White Spruce Populations Research Articles

Contrasting physiological strategies explain heterogeneous responses to severe drought conditions within local populations of a widespread conifer

Understanding how trees prioritize carbon gain at the cost of drought vulnerability under severe drought conditions is crucial for predicting which genetic groups and individuals will be resilient to future climate conditions. In this study, we investigated variations in growth, tree-ring anatomy as well as carbon and oxygen isotope ratios to assess the sensitivity and the xylem formation process in response to an episode of severe drought in 29 mature white spruce (Picea glauca [Moench] Voss) families grown in a common garden trial. During the drought episode, the majority of families displayed decreased growth and exhibited either sustained or increased intrinsic water-use efficiency (iWUE), which was largely influenced by reduced stomatal conductance as revealed by the dual carbon‑oxygen isotope approach. Different water-use strategies were detected within white spruce populations in response to drought conditions. Our results revealed intraspecific variation in the prevailing physiological mechanisms underlying drought response within and among populations of Picea glauca. The presence of different genetic groups reflecting diverse water-use strategies within this largely-distributed conifer is likely to lessen the negative effects of drought and decrease the overall forest ecosystems' sensitivity to it.

The dynamics of a changing Lutz spruce (Picea×lutzii) hybrid zone on the Kenai Peninsula, Alaska

We investigated the genetic makeup of Lutz spruce, a natural hybrid between white and Sitka spruce on the Kenai Peninsula, Alaska. Microsatellites indicate 72% of individuals sampled had predominantly white spruce ancestry, whereas 14% had predominantly Sitka spruce ancestry; some individuals classified as white spruce had Sitka spruce plastid genotypes. As Picea mitochondria are maternally inherited and plastids are paternally inherited, it appears that white spruce was the ancestral seed parent of nearly all spruce on the western peninsula, whereas Sitka spruce alleles originated from pollen. Pollen records show that white spruce colonized the western peninsula ∼8500 YBP from glacial refugium, whereas Sitka spruce arrived on the eastern peninsula ∼4000 YBP after migrating up the Pacific coast. Our data suggest that Sitka spruce colonization west of the Kenai Mountains may have occurred not via seed dispersal but by long distance transport of wind-borne pollen and subsequent hybridization with established white spruce populations. Hybridization was an important mechanism that allowed Sitka spruce to expand the leading edge of its range in response to historical climate change. As the climate continues to warm, climate envelope modeling suggests Lutz spruce may ultimately displace white spruce on the western peninsula even as Sitka spruce is constrained to the eastern peninsula where it will continue to hybridize.

Genetic basis of growth reaction to drought stress differs in contrasting high-latitude treeline ecotones of a widespread conifer.

Climate change is increasing the frequency and intensity of drought events in many boreal forests. Trees are sessile organisms with a long generation time, which makes them vulnerable to fast climate change and hinders fast adaptations. Therefore, it is important to know how forests cope with drought stress and to explore the genetic basis of these reactions. We investigated three natural populations of white spruce (Picea glauca) in Alaska, located at one drought-limited and two cold-limited treelines with a paired plot design of one forest and one treeline plot. We obtained individual increment cores from 458 trees and climate data to assess dendrophenotypes, in particular the growth reaction to drought stress. To explore the genetic basis of these dendrophenotypes, we genotyped the individual trees at 3000 single nucleotide polymorphisms in candidate genes and performed genotype-phenotype association analysis using linear mixed models and Bayesian sparse linear mixed models. Growth reaction to drought stress differed in contrasting treeline populations. Therefore, the populations are likely to be unevenly affected by climate change. We identified 40 genes associated with dendrophenotypic traits that differed among the treeline populations. Most genes were identified in the drought-limited site, indicating comparatively strong selection pressure of drought-tolerant phenotypes. Contrasting patterns of drought-associated genes among sampled sites and in comparison to Canadian populations in a previous study suggest that drought adaptation acts on a local scale. Our results highlight genes that are associated with wood traits which in turn are critical for the establishment and persistence of future forests under climate change.

