
PC
N
N
PT
TS
PC
N
PT
CB
N
CB
N
N
FW
FW
PT
FW
N
N
N
TS
N
N
N
LM
LM
L
H
LM
H
H
LM
H
H
LM
H
M
LM
H
H
H
H
H
H
H
H
H
N
H
MH
M
L
LM
H
L
LM
M
H
H
H
MH
MH
H
LM
H
L
N
H
H
H
H
LM
L
L
H
LM
H
LM
A
A
P
A
P
A
P
A
A
A
A
P
A
A
A
A
A
A
A
A
P
P
P
A
A
A
A
P
A
A
P
A
A
A
A
A
P
P
A
A
A
A
A
P
P
P
P
A
A
P
A
A
A
A
A
A
A
P
P
P
P
A
P
P
A
H
P
A
P
P
P
P
P
P
A
P
P
A
P
A
A
H
A
P
P
A
P
P
P
P
P
P
A
A
P
A
A
A
A
P
P
A
A
P
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
P
A
A
A
A
A
A
A
A
A
H
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
P
A
P
A
A
A
P
A
A
A
A
A
P
P
A
A
P
A
A
A
P
A
P
P
P
A
A
P
P
A
A
H
P
A
P
P
P
A
A
A
A
P
A
P
P
A
A
P
A
A
A
A
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
172
F27-23-352
F27-23-342
F27-23-245
F27-23-192
F27-23-168
F27-23-160
F27-23-155
F27-23-149
F27-22-331
F27-22-276
F27-22-125
F27-22-124
F27-18-440
F27-18-439
F27-18-394
F27-18-387
F27-17-985
F27-17-799
F27-17-646
F27-17-563
F27-17-526
F27-17-458
F27-14-61
F27-14-180
F27-14-177
F27-14-176
F27-13-209
E28-25-100
E28-16-84
E27-6-11
E27-15-62
E27-15-58
E27-15-16
E27-15-116
N
F
F
M
N
M
N
N
N
N
F
N
N
N
N
N
N
F
F
N
N
F
F
N
N
N
F
N
N
N
M
N
F
F
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
FM
PSH
PSH
PSH
CO
PSH
PSH
PRS
DSS
PR
PRE
CO
PRE
PR
DSH
DSE
PRE
DDS
SH
CO
PRS
DSH
DSH
CO
DSH
DS
SH
CO
DS
DS
DSS
SH
DSH
SH
CO
CO
CO
CO
CO
CO
CO
CO
CO
ME
LT
CO
ME
ME
CO
CO
ME
CO
CO
CO
CO
CO
CO
CO
CR
LT
CO
CO
DS
ME
CO
CO
CO
CO
CO
R
L
R
R
L
L
R
L
N
N
R
N
R
R
L
N
L
R
R
L
L
R
R
L
L
L
L
L
L
L
R
R
R
R
3
3
2
3
3
0
3
4
3
0
3
0
3
4
4
0
0
4
3
2
4
2
0
4
4
4
3
4
4
4
4
4
4
3
4
4
4
3
4
4
4
3
4
4
3
4
4
3
0
4
3
4
3
4
3
3
0
3
3
4
3
3
3
3
4
3
4
3
D
D
G
D
D
D
D
G
G
G
G
N
G
D
G
G
G
G
G
D
G
G
G
D
D
G
G
D
D
G
G
G
D
G
SEX
N
F
N
N
M
N
M
N
M
N
N
M
N
N
N
N
ELE
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
POR
DSH
CO
CO
DSH
CO
DSH
CO
DSS
CO
PRE
DFD
CO
DSH
PSH
DF
PSH
SEG
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
SD PFUS DFUS BR1
R
4
3C
L
3
3C
L
3
3C
L
4
3D
R
3
3C
R
4
3C
R
3
3C
L
4
3D
L
3
3C
R
0
4C
R
0
3C
R
3
3C
R
4
3C
L
3
4D
R
0
0C
R
3
4C
G
G
N
G
G
G
G
D
N
N
D
N
D
G
N
N
D
D
D
G
D
D
N
N
N
N
D
G
N
D
N
D
G
N
P
P
P
P
P
P
P
P
P
P
P
A
P
P
P
P
P
P
P
P
P
P
P
A
A
P
P
P
A
P
P
P
P
P
FW
PC
FW
FW
PC
TS
FW
FW
FW
FW
FW
N
FW
FW
TS
FW
PC
PT
FW
PC
FW
PC
CB
N
N
CB
PC
TS
N
FW
CT
PC
CB
FW
N
FW
CB
PC
FW
CB
N
PC
PC
N
PC
N
PC
PC
FW
N
FW
CB
PC
FW
PC
FW
SP
N
N
PT
FW
FW
N
N
SP
N
PC
PC
N
PT
N
N
N
FW
N
N
TS
N
CB
N
N
SP
PC
N
TS
SP
N
N
N
TS
FW
N
N
TS
N
PC
N
N
PT
N
SP
N
M
H
H
L
M
H
M
MH
LM
LM
MH
N
LM
MH
LM
LM
H
H
L
M
M
H
H
N
N
H
LM
H
N
M
H
LM
H
LM
A
P
A
A
A
P
A
A
A
A
P
A
A
P
A
A
A
A
P
A
A
P
P
P
A
A
P
A
A
A
P
A
P
A
A
P
P
P
A
P
A
P
A
A
P
A
A
A
A
A
A
P
P
A
P
P
P
A
A
P
P
A
A
P
P
P
P
P
A
A
A
P
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
P
P
P
P
P
P
P
A
A
A
P
A
A
H
A
A
A
A
P
A
A
H
P
A
A
A
P
A
A
A
A
P
P
P
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
BR2
D
N
D
N
N
D
N
N
N
N
N
N
N
N
G
D
MOD
P
P
P
A
P
P
P
A
P
P
P
P
P
A
P
P
C1
CB
FW
FW
N
PC
PT
PC
N
FW
FW
FW
FW
CB
N
PT
PC
C2
PT
PC
N
N
FW
CB
N
N
PC
N
PC
N
PT
N
N
FW
C3
TS
N
N
N
N
N
N
N
N
N
N
N
FW
N
N
N
CU
H
L
L
N
LM
H
L
N
LM
L
L
L
H
N
H
L
LM1
H
P
A
A
P
H
P
A
P
A
H
P
H
A
A
P
LM2
P
P
A
A
P
P
P
A
P
A
P
P
P
A
P
P
LM3
A
A
P
A
A
A
A
A
P
A
A
A
P
A
A
A
LM4
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM5
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM6
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
Tibia
BN #
F28-9-442
F28-9-407
F28-9-220
F28-8-883
F28-8-84
F28-8-733
F28-8-522
F28-8-44
F28-8-199
F28-8-193
F28-8-192
F28-8-106
F28-7-69
F28-7-612
F28-7-507
F28-7-368
173
F28-7-199
F28-5-125
F28-4-826
F28-4-812
F28-4-773
F28-4-671
F28-4-606
F28-4-587
F28-4-578
F28-4-570
F28-4-549
F28-4-433
F28-4-379
F28-4-250
F28-4-233
F28-4-204
F28-4-179
F28-4-177
F28-3-96
F28-3-87
F28-3-505
F28-3-459
F28-3-378
F28-3-322
F28-3-223
F28-3-218
F28-3-174
F28-3-150
F28-3-136
F28-3-135
F28-25-108
F28-2-355
F28-2-347
F28-2-302
F28-2-264
F28-13-533
F28-13-422
F28-13-352
F28-13-199
F28-12-450
F28-12-435
F28-12-348
F28-12-345
F28-12-247
F28-12-211
F28-12-208
F27-24-845
F27-24-700
F27-24-632
F27-24-589
F27-24-574
F27-24-389
F27-24-324
F27-24-311
F27-24-308
F27-24-247
N
N
N
F
N
N
N
N
N
M
N
F
N
M
N
N
N
F
N
N
N
N
N
N
N
N
N
F
M
F
N
N
N
N
F
N
M
N
N
N
N
N
N
N
F
M
N
N
N
N
N
N
N
N
F
N
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
PSH
DSE
CO
CO
DF
DSH
DSH
DSH
DSH
CO
DSH
CO
DSH
CO
DSH
DSH
DSH
CO
PRS
CO
PSH
DSH
DSH
CO
DSH
DSH
DSH
CO
CO
CO
SH
DSH
DSH
DSH
