Atlantic salmon colonization of the Russian Arctic coast: pioneers from North America

Previous studies of the ESTD * isozyme locus in the Atlantic salmon show the * 80 allele to be absent across the species' European range, with the exception of northern Russia, whereas the allele is nearly fixed in North American populations. The allele was found in samples from 15 out of 18 rivers on the Kola Peninsula and White sea coast and had frequencies that ranged from 0·017 to 0·363. Typing of fish in nine of these rivers for mtDNA variation in the ND1 gene region found variation characteristic of North American salmon in three Kola Peninsula populations and is the only part of Europe where such variants have been detected. The study area was completely glaciated during the late Pleistocene period and the restriction of the European distribution of these ESTD * and mtDNA variants to this area suggests that salmon that colonized the Russian Arctic coast rivers included fish of North American origin after the Pleistocene glacier had retreated.     

Parallel and nonparallel genome‐wide divergence among replicate population pairs of freshwater and anadromous Atlantic salmon

Little is known about the genetic basis differentiating resident and anadromous forms found in many salmonid species. Using a medium‐density SNP array, we documented genomic diversity and divergence at 2336 genetically mapped loci among three pairs of North American anadromous and freshwater Atlantic salmon populations. Our results show that across the genome, freshwater populations have lower diversity and a smaller proportion of private polymorphism relative to anadromous populations. Moreover, differentiation was more pronounced among freshwater than among anadromous populations at multiple spatial scales, suggesting a large effect of genetic drift in these isolated freshwater populations. Using nonhierarchical and hierarchical genome scans, we identified hundreds of markers spread across the genome that are potentially under divergent selection between anadromous and freshwater populations, but few outlier loci were repeatedly found in all three freshwater–anadromous comparisons. Similarly, a sliding window analysis revealed numerous regions of high divergence that were nonparallel among the three comparisons. These last results show little evidence for the parallel evolution of alleles selected for in freshwater populations, but suggest nonparallel adaptive divergence at many loci of small effects distributed through the genome. Overall, this study emphasizes the important role of genetic drift in driving genome‐wide reduction in diversity and divergence in freshwater Atlantic salmon populations and suggests a complex multigenic basis of adaptation to resident and anadromous strategies with little parallelism.     

Expression of olfactory receptors in different life stages and life histories of wild Atlantic salmon (Salmo salar)

It has been hypothesized that salmonids use olfactory cues to return to their natal rivers and streams. However, the key components of the molecular pathway involved in imprinting and homing are still unknown. If odorants are involved in salmon homing migration, then olfactory receptors should play a critical role in the dissipation of information from the environment to the fish. Therefore, we examined the expression profiles of a suite of genes encoding olfactory receptors and other olfactory-related genes in the olfactory rosettes of different life stages in two anadromous and one non-anadromous wild Atlantic salmon populations from Newfoundland, Canada. We identified seven differentially expressed OlfC genes in juvenile anadromous salmon compared to returning adults in both populations of anadromous Atlantic salmon. The salmon from the Campbellton River had an additional 10 genes that were differentially expressed in juveniles compared to returning adults. There was no statistically significant difference in gene expression of any of the genes in the non-anadromous population (P < 0.01). The function of the OlfC gene products is not clear, but they are predicted to be amino acid receptors. Other studies have suggested that salmon use amino acids for imprinting and homing. This study, the first to examine the expression of olfactory-related genes in wild North American Atlantic salmon, has identified seven OlfC genes that may be involved in the imprinting and homeward migration of anadromous Atlantic salmon.     

Stream flow, salmon and beaver dams: roles in the structuring of stream fish communities within an anadromous salmon dominated stream

