The Pennsylvania State University
The Graduate School
College of the Liberal Arts
ADAPTATION, TRANSLOCATION, AND BREED SPECIALIZATION: EXPLORING THE
POTENTIAL TO IDENTIFY AND EXPLAIN BREED DEVELOPMENT IN EUROPEAN
AND NATIVE AMERICAN DOMESTIC DOGS (CANIS FAMILIARIS).
A Dissertation in
Anthropology
by
Martin Hughes Welker
2018 Martin Hughes Welker
Submitted in Partial Fulfillment
of the Requirements
for the Degree of
Doctor of Philosophy
August 2018
The dissertation of Martin Hughes Welker was reviewed and approved* by the following:
Sarah B. McClure
Associate Professor of Anthropology
Dissertation Advisor
Chair of Committee
Douglas J. Kennett
Professor of Anthropology
Head of the Department of Anthropology
Douglass W. Bird
Associate Professor of Anthropology
Ellen Stroud
Associate Professor of History
*Signatures are on file in the Graduate School
iii ABSTRACT
The domestication of animal species was followed by the development of breeds, or distinct populations that are both homogenous in appearance and differentiable from others of the same species (Clutton-Brock 2012). Breeds emerge through a combination of natural and artificial selection acting upon animal species in close association with human populations. In many cases (e.g., Clydesdale horses, Black Angus cattle, Border Collies) breed development can reflect the role particular populations play within human communities. As a result, understanding when, where, and how morphologically distinct populations of domesticates emerge has significant potential for furthering our understanding of human-animal relationships and interaction.
Identifying and understanding the domestication process, the roles of domestic plant and animal species in agricultural and forager societies, and the ecological impacts of domesticates on human societies remain among archaeology’s “Grand Challenges”
(Kintigh et al. 2014). Extensive archaeological investigation has focused on developing techniques for identifying physical attributes associated with domestication (e.g., Benecke
1987; Crockford 1997; Evin et al. 2013; Lawrence and Bossert 1967; Olsen 1985; Zeder
2001, 2006; Zeder and Hesse 2000; Zeder et al. 2006). Further discussion has centered upon the definition of distinct domestication pathways which explain the emergence of various domesticates or cohabiting organisms adapted to living in close association with human beings (Zeder 2012). In comparison, the long-term management of species, and resulting genotypic and phenotypic impacts have received significantly less study, despite
iv the presence of breeds and breed improvement strategies in Europe since at least the Iron
Age.
This dissertation uses domestic dogs (Canis familiaris) as a case study for investigating the potential of detecting and using population or breed level differences within archaeological assemblages to better understand interaction between humans and domesticates. Domestic dogs are a logical case species with which to start for several reasons. First, they are among the most plastic of extant species with the most exaggerated phenotypic differentiation. Their resultant morphological characteristics are highly exaggerated and more easily recognizable using the methods employed here. Second, in undertaking this study, I also identify challenges that arise from natural morphological variation which will be faced by future researchers seeking to identify breeds in other species in order to explore aspects of human culture.
Dogs were domesticated by 12,000-14,000 years BP in Eurasia (Castroviejo-Fisher et al. 2011; Larson et al. 2012; Leonard et al. 2002; van Asch et al. 2013). Since their domestication, dogs have thrived in foraging and agricultural human societies, become globally distributed, and have adapted to a suite of specialized roles as hunting aids, weapons of war, food, sources of fiber for clothing and textiles, livestock protection, and beasts of burden (Allen 1920; Crockford 2000; Horard-Herbin et al. 2014; Lupo 2017;
Schwartz 1998). Adaptation to these roles has drastically impacted domestic dogs’ physical form in size (e.g. chihuahuas and great danes), physical proportions (e.g. dachshunds and whippets), and behavioral patterns (Mehrkam and Wynne 2014). Archaeological identification of dog breeds relies on recognizing these patterns of extreme morphological
v variability in domestic dogs and has resulted in numerous techniques for detecting and characterizing physical attributes from canid skeletal remains (Clark 1996; Harcourt 1974;
Losey et al. 2015; Losey et al. 2017; Worthington 2008).
My dissertation builds upon this foundation to address gaps in our knowledge concerning both the development of breeds along with the implications of breed’s existence and use in past societies. This dissertation includes three chapters that address the emergence of breeds, their adaptation to specialized roles within human societies, and their potential to serve as markers of human migration. I integrate datasets drawn from biomorphometric analyses, ethnographic and historic documents, and the Field Processing
Model from Human Behavioral Ecology to address these questions. Morphometric data collected on European and Native American dog populations, was used to systematically test Colonial European’s preference for large working breeds and their use as markers of human migration. Ethnographic, experimental, and archaeological data is used to test ethnographic sources and historic accounts concerning Intermountain and Great Plains dogs’ load capacity when using travois (simple drag sleds), and pannier-style pack. Finally, the Field Processing Model from Human Behavioral Ecology was used to investigate the conditions that encourage or discourage using dogs to haul or carry goods and resources and highlight dogs’ significance to Intermountain and Plains residential mobility.
By developing an understanding of when and where breeds have emerged we can begin to understand both the roles that domesticates played in human societies and the cultural and environmental criteria that make breed development possible. A theoretical framework drawn from Human Behavioral Ecology is useful for identifying and
vi explaining the decision to invest in breed development while following established protocols (Von den Driesch 1976) for highly reproducible methods such as biomorphometrics (measuring skeletal landmarks) is vital to recognizing breeds within the archaeological record and investigating how they were used in the past. The Direct
Historical Approach applied and a theoretical framework drawn from Human Behavioral
Ecology provide powerful tools for identifying and explaining the decision to invest in breed development and associated technological innovations such as the travois. Using these tools, this dissertation generates a series of important contributions to anthropological research including: 1) exploring the potential for breed selection and the creation of distinctive phenotypes to serve as a signature of colonization; 2) testing the capacity of Intermountain and Plains dogs to transport loads of as much as 100lbs; 3) finding evidence that dogs’ roles in transport on the Great Plains and in the Intermountain
West may extend back to over 3,000 years BP; and 4) showing that draught dogs’ functioned as a significant element in highly adapted system of strategic residential mobility used by Intermountain and Plains foragers to counter environmental uncertainty.
vii TABLE OF CONTENTS
LIST OF FIGURES ...... ix
LIST OF TABLES ...... xi
ACKNOWLEDGEMENTS ...... xii
Chapter 1 Introduction ...... 1
The Promise and Peril of Breed Development: Identifying and Studying Domesticates and Breeds in the Archaeological Record ...... 4 Breed Taxonomy ...... 7 Population Size and Breed Development ...... 10 Evolutionary Change ...... 13 Natural and Artificial Morphological Patterns and Scientific Taxonomy ...... 19 Natural Morphological Variation ...... 19 Domestication Syndrome and Morphological Changes in Domesticated Animal Species ...... 24 Domesticates as species, integrating natural and artificial selection ...... 29 When Will Breeds be Detected in the Archaeological Record ...... 32 Dogs as a Case Study ...... 34 Dissertation Organization and Research Questions ...... 35
Chapter 2 Breed Selection and Colonization: Exploring the Potential for Breeds to Act as Markers of Population Movement ...... 41
Introduction ...... 42 Background: ...... 44 Colonial Dogs: ...... 44 Dogs in England and Europe: ...... 45 Native American Dogs: ...... 46 Materials and Methods: ...... 47 Results: ...... 53 Discussion: ...... 57 Conclusion: ...... 60 Acknowledgments: ...... 61
Chapter 3 The Birch Creek Canids and Dogs as Transport Labor in the Intermountain West ...... 62
Introduction: ...... 63 Domestic Dog Morphology, Mobility and Labor ...... 67 Data and Methods ...... 72 Results ...... 76 Discussion: The Birch Creek Dogs in Broader Context ...... 86 Conclusions ...... 96
viii Acknowledgements: ...... 98
Chapter 4 Traveling with the Pack: The Field Processing Model and the Role of Dogs in Intermountain and Plains Residential Mobility ...... 99
Introduction: ...... 100 Dogs in Foraging Societies - Hunting, Provisioning, and Transport ...... 103 Firewood: ...... 106 Shelter and Household Belongings: ...... 107 Bison Meat: ...... 108 Modeling Dogs as Draught Animals: ...... 110 Results: ...... 117 Discussion: ...... 119 Conclusion: ...... 124
Chapter 5 Conclusions and Future Directions ...... 127
Future Directions: ...... 132
Appendix A Shoulder Height in Native American, Colonial, and European Dogs ...... 136
Appendix B Body Mass Estimates for Intermountain, Plains and Great Basin Dog Remains ...... 146
References Cited ...... 164
ix LIST OF FIGURES
Figure 1-1: A graphical representation of the three pathways to domestication defined by Zeder (2012)...... 6
Figure 1-2: A graphical representation breed emergence and specialization processes...... 9
Figure 1-3: The effects of Bergmann-Allen’s and Rensch’s Rules...... 21
Figure 1-4: A comparison of sexual size dimorphism in wild and domestic sheep (Ovis spp.) and goats (Capra spp.). Data from Polák and Frynta 2009...... 26
Figure 2-1: The location of the Fortress of Louisbourg (top left). Fortress of Louisbourg site overview (top right). Locations where domestic dog remains used in this analysis were recovered by Marwitt (1966) and Silver (1969) (bottom)...... 48
Figure 2-2: Excavation photos for the Fortress of Louisbourg dogs (Marwitt 1966; Silver 1969), and photos showing the severe displaced fracture suffered by dog 1B3B87...... 49
Figure 2-3: The distribution of dogs for which morphometric data exists and date to the Colonial Period (Numbering follows Table 2-3)...... 51
Figure 2-4: Shoulder height (in cm) in Native American, European, and Colonial dog populations...... 54
Figure 2-5: Native American dog shoulder height (in cm) plotted against time for the period 0-1800 AD...... 56
Figure 2-6: European dog shoulder height (in cm) plotted against time for the period 0- 1800 AD...... 56
Figure 3-1: A map depicting the location of sites used in this analysis which are numbered as follows: 1. Veratic Rockshelter, 2. Braden Site, 3. Jaguar Cave, 4. Fishing Bridge Campground, 5. Stillwater Marsh, 6. Pyramid Lake, 7. Vista Site, 8. Hogup Cave, 9. Danger Cave, 10. Caldwell Village, 11. Pharo Village, 12. Larson Site, 13. Lower Grand Site, 14. Potts Site, 15. Pretty Head Site, 16. White Buffalo Robe Site, 17. Big Hidatsa Site, 18. Barcal Site, 19. Bellwood Site, 20. Burkett Site, 21. Clarks Site, 22. Gray Site, 23. Hill Site, 24. Horse Creek Site, 25. Linwood Site, 26. Palmer Site, 27. Write Site...... 65
Figure 3-2: A panel of photos depicting mandibles a) IMNH-18803, b) IMNH- 18418/IMNH-19210, c) IMNH-19636, d) IMNH-19641, and reconstructed crania e/f) IMNH-19641 which exhibits an unhealed cranial fracture...... 79
Figure 3-3: (a) Mandibular crowding indices for archaeologically reported domestic dogs (see Clark 1996), (b) M1 length for the Veratic Rockshelter dogs and reported values
x for male (MNI=62) and female (MNI=47) wolves, and male (MNI=99) and female (MNI=99) coyotes (Nowak 1979)...... 81
Figure 3-4: The most commonly used measurements used in this analysis for estimating body mass (Based on von den Driesch 1976)...... 82
Figure 3-5: Body mass in kg for the Veratic Rockshelter dogs, male (MNI=24) and female (MNI=25) wolves (Mech 2006), and an unknown number of male and female western coyotes (Way 2007)...... 84
Figure 3-6: A boxplot showing the range of body mass calculated for dogs found in each region following Losey et al. (2015) and Losey et al (2017). Dotted lines depict the upper and lower limits of body mass estimates generated for mandibles from sites in Nebraska which Bozell (1988) attributed to the smaller “Plains-Indian dog”...... 89
Figure 3-7: A histogram showing the distribution of body mass estimates generated for archaeological dog remains from the Great Plains...... 92
Figure 3-8: The Plains sample through time showing the timing of Spanish exploration on the Southern Plains, and the periods of Iroquois and European managed fur trade in the North...... 95
Figure 4-1: A graphical representation of the Field Processing Model as adapted to travois transport. U=caloric value, S=processing for a human load, P= additional processing needed for larger travois loads...... 112
Figure 4-2: The loads evaluated in this study. *Note that because human and long- distance dog only travois loads are equivalent to one another this load does not meet condition one of the Field Processing Model...... 114
xi LIST OF TABLES
Table 1-1: Minimum effective population sizes by species (Leroy et al. 2013)...... 12
Table 1-2. Adherence of various taxa to Bergmann’s and Rensch’s Rules...... 24
Table 2-1. Equations used to estimate shoulder height from long bone greatest length (GL) measurements by element (Harcourt 1974)...... 50
Table 2-2. Louisbourg dog measurements and shoulder height estimates...... 52
Table 2-3. Colonial dog shoulder height estimates and dates...... 53
Table 2-4. Comparative analysis results...... 54
Table 3-1: Veratic Rockshelter domestic dog specimens...... 77
Table 3-2: Radiocarbon data for dated specimens (samples processed by Direct AMS and University of Utah)...... 80
Table 3-3: Identifying metrics for domestic dog specimens...... 80
Table 3-4: Veratic Rockshelter dog body mass estimates, short- and long-distance travois, and pack load capacities...... 83
Table 3-5: Veratic Rockshelter dog body mass estimates, short- and long-distance travois, and pack load capacities...... 87
Table 3-6: Comparative sample body mass estimates, and travois and pack load capacity estimates...... 91
Table 4-1: Open fire fuelwood consumption in various societies...... 107
Table 4-2: Available weight data on Plains tipi...... 108
Table 4-3: Caloric value, encounter rates, and processing time for resources available to Intermountain and Plains foragers (Data from Byers and Ugan (2005), Simms (1985, 1987), Ugan (2005)...... 115
Table 4-4: Short- and long-distance handling time (in hours), loads required, and kcal/load data for bison, deer, and pronghorn antelope...... 120
Table 4-5: Travel time and distance transition points from human loads to travois loading strategies...... 121
Table 4-6: Herd density for bison, elk, mule deer, mountain sheep, and pronghorn antelope...... 124
xii ACKNOWLEDGEMENTS
To say the path to this dissertation was wandering would be an understatement.
Rather, the research encapsulated in this dissertation is but one of many projects I have started and which stand in various stages of completion. It is, however, fitting that the culmination of my journey through graduate school has returned me to my familial, geographical, and even theoretical roots in one way or another. As a bright-eyed and bushy-tailed young graduate student I aspired to study the European Neolithic. In reality,
I did anything but this. Following the steps of my ancestors I moved to central
Pennsylvania a stone’s throw from the farms my paternal ancestors have worked since the 1700’s. I undertook research at the Fortress of Louisbourg which my maternal ancestor Neil McPhee (Private, Fraser’s Highlanders) helped capture in the French and
Indian War. Though I traveled to Europe numerous times and spent incalculable hours analyzing bone from Neolithic sites, material from Idaho and the Intermountain West became a central feature in my dissertation research. And, I retraced the paw-prints of the familiar intellectual pathway that is Human Behavioral Ecology to accomplish these studies. Lessons learned.
The guidance and support of my mentors past and present has been exceptional and deeply appreciated. I am profoundly grateful to Drs. Sarah McClure, Douglas
Kennett, Douglas Bird, and Ellen Stroud who agreed to serve on my dissertation committee despite the evolving nature of my dissertation research. In particular I must thank Sarah McClure who has provided me with her unwavering support through the best and worst of my experiences in graduate school, and who has enabled me to continue
xiii entertaining my fanciful notions of studying the Neolithic even as my research led me further and further away. The Penn State Zooarchaeology lab will forever be my happy place and sanctuary.
I must also acknowledge the enduring impact that Drs. Patricia Lambert, David
Byers, Kenneth Cannon, Molly Cannon, and Steven Simms and the rest of the faculty at
Utah State University had, and continue to have, on my development as an anthropologist. I would not be the person I am without the field, lab, and classroom experiences they afforded me as an undergraduate student. Their continued support throughout my graduate career has been extraordinary. I am forever indebted to their tutelage and can only hope to someday repay them for the kindness and unwavering encouragement.
I would be remiss if I did not also extend my deep-felt thanks to the numerous friends and colleagues including Emily Zavodny, Claire Ebert, Jelena Jovic, Emil
Podrug, Richard George, Sandra Koch, Kristina Douglass, Diana Azevedo, Britt
McNamara, Nick Triozzi, Brendan Culleton, Trina and David Axford, Theresa Pitts-
Singer, and Ellen Klomps, who supported me throughout. In particular I must thank
Sandra Koch, my co-brewmaster organizer of the Monday morning writing group which kept me on task, and reader of endless drafts in various stages of legibility.
I am also sincerely grateful to Jason Beale, Abby Hileman, Abby Flemming and the rest of the Shaver’s Creek Environmental Center staff who allowed me to escape my life as a graduate student every Monday afternoon to join their menagerie. Matilda, Echo,
Jamaica, Cook, Jerudi, Chauntey, Tea, Tussey, Neo, Rufus, Susquahanna, and the rest of
xiv the flock provided endless hours of excitement, fun, and at times frustration, for which I will be forever thankful. To my girl Matilda in particular, you added the spark to my week. May the rats be forever plentiful, the trainers forever gullible, the shoelaces forever within reach, and your trust-bank never overdrawn.
Finally, I cannot thank my parents Joanne Hughes and Dennis Welker enough.
You guys are the best of friends, the most supportive of colleagues, and superb linemates on the ice. You have supported me through thick and thin and for that I can never repay you. I will be forever grateful for your sage advice and for making me who I am today.
Chapter 1
Introduction
Domesticated plant and animal species have played a central role in the livelihood, economy, and mobility of human communities around the world, providing both primary
(meat, blood, bone, seed crops, lumber) and secondary (milk, labor, wool, sap, fruit) products. The significance of domesticates in human cultures is reflected in their dietary importance, but also in the proliferation of breeds (e.g., Clydesdale horses) and populations
(oxen, mules) adapted to highly specialized roles within human society, the technological innovations used to maximize load size, and the training regimen used to accustom animals to these roles. The ability to identify breeds and the roles that domesticates play in human societies is therefore an important aspect to develop a holistic understanding of human- environmental and human-animal interactions in the past. A range of issues related to these interactions, including domestication, agricultural or pastoral intensification, migration, and the development of social complexity, remain among the “grand challenges” for archaeology as a field (Kintigh et al. 2014). Zooarchaeological data concerning the appearance of domesticates and changes in population structure or size have served both as type fossils demarcating the arrival or adoption of domesticates and changes in livestock management. The ability to identify archaeological animal populations that can be distinguished from other populations of the same species and changes in population
2 characteristics provides opportunities to both identify and trace population origins with greater detail and assess species’ role within human communities. Preliminary evidence suggests such populations may be used in ways similar to type fossils or artifacts used to create archaeological cultures (e.g., Bennet et al. 2016; Horard-Herbin et al. 2014). In this way, morphologically distinct populations may be thought of as archaeological breeds and act in the same way that “archaeological cultures” (Childe 1929; Kidder 1924; Steward
1942, 1955, 1972) based on elements of material culture (e.g. ceramics) are used to identify prehistoric populations and contribute to our understanding of past human behavior.
Breeds are domesticated animal populations that have been bred by humans to possess uniform heritable characteristics that distinguish them from other animals belonging to the same species (Clutton-Brock 2012:3). As a result, characterizing and comparing physical morphology within populations in the past is key to breed identification. Modern breeds are commonly defined using highly specific standards describing size, physical proportions, coloration, coat patterning, and parentage. Though detailed standards emerged in the 18th-19th centuries AD, imagery and historic descriptions support the existence of geographically or culturally defined breeds in many domesticates since at least the end of the Neolithic (Sherratt 1981, 1983). Archaeological evidence supports the presence of morphologically distinct populations (Clutton Brock 2012;
Horard-Herbin et al. 2014; Kazantzis and Albarella 2016) the extraction of secondary products, many of which became the basis of specialized breeds, throughout the Neolithic
(Bogaard 2004; Bogucki 1993; Milisauskas and Kruk 1991; Vigne and Helmer 2007, see also Greenfield 2010). Skeletal remains are among the most salient and best preserved archaeological evidence documenting evolutionary processes and speciation events in the
3 past, but also breed development and the roles domesticates filled within human communities. Biomorphometrics -the measurement of skeletal landmarks- is a highly reproducible, cost effective, and relatively quick means of investigating and characterizing physical morphology within populations.
This dissertation builds upon existing theoretical and methodological studies to explore the ways in which morphologically distinct populations can be detected within the archaeological record, and how their recognition may contribute to our understanding of the human condition. Morphological variation within domesticates emerges from the layering of characteristics resulting from artificial selection upon adaptations to environmental conditions resulting from natural selection. As a result, the study of breeds and their antiquity cannot be accomplished without first understanding the evolutionary processes that produced morphological variation within taxa, the domestication process, the role of domesticates in human society, and also defining what is accepted as a “breed” in the archaeological record. To accomplish these goals and better understand morphological variation in domesticates I integrate zoological, ecological, and archaeological research concerning domestication, the roles of domestic livestock within human societies, ecogeography, and the development of archaeological breeds. From this foundation I undertake several case studies to explore the ways in which morphologically distinct breeds or populations of domestic dog (Canis familiaris) can be used to better understand migrations and the behavioral processes that contribute to breed development.
4 The Promise and Peril of Breed Development: Identifying and Studying Domesticates and
Breeds in the Archaeological Record
In order to understand the significance of breed development and understand why their identification is important for archaeological research one needs to understand the process by which breeds, or morphologically distinct populations emerge (Clutton-Brock
2012:3). Domestication is the process whereby wild organisms are habituated to, and propagated within, human societies. As such domestication includes not only the initial taming of species, but also directed breeding, management, and other aspects of human animal cohabitation. Breeds are populations of domestic animals which share an internally homogenous phenotype, which is distinct from other members of the same species (Clutton-Brock 2012:3). Theoretically breeds have been present since two different populations of any domesticated animal species were brought under human management; however, they are rarely discussed within the archaeological literature. At this point it is necessary to further explore the concept of breeds, what they represent, how they develop, to what degree they were present in the past, and why they are significant to the study of past animal populations.
Domestication and Taming Wild Species
Domestication has occurred repeatedly in many regions of the world referred to as
“domestication centers” (Zeder and Hesse 2000). Extensive research efforts have concentrated on exploring how and why domestication events occur and identifying characteristics that primed species for domestication (Larson et al. 2005; Zeder 2012;
Zeder and Hesse 2000). These studies generally conclude that the progenitors of many
5 domesticated animal species share a propensity for being social herd or pack animals exhibiting established social hierarchies (Hale 1969; Price 1984, 2002; Zeder 2012)
(Figure 1-1). The presence of social hierarchies within these species enabled human beings to occupy existing positions of dominance (Hale 1969; Price 1984, 2002; Zeder
2012). In addition, species chosen for domestication frequently have short flight distances allowing humans to more easily accustom them to management (Hale 1969; Price 1984,
2002; Zeder 2012, Figure 1-1).
Domestication events followed unique trajectories but are believed to have proceeded through the relaxation of natural selection in concert with intensive artificial selective pressures focused upon largely behavioral characteristics encouraging reduced fight or flight responses in wild taxa (Zeder 2012). Zeder (2012) has organized domestication events into three “pathways” termed commensal, prey, and directed domestication. Commensal domesticates include dogs, cats, pigs, and chickens, and are those which are believed to have in some way initiated their own domestication usually by feeding on refuse near human settlements (Coppinger and Coppinger 2001; Morey
1994; Zeder 2012). Prey domesticates are those domesticated to serve as a source of food, and include most major livestock species including cattle, sheep, and goats. These species were likely subject to an extensive period of semi-management prior to domestication as human hunters followed herds around their natural habitats (Zeder 2008, 2011). Directed domesticates include horses and donkeys and are those who were domesticated with the goal of procuring an alternative product to meat protein. Such domesticates were likely exploited prior to domestication as a source of meat protein but were domesticated
6 primarily to provide secondary products including labor, fiber, feathers, or milk (Zeder
2012).
Figure 1-1: A graphical representation of the three pathways to domestication defined by Zeder (2012).
Breeds are defined as domesticated animal populations that exhibit a uniform set of physical characteristics selected for by human communities that distinguish these animals from others within the same species, and that are passed down through generations
(Clutton-Brock 2012:3). Clutton-Brock (2012) posits that the development of distinct breeds resulted from intentional selection for preferred behaviors, useful traits, or aesthetic characteristics. Selective breeding of this type requires not only control over mating behavior, but collective action in directed breeding efforts and specific roles or characteristics that were found to be useful or aesthetically pleasing. Human breeders rely upon regulated mating and culling efforts to manipulate the frequency of desired
7 characteristics within populations. Though this process is acknowledged in the archaeological literature, the details and implications of breed development, including the population size necessary for maintaining population viability or the timescale upon which breeds emerge, have not been widely studied.
Breed Taxonomy
The concept of breeds is both complex and multifaceted. Breeds distinctions are created within a culturally defined taxonomic system based upon biological variability resulting from a combination of both natural selection and investment on the part of human beings who define the characteristics they wish to see within their herds (Rege 1999, 2003;
Rege and Tawah 1999; Sponenberg et al. 2014). Breeds are defined most broadly as populations of domesticates sharing physical attributes which distinguish them from other populations of the same species (Clutton-Brock 2012: 3). The concept of breeds is most developed within Western cultural traditions where breeds are generally formally defined and evaluated against a detailed list of physical characteristics termed a breed standard.
Many such breed standards are listed in herd books or similar documents that may also document the pedigree of “purebred” individuals whose parentage includes only members of a single breed (Sponenberg et al. 2014). However, this concept of breed formalization is a product of 18th century European efforts to improve livestock or adapt them to very specific, and highly specialized roles within human communities (Rege 1999, 2003; Rege and Tawah 1999; Sponenbert et al. 2014).
8 These breeds are generally articulated with a taxonomic system which groups closely related breeds, those from similar geographic origins (e.g., Continental vs British cattle), or those developed for specific purposes (e.g., Herding vs. Toy dogs) into groups or classes (American Kennel Club, Rege 1999, 2003; Rege and Tawah 1999; Sponenberg et al. 2014). Breeds may also be subdivided into varieties or strains defined by variation that is not significant enough to warrant a new breed classification. In chickens, varieties are commonly defined by color or comb type. For example, a breed like the Wyandotte is available in several colors (e.g., white, buff, Columbian and silver-laced) which are controlled by genes that can be reliably produced in offspring (Hutt 2003; Jacob 2018).
The concept of formally defined breeds carries little weight outside of industrialized
Western cultures where intentional breeding programs have been used to develop livestock populations which are adapted to specific roles, a process termed “improving” breeds
(Rege 2003). This concept of breeds is problematic for archaeologists who commonly work with populations of domesticates which predate the formalization of breeds in this manner; but also because similar population divergence can result from either geographic distance or selective breeding (Rege 2003). Distinct differences are found between populations of domesticates in developing nations (Rege 1999, 2003; Rege and Tawah 1999), though these are rarely as formalized as those found in Western European cultures they are significant enough that the societies with which they live commonly create taxonomic distinctions between these populations (Rege 1999, 2003; Rege and Tawah 1999). In the literature these populations are referred to variably, and at times interchangeably, as
“breeds”, “landraces”, or “types” (Allen 1920; Banerjee et al. 2000; Rege 1999, 2003; Rege and Tawah 1999).
9 For archaeological purposes a less-formalized concept of breed identity is likely to be of greater use. Formalized or improved Western breeds developed primarily during the
18th Century and are unlikely to be found in many archaeological assemblages. In addition, formalized breeds developed from what are commonly referred to as “heritage breeds” or less specialized, though still distinctive populations of livestock akin to those in non-
Western traditions. These heritage breeds, landraces, or types have evolved under human management, but are generally more keenly attuned to survival in local environmental conditions (see Figure 1-2), than to fulfilling specific, highly specialized roles within human society (e.g., though cattle may be present, it is unlikely that specialized dairy or beef cattle breeds will be found in this situation).
Figure 1-2: A graphical representation breed emergence and specialization processes.
Irrespective of formalization, breeds, types, or landraces are only maintained when several conditions must be met. These include the management of a large enough breeding population to sustain populations over sustained periods of time, the prevention of significant genetic input from outside populations, and often geographical isolation and/or
10 exposure to disparate environmental conditions (Sponenberg et al. 2014). Acknowledging these conditions and using these to explore past animal populations may tell us something about the breed development, when and where we may expect to find breeds, and what their presence tells us about the communities in which they emerged.
Population Size and Breed Development
Breeds, types, or strains are created within human communities by genetically isolating a population of domestic animals through either geographic distance, or artificial selection and selective breeding (Rege 1999, 2003, Rege and Tawah 1999). Though archaeologists recognize that such processes were likely occurring in the past, those studying domestication have rarely discussed population sizes and questions of ownership in their analyses. Indeed, our understanding of the intersection between ownership, population size, and breed development is drawn from ethnographic studies and livestock breeders. Anthropologists working in African pastoral societies have generated substantial datasets regarding herd ownership, size, and demographics (e.g., Dyson-Hudson 1966;
Evans-Pritchard 1940; Schneider 1957; Sutter 1987; Williamson and Payne 1965).
Breeders and population geneticists have become increasingly aware of the dangers posed by inbreeding within genetically limited populations common to industrialized beef and dairying operations (see Leroy et al. 2013). These conditions are commonly discussed within the context of population bottlenecks when populations decline rapidly in response to environmental conditions. Under these conditions, limited phenotypic and genotypic diversity within populations reduce the likelihood that advantageous characteristics or
11 genetic mutations occur (Frankham et al. 1999; Visscher et al. 2001). Domesticates, and their composite breeds and herds, are thus populations whose size is artificially limited by human management.
Though anthropologists Dahl and Hjort (1976) have harnessed much of the available ethnographic information in their mathematical examination of herd demographics in African pastoral societies, few archaeologists have discussed likely herd sizes or composition in any detail. Bogucki (1988) remains one of the few archaeologists who has attempted to estimate herd size and composition but has done so on a purely theoretical level (though see Halstead 1996). He suggests that herds will ideally emphasize mature and immature female cattle, followed by castrates since these classes provide the majority of meat, milk, and traction which can be extracted from a herd. Herds will additionally include at least one mature and a few immature males which are needed to propagation (Bogucki 1988). Using ratios of these classes relative to one another Bogucki asserts that between 30-50 cattle are needed for herds to be economically beneficial.
Population geneticists and breeders have employed various measures of genetic relatedness, sex ratios, and generation length, to produce estimates of minimum population size for the purposes of breed or species conservation termed minimum viable populations or minimum effective populations (Leroy et al. 2013). These measures are informed by biological requirements and genetic diversity and are generally used to estimate the number of individuals needed to protect against the risks of population bottlenecks. Estimated effective population sizes for the major domesticates including cattle, sheep, horses, and dogs (Table 1-1) range from 70 to several thousand individuals depending upon the methods used (Leroy et al. 2013).
12 Few attempts have been made to correlate ethnographically observed or historically documented populations to these measures. Dahl and Hjort (1976) and Cribb (1984) provide significant contributions towards understanding herd size and demographics through quantitative models; however, these generally do not make a distinction between animals owned by individuals or families, and the larger, community or village level, herd.
A survey of the ethnographic and historic record shows that few individual preindustrial subsistence farmers owned herds large enough to be sustainable (Pruitt 1984; Schneider
1957; Shammas 1982; Sutter 1987). Early American colonists in North America are believed to have owned an average of under four cattle per family (Pruitt 1984; Shammas
1982). Even African pastoralists, who are largely dependent upon their livestock, own an average of 43 cattle per family, with 70% of families owning fewer than 50 (Sutter 1897).
These data imply that managing and sustaining viable populations of domesticates commonly requires cooperative investment on a community or inter-community level.
Table 1-1: Minimum effective population sizes by species (Leroy et al. 2013).
