A method of analyzing cultivar x location x year experiments: a new stability parameter

Summary Assessment of cultivar performance in a cultivar x location x year experiment is often difficult because of the presence of a location x year interaction. Our objective is to demonstrate a method on separation of environment effects (location x year) into predictable and unpredictabel components. The analysis consists of two parts: (1) a regression analysis based on location effects (averaged over years), assuming that the location means represent predictable environmental variation; and (2) the estimation of stability (denoted type 4) based on the years within location mean squares, assuming that years within location represent unpredictable environmental variation. From the regression analysis in (1), a breeder can determine the optimum range of locations in which a cultivar is well suited, and from (2) he can choose the most stable cultivars. The advantage of type 4 stability is that it is independent of the other cultivars included in the test and of the regression coefficient estimated for predictable variation. Three sets of published data are used to illustrate the analysis. Type 4 stability is compared with type 3 stability (deviation mean square from regression on environmental index) for genetic consistency. The analyses suggest that type 4 stability is consistent and is therefore a potential genetic parameter, but type 3 stability is not.Contribution No. I-806 from Engineering and Statistical Research Centre, Research Branch, Agriculture Canada, Ottawa, K1A OC6, Canada     

Assessment of a method for cultivar selection based on regional trial data

Summary In this study a method for analyzing regional trial data is investigated for its effectiveness in cultivar selection. The method is a synthesis of three procedures: (1) regression analysis for genotype × environment (GE) interaction, and subsequent cluster analysis for grouping cultivars for similarity of response; (2) superiority measure analysis of cultivar performance based on the distance mean square between the test cultivar and the maximum response; (3) type 4 stability analysis for three-way classification data (cultivar × location × year), to measure a cultivar's stability. Each of these three procedures is aimed at different aspects of the selection problem: the first obtains an overview of the types of cultivar response; the second measures a cultivar's general adaptability within the region; the third assesses a cultivar's ability to withstand unpredictable variation, namely that caused by year effects. Four sets of published data, each originally analyzed by a univariate or a multivariate approach, were used as examples. The results suggest that the present method provides direct and useful information for selection purposes. The superiority measure, which is the core of the method, can be used even if the data do not fit the linear model for GE interaction. Plotting the data with a fixed standard represented by the maximum response provides a simple visual tool for identifying environments where a cultivar performs well.Contribution No. C-035 from Research Program Service, Research Branch, Agriculture Canada, Ottawa, Ontario, K1A 0C6     

Genetic properties of four types of stability parameter

Summary The genetic properties of four types of stability parameter for individual genotypes were investigated using a set of diallel cross data (28 genotypes x four locations x 3 years). The specific parameters studied were: the variance of a genotype across environments (T1); the genotype x environment (GE) interaction effect for a genotype, squared and summed across all environments (T2); the residual mean square (MS) of deviations from the regression of a genotype on an environmental index (T3); and years within locations MS for a genotype, averaged over all locations (T4). Each stability statistic was fitted to the additive model, based on the assumption that if the stability parameter is heritable, stability of F₁ is most likely to be the average stability of its parents. The results showed that T1 and T4 were additive, but T2 and T3 were not. A study of the consistency of stability rankings between two seeding rates over the same set of environments showed a similar pattern. It appears that stability parameters of types 1 and 4 are heritable, and thus useful for selection, while those of types 2 and 3 are nonheritable, and thus not useful.Contribution No. R-029     

A comparative evaluation of two methods of selecting locations used for testing spring wheat cultivars

Summary One of the considerations of regional cultivar evaluation programs is to optimize the number of locations used for testing. Although optimization of numbers of locations using cluster analysis has been previously attempted, no objective comparison of methods has yet been made. A new clustering method that uses the pairwise contribution of locations to the cultivar x location mean square as the distance measure (LB) was compared to another method that employs diallel correlations as the distance measure (CL). Data from six spring wheat (Triticum aestivum L. em Thell) cultivars grown at 13 locations for five years were used in the initial cluster analysis. Another set of data, from a separate year, consisting of yields of the original 6 cultivars and a set of 12 independent cultivars was then used to check the validity of the original groupings and to compare the two methods. When the 6 original cultivars were considered, the LB technique was found to be superior to the CL. When the 12 independent cultivars were used, neither method was considered to be superior. Because of the lack of flexibility on the part of the LB method, neither technique could be deemed as fully adequate.Contribution Number 842 of the Department of Plant Science, University of Manitoba, Winnipeg, Manitoba, Canada     