The current paradigm of fish community distribution is one of a downstream increase in species richness by addition, but this concept is based on a small number of streams from the mid-west and southern United States, which are dominated by cyprinids. Further, the measure of species richness traditionally used, without including evenness, may not be providing an accurate reflection of the fish community. We hypothesize that in streams dominated by anadromous salmonids, fish community diversity will be affected by the presence of the anadromous species, and therefore be influenced by those factors affecting the salmonid population. Catamaran Brook, New Brunswick, Canada, provides a long-term data set to evaluate fish community diversity upstream and downstream of an obstruction (North American beaver Castor canadensis dam complex), which affects distribution of Atlantic salmon Salmo salar. The Shannon Weiner diversity index and community evenness were calculated for sample sites distributed throughout the brook and over 15 years. Fish community diversity was greatest upstream of the beaver dams and in the absence of Atlantic salmon. The salmon appear to depress the evenness of the community but do not affect species richness. The community upstream of the beaver dams changes due to replacement of slimy sculpin Cottus cognatus by salmon, rather than addition, when access is provided. Within Catamaran Brook, location of beaver dams and autumn streamflow interact to govern adult Atlantic salmon spawner distribution, which then dictates juvenile production and effects on fish community. These communities in an anadromous Atlantic salmon dominated stream do not follow the species richness gradient pattern shown in cyprinid-dominated streams and an alternative model for stream fish community distribution in streams dominated by anadromous salmonids is presented. This alternative model suggests that community distribution may be a function of semipermeable obstructions, streamflow and the distribution of the anadromous species affecting resident stream fish species richness, evenness, biomass and production.     

Population structure of Atlantic salmon (Salmo salar L.): a range-wide perspective from microsatellite DNA variation

Atlantic salmon (n = 1682) from 27 anadromous river populations and two nonanadromous strains ranging from south-central Maine, USA to northern Spain were genotyped at 12 microsatellite DNA loci. This suite of moderate to highly polymorphic loci revealed 266 alleles (5-37/locus) range-wide. Statistically significant allelic and genotypic heterogeneity was observed across loci between all but one pairwise comparison. Significant isolation by distance was found within and between North American and European populations, indicating reduced gene flow at all geographical scales examined. North American Atlantic salmon populations had fewer alleles, fewer unique alleles (though at a higher frequency) and a shallower phylogenetic structure than European Atlantic salmon populations. We believe these characteristics result from the differing glacial histories of the two continents, as the North American range of Atlantic salmon was glaciated more recently and more uniformly than the European range. Genotypic assignment tests based on maximum-likelihood provided 100% correct classification to continent of origin and averaged nearly 83% correct classification to province of origin across continents. This multilocus method, which may be enhanced with additional polymorphic loci, provides fishery managers the highest degree of correct assignment to management unit of any technique currently available.     

Formation of population genetic structure following the introduction and establishment of non-native American shad (<Emphasis Type="Italic">Alosa sapidissima</Emphasis>) along the Pacific Coast of North America

Biological invasions provide opportunities to examine contemporary evolutionary processes in novel environments. American shad, an anadromous fish native to the Atlantic Coast of North America, was introduced to California in 1871 and established spawning populations along the Pacific Coast that may provide insights into the dynamics of dispersal, colonization, and the establishment of philopatry. Using 13 neutral microsatellite loci we genotyped anadromous, freshwater resident and landlocked American shad from 14 locations along the US Pacific Coast to resolve population genetic structure. We observed significant differences in multilocus allele frequency distributions in nearly all (61/66; 92%) pairwise comparisons of non-native anadromous, freshwater resident and landlocked populations, and detected significant genetic differentiation for most (55/66; 83%) of these comparisons. Genetic divergence between landlocked and anadromous populations is due to genetic drift in isolation because of a physical migration barrier. However, some reproductive isolating mechanism maintains genetic differentiation between sympatric populations in the Columbia River exhibiting alternative life history strategies (i.e. anadromous vs. ‘freshwater-type’). Non-native populations possessed genetic variants that were not observed in the species’ native range and were strongly differentiated from Atlantic Coast populations (\({\text{G}}^{{\prime }}_{\text{ST}}\)?=?0.218). Our results indicate that philopatry became established shortly after dispersal and colonization along the Pacific Coast. This study contributes to our understanding of dynamic evolutionary processes during invasions.     

Who are the missing parents? Grandparentage analysis identifies multiple sources of gene flow into a wild population

