Distribution and population genetic structure of river- and sea-type sockeye salmon in western North America

Abstract– In contrast to the well-known "lake-type" sockeye salmon, two additional anadromous life-history types have been recognized within the species: 'river-type' sockeye salmon whose juveniles spend 1 or 2 years in off-channel river habitats prior to migrating to sea, and "sea-type" sockeye salmon that initially rear in similar river habitats yet migrate to sea as underyearlings. Persistent populations of river-/sea-type sockeye salmon occur in small numbers throughout the species'range in North America but are usually associated with glacier-fed rivers. We found published and unpublished records showing that riverine-spawning sockeye salmon occur in 11 rivers in western Washington, USA, that don't have access to juvenile lake-rearing habitat. Evidence of persistent spawning was strongest for the Nooksack and Skagit rivers in northern Puget Sound. We analyzed allozyme frequency differentiation in 26 laketype and 12 river-/sea-type populations of sockeye salmon in North America, ranging from northern Puget Sound, Washington (including 3 in the Nooksack and Skagit rivers) to northern Southeast Alaska. Across this 2000 km range, river-/sea-type sockeye salmon showed very little genetic differentiation between populations, much less than that displayed by the highly divergent lake-type sockeye salmon. Genetic similarity among river-/sea-type sockeye salmon in this study is likely a result of common ancestry and a high level of historical gene flow among river-/sea-type sockeye salmon populations.     

Freshwater habitat associations between pink (Oncorhynchus gorbuscha), chum (O. keta) and Chinook salmon (O. tshawytscha) in a watershed dominated by sockeye salmon (O. nerka) abundance

To understand the interplay between habitat use and contemporary anadromous Pacific salmon, Oncorhynchus spp., distributions we explored the habitat associations of three species, pink (O. gorbuscha), chum (O. keta) and Chinook salmon (O. tshawytscha) in streams of the Wood River system of Bristol Bay, Alaska, where sockeye salmon (O. nerka) are numerically dominant. We developed models to investigate the occurrence of nondominant salmon in relation to habitat characteristics and sockeye salmon density, using four decades of salmon presence and abundance data. The frequency of occurrence and abundance of nondominant species increased with watershed drainage area and stream depth and decreased with sockeye salmon density. The range of occurrence varied from nonexistent to perennial for the other species in sockeye‐dominated streams. Increasing watershed area resulted in larger stream habitat area and deeper habitats, allowing for the sympatric occurrence and persistence of all salmon species. The relationships between habitat and the presence of these Pacific salmon help define their requirements but also remind us that the patterns of presence and absence, within the overall ranges of salmon species, have yet to be fully understood.     

Distribution and feeding of juvenile Pacific salmon in freshwater tidal creeks of the lower Fraser River, British Columbia

Abstract This study examined juvenile salmonid use of a freshwater tidal creek system draining a wetland on the floodplain of the lower Fraser River, British Columbia, Canada. Chum, Oncorhynchus keta (Walbaum), chinook, O. tshawytscha (Walbaum), and sockeye, O. nerka (Walbaum), salmon fry were abundant in the tidal creeks in spring. The fry were found in non-natal habitat up to 1.5 km from the main channel of the river. The salmon fry ate dipteran adults, larvae and pupae, cyclopoid and harpacticoid copepods, and Collembola. Mysids Neomysis mercedis Holmes (Walbaum), and amphipods, Crangonyx richmondensis occidentalis (Hubricht and Harrison), were also consumed. The upper reaches of an undisturbed creek were the winter rearing habitat for presmolt coho salmon, O. kisutch (Walbaum), where this species ate dipteran pupae and larvae as well as a freshwater isopod, Asellus communis Say (Walbaum).     