Editorial 2022

Editorial 2022

Population structure and the influence of microenvironment and genetic similarity on individual growth at Alaskan white spruce treelines

Knowledge on the adaptation of trees to rapid environmental changes is essential to preserve forests and their ecosystem services under climate change. Treeline populations are particularly suitable for studying adaptation processes in trees, as environmental stress together with reduced gene flow can enhance local adaptation. We investigated white spruce (Picea glauca) populations in Alaska on one moisture-limited and two cold-limited treeline sites with a paired plot design of one forest and one treeline population each, resulting in six plots. Additionally, one forest plot in the middle of the distribution range complements the study design. We combined spatial, climatic and dendrochronological data with neutral genetic marker of 2203 trees to investigate population genetic structure and drivers of tree growth. We used several individual-based approaches including random slope mixed-effects models to test the influence of genetic similarity and microenvironment on growth performance. A high degree of genetic diversity was found within each of the seven plots associated with high rates of gene flow. We discovered a low genetic differentiation between the three sites which was better explained by geographic distances than by environmental differences, indicating genetic drift as the main driver of population differentiation. Our findings indicated that microenvironmental features had an overall larger influence on growth performances than genetic similarity among individuals. The effects of climate on growth differed between sites but were smaller than the effect of tree size. Overall, our results suggest that the high genetic diversity of white spruce may result in a wider range of phenotypes which enhances the efficiency of selection when the species is facing rapid climatic changes. In addition, the large intra-individual variability in growth responses may indicate the high phenotypic plasticity of white spruce which can buffer short-term environmental changes and, thus, allow enduring the present changing climate conditions.

Moisture deficits limit growth of white spruce in the west-central boreal forest of North America

Climate change may cause reduced forest productivity and higher tree mortality due to water deficits that result from increased evapotranspiration. Such limitations may occur in some areas of the North American boreal forest where precipitation is low and warming trends are high. This paper analyzes climatic factors that limit the growth of a commercially important and widespread boreal tree species, white spruce (Picea glauca (Moench) Voss), based on a range-wide dendroclimatological analysis of 9795 trees from 227 sample sites across the North American boreal forest. A bootstrapped response function analysis was conducted for monthly temperature and precipitation variables, and a multivariate regression tree analysis was used to group white spruce populations with similar response coefficients where climatic factors explained 46% of the total variance in response coefficients. The results of this study show that white spruce populations in the west-central boreal forest of North America are the most precipitation-limited group and therefore likely to be most susceptible to climate change. Populations both to the north and south of this group appear less vulnerable, and eastern populations generally do not experience growth-limiting moisture deficits. Regional climate change projections suggest that moisture limitations will become more pronounced in the future, especially for southwest and west-central boreal tree populations.

Adaptive limitations of white spruce populations to drought imply vulnerability to climate change in its western range.

A cost‐effective climate change adaptation strategy for the forestry sector is to move seed sources to more northern and higher elevation planting sites as part of ongoing reforestation programs. This is meant to match locally adapted populations with anticipated environments, but adaptive traits do not always show population differences suitable to mitigate climate change impacts. For white spruce, drought tolerance is a critical adaptive trait to prevent mortality and productivity losses. Here, we use a 40‐year‐old provenance experiment that has been exposed to severe drought periods in 1999 and 2002 to retrospectively investigate drought response and the adaptive capacity of white spruce populations across their boreal range. Relying on dendrochronological analysis under experimentally controlled environments, we evaluate population differences in resistance, resilience, and recovery to these extreme events. Results showed evidence for population differentiation in resistance and recovery parameters, but provenances conformed to approximately the same growth rates under drought conditions and had similar resilience metrics. The lack of populations with better growth rates under drought conditions is contrary to expectations for a wide‐ranging species with distinct regional climates. Populations from the wettest environments in the northeastern boreal were surprisingly drought‐tolerant, suggesting that these populations would readily resist water deficits projected for the 2080s, and supporting the view that northeastern Canada will provide a refugium for boreal species under climate change. The findings also suggest that white spruce is sensitive to growth reductions under climate change in the western boreal. The study highlights that population differentiation in adaptive capacity is species‐ and trait‐specific, and we provide a counterexample for drought tolerance traits, where assisted migration prescriptions may be ineffective to mitigate climate change impacts. For resource managers and policy makers, we provide maps where planning for widespread declines of boreal white spruce forests may be unavoidable.

