The relationship between temperature and the size and age of larvae and peri‐metamorphic stages of Pleuronectes flesus

Habitat use of 0+ year pike Esox lucius (mean 20–31 mm L _T) was evaluated in artificial ponds (surface area = 16 m²) in relation to water transparencies (low and high), habitat complexity (low and high), food availability (zooplankton) and cannibalism (mortality in the ponds). Pike could choose freely between a structured area, an open water area and a structure and open water interface area. Habitat use shifted towards the structured habitat type in the presence of high habitat complexity and low water transparency. Both findings could be explained as a consequence of territorial behaviour. When cannibalism was intense, the proportions of pike using the structures were reduced and the largest pike, and hence the potential cannibals, were found inside the structures. Since the smallest pike were constrained to the unsheltered, and hence probably more hazardous, habitat outside the structures, this study suggests that mortality due to intra‐cohort cannibalism may occur in an indirect as well as a direct way.     

Otolith-based analysis of survival and size‐selective mortality of stocked 0+ year pike related to time of stocking

The effect of time of stocking on the extent and source of mortality in 0+ year pike Esox lucius was investigated in a lake (20 ha) and a drainable pond (0·5 ha) using pike with alizarin marked otoliths. The results indicated that pike stocked late relative to the recruitment of native 0+ year pike fell victim to cannibalism from these larger individuals. This resulted in very low survival through the first growing season (<2%). Pike stocked early in the season exhibited significantly higher survival (>12%). Analyses of the size distribution of the alizarin marks from these fish revealed that the largest 0+ year pike exhibited by a factor of 3·3, higher survival than the average 0+ year pike in the lake. In order for large 0+ year pike to exhibit such high relative fitness a minimum of 69·7% of the original population must suffer size dependent mortality. In the pond the survival of the largest pike was by a factor of 4·2 higher than the average pike, and the corresponding size dependent mortality was 76·4%. The substantial size‐dependent mortality was most probably due to intra‐cohort cannibalism or habitat segregation between large and small 0+ year pike that exposes the small pike to predatory fishes in the open lake. Cannibalism exerts a major influence on the survival of 0+ year pike post‐stocking although the magnitude and origin differ in relation to stocking time.     

Stocking success of newly hatched pike evaluated by radioactive strontium (85Sr) marking

The stocking of newly hatched pike Esox lucius labelled with radioactive strontium (⁸⁵Sr) and later electrofishing of the same area was found to be a workable tool in assessing the success of pike stockings. ⁸⁵Sr‐labelled individuals constituted 96% of the 0+year pike catch from late summer electrofishing in the stocked littoral areas of the heavily regulated Lokka Reservoir. The density of 0+year pike reached 2·1 individuals 100 m⁻² in the stocked areas in comparison to 0·3 individuals 100 m⁻² in the reference areas. In the three other stocked lakes the proportion of labelled pike in the catch ranged from 0 to 42% in late summer. The results suggest that pike stockings are profitable in filling empty or nearly empty 0+year pike habitats of regulated lakes or other waterbodies, where the natural reproduction of pike has declined due to human activities.     

Changes in the fish community and water quality during seven years of stocking piscivorous fish in a shallow lake

