Background Dynamic laryngeal collapse (DLC) associated with poll flexion is the most common disorder of the upper respiratory tract (URT) in the Norwegian-Swedish Coldblooded Trotter (NSCT). The disorder, which has also been diagnosed in other breeds of trotters and gaited horses, appears to be related to anatomic phenotypes and only occurs during poll flexion when the horse is exercised 'on the bit'. Objectives Identify genomic regions associated with DLC in the NSCT by combining a rigorous phenotyping protocol with genomic data from a high-density equine genotyping array. Study design Prospective case/control study. Methods High-speed treadmill endoscopy was used to phenotype horses (n = 61) for DLC, distinguishing between cases and controls. Genome-wide association (GWA) analysis of DLC status was then performed using a principal component approach (PCA) with haplotype analyses subsequently performed for regions containing single-nucleotide polymorphisms (SNPs) above the suggestive genome-wide significance (GWS) threshold (P<1.0 x 10(-5)). Results One region containing 10 SNPs (Equus caballus chromosome [ECA] 7: 89,601,935-94,647,192) was above the suggestive GWS threshold. Two inferred haplotypes in this region demonstrated significant differences (P<0.001) between cases and controls, with the most frequent haplotype resulting in a significantly increased risk of DLC. Main limitations Small sample size as a result of stringent phenotyping protocols. Conclusions The current study highlights a candidate genomic region on ECA7 as potentially important with regard to the manifestation of DLC. Further exploration of this region and the genes included within it will bring veterinarians and researchers closer to fully understanding the biological mechanisms underlying DLC in horses.
Females' dispersion during the mating season has been regarded as being determined primarily by the distribution of food resources. However, females' distribution and behaviour may also be affected by the males' availability during rut. Indeed, it is challenging to disentangle female dispersion for food from female mate choice. We present the results of female reindeer (Rangifer tarandus) behaviour in two manipulated herds during the peak week of the rut: one without males (MA) and one with males present (MP). Presence of males did not influence mean typical group size of females (MP herd: 10.14, MA herd: 11.85). However, females in the MA herd travelled longer daily distances (1.8km) compared to females in the MP herd (1.3km). The proportion of the time females spent feeding (~ 75%), walking (~ 16%), and standing (~ 8%) did not differ between the herds, whereas mating-related behaviour was, as expected, significantly higher in the MP herd, although it accounted for only 1.4% of their observed activity. The high proportion of time spent feeding indicates that females' movement is driven primarily by maximizing forage intake. No difference in the females' somatic body weight change during rut between the two herds indicates low cost of female mating related activities during rut. Contrary to our predictions, the results showed that female reindeer behaviour during the mating season is not affected by male availability, male mating or courtship behaviour, but is primarily driven by distribution of food resources.
Cucujus cinnaberinus is an obligate saproxylic beetle distributed in Europe and considered as near threatened in its entire range (IUCN red list). Ten polymorphic microsatellites were characterized among 26 and 45 individual samples obtained from dead trees in the Czech Republic and Norway respectively. All loci were polymorphic in both samples with number of alleles per locus ranged from two to eleven in the Norwegian sample and from two to six in the Czech sample. The genetic difference between the Czech and the Norwegian material were highly significant (P < 0.0001). For several trees the sampled genotypes gave evidence that more than two parents were responsible for the breeding. These microsatellite loci should prove useful in the study of population structure including mate and dispersal patterns for this species.
For centuries, domestic horses have represented an important means of transport and served as working and companion animals. Although their role in transportation is less important today, many horse breeds are still subject to intense selection based on their pattern of locomotion. A striking example of such a selected trait is the ability of a horse to perform additional gaits other than the common walk, trot and gallop. Those could be four-beat ambling gaits, which are particularly smooth and comfortable for the rider, or pace, used mainly in racing. Gaited horse breeds occur around the globe, suggesting that gaitedness is an old trait, selected for in many breeds. A recent study discovered that a nonsense mutation in DMRT3 has a major impact on gaitedness in horses and is present at a high frequency in gaited breeds and in horses bred for harness racing. Here, we report a study of the worldwide distribution of this mutation. We genotyped 4396 horses representing 141 horse breeds for the DMRT3 stop mutation. More than half (2749) of these horses also were genotyped for a SNP situated 32 kb upstream of the DMRT3 nonsense mutation because these two SNPs are in very strong linkage disequilibrium. We show that the DMRT3 mutation is present in 68 of the 141 genotyped horse breeds at a frequency ranging from 1% to 100%. We also show that the mutation is not limited to a geographical area, but is found worldwide. The breeds with a high frequency of the stop mutation (>50%) are either classified as gaited or bred for harness racing.