Survival, growth and cold hardiness tradeoffs in white spruce populations: Implications for assisted migration

Human assisted movement of reforestation stock poleward or upward in elevation has been proposed as a tool to address climate change in regular reforestation programs. However, moving warm adapted seed sources to colder environments could carry the risk of frost damage if seed sources are moved too far. Here, we assess genetic differentiation in growth potential, survival and cold hardiness of a wide-ranging tree species, white spruce (Picea glauca [Moench] Voss). We use data from a 34-year-old common garden experiment planted in the approximate center of the species range. Cold hardiness was negatively related to growth and positively to survival. Generalized additive models identified mean coldest month temperature and latitude (as a proxy for the day length regime) as an exceptionally good predictor for the onset of cold hardiness (R2adj = 0.91). The results suggest that day length, an environmental factor that is not influenced by climate change, is an important factor controlling the timing of the onset of hardiness. Survival was only moderately well predicted, primarily by precipitation of the provenance origin (R2adj = 0.36) indicating that other adaptive traits besides cold hardiness should be considered in assisted migration prescriptions. Survival of seed sources appears primarily compromised when transplanting sources from wet origins to a dry location. Nevertheless, acceptable migration distances without significant tradeoffs were up to 500 km north and 1500 km west towards a central planting location.

Restructuring tree provenance test data to conform to reciprocal transplant experiments for detecting local adaptation

Summary Local adaptation is a fundamental assumption in delineating seed zones and developing seed transfer guidelines to safeguard climatic adaptation of tree and plant species during forest regeneration and ecological restoration. It is also broadly assumed for forest tree species that show genetic differentiation among geographic populations, especially for those with widespread natural distributions that occur in distinct environments. However, due to a scarcity of suitable data, the inference of local adaptation has rarely been validated for forest tree species through reciprocal transplant experiments (RTEs). In this study, we illustrate a novel approach to restructure tree provenance test data to conform to RTEs and use recently proposed statistical models to detect local adaptation, using white spruce Picea glauca (Moench) Voss as an example. Our research demonstrates how similar studies can be conducted to validate local adaptation in other forest species and populations, for which RTEs are lacking, but where large, high‐quality provenance test data sets are available. Contrary to common belief, our results show that local adaptation is absent in survival and height for white spruce populations from Ontario, Canada, although they have evolved in considerably different climatic habitats, and exhibit substantial and clinal genetic differentiation and significant genotype‐by‐environment interactions in these two adaptive traits. Synthesis and applications. Our results show that the common assumption of local adaptation in forest tree species may not necessarily be correct within significant portions of their natural range. In forest genetic studies, population differentiation in adaptive traits has often been attributed to local adaptation without rigorous validation. Our results caution against such interpretation of experimental results. In the absence of local adaptation as shown by reciprocal transplant experiments, assisted migration may be considered as an option for enhancing forest adaptation to climate change.

Fine-scale geographic variation in photosynthetic-related traits of Picea glauca seedlings indicates local adaptation to climate.

Climate-related variations in functional traits of boreal tree species can result both from physiological acclimation and genetic adaptation of local populations to their biophysical environment. To improve our understanding and prediction of the physiological and growth responses of populations to climate change, we studied the role of climate of seed origin in determining variations in functional traits and its implications for tree improvement programs for a commonly reforested boreal conifer, white spruce (Picea glauca (Moench) Voss). We evaluated growth, root-to-shoot ratio (R/S), specific leaf area (SLA), needle nitrogen (N(mass)), total non-structural carbohydrates (NSC) and photosynthetic traits of 3-year-old seedlings in a greenhouse experiment using seed from six seed orchards (SO) representing the different regions where white spruce is reforested in Québec. Height and total dry mass (TDM) were positively correlated with photosynthetic capacity (A(max)), stomatal conductance (g(s)) and mesophyll conductance (g(m)). Total dry mass, but not height growth, was strongly correlated with latitude of seed origin (SO) and associated climate variables. A(max), g(s), g(m) and more marginally, photosynthetic nitrogen-use efficiency (PNUE) were positively associated with the mean July temperature of the SO, while water use efficiency (WUE) was negatively associated. Maximum rates of carboxylation (V(cmax)), maximum rates of electron transport (J(max)), SLA, N(mass), NSC and R/S showed no pattern. Our results did not demonstrate a higher Amax for northern seed orchards, although this has been previously hypothesized as an adaptation mechanism for maintaining carbon uptake in northern regions. We suggest that gs, gm, WUE and PNUE are the functional traits most associated with fine-scale geographic clines and with the degree of local adaptation of white spruce populations to their biophysical environments. These geographic patterns may reflect in situ adaptive genetic differences in photosynthetic efficiency along the cline.