SUMMARY 1. Piscivores (annual stocking of 1000 individuals ha^?1 of 0+ pike and a single stocking of 30 kg ha^?1 of large 20–30 cm perch) were stocked in seven consecutive years in a shallow eutrophic lake in Denmark. The stocking programme aimed at changing food‐web structure by reducing zooplanktivorous and benthivorous fish, with resultant effects on lower trophic levels and ultimately water quality. 2. The fish community and water quality parameters (Secchi depth, concentrations of total phosphorus, chlorophyll a and suspended solids) were monitored between 1996 and 2000 and relationships were evaluated between predatory fish and potential prey and between zooplanktivorous or benthivorous fish and water quality parameters. In addition, potential consumption of piscivorous fishes was calculated. 3. The density of fish feeding on larger zooplankton or benthos (roach >15 cm, crucian carp >15 cm) declined distinctly during the study period. This effect was attributed to predation by large (>50 cm) pike. Based on scale readings, we cautiously suggest that the stocking of 0+ pike boosted the adult pike population to produce an unexpected impact in later years. Conversely, no direct impact of stocked 0+ pike was detected on 0+ roach. 4. A major decline in the recruitment strength of 0+ roach was observed in 2000. A combination of (i) the indirect effect of large pike preying on adult roach, with negative effect on roach reproduction and (ii) the direct predation effect of 0+ pike and/or 1+ and 2+ perch recruited in the lake, provides the most likely explanation of this phenomenon. 5. A marked increase in Secchi depth in 2000 and declining trends in suspended solids, chlorophyll‐a and total phosphorus concentrations were observed. These changes may also be attributable to changes in the fish community, although the relationships were not straightforward. 6. This 7‐year study indicates that piscivorous fish may be a significant structuring force in shallow eutrophic lakes, suggesting that stocking piscivores can increase predation pressure on cyprinids. However, the general lack of impact of 0+ pike points to the need of refining current stocking practices in several countries across Europe.     

Mortality and production of 0+ perch, Perca fluviatilis L., in two Scottish lakes

Mortality of 0+ Eurasian perch was measured directly from the decline in catch in a high-speed plankton sampler and described by a negative exponential model. A mean mortality of 0.063 day ¹ (range=0.020–0.089) was calculated; for 0+ juveniles it was 0.021 day ¹. Mean larval production was 0.85–1.30 mg dry weight m⁻² day⁻¹ in Loch Kinord and 0.08–0.12 in Loch Davan. Annual production of 0+ juveniles was 1.97–7.55 kg wet weight ha ⁻¹ at L. Kinord and 1.01 kg at L. Davan. The contribution of 0+ perch, including the egg stages, to total perch production was 88–96%. Availability of food items on transition to exogenous feeding and through the larval period did not limit survival. Cannibalism by adults accounted for most of larval mortality, and predation by pike and adult perch were responsible for the majority of juvenile losses. The population structure of adult perch, for 12 year classes, was stable at L. Kinord, while year classes at L. Davan exhibited fluctuations in abundance and in two sampling years were eliminated in the first summer. Survival in the eleutheroembryonic phase was variable at L. Davan and observed to be low compared with L. Kinord. It is suggested that this was a result of water turbulence on the exposed and shallow spawning areas at L. Davan.     

The population biology of pike, Esox lucius L., in two gravel pit lakes, with special reference to early life history

The population biology of pike in two gravel pit lakes of contrasting habitat type were studied. The density of pike (1 + and over) in November 1987 in the Main Lake was 7.7 fish ha⁻¹ (19 kg ha⁻¹) and in St Peter's was 71 fish ha'(53 kg ha⁻¹). The age distribution showed that recruitment was consistent in St Peter's but variable in the Main Lake. The main factor causing this difference in recruitment is abundance of aquatic vegetation. Survival up to November 1987 of 0-group pike in the Main Lake was 0.00175% from potential egg deposition, producing 6.6 fish ha⁻¹ (0.83 kg ha⁻¹). In St Peter's survival was 0.00315%, producing 23 fish ha⁻¹ (3.04 kg ha ¹).     

Density related growth of grass carp, Ctenopharyngodon idella (Val.) in managed small impoundments in Alabama

Growth data for grass carp, Ctenopharyngodon idella , in managed small impoundments were examined for density related effects. Ponds ranged in area from 0.02 to 5.0 ha; various species stocking combinations were used. Growth during the first year of life was strongly affected by density. Total standing stock increased and average size decreased with increasing density. This growth–density relationship was more variable among fish in the second year of life, primarily because of differences in availability of preferred food in different ponds. Survival tended to vary during the first year of life depending on the size at stocking but was uniformly high in the second year.     

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.     

