Reindeer Domestication Research Articles

Whole-genome sequencing provides novel insights into the evolutionary history and genetic adaptation of reindeer populations in northern Eurasia

Domestic reindeer (Rangifer tarandus) play a vital role in the culture and livelihoods of indigenous people across northern Eurasia. These animals are well adapted to harsh environmental conditions, such as extreme cold, limited feed availability and long migration distances. Therefore, understanding the genomics of reindeer is crucial for improving their management, conservation and utilisation. In this study, we have generated a new genome assembly for the Fennoscandian domestic reindeer with high contiguity, making it the most complete reference genome for reindeer to date. The new genome assembly was utilised to explore genetic diversity, population structure and selective sweeps in Eurasian Rangifer tarandus populations which was based on the largest population genomic dataset for reindeer, encompassing 58 individuals from diverse populations. Phylogenetic analyses revealed distinct genetic clusters, with the Finnish wild forest reindeer (Rangifer tarandus fennicus) standing out as a unique subspecies. Divergence time estimates suggested a separation of ~ 52 thousand years ago (Kya) between the northern European Rangifer tarandus fennicus and Rangifer tarandus tarandus. Our study identified four main genetic clusters: Fennoscandian, the eastern/northern Russian and Alaskan group, the Finnish forest reindeer, and the Svalbard reindeer. Furthermore, two independent reindeer domestication processes were inferred, suggesting separate origins for the domestic Fennoscandian and eastern/northern Russian reindeer. Notably, shared genes under selection, including retroviral genes, point towards molecular domestication processes that aided adaptation of this species to diverse environments.

Investigating the domestication and early management of reindeer (Rangifer tarandus) in the Sámi archaeological context from teeth geometric morphometrics

For centuries, reindeer herding has been an integral part of the subsistence, lifeways, economy and cosmology of the Sámi of northern Fennoscandia. Despite its importance, the timing and details of early reindeer domestication are still highly debated. Identifying domesticated individuals in the archaeological record remains complicated due to the presence of two interbreeding subspecies in Fennoscandia and a mixed socio-economic organisation by Sámi populations, which was mainly a combination of wild reindeer hunting and small-scale reindeer herding. This study proposes methodological improvement for identifying domestic individuals using 2D landmark and sliding semi-landmark based geometric morphometrics on the isolated lower molars of 389 modern specimens, and 90 teeth from four archaeological sites in Finnish Lapland. Our results indicate that despite the significant impact of wear on overall tooth morphology, our protocol is very useful for identifying subspecies (classification accuracy of the two species is between 78 and 91% depending on the wear class) and understanding the morphological changes induced by the domestication process. We suggest that the morphological variation observable among modern populations has been impacted by recent changes in herding strategies in northern Fennoscandia, and that the archaeological domesticated reindeer populations were relatively different, probably due to selection by the Sámi. This study also highlights the importance of using other direct evidence or contextual archaeological data to better trace the early evidence of a domesticated reindeer economy in northern Fennoscandia, and aid in reconstructing the socio-economic changes in Sámi populations over time.

The Archaeology of Reindeer Domestication and Herding Practices in Northern Fennoscandia

Animal domestication is a profound change for human societies, economies, and worldviews. The shifting definitions of animal domestication reflect its varying and process-like nature. Reindeer is one of the species whose domestication is not easily pinned down using standard definitions and research methodologies of animal domestication. In recent years, advances in archaeological methodology and the conceptual understanding of animal domestication have opened new avenues for research on this topic. This review summarizes recent research on the archaeology of reindeer domestication among the Indigenous Sámi of northern Fennoscandia. It compiles a chronological framework of reindeer domestication with an emphasis on the development of reindeer-herding practices and human–reindeer relationships. I argue that while a major transition to reindeer herding occurred among the Sámi from the 15th century onward, small-scale reindeer herding characterized by interspecies sociality, cooperation, and care developed earlier during the Late Iron Age, with regional variations in the timing and details of the events. By focusing on reindeer-herding practices and the human–reindeer relationships embedded in them, I also argue that reindeer domestication, and animal domestication in general, is a relationship constructed and constantly renegotiated in everyday interactions with the animals.