CO
DF
CO
DF
CO
DSH
DF
DSE
DF
DSH
CO
CO
CO
SH
SH
DSE
PRE
CO
DSH
CO
CO
DSH
CO
LT
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
PL
CO
CO
CO
CO
CO
CO
CO
L
R
L
R
L
R
L
L
R
R
L
L
L
L
L
L
L
R
R
R
L
R
R
L
R
R
R
L
R
R
N
R
L
L
R
R
L
L
L
L
R
R
R
R
L
L
R
N
N
L
R
R
R
R
R
L
3
4
3
3
0
4
4
4
4
3
4
3
4
2
4
4
4
3
3
3
3
4
4
2
4
4
3
3
3
3
4
4
4
4
3
4
3
0
2
4
0
4
0
4
3
3
3
4
4
4
0
3
4
2
3
4
4
0
3
3
0
3
3
3
3
3
3
3
3
3
3
3
3
3
4
3
4
3
3
3
3
3
3
3
3
3
4
3
3
3
3
0
3
0
3
3
0
0
0
3
3
3
3
4
4
0
4
3
3
3
3
3
C
D
D
C
C
C
C
D
C
D
C
C
C
C
C
C
C
C
C
C
C
D
C
C
C
C
D
C
C
C
D
C
C
C
C
C
C
C
C
C
C
D
C
C
C
C
D
C
D
D
D
C
C
C
C
C
D
N
N
N
D
D
D
N
D
N
D
D
D
N
D
D
N
N
G
D
D
N
D
D
N
D
N
N
N
N
N
D
D
N
D
D
D
N
D
N
D
N
N
D
D
N
N
D
N
N
N
D
N
D
D
N
P
A
A
P
P
P
P
A
P
A
P
P
P
P
P
P
P
P
P
P
P
A
P
P
P
P
A
P
P
P
A
P
P
P
P
P
P
P
P
P
P
A
P
P
P
P
A
P
A
A
A
P
P
P
P
P
PC
N
N
PC
CB
PT
PT
N
PT
N
PT
FW
CB
FW
PT
CB
CB
PC
FW
FW
PC
N
CB
FW
CB
CB
N
FW
FW
FW
N
CB
CB
CB
FW
CB
FW
CB
FW
PT
FW
N
CB
PT
FW
FW
N
FW
N
N
N
FW
CB
FW
PC
CB
FW
N
N
N
N
CB
CB
N
CB
N
CB
TS
PT
N
CT
PT
PT
N
N
N
N
N
PT
N
PT
PT
N
N
N
N
N
PT
FW
PT
N
PT
PC
PT
N
FW
N
N
SP
CB
N
TS
N
N
N
N
N
PC
PT
N
N
PT
N
N
N
N
N
N
SP
N
SP
N
N
N
SP
N
N
SP
N
N
N
N
N
N
N
N
TS
N
N
N
N
N
N
SP
SP
N
N
SP
N
SP
N
CB
N
N
N
N
N
PC
N
N
N
N
N
N
SP
N
N
SP
L
N
N
L
L
H
H
N
H
N
H
L
H
LM
H
H
H
L
LM
LM
L
N
H
LM
H
H
N
L
LM
L
N
H
H
H
L
H
L
H
L
H
H
N
H
H
L
L
N
H
N
N
N
LM
H
L
L
H
P
A
A
P
A
A
H
A
H
P
H
P
H
P
A
H
P
P
A
A
H
A
A
A
H
H
P
P
P
P
A
H
H
H
P
A
P
H
A
A
H
A
H
H
P
P
A
A
A
A
A
P
H
H
P
H
P
A
A
P
A
P
P
A
P
P
A
P
P
P
A
P
P
A
A
P
P
A
A
P
P
P
P
P
P
P
A
P
P
P
P
P
P
A
P
P
A
A
P
P
P
P
A
A
A
A
A
P
A
P
P
P
A
A
A
A
A
A
A
A
A
A
A
P
A
A
A
A
P
P
A
A
A
P
A
A
A
P
A
A
A
A
A
A
A
P
P
A
P
A
A
A
A
A
A
A
A
P
A
A
A
A
A
A
A
P
A
P
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
174
F27-24-244
F27-24-230
F27-24-215
F27-24-207
F27-24-206
F27-24-205
F27-23-965
F27-23-562
F27-23-513
F27-23-447
F27-23-437
F27-23-416
F27-23-353
F27-23-322
F27-23-319
F27-23-278
F27-23-235
F27-23-229
F27-23-159
F27-23-150
F27-23-112
F27-22-327
F27-22-284
F27-19-376
F27-19-340
F27-18-631
F27-18-301
F27-18-280
F27-18-262
F27-18-246
F27-18-245
F27-18-214
F27-18-124
F27-18-120
F27-18-119
F27-17-655
F27-17-577
F27-17-496
F27-17-473
F27-14-104
F27-13-494
F27-13-462
F27-13-445
F27-13-392
F27-13-123
F27-12-220
F27-12-164
E28-5-11
E28-25-54
E28-25-112
E28-16-8
E28-16-29
E27-5-34
E27-16-18
E27-15-55
E27-15-171
M
N
N
N
F
M
N
N
N
N
F
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
M
F
N
N
N
N
N
N
N
N
N
M
N
N
N
N
N
N
F
N
N
N
N
N
N
M
N
F
N
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
TA
CO
CO
CO
DSH
CO
CO
CO
DSH
DSH
DSH
CO
DSH
DSH
DSH
DSH
DSH
DSH
DF
CO
CO
PSH
CO
PRS
DSH
DSH
SH
DSH
CO
CO
CO
DSH
DSE
DSH
FK
DSH
CO
DSH
CO
CO
DSH
DF
PRE
PRS
DSH
DF
CO
DF
DSS
DSS
PRS
CO
DSE
CO
DF
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
AM
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
DS
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
L
L
L
R
L
R
L
L
L
L
L
R
L
R
R
R
L
R
R
R
R
N
R
L
L
N
L
R
L
R
R
L
L
R
R
L
L
R
R
R
L
L
R
R
N
L
R
R
L
L
N
L
L
R
R
R
3
3
3
4
3
3
3
4
4
4
3
4
3
4
4
4
4
4
2
2
3
4
4
4
4
4
4
2
3
3
4
4
4
4
4
3
4
2
3
4
0
0
3
4
0
3
0
4
4
3
3
4
2
0
3
3
3
3
3
3
3
3
3
2
3
3
3
0
3
3
3
3
3
4
3
3
4
3
3
3
3
4
3
3
3
3
3
0
3
4
3
3
2
3
3
3
0
4
4
3
0
3
0
3
3
4
3
0
3
0
3
3
C
C
C
C
D
C
C
C
C
D
C
G
D
C
C
D
D
C
C
D
C
C
C
D
C
C
C
C
C
D
C
D
C
G
C
D
D
C
C
C
C
C
C
D
D
N
C
G
D
C
D
N
D
C
C
C
N
D
D
N
N
N
D
D
D
N
D
D
N
D
D
N
N
G
D
G
G
D
G
N
D
G
D
D
N
N
N
N
N
N
N
N
N
D
D
D
D
D
D
N
G
N
N
N
N
D
N
N
N
G
N
D
P
P
P
P
A
P
P
P
P
A
P
A
A
P
P
A
A
P
P
A
P
P
P
A
P
P
P
P
P
A
P
A
P
A
P
A
A
P
P
P
P
P
P
A
A
A
P
A
A
P
A
A
A
P
P
P
FW
FW
FW
CB
N
FW
FW
CB
CB
N
FW
N
N
CB
CB
N
N
CB
FW
N
FW
PT
CB
N
CT
CB
PT
FW
FW
N
CB
N
CB
N
CB
N
N
FW
FW
PT
PT
FW
PC
N
N
N
CB
N
N
FW
N
N
N
CB
FW
FW
PC
PC
N
PT
N
N
PC
PT
PT
N
N
N
N
PT
PT
N
N
PT
PC
N
N
CB
PT
N
PT
PT
CB
PC
PC
N
FW
N
PT
N
FW
N
N
N
PC
CB
CB
N
FW
N
N
N
N
N
N
N
N
N
N
PT
PC
N
N
N
N
SP
N
N
N
TS
N
N
N
N
N
SP
FW
N
N
N
N
N
N
SP
N
N
N
N
SP
N
N
N
PT
N
FW
N
PT
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
LM
LM
L
H
N
L
LM
H
H
N
LM
N
N
H
H
N
N
H
LM
N
L
H
H
N
H
H
H
L
L
N
H
N
H
N
H
N
N
L
LM
H
H
L
L
N
N
N
H
N
N
L
N
N
N
H
L
LM
P
P
P
H
P
P
P
A
H
A
P
A
H
H
H
A
A
A
H
H
P
A
A
A
A
A
H
H
P
A
H
A
H
A
H
H
A
H
H
H
A
A
P
A
A
P
H
A
A
P
A
A
P
A
P
A
P
P
P
P
P
P
P
A
P
P
A
A
P
P
P
A
A
P
A
P
P
A
A
P
P
A
P
P
P
A
P
A
P
A
P
P
A
P
P
P
A
A
A
A
A
P
P
A
A
P
A
A
P
P
P
P
A
A
A
A
A
A
A
A
A
A
A
A
P
A
