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 ..................................................................................