The relationship between a summer fry (0+) abundance index, derived from semi-quantitative electrofishing, and egg deposition of Atlantic salmon, in the River Bush, Northern Ireland

Atlantic salmon survival in the R. Bush (N. Ireland) from egg to summer 0+ was inversely density-dependent on egg deposition ( P <0.05). A stock-recruitment relationship derived from egg deposition and summer 0+ abundance index data was compared to that derived from adult and smolt counts based on total trapping. Fitted Ricker curves indicated maximum recruitment at around 2.35 million eggs and 2.46 million eggs for 0+ index and smolt count methods, respectively. Salmon 0+ abundance index data from semi-quantitative electrofishing could be obtained with relatively little effort, and used to derive whole-river stock-recruitment relationships on rivers where only adult count or some other estimator of parental stock is available. The derivation and expression of spawning targets from stock/recruitment relationships is discussed with reference to the R. Bush data.     

Seasonal variation in density of juvenile Atlantic salmon in fluvial shoreline habitats of a large subarctic river

The densities of Atlantic salmon fry (0+ years) and parr (1+ years and older) in shoreline habitats of the large River Teno watercourse and its tributary, the River Utsjoki generally fluctuated considerably, showing an increase from early summer towards late August and a subsequent decline towards autumn. The seasonal pattern of variation in density was more distinct for parr than for fry. In the period between late July and early September, parr density followed a sinusoidal curve, being highest in late August and lowest in early August and in September. Fry density had a weaker seasonal profile than parr, being highest in late August and in early September. Frequency distributions of the parr age groups (1+, 2+ and 3+ years) were mainly independent of the sampling month.     

Periodic habitat loss alters the competitive coexistence between brown trout and bullheads in a small stream over 34 years

1. Changes in the population density of juvenile sea trout Salmo trutta L. and bullheads Cottus gobio L. were compared in a small stream over 34 years. Both species have a similar diet and obviously live in the same general habitat. Habitat loss was most marked in seven summer droughts: severest in 1976, 1983, 1984, 1995, and less severe but followed by autumn droughts in 1969, 1989 and 1993. The contrasting effects of habitat loss on the two species were examined. 2. For both species, the Ricker curvilinear model significantly fit (P < 0.001) the relationship between initial egg density and survivor density for successive life stages, even though egg densities were much lower for bullheads than trout. These analyses provided evidence for density-dependent population regulation and also identified extreme outliers, most being for year-classes affected by summer droughts. 3. The variable effects of changes in habitable area (= % wettable area in sampling section) were quantified by using the residuals, each residual being the absolute value expressed as a percentage of the expected value from the Ricker curve. Significant relationships between the residuals and habitable area showed that habitat loss had a marked effect on survivor density, this being negative for 0+ and 1+ trout, and positive for 0+, 1+ and 2+/3+ bullheads. 4. Therefore, during periods of habitat loss in the summer months, bullhead density increased at the expense of trout density. Low flows and a decrease in wettable area were associated with a marked reduction in habitat quality for drift-feeding trout and an increase in habitat quality, and perhaps also quantity, for benthic-feeding bullheads. This case study shows that, during a major perturbation, the relationship between the densities of two species can change markedly in favour of the less numerous species. The competitive coexistence between the two species is therefore a dynamic process that changes through time with periodic changes in the environment.     

Stock and recruitment and inversely density-dependent growth of salmon, Salmo salar L., in a Scottish stream

For the 6 years for which detailed data are readily available, estimates of the survival of emergent fry of Atlantic salmon, Salmo salar L., to the first and second autumns at a site on the Shelligan Burn are consistent with the dome-shaped Ricker model with about 11 emergent fry m⁻² maximizing recruitment. The data are not satisfactorily fitted by the asymptotic Beverton and Holt model. A possible mechanism, which results from the observed inversely density-dependent growth, is discussed briefly.     

