Taxonomy and origin of reindeer

Reindeer and caribou was probably the key species for the human immigration and colonization in the Arctic and sub-Arctic by the retreat of the ice in the last glacial period. The close connection between human and reindeer has contributed to great interest and variation in reindeer taxonomy and origin. Through the history several both species, subspecies and types of reindeer and caribou have been described. The early taxonomy of the species is marked by comparisons of individual specimen using traits as body size, skin colour or antler formations - characteristics known to be highly variable and subjected to environmental and nutritional level. During the mid 1900s the taxonomy was more based on variation of morphological traits among populations by analysing a large series of specimens representative of the various geographic populations and a consensus of classification of several subspecies, all belonging to the same species, evolved. During late 1900 the development of modern molecular techniques procured tools for revealing genetic structure of populations reflecting different origin and isolation rather than environmental influences. The genetic structure revealed a major genetic dichotomy between American woodland caribou on the one hand and all other types of reindeer and caribou on the other which gave evidence that the ancestors of present woodland caribou had survived and evolved in ice free refugium south to the glacier in North America and the ancestors of all other types of reindeer and caribou had evolved separated from these in refugium in Eurasia and Beringia. The ancestors of present reindeer in Scandinavia appear furthermore to have evolved from different populations separated during the last glaciation period and the colonization and origin of present wild and domestic reindeer will be discussed in this perspective.Taksonomi og opprinnelse til reinAbstract in Norwegian / Sammendrag: Rein og caribou har hatt stor betydning for det moderne menneskets utvikling og kolonisering av nordlige Eurasia og Amerika etter siste istid. Den nære sammenhengen mellom mennesker og rein har bidratt til stor interesse og variasjon i oppfatningen av reinens taksonomi og opprinnelse. Et utall av både arter, underarter og raser av rein er beskrevet opp gjennom historien. Tidlig taksonomi av rein bar preg av å være basert på enkeltobservasjoner og på morfologiske karakterer som kroppsstørrelse, pelsfarge og størrelse og form på gevir, karakterer som i stor grad påvirkes av miljø og næringsforhold. Først på midten av 1900 tallet ble taksonomien i større grad basert på ulike morfologiske trekk som viste variasjon mellom bestander av rein, og en fikk bl.a. en forståelse av at alle underarter og former av rein og caribou tilhørte samme art. Med utviklingen av den moderne molekylærbiologien på slutten av 1900-tallet fikk en tilgang til verktøy som avdekket genetiske strukturer som reflekterer ulik opprinnelse og utvikling mer enn miljømessig påvirkning. Den genetiske strukturen som ble avdekket, viste liten overensstemmelse med oppdelingen i underarter som var basert på morfologiske trekk. Molekylærgenetiske struktur viser et hovedskille mellom amerikansk woodland caribou på den ene siden og all annen rein og caribou på den andre siden, noe som reflekterer at forfedrene til woodland caribou levde og utviklet seg i isfrie områder sør for iskanten i Nord-Amerika, mens forfedrene til andre typer rein levde atskilt fra disse i isfrie områder i Eurasia og Beringia. Forfedrene til dagens rein i Fennoskandia syntes også å ha utviklet seg fra atskilte bestander av rein som kan føres tilbake til slutten av siste istid. Innvandring og opprinnelse til dagens vill- og tamrein i Fennoskandia vil bli belyst i dette perspektiv