Site fidelity of spawning sockeye salmon (Oncorhynchus nerka W.) in the presence and absence of olfactory cues

Abstract –  We examined the site fidelity of spawning adult sockeye salmon ( Oncorhynchus nerka ) by tagging and releasing fish in the same stream reach (controls) and displacing them among different but nearby sites ( c . 50 m away). Three sites – two above a stream junction ('upper' reach and 'pond') and one below ('lower' reach) – allowed us to compare the behavior of salmon in the presence and absence of olfactory cues and habitat similarity. Most controls of both sexes (90%) remained in the immediate vicinity of the tagging and release site. When displaced downstream, where the odors of both the upper reach and the pond were detectable, most salmon returned to their former site (65%). Displaced sockeye were more likely to return to the pond from the lower reach than from the upper one ( P  = 0.05), consistent with olfactory orientation and the hypothesis that salmon prefer certain habitats. Salmon displaced from the upper to the lower reach were much more likely to return than those displaced to the pond ( P  < 0.01), consistent with the role of odors in orientation and inconsistent with the habitat choice hypothesis.     

Long-term changes in rearing habitat and downstream movement by juvenile sockeye salmon (Oncorhynchus nerka) in an interconnected Alaska lake system

Abstract –  In some populations the phenomenon of partial migration develops where some individuals stay in a given habitat rather than move with the migratory component. Depending on the selective pressures, the individuals that stay may be larger, smaller or similar in size to those that move. Freshwater movements of juvenile sockeye salmon ( Oncorhynchus nerka Walbaum) fry vary among and within populations, and can be complex, especially in interconnected lake systems. We examined variation of movement patterns by a sockeye salmon population in an interconnected lake system during a period of rapid natural habitat change and found that fry migrating downstream were shorter, had lower body condition, and were more likely ill and moribund compared with fish remaining in the lake. However, otolith microstructure measurements indicated that emigrants did not grow significantly slower than residents prior to downstream movement. We show that patterns (i.e., demography of migrants, timing of movement) of downstream movement have changed since the 1970s, corresponding to changes in rearing habitat. Our findings parallel the results with other salmonid species and are generally consistent with the paradigm that density-dependent interactions from declining habitat availability or quality result in the downstream movement of competitively inferior individuals, although the mechanisms governing downstream migration are unclear in this system.     

Influence of the marine abundance of pink (Oncorhynchus gorbuscha) and sockeye salmon (O. nerka) on growth of Ozernaya River sockeye

The length and weight of Russian sockeye ( Oncorhynchus nerka ) returning to the Ozernaya River (Kamchatka) was substantially reduced in years when the ocean abundances of Kamchatkan pink ( O. gorbuscha ) and sockeye salmon were high. We found that the density-dependent reduction in sockeye growth on a per-capita basis was greater for sockeye than for pink salmon. However, the overall effect of pink salmon abundance on sockeye growth was greater because of the higher numerical abundance of pink salmon. The strongest statistical relationships were found for sockeye from separate age groups; pooled data combining all age classes were statistically insignificant. We estimate that, if pink salmon were absent, the most strongly affected age group of sockeye salmon (2.1 males) would weigh twice as much at maturity than if pink salmon populations from eastern and western Kamchatka were both simultaneously at peak observed abundances. Trophic competition in the ocean between pink and sockeye salmon can therefore have a significant influence on the productivity of sockeye populations for the most strongly affected age groups. These effects are large enough that they should be explicitly considered in the management of salmon populations.     

Body size trends along vertical and thermal gradients of chum salmon in the Bering Sea during summer

Relationships between the vertical distribution and thermal habitat, and body size of chum salmon Oncorhynchus keta were studied in the Bering Sea in summer using trawl surveys at various depths. Chum salmon abundance decreased with increasing depth, but the patterns of decrease differed between size groups. The abundance of small salmon fell rapidly with depth, whereas that of large salmon decreased gradually to 40 m depth, and abruptly below that. The average fork length of chum salmon collected from each trawl correlated positively with trawl net depth and negatively with water temperature. Since the optimal temperature for growth decreases with body size in this species, the observed body size‐related vertical habitat use by chum salmon may indicate size‐dependent thermal preferences.     