Population Differentiation and Climatic Adaptation for Growth Potential of White Spruce (Picea glauca) in Alberta, Canada

Abstract Genetic differentiation among white spruce populations in Alberta, Canada, was studied using time series data of height and diameter and a climatic index developed by principal component analysis. The objectives were to discern patterns of variation for growth potential and predicted optimum climate; compare optimum climate between populations, between height and diameter at the same age and between height or diameter at different ages; and to see if optimum climate differed from the climate inhabited by populations. Using cluster analysis we found that: (1) populations from mid-latitudes (54° - 57°N) and mid-elevations (600 - 800 m) were grouped together and exhibited high growth potential; populations from north of 57°N were grouped with those from elevations higher than 900m in the Rocky Mountains and exhibited low growth potential; and (2) With minor exceptions, populations from similar climates or geography were grouped together in terms of predicted optimum climate. (3) Analysis of variance showed that optimum climate differed significantly (P < 0.05) among populations; among heights at different ages; among diameters at different ages and between height and diameter at the same ages. However, there was no consistent trend in the direction of change in optimum climate with tree age. (4) The range of climate inhabited by the populations (PI₁ = -5.792 to 4.483) was much wider than the range of their predicted optimum climate (P̂Ō₁ = -1.001 to 0.842), which suggests that in terms of growth potential some populations inhabit sub-optimal climates. Implications of the results on management of white spruce in Alberta are discussed.

Phylogeography of white spruce (Picea glauca) in eastern North America reveals contrasting ecological trajectories

AbstractAim The main objective of this study was to investigate the origin of white spruce (Picea glauca) range discontinuities in eastern North America. A first analysis aimed at uncovering the glacial origin and post‐glacial migration route of white spruce in the eastern part of the range. A second analysis aimed at evaluating whether disjunct white spruce populations in central Québec result from a recent expansion or are remnant stands following a northern range contraction.Location The first analysis covered the North American transcontinental boreal distribution of white spruce, and the second focused on the eastern part of its range.Methods A total of 589 P. glauca individuals from 51 populations, 24 of which were from a previous published source, were analysed for variation in three chloroplast DNA regions (trnT/L, trnL/F and ndhK/C). At the continental scale, haplotypic diversity, zones of genetic discontinuities, clustering analysis and analysis of molecular variance (AMOVA) were employed. In the eastern part of the range, haplotypic diversity was assessed with a permutation procedure, demographic history with mismatch analyses, and genetic structure with AMOVA.Results At the continental scale, various lines of evidence (genetic‐diversity hotspots, endemic haplotypes, zones of genetic discontinuities, and pattern of molecular variation) show that different lineages migrated from three glacial refugia (Beringian, Mississippian, east Appalachian). Strong genetic discontinuities, low diversity and lack of evidence for demographic expansion were found among stands from isolated high hills in central Québec.Main conclusions White spruce in the eastern part of its range has an east Appalachian origin, a lineage genetically distinct from populations west of the Great Lakes. Previous studies have shown that the northward migration of white spruce is still ongoing as the northernmost populations expand into the tundra. In contrast, mixed white spruce and balsam fir (Abies balsamea) stands from isolated high hills in central Québec have already reached their Holocene maxima and are currently declining. This range contraction is probably a result of increased wildfire disturbance, which gradually creates a pattern of regional white spruce isolation and replacement by fire‐prone black spruce (Picea mariana) forest.

Scanning the genome for gene single nucleotide polymorphisms involved in adaptive population differentiation in white spruce