Effective Population Size Low High Cattle 91 21,648 Sheep 68 1,502 Horse 80 1,906 Dog 93 1,472
13 Evolutionary Change
Evolutionary change is frequently portrayed as a slow-moving process occurring over millennia; however, numerous examples have indicated that significant phenotypic and genotypic change can be achieved over relatively short timeframes (e.g., Johnson and
Selander 1964; Lamichhaney et al. 2018; Stuart et al. 2014). The speed with which alterations to population genotype or phenotype occur are linked to the strength of selective pressure – how advantageous a particular characteristic is under the environmental conditions an organism is exposed to. Within population genetics, selective pressure is measured using a selection coefficient generated by measuring selective pressure against a particular genotype on a scale of 0 to 1 where values close to 0 indicate high fitness
(positive selective pressure) and values close to 1 indicate low fitness (negative selective pressure). Long-term observation of species such as Darwin’s finches on the Galapagos
Islands, Ecuador, has revealed that phenotypic changes in beak size can occur within only a few decades (Lamichhaney et al. 2018). These studies indicate that significant and detectable phenotypic changes can appear in relatively short periods of time.
Human management mitigates the influence of natural selection acting upon species and expose them instead to artificial selective pressure, which commonly exhibits much stronger selective coefficients than those found in many natural systems. As a result, though both natural and artificial selection continue to influence domesticates and breeds under human management, artificial selection becomes the primary driver of phenotypic and genotypic change (Zeder 2012). Extensive experiments involving the managed breeding of captive foxes indicate that traits associated with domesticated dogs including
14 floppy ears, having piebald (patched black and white) coats or brown spots, barking, and shortened snouts emerged within only 40 generations bred between 1965 and 2002
(Belyaev et al. 1985; Trut 1999; Trut et al. 2004; Trut et al. 2009).
The speed with which phenotypic changes emerge within species is a significant though poorly understood element of domestication. Archaeologists working from skeletal remains face a number of significant challenges in identifying morphologically distinct populations in the archaeological record. Among these is the fact that zooarchaeologists rarely work with the remains of individual animals, but instead from material belonging to many comingled individuals, and possibly reflecting relatively long spans of time.
Taphonomic processes also present challenges as they frequently result in both small sample sizes and poor preservation. Overcoming these challenges requires greater understanding of the processes that contribute to morphological variation, and the ways in which domestication and sustained artificial selection may influence species morphotypes.
In addition, archaeologists must acquire sufficiently large samples to accurately characterize phenotypic variation within populations. Zoology and ecology have much to offer in these respects as a source of theoretical knowledge and comparative data. Despite these challenges, recent studies have shown that although defined breed standards (e.g., those of the American Kennel Club) were established as recently as the 18th and 19th centuries AD, distinct morphotypes in cattle (Kazantiz and Albarella 2016) and dogs
(Horard-Herbin et al. 2014) emerged as early as the Neolithic. Documentary evidence suggests that people were aware of phenotypic variation within domesticates, actively selecting for specific characteristics by the Roman period in Europe, but likely much earlier
(MacKinnon 2010).
15
Breed Visibility and Detection
These methods are variably called biomorphometrics, zoomorphometrics, geomorphometrics, or simply morphometrics. The term “morphometrics” itself was coined by zoologist Robert E. Backlith (1957) and refers to a collection of methods used for studying shape, size, and physical trait variation in organisms (Elewa 2010). Morphometric analyses of one sort or another have been used since at least the 5th century BC (Elewa
2010) and served as the foundation for theoretical developments on speciation (e.g.,
Linnaeus 1735) and evolution (e.g., Darwin 1859) and studying trait inheritance (Mendel
1866). Quantitative approaches in morphometrics emerged in the late 1800s and early
1900s by Karl Pearson, Francis Galton, and W. F. R. Weldon. Since that time morphological analyses have been adapted for use in ecology, biology, zoology, paleontology, and anthropology. Within archaeology and paleozoology morphometrical analyses focus upon skeletal remains and have been used to investigate diverse topics including predation pressure and over-fishing (Audzijonyte et al. 2013; Azevedo 2015;
Roy et al. 2003), game management and stocking (Wolverton et al. 2007), and conservation biology (Lyman 2009; Wolverton and Lyman 2012; Wolverton et al. 2016).
A survey of the literature illustrates the adaptability of morphometrics as a method.
Morphometric analyses performed on domesticates have focused on identifying skeletal characteristics indicative of initial taming of species and early domestication (Ameen et al.
2017; Clark 1996; Evin et al. 2013), and changes in physical morphology associated with their role in human societies (de Cupere et al. 2000; Groot 2005; Lin et al. 2016; Salmi and
Niinimäke 2016; Telldahl 2012; Thomas 2008). Despite the potential for breed
16 identification to contribute to our understanding of human-animal partnerships in the past, archaeologists have rarely discussed breed development, likely because they are wary of equivocating morphological variability within assemblages with the development of more than one morphologically distinct form or population (though see Albarella 1997; Grigson
1974, 1975, 1976, 1978, 1980). A degree of caution is warranted. Archaeologists are frequently confronted with comingled remains representing an unknown number of individuals, which produce relatively few repetitions of any particular element/feature/or measurable landmark, and often represent deposition over extended periods of time. As a result, morphological signatures generated from these remains do not necessarily reflect any single individual, but rather a cobbled together impression of population level variation.
Determining when archaeological evidence is strong enough to support the identification of morphologically distinct populations is complicated by several sources of natural and artificial morphological variability including sexual dimorphism, mixed deposition of wild and domestic individuals of a given taxa, translocation and adaptation to local environmental conditions, selective culling, and changes in management practices.
These challenges complicate decisions concerning which datasets provide meaningful comparisons. Successfully employing morphometric analyses to identify breeds and further our understanding of the human condition requires that we be able to identify and account for these variables in archaeological analyses.
Archaeologists have developed several methods for determining the sex of remains recovered from archaeological deposits. Some species have specific elements, or element morphology characteristic, of only one sex (e.g. bacculi in dogs or antlers in deer; see
17 Hatting 2017; Ruscillo 2003 2006). Notably, the absence of bacculi, antler, or horn may result from taphonomic processes and as a result such methods are not necessarily reliable indicators of sex. Accounting for sexual dimorphism is important for studying breed development and origins, especially for strongly dimorphic species (see Zeder 2001).
Archaeological analyses have commonly addressed this issue by plotting morphometric data on simple bivariate plots or using more advanced algorithmic analyses (see Telldahl et al. 2012). Clusters of larger and smaller individuals revealed by such plots are assumed to reflect the two sexes. Unfortunately, not all elements are equally sexually dimorphic, and as a result it can be difficult to reliably differentiate some skeletal elements. Further, size or physical characteristic variability may be introduced by cultural processes such as castration (e.g. oxen) or inter-species mating (e.g. mules).
Translocation and phenotypic change resulting from selection are more difficult to identify. Translocation is the process of introducing taxa to new environments.
Domesticates were displaced from the habitats to which they had become adapted through natural selection as a result of human migration and trade. The initial translocation of domesticates once identified is relatively easy to spot and has been widely used in identifying the arrival of agricultural populations (Bradley and Magee 2006; Edwards et al. 2007; Horard-Herbin et al. 2014; Zeder 2008; Zilhão 1993, 2001). Subsequent population movement and translocation events are more difficult to recognize; however, archaeological analyses have successfully explored human migration, livestock management, selective breeding, and trade revealing tantalizing new lines of evidence.
Horard-Herbin et al. (2014) shows that Paleolithic and Mesolithic foragers kept a phenotypically heterogenous population of dogs which was replaced by a highly
18 homogenous dog population introduced by incoming Neolithic farmers. Dog phenotypic diversity increased at the end of the Neolithic (de Grossi Mazzorin and Tagliacozzo 2000;
Harcourt 1974; Horard-Herbin et al. 2014) and by the time of the Roman Empire as many as 30 distinct forms existed (Bennett et al. 2016; Horard-Herbin et al. 2014). Allen (1920),
Haag (1948), Crockford (1997), and others have developed substantial datasets regarding breeds of Native American dog. Further datasets concerning the cranial morphology of modern domestic breeds have also been used for studying similarities between archaeological remains and modern dogs (Phillips et al. 2009; Tourigny et al. 2015).
Analyses of cattle, sheep, and goats have also produced significant contributions focusing on different cattle populations (Kazantzis and Albarella 2016) and phases of Roman,
Medieval, and Post-Medieval breed improvement (Albarella 1997; Davis 2008; Davis and
Beckett 1999; MacKinnon 2010; Van and Grimm 2010).
Unfortunately, progress in this area within archaeology has been accomplished largely in isolation from significant advancements by biologists, zoologists, and ecologists.
As a result, archaeological studies seeking to understand domestication and morphological variation have been divorced from a significant body of knowledge concerning the phenotypic expression of extant species, and the ways in which such morphological variation adapts taxa to their environmental conditions. Integrating theoretical and methodological knowledge from these fields will enable the refinement and development of methods with which to further our understanding human migration, trade, socio-political and cultural evolution, and of domesticates’ roles within human societies.
19 Natural and Artificial Morphological Patterns and Scientific Taxonomy
The archaeological record is inherently incomplete. Skeletal remains are commonly disassociated from soft tissue and hair, comingled, and influenced by additional taphonomic processes. The high cost of genetic analyses places detailed analyses of genotypic history out of reach for many archaeologists. Archaeological research into breed development is therefore reliant upon detecting morphological change in domesticates. In order to accomplish this effectively we must understand the underlying processes that result in natural patterns of morphological variability and the layered-on changes resulting from artificial selective processes.
Natural Morphological Variation
The physical appearance and characteristics of species emerge through long-term evolutionary adaptations capturing the interaction between taxa and their environments and are tied to genetics, life history, nutrition, and sex. Though all members of a given species share extreme similarities in morphology, variation within, or between, populations belonging to the same species arise as populations adapt to local level environmental conditions over long periods of time. Individuals exhibiting advantageous characteristics pass these along to their offspring. Over time these advantageous characteristics may become fixed within populations. Many of these factors have been explored previously in ecological and zoological studies and have been used to develop and test several ecogeographic rules describing widely observed patterns in morphological variation. These
20 rules pertain to trends in species’ or population level trends in body size or appearance and include:
Allen’s Rule: Organisms living near the equator are more likely to exhibit longer, more slender limbs than those living in colder climates (Allen 1877).
Bergmann’s Rule: Within a widely distributed geographic taxonomic clade, species and populations will be distributed such that smaller-bodied species or populations will be found in warmer environments, while larger-bodied species or populations will be found in colder ones (Bergmann 1847).
Gloger’s Rule: Populations of taxa living near the equator where individuals are exposed to higher temperatures, humidity, and greater ultraviolet radiation tend to be more heavily pigmented, while less pigmented forms will be found in cooler and drier environments near the poles (Gloger 1833).
Allen’s, Bergmann’s and Gloger’s Rules document variation in populations frequently correlated to latitudinal gradients with larger-bodied, stockier, and paler populations found near the poles and slimmer, longer limbed, often darker populations found closer to the equator (Figure 1-3). Allen’s and Bergmann’s Rules describe correlations between body proportions which are believed to most effectively maintain body temperature. Body temperature regulation presents significant energetic challenges in climates where average daily temperatures are frequently colder or warmer than normal
21 body temperature. Larger bodies with stockier limbs minimize the surface area through which heat may be lost, while maximizing internal volume thereby enabling populations living in colder environments to more efficiently conserve body heat (Blackburn et al.
1999; Bergmann 1847). In warmer climes keeping the body from overheating may have resulted in longer limbs and smaller body-sizes as this confers a selective advantage by allowing individuals to regulate temperature through increased surface area (Blackburn et al. 1999; Bergmann 1847).
Figure 1-3: The effects of Bergmann-Allen’s and Rensch’s Rules.
Coat color is most commonly explained as a camouflage adaptation in the scientific literature (see Caro 2005) but has been linked to a variety of other adaptive processes.
Studies have found that Gloger’s Rule frequently explains species’ predominant coloration but does not necessarily explain aspects of coloration related to age, sex, or individual specific characteristics (Roulin and Randin 2015). Gloger’s Rule asserts that taxa with lighter pigmentation will more commonly be found in cooler and drier environments -often those near the poles, while those of darker pigmentation will be located in warmer and wetter environments including those found near the equator (Gloger 1833). A classic
22 example of Gloger’s Rule is found in hare species. When confronted with a predator, hares will freeze in place, relying on their coloration to blend into the background before bolting as the predator approaches. The arctic hare (Lepus arcticus) exhibits white coloration which blends into snow, while other hare species including the European hare (Lepus europaeus) or the Antelope jackrabbit (Lepus alleni) are brown or tan in color (Millien et al. 2006). Similar findings have been reported in wolves (Gipson et al. 2002), a variety of bird species (Burtt and Ichida 2004; Delhey 2017), and also plant species (Koski and
Ashman 2015).
One of the strongest and most widely studied deviations from camouflage-based coat coloration is the potential for coloration to convey thermodynamic or energetic advantages. However, the thermodynamic properties of hair and feather pigmentation and form are still poorly understood (see Walsberg 1983). Experimental comparisons of black and white goats from Bedouin herds in the Negev and Sinai deserts have shown that black goats lose significantly more heat through evaporation than their white counterparts despite gaining more heat from their environment (Finch et al. 1980). Similarly, Hassanalian et al.
(2017) have recently reported that birds’ widely shared adherence to dark coloration on the top surfaces of wings, and light surfaces on the lower surface increase the energetic efficiency of flight by selectively influencing air speed and creating lift.
Trends in overall body size and appearance can be compounded by morphological differences between male and female individuals of a given species. Sexual size dimorphism, or differences in the mean body size between male and female individuals of a single species, characterizes nearly all species. Sexual dimorphism can take many forms including differences in coat color or patterning, horn/antler presence, size, or body size or
23 proportions. Distinctions can be made between male and female individuals of many species by the presence of sex specific elements/organs (e.g. bacculi in dogs, antlers in deer) or element morphology (Hatting 2017; Ruscillo 2003, 2006). Patterning in sexual dimorphism is described in:
Rensch’s Rule: Sexual size dimorphism becomes increasingly pronounced as species increase in size when males are larger but decreases in species where females are larger
(Abouheif and Fairbairn 1997; Rensch 1960).
Rensch’s Rule is traditionally tested by log transforming and plotting male and female body size against one another. The slope of trend lines generated for these comparisons provide a measure of sexual dimorphism with those significantly greater than, or less than one indicating strong dimorphism (Abouheif and Fairbairn 1997; Bidau and
Martinez 2016, 2017; Frynta 2012). Applications of Rensch’s Rule have found that 85% of mammalian species conform, but that sexual dimorphism is most extreme among members of a single species near the equator (Bidau and Martinez 2016). Additional studies have shown that males are frequently more variable in their size and morphology than females (McPherson and Chenoweth 2012; Zeder 2001).
Widespread conformity to ecogeographical rules among taxa (Table 1-2) makes them a useful foundation to develop hypotheses regarding general species morphology and the effects of domestication. Deviation from these rules most commonly occurs on a local level and is frequently explained by local environmental conditions. The island effect describes a widely observed phenomenon in which large bodied taxa living on islands
24 decrease in average size, while smaller bodied taxa increase (see Foster 1963, 1964, 1965;
Lomolino 2005; Van Valen 1973). Predation intensity has also been linked to changes in body size (Meachen and Samuels 2012). Coyotes (Canis latrans) abruptly decreased in size following the end of the Pleistocene (Meachen and Samuels 2012) due to increased predation by wolves (Canis lupus) whose population increased to fill ecological niches previously filled by extinct dire wolves (Canis dirus) (Meachen and Samuels 2012).
Table 0-2. Adherence of various taxa to Bergmann’s and Rensch’s Rules.
Bergmann's Rule Resch's Rule* Taxa % Species Tested % Species Tested Birds 76 100 85 686 Lizards 30 50 56 151 Snakes 21 33 0 8 Turtles 83 23 13 71 Mammals 71 149 62 (78) 673 (467) Salamanders 72 18 na na Frogs/Toads 62 16 0 650 Fishes 28 18 na na *Estimated from Abouheif and Fairbairn (1997) () when limited to species larger than rabbits
Domestication Syndrome and Morphological Changes in Domesticated Animal Species
Early domestication processes are believed to have been characterized by sustained selection for behavioral characteristics, especially reduced fight or flight responses and docility; however, extensive changes to species’ physical attributes and morphology did occur (Zeder 2012). Domesticates commonly exhibit weakened eyesight, smaller body size, smaller brain size, and neotony –the retention of juvenile characteristics into adulthood (Zeder 2012). Domesticated species also commonly exhibit piebald, or black
25 and white, coloration not common to their wild progenitors (Zeder 2012). The emergence of black and white coat color within domesticates may have been encouraged because such individuals were recognizable and unique. Together, these physical characteristics have been termed the “domestication syndrome” (Wilkins et al. 2014; Zeder 2015).
Behavioral characteristics are generally hard to observe from the archaeological record. As a result, archaeological studies emphasize detection of changes in domesticates’ physical form visible in their skeletal remains. Alterations to domesticates’ phenotype are layered upon existing patterns of morphological variation found in domesticated species’ wild predecessors. Morphological changes in domesticates were intended to improve their manageability, tailor domesticates’ to specific roles within human societies, or adapt domesticates to breeders’ aesthetic ideals (Clutton-Brock 1981; Zeder 2012, 2015). The long history of human management of domesticates ensures that these changes are often tiered and multifaceted. The result of these processes was the development and formation of breeds that we recognize today.
Archaeological studies of early domesticates frequently assert that the early phases of domestication were characterized by an overall reduction in body-size reflecting efforts to produce animals who were easier to manage (Bökönyi 1976; Degerbøl and Fredskild
1970; Meadow 1989; Uerpmann 1978). The ecological and archaeological literature show that this situation may be complicated by several factors. First, the apparently diminished size of early domestic goats may have resulted from the smaller size of individual animals but may also result from changes in the sex ratio of wild to domestic herds, which frequently have fewer large males and thus smaller average sizes (Zeder 2001). Secondly, tests of Rensch’s Rule regarding sexual size dimorphism in modern domesticates have
26 discovered that domesticates frequently exhibit reduced sexual size dimorphism than their wild progenitors (Polák and Frynta 2009; 2010, see Figure 1-4). Sexual size dimorphism among wild ovicaprids (sheep and goats) results from delayed sexual maturation in males
(Côté and Festa Bianchet 2001) coupled with influences from nutrition and individual life history (Toïgo et al. 1999; LeBlanc et al. 2001). Polák and Frynta (2009) attribute decreased sexual dimorphism in domestic ovicaprids to a combination of 1) relaxed intrasexual selection and the reduced importance of male combat under human management, 2) artificial selection against males achieving sizes large enough that they would be difficult to manage, and 3) reduced sexual segregation through the pasturing of mixed herds including males and females.
Figure 1-4: A comparison of sexual size dimorphism in wild and domestic sheep (Ovis spp.) and goats (Capra spp.). Data from Polák and Frynta 2009.
27 It should also be noted that archaeologists seeking to test reductions in body mass or body size among early domesticates have rarely considered the potential effects of natural variation among wild populations when seeking comparative samples. As an example, in a foundational article on the subject Degerbøl and Fredskild (1970) demonstrate that cattle recovered in Neolithic assemblages from Denmark are smaller than
Danish aurochs. The smaller body size of Neolithic domestic cattle relative to Danish aurochs has been widely accepted as evidence that domestic cattle truly are smaller than wild aurochs (see Degerbøl and Fredskild 1970). However, Wright and Viner-Daniels
(2015) have identified a gradient of size variation in European aurochs, supporting
Bergmann’s Rule with Northern European aurochs being significantly larger than those recovered from contexts in Spain, Portugal, or Italy. Because cattle were domesticated from likely smaller aurochs present in the Fertile Crescent, changes in body size between wild and domestic cattle are probably exaggerated through comparison to already larger aurochs populations. A more effective test of body size reduction would be to compare domestic cattle to Middle Eastern aurochs.
These data suggest a complicated suite of processes may be behind apparent reductions in body size in domesticates. Overall body mass reduction is clearly present in some species. Early domestic dogs recovered from many regions of the world are significantly smaller than wolves (Davis and Valla 1978; Degerbøl 1961; Germonpré et al.
2009; Morey 1994; Pionnier-Capitan et al. 2011). In both cases reductions in body size are readily apparent by the Neolithic. Smaller bodied ovicaprids may have been subjected to less overall body mass reduction, and modern herds do suggest that higher numbers of females would have been kept as they are more significant to the perpetuation of the herd
28 than males, but also as a source of milk (Bogucki 1988 McClure et al. in review). However, efforts to reduce size may have been especially focused on males which are frequently larger and harder to manage (Parés-Casanova 2015; Polák and Frynta 2009; Zeder 2001).
In this way patterns attributed by Zeder (2001) solely to the increasing number of females in herds, may also have involved the decreasing size of rams present in herds.
Changes have also been widely noted in many domesticates’ coat color. The piebald, black and white, coat color common to many domesticates is not widely found in their wild progenitors (Hemmer 1990:121-130; Keeler et al. 1968; Zeder 2012).
Mummified dog remains from the American Southwest have revealed black and white coats testifying to the widespread nature of this characteristic (Olsen 1974). Coat color is controlled by a complex interaction of many genes (Montoliu, Oetting & Bennett 2009).
Genetic analysis of horses has revealed that coat color variants emerged relatively quickly in horses following their domestication (Ludwig et al. 2009). Taming experiments with rats and foxes have also recreated the emergence of patched white and black coat coloration
(Albert et al. 2009; Trut et al. 2004, 2009), but have found no close association between genes linked to pigmentation and those linked to tameness (Albert et al. 2009). Coat color and breed emergence are highly linked. Many modern breeds are characterized by homogeneity in their coat color, or the patterning of their coat coloration (e.g., red Devon and brown Swiss cattle, yellow and chocolate Labradors).
In addition to the effects of natural evolutionary processes, and the phenotypic effects of the domestication syndrome, artificial selection has created unique morphotypes not found in natural populations and led to significant phenotypic changes in domesticated species long after their domestication. There are a number of completely artificial
29 phenotypes that exist only in artificial systems. The most widespread of these are created through castration. Castration alters the production of hormones in males and is often used to make male herd members, who are not needed for breeding purposes, more docile and allowing them to be maintained for meat, traction, or other purposes. Castrated male cattle, called oxen, do not achieve the full adult body size of male cattle, and have differential deposition of fat and muscle tissue. Oxen have been widely used in preindustrial societies for plowing fields and hauling cargo. Müller (1964) asserts that castration has been used in cattle management since the Neolithic, though carts or plows appear to develop in the Late
Neolithic or Bronze Age (Anthony 1995, 2010; Kristiansen and Larsson 2005; Milisauskas and Kruk 1991). Draught oxen are a morphologically intermediate size class falling between male and female cattle in assemblages when morphometric data are plotted
(Thomas et al. 2013). Similar results have been achieved with wethers, or castrated male sheep (Armitage and Goodall 1977; Davis 2000).
Domesticates as species, integrating natural and artificial selection
As distinct populations are subjected to relaxed natural selection and increased artificial selection relative to their wild counterparts, they attain morphotypes not found among wild populations. Populations belonging to any given species will frequently diverge from one another in physical form and genetic composition when isolated and exposed to differential selective pressures (e.g., Darwin’s finches). Species are commonly defined as populations that cannot interbreed. Many domesticates can interbreed with their wild progenitors (Clutton-Brock 2012) and may even be encouraged to do so (Buffalo Bird
30 Woman in Wilson 1924). As a result, some classify domesticates as sub-species or forms of their wild progenitors (Dennler de la Tour 1959, 1968; Herre and Röhrs 1990, e.g., Canis lupus familiaris). Others suggest that domestic populations require special treatment because sustained human interference frequently propagates those individuals which would be removed from natural populations, and results in significant genotypic and phenotypic changes in domestic species (e.g., Harlan and de Wett 1971). For the purposes of this dissertation I adopt the latter system, classifying domesticates as species.
I classify domesticates as distinct species because, though many domesticates can interbreed with their wild progenitors, they are also morphologically and behaviorally distinct. Artificial selection occurs alongside natural selection but acts as the dominant driver of population adaptation in domesticates (Driscoll et al. 2009; Giovambattista 2001;
Trut 1999). The relaxation of natural selective pressure acting upon domesticates has likely reduced the degree to which translocated populations respond to local environmental conditions; however, artificial selection frequently acts by exaggerating characteristics within a range of possibilities already defined by existing natural selection (Drake and
Klingenberg 2010; von Schantz et al. 1995). Anthropogenic selection for specific behaviors or characteristics may mitigate the natural selective pressure acting upon populations; however, the modification of any species through anthropogenic selection rarely produces domesticated animal species incapable of their own survival.
A result of these processes is that it is unclear how strongly domesticates adhere to ecogeographical rules such as Allen’s and Bergmann’s Rules. Domestic populations’ phenotype can change rapidly at the whim of human managers. To date, few analyses have attempted to test this assumption. Jordana et al. (1995) suggest that the morphology of 22
31 horse breeds follow expectations drawn from Allen’s and Bergmann’s Rules. Similarly,
Cassaing et al. (2011) tentatively suggest that house mice, a commensal domesticate, recovered from Gallo-Roman to Medieval sites in Europe and Africa appear to follow
Bergmann’s Rule. Unfortunately, their analysis relies solely on teeth, which form early in development and are not very responsive to environmental conditions (Cassaing et al.
2011). In addition, unique morphotypes may be obtained by hybridizing or cross-breeding related species, for example horses and donkeys. Offspring from these cross-breeds are generally infertile, in the case of mules and hinnies this is caused by imperfect alignment of the chromosomes. Mules and hinnies are rare in the wild but have at times been encouraged within human management. Mules were especially popular as pack animals in the Roman army and remained in common use as pack animals into the 1700-1800’s because they are generally as large as horses but more resilient (Clutton-Brock 1981).
Classifying domesticates as unique species acknowledges the significant impacts of human interference in domesticated species’ existence and morphology, but also results in a more simplistic and straightforward classification scheme for breeds. Populations of wild plant and animal species that belong to the same species but exhibit small and persistent morphological differences are classified as subspecies (Clutton-Brock 1981;
Darwin 1859). Breeds are similarly defined as those animal populations that have been bred by humans to possess uniform characteristics that are passed down through generations and distinguish the animals from other animals within the same species
(Clutton-Brock 2012:3). The defining criteria for both domesticates and breeds is thus some level of human interference in natural selection, and subspecies and breeds are
32 effectively equal taxonomic classifications describing phenotypically, and at times genotypically, distinct populations that are recognized as belonging to a single species.
When Will Breeds be Detected in the Archaeological Record
Breeds are defined on the basis of a shared physical morphology which differentiates them from other members of the same species. Many characteristics used to differentiate one breed from another are difficult to detect in an archaeological record obscured by both taphonomic processes and a lack of information regarding the number of breeds present. Archaeologists seeking to identify morphologically distinct populations of domesticates and use these in understanding past communities are largely limited to what is visible in skeletal remains.
As a result, efforts to identify breeds of morphologically distinct populations from the archaeological record will only be possible under certain conditions. First, when distinctive or extreme morphological traits exist in populations which are reliably identifiable in archaeological assemblages (Bennett et al. 2016). For example, Bennett (et al. 2016) identifies several breeds or types in the Vindolanda assemblage based on physical morphology including small dachshund-like humeri. Second, where skeletal differences on a population level exceed sex specific or Bergmann-Allen’s rule-type regional differences. This kind of variation has been used by Albarella (1997) and
Mackinnon (2010) to identify distinct morphotypes of cattle from Medieval and Post- medieval England and the Netherlands and Roman Period Italy.
33 A common characteristic of these studies is the availability of large spatially and temporally controlled samples. Though these samples do identify distinct populations from skeletal material, such archaeological “breeds” may imperfectly correspond to culturally defined breeds based upon characteristics like coat color. Archaeological studies are further limited by characterizing only a small number of morphological characteristics. In addition, few databases containing morphometric data on individuals of known breed exist (though see Phillips et al. 2016, Foster personal communication).
This does however raise an important issue – when will multiple breeds be present, and across what geographical or temporal scale should such analyses be undertaken?
Analysts rarely know how many breeds or types are present within the skeletal assemblage they are working with, but they have a dataset limited by both taphonomic processes and a relative poor comparative dataset on modern breeds to work with. These are issues which are as yet poorly resolved; however, many communities in developing nations possess relatively few distinct local breeds of any one livestock species (see Rege 1999, 2003; Rege and Tawah 1999; Sponenberg et al. 2014). This implies that many archaeological contexts will contain one or a few locally adapted breeds, rather than many. Further, by considering
Rensch’s and Bergmann-Allen’s Rules we may begin to identify locally adapted breeds, landraces, or types adapted to specific environmental conditions and use these to explore aspects of human subsistence, migration, and culture.
34 Dogs as a Case Study
Dogs are well positioned to serve as a case study for understanding domestication and breed development for several reasons. Dogs are the earliest known domesticated animal species having been domesticated in at least three domestication centers in Europe,
East Asia, and Southwest Asia sometime before 14,000 years BP (Castroviejo-Fisher et al.
2011; Larson et al. 2012; Leonard et al. 2002; van Asch et al. 2013). Dogs became globally distributed in foraging, pastoral, and agricultural societies and have adapted to specialized roles as hunting aids, weapons of war, food, sources of fiber for clothing and textiles, livestock protection, and beasts of burden providing numerous opportunities for breed development (Allen 1920; Crockford 2000; Horard-Herbin et al. 2014; Lupo 2017;
Schwartz 1998). Adaptation to these roles occurred relatively early, by the Roman period or earlier in Europe (Bennett et al. 2016) and has drastically impacted domestic dogs’ physical form producing differences in size (e.g. chihuahuas and great danes), physical proportions (e.g. dachshunds and whippets), and behavioral patterns (Mehrkam and Wynne
2014). In addition, canids in general exhibit relatively weak sexual dimorphism with domestic dogs averaging a score of only 1.18 for body mass, and 1.06 for shoulder/wither height (Bidau and Martinez 2016; Frynta et al. 2012; Sutter et al. 2008).
Dog’s early domestication, global dispersion, and abundant roles within human societies, are complemented by the attention they have received from archaeologists in many regions of the world. Extensive interest in domestic dogs among archaeologists has resulted in the development of numerous techniques for detecting and characterizing physical attributes including wither height (Clark 1996; Harcourt 1974), robusticity (de
35 Grossi Mazzorin and Tagliacozzo 2000), body mass (Losey et al. 2015; Losey et al. 2017;
Van Valkenburgh 1990; Wing 1978), and breed affinity (Phillips et al. 2008; Tourigney et al. 2016) on a scale not seen in any other domesticate. The history of research focused on dog domestication and human cohabitation by past and current archaeologists has also caused the development of datasets that dwarf those available for other species, domesticated or otherwise (see Crockford 1997; Haag 1949; Harcourt 1974; Worthington
2008).
Dissertation Organization and Research Questions
This dissertation builds upon the foundation established by prior studies to address gaps in our knowledge concerning both the implications of breed development and use in past societies. This dissertation includes three papers dovetailing datasets drawn from biomorphometric analyses, ethnographic and historic documents, and theoretical models drawn from Human Behavioral Ecology (HBE). These chapters will individually address the emergence of breeds, their adaptation to specialized roles within human societies, and their potential to serve as markers of human migration. Through developing an understanding of when and where breeds have emerged we can begin to understand both the roles that domesticates have played in human societies and the cultural and environmental criteria that make breed development possible, questions that are integrally linked to some of archaeology’s grand challenges as defined by Kintigh et al. (2014).
The domestic dog is uniquely positioned to serve as a test species for this research for many reasons. Dogs were among the earliest and most widely distributed domesticates
36 (Castroviejo-Fisher et al. 2011; Larson et al. 2012; Leonard et al. 2002; van Asch et al.
