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An interesting question connected to the history of the domestic dog is the origin of the Australian dingo. The dingo is a wild canid, morphologically resembling South Asian domestic dogs, which according to the archaeological record arrived in Australia between 3500 and 12,000 years ago (Clutton-Brock, 1995). However, the precise ancestry and time of arrival in Australia of the dingo have not been known, nor whether, on its arrival, it was domesticated or half-domesticated before becoming feral, or a truly wild dog. In a study of 211 dingoes, sampled around Australia, 582 base pairs of the mtDNA CR were analysed and compared to dog and wolf data, giving very distinct results (Savolainen et al., 2004). All dingo sequences clustered in dog clade A, forming a single internal cluster of mtDNA types unique to dingoes, around a central type, A29, found in both dingoes and dogs (Fig. 2.4). Among domestic dogs, A29 was found only east of the Himalayas, together with most types in that part of clade A. This indicates that the dingo pop-ulation originates from a single introduction of a small poppop-ulation of domestic dogs coming from East Asia, carrying mtDNA type A29. The mean distance among the dingo sequences to A29 indicates their arrival to Australia ~5000 years ago. The dingo has probably remained isolated since then, and represents a unique isolate of early undifferentiated dogs.

Summary

Studies of physical and behavioural characteristics, and of molecular genetic markers, give consistent evidence that the domestic dog originates from the wolf, and the archaeological record indicates that the wolf was domesticated approxi-mately 15,000 years ago. Population genetic data, based on analysis of mtDNA, shows that all dogs originate from a common gene pool, and indicates that they originated at a single domestication event in East Asia, from which they spread to all continents, including Australia where the domestic dogs developed into the feral dingo. The mtDNA data give only two clear indications of crossbreeding between dog and wolf, indicating that crossbreeding between male dogs and female wolves has contributed only marginally to the dog gene-pool. Studies of

Y-chromosomal and autosomal haplotypes, which will no doubt be initiated within a few years, are necessary to test the validity of the conclusions obtained from the mtDNA data, and will, by giving the male side of the story, help to create an even more detailed picture of the evolution of the dog.

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A29

Fig. 2.4. Minimum-spanning network of the main dog clade, A (Savolainen et al., 2004). mtDNA types (circles) and hypothetical intermediates (solid dots) are separated by one mutational step (substitutions, indels not shown). White circles are mtDNA types found in dogs only; light grey circles are types found in dingoes only; dark grey circles are types found in both dingoes and dogs. Squares denote wolves. Areas of grey circles are proportional to frequencies among dingoes, but the area of A29 is reduced by 50%.

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Introduction

Modern dog breeds demonstrate a stunning level of morphological and behav-ioural diversity. The magnitude of morphological differences between dog breeds is greater than that between all wild species in Canidae, the dog family (Wayne, 1986a, b). This diversity led Charles Darwin and Konrad Lorenz to speculate that perhaps dogs derived from more than one wild ancestral species. However, molecular genetic data has shown conclusively that domestic dogs derive only from the grey wolf (Canis lupus) and not any other wild canid (Seal et al., 1970;

Sarich, 1977; Ferrell et al., 1978; Wurster-Hill and Centerwall, 1982; Wayne and O’Brien, 1987; Wayne et al., 1987a, b, 1997; see Chapter 2 in this volume). This brings us to the next, obvious question: where did all of the diversity we observe in dogs today come from?

Much has been learnt about the origin of dogs thanks to many years of archaeological research. This research is based on the study of all sorts of material that could indicate prehistoric associations between dogs and humans, from bones to art. During the last 10 years, the development of molecular genetics techniques has opened a new field of research, complementary to the archaeological investigation, which can contribute to a better understanding of the domestication of the dog. In this chapter we describe some molecular techniques and some studies that have employed molecular techniques to clarify some aspect of the process by which dogs were domesticated and modern dog breeds were formed. However, as we will see, these studies have not yet led to a complete understanding of the process and many questions remain to be answered.

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