Population structure,growth and fecundity of perch (Perca fluviatilis L.) in an acidified river system in Southern Norway

Population structure, density and reproductive potential of perch (Perca fluviatilis L.) from 4 acidified lakes in a river system in Southern Norway were investigated. The upper 3 lakes were most affected by acidification and the number of perch caught per unit effort indicate three-fold increase in density from the upper Gjerstadvann to the lower Brøbørvann. Low density of perch in the upper lakes is explained by; 1) Abnormal mortality occurring in episodes due to acidification, probably caused by aluminium toxicity at pH = 5.2–5.4. 2). Acid water (pH ≤ 5.0) during spawning and development of eggs and larvae, resulting in recruitment failure. Juvenile mortality of perch may depend on the density of the parental stock. In the 3 upper lakes, the density of adult perch probably was too low to produce significant juvenile mortality, and in these lakes the recruitment probably depended more on the May water quality. Improved water quality increases egg hatching and survival and also benefits zooplankton production, the main food for perch during the first summer. Low population density has reduced competition for food and thus improved growth of perch in the 3 upper lakes compared to the lower lake. The perch in Brøbørvann mature at higher age and have lower individual fecundity than perch in Gjerstadvann, an effect of both slower growth and lower length specific fecundity in Brøbørvann. In Gjerstadvann, the perch therefore have a higher reproductive potential relative to the stock density, and may, when the water quality is good enough, give rise to strong year-classes and more frequent year-class fluctuations than perch in the less acid Brøbørvann.     

The population dynamics of pike,Esox lucius,and perch,Perca fluviatilis,in a simple predator-prey system

Synopsis The population dynamics and predator-prey relationship of pike, Esox lucius, and perch, Perca fluviatilis, were examined in simple fish communities in two adjacent shallow lakes, Lochs Kinord and Davan, Deeside, Scotland. Few perch survive to age 3 but Z is low for fish > 3 years and perch live up to 17 years. Population fecundity remained relatively high and constant in perch because of the multi-age spawning stock and the presence of older more fecund perch. Growth rates of perch in both lochs are relatively high as a consequence of low stock abundance. The N, B, and P of adult perch were unusually low. The age range of pike, and N, B, P, and growth were in the range of values reported elsewhere. There was little variation in the strength of pike year classes and the importance of cannibalism and low occurrence of alternative prey in the lochs suggest that the populations were self-regulating. Cannibalism by adults was responsible for most of mortality in perch larvae, and predation by pike and adult perch was responsible for the majority of juvenile losses. This conclusion is supported by the high biomass ratios of pike:juvenile perch of 1.0–30.8. While the number of adult fish was almost equal, the biomass of adult pike was 2–3 × that of perch in Kinord and 6 × in Davan. In L. Kinord, where year class strength was stable, high predation pressure from perch and pike reduced perch abundance rather than eliminated year classes. Perch year classes fluctuated in abundance in L. Davan and were eliminated in the first summer in two sampling years. The pike, and particularly the perch populations, have features characteristic of fish communities in unperturbed ecosystems: namely, a wide range of age classes, stability in biomass with variation dampened by longevity, and low production.     

Effects of temperature on life history variables in perch

Population structure, recruitment, mortality, life expectancy, age at maturity, gonad development, fecundity, net reproductive rate and energy storage were studied in a perch population exposed to thermal discharge. The experiment was conducted during 1978–1990 in an artificial enclosure at a nuclear power plant on the Baltic Sea coast. Abundance increased after the plant started operating, although the proportion of larger perch dropped to a low level. Recruitment improved, but adult mortality increased. This effect was accentuated as the fishes matured very early and at a very small size, reducing their ability to support a growing gonad in high winter temperatures. During the spawning period, condition dropped to very low levels accompanied by increased mortality. Surviving fish delayed the next spawning by one or more years and reduced their fecundity. Body condition thus improved in large perch at the end of the study period. Life-time fecundity was reduced, and reproductive performance shifted to youner ages.     