In order to increase the size of declining salmonid populations, supplementation programmes intentionally release fish raised in hatcheries into the wild. Because hatchery-born fish often have lower fitness than wild-born fish, estimating rates of gene flow from hatcheries into wild populations is essential for predicting the fitness cost to wild populations. Steelhead trout (Oncorhynchus mykiss) have both freshwater resident and anadromous (ocean-going) life history forms, known as rainbow trout and steelhead, respectively. Juvenile hatchery steelhead that 'residualize' (become residents rather than go to sea as intended) provide a previously unmeasured route for gene flow from hatchery into wild populations. We apply a combination of parentage and grandparentage methods to a three-generation pedigree of steelhead from the Hood River, Oregon, to identify the missing parents of anadromous fish. For fish with only one anadromous parent, 83% were identified as having a resident father while 17% were identified as having a resident mother. Additionally, we documented that resident hatchery males produced more offspring with wild anadromous females than with hatchery anadromous females. One explanation is the high fitness cost associated with matings between two hatchery fish. After accounting for all of the possible matings involving steelhead, we find that only 1% of steelhead genes come from residualized hatchery fish, while 20% of steelhead genes come from wild residents. A further 23% of anadromous steelhead genes come from matings between two resident parents. If these matings mirror the proportion of matings between residualized hatchery fish and anadromous partners, then closer to 40% of all steelhead genes come from wild trout each generation. These results suggest that wild resident fish contribute substantially to endangered steelhead 'populations' and highlight the need for conservation and management efforts to fully account for interconnected Oncorhynchus mykiss life histories.     

Genetic population structure of Norwegian brown trout

Biochemical genetic variation in populations of anadromous and resident brown trout, Salmo trutta L., was studied. Altogether 50 Norwegian populations were screened for 32 enzyme loci. Genetic polymorphism was found at the following 11 loci: AAT-4 * (E.C. 2.6.1.1), CK-1 * (E.C. 2.7.3.2), G3PDH-2 * (E.C. 1.1.1.8), IDHP-2 * (E.C. 1.1.1.42), LDH-5 * (E.C. 1.1.1.27), MDH-2 * (E.C. 1.1.1.37), MDH-3/4 * (E.C. 1.1.1.37), MEP-2 * (E.C. 1.1.1.40), GPI-2 * (E.C. 5.3.1.9). GPI-5 * (E.C. 5.3.1.9) and PGM-1 * (E.C. 5.4.2.2), giving an overall polymorphism of 34%, ranging from 3.7 to 29.6% among individual populations. The average calculated heterozygosity ranged from 1.4 to 10.2% among populations. Genetic heterogeneity was observed among anadromous populations, and significant differences in allelic frequencies were found between anadromous populations in neighbouring watercourses, among resident populations and between anadromous and resident populations inhabiting the same watercourses. Significant heterogeneity was also found among 12 populations from Lake Mjøsa, with a major division between the western and eastern populations of the lake. Differences in allelic frequencies were found between wild stocks and their hatchery derivatives, and between different hatchery derivatives originating from the same wild population. In some cases release of hatchery populations into wild stocks may have influenced the genetic characteristics of wild stocks. The data support the hypothesis of eastern as well as western postglacial colonization lines for Norwegian brown trout.     

Differences in growth between offspring of anadromous and freshwater brown trout Salmo trutta

In this study, individual growth of juvenile offspring of anadromous and freshwater resident brown trout Salmo trutta and crosses between the two from the River Imsa, Norway, was estimated. The juveniles were incubated until hatching at two temperatures (±S.D. ), either 4.4 ± 1.5°C or 7.1 ± 0.6°C. Growth rate was estimated for 22 days in August–September when the fish on average were c. 8 g in wet mass, and the estimates were standardized to 1 g fish dry mass. Offspring of anadromous S. trutta grew better at both 15 and 18°C than offspring of freshwater resident S. trutta or offspring of crosses between the two S. trutta types. This difference appears not to result from a maternal effect because anadromous S. trutta grew better than the hybrids with anadromous mothers. Instead, this appears to be an inherited difference between the anadromous and the freshwater resident fish lending support to the hypothesis that anadromous and freshwater resident S. trutta in this river differ in genetic expression. Egg incubation temperature of S. trutta appeared not to influence the later growth as reported earlier from the studies of Atlantic salmon Salmo salar.     

Growth and survival of Atlantic salmon (Salmo salar) in Llyn Dwythwch, North Wales

SUMMARY. Atlantic salmon fry have been annually stocked into Llyn Dwythwch, North Wales, since 1969, in an attempt to increase the natural stocks of the area. The growth and survival of 1- and 2-year-old salmon were investigated, and compared with that of other lake-reared populations and also with salmon in the natural stream environment. Lake-reared salmon follow the same patterns of slow and rapid growth as found for river fish, but the growth rate was superior in the former. The variation in length – weight relationship with age and sex was investigated. Survival rates in general compared favourably with the survival in rivers, with high mortality rates of salmon in Llyn Dwythwch resulting from predation at spring stocking by the resident brown trout. This was later avoided by stocking larger fish in the autumn.     