The protein-coding region of melanocortin 1 receptor (MC1R) was sequenced to identify potential variation affecting coat color in reindeer (Rangifer tarandus). A T→C sequence variation at nucleotide position 218 (c.218T>C) causing an amino acid (aa) change from methionine to threonine at aa position 73 (p.Met73Thr) was identified. In addition, a T→G sequence variation was found at nucleotide position 839 (c.839T>G), causing phenylalanine to be exchanged by cysteine at aa position 280 (p.Phe280Cys). The two sequence variants (c.218C and c.839G) were found to be closely associated with a darker belly coat compared with animals not having any of these two variants. The aa acid change p.Met73Thr affects the same position as p.Met73Lys previously reported to give constitutive activation of MC1R in black sheep (Ovis aries), whereas p.Phe280Cys is identical to one of two variants previously reported to be associated with dark coat color in Arctic fox (Alopex lagopus), supporting that the two variants found in reindeer are functional. The complete absence of Thr73 and Cys280 among the 51 wild reindeer analyzed provides some evidence that these variants are more common in the domestic herds.
REASONS FOR PERFORMING STUDY:The pathogenesis of osteochondrosis (OC) and palmar/plantar first phalanx osteochondral fragments (POFs) is multifactorial, but specific knowledge of heritability is limited.OBJECTIVES:To improve the precision of heritability estimates and to estimate the genetic correlation between tarsocrural OC and POFs in Standardbred trotters. Further aims were to examine whether the prevalence of OC/POFs was different in the American and French lineages that have contributed to the Norwegian population, and if the prevalence was affected by heterozygosity.STUDY DESIGN:Retrospective cohort study.METHODS:Categorical data on tarsocrural OC and POFs from 2 radiographic studies performed in 1989 and 2007/2008 (n = 1217) were analysed with sire threshold models that included 230 sires.RESULTS:Heritability of OC at the distal intermediate ridge of the tibia and/or the lateral trochlear ridge of the talus was estimated at 0.29 ± 0.15. For OC at the distal intermediate ridge of the tibia only, the estimate was 0.40 ± 0.17. Heritability of POFs in all 4 limbs was estimated at 0.23 ± 0.13; for metatarsophalangeal POFs this was 0.26 ± 0.13 and for medial metatarsophalangeal POFs 0.32 ± 0.14. Estimates of genetic correlation between OC and POFs ranged from 0.68 ± 0.27 to 0.73 ± 0.28 but were not significantly different from a zero-genetic correlation. Effects of lineages or heterozygosity were not observed.CONCLUSIONS AND POTENTIAL RELEVANCE:This study confirmed a moderate to high heritability of tarsocrural OC and POF, providing further evidence of the heritable nature of these diseases. Examination of specific lesions yielded the highest heritability; therefore, breeding programmes and future genome-analysis studies should focus on predilection sites rather than the entire disease complex.
Palmar/plantar osteochondral fragments (POF) in fetlock joints commonly affect and influence the athletic performance of horses. In this study, we used the Equine SNP50 BeadChip® to perform a genome-wide association study of metatarsophalangeal POF in 176 Norwegian Standardbred trotter yearlings. Putative quantitative trait loci (QTL) for medial and/or lateral POF, and medial POF only were identified on ECA1, 2, 7, 9 and 31, whereas for lateral POF, only on ECA7, 11, 27 and X. The moderate number of QTL evidences a complex inheritance and suggests various genes controlling POF development in medial and lateral locations.
Rates of introgression from non-indigenous into native populations are increasing worldwide, often as a result of anthropogenic translocation events. In ungulates translocations have been common, especially among deer. European red deer consists of two distinct lineages, one western and one eastern. These probably originate from different glacial refuges, but it is unknown to what extent they hold different adaptations. Here we address dispersal and introgression into the Norwegian mainland population from an introduced island stock consisting of an admixture of both European lineages. The last decade this stock has grown considerably in number and dispersal could be expected to have increased. We therefore used samples separated by a 5 year interval from Otterøya, adjacent mainland areas and a more distant sub-population. Bayesian assignment analysis verified the genetic structure and identified dispersal between the Otterøya stock and the adjacent mainland coastal areas. Three individuals (two newly sampled) with second or third generation non-indigenous origin were found among the adjacent mainland samples (5 and 3 %, respectively). Two individuals with first and second generation mainland-origin were found on Otterøya (old samples). This suggests some non-indigenous introgression from Otterøya into the mainland Norwegian population.