Prespawning migratory behaviour and spawning success of sea-ranched Atlantic salmon, Salmo salar L., in the River Gudenaa, Denmark

The migratory behaviour of sea-ranched Atlantic salmon, Salmo salar L., was analysed by radio-telemetry in the River Gudenaa, Denmark. The main objectives were to: (1) estimate mortality of returning adults through the fjord; (2) observe rate of progression and migratory pattern in the fjord and river; and (3) record whether spawning occurs in the river. Forty-two returning salmon (19 males and 23 females of total body length from 60–97 cm) reared and released as smolts, were caught and equipped with external radio transmitters in the outer estuary of the River Gudenaa in 1994 and 1995. Of the tagged salmon, 18 (43%) were caught in the estuary, four (10%) were not recorded after release and 20 (47%) entered the river. The mean rate of progression through the fjord was 7.6 km d−1 (range 1.4–18.2) in 1994 and 5.4 km  d−1 (range 1.6–17.1) in 1995. Eleven salmon were alive at the onset of the spawning period. Eight were retrieved dead from the river during or after the spawning period; four with empty gonads assumed to be successful spawners, and four with intact gonads. In 1994, unsuccessful spawners (found dead with intact gonads) entered the river earlier and had a longer total migration distance in the river compared to successful spawners. This suggests that spawning success of sea-ranched salmon is associated with time of river entry and river migration length.     

Prey partitioning and use of insects by juvenile sockeye salmon and a potential competitor,threespine stickleback,in Afognak Lake,Alaska

Freshwater growth of juvenile sockeye salmon (Oncorhynchus nerka) depends upon the quality and quantity of prey and interactions with potential competitors in the foraging environment. To a large extent, knowledge about the ecology of lake‐rearing juvenile sockeye salmon has emerged from studies of commercially important runs returning to deep nursery lakes, yet information from shallow nursery lakes (mean depth ≤ 10 m) is limited. We examined seasonal and ontogenetic variation in diets of juvenile sockeye salmon (N = 219, 30–85 mm) and an abundant potential competitor, threespine stickleback (Gasterosteus aculeatus; N = 198, 42–67 mm), to understand their foraging ecology and potential trophic interactions in a shallow Alaska lake. This study revealed that adult insects made up 74% of all sockeye salmon diets by weight and were present in 98% of all stomachs in Afognak Lake during the summer of 2013. Diets varied temporally for all fishes, but small sockeye salmon (<60 mm) showed a distinct shift in consumption from zooplankton in early summer to adult insects in late summer. We found significant differences in diet composition between sockeye salmon and threespine stickleback and the origin of their prey indicated that they also separated their use of habitat on a fine scale; however, the two species showed overlap in size selectivity of zooplankton prey. Considering that aquatic insects can be a primary resource for juvenile sockeye salmon in Afognak Lake, we encourage the development of nursery lake carrying capacity models that include aquatic insects as a prey source for sockeye salmon.     

Heritability estimates for processing and quality traits in common carp (Cyprinus carpio L.) using a molecular pedigree

Heritability estimates with a microsatellite parentage assignment based pedigree are of special interest in common carp cultured under traditional pond conditions. This method reduces common environment effects as all families can be grown immediately after hatching mixed in the same pond. We applied this method to study genetics of growth and processing traits in common carp at market size (1.5 kg and more). The experimental progeny was established by crossing 147 two-year old males and 8 females (six to eight-year old) of Hungarian synthetic mirror carp population. The fish grew up through three vegetation seasons and at the end a sample of 331 fish was examined for biometrical traits (standard length, body weight, relative head length, relative body height, relative body width), percent fat and processing traits (% processed body, % fillets with skin, % fillets without skin). It was shown that sex had a significant effect on most traits: females were larger and fatter than males, and they had both higher percent processed body and percent fillet with skin. Standard length, body weight, percent fat and relative head length had a high heritability (> 0.5), while relative body height, relative body width, percent processed body and fillet yields had a medium heritability (0.2–0.5). We found relatively high positive genetic correlations between body size (standard length and body weight) and percent fat (0.71 and 0.59, respectively), favourable genetic correlations between body size and percent processed body (0.69 for standard length and 0.74 for body weight) and between body size and fillet yields (0.50–0.77). Genetic correlations between body size and body shape (relative head length, relative body height and relative body width) were weak to moderate, thus selection for better growth should have little impact towards more rotund shape. Relative head length had strong negative correlation (− 0.7 to − 0.9) with percent fat, percent processed body and percent fillet yields. This means that indirect selection for reduced relative head length should be effective in improving of fillet yield.     