Conifers are characterized by a large genome size and a rapid decay of linkage disequilibrium, most often within gene limits. Genome scans based on noncoding markers are less likely to detect molecular adaptation linked to genes in these species. In this study, we assessed the effectiveness of a genome-wide single nucleotide polymorphism (SNP) scan focused on expressed genes in detecting local adaptation in a conifer species. Samples were collected from six natural populations of white spruce (Picea glauca) moderately differentiated for several quantitative characters. A total of 534 SNPs representing 345 expressed genes were analysed. Genes potentially under natural selection were identified by estimating the differentiation in SNP frequencies among populations (FST) and identifying outliers, and by estimating local differentiation using a Bayesian approach. Both average expected heterozygosity and population differentiation estimates (HE = 0.270 and FST = 0.006) were comparable to those obtained with other genetic markers. Of all genes, 5.5% were identified as outliers with FST at the 95% confidence level, while 14% were identified as candidates for local adaptation with the Bayesian method. There was some overlap between the two gene sets. More than half of the candidate genes for local adaptation were specific to the warmest population, about 20% to the most arid population, and 15% to the coldest and most humid higher altitude population. These adaptive trends were consistent with the genes’ putative functions and the divergence in quantitative traits noted among the populations. The results suggest that an approach separating the locus and population effects is useful to identify genes potentially under selection. These candidates are worth exploring in more details at the physiological and ecological levels.

Recent advance of white spruce (Picea glauca) in the coastal tundra of the eastern shore of Hudson Bay (Québec, Canada)

AbstractAim The species‐specific response of tree‐line species to climatic forcing is a crucial topic in modelling climate‐driven ecosystem dynamics. In northern Québec, Canada, black spruce (Picea mariana) is the dominant species at the tree line, but white spruce (Picea glauca) also occurs along the maritime coast of Hudson Bay, and is expanding along the coast and on lands that have recently emerged because of isostatic uplift. Here we outline the present distribution, structure, dynamics and recent spread of white spruce from the tree line up to its northernmost position in the shrub tundra along the Hudson Bay coast. We aimed to obtain a minimum date of the arrival of the species in the area and to evaluate its dynamics relative to recent climate changes.Location White spruce populations and individuals were sampled along a latitudinal transect from the tree line to the northernmost individual in the shrub tundra along the Hudson Bay coast and in the Nastapoka archipelago in northern Québec and Nunavut, Canada (56°06′–56°32′ N).Methods White spruce populations were mapped, and the position, dimension, growth form and origin (seed or layering) of every individual recorded. Tree‐ring analyses of living and dead trees allowed an estimation of the population structure, past recruitment, growth trends and growth rate of the species. A macrofossil analysis was performed of the organic horizon of the northernmost white spruce stands and individuals. Radiocarbon dates of white spruce remains and organic matter were obtained. The rate of isostatic uplift was assessed by radiocarbon dating of drifted wood fragments.Results The first recorded establishment of white spruce was almost synchronous at all sites and occurred around ad 1660. Spruce recruitment was rather continuous at the tree line, while it showed a gap in the northern shrub tundra during the first decades of the 19th century. A vigorous, recent establishment of seedlings was observed in the shrub tundra; only wind‐exposed, low krummholz (stunted individuals) did not show any sexual regeneration. A period of suppressed growth occurred from the 1810s to the 1850s in most sites. A growth increase was evident from the second half of the 19th century and peaked in the 1880s and the 20th century. A shift from stunted to tree growth form has occurred since the mid‐19th century. No sample associated with white spruce remains gave a date older than 300 14C years bp [calibrated age (cal.) ad 1430–1690].Main conclusions White spruce probably arrived recently in the coastal tundra of Hudson Bay due to a delayed post‐glacial spread. The arrival of the species probably occurred during the Little Ice Age. The established individuals survived by layering during unfavourable periods, but acted as nuclei for sexual recruitment almost continuously, except in the northernmost and most exposed sites. Warmer periods were marked by strong seedling recruitment and a shift to tree growth form. Unlike white spruce, black spruce showed no evidence of an ongoing change in growth form and sexual recruitment. Ecological requirements and recent history of tree‐line species should be taken into account in order to understand the present dynamics of high‐latitude ecosystems.

The impact of climate change on growth of local white spruce populations in Québec, Canada

In the near future, forest tree species growing in eastern Canada are expected to be affected by climate change due to an increase of human-induced greenhouse gas emissions. In this study, models were developed to estimate the impact of climate change on growth in white spruce ( Picea glauca (Moench) Voss). Data were collected in a genecological test, replicated in three locations, and involving 45 distinct geographical seed sources, most of them represented by five open-pollinated families. Transfer models predicting the performance of seed sources were developed, based on temperature and precipitation differentials between the geographical origin of seed sources and the experimental site locations. These models were validated using data collected in a second genecological test series. We found that white spruce populations located within the sampled area were optimally adapted to their local environment for thermal conditions but not for moisture conditions; populations that originated from sites receiving more precipitation generally showed higher tree growth than the local sources. We predict that the adaptive lag currently related to precipitation will increase under global warming conditions. Simulations of growth under various scenarios of climate change indicated that it would be diminished tangibly under more intense warming. However, for a given temperature increase, the relative loss in growth will be less if precipitation is reduced than if it increases. Consequently, predictions based solely on temperature change appear inaccurate, and more effort should be directed toward better anticipating the magnitude and the direction of changes in precipitation patterns at the regional scale. The necessity of human intervention to assist tree migration under climate change is examined.