Wildlife species exposed to habitat fragmentation are often in need of a conservation effort. The African buffalo (Syncerus caffer) is one of the key species in the Serengeti ecosystem as they form a large part of the herbivore biomass, providing ecotourism and valuable trophies. The ecosystem is a part of Tanzanias protected areas and is administrated under different management practices. Among these, we have analysed the genetic structure of buffalo (n = 68) from the Serengeti National Park (SNP), the Ngorongoro conservation area (NCA) and the Maswa game reserve (MGR). Both the sequence variation in a 493 base pair fragment of the mitochondrial D-loop and the allele frequency-distribution in 15 microsatellites suggest genetic structuring of the buffalo populations within the ecosystem. Both the allele frequency-distribution and the amount of genetic variation were high and similar in SNP and MGR, suggesting a high degree of gene flow between these locations. By comparison, the NCA buffaloes had significantly lower genetic variation and were genetically differentiated from SNP and MGR. Approximate Bayesian computation estimates suggest that the observed genetic structure is of a recent origin, indicating that the recent increases in developmental activity in the region may have influenced the genetic structure of the buffalo within the Serengeti ecosystem.
Knowledge about changes in behavioural traits related to wildness and tameness is for most mammals lacking, despite the increased trend of using domestic stock to re-establish wild populations into historical ranges. To test for persistence of behavioural traits of wild reindeer (Rangifer tarandus L.) exposed to hunting, we sampled DNA, vigilance and flight responses in wild reindeer herds with varying domestic ancestry. Analyses of 14 DNA microsatellite loci revealed a dichotomous main genetic structure reflecting their native origin, with the Rondane reindeer genetically different from the others and with least differentiation towards the Hardangervidda reindeer. The genetic clustering of the reindeer in Norefjell-Reinsjofjell, Ottadalen and Forollhogna, together with domestic reindeer, supports a predominant domestic origin of these herds. Despite extensive hunting in all herds, the behavioural measures indicate increasing vigilance, alert and flight responses with increasing genetic dissimilarity with domestic herds. Vigilance frequency and time spent vigilant were higher in Rondane compared to Hardangervidda, which again were higher than herds with a domestic origin. We conclude that previous domestication has preserved a hard wired behavioural trait in some reindeer herds exhibiting less fright responses towards humans that extensive hunting has, but only slightly, altered. This brings novel and relevant knowledge to discussions about genetic diversity of wildlife in general and wild reindeer herds in Norway in specific.
Despite the importance of the reindeer husbandry in the subsistence of many northern cultures the origin and spread of domestic reindeer is still highly debated. Recent analyses of mitochondrial DNA in reindeer herds across Eurasia revealed distinct geographic structures of domestic reindeer in Fennoscandia and Russia, pointing towards independent origins of domestic reindeer in these areas. A high degree of haplotype sharing between the extant wild reindeer population in the Hardangervidda mountain region in southern Norway and the domestic herds in Fennoscandia indicate that this population could have contributed in the early domestication or augmented the domestic population. However, genetic analyses of excavated reindeer remains from the early medieval period from the Hardangervidda region revealed no haplotype sharing with extant domestic reindeer, demonstrating that reindeer from Hardangervidda did not contribute in the early domestication process in Fennoscandia. The substantial temporal genetic alteration observed in this population is related to introgression of domestic reindeer into the wild gene pool in the 19th century during periods when reindeer husbandry was practiced in this mountain region.
