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Abstract Climate warming at the end of the last glacial period had profound effects on the distribution of cold-adapted species. As their range shifted towards northern latitudes, they were able to colonise previously glaciated areas, including remote Arctic islands. However, there is still uncertainty about the routes and timing of colonisation. At the end of the last ice age, reindeer/caribou (Rangifer tarandus) expanded to the Holarctic region and colonised the archipelagos of Svalbard and Franz Josef Land. Earlier studies have proposed two possible colonisation routes, either from the Eurasian mainland or from Canada via Greenland. Here, we used 174 ancient, historical and modern mitogenomes to reconstruct the phylogeny of reindeer across its whole range and to infer the colonisation route of the Arctic islands. Our data shows a close affinity among Svalbard, Franz Josef Land and Novaya Zemlya reindeer. We also found tentative evidence for positive selection in the mitochondrial gene ND4, which is possibly associated with increased heat production. Our results thus support a colonisation of the Eurasian Arctic archipelagos from the Eurasian mainland and provide some insights into the evolutionary history and adaptation of the species to its High Arctic habitat.

Abstract Reindeer, called caribou in North America, has a circumpolar distribution and all extant populations belong to the same species (Rangifer tarandus). It has survived the Holocene thanks to its immense adaptability and successful coexistence with humans in different forms of hunting and herding cultures. Here, we examine the paternal and maternal history of Rangifer based on robust Y‐chromosomal and mitochondrial DNA (mtDNA) trees representing Eurasian tundra reindeer, Finnish forest reindeer, Svalbard reindeer, Alaska tundra caribou, and woodland caribou. We first assembled Y‐chromosomal contigs, representing 1.3 Mb of single‐copy Y regions. Based on 545 Y‐chromosomal and 458 mtDNA SNPs defined in 55 males, maximum parsimony trees were created. We observed two well separated clades in both phylogenies: the “EuroBeringian clade” formed by animals from Arctic Islands, Eurasia, and a few from North America and the “North American clade” formed only by caribou from North America. The time calibrated Y tree revealed an expansion and dispersal of lineages across continents after the Last Glacial Maximum. We show for the first time unique paternal lineages in Svalbard reindeer and Finnish forest reindeer and reveal a circumscribed Y haplogroup in Fennoscandian tundra reindeer. The Y chromosome in domesticated reindeer is markedly diverse indicating that several male lineages have undergone domestication and less intensive selection on males. This study places R. tarandus onto the list of species with resolved Y and mtDNA phylogenies and builds the basis for studies of the distribution and origin of paternal and maternal lineages in the future.

Summary: Typically much smaller in number than their mainland counterparts, island populations are ideal systems to investigate genetic threats to small populations. The Svalbard reindeer (Rangifer tarandus platyrhynchus) is an endemic subspecies that colonized the Svalbard archipelago ca. 6,000–8,000 years ago and now shows numerous physiological and morphological adaptations to its arctic habitat. Here, we report a de-novo chromosome-level assembly for Svalbard reindeer and analyze 133 reindeer genomes spanning Svalbard and most of the species’ Holarctic range, to examine the genomic consequences of long-term isolation and small population size in this insular subspecies. Empirical data, demographic reconstructions, and forward simulations show that long-term isolation and high inbreeding levels may have facilitated the reduction of highly deleterious—and to a lesser extent, moderately deleterious—variation. Our study indicates that long-term reduced genetic diversity did not preclude local adaptation to the High Arctic, suggesting that even severely bottlenecked populations can retain evolutionary potential.