A
A
P
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
P
A
A
A
A
A
A
P
A
A
A
A
A
A
A
P
P
A
A
A
A
A
A
A
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
175
Metatarsal
BN #
F28-9-469
F28-9-460
F28-9-224
F28-9-141
F28-8-43
F28-8-227
F28-8-222
F28-8-205
F28-8-200
F28-7-99
F28-7-514
F28-7-495
F28-7-353
F28-7-32
F28-7-13
F28-5-89
F28-5-84
F28-5-25
F28-5-21
F28-5-106
F28-4-584
F28-4-574
F28-4-562
F28-4-55
F28-4-525
F28-4-399
F28-4-394
F28-4-227
F28-4-1631
F28-4-1543
F28-3-97
F28-3-492
F28-3-460
F28-3-430
F28-3-345
F28-3-341
F28-3-325
F28-3-290
F28-3-171
F28-3-163
F28-3-156
F28-3-114
F28-24-519
F28-2-408
F28-2-288
F28-13-436
F28-13-392
F28-12-96
F28-12-3083
F28-12-303
F28-12-252
F28-12-215
F27-9-193
F27-9-160
SEX
F
N
N
N
N
N
F
F
F
N
N
F
N
M
F
M
N
N
N
N
N
F
N
F
N
F
M
M
N
N
F
N
M
M
F
N
M
N
N
N
N
N
M
N
F
N
N
N
M
F
N
N
M
F
ELE
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
POR
CO
FK
CO
PRS
PRS
CO
CO
CO
CO
PSH
DFP
CO
CO
CO
CO
CO
PSH
CO
FK
CO
PRS
CO
DSS
CO
PSH
CO
CO
CO
CO
DSH
CO
CO
CO
CO
CO
DSH
CO
CO
DFP
PSH
DSH
CO
CO
DSH
CO
CO
DFP
DFP
CO
CO
CO
DS
CO
CO
SEG
CO
PM
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
AM
CO
AM
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
AM
CO
CO
CO
CO
CO
CO
CO
SD
R
N
L
R
R
L
R
R
R
L
R
L
N
L
L
R
R
R
N
R
L
L
R
L
L
R
L
L
R
L
L
L
R
L
R
L
R
R
L
R
R
L
R
R
L
L
L
L
L
L
L
L
L
L
PFUS DFUS BR1
NA
3C
NA
4G
NA
2D
NA
4G
NA
4D
NA
3D
NA
3N
NA
3D
NA
3D
NA
3D
NA
0D
NA
3D
NA
3D
NA
3D
NA
3D
NA
3D
NA
4D
NA
3D
NA
4G
NA
3D
NA
4D
NA
3N
NA
3C
NA
3D
NA
4D
NA
3D
NA
3N
NA
3D
NA
3D
NA
2D
NA
3D
NA
3D
NA
3D
NA
3D
NA
3D
NA
4C
NA
3N
NA
3D
NA
0D
NA
4D
NA
3G
NA
0D
NA
3D
NA
3D
NA
3N
NA
3D
NA
0D
NA
0D
NA
3D
NA
3D
NA
3D
NA
3D
NA
3N
NA
3D
BR2
D
D
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
D
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
MOD
P
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
P
A
A
A
A
A
A
A
A
A
A
A
A
P
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
C1
PC
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
PC
N
N
N
N
N
N
N
N
N
N
N
N
CB
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
C2
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
PT
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
C3
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
TS
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
CU
L
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
L
N
N
N
N
N
N
N
N
N
N
N
N
H
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
LM1
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM2
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM3
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM4
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM5
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM6
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
176
F27-25-99
F27-25-102
F27-24-684
F27-24-610
F27-24-600
F27-24-597
F27-24-481
F27-24-363
F27-24-357
F27-24-314
F27-24-255
F27-24-254
F27-23-967
F27-23-956
F27-23-943
F27-23-936
F27-23-907
F27-23-730
F27-23-675
F27-23-534
F27-23-483
F27-23-428
F27-23-393
F27-23-387
F27-23-336
F27-23-263
F27-23-263
F27-23-244
F27-23-238
F27-23-236
F27-23-230
F27-23-221
F27-23-188
F27-22-340
F27-22-289
F27-19-366
F27-19-363
F27-18-844
F27-18-781
F27-18-573
F27-18-438
F27-18-343
F27-18-243
F27-18-105
F27-18-104
F27-17-950
F27-17-758
F27-17-723
F27-17-679
F27-17-640
F27-17-578
F27-17-571
F27-17-543
F27-17-489
F27-17-476
F27-17-345
F27-17-280
N
N
N
N
M
N
N
M
N
F
M
N
N
F
N
N
N
F
N
N
N
N
N
N
N
F
F
N
N
M
F
F
M
N
F
M
M
N
N
F
N
F
N
N
M
N
N
N
M
M
F
M
N
N
F
F
F
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
CO
CO
DSH
DSH
CO
DSH
PRS
CO
PRS
CO
CO
CO
CO
CO
CO
CO
DDS
CO
PSH
PSH
PRS
CO
PSH
CO
CO
CO
CO
CO
PR
CO
CO
CO
CO
DFP
CO
CO
CO
PRS
CO
CO
DSH
CO
CO
DFP
CO
CO
PRS
CO
CO
CO
CO
CO
DFP
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
PR
CO
CO
CO
CO
CO
CO
CO
CO
CO
ME
CO
PR
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
DS
CO
CO
CO
PR
CO
CO
CO
CO
CO
DS
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
R
R
L
N
R
L
R
R
R
L
L
L
L
R
L
L
N
L
R
L
R
L
L
L
R
L
R
L
L
L
L
R
R
N
R
L
R
L
L
R
L
L
L
N
R
L
L
R
R
R
R
L
L
R
R
L
R
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
3
3
4
4
3
3
4
3
4
3
3
2
2
3
3
3
0
3
4
4
4
3
4
3
2
3
3
3
4
3
3
3
3
0
3
3
3
4
3
3
4
3
3
0
3
2
4
3
3
3
3
3
0
3
3
3
3
N
D
D
C
D
D
D
N
C
N
D
D
D
C
D
D
C
D
D
D
D
C
D
D
C
D
D
D
G
D
D
D
D
G
D
D
D
G
D
D
C
N
D
C
N
D
G
D
D
D
D
