Assessing current visual tooth wear age estimation methods for Rangifer tarandus using a known age sample from Canada.

Age estimation is crucial for investigating animal populations in the past and present. Visual examination of tooth wear and eruption is one of the most common ageing methods in zooarchaeology, wildlife management, palaeontology, and veterinary research. Such approaches are particularly advantageous because they are non-destructive, can be completed using photographs, and do not require specialized training. Several tooth wear and eruption methods have been developed for Rangifer tarandus, a widely distributed and long-utilized species in the North. This paper evaluates the practicality and effectiveness of three existing visual tooth wear and eruption methods for this species using a large known-age sample from several caribou populations in northern Canada (Bluenose East, Bluenose West, Dolphin-Union, Qamanirjuaq, and Beverly herds). These methods are evaluated based on: (1) the amount of error and bias between estimated and actual ages, (2) suitable and interpretable results, (3) user-friendly and unambiguous procedures, and (4) which teeth and visual features of those teeth are used to record wear and eruption status. This study finds that the three evaluated methods all have variable errors and biases, and two show extensive biases when applied to older individuals. Demographic data is simpler to generate and more flexible to report when methods allow age to be estimated as a continuous or discrete variable, rather than as age ranges. The dentition samples used by two of the previously developed methods impact their applicability to other populations of Rangifer. In one existing method, individuals were unavailable from some age ranges leaving gaps when assigning ages. For another Rangifer-ageing method, the population utilized was too distinct in morphology or diet to be used with the Canadian caribou analyzed here. Additional refinement of tooth wear and eruption ageing methods will benefit zooarchaeological research on reindeer and caribou remains.

The evolution of dog diet and foraging: Insights from archaeological canids in Siberia.

Research on the evolution of dog foraging and diet has largely focused on scavenging during their initial domestication and genetic adaptations to starch-rich food environments following the advent of agriculture. The Siberian archaeological record evidences other critical shifts in dog foraging and diet that likely characterize Holocene dogs globally. By the Middle Holocene, body size reconstruction for Siberia dogs indicates that most were far smaller than Pleistocene wolves. This contributed to dogs' tendencies to scavenge, feed on small prey, and reduce social foraging. Stable carbon and nitrogen isotope analysis of Siberian dogs reveals that their diets were more diverse than those of Pleistocene wolves. This included habitual consumption of marine and freshwater foods by the Middle Holocene and reliance on C4 foods by the Late Holocene. Feeding on such foods and anthropogenic waste increased dogs' exposure to microbes, affected their gut microbiomes, and shaped long-term dog population history.

Modern Siberian dog ancestry was shaped by several thousand years of Eurasian-wide trade and human dispersal.

Dogs have been essential to life in the Siberian Arctic for over 9,500 y, and this tight link between people and dogs continues in Siberian communities. Although Arctic Siberian groups such as the Nenets received limited gene flow from neighboring groups, archaeological evidence suggests that metallurgy and new subsistence strategies emerged in Northwest Siberia around 2,000 y ago. It is unclear if the Siberian Arctic dog population was as continuous as the people of the region or if instead admixture occurred, possibly in relation to the influx of material culture from other parts of Eurasia. To address this question, we sequenced and analyzed the genomes of 20 ancient and historical Siberian and Eurasian Steppe dogs. Our analyses indicate that while Siberian dogs were genetically homogenous between 9,500 to 7,000 y ago, later introduction of dogs from the Eurasian Steppe and Europe led to substantial admixture. This is clearly the case in the Iamal-Nenets region (Northwestern Siberia) where dogs from the Iron Age period (∼2,000 y ago) possess substantially less ancestry related to European and Steppe dogs than dogs from the medieval period (∼1,000 y ago). Combined with findings of nonlocal materials recovered from these archaeological sites, including glass beads and metal items, these results indicate that Northwest Siberian communities were connected to a larger trade network through which they acquired genetically distinctive dogs from other regions. These exchanges were part of a series of major societal changes, including the rise of large-scale reindeer pastoralism ∼800 y ago.

Origins and genetic legacy of prehistoric dogs.

