Density‐dependent territory size and individual growth rate in juvenile Atlantic salmon (Salmo salar)

Whether territoriality regulates population size depends on the flexibility of territory size, but few studies have quantified territory size over a broad range of densities. While juvenile salmonids in streams exhibit density‐dependent mortality and emigration, consistent with space limitation, there has been relatively little study of how territory size and individual growth rate change over a broad range of densities, particularly in field experiments. Consequently, we manipulated the density (range = 0.25–8 m^?2) of young‐of‐the‐year (YOY) Atlantic salmon (Salmo salar) in mesh enclosures erected in a natural stream to test whether (i) territory size is fixed, decreases continuously or decreases towards an asymptotic minimum size as density increases; and (ii) individual growth rate decreases as a negative power curve with density as in observational field studies. Territory size decreased with increasing density, consistent with an asymptotic minimum size of about 0.13 m² for a 5‐cm fish. Individual growth rate also decreased with density, although the magnitude of decrease was steeper than in observational studies. Our results suggest a limit to how small territories can be compressed, which will set the upper limit to the local density in a habitat. The density‐dependent changes in territory size and individual growth rate will both play a role in the regulation of stream salmonid populations.     

Alternative use of food resources causes intra-cohort variation in the size distribution of young-of-the-year perch (Perca fluviatilis)

Abstract –  Body sizes of young-of-the-year (YOY) perch ( Perca fluviatilis ) at the end of their first summer are extremely variable and range in different studies between 4 and 15 cm. To analyse whether size divergences in YOY perch may be attributed to alternative use of food resources, adult perch were stocked into two previously fishless ponds and growth, size distribution and food intake of the YOY perch were recorded. In addition to perch, adult bream ( Abramis brama ) were introduced to produce juvenile bream that could serve as a food resource for YOY perch. The body sizes of YOY perch at the end of the experiment ranged from 32 to 168 mm with a bimodal size distribution. The combination of stomach content analyses and stable isotope signatures revealed that the small size cohort were planctivorous/benthivorous while the large size cohort was piscivorous/cannibalistic. Results implicated that different feeding behaviour contributed to the size divergences in YOY perch and that the extreme growth of the large size cohort was induced by piscivory.     

Long-term changes in the smolt size and age of Atlantic salmon, Salmo salar L., in a northern Baltic river related to parr density, growth opportunity and postsmolt survival

Abstract –  The size of wild Atlantic salmon ( Salmo salar L.) smolts in 1972–2004 was studied in relation to parr density, smolt age, growth opportunity and postsmolt survival in the Simojoki River. There was a significant negative regression between the annual mean smolt size and the density of wild >1 year parr in the previous autumn, but not between the annual mean smolt size and age. The density of reared parr released into the river or the growth opportunity, based on the day length and air temperature during the previous summer, did not affect the size of wild smolts. The data on postsmolt survival based on recaptures of Carlin-tagged smolts showed a significant positive relationship ( P  < 0.01) between the survival of postsmolts and the annual size of wild smolts. It is hypothesised that the increased density of wild >1 year parr could have contributed to the decreased smolt size since the 1990s, and the reduced size of wild smolts could be included among the factors resulting in their declined postsmolt survival in the Baltic Sea.     

Zooplanktivore fish body growth responses to browning-induced light limitation vary over ontogeny,but not with fish density

Ongoing climate change is leading to browning of many lakes and coastal areas, which can impair fish body growth and biomass production. However, whether and how effects of light limitation caused by browning on fish body growth vary over early ontogeny is unknown. In this study, we set up a mesocosm experiment to test whether roach (Rutilus rutilus) body growth responses to browning depend on body size, and if findings are robust over roach densities. We also studied a potential mechanism for size-specific responses by conducting an aquaria experiment to test if size-specific prey selectivity in roach changes with browning. We found that roach body growth responses to browning-induced light limitation vary over ontogeny (independent of roach density), negatively affecting body growth of young-of-the-year (YOY) but not of 1-year-old individuals. We also show that this difference in growth response is likely a consequence of browning-induced alterations in zooplankton community composition and variation in prey selectivity between YOY and 1-year-old fish. This suggests that we should account for the diverse effects of browning over fish ontogeny, mediated via altered prey composition and ontogenetic changes in prey preference, when assessing overall impacts of browning on aquatic ecosystems.     