2013) and have become adapted to many highly specific roles within forager, agricultural, and pastoral communities (Allen 1920; Crockford 2000; Horard-Herbin et al. 2014; Lupo
2017; Schwartz 1998). As a result of these factors dogs have come to exhibit an extraordinary diversity of physical characteristics, but also exhibit relatively weak sexual dimorphism (Bidau and Martinez 2016; Frynta et la. 2012; Sutter et al. 2008). Historical and ethnographic accounts provide an invaluable record of dogs’ physical characteristics and roles in European and Native American communities in the recent past. I acknowledge the strength of this record and employ a direct historical approach (Deal 2017; Lyman and
O’Brien 2001; Steward 1942; Strong 1940) to develop hypotheses about the physical attributes and roles of domestic dogs in past societies which can be quantitatively tested using archaeological remains dating to both the historic period and those of greater antiquity.
Biomorphometrics –the measurement of skeletal landmarks– is one of the strongest and most reproducible methods available to archaeologists (Albarella 1994) and provides a powerful means of characterizing and quantitatively studying variation in physical shape and proportion. By following established protocols for biomorphometric analyses (Haag
1948; von den Driesch 1976) I have assembled large datasets to characterize the degree of physical variability in dog populations. These data are then integrated with experimental and ecological data to quantitatively test hypotheses based directly upon historic and ethnographic sources.
With this dissertation I work from the confluence of ethnographic and historic sources and archaeological data to formulate hypotheses about the physical attributes and
37 role of dogs within past societies, and through these the development of dog breeds. I then address these questions by characterizing the physical variability and capabilities of dog populations in past European and Native American communities using morphometric data collected on domestic dog skeletal remains in archaeological assemblages. By integrating the direct historical approach with quantitative and highly reproducible methods, this dissertation takes a significant first step towards utilizing breed development as a tool for understanding the legacy effects of long-term cohabitation between humans and domesticated animals. The questions addressed by this dissertation include:
1) Can dog populations originating in Europe and North America be morphologically
differentiated from one another and thereby serve as markers of migration and
colonization processes?
The introduction of domesticated species to new environments has been used to identify colonization events in the archaeological record, but rarely provides the opportunity to investigate colonists’ selection of particular breeds, stock, or subsequent adaptations to environmental conditions. Chapter Two employs morphometrics, the measurement of skeletal landmarks, and analysis of historic documentation to explore the effects of divergent selective breeding processes on the physical characteristics of domestic dogs (Canis familiaris) in Native American and European societies. Specifically, this analysis investigates whether or not European colonists’ preference for large working dogs resulted in the importation of dogs exhibiting identifiable morphological characteristics which can serve as markers of population movement and interaction. Historic sources
38 suggest that European colonists selectively imported large dog breeds capable of defending settlements and livestock, acting as war dogs, and aiding hunters. Colonists’ dogs are reputed to have been significantly larger than Native American dogs. This study compares standardized measurements taken on bones to estimate shoulder height. We find that dog populations in England and North America spanning the period 0-1800 AD exhibit an almost identical average stature; however, the range of shoulder height variation in
European dogs is far greater than seen in eastern Native American dog populations. Dogs in colonial American contexts are statistically larger than both Native American and
European dog populations, supporting documentary accounts that colonists expressed a strong selective preference for large body size, and selectively imported some of the largest dog breeds available in Europe. This analysis contributes to a small, but growing body of research (Bennet et al. 2016; Horard-Herbin et al. 2014) which shows that morphologically distinct populations of animals associated with specific populations may be used to trace migration and population interaction in the past.
2) Can dogs bred for and used as draught animals by Plains and Intermountain West
Native American communities be identified and differentiated from other Native
American dog breeds, and can recognition of these dogs as a distinct population be
used to investigate how long dogs have been used in Native American transport
activities?
Domestic dogs (Canis familiaris) specifically bred for large size and stamina necessary for load transport were central features of Native American mobility in the
39 Intermountain West and on the Great Plains. Ethnographic accounts highlight their use in hauling travois, simple drag-sleds, and carrying pannier-style packs estimated to weigh as much as 45.36kg (100lb). Chapter Three employs a novel application of body-mass estimation coupled with experimental and modern reference data to show that larger dogs recovered from the Birch Creek Valley in Idaho and sites on the Great Plains were capable of hauling ethnographically reported loads. Furthermore, dating of at least one of these specimens indicates that dogs of similar capability to those documented in historic times have existed in the region for over 5,000 years, suggesting dogs’ role in transport may be at least this old. These data have important implications for our understanding of prehistoric mobility in the Intermountain West and Plains.
3) Under what conditions does it make sense to use a dog in transporting goods?
Dogs in Native American communities from the Great Plains, Intermountain West,
Southwest, and Arctic were adapted to specialized roles in transport through intensive culling of smaller pups, interbreeding with wolves, and investment in transport technology.
Poor preservation of the transport technology (travois, sleds, pannier-style packs), and, until recently, an inability to assess which dogs were capable of ethnographically recorded loads, has inhibited archaeological attempts to identify when such roles began. Further, it is as yet unclear why such roles developed in some communities and not others. Chapter
Four draws upon ethnographic sources, historical accounts, and archaeological evidence to frame Intermountain West and Great Plains dog-based transport within the Field
Processing Model drawn from Human Behavioral Ecology (Barlow and Metcalfe 1996;
40 Bettinger 2009; Bird and Bliege Bird 1997; Metcalfe and Barlow 1992; Price 2016). In doing so this analysis explores both when dogs should be used in transport, and over what distances their use is beneficial, but also represents a foray into the use of HBE to investigate domestication and animal management.
Finally, Chapter Five serves to draw these lines of evidence together and outline the implications and future directions for this research. The emergence of breeds in prehistoric and historic societies reflects significant investment on a community level and may provide insight into the cultural processes and characteristics associated with sustained animal management and niche construction processes. Dogs are the most morphologically variable and most widely dispersed domesticated species in the world.
This morphological variability makes them a valuable case study with which to explore the potential to recognize breed recognition and development in archaeological assemblages, and through doing so contribute to our understanding of the human past.
Together this dissertation explores several avenues through which breed development can be explained and breed recognition can contribute to our understanding of human migration and trade networks.
Chapter 2
Breed Selection and Colonization: Exploring the Potential for Breeds to Act as Markers of
Population Movement1
The introduction of domesticated species to new environments has been used to identify colonization events in the archaeological record, but rarely provides the opportunity to investigate colonists’ selection of particular breeds or stock. This analysis employs morphometrics, the measurement of skeletal landmarks, supported by historic documentation to explore the intersection between colonists’ breed preferences and species translocation. In doing so, this analysis provides insight into the differential effects of human selective breeding on domestic dogs’ (Canis familiaris) physical characteristics in Native American and European societies. Historic sources suggest that European colonists selectively imported large dog breeds capable of defending settlements and livestock, acting as war dogs, and aiding hunters.
Colonists’ dogs are reputed to have been significantly larger than Native American dogs. This study compares standardized measurements taken on bones to estimate shoulder height. We find that dog populations in England and North America spanning the period 0-1800 AD exhibit an almost identical average stature; however, the range of shoulder height variation in European dogs is far greater than seen in eastern Native American dog populations. Dogs in colonial
1 Co-authored with Rebecca Dunham, Parks Canada senior archaeologist at the Fortress of Louisbourg National Historic Site, Louisbourg, NS, Canada
42 American contexts are statistically larger than both Native American and European dog populations, supporting documentary accounts that colonists selectively imported breeds from the largest available in Europe.
Introduction
Archaeological migration events are commonly identified using combinations of artifacts associated with a particular population or culture termed “archaeological cultures”. Despite this, understanding and contextualizing migration events remains among the “Grand Challenges” for archaeology (Kintigh et al. 2014). The appearance of domesticated plants and livestock in archaeological assemblages is employed in similar ways as a widely accepted hallmark demarcating the arrival of agriculturalists during the European Neolithic. Morphometric methods provide a means of achieving a greater resolution in such analyses. Morphologically distinct domestic dog (Canis familiaris) populations have successfully been used to trace migration by
Neolithic farmers (Horard-Herbin et al. 2014). In both cases independent evolutionary trajectories produced distinctive physical characteristics which can be used to identify introduced dog populations. Because dogs existed in both the New and Old Worlds prior to the Columbian
Exchange (Bökönyi 1974; Brown et al. 2013; Clutton-Brock 1984; Tchernov & Valla 1997;
Tamm et al. 2007; Witt et al. 2015) because they provide an opportunity to test whether
European and Native American dog remains can be used in a similar manner.
A preference for large breeds appears to have remained consistent throughout the early colonial period (Mastromarino 1986; Meacham 2011; Varner and Varner 1983). Defined breed standards for dogs and other livestock emerged in the 18th and 19th centuries (Arman 2007);
43 however, historic documents indicate European colonists in North America preferred Old World dogs (Derr 2004), especially bloodhounds, greyhounds, and mastiffs that could be used to hunt and defend livestock from New World predators (Jefferson 1791; Lafayette 1785; Washington
1785a, 1785b). European colonists also recognized that large dogs were an intimidating deterrent against rival colonists and potentially hostile native peoples and continued to use mastiffs as war dogs until the 1750s (Mastromarino 1986; Meacham 2011; Varner and Varner 1983).
Comparison of dogs’ shoulder height (Clark 1996; Harcourt 1974) generated from greatest length measurements taken on long bones in faunal assemblages from North American colonies and Europe are used to test colonists’ preference for large dogs, while comparisons of colonial and Native American dog bones assess whether European dogs, especially those brought over by the colonists, were truly larger than indigenous dogs in Eastern North America. We present morphometric data on domestic dog skeletons from North American colonial period assemblages, including 18th century dog skeletons from the King’s Bastion of the Fortress of
Louisbourg (Cape Breton, Canada). These are compared to a dataset of shoulder height estimates from Native American and European dogs dating from 0-1800 AD compiled from the published and unpublished literature. This analysis explores whether the historically documented European preference for Old World breeds led to detectable differences in shoulder height between dogs in colonial and Native American contexts in Eastern North America.
44 Background:
Colonial Dogs:
Historic documents indicate that European colonists introduced dogs to the New World to herd livestock, defend settlements, track game and runaway fugitives or slaves, and act as war dogs for intimidating, tracking down and attacking hostile Native Americans and other European colonists (Derr 2004; Mastromarino 1986). Significant bodies of documentary evidence for this period indicate greyhounds, bloodhounds, and mastiffs were among the most commonly referenced breeds (Hawes 1633; Jefferson 1791; Lafayette 1785; Meacham 2011; Washington
1785a, 1785b). Mastiffs accompanied English colonists who established Jamestowne in 1585 and pilgrims aboard the Mayflower in 1620 (Derr 2004). Ownership of specific types of dog was likely limited to wealthy merchants and landholders who used their contacts in Europe to acquire dogs of known breeding lineages (Hawes 1633; Jefferson 1791; Lafayette 1785; Meacham 2011;
Washington 1785a, 1785b). Poorer colonists frequently owned mongrels of indeterminate breed, but still of European origin and likely of large size (Meacham 2011).
Spanish and English use of war dogs is well established. Columbus employed war dogs against the Caribbean Taino (Derr 2004). Dogs also accompanied the Spanish De Soto (1539-
1543) and Coronado expeditions (1540-1542) (Derr 2004; Purdy 1977) and were used to defend the New England colonies during King Philip’s (1675-1677) and Queen Anne’s (1702-1713)
Wars (Bodge 1891; Derr 2004). As late as the French and Indian (Seven Years) War (1754-
1763) military leaders discussed the possibility of arming English soldiers with mastiffs to track down the hostile Shawnee and Delaware (Bouquet 1764; Hughes 1763). European colonists
45 frequently sought to prevent Native Americans from acquiring European dogs in order to maintain the dog’s role as a deterrent. English colonial governments including Virginia and
Connecticut banned the trading of “English dogs of quality” to the Native Americans in 1619 and 1650, fining between five shillings and ten pounds per offence (Andrus 1830; Derr 2004).
Attempts to manage the trade of livestock, including dogs, to Native Americans ended in the late
17th century (Derr 2004). Settlers on the frontier are known to have eventually interbred Native
American dogs and European breeds producing the “yellow cur”, a hunting dog which is mentioned in records from the frontier by the 1770s (Derr 2004).
Dogs in England and Europe:
Dogs were domesticated sometime before 15,000 BC (Benecke 1987; Clutton-Brock
1984; Pionnier-Capitan et al. 2011; Sablin & Khopachev 2002; Tchernov & Valla 1997).
Chronological comparisons of shoulder height in European populations reveal dogs in early agricultural societies during the Neolithic (8,000-4,000 BC) to be 40 – 50 cm in height (Clark
1996, 2006; Harcourt 1974; Horard-Herbin et al. 2014). Dogs of this size persisted through the
European Bronze and Early Iron Ages (Clark 1996, 2006; Harcourt 1974; Horard-Herbin et al.
2014). Increasing diversity in shoulder height emerged during the Late Iron and Roman period and is frequently linked to Roman trade and occupation (Bennett et al. 2016; Horard-Herbin et al. 2014).
Though dogs have been under intensive artificial selection for characteristics suited to specific purposes in human society for thousands of years, the definition of breed standards and development of most modern breeds occurred in the 18th-19th centuries (Larson et al. 2012; Pang
46 et al. 2009; Savolainen et al. 2002). Prior to the development of modern breeds, dogs in England were divided into three functional classes: game (hunting dogs), homely (working dogs), and currish (mixed breeds) (Caius 1880). Although small breeds were known (Harcourt 1974;
Horard-Herbin et al. 2014), O’Connor (1992) notes that many Medieval and Post-Medieval dogs were similar in stature to the modern border collie (60-66 cm, American Kennel Club, 1977).
These dogs were used to guard homes or livestock, control pests, hunt wild game, and act as pets. Cut marks identified on dogs from medieval and post-medieval contexts in England are believed to indicate knackering, the use of dead dogs as food for other livestock (Thomas &
Lacock 2000; Wilson & Edwards 1993), or the collection of hides for tanning (Albarella 1999;
Thomas 2005).
Native American Dogs:
The oldest dog remains in North America date to between 9,000 and 10,000 BP (Grayson
1988). Some have suggested the absence of dog remains older than this date implies independent domestication from North American wolves (Leonard et al. 2002); however, aDNA analysis has shown that Native American dogs are descended from East Asian wolves and belong to at least
14 founder haplogroups associated with a unique genetic clade (Leonard et al. 2002; Tamm et al.
2007; Fagundes et al. 2008; Achilli et al. 2008, 2013; Perego et al. 2009; Malhi et al. 2010;
Hooshiar Kashani et al. 2012). Genetic diversity recorded in aDNA analyses reveals Native
American dog populations were subject to varying levels of selective pressure and at times interbred with North American wolf populations on Native American dog populations (Leonard et al. 2002; Witt et al. 2015).
47 Native American dogs served as sources of hair fiber and food, aided hunters, and pulled travois and sleds (Schwartz 1998). Morphometric analyses indicate that Native American dogs generally increase in size through time (Haag 1948). Historical descriptions of Native American dogs highlight pointed ears and wolfish coloration, and suggest many were relatively small, terrier-sized, animals (Lewis 1806). Documentary evidence reveal widespread efforts to extirpate indigenous breeds by European colonists (Derr 2004). These, and intentional and accidental interbreeding along the frontier (Derr 2004) contributed to loss of genetic integrity in Native
American dog populations. This interbreeding is evidenced by the close relatedness found when comparing modern New-World and European breeds (Leonard et al. 2002).
Materials and Methods:
Three mostly complete dog skeletons (provenience: 1B3B87, 1B43D1, and 1B4T8,
Figures 2-1 and 2-2) were recovered during the excavation of 18th century French or British contexts at the Fortress of Louisbourg (Howard 1962; Marwitt 1966). All three dogs appear to be intentional internments of dogs in the casemate or moat of the King’s Bastion Barracks and cannot be attributed with confidence to either French or English occupations of the site (Howard
1962; Marwitt 1966). Dogs 1B3B87 and 1B43D1 possessed complete, permanent dentitions and full epiphyseal fusion, making them over a year-and-a-half at death (Silver 1969). No cut marks, intentional fracturing, or burning were noted; however, 1B3B87 suffered a displaced, complete fracture of the left femur which had never been properly set and healed crookedly (Figure 2-2).
48
Figure 2-1: The location of the Fortress of Louisbourg (top left). Fortress of Louisbourg site overview (top right). Locations where domestic dog remains used in this analysis were recovered by Marwitt (1966) and Silver (1969) (bottom).
Morphometric measurements were collected on adult individuals 1B3B87 and 1B43D1 using an osteometric board and digital calipers following von den Driesch (1976). Shoulder height estimates were calculated using greatest length measurements taken on complete long bones for both specimens following Harcourt (1974) (Table 2-1). 1B4T8 was under seven months old with incomplete permanent dentition and no fused epiphyses (Silver 1969) at death.
Measurements taken on an unfused right radius generates a shoulder height of 33.63cm;
49 however, equations designed for estimating height in adult dogs (Harcourt 1974) may not accurately predict the size of juvenile animals (Table 2-2). As a result, IB4T8 was not used in this analysis.
Figure 2-2: Excavation photos for the Fortress of Louisbourg dogs (Marwitt 1966; Silver 1969), and photos showing the severe displaced fracture suffered by dog 1B3B87.
50
Table 2-1. Equations used to estimate shoulder height from long bone greatest length (GL) measurements by element (Harcourt 1974). Element Equation Humerus (3.43*GL) -26.54 Ulna (3.18*GL) +19.51 Radius (2.78*GL) +6.21 Fore Limb (Humerus + Radius) (1.65*GL) -4.32 Femur (3.14*GL) -12.96 Tibia (2.92*GL) +9.41 Hind Limb (Femur +Tibia) (1.52*GL) -2.47
A comparative dataset of dogs from Spanish and English colonies (n=10), Native American dogs from eastern North America (n=95), and European dogs (n=165) from contexts dating to between 0-1800 AD was assembled (Data S1). Shoulder height estimates were generated using conversion factors for greatest length measurements of long bones (Harcourt 1974). The colonial dogs span the period 1639-1800 (Table 2-3, see also Figure 2-3).
Though this sample largely post-dates active efforts to ban the trade of dogs between
Europeans and Native Americans, documentary records suggest preference for Old World breeds remained high (Hawes 1633; Jefferson 1791; Lafayette 1785; Meacham 2011; Washington
1785a, 1785b). All dogs in the Colonial sample except those from the Sylvester Manor and St.
Augustine recovered from intentional internments (Bostwick 1980; Sportman 2003). A complete adult female dog was recovered from a destruction feature at Ferryland, Newfoundland, known to have occurred in 1696 (Gaulton personal communication). Additional specimens were recovered from a palisade feature at Fort Shirley (Welker under review), and a feature under the floor of the Anderson Armory in Colonial Williamsburg (Wagner et al. 2014).
51 The Native American sample is drawn largely from the dataset compiled by Worthington
(2008) for the southeastern U.S. with additional published data to include areas north to Canada
(Bathurst & Barta 2004; Lawler et al. 2016), and south to the West Indies (Grouard et al. 2013) and Mexico (Galacia et al. 2001). The European sample is composed of dogs drawn from the
Animal Bone Metric Archive Project (ABMAP) managed by the University of Southampton and other published and unpublished sources. European and North American samples were plotted chronologically to ensure that neither sample was unduly influenced by significant changes in dog height over the sampling period.
Figure 2-3: The distribution of dogs for which morphometric data exists and date to the Colonial Period (Numbering follows Table 2-3).
52 Table 2-2. Louisbourg dog measurements and shoulder height estimates. Specimen Element Greatest Shoulder Length (mm) Height (cm) 1B3B87 Left Humerus 176 57.71 Right Radius 175 57.60 Right Femur 195 59.93 Right Tibia 195 57.88 Left Tibia 198 56.52 Right Hind Limb 390 59.03 Average 58.11 1B43D1 Right Humerus 177 58.06 Right Femur 202 62.13 Right Tibia 207 61.39 Right Hind Limb 409 61.92 Average 60.87 1B478* Right Radius 111.89 33.63 * Estimated from unfused shaft alone –not used in analysis.
To study the morphometric differences in dogs from European, Colonial, and Native
American contexts, historic documentation was used to develop several expectations which could be tested with the data available for this analysis. Historic documentation suggests
European colonists intentionally selected large breeds to import into North America, and that these were frequently larger than Native American dogs in Eastern North America (Derr 2004).
If this is the case, shoulder height estimates from colonial dogs should be larger than those from
Native American sites. Additionally, because European colonists were selecting working dogs capable of fulfilling specific guarding, hunting, and military roles, these dogs may be expected to fall within the size range present in the source population in Europe but may disproportionately reflect the high end of the available range. ANOVA analysis performed in R was used to test for statistically significant differences in shoulder height within and between Colonial, European, and Native American dogs. An additional two-sample Kolmogorov-Smirnov test performed in R was used to assess whether dogs from these samples could have been drawn from the same
53 distribution –that is, could dogs exhibiting the shoulder heights found in the Native American,
European, and Colonial samples have been randomly selected from a single population.
Table 2-3. Colonial dog shoulder height estimates and dates. Shoulder Specimen Site Date Height (cm) Source Sylvester Manor 1652-? 60* Sportman 2003 Ferryland 1696 64.02 Gaulton & Burchell personal communication St. Augustine 1700-1782 52.50 Bostwick 1980 1B3B87 Fortress of Louisbourg 1750-1768 58.11 This Analysis 1B43D1 Fortress of Louisbourg 1750-1768 60.87 This Analysis Fort Shirley 1753-1756 60.09 Welker et al. under review Anderson’s Armory 1775-1800 49.31 Wagner et al. 2014 Anderson’s Armory 1775-1800 56.45 Wagner et al. 2014 Anderson’s Armory 1775-1800 32.29 Wagner et al. 2014 Anderson’s Armory 1775-1800 53.41 Wagner et al. 2014 Anderson’s Armory 1775-1800 63.28 Wagner et al. 2014 Anderson’s Armory 1775-1800 66.33 Wagner et al. 2014 * Estimated based on description
Results:
Shoulder height estimates generated from archaeological materials support the hypotheses that European colonists’ preference for large working dogs resulted in a distinctive pattern of large dogs in colonial period assemblages. Adult dogs 1B3B87 and 1B43D1 recovered from Louisbourg stood between 58 and 61 cm in height (Table 2-2). Shoulder height estimates generated on dogs from 17th and 18th century colonial contexts in North America reveal that these dogs were similarly large, standing 49.31 cm or taller at the shoulder in all but one case
(Table 2-3, Figure 2-3). Colonial dogs exhibit a markedly different pattern in shoulder height than that seen in either the Native American or European samples (Table 2-4, Figure 2-4).
ANOVA analysis indicates statistically significant differences between Colonial and Native
American dogs (F = 45.43, df = 95, p < 0.000), supporting assertions that colonial dogs were
54 larger than contemporaneous Native American dogs in eastern North America (Bodge 1891;
Derr 2004; Mastromarino 1986). ANOVA analysis also identifies statistically significant differences between Colonial and European dogs (F = 16.31, df = 177, p < 0.000) indicating the colonial dogs are on average significantly larger than the general European dogs.
Table 2-4. Comparative analysis results. Sample Height (cm) Native Min. 30.05 American Avg. 43.78 Max. 61.1 Colonial Min. 32.29 Avg. 56.39
Max. 66.33 European Min. 23.01
Avg. 43.01 Max. 74.00
Figure 2-4: Shoulder height (in cm) in Native American, European, and Colonial dog populations.
55 As expected, Colonial dogs fall into the upper range of shoulder heights present within the European sample (24.64 to 74 cm). The average colonial dog (56.39 cm) is almost 12 cm larger than the average European dog (43.01 cm) (Table 2-4). Colonial dogs are also 12 cm larger than the average Native American dog (43.78 cm) supporting assertions that colonial dogs were larger than those in Native American societies (see Derr 2004). These data also suggest that the average European and Native American dogs were similar in size (Table 2-4, Figure 2-4), a conclusion supported by ANOVA analysis which reveals no statistically significant differences in average shoulder height between European and Native American dogs (F = 0.48, df = 274, p =
0.489). Comparison of the range of shoulder height found in Native American (34.86 to 57.84 cm) and European dog populations reveals that Native American dogs exhibit a smaller range in shoulder height than their European counterparts. Plotting the Native American and European samples chronologically shows that no significant changes in size in Native American dog populations occurred over this period (Figure 2-5), though some evidence suggests the very largest of the English dogs developed relatively late in the study period (Figure 2-6). Notably, the largest of the Native American dogs which was recovered from the Squantum Site in
Massachusetts has been directly dated to AD 1649-1946, as a result it may in fact be a European dog (Chilton et al. 2001). Removal of the Squantum dog from the Native American sample does not appreciably change the results of ANOVA analysis (F = 50.15, df = 94, p < 0.000).
An additional two sample Kolmogorov-Smirnov test was performed to assess whether the
Colonial, European, and Native American samples used in this study could have been randomly drawn from a single population. Comparison of Colonial and Native American (p < 0.000) and
European and Native American (p < 0.000) samples indicates these were drawn from significantly different source populations. A comparison between Colonial and European dogs (p
56 < 0.000) reveals that the chances of selecting only large dogs from the European population to be exceedingly low, unless European colonists were intentionally selecting for large body size in the dogs they chose to introduce to the Americas.
Figure 2-5: Native American dog shoulder height (in cm) plotted against time for the period 0-1800 AD.
Figure 2-6: European dog shoulder height (in cm) plotted against time for the period 0-1800 AD.
57 Discussion:
Much of what we know about post-Columbian species translocation between the Old and
New Worlds, including European dogs, is based on documentary sources (e.g., Derr 2004; Rouse
1973). These sources provide valuable foundational knowledge regarding the origins of livestock, and the challenges faced by colonists in transporting and establishing domesticates in the New World. Though historical accounts are subject to the biases and opinions of the colonists who wrote them, historical archaeology is uniquely suited to testing historical accounts and addressing some of archaeology’s most enduring challenges including the interaction between migrant populations and unfamiliar environments (Cleland and Fitting 1978; Deagan 1988;
Kintigh et al. 2014; Schuyler 1988). Zooarchaeological data from historical contexts enables testing of poorly documented aspects of the relationship between colonists and their domesticates including the genetic origins, selection of species and breeds for importation, and colonial management practices (Cossette & Horard-Herbin 2003; Reitz 1992; Reitz & Ruff 1994).
The European colonization of North America brought into contact two previously isolated populations of domestic dog. The unique evolutionary history of these dogs within
Native American and European societies shaped these populations in distinct ways which may be detectable through comparison of Old and New-World dogs. Historical records suggest European colonists expressed a strong preference for large working dogs, and imported these when possible (e.g., Bouquet 1764; Hughes 1763; Lafayette 1785; Washington 1785a, 1785b;
Jefferson 1791). The data presented here support preferential selection for larger working dogs by colonists in eastern North America (Figure 2-4). Shoulder height estimates for dogs from the
Fortress of Louisbourg and other colonial contexts suggest French, British, and Spanish colonists
58 selectively imported larger breeds. In addition to supporting historical documentation, the results of this analysis suggest that the selective pressures acting upon European and eastern North
American dog populations resulted in divergences in physical attributes including shoulder height, but potentially other characteristics as well. The identification of these characteristics may enable significant contributions to our understanding of the effects of long-term artificial selection, and our ability to track animal translocation events.
Differences in stature may prove useful in some contexts; however, overlap in stature between smaller European and larger Native American dogs indicates that stature alone will not reliably differentiate all European and Native American dogs. European dogs’ exaggerated range of shoulder height variability likely reflects intensive selective breeding programs designed to suit dogs to specific roles in European society including ratting, hunting, livestock management, warfare, and for use as lapdogs. This range of shoulder height variability in
European dog populations made it possible for European colonists to import large dogs to the
Americas. Less shoulder height variability in Native American dogs from eastern North America does not preclude selection for specific purposes, (e.g. consumption, hunting, fiber, or traction) but suggests that those roles did not require the extreme differences in body size and shape found in European dogs (Figure 2-4). Additional traits including the congenital absence of permanent premolar (P1) observed in many North American dogs (Crockford 1997; Warren 2000), and the application of multivariate morphometric methods detailing aspects of cranial shape (Phillips et al. 2009; Tourigny et al. 2015) will likely prove useful in future analyses.
Genetic evidence indicates interbreeding between Native American and European dogs has resulted in a significant replacement of some Native American dog populations (Leonard et al. 2002; Tamm et al. 2007; Fagundes et al. 2008; Achilli et al. 2008, 2013; Perego et al. 2009;
59 Malhi et al. 2010; Hooshiar Kashani et al. 2012). The date of interbreeding remains unknown; however, the documentary history of dogs in the American colonies emphasizes their use in warfare and intimidation (Derr 2004; Mastromarino 1986). Institutionalized laws in at least
English colonies sought to prevent Native Americans from acquiring large European dogs throughout much of the 17th Century (Andrus 1830; Derr 2004). Comparisons of shoulder height in Native American dogs in Eastern North America (Figure 3-5) reveal no significant change in
Native American dogs through time; however, no detailed analysis of eastern North American dogs has yet been completed.
Physically large guard dogs also played important roles in the introduction of other domesticates. Colonists were confronted with increased rates of predation by wolves and bears, as well as uniquely North American predators like coyotes and mountain lions (Kennedy 2014).
Wolves and bears had been on the decline in England since the Norman invasion in 1066 AD and were extirpated from much of the British Isles by 1400 AD (Pluskowski 2010), though wolves at least persisted in parts of the British Isles until the 18th century (Derr, 2004; Larson et al., 2012). The challenges associated with introducing, managing, and defending livestock likely encouraged selection for larger, self-sufficient, livestock species like cattle (Rouse 1973) which were often allowed to roam freely in the forests surrounding settlements (Bitterman and
McCallum 2016; Cronon 1983). Predation on livestock caused colonists like John Winthrop Jr. to import Irish greyhounds in 1633 to defend his herds (Hawes 1633; Winthrop 1633). George
Washington and Thomas Jefferson were still seeking “Irish wolf dogs” and various breeds of
English and French shepherd dogs as late as 1787 (Jefferson 1791; Lafayette, 1785; Washington
1785a, 1785b).
60 Historical accounts such as these make it clear that relationships existed between environmental conditions, selection of livestock species, and the imported domesticates.
However, these have frequently gone unexplored in historical archaeology. Significant contributions have been made to the study of New World cattle from archaeological contexts
(Cossette & Horard-Herbin 2003; Reitz 1992; Reitz & Ruff 1994; Speller et al. 2013) and pigs
(Guiry et al. 2012), but other domestic species introduced by Europeans have not received similar attention. Biomorphometrics has potential applications for understanding selective pressure, breed differences, and breed improvement (Albarella 1997, 1999, 2002; Phillips et al.
2009; Tourigney et al. 2015). Additional morphometric analyses are needed to expand existing datasets, and address questions raised by previous authors, including whether colonists’ inability to feed and shelter livestock resulted in stunted growth (Cossette & Horard-Herbin 2003), and what timescale is required for the construction of agricultural and pastoral niches (Cronon 1983;
Cossette & Horard-Herbin 2003; Laland & O’Brien 2010; McClure 2015). The joint application of morphometrics, ancient DNA, and isotopic methods provide promising avenues for historical archaeologists seeking to address these questions.
Conclusion:
European colonists intentionally selected dogs possessing specific characteristics, including large size required to defend settlements and livestock, for importation to North
America. These criteria resulted in a disproportionate representation of large dogs from the
European parent population in North America. Though historical research has focused on dogs in the Spanish and English colonies, shoulder height estimates generated on two dogs from the
61 Fortress of Louisbourg suggest that a preference for large dogs existed in French colonial societies as well. These data support claims that dogs in the colonies were larger than those associated with many Native American groups in eastern North America. As shown through this analysis, archaeological data have the potential to corroborate and expand our understanding of colonial introductions.
These data have significant implications for understanding the introduction of European dog populations to North America, and interaction of these and Native American dog populations. Morphometric analyses of dog remains from Colonial assemblages using methods developed by Philips et al. (2009) may reveal close similarity between these remains and morphometric profiles developed for large working dogs in post-medieval and modern populations. Similarly, if European dogs introduced to North America were predominantly large breeds, their appearance in, or contribution to, Native American dog populations may be marked by significant changes in size. By developing these and other lines of evidence will significantly advance our understanding of dog populations in North America and the human communities associated with them.
