Sea ice loss increases genetic isolation in a high Arctic ungulate metapopulation.

Sea ice loss may have dramatic consequences for population connectivity, extinction-colonization dynamics, and even the persistence of Arctic species subject to climate change. This is of particular concern in face of additional anthropogenic stressors, such as overexploitation. In this study, we assess the population-genetic implications of diminishing sea ice cover in the endemic, high Arctic Svalbard reindeer (Rangifer tarandus platyrhynchus) by analyzing the interactive effects of landscape barriers and reintroductions (following harvest-induced extirpations) on their metapopulation genetic structure. We genotyped 411 wild reindeer from 25 sampling sites throughout the entire subspecies' range at 19 microsatellite loci. Bayesian clustering analysis showed a genetic structure composed of eight populations, of which two were admixed. Overall population genetic differentiation was high (mean FST  = 0.21). Genetic diversity was low (allelic richness [AR] = 2.07-2.58; observed heterozygosity = 0.23-0.43) and declined toward the outer distribution range, where populations showed significant levels of inbreeding. Coalescent estimates of effective population sizes and migration rates revealed strong evolutionary source-sink dynamics with the central population as the main source. The population genetic structure was best explained by a landscape genetics model combining strong isolation by glaciers and open water, and high connectivity by dispersal across winter sea ice. However, the observed patterns of natural isolation were strongly modified by the signature of past harvest-induced extirpations, subsequent reintroductions, and recent lack of sea ice. These results suggest that past and current anthropogenic drivers of metapopulation dynamics may have interactive effects on large-scale ecological and evolutionary processes. Continued loss of sea ice as a dispersal corridor within and between island systems is expected to increase the genetic isolation of populations, and thus threaten the evolutionary potential and persistence of Arctic wildlife.

Opposing fitness consequences of habitat use in a harvested moose population.

Landscape changes are happening at an unprecedented pace, and together with high levels of wildlife harvesting humans have a large effect on wildlife populations. A thorough knowledge of their combined influence on individual fitness is important to understand factors affecting population dynamics. The goal of the study was to assess the individual consistency in the use of risky habitat types, and how habitat use was related to fitness components and life-history strategies. Using data from a closely monitored and harvested population of moose Alces alces, we examined how individual variation in offspring size, reproduction and survival was related to the use of open grasslands; a habitat type that offers high-quality forage during summer, but at the cost of being more exposed to hunters in autumn. The use of this habitat type may therefore involve a trade-off between high mortality risk and forage maximization. There was a high repeatability in habitat use, which suggests consistent behaviour within individuals. Offspring number and weight were positively related to the mothers' use of open grasslands, whereas the probability of surviving the subsequent harvest season was negatively related to the use of the same habitat type. As a consequence, we found a nonsignificant relationship between habitat use and lifetime fitness. The study suggests that harvesting, even if intended to be nonselective with regard to phenotypes, may be selective towards animals with specific behaviour and life-history strategies. As a consequence, harvesting can alter the life-history composition of the population and target life-history strategies that would be beneficial for individual fitness and population growth in the absence of hunting.

Genomic measures of inbreeding in the Norwegian-Swedish Coldblooded Trotter and their associations with known QTL for reproduction and health traits.

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 < 0.001) and ranged from 0.27 to 0.56, with FPED and FROH from 2000 to 2009 increasing by 1.48 and 3.15%, respectively. Comparisons of ROH between individuals yielded 1403 regions that were present in at least 95% of the sampled horses. The average percentage of a single chromosome covered in ROH ranged from 9.84 to 18.82% with chromosome 31 and 18 showing, respectively, the largest and smallest amount of homozygosity. CONCLUSIONS: Genomic inbreeding coefficients were higher than pedigree inbreeding coefficients with both methods showing a gradual increase in inbreeding level in the NSCT breed between 2000 and 2009. Opportunities exist for the NSCT industry to develop programs that provide breeders with easily interpretable feedback on regions of the genome that are suboptimal from the perspective of genetic merit or that are sensitive to inbreeding within the population. The use of molecular data to identify genomic regions that may contribute to inbreeding depression in the NSCT will likely prove to be a valuable tool for the preservation of its genetic diversity in the long term.

Recalibrating Equus evolution using the genome sequence of an early Middle Pleistocene horse.