D
D
D
D
D
D
N
N
N
D
N
N
N
N
D
N
N
N
N
D
N
N
D
N
N
N
N
D
N
N
D
N
N
N
N
N
N
N
N
C
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
A
A
A
P
A
A
A
A
P
A
A
A
A
P
A
A
P
A
A
A
A
P
A
A
P
A
A
A
A
A
A
A
A
P
A
A
A
A
A
A
P
A
A
P
A
A
A
A
A
A
A
A
A
A
A
A
A
N
N
N
CB
N
N
N
N
FW
N
N
N
N
FW
N
N
PC
N
N
N
N
PC
N
N
FW
N
N
N
N
N
N
N
N
PC
N
N
N
N
N
N
CB
N
N
FW
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
PT
N
N
N
N
N
N
N
N
N
N
N
N
FW
N
N
N
N
FW
N
N
N
N
N
N
N
N
N
N
N
FW
N
N
N
N
N
N
PT
N
N
PC
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
TS
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
TS
N
N
PT
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
H
N
N
N
N
L
N
N
N
N
L
N
N
H
N
N
N
N
LM
N
N
L
N
N
N
N
N
N
N
N
H
N
N
N
N
N
N
H
N
N
H
N
N
N
N
N
N
N
N
N
N
N
N
N
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
177
F27-14-101
F27-13-466
F27-13-425
F27-13-384
F27-12-323
F27-12-221
F27-12-205
E28-25-76
E28-25-52
E28-25-42
E27-6-43
E27-16-20
E27-16-17
N
N
N
F
M
N
F
N
N
F
N
N
F
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
MT
CO
DFP
CO
CO
CO
DFP
CO
PSH
DSS
CO
DFP
DFP
CO
CO
CO
CO
CO
CO
CO
CO
CO
DS
CO
CO
CO
CO
L
L
L
R
L
L
R
L
L
R
R
L
R
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
3
0
3
3
3
0
3
4
3
3
0
0
3
C
D
D
D
D
C
N
D
D
D
C
N
D
D
N
N
N
N
D
N
N
N
N
N
N
N
P
A
A
A
A
P
A
A
A
A
P
A
A
FW
N
N
N
N
PC
N
N
N
N
PC
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
L
N
N
N
N
L
N
N
N
N
L
N
N
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
C1
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
FW
FW
N
N
FW
N
N
N
PC
N
N
PC
N
N
N
N
N
FW
N
N
C2
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
C3
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
CU
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
L
LM
N
N
L
N
N
N
L
N
N
L
N
N
N
N
N
L
N
N
LM1
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM2
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM3
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM4
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM5
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM6
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
Astragalus
BN #
NO NUMBER
G27-23-2
G27-20-01
F28-9-410
F28-9-222
F28-8-771
F28-8-737
F28-8-723
F28-8-47
F28-8-405
F28-8-300
F28-8-194
F28-8-107
F28-8-1007
F28-7-97
F28-7-43
F28-7-399
F28-7-398
F28-7-380
F28-5-69
F28-5-32
F28-5-19
F28-5-18
F28-5-155
F28-4-89
F28-4-88
F28-4-605
F28-4-586
F28-4-580
F28-4-550
F28-4-529
F28-4-435
F28-4-434
F28-4-382
F28-4-335
F28-4-219
F28-4-000
F28-3-580
F28-3-556
SEX
F
M
N
F
F
F
F
M
F
M
M
F
F
N
M
F
F
F
F
F
N
F
M
F
M
M
F
M
F
F
M
M
F
F
M
M
F
N
M
ELE
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
POR
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
SEG
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
SD
R
L
L
L
L
R
L
R
R
L
R
R
L
R
L
R
L
L
L
L
L
R
L
R
R
R
L
L
R
L
L
L
L
R
L
R
L
L
L
PFUS
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
DFUS
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
BR1
D
D
D
D
D
D
D
D
N
D
N
N
N
D
N
D
D
D
N
C
C
N
D
C
N
D
D
C
D
N
C
D
D
D
D
D
C
D
D
BR2
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
D
D
N
N
N
N
N
N
D
N
N
D
N
N
N
N
N
D
N
N
MOD
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
P
P
A
A
P
A
A
A
P
A
A
P
A
A
A
A
A
P
A
A
178
F28-3-537
F28-3-530
F28-3-463
F28-3-462
F28-3-461
F28-3-420
F28-3-329
F28-3-321
F28-3-316
F28-3-293
F28-3-256
F28-3-230
F28-3-225
F28-23-958
F28-18-303
F28-13-319
F28-13-228
F28-13-220
F28-12-346
F28-12-278
F28-12-212
F28-12-156
F27-73-411
F27-26-286
F27-24-865
F27-24-637
F27-24-507
F27-24-485
F27-24-344
F27-24-321
F27-24-253
F27-24-251
F27-23-935
F27-23-728
F27-23-585
F27-23-558
F27-23-514
F27-23-505
F27-23-463
F27-23-462
F27-23-368
F27-23-313
F27-23-308
F27-23-170
F27-22-339
F27-22-328
F27-22-325
F27-22-285
F27-19-515
F27-19-378
F27-19-377
F27-19-267
F27-18-845
F27-18-832
F27-18-576
F27-18-417
F27-18-368
F27-18-363
F27-18-287
F27-18-215
M
F
F
F
M
M
M
M
F
N
F
F
F
N
M
M
M
F
M
F
F
F
F
M
F
M
N
F
F
M
F
M
N
M
F
F
M
M
F
M
M
N
F
N
M
N
N
N
M
M
M
M
F
F
F
M
F
F
M
F
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
R
L
L
L
R
L
L
R
R
R
L
L
R
L
L
L
L
L
R
R
L
R
L
R
R
R
L
L
L
R
L
R
R
L
R
R
L
R
R
R
L
R
R
L
R
R
R
R
L
L
L
R
L
L
R
R
L
R
R
L
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
C
D
D
N
N
D
C
G
N
D
C
C
N
D
D
D
D
D
C
D
D
C
D
D
D
C
C
D
N
N
D
N
C
D
D
D
D
C
D
D
C
D
D
C
D
D
D
D
D
D
D
N
C
N
D
D
N
D
N