Cultivar x environment interactions in sorghum,Sorghum bicolor (L.) moench

Summary The most significant and important interactions in 2, and 3-year sorghum zonal trials were location x year, cultivar x location and cultivar x location x year. Cultivar x year interaction was significant in one out of four ecologic zones used in the trials. The variance components were relatively small with the second order interaction variance component larger than the first order types involving cultivars. Environmental variance was the largest in all four zones.Computations on theoretical standard error of cultivar means suggest that six replications, eight locations and four years is an ideal testing procedure in the Northern Guinea Savanna, while in the Sudan Savanna, the combination is better with four replications, eight locations and three years. The true value or performance of a variety is most effectively obtained by increasing the number of years, while increasing number of replications is the least effective.     

Comparison of unpredictable environmental variation generated by year and by seeding-time factors for measuring type 4 stability

Summary Type 4 stability has been proposed to measure a cultivar's homeostatic property to resist unpredictable environmental variation. The requirement for calculating this stability is that the experiment must contain a time factor in addition to the cultivar x location factors. Of the two time factors, year and seeding-time, the latter is less attractive biologically because it represents only a part of the broader context of unpredictable variation represented by years, but it is attractive in terms of shortening the test period. Investigation of historical data from the Eastern Cooperative trials and the Ontario Production trials in Canada indicates that the unpredictable variation generated by seeding-time was about half that generated by year. Although type 4 stability measured by both factors appears to be the same, the stability measured by seeding-time is more prone to variation. The implication is that complete substitution of the year factor by seeding-time is not appropriate, but use of both factors in combination may be sensible.Contribution No. C-055 from the Engineering and Statistical Research Centre, Research Branch, Agriculture Canada, Ottawa, K1A 0C6, Canada     

作物品种区域试验试点区辨力评价方法的比较(英文)

试点区辨力是作物品种区域试验试点选择的重要依据,可以用不同的方法进行评价。本研究分别对西南玉米区域试验2003和2004两年的产量数据进行联合方差分析,然后用变异系数(CV)、遗传变异系数(GCV)、环境区分指数(DI)、B型生态价(WE)、回归系数(RC)、基于AM MI分析结果的试点互作主成分点到原点的距离(DA)、AMMI稳定性值(ASV)和基于GGE双标图分析的试点到原点的距离(DG)对西南玉米区域试验试点的区辨力进行评价,进而用秩次相关分析和主成分分析揭示试点区辨力评价方法间的内在关系。结果表明,两年试验品种间、试点间均存在极显著差异,且品种和试点间互作也极显著;8个统计数也许可以用来度量试点的区辨力,但是不同统计数对同一年度内试点的区辨力评价结果不完全一致;统计数CV、GCV、WE、DA和DG间都存在极显著的秩次相关,但是统计数DG也许更有利于评价试点的区辨力。     

Metabolite profiling of maize grain: differentiation due to genetics and environment

A comparative metabolite profiling approach based on gas chromatography-mass spectrometry (GC/MS) was applied to investigate the impact of genetic background, growing location and season on the chemical composition of maize grain. The metabolite profiling protocol involved sub-fractionation of the metabolites and allowed the assessment of about 300 distinct analytes from different chemical classes (polar to lipophilic), of which 167 could be identified. A comparison, over three consecutive growing seasons, of the metabolite profiles of four maize cultivars which differed in their maturity classification, was carried out using principal component analysis (PCA). This revealed a strong separation of one cultivar in the first growing season, which could be explained by the immaturity of the kernels of this cultivar compared with others in the field trial. Further evaluations by pair-wise comparison using Student’s t-test and analysis of variance (ANOVA) showed that the growing season was the most prominent impact factor driving variation of the metabolite pool. An increased understanding of metabolic variation was achieved by analysis of a second sample set comprising one cultivar grown for 3 years at four locations. The applied GC/MS-based metabolite profiling demonstrated the natural variation in maize grain metabolite pools resulting from the interplay of environment, season, and genotype.     