The osteometric identification of castrated reindeer (Rangifer tarandus) and the significance of castration in tracing human-animal relationships in the North

Reindeer are the only domestic cervid and have formed the cosmologies and practical daily lives of numerous peoples in the Northern Hemisphere for thousands of years. The questions of when, how, and where reindeer domestication originated and how it developed remain one of the scientific enigmas of our time. The practice of reindeer castration is an essential feature of all communities practicing reindeer herding today. It has probably been one of the most important interventions in the reindeer’s life cycle and biology that marked the start of domesticating human-reindeer relationships long ago. Castration is and has been essential for reindeer taming, control, training, herd management, and ritual practices. Unsuitably, to this present day, there are no methods zooarchaeologists can employ to distinguish a reindeer gelding from a reindeer bull in the archaeological record. In this current paper, we outline a new method that presents the possibility of differentiating between full males, castrated males, and females based on osteometric features. We measured the leg bones and pelvis of the complete or partial skeletons of 97 adult modern domestic reindeer individuals to determine the precise effects castration has on skeletal size and morphology. We explored our osteometric dataset with different statistical methods. We found a clear separation of the two male groups in the radioulna, humerus, and femur but in the tibia and metapodials to a lesser extent. Osteometric depth and width were generally more affected than the longitudinal axis. Females were easily distinguishable from castrates and full males based on nearly every bone measurement. Our analysis shows that reindeer castration can be proven through osteometric analysis.

Cross-sectional properties of reindeer long bones and metapodials allow identification of activity patterns

Habitual loading patterns of domesticated animals may differ due to human influence from their wild counterparts. In the early stages of human-reindeer interaction, cargo and draft use was likely important, as well as corralling tame reindeer. This may result to changes in loading as increased (working) or decreased (captive) loading, as well as foraging patterns (digging for lichen from under the snow versus fed working and/or captive reindeer). Our aim is to study whether differences in activity modify variation in bone cross-sectional properties and external dimensions. Our material consists of donated skeletons of modern reindeer: 20 working reindeer (19 racing and one draft), 24 zoo reindeer, and sample of 78 free-ranging/wild reindeer as a reference group. We used general linear modelling to first establish the total variation in cross-sectional properties among wild and free-ranging reindeer, and then to infer how differences in loading modify observed variation among zoo and working reindeer. According to our results, direction of greater bone quantity as well as external dimensions in of radioulna of female reindeer differs from female reference group, likely relating to foraging behavior. External dimensions of humerus differ in working and zoo male reindeer compared to male reference group. Increased robusticity of long bones, especially of tibia among working male reindeer, may indicate increased loading, and increased cortical area of long bones may indicate sedentary lifestyle among female reindeer. The results of this study can be used to understand early stages of reindeer domestication by observing reindeer activity patterns from archaeological material.

Impact of selection and domestication on hindlimb bones of modern reindeer populations: Archaeological implications for early reindeer management by Sámi in Fennoscandia

ABSTRACT For centuries, reindeer herding has been an integral part of the subsistence and culture among the Sámi of northern Fennoscandia. Despite the importance of this husbandry in their history, the timing and details of early reindeer domestication are still highly debated. Indeed, identifying domesticated individuals in the archaeological record remains complicated because reindeer are still considered to be in the early phases of the domestication process. In this work, we propose solutions for identifying domestic individuals using 3D geometric morphometrics on isolated elements from the long bones of the hindlimb in modern reindeer populations. These bones are important for understanding both the mobility of reindeer and the effect of load carrying or draught. A good level of distinction between the size and shape variables of these bones was found among subspecies, sex and lifestyles. This demonstrates that the long bones of the hindlimb can provide information on changes in locomotor behaviour induced by the domestication process, such as control and reduction of reindeer mobility by humans. This also demonstrates that analysis in geometric morphometrics is useful for exploring the use of draught reindeer in early Sámi reindeer herding and the implications for understanding reindeer domestication and early reindeer herding strategies.