Life‐history differences of juvenile Chinook salmon Oncorhynchus tshawytscha across rearing locations in the Shasta River,California

Salmon from different locations in a watershed can have different life histories. It is often unclear to what extent this variation is a response to the current environmental conditions an individual experiences as opposed to local‐scale genetic adaptation or the environment experienced early in development. We used a mark–recapture transplant experiment in the Shasta River, CA, to test whether life‐history traits of juvenile Chinook salmon Oncorhynchus tshawytscha varied among locations, and whether individuals could adopt a new life history upon encountering new habitat type. The Shasta River, a Klamath River tributary, has two Chinook salmon spawning and juvenile rearing areas, a lower basin canyon (river km 0–12) and upper basin spring complex (river km 40–56), characterised by dramatically different in‐stream habitats. In 2012 and 2013, we created three experimental groups: (i) fish caught, tagged and released in the upper basin; (ii) fish caught at the river mouth (confluence with the Klamath River, river km 0), tagged and released in the upper basin; and (iii) fish caught at the river mouth, tagged and released in the lower basin. Fish released in the upper basin outmigrated later and at a larger size than those released in the lower basin. The traits of fish transplanted to the upper basin were similar to fish originating in the upper basin. Chinook salmon juvenile life‐history traits reflected habitat conditions fish experienced rather than those where they originated, indicating that habitat modification or transportation to new habitats can rapidly alter the life‐history composition of populations.     

Development of a Piscirickettsia salmonis immersion challenge model to investigate the comparative susceptibility of three salmon species

Piscirickettsia salmonis, the aetiological agent of salmonid rickettsial septicaemia (SRS), is a global pathogen of wild and cultured marine salmonids. Here, we describe the development and application of a reproducible, standardized immersion challenge model to induce clinical SRS in juvenile pink (Oncorhynchus gorbuscha), Atlantic (Salmo salar) and sockeye salmon (O. nerka). Following a 1‐hr immersion in 10⁵ colony‐forming units/ml, cumulative mortality in Atlantic salmon was 63.2% while mortality in sockeye salmon was 10%. Prevalence and levels of the bacterium in kidney prior to onset of mortality were lower in sockeye compared with Atlantic or pink salmon. The timing and magnitude of bacterial shedding were estimated from water samples collected during the exposure trials. Shedding was estimated to be 82‐fold higher in Atlantic salmon as compared to sockeye salmon and peaked in the Atlantic salmon trial at 36 d post‐immersion. These data suggest sockeye salmon are less susceptible to P. salmonis than Atlantic or pink salmon. Finally, skin lesions were observed on infected fish during all trials, often in the absence of detectable infection in kidney. As a result, we hypothesize that skin is the primary point of entry for P. salmonis during the immersion challenge.     