Effect of habitat types on survival, spatial distribution and production of an allopatric cohort of Atlantic salmon, Salmo salar L., under conditions of low competition

An allopatric cohort of Atlantic salmon, Salmo salar L., introduced to a small previously fishless stream was studied from parr to the smolt stages. In May 3900 0+ parr (mean total length 30mm) were planted at three different densities in habitats with slow, intermediate and fast water velocities. During the first year, high mortality occurred during the first 7 weeks after planting in May, and in September–October. Survival from May 1985 to April 1986, before the smolt emigration, was24.8%. The smolt yield 1 year after planting was 15.5%. It is suggested that the high survival was caused by low competition. Most of the redistribution of the fish took place during the first months. Type of planting habitat affected the timing of redistribution. The parr left slow-flowing, deep habitats with fine substrate soon after planting, while redistribution was slowest in the fastest flowing habitats with coarse substrate. The observed avoidance of slow, deep habitat types in the absence of interspecific competition, suggests that this may be a fixed behavioural response, and not due to competition. Long movements, up to 800 m, were recorded only within the first 7 weeks after planting. The effect of planting densities on population density was most pronounced immediately after planting in the fast and also intermediate habitats. Planting density effects declined and were not detectable after 1 year. The effect of habitat type on fish numbers and biomass was pronounced irrespective of planting densities. Growth was fastest in the intermediate habitat, and at the lower planting densities. Production was 7.2 g m⁻² the first summer-autumn. Due to smolt emigration, few fish remained in the stream the second summer-autumn, and the production was 1.0 g m⁻².     

Population regulation in adult, but not juvenile, resident trout (Salmo trutta) in a Lake District stream

1. Unlike a neighbouring sea-trout population that showed strong density-dependent survival, a resident trout population ( Salmo trutta L.) showed simple proportionate survival in the early life-stages. However, this persistent population fluctuated within narrow limits. Mature adults, especially during spawning, were the only possible life-stage left in which regulation might occur. 2. An October census, just prior to spawning, was made at five sites (total area 300 m ² ) from 1965 to 1983. Gravel nests (redds) associated with females of known size were excavated outside these sites to obtain a power-function relationship between egg density per redd and female length (range 181–280 mm, n = 26). This relationship and the census data for females (range 186–284 mm) were used to estimate egg densities in each year-class. 3. The census data for the early life-stages (0+, 1+, 2+ trout) confirmed proportionate survival with no evidence for density-dependent regulation. In contrast, the number of spawning females produced in each year-class was strongly density dependent on the initial number of females that laid eggs at the start of the year-class. Similarly, total egg production in each year-class was density dependent on initial egg density. 4 Both relationships were well described by the Ricker and Beverton-Holt stock-recruitment models (P < 0.001) and the goodness-of-fit was similar for both models. This study is probably the first to provide clear evidence for fish population regulation in the adult, rather than the juvenile, stage.     

Effects of life history variation on size and growth in stream-dwelling Atlantic salmon

A large size variation amongst life histories for stream-dwelling Atlantic salmon Salmo salar was found and the relative effect of life histories on size varied over time. As early as December (age 0+ years), fish that later smolted at age 2+ years were significantly larger than fish that did not smolt at age 2+ years. In contrast, there were no mass differences at age 0+ years between fish that would mature or not at age 1+ years (October). The mass differences between smolts and non-smolts persisted until smolting, and differences between mature and immature fish first appeared in May (age 1+ years). Following September (age 1+ years), there was also a significant interaction between smolting and maturity. Previously mature and immature age 2+ year smolts were not significantly different in size, but immature age 2+ year non-smolts were much lighter than mature age 2+ year non-smolts. Based on mass differences, the apparent 'decision' to smolt occurred c . 5 months before (winter, age 0+ years) the decision to mature (late spring, age 1+ years). In addition to strong seasonal growth variation, sizes of freshwater Atlantic salmon were largely structured by the complex interaction between smolt-age and maturity.     