Field observations and laboratory experiments on growth and lipid content of young-of-the-year perch

Abstract –  Differences in growth and allocation of energy of young-of-the-year (YOY) perch ( Perca fluviatilis ) were studied in relation to both quantity and quality of food. Feeding on fish and Mysidacea, the mean size of YOY perch after the first growing season in Lake Pfeiffer was 152 mm. In contrast, YOY perch grew to 107 mm in Lake Speldrop and 83 mm in Lake Reeser Meer while mainly feeding on zooplankton. In correlation with food uptake and growth, the lipid content of YOY perch was significantly highest in Lake Pfeiffer and lowest in Lake Reeser Meer, assuming that YOY perch from Lake Pfeiffer should be able to survive starvation periods during their first winter better then those from Lake Speldrop and Lake Reeser. While feeding on fish, mean highest growth rates were 1.31 mm·day⁻¹ in the laboratory. Growth in weight started when food uptake exceeded 66 J·g WW⁻¹·day⁻¹, while an increase of lipid content required food input of 175 J·g WW⁻¹·day⁻¹. Increased growth potential associated with the availability of energetically profitable food is assumed as an important factor that could have far-reaching consequences for the fish community.     

Density‐dependent diet composition of juvenile Atlantic salmon (Salmo salar)

Abstract – Decreases in body growth with increasing population density may be caused by reduced prey delivery rate. However, changes in food quality because of an increasing inclusion of suboptimal prey in the diet may also contribute to such effects. Here, we test for density‐dependent diet composition by creating spatial variation in Atlantic salmon young‐of‐the‐year (YOY) density in three replicate streams and obtain detailed information on individual positions (±1 m) and diet. Diet breadth with respect to prey size increased with increasing local density for the two most common prey types (Chironomidae and Ephemeroptera). For the largest prey type (Ephemeroptera), there was also an increase in mean prey size with increasing density, suggesting that YOY salmon preferentially utilise the smaller prey among those available. According to optimal foraging theory, changes in diet with increasing local density are likely to come at an energetic cost and hence may contribute to the commonly observed density‐dependent growth of juvenile salmonids.     

Deep habitats are important for juvenile Atlantic salmon Salmo salar L. in large rivers

Linnansaari T, Keskinen A, Romakkaniemi A, Erkinaro J, Orell P. Deep habitats are important for juvenile Atlantic salmon Salmo salar L. in large rivers. Ecology of Freshwater Fish 2010: 19: 618–626. © 2010 John Wiley & Sons A/S Abstract – Juvenile Atlantic salmon were studied by underwater video surveillance and self contained underwater breathing apparatus‐diving in deep (i.e. >1.0 m), fast flowing areas of two large river systems (River Teno, River Tornionjoki) in northern Finland. Both video and diving data indicated that young‐of‐the‐year (YOY) salmon (0+) and salmon parr (>0+) readily utilised habitats deeper than 1 m. Young‐of‐the‐year salmon and parr were observed through a range of 0.5–1.9 m and 0.4–2.2 m, respectively. A negative linear relationship between the mean abundance of YOY salmon and mean depth was noted from the diving transects. Salmon parr were similarly abundant throughout the range of depths studied. Video data showed that deep habitats were used throughout the summer (June–August). It was concluded that deep, fast‐flowing areas in large rivers may constitute a significant habitat resource for juvenile salmon that has not been traditionally accounted for when estimating salmon production.     

Historical changes in age structure of Atlantic salmon, Salmo salar L., in the River Eo, northern Spain

Abstract Variations in the age structure and number of Atlantic salmon, Salmo salar L., running into the River Eo in northern Spain were assessed from catch records of sport fisheries for the period 1949 to 1991. The analysis focused on two periods (1951–1960 and 1981–1991) for which more complete data on fish number, size and age were available. Over the whole period, no evidence for a decline in the total salmon run was found, but the last few years revealed significant changes in both the number and age structure of salmon caught. The catches were mainly composed of multi-sea-winter salmon, and this did not change from the 1950s to the 1980s as the proportion of grilse remained unchanged. However, since 1980, there was a marked decline in both the proportion of 3-sea-winter salmon (versus 2-sea-winter fish) and the frequency of previous spawners among the multi-sea-winter salmon. These changes were probably linked because most of the previous spawners re-entered the river as 3-sea-winter or older salmon. River age of salmon caught in the 1950s was low and constant (about 90% of the catch had smolted at one year) but the mean age of smolting increased in the 1980s.     