Acknowledgments:
We would like to thank volunteer Joanne Hughes for her contributions of time and effort. Thanks are also due to those whose biomorphometric data was used in this analysis, especially Drs.
Barry Gaulton and Meghan Burchell (Memorial University) for contributing measurements on a colonial dog from Ferryland, Newfoundland.
Chapter 3
The Birch Creek Canids and Dogs as Transport Labor in the Intermountain West2
Domestic dogs (Canis familiaris) are ethnographically documented as being central features of Intermountain West and Great Plains camps. Some of these dogs were bred specifically for large size and stamina and were used in hauling travois and carrying pannier-style packs. Ethnographic accounts frequently highlight dogs’ importance in
Intermountain West and Plains mobility and report loads as heavy as 45.36 kg (100 lbs).
We calculate body mass from skeletal morphometric data and use these to estimate prehistoric and historic dogs’ load capacity with travois and pannier-style packs in the
Intermountain West, Great Plains, and Great Basin. These data indicate that large dogs recovered archaeologically from the Birch Creek Valley in Idaho and sites on the Great
Plains were capable of ethnographically recorded loads. Further, direct dating of the
Birch Creek dogs indicates that dogs of this size have been present in the Intermountain
West for over 3000 years. These data have important implications for our understanding of prehistoric mobility in the Intermountain West and Plains and suggest that the use of dogs in transporting cargo may have begun as much as 5000 years ago.
2 Co-authored with Dr. David Byers, Utah State University. Under Review at American Antiquity
63 Introduction:
Domestic dogs (Canis familiaris) filled many important roles within Native
American communities including assisting in hunting and camp security, acting as a food source, and providing hair for cordage (Schwartz 1997; Snyder 1991, 1995). Perhaps most importantly, however, they provided a source of labor by hauling, pulling, or carrying cargo in the Arctic, Intermountain West, Great Plains, and Southwest (Allen 1920; Crockford
1997; Driver and Massey 1957; Latham 2016). Unfortunately, though dogs’ importance in transporting resources, belongings, and trade goods during the historic period is clearly demonstrated by ethnographic accounts (Haines 1938:116), the antiquity of dogs’ involvement in North American transport activities remains unknown. Though direct evidence of dog sledding has been recovered from early Holocene contexts in Siberia
(Pitulko and Kasparov 2017), and some have speculated about dogs’ role in Paleo-Indian migration and megafaunal extinction (Fiedel 2005), evidence for dog-based transport in the Americas is limited and at times contentious. Integrating datasets drawn from ethnographic, archaeological, and biological sources provide a powerful and more reliable means of exploring the antiquity of dogs’ role as beasts of burden in North America.
Historic accounts clearly document the significance of dogs to historic Native
American mobility; however, limited direct archaeological evidence has become widely accepted as a measure of the antiquity of these roles in North America. Possible travois fragments have been reported from sites in Wyoming and Montana (Gebhard et al. 1964;
Grey 1963), but these are neither common nor definitively linked to travois technology.
Others have reported on skeletal modifications thought to be associated with dogs’ role in
64 transport including intentionally broken canines similar to modifications seen in Arctic sled dogs (Walker and Frison 1982) and deformation of vertebral spinous processes; however, such vertebral deformations at least have been found in many wolf populations never used in draught roles (Latham 2016; Lawler et al. 2016). Additionally, though dogs are reported in large numbers from Intermountain West and Plains communities (Brackenridge 1906;
Catlin 1973; Hulkrantz 1954, 1956, 1967; Kurz 1937; Lowie 1963; Russell 1964; Wilson
1924), a relatively limited sample of archaeological specimens have been reported on in detail from the Intermountain West and neighboring Great Basin (but see Haag 1956; Lupo and Janetski 1994; Yohe and Pavesic 2000; Lawrence 1967, 1968; Swanson 1972). Further, many dog remains from the Plains clearly date to the historic period (e.g., Bozell 1988;
Morey 1986) when genetic analyses indicate the widespread replacement of Native
American dogs with European dogs (Leonard et al. 2002).
In this paper, we report on the domestic dog remains from two sites in the Birch
Creek drainage of Idaho, Veratic (10CL3) and Bison (10CL10) shelters. Others have previously reported on canid remains from Veratic Rockshelter, a deeply stratified site in the Birch Creek Valley (Swanson 1972; Lawrence 1967, 1968, Figure 3-1). Reanalysis of the Birch Creek faunal materials, today housed at the Idaho Museum of Natural History
(IMNH) has, however, identified a sizable collection of canid remains which we report on here with the goal of better understanding the role of dogs in human lifeways in the
Intermountain West. We do so by asking two questions: do dogs in the Birch Creek assemblages possess the physical characteristics necessary to transport ethnographically reported travois and pack loads and, if so, can their antiquity be used to assess changes in dog size (and potentially their role in transport)?
65
Figure 3-1: A map depicting the location of sites used in this analysis which are numbered as follows: 1. Veratic Rockshelter, 2. Braden Site, 3. Jaguar Cave, 4. Fishing Bridge Campground, 5. Stillwater Marsh, 6. Pyramid Lake, 7. Vista Site, 8. Hogup Cave, 9. Danger Cave, 10. Caldwell Village, 11. Pharo Village, 12. Larson Site, 13. Lower Grand Site, 14. Potts Site, 15. Pretty Head Site, 16. White Buffalo Robe Site, 17. Big Hidatsa Site, 18. Barcal Site, 19. Bellwood Site, 20. Burkett Site, 21. Clarks Site, 22. Gray Site, 23. Hill Site, 24. Horse Creek Site, 25. Linwood Site, 26. Palmer Site, 27. Write Site.
Identifying domestic dogs in the Birch Creek and similar archaeological contexts and estimating their body size is important, especially if those remains predate ethnographic observations. Contextualized within a direct historical approach grounded in ethnographic accounts of dogs in native communities, if the Birch Creek canid remains derive from domestic animals, then this information has implications for understanding the lifeways of associated prehistoric peoples. The development of large dogs adapted to travois and pack transport may provide clues to the invention and intensification of transport technology, changes in mobility, and integration of the Intermountain West’s
66 occupants into trade networks which moved obsidian across sometimes great distances
(Griffin et al. 1969; Hatch et al. 1990). Conversely, if prehistoric dogs lack the robusticity of their more recent counterparts, then their presence in archaeofaunas points to other roles such as hunting or camp protection. In this paper, we test the simple, but as of today unevaluated, idea that prehistoric dogs could have hauled loads similar to those documented in ethnographic accounts.
To test this hypothesis, we first evaluate the collection of canid remains from
Veratic and Bison shelters for the presence of domestic dogs. We do so using characteristics commonly deployed to distinguish domestic dog remains from those of coyote (C. latrans) and wolf (C. lupus). These include tooth and root structure, the congenital absence of mandibular premolar one (P1), length of mandibular molar one (M1), the shape of the ascending ramus and tooth crowding (see Benecke 1987; Clark 1996;
Crockford 1997; Haag 1948; Krantz 1959; Lawrence and Bossert 1967; Lawrence 1968;
Morey and Wiant 1992; Olsen 1985; Young and Jackson 1951). We estimate body mass for domestic dog remains identified in the Birch Creek assemblage using regression formula developed by Losey et al. (2015) and Losey et al. (2017). Experimental data on travois travel (Henderson 1994) and modern pack dogs are then used to discern whether the Birch Creek dogs possessed the physical characteristics necessary to pull the travois loads reported in ethnographic and historic accounts, or otherwise contribute to
Intermountain cultures as beasts of burden. Finally, we discuss our results within the context of body mass estimates for domestic dogs from other Intermountain West, Great
Basin, and Western Great Plains archaeological sites.
67 Domestic Dog Morphology, Mobility and Labor
Domestic dogs were multifunctional contributors to prehistoric and historic
Intermountain West and Great Plains Native American communities. Ethnographic sources indicate that at least two types of dog were present in both the Intermountain West and the
Great Plains, that these assisted in hunting, guarding camps, and that the larger type was most commonly employed in transport activities. Although there are numerous ethnographic accounts of dogs as beasts of burden, lacking are the middle-range studies necessary for linking archaeological remains with their transport capacities, and the availability of dog labor in past communities. Ethnographic sources provide a source of information concerning expected variability present within archaeological remains, as well as the activities and capabilities of archaeological dogs. Data concerning dogs’ physical capabilities drawn from various recreational or experimental transport activities will be used to parameterize the dogs’ load capacity and evaluate the reliability of ethnographic analogies used in interpreting archaeological remains.
Ethnographic and archaeological material from the Intermountain West and Plains suggest dogs in these regions came in two sizes, small dogs which assisted in hunting small game, and large dogs who hauled travois and carried packs, but also chased down or directed mountain sheep, antelope, and bison into corrals and traps following vocal commands (Hultkrantz 1967; Kurz 1937; Murphy and Murphy 1986; Scheiber and Finley
2010; Shimkin 1937-1938). Russell (1964) reports seeing more than two dogs for every individual in a Shoshonean band he encountered in Yellowstone National Park in 1834.
68 Similar numbers of between four and six dogs are reported for Plains families (Buffalo
Bird Woman in Wilson 1924; Catlin 1973; Kennedy and Stevens 1972; Lowie 1963).
Allen’s (1920) ethnographic and historic descriptions of Plains-Indian and Sioux dogs document “wolf-like” animals with erect ears, and tawny, black, grey, or white coloration. Allen (1920) indicates prehistoric dogs in these regions fell into two “breeds,” a small-medium sized “Plains-Indian” dog found widely distributed from the Plains to the
Pacific coast and Canada, to as far south as Mexico (Allen 1920) and a larger “Sioux” dog found primarily on the Northern Plains. Morphometric investigation of archaeological specimens, including two dog skeletons recovered from the Fishing Bridge Campground in Yellowstone National Park, shows some Intermountain West dogs were between coyotes and wolves in size, with crania similar in width to wolves, but with shortened muzzles and massive jaws (Haag 1956; Lawrence 1967, 1968; Yohe and Pavesic 2000). Data from specimens in Wyoming (Walker and Frison 1982) and the Plains (Bozell 1988; Morey
1987) also support the presence of at least two dog types which are differentiable by size.
Ethnographic accounts also provide a window into the management and labor capacities of Native American domestic dogs. Great Plains foragers, for example, adapted dogs to specialized roles by culling smaller pups and reportedly interbreeding domestic animals with wolves (Canis lupus) (Buffalo-bird-woman in Wilson 1924; McFarlane
1905). Doing so encouraged the formation of dog populations large in size and capable of pulling significant loads on simple drag-sleds called travois, and carried folded rawhide, or parfleche-style, packs. Estimates for dog load capacity using travois vary widely from as little as 13.61 kg (30 lb) to loads of as much as 27.22-45.36 kg (60-100 lb) (Harman
1957; Hind 1971; Maximillian 1982; Wolf-chief in Wilson 1924; Grinnel 1962; Weltfish
69 1965; Winship 1896). Similar observations document pack weights of 15.88-22.68 kg (35-
50 lb) for Plains dogs (Castañeda 1904; Kurz 1937). In the Intermountain West, Shoshone,
Nez Perce, and other Native groups kept both small dogs who assisted in hunting small mammals and larger Plains-like dogs who reportedly acted as beasts of burden.
Ethnographic accounts imply the latter were capable of hauling travois weighing 31.75 kg
(70 lb) across level terrain and carrying paired parfleche-style packs weighing 22.68 kg (50 lb) through mountainous territory (Hultkrantz 1954, 1956, 1967; Kurz 1937; Lowie 1955;
Nabokov and Loendorf 2004; Russell 1964).
We could locate only one directly relevant experimental study investigating dogs’ potential to act as beasts of burden. Henderson’s (1994) experimental replication of travois travel using a modern Alaskan husky indicates a 25.4 kg dog could haul at least 27.2 kg, or 107% of its body mass, over short distances of a few kilometers, but was more comfortable with loads ranging of 11.8-13.6 kg (40-50% of body mass) on longer trips covering as much as 27 km in 7 hours. Henderson’s (1994) recreation of travois travel allows us to contextualize the available ethnographic data and supports a distinction between long and short distance loading strategies documented by ethnographic sources
(Bradley 1923; Wolf-chief in Wilson 1924). Ethnographic reports indicate that travois loads were also impacted by environmental conditions, being lighter in summer than in winter when cooler temperatures kept dogs from overheating, and snow both reduced friction on the travois and provided dogs with water (Buffalo-bird-woman in Wilson 1924;
Wolf-chief in Wilson 1924). Henderson’s (1994) experiment further highlights the difficulties of employing travois in densely vegetated environments that frequently
70 entangled the travois, or sloped environments that forced the dog to tack side-to-side when going uphill and caused the travois to ride up over the dog’s head when descending.
Whereas archaeological research into dog labor is scarce, several studies tied to different goals provide useful data, especially for sled dogs. One study demonstrated that sled teams of dogs, averaging 39 kg in body mass and acting as a group, could pull as much as 115 kg a piece, but required frequent rest periods because of high load weight (Taylor
1955; cited by Bostelmann 1976). Loads of 45 kg per animal (115% of body mass) were found more reasonable (Taylor 1955; cited by Bostelmann 1976), and the most efficient load for rapid transport sled dogs was only 23 kg (58.97% of body mass) (U.S. War
Department 1994). These data indicate sled dogs are capable of moving extremely large loads (e.g. 115 kg, or 294.87% of body mass), but do so in teams, and under snowy or icy conditions which reduce friction acting upon their loads. As with Henderson’s (1994) findings, these data also document two loading strategies, one maximizing load size (115% of body mass), and another intended to maximize transit speed (60% of body mass).
Notably, the load capacity of individual dogs in both loading strategies are reasonably close to those identified by Henderson (1994) for travois travel.
Modern data are also available for dogs carrying packs. A survey of dog backpacking guides, blogs, advertisements, and equipment found many recommend no more than 25-30% of a dog’s body mass be carried in packs (Balogh 2017; Green 2017;
Samoyed Club of America 2017; Terrill 2012). The U.S. War Department (1994) indicates that pack dogs averaging 35 kg or more are capable of carrying loads of up to 23 kg
(65.71% body mass) for a few days without harm but recommends that loads average 16 kg (45.71% of body mass). Notably, modern recommendations may be tempered by
71 ideological notions concerning humane treatment and animal abuse. Dogs in the
Intermountain West reportedly carried as much as 22.68 kg (50 lb) (Kurz 1937) in packs while those on the Plains carried 15.88-22.68 kg (35-50 lb) (Castañeda 1904). Though high loads of 22.68 kg may suggest loads exceeding modern recommendations, smaller loads reported on the Plains may indicate that load size was again linked to distance, speed, or environmental conditions.
These data present both insight into the role of dogs as a source of labor, and an opportunity to evaluate the likelihood that ethnographic and historic accounts accurately document these animal’s transport capabilities. Dogs would likely only have pulled travois in certain circumstances, such as passage over open ground. Other circumstances, such as rugged, densely vegetated terrain, would have encouraged the use of packs. Regardless of the apparatus used to facilitate load transport, Henderson’s (1994) experimental data, ethnographic sources, and modern reference data present a set of parameters for linking dog body mass with load capacity, one we explore through analysis of both the Birch Creek canids and a broader dataset drawn from published accounts of dog remains from the
Intermountain West and adjacent regions.
Henderson’s (1994) experimental data and modern data provide a useful frame of reference for understanding the prehistoric labor utility of dogs as a ratio of load weight to total body mass. Given Henderson’s (1994) study in combination with the research on sled and pack dog load capacities, we suggest 107% and 45% of body mass are reasonable estimates for dogs’ short- and long-distance travois load capacities. A load of 30% of body mass used by modern backpackers is used as a reasonable estimate of dog’s pack load capacity since the entire load of packs is placed on a dog’s back, rather than braced on the
72 ground (Balogh 2017; Green 2017; Samoyed Club of America 2017; Terrill 2012).
Notably, modern estimates of dogs’ load capacity, especially concerning packs for which no experiment was available, may be influenced by modern notions regarding the humane treatment of animals, and higher loads may have been achieved in the past. To test ethnographic observations against potential prehistoric dog labor capacity we estimate travois and pack load capacities as ratios of 107%, 45%, and 30% from body mass estimates generated following methods developed by Losey et al. (2015) and Losey et al.
(2017).
Data and Methods
The methods employed in this analysis address two goals; 1) the identification of domestic dog remains in the Bison and Veratic Rockshelter assemblages, and 2) determining the potential for any Birch Creek dogs, as well as a sample of dogs from a broader regional context, to have hauled ethnographically recorded travois and pack load sizes. The Bison Rockshelter and Veratic Rockshelter assemblages were excavated between 1960-1961 (Swanson 1972). Deeply stratified deposits from these rockshelters reflect intermittent occupation associated with seasonal hunting expeditions since at least
9950-9500 cal BP (Keene in press) and have contributed to the development of well-dated, regional, projectile point chronologies (Butler 1978; Holmer 1986, 2009; Keene in press).
Swanson’s excavations recovered a large faunal assemblage (Swanson 1972) containing a number of canid specimens. Lawrence (1967, 1968) reported on a subset of the Birch Creek canids including a cranium (IMNH-19613), maxillary fragments (IMNH-18725, and
73 IMNH-18802), and mandibles (IMNH-18803, IMNH-18418, IMNH-19636, IMNH-
19637). These were identified based on tooth size, paracone and metacone development, and the relatively weak development of tooth roots (Lawrence 1968). In these analyses,
Lawrence identified at least two types of domestic dogs distinguished largely by size in both Jaguar Cave and the Veratic Rockshelter (Lawrence 1967, 1968). Both types exhibit characteristically short and broad muzzles, and massively deep mandibles (Lawrence 1967,
1968).
In this study, we revisit Lawrence’s (1968) identifications, and present new findings on five specimens she did not evaluate. Canid remains from the Bison and Veratic
Rockshelters were analyzed to verify taxonomic identification and collect morphometric measurements. All measurements follow von den Driesch (1976) and we identify the specific metrics by element and measurement number as presented in this reference. Within our reanalysis, we employed the following four lines of complementary evidence from the
Birch Creek canid remains that others have used to distinguish domestic dogs from wolves and other wild canids. The congenital absence of P1 was observed in 82% of domestic dog mandibles examined by Crockford (1997). Consequently, zooarchaeologists often use this attribute to identify North American domesticated dogs (Allen 1920; Haag 1948; Lupo and
Janetski 1994). M1 length has also been used to identify domestic canids (Lupo and Janetski
1994). Wolf populations in the Western U.S. commonly exhibit molars of at least 25.1 mm in length for females, and 26 mm for males (Nowack 1979), while coyotes exhibit molar lengths of 18.5 mm for females and 19.6 mm for males (Nowack 1979). Domestic dog M1 length commonly falls between those for wolves and coyotes, though some overlap is possible (Crockford 1997; Lupo and Janetski 1994). A specific mandiblular morphology,
74 in this case a notable caudal curvature of the ascending ramus, often presents in domestic dogs, but not wild canids (Benecke 1987; Olsen 1985). Finally, although Ameen et al.
(2017) have recently called into question the reliability of tooth crowding, such indices for both mandibles and maxilla containing full adult dentition have also shown utility in sorting wild from domestic canids (Clark 1996; Clutton-Brock 1963; Degerbøl 1963; van
Wijngaarden-Bakker 1974). Tooth crowding indices are generated by dividing the summed length of the permanent premolars, by the length of the permanent premolar row measured from the anterior surface of the P1 to the posterior surface of P3. This metric indicates that crowding values for domestic dogs fall between 86.3 and 103 in mandibles and 79.4 and
109 in maxilla, while scores below this range are most commonly found in wild canids
(Clark 1996; Clutton-Brock 1963; Degerbøl 1963; van Wijngaarden-Bakker 1974).
We evaluate the size and load capacity of the Veratic Rockshelter canids through estimation of their body mass in kilograms. Zooarchaeologists have developed several methods for investigating the size of archaeological dogs, including shoulder height
(Harcourt 1974) and body mass (Wing 1978; Van Valkenburgh 1990). Unfortunately, differences in the skeletal landmarks chosen and the way in which selected landmarks are measured between various morphometric measurement systems (e.g. von den Driesch
1976; Haag 1948; Lawrence 1967, 1968), and the differential preservation of specimens, mean that data compiled from the literature are often difficult to compare. Furthermore, techniques for estimating body mass have in the past been hampered by small sample sizes, and in some cases the lack of domestic dogs from study collections (see Wing 1978; Van
Valkenburgh 1990).
75 Body mass, the amount of matter an organism is composed of, may be estimated from skeletal remains and is therefore frequently used by biologists and paleobiologists to approximate body size (Anyonge and Roman 2006; Campione and Evans 2012; Damuth and MacFadden 1990; Legendre and Roth 1988; Thackeray and Kieser 1992). Body mass has been strongly correlated with a variety of ecological characteristics including aspects of life history, home range size, population density and growth, functional morphology, and metabolism, and as a result it has been widely used in the study of both extant and extinct species (see Anyonge and Roman 2006; Campione and Evans 2012; Damuth and
MacFadden 1990; Legendre and Roth 1988; Thackeray and Kieser 1992). Such studies frequently test for proportional relationships between the metrics of interest and the body masses of various species. Prior studies have revealed that elements involved in biomechanical loading and functional stressors, especially long bones provide some of the most reliable estimates of body mass (Campione and Evans 2012; Figueirido et al. 2011).
However, numerous studies have revealed high correlation coefficients when comparing mandibular characteristics, especially the length of carnassial teeth, and body mass
(Legendre and Roth 1988; Thackeray and Kieser 1992)
Because body mass can be calculated from morphometric data collected on a variety of skeletal elements, it provides an avenue for studies of archaeological remains inhibited by small sample sizes or differential preservation. Losey et al. (2015) have developed a set of logarithmic regression formula for 20 cranial and 20 mandibular measurements, and 29 long-bone measurements (Losey et al. 2017) which generate estimates of body mass in kilograms from a sample of 36 domestic dogs including 22 Inuit sled dogs, and 108 wolves of known body mass. In the case of archaeological canids, high
76 similarity in post-cranial skeletal anatomy in wild and domestic canids means that archaeologists are often forced to rely upon crania and mandibles to identify domestic dog remains. As with previous studies, Losey et al. (2015) and Losey et al. (2017) find that mandibular measurements frequently generate reliable and accurate estimates of body mass, but that individual measurements predict wolf and dog body mass with different levels of accuracy. We calculated body mass estimates in kilograms using morphometric measurements following von den Driesch (1976) and regression formula shown by Losey et al. (2015) and Losey et al. (2017) to most accurately predict body mass in domestic dogs.
Table 3-2 presents the generic formulae, associated regression coefficients may be found in the Supplemental Data. Finally, we use the resulting datasets to calculate both short
(107% body mass) and long (45% body mass) distance travois loads, and pack (30% body mass) capacities to test the ethnographic observations reviewed above.
Results
Nineteen specimens representing a minimum of four adult domestic dogs were identified in the Veratic Rockshelter assemblage. No specimens clearly belonging to domestic dogs were identified in the Bison Rockshelter materials, and no juvenile remains were recovered from either site. The 19 domestic dog specimens identified here include pieces of at least three crania, eight maxillae, and seven mandibles (Table 3-1). An additional 68 specimens were identified as coyote (Canis latrans), and another 67 represent unidentified canids. We do not report any further on the coyote and other canid specimens here. Four of the domestic dog specimens were directly dated (Table 3-2) and returned
77 median ages ranging from 5226 to 387 cal BP, indicating dogs were part of local adaptation from the at least the middle Archaic to the Protohistoric periods in the Birch Creek Valley
(Plew 2016).
Table 3-1: Veratic Rockshelter domestic dog specimens. Specimen Number Occupation Level/Depth Below Datum Element IMNH-19613 Occupation V 14 Cranium IMNH-19636 Occupation V 14 R Mandible IMNH-19637 Occupation V 14 L Mandible IMNH-25420 Occupation III/IV 65-75 cm R Mandible IMNH-18418/19210 Occupation IV 21 L Mandible IMNH-19551 Occupation III 24-25 Cranial Fragment IMNH-19566 Occupation III 24-25 R Maxilla IMNH-19567 Occupation III 24-25 L Maxilla IMNH-18880/18816 Occupation III 25 Cranium IMNH-18803 Occupation III 25 R Mandible IMNH-18724 Occupation III 25 L Mandible IMNH-18425 Occupation III 25 L Maxilla IMNH-26128 Occupation III 25 R Mandible IMNH-18804 Occupation III 26 Cranial Fragment IMNH-18805 Occupation III 26 R Maxilla IMNH-18802 Occupation III 26 R Maxilla IMNH-19617 Occupation II Feature 4 Maxilla IMNH-26344 Occupation II 110-120 cm L Maxilla IMNH-27016 Unknown Maxilla
A number of characteristics were used in identifying domestic dog remains in the
Birch Creek assemblages. The first premolar was congenitally absent in mandibles IMNH-
18803, and IMNH-18724, a characteristic common to Native American domestic dogs
(Crockford 1997, Table 3-3; Figure 3-2a, b). M1 length in mandibles IMNH-19636 and
IMNH-19637 (Table 3-3; Figure 3-2c, d) is larger than the known range for coyotes, but
78 smaller than that for wolves and indicating that these specimens derive from domestic animals (Figure 3-3a). Mandibular tooth crowding indices calculated for IMNH-19636
(99.648) and IMNH-19367 (93.798) also fall well within the range for domestic dogs
(Clark 1996; Clutton-Brock 1979; Degerbøl 1963; van Wijngaarden-Bakker 1974, Figure
3-3b). Additionally, caudal curvature is observed on the ascending ramus of these mandibles, further supporting their identification as domestic dogs (Benecke 1987; Olsen
1985). Though mandible (IMNH-26128) with an M1 length of 27.88 mm may represent a wolf, it is morphologically very similar to other domestic dogs identified in the assemblage and is treated as one for this analysis.
The Birch Creek specimens display several cultural bone modifications. While no cut marks were identified on any of the bones; several domestic dog, coyote, and unidentified canid specimens exhibited burning (NISP= 32, 21.33%) and spiral fractures
(NISP= 12, 8.00%), indicating that these animals may have been processed as food (Snyder
1991, 1995). An unhealed depressed fracture in the right frontal bone of cranium IMNH-
19613 (Figure 3-2f) suggests the prehistoric inhabitants of Veratic Shelter dispatched at least one of the animals in our sample. Similar injuries to the frontal bone of canid crania have been reported at the Vore Site (Walker 1975; Walker and Frison 1982), sites on the
Great Plains (Morey 1986) and elsewhere in North America and Russia (Losey et al. 2014;
Park 1987). These studies have revealed that both healed and unhealed fractures to the frontal bone, like that found on cranium IMNH-19613, are significantly more common in domestic dogs than wolves, and attributed to blows from humans or fights with other dogs
(Losey et al. 2014; Park 1987).
79
Figure 3-2: A panel of photos depicting mandibles a) IMNH-18803, b) IMNH-18418/IMNH- 19210, c) IMNH-19636, d) IMNH-19641, and reconstructed crania e/f) IMNH-19641 which exhibits an unhealed cranial fracture.
Table 3-2: Radiocarbon data for dated specimens (samples processed by Direct AMS and University of Utah). % IMNH Collagen 14C Age Cal. 2 σ Median Cal Site Spec. # Identification Provenience Occupation Element Yield BP Range BP BP 10CL10 27491 Canis sp. (Large) Level 3 Occupation V Femur 9.3 332 ± 25 309-468 387 10CL3 19636 Canis familiaris Level 14 Occupation V Mandible 3.9 2932 ± 34 2966-3174 3084 10CL3 18724 Canis familiaris Level 25 Occupation III Mandible 12.9 3132 ± 32 3249-3444 3357 Canis cf. 10CL3 26128 familliaris Level 25 Occupation III Mandible 13.9 4569 ± 40 5052-5444 5226
Table 3-3: Identifying metrics for domestic dog specimens.
2 Crowding M1 Ascending Correlation Body-mass kg 1 2 2 Specimen Level Element Index length P1 absent Ramus Measurement Coefficient Constant (lbs) IMNH-19637 14 L Mandible 99.8 24.62 No Curved Caudally VDD M7 3.041 -4.437 28.53 (62.90) IMNH-19636 14 R Mandible 99.65 24.59 No Curved Caudally VDD M7 3.041 -4.437 28.32 (62.43) IMNH-19613 14 Crania 86.73 NA NA NA VDD C8 2.604 3.844 24.31 (53.60) IMNH-18724 25 L Mandible NA NA Yes NA VDD M12 3.161 -3.579 27.48 (60.59) IMNH-18803 25 R Mandible NA NA Yes Curved Caudally VDD M7 3.041 -4.437 21.88 (48.23) IMNH-26128 25 R Mandible NA 27.88 NA NA VDD M19 3.086 -3.036 33.52 (73.90) 1Mandibular Crowding Index: (P +P +P +P )/(Length P to P )*100 1 2 3 4 1 4 3Following Losey et al. (2014) Bodymass=Log (Measurement)*Correlation Coefficient-Constant 10
Figure 3-3: (a) Mandibular crowding indices for archaeologically reported domestic dogs (see Clark 1996), (b) M1 length for the Veratic Rockshelter dogs and reported values for male (MNI=62) and female (MNI=47) wolves, and male (MNI=99) and female (MNI=99) coyotes (Nowak 1979).
Only domestic dog cranium IMNH-19613, and mandibles IMNH-18724, IMNH-
18803, IMNH-19636, and IMNH-19637 could be used to generate body mass estimates using the methods described in Losey et al. (2015) and Losey et al. (2017). Table 3-4 presents these data. Occupational level 25 produced both large (IMNH-18724) and small
(IMNH-18803) dogs. Mandibular measurement 12 (Figure 3-4) for the former specimen resulted in a body mass estimate of 27.48 kg, while mandibular measurement seven (Figure
3-4) for the latter generates a body mass estimate of 21.88 kg. Dogs from the level 25 assemblage average 24.68 kg.
82
Figure 3-4: The most commonly used measurements used in this analysis for estimating body mass (Based on von den Driesch 1976).
Three additional body mass estimates were generated for specimens recovered from level 14. These include one cranium (IMNH-19613) and two mandibles (IMNH-19636,
IMNH-19637) (Figure 3-4). These latter two specimens likely represent a single individual.
Cranial measurement eight (Figure 3-4) taken on IMNH-19613 resulted in a body mass
83 estimate of 24.31 kg. Finally, mandibular measurement seven (Figure 3-4) produced body mass for mandibles IMNH-19636 and IMNH-19637 of 28.32 kg and 28.53 kg, respectively. Together these data indicate the dogs recovered from the Veratic Rockshelter ranged between 21.88-28.53 kg (Table 3-4, Figure 3-5), making them similar in body mass to modern Siberian huskies (American Kennel Club 2017). Dogs recovered from Veratic
Shelter occupational level 25 average 24.68 kg. Dogs recovered from occupational level
14 average 27.05 kg and are on-average larger animals than those found in the older level
25 sample.
Table 3-4: Veratic Rockshelter dog body mass estimates, short- and long-distance travois, and pack load capacities. Travois Load Sample Body-mass (lb) Short (lb) Long (lb) Pack Load IMNH- 27.48 (60.58) 29.41 (64.82) 12.37 (27.26) 8.24 (18.18) 18724 IMNH- 21.88 (48.23) 23.41 (51.61) 9.85 (21.71) 6.56 (14.47) 18803 IMNH- 28.53 (62.90) 30.53 (67.30) 12.84 (28.30) 8.56 (18.87) 19637 IMNH- 28.32 (62.43) 30.30 (66.81) 12.74 (28.10) 8.50 (18.73) 19636 IMNH- 24.31 (53.60) 26.02 (57.35) 10.94 (24.12) 7.29 (16.08) 19613
Our analysis shows the Birch Creek dogs display a range of body masses, which can be interpreted in at least two different ways. First, the presence of both small and large dogs could reflect different types or breeds, as identified in the ethnographic literature. If this is the case, the smaller of these populations likely assisted hunting parties, while larger
84 animals served in transport activities (Hultkrantz 1954, 1956, 1967). Notably, this interpretation might imply a relatively recent origin for dogs’ draught roles.