The annual reproductive cycle of pike, Esox lucius L., in two Scottish lakes

Histological changes in the ovaries of pike from two shallow, adjacent, Scottish lakes are described and related to a macroscopic scale of maturity stages and to changes in gonad weight. There was no evidence of pre-ovulatory degeneration although occasional residual oocytes were present in spent ovaries. The gonadosomatic index (GSI) was high in males from October to March, and in females it increased through the winter months to a maximum immediately prior to spawning. Maximum ovarian weight was 8.3–9.0% of total body weight compared with only 0.9–1.1% for testes. These GSIs were low compared with other reports for the British Isles, suggesting lower reproductive output. Spawning occurred in the first two weeks of April at a water temperature of 6.0–7.5° C. Fecundity was described from a regression of egg number (F) on fork length (f.l.): log₁₀F= -1.51 + 3.33 log₁₀ f.l. (cm). Mean egg diameter after preservation varied from 1.1 to 2.4 mm over the size range examined. Males were mature at age 2 at a length of ≥ 27 cm, while females first spawned at age 3 at a length of ≤ 28 cm, although a minority were mature at age 2. The sex ratio was 1.2 males: 1 female at Kinord and 1.4 : 1 at Davan. Greater egg production at Davan, 50 678 ha^?1, compared with Kinord, 33 557 ha^?1, was due to higher individual fecundity-at-age resulting from better growth. Fecundity (19.29 × 10³ calculated for a pike of 55 cm F.L.), relative fecundity of only 10–24 eggs per gram wet weight of adult, and GSI values were low compared with pike in North America and Eurasia. Food supply may restrict reproductive output of pike in Lochs Kinord and Davan. The abundance ratio of predator: prey is high with the main prey species, perch, present in low numbers, and the only alternative species is eel in these simple fish communities.     

Population regulation in lake whitefish, Coregonus clupeaformis (Mitchill)

Mechanisms that might compensate for changes in mortality are well known but documentation of their operation and determination of their relative capacities is difficult. In this study the potential of lake whitefish to compensate for fishing mortality is quantified using a conventional fishery stock assessment model. The mechanisms examined are proportion of females, fecundity, survival of eggs and larvae, decrease in age at maturity and growth. The change in each of these parameters necessary to maintain constant recruitment with increasing fishing pressure is determined. Decrease in age of maturity and increased survival of larvae and eggs have the greatest potential for compensation. The total capacity for compensation appears large even when the mechanisms are considered alone. A review of several lake whitefish population studies indicates a large variation in the vital statistics. There is a close relation between total mortality and growth. There is also a close relation between growth and age at maturity. The analyses indicate that increased fishing results in increased growth which then results in a lower age at maturity. This feedback mechanism enables lake whitefish to respond to changes in environmental conditions and on average to maintain a birth rate in balance with mortality.     

Age, growth and reproduction of Cobitis paludica in a seasonal stream

In the Iberian endemic loach Cobitis paludica , in a small seasonal tributary of the Guadalquivir River, 65–70% of the total growth in length occurred in the first year of life. The maximum ages observed were 3+ years in males and 4+ years in females. Both sexes matured at the beginning of their second year of life. There was a significant difference from 1: 1 in the overall sex ratio of 412 males to 674 females. Spawning began in late March and ended in July. Cobitis paludica is a multiple spawner that releases a minimum of two batches of eggs per female each year. Once spawning had started, however, there was no recruitment from the stock of transparent oocytes to the mature stock. The number of eggs decreased from the first batch to the second, but there was no significant difference in the mean egg diameter between batches. Compared with other populations of this species, the present population, from a low latitude, is characterized by a low number of age groups, fast growth, early maturity and high fecundity in multiple spawnings. These life-history characteristics are typical of species in unstable environments, where adult mortality is high, variable or unpredictable.     

Experimental manipulations of the fish populations in Windermere

A large scale experiment has been carried out on the perch and pike populations of Windermere in the English Lake District for nearly 40 years, and the objectives have changed over this period. Initially the aim was to manage the trout population, but the work has produced data of interest in predator-prey relationships and pike management. Perch were first heavily fished and then monitored for size, and numbers by trapping. Large pike have been culled by gill netting. The effects of pike gill netting have been to increase the growth rate and decrease the average age in the population. These changes in the pike population structure, together with climatic variations, have modified the perch population. The perch did not recover from the heavy fishing for many years and the population numbers remained low due to pike predation supressing recruitment. It was not until a large cohort, mainly controlled by the weather, was produced that the population increased. Cannibalism by these perch later combined with the pike predation to suppress future cohorts. There have also been as yet unexplained changes in the perch mortality rates. The experiments have led to a greater understanding of the relationship between pike and perch populations, but the use of this knowledge for management depends on what the objectives are.     

Cannibalism in pike fry, Esox lucius L.: some experiments with fry densities

Pike fry were kept in 0.054-m³ tanks at densities of 50, 100 and 150 (277, 555, 833 fry m⁻²) for seven weeks during which the development of social and feeding behaviour was observed. Zooplankton, macro-invertebrates and perch fry were provided sequentially as food; the pike fry were allowed to feed ad libitum.
Zooplanktivorous fry stopped growing at 22 mm whilst, in the presence of abundant suitable alternative prey, 1–4% turned cannibalistic at 5 weeks of age; cannibals subsequently grew rapidly (mean 1.88 mm day⁻¹). Cannibalism ensued in all tanks when the ratio of predator size: prey size was c .2:1.
Fry tended to space-out evenly in the tanks with no overt aggression or territoriality. Behaviour was typified by remaining still for long periods, particularly subsequent to the onset of cannibalism. Cannibals were attracted by fry movements which often initiated attacks.
Daily per capita mortality rates showed no crowding effects before cannibalism but significant density-dependent mortality due to cannibals. Consumption rates of cannibals varied between 0.63 and 6.0 fry per cannibal per day. Cannibals accounted for 54–96% of daily mortality in the experimental tanks. These results are discussed in relation to proposed mechanisms of pike population density regulation.     