An integrated approach to oilseed rape cultivar selection using phenotypic stability

Summary Various methods of evaluating phenotypic stability have been proposed; however, no single method can adequately describe cultivar performance. The objectives of this study were to integrate a number of methods of evaluating stability and to use this approach for cultivar selection. These objectives were considered in the context of the broad-based oilseed rape cultivar (Brassica napus spp. oleifera) evaluation system currently used in western Canada. Regression analysis was used to assess cultivar response to environments. Cluster analysis was used to assemble cultivars into groups with similar regression coefficients (b _i ) and mean yield. Three parametric stability parameters, years within locations mean square (MS; Y/L), Shukla's stability variance ( _i ² ), and Francis and Kannenberg's coefficient of variability (CV _i ), were compared to determine which method would be most suitable for selection of oilseed rape cultivars from within clustered groups. Yield data from three cultivars and six breeding lines that had been tested for 2 years at 26 locations in the Western Canola Cooperative Test A were used for all calculations. The cluster analysis was successful in identifying commercially acceptable breeding lines. The parameter MS _i Y/L was considered to be more appropriate than either CV _i or _i ² , because it measured only the unpredictable portion of the genotype x environment interaction and was independent of the other cultivars in the test. The use of cluster analysis to group entries with similar b _i values and mean yields, followed by selection for stability within groups, was advocated.Contribution No. 846 of the Plant Science Department, University of Manitoba     

Grouping genotypes by a cluster method directly related to genotype-environment interaction mean square

Summary To facilitate the interpretation of data from a genotype by environment (GE) experiment when the GE interaction is large, a cluster method is proposed to group genotypics according to their response to the environments. The dissimilarity index between a pair of genotypes is defined in terms of distance adjusted for the average effects of genotypes, and Sokal and Michener's (1958) unweighted pair-group method is used in the clustering algorithm. The new index, constructed in each cluster cycle for any group, is shown to be equivalent to within group GE interaction mean square under 2-way ANOVA. Thus, if the F-value is used as an empirical stopping criterion for clustering, there will be no significant GE interaction within groups and the genotypes within the groups can be compared by their average effects. The method of analysis is illustrated by an example.Contribution no. I-348 from the Engineering and Statistical Research Institute     

Environmental response patterns in commercial classes of common bean (Phaseolus vulgaris L.)

Summary The yield data of 39 cultivars of diverse commercial classes of beans (Phaseolus vulgaris L.) planted in seven locations in Michigan were subjected to cluster and canonical variate analyses. The essential findings and conclusions can be summarized as follows: (1) Cluster analysis classified the cultivars into sub-sets or clusters almost identically coinciding with their commercial class designation. Canonical variate analysis completely confirmed the sub-groupings. Within class similarities were attributed to a narrow genetic base resulting from a common genetic relationship, or at least sharing of a common gene pool. (2) It was found that two clusters could possess almost identical mean (cluster mean) yields, and deviate in opposite directions over the same range of environments. (3) When total genotype × environmental interaction variance was partitioned into between and within clusters, the cluster × environment portion constituted 80% of the total. (4) These results imply that if the behavior of a given cultivar across a series of environments is known, the behavior of all other members of the class across a similar range of environments would be predictable.Journal Article No. 10329 of the Michigan Agricultural Experiment Station     

Multitrait selection system using populations with a small number of interploid (4x-2x) hybrid seedlings in potato: Degree of high-parent heterosis for yield and frequency of clones combining quantitative agronomic traits

One of the most attractive features of the 4x-2x breeding scheme via unilateral sexual polyploidization (USP) is the possibility of obtaining heterotic clones with a combination of desirable traits based upon selection in a small number of hybrid seedlings. A set of experiments was carried out to verify this characteristic of the USP scheme using 42 families (with 20 plants each) derived from 2x Phureja-haploid Tuberosum or haploid Tuberosum-Solanum chacoense hybrids as male parents. These clones were 2n-pol-len-producers by either first-division restitution with crossing over (FDR-CO) or without crossing over (FDR-NCO). The 4x parents were eight cultivars of USA and European origin. A total of 168 out 840 clones was initially selected (four clones/family) based on general tuber appearance (GTA). An additional round of selection (not taking into account the parentage of the clone) for total tuber yield (TTY) reduced the sample to 96 clones. These selected clones and the original 4x parents were evaluated in two locations at Wisconsin (Hancock – E?1, and Rhinelander – E?2). The average high-parent heterosis values for TTY ranged from 27.8% (E?2) to 48% (E?1). The population of experimental clones also had a TTY range wider than that of the 4x parents. The best clone gave a yield of 101.6% and 63.2% over the best 4x parent at E?1 and E?2, respectively. For TTY, a significant genotype×environment (G×E) interaction for the experimental clones was found when a combined analysis of variance for both locations was carried out. However, the G×E interaction was not significant when only the 4x-parent group was considered. Clones derived from 4x European cultivars had higher GTA scores than clones derived from 4x USA cultivars. With two culling levels being set on the TTY≥4x parent group mean and the GTA≥three, about 56% (E?1) and 48% (E?2) of the clones would be retained for further evaluation. These percentages of selected clones are much higher than those reported using conventional 4x-4x crosses. Our results indicate that a USP strategy with 4x-2x (FDR) crosses would be more effective than intra-Tuberosum crosses in generating heterotic clones with a combination of desirable quantitative traits using populations with a small number of hybrid seedlings.     