Figure 3-5: Body mass in kg for the Veratic Rockshelter dogs, male (MNI=24) and female (MNI=25) wolves (Mech 2006), and an unknown number of male and female western coyotes (Way 2007).
Alternatively, the size differences we identify may more simply reflect sexual dimorphism. Canids generally adhere to patterns of sexual dimorphism measured by comparing adult male and female shoulder height and body mass (Frynta et al. 2012).
Though wolves, the progenitor species for domestic dogs exhibit the most sexual dimorphism of any wild canid (Frynta et al. 2012), the degree of sexual dimorphism in domestic dogs is highly variable (Bidau and Martinez 2017; Frynta et al. 2012). Male domestic dogs are generally 1.10-1.46 times larger than females, but across most breeds males average only 1.15 times larger (Bidau and Martinez 2017). Comparison of body
85 mass estimates generated on dogs from the Veratic Rockshelter reveals that the largest dog
(28.53 kg) is 1.30 times larger than the smallest (21.88 kg) indicating that body mass differences could reflect sexual dimorphism but would be on high end of the known range for domestic dogs. A student’s T test reveals no statistically significant difference between these populations (df=3, p=0.4497, t=0.8670). Together, we take these data to indicate that the dogs from the Veratic Rockshelter likely represent a single sexually dimorphic population.
As discussed above, we employ estimates of 107% and 45% of body mass based on Henderson’s (1994) experimental replication of travois travel to approximate long and short distance travois load sizes. An estimate of 30% of body mass, the upper limit of many modern recommendations regarding the load size of backpacking dogs, provides a proxy for pack-load capacity. Assuming prehistoric dogs had similar capabilities as modern dog breeds, including the husky used in Henderson’s (1994) analysis, a dog 24.68 kg in size would have been capable of pulling 26.41 kg over short distances, 11.11 kg over long distances, and carrying 7.41 kg by pack (Table 3-4). Dogs from level 14 averaging 27.05 kg would have been capable of pulling 28.95 kg over short distances, 12.17 kg over longer distances, and carrying 8.12 kg by pack. The largest body mass estimates (IMNH-19637,
IMNH19636; possibly one individual), approximate the travois loads reported by Kurz
(1937) for dogs in the Intermountain West, and are also consistent with short haul load limits derived from our literature review. These data suggest this individual would have been able to move relatively heavy loads for distances likely in the range of a few kilometers across level and open terrain. No estimates of dogs’ pack load capacity reach the weights of 22.68 kg (50 lb) recorded by European observers (Kurz 1937), implying
86 either ethnographic observers over-estimated pack load sizes, or loads carried by Native
American dogs exceeded modern recommendations.
Discussion: The Birch Creek Dogs in Broader Context
This paper sought to identify domestic dogs in the Birch Creek assemblage and assess whether these animals may have been capable of transporting the same loads as their ethnographic counterparts. In response to these research questions, our analyses identified at least four domestic dogs in the Birch Creek collections. We also found the Birch Creek dogs vary in size, with some approximating sizes capable of pulling short distance travois loads of more than 22.6 kg (50 lb) similar in size to those mentioned in ethnographic and historic documents, while others appear much smaller. Further, directly dating dog bone has shown that large dogs have been present in the Intermountain West for at least 3,000 years.
To place the Birch Creek dogs into a broader context and investigate archaeological load capacities of prehistoric dogs more generally, we compare calculated body masses and load capacities for the Birch Creek animals with similar values calculated for archaeological domestic dogs from nearby regions for which morphometric data were available (Table 3-5, Figure 3-1, see also Appendix B). These include 10 dogs from the
Intermountain West (including the Birch Creek dogs), 10 from the Great Basin, and 115 from Great Plains contexts. Here again, we use Losey et al. (2015) and Losey et al.’s
(2017) regression formulae to derive body mass estimates for the specimens in our regional sample. Short and long-distance travois loads and pack load capacities are set respectively
87 to 107%, 45%, and 30% of calculated body mass. Cranial measurement 13a and
mandibular measurements 8, 9, and 19 were most commonly used in generating body mass
estimates for the comparative sample.
Table 3-5: Veratic Rockshelter dog body mass estimates, short- and long-distance travois, and pack load capacities.
Body mass Region Site Estimates Source Veratic Rockshelter 5 This analysis; Keene (in press); Lawrence (1967, 1968) Intermountain Braden Site 2 Yohe and Pavasec (2000) West Jaguar Cave 2 Lawrence (1967, 1968); Gowlett et al. 1987 Fishing Bridge 2 Haag (1970) Stillwater Marsh 2 Schmitt and Sharp (1980), Rhode et al. (2000) Pyramid Lake 1 Schmitt and Sharp (1980) Vista Site 2 Dansie and Schmitt (1986); Delacorte (1997) Great Basin Danger Cave 1 Grayson (1988) Hogup Cave 1 Haag (1970) Caldwell Village 2 Haag (1966); Ambler (1966) Pharo Village 1 Haag (1968) Larson 3 Morey (1986) Lower Grand 1 Morey (1986) Potts 2 Morey (1986) Pretty Head 1 Morey (1986) W. B. Robe 2 Morey (1986) Big Hidatsa 3 Morey (1986) Barcal 24 Bozell (1988) Bellwood 5 Bozell (1988) Great Plains Burkett 20 Bozell (1988) Clarks 1 Bozell (1988) Gray 26 Bozell (1988) Hill 8 Bozell (1988) Horse Creek 4 Bozell (1988) Linwood 2 Bozell (1988) Palmer 1 Bozell (1988) Wright 12 Bozell (1988)
*See Appendix 2 for raw data
88 Archaeological specimens are rarely reported from Intermountain West and Great
Basin contexts (but see Lupo and Janetski 1994; Yohe and Pavesic 2000), and this situation limits our comparative sample for these regions (Table 3-5). In addition to the Birch Creek dogs, the Intermountain West sample includes six dogs from the Braden Site, Jaguar Cave in Idaho, and the Fishing Bridge Campground in Yellowstone National Park (Haag 1956;
Lawrence 1967, 1968; Yohe and Pavesic 2000). Ten specimens derive from Great Basin contexts, including Stillwater Marsh, the Vista Site, and Pyramid Lake in Nevada, as well as Danger and Hogup Caves and the Caldwell and Pharo Village Sites in Utah (Dansie and
Schmitt 1986; Grayson 1988; Haag 1966, 1968, 1970; Schmitt and Sharp 1990). In contrast, the Great Plains sample includes dogs from ten sites in Nebraska (the Burkett,
Gray, Wright, Barcal Hill, Horse Creek, Linwood, Bellwood, Palmer, and Clarks Sites;
Bozell 1988) and six sites in North and South Dakota (the Larson, Lower Grand, Potts,
Pretty Head, White Buffalo Robe, Big Hidatsa Sites; Morey 1986).
The Birch Creek dogs ranging from 21.87-28.53 kg represent animals larger than dogs from other Intermountain West sites, which range from 11.01-20.78 kg in our sample.
These data may again indicate that variability in size exhibited by dogs in the Veratic
Rockshelter reflects sexual dimorphism rather than breed differences. Great Basin dogs closely resemble smaller Intermountain West dogs (9.63-21.09 kg), averaging only 15.18 kg. While Great Plains dogs demonstrate high variability (5.31 - 39.47 kg) and include the largest and smallest dogs in the comparative sample (Figure 3-6), the overall sample does not exhibit the bimodal distribution as predicted by ethnographic accounts (Figure 3-7).
Smaller dogs in all three regions resemble Plains Indian dogs (Bozell 1988) in size, regardless of whether or not these animals reflect region specific breeds. Dogs from the
89 Intermountain West and Great Plains exhibit a similar average body mass (Table 3-6;
Figure 3-6); however, the largest dog from the Intermountain West is nearly ten kilograms smaller than the largest Plains dog (Table 3-6).
Figure 3-6: A boxplot showing the range of body mass calculated for dogs found in each region following Losey et al. (2015) and Losey et al (2017). Dotted lines depict the upper and lower limits of body mass estimates generated for mandibles from sites in Nebraska which Bozell (1988) attributed to the smaller “Plains-Indian dog”.
Though regional populations appear distinct on the basis of size, statistical analyses failed to find any statistically significant size differences between these populations. An
90 ANOVA test for difference found no statistically significant between-group differences in body mass (df=133, F=2.138, p=0.122) (Figure 3-6). A Tukey HSD test which compares all possible combinations of mean values from the previous ANOVA analysis also finds no statistically significant difference in mean body mass between the Great Plains and
Great Basin (p=0.102), Intermountain West and Great Basin (p=0.273), or the
Intermountain West and Great Plains (p=1.000). Though these results suggest that dogs from the three regions could derive from a single highly variable population as defined by size alone, the largest dog included in this sample is 4.15 times larger than the smallest dog, far exceeding the known range of sexual dimorphism in domestic dogs (Bidau and
Martinez 2017; Frynta et al. 2012). Bozell (1988) has already shown that mandibles from sites in Nebraska cluster into several size-based classes which he interprets as a combination of sexual dimorphism and breed differences. More detailed analyses may be capable of detecting further divisions within and between these datasets.
Though we were unable to statistically identify different populations within our sample, we can nonetheless sort these animals based on load capacity. Only the largest
Plains and Intermountain West dogs accounting for only 21.43% (n=27 dogs) of the sample were found to be capable of hauling ethnographically recorded travois loads of
27.22-45.36 kg (60-100 lb) for short trips, and 13.61 kg (30 lb) on longer ones (Table 3-
6). Although Allen (1920) states that smaller Plains-Indian dogs were employed in travois transport, the average Plains Indian dog identified by Bozell (1988) would have had a short distance load of only 14.18 kg (31.26 lb). These data imply European observers accurately estimated travois loads observed in Native American communities, but frequently emphasize more impressive 27.22-45.36 kg (60-100 lb) loads transported
91 Table 3-6: Comparative sample body mass estimates, and travois and pack load capacity estimates. over relatively short distances in day-to-day activities. No regional sample reached pack Travois Body- Sample Short (lbs) Long (lbs) Pack (lbs) Metric mass Intermountain Min 11.01 11.78 (25.98) 4.96 (10.93) 3.30 (7.28) West n=11 Avg 19.83 21.22 (46.79) 8.92 (19.68) 5.95 (13.12) Max 27.48 29.41 (64.83) 12.37 (27.26) 8.24 (18.96) Great Basin Min 9.62 10.30 (22.71) 4.33 (9.55) 2.88 (6.37) n=10 Avg 15.18 16.24 (35.81) 6.83 (15.06) 4.55 (10.04) Max 21.09 22.56 (49.74) 9.49 (20.92) 6.33 (13.95) Great Plains Min 5.36 5.73 (12.65) 2.41 (5.32) 1.61 (3.55) (Total) n=115 Avg 19.86 21.25 (46.86) 8.94 (19.71) 5.96 (13.14) Max 39.47 42.23 (93.11) 17.76 (39.16) 11.84 (26.11) Plains Indian dogs Min 5.36 5.73 (12.65) 2.41 (5.32) 1.61 (3.55) Bozell (1988) Avg 13.25 14.18 (31.26) 5.96 (13.15) 4.04 (8.90) n=41 Max 21.41 22.91 (50.51) 9.63 (21.24) 6.42 (14.16) Sioux dogs Min 14.28 15.27 (33.67) 6.42 (14.16) 4.28 (9.44) Bozell (1988) Avg 23.14 24.95 (55.00) 10.49 (23.13) 6.99 (15.42) n=62 Max 39.47 42.23 (93.12) 17.76 (39.16) 11.84 (26.11) Bold values meet ethnographic expectations load estimates found in ethnographic observations indicating that the pack loads carried by Native American dogs likely exceeded modern recommendations. The U. S. War
Department (1994) reports dogs are capable of carrying loads exceeding 45%, or even
65% of body mass, though it is unclear how far these loads were carried. If loads of 45% of body mass were achieved in Native American societies, the largest archaeological
Plains dog would meet ethnographic expectations with a load of 17.76kg (40 lb) (Table
3-6).
92
Figure 3-7: A histogram showing the distribution of body mass estimates generated for archaeological dog remains from the Great Plains.
The fact that only the largest dogs in the Veratic Shelter and Great Plains samples were capable of pulling long and short distance travois loads, in combination with the absence of large dogs from the Great Basin where the travois was not used, has implications for understanding how these dogs were used. In this case, selective management including culling pups from litters was likely important in adapting dogs to regionally specific transport goals whether or not Native American communities were selectively breeding large dogs. Though removal of small pups from litters may not have had the same selective strength as intentionally breeding large male and female individuals, removal of smaller pups from litters could translate into selection for large size simply by increasing the frequency of large animals within the population.
93 Sampling and methodological limitations inhibit our ability to document when
Native American societies began using the travois, or other dog-based transport strategies. Previous scholars have asserted a strong concordance between the use of tipi and the dogs in transport (Driver and Massey 1957:298). The advent of tipi on the
Northern Plains has been placed at some time before 900 AD (Brasser 1982; Wissler
1910), and dogs exhibiting pathological signs of use in traction have been recovered from contexts dated to between 4,000-5,000 BP at the Gray Site in Saskatchewan (Millar
1978:365-369) though morphological data was not available for this sample.
The dataset of dogs from these regions for which morphometric data is available is heavily skewed towards the historic period and inhibits our ability to trace when large dog types in the Intermountain West and Plains developed. Direct dating of dog bone from the Veratic Rockshelter shows that several dogs capable of ethnographically reported loads date to 3000 BP or older indicating that dogs capable of pulling or carrying ethnographically recorded loads have existed in this region for several thousand years.
These data suggest that dogs large enough to transport ethnographically reported loads were present in the Intermountain West and Plains prior to the introduction of
European dog breeds. These data do not, however, preclude the possibility that dogs in historic periods were either large breeds introduced by European colonists, or crosses between indigenous and European stock. A combination of historic sources and archaeological data has shown that European dogs introduced during the historic period were generally significantly larger than many dogs found in the eastern U.S. and Canada
94 (Chapter 2) making it possible that these dogs would have been desirable for transport purposes.
Several impediments make a solely European origin for large dogs in Plains communities unlikely. Among these are inconsistencies in timing and geography, and culturally instituted bans on the trade of European dogs to Native American communities.
Dogs’ use in transport is believed to have begun in the northeastern Plains and diffused from there to the southern Plains (Brasser 1982; Driver and Massey 1957; Wissler 1922).
This pattern is accompanied by an overall decrease in dog size from north to south (see
Allen 1920). Encounters between Plains and European communities began in the south during the 1500’s with Spanish explorers at which time dogs were already reported as being used in travois transport (Castañeda 1904). As a result, Hudson Bay Company and
French fur traders operating in the Northeast are the most likely sources for large
European dog breeds to the Northern Plains. However, interaction between European traders and travois using Native American groups is not known until the late 1600’s or early 1700’s at the latest (see Francis and Morantz 2983; Nassaney 2015).
The French and English fur trade was largely undertaken by fishermen operating at the mouth of the St. Lawrence river during the 1500’s (Nassaney 2015). Quebec, established in 1608, marks the first permanent inland French settlement (Nassaney 2015).
The Iroquois confederacy, who are not known to have used travois, managed the fur trade until the 1640s when local sources of furs became over exploited (Francis and Morantz
1983; Nassaney 2015). Though French traders began venturing into the interior themselves in the 1650s in search of new trading opportunities, permanent French settlement remained east of the Great Lakes until Fort St. Joseph was established in 1691
95 (Nassaney 2015). Trade through Hudson’s Bay did not become a regular occurrence until after 1668 (Francis and Morantz 1983). Together, this timeline places major contact between French communities and Northern Plains groups using travois at some time after
1650, and likely much later. In addition, many colonies banned the trade of European dogs to Native American groups through much of the 1600’s (Andrus 1830; Derr 2004).
Figure 3-8: The Plains sample through time showing the timing of Spanish exploration on the Southern Plains, and the periods of Iroquois and European managed fur trade in the North.
When dogs from the Plains sample are plotted across time (Figure 3-8) it is clear that at least one dog capable of ethnographically reported loads in the Plains sample predates even Spanish exploration of the Southern Plains. Further, in order for European dogs capable of ethnographically reported loads to have been present at the time of
Spanish exploration in the south to have spread from the Northeast they would have
96 needed to be introduced well before the 1540’s. At this time English and French fur traders were still operating at the mouth of the St. Lawrence River (Francis and Morantz
1983; Nassaney 2015). Such traders did not establish permanent trading facilities at
Quebec until the 1608 and did not begin operating on their own in the Great Lakes until the 1640s or later, a century or more after dogs were observed transporting cargo on the
Southern Plains (Castañeda 1904).
Conclusions
Our findings provide an important test of ethnographic accounts documenting dog transport loads. First contact between European and many Plains and Intermountain West cultures occurred during a period of significant cultural change initiated by the replacement of indigenous dog breeds with European stock, but also the introduction of horses and guns which significantly impacted mobility and social organization in these areas (Hämäläinen 2003; Mitchell 2015; Wissler 1914). Henderson’s (1994) experiment showed that dogs hauling travois loaded with around 45% of their body mass were capable of long distance travel of as much as 27 km at speeds consistent with those of a walking person. Though Henderson’s (1994) data document the capabilities of only one breed, the Alaskan husky, they remain the only experimental dataset available. Assuming these data capture the capabilities of prehistoric Native American dogs, the data presented here indicate that at least some Intermountain West and Great Plains dogs were capable of achieving loads of the size reported in ethnographic sources. Ethnographic accounts report that Intermountain West and Great Plains families frequently owned
97 between four and six dogs (Catlin 1973; Lowie 1963; Russell 1964; Wilson 1924) meaning they could provide a large pool of collective labor for historic societies.
Despite more than a century of research, many questions linked to Native
American dogs, changes in dogs’ physical characteristics through time, and the initiation and influence of selective pressure on dogs capable of hauling larger loads have yet to be fully explored. Though such studies have at times categorized Native American dogs into breeds (e.g., Allen 1920; Crockford 1997; Gleeson 1976; Olsen 1976; Worthington 2008) such efforts have been critiqued for oversimplifying population diversity (Lawler et al.
2016). This research has revealed archaeological dogs could have fulfilled a diverse suite of roles within Native American society including labor, to hunting, and even fiber exploitation (Allen 1920; Crockford 1997; Worthington 2008). Here we have attempted to contextualize archaeological remains within ethnographic and biological data concerning the roles and capabilities of dogs within Intermountain West, Great Plains, and Great Basin communities. These ethnographic sources indicate at least two types sizes of dog were present in the Intermountain West (Hultkrantz 1954, 1956, 1967). The largest of these was used for transporting goods and provisions using travois and packs
(Hultkrantz 1954, 1956, 1967) and was likely under some level of selective pressure for the size and stamina needed for transporting heavy loads. Smaller dogs were used predominantly for hunting rodents and other small game (Hulkrantz 1956) and Great
Basin (Lowie 1924, 1939; Steward 1933).
Additional morphometric data on dogs are needed to improve our understanding of dog populations in the Intermountain West, Great Plains, and Great Basin. Though numerous archaeological dogs from the Great Basin and the Intermountain West regions
98 are known, they are often incompletely documented (e.g., Cressman et al. 1950; Stanford
1978). Similarly, dogs of greater antiquity are needed from the Great Plains in order to assess when selection for large bodied dogs capable of hauling and carrying useful loads was initiated. Identifying when large bodied dogs appear within the archaeological record may provide an avenue for identifying the origins and antiquity of dogs’ use in transport activities. Consequently, existing collections warrant further investigation and more detailed publication and we hope to pursue such lines of research in the future.
Acknowledgements:
We would like to thank the Bureau of Land Management Idaho Fall Field Office and the
Idaho Museum of Natural History for permitting access to the Birch Creek Site assemblage. We give further thanks to Rob Bozell for providing us with the original morphometric data on Plains dogs, Penn State undergraduate Madie Daisley for her assistance in data entry, and Dr. Sarah McClure for her input in the development of this analysis.
Chapter 4
Traveling with the Pack: The Field Processing Model and the Role of Dogs in
Intermountain and Plains Residential Mobility
Domestic dogs feature prominently in ethnographic and historic accounts of
Native American communities in the Intermountain West and Great Plains for their role in transporting goods on simple drag-sleds called travois or in pannier-style packs.
However, dogs are reported to have been used predominantly to transport a limited number of goods including firewood, shelters and household belongings, trade goods, and bison meat following large communal hunts. The Field Processing Model, a fundamental model from Human Behavioral Ecology, shows that a combination of processing and transport costs make dogs’ use in transporting many food items untenable, except in cases where many trips are necessary and processing costs are relatively low. These data indicate that dogs were not adapted to use as draught animals for the daily acquisition of food, but rather to facilitate the transport of equipment, shelter, and food reserves that offset risks of resource shortfall. Studies have shown hunter-gatherer investment in technology and degree of residential mobility are strongly correlated with environmental unpredictability (Collard et al. 2011; Kelly 2007). Draught dogs and transport technology are thus elements of a highly specialized adaptive system used by Intermountain and
Plains foragers to counter challenging environmental conditions whose use enabled the transport of valuable shelter, equipment, resources, and preserved food supplies.
100 Introduction:
Dogs filled many roles in past human societies as hunting aids, sources of fiber for textiles, food, and as beasts of burden by highly mobile foragers in the American and
Eurasian Arctic, and the North American Intermountain West and Great Plains (Allen
1920; Buskirk 1986; Crockford 1997; Hultkrantz 1967; Kelly 1934; Pitulko and
Kasparov 2017; Schaeffer 1978; Spier 1933; Talayesva 1942). Archaeological evidence for dogs’ role as draught animals is relatively scarce. Sled fragments recovered from the
Zhokhov Site, Siberia suggest dogs were used as draught animals by at least 9,000 BP
(Pitulko and Kasparov 2017). Evidence for dogs’ role as beasts of burden in North
America is uncommon, and largely constrained to ethnographic accounts and fragments of wood identified as a travois, a simple drag-sled hauled by dogs in Intermountain and
Plains communities, recovered from historic contexts (Buffalo-Bird-Woman in Wilson
1924; Gebhard et al. 1964; Grey 1963). Despite the absence of archaeological evidence, is seems likely that dogs’ role in Intermountain and Plains transport developed in the
Northeastern Plains at some date before 900 AD (Brasser 1982; Driver and Massey
1957:298; Wissler 1910) and was already well developed in the Southern Plains by the
1540s (Castañeda 1904).
The most complete ethnographic sources concerning Plains travois dogs are a series of interviews with three Hidatsa informants, Buffalo-Bird-Woman and her brothers
Henry Wolf Chief and Edward Goodbird (reported by Gilbert Wilson in 1924). Buffalo-
Bird-Women and her brothers were traditional Hidatsa living on the Fort Berthold
Reservation in North Dakota. Wilson’s report of these interviews (Wilson 1924) includes
101 a highly detailed compilation of traditional tales coupled with descriptions of pre- reservation life, and the use and management of both horses and dogs by the Hidatsa.
Additional information may be drawn from the published journals and accounts of early explorers, army officers, and ethnographers ranging in date from the 1540’s (see
Castañeda 1904), to the mid1900’s (Harman 1957; Hind 1971; Maximillian 1982;
Grinnel 1962; Russell 1964; Weltfish 1965; Winship 1896).
These sources assert that selective management including the culling of small pups and castration of male dogs not wanted for breeding purposes was used to increase dog size and stamina (Buffalo-Bird-Woman in Wilson 1924). These management practices adapted dogs in Plains and Intermountain West communities to transporting loads as large as 45kg (100 lb) using travois and pannier-style packs. Henderson (1994) reports that dogs’ load capacity is highly correlated with body size, local environmental conditions, and distance. Welker and Byers (under review; Chapter 3) have recently used Henderson’s (1994) data to show that some dogs found in Intermountain West and
Plains assemblages were capable of ethnographically reported loads, but only over short distances. Draught dogs’ prominence in ethnographic accounts suggests they were adapted for transport roles with great effect but does little to explain the conditions that led to dogs’ use in transport, or investment in transport technology by Intermountain and
Plains foragers but not other groups.
Collard et al. (2011) and Kelly (2007) assert that both investment in technology and residential mobility among hunter-gatherers are strongly correlated with environmental unpredictability and environments characterized by low primary biomass, including the Arctic and the North American Great Plains. These data imply that the
102 adaptation of dogs to draught roles may have emerged as a response to foragers’ use of residential mobility to escape unfavorable localized environmental conditions or target short lived resource hot-spots. However, ethnographic accounts document dogs’ use in transporting a limited number of largely non-food goods including shelters (poles, bison hides, etc.), firewood, bison meat, and trade goods such as furs, obsidian, and pemmican
(Ewers 1945, 1955; Griffin et al. 1969; Hatch et al, 1990; Schaeffer 1978; Wissler 1910).
With the exception of bison meat, dogs’ role in transport appear to have been heavily linked to residential mobility, camp management, and other activities predominantly associated with Intermountain and Plains women (Buffalo Bird Woman in Wilson 1924;
Henderson 1994). This is further supported by the assertion that women were charged with managing and loading camp dog populations (Buffalo Bird Woman in Wilson 1924;
Henderson 1994).
In this paper I investigate the costs and benefits of using dogs in Plains-style travois transport activities and evaluate why dogs are not more commonly linked to food transport. To accomplish this, I draw upon data on the abundance, weight, and processing costs for various food resources in the Intermountain West, Great Basin, and Great
Plains. These data are used to assess the costs and benefits of using dogs to transport these resources using the Field Processing Model (Bettinger 2009; Bird and Bliege Bird
1997; Metcalfe and Barlow 1992; Price 2017) drawn from Human Behavioral Ecology.
103 Dogs in Foraging Societies - Hunting, Provisioning, and Transport
Dogs are widely documented in the acquisition and transport of prey during hunting expeditions (Buskirk 1986; Hultkrantz 1967; Kelly 1934; Lupo 2011; Spier
1933; Talayesva 1942). Dogs in Intermountain West and Plains communities were used in catching, retrieving, or driving rabbits, squirrel, groundhog, quail, various waterfowl, antelope, mule deer, mountain sheep, and bison (Buskirk 1986; Hultkrantz 1967; Kelly
1934; Spier 1933; Talayesva 1942). Dogs frequently accompany African Bofi and Aka men, women, and children on foraging expeditions where they are used to drive prey out of thickets, dig prey out of logs, and track larger prey (Lupo 2011). Though dogs accompany the majority of Bofi and Aka hunts and are associated with shorter hunt times, dogs are also correlated with an overall reduction in the number of prey acquired (Lupo
2011).
The most appreciable cost to keeping domestic animals, including dogs, is their provisioning. Ethnographic sources suggest that when dogs are provisioned by humans, they are commonly fed portions of carcasses deemed inedible or undesirable (Ewers
1955; Lupo 2011). However, a significant body of literature indicates that dogs in foraging societies subsist largely on scraps scavenged from camp garbage (Lupo 2011).
These data also point to a strong connection between hunting success and dogs receiving food directly from human foragers (Lupo 2011) and imply that dogs in many foraging societies are semi- or predominantly self-sufficient.
In addition to acting as hunting aids and maintaining camp cleanliness by consuming garbage, dogs were adapted and trained to provide labor in the Eurasian and
104 North American Arctic, and the North American Intermountain West, Great Plains, and
Southwest (Allen 1920; Buskirk 1986; Ewers 1945, 1955; Hultkrantz 1967; Kelly 1934;
Spier 1933; Talayesva 1942). Ethnographic records from the Plains document long-term selective management including the culling of small pups, and the development of transport technology (Buffalo Bird Woman in Wilson 1924; Ewers 1945, 1955). When animals are used for draught purposes an additional cost is incurred in the form of developing and manufacturing transport technology, and training animals to its use.
Transport dogs in Intermountain and Plains communities were frequently managed and trained by women who manufactured the travois and packs used in transport (Ewers
1955), and were associated with camp relocation, firewood collection, and other tasks predominantly performed by women (Buffalo Bird Woman, in Wilson 1924).
Ewers (1955) classifies travois into two general forms, improvised travois produced by tying two shelter poles together, and more permanent travois which served as sun shades and meat drying racks in camp (see also Forde 1902). Wissler (1910) suggests permanent travois may be more commonly used with dogs. Ethnographic sources indicate that travois production included cutting down two trees and leaving them dry (Buffalo Bird Woman in Wilson 1924). These poles were then scraped clean of bark and branches, tied together at one end, and a patch of leather or a wooden platform for cargo was attached (Buffalo Bird Woman in Wilson 1924). Ewers (1955) reports that the poles for horse travois produced by the Blackfeet were between 4-5 inches in diameter wrapped in rawhide to prevent splitting. Wissler (1910) reports that the poles in a dog travois he examined were 2.2 meters in length. Extensive experiments have shown that stone axes can reliably cut down small trees within 10-30 minutes (Dickson 1976;
105 Mathieu and Meyer 1997). Together, these data suggest that travois production may have taken only a few hours of active labor investment spread out over a period of several days. Well-made travois could reportedly be used for at least a year (Ewers 1955; Wilson
1924:218).
Training a dog to perform as a draught animal reportedly only involved tying the dog into a travois harness for several days until it became accustomed to the travois
(Buffalo Bird Woman, in Wilson 1924). Travois dogs are capable of transporting loads as high as 107% of their body mass (27.22-45.36kg; 60-100lb) over a few kilometers
(Henderson 1994). Dogs’ load capacity drops to only 13.61-22.68kg (45% of body mass,
30-50lb) on longer trips (Henderson 1994). Intermountain and Plains households kept between four and six dogs each (Kennedy and Stevens 1972; Russell 1964) implying a maximum collective short distance load capacity of 272 kg. Blackfeet informants reported to Ewers (1955) that a train of pack dogs could walk only 9.7-14.5 km (6-9 mi.) per day. Given that transport dogs and humans travel at similar average speeds, and humans average approximately 5 km an hour (Browning et al. 2006; Levine and
Norenzayan 1999; Mohler et al. 2007; Ralston 1958), this translates to only two or three hours of active transport time per day. Further, Henderson (1994) reveals that travois were only usable in flat, relatively unforested environments.
Involvement in hunting or transport activities would have prevented dogs from foraging for themselves while performing their roles, though they may have been fed when acting as draught animals. On the Plains, draught dogs are associated with the collection and transportation of firewood, shelters and household goods, and meat from communal bison hunts. Of these, only bison meat is a food item that could have been
106 used in feeding dogs involved in travois transport. As a result, dogs must have been fed either from stores carried with them, from game hunted during the trip, or when they were permitted to forage for themselves on days when they engaged in transport activities to satisfy their needs. To better understand the costs and benefits of transporting these items I explore the specific characteristics of the three most common goods transported by dogs.
Firewood:
Fuelwood requirements are determined by cooking and heating and are therefore linked to local environmental conditions and seasonality. In summer months, cooking would likely have accounted for much of the fuelwood used by Intermountain and Plains foragers. Data from several fuelwood studies suggest between 2.8-12.5 kg may be necessary for daily cooking when using open fires (Ichikawa 2001; Sangay 2011) (Table
4-1). Fuel for heat, particularly during cold winter months, would have added to the quantity of fuelwood required. Experimental studies in Finland have revealed that as much as 87.6 kg of firewood were needed per day to maintain comfortable temperatures inside traditional Sami tents of similar construction to Plains tipi (Östlund et al. 2013)
(Table 4-1). Plains communities accommodated fuelwood needs by selectively camping around springs and along river valleys where firewood could be collected by breaking dead branches off of trees using ropes, poles, or axes (Wissler 1910). Bison dung was also used when firewood was not readily available (Wedel 1963; Wissler 1910).
107 Table 4-1: Open fire fuelwood consumption in various societies.
Season Observations Average/day (kg) Error Bhutan (Camps) Summer 9 2.80 0.3 Sangay 2011 Bhutan (Stone house) Summer 3 4.70 1.2 Sangay 2011 Mbuti Summer NA 12.50 2.5 Ichikawa 2001 Sami Hut Experiment Winter 3 87.60 19.3 Ostlund et al. 2013
Shelter and Household Belongings:
The weight and number of belongings Intermountain and Plains households
owned is of great interest when studying the role dogs may have played in their transport.