Abstract We have analyzed DNA microsatellites and the mitochondrial control region in reindeer from 31 different husbandry areas in Norway, Sweden, and Finland in order to better understand the processes that underlie the genetic variability of the Nordic domestic herds. The distinct differentiation found in the nuclear markers but less so in the mitochondrial marker gives evidence of an origin from a common ancestral population which later evolved into the two main gene pools characterizing the nuclear genomes of domestic reindeer in Finland and most of Sweden and Norway. Analyses of temporal trends in effective population size give evidence of a rapid increase in number of reindeer before the population growth associated with the pastoral transition. This implies that the ancestry of contemporary domestic reindeer lay among a rapidly growing wild population possibly located in the boreal areas of eastern Fennoscandia or European Russia. The evolution of reindeer husbandry in Finland, perhaps with input from European Russia, which later spread to northern Norway could explain the shared genomic pattern observed in these areas today. The structured selection of productive female‐centered herds may explain the genetic structure in other parts of Norway and in Sweden. The further substructuring of the Swedish/ Norwegian gene pool appears to follow the traditional language borders with the South Sámi language dominating the southern and the Central Sámi language in the more northern genetic subclusters. This suggests that traditional knowledge, cultural identities, and herd migrations have contributed to shape the genetic structure seen today. Ecological gradients are more pronounced within as compared to between the genetic clusters, giving further evidence that historical and social–cultural processes are important drivers for the genetic differentiations found in domestic reindeer across the Nordic countries.

Abstract 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.

Abstract Background Since the 1950s, the Norwegian–Swedish Coldblooded trotter (NSCT) has been intensively selected for harness racing performance. As a result, the racing performance of the NSCT has improved remarkably; however, this improved racing performance has also been accompanied by a gradual increase in inbreeding level. Inbreeding in NSCT has historically been monitored by using traditional methods that are based on pedigree analysis, but with recent advancements in genomics, the NSCT industry has shown interest in adopting molecular approaches for the selection and maintenance of this breed. Consequently, the aims of the current study were to estimate genomic-based inbreeding coefficients, i.e. the proportion of runs of homozygosity (ROH), for a sample of NSCT individuals using high-density genotyping array data, and subsequently to compare the resulting rate of genomic-based F (FROH) to that of pedigree-based F (FPED) coefficients within the breed. Results A total of 566 raced NSCT were available for analyses. Average FROH ranged from 1.78 to 13.95%. Correlations between FROH and FPED were significant (P

Chronic wasting disease (CWD) persists in cervid populations of North America and in 2016 was detected for the first time in Europe in a wild reindeer in Norway. We report the detection of CWD in 3 moose (Alces alces) in Norway, identified through a large scale surveillance program. The cases occurred in 13–14-year-old female moose, and we detected an abnormal form of prion protein (PrPSc) in the brain but not in lymphoid tissues. Immunohistochemistry revealed that the moose shared the same neuropathologic phenotype, characterized by mostly intraneuronal deposition of PrPSc. This pattern differed from that observed in reindeer and has not been previously reported in CWD-infected cervids. Moreover, Western blot revealed a PrPSc type distinguishable from previous CWD cases and from known ruminant prion diseases in Europe, with the possible exception of sheep CH1641. These findings suggest that these cases in moose represent a novel type of CWD.

Abstract Background Although harness racing is of high economic importance to the global equine industry, significant genomic resources have yet to be applied to mapping harness racing success. To identify genomic regions associated with harness racing success, the current study performs genome-wide association analyses with three racing performance traits in the Norwegian-Swedish Coldblooded Trotter using the 670 K Axiom Equine Genotyping Array. Results Following quality control, 613 horses and 359,635 SNPs were retained for further analysis. After strict Bonferroni correction, nine genome-wide significant SNPs were identified for career earnings. No genome-wide significant SNPs were identified for number of gallops or best km time. However, four suggestive genome-wide significant SNPs were identified for number of gallops, while 19 were identified for best km time. Multiple genes related to intelligence, energy metabolism, and immune function were identified as potential candidate genes for harness racing success. Conclusions Apart from the physiological requirements needed for a harness racing horse to be successful, the results of the current study also advocate learning ability and memory as important elements for harness racing success. Further exploration into the mental capacity required for a horse to achieve racing success is likely warranted.