Temporal and spatial variation in growth of juvenile Atlantic salmon

Spatial and temporal variation in length‐at‐age and environmental factors affecting variation in growth rate of juvenile Atlantic salmon Salmo salar were studied using data from a long‐term study in the River Stjørdalselva, central Norway. Mean annual instantaneous growth rate among 1+ and 2+year juvenile Atlantic salmon varied between 0·59 and 1·50 g g⁻¹ year⁻¹ and mean instantaneous daily growth rate of young‐of‐the‐year (YOY) varied between 0·013 and 0·033 g g⁻¹ day⁻¹. Between year variation in growth was larger than the within year intra‐watercourse spatial variation. For YOY and 1+year Atlantic salmon, a major part of the observed between year variation in growth rates was explained by variation in mean daily water discharge and spring temperature. For 2+year juvenile Atlantic salmon, mean daily water discharge and cohort density were the only variables to significantly explain variation in growth rates. A large part of the within water‐course spatial variation could not be explained by temperature variations and juvenile Atlantic salmon in the uppermost areas of the river, experiencing the lowest ambient temperatures during the growth period, displayed the highest growth rates. Within the baselines set by temperature, biotic and abiotic factors connected to water flow regime and variation in food availability are suggested to be a major determinants of the temporal and spatial variation in juvenile Atlantic salmon growth rates.     

Piscivory of 0+ pike (Esox lucius L.) in a small eutrophic lake and its implication for biomanipulation

Stocking 0+ pike in shallow eutrophic lakes as an indirect way to facilitate clear water by increased phytoplankton grazing by zooplankton, has been used in several studies with varying success. Lack of piscivory could be one reason for poor impact of the pike stockings, on reducing the 0+ cyprinid fish population. In relation to this, we present data on food and feeding habits of 936 0+ pike (Esox lucius L.) (2–18 cm) sampled during five years in a small eutrophic lake undergoing biomanipulation. Overall, the food of 0+ pike followed a sequence of diet shifts including microcrustacea, insects, macrocrustacea and finally vertebrates (fish) with increase in pike length. Despite a high abundance of potential prey fish in the lake, in general less than 50% of the pike were piscivorous until a length of 10.1–13.7 cm. As piscivory increased, the principal prey fish (0+ roach) apparently displayed antipredatorial behaviour which decreased their availability for 0+ pike. We conclude that lack of piscivory caused by e.g. anti predator tactics by prey fish, may reduce the impact of 0+ pike stockings in lake restoration projects.

     

A pike management strategy for a trout fishery

The efficiency of pike removal from a chalkstream trout fishery, by an angling society using electrofishing, was estimated over five successive autumns using mark-recapture methods. It averaged c . 50% for pike aged 1+ years or more. Numbers of 3+ and older pike decreased from 3.7 ha^-1 (1980) to 1.4 ha^-1 (1984), but there was no corresponding reduction in recruitment success. Both pike immigration and an inverse relationship between natural and fishing mortality could have reduced the effect of the pike culls. The mean weight of 0+ pike increased from 19.4 g to 48.2 g over the 5 years, but there was no change in the growth rates of older pike. Only the larger pike were able to eat the larger trout stocked, and the reduction in numbers of 3 + and older pike appears to have reduced the predation pressure on the trout population. Angling effort increased more than three-fold over the 5 years studied and rod-catches more than doubled, but catch-per-unit-effort was extremely variable between years. The increased rod catch occurred despite the decrease in numbers of large (> 33 cm) trout stocked in the spring and early summer. Compared with pre-cull years, the angling society saved about ş850 (present day prices) by stocking fewer trout.     

Evaluating stocking of YOY pike Esox lucius as a tool in the restoration of shallow lakes

1. Stocking of piscivores in shallow, eutrophicated lakes to reduce cyprinid densities is a common approach in lake restorations. Young‐of‐the‐year (YOY) pike Esox lucius are frequently used to reduce cyprinid densities, but their effectiveness is equivocal. This study uses a simple model to assess the efficacy of pike stocking as a lake restoration tool. 2. The model evaluates YOY pike consumption in relation to different survival and growth patterns, timing of stocking, degree of piscivory of pike as well as YOY cyprinid growth patterns and productivity. The parameters used in the model are obtained from field surveys in eight study lakes stocked with YOY pike as well as from the literature. 3. Our model showed that all parameters studied were important for predicting the effects of pike stocking on cyprinids. In particular, body size at stocking, cyprinid production and pike survival were good predictors of biomanipulation success. 4. However, our model showed that only under very specific circumstances will stocking of YOY pike result in a reduction of YOY cyprinid densities large enough to affect water quality. 5. The findings of this study question the efficacy of pike stocking as an appropriate and reliable tool for restoration programmes in shallow lakes, and indicate that when used efforts should be made to optimize the timing of stocking in relation to YOY cyprinid production and to increase the stocking body size of the pike.     