Refugial origin and postglacial colonization of holarctic reindeer and caribou

The classification and colonization of reindeer and caribou (Rangifer tarandus) was assessed from analysis of both proteins, nuclear DNA and mitochondrial DNA. I demonstrate that the current subspecies designations are not compatible with the differentiation at these markers, suggesting that the morphological differences among extant subspecies did not evolve in separate glacial refugia. Thus, morphological differences among extant subspecies probably evolved as adaptive responses to post-glacial environmental changes. An exception to this is the North American woodland caribou, where all three marker systems support a subspecies-specific refugium as the ancestral origin of these animals. Three major mtDNA haplogroups reported, represent three separate origins of the species during the last glaciation. The most influential origin has contributed to the gene pool of all extant subspecies, suggesting the existence of a large and continuous glacial population ranging across extensive areas of tundra in Eurasia and Beringia. The North American tundra forms (R.t. granti and groenlandicus) and the arctic forms (R.t platyrhynchus, R.t pearyi and R.t eogroenlandicus) almost exclusively comprise haplotypes of such an origin. Another small and isolated refugium seems to have arisen in western Eurasia in close connection to the extensive ice sheet that covered Fennoscandia. The two Eurasian subspecies R.t. tarandus and R.t. fennicus appear to have a diphyletic origin as both the putatively small and isolated Eurasian refugium and the large Beringia refugium have contributed to their gene pools. A third distinct and geographically well-defined refugial area was probably located south to the extensive North American continental ice sheet from where the ancestors of the present North American woodland caribou (R.t. caribou) likely originated.Abstract in Norwegian / Sammendrag: Systematisk inndeling og kolonisering av rein (Rangifer tarandus) ble bestemt ved å analysere for variasjon i genetiske markører som proteiner, kjerneDNA og mitokondrieDNA. Dagens oppdeling av rein i underarter viser liten overensstemmelse med variasjonsmønsteret i de undersøkte markørene, noe som viser at de morfologiske forskjellene som karakteriserer dagens underarter ikke har utviklet seg i atskilte refugier i løpet av siste istid. Unntak fra dette er nordamerikansk skogsrein (woodland caribou-R.t. caribou) hvor alle tre markørsystemene indikerer at denne har utviklet seg i et refugium forskjellig fra andre underarter. De tre registrerte hovedhaplogruppene i mitokondrie-DNA representerer tre atskilte opprinnelser av rein i løpet av siste istid. Den mest innflytelsesrike av disse bidro vesentlig til genbanken til alle dagens underarter av rein, noe som tyder på at det under siste istid eksisterte en stor reinpopulasjon med kontinuerlig utbredelse gjennom store deler av tundraen i Eurasia og Beringia. De nordamerikanske tundrareintypene (R.t. granti og R.t. groenlandicus), samt de arktiske typene (R.t. platyrhunchus, R.t. pearyi og R.t. eogroenlandicus) består nærmest utelukkende av haplotyper med denne opprinnelse. Et annet lite og isolert refugium syntes å ha oppstått i Vest-Europa i nærheten av den omfattende isbreen som dekket Fennoskandia. De to europeiske underarter, R.t. tarandus og R.t. fennicus, syntes å ha en todelt opprinnelse med genetisk påvirkning fra både det antatt lille og isolerte refugiet i Eurasia samt fra det store Beringia refugiet. Et tredje geografisk distinkt refugium var antagelig lokalisert sør for den omfattende isbreen i Nord Amerika hvorfra forfedrene til dagens nordamerikanske skogsrein (R.t. caribou) har sin mest sannsynlige opprinnelse

DNA from Ancient Reindeer Antler as Marker for Transport Routes and Movement of Craftspeople, Raw Material and Products in Medieval Scandinavia

This pilot project is a joint venture between natural and cultural scientists that share a common interest in exploiting whether available DNA technology makes it possible to trace back archaeologically found reindeer antler from medieval urban comb production sites to its original provenance. The provenancing of reindeer antler, used in the production of combs and other personal accessories during the Middle Ages, may be a key factor in the study of the identity and the organization of medieval combmakers. Hereunder routes of transportation used by these craftspeople and their products are important. Provenancing will also enhance the understanding of the social and economic importance of the different reindeer mass trapping systems in medieval Scandinavia.publishedVersio

Transferrin variation and evolution of Alaskan reindeer and caribou, Rangifer tarandus L.

Polyacrylamide gel electrophoresis was used to analyse transferrin variation in wild caribou (Rangifer tarandus granti) and domestic reindeer (R.t. tarandus) from Alaska. Eighteen alleles were detected in caribou and ten alleles were detected in reindeer. The most common allele was Tf 1 with a frequency of 0.304 and 0.408 in caribou and reindeer, respectively. The allele frequency distributions were significantly different in reindeer and caribou. This finding, together with the absence in reindeer of nine alleles present in caribou, suggests that little genetic exchange has taken place between caribou and reindeer in Alaska. The allele frequency distribution in Alaska caribou and reindeer are compared with those for other populations of caribou and reindeer. This comparison indicates that Alaskan caribou as well as Eurasian reindeer have evolved from a common ancesteral population different from the ancesteral population of Peary cairbou (R.t. pearyi) and Svalbard reindeer (R.t. platyrhynchus)