Earliest archaeological evidence for domesticated reindeer economy among the Sámi of Northeastern Fennoscandia AD 1300 onwards

The domestication of the reindeer among the Sámi of Northern Fennoscandia is a pressing question for the archaeology of the area and has wider relevance to animal domestication studies globally. Despite considerable research activity, many details of reindeer domestication and early reindeer management remain unclear. This paper explores the use of draught reindeer in early Sámi reindeer herding and the implications for understanding reindeer domestication and early reindeer herding strategies. Faunal assemblages from three Sámi dwelling sites in Northeastern Fennoscandia (AD 1300–1800) were subjected to radiocarbon dating and analysis of palaeopathological lesions, entheseal changes and osteometric measurements. The results suggest that working reindeer were present in the archaeological assemblages from AD 1300 onwards. This is the earliest direct evidence of draught reindeer use by the Sámi. It predates the earliest unequivocal historical sources on draught reindeer use, and confirms the hypothesis that draught reindeer were important in early reindeer herding. Our results show that that small-scale reindeer herding was integrated into the subsistence strategy of the Sámi of Northeastern Fennoscandia earlier than previously suggested. Furthermore, the results imply that training and working together with reindeer were ways of constructing the domestication relationship between the Sámi and reindeer.

Transnational Landscapes of Sámi Reindeer: Domestication and Herding in Northernmost Europe 700–1800 A.D.

ABSTRACT This paper presents the development of Sámi reindeer domestication and pastoralism in northwestern Sápmi, the homeland of the indigenous Sámi people, based on recent surveys in the Gilbbesjávri region, Finland. We have documented about 99 percent of known sites in the study area and present the first radiocarbon dates from herding sites in this part of Finnish Sápmi. Since Sápmi is transnational, archaeological data from the adjoining countries are included in the analyses. The features connected to the early use of domesticated reindeer appear by the 9th century A.D. in a mixed hunter-herder economy and suggest a tethered residential but dynamic logistical mobility. Major changes in site locations and features link with the initiation of nomadic pastoralism by the 15th century A.D., with a high residential mobility. The expansion of both early herding and mobile pastoralism appear to have been conscious, indigenous Sámi responses to wider socio-economic and environmental developments. “The Sámi have always lived in these parts, here in the Sámi homeland.” Sámi author and reindeer herder Johan Turi (2011 [1910], 10)

Reindeer from Sámi offering sites document the replacement of wild reindeer genetic lineages by domestic ones in Northern Finland starting from 1400 to 1600 AD

Reindeer herding emerged among the indigenous Sámi of Northern Fennoscandia between ca. 800 and 1500 CE. While the details of the reindeer domestication process are still actively debated, it has been hypothesized that the transition to reindeer herding affected Sámi ritual practice, especially animal offerings given at various sacred sites. To explore this hypothesis, we analyzed DNA from reindeer bone samples dating to ca. 1200–1700 CE from Sámi offering sites located in present-day Northern Finland as well as from samples originating from ancient dwelling site in Southern Finland and Kola Peninsula, Russia. The results show that haplotypes related to wild Finnish forest reindeer (Rangifer tarandus fennicus) began to be replaced by haplotypes common in modern domesticated reindeer in the faunal assemblages from offering sites starting between 1400 and 1600 CE. The results suggest that, although the role of reindeer herding in the economy of the Sámi communities varied greatly, the transition to reindeer herding may have affected ritual practices, testifying to a shared way of reciprocating with the land and animals.