However, ethnographic accounts largely postdate the introduction of the horse which
could transport larger loads more quickly and is believed to have encouraged significant
increases in Plains shelter size and the quantity of household goods, in addition to
influencing mobility and warfare (Dempsey 1986; Hämäläinen 2003; Mitchell 2015;
Wissler 1914). Voget (2001) reports that five horses were needed to transport a
household’s shelter and belongings, of these two or three were required to transport a tipi
of the type used following the introduction of the horse. Dogs could reportedly pull
between two and four poles from a historic tipi each during camp moves (Ewers 1953).
Ethnographic sources indicate shelters of the kind used prior to horses’
introduction were heavy enough that two dogs were required to transport them (Ewers
2012; Flannery 1953). Kehoe (1960:462) reports that the floor diameter of tipi, estimated
from rings of stone used to hold down tipi covers, increases from an average of 3.14 m on
sites predating the horse to nearly 9.14 m on sites dating to the 1870s. Ewers (2012) notes
that the typical small shelter cover consisted of 6-8 buffalo hides weighing 34.02 kg (75
108 lb) (Table 4-2). Laubin and Laubin (2012) provide estimates for two Plains tents. A
shelter similar to the small tent Ewers (2012) described reportedly weighed 45.36 kg (100
lb) (Table 4-2), implying the poles weighed 11.34 kg (25 lb). Larger historic period tipi
averaged three or four poles, but could include as many as 17 poles, each weighing
approximately 7 kg (15 lb). Together, these data indicate that a historic period tipi could
weigh over 154 kg (340 lb) (Table 4-2) (Laubin and Laubin 2012; Nabokov and Easton
1989; Otelaar 2003).
Table 4-2: Available weight data on Plains tipi.
Period Culture Tent/Object Weight Blackfoot Cover (6-7 buffalo hides) 22.68 kg (50 lbs) Ewers (1955) Blackfoot Cover (6-8 buffalo hides) 34.02 kg (75 lbs) Ewers (2012) Pre-Horse 11.34 kg Laubin and Laubin NA Poles (25lbs)* (2012) 45.36 kg (100 NA Whole tent lbs) Flannery (1953) 154 kg (340 Laubin and Laubin Post-Horse NA Poles (Tipi)-post horse lbs)* (2012) *Estimated based on description
Bison Meat:
Bison, a staple food in Plains diets, were acquired in large numbers from
communal hunts which could include between 4-500 foragers and generate between 100-
500 bison (Denig 1930; North 1859). Women, children and dogs assisted hunters in
communal bison drives by directing herds towards jumps and corrals and were commonly
tasked with the butchery and transport of bison meat afterwards (Denig 1930). Plains
109 women processed and transported large quantities of meat from communal hunts to camps located some distance away to avoid scavengers, flies, and the smell of rotting carcasses (Denig 1930; Laubin and Laubin 2012; White 1953). O’Connell et al. (1990) report that African foragers were observed hunting and transporting large game within a two hour foraging radius (Binford 1979, 1980, 1982; see also Morgan 2008). This distance corresponds well with the short distance loading strategy reported by Henderson
(1994) and the travel time on Great Basin foraging expeditions (Stewart 1933, 1938).
Acquisition of meat in quantities larger than can be consumed before spoilage occurs is effectively useless unless it can be preserved. Plains foragers coped with this challenge by jerking or smoking meat (Laubin and Laubin 2012). Jerking or smoking meat prevents spoilage and reduces the weight of meat by 60-80%, making it far lighter and easier to transport (Laubin and Laubin 2012; Lee 1979:223). Laubin and Laubin
(2012) report that two Plains women could process an entire steer carcass for jerking in a single day; however, Reidhead (1976:82-83) reports the actual jerking or smoking took
35 hours per 100 kg of meat. These data suggest Plains women were capable of preserving large quantities of meat, but that their abilities were constrained by the amount of meat which could be processed and transported before spoilage occurred. Dogs were likely crucial to both removing meat from the kill site before it spoiled, as well as transporting jerked meat when camps were relocated.
A notable feature of firewood, household belongings, and bison meat is that they occur in dense accumulations –firewood because wood was extracted from extant stands of trees, shelter and household belongings as a result of human mediated accumulation,
110 and bison meat because large numbers of animals were slaughtered in single events and the meat had to be processed before it rotted or attracted scavengers. Furthermore, all are heavy or bulky and costly to transport. These data suggest an important characteristic of resources that dogs were used to transport may be related to the density or size of initial loads available at a single location. To investigate this relationship further and assess why dogs are not more widely connected to the transport of food resources I apply a variant of the Field Processing Model from Human Behavioral Ecology developed by Bettinger
(2007:97).
Modeling Dogs as Draught Animals:
Evaluating when dogs are beneficial in transport necessitates understanding the implicit relationship between the relative costs and benefits of acquiring, processing, and transporting resources. Human Behavioral Ecology is a body of theory that seeks to understand human behavior and decision-making processes from an evolutionary and ecological perspective. Within Human Behavioral Ecology predictions about human behavior are frequently made by mathematically modeling specific behaviors under prescribed conditions. The Field Processing Model (Bettinger 2009; Metcalfe and Barlow
1992; Price 2017) is generally used to evaluate the relationship between transport distance, load quality, and processing time in order to predict when foragers will refine loads by removing waste material (shells, portions of a carcass low in meat, etc.) in the field before transport to base camps (Figure 4-1). For example, foragers traveling short distances can make many trips at relatively low cost (travel time) and may not be
111 motivated to shell nuts before transport since this can be done in camp. However, a forager who must travel farther, and can therefore make only a limited number of trips, may choose to remove waste material (e.g., nut shells) and either include more shelled nuts in their load, or reduce the weight of loads delivered to camp.
When considering travois transport, we are not interested in knowing when foragers will decide to process a load. Rather, we want to know when foragers will choose to increase the size of the load they transport by using dogs and thereby reduce the number of trips that must be made to acquire a resource too large for them to transport in one trip. In order for the Field Processing Model to function, two conditions must be met by the resources in question. These are:
1) The processed load must produce a higher caloric value. That is, a forager
processing in the field must receive higher caloric returns per load than he/she
would moving unprocessed loads.
2) The unprocessed load must generate at least equivalent returns per unit of
foraging time. This means a forager must be able to acquire a full load without
processing at least as quickly as they can collect a load when processing.
In addition, I make several assumptions specific to travois transport. First, foragers and dogs will transport only complete loads. Second, when resources exceed foragers’ ability to transport in a single trip, they will continue to make trips until the whole resource has been collected. Third, even though a forager must accompany the dog or dogs on
112 foraging trips, the dog and forager are never simultaneously transporting a load unless specified.
Figure 4-1: A graphical representation of the Field Processing Model as adapted to travois transport. U=caloric value, S=processing for a human load, P= additional processing needed for larger travois loads.
Experimental and ethnographic data are used here to establish load sizes for seven transport strategies. These include a forager working alone, a dog working alone, a forager and a dog transporting simultaneously, and a forager working with two dogs
(Figure 4-2). Human load capacity is reported to be between 10-30 kg for Hadza and
!Kung foragers in Africa (Lee 1979: 193,194; O’Connell et al. 1988) and set at 20 kg for the purposes of this model. Dog travois loads reported by Henderson (1994) are separated into short-distance (27.22-45.36 kg, 60-100 lb, 107% of body mass) and long-distance
113 (13.61-22.68 kg, 30-50 lb, 45% of body mass) load sizes. Though O’Connell et al. (1990) indicate that most large game transported to camps is captured less than an hour from camp, both short- and long-distance loads are calculated for all dog loading strategies in order to assess dogs’ potential in both short- and long-distance transport (Figure 4-2). I set these at 35 kg and 20 kg and respectively for this model.
In order to evaluate when a forager will transition from one loading strategy to another using the Field Processing Model one needs to compare cost and caloric value of each. To this end, processing costs and caloric return rates for taxa available to
Intermountain and Great Plains foragers were drawn from published and unpublished sources (Byers and Ugan 2005; Simms 1985, 1987; Ugan 2005; Table 4-3). These include some resources exceeding individual human carrying capacity such as bison, deer, mountain sheep, and pronghorn antelope, and others that individually are well within human carrying capacity such as rabbits, ground squirrels, and ducks. Of these, mountain sheep are unlikely to have been transported using travois because they occur in rugged high elevation conditions, which are ill-suited to travois use for much of the year
(see Henderson 1994). Ugan (2005) has shown that small-bodied taxa, which must be acquired in large numbers, are unlikely to have been appealing for travois transport because the cost of individually processing (i.e., skinning, etc.) each makes them economically inefficient and costly per kg of meat relative to larger game. Further, ethnographic accounts suggest that when small game such as rabbits were acquired in large numbers it was by groups who individually received loads well within their carrying capacity (Lowie 1924,1936; Palmer 1896; Speck et al. 1946; Speck and
Schaeffer 1950).
114
Figure 4-2: The loads evaluated in this study. *Note that because human and long-distance dog only travois loads are equivalent to one another this load does not meet condition one of the Field Processing Model.
Table 4-3: Caloric value, encounter rates, and processing time for resources available to Intermountain and Plains foragers (Data from Byers and Ugan (2005), Simms (1985, 1987), Ugan (2005).
Live Handling Return Rate Energy Weight Edible Edible Total Time Handling Time Encounter Encounter on-encounter Resource Scientific Name (kcal/kg) (kg/ind) Fraction Weight kcal (hr) (hr/kg) Rate (ind/hr) Rate (kg/hr) (kcal/hr) Bison Bison bison 1450 354 0.6 212.4 307980 4 0.0188 0.00324 1.747 40291
Deer Odocoileus hemionus 1200 85 0.6 51 61200 2.517 0.0493 0.01899 0.9685 24318
Mtn Sheep Ovis canadensis 1200 75 0.6 45 54000 2.517 0.0559 0.02086 0.9387 21457
Pronghorn Antelocapra americana 1140 56.5 0.6 33.9 38646 2.017 0.0595 0.0258 0.8745 19163
Hare Lepus spp. 1140 2.42 0.6 1.452 1655.28 0.125 0.0861 0.27399 0.3978 13242
Cottontail Sylvilagus spp. 1140 1.1 0.6 0.66 752.4 0.083 0.1263 0.49494 0.3267 9029
Gopher Thomomys sp. 1200 0.25 0.85 0.2125 255 0.042 0.1961 1.50362 0.3195 6120
Lg Squirrel Marmotta spp. 1200 0.35 0.85 0.2975 357 0.058 0.1961 1.16827 0.3476 6120 (Ground) Sage Grouse Centrocercus urophasianus 1340 1.5 0.7 1.05 1407 0.258 0.246 0.8125 0.8531 5446
Sm Squirrel Marmotta spp. 1200 0.2 0.85 0.17 204 0.042 0.2451 1.77755 0.3022 4896 (Ground) Ducks Anas spp. 1230 1 0.7 0.7 861 0.258 0.369 0.8125 0.5688 3333
Having identified bison, deer, and pronghorn antelope as the most likely candidates for travois transport, I calculate the per load caloric returns for each transported under each of seven human and travois loading strategies (Figure 4-2).
Though one could incorporate the manufacturing cost for travois into this model, it was decided that doing so would treat travois transport as if foragers manufactured a new travois each time. This assumption is unsupported by ethnographic sources, which document the reuse of travois as well as the use of travois poles in tipi (Ewers 1945,
1955). The caloric value of loads, also termed return rate or load utility, for each strategy is estimated as the caloric value per kg of a taxa multiplied by number of kg per load and divided by the processing costs:
푘푐푎푙 푝푒푟 푘푔 ∗ 푙표푎푑 푠푖푧푒 푢푗푖 = 푝푗푖 where uji = load utility pji = resource processing time*
* the resource processing cost used here is the estimated total processing time for
the whole resource to account for our foragers’ intent to acquire the whole
carcass.
These data are integrated into a variant of the Field Processing Model generalized to both large resources requiring field processing, or smaller resources collected in quantities provided by Bettinger (2007:97) with the formula:
푢푗 − 1 푇(푗 − 1 푡표 푗) = (푝푗) ( ) − 푠푒 푢푗 − (푢푗 − 1)
117 where
T= Travel Time (hrs) u j= Resource Utility (kcal/load) at stage j u(j-1)= Resource Utility (kcal/load) at stage j-1 pj=Time (hrs) required to process to stage j s=Time (hrs) required to procure a full unprocessed load e= coefficient of resource concentration (1 is dense, 0 is dispersed)
This formula predicts the round-trip travel time, or transition threshold, at which transporting loads using the two strategies modeled are equally effective, and beyond which a forager should decide to invest in the more-costly option. This time is then divided by the number of loads required to transport the entire animal in order to identify the travel time in minutes for individual loads. Multiplying the travel time by individual loads by an estimate of travel speed, which is similar for human and dogs carrying loads and set at 5 km per hour, allows us to estimate the per load travel distance at which point foragers are expected to transition from one loading strategy to another (Browning et al.
2006; Mohler et al. 2007; Levine and Norenzavan 1999; Ralston 1958).
Results:
The Field Processing Model is used to estimate the travel time at which dog-based transport becomes economically beneficial given the caloric value and processing costs of two loading strategies relative to one another. Though the Field Processing Model is
118 typically used to assess when foragers will choose to remove low-value portions of their load (e.g., Barlow and Metcalfe 1996; Bettinger et al. 1997; Bird and Bliege Bird 1997;
Metcalfe and Barlow 1992; Price 2017; Zeanah 2000), we use this model instead to predict when foragers will choose to change the size of the loads transported. When considering bison, deer, and pronghorn, the Field Processing Model predicts travois use in all but long-distance transport by dogs alone. In this case, dog and human loads approximate one another (20 kg) and no significant benefit is incurred by choosing dog- based transport over human transport.
When transporting bison over short distances, the Field Processing Model predicts foragers will adopt travois technology at only four minutes (0.413 km), to supplement their own load with either one or two dogs at three minutes (0.279 km and 0.228 km) respectively (Table 4-4, 4-5). Similar results are generated using long-distance loads calculated for humans working with one dog (four minutes, 0.355 km) and two dogs
(three minutes, 0.266 km), These results imply that the relative cost of producing and transporting larger loads is worthwhile if the bison can be returned to camp in fewer trips
(see Table 4-4, 4-5).
When applied to deer (see Table 4-4, 4-5), the Field Processing Model predicts transition times for short distance transport ranging between 54 minutes (4.51 km) for dogs alone, and foragers using two dogs at 16 minutes (1.42 km). Long distance transitions can only be modeled for foragers working with one and two dogs, at 46 minutes (3.86 km) and 29 minutes (2.47 km). For pronghorn antelope (Table 4-4, 4-5) the short distance transition time fall at 1 hour 34 minutes (5.8 km) for dogs alone, 40
119 minutes (3.19 km) and 19 minutes (1.6 km). For long distance foragers should work with dogs at 1 hour 11 minutes (4.96 km) and 35 minutes (2.98 km).
Notably, the transition distances at which the Field Processing Model predicts travois use for deer and pronghorn frequently exceed the 2 km distance Henderson
(1994) suggests as a limit for short distance transport. The most obvious dichotomy between bison, deer, and pronghorn is the number of necessary trips. Bison are large, and even when foragers supplement their own labor with two dogs require more than two trips to bring back to camp. Deer and pronghorn exceed the capacity of a human or dog working individually but can be transported in one trip by human foragers and dogs working together. As a result, the per trip time and distance are not spread out across several loads and the relative cost of a trip remains high.
Discussion:
Ethnographic accounts of travois use by Plains and Intermountain communities emphasize the role of dogs in transporting non-food items including firewood, shelter, trade goods, and at times bison meat from communal drives (Ewers 1945, 1955; Griffin et al. 1969; Hatch et al, 1990; Wissler 1910). The Field Processing Model shows that travois are only economically beneficial as a means of transport for food taxa large enough to partition transport costs across numerous trips. As a result, it seems unlikely that travois were employed in food acquisition and transport except in very specific circumstances such as large-scale bison hunts, and likely only over relatively flat terrain
(Henderson 1994).
Table 4-4: Short- and long-distance handling time (in hours), loads required, and kcal/load data for bison, deer, and pronghorn antelope.
Short Distance Human (20 kg) Dog Only (35kg) Human & Dog (55kg) Human &Two Dogs (90 kg) handling/load # Loads kcal/load handling/load # Loads kcal/load handling/load # Loads kcal/load Handling/Load # Loads kcal/Load
Bison 0.377 10.620 7250.000 0.659 6.069 12687.500 1.036 3.862 19937.500 1.695 2.360 32625.000 Deer 0.986 2.550 9535.161 1.726 1.457 16686.532 2.712 0.927 26221.692 4.437 0.567 42908.224 Pronghorn 1.190 1.695 11303.917 2.083 0.969 19781.854 3.273 0.616 31085.771 5.355 0.377 50867.625 Long Distance Human (20 kg) Dog Only (20kg)* Human & Dog (40kg) Human &Two Dogs (60 kg) Handling/Load # Loads kcal/Load Handling/Load # Loads kcal/Load Handling/Load # Loads kcal/load Handling/Load # Loads kcal/Load
Bison 0.377 10.620 7250.000 0.377 10.620 7250.000 0.753 5.310 14500.000 1.130 3.540 21750.000 Deer 0.986 2.550 9535.161 0.986 2.550 9535.161 1.972 1.275 19070.322 2.958 0.850 28605.483 Pronghorn 1.190 1.695 11303.917 1.190 1.695 11303.917 2.380 0.848 22607.833 3.570 0.565 33911.750 *Long distance dog only loads do not meet model condition because the caloric value of dog only and human loads are identical
121
Table 4-5: Travel time and distance transition points from human loads to travois loading strategies.
Short Distance Total Transition Time (hr) Per Trip Transition Time (hr) Per Trip Transition Distance (km) Dog Only Human & Human & 2 Dog Only Human & Human & 2 Dog Only Human & Human & 2 Dog Dogs Dog Dogs Dog Dogs Bison 0.502 0.215 0.108 0.083 0.056 0.046 0.414 0.279 0.228 Deer 1.315 0.563 0.282 0.902 0.563 0.282 4.511 2.817 1.409 Pronghorn 1.587 0.680 0.340 1.587 0.680 0.340 7.933 3.400 1.700 Long Distance Total Transition Time (hr) Per Trip Transition Time (hr) Per Trip Transition Distance (km) Dog Only Human & Human & 2 Dog Only Human & Human & 2 Dog Only Human & Human & 2 Dog Dogs Dog Dogs Dog Dogs Bison N/A 0.377 0.188 N/A 0.071 0.053 N/A 0.355 0.266 Deer N/A 0.986 0.493 N/A 0.773 0.493 N/A 3.867 2.465 Pronghorn N/A 1.190 0.595 N/A 1.190 0.595 N/A 5.950 2.975
This application of the Field Processing Model has tested the costs of travois transport against the utility of individual animals. Ugan (2005) reports that the most significant cost to collecting resources in large numbers is the associated increase in handling time incurred with each new individual, especially when processing costs cannot be performed simultaneously. These costs are such that, though locally abundant resources (e.g. rabbits) can be collected in mass, the time needed to process (skin) large numbers may actually make them unattractive to foragers. Processing costs are especially challenging for small-bodied prey, which can require 25-800% more processing per kilogram than large game (Simms 1985, 1987). These costs are likely amplified by the technological investment required for manufacturing nets, baskets, beaters, and other tools used in their acquisition, and therefore, only appealing under very specific circumstances (see Ugan 2005; Ugan et al. 2003).
Ethnographic sources report that bison, deer, pronghorn antelope, and occasionally elk, were hunted in groups through a variety of communal hunting methods
(Labelle and Pelton 2013; Lubinski 1997, 1999), but dogs are only reported to have been used in transporting meat from bison recovered from communal hunts (see Schaeffer
1978). The low processing cost per kg of meat achievable from deer and pronghorn- antelope relative to small-bodied taxa may have enabled foragers to overcome the travel and processing limitations identified by the Field Processing Model when caught in numbers. However, this will only be possible if these taxa occur in dense enough populations to generate multiple animals for transport in one hunting trip. Elk and deer occur in smaller herds than bison or pronghorn making it less likely that they would have been acquired in large enough numbers to encourage travois use (Table 4-6). Though
123 pronghorn antelope do occur in herds of between 20-60 individuals (Beuchner 1950), this is only the case during the fall/early winter when smaller herds congregated (Table 4-6), meaning large numbers of pronghorn antelope were only available on a seasonal basis.
Notably, ethnographic reports indicate that antelope meat was not well liked, and that pronghorn were hunted primarily for their hides (Brumley 1984:109; Fowler 1989:14-19;
Irving 1837:214; Maximilian von Wied 1906:298; Turney-High 1937:119; Wissler
1910:38), a fact which may have limited the need for travois transport even when they were caught in large numbers.
Despite the absence of direct caloric values, a similar cost benefit relationship may be inferred discussed for travois use in transporting non-food items mentioned in ethnographic sources. Some of these (e.g., fuelwood) would be required on a daily basis, while others (e.g., shelter, household belongings, and bison meat) would only be transported when a communal bison hunt occurred or when camp was relocated. Plains communities reportedly moved camp between 10 and 40 times per year at contact
(Gussow 1964; Nabokov 1967; Rogers 1963, 1967a, b, 1972). Estimated fuelwood needs for one family’s summer cooking could have been easily transported by humans in a single load (2.8-12.5 kg); however, if winter heating costs were similar to those needed to heat Sami tents (87.5 kg), five trips would have been needed. Using dogs drops this to only three trips, and a single forager working with two dogs could have transported this much wood in a single trip.
Bison meat and household belongings less commonly transported by travois provide similarly substantial benefits. A single 212.4 kg bison when jerked would weigh only 42.48 kg (Laubin and Laubin 2012). This is a load within the capability of one or
124 Table 4-6: Herd density for bison, elk, mule deer, mountain sheep, and pronghorn antelope.
Scientific Name Herd density Source Bison Bison bison 5-24 (2-220*) Soper 1941 Elk Cervus canadensis 3-13.5 Coughenout and Singer 1996; Singer et al. 1997 Mule deer Odocoileus hemionus 1-24 Kucera 1978 Mountain sheep Ovis canadensis 0.8-2.6 Jorgenson et al. 1997; Festa-Bianchet et al. 1995 Pronghorn antelope Antelocapra americana 6-20 (20-60*) Buechner 1950 *Large herds of bison and antelope form during the late fall/winter following the rut (Buechner 1950; Soper 1941:392).
two dogs and could provide between 35-84 meals, depending upon fat content (Morris et
al. 1981). The quantity and weight of tents and household belongings are poorly
documented; however, Hansen (2011) reports that it could take as much as two years to
acquire the hides and sinew needed to produce a tipi cover. Tipi production and the
associated cost of processing bison, tanning hides, sewing hides together, and cutting
poles could require hundreds of hours even when tipi poles doubled as travois poles (see
Reilly 2015; Rosoff 2011). Measured against these benefits, the cost of spending two
hours manufacturing a travois could easily have been recouped.
Conclusion:
Ethnographic sources highlight the importance of dogs in hauling cargo on travois
to Intermountain and Plains communities, but rarely describe their use in transporting
food items (Ewers 1945, 1955; Griffin et al. 1969; Hatch et al, 1990; Wissler 1910).
Using the Field Processing Model from Human Behavioral Ecology we investigated
whether or not using dogs would have improved the efficiency with which large food
125 items such as bison, deer, and pronghorn antelope could have been transported. Though the Field Processing Model is generally used to assess when low-value portions of a load will be removed prior to transport, we adapt it to predict the travel time at which foragers will choose to increase load size using one of seven loading strategies.
This analysis reveals that dog-based travois transport becomes economically beneficial in food transport when per kg processing costs are relatively low and several trips are required. Of the resources tested here, bison are the only one that satisfies these criteria. While elk, deer, mountain sheep, and pronghorn antelope are all large enough to have required field processing, they are unlikely to have been transported using travois for a number of reasons. First, elk were rare in the Intermountain West until the 1800s when European settlers began hunting wolves in the region (White et al. 1998). Similarly, mountain sheep occur in environments that are unfavorable to travois use (see Henderson
1994), deer occur in small herd sizes, and pronghorn antelope were reportedly not favored for their meat (see Brumley 1984:109; Fowler 1989:14-19; Irving 1837:214;
Maximilian von Wied 1906:298; Turney-High 1937:119; Wissler 1910:38). Small-bodied taxa would likely also have been unappealing for travois transport because high individual processing costs would have made them unappealing in the numbers needed to require travois transport (see Ugan 2005).
These data suggest the primary benefit in dog-based travois transport was the ability to supplement human labor. Technological investment and residential mobility among foragers are strongly correlated with environmental unpredictability and the risk of resource shortfall (Collard 2011; Kelly 2007). Plains and Arctic foragers expended significant effort and resources in developing large dogs capable of hauling belongings
126 and developing technology which could be used to increase dogs’ load capacity (Buffalo
Bird Woman in Wilson 1924). Notably, foragers in these two regions also exhibit high residential mobility moving camp between 10-60 times a year (Gussow 1964; Nabokov
1967; Rogers 1963, 1967a, b, 1972, see data compiled by Kelly 2007). Combining dog- based transport and high residential mobility enabled Intermountain and Plains foragers to transport equipment, shelter, and food stores used to offset unpredictable environmental conditions, while remaining mobile enough to target patchy or seasonally abundant resources (Kelly 2007).
The archaeological signature of dog-based transport is scarce and likely strongly influenced by historic period ethnographic sources frequently describing societies already influenced by the horse (Dempsey 1986; Hämäläinen 2003; Mitchell 2015; Wissler
1914). Various analyses have drawn a strong link between the development of the tipi in the Northeastern Plains and the use of travois and have placed these cultural adaptations at some time before 900 AD (Brasser 1982; Driver and Massey 1957:298; Wissler 1910).
Dogs exhibiting pathologies attributed to travois travel and dating to between 4,000-
5,000 BP have been reported from the Gray Site in Saskatchewan (Millar 1978:365-369).
Recent analysis indicates that dogs from Intermountain and Plains assemblages were capable of loads reported in ethnographic reports, and that dogs of this size have existed for over 3,000 years (Welker and Byers under review; Chapter 3). Together with the improved understanding of travois transport provided by this analysis, these results suggest residential mobility and draught dogs were key components of an adaptive strategy which may have been used by foragers in these regions for thousands of years.
Chapter 5
Conclusions and Future Directions
Domesticated plants and animals have been an intrinsic component of life in human communities throughout the Holocene. Domesticates have filled numerous, at times highly specialized, roles within human societies and are central elements in many of the most enduring questions facing the field of anthropology (Kintigh et al. 2014). The ability to establish, manage, and rely upon domesticated crops and livestock was vital to many human migration events, the colonization of new and unfamiliar landscapes, the rise of social complexity, and urbanization (Campana et al. 2010; Crabtree 1990;
DeFrance 2009; Gumerman 1997; Landon 1997; Zierden and Reitz 2016; Welker et al. under review; Zeder 1991). Domesticates’ significance within these processes and adaptation to roles within human communities drove the creation of anthropogenically constructed environmental conditions (see McClure 2015). Existence within these environmental conditions and long-term cohabitation and management contributed to the formation of morphologically and genetically distinct populations variably termed breeds, landraces, or types (Clutton-Brock 2012:3; Rege 1999; Rege and Tawah 1999;
Sponenberg et al. 2014).
The concept of breeds has been complicated by breed formalization or improvement efforts used since the 18th Century by Western livestock managers to create highly-specialized livestock breeds (Rege 1999; Rege and Tawah 1999; Sponenberg et al.
128 2014). Such formalized breeds are rare outside of modern, Western, cultures (Rege 1999;
Rege and Tawah 1999; Sponenberg et al. 2014). Archaeologists are instead more likely to encounter the progenitors of these breeds, populations similar to the contemporary heritage breeds, landraces, or types from which formalized breeds developed, that still characterize livestock populations in many developing nations. These populations are commonly more heavily influenced by natural selection and therefore adapted to local environmental conditions (Sponenberg et al. 2014). As a result, these populations tend to be geographically defined, and are frequently unspecialized (i.e. it is less likely there will be distinct dairy and beef breeds) (See Rege 1999; Rege and Tawah 1999; Sponenberg et al. 2014).
Archaeologists have expended significant energy investigating the initial domestication of plant and animal taxa and have developed numerous methods that permit the differentiation of domesticates from their wild progenitors (Ameen et al. 2017;
Clark 1996; Evin et al. 2013). Archaeological analyses have employed these methods in concert with other datasets to investigate the social, ecological, and economic conditions that led to domestication (Coppinger and Coppinger 2001; Hale 1969; Morey 1994; Price
1984, 2002; Larson et al. 2005; Zeder 2012; Zeder and Hesse 2000). Further research has centered on exploring animals’ roles within human society as sources of traction, food, and other products (de Cupere et al. 2000; Groot 2005; Lin et al. 2016; Salmi and
Niinimäke 2016; Telldahl 2012; Thomas 2008; Zeder 2012); however, this research has rarely focused on identifying the conditions under which morphologically distinct populations of domestic animals emerged, or how identifiable breeds are within archaeological assemblages.
129 Phenotypic changes in domestic plant and animal populations result from the intersection of natural and artificial selection (Clutton Brock 2012:3; Trut et al. 2004;
Trut et al. 2009; Zeder 2012). As a result, breed development is a legacy effect of enduring biological, cultural, and ecological processes and represents a record of human and animal interaction that may be used to understand aspects of this relationship through time. Studies focused on the long-term history of domesticates in association with human societies have shown great promise. Characterizing and tracking changes in phenotypic expression within and between populations can provide a powerful source of evidence for population movement and interaction, and the changing roles of domesticates within human communities (Albarella 1997; Bennett et al. 2016; Davis 2008; Davis and Beckett
1999; Horard-Herbin et al. 2014; MacKinnon 2010; Van and Grimm 2010). However, achieving this level of understanding requires substantial datasets of morphometric or genetic data, and an understanding of ecological theory and the natural morphological variation underlying breed level characteristics.
Domestic dogs are a useful test species because of an advantageous combination of ecological, biological, and archaeological factors. Dogs exhibit relatively limited sexual dimorphism (Bidau and Martinez 2016; Frynta et la. 2012; Sutter et al. 2008) and, as a result of early domestication, global distribution, and adaptation to many roles within human societies, have developed morphological variability to a degree not found in any other domesticate (Allen 1920; Crockford 2000; Horard-Herbin et al. 2014; Lupo 2017;
Schwartz 1998). This morphological variability is captured in numerous ethnographic and historic records. The combination of these records with abundant archaeological remains and specialized methodological techniques for characterizing and quantitatively
130 comparing dog morphology make dogs an excellent case study for investigating population level morphological variability and associated elements of human culture (see
Clark 1996; de Grossi Mazzorin and Tagliacozzo 2000; Harcourt 1974; Losey et al. 2015;
Losey et al. 2017; Phillips et al. 2008; Tourigney et al. 2016; Van Valkenburgh 1990;
Wing 1978).
This dissertation explored the potential to identify morphologically distinct populations of domestic dog within the archaeological record and harness this recognition to better understand dogs’ roles within Native American and European communities in the Americas. In doing so, this dissertation revealed notable advances in understanding the ways in which cultural preferences and uses in colonial and Native American communities shaped the phenotype of archaeological dog populations. Capitalizing on the strategic position afforded by dogs’ biology and the abundance of archaeological data, I have explored the potential for understanding both specific decision-making criteria that resulted in the development of and transplantation of specific dog populations, as well as the long-term legacy effects of domestication on population-level morphological characteristics.
Using a combination of morphometric data, ethnographic sources, and quantitative modeling, this dissertation investigated the potential to use morphologically distinct populations to investigate the human past. Chapter Two employed data on
Native American and European dogs to show that that the environmental and cultural conditions influencing Native American and European dog populations resulted in morphologically distinct populations. Further, a widespread preference for large working dogs among European colonists resulted in an over-representation of mastiffs, hounds,
131 and other dogs used in warfare, hunting, and defending livestock and settlements. This analysis articulates both the selection processes underlying which dogs were introduced to North America by colonists and presents evidence that colonists’ preferences resulted in a distinct and identifiable population of dogs in North American colonial contexts.