N
D
N
N
N
N
N
D
N
N
N
D
N
N
N
N
N
N
N
D
N
N
N
N
N
N
D
D
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
P
A
A
A
A
A
P
A
A
A
P
P
A
A
A
A
A
A
P
A
A
P
A
A
A
P
P
A
A
A
A
A
P
A
A
A
A
P
A
A
P
A
A
P
A
A
A
A
A
A
A
A
P
A
A
A
A
A
A
A
PC
N
N
N
N
N
PC
N
N
N
PC
PC
N
N
N
N
N
N
FW
N
N
PC
N
N
N
PC
PC
N
N
N
N
N
FW
N
N
N
N
TS
N
N
FW
N
N
FW
N
N
N
N
N
N
N
N
PC
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
FW
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
PC
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
L
N
N
N
N
N
L
N
N
N
L
L
N
N
N
N
N
N
L
N
N
L
N
N
N
L
LM
N
N
N
N
N
LM
N
N
N
N
L
N
N
L
N
N
LM
N
N
N
N
N
N
N
N
L
N
N
N
N
N
N
N
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
179
F27-17-880
F27-17-820
F27-17-643
F27-17-638
F27-17-594
F27-17-525
F27-17-494
F27-17-484
F27-17-1011
F27-14-106
F27-13-447
F27-13-388
F27-13-129
F27-12-334
F27-12-278
E27-6-56
E27-6-45
E27-6-25
E27-5-49
E27-5-31
E27-15-46
M
F
N
F
N
F
F
F
M
M
F
F
N
M
F
F
F
N
N
M
F
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
AS
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
L
R
L
L
R
L
R
R
L
R
R
R
L
R
L
L
L
R
L
L
R
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
ELE
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
POR
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
SEG
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
SD PFUS DFUS
N
3 NA
R
3 NA
L
3 NA
L
3 NA
L
3 NA
L
3 NA
R
0 NA
L
0 NA
R
3 NA
R
3 NA
R
3 NA
R
4 NA
L
2 NA
L
3 NA
R
3 NA
R
2 NA
R
0 NA
L
3 NA
R
2 NA
L
3 NA
R
3 NA
L
3 NA
R
3 NA
L
3 NA
L
3 NA
L
3 NA
R
3 NA
L
3 NA
N
3 NA
L
3 NA
L
3 NA
N
D
C
C
D
N
D
N
D
D
C
D
D
D
D
C
D
D
C
D
N
N
N
D
D
N
N
N
N
N
N
N
N
N
N
N
N
N
N
D
N
N
A
A
P
P
A
A
A
A
A
A
P
A
A
A
A
P
A
A
P
A
A
N
N
PC
FW
N
N
N
N
N
N
FW
N
N
N
N
PC
N
N
PC
N
N
N
N
FW
N
N
N
N
N
N
N
PT
N
N
N
N
FW
N
N
FW
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
LM
L
N
N
N
N
N
N
L
N
N
N
N
L
N
N
LM
N
N
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
BR1
C
D
D
D
N
D
N
D
D
D
N
D
D
N
D
D
D
C
D
D
D
D
D
D
N
D
N
N
D
D
D
BR2
D
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
D
N
N
N
N
N
N
N
N
N
N
N
N
N
MOD
P
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
P
A
A
A
A
A
A
A
A
A
A
A
A
A
C1
FW
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
PC
N
N
N
N
N
N
N
N
N
N
N
N
N
C2
SP
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
C3
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
CU
H
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
L
N
N
N
N
N
N
N
N
N
N
N
N
N
LM1
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM2
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM3
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM4
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM5
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM6
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
Calcaneus
BN #
NO NUMBER
G27-20-7
G27-20-5
F28-9-257
F28-9-230
F28-8-85
F28-8-64
F28-8-2001
F28-7-613
F28-7-499
F28-7-406
F28-73-45
F28-5-28
F28-5-17
F28-5-156
F28-4-87
F28-4-707
F28-4-604
F28-4-579
F28-4-530
F28-4-447
F28-4-432
F28-4-268
F28-4-249
F28-4-223
F28-3-775
F28-3-70
F28-3-396
F28-3-369
F28-3-362
F28-3-357
SEX
N
F
N
F
F
F
N
F
N
N
F
F
M
F
F
N
N
N
F
N
F
F
F
F
F
N
F
M
N
N
N
180
F28-3-319
F28-3-312
F28-3-244
F28-3-231
F28-3-180
F28-3-121
F28-2-402
F28-2-395
F28-2-316
F28-2-268
F28-2-212
F28-18-302
F28-13-539
F28-13-301
F28-13-226
F28-12-333
F28-12-273
F28-12-161
F27-9-136
F27-25-87
F27-25-30
F27-25-116
F27-24-866
F27-24-86
F27-24-820
F27-24-628
F27-24-626
F27-24-529
F27-24-408
F27-24-279
F27-24-260
F27-24-256
F27-24-243
F27-24-225
F27-24-214
F27-24-174
F27-23-993
F27-23-987
F27-23-641
F27-23-637
F27-23-582
F27-23-581
F27-23-572
F27-23-537
F27-23-504
F27-23-500
F27-23-493
F27-23-481
F27-23-420
F27-23-388
F27-23-375
F27-23-318
F27-23-316
F27-23-239
F27-23-223
F27-23-220
N
F
F
F
F
F
F
F
F
F
F
F
N
F
F
F
F
N
F
F
F
F
N
F
F
F
N
F
F
F
N
M
F
F
F
M
N
F
N
F
N
N
F
N
M
F
N
F
N
N
F
F
N
M
F
F
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CR
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
L
L
R
L
L
R
R
R
R
L
L
R
R
L
L
L
R
L
L
L
R
L
R
L
R
R
L
R
L
L
L
R
R
L
R
R
R
R
N
R
L
L
R
R
R
R
L
L
R
L
L
R
R
L
R
R
0
3
3
3
0
3
3
0
3
3
0
3
0
3
0
3
3
3
3
0
3
3
3
0
3
3
2
3
3
3
4
3
3
3
3
3
3
0
3
3
3
4
3
3
0
2
3
3
3
3
3
3
3
3
3
3
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
C
D
C
N
D
C
C
D
D
C
D
D
D
C
D
N
N
D
N
D
N
C
C
N
D
N
D
C
D
D
C
N
N
N
N
C
D
D
D