The rich fossil record of equids has made them a model for evolutionary processes. Here we present a 1.12-times coverage draft genome from a horse bone recovered from permafrost dated to approximately 560-780 thousand years before present (kyr BP). Our data represent the oldest full genome sequence determined so far by almost an order of magnitude. For comparison, we sequenced the genome of a Late Pleistocene horse (43 kyr BP), and modern genomes of five domestic horse breeds (Equus ferus caballus), a Przewalski's horse (E. f. przewalskii) and a donkey (E. asinus). Our analyses suggest that the Equus lineage giving rise to all contemporary horses, zebras and donkeys originated 4.0-4.5 million years before present (Myr BP), twice the conventionally accepted time to the most recent common ancestor of the genus Equus. We also find that horse population size fluctuated multiple times over the past 2 Myr, particularly during periods of severe climatic changes. We estimate that the Przewalski's and domestic horse populations diverged 38-72 kyr BP, and find no evidence of recent admixture between the domestic horse breeds and the Przewalski's horse investigated. This supports the contention that Przewalski's horses represent the last surviving wild horse population. We find similar levels of genetic variation among Przewalski's and domestic populations, indicating that the former are genetically viable and worthy of conservation efforts. We also find evidence for continuous selection on the immune system and olfaction throughout horse evolution. Finally, we identify 29 genomic regions among horse breeds that deviate from neutrality and show low levels of genetic variation compared to the Przewalski's horse. Such regions could correspond to loci selected early during domestication.

Novel Type of Chronic Wasting Disease Detected in Moose (Alces alces), Norway.

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.

A genome-wide association study for harness racing success in the Norwegian-Swedish coldblooded trotter reveals genes for learning and energy metabolism.

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.

Fitness correlates of age at primiparity in a hunted moose population.

Trade-offs between fitness-related traits are predicted from the principle of resource allocation, where increased fecundity or parental investment leads to reduced future reproduction or survival. However, fitness traits can also be positively correlated due to individual differences (e.g. body mass). Age at primiparity could potentially explain variation in individual fitness either because early primiparity is costly, or it may lead to higher lifetime reproductive success. Based on long-term monitoring and genetic parentage assignment of an island population of moose, we quantified reproductive performance and survival, and examined whether early maturing females have higher total calf production than late maturing females. We explored if harvesting of calves affected the subsequent reproductive success of their mothers, i.e. also due to a post-weaning cost of reproduction, and whether there are any intergenerational effects of female reproductive success. There was a positive relationship between current and future reproduction. The probability to reproduce was lower for females that were unsuccessful the year before, indicating a strong quality effect on productivity. Females that started to reproduce as 2-year olds had a slightly higher total calf production compared to those starting at age three or four. High-performing mothers were also correlated with daughters that performed well in terms of reproductive success. Our results suggest that the observed individual heterogeneity in fitness could be associated with differences in age at primiparity. This heterogeneity was not affected by reproductive costs associated with tending for a calf post-weaning.

Genome-wide analysis reveals selection for important traits in domestic horse breeds.

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.

A high density SNP array for the domestic horse and extant Perissodactyla: utility for association mapping, genetic diversity, and phylogeny studies.

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.

Population properties affect inbreeding avoidance in moose.

Mechanisms reducing inbreeding are thought to have evolved owing to fitness costs of breeding with close relatives. In small and isolated populations, or populations with skewed age- or sex distributions, mate choice becomes limited, and inbreeding avoidance mechanisms ineffective. We used a unique individual-based dataset on moose from a small island in Norway to assess whether inbreeding avoidance was related to population structure and size, expecting inbreeding avoidance to be greater in years with larger populations and even adult sex ratios. The probability that a potential mating event was realized was negatively related to the inbreeding coefficient of the potential offspring, with a stronger relationship in years with a higher proportion or number of males in the population. Thus, adult sex ratio and population size affect the degree of inbreeding avoidance. Consequently, conservation managers should aim for sex ratios that facilitate inbreeding avoidance, especially in small and isolated populations.

Moose body mass variation revisited: disentangling effects of environmental conditions and genetics.