Diet overlap between young-of-the-year perch, Perca fluviatilis L., and burbot, Lota lota (L.), during early life-history stages

Abstract –  The diet overlap between young-of-the-year (YOY) perch and burbot in the pelagic zone of Lake Constance during spring and summer was investigated in relation to gape size limitation. Because perch were larger and grew faster than burbot during their early life history, perch overcame gape size limitation for various zooplankton taxa earlier than burbot. The interspecific diet overlap between perch and burbot decreased continuously until June, but increased slightly, when burbot became able to feed on large daphnids by the beginning of July. All zooplankton taxa could be found within perch stomachs by the middle of June, when perch overcame gape size limitation for large cladocerans. However, there was an increasing tendency for individual diet specialisation of perch, as the similarity between individual perch stomach contents decreased. In contrast, the similarity between individual burbot stomach contents remained at almost 50% until the end of August, indicating that all burbot rely on cyclopoid copepods during their entire pelagic life-history stage. Because by July YOY perch are more abundant by one order of magnitude in the pelagic zone than burbot, YOY perch may be more affected by intraspecific competition than by interspecific competition with burbot. Burbot, on the other hand, may evade strong competition with YOY perch by performing diel vertical migrations, thus being restricted to feed on migrating zooplankton prey.     

Reproductive strategies in small populations: using Atlantic salmon as a case study

Abstract –  Wild salmonid populations with only a few breeding adults may not exhibit a significant reduction in genetic variability compared with larger populations. Such an observation suggests that effective population sizes are larger than population size estimates based on direct adult counts and/or the mating strategy maximises outbreeding, contributing to increased heterozygosity. In the case of wild Atlantic salmon Salmo salar populations, stratification by age classes and sexes on the spawning grounds avoids inbreeding and increases genetic variability. We studied the breeding composition of four Spanish salmon populations. Over a 7-year period we concluded that the probability of within-cohort mating is very low: females generally reproduce after two sea-winters whereas males reproduce mostly as one sea-winter ( grilse ) and/or mature parr. Considering different levels of contribution of mature parr to spawning derived from field surveys, we developed a simple model for estimating effective population sizes and found that they doubled with 65% parr contribution expected for rivers at this latitude (43°N), and ranged from 100–800 individuals. The effect of between-cohort mating was modelled considering different ranges of differences in allele frequencies between cohorts and resulted in 28–50% increases in heterozygosity when considering a 65% parr contribution. The complex mating strategy of Atlantic salmon contributes to explain the high levels of genetic variability found for small populations of this species. This model can probably be extended to other animal species with mating strategies involving different cohorts.     

Annual variability in survival of sea-ranched Baltic salmon, Salmo salar L: significance of smolt size and marine conditions

Abstract Carlin tagging data for 1980–1991 were used to examine the influence of smolt size and feeding conditions on the post-smolt survival of ranched River Neva salmon, ( Salmo salar L.), in the the Gulf of Finland, which is the native feeding area of the stock, and in the Bothnian Sea, where the stock has been introduced. Because of better feeding conditions, the survival rates were higher and less variable in the Gulf of Finland than in the Bothnian Sea. In the Bothnian Sea, the annual variability in survival decreased and the mean value increased with increasing smolt size from the smallest (14–16 cm) to the largest (28–30 cm) size classes. The survival was positively correlated with growth rates, food resources and sea-surface temperatures. This suggests that in the Bothnian Sea the annual variability in survival is mainly because variable marine conditions affect growth rates, and, thus, the vulnerability of the post-smolts to size-dependent predation. In the Gulf of Finland, the survival advantage of large initial size and rapid growth was counteracted by size-selective post-smolt mortality from fishing. The increase in the survival rate with increasing smolt size levelled off at 22 cm, and the correlations between survival, growth and the indices of feeding conditions were mostly insignificant. For large smolts, some negative correlations were recorded, suggesting that the relative significance of mortality from fishing may even exceed that of size-dependent natural mortality. The implications of the results for management are discussed.     

Egg size differentiation among sympatric demes of brown trout: possible effects of density-dependent interactions among fry

Abstract –  This study investigates differentiation in egg size among five sympatric brown trout ( Salmo trutta (L.)) demes. We explore a hypothesis predicting high density-dependent interactions among juveniles to favour large eggs by sampling closely located (<100 m) deme pairs with low and high fry abundances. A mancova model fitted the egg size versus egg number relationship as a function of large-scale spatial habitat heterogeneity (basin) and maternal phenotype revealed that demes have significantly different egg size versus fecundity relationships and that the differentiation is mainly due to interdeme variation in egg size. Fry density was significantly and positively associated with egg size and a post-hoc test indicated egg size to be significantly greater in high-density than low-density tributaries. The data is consistent with the density-dependent hypothesis and suggest that reproductive investment can diverge over small geographic distances, potentially in response to environments favouring greater investment in offspring quality.     