Contrasting evolutionary forces driving population structure at expressed sequence tag polymorphisms, allozymes and quantitative traits in white spruce.

Patterns of variation in quantitative characters and genetic markers were compared among six regional populations of white spruce [Picea glauca (Moench) Voss]. Although some phenotypic characters were correlated with latitude (r = 0.791), longitude (r = -0.796) and precipitation during the growing season (r = 0.789), variability at genetic markers was not correlated with geographical or bioclimatic variables, and followed neutral expectations. Estimates of genetic diversity and population differentiation for 14 allozymes (translated regions of coding genes) were essentially indistinguishable from those observed for 11 expressed sequence tag polymorphisms (ESTPs) from untranslated regions of coding genes. Variation among populations for quantitative traits such as eighth year height (Q(ST) = 0.082), thirteenth year height (Q(ST) = 0.069), total wood density (Q(ST) = 0.102) and date of budset (Q(ST) = 0.246), was greater than for allozymes (G(ST) = 0.014) and ESTPs (G(ST) = 0.019). These trends suggest a strong adaptive response in quantitative traits, contrasting to allozymes and ESTPs where no selective response could be detected and where populations appeared to be essentially in a migration-drift equilibrium.

Comparisons of genetic diversity in white spruce (Picea glauca) and jack pine (Pinus banksiana) seed orchards with natural populations

Genetic diversity within a white spruce (Picea glauca (Moench) Voss) seed orchard (consisting of 40 clones) and a jack pine (Pinus banksiana Lamb.) seed orchard (31 clones) was assessed and compared with genetic diversity in natural populations within the source area for the orchards. Genetic diversity was determined at 18 allozyme loci for seven white spruce populations and 27 loci for five jack pine populations, and the two orchards. Gene diversity maintained within the seed orchards (He = 0.157 for white spruce and 0.114 for jack pine) was similar to that found within the source area for each species (He = 0.164 and 0.114 for white spruce and jack pine, respectively). However, nine white spruce alleles and 12 jack pine alleles identified in the source area were not present in the seed orchards. These alleles occurred at low frequencies in the natural populations (mean frequency = 0.023 and 0.014 for white spruce and jack pine, respectively). Mean genetic identities between the seed orchards and their natural populations were high (>0.99), indicating that common allele occurrences and frequencies were similar between the orchards and their source area. One allele in the white spruce orchard and two in the jack pine seed orchard did not occur within the natural population samples. Simulations indicated that randomly reducing the number of clones within the seed orchards would decrease allelic richness slightly but would have little effect on overall gene diversity.

Interannual growth variation in fish and tree rings

Interannual growth variations were compared among neighbouring populations of lake sturgeon (Acipenser fulvescens) and white spruce (Picea glauca), white pine (Pinus strobus), and red pine (Pinus resinosa). Measures of growth were obtained by removing long-term trends from widths of rings in the hard tissues of both aquatic and terrestrial organisms and assembling these measures into growth chronologies. Interannual growth variations were negatively correlated (r) between sturgeon and nearby tree population chronologies for those fish populations that displayed strong interseries correlation (mean r). The three sturgeon population chronologies developed from individuals that displayed the lowest interseries correlation coefficients failed to display significant correlation with tree growth. The results of this investigation indicate that the negative relationships between fish and tree growth may be related to annual fluctuations in air temperature. In general, fish population chronologies displayed positive correlation with measures of air temperature during the current season of growth, while tree population chronologies displayed negative correlation with air temperatures from either the current or the previous season of growth.