The effect of single versus multiple sets on strength

Research has previously been divided on whether performing resistance training with a single set per training session is as effective for increasing strength as training with multiple sets. The purpose of this study was to determine the effect of single sets versus multiple sets on strength. Forty subjects were randomly assigned into 1 of 3 groups: control (C; n = 8), single set (SS; n = 14), or multiple sets (MS; n = 18) to perform 8 maximal knee extensions at 60 degrees .s(-1) on a Biodex System 3 isokinetic dynamometer twice a week for 8 weeks. The SS group performed 1 set while the MS group performed 3 sets. All groups were pre-, mid- (4 weeks), and posttested at 60 degrees x s(-1). Strength was expressed as peak torque (PT). A 3 x 3 x 2 (time x group x sex) mixed factor repeated measures analysis of variance (ANOVA) revealed no interaction involving sex, but there was an interaction of group by time. The MS group exhibited a significant (p < 0.05) increase in PT (pre = 171.39 +/- 61.98 Nm; mid = 193.08 +/- 66.23 Nm) between the pretest and the midtest while the SS (pre = 163.45 +/- 56.37 Nm; mid = 172.60 +/- 61.78 Nm) and C groups (pre = 135.997 +/- 54.31 Nm; mid = 127.66 +/- 53.12 Nm) did not change. Strength did not change between the midtest and the posttest for any group. It was concluded that performing 3 sets of isokinetic knee extensions was more effective than performing a single set for increasing peak torque. These results seem to indicate that for increasing strength of the quadriceps, performing multiple sets is superior to performing a single set of resistance exercise.     

Use of the additive main effects and multiplicative interaction model in QTL mapping for adaptation in barley

The additive main effects and multiplicative interaction (AMMI) model has emerged as a powerful analytical tool for genotype x environment studies. The objective of the present study was to assess its value in quantitative trait locus (QTL) mapping. This was done through the analysis of a large two-way table of genotype-by-environment data of barley (Hordeum vulgare L.) grain yields, where the genotypes constituted a genetic population suitable for mapping studies. Grain yield data of 150 doubled haploid lines derived from the Steptoe x Morex cross, and the two parental lines, were taken by the North American Barley Genome Mapping Project (NABGMP) at 16 environments throughout the barley production areas of the USA and Canada. Four regions of the genome were responsible for most of the differential genotypic expression across environments. They accounted for approximately 50% of the genotypic main effect and 30% of the genotype x environment interaction (GE) sums of squares. The magnitude and sign of AMMI scores for genotypes and sites facilitate inferences about specific interactions. The parallel use of classification (cluster analysis of environments) and ordination (principal component analysis of GE matrix) techniques allowed most of the variation present in the genotype x environment matrix to be summarized in just a few dimensions, specifically four QTLs showing differential adaptation to four clusters of environments. Thus, AMMI genotypic scores, when the genotypes constituted a population suitable for QTL mapping, could provide an adequate way of resolving the magnitude and nature of QTL x environment interactions.Ignacio Romagosa was on sabbatical leave from the University of Lleida and the Institut de Recerca i Tecnologia Agroalimentàries, Lleida, Spain, when this study was conducted     

Prediction of protein subcellular locations by support vector machines using compositions of amino acids and amino acid pairs