A descriptive and experimental study of microhabitat use by young-of-the-year benthic stream fishes

Abstract– We studied microhabitat use of young-of-the-year (YOY) benthic stream fishes ( Cottus, Etheostoma, Percina ) using underwater surveys conducted in the Little River of Tennessee, United States. At the upstream site, only Cottus was present and its density was low. Further downstream all genera were present, and YOY density was high and concentrated in deep water habitats. Microhabitat use by the species differed from microhabitat availability, with most species using deep, slow-moving water with silt-free, coarse substrates that lacked riverweed, Podostemum ceratophyllum. Despite differences in microhabitat use, there was substantial overlap among species. Because YOY were positively associated with coarse substrates and coarse substrates were uncommon in deep water, we tested their importance by adding cobbles to 3 deep water habitats. We found that YOY benthic fishes colonized areas with cobbles but not sand. After 2 months, however, many of the cobble areas were covered by sediment. These data suggest that YOY fish are substrate-limited in deep water habitats and that the hydrological properties of pools preclude establishment of permanent instream cover. It is unclear why YOY occur in areas with little instream cover.     

Assessment of winter size‐selective mortality of young‐of‐the‐year Atlantic salmon (Salmo salar) using otolith microstructure analysis*

Abstract – The objectives of this study were, first, to assess the usefulness of otolith microstructure analysis to examine winter size‐selective mortality of young‐of‐the‐year (YOY) Atlantic salmon and, secondly, to validate various hypotheses relating to the dynamics of two populations with different winter survival. By examining otolith microstructure, we back‐calculated body size at hatching and at emergence of YOY salmon sampled in fall 2000 and in early summer 2001 on the Petite Cascapédia River and the Bonaventure River (Québec, Canada). The results of the study did not reveal any size‐selective mortality of YOY salmon in the Petite Cascapédia River, while in the Bonaventure River, size‐selective mortality of the smaller individuals of the cohort was detected. This case study allowed not only a better comprehension of the population dynamics of those rivers but demonstrated the usefulness of otolith analysis to detect winter size‐selective mortality under a natural environment.     

The effects of fluvial processes and habitat heterogeneity on distribution, growth and densities of juvenile Atlantic salmon (Salmo salar L.), with consequences on abundance of the adult fish

Abstract – The required freshwater habitats of juvenile Atlantic salmon ( Salmo salar ) are, in general, well known, but vary in quality, related to interacting effects of several variables, which may depend on different parts of a river system. Examples are given of ranges of densities and growth that can be found at various sites in eastern Canada, illustrating the biological and physico-chemical factors affecting production of juvenile salmon. Relative growth rates can indicate habitat quality and population densities. Salmon parr have negative effects on brook trout in riffle habitats. The effects of migrations within the river and of changes with stream succession on juvenile salmon production are illustrated with examples from a Newfoundland river. Migration of age-classes can be quantified from 'self-thinning' curves. Lakes have enhancing effects on downstream fluvial habitats, and, at least in Newfoundland, and probably in many boreal areas, the lacustrine proportion of the basin can be used as an index for deriving estimates of required spawning escapement. The factors described should be taken into account for more refined estimates of river production and management of the salmon resource.     

Effects of water temperature and wind on year-class success of Tanner crabs in Bristol Bay, Alaska

We investigated five a priori hypotheses on factors affecting year-class success of commercially exploited Tanner crabs, Chionoecetes bairdi , in Bristol Bay, Alaska, through correlation analysis and multiple regression modelling. Estimates of recruitment from Zheng et al .' s (1998 ; Can. Spec. Publ. Fish. Aquat. Sci. 125:97–105) length-based analysis of assessment survey and commercial catch data were used to index year-class strength. This work extends results of an earlier study ( Rosenkranz et al ., 1998 ; Alaska Fish. Res. Bull. 5:18–24), which reported positive correlations between Tanner crab year-class size and north-east (NE) winds during the spring larval period, by considering the effects of nondirectional wind speed, bottom and surface water temperature, and abundance of the potential predators sockeye salmon ( Oncorhynchus nerka ) and Pacific cod ( Gadus macrocephalus ). No relationships were found between year-class size and mean wind speed or predator abundance, but positive correlations were found with bottom temperatures during gonadal development and egg incubation. Linear regression models with the independent variables NE wind and bottom temperature accounted for about half the variability in the year-class strength index ( r ²=0.50 for males, r ²=0.48 for females). Anomalously cold bottom temperatures may adversely affect the Tanner crab reproductive cycle, and NE winds may promote coastal upwelling while advecting larvae to regions of fine sediments favourable for survival upon settling. The role of Bering Sea oceanography on decadal-scale variability in Tanner crab population dynamics could not be resolved with the relatively short (∼ two decades) time series of stock assessments.     