Dogs were the first domestic animal, but little is known about their population history and to what extent it was linked to humans. We sequenced 27 ancient dog genomes and found that all dogs share a common ancestry distinct from present-day wolves, with limited gene flow from wolves since domestication but substantial dog-to-wolf gene flow. By 11,000 years ago, at least five major ancestry lineages had diversified, demonstrating a deep genetic history of dogs during the Paleolithic. Coanalysis with human genomes reveals aspects of dog population history that mirror humans, including Levant-related ancestry in Africa and early agricultural Europe. Other aspects differ, including the impacts of steppe pastoralist expansions in West and East Eurasia and a near-complete turnover of Neolithic European dog ancestry.

Temporal and structural genetic variation in reindeer (Rangifer tarandus) associated with the pastoral transition in Northwestern Siberia.

Just as the domestication of livestock is often cited as a key element in the Neolithic transition to settled, the emergence of large-scaled reindeer husbandry was a fundamental social transformation for the indigenous peoples of Arctic Eurasia. To better understand the history of reindeer domestication, and the genetic processes associated with the pastoral transition in the Eurasian Arctic, we analyzed archaeological and contemporary reindeer samples from Northwestern Siberia. The material represents Rangifer genealogies spanning from 15,000 years ago to the 18th century, as well as modern samples from the wild Taimyr population and from domestic herds managed by Nenetses. The wild and the domestic population are the largest populations of their kind in Northern Eurasia, and some Nenetses hold their domestic reindeer beside their wild cousins. Our analyses of 197 modern and 223 ancient mitochondrial DNA sequences revealed two genetic clusters, which are interpreted as representing the gene pools of contemporary domestic and past wild reindeer. Among a total of 137 different mitochondrial haplotypes identified in both the modern and archaeological samples, only 21 were detected in the modern domestic gene pool, while 11 of these were absent from the wild gene pool. The significant temporal genetic shift that we associate with the pastoral transition suggests that the emergence and spread of reindeer pastoralism in Northwestern Siberia originated with the translocation and subsequent selective breeding of a special type of animal from outside the region. The distinct and persistent domestic characteristics of the haplotype structure since the 18th century suggests little genetic exchange since then. The absence of the typical domestic clade in modern nearby wild populations suggests that the contemporary Nenets domestic breed feature an ancestry from outside its present main distribution, possibly from further South.

Storing fish?: a dog's isotopic biography provides insight into Iron Age food preservation strategies in the Russian Arctic.

Analysis of individual animal bodies can provide numerous useful insights in archeology, including how humans provisioned such animals, which in turn informs on a variety of other past behaviors such as human dietary patterns. In this study, we conducted stable carbon (δ 13C) and nitrogen (δ 15N) isotope analysis of collagen and keratin from four types of tissues from a dog burial at the Ust'-Polui site in the Iamal region of Arctic Russia. Ust'-Polui is an Iron Age site located on the Lower Ob River, a major northern fishery characterized by extreme seasonal shifts in fish presence. During a 6-month period stretching over the coldest months of the year, fish are nearly entirely absent in the Lower Ob River. Despite this, the stable isotope compositions of the dog's bone and dentine collagen and hair and nail keratin all indicate a monotonous diet focusing on local fish. This pattern indicates the dog was provisioned year-round with fish. This was likely accomplished by mass harvesting of fish using nets or traps. Such fish were then processed and frozen for consumption during the non-fishing season. These findings suggest that people in the Ust'-Polui region also relied to some extent on fish throughout the year. Stored fish likely provided a dietary buffer for uneven returns from reindeer and bird hunting, both of which also are well-evidenced at the site.

Specialized sledge dogs accompanied Inuit dispersal across the North American Arctic.

Domestic dogs have been central to life in the North American Arctic for millennia. The ancestors of the Inuit were the first to introduce the widespread usage of dog sledge transportation technology to the Americas, but whether the Inuit adopted local Palaeo-Inuit dogs or introduced a new dog population to the region remains unknown. To test these hypotheses, we generated mitochondrial DNA and geometric morphometric data of skull and dental elements from a total of 922 North American Arctic dogs and wolves spanning over 4500 years. Our analyses revealed that dogs from Inuit sites dating from 2000 BP possess morphological and genetic signatures that distinguish them from earlier Palaeo-Inuit dogs, and identified a novel mitochondrial clade in eastern Siberia and Alaska. The genetic legacy of these Inuit dogs survives today in modern Arctic sledge dogs despite phenotypic differences between archaeological and modern Arctic dogs. Together, our data reveal that Inuit dogs derive from a secondary pre-contact migration of dogs distinct from Palaeo-Inuit dogs, and probably aided the Inuit expansion across the North American Arctic beginning around 1000 BP.