Predicting fluctuations in the size of newly emerged sea-trout fry in a Lake District stream

The objective was to predict interannual fluctuations in the size of sea-trout fry when they emerged from the redd, using models developed from field data for 70 excavated redds (≥three per year), and from experimental data on egg and alevin development at 30 constant temperatures in the laboratory (range 1·5—10·5) C with 100 naturally fertilized eggs at each temperature). Egg weight increased with female length and also with the number of eggs laid in a redd, both relationships being well described by a power function. Early spawners were the largest females laying the largest and most numerous eggs, whilst late spawners were the smallest females laying the smallest and least numerous eggs, with middle spawners being intermediate between these two extremes. Mean values for egg weight and number of eggs per redd were obtained for these three groups. The numbers of early, middle and late spawners for each year of a 30-year study and the mean values from the excavated redds were used to estimate weighted means for the number of eggs per unit area and egg weight. Mean values varied considerably between years (30-year ranges: 518–7964 eggs per 60 m2; 112–138 mg wet weight). In the laboratory, mean weights of newly hatched alevins and newly emerged fry were both related positively to mean egg weights. Alevin and fry mean weights were independent of the number of days required for 50% of the eggs to hatch or fry to emerge. Models described in a previous paper formed the basis of those used to predict fry weights over the emergence period. Model predictions were validated by field data for the whole emergence period in 8 years (1967–1971, 1974, 1975, 1980), and by pre-fry weights on single dates in 21 years (1967–1987). As pre-fry densities on these single dates were very similar to egg densities for the same year class, mortality in the egg and alevin stages was very low. The chief objective was therefore fulfilled, and the extent of interannual fluctuations for the 30-year study showed some variation in mean fry weight (30-year ranges: 153–193 mg for both the whole emergence period and the date on which 50% of fry emerged) but a progressive decrease in fry weight through the emergence period. Possible reasons for this variation are discussed, and it is concluded that the size of the female spawners is the dominant factor.     

Linking climate change projections for an Alaskan watershed to future coho salmon production

Climate change is predicted to dramatically change hydrologic processes across Alaska, but estimates of how these impacts will influence specific watersheds and aquatic species are lacking. Here, we linked climate, hydrology, and habitat models within a coho salmon (Oncorhynchus kisutch) population model to assess how projected climate change could affect survival at each freshwater life stage and, in turn, production of coho salmon smolts in three subwatersheds of the Chuitna (Chuit) River watershed, Alaska. Based on future climate scenarios and projections from a three‐dimensional hydrology model, we simulated coho smolt production over a 20‐year span at the end of the century (2080–2100). The direction (i.e., positive vs. negative) and magnitude of changes in smolt production varied substantially by climate scenario and subwatershed. Projected smolt production decreased in all three subwatersheds under the minimum air temperature and maximum precipitation scenario due to elevated peak flows and a resulting 98% reduction in egg‐to‐fry survival. In contrast, the maximum air temperature and minimum precipitation scenario led to an increase in smolt production in all three subwatersheds through an increase in fry survival. Other climate change scenarios led to mixed responses, with projected smolt production increasing and decreasing in different subwatersheds. Our analysis highlights the complexity inherent in predicting climate‐change‐related impacts to salmon populations and demonstrates that population effects may depend on interactions between the relative magnitude of hydrologic and thermal changes and their interactions with features of the local habitat.     