Brown trout Salmo trutta released to support recreational fishing in a Norwegian fjord

One- and two-year-old hatchery reared juveniles of seven freshwater resident and anadromous populations of Scandinavian brown trout were released in the outer and inner part of the Oslofjord and in the River Akerselva, flowing through the city of Oslo. Recapture rates were highest (mean 20.3%) for river released fish and lowest for those released in the outer (16.8%) and inner (12.1%) fjord. In general, recapture rate increased with fish size at release (r=0.76). When released in fresh water, most of the recaptures were from fresh water and when released in the fjord, most recaptures were from the fjord. In general, freshwater resident stocks showed a higher degree of freshwater residency than anadromous stocks. However, mean migratory distance was longer for freshwater resident than anadromous fish. Trout moved longer distances at sea when released in the outer than in the inner fjord. Specific growth rate and size at recapture varied among release sites and stocks; they were highest for fish released in the outer fjord and lowest for river-released trout. There was no consistent difference in sea growth between freshwater resident and anadromous stocks. Estimated total yield was highest for fish released in the outer fjord, whereas there was no significant difference in yields between trout released in the river and the inner Oslofjord.     

Occurrence and genetic typing of infectious hematopoietic necrosis virus in Kamchatka, Russia

Infectious hematopoietic necrosis virus (IHNV) is a well known rhabdoviral pathogen of salmonid fish in North America that has become established in Asia and Europe. On the Pacific coast of Russia, IHNV was first detected in hatchery sockeye from the Kamchatka Peninsula in 2001. Results of virological examinations of over 10,000 wild and cultured salmonid fish from Kamchatka during 1996 to 2005 revealed IHNV in several sockeye salmon Oncorhynchus nerka populations. The virus was isolated from spawning adults and from juveniles undergoing epidemics in both hatchery and wild sockeye populations from the Bolshaya watershed. No virus was detected in 2 other watersheds, or in species other than sockeye salmon. Genetic typing of 8 virus isolates by sequence analysis of partial glycoprotein and nucleocapsid genes revealed that they were genetically homogeneous and fell within the U genogroup of IHNV. In phylogenetic analyses, the Russian IHNV sequences were indistinguishable from the sequences of North American U genogroup isolates that occur throughout Alaska, British Columbia, Washington, and Oregon. The high similarity, and in some cases identity, between Russian and North American IHNV isolates suggests virus transmission or exposure to a common viral reservoir in the North Pacific Ocean.     

Variations in juvenile growth, energy allocation and life-history strategies of two populations of Arctic charr in North Norway

Within the populations of Arctic charr Salvelinus alpinus in Storvatn and Rungavatn, Norway, fish that attained the largest size in the late parr stage migrated as 4+ smolts, fish that attained a medium size became 5+ smolts, and the smallest parr became lake residents. Within the last 2 years of the parr stage, those that became anadromous had a lower growth rate during the winter and early summer than those that became resident, and vice versa in late summer. Thus, anadromous fish grew faster in the parr stage, but in years prior to migration, their growth pattern seemed to differ from that of parr that became resident. Due to early size differences between parr that became anadromous or resident, it is suggested that some of the basis for the decision to smoltify or not in charr depends on the growth rate through the whole parr stage, but with significant modification by the local environment. Rungavatn parr had a significantly higher population density (5 ×), higher lipid content and mortality, an earlier maturation, a lower rate of growth, and only one-third the degree of anadromy than those from Storvatn. It is suggested that the two charr populations have evolved different life-history adaptations due to unequal growth, energy allocation and competition opportunities in the lakes.     

Landscape genetics and hierarchical genetic structure in Atlantic salmon: the interaction of gene flow and local adaptation

Disentangling evolutionary forces that may interact to determine the patterns of genetic differentiation within and among wild populations is a major challenge in evolutionary biology. The objective of this study was to assess the genetic structure and the potential influence of several ecological variables on the extent of genetic differentiation at multiple spatial scales in a widely distributed species, the Atlantic salmon, Salmo salar . A total of 2775 anadromous fish were sampled from 51 rivers along the North American Atlantic coast and were genotyped using 13 microsatellites. A Bayesian analysis clustered these populations into seven genetically and geographically distinct groups, characterized by different environmental and ecological factors, mainly temperature. These groups were also characterized by different extent of genetic differentiation among populations. Dispersal was relatively high and of the same magnitude within compared to among regional groups, which contrasted with the maintenance of a regional genetic structure. However, genetic differentiation was lower among populations exchanging similar rates of local as opposed to inter-regional migrants, over the same geographical scale. This raised the hypothesis that gene flow could be constrained by local adaptation at the regional scale. Both coastal distance and temperature regime were found to influence the observed genetic structure according to landscape genetic analyses. The influence of other factors such as latitude, river length and altitude, migration tactic, and stocking was not significant at any spatial scale. Overall, these results suggested that the interaction between gene flow and thermal regime adaptation mainly explained the hierarchical genetic structure observed among Atlantic salmon populations.     