Abstract Infection patterns linked to age and sex are crucial to predict the population dynamic effects of diseases in long‐lived species. How such demographic patterns of infection arise is often multifactorial, although the cause is commonly seen as a combination of immune status as well as variation in pathogen exposure. Prion diseases are particularly interesting, as they do not trigger an adaptive immune response; hence, differences in pathogen exposure linked to behavior could be the prime determinant of the pattern of infection. In cervids, the fatal prion disease, chronic wasting disease (CWD), is spreading geographically, with economic and cultural consequences in affected areas in North America, and all infected individuals eventually die from disease‐associated sequelae if they live long enough. Understanding the causes of the demographic pattern of infection with CWD is therefore urgent but is limited by the fact that reported data primarily come from related deer species in North America. The recent (detected 2016) emergence of CWD among wild alpine reindeer (Rangifer tarandus) in Norway with a different social organization, that is, no home range behavior and no matrilineal female groups, offers an opportunity to advance our understanding of how behavior influences the infection patterns. Testing of 1081 males and 1278 females detected 19 animals positive for abnormal prion protein in brain and/or lymphatic tissues. No calves and only one male yearling were infected, with the remaining positives being adults (representing 1.5% of adult males and 0.5% of adult females). We found a strong sex‐biased infection pattern in reindeer (with infection 2.7 times more likely in adult males), which is similar to the results reported in mule deer and white‐tailed deer. The hazard of being detected as positive increased with age in males. There was no close genetic relatedness among positive animals. The results were consistent with the within‐group contact of males being a possible major route of transmission. We discuss the demographic pattern of infection with CWD in view of the lack of stable home range behavior and other key behavioral traits of reindeer relevant to understanding pathogen exposure in general.

In reindeer husbandry, animal breeding could offer tools for improving productivity by selection. The traits affecting meat production efficiency are primarily related to calf growth and survival, and to dam’s longevity and lifetime ability to raise heavy calves. Information on genetic (co)variation in these traits is scarce but needed in estimating the potential and effectiveness of selection as well as biological constraints. The objectives of the study were to estimate these genetic parameters from the data of an experimental reindeer herd at Kutuharju (Inari, Finland). Heritabilities (h2) and genetic correlations (rg) among the average daily gain of calves (ADG), dams’ age at maturity, individual fitness (λind) and the cumulative sum of her calves’ weaning weight over seven years (WW7) were studied with AS-Reml software using the dataset from the experimental herd. The pedigree included also sire information to allow the separation of the maternal effects. Direct and maternal heritabilities of ADG were moderate (0.24±0.09 and 0.18±0.05, respectively) and the direct-maternal correlation was highly negative (-0.73±0.17). Indeed, selection on growth necessitates information on both calf and dam. The genetic correlation of growth with birth date and birth weight could not be detected with the data. The age at maturity and λind had a small heritable component (0.07±0.12 and 0.10±0.06, respectively), whereas the heritability value of WW7 was 0.23±0.07. Reindeer herders’ empirical selection on calf’s autumn weight favours fast growth (rg between growth and autumn weight = 0.35±0.24). Dam’s weight in her first autumn was strongly correlated with her lifetime production expressed by her individual fitness (rg = 0.71±0.23) and the cumulative sum of her calves’ weaning weight (rg = 0.63±0.12). Hence, the early information on the dam (her weight in her first autumn or her first calf’s autumn weight) works as useful selection criteria for the traits related to lifetime production.

The factors causing variation in birth date and birth weight were analysed from the data from an experimental reindeer herd consisting of 1136 calves with parental information. The traits had coefficient of variation of 37 and 14%, respectively. The variation in both traits was affected by year and sex (male calves heavier) and by management factors, such as the age distribution of females and males. Early calving results from the use of older breeding males and is most apparent in prime age females. In both traits the heritability was moderate (0.23 and 0.24) with a high proportion of maternal genetic variation (0.23) in birth weight. The North Atlantic Oscillation (NAO) indices, summarizing the major weather conditions prior to rut, explained part of the annual variation in the traits. The amount of total genetic variation in relation to trait mean, or the evolvability, was 21% in birth date and 10% in birth weight indicating that selection could successfully be used to improve herd productivity and that the traits have substantial potential for adapting to possible changing environmental conditions. The results on genetic correlations imply that selection on calf’s birth weight leads on one hand to calves being born earlier and on the other hand to dams with later parturition.