Reindeer Imagery in the Making at Ust’-Polui in Arctic Siberia

The history of reindeer domestication is a critical topic in the study of human-animal relationships across Northern Eurasia. The Iamal-Nenets region of Arctic Siberia, now a global centre of reindeer pastoralism, has been the subject of much recent research on reindeer domestication. However, tracking the beginnings of reindeer domestication in this region and elsewhere in Eurasia has proved challenging. Archaeological imagery is an under-utilized source of information for exploring animal domestication. In this paper we explore the abundant reindeer imagery found at the Iron Age site of Ust’-Polui in Iamal, dating from ~260 bce to ce 140. While reindeer were hunted in Siberia long before the occupation of Ust’-Polui, portable reindeer imagery appears abruptly at this time period, co-occurring at the site with equipment thought to be for training transport reindeer. Training and working with transport reindeer required long-term engagement with specific animals that became well known and precious to their human keepers. Creating, utilizing and depositing the reindeer imagery objects at Ust’-Polui was a way of acknowledging critical new working relationships with specific domestic reindeer.

Identifying early stages of reindeer domestication in the archaeological record: a 3D morphological investigation on forelimb bones of modern populations from Fennoscandia

Reindeer herding probably developed during the Late Iron Age onwards and is still an important part of the subsistence and culture of many peoples in northern Eurasia. However, despite the importance of this husbandry in the history of these Arctic people, the period and place of the origin as well as the spread of domestic reindeer is still highly debated. Besides the existence of different breeding methods in these territories, identifying domesticated individuals in the archaeological record is complicated because reindeers are considered to still be in the early phases of the domestication process. Indeed, the traditional morphological markers used in zooarchaeology to decipher the domestication syndrome are hardly perceptible in these early stages. In this work, we propose solutions for identifying domestic reindeer bones using 3D geometric morphometrics on isolated elements from the long bones of the forelimb (i.e. humerus, radio-ulna and metacarpal). These bones are important to understand both the feeding behaviour and the mobility of reindeer, and the potential effect of load-carrying or draught in the case of domestic reindeer. We analysed 123 modern specimens from Fennoscandia, including the two interbreeding subspecies currently present in these territories: mountain reindeer (Rangifer tarandus tarandus) and forest reindeer (R.t. fennicus); and where the sex and the lifestyle were known (i.e. free-ranging, racing or draught and captive individuals). A good level of discrimination between the size and shape variables of the bones of the forelimb was found among both subspecies and sexes. Moreover, individuals bred in captivity had smaller bone elements and a thinner and more slender morphology than free-ranging individuals. This demonstrates that the long bones of the forelimb can provide information on changes in feeding and locomotor behaviour prompted by the domestication process, like control and/or reduction of mobility and food of individual reindeer by humans. This also demonstrates that analysis in 3D geometric morphometrics is useful in detecting reindeer incipient domestication markers. Our results can be used by archaeologists to trace the early stages of domestication from fossil reindeer remains, and aid in reconstructing the socio-economic changes of past Arctic populations over time.

Genome sequence and comparative analysis of reindeer (Rangifer tarandus) in northern Eurasia

Reindeer are semi-domesticated ruminants that have adapted to the challenging northern Eurasian environment characterized by long winters and marked annual fluctuations in daylight. We explored the genetic makeup behind their unique characteristics by de novo sequencing the genome of a male reindeer and conducted gene family analyses with nine other mammalian species. We performed a population genomics study of 23 additional reindeer representing both domestic and wild populations and several ecotypes from various geographic locations. We assembled 2.66 Gb (N50 scaffold of 5 Mb) of the estimated 2.92 Gb reindeer genome, comprising 27,332 genes. The results from the demographic history analysis suggested marked changes in the effective population size of reindeer during the Pleistocene period. We detected 160 reindeer-specific and expanded genes, of which zinc finger proteins (n = 42) and olfactory receptors (n = 13) were the most abundant. Comparative genome analyses revealed several genes that may have promoted the adaptation of reindeer, such as those involved in recombination and speciation (PRDM9), vitamin D metabolism (TRPV5, TRPV6), retinal development (PRDM1, OPN4B), circadian rhythm (GRIA1), immunity (CXCR1, CXCR2, CXCR4, IFNW1), tolerance to cold-triggered pain (SCN11A) and antler development (SILT2). The majority of these characteristic reindeer genes have been reported for the first time here. Moreover, our population genomics analysis suggested at least two independent reindeer domestication events with genetic lineages originating from different refugial regions after the Last Glacial Maximum. Taken together, our study has provided new insights into the domestication, evolution and adaptation of reindeer and has promoted novel genomic research of reindeer.

