Archaeological dogs from the Early Holocene Zhokhov site in the Eastern Siberian Arctic

https://doi.org/10.1016/j.jasrep.2017.04.003Get rights and content

Highlights

  • Early Holocene-aged faunal remains of domestic dogs are analyzed.

  • Two craniomorphology indices serve as reliable dog domestication criteria.

  • Sled dog breed existed in the Arctic as early as 9000 years ago.

  • Sled dogs could have been used in Siberia starting around 15,000 years ago.

Abstract

Excavations on the Zhokhov Island yielded a small but meaningful collection of Early Holocene canine faunal remains (MNI = 13). To clarify the morphological characteristics of the Zhokhov dogs' skulls, they were compared to the modern skulls of the Eastern Siberian wolves. This analysis shows, that around 9000 years ago a fully formed, as a species, domesticated dog was present at the Zhokhov site and used by the inhabitants of the area while hunting and as a draft animal. Presence of a developed form of dog sledding has been securely established for this culture. The Zhokhov site osteological dog remains analysis showed that the comparison of two proportional ratios (the ratio of skull height in the cranial region to total skull length and the ratio of snout height at Рm1 – Рm2 to total skull length) is a universal and reliable criterion for distinguishing wolf skulls from wolf-like dog skulls. This method prevents the influence of any single measurement on the identification of any individual as a dog or a wolf. We propose to name them indices DD1 and DD2, accordingly (for Dog Domestication). We also show that a sled dog and hunting dog standard existed, which significantly changes the current ideas about the timing of specialized dog breed formation via directed selection. Full completion of this process marks the third stage of domestication. Two earlier stages precede it in time and space (natural selection via the feeding behavior within specific ecological niche, formed by the human activity, followed by artificial selection based on the animal's ability to be tamed). We suggest that sled dogs could have been used in Siberia around 15,000 years ago.

Introduction

By virtue of a successful internal logic, the binary plant and animal nomenclature, developed by Carl Linnaeus in the 1730s, turned out to be quite compatible with Darwin's evolutionary theory, which emerged almost 150 years later, and continues to be viable to this day. Interspecies differences in Linnaeus' system are based on rigorously evaluated (counted and measured) differences in the organisms' morphology. The Canidae family contains several wild species (grey wolf, red wolf coyote, jackal), but only one domesticated species: domesticated dog, Canis familiaris. This last species, which includes all domestic dogs, proved to be the most successful: despite its relatively short evolutionary history it inhabited the entire planet and numbers in tens or even hundreds of millions individual dogs, while their wild relatives are only encountered in limited areas in much smaller numbers. Moreover, wolves, for example, currently need protection to prevent their extinction (Coppinger and Coppinger, 2001).

Domesticated dogs flourish with human support, demonstrating unparalleled plasticity, which allows creating a practically infinite number of breeds, most of which were formed in the last 200 years. Interestingly, after receiving the status of a breed, the grouping is strictly guarded from cross-breeding during following selective breeding. In that sense, each breed could be classified as a species, since it completely matches the classical species definition, advanced by Mayr (1942), except for the geographical segregation, since dog breeds are exterritorial.

As early as the 19th century, Rütimeyer (1861), and then Inostrantsev (1882) and Anuchin (1882) showed that domesticated dogs are present in the fossil record. Species-level identification of the remains was possible, among other reasons, because they belonged to animals with a well-expressed morphology, i.e. they were an established form, which existed in northern Europe 4000–5000 years ago. Dogs have long held a special place in human culture as a partner and an assistant (Serphel, 2002) and perhaps are the first domesticated animal. Obviously, their appearance was preceded by a somewhat lengthy period of species formation. Various opinions regarding domestication generally boil down to the idea of gradual transformation via initial taming of the wolf. However, Coppinger and Coppinger (2001) rightly notes, that it is populations, not individuals, which are necessary for speciation.

Thus, the questions of dog domestication timing is rather important and has seen a great deal of discussion over the last 10–15 years in the context of studying canine remains from the Upper Paleolithic sites in Eurasia, including those dating to the pre-LGM. In this regard, analyzing securely identified faunal remains of Early Holocene (Mesolithic) dogs is an interesting and critical objective, since it can provide the means and the method for reliable identification of the domestic dog form Canis familiaris.

Domestic dog, fully formed as a species, is ubiquitous in the archaeological record (e.g.: Clutton-Brock and Noe-Nygaard, 1990, Detry and Cardoso, 2010, Losey et al., 2011, Morey, 1992, Morey, 2010, Shye, 2013), mainly starting in Mid-Holocene sites, where the remains of these animals are numerous, including in Siberia (Losey et al., 2011, Morey, 2014, Pitulko and Kasparov, 1996). Secure finds, corresponding to the Pleistocene-Holocene boundary or older are extremely rare and usually come from the Mediterranean. In the European part of the continent, oldest securely dated (around 14,000 years ago) and morphologically recognizable dog remains were found at Bonn-Oberkassel (Morey, 2010), while in Siberia, they are only present in the dog burial at the Ushki-1 site (Kamchatka Peninsula), in Level VI. This horizon dates from 12,355 to 11,955 cal. years BP (Goebel et al., 2003), but species identification remains somewhat questionable since it was made by N.K. Vereschagin based on a photo of a cleaned set of poorly preserved buried animal remains (Dikov, 1977). More recently, four separate finds of fossil canids that come from different locations of North-East Siberia are reported by Germonpré et al. (2017), one of them is proto-historic dog, another skull clearly belongs to Pleistocene wolves, the third one is claimed as ‘Paleolithic dog’ while the last one remained questionable. This expands the reference group but does not bring us closer to the end of the debate.