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.

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.

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 rein Abstract 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.

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.

Polycrylamide gel electrophoresis was used to analyse tranferrrin variation in caribou populations from Manitoba, Ontario, Québec/Labrador, and from Baffin Island, Northwest Territories in eastern Canada. The transferrin allele frequencies in these populations were compared with those previously reported for Canadian barren-ground caribou, Rangifer tarandus groenlandicus, Alaska caribou, R.t. grand, Peary caribou, R.t. pearyi, Svalbard reindeer, R.t. pla-tyrhynchus, and Eurasian tundra reindeer, R.t. tarandus. A total of twenty different alleles was detected in the analysed material, considerable genetic heterogeneity being detected among regions. Three alleles that were relatively common in caribou from Ontario, Manitoba and Québec/Labrador, were not present in R.t. grand, R.t. pearyi, R.t. tarandus or R.t. platyrhynchus, and present only at very low frequencies 'm R.t. groenlandicus. These findings, together with genetic identity analyses, suggest that the caribou in Manitoba, Ontario, and Québec/Labrador are mainly of the R.t. caribou type, and that little interbreeding has occurred with other subspecies. The large genetic distance in the transferrin locus between R.t. caribou and other subspecies of reindeer/caribou suggests that, during the Wisconsin glaciation the ancestral populations of R.t. caribou survived in a refugium different from that of the ancestral populations of the other subspecies. Significant genetic differences between Baffin Island caribou and all other populations were mainly due to the presence of one allele that was in high frequency in Baffin Island caribou, but that was absent, or present in very low frequencies, in all other reindeer/caribou populations. The genetic differences between Baffin Island caribou and the other subspecies were greater than the differences between several of the currently recognized subspecies.

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.

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).

Polyacrylamide gel electrophoresis was used to analyse transferrin variation in herds of semi-domestic reindeer from Scandinavia. The results are compared with previously reported values for other populations of both semi-domestic and wild reindeer using the same techniques as in the present study. In all populations the number of alleles was high, ranging from seven to eleven, and the heterozygosity was correspondingly high, with a mean of 0.749. This high genetic variation in all populations suggests that inbreeding is not widespread among Scandinavian reindeer. The pattern of allele frequency distribution indicates a high degree of genetic heterogeneity in the transferrin locus, both between the different semi-domestic herds and between the different wild populations. The mean value of genetic distance was 0.069 between semi-domestic herds and 0.091 between wild populations. Between semi-domestic and wild populations the genetic distance was particularly high, with a mean of 0.188. This high value was mainly due to a different pattern in the distribution of the two most common transferrin alleles: Tfu was most common among semi-domestic herds, while TfEI was most common among wild populations. These differences in transferrin allele distribution are discussed in relation to possible different origins of semi-domestic and wild reindeer in Scandinavia, or alternatively, to different selection forces acting on transferrin genotypes in semi-domestic and wild populations. Transferrin-variasjon og genetisk struktur hos rein i Skandinavia. Abstact in Norwegian / Sammendrag: Transferrin-variasjon i tamreinflokker ble analysert ved hjelp av polyacrylamid gel elektroforese. Resultatene er sammenlignet med verdier som tidligere er beskrevet for både tamrein og villrein hvor det ble benyttet samme metode som i denne undersøkelsen. I alle populasjonene ble det registrert et høyt antall alleler (7-11) og heterozygositeten var tilsvarende høy med en middelverdi på 0.749. Denne høye graden av genetisk variasjon i alle undersøkte populasjoner indikerer at det ikke foregår mye innavl blant rein i Skandinavia. Utbredelsen av de enkelte allelene viste høy grad av genetisk oppdeling i transferrin-locuset mellom flokker av både tamrein og villrein. Middelverdien for genetisk avstand var 0.069 mellom tamreinflokker og 0.091 mellom villreinflokker. Særlig stor genetisk avstand (middelverdi 0.188) ble funnet mellom tamrein og villrein. Denne store forskjellen skyldes i stor grad forskjellig mønster i utbredelsen av de to vanligste allelene: Tf' var mest vanlig blant tamrein og Tf1' var mest vanlig blant villrein. Denne forskjellen er diskutert i relasjon til forskjellig opprinnelse av tamrein og villrein og alternativt, i relasjon til forskjellig seleksjonskrefter som virker på transferrin genotyper i tamrein og villrein.