The effect of stocking with 0+ year age‐class Atlantic salmon Salmo salar fry: a case study from the River Bush,Northern Ireland

An enhancement programme based on stocking 0+ year age‐class Atlantic salmon Salmo salar, conducted in the River Bush, Northern Ireland, U.K. over the period 1996–2005, was reviewed with reference to the performance and biological characteristics of wild fish. Wild ova to 0+ year fry (summer) survival was c. 8% with subsequent wild 0+ year fry‐to‐smolt survival c. 9%. Stocked unfed 0+ year juveniles gave c. 1% survival to smolt whilst fed 0+ year S. salar stocked in late summer exhibited survival at c. 5%. Stocking with unfed and fed fry contributed to increased smolt production and helped attain local management objectives between 2001 and 2005. Significant differences in biological characteristics were observed between wild and stocked‐origin fish. Wild‐smolt cohorts were dominated by 2+ year age‐class fish on the River Bush whilst smolts originating from fed fry mostly comprised younger 1+ year individuals. The mean mass of 1+ year smolts derived from stocked fed fry was significantly lower than that of wild 1+ year smolts, although these differences were not evident between older age classes. Differences in run timing between wild smolts and smolts derived from stocked fry were also apparent with the stocked‐origin fish tending to run earlier than wild fish. Although the stocking exercise was useful in terms of maximizing freshwater production, concerns over the quality of stocked‐origin recruits and the long term consequences for productivity are highlighted.     

Intraspecific competition and density dependence in an Ephestia kuehniella–Venturia canescens laboratory system

A model host-parasitoid system of Ephestia kuehniella and Venturia canescens was used to examine the influence of host and parasitoid density on host and parasitoid life-history parameters via a two-way factorial experimental design (5 initial host densities×3 parasitoid densities). In the absence of parasitoids, E. kuehniella experienced scramble-type competition with reduced growth, diminished adult size and a subsequent fecundity trade-off for mortality. The mortality that did occur was confined to the late larval and pupal stages. In the presence of parasitoids attacking the late larval stage, competition changed from scramble for food to contest for enemy-free space, with hosts escaping parasitism being small with low fecundity and reduced egg size, and with parasitoid adult size inversely dependent on host density. Total insect emergence (host+parasitoid), a measure of the influence of host resource competition on survivorship, exhibited a threshold effect as a function of initial host density; the threshold value was increased to a higher initial host density in the presence of parasitoids. Models of host self-limitation were fitted to the data, with the generalized Beverton-Holt model that incorporates a threshold effect providing the best fit, and the Ricker model with no threshold providing a very poor fit to the data.     

Latitudinal clines in life-history variables of anadromous brown trout in Europe

Based on published data, we reviewed clinal variations in life-history characteristics of anadromous brown trout Salmo trutta from 102 European rivers at latitudes between 54 and 70° N. Growth rate in fresh water, mean smolt age, mean sea age at first sexual maturity, proportion of repeal spawners among adults, longevity, and length of adult life span exhibited latitudinal clines. Brown trout grew faster in fresh water, smolted and matured younger, lived fewer years but spawned more times in the south than in the north. The life-history traits studied were often correlated. Longevity, smolt age and sea age at maturity were negatively and smolt length and proportion of repeat spawners among adults were positively correlated with growth rate in fresh water. Longevity was positively correlated with smolt age and sea age at maturity. The latter also increased with increasing smolt age. None of these significant correlations among life history variables, except for those between smolt age and parr growth and proportion of repeat spawners and parr growth, are latitudinal effects. We do not know to what extent the latitudinal variation in life–history traits reflects phenotypic plasticity and to what extent it is caused by inherited differences among populations.     

Changes in smolt traits of Atlantic salmon (Salmo salar Linnaeus, 1758) and linkages to parr density and water temperature

Smolt traits (length, age) and timing of smolt migration of wild Atlantic salmon, Salmo salar L., were investigated in the Simojoki River, northern Baltic Sea. The aim was to determine whether they responded to changes in parr length, parr density and temperature from 2000 to 2014. Annual electrofishing surveys and smolt numbers determined parr densities by springtime trapping in the river mouth. During the smolt trapping period captured parr and smolts were aged from scales. Water temperature was measured daily. Mean length decreased from 137 mm (TL) to 129 mm among 2‐year‐old smolts, and from 150 mm to 139 mm among 3‐year‐olds. Median date of the smolt migration was 10 days earlier, from early June to late May during the study period, linked to the rise in air temperature in May at the nearby Kemi‐Tornio airport. However, the median day temperature and the mean daily water temperatures during the second (Q₂) and third (Q₃) migration quartiles did not change. This implied that migration began when a suitable water temperature was reached, independent of the date.     