A metapopulation perspective for salmon and other anadromous fish

Salmonids are an important component of biodiversity, culture and economy in several regions, particularly the North Pacific Rim. Given this importance, they have been intensively studied for about a century, and the pioneering scientists recognized the critical link between population structure and conservation. Spatial structure is indeed of prime importance for salmon conservation and management. At first glance, the essence of the metapopulation concept, i.e. a population of populations, widely used on other organisms like butterflies, seems to be particularly relevant to salmon, and more generally to anadromous fish. Nevertheless, the concept is rarely used, and barely tested. Here, we present a metapopulation perspective for anadromous fish, assessing in terms of processes rather than of patterns the set of necessary conditions for metapopulation dynamics to exist. Salmon, and particularly sockeye salmon in Alaska, are used as an illustrative case study. A review of life history traits indicates that the three basic conditions are likely to be fulfilled by anadromous salmon: (i) the spawning habitat is discrete and populations are spatially separated by unsuitable habitat; (ii) some asynchrony is present in the dynamics of more or less distant populations and (iii) dispersal links populations because some salmon stray from their natal population. The implications of some peculiarities of salmon life history traits, unusual in classical metapopulations, are also discussed. Deeper understanding of the population structure of anadromous fish will be advanced by future studies on specific topics: (i) criteria must be defined for the delineation of suitable habitats that are based on features of the biotope and not on the presence of fish; (ii) the collection of long‐term data and the development of improved methods to determine age structure are essential for correctly estimating levels of asynchrony between populations and (iii) several key aspects of dispersal are still poorly understood and need to be examined in detail: the spatial and temporal scales of dispersal movements, the origin and destination populations instead of simple straying rates, and the relative reproductive success of immigrants and residents.     

Diverse juvenile life‐history behaviours contribute to the spawning stock of an anadromous fish population

Habitat quality often varies substantially across space and time, producing a shifting mosaic of growth and mortality trade‐offs across watersheds. Traditional studies of juvenile habitat use have emphasised the evolution of single optimal strategies that maximise recruitment to adulthood and eventual fitness. However, linking the distribution of individual behaviours that contribute to recruitment at the population level has been elusive, particularly for highly fecund aquatic organisms. We examined juvenile habitat use within a population of sockeye salmon (Oncorhynchus nerka) that spawn in a watershed consisting of two interconnected lakes and a marine lagoon. Otolith microchemical analysis revealed that the productive headwater lake accounted for about half of juvenile growth for those individuals surviving to spawn in a single river in the upper watershed. However, 47% of adults had achieved more than half of their juvenile growth in the downstream less productive lake, and 3% of individuals migrated to the estuarine environment during their first summer and returned to freshwater to overwinter before migrating back to sea. These results describe a diversity of viable habitat‐use strategies by juvenile sockeye salmon that may buffer the population against poor conditions in any single rearing environment, reduce density‐dependent mortality and have implications for the designation of critical habitat for conservation purposes. A network of accessible alternative habitats providing trade‐offs in growth and survival may be important for long‐term viability of populations.     

Dynamic in-lake spawning migrations by female sockeye salmon

Abstract –  Precise homing by salmon to natal habitats is considered the primary mechanism in the evolution of population-specific traits, yet few studies have focused on this final phase of their spawning migration. We radio tagged 157 female sockeye salmon ( Oncorhynchus nerka ) as they entered Lake Clark, Alaska, and tracked them every 1–10 days to their spawning locations. Contrary to past research, no specific shoreline migration pattern was observed (e.g., clockwise) nor did fish enter a tributary unless they spawned in that tributary. Tributary spawning fish migrated faster (mean = 4.7 km·day⁻¹, SD = 2.7, vs. 1.6 km·day⁻¹, SD = 2.1) and more directly (mean linearity = 0.8, SD = 0.2, vs. 0.4, SD = 0.2) than Lake Clark beach spawning fish. Although radio-tagged salmon migrated to within 5 km of their final spawning location in an average of 21.2 days (SD = 13.2), some fish migrated five times the distance necessary and over 50 days to reach their spawning destination. These results demonstrate the dynamic nature of this final phase of migration and support studies indicating a higher degree of homing precision by tributary spawning fish.     