Intense selective pressures applied over short evolutionary time have resulted in homogeneity within, but substantial variation among, horse breeds. Utilizing this population structure, 744 individuals from 33 breeds, and a 54,000 SNP genotyping array, breed-specific targets of selection were identified using an F(ST)-based statistic calculated in 500-kb windows across the genome. A 5.5-Mb region of ECA18, in which the myostatin (MSTN) gene was centered, contained the highest signature of selection in both the Paint and Quarter Horse. Gene sequencing and histological analysis of gluteal muscle biopsies showed a promoter variant and intronic SNP of MSTN were each significantly associated with higher Type 2B and lower Type 1 muscle fiber proportions in the Quarter Horse, demonstrating a functional consequence of selection at this locus. Signatures of selection on ECA23 in all gaited breeds in the sample led to the identification of a shared, 186-kb haplotype including two doublesex related mab transcription factor genes (DMRT2 and 3). The recent identification of a DMRT3 mutation within this haplotype, which appears necessary for the ability to perform alternative gaits, provides further evidence for selection at this locus. Finally, putative loci for the determination of size were identified in the draft breeds and the Miniature horse on ECA11, as well as when signatures of selection surrounding candidate genes at other loci were examined. This work provides further evidence of the importance of MSTN in racing breeds, provides strong evidence for selection upon gait and size, and illustrates the potential for population-based techniques to find genomic regions driving important phenotypes in the modern horse.

Although there is little doubt that the domestication of mammals was instrumental for the modernization of human societies, even basic features of the path towards domestication remain largely unresolved for many species. Reindeer are considered to be in the early phase of domestication with wild and domestic herds still coexisting widely across Eurasia. This provides a unique model system for understanding how the early domestication process may have taken place. We analysed mitochondrial sequences and nuclear microsatellites in domestic and wild herds throughout Eurasia to address the origin of reindeer herding and domestication history. Our data demonstrate independent origins of domestic reindeer in Russia and Fennoscandia. This implies that the Saami people of Fennoscandia domesticated their own reindeer independently of the indigenous cultures in western Russia. We also found that augmentation of local reindeer herds by crossing with wild animals has been common. However, some wild reindeer populations have not contributed to the domestic gene pool, suggesting variation in domestication potential among populations. These differences may explain why geographically isolated indigenous groups have been able to make the technological shift from mobile hunting to large-scale reindeer pastoralism independently. [ABSTRACT FROM AUTHOR]

Evolutionary models of sex ratio adjustment applied to mammals have ignored that females may gain indirect genetic benefits from their mates. The differential allocation hypothesis (DAH) predicts that females bias the sex ratio of their offspring towards (more costly) males when breeding with an attractive male. We manipulated the number of available males during rut in a polygynous ungulate species, the reindeer (Rangifer tarandus), and found that a doubling of average male mass (and thus male attractiveness) in the breeding herd increased the proportion of male offspring from approximately 40 to 60%. Paternity analysis revealed indeed that males of high phenotypic quality sired more males, consistent with the DAH. This insight has consequences for proper management of large mammal populations. Our study suggests that harvesting, by generating a high proportion of young, small and unattractive mates, affects the secondary sex ratio due to differential allocation effects in females. Sustainable management needs to consider not only the direct demographic changes due to harvest mortality and selection, but also the components related to behavioural ecology and opportunities for female choice. [ABSTRACT FROM AUTHOR]