Evolutionary responses of Drosophila melanogaster life history to differences in larval density

The study examined the effects of evolution at two different larval densities on pre-adult and adult fitness traits. Five replicate selection lines each were cultured at either 50 or 150 larvae per vial, avoiding selection on development time, age at breeding or for adaptation to adult density, one or more of which factors has been a confounding variable in previous studies. Low density selection lines evolved extended development times at both growth densities. The extended development times were associated with greater adult body size at the lower growth density only, and particularly in females. The lines did not differ significantly in larval competitive ability at either growth density. At neither growth density did the early adult fertility of females or the lifespan of either sex differ between the lines from the two selection regimes, but at the lower growth density the late fertility of low density line females was significantly enhanced. The results suggest that larval density does have important effects on the expression and resolution of life history trade-offs in Drosophila melanogaster, but that these may be somewhat different from those reported in previous studies.     

The survival, growth and diet of pike fry, Esox lucius L., stocked at different densities in experimental ponds

Yolk-sac pike fry were stocked at densities of 0.74 – 81.4 m⁻² in two ponds, each divided into eight sectors (mean area 155.8 m²). Growth and survival were recorded from May to December 1985. The growth rates were variable within each sector. The size-range of sampled fish increased throughout the year, but showed no significant correlation with density. Fry survival was initially density-independent but switched by late June/July to density-dependence, ranging from 0.5 to 43.6% of initial stocking numbers. The highest mean daily mortality rates occurred during May-July. The final survival in December ranged between sectors from 0.059 to 11.25% of the starting stock densities. The final biomass per unit area of pike surviving in December was not related to initial stocking density. In Pond 1 the mean biomass produced was 2.21 gm⁻² and in Pond 2 was 3.49gm⁻².
Pike fry < 30 mm fed only on invertebrates; those 30–100 mm took a wide range of invertebrates, cyprinids. sticklebacks and other pike. Cannibalism occurred at most densities between 5.45 and 81.4 fish m⁻².
Where attempts are made to increase pike production in managed populations by releasing small fry, an upper stock density of 5 fry m⁻² is advised if large, density-dependent mortalities are to be avoided.     

Marine post-smolt growth and age at maturity of Atlantic salmon

The annual variation in sea-age of maturation for a hatchery dependent stock of Atlantic salmon was compared to variation in post-smolt growth as evidenced by circuli spacing patterns. The proportion of returns of 1-seawinter (1 SW) and 2 SW salmon and the fraction of the smolt year class or cohort that maturated as 1 SW fish, were compared to seasonal growth indices determined from circuli spacing on the scales of smolt class survivors returning as 1 SW and 2 SW spawners. Using image processing techniques, we extracted inter-circuli distances from scales from 2244 recaptured fish. Spacing data for the first year at sea were collected and then expressed as seasonal growth indices for the spring period, when post-smolts first enter the ocean; the summer, when growth appears maximal; and winter, when growth appears to be at a minimum. In general, circuli spacings were wider for 1 SW than for the 2 SW returns of the same smolt cohort. The 1 SW fraction was significantly and positively correlated with late summer growth, suggesting that growth during this season is pivotal in determining the proportion of a smolt class that matures early.     

Is there a threshold size regulating seaward migration of brown trout and Atlantic salmon?