Genomic population structure of freshwater‐resident and anadromous ide (Leuciscus idus) in north‐western Europe

Climate change experts largely agree that future climate change and associated rises in oceanic water levels over the upcoming decades, will affect marine salinity levels. The subsequent effects on fish communities in estuarine ecosystems however, are less clear. One species that is likely to become increasingly affected by changes in salinity is the ide (Leuciscus idus). The ide is a stenohaline freshwater fish that primarily inhabits rivers, with frequent anadromous behavior when sea salinity does not exceed 15%. Unlike most other anadromous Baltic Sea fish species, the ide has yet to be subjected to large‐scale stocking programs, and thus provides an excellent opportunity for studying the natural population structure across the current salinity gradient in the Danish Belts. To explore this, we used Genotyping‐by‐Sequencing to determine genomic population structure of both freshwater resident and anadromous ide populations in the western Baltic Sea region, and relate the results to the current salinity gradient and the demographic history of ide in the region. The sample sites separate into four clusters, with all anadromous populations in one cluster and the freshwater resident populations in the remaining three. Results demonstrate high level of differentiation between sites hosting freshwater resident populations, but little differentiation among anadromous populations. Thus ide exhibit the genomic population structure of both a typical freshwater species, and a typical anadromous species. In addition to providing a first insight into the population structure of north‐western European ide, our data also (1) provide indications of a single illegal introduction by man; (2) suggest limited genetic effects of heavy pollution in the past; and (3) indicate possible historical anadromous behavior in a now isolated freshwater population.     

New polymorphic microsatellite markers for the masu salmon (Oncorhynchus masou masou) from Korea and their application to wild and hatchery populations

Masu salmon, Oncorhynchus masou masou, is an economically important fish species in the Far East and occurs in two life history forms: sea-run migratory (anadromous) and freshwater resident (non-anadromous). The non-anadromous form has recently become a popular freshwater food and game fish during a well-known Korean winter festival. However, the genetic background of this species remains largely unknown, partly due to a lack of molecular genetic markers. In this study, we developed new polymorphic microsatellite markers for masu salmon using next-generation sequencing technology. From 40 primer sets, 11 primer sets (27.5% of the primer sets selected) were successfully amplified with 106 alleles (range 2–9) in 64 individuals from different populations: two wild and one hatchery. Observed and expected heterozygosities ranged from 0.304 to 0.947 and 0.278 to 0.865, respectively. Significant departures from the Hardy–Weinberg equilibrium were detected for four markers (OMM11, OMM17, OMM28, and OMM33) in a single population. All pair-wise F_ST values were highly significant between the wild and hatchery populations (range 0.084–0.183, P < 0.0001). We identified a set of robust microsatellite markers that worked well even in formalin-fixed samples, which will be suitable for biogeographical and population structure analyses of the masu salmon.     

Ecological Forms of Black Sea Brown Trout (<Emphasis Type="Italic">Salmo trutta labrax</Emphasis>) in the Mzymta River as Manifestation of Ontogenetic Plasticity

Populations of brown trout in the Mzymta River and its tributaries include anadromous (mainly female) and resident (mainly males) fish. Some resident males in the basin of the Mzymty River attain sexual maturity at the age 1+, and resident females mature at the age 2+ or 3+. The maximum age of resident fish is 4+ in the samples studied. Migrations of anadromous brown trout to the sea occur at the ages 1+, 2+, or 3+. Future spawners spend from 1 to 4 years at feeding grounds in the sea. Smolts of the population are characterized by performing not only spring but also autumn migrations to the sea. One smolt specimen has been detected upstream from the dam in the river where spawners of anadromous brown trout do not migrate; this means that the capability for sea migrations persists long in the population represented only by resident specimens of brown trout. The diversity of life cycles and ecological forms in populations of brown trout is not lower than in populations of brown trout in Northern and Western Europe. The comparison of the data obtained with published data makes it possible to come to the conclusion about the high plasticity of ontogenesis of Black Sea brown trout.     