Large mammals in seasonal environments have a pattern of high-reproductive synchrony in spring, but how the timing of reproduction affects resource allocation decisions at different stages of the reproductive cycle remains largely unexplored. By manipulating the timing of conception in reindeer (Rangifer tarandus), we tested how the timing of conception affected sex ratio, gestation length and weight development of mother and offspring. Females that conceived at their first ovulation within the rut had a 60.5% probability of producing a male; in contrast, females that conceived a cycle later had a 31.3% probability of producing a male. Late conceiving females had gestation times that were 10 days shorter and the calves were 0.6kg (9.2%) lighter at birth and 7.4kg (14.7%) lighter in autumn. Over the year, female weight changes was similar between the groups suggesting reindeer follow a bet-hedging strategy; reducing the quality of this year's offspring to ensure their own future reproduction and survival. Harvesting is often selective leading to skewed sex ratios and age structure, which may influence the timing of reproduction due to females hesitation to mate with young males. Whenever this hesitation is strong enough to increase the frequency of recycling, harvesting is likely to have profound life history consequences. [ABSTRACT FROM AUTHOR]

In sexually dimorphic ungulates, male reproductive success depends on fighting with other males for access to females during a brief rutting season. Large body size is necessary for success in intrasexual competition, and a few large-sized males are often able to monopolize access to female groups. Earlier studies have reported that reproductive effort increases with age until prime-age is reached, and one study that population density lowered effort in (older) males. No study has directly assessed whether there is within-age-class variation in effort resulting from varying levels of intra-male competition. It is reported here the weight loss during the rutting season of 54 individual male reindeer Rangifer tarandus coming from eight herds with varying density (3.3–6.0 deer/km2) and sex ratio (4–28% males). In agreement with earlier studies, reproductive effort was lower for young (1- to 2-year-old) than for prime-aged (3- to 5-year-old) males both on an absolute and relative scale. Among 1-year-old males (n=33), effort was lower as sex ratio became closer to even, but density during the rutting season had no effect. This suggests that yearling males take a more active role when prime-aged males are absent. In addition to the insight into male ungulate life history, understanding male rutting behaviour may also have implications for population dynamics. [ABSTRACT FROM AUTHOR]

The Swedish-Norwegian Coldblooded trotter (CBT) is a local breed in Sweden and Norway mainly used for harness racing. Previous studies have shown that a mutation from cytosine (C) to adenine (A) in the doublesex and mab-3 related transcription factor 3 (DMRT3) gene has a major impact on harness racing performance of different breeds. An association of the DMRT3 mutation with early career performance has also been suggested. The aim of the current study was to investigate this proposed association in a randomly selected group of CBTs. 769 CBTs (485 raced, 284 unraced) were genotyped for the DMRT3 mutation. The association with racing performance was investigated for 13 performance traits and three different age intervals: 3 years, 3 to 6 years, and 7 to 10 years of age, using the statistical software R. Each performance trait was analyzed for association with DMRT3 using linear models. The results suggest no association of the DMRT3 mutation with precocity (i.e. performance at 3 years of age). Only two traits (race time and number of disqualifications) were significantly different between the genotypes, with AA horses having the fastest times and CC horses having the highest number of disqualifications at 3 years of age. The frequency of the AA genotype was significantly lower in the raced CBT sample compared with the unraced sample and less than 50% of the AA horses participated in a race. For the age intervals 3 to 6 and 7 to 10 years the AA horses also failed to demonstrate significantly better performance than the other genotypes. Although suggested as the most favorable genotype for racing performance in Standardbreds and Finnhorses across all ages, the AA genotype does not appear to be associated with superior performance, early or late, in the racing career of CBTs.