Competitive effects between rainbow trout and Atlantic salmon in natural and artificial streams

Competition with non‐native species may impede the restoration of native species, but differences in competitive abilities among intraspecific native populations may make some populations more suitable for reintroduction than others. Here, juvenile Atlantic salmon (Salmo salar) from two allopatric populations (LaHave and Sebago) being used for reintroduction into Lake Ontario were placed into two natural stream sites differing in the presence of ecologically similar rainbow trout (Oncorhynchus mykiss). We assessed the effects of competition in the natural streams on fitness‐related traits and habitat use of the Atlantic salmon. We then compared these effects to those observed in artificial streams from a previous study. Atlantic salmon in natural streams had reduced fitness‐related traits that were associated with suboptimal microhabitats in the presence of rainbow trout, but utilised optimal microhabitats in their absence. In the presence of rainbow trout, the two Atlantic salmon populations exhibited comparable recapture proportions to each other, but the individuals from the Sebago population had better performance (body size and condition) than those from the LaHave population. Responses of both Atlantic salmon populations to competition with rainbow trout were generally similar in both direction and magnitude when compared to results from the artificial stream study. The combined results suggest that native species restoration efforts should be focused on candidate populations that are ecologically suitable to reintroduction environments, as well as on suitable habitats that do not contain exotic competitors. Moreover, this study highlights the value of controlled experiments in artificial environments for predicting fitness‐related performance in natural environments.     

黄鳍金枪鱼5月龄幼鱼形态性状对体质量的相关性及通径分析

该研究以5月龄黄鳍金枪鱼(Thunnus albacores)幼鱼为实验对象,测量并探索其形态性状与体质量的相关性.结果显示,体质量与各形态性状均呈极显著正相关(P<0.01);通径分析发现胸鳍长对体质量直接作用最大(0.507),其次为全长,而上颚长主要通过全长、头长和胸鳍长间接影响体质量;决定系数分析显示,全长、头...     

Diurnal stream habitat use of juvenile Atlantic salmon, brown trout and rainbow trout in winter

Abstract  The diurnal winter habitat of three species of juvenile salmonids was examined in a tributary of Skaneateles Lake, NY to compare habitat differences among species and to determine if species/age classes were selecting specific habitats. A total of 792 observations were made on the depth, velocity, substrate and cover (amount and type) used by sympatric subyearling Atlantic salmon, subyearling brown trout and subyearling and yearling rainbow trout. Subyearling Atlantic salmon occurred in shallower areas with faster velocities and less cover than the other salmonid groups. Subyearling salmon was also the only group associated with substrate of a size larger than the average size substrate in the study reach during both winters. Subyearling brown trout exhibited a preference for vegetative cover. Compared with available habitat, yearling rainbow trout were the most selective in their habitat use. All salmonid groups were associated with more substrate cover in 2002 under high flow conditions. Differences in the winter habitat use of these salmonid groups have important management implications in terms of both habitat protection and habitat enhancement.     

2010年乌苏里江大麻哈鱼的群体结构及其变动

2010年10月4日~21日,对乌苏里江大麻哈鱼（Oncorhynchus keta）洄游繁殖群体渔业资源及其种群结构变动进行了调查研究,共统计标本9160尾。初步结果显示,大麻哈鱼的日洄游数量受水温和溶解氧的影响较大,出现2次洄游高峰期,平均日单船捕捞量为64尾。体长为55~65cm的个体占群体中的74%,其中体长为55~60cm的雌（36.82%）、雄（36.58%）个体在群体中所占比例接近,在体长小于55cm的群体中,雄性（27.17%）个体在本群体中所占比例高于雌性（12.75%）,体长大于60cm的群体中,雌性（48.61%）在本群体中所占比例高于雄性（34.25%）。体质量为2 000~3 000g的个体占群体中的57%,体质量大于2 500g的群体中,雌性（61.73%）个体在本群体中所占比例高于雄性（46.06%）。雌、雄性比为1.09：1,其中体重大于2 500g的雌性在群体所占比例最高为36.06%,其次为小于2 500g的雄性,占28.8%。对比历史数据表明,该渔业产量为近15年的最高值,总体资源量呈增多趋势,群体中个体趋于小型化,雌性个体在群体中处于主导地位。