Identification of working reindeer using palaeopathology and entheseal changes

ObjectiveThis paper explores the potential of analyzing pathological lesions and entheseal changes in the identification of working reindeer. MethodsThe skeletons of 26 modern working reindeer from Siberia and Northern Finland are analyzed for pathological lesions and entheseal changes. ResultsWorking results in elevated frequencies of pathological lesions, specifically joint disease in cervical and thoracic vertebrae, humeri, os coxae and proximal phalanges. Entheseal scores indicate the intensified use of shoulder flexors and extensors, and possibly elbow, hip and knee flexors and extensors in working reindeer. ConclusionPatterns of skeletal changes can be used in the identification of working reindeer from the past. SignificanceThis study provides first evidence that pathological lesions and entheseal changes can be used to assess draught and cargo use of reindeer. Therefore, the methods presented in this study provide an opportunity to scrutinize past reindeer herding practices, reindeer domestication and human-reindeer cooperation. LimitationsMethods need to be applied with caution due to the multifactorial etiologies of pathological lesions and entheseal changes. Suggestions for future researchWe suggest that in future studies, these methods are applied to archaeological material accompanied by osteometric and contextual analyses.

Domestication as Enskilment: Harnessing Reindeer in Arctic Siberia

The study of reindeer domestication provides a unique opportunity to examine how domestication involves more than bodily changes in animals produced through selection. Domestication requires enskilment among humans and animals, and this process of pragmatic learning is dependent on specific forms of material culture. Particularly with the domestication of working animals, the use of such material culture may predate phenotypic and genetic changes produced through selective breeding. The Iamal region of Arctic Siberia is generating an increasingly diverse set of archeological data for reindeer domestication that evidences such processes. Three early sites, Ust’-Polui, Tiutei-Sale I, and Iarte VI, contain artifacts proposed to be parts of headgear worn by transport reindeer, the earliest dating to just over 2000 years ago. Contemporary Nenets reindeer herders scrutinized replicas of these archeological objects, and comparisons with historic reindeer harness parts from Arctic Russia were also made. Nenets consistently interpreted barbed L-shaped antler pieces from Iamal as parts of headgear for training young reindeer in pulling sleds. Some types of swivels were also interpreted as transport reindeer headgear. Based on these consultations with Nenets and observations of their ongoing reindeer domestication practices, we argue that material things such as headgear, harnesses, and sleds are not merely technological means of using or controlling reindeer in transportation but instead were part of the meshwork within which some reindeer became enskilled to being domestic. Domestication of reindeer and other animals involves ongoing efforts, landscapes, and made things, all of which form the environment within which domestic relationships emerge.

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 Taĭmyr 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.

Zooarchaeological and stable isotope evidence of Sámi reindeer offerings

This paper presents new osteometric and stable isotope evidence of Sámi reindeer offerings. Previous archaeological studies have shown that reindeer domestication and intensification of reindeer herding transformed Sámi indigenous religion. However, because of the methodological challenges in the identification of wild and domesticated reindeer in the archaeological record, the exact nature of the relationship between people and offered reindeer has remained elusive. To address this problem, we analyze zooarchaeological and stable isotope data from thirteen Sámi offering sites situated in Finland and Sweden and dating to c. 1200–1700 CE. We employ zooarchaeological analysis of age, sex and size and explore the possibilities of these analyses to identify domestication and other characteristics of reindeer selected for offering. Analyses of stable isotopes of carbon, nitrogen and sulphur are utilized to identify human influence on reindeer feeding patterns and mobility. Our results show that many kinds of reindeer with different engagements with people were offered. The results confirm that people had different motives for giving offerings and that a simple dichotomy of wild/domesticated does not adequately reflect the range of relationships the Sámi had with reindeer.