Large-scale geographical variation in phenotypic traits within species is often correlated to local environmental conditions and population density. Such phenotypic variation has recently been shown to also be influenced by genetic structuring of populations. In ungulates, large-scale geographical variation in phenotypic traits, such as body mass, has been related to environmental conditions and population density, but little is known about the genetic influences. Research on the genetic structure of moose suggests two distinct genetic lineages in Norway, structured along a north-south gradient. This corresponds with many environmental gradients, thus genetic structuring provides an additional factor affecting geographical phenotypic variation in Norwegian moose. We investigated if genetic structure explained geographical variation in body mass in Norwegian moose while accounting for environmental conditions, age and sex, and if it captured some of the variance in body mass that previously was attributed to environmental factors. Genetic structuring of moose was the most important variable in explaining the geographic variation in body mass within age and sex classes. Several environmental variables also had strong explanatory power, related to habitat diversity, environmental seasonality and winter harshness. The results suggest that environmental conditions, landscape characteristics, and genetic structure should be evaluated together when explaining large-scale patterns in phenotypic characters or life history traits. However, to better understand the role of genetic and environmental effects on phenotypic traits in moose, an extended individual-based study of variation in fitness-related characters is needed, preferably in an area of convergence between different genetic lineages.

Inferences about the population history of Rangifer tarandus from Y chromosome and mtDNA phylogenies.

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.

Ancient reindeer mitogenomes reveal island-hopping colonisation of the Arctic archipelagos.

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.

Mismatch between calf paternity and observed copulations between male and female reindeer: Multiple mating in a polygynous ungulate?

In polygynous systems, such as that exhibited by reindeer Rangifer tarandus, mate choice can be difficult to disentangle from male intrasexual competition because male behavior may constrain female choice. Multiple mating may provide an avenue for female mate choice, though it is difficult to identify using behavioral estimators alone. Molecular techniques address this issue by affording ecologists an opportunity to reassess mating systems from a genetic perspective. We assessed the frequency and possible explanations for multiple mating in reindeer using a genetic approach to determine the success of observed copulations in a semi-domesticated herd in Kaamanen, Finland. Behavioral and genetic data were synthesized with population characteristics over a 7-year period to test the hypothesis that, if present, polyandry in reindeer is driven by sexual harassment from sub-dominant males. We observed multiple mating in 42% of females, with as many as 60% exhibiting multiple mating in certain years. We found no evidence that multiple mating resulted from sexual harassment by sub-dominant males, suggesting that it is likely a deliberate strategy among females. Conversion rate of copulations into paternities varied with male size, with smaller males more likely to experience mismatch than larger males. Female preference for larger males persisted despite the occurrence of multiple mating, possibly suggesting a mechanism for cryptic post-copulatory selection. We suggest further research to delineate the possible influence of cryptic post-copulatory selection and multiple mating to defend against infertility in exhausted males.

Evidence suggesting that reindeer mothers allonurse according to the direct reciprocity and generalized reciprocity decision rules.

Allonursing is the nursing of the offspring of other mothers. Cooperation is an emergent property of evolved decision rules. Cooperation can be explained by at least three evolved decision rules: 1) direct reciprocity, i.e. help someone who previously helped you, 2) kin discrimination, i.e. preferentially direct help to kin than to non-kin, and 3) generalized reciprocity, i.e. help anyone if helped by someone. We assessed if semi-domesticated reindeer, Rangifer tarandus, mothers allonursed according to the decision rules of direct reciprocity, generalized reciprocity and kin discrimination over 2 years. To assess if reindeer mothers allonursed according to the direct reciprocity decision rule, we predicted that mothers should give more help to those who previously helped them more often. To assess if reindeer mothers allonursed according to the kin discrimination decision rule, we predicted that help given should increase as pairwise genetic relatedness increased. To assess if reindeer mothers allonursed according to the generalized reciprocity decision rule, we predicted that the overall number of help given by reindeer mothers should increase as the overall number of help received by reindeer mothers increased. The number of help given i) increased as the number of help received from the same partner increased in the 2012 group but not in both 2013 groups, ii) was not influenced by relatedness, and iii) was not influenced by an interaction between the number of help received from the same partner and relatedness. iv) The overall number of help given increased as the overall number of help received increased. The results did not support the prediction that reindeer mothers allonursed according to the kin discrimination decision rule. The results suggest that reindeer mothers may allonurse according to the direct reciprocity and generalized reciprocity decision rules.

Adaptation to the High-Arctic island environment despite long-term reduced genetic variation in Svalbard reindeer.

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.

Reconstructing the history of a fragmented and heavily exploited red deer population using ancient and contemporary DNA.