MOTIVATION: The subcellular location of a protein is closely correlated to its function. Thus, computational prediction of subcellular locations from the amino acid sequence information would help annotation and functional prediction of protein coding genes in complete genomes. We have developed a method based on support vector machines (SVMs). RESULTS: We considered 12 subcellular locations in eukaryotic cells: chloroplast, cytoplasm, cytoskeleton, endoplasmic reticulum, extracellular medium, Golgi apparatus, lysosome, mitochondrion, nucleus, peroxisome, plasma membrane, and vacuole. We constructed a data set of proteins with known locations from the SWISS-PROT database. A set of SVMs was trained to predict the subcellular location of a given protein based on its amino acid, amino acid pair, and gapped amino acid pair compositions. The predictors based on these different compositions were then combined using a voting scheme. Results obtained through 5-fold cross-validation tests showed an improvement in prediction accuracy over the algorithm based on the amino acid composition only. This prediction method is available via the Internet.     

Genetic diversity among barley cultivars assessed by sequence-specific amplification polymorphism

We analyzed the genetic structure and relationships among barley cultivars (Hordeum vulgare L.) with sequence-specific amplification polymorphisms (S-SAPs). Polymorphisms were identified in 824 individual barley plants representing 103 cultivars (eight plants per cultivar) widely grown in Canada and the United States, using PCR primers designed from the long terminal repeat of the barley retrotransposon BARE-1 and a subset of four selective MseI primers. From the 404 bands scored, 150 were polymorphic either within or between cultivars. Genetic structure assessed with analysis of molecular variance attributed the largest component of variation to the within groups of cultivars (69–86%). Within-cultivar genetic variation was estimated as average gene diversity over loci and ranged from 0 (completely homogenous) to 0.076 (most heterogeneous cultivar). Only 17 out of 103 cultivars (16%) were judged to be homogenous by this criterion. Relationships among cultivars were analyzed by cluster analysis using unweighted pair-groups using arithmetic averages and found groups similar to those determined by agriculturally significant phenotypic traits such as spike morphology (two-rowed or six-rowed), cultivar type (malting or feed), seed characteristic (hull-less or hulled), and growth habit (winter or spring), with minor overlaps. Discriminant analysis of groups determined by these phenotypic traits fully supported the different groups with minor overlaps between the malting/feed. S-SAP markers generated from retrotransposons such as BARE-1 are invaluable tools for the study of genetic diversity in organisms with a narrow genetic base such as barley. In this study, S-SAP analysis revealed significant amounts of cryptic variation in closely related cultivars including somaclonal variation, which could not be inferred by the pedigree analysis.     

Density and growth rates of spider mites in relation to phenological stages of soybean cultivars in Brazil

The population fluctuations of spider mites on different soybean cultivars were studied in two experiments performed in the municipalities of São Sepé (experiment 1) and Santa Maria (experiment 2) in the state of Rio Grande do Sul, Brazil. The number of cultivars was 20 in São Sepé and 25 in Santa Maria, and at each location leaves were sampled from 15 plants per cultivar every week throughout the entire plant developmental cycle, amounting to 11 and 10 samplings in experiment 1 and 2, respectively. The statistical analysis revealed that mite densities varied significantly with cultivar and time. Besides, the interaction between soybean cultivars and sampling times was significant at both locations. This indicates that the populations did not vary in the same way among cultivars over time, which is attributed to differences among the cultivars with respect to their phenology and response to spider mites. Equations for describing the mite population dynamics were estimated for each cultivar, mostly by fitting cubic equations to the weekly growth rates (per capita changes) in mite densities. We also found that the highest growth rates for mites on soybean cultivars in both locations occurred after the beginning of flowering, when the plants shifted from the vegetative to reproductive stages, about 3 weeks before reaching the peak densities.     

Effect of population structure on protein-yield improvements in spring wheat (Triticum aestivum L. em Thell)