Many species with currently continuously distributed populations have histories of geographic range shifts and successive shifts between decline or fragmentation, growth and spatial expansion. The moose (Alces alces) colonised Scandinavia after the last ice age. Historic records document a high abundance and a wide distribution across Norway in the middle ages, but major decline and fragmentation in the eighteenth and nineteenth centuries. After growth and expansion during the twentieth century, the Norwegian population is currently abundant and continuously distributed. We examined the distribution of genetic variation, differentiation and admixture in Norwegian moose, using 15 microsatellites. We assessed whether admixture has homogenised the population or if there are any genetic structures or discontinuities that can be related to recent or ancient shifts in demography or distribution. The Bayesian clustering algorithm STRUCTURE without any spatial information showed that there is currently a genetic dichotomy dividing the population into one southern and one northern subpopulation. Including spatial information, the Bayesian clustering algorithm TESS, which considers gradients of genetic variation and spatial autocorrelation, suggests that the population is divided into three subpopulations along a latitudinal axis, the southern one identical to the one identified with STRUCTURE. Present convergence zones of high admixture separate the identified subpopulations, which are delimited by genetic discontinuities corresponding to geographic barriers against dispersal, e.g. wide fiords and mountain ranges. The distribution of the subpopulations is supported by spatial autocorrelation analysis. However, some loci are not in Hardy–Weinberg equilibrium and the STRUCTURE analysis suggests that a lower hierarchical structure may exist within the southernmost subpopulation. No bottlenecks or founder events are indicated by the levels of genetic variation, rather a high degree of private alleles in the northern subpopulations indicates introgression. Coalescent-based Approximate Bayesian Computation estimates unambiguously suggest that the genetic structure is a result of an ancient divergence event and a more recent admixture event a few centuries ago. This indicates that the central Scandinavian subpopulation constitutes a relatively recent convergence zone of secondary contact.
REASONS FOR PERFORMING STUDYDevelopmental orthopaedic diseases (DOD) such as osteochondrosis (OC)/osteochondrosis dissecans (OCD), palmar/plantar osteochondral fragments (POF), ununited palmar/plantar eminences (UPE) and dorsoproximal first phalanx fragments are well recognised in the horse. Aetiopathogeneses are controversial and molecular genetic screening of DNA has recently been employed for their elucidation. Precise phenotypic definition and knowledge of breed-specific prevalence and interrelations are essential for the interpretation of following genomic studies in Standardbred trotters.OBJECTIVESTo assess the prevalence, trend of development and interrelation of DOD in tarsocrural, metacarpophalangeal (MCP) and metatarsophalangeal (MTP) joints in Standardbred trotters.METHODSThe tarsocrural and MCP/MTP joints of 464 Norwegian Standardbred yearlings were radiographed and the prevalence and interrelation of osteochondral lesions calculated.RESULTSOsteochondral lesions were diagnosed in 50.7% of the horses. The prevalence of tarsocrural OC/OCD at the distal intermediate ridge of the tibia (DIT) and the lateral trochlear ridge of the talus (LTT) was 19.3%. The prevalence of OC/OCD in MCP joints was 3.6%, whereas those of POF and UPE in MCP/MTP joints were 23.1 and 3.9%, respectively. Interrelation was evident for 1) most equivalent lesions in joint homologues, 2) OCD DIT and OCD LTT and 3) POF and UPE. Lesions in hock and fetlock joints were generally not significantly associated.CONCLUSIONSThe prevalence of tarsocrural OC/OCD in Norwegian Standardbreds is apparently increasing, whereas that of other articular DOD appears stable. Association analyses verify bilateralism for most equivalent lesions and suggest aetiological resemblance also between other lesions. The absence of a significant association between tarsocrural OCD and POF implies that the lesions must be considered statistically different disorders.POTENTIAL RELEVANCEThe prevalence results emphasise that DOD should be considered in Standardbred breeding regimens (e.g. by sire selection subsequent to progeny testing). Also, improved phenotypic definitions will help elucidate the true causal genes in following genomic studies.
The Norwegian red deer population (Cervus elaphus) was from the mid eighteenth to the early twentieth century drastically reduced in size and distribution but has the last century expanded both demographically and spatially. We have investigated genetic variation, differentiation and admixture in this spatially expanding ungulate population, using 14 microsatellites. The present genetic structure is moderate to strong with an average F ST = 0.08. Low M-ratios indicate loss of genetic variation in all localities and signals of a recent bottleneck was identified in 14 of 15 localities. Genetic distances between the localities indicate two main routes of dispersal during expansion, from the north–west and south–west, respectively. Bayesian assignment tests verify a break of the dataset in two, and demonstrate 99.9% probability for the existence of five sub-populations, which coincide well with five relict populations described by historic records. Computer simulations suggest that the observed genetic differentiation is recent rather than ancient, and that it may be explained by models of fragmentation or of founder events and subsequent merging rather than by models of recent bottlenecks in some particular demes within an ancient genetic structure.