BACKGROUND: Red deer (Cervus elaphus) have been an important human resource for millennia, experiencing intensive human influence through habitat alterations, hunting and translocation of animals. In this study we investigate a time series of ancient and contemporary DNA from Norwegian red deer spanning about 7,000 years. Our main aim was to investigate how increasing agricultural land use, hunting pressure and possibly human mediated translocation of animals have affected the genetic diversity on a long-term scale. RESULTS: We obtained mtDNA (D-loop) sequences from 73 ancient specimens. These show higher genetic diversity in ancient compared to extant samples, with the highest diversity preceding the onset of agricultural intensification in the Early Iron Age. Using standard diversity indices, Bayesian skyline plot and approximate Bayesian computation, we detected a population reduction which was more prolonged than, but not as severe as, historic documents indicate. There are signs of substantial changes in haplotype frequencies primarily due to loss of haplotypes through genetic drift. There is no indication of human mediated translocations into the Norwegian population. All the Norwegian sequences show a western European origin, from which the Norwegian lineage diverged approximately 15,000 years ago. CONCLUSIONS: Our results provide direct insight into the effects of increasing habitat fragmentation and human hunting pressure on genetic diversity and structure of red deer populations. They also shed light on the northward post-glacial colonisation process of red deer in Europe and suggest increased precision in inferring past demographic events when including both ancient and contemporary DNA.

Population connectivity: dam migration mitigations and contemporary site fidelity in arctic char.

BACKGROUND: Animal feeding and spawning migrations may be limited by physical barriers and behavioral interactions. Dam constructions (e.g. hydropower) commonly include gateways for fish migrations to sustain ecological connectivity. Relative genetic impacts of fish passage devices versus natural processes (e.g. hybrid inferiority) are, however, rarely studied. We examined genetic (i.e. microsatellite) population connectivity of highly migrating lake-dwelling Arctic char (Salvelinus alpinus), introduced 20 generations ago, across and within two subalpine lakes separated by a dam with a subterranean tunnel and spill gates after 7 generations. Due to water flow regime, the time window for fish migration is highly restricted. RESULTS: Char populations, with similar genetic structuring and diversity observed across and within lakes, were admixed across the dam with fishways during feeding. For spawning, however, statistically significant, but very low population differentiation (θ; 0.002 - 0.013) was found in nine out of ten reproductive site comparisons, reflecting interactions between extensive migration (mean first generation (F0) = 10.8%) and initial site fidelity. Simulations indicated that genetic drift among relatively small effective populations (mean N(e) = 62) may have caused the observed contemporary differentiation. Novel Bayesian analyses indicated mean contributions of 71% F0 population hybrids in spawning populations, of which 76% had maternal or paternal native origin. CONCLUSIONS: Ecological connectivity between lakes separated by a dam has been retained through construction of fishways for feeding migration. Considerable survival and homing to ancestral spawning sites in hybrid progeny was documented. Population differentiation despite preceding admixture is likely caused by contemporary reduced reproductive fitness of population hybrids. The study documents the beginning stages of population divergence among spatial aggregations with recent common ancestry.

Identification of differentially expressed genes associated with osteochondrosis in standardbred horses using RNA arbitrarily primed PCR.

The aim of this study was to investigate genes for differential expression in cartilage of foals predisposed to osteochondrosis (OC). Tissue was sampled from the cranial part of the distal intermediate ridge of the tibia in the tarso-crural joint. Foals were considered predisposed to OC when parents had OC at the distal intermediate ridge of the tibia. RNA was isolated and subjected to arbitrarily primed PCR (RAP-PCR) followed by fingerprinting to screen for differentially expressed genes. By verification of results from the RAP-PCR fingerprint screening using real-time RT-PCR, we identified two genes not previously correlated with OC as differentially expressed. The two genes, which were identical to TLK2 and an equine EST, are good targets for future research on OC.

Stabilizing selection and adaptive evolution in a combination of two traits in an arctic ungulate.

Stabilizing selection is thought to be common in wild populations and act as one of the main evolutionary mechanisms, which constrain phenotypic variation. When multiple traits interact to create a combined phenotype, correlational selection may be an important process driving adaptive evolution. Here, we report on phenotypic selection and evolutionary changes in two natal traits in a semidomestic population of reindeer (Rangifer tarandus) in northern Finland. The population has been closely monitored since 1969, and detailed data have been collected on individuals since they were born. Over the length of the study period (1969-2015), we found directional and stabilizing selection toward a combination of earlier birth date and heavier birth mass with an intermediate optimum along the major axis of the selection surface. In addition, we demonstrate significant changes in mean traits toward earlier birth date and heavier birth mass, with corresponding genetic changes in breeding values during the study period. Our results demonstrate evolutionary changes in a combination of two traits, which agree closely with estimated patterns of phenotypic selection. Knowledge of the selective surface for combinations of genetically correlated traits are vital to predict how population mean phenotypes and fitness are affected when environments change.