Chapters Three and Four shifted the focus from investigating dogs as cultural artifacts or type specimens akin to archaeological cultures, to investigating the selection processes underlying the adaptation of domesticates to specialized roles. Dogs have been used for transport by Arctic foragers in Eurasia and North America, as well as those in the Intermountain West and Great Plains (Allen 1920; Buskirk 1986; Hultkrantz 1967;
Kelly 1934; Pitulko and Kasparov 2017; Spier 1933; Talayesva 1942). Ethnographic accounts of Intermountain and Plains communities prominently feature draught dogs but mention a relatively limited number of mostly non-food goods that were transported
(Allen 1920; Buskirk 1986; Hultkrantz 1967; Kelly 1934; Spier 1933; Talayesva 1942).
Chapter Three combines morphometric, experimental, and ethnographic data to investigate the potential for dogs to haul cargo in the Intermountain West and Great
Plains. This analysis shows that Native American selective management strategies were not only successful in biologically adapting dogs to draught through encouraging large body size and stamina, but also revealed evidence that these dogs were capable of loads reported in ethnographic sources. In addition, directly dated dog bone from the Veratic
Rockshelter in Idaho revealed and that large dogs capable of ethnographically reported loads may have existed in the Americas for over 3,000 years.
Chapter Four built upon our improved understanding of travois transport.
Ethnographic accounts report dogs’ use in transporting a limited suite of largely non-food
132 items (shelter and household belongings, firewood, trade goods, and bison meat following communal hunts. Using a modified form of the Field Processing Model from
Human Behavioral Ecology provided by Bettinger (2007:97) I explored the economic costs and benefits of using travois in food transport. This analysis suggests that the combination of processing costs involved in refining animal carcasses to a portable size and manufacturing costs associated with producing travois make their use impractical for many food resources. The exception to this rule was bison which are large enough that the larger loads possible with dog-based transport resulted in significantly reduced transport time. These findings are significant because they indicate that dog-based transport likely developed as an adaptation to unpredictable environments characterized a patchy distribution of resources and a relatively high risk of shortfall. Moving from patch-to-patch in this manner enables foragers to take advantage of seasonal or locational abundances but raised a series of logistical challenges associated with transporting shelter, equipment, and food stores. Investment in developing large dogs capably of transporting shelter, food stores, and other goods enabled Intermountain and Plains foragers to construct a portable niche and protect against environmental risk (Collard et al. 2011; Kelly 2007).
Future Directions:
These analyses have established a foundation from which to undertake future studies but have only scratched the surface of an under-explored and promising field of study. The confluence of the direct historical approach, evolutionary and ecological
133 theory, and abundant archaeological records is a fertile field in which to address a variety of significant questions and have a lasting impact on the field. This dissertation lays the groundwork for a series of future studies which I have already begun to plan out and undertake.
In developing datasets for Chapters Two and Three it has become clear that some regions and/or temporal phases lack datasets large enough to characterize population-level morphology. Small sample sizes are in some cases directly linked to the absence of archaeological specimens; however, they may also reflect either analysts’ inability to differentiate dogs from con-specific wild canids including coyotes (Canis latrans) and wolves (Canis lupus), or analysts not collecting and reporting morphometric data. I have begun to address these sampling issues in several ways. First, I am developing promising new methods capable of detecting differences in skeletal morphology between domestic dogs, coyotes and wolves, which have contributed to regionally poor datasets on dogs in North America. Preliminary results for a new method for the mandible using morphometric data collected on 440 dog, coyote, wolf, and fox mandibles indicate that the degree of curvature on the underside of mandibles may be useful in differentiating domestic dog mandibles from those of other canids.
I am also actively expanding datasets developed in Chapters Two and Three in order to undertake a continental level spatial and temporal analysis of Native American dog morphology. Morphometric data on domestic dogs from Native American sites
(n>650 specimens) are being compiled from the published and grey literature. This analysis revisits widely held assumptions about diversity in Native American domestic dogs based largely on the work of Allen (1920), and last investigated on a continental
134 scale by Haag (1948). By building upon Haag’s dataset with specimens from the published record and grey literature, I will explore both spatial and temporal trends in dog phenotype. Doing so provides opportunities to evaluate the diversity of dog forms in
North America, but also the emergence of particular dog types correlated with elements of human culture including agriculture, social complexity, sedentism and mobility.
Finally, analyses are needed to explore the potential for similar studies in other domesticates. I have begun to investigate the potential for using census records and morphometric data to study herd demographics and population origins in domestic cattle
(Bos taurus) introduced to North America by European colonists. Historical records including shipping manifests, census records, and probate inventories provide an invaluable source of data for assessing individual and community level herd demographics. These are issues of interest to archaeologists working on any agro-pastoral or pastoral society, but they are frequently inaccessible to those working in prehistory.
Furthermore, morphometric studies on cattle have produced tantalizing evidence that cattle from different source populations in Europe may be differentiated from one another
(e.g. Reitz and Ruff 1994; Cossette and Horard-Herbin 2003). Unfortunately, these studies suffer from several critical weaknesses. They have relied upon small datasets that may not accurately capture variability within these populations and have not accounted for sexual dimorphism and other elements livestock biology. Further, they have not developed datasets for European populations with which to test morphological patterns observed in North American assemblages.
Application of the morphometric methods employed in this dissertation also has significant potential for investigating long-term interactions between humans and wild
135 taxa. These relationships have garnered increasing attention as climate change has gained prominence within the public awareness. The archaeological record and morphometric analyses provide a means of extending our understanding of these relationships from the present into the past. Morphometric analyses have already been used to investigate topics ranging from predation pressure and over-fishing (Audzijonyte et al. 2013; Azevedo
2015; Roy et al. 2003), to game management and stocking (Wolverton et al. 2007), and conservation biology (Lyman 2009; Wolverton and Lyman 2012; Wolverton et al. 2016).
In the future I hope to engage with these issues using the lessons and experience afforded by this dissertation and associated research.
Appendix A
Shoulder Height in Native American, Colonial, and European Dogs
Date Shoulder Specimen Group Site Region (BP) Date Note State/County Culture Sex Age Height Range SHA VRG Citation
217 1 Tallahassee SE 250 Tennessee H M 52.30 – 53.31 52.81 Parmalee & Bogan 1978
218 1 Chota SE 250 Tennessee H M 34.18 – 35.54 34.86 Parmalee & Bogan 1978
219 1 Chota SE 250 Tennessee H M 55.63 – 57.94 56.79 Parmalee & Bogan 1978
114 1 Fig Springs (1) SE 318 Florida H M 56.10 – 61.74 58.81 Weisman 1991
192 1 Oliver (1) SE 365 Mississippi M F? 35.90-40.36 38.39 Yerkes 2000
236 1 Oliver (1) SE 365 Mississippi M U 40.90 – 42.88 41.84 Yerkes 2000
212 1 Citico SE 400 Tennessee M U 47.99 47.99 Parmalee & Bogan 1978
213 1 Toqua SE 400 Tennessee M U 53 53 Parmalee & Bogan 1978
215 1 Toqua SE 400 Tennessee M M 47.16 47.16 Parmalee & Bogan 1978 1540 (+/- 1 Cleveland NE 410 90) Ontario PH M 45.909-47.743 46.5 Bathurst & Barta 2004
85 1 Morris Village (1) SE 450 Kentucky M M 41.09 - 41.70 41.4 Haag 1948
223 1 Key Marco SE 494 Florida M U 43.61 43.61 Wing 1965
224 1 Key Marco SE 494 Florida M U 44.55 44.55 Wing 1965
225 1 Key Marco SE 494 Florida M U 44.23 44.23 Wing 1965
226 1 Key Marco SE 494 Florida M U 43.87 43.87 Wing 1965
227 1 Key Marco SE 494 Florida M U 41.82 41.82 Wing 1965
137 Date Shoulder Specimen Group Site Region (BP) Date Note State/County Culture Sex Age Height Range SHA VRG Citation
228 1 Key Marco SE 494 Florida M U 41.82 41.82 Wing 1965
229 1 Key Marco SE 494 Florida M U 43.81 43.81 Wing 1965
230 1 Key Marco SE 494 Florida M U 42.97 42.97 Wing 1965
231 1 Key Marco SE 494 Florida M U 44.56 44.56 Wing 1965
232 1 Key Marco SE 494 Florida M U 43.29 43.29 Wing 1965
233 1 Key Marco SE 494 Florida M U 53.03 53.03 Wing 1965
234 1 Key Marco SE 494 Florida M U 49.71 49.71 Wing 1965
235 1 Key Marco SE 494 Florida M U 53.47 53.47 Wing 1965
2 1 Armorel (1) SE 500 Arkansas M M 42.80 – 48.16 45.44 Mitchie & Mulvihill 1999
140 1 Greenhouse SE 550 Kentucky M U 38.9 38.9 Haag 1948 Hillside Burial 1 Emmons IL 570 600-540 Fulton Co, IL M F? A 43.3 Lawler et al. 2016
Burial 3 1 Emmons IL 570 600-540 Fulton Co, IL M F? A 41.9 Lawler et al. 2016 Lake Jackson Mound 113 1 (1) SE 592 Florida M M 40.80 – 44.56 42.84 Jones 1994
151 1 Henry Island SE 600 Alabama M U 42.35-43.43 42.89 Haag 1948
221 1 Ramon Santana WI 600 Dominican Republic M U 35.84 – 36.14 35.97 Lawrence 1977
222 1 Ramon Santana WI 600 Dominican Republic M U 35.08 – 35.66 35.35 Lawrence 1977
1 Cueva del Pirul MesoAmerica 600 700-500 Mexico F 38 Valdez Azúa et al. 2000
87 1 Fewkes (1) SE 650 Tennessee M M 41.24 – 41.25 41.25 Peres 2002
191 1 Chucalissa (1) SE 650 Tennessee M U 41.21-42.02 41.63 Parmalee 1960
237 1 Tubbs NE 650 Connecticut W M 44.61 – 49.24 47.61 Handley 2000
243 1 Meeting House:1 NE 650 Rhode Island W F 51.03 – 52.47 51.62 Handley 2000
259 1 Lambert Farm NE 650 Rhode Island W M 45.65 – 47.91 47.09 Handley 2000
1 1 Etowah (1) SE 670 Georgia M M 40.60 – 45.80 43.72 Larson 1978
93 1 Etowah SE 670 Georgia M M 45.90 – 49.02 47.24 Larson 1978
4 1 Ausmus Farm (1) SE 700 Tennessee M U 33.90 – 39.52 36.14 Webb 1938
138 Date Shoulder Specimen Group Site Region (BP) Date Note State/County Culture Sex Age Height Range SHA VRG Citation
13 1 Hampton (1) SE 900 Tennessee M M 36.80 – 40.75 39.02 Webb & Wilder 1951
14 1 Hampton (1) SE 900 Tennessee M M 45.84 – 41.10 43.45 Webb & Wilder 1951
98 1 Austin (1) SE 900 Mississippi M F 33.10 – 37.32 35.7 Connaway (N.D.)
99 1 Austin (1) SE 900 Mississippi M M 39.30 – 42.70 41.5 Connaway (N.D.)
100 1 Austin (1) SE 900 Mississippi M M 39.80 – 41.38 40.47 Connaway (N.D.)
101 1 Austin (1) SE 900 Mississippi M M 33.10 – 39.54 37.31 Connaway (N.D.)
102 1 Austin (1) SE 900 Mississippi M U 38.60 – 42.77 40.94 Connaway (N.D.)
103 1 Austin (1) SE 900 Mississippi M M 38.00 – 43.29 40 Connaway (N.D.)
106 1 Trail site (1) SE 1000 Florida W U 37.48 – 37.95 37.78 Carr 2003
220 1 El Carril WI 1030 Caicos W U 37.32 37.32 Wing 2001
110 1 Palmer (1) SE 1100 Florida W U 43.20 – 47.90 45.1 Bullen and Bullen 1976
111 1 Palmer (1) SE 1100 Florida W F 28.47 – 31.60 30.05 Bullen and Bullen 1976
112 1 Palmer (1) SE 1100 Florida W U 41.50 – 44.68 43.3 Bullen and Bullen 1976
Burial 4 1 Apple Creek IL 1225 1550-900 Greene CO, IL LW F? J 35.1 Lawler et al. 2016
Burial 1 1 Apple Creek IL 1225 1550-900 Greene CO, IL LW F? A 41.6 Lawler et al. 2016
Burial 3 1 Apple Creek IL 1225 1550-900 Greene CO, IL LW F? A 42.3 Lawler et al. 2016
Burial 5 1 Apple Creek IL 1225 1550-900 Greene CO, IL LW F? A 41.6 Lawler et al. 2016
184 1 Miller Cave SE 1300 Missouri W F 40.51-42.10 41.33 Darwent & Gilliland 2001
1 Guadalupe MesoAmerica 1300 1500-1100 Mexico M 42.4 Rodríguez et al. 2001
1 Guadalupe MesoAmerica 1300 1500-1100 Mexico F 37.7 Rodríguez et al. 2001
183 1 Kersey SE 1625 Missouri W M 49.12-51.77 50.44 Darwent & Gilliland 2001
84 1 Bell Shelter (1) SE 1650 Kentucky W M 39.90 – 42.99 41.68 Haag 1948
139 1 Stearns site SE 1650 Alabama W U 40.47 41.77 Haag 1948
144 1 Riley SE 1650 Alabama W U 44.23 44.23 Haag 1948
153 1 Deposit Landing SE 1650 Alabama W M 41.72 41.72 Haag 1948
139 Date Shoulder Specimen Group Site Region (BP) Date Note State/County Culture Sex Age Height Range SHA VRG Citation
154 1 Deposit Landing SE 1650 Alabama W M 42.66 42.66 Haag 1948
238 1 Granniss Island NE 1650 Connecticut W F 49.79 – 52.62 50.85 Handley 2000
239 1 Granniss Island NE 1650 Connecticut W M 56.00 – 57.00 56.47 Handley 2000
240 1 Granniss Island NE 1650 Connecticut W F 40.32 – 42.98 41.57 Handley 2000
244 1 South Truro NE 1650 Massachusetts W M 51.17 – 54.10 52.6 Handley 2000
245 1 Quincy/Squantum* NE 1650 Massachusetts W M 56.00 – 65.90 61.1 Handley 2000
251 1 Turner Farm NE 1650 Maine W M 54.43 – 54.63 54.53 Handley 2000
252 1 Port Washington NE 1650 Long Island W M 48.57 – 49.83 49.26 Handley 2000
253 1 Port Washington NE 1650 Long Island W M 48.89 – 50.80 49.64 Handley 2000
254 1 Port Washington NE 1650 Long Island W F 45.33 – 47.04 46.07 Handley 2000
255 1 Port Washington NE 1650 Long Island W M 48.83 – 50. 37 49.67 Handley 2000
256 1 Port Washington NE 1650 Long Island W F 38.76 – 39.54 39.08 Handley 2000
257 1 Port Washington NE 1650 Long Island W M 45.91 – 46.54 46.23 Handley 2000
258 1 Port Washington NE 1650 Long Island W F 40.43 – 43.26 41.72 Handley 2000 Whaleback Shell 261 1 Heap NE 1650 Maine W M 50.51 – 51.88 51.2 Handley 2000
119 1 Sorce, Visques (1) WI 1673 Puerto Rico W F 39.00 – 43.71 41.12 Narganes 1982
121 1 Sorce, Visques (1) WI 1673 Puerto Rico W F? 43.90 – 45.52 44.98 Narganes 1982 1700 (+/- 100), 1770 1 Morel 2727 F90-01-1 WI 1735 (+/-100) Guadeloupe 40.8 Grouard et al. 2013 1700 (+/- 100), 1770 1 Morel 2727 F90-01-2 WI 1735 (+/-100) Guadeloupe 40.7 Grouard et al. 2013 1700 (+/- 100), 1770 1 Morel 2727 F90-01-3 WI 1735 (+/-100) Guadeloupe 40 Grouard et al. 2013 1700 (+/- 100), 1770 1 Morel 2728 F91-11 WI 1735 (+/-100) Guadeloupe 42.7 Grouard et al. 2013 1700 (+/- 100), 1770 1 Morel 2729 F90-16 WI 1735 (+/-100) Guadeloupe 39.5 Grouard et al. 2013
140 Date Shoulder Specimen Group Site Region (BP) Date Note State/County Culture Sex Age Height Range SHA VRG Citation 1700 (+/- 100), 1770 1 Morel F281 WI 1735 (+/-100) Guadeloupe 37.5 Grouard et al. 2013 1700 (+/- Morel 5237 CLERC 100), 1770 1 851-4-2 WI 1735 (+/-100) Guadeloupe 44 Grouard et al. 2013 1700 (+/- Morel 6263 CLERC 100), 1770 1 1012 WI 1735 (+/-100) Guadeloupe 39.5 Grouard et al. 2013
117 1 Cayon (1) WI 1800 St. Kitts W U 42.60 – 56.43 45.1 Wing 1998
Burial 6 1 Apple Creek IL 1800 2000-1600 Greene CO, IL H F? A 37.1 Lawler et al. 2016
95 1 G.S. Lewis West (1) SE 1950 South Carolina W F? 41.09 – 44.56 43.25 Anderson (N.D.)
96 1 G.S. Lewis West (1) SE 1950 South Carolina W M 43.20 – 46.60 45.45 Anderson (N.D.) Grouard et al. (2013); 1 Seaview WI 2132 2015-2250 Barbuda 45.2 Perdikaris et al. 2008
83 1 15Jo9 SE 2150 Kentucky W F? 39.80 – 43.57 41.69 Webb 1942 Westmoreland-Barber Faulkner & Graham 1965, 3 1 (1) SE 2285 Tennessee W M 38.10 – 40.40 39.47 1966; Chapman 1988
248 1 Frenchman’s Bay:1 NE 2300 Maine W F 43.29 – 50.20 48 Handley 2000 Ensworth/Chase 89 1 Devon (1) SE 3370 Tennessee A F 35.30 – 38.30 36.68 Deter-Wolf 2004
132 1 Read site SE 3400 Kentucky A M 42.04 42.04 Webb 1950 Cedar Creek 105 1 Reservoir (1) SE 3450 Alabama A M 38.50 – 40.70 39.67 Futato 1983 Lewis & Kneberg 1947, 35 1 Cherry (1) SE 3700 Tennessee A F 34.60 – 37.57 36.13 1959 Lewis & Kneberg 1947, 36 1 Cherry (1) SE 3700 Tennessee A M 40.40 – 42.66 41.54 1959 Lewis & Kneberg 1947, 37 1 Cherry (1) SE 3700 Tennessee A M 39.70 – 41.72 40.47 1959 Lewis & Kneberg 1947, 38 1 Cherry (1) SE 3700 Tennessee A M 40.70 – 42.80 41.8 1959 Lewis & Kneberg 1947, 39 1 Cherry (1) SE 3700 Tennessee A U 41.53 – 39.10 40.47 1959 Lewis & Kneberg 1947, 40 1 Cherry (1) SE 3700 Tennessee A F? 36.79 – 39.40 38.36 1959 Lewis & Kneberg 1947, 41 1 Cherry (1) SE 3700 Tennessee A U 40.07 40.07 1959
193 1 Indian Knoll SE 4282 Kentucky A M 41.72 41.72 Haag 1948
194 1 Indian Knoll SE 4282 Kentucky A F? 31.52-34.95 33.24 Haag 1948
141 Date Shoulder Specimen Group Site Region (BP) Date Note State/County Culture Sex Age Height Range SHA VRG Citation
195 1 Indian Knoll SE 4282 Kentucky A F 35.37-41.41 38.39 Haag 1948
196 1 Indian Knoll SE 4282 Kentucky A M 37.95 37.95 Haag 1948
197 1 Indian Knoll SE 4282 Kentucky A F? 34.26-35.91 35.09 Haag 1948
198 1 Indian Knoll SE 4282 Kentucky A F? 40.78-41.77 41.24 Haag 1948
199 1 Indian Knoll SE 4282 Kentucky A F? 40.47 40.47 Haag 1948
200 1 Indian Knoll SE 4282 Kentucky A M 37.33-37.60 37.47 Haag 1948
201 1 Indian Knoll SE 4282 Kentucky A U 34.95-35.44 35.2 Haag 1948
202 1 Indian Knoll SE 4282 Kentucky A M? 38.57-38.58 38.58 Haag 1948
203 1 Indian Knoll SE 4282 Kentucky A M 42.60-42.98 42.75 Haag 1948
204 1 Indian Knoll SE 4282 Kentucky A F 36.07 36.07 Haag 1948
205 1 Indian Knoll SE 4282 Kentucky A M 38.90-40.93 39.92 Haag 1948
206 1 Indian Knoll SE 4282 Kentucky A M 41.72 41.72 Haag 1948
207 1 Indian Knoll SE 4282 Kentucky A M 41.25 41.25 Haag 1948
208 1 Indian Knoll SE 4282 Kentucky A F 38.58-39.68 39.13 Haag 1948
209 1 Indian Knoll SE 4282 Kentucky A F 38.90-39.26 39.08 Haag 1948
210 1 Indian Knoll SE 4282 Kentucky A U 38.01 38.01 Haag 1948
211 1 Indian Knoll SE 4282 Kentucky A M 47.06 – 47.88 47.47 Haag 1948
6 1 Danville Ferry (1) SE 4450 Tennessee A M 38.20 – 40.75 39.2 Chapman 1988
7 1 Danville Ferry (1) SE 4450 Tennessee A M 39.52 – 43.80 42.15 Chapman 1988
241 1 Nevin site:1 NE 4450 Maine A M 45.62 – 47.69 46.8 Handley 2000
249 1 Turner Farm NE 4450 Maine A M 43.75 – 45.09 44.3 Handley 2000
250 1 Turner Farm NE 4450 Maine A F 45.87 – 46.50 46.15 Handley 2000
260 1 Ruth Moore NE 4450 Maine A M 45.12 – 46.68 45.8 Handley 2000
42 1 Carlston Annis (1) SE 4578 Kentucky A F 37.13 – 38.90 37.89 Webb 1950
43 1 Carlston Annis (1) SE 4578 Kentucky A F 40.00 – 42.34 41.46 Webb 1950
142 Date Shoulder Specimen Group Site Region (BP) Date Note State/County Culture Sex Age Height Range SHA VRG Citation
44 1 Carlston Annis (1) SE 4578 Kentucky A M 39.60 – 41.40 40.78 Webb 1950
45 1 Carlston Annis (1) SE 4578 Kentucky A M 38.16 – 40.75 39.3 Webb 1950
46 1 Carlston Annis (1) SE 4578 Kentucky A M 39.80 – 42.66 41.48 Webb 1950
47 1 Carlston Annis (1) SE 4578 Kentucky A M 40.70 – 44.75 43.21 Webb 1950
48 1 Carlston Annis (1) SE 4578 Kentucky A M 36.90 – 41.77 40.18 Webb 1950
49 1 Carlston Annis (1) SE 4578 Kentucky A M 42.80 – 44.70 44.03 Webb 1950
50 1 Carlston Annis (1) SE 4578 Kentucky A F 35.10 – 36.30 35.62 Webb 1950
51 1 Carlston Annis (1) SE 4578 Kentucky A M 43.20 – 45.90 44.43 Webb 1950
155 1 Perry SE 4764 Alabama A M 37.95-38.43 38.19 Haag 1948
156 1 Perry SE 4764 Alabama A M 38.15-40.47 39.31 Haag 1948
157 1 Perry SE 4764 Alabama A F 42.88-43.14 43.01 Haag 1948
158 1 Perry SE 4764 Alabama A F 41.09 41.09 Haag 1948
159 1 Perry SE 4764 Alabama A U 39.52 39.52 Haag 1948
160 1 Perry SE 4764 Alabama A M 43.61-43.99 43.8 Haag 1948
161 1 Perry SE 4764 Alabama A F 35.76-36.48 36.12 Haag 1948
272 1 Perry SE 4764 Alabama A M 39.21 – 41.77 40.49 Haag 1948
64 1 Kirkland (1) SE 4945 Kentucky A M 40.60 – 42.97 41.8 Webb and Haag 1940
65 1 Kirkland (1) SE 4945 Kentucky A F 35.70 – 37.30 36.53 Webb and Haag 1940
22 1 Eva (1) SE 4950 Tennessee A M 37.13 – 38.06 37.8 Lewis & Lewis 1961
23 1 Eva (1) SE 4950 Tennessee A M 38.40 – 41.77 40.47 Lewis & Lewis 1961
24 1 Eva (1) SE 4950 Tennessee A M 47.66 – 49.30 48.49 Lewis & Lewis 1961
25 1 Eva (1) SE 4950 Tennessee A M? 36.79 – 38.52 37.56 Lewis & Lewis 1961
26 1 Eva (1) SE 4950 Tennessee A U 36.10 – 38.58 37.73 Lewis & Lewis 1961
27 1 Eva (1) SE 4950 Tennessee A M 37.60 – 45.66 43.9 Lewis & Lewis 1961
28 1 Eva (1) SE 4950 Tennessee A M? 34.00 – 39.78 37.2 Lewis & Lewis 1961
143 Date Shoulder Specimen Group Site Region (BP) Date Note State/County Culture Sex Age Height Range SHA VRG Citation
29 1 Eva (1) SE 4950 Tennessee A M 38.40 – 49.10 41.45 Lewis & Lewis 1961
30 1 Eva (1) SE 4950 Tennessee A U 34.90 – 38.90 36.79 Lewis & Lewis 1961
31 1 Eva (1) SE 4950 Tennessee A M 43.40 – 46.00 44.76 Lewis & Lewis 1961
32 1 Eva (1) SE 4950 Tennessee A M 38.50 – 42.66 40.17 Lewis & Lewis 1961
33 1 Eva (1) SE 4950 Tennessee A M 37.20 – 40.56 39.41 Lewis & Lewis 1961
69 1 Ward (1) SE 4960 Kentucky A F? 37.80 – 42.00 39.85 Webb and Haag 1940
70 1 Ward (1) SE 4960 Kentucky A M 41.80 – 45.05 43.75 Webb and Haag 1940
72 1 Ward (1) SE 4960 Kentucky A M 39.60 – 42.66 41.53 Webb and Haag 1940
73 1 Ward (1) SE 4960 Kentucky A U 35.70 – 38.90 37.98 Webb and Haag 1940
74 1 Ward (1) SE 4960 Kentucky A M 46.43 – 44.90 45.67 Webb and Haag 1940
75 1 Ward (1) SE 4960 Kentucky A F 38.03 – 39.88 38.86 Webb and Haag 1940
52 1 Chiggerville (1) SE 5000 Kentucky A M 39.20 – 42.97 41.3 Webb & Haag 1939
53 1 Chiggerville (1) SE 5000 Kentucky A M 36.10 – 43.57 41.13 Webb & Haag 1939
54 1 Chiggerville (1) SE 5000 Kentucky A M 39.40 – 42.04 40.8 Webb & Haag 1939
55 1 Chiggerville (1) SE 5000 Kentucky A M 38.27 – 39.54 38.63 Webb & Haag 1939
56 1 Chiggerville (1) SE 5000 Kentucky A M 34.60 – 36.76 35.66 Webb & Haag 1939
57 1 Chiggerville (1) SE 5000 Kentucky A M 41.30 – 43.61 42.93 Webb & Haag 1939
58 1 Chiggerville (1) SE 5000 Kentucky A M 37.30 – 39.19 37.99 Webb & Haag 1939
59 1 Chiggerville (1) SE 5000 Kentucky A F 41.60 – 44.25 42.94 Webb & Haag 1939
134 1 Flint River SE 5000 Alabama A M 48.44-48.94 48.69 Haag 1948
145 1 Whitesburg Bridge SE 5000 Alabama A M 45.18 45.18 Haag 1948
66 1 Barret (1) SE 5070 Kentucky A F 37.57 – 36.10 36.88 Webb and Haag 1947
67 1 Barret (1) SE 5070 Kentucky A U 35.40 – 37.48 36.93 Webb and Haag 1947
68 1 Barret (1) SE 5070 Kentucky A F? 38.40 – 41.38 40.04 Webb and Haag 1947 5111 (+/- 1 Hiscock NE 5111 150) New York U 46.17 46.17 Thomas 2003
144 Date Shoulder Specimen Group Site Region (BP) Date Note State/County Culture Sex Age Height Range SHA VRG Citation Lewis & Kneberg 1947, 8 1 Big Sandy (1) SE 5450 Tennessee A F? 32.60 – 39.21 36.46 1959 Lewis & Kneberg 1947, 9 1 Big Sandy (1) SE 5450 Tennessee A F 39.82 – 41.07 40.61 1959
60 1 Jackson Bluff (1) SE 5450 Kentucky A U 37.60 – 39.30 38.2 Stout & Baugh 1938
61 1 Jackson Bluff (1) SE 5450 Kentucky A F? 41.09 – 44.55 42.61 Stout & Baugh 1938
62 1 Jackson Bluff (1) SE 5450 Kentucky A M 43.31 – 45.60 44.2 Stout & Baugh 1938
63 1 Jackson Bluff (1) SE 5450 Kentucky A F? 39.78 – 42.90 41.37 Stout & Baugh 1938
76 1 Baker (1) SE 5450 Kentucky A U 40.70 – 43.10 42.07 Haag 1948
77 1 Baker (1) SE 5450 Kentucky A F? 36.60 – 40.65 38.86 Haag 1948
78 1 Baker (1) SE 5450 Kentucky A M 36.90 – 39.00 38.48 Haag 1948
79 1 Baker (1) SE 5450 Kentucky A M 42.70 – 44.82 43.79 Haag 1948
80 1 Baker (1) SE 5450 Kentucky A M 40.10 – 43.43 41.6 Haag 1948
86 1 Butterfield (1) SE 5450 Kentucky A M 44.74 - 46.05 45.4 Webb and Haag 1947
136 1 Little Bear Creek SE 5450 Alabama A M 39.21-43.99 41.6 Haag 1948
137 1 Little Bear Creek SE 5450 Alabama A U 42.35 42.35 Haag 1948
146 1 Bear Creek Cave SE 5450 Alabama A U 39.52 39.52 Haag 1948
147 1 Flint Shop SE 5450 Alabama A U 37.32 37.32 Haag 1948
149 1 Mason Island SE 5450 Alabama A M 42.04-43.43 42.74 Haag 1948 Randolph Co, 6090 1 Modoc Shelter IL 5525 5720-5330 Illinois A F? A 38.5 Lawler et al. 2016
162 1 Mulberry Creek SE 6950 Alabama A U 41.09 41.09 Haag 1948
163 1 Mulberry Creek SE 6950 Alabama A U 39.54 39.54 Haag 1948
164 1 Mulberry Creek SE 6950 Alabama A M 41.41-42.04 41.73 Haag 1948
166 1 Mulberry Creek SE 6950 Alabama A M 46.49 46.49 Haag 1948
167 1 Mulberry Creek SE 6950 Alabama A M 44.86-45.66 45.26 Haag 1948
185 1 Dust Cave SE 6950 Alabama A F 36.11-39.79 38.26 Walker et al. 2005
186 1 Dust Cave SE 6950 Alabama A F 37.15-38.12 37.8 Walker et al. 2005
145 Date Shoulder Specimen Group Site Region (BP) Date Note State/County Culture Sex Age Height Range SHA VRG Citation
187 1 Dust Cave SE 6950 Alabama A U 43.24-44.73 44.13 Walker et al. 2005
188 1 Dust Cave SE 6950 Alabama A F 44.89-45.87 45.38 Walker et al. 2005 Randolph Co, Burial 2 1 Modoc Shelter IL 8380 8560-8200 Illinois A M A 45.4 Lawler et al. 2016 10130- 2256 1 Koster IL 9755 9680 Greene CO, IL A F? A 44.7 Lawler et al. 2016 10130- 222 1 Koster IL 9755 9681 Greene CO, IL A F? A 38.8 Lawler et al. 2016 10130- 2407 1 Koster IL 9755 9682 Greene CO, IL A M A 45.4 Lawler et al. 2016 10190- Burial 1 Stilwell IL 9775 9630 Calhoun CO, IL A F? A 45 Lawler et al. 2016
12 1 Altatoma Dam SE W M 39.40 – 42.66 40.7 Weyanoke Old Town 1 (44PG51) SE Virginia 42 42 Blick 2005
1 CBT US5002 WI Guadeloupe 42.8 Grouard et al. (2013)
1 GMBT US1002/1003 WI Guadeloupe 44.1 Grouard et al. (2013)
1 GMBT US1008 WI Guadeloupe 42.6 Grouard et al. (2013)
1 Basse-Terre WI Guadeloupe 43.1 Grouard et al. (2013) Dominican Republic 1 No1 WI Dominican Republic 37.3 Grouard et al. (2013) Dominican Republic 1 No2 WI Dominican Republic 36 Grouard et al. (2013) Dominican Republic 1 No3 WI Dominican Republic 35.5 Grouard et al. (2013) New Mexico Governador Site 1 41174 WI Dominican Republic 34.7 Grouard et al. (2013)
1 Dominican Republic WI Dominican Republic 35.4 Grouard et al. (2013)
For key to coding see Worthington 2008.