SummaryOsteochondrosis (OC), a disturbance in the process of endochondral ossification, is by far the most important equine developmental orthopaedic disease and is also common in other domestic animals and humans. The purpose of this study was to identify quantitative trait loci (QTL) associated with osteochondrosis dissecans (OCD) at the intermediate ridge of the distal tibia in Norwegian Standardbred (SB) using the Illumina Equine SNP50 BeadChip whole‐genome single‐nucleotide polymorphism (SNP) assay. Radiographic data and blood samples were obtained from 464 SB yearlings. Based on the radiographic examination, 162 horses were selected for genotyping; 80 of these were cases with an OCD at the intermediate ridge of the distal tibia, and 82 were controls without any developmental lesions in the joints examined. Genotyped horses descended from 22 sires, and the number of horses in each half‐sib group ranged from 3 to 14. The population structure necessitated statistical correction for stratification. When conducting a case–control genome‐wide association study (GWAS), mixed‐model analyses displayed regions on chromosomes (Equus callabuschromosome – ECA) 5, 10, 27 and 28 that showed moderate evidence of association (P ≤ 5 × 10−5; thisP‐value is uncorrected i.e. not adjusted for multiple comparisons) with OCD in the tibiotarsal joint. Two SNPs on ECA10 represent the most significant hits (uncorrectedP = 1.19 × 10−5in the mixed‐model). In the basic association (chi‐square) test, these SNPs achieved statistical significance with the Bonferroni correction (P = 0.038) and were close in the permuted logistic regression test (P = 0.054). Putative QTL on ECA 5, 10, 27 and 28 represent interesting areas for future research, validation studies and fine mapping of candidate regions. Results presented here represent the first GWAS of OC in horses using the recently released Illumina Equine SNP50 BeadChip.
We report a high-quality draft sequence of the genome of the horse (Equus caballus). The genome is relatively repetitive but has little segmental duplication. Chromosomes appear to have undergone few historical rearrangements: 53% of equine chromosomes show conserved synteny to a single human chromosome. Equine chromosome 11 is shown to have an evolutionary new centromere devoid of centromeric satellite DNA, suggesting that centromeric function may arise before satellite repeat accumulation. Linkage disequilibrium, showing the influences of early domestication of large herds of female horses, is intermediate in length between dog and human, and there is long-range haplotype sharing among breeds.
Traditional reindeer herding of northern Fennoscandia has been based on seasonal movements independent of national borders. At the beginning of the 19th century, these yearly movements of reindeer were excessive, but during that century the borders between the Fennoscandian countries were closed. By analysing a 190-base pair fragment of the mitochondrial DNA control region in 79 museum samples, we show that the reindeer of northern Fennoscandia were one homogenous population shortly after the national borders were closed. However, anthropogenic activity has effectively ended genetic exchange within northern Fennoscandia and has made the reindeer population within this region heterogeneous. Genetic input of eastern origin is also suggested within the extant Russian reindeer of the Kola Peninsula.