History ofTaenia saginataTapeworms in Northern Russia

Taenia saginata is the most common species of tapeworm infecting humans. Infection is acquired by eating cysticercus larvae in undercooked beef. A closely related species, T. asiatica, is found in eastern and southeastern Asia. The larvae of T. asiatica develop in viscera of pigs. In northern Russia, there is a third member of this morphologically indistinguishable group. Cysticerci of so-called northern T. saginata are found in cerebral meninges of reindeer, and the unique life cycle is dependent on a native custom of eating raw reindeer brain. We report the winding history of this mysterious tapeworm from the first reports to the present time. In addition, we confirm the position of this parasite as a strain of T. saginata by analyzing a mitochondrial DNA sequence of an archival specimen. The origin of this strain might date back to reindeer domestication and contacts between cattle-herding and reindeer-herding peoples in Asia.

The Complex Admixture History and Recent Southern Origins of Siberian Populations.

Although Siberia was inhabited by modern humans at an early stage, there is still debate over whether it remained habitable during the extreme cold of the Last Glacial Maximum or whether it was subsequently repopulated by peoples with recent shared ancestry. Previous studies of the genetic history of Siberian populations were hampered by the extensive admixture that appears to have taken place among these populations, because commonly used methods assume a tree-like population history and at most single admixture events. Here we analyze geogenetic maps and use other approaches to distinguish the effects of shared ancestry from prehistoric migrations and contact, and develop a new method based on the covariance of ancestry components, to investigate the potentially complex admixture history. We furthermore adapt a previously devised method of admixture dating for use with multiple events of gene flow, and apply these methods to whole-genome genotype data from over 500 individuals belonging to 20 different Siberian ethnolinguistic groups. The results of these analyses indicate that there have been multiple layers of admixture detectable in most of the Siberian populations, with considerable differences in the admixture histories of individual populations. Furthermore, most of the populations of Siberia included here, even those settled far to the north, appear to have a southern origin, with the northward expansions of different populations possibly being driven partly by the advent of pastoralism, especially reindeer domestication. These newly developed methods to analyze multiple admixture events should aid in the investigation of similarly complex population histories elsewhere.

Animal offerings at the Sámi offering site of Unna Saiva – Changing religious practices and human–animal relationships

Unna Saiva is a Sámi offering site situated in Gällivare in Northern Sweden. The site was excavated in the early 20th century. It yielded a large number of finds, including objects of silver, pewter and other metals, coins, and animal bones. The metal objects and coins date mainly to the late 10th century and 11th century AD, whereas the animal bone finds date from the 6th to the 17th centuries AD. Zooarchaeological analysis, radiocarbon datings of animal bones and stable isotope analyses conducted in this study reveal new information about religious ritual, religious change, and human–animal relationships among the Sámi. We argue that there was a change in the offering tradition, intertwining with changes in the subsistence economy and especially reindeer domestication. Our results indicate that reindeer domestication, acknowledged to have had a major impact on social organization and economy, was also a major factor in the transformation of Sámi indigenous religion. However, the underlying nature of the offering tradition remained consistent although the focal species of economic and religious interest changed.

Sacrifice as the ideal hunt: a cosmological explanation for the origin of reindeer domestication

The Siberian Northeast shows striking parallels between the cosmologies of hunters and reindeer herders. What may this tell us about the transformation from hunting to pastoralism? This article argues for a structural identity between hunting and sacrifice, and for the domestication of the reindeer as the result of hunters' efforts to use sacrifice to control the accidental variables of the hunt. Hunters can practise their ethos of ‘trust’ with prey only through highly controlled ritual enactments. We describe two: the famous bear festival of the Amur Gulf region and the consecrated reindeer of the Eveny. Both express the same overall logic by which sacrifice functions as an ideal hunt. The animal is involved in a relation not of domination but of trust, while also undergoing a process of taming. We therefore suggest that the reindeer's domestication may be based not only on ecological or economic adaptations, but also on cosmology.