Appendix B
Body Mass Estimates for Intermountain, Plains and Great Basin Dog Remains
Short Distance Long Distance Pannier Pack Body Mass Load (1.07) Load (0.45) load (0.3) Dog Reference Specimen AD/BC (Date # Site Date C14 Date Measurement Length kg lbs kg lbs kg lbs kg lbs Reference)
Veratic IMNH- Rockshelter
19613 (Level 14) VDDc8 101.95 24.31 53.60 26.02 57.35 10.94 24.12 7.29 16.08 This study
Veratic IMNH- Rockshelter 19636 (Level 14) 2932±34 VDDm7 86.41 28.32 62.43 30.30 66.81 12.74 28.10 8.50 18.73 This study
Intermountain West Intermountain
Veratic IMNH- Rockshelter 19637 (Level 14) VDDm7 86.62 28.53 62.90 30.53 67.30 12.84 28.30 8.56 18.87 This study
147
Short Distance Long Distance Pannier Pack Body Mass Load (1.07) Load (0.45) load (0.3) Dog Reference Specimen AD/BC (Date # Site Date C14 Date Measurement Length kg lbs kg lbs kg lbs kg lbs Reference)
Veratic IMNH- Rockshelter 18724 (Level 25) 3132±32 VDDm12 38.68 27.48 60.59 29.41 64.83 12.37 27.26 8.24 18.18 This study
Veratic IMNH- Rockshelter 18803 (Level 25) VDDm7 79.38 21.88 48.23 23.41 51.61 9.85 21.71 6.56 14.47 This study Yohe and Braden Pavesic Dog 1 Braden Site 6590±90 VDDm12 33.1 16.79 37.03 17.97 39.62 7.56 16.66 5.04 11.11 2000 Yohe and Braden Pavesic Dog 2 Braden Site Humerus BD 29.3 13.35 29.42 14.28 31.48 6.01 13.24 4.00 8.83 2000 Lawrence 1967 (Gowlett MCZ et al. 51769 Jaguar Cave 3220±80 VDDm19 67.75 14.34 31.61 15.34 33.82 6.45 14.22 4.30 9.48 1987) Lawrence 1967 (Gowlett MCZ et al. 51770 Jaguar Cave 940±80 VDDm19 63.2 11.01 24.28 11.78 25.98 4.96 10.93 3.30 7.28 1987)
148
Short Distance Long Distance Pannier Pack Body Mass Load (1.07) Load (0.45) load (0.3) Dog Reference Specimen AD/BC (Date # Site Date C14 Date Measurement Length kg lbs kg lbs kg lbs kg lbs Reference)
Fishing Bridge 500-1000 Dog 1 Campground AD VDDc2 185 20.78 45.81 22.24 49.02 9.35 20.62 6.23 13.74 Haag 1956
Fishing Bridge 500-1000 Dog 2 Campground AD VDDc2 153 11.38 25.10 12.18 26.85 5.12 11.29 3.42 7.53 Haag 1956 Schmitt Stillwater and Sharp 26Ch1049* Marsh VDDm8 66 12.98 28.62 13.89 30.62 5.84 12.88 3.89 8.59 1980 Schmitt and Sharp 1980 (Rhode, Adams,
Stillwater Elston 26Ch1048* Marsh 800±90 VDDm8 66 12.98 28.62 13.89 30.62 5.84 12.88 3.89 8.59 2000) Schmitt
Great Basin Great Pyramid and Sharp 26Wa275* Lake VDDm8 63 10.88 23.99 11.64 25.67 4.90 10.80 3.26 7.20 1980 Dansie and Specimen Schmitt 356 Vista Site 1320±320 VDDm8 63.6 11.28 24.87 12.07 26.61 5.08 11.19 3.38 7.46 1986 Dansie Specimen and 718-3 Vista Site 1320±320 Humerus BD 34.05 19.58 43.17 20.95 46.19 8.81 19.42 5.87 12.95 Schmitt
149
Short Distance Long Distance Pannier Pack Body Mass Load (1.07) Load (0.45) load (0.3) Dog Reference Specimen AD/BC (Date # Site Date C14 Date Measurement Length kg lbs kg lbs kg lbs kg lbs Reference) 1986 (Delacorte 1997) UU- 23684/DC- Grayson 87 Danger Cave 9500±500 VDDm8 61 9.63 21.23 10.30 22.71 4.33 9.55 2.89 6.37 1988
FS210-37 Hogup Cave 7815±350 VDDm7 85 20.79 45.84 22.25 49.05 9.36 20.63 6.24 13.75 Haag 1970 Haag 1966 Caldwell AD 1050- (Ambler Dog 1 Village 1200 VDDm9 72.8 18.83 41.52 20.15 44.43 8.47 18.68 5.65 12.46 1966) Haag 1966 Caldwell AD 1050- (Ambler Dog 2 Village 1200 VDDm9 67 13.74 30.30 14.71 32.42 6.19 13.64 4.12 9.09 1966) Pharo AD 350- Village 1500 VDDm9 75 21.09 46.49 22.56 49.74 9.49 20.92 6.33 13.95 Haag 1968
Larson AD 1675- Morey
Dog C (39WW2) 1780 VDDc13a 98 21.38 47.13 22.88 50.43 9.62 21.21 6.41 14.14 1986
Larson AD 1675- Morey Dog E (39WW2) 1780 VDDc13a 104 25.84 56.98 27.65 60.97 11.63 25.64 7.75 17.09 1986
Great Plains Great
Larson AD 1675- Morey Dog G (39WW2) 1780 VDDc13a 99 22.08 48.69 23.63 52.09 9.94 21.91 6.63 14.61 1986
150
Short Distance Long Distance Pannier Pack Body Mass Load (1.07) Load (0.45) load (0.3) Dog Reference Specimen AD/BC (Date # Site Date C14 Date Measurement Length kg lbs kg lbs kg lbs kg lbs Reference)
Lower Grand AD 1200- Morey Dog Q (39CO14) 1500 VDDc13a 99 22.08 48.69 23.63 52.09 9.94 21.91 6.63 14.61 1986
Potts AD 1550- Morey Dog U (39CO19) 1700 VDDc13a 100 22.80 50.27 24.40 53.79 10.26 22.62 6.84 15.08 1986
Potts AD 1550- Morey Dog V (39CO19) 1700 VDDc13a 102 24.29 53.55 25.99 57.30 10.93 24.10 7.29 16.07 1986
Pretty Head AD 1150- Morey Dog W (39LM232) 1500 VDDc13a 95 19.36 42.68 20.71 45.67 8.71 19.21 5.81 12.80 1986
White Buffalo Robe AD 1200- Morey Dog A' (32ME7) 1400 VDDc13a 107 28.30 62.39 30.28 66.76 12.74 28.08 8.49 18.72 1986
White Buffalo Robe AD 1200- Morey Dog B' (32ME7) 1400 VDDc13a 92 17.47 38.53 18.70 41.22 7.86 17.34 5.24 11.56 1986
151
Short Distance Long Distance Pannier Pack Body Mass Load (1.07) Load (0.45) load (0.3) Dog Reference Specimen AD/BC (Date # Site Date C14 Date Measurement Length kg lbs kg lbs kg lbs kg lbs Reference)
Big Hidatsa AD 1800- Morey Dog C' (32ME12) 1830 VDDc13a 105 26.65 58.74 28.51 62.86 11.99 26.44 7.99 17.62 1986
Big Hidatsa AD 1800- Morey Dog D' (32ME12) 1830 VDDc13a 116 36.62 80.74 39.19 86.39 16.48 36.33 10.99 24.22 1986
Big Hidatsa AD 1800- Morey Dog E' (32ME12) 1830 VDDc13a 96 20.02 44.13 21.42 47.22 9.01 19.86 6.01 13.24 1986
Barcal AD 1700- Bozell 61 (25BU4) 1750 VDDm19 26.8 23.51 51.83 25.15 55.46 10.58 23.32 7.05 15.55 1988
Barcal AD 1700- Bozell 65 (25BU4) 1750 VDDm19 26.6 22.97 50.64 24.58 54.19 10.34 22.79 6.89 15.19 1988
Barcal AD 1700- Bozell 66 (25BU4) 1750 VDDm19 28.4 28.12 61.98 30.08 66.32 12.65 27.89 8.43 18.59 1988
Barcal AD 1700- Bozell 67 (25BU4) 1750 VDDm19 28.2 27.51 60.65 29.43 64.89 12.38 27.29 8.25 18.19 1988
Barcal AD 1700- Bozell 69 (25BU4) 1750 VDDm19 25.4 19.92 43.92 21.32 46.99 8.96 19.76 5.98 13.18 1988
152
Short Distance Long Distance Pannier Pack Body Mass Load (1.07) Load (0.45) load (0.3) Dog Reference Specimen AD/BC (Date # Site Date C14 Date Measurement Length kg lbs kg lbs kg lbs kg lbs Reference)
Barcal AD 1700- Bozell 73 (25BU4) 1750 VDDm19 29.3 30.96 68.25 33.12 73.02 13.93 30.71 9.29 20.47 1988
Barcal AD 1700- Bozell 60 (25BU4) 1750 VDDm19 25.8 20.91 46.09 22.37 49.31 9.41 20.74 6.27 13.83 1988
Barcal AD 1700- Bozell 62 (25BU4) 1750 VDDm19 24.0 16.72 36.87 17.89 39.45 7.53 16.59 5.02 11.06 1988
Barcal AD 1700- Bozell 68 (25BU4) 1750 VDDm19 26.9 23.78 52.43 25.44 56.10 10.70 23.59 7.13 15.73 1988
Barcal AD 1700- Bozell 72 (25BU4) 1750 VDDm19 25.2 19.44 42.86 20.80 45.86 8.75 19.29 5.83 12.86 1988
Barcal AD 1700- Bozell 74 (25BU4) 1750 VDDm19 25.3 19.68 43.39 21.06 46.42 8.86 19.52 5.90 13.02 1988
Barcal AD 1700- Bozell 75 (25BU4) 1750 VDDm19 25.1 19.20 42.34 20.55 45.30 8.64 19.05 5.76 12.70 1988
Barcal AD 1700- Bozell 77 (25BU4) 1750 VDDm19 24.2 17.16 37.83 18.36 40.47 7.72 17.02 5.15 11.35 1988
Barcal AD 1700- Bozell 78 (25BU4) 1750 VDDm19 24.8 18.50 40.80 19.80 43.65 8.33 18.36 5.55 12.24 1988
153
Short Distance Long Distance Pannier Pack Body Mass Load (1.07) Load (0.45) load (0.3) Dog Reference Specimen AD/BC (Date # Site Date C14 Date Measurement Length kg lbs kg lbs kg lbs kg lbs Reference)
Barcal AD 1700- Bozell 79 (25BU4) 1750 VDDm19 25.3 19.68 43.39 21.06 46.42 8.86 19.52 5.90 13.02 1988
Barcal AD 1700- Bozell 82 (25BU4) 1750 VDDm19 25.5 20.16 44.45 21.58 47.57 9.07 20.00 6.05 13.34 1988
Barcal AD 1700- Bozell 59 (25BU4) 1750 VDDm19 23.2 15.06 33.21 16.12 35.53 6.78 14.94 4.52 9.96 1988
Barcal AD 1700- Bozell 63 (25BU4) 1750 VDDm19 22.8 14.28 31.47 15.27 33.67 6.42 14.16 4.28 9.44 1988
Barcal AD 1700- Bozell 64 (25BU4) 1750 VDDm19 22.2 13.15 28.99 14.07 31.01 5.92 13.04 3.94 8.70 1988
Barcal AD 1700- Bozell 70 (25BU4) 1750 VDDm19 23.2 15.06 33.21 16.12 35.53 6.78 14.94 4.52 9.96 1988
Barcal AD 1700- Bozell 71 (25BU4) 1750 VDDm19 22.8 14.28 31.47 15.27 33.67 6.42 14.16 4.28 9.44 1988
Barcal AD 1700- Bozell 76 (25BU4) 1750 VDDm19 19.3 8.54 18.82 9.13 20.13 3.84 8.47 2.56 5.65 1988
Barcal AD 1700- Bozell 80 (25BU4) 1750 VDDm19 21.6 12.08 26.64 12.93 28.50 5.44 11.99 3.62 7.99 1988
154
Short Distance Long Distance Pannier Pack Body Mass Load (1.07) Load (0.45) load (0.3) Dog Reference Specimen AD/BC (Date # Site Date C14 Date Measurement Length kg lbs kg lbs kg lbs kg lbs Reference)
Barcal AD 1700- Bozell 81 (25BU4) 1750 VDDm19 25.1 19.20 42.34 20.55 45.30 8.64 19.05 5.76 12.70 1988
Bellwood AD 1795- Bozell 97 (25BU2) 1800 VDDm19 22.8 14.28 31.47 15.27 33.67 6.42 14.16 4.28 9.44 1988
Bellwood AD 1795- Bozell 98 (25BU2) 1800 VDDm19 23.1 14.86 32.77 15.90 35.06 6.69 14.75 4.46 9.83 1988
Bellwood AD 1795- Bozell 99 (25BU2) 1800 VDDm19 22.0 12.79 28.19 13.68 30.16 5.75 12.68 3.84 8.46 1988
Bellwood AD 1795- Bozell 100 (25BU2) 1800 VDDm19 18.4 7.37 16.24 7.88 17.38 3.31 7.31 2.21 4.87 1988
Bellwood AD 1795- Bozell 101 (25BU2) 1800 VDDm19 24.0 16.72 36.87 17.89 39.45 7.53 16.59 5.02 11.06 1988
Burkett AD 1500- Bozell 28 (25NC1) 1650 VDDm19 27.8 26.32 58.03 28.16 62.09 11.84 26.11 7.90 17.41 1988
155
Short Distance Long Distance Pannier Pack Body Mass Load (1.07) Load (0.45) load (0.3) Dog Reference Specimen AD/BC (Date # Site Date C14 Date Measurement Length kg lbs kg lbs kg lbs kg lbs Reference)
Burkett AD 1500- Bozell 34 (25NC1) 1650 VDDm19 27.2 24.61 54.25 26.33 58.05 11.07 24.41 7.38 16.28 1988
Burkett AD 1500- Bozell 35 (25NC1) 1650 VDDm19 29.5 31.61 69.70 33.83 74.57 14.23 31.36 9.48 20.91 1988
Burkett AD 1500- Bozell 37 (25NC1) 1650 VDDm19 30.8 36.11 79.62 38.64 85.19 16.25 35.83 10.83 23.88 1988
Burkett AD 1500- Bozell 44 (25NC1) 1650 VDDm19 25.5 20.16 44.45 21.58 47.57 9.07 20.00 6.05 13.34 1988
Burkett AD 1500- Bozell 27 (25NC1) 1650 VDDm19 24.0 16.72 36.87 17.89 39.45 7.53 16.59 5.02 11.06 1988
Burkett AD 1500- Bozell 29 (25NC1) 1650 VDDm19 23.8 16.30 35.93 17.44 38.44 7.33 16.17 4.89 10.78 1988
Burkett AD 1500- Bozell 31 (25NC1) 1650 VDDm19 24.2 17.16 37.83 18.36 40.47 7.72 17.02 5.15 11.35 1988
Burkett AD 1500- Bozell 32 (25NC1) 1650 VDDm19 25.1 19.20 42.34 20.55 45.30 8.64 19.05 5.76 12.70 1988
Burkett AD 1500- Bozell 33 (25NC1) 1650 VDDm19 25.9 21.16 46.64 22.64 49.91 9.52 20.99 6.35 13.99 1988
156
Short Distance Long Distance Pannier Pack Body Mass Load (1.07) Load (0.45) load (0.3) Dog Reference Specimen AD/BC (Date # Site Date C14 Date Measurement Length kg lbs kg lbs kg lbs kg lbs Reference)
Burkett AD 1500- Bozell 39 (25NC1) 1650 VDDm19 26.5 22.71 50.06 24.30 53.56 10.22 22.53 6.81 15.02 1988
Burkett AD 1500- Bozell 42 (25NC1) 1650 VDDm19 25.5 20.16 44.45 21.58 47.57 9.07 20.00 6.05 13.34 1988
Burkett AD 1500- Bozell 43 (25NC1) 1650 VDDm19 25.7 20.66 45.54 22.10 48.73 9.30 20.49 6.20 13.66 1988
Burkett AD 1500- Bozell 46 (25NC1) 1650 VDDm19 23.2 15.06 33.21 16.12 35.53 6.78 14.94 4.52 9.96 1988
Burkett AD 1500- Bozell 30 (25NC1) 1650 VDDm19 23.4 15.47 34.10 16.55 36.49 6.96 15.34 4.64 10.23 1988
Burkett AD 1500- Bozell 36 (25NC1) 1650 VDDm19 24.6 18.05 39.79 19.31 42.57 8.12 17.90 5.41 11.94 1988
Burkett AD 1500- Bozell 38 (25NC1) 1650 VDDm19 22.2 13.15 28.99 14.07 31.01 5.92 13.04 3.94 8.70 1988
Burkett AD 1500- Bozell 40 (25NC1) 1650 VDDm19 24.6 18.05 39.79 19.31 42.57 8.12 17.90 5.41 11.94 1988
Burkett AD 1500- Bozell 41 (25NC1) 1650 VDDm19 19.4 8.67 19.12 9.28 20.46 3.90 8.60 2.60 5.74 1988
157
Short Distance Long Distance Pannier Pack Body Mass Load (1.07) Load (0.45) load (0.3) Dog Reference Specimen AD/BC (Date # Site Date C14 Date Measurement Length kg lbs kg lbs kg lbs kg lbs Reference)
Burkett AD 1500- Bozell 45 (25NC1) 1650 VDDm19 24.1 16.94 37.35 18.13 39.96 7.62 16.81 5.08 11.20 1988
Clarks AD 1820- Bozell 103 (25PK1) 1849 VDDm19 22.6 13.89 30.63 14.86 32.77 6.25 13.78 4.17 9.19 1988
Gray AD 1600- Bozell 8 (25CX1) 1670 VDDm19 30.4 34.69 76.47 37.11 81.82 15.61 34.41 10.41 22.94 1988
Gray AD 1600- Bozell 23 (25CX1) 1670 VDDm19 31.7 39.47 87.02 42.23 93.11 17.76 39.16 11.84 26.11 1988
Gray AD 1600- Bozell 1 (25CX1) 1670 VDDm19 29.0 29.99 66.11 32.09 70.74 13.49 29.75 9.00 19.83 1988
Gray AD 1600- Bozell 14 (25CX1) 1670 VDDm19 28.1 27.21 59.98 29.11 64.18 12.24 26.99 8.16 18.00 1988
Gray AD 1600- Bozell 16 (25CX1) 1670 VDDm19 30.4 34.69 76.47 37.11 81.82 15.61 34.41 10.41 22.94 1988
Gray AD 1600- Bozell 17 (25CX1) 1670 VDDm19 29.5 31.61 69.70 33.83 74.57 14.23 31.36 9.48 20.91 1988
Gray AD 1600- Bozell 18 (25CX1) 1670 VDDm19 27.9 26.62 58.68 28.48 62.78 11.98 26.40 7.98 17.60 1988
158
Short Distance Long Distance Pannier Pack Body Mass Load (1.07) Load (0.45) load (0.3) Dog Reference Specimen AD/BC (Date # Site Date C14 Date Measurement Length kg lbs kg lbs kg lbs kg lbs Reference)
Gray AD 1600- Bozell 20 (25CX1) 1670 VDDm19 26.3 22.18 48.90 23.73 52.32 9.98 22.01 6.65 14.67 1988
Gray AD 1600- Bozell 24 (25CX1) 1670 VDDm19 27.9 26.62 58.68 28.48 62.78 11.98 26.40 7.98 17.60 1988
Gray AD 1600- Bozell 25 (25CX1) 1670 VDDm19 28.8 29.36 64.72 31.41 69.25 13.21 29.12 8.81 19.42 1988
Gray AD 1600- Bozell 2 (25CX1) 1670 VDDm19 23.0 14.67 32.33 15.69 34.59 6.60 14.55 4.40 9.70 1988
Gray AD 1600- Bozell 3 (25CX1) 1670 VDDm19 27.5 25.46 56.12 27.24 60.05 11.45 25.25 7.64 16.84 1988
Gray AD 1600- Bozell 4 (25CX1) 1670 VDDm19 29.4 31.28 68.97 33.47 73.80 14.08 31.04 9.39 20.69 1988
Gray AD 1600- Bozell 6 (25CX1) 1670 VDDm19 24.7 18.28 40.29 19.55 43.11 8.22 18.13 5.48 12.09 1988
Gray AD 1600- Bozell 9 (25CX1) 1670 VDDm19 24.5 17.82 39.29 19.07 42.04 8.02 17.68 5.35 11.79 1988
Gray AD 1600- Bozell 10 (25CX1) 1670 VDDm19 25.8 20.91 46.09 22.37 49.31 9.41 20.74 6.27 13.83 1988
159
Short Distance Long Distance Pannier Pack Body Mass Load (1.07) Load (0.45) load (0.3) Dog Reference Specimen AD/BC (Date # Site Date C14 Date Measurement Length kg lbs kg lbs kg lbs kg lbs Reference)
Gray AD 1600- Bozell 11 (25CX1) 1670 VDDm19 25.5 20.16 44.45 21.58 47.57 9.07 20.00 6.05 13.34 1988
Gray AD 1600- Bozell 12 (25CX1) 1670 VDDm19 26.5 22.71 50.06 24.30 53.56 10.22 22.53 6.81 15.02 1988
Gray AD 1600- Bozell 21 (25CX1) 1670 VDDm19 26.6 22.97 50.64 24.58 54.19 10.34 22.79 6.89 15.19 1988
Gray AD 1600- Bozell 5 (25CX1) 1670 VDDm19 19.3 8.54 18.82 9.13 20.13 3.84 8.47 2.56 5.65 1988
Gray AD 1600- Bozell 7 (25CX1) 1670 VDDm19 23.9 16.51 36.40 17.67 38.95 7.43 16.38 4.95 10.92 1988
Gray AD 1600- Bozell 13 (25CX1) 1670 VDDm19 20.4 10.13 22.33 10.84 23.89 4.56 10.05 3.04 6.70 1988
Gray AD 1600- Bozell 15 (25CX1) 1670 VDDm19 22.7 14.08 31.05 15.07 33.22 6.34 13.97 4.22 9.31 1988
Gray AD 1600- Bozell 19 (25CX1) 1670 VDDm19 23.6 15.88 35.01 16.99 37.46 7.15 15.75 4.76 10.50 1988
Gray AD 1600- Bozell 22 (25CX1) 1670 VDDm19 24.2 17.16 37.83 18.36 40.47 7.72 17.02 5.15 11.35 1988
160
Short Distance Long Distance Pannier Pack Body Mass Load (1.07) Load (0.45) load (0.3) Dog Reference Specimen AD/BC (Date # Site Date C14 Date Measurement Length kg lbs kg lbs kg lbs kg lbs Reference)
Gray AD 1600- Bozell 26 (25CX1) 1670 VDDm19 26.0 21.41 47.20 22.91 50.50 9.63 21.24 6.42 14.16 1988
AD 1775- Bozell 89 Hill (25WT1) 1815 VDDm19 26.0 21.41 47.20 22.91 50.50 9.63 21.24 6.42 14.16 1988
AD 1775- Bozell 83 Hill (25WT1) 1815 VDDm19 19.4 8.67 19.12 9.28 20.46 3.90 8.60 2.60 5.74 1988
AD 1775- Bozell 84 Hill (25WT1) 1815 VDDm19 21.2 11.40 25.14 12.20 26.90 5.13 11.31 3.42 7.54 1988
AD 1775- Bozell 85 Hill (25WT1) 1815 VDDm19 21.6 12.08 26.64 12.93 28.50 5.44 11.99 3.62 7.99 1988
AD 1775- Bozell 86 Hill (25WT1) 1815 VDDm19 22.6 13.89 30.63 14.86 32.77 6.25 13.78 4.17 9.19 1988
AD 1775- Bozell 87 Hill (25WT1) 1815 VDDm19 16.6 5.36 11.82 5.74 12.65 2.41 5.32 1.61 3.55 1988
AD 1775- Bozell 88 Hill (25WT1) 1815 VDDm19 20.2 9.82 21.66 10.51 23.18 4.42 9.75 2.95 6.50 1988
AD 1775- Bozell 90 Hill (25WT1) 1815 VDDm19 18.5 7.49 16.51 8.01 17.67 3.37 7.43 2.25 4.95 1988
161
Short Distance Long Distance Pannier Pack Body Mass Load (1.07) Load (0.45) load (0.3) Dog Reference Specimen AD/BC (Date # Site Date C14 Date Measurement Length kg lbs kg lbs kg lbs kg lbs Reference)
Horse Creek AD 1809- Bozell 94 (25NC2) 1839 VDDm19 24.1 16.94 37.35 18.13 39.96 7.62 16.81 5.08 11.20 1988
Horse Creek AD 1809- Bozell 91 (25NC2) 1839 VDDm19 23.3 15.26 33.65 16.33 36.01 6.87 15.14 4.58 10.10 1988
Horse Creek AD 1809- Bozell 92 (25NC2) 1839 VDDm19 23.3 15.26 33.65 16.33 36.01 6.87 15.14 4.58 10.10 1988
Horse Creek AD 1809- Bozell 93 (25NC2) 1839 VDDm19 21.7 12.26 27.02 13.11 28.91 5.51 12.16 3.68 8.11 1988
Linwood AD 1770- Bozell 95 (25BU1) 1853 VDDm19 23.1 14.86 32.77 15.90 35.06 6.69 14.75 4.46 9.83 1988
Linwood AD 1770- Bozell 96 (25BU1) 1853 VDDm19 23.2 15.06 33.21 16.12 35.53 6.78 14.94 4.52 9.96 1988
Palmer AD 1770- Bozell 102 (25HW1) 1844 VDDm19 27.2 24.61 54.25 26.33 58.05 11.07 24.41 7.38 16.28 1988
162
Short Distance Long Distance Pannier Pack Body Mass Load (1.07) Load (0.45) load (0.3) Dog Reference Specimen AD/BC (Date # Site Date C14 Date Measurement Length kg lbs kg lbs kg lbs kg lbs Reference)
Wright AD 1650- Bozell 48 (25NC3) 1750 VDDm19 30.2 33.99 74.93 36.37 80.17 15.29 33.72 10.20 22.48 1988
Wright AD 1650- Bozell 50 (25NC3) 1750 VDDm19 30.5 35.04 77.25 37.49 82.65 15.77 34.76 10.51 23.17 1988
Wright AD 1650- Bozell 57 (25NC3) 1750 VDDm19 28.8 29.36 64.72 31.41 69.25 13.21 29.12 8.81 19.42 1988
Wright AD 1650- Bozell 47 (25NC3) 1750 VDDm19 25.6 20.41 44.99 21.84 48.14 9.18 20.25 6.12 13.50 1988
Wright AD 1650- Bozell 49 (25NC3) 1750 VDDm19 23.6 15.88 35.01 16.99 37.46 7.15 15.75 4.76 10.50 1988
Wright AD 1650- Bozell 51 (25NC3) 1750 VDDm19 26.2 21.92 48.33 23.46 51.71 9.86 21.75 6.58 14.50 1988
Wright AD 1650- Bozell 53 (25NC3) 1750 VDDm19 24.7 18.28 40.29 19.55 43.11 8.22 18.13 5.48 12.09 1988
Wright AD 1650- Bozell 54 (25NC3) 1750 VDDm19 25.0 18.97 41.82 20.30 44.75 8.54 18.82 5.69 12.55 1988
Wright AD 1650- Bozell 58 (25NC3) 1750 VDDm19 26.1 21.66 47.76 23.18 51.11 9.75 21.49 6.50 14.33 1988
163
Short Distance Long Distance Pannier Pack Body Mass Load (1.07) Load (0.45) load (0.3) Dog Reference Specimen AD/BC (Date # Site Date C14 Date Measurement Length kg lbs kg lbs kg lbs kg lbs Reference)
Wright AD 1650- Bozell 52 (25NC3) 1750 VDDm19 24.0 16.72 36.87 17.89 39.45 7.53 16.59 5.02 11.06 1988
Wright AD 1650- Bozell 55 (25NC3) 1750 VDDm19 17.6 6.42 14.16 6.87 15.15 2.89 6.37 1.93 4.25 1988
Wright AD 1650- Bozell 56 (25NC3) 1750 VDDm19 23.6 15.88 35.01 16.99 37.46 7.15 15.75 4.76 10.50 1988 *Specimen measurement estimated from available sources.
164
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Vita
Martin Hughes Welker
Education Aug. 2018 Ph.D. in Anthropology, The Pennsylvania State University, State College, PA. Dissertation Chair: S. B. McClure, Ph.D. Dec. 2015 M.A. in Anthropology, The Pennsylvania State University, State College, PA. Thesis Adviser: S. B. McClure, Ph.D. Dec. 2015 Certificate of Online Teaching, The Pennsylvania State University, State College, PA. May 2013 B.S. magna cum laude in Anthropology with Departmental Honors and a Museum Studies Certificate Utah State University, Logan, UT. Dean’s List: Jan. 2010-May 2013.
Grants and Fellowships 2016 M. H. Welker Frontier Adaptation and Dietary Patterning Among French and British Colonists in North America (1625-1790). Penn State Department of Anthropology Post-Comprehensive Hill Fellowship ($3000). 2016 M. H. Welker Identifying Environmental and Cultural Influences on Dietary Practices at the Historic Fortress of Louisbourg (1713-1768), Cape Breton, Canada. Penn State Department of Anthropology Pre-Comprehensive Hill Fellowship Pilot for Dissertation Research ($1325). 2015 E. Zavodny and M. H. Welker A Pretense of Defense? Hillfort Function in Bronze and Iron Age Lika, Croatia. The Explorer’s Club, Student Grant ($1000).
Peer Reviewed Publications (* Student author) Welker, M.H. and D. A. Byers (submitted) The Birch Creek Canids and Dogs as Transport Labor in the Intermountain West. American Antiquity. Welker, M. H., S. M. Billings*, J. A. Burns, and S. B. McClure (submitted) Roads and Military Provisioning During the French and Indian War (1754- 1763): The Faunal Remains of Fort Shirley, PA in Context. Open Quaternary. Zavodny, E., S. B. McClure, M. H. Welker, B. J. Culleton, J. Balen, and D. J. Kennett (submitted) Stable Isotope Markers of Herd Management and the Development of Transhumance in Bronze-Iron Age Lika. Journal of Archaeological Science: Reports McClure, S. B. and M. H. Welker (2017) Farming with Animals: The Cardial Neolithic and the Role of Domesticated Animals in the Western Mediterranean. In Times of Neolithic Transition along the Western Mediterranean. Springer Book. Lambert, P. M. and M. H. Welker (2017) Traumatic Injury Risk and Agricultural Transitions: A View from the American Southeast and Beyond. American Journal of Physical Anthropology 162(1): 120-142.