D
D
C
C
D
D
N
D
N
D
D
D
C
D
N
N
N
D
N
N
N
N
N
N
N
N
D
N
N
N
N
N
N
N
N
N
N
N
N
D
N
N
N
N
D
N
N
D
N
N
N
N
N
N
N
N
N
N
N
D
N
N
N
N
N
N
N
N
N
N
N
N
N
P
A
P
A
A
P
P
A
A
P
A
A
A
P
A
A
A
A
A
A
A
P
P
A
A
A
A
P
A
A
P
A
A
A
A
P
A
A
A
A
A
P
P
A
A
A
A
A
A
A
A
P
A
A
A
A
FW
N
PC
N
N
TS
PC
N
N
PC
N
N
N
PC
N
N
N
N
N
N
N
PC
TS
N
N
N
N
SP
N
N
FW
N
N
N
N
TS
N
N
N
N
N
SP
PC
N
N
N
N
N
N
N
N
PC
N
N
N
N
TS
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
FW
N
N
N
N
N
N
N
N
N
N
N
N
N
FW
N
N
N
N
N
N
N
N
N
N
N
N
N
N
PC
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
PT
N
N
N
N
N
N
N
N
N
N
N
N
N
N
M
N
L
N
N
L
L
N
N
L
N
N
N
L
N
N
N
N
N
N
N
L
L
N
N
N
N
LM
N
N
M
N
N
N
N
L
N
N
N
N
N
H
L
N
N
N
N
N
N
N
N
L
N
N
N
N
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
181
F27-23-189
F27-23-187
F27-23-185
F27-23-1001
F27-19-291
F27-18-848
F27-18-577
F27-18-435
F27-18-434
F27-18-285
F27-17-929
F27-17-620
F27-17-475
F27-17-282
F27-14-139
F27-14-110
F27-13-382
F27-13-267
F27-13-137
F27-12-300
F27-11-536
E28-5-18
E28-25-75
E28-16-171
E28-15-49
E28-15-32
E27-15-60
F
F
F
F
F
F
F
F
F
N
M
F
F
N
F
F
F
M
N
F
N
N
N
N
F
F
F
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CL
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CR
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
L
L
L
R
R
L
R
L
R
R
L
L
R
L
R
R
R
R
L
R
R
R
L
L
L
R
L
POR
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
SEG
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
SD
ME
ME
ME
LT
LT
ME
ME
ME
ME
LT
LT
ME
LT
ME
LT
LT
LT
LT
LT
LT
ME
ME
ME
ME
3
3
0
3
0
0
3
3
2
3
3
4
2
4
0
0
3
3
3
2
3
0
3
3
3
0
3
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
C
N
D
D
D
N
D
D
D
C
D
C
D
D
D
N
D
D
C
D
D
D
D
C
N
N
N
N
N
N
N
N
N
N
N
N
D
N
D
N
N
N
N
N
N
N
N
N
N
N
D
N
N
N
P
A
A
A
A
A
A
A
A
P
A
P
A
A
A
A
A
A
P
A
A
A
A
P
A
A
A
PC
N
N
N
N
N
N
N
N
PC
N
PC
N
N
N
N
N
N
FW
N
N
N
N
PC
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
L
N
N
N
N
N
N
N
N
L
N
L
N
N
N
N
N
N
LM
N
N
N
N
L
N
N
N
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
DFUS
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
BR1
C
D
C
D
N
C
D
D
N
C
N
D
N
C
N
N
N
N
D
N
C
N
N
N
BR2
N
N
D
N
N
D
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
MOD
P
A
P
A
A
P
A
A
A
P
A
A
A
P
A
A
A
A
A
A
P
A
A
A
C1
FW
N
PC
N
N
FW
N
N
N
FW
N
N
N
FW
N
N
N
N
N
N
PC
N
N
N
C2
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
C3
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
CU
LM
N
L
N
N
L
N
N
N
L
N
N
N
LM
N
N
N
N
N
N
L
N
N
N
LM1
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM2
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM3
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM4
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM5
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
LM6
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
Third Phalanx
BN #
F29-3-282
F28-9-714
F28-9-513
F28-9-480
F28-9-479
F28-9-472
F28-9-402
F28-9-392
F28-9-390
F28-9-367
F28-9-339
F28-9-338
F28-9-327
F28-9-267
F28-9-254
F28-9-243
F28-9-125
F28-8-958
F28-8-934
F28-8-920
F28-8-919
F28-8-880
F28-8-87
F28-8-83
SEX
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
ELE
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PFUS
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
182
F28-8-77
F28-8-760
F28-8-724
F28-8-61
F28-8-438
F28-8-433
F28-8-385
F28-8-377
F28-8-373
F28-8-369
F28-8-163
F28-8-108
F28-7-75
F28-7-606
F28-7-510
F28-7-505
F28-7-462
F28-7-459
F28-7-40
F28-7-25
F28-7-116
F28-7-107
F28-6-59
F28-6-120
F28-6-116
F28-5-60
F28-5-29
F28-5-198
F28-5-157
F28-5-13
F28-4-813
F28-4-762
F28-4-760
F28-4-745
F28-4-738
F28-4-683
F28-4-662
F28-4-655
F28-4-654
F28-4-648
F28-4-647
F28-4-645
F28-4-623
F28-4-593
F28-4-591
F28-4-538
F28-4-490
F28-4-466
F28-4-431
F28-4-416
F28-4-360
F28-4-347
F28-4-338
F28-4-306
F28-4-228
F28-4-197
F28-4-1663
F28-4-1653
F28-4-1611
F28-4-1591
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
LT
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
LT
LT
ME
LT
LT
ME
ME
ME
LT
ME
LT
ME
ME
LT
LT
ME
ME
LT
ME
ME
ME
ME
LT
LT
ME
LT
ME
LT
ME
ME
ME
LT
ME
LT
ME
ME
LT
ME
LT
LT
ME
ME
LT
LT
LT
ME
LT
ME
ME
LT
ME
LT
ME
LT
ME
ME
LT
ME
LT
LT
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
N
C
D
N
C
C
N
C
D
C
C
N
N
D
D
N
N
C
N
C
C
C
C
N
D
D
C
C
C
C
C
C
N
D
C
N
N
C
D
D
D
N
N
N
N
N
N
D
D
N
N
N
N
D
D
D
C
C
C
C
N
D
N
N
D
D
N
D
N
N
D
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
D
D
N