We investigated the influence of variation in body size and growth rate on age of smolting in Atlantic salmon and brown trout in four different Norwegian rivers. In Atlantic salmon smolt ages varied between 2 and 6 years, and in brown trout between 2 and 7 years. Smolt age was negatively correlated with parr growth, and positively correlated with smolt size. Age at smolting was more variable in the two northern than the two southern rivers. Smolt sizes and ages were also more variable in brown trout than in Atlantic salmon. Based on the observed variation in smolt size and age, we reject the hypothesis that a threshold size alone regulates age at smolting. Within populations smolt age depends on growth rate so that fast-growing parr smolted younger and smaller than slow-growing parr. We hypothesize that smolt size and age is a trade-off between expected benefits and costs imposed by differences in individual growth rate.     

Interspecific relationships between emerging Atlantic salmon, Salmo salar, and coho salmon, Oncorhynchus kisutch, juveniles

Interspecific relationships between Atlantic salmon and coho salmon were studied at early life stages in laboratory and semi-natural stream channels. During emergence, the survival and dispersal patterns were similar for the two species in single or mixed populations. Survival of Atlantic salmon fry was reduced in the presence of older coho fry. However, no predation was observed. Microdistribution differed between the two species, with Atlantic salmon fry more numerous in riffles when coho were present.
Coho juveniles had a pelagic and gregarious distribution, in contrast to the benthic behaviour of the Atlantic salmon. In laboratory streams, Atlantic salmon fry moved out or adopted a subordinate cryptic behaviour which allowed them to escape predation while negatively affecting their growth.     

Recruitment variability of resident brown trout in peripheral populations from southern Europe

1. Population regulation was studied for seven consecutive years (1992–98) in five rivers at the periphery of the distribution of Salmo trutta, where the fish were living under environmental constraints quite different from those of the main distribution area. 2. Recruitment is naturally highly variable and the populations had been earlier classified as overexploited. Thus we expected that densities of young trout in most populations would be too low for density‐dependent mortality to operate. We tested this by fitting the abundance of recruits to egg densities over seven consecutive years (stock–recruitment relationship), and used the results to judge whether exploitation should be restricted in the interests of conserving the populations. 3. The density of 0+ trout in early September, as well as the initial density of eggs and parents, varied greatly among localities and years. The data for all populations fitted the Ricker stock–recruitment model. The proportion of variance explained by the population curves varied between 32% and 51%. However, in most cases the observations were in the density‐independent part of the stock–recruitment curve, where densities of the recruits increased proportionally with egg densities. 4. Our findings suggest that recruitment densities in most rivers and years were below the carrying capacity of the habitats. Although density‐dependent mechanisms seemed to regulate fish abundance in some cases, environmental factors and harvesting appeared generally to preclude populations from reaching densities high enough for negative feedbacks to operate. The findings thus lend support to Haldane’s (1956) second hypothesis that changes in population density are primarily due to density‐independent factors in unfavourable areas and areas with low density due to exploitation. Exploitation should be reduced to allow natural selection to operate more effectively.     

The critical-period concept for juvenile survival and its relevance for population regulation in young sea trout, Salmo trutta

An example of density-dependent regulation is provided by a long-term investigation (1966-present) of a population of migratory trout (estuarine and sea trout), Salmo trutta L., in a Lake District stream. Evidence for the concept of a critical period for the survival of young fish is briefly reviewed and found to be rather equivocal. The concept is, however, relevant to the trout population. Loss rates were high before but low after a critical survival time ( t_c days after fry emergence) that varied between year-classes (range 33-70 days) and was inversely density-dependent on egg density. Survivor density and loss rates were strongly density-dependent on egg density before t _c, but proportionate survival with stable loss-rates occurred after t _c. Some trout established feeding territories soon after emergence and the number of fish without territories decreased from a high initial value to a negligible value at t _c. Fish size at t_c was not constant but increased as t _c increased. The range of t _c for the different year-classes was similar to that for survival times of unfed fry in the laboratory. A new stock-recruitment model, incorporating t _c, has been developed for the trout population and shown to be related to the model (Ricker curve) used in the long-term study. The critical time can also be regarded as the critical age for survival in young trout; this concept may be relevant to other fish species.