Summary In a study designed to develop a more efficient breeding method for concurrent protein-yield improvements in wheat (Triticum aestivum L. em. Thell), 7 base populations [2 F₂'s, 1 intermated F₂ (IF₂) and 4 partial backcross (PBC) populations] developed from biparental crosses involving 2 Canadian hard red spring (CHRS) and 2 Canadian utility (CU) wheat cultivars were evaluated in Winnipeg, Manitoba, Canada. The IF₂ and PBC populations were generated for comparison with conventional F₂ populations and to determine which of the 4 methods of population development would provide a more efficient means of producing potentially superior genetic recombinants. Parameters pertaining to means, variances, correlations, heritabilities and frequencies of desirable and undesirable progenies were used to evaluate the limitations to genetic gain that may be expected from selection for GY and GPC in F₂, IF₂, CHRS-PBC and CU-PBC populations. Analysis of protein and yield data from 105 S₁ lines derived from each of the 7 populations showed the CU-PBC's to have the highest grain yield (GY) and the lowest grain protein concentration (GPC) means; and the CHRS-PBC's, the lowest GY and the highest GPC means. The F₂ and IF₂ populations were intermediate for both characteristics. Populations developed from the same biparental cross did not differ significantly with respect to the majority of genetic parameters. However, desirable progenies combining high GY with high GPC were more frequent in the CU-PBC, and least frequent in the CHRS-PBC populations. The observed superiority of the CU-PBC populations appeared to be related to the advantage the system has in preserving the genetic integrity of a proven cultivar, while adding desirable genetic factors from another cultivar, thus capitalizing on introgression and upgrading simultaneously.Contribution No. 549 from Agriculture Canada, Lacombe Research Station. Research was supported by a grant from the Natural Sciences and Engineering Research Council of Canada     

Isolate Virulence and Cultivar Response in the Winter Wheat: Pyrenophora tritici-repentis (Tan Spot) Pathosystem in Oklahoma

Prevalence of tan spot of wheat caused by the fungus Pyrenophora tritici-repentis has become more prevalent in Oklahoma as no-till cultivation in wheat has increased. Hence, developing wheat varieties resistant to tan spot has been emphasized, and selecting pathogen isolates to screen for resistance to this disease is critical. Twelve isolates of P. tritici-repentis were used to inoculate 11 wheat cultivars in a greenhouse study in split-plot experiments. Virulence of isolates and cultivar resistance were measured in percent leaf area infection for all possible isolate x cultivar interactions. Isolates differed significantly (P < 0.01) in virulence on wheat cultivars, and cultivars differed significantly in disease reaction to isolates. Increased virulence of isolates detected increased variability in cultivar response (percent leaf area infection) (r = 0.56, P < 0.05) while increased susceptibility in cultivars detected increased variance in virulence of the isolates (r = 0.76, P < 0.01). A significant isolate × cultivar interaction indicated specificity between isolates and cultivars, however, cluster analysis indicated low to moderate physiological specialization. Similarity in wheat cultivars in response to pathogen isolates also was determined by cluster analysis. The use of diverse isolates of the fungus would facilitate evaluation of resistance in wheat cultivars to tan spot.     

Genetic structure among continental and island populations of gyrfalcons

Little is known about the possible influence that past glacial events have had on the phylogeography and population structure of avian predators in the Arctic and sub-Arctic. In this study, we use microsatellite and mitochondrial control region DNA variation to investigate the population genetic structure of gyrfalcons (Falco rusticolus) throughout a large portion of their circumpolar distribution. In most locations sampled, the mtDNA data revealed little geographic structure; however, five out of eight mtDNA haplotypes were unique to a particular geographic area (Greenland, Iceland, or Alaska) and the Iceland population differed from others based on haplotype frequency differences (F(ST)). With the microsatellite results, significant population structure (F(ST), principal components analysis, and cluster analysis) was observed identifying Greenland and Iceland as separate populations, while Norway, Alaska and Canada were identified as a single population consistent with contemporary gene flow across Russia. Within Greenland, differing levels of gene flow between western and eastern sampling locations was indicated with apparent asymmetric dispersal in western Greenland from north to south. This dispersal bias is in agreement with the distribution of plumage colour variants with white gyrfalcons in much higher proportion in northern Greenland. Lastly, because the mtDNA control region sequence differed by only one to four nucleotides from a common haplotype among all gyrfalcons, we infer that the observed microsatellite population genetic structure has developed since the last glacial maximum. This conclusion is further supported by our finding that a closely related species, the saker falcon (Falco cherrug), has greater genetic heterogeneity, including mtDNA haplotypes differing by 1-16 nucleotide substitutions from a common gyrfalcon haplotype. This is consistent with gyrfalcons having expanded rapidly from a single glacial-age refugium to their current circumpolar distribution. Additional sampling of gyrfalcons from Fennoscandia and Russia throughout Siberia is necessary to test putative gene flow between Norway and Alaska and Canada as suggested by this study.