Chernobyl-derived radiocaesium in fish as dependent on water quality and lake morphometry

¹³⁷Cs concentrations in perch Perca fluviatilis , pike Esox lucius and roach Rutilus rutilus obtained from lakes of different size and water quality in an area which received about 10–67 kBq m⁻²¹³⁷Cs, were compared with environmental data. Radiocaesium concentrations were highest in pike, and were about two to three times higher in the pike and perch than in the roach. The largest perch had about four times more ¹³⁷Cs than the smallest ones, but the activities in the pike and roach were independent of fish size. All of the water quality parameters examined correlated with ¹³⁷Cs concentrations in the fish, but the concentrations of ¹³⁷Cs in the bottom sediment did not. ^l37Cs fallout and water retention time in lakes showed a positive correlation with fish ¹³⁷Cs, while the size of the catchment area, phosphorus content of the water, water colour, pH and electrical conductivity showed a negative one. Thus there was more ¹³⁷Cs in the fish from the oligotrophic lakes than from the eutrophic ones, from the less humic lakes than from the more humic ones, from the more acid than from the less acid ones and from the lakes containing less potassium than from those containing more potassium. The depth, area and volume of the lake did not affect radiocaesium in the fish. Regression equations are presented for predicting ¹³⁷Cs in fish on the basis of environmental data. Fallout, total phosphorus and pH together had the highest influence in multiple regression models but were less significant in the roach than in the perch and pike.     

Mackerel icefish size and age differences and long‐term change at South Georgia and Shag Rocks

This study analysed the total length ( L _T)‐frequency distribution of mackerel icefish Champsocephalus gunnari at South Georgia and Shag Rocks from nine bottom trawl surveys at South Georgia and eight at Shag Rocks between 1987 and 2002. The estimated mean L _T of age‐classes 1+, 2+, 3+ and 4+ years during January were, respectively, 14·7, 23·5, 29·8 and 35·1 cm at South Georgia. Age‐classes 1+, 2+ and 3+ years were 18·3, 26·2 and 33·8 cm at Shag Rocks. The derived Bertalanffy growth parameters for South Georgia were: L _∞ = 51·7 cm, k  = 0·27 and t ₀ = −0·26. The mean L _T of each age‐class of C. gunnari at Shag Rocks was significantly larger than at South Georgia, equivalent to c . 5 months growth, although the annual growth in L _T was similar. This is further evidence that C. gunnari hatched earlier at Shag Rocks. At South Georgia, the mean L _T of age‐classes 1+ and 3+ years were correlated, and significantly decreased between 1987 and 2002, and were smaller following warmer summers. This decrease in the size of C. gunnari may be the result of reduced food availability linked to climate warming.     

Large-scale synchrony and inter-annual variability in roach recruitment in the Rhône River: the relative role of climatic factors and density-dependent processes

1. We report patterns of temporal variation in the recruitment of roach ( Rutilus rutilus ). The data consist of the annual abundance of the first 2 year-classes, 0+ and 1+ fish, at four sites in the Rhône River (France) between 1987 and 1997. Over this 11-year period both 0+ and 1+ fish fluctuated strongly.
2. Cross-correlation indicated high spatial synchrony in 0+ dynamics, although correlations among sites in 1+ dynamics were weaker. No clear pattern was apparent in the relationship between the level of synchrony and distance between pairs of sites.
3. The spatial synchrony in 0+ fish could be attributable to large-scale variations in weather, influencing water temperature. Total body length of 0+ roach was correlated with water temperature (expressed in degree-days over 12 °C), and water temperature was the main factor explaining inter-annual variation in 0+ cohort size. Monthly variation in abiotic factors (measured by standard deviation in water temperature and discharge) did not influence 0+ fluctuations. Correlations with June water temperature suggest that year-class strength was mainly determined by abiotic factors during the first few months of life.
4. The absence of spatial synchrony in 1+ fluctuations suggests little correlation between survival and abiotic conditions during the first year of life, other factors influencing survival.
5. Survival in the first year was density-dependent. Intraspecific competition within the 0+ cohort could thus influence the fluctuations in recruitment to older age-classes.
6. The implications of age- or stage-dependent synchrony in temporal variation for species with complex life histories are discussed. Studying spatial synchrony for the different life history stages could enhance our understanding of the population dynamics of spatially structured species.     