Spatiotemporal change in growth of two populations of Asian chum salmon in relation to intraspecific interaction

Spatiotemporal changes in growth patterns of chum salmon Oncorhynchus keta that returned to the Ishikari (Japan) and Namdae (Korea) rivers in 1984–1998 were investigated using scale analysis. Juvenile chum salmon from both populations left coastal marine areas after spring at a size of over 8 cm fork length (FL). In summer, juvenile salmon from the Namdae River entered the Okhotsk Sea at a larger FL than did Ishikari River juveniles. There were no significant differences in annual growth between populations of 1-, 2-, and 4-year-old fish. For 3-year-old fish, however, Namdae River salmon had significantly higher synchronous and sympatric growth than did Ishikari River salmon. Mean FL of adults was also larger in Namdae River salmon than in Ishikari River salmon. Analysis of covariance (ANCOVA) results showed (1) negative linear relations between FL and catch, (2) homogeneous slopes of those relations at regional and species levels, and (3) nonhomogeneous slopes at the population level, indicating that density-dependent effects on growth were most significant at this level. We concluded that growth of chum salmon was concurrently influenced by stronger effects of intrapopulation competition and weaker effects of inter- and intraspecific interactions in the Bering Sea.     

Influence of European sculpin, Cottus gobio, on Atlantic salmon Salmo salar, recruitment and the effect of gravel size on egg predation – implications for spawning habitat restoration

Abstract  The study examined if recruitment of juvenile Atlantic salmon, Salmo salar L., was influenced by the presence of European sculpin, Cottus gobio L., and if the spawning substrate size used by salmon influences sculpin predation on salmon eggs. A combination of information on spawning site selection by female salmon and associated densities of juvenile salmon indicated that recruitment of juvenile salmon was 10 times lower in areas where sculpin was present than areas without sculpin. Predation rate on salmon eggs was found to be dependent on substrate size in artificial redds created in stream aquaria using four different sizes of substrate (13, 23, 37 and 62 mm). Predation rate averaged 83% in the aquaria with the largest substrate size, whereas a rate of only 2–3% was observed using smaller substrate sizes. Sculpin may thus be an important factor influencing the recruitment of juvenile salmon. Selecting small enough gravel sizes during restoration of salmon spawning habitat could therefore be important to minimise egg predation.     

Effects of turbidity on feeding of the young-of-the-year pikeperch (Sander lucioperca) in fishponds

The effect of water turbidity on the prey selection and consumption of the young-of-the-year (YOY) pikeperch in the planktivorous feeding stage was studied. Attention was paid particularly to the question of how the food selectivity depends on the size of YOY pikeperch and how the turbidity affects feeding in different size classes. Studies were carried out in ponds of two fish farms in Estonia over 4 years. Small cladocerans were the most preferred prey in the smallest pikeperch size class. In larger size classes, the most selected prey were the large cladocerans. Water turbidity affected the prey selection of the planktivorous pikeperch significantly. In more turbid environments, the larger zooplankters were more positively selected than the smaller ones. Turbidity decreased both total zooplankton consumption and Fulton's condition factor of fish only in the largest size class of pikeperch. The effect of turbidity on foraging and growth, and thus on the size of juvenile pikeperch of a particular year class is substantial under conditions where juveniles cannot shift from planktivory to piscivory.     

Spatial behaviour of young-of-the-year northern pike (Esox lucius L.) in a temporarily flooded nursery area

Abstract –  Unlike adult northern pike ( Esox lucius ), small-bodied young-of-the-year (YOY) individuals cannot be tracked by conventional telemetry, and virtually no data exist on their spatial behaviour. Here, we monitored 192 individuals released into a nursery area (0.47 ha, subdivided into 253 cells of 4 m × 5 m) that dried out progressively using a portable passive integrated transponder detector. Among them, 66 spent more than 5 days in the nursery area and were localised at least twice, allowing spatial behaviour analyses. The mean radial distance moved averaged 14.3 m (±8.4 SD, range 2.2–41.0) and the average daily movement was 8.0 m·day⁻¹ (±5.3 SD, range 0.6–31.1). Pike abundance in a cell significantly correlated with the abundance observed in adjacent cells within 32 m, confirming a patchy distribution. A selective use of the deepest area was measured while no effect of vegetation cover was observed at the cell scale. The existence of YOY pike aggregations and the positive correlation between water depth and their spatial distribution might, however, facilitate predation and/or cannibalism during this critical period. This also suggests that the choice of release habitats requires particular attention during restocking procedures.