Population genetic variation, structure, and evolution in Engelmann spruce, white spruce, and their natural hybrid complex in Alberta

Genetic variation, structure, and evolution of 12 populations of putative Engelmann spruce (Picea engelmanii Parry), white spruce (Picea glauca (Moench) Voss), and Engelmann - white spruce natural hybrids from the sympatric areas and two populations of white spruce from the allopatric areas in Alberta were examined using 23 allozyme loci coding for 13 enzymes in needles. Although most of the alleles were widespread, unique alleles were found in 9 of the 14 populations. No species-specific allele was observed. However, allele frequency differences were observed between the putative Engelmann and white spruce populations at a number of loci. Frequencies of 13 alleles showed significant correlation with altitude, those of 11 alleles showed correlation with latitude, and those of 3 alleles showed correlation with longitude. On average, 66.2% (99% criterion) of the loci were polymorphic, the number of alleles per locus was 1.88, the number of alleles per polymorphic locus was 2.88, and the observed and expected heterozygosities were 0.063 and 0.184, respectively. Genetic variability of allopatric white spruce, putative Engelmann, sympatric white spruce, and hybrid populations was quite comparable. The mean FSTestimate was 0.123 for the total populations. Canonical discriminant functions separated four putative Engelmann spruce populations from the fifth putative Engelmann spruce population and from the allopatric and sympatric white spruce and sympatric putative hybrid populations. A cluster analysis from genetic distances generally separated allopatric and sympatric white spruce populations from the putative Engelmann spruce and hybrid populations. A Wagner tree of the 14 populations produced two main branches; one branch consisting of two allopatric and two sympatric white spruce and one putative hybrid populations, and another branch consisting of the remaining nine spruce populations representing putative Engelmann spruce, putative hybrids, and sympatric white spruce. Putative hybrids showed lower distances to white spruce from the canonical discriminant analysis, whereas they showed lower distances to putative Engelmann spruce from the genetic distance analysis. High allozyme genetic identities between putative Engelmann and white spruce and allelic differentiation related to altitude in Alberta suggest that Engelmann spruce could at best be considered as a subspecies of Picea glauca, with white spruce named as Picea glauca ssp. glauca and Engelmann spruce named as Picea glauca ssp. engelmannii.Key words: Picea glauca, Picea engelmannii, biosystematics, natural hybridization, species differentiation, genetic divergence.

Population genetic variation, structure, and evolution in Engelmann spruce, white spruce, and their natural hybrid complex in Alberta

Genetic variation, structure, and evolution of 12 populations of putative Engelmann spruce ( Picea engelmanii Parry), white spruce (Picea glauca (Moench) Voss), and Engelmann - white spruce natural hybrids from the sympatric areas and two populations of white spruce from the allopatric areas in Alberta were examined using 23 allozyme loci coding for 13 enzymes in needles. Although most of the alleles were widespread, unique alleles were found in 9 of the 14 populations. No species-specific allele was observed. However, allele frequency differences were observed between the putative Engelmann and white spruce populations at a number of loci. Frequencies of 13 alleles showed significant correlation with altitude, those of 11 alleles showed correlation with latitude, and those of 3 alleles showed correlation with longitude. On average, 66.2% (99% criterion) of the loci were polymorphic, the number of alleles per locus was 1.88, the number of alleles per polymorphic locus was 2.88, and the observed and expected heterozygosities were 0.063 and 0.184, respectively. Genetic variability of allopatric white spruce, putative Engelmann, sympatric white spruce, and hybrid populations was quite comparable. The mean FST estimate was 0.123 for the total populations. Ca- nonical discriminant functions separated four putative Engelmann spruce populations from the fifth putative Engelmann spruce population and from the allopatric and sympatric white spruce and sympatric putative hybrid populations. A cluster analysis from genetic distances generally separated allopatric and sympatric white spruce populations from the putative Engelmann spruce and hybrid populations. A Wagner tree of the 14 populations produced two main branches; one branch consisting of two allopatric and two sympatric white spruce and one putative hybrid populations, and an- other branch consisting of the remaining nine spruce populations representing putative Engelmann spruce, putative hy- brids, and sympatric white spruce. Putative hybrids showed lower distances to white spruce from the canonical discriminant analysis, whereas they showed lower distances to putative Engelmann spruce from the genetic distance analysis. High allozyme genetic identities between putative Engelmann and white spruce and allelic differentiation re- lated to altitude in Alberta suggest that Engelmann spruce could at best be considered as a subspecies of Picea glauca, with white spruce named as Picea glauca ssp. glauca and Engelmann spruce named as Picea glauca ssp. engelmannii.