Temporal dynamics in epipelic, pelagic and epiphytic algal production in a clear and a turbid shallow lake

SUMMARY 1. Pelagic and epipelic microalgal production were measured over a year in a pre-defined area (depth 0.5 m) in each of two lakes, one turbid and one with clear water. Further estimates of epiphytic production within reed stands were obtained by measuring production of periphyton developed on artificial substrata.
2. Total annual production of phytoplankton and epipelon was 34% greater in the turbid lake (190 g C m⁻² year⁻¹) than in the clearwater lake (141 g C m⁻² year⁻¹). However, the ratio of total production to mean water column TP concentration was two fold greater in the clearwater lake.
3. Phytoplankton accounted for the majority of the annual production (96%) in the turbid lake, while epipelic microalgal production dominated (77%) in the clear lake. The relative contribution of epipelic algae varied over the year, however, and in the turbid lake was higher in winter (11–25%), when the water was relatively clear, than during summer (0.7–1.7%), when the water was more turbid. In the clearwater lake, the relative contribution of epipelon was high both in winter, when the water was most clear, and in mid-summer, when phytoplankton production was constrained either by nutrients or grazing.
4. Compared with pelagic and epipelic primary production, epiphytic production within a reed stand was low and did not vary significantly between the lakes.
5. The study supports the theory of a competitive and compensatory trade-off between primary producers in lakes with contrasting nutrient concentrations, resulting in relatively small differences in overall production between clear and turbid lakes when integrating over the season and over different habitats.     

The reproductive success of pike, Esox lucius: aspects of fecundity, egg density and survival

Power-law relationships have been estimated between fecundity and fish length, weight and age for pike from two gravel pit lakes (Main Lake and St Peter's Lake) in Buckinghamshire, England. Relative fecundity, estimated at 17.6 eggs g⁻¹ for Main Lake pike and 19.4 g⁻¹ for St Peter's Lake pike, did not differ significantly between the lakes. Population fecundity for the Main Lake was estimated at 10.6 million eggs and 6.4 million eggs in 1986 and 1987, respectively.
Experimental work involving samples of eggs from 18 artificially fertilized Main Lake female pike showed a significant relationship between egg diameter and female length, and a significant difference between mean egg diameters for 41-cm and 101-cm females. No significant relationships were found between mean egg size and mean fry size, nor between fry length and adult length. Mean size differences between newly hatched, 18-day-old and 41-day-old fry from the 41-cm and 101-cm females were not significant.
Samples of eggs taken at 2-day intervals from 6 April to 5 May in a pike spawning area of the Main Lake revealed clumped distributions of eggs, with average egg densities ranging from 1 to 47 eggs per 0.071 m² (14–671 m ²) on sand and silt substrata. The maximum egg density of 51 per 0.071 m² (729 m⁻²) was found on flooded grass. The stages of development of the eggs were identified over the sampling period. Survival from stage 1 to fry was estimated at 3–6%. Egg losses were estimated at 9–10% day⁻¹.
Egg survival experiments gave overall hatching success rates of 11 % on sand/silt compared with 2.5% on aquatic plants (1986), and 18.9% on undisturbed sand/silt compared with 7.9% on disturbed sand/silt (1987). The effect of siltation on egg hatching is discussed.     