It appears that Late Pleistocene canine remains from some European Upper Paleolithic sites (Les Grottes de Goyet, Przhedmost, Eliseyevichi, Mezin and others) could represent the early forms of domestic dogs, and the age of some finds could reach 30,000 years ago (Germonpré et al., 2009, Sablin and Khlopachev, 2002). We should note, that, although these authors do not overindulge in citing their predecessors, as early as a century ago I.T. Savenkov (1892) wrote about the possible presence of a domesticated wolf/dog form in the Late Paleolithic Siberia (based on the Afontova Gora faunal remains identifications made by I.D. Cherskiy). Later, V.I. Gromov (1932) worked with and wrote about the materials from newer excavations of the same site. The Afontova Gora faunal remains in question, according to N.D. Ovodov (1999), display significant similarity to dog remains, however their age most likely does not exceed 15,000 years (Astakhov, 1999), which corresponds to the age estimates of the earliest reliable dog remains (Morey, 2010).

Arguments for early appearance of a domestic dog form (see for example Germonpré et al., 2009, Sablin and Khlopachev, 2002), have numerous opponents (Boudadi-Maligne and Escarguel, 2014, Crockford and Kuzmin, 2012, Morey, 2014), who rightly point out a number of circumstances, exacerbating the shortage of data related to the fossil forms of ancient dogs. Among these, first of all, is the high polymorphism of the species, which makes many popular approaches problematic from the biometrics standpoint (Boudadi-Maligne and Escarguel, 2014). In addition, in some research (e.g. Germonpré et al., 2009, Germonpré et al., 2012), a certain amount of data manipulation takes place during the reference collection selection, and it was shown (Morey, 2014), that a different set of reference material would not allow identifying any “ancient” dogs, so classified by the authors, as reliable fossil examples of domestic dogs; in fact, they would fit well into the wolf sample. These results are not surprising, since grey wolves display significant interspecies polymorphism (interpopulation, intrapopulation and geographical variability, as well as a distinct sexual dimorphism, which can be hard to discover in a limited excavation sample).

In this regard, it is quite interesting, that, at least for Eastern Beringia, the existence of an extinct Pleistocene wolf form has been established; it was replaced by a modern grey wolf population after 15,000 years ago. From the craniofacial morphology standpoint, this was a specialized ecomorph with a relatively short snout and powerful M1, which occupied the ecological niche similar to that of the spotted hyena. The feeding structure of this carnivore assumed the presence of large herbivores, which were hunted but also available as carrion. Leonard et al. (2007) have shown that genetic lineages of East Beringian Pleistocene wolves do not continue in the contemporary genetic material, and this conclusion could be true for other areas as well.

Section snippets

Materials and methods

Fossil dog osteological material presented here was obtained during the Zhokhov site investigations in 1989, 1990, and 2000–2005 (Pitulko, 1998, Pitulko, 2011). These excavations opened up a total area of 571 m2. In addition to varied and numerous archaeological material (Pitulko, 1998), the work yielded a large faunal collection (N = 54,850, NISP = 21,994), dominated by northern reindeer and polar bear remains (Pitulko et al., 2015). Several occupations of the site span from 8250 to 7800 14С years

Reconstructing the use of dogs in Zhokhov site

The results we obtained suggest that around 8000 14С years ago the Zhokhov site was occupied, in addition to people, by a quite developed as a species domestic dog. In size, it was similar to the modern sled and hunting husky-like dogs of the Arctic Siberia.

To evaluate approximate weight of the individuals whose skulls were investigated here, we used the method of calculating the weight of dogs and some other Canidae representatives, based on the correlation established between the body mass and

Discussion

The questions about the domestic dog, its place in ancient human culture, domestication methods, the timing and location of this event are actively discussed both by biologists (primarily in the context of evolutionary theory) and by archaeologists. In fact, this animal was an important data source even for Charles Darwin. Not much progress towards the answers has been made since then; however, it is completely clear that the interval of species formation is insignificant on the geological

Conclusion

In this article we discussed the main scenarios of Canis familiaris evolution and domestic dog's use in human economies during the Early Holocene in the Siberia Arctic, as illuminated by the faunal material from the Zhokhov site. In our view, available data suggest that after splitting from the common ancestor with the grey wolf, the Canis familiaris evolution appears to have taken place over three phases and included the following three stages of natural and artificial selection: (1) natural

Acknowledgements

The material discussed here was obtained during the Zhokhov site investigations, initially as a part of the А-162 expedition of the Arctic and Antarctic Research Institute (St. Petersburg, Russia) in 1989 and 1990, and then during the research project “Zhokhov-2000” in 2000–2005. These field investigations were supported by the private Rock Foundation research fund (New York). The present study was completed with the support of the Russian Science Foundation (project N 16-18-10265 RNF granted

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