The demographic history of Atlantic salmon (Salmo salar) across its distribution range reconstructed from approximate Bayesian computations*

Understanding the dual roles of demographic and selective processes in the buildup of population divergence is one of the most challenging tasks in evolutionary biology. Here, we investigated the demographic history of Atlantic salmon across the entire species range using 2035 anadromous individuals from North America and Eurasia. By combining results from admixture graphs, geo‐genetic maps, and an Approximate Bayesian Computation (ABC) framework, we validated previous hypotheses pertaining to secondary contact between European and Northern American populations, but also identified secondary contacts in European populations from different glacial refugia. We further identified the major sources of admixture from the southern range of North America into more northern populations along with a strong signal of secondary gene flow between genetic regional groups. We hypothesize that these patterns reflect the spatial redistribution of ancestral variation across the entire North American range. Results also support a role for linked selection and differential introgression that likely played an underappreciated role in shaping the genomic landscape of species in the Northern hemisphere. We conclude that studies between partially isolated populations should systematically include heterogeneity in selective and introgressive effects among loci to perform more rigorous demographic inferences of the divergence process.     

Trade-off between egg mass and egg number in brown trout

Individual egg mass and fecundity increased with somatic mass in first time and repeat spawning wild anadromous and freshwater resident brown trout Salmo trutta . The egg mass was larger for similar-sized trout in south (58° N) than mid Norway (63° N), whereas fecundity was higher in mid- than in south Norway, making total gonadal investment similar in the two areas. Repeat spawners had heavier eggs than similar-sized first time spawners. The egg mass of residents was c. 10% larger than that of similar-sized first time spawning anadromous trout. Common garden experiments with offspring of wild anadromous trout showed no significant correlation between egg and somatic mass in first time spawners in two of the three populations studied. In the third population, a slight positive correlation was found. Similar results were found for repeat spawners. In the three populations, fecundity increased significantly with somatic mass in both first time and repeat spawners. Wild and hatchery-reared trout showed negative correlation between egg mass and fecundity when the effect of body size was excluded, indicating a trade-off between the two parameters. In wild trout, this was caused by variation among populations, whereas in hatchery fish, within population variation was observed in egg mass over fecundity. Furthermore, the egg mass of first time and repeat spawners were positively correlated, when adjusted for fish size. Size-specific gonadal investment was significantly higher in wild anadromous than resident trout. There was no significant difference in gonadal investment between first time and repeat spawners in wild anadromous trout. However, in the hatchery-reared trout, gonadal investment was significantly higher at repeat than first time maturation. The hatchery trout did not spawn naturally, but were artificially stripped. Among populations, a part of the variation in egg mass and fecundity is phenotypically plastic, a part appears genetically determined.     

Migration of Pacific Rim Chum Salmon on the High Seas: Insights from Genetic Data

Wild stocks of chum salmon, Oncorhynchus keta, have experienced recent declines in some areas of their range. Also, the release of hatchery chum salmon has escalated to nearly three billion fish annually. The decline of wild stocks and the unknown effects of hatchery fish combined with the uncertainty of future production caused by global climate change have renewed interest in the migratory patterns of chum salmon on the high seas. We studied the composition of high-seas mixtures of maturing and immature individuals using baseline data for 20 allozyme loci from 356 populations from throughout the Pacific Rim. Composition estimates were made from three time series. Two of these time series were from important coastal migratory corridors: the Shumagin Islands south of the Alaska Peninsula and the east coast of the Kamchatka Peninsula. The third was from chum salmon captured incidentally in the Bering Sea trawl fishery for walleye pollock. We also analyzed geographically dispersed collections of chum salmon captured in the month of July. The time series show dynamic changes in stock composition. The Shumagin Island corridor was used primarily by Northwest Alaskan and Asian populations in June; by the end of July stocks from the Alaska Peninsula and southern North America dominated the composition. The composition along the Kamchatka coast changed dramatically from primarily Russian stocks in May to primarily Japanese stocks in August; the previously undocumented presence of stocks from the Alaska Peninsula and Gulf of Alaska was also demonstrated. Immature chum salmon from throughout the Pacific Rim, including large proportions of southern North American stocks, contributed to the Bering Sea bycatch during the months of September and October. The migration routes of North American stocks is far more widespread than previously observed, and the Bering Sea is an important rearing area for maturing and immature chum salmon from throughout the species' range.