Dispersal and gene flow are important mechanisms affecting the dynamics of vectors and their pathogens. Here, patterns of genetic diversity were analyzed in many North European populations of the tick, Ixodes ricinus. Population sites were selected within and between areas separated by geographical barriers in order to evaluate the importance of tick transportation by birds in producing genetic connectivity across open sea and mountain ranges. The phylogenetic analyses of the mitochondrial control region and the cytochrome b gene revealed two distinct clades with supported sub-clades, with three genetic lineages: GB and WNo associated with Great Britain and western Norway respectively, and Eu with a wider distribution across continental Europe in agreement with much lower efficiency of tick dispersal by birds than by large mammals. The results suggest different ancestry of I. ricinus colonizing Britain and the rest of northern Europe, possibly from different glacial refuges, while ticks from western Norway and continental Europe share a more recent common ancestry. Demographic history modeling suggests a period of strong increase in tick abundance coincident with progression of the European Neolithic culture, long after their post-glacial colonization of NW Europe.

In light of current debates on global climate change it has become important to know more on how large, roaming species have responded to environmental change in the past. Using the highly variable mitochondrial control region, we revisit theories of Rangifer colonization and propose that the High Arctic archipelagos of Svalbard, Franz Josef Land, and Novaia Zemlia were colonized by reindeer from the Eurasian mainland after the last glacial maximum. Comparing mtDNA control region sequences from the three Arctic archipelagos showed a strong genetic connection between the populations, supporting a common origin in the past. A genetic connection between the three archipelagos and two Russian mainland populations was also found, suggesting colonization of the Eurasian high Arctic archipelagos from the Eurasian mainland. The age of the Franz Josef Land material (>2000 years before present) implies that Arctic indigenous reindeer colonized the Eurasian Arctic archipelagos through natural dispersal, before humans approached this region.

Horses were domesticated from the Eurasian steppes 5,000-6,000 years ago. Since then, the use of horses for transportation, warfare, and agriculture, as well as selection for desired traits and fitness, has resulted in diverse populations distributed across the world, many of which have become or are in the process of becoming formally organized into closed, breeding populations (breeds). This report describes the use of a genome-wide set of autosomal SNPs and 814 horses from 36 breeds to provide the first detailed description of equine breed diversity. F(ST) calculations, parsimony, and distance analysis demonstrated relationships among the breeds that largely reflect geographic origins and known breed histories. Low levels of population divergence were observed between breeds that are relatively early on in the process of breed development, and between those with high levels of within-breed diversity, whether due to large population size, ongoing outcrossing, or large within-breed phenotypic diversity. Populations with low within-breed diversity included those which have experienced population bottlenecks, have been under intense selective pressure, or are closed populations with long breed histories. These results provide new insights into the relationships among and the diversity within breeds of horses. In addition these results will facilitate future genome-wide association studies and investigations into genomic targets of selection.

An equine SNP genotyping array was developed and evaluated on a panel of samples representing 14 domestic horse breeds and 18 evolutionarily related species. More than 54,000 polymorphic SNPs provided an average inter-SNP spacing of ∼43 kb. The mean minor allele frequency across domestic horse breeds was 0.23, and the number of polymorphic SNPs within breeds ranged from 43,287 to 52,085. Genome-wide linkage disequilibrium (LD) in most breeds declined rapidly over the first 50-100 kb and reached background levels within 1-2 Mb. The extent of LD and the level of inbreeding were highest in the Thoroughbred and lowest in the Mongolian and Quarter Horse. Multidimensional scaling (MDS) analyses demonstrated the tight grouping of individuals within most breeds, close proximity of related breeds, and less tight grouping in admixed breeds. The close relationship between the Przewalski's Horse and the domestic horse was demonstrated by pair-wise genetic distance and MDS. Genotyping of other Perissodactyla (zebras, asses, tapirs, and rhinoceros) was variably successful, with call rates and the number of polymorphic loci varying across taxa. Parsimony analysis placed the modern horse as sister taxa to Equus przewalski. The utility of the SNP array in genome-wide association was confirmed by mapping the known recessive chestnut coat color locus (MC1R) and defining a conserved haplotype of ∼750 kb across all breeds. These results demonstrate the high quality of this SNP genotyping resource, its usefulness in diverse genome analyses of the horse, and potential use in related species.