Early life characteristics of pike, Esox lucius, in rearing ponds: temporal survival pattern and ontogenetic diet shifts

Survival, biomass and diet of pike, Esox lucius , larvae and juveniles were studied over 3 years by stocking free embryos at a density of six fish m⁻² in 12 drainable outdoor ponds. The ponds were sequentially drained at six larval/juvenile developmental stages, up to a total length (T.L.) of 139 mm. The mean rate of survival at harvest decreased irregularly over time and the highest mortality rates were recorded during the early larval period (13 to 27 mm t.l .) and two intervals of the juvenile period (46 to 99 mm T.L. and 121 to 139 mm t.l .). Mean biomass increased dramatically between 46 mm T.L. (19.8 kg ha⁻¹) and 121 mm T.L. (181.8 kg ha⁻¹) and stabilized between 121 and 139 mm t.l . Sharp increases in the mean weight coefficient of variation were recorded during the early larval period (13 to 27 mm t.l .) and between 74 and 121 mm t.l . (development of cannibalism). Diet breadths were relatively narrow in pike larvae and reached maximum levels in 99 mm t.l . juveniles. Average-sized pike exhibited a sequence of size-dependent shifts from a diet composed primarily (in terms of weight) of micro-crustaceans (at 13 mm t.l .), to chironomid larvae (at 74 to 99 mm t.l .), and finally macrocrustaceans (at 121 to 139 mm t.l .). Cannibalism was detected first among the largest fish at the 74 mm t.l . stage. Between-year diet similarity at various developmental stages was consistently high. In 70 mm t.l . fish harvested from different ponds, we found significant among-pond differences in diet composition; however, similar trends of diet changes in relation to fish size were observed from pond to pond. Our results are discussed in light of existing knowledge of young pike trophic ecology and current aquaculture practices.     

Swimming performance and metabolism of 0+ year Thymallus arcticus

The prolonged swimming speed and metabolic rate of 0+ year Arctic grayling Thymallus articus were examined with respect to current velocity, water temperature and fish size, and compared to conditions fish occupy in the river. Oxygen consumption (mg O₂ h⁻¹) increased with fish mass and temperature (6–23° C), with a steep increase in metabolic rate between 12 and 16° C. Absolute prolonged swimming speed (cm s⁻¹) increased rapidly with fish size (total length, L _T, and mass), however, fish in the natural stream habitat occupied current velocities between 15 and 25 cm s⁻¹ or 4  L _T s⁻¹, approximately half their potential prolonged swimming speed (10  L _T s⁻¹).     

The consequences of a drastic fish stock reduction in the large and shallow Lake Wolderwijd,The Netherlands. Can we understand what happened?

In 1990 an experiment started in the large and shallow lake Wolderwijd (2700 ha, mean depth 1.5 m) to improve the water quality. About 75% of the fish stock was removed (425 000 kg fish). The fish was mainly composed of bream and roach. In May 600000 young pikes (3–4 cm) were introduced.In May 1991 the water became very clear (Secchi depth 1.8 m) during a spring bloom of large Daphnia. Then the grazing by zooplankton was eight times higher than the primary production of algae and the total suspended matter concentration became very low. Compared to the situation before the fish reduction, the grazing had increased only slightly, while the primary production had decreased significantly in early spring. The fish stock reduction might have contributed to the reduction in primary production by a reduced internal nutrient load. The clear water period lasted six weeks. Daphnia disappeared in July due to food limitation, the algal biomass increased and the Secchi depth became 50 cm. Daphnia did not recover during summer, due to predation that was not caused by 0 + fish but by the mysid shrimp Neomysis integer. Neomysis could develop abundantly, because of the reduced biomass of the predator perch. The production of young fish had been low because of the cold spring weather. The cold weather was probably also responsible for the slow increase in density of macrophytes. After 1991, perch probably can control Neomysis. Due to lack of spawning places and shelter for 0 + pike, pike was probably not able to control the production of 0 + fish. In a lake of this scale, it will not be easy to get more than 50% coverage of macrophytes, which seems necessary to keep the algal biomass low by nutrient competition. Therefore, we expect also in the future a decrease in transparency in the summer. Locally, especially near Characeae, the water might stay clear.