HereditasVolume 102, Issue 2 p. 199-206 Open Access Genetic differences at the transferrin locus in Norwegian semidomestic and wild reindeer (Rangifer tarandus L.) K. H. RØED, K. H. RØED Department of Zoology, Agricultural University, NorwaySearch for more papers by this author K. H. RØED, K. H. RØED Department of Zoology, Agricultural University, NorwaySearch for more papers by this author First published: August 1985 https://doi.org/10.1111/j.1601-5223.1985.tb00616.xCitations: 7AboutPDF ToolsExport citationAdd to favoritesTrack citation ShareShare Give accessShare full text accessShare full-text accessPlease review our Terms and Conditions of Use and check box below to share full-text version of article.I have read and accept the Wiley Online Library Terms and Conditions of UseShareable LinkUse the link below to share a full-text version of this article with your friends and colleagues. Learn more.Copy URL Abstract Blood samples from 928 reindeer from five semi-domestic and three wild populations of reindeer, Rangifer tarandus L., from southern Norway were analysed for transferrin variability by polyacrylamide gel electrophoresis. A total of 12 alleles were detected, and in all populations the number of alleles was high, ranging from eight to eleven. The mean value of heterozygosities was approximately the same for semi-domestic (0.778) and wild (0.770) populations. The pattern of allele frequency distribution indicates a high degree of genetic heterogeneity in the transferrin locus and each population revealed significant differences for at least one allele between all pairs of combinations. Using a hierarchical approach, 45% of the heterogeneity between populations was explained by dividing into semi-domestic and wild animals. The major contributor to the divergence was the Tfct allele which changed in frequency from a mean (±SD) of 0.331 (±0.040) in semidomestic herds, to 0.167 (±0.045) in wild populations. These results suggest that the different selection strategies in the management of semi-domestic and wild reindeer influence the transferrin allele frequencies. References Ashton, G. C. 1958. B-globulin polymorphism and early foetal mortality in cattle. Nature 183: 404– 405. Ashton, G. C. 1965. Cattle serum transferrins: a balanced polymorphism Genetics 52: 983– 997. Brænd, M. 1964. Genetic studies on serum transferrins in reindeer. Hereditas 52: 181– 188. Cavalli-Sforza, L. 1966. Population structure and human evolution. Proc. R. Soc. London Ser. B 164: 362– 379. Chen, S. H. and Sutton, H. E. 1967. Bovine transferrins: Sialic acid and the complex phenotype. Genetics 56: 425– 430. Diezel, W., Kopperschläger, G. and Hofmann, E. 1972. 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Effective population size, generation interval, and potential loss of genetic variability in game species under different hunting regimes. Oikos 36: 257– 266. Selander, R. K. 1976. Genetic variation in natural populations.– In Molecular Evolution (Ed. F. J. Ayala), Sinauer Ass., Sunderland , Mass. , p. 21– 45. Shubin, P. N. 1977. Electrophoretic study of proteins in two races of reindeer. Izv. Akad. Nauk CCP, Ser. Biol. 6: 819– 828 (in Russian). Shubin, P. N. and Ionova, T. A. 1981. Genetic interrelation between domestic and wild reindeer, Rangifer tarandus L. Biologischeskie Problemy Severa (IX Simpozium) Ch. 11, Syktyvkar 1981 (in Russian). Shubin, P. N. and Matyukov, V. S. 1982. Genetic differentiation of reindeer populations. Genetika 18: 2030– 2035 (in Russian). Skjenneberg, S. and Slagsvold, L. 1968. Reindriften. Scandinavian University Books. Universitetsforlaget, Oslo/Bergen/Tromsø (in Norwegian). Skogland, T. and Mølmen, Ø. 1980. 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Røed, K. H. 1987. Transfenin variation and body size in reindeer. Rangifer tarandus L. - Hereditas 106: 67–71. Lund, Sweden. ISSN 0018–0661. Received March 10, 1986 Associations between transferrin variation and individual weight of reindeer calves and yearlings sampled in two subsequent years were investigated. Polyacrylamide gel electrophoresis was used to analyse the transferrin variation. Animals heterozygous or homozygous at the transferrin locus did not show any significant differences in individual dressed weight. The most common allele, TfET, was positively associated with high weight among male calves but not among yearlings. The second most common allele, TfEJ, was positively associated with high weight among male yearlings but not among calves. This indicates that different transferrin alleles in reindeer have different selective advantages over time, which suggests that balancing selection is important in maintaining transferrin variation in reindeer.
Polyacrylamide gel electrophoresis was used to score genetic variation at 35 loci in wild reindeer, Rangifer tarandus L., sampled from five different areas within two mountain regions in southern Norway. The amount of genetic variability was within the range reported in mammals in general. The proportion of polymorphic loci at the 99% criterion for polymorphism varied from 0.114 to 0.200, while the average heterozygosity varied from 0.041 to 0.053 in the five populations investigated. The mean value of genetic distance between populations was 0.0017, and highly significant allele frequency differences were observed in reindeer from the two mountain regions, as well as within regions. This finding suggests that the amount of gene flow between local populations of wild reindeer can be highly restricted even in the presence of migration. The amount of genetic variability in wild reindeer is compared with previously reported values for semidomestic reindeer investigated with the same loci and techniques as in the present study. The wild reindeer reveal slightly more genetic variability than do semi-domestic reindeer and this is discussed in relation to the importance of genetic variability in the early domestication process.