Spondylosis deformans as an indicator of transport activities in archaeological dogs: A systematic evaluation of current methods for assessing archaeological specimens.

Over the past several decades archaeologists have used the spinal pathology spondylosis deformans as an indicator that archaeological dogs were used to pull or carry loads. This interpretive approach is largely based upon observations of prehistoric dog remains and archaeologist's interpretations of veterinary literature on recent sled dogs and other draft animals. However, no comparative large-scale studies of the occurrence of spondylosis deformans in wild canids, transport dogs, and dogs never involved in pulling or carrying loads have been published. To evaluate the reliability of spondylosis deformans in archaeological dogs as an indicator of participation in transport activities, 136 modern non-transport dogs, 19 sled dogs, and 241 wolves were systematically analyzed for the occurrence of spondylosis deformans. Our results indicate this pathology is not a reliable skeletal indicator of dog transport because the disease is prevalent in both dogs and wolves, regardless of their occupational histories. Numerous factors correlate with the occurrence and manifestation of this disease in canids, including age, body size, sex, and inbreeding. As such, it remains extremely challenging to identify specific etiologies for spondylosis deformans in archaeological specimens.

Three-Dimensional Geometric Morphometric Analysis of Fossil Canid Mandibles and Skulls.

Much of the fossil record for dogs consists of mandibles. However, can fossil canid mandibles be reliably identified as dogs or wolves? 3D geometric morphometric analysis correctly classifies 99.5% of the modern dog and wolf mandibles. However, only 4 of 26 Ust'-Polui fossil mandibles, a Russian Arctic site occupied from 250BCE to 150CE, were identified as dogs and none of the 20 Ivolgin mandibles, an Iron Age site in southern Russia, were identified as dogs. Three of the Ust'-Polui mandibles and 8 of the Ivolgin mandibles were identified as wolves. In contrast, all 12 Ivolgin skulls and 5 Ust'-Polui skulls were clearly identified as dogs. Only the classification of the UP6571 skull as a dog (Dog Posterior Probability = 1.0) was not supported by the typical probability. Other evidence indicates these canids were domesticated: they were located within human dwellings, remains at both sites have butchery marks indicating that they were consumed, and isotope analysis of canid and human remains from Ust'-Polui demonstrate that both were consuming freshwater protein; indicating that the humans were feeding the canids. Our results demonstrate that the mandible may not evolve as rapidly as the cranium and the mandible is not reliable for identifying early dog fossils.

A Second Mortuary Hiatus on Lake Baikal in Siberia and the Arrival of Small-Scale Pastoralism.

The spread of pastoralism in Asia is poorly understood, including how such processes affected northern forager populations. Lake Baikal's western shore has a rich Holocene archaeological record that tracks these processes. The Early Bronze Age here is evidenced by numerous forager burials. The Early Iron Age (EIA) is thought to mark the arrival of pastoralists, but archaeological remains from this period have received little analysis. New radiocarbon dates for EIA human remains from 23 cemeteries indicate that no burials were created along this shore for ~900 years. This period, from ~3670 to 2760 cal. BP, spans from the end of the Early Bronze Age to the advent of the EIA. The burial gap may mark disruption of local foraging populations through incursions by non-local pastoralists. Radiocarbon dates on faunal remains indicate that domestic herd animals first appear around 3275 cal. BP, just prior to the first EIA human burials. Stable carbon and nitrogen isotope analysis of human remains and zooarchaeological data indicate that domestic fauna were minor dietary components for EIA people. Like preceding foragers, the EIA groups relied extensively on Baikal's aquatic food sources, indicating that the scale of pastoralism during this period was relatively limited.

Spondylosis deformans in three large canids from the Gravettian Predmostí site: Comparison with other canid populations.