N
D
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
D
A
P
A
A
P
P
A
P
A
P
P
A
A
A
A
A
A
P
A
P
P
P
P
A
A
A
P
P
P
P
P
P
A
A
P
A
A
P
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
A
P
P
P
P
N
FW
N
N
FW
FW
N
PC
N
PC
FW
N
N
N
N
N
N
FW
N
FW
FW
FW
FW
N
N
N
FW
FW
PC
FW
FW
PC
N
N
PC
N
N
PC
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
FW
FW
FW
PC
N
N
N
N
TS
N
N
N
N
N
PC
N
N
N
N
N
N
N
N
N
N
PC
N
N
N
N
N
N
N
PC
N
N
N
N
FW
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
PC
N
N
FW
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
L
N
N
M
L
N
L
N
L
LM
N
N
N
N
N
N
L
N
L
L
LM
L
N
N
N
L
L
L
L
LM
LM
N
N
LM
N
N
L
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
L
L
L
LM
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
183
F28-4-1559
F28-3-85
F28-3-84
F28-3-782
F28-3-605
F28-3-577
F28-3-566
F28-3-553
F28-3-552
F28-3-527
F28-3-517
F28-3-508
F28-3-507
F28-3-506
F28-3-494
F28-3-444
F28-3-432
F28-3-426
F28-3-413
F28-3-398
F28-3-385
F28-3-380
F28-3-349
F28-3-315
F28-3-291
F28-3-260
F28-3-249
F28-3-244
F28-3-219
F28-3-181
F28-3-118
F28-3-107
F28-2-396
F28-2-385
F28-2-373
F28-2-340
F28-2-308
F28-2-276
F28-13-273
F28-13-266
F28-13-265
F28-12-479
F28-12-470
F28-12-427
F28-12-241
F28-12-216
F28-12-189
F28-12-164
F27-9-179
F27-9-177
F27-27-959
F27-25-69
F27-25-43
F27-25-135
F27-25-129
F27-25-113
F27-25-112
F27-25-109
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
LT
LT
LT
LT
ME
ME
LT
LT
LT
LT
LT
ME
ME
ME
LT
LT
LT
ME
LT
ME
LT
ME
ME
ME
ME
ME
ME
LT
ME
LT
LT
ME
LT
ME
LT
ME
LT
LT
ME
ME
LT
ME
ME
ME
LT
ME
LT
LT
ME
LT
LT
LT
LT
ME
LT
ME
LT
LT
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
C
C
N
G
C
C
D
D
D
N
N
N
N
D
N
D
D
D
C
C
N
D
C
C
N
N
D
C
N
D
N
N
D
N
D
C
D
N
D
N
N
D
C
N
N
C
D
D
D
N
C
N
N
D
N
N
C
N
D
N
N
D
N
N
N
N
N
N
N
N
N
N
N
N
N
N
D
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
D
N
N
N
N
N
N
N
N
N
N
N
N
P
P
A
A
P
P
A
A
A
A
A
A
A
A
A
A
A
A
P
P
A
A
P
P
A
A
A
P
A
A
A
A
A
A
A
P
A
A
A
A
A
A
P
A
A
P
A
A
A
A
P
A
A
A
A
A
P
A
PC
PC
N
N
FW
FW
N
N
N
N
N
N
N
N
N
N
N
N
PC
PC
N
N
FW
FW
N
N
N
PC
N
N
N
N
N
N
N
PC
N
N
N
N
N
N
PC
N
N
PC
N
N
N
N
FW
N
N
N
N
N
FW
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
PC
N
N
N
FW
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
L
L
N
N
L
L
N
N
N
N
N
N
N
N
N
N
N
N
L
L
N
N
L
LM
N
N
N
LM
N
N
N
N
N
N
N
L
N
N
N
N
N
N
L
N
N
L
N
N
N
N
L
N
N
N
N
N
L
N
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
184
F27-25-108
F27-24-731
F27-24-704
F27-24-457
F27-24-433
F27-24-426
F27-24-410
F27-24-402
F27-24-384
F27-24-383
F27-24-376
F27-24-374
F27-24-372
F27-24-353
F27-24-348
F27-24-342
F27-24-291
F27-24-271
F27-24-263
F27-24--220
F27-24-218
F27-23-949
F27-23-940
F27-23-939
F27-23-883
F27-23-764
F27-23-760
F27-23-755
F27-23-741
F27-23-740
F27-23-738
F27-23-724
F27-23-723
F27-23-722
F27-23-714
F27-23-706
F27-23-700
F27-23-625
F27-23-559
F27-23-526
F27-23-453
F27-23-406
F27-23-335
F27-23-332
F27-23-331
F27-23-324
F27-23-317
F27-23-307
F27-23-289
F27-23-247
F27-23-243
F27-23-194
F27-23-193
F27-23-183
F27-23-182
F27-23-179
F27-23-173
F27-23-107
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
LT
ME
ME
ME
ME
LT
ME
ME
LT
ME
LT
ME
LT
LT
ME
LT
LT
LT
LT
LT
ME
ME
LT
LT
ME
ME
ME
ME
LT
LT
ME
ME
ME
LT
ME
LT
LT
ME
ME
ME
ME
ME
ME
ME
ME
LT
ME
ME
LT
LT
ME
ME
LT
LT
ME
LT
ME
ME
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
D
C
D
N
D
N
D
N
C
D
D
D
C
N
D
C
D
D
C
N
C
C
D
C
C
D
C
C
C
C
C
D
C
C
C
C
C
N
N
N
D
C
D
N
N
N
C
C
C
N
C
N
N
C
N
N
D
N
N
N
N
N
N
N
N
N
N
N
N
N
D
N
N
N
N
N
D
N
D
N
N
N
D
N
N
N
G
D
N
N
N
N
D
D
N
N
N
N
N
N
N
N
N
N
N
N
D
N
N
N
N
D
N
N
N
N
A
P
A
A
A
A
A
A
P
A
A
A
P
A
A
P
A
A
P
A
P
P
A
P
P
A
P
P
P
P
P
A
P
P
P
P
P
A
A
A
A
P
A
A
A
A
P
P
P
A
P
A
A
P
A
A
A
A
N
FW
N
N
N
N
N
N
FW
N
N
N
FW
N
N
PC
N
N
FW
N
PC
PC
N
PC
FW
N
FW
FW
FW
FW
PC
N
PC
FW
FW
FW
FW
N
N
N
N
FW
N
N
N
N
FW
FW
FW
N
FW
N
N
FW
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
PT
N
N
FW
N
TS
N
N
N
N
N
N
FW
N
FW
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
FW
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
L
N
N
N
N
N
N
L
N
N
N
LM
N
N
L
N
N
M
N
L
LM
N
LM
LM
N
L
L
L
LM
L
N
L
L
LM
L
LM
N
N
N
N
LM
N
N
N
N
L
L
L
N
L
N
N
L
N
N
N
N
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
185
F27-22-399
F27-22-383
F27-22-378
F27-22-312
F27-22-310
F27-22-285
F27-22-285
F27-22-270
F27-22-257
F27-22-255
F27-19-328
F27-19-288
F27-18-821
F27-18-807
F27-18-725
F27-18-626
F27-18-623