Transferrin variation and evolution of Canadian barren-ground caribou

Blood samples were obtained from 95 barren-ground caribou (Rangifer tarandus groenlandicus) of the Beverly herd in Northwest Territories, Canada. Polyacrylamid gel electrophoresis was used to score for genetic variation in the locus coding for transferrin. The pattern of allele frequency distribution are compared with previously reported values of Eurasian tundra reindeer (R.t. tarandus), Alaska caribou (R.t. granti), Peary caribou (R.t. pearyi), and Svalbard reindeer (R.t. platyrhynchus). In the Beverly herd a total of 21 different transferrin alleles were detected. The amount of genetic variation was higher in the Canadian barren-ground caribou than what has been detected in other subspecies of reindeer/caribou. Highly gene-tical differences in the allele frequencies were detected between the Canadian barren-ground caribou and the other subspecies. The genetic identity analyses indicates approximately the same amount of genetic differentiation when the Canadian barren-ground caribou are compared with Alaska caribou as with the Peary caribou. The allele frequency pattern could be explained by a possible origin of the Canadian barren-ground caribou from an ancestral population which was genetical influenced by animals surviving the We-ichselian glaciation in refugia both in high Arctic, in Beringia, and south of the ice sheet

Maintaining genetic integrity of coexisting wild and domestic populations : Genetic differentiation between wild and domestic Rangifer with long traditions of intentional interbreeding

The funding for the fieldwork and laboratory work for this study was provided by the ERC Advanced Grant 295458 Arctic Domus (PI D.G. Anderson). The writing and analysis was supported by ESRC ES-M0110548-1 JPI HUMANOR (PI D.G. Anderson). The sample set for Lake Nichatka was collected and deposited under a research programme of the Norwegian Institute for Nature Research. We thank Liv Midthjell for skilful laboratory analyses, Konstantin Klokov for help sourcing statistics on Russian reindeer populations, and Jan Heggenes for useful comments on an earlier version of this paper. A full list of project participants is in Appendix 2.Peer reviewedPublisher PD

Sex-biased dispersal in a northern ungulate population

In most mammals dispersal is male-biased and in many polygynous ungulates female philopatry and matrilineal grouping involve small-scale genetic structure. We have through sex-related differences in microsatellite allele distribution addressed sex-biased dispersal in a spatially expanding northern ungulate population. The Norwegian red deer population (Cervus elaphus atlanticus) has the last hundred years grown substantially and expanded spatially after a major decline from 300 to 100 years ago. Previous Bayesian analyses suggest a present division of genetic variation into five geographically separated subpopulations. Among these subpopulations the overall Fst values were 0.067 (SE=0.014) for males and 0.094 (SE=0.017) for females. Pairwise Fst values were significantly higher for females than males, demonstrating a stronger genetic structure among females, and that dispersal has been lower in females than males. Accordingly, a higher number of male than female first generation dispersers were identified among the five subpopulations using Bayesian assignment with prior population information, but significantly so only with relaxed stringency levels of assignment. The identified male-biased dispersal distances varied from 30 to 300 kilometers suggesting male biased dispersal on a large scale in red deer

Genetic structure and origin of semi-domesticated reindeer

This chapter discusses the genetic variation and structure of semi-domesticated reindeer, as well as the genetic effects of the domestication process in connection with the pastoral transition from hunters to herders. The research material analysed includes data on the ancestors of the Fennoscandian semi-domesticated reindeer, as well as on genetic diversity in extant reindeer. The study provides a better understanding of when and how domestication occurred, as well as the implications for future adaptation to a changing environment.publishedVersio

Habituation responses in wild reindeer exposed to recreational activities

Displacement is the effect most often predicted when recreational activities in wild reindeer (Rangifer tarandus tarandus) areas are discussed. Wild reindeer in Blefjell (225 km2) are exposed to humans more frequently than in Hardangervidda (8200 km2), from which the Blefjell herd originate. We recorded fright and flight response distances of groups of reindeer in both herds to a person directly approaching them on foot or skis during winter, summer, and autumn post-hunting and rutting season in 2004-2006. The response distances sight, alert, flight initiation and escape were shorter in Blefjell than in Hardangervidda while the probability of assessing the observer before flight tended to be greater in Blefjell. To test whether these results could be due to habituation or genetic influence of semi-domestic reindeer previously released in the Blefjell region, we compared the genetic variation of the Blefjell reindeer with previously reported variation in semi-domestic reindeer and in the wild reindeer from Hardangervidda. Microsatellite analyses revealed closer genetic ancestry of the Blefjell reindeer to the wild Hardangervidda reindeer and not to the semi-domestic reindeer at both the herd and the individual level. We conclude that the decreased flight responses in Blefjell reindeer appear to be a habituation response to frequent human encounters rather than traits inherited from a semi-domestic origin