Spondylosis deformans is a common degenerative condition of the spinal column, especially in modern domestic dogs. The presence and severity of lesions are related to age and physical activity, but they can be influenced by genetics, with some modern breeds being particularly predisposed. Spondylosis deformans also has been reported in prehistoric dogs. Here, we describe three affected vertebrae, likely from three individuals, in a large canid assemblage from the Gravettian Predmostí site, Czech Republic. We compare the proportion of affected individuals from the Predmostí assemblage with that of affected individuals among recent wild Northern wolves, inbred wolves, captive wolves, and recent Northern dogs. The proportion of affected individuals among the captive wolves differs significantly from the other wolf groups. The proportion in the Predmostí assemblage does not differ significantly from that of the wild wolf groups.

A 9,000 Year History of Seal Hunting on Lake Baikal, Siberia: The Zooarchaeology of Sagan-Zaba II.

Sagan-Zaba II, a habitation site on the shore of Siberia's Lake Baikal, contains a record of seal hunting that spans much of the Holocene, making it one of the longest histories of seal use in North Asia. Zooarchaeological analyses of the 16,000 Baikal seal remains from this well-dated site clearly show that sealing began here at least 9000 calendar years ago. The use of these animals at Sagan-Zaba appears to have peaked in the Middle Holocene, when foragers used the site as a spring hunting and processing location for yearling and juvenile seals taken on the lake ice. After 4800 years ago, seal use declined at the site, while the relative importance of ungulate hunting and fishing increased. Pastoralists began occupying Sagan-Zaba at some point during the Late Holocene, and these groups too utilized the lake's seals. Domesticated animals are increasingly common after about 2000 years ago, a pattern seen elsewhere in the region, but spring and some summer hunting of seals was still occurring. This use of seals by prehistoric herders mirrors patterns of seal use among the region's historic and modern groups. Overall, the data presented in the paper demonstrate that Lake Baikal witnessed thousands of years of human use of aquatic resources.

Craniomandibular trauma and tooth loss in northern dogs and wolves: implications for the archaeological study of dog husbandry and domestication.

Archaeological dog remains from many areas clearly show that these animals suffered tooth fractures, tooth loss, trauma, and dental defects during their lives. Relatively little research has explored the meanings of these patterns, particularly for ancient dog remains from small-scale societies of the North. One limiting issue is the lack of comparative data on dental health and experiences of trauma among northern wolves and dogs. This paper examines tooth loss, tooth fracture, enamel hypoplasia, and cranial trauma in a large sample of historic dog and wolf remains from North America and Northern Russia. The data indicate that the dogs more commonly experienced tooth loss and tooth fracture than the wolves, despite reportedly being fed mostly soft foods such as blubber and fish. The higher rates observed in the dogs likely is a result of food stress and self-provisioning through scavenging. The ability to self-provision was likely important for the long-term history of dog use in the north. Dogs also more commonly experienced cranial fractures than wolves, particularly depression fractures on their frontal bones, which were likely the result of blows from humans. Hypoplastic lesions are rare in both wolves and dogs, and probably result from multiple causes, including food stress, disease, and trauma.

Burying dogs in ancient Cis-Baikal, Siberia: temporal trends and relationships with human diet and subsistence practices.

The first objective of this study is to examine temporal patterns in ancient dog burials in the Lake Baikal region of Eastern Siberia. The second objective is to determine if the practice of dog burial here can be correlated with patterns in human subsistence practices, in particular a reliance on terrestrial mammals. Direct radiocarbon dating of a suite of the region's dog remains indicates that these animals were given burial only during periods in which human burials were common. Dog burials of any kind were most common during the Early Neolithic (∼7-8000 B.P.), and rare during all other time periods. Further, only foraging groups seem to have buried canids in this region, as pastoralist habitation sites and cemeteries generally lack dog interments, with the exception of sacrificed animals. Stable carbon and nitrogen isotope data indicate that dogs were only buried where and when human diets were relatively rich in aquatic foods, which here most likely included river and lake fish and Baikal seal (Phoca sibirica). Generally, human and dog diets appear to have been similar across the study subregions, and this is important for interpreting their radiocarbon dates, and comparing them to those obtained on the region's human remains, both of which likely carry a freshwater old carbon bias. Slight offsets were observed in the isotope values of dogs and humans in our samples, particularly where both have diets rich in aquatic fauna. This may result from dietary differences between people and their dogs, perhaps due to consuming fish of different sizes, or even different tissues from the same aquatic fauna. This paper also provides a first glimpse of the DNA of ancient canids in Northeast Asia.