F27-18-560
F27-18-48
F27-18-428
F27-18-421
F27-18-412
F27-18-408
F27-18-402
F27-18-397
F27-18-366
F27-18-365
F27-18-322
F27-17-977
F27-17-954
F27-17-883
F27-17-867
F27-17-852
F27-17-842
F27-17-815
F27-17-777
F27-17-733
F27-17-727
F27-17-725
F27-17-669
F27-17-648
F27-17-627
F27-17-597
F27-17-593
F27-17-528
F27-17-507
F27-17-480
F27-17-439
F27-17-383
F27-17-1502
F27-15-151
F27-14-185
F27-14-149
F27-13-486
F27-13-460
F27-13-439
F27-13-377
F27-13-319
F27-13-260
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
LT
CO
CO
CO
CO
CO
CO
CO
CO
LT
ME
LT
LT
LT
ME
ME
ME
LT
ME
ME
ME
LT
ME
ME
LT
ME
ME
ME
LT
LT
LT
LT
LT
ME
LT
ME
ME
ME
ME
LT
LT
LT
ME
LT
ME
ME
LT
LT
LT
ME
ME
ME
ME
ME
LT
LT
ME
LT
LT
LT
ME
LT
ME
ME
LT
ME
LT
LT
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
N
N
C
N
C
C
N
C
N
N
N
N
C
C
N
D
N
C
N
C
C
C
C
N
C
C
N
C
N
D
C
D
C
D
N
N
C
C
G
N
N
D
D
D
C
N
D
C
D
C
N
C
N
N
N
N
C
N
N
N
N
N
N
D
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
D
N
N
N
D
N
N
N
N
N
D
N
D
N
N
N
N
N
N
N
N
N
N
N
D
N
N
N
N
D
N
N
N
N
N
N
N
N
N
A
A
P
A
P
P
A
P
A
A
A
A
P
P
A
A
A
P
A
P
P
P
P
A
P
P
A
P
A
A
P
A
P
A
A
A
P
P
A
A
A
A
A
A
P
A
A
P
A
P
A
P
A
A
A
A
P
A
A
N
N
FW
N
FW
FW
N
FW
N
N
N
N
FW
PC
N
N
N
FW
N
FW
FW
TS
FW
N
FW
FW
N
FW
N
N
PC
N
FW
N
N
N
PC
FW
N
N
N
N
N
N
PC
N
N
PC
N
FW
N
FW
N
N
N
N
PC
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
PC
N
PC
N
N
PT
N
N
N
N
PC
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
FW
N
N
FW
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
L
N
L
LM
N
L
N
N
N
N
L
L
N
N
N
L
N
LM
LM
L
LM
N
LM
LM
N
LM
N
N
LM
N
L
N
N
N
L
L
N
N
N
N
N
N
L
N
N
LM
N
L
N
LM
N
N
N
N
L
N
N
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
186
F27-12-324
F25-7-24
E28-6-13
E28-25-90
E28-25-9
E28-25-121
E28-16-92
E28-16-68
E28-16-58
E28-15-45
E28-15-3
E27-6-33
E27-6-18
E27-5-39
E27-5-38
E27-15-55
E27-15-186
E27-15-158
E27-15-130
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
PHT
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
CO
LT
LT
ME
ME
LT
ME
ME
ME
LT
LT
ME
LT
LT
LT
ME
ME
ME
ME
LT
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
N
D
N
C
C
N
N
D
D
D
N
N
N
N
N
D
N
N
N
N
N
N
N
D
N
N
N
N
N
N
N
N
N
N
N
N
N
N
A
A
A
P
P
A
A
A
A
A
A
A
A
A
A
A
A
A
A
N
N
N
FW
FW
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
N
L
L
N
N
N
N
N
N
N
N
N
N
N
N
N
N
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
187
APPENDIX E: ANALYSIS OF SEX FOR SPECIFIC
SKELETAL ELEMENTS
Humerus
Figure E.1: Sex analysis scatter plot for the humerus using measurements 11 and 7.
188
Radius-Ulna
Figure E.2: Sex analysis scatter plot for the radius-ulna using measurements 9 and 4.
Figure E.3: Sex analysis scatter plot for the radius-ulna using measurements 11 and 7.
189
Metacarpal
Figure E.4: Sex analysis scatter plot for the metacarpal using measurements 1 and 5.
Figure E.5: Sex analysis scatter plot for the metacarpal using measurements 2 and 5.
190
Figure E.6: Sex analysis scatter plot for the metacarpal using measurements 3 and 5.
Figure E.7: Sex analysis scatter plot for the metacarpal using measurements 4 and 5.
191
Femur
Figure E.8: Sex analysis scatter plot for the femur using measurements 17 and 10.
Figure E.9: Sex analysis scatter plot for the femur using measurements 8 and 10.
192
Figure E.10: Sex analysis scatter plot for the femur using measurements 8 and 17.
Tibia
Figure E.11: Sex analysis scatter plot for the tibia using measurements 1 and 2.
193
Figure E.12: Sex analysis scatter plot for the tibia using measurements 1 and 6.
Figure E.13: Sex analysis scatter plot for the tibia using measurements 2 and 6.
194
Figure E.14: Sex analysis scatter plot for tibia using measurements 1 and 7.
Figure E.15: Sex analysis scatter plot for the tibia using measurements 2 and 7.
195
Metatarsal
Figure E.16: Sex analysis scatter plot for the metatarsal using measurements 1 and 5.
Figure E.17: Sex analysis scatter plot for the metatarsal using measurements 2 and 5.
196
Figure E.18: Sex analysis scatter plot for the metatarsal using measurements 3 and 5.
Figure E.19: Sex analysis scatter plot for the metatarsal using measurements 4 and 5.
197
Astragalus
Figure E.20: Sex analysis scatter plot for the astragalus using measurements 3 and 5.
Calcaneus
Figure E.21: Sex analysis scatter plot for the calcaneus using measurements 5 and 4.
198
Figure E.22: Sex analysis scatter plot for the calcaneus using measurements 6 and 7.
199