Evaluation of pulse fishing for the walleye, Stizostedion vitreum vitreum, in Henderson Lake, Ontario

A total of 3226 walleye were removed from Henderson Lake, Ontario over 3 years (1980–2), causing the stock to collapse. This removal tested the applicability of 'pulse' fishing as a management alternative, and also provided an opportunity to determine which population characteristics might be monitored to serve as predictors of stock collapse in this species.
Following initial exploitation, increased length at age occurred only within younger age-lasses. To use this response as a predictor, one must first obtain pre-exploitation length-at-age data through using the proper gear to sample small fish. Abrosov's mean age to mean age at maturity index may also forewarn of stress (1.2 critical t value for Henderson Lake), but poor recruitment usually invalidates this index, since the lower angling vulnerability of large fish biases mean age calculations. While annual production estimates were a good indicator of the collapse, their determination required much effort. Petersen population estimates produced much more realistic estimates of population size than Schumacher–Eschmeyer estimates, but both estimates not only require much effort but also are influenced by changes in recruitment.
Poor predictors of the walleye collapse included: catch-per-unit-effort data, which, while giving some idea of fish density, were not good indices of exploitation stress; condition factors, which were not correlated to fish abundance; fecundity increases, which suffered far too much of a retarded response in Henderson Lake to serve as a predictor of stock collapse. The inability to determine accurately sex ratios of measure recruitment made these parameters unless.
At least over the short term, neither northern pike nor white sucker populations have increased following the walleye collapse. As yet, walleye have not returned to their former abundance, so the operational usefulness of 'pulse' fishing remains an unknown.     

State-dependent invasion windows for prey in size-structured predator-prey systems: whole lake experiments

1. In size-structured communities where individuals grow in size over their life cycle, interactions between species will shift between competitive and predatory interactions depending on size relationships. The outcome of interactions will subsequently depend on the strength of competitive and predatory interactions, respectively. 2. In a whole lake experiment including four experimental lakes, it was tested under which conditions the competing prey, roach Rutilus rutilus, could successfully recruit into systems previously occupied by the predator, perch Perca fluviatilis. Two replicated introduction experiments were carried out 3 years apart. 3. Roach were able to successfully recruit into three of the four experimental lakes of which two were also inhabited by the top predator pike Esox lucius. Resource levels were unrelated to whether roach could successfully recruit into the systems as recruiting roach in all years were feeding close to their maximum rate. 4. High population fecundity of roach and low predation pressure by perch combined were necessary ingredients for successful recruitment and the presence of only one of these conditions did not result in successful recruitment. 5. It is hypothesized that, although roach were able to successfully recruit into one lake with only perch present in addition to the two lakes that also inhabited pike, long-term coexistence of roach and perch depends on the presence of another top predator (e.g. pike) selectively preying on perch. This hypothesis was supported by data on co-occurrence of perch and roach in different lakes. 6. Overall, the results are in accordance with expectation of size-structured life-history omnivory theory suggesting that coexistence between top predator and intermediate consumer is fragile.     

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.     

Mark-recapture experiments on fish in Windermere, 1943–1982

Marking experiments on perch, Perca fluviatilis L., pike, Esox lucius L., and charr, Salvelinus alpinus L., have been carried out in Windermere. Numbered tags were used for individual identification, except in some short term experiments on charr. A total of 13 182 perch and 4696 pike were tagged; 2066 charr were tagged subcutaneously and marked with a fin clip. In the short-term experiments 2015 charr were marked with a fin clip or punched hole. Perch were recaptured up to 8 years after tagging, pike 12 years and charr 5 years. Estimates of numbers and mortality were not satisfactory, except for limited sections of the populations of pike and charr. The main reasons for the unsatisfactory results were: for perch, unknown mortality at time of tagging; for pike, selectivity of the gillnets and in some years low numbers tagged; for charr, restriction to few spawning sites and in some places low numbers recaptured. The experiments provided useful information on movements and growth, which could not have been obtained in any other way.     

FREQUENCY-DEPENDENT FITNESS CONSEQUENCES OF INTRASPECIFIC NEST PARASITISM IN SNOW GEESE

The reproductive efficiency, defined as the number of breeding recruits produced per egg laid; of intraspecific nest parasites; of hosts in parasitized nests; and of unparasitized nesting females, was measured for 14 years for lesser snow geese Anser caerulescens caerulescens nesting near Churchill, Manitoba, Canada. Relative efficiencies were 0.71–0.88, 0.91, and 1.0 for eggs of parasites, hosts, and unparasitized birds, respectively. Differences in the hatching probabilities of the three classes of eggs produced the efficiency differences. Parasitic success was limited by the parasites' failure to place more eggs than expected by chance into nests at the appropriate time relative to host incubation. Host nesting success was lower when more than one parasitic egg was added to the clutch. No differences in gosling survival and breeding recruitment probabilities were detected among any categories of goslings. Thus, hatching parasitic young are at no disadvantage relative to parental young, and there is no support for the hypothesis that increased success of host young at later stages of reproduction might offset negative effects at the egg stage. The hatching efficiency of parasitic eggs declined more rapidly than that of parental eggs as the parasitism rate increased. Efficiencies were similar when 3–4% of the eggs laid per year were parasitic, but relative parasitic efficiency was significantly lower when the parasitism rate was 8–9% or more. Using ancillary information and assumptions about the fecundity, viability, and behavioral flexibility of parasitic and parental females, we conclude that intraspecific nest parasitism could compete with nesting as a reproductive strategy in this population. The conditional use of parasitism by a large component of the population in certain years, however, combined with negative-frequency dependent success, limits the potential spread of a purely parasitic strategy in this population.     

Age and length at maturity of the female spiny dogfish,Squalus acanthias,in the Strait of Georgia,British Columbia,Canada

Synopsis A total of 3068 female spiny dogfish were examined to determine the age and length at maturity. The median age at maturity for females was 35.5 years with 95% confidence limits between 35.0 and 35.9 years. It was found that slower growing dogfish tended to mature at a smaller size. A deterministic model incorporating fecundity, growth and reproduction was used to examine the reproductive style of spiny dogfish. The age at maturity reported corresponds to the level that theoretically maximizes lifetime reproductive output for a cohort.     

Population dynamics of perch, Perca fluviatilis L. in Slapton Ley, Devon

Perch, Percafluviatilis L. were studied in Slapton Ley which is a eutrophic freshwater lagoon in south Devon, England. The perch were caught in traps from February 1970 to June 1971. The traps selected males during the spawning season. The water temperature did not affect the numbers of trapped fish directly, but the sight of perch in the traps appeared to attract others. Sex ratios were found to be equal in the first 2 years of life but older year classes were predominantly female. The population consisted of young individuals. Males were found to mature in their first year and most females matured in their second. The fecundity varied from an average of 1000 eggs in perch n i the 9.75 cm length group to 30,500 eggs in fish in the 27.75 cm group. A few shoals of perch moved throughout the Ley although the majority of fish stayed in the same area. Population estimates showed a range from 10,000 to 95,000 for fish 9.75 cm and above. The large variations in population estimates and survival rates were attributed to the necessity of using traps which had variable catches and to the harmful effects of handling the perch. Chironomids were important in the diet of perch of all sizes. Plankton and Asellus were other important foods, the latter especially during the winter months. Perch 11.5 cm to 13.9 cm selected the widest range of food organisms. Only perch 14.0 cm and above ate fish.     

Biotic and abiotic influences on the recruitment of male perch in Windermere, U.K.

Perch Perca fluviatilis recruitment (based on the catch per unit effort, CPUE, of 2 or 3 year male perch) varied greatly from 1941 to 1995 in Windermere, U.K., with year class strengths strongly synchronous between the two basins of this temperate, mesotrophic lake. Statistically significant modified Ricker (Saila‐Lorda) stock‐recruitment curves could be fitted to data from three of four sampling sites, while the presence of widespread disease and late summer water temperature were also important explanatory variables. Multiple‐regression analyses revealed no influences of pike Esox lucius recruitment, zooplankton abundance, or the climatic influences ( i.e . North Atlantic Oscillation and displacement of the summer position of the Gulf Stream) on perch recruitment over and above associated influences from temperature. Fitted models realistically explained up to c . 60% of the observed variation in perch recruitment.     

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.     

Sexual maturation of cod (Gadus morhua L.) in the southern Baltic (1990–2006)

Baltic cod, like other species, is susceptible to inter‐annual fluctuations in sexual maturation, depending on the length, age, sex, extent of the habitat area, and stock abundance of the cod population. Maturity is one of the biological indicators used to detect changes in a stock that can be caused by fishing. To address these issues specifically for the eastern Baltic cod stock, long‐term data (1990–2006) from Polish research vessels in the southern Baltic were examined. To date, the ICES has used the same maturity ogives over extended periods and assumed invariant sex ratios for the assessment of eastern Baltic cod. The combined maturity ogives calculated in the present study were markedly lower, particularly for age groups 2–4 (5), in all periods, than those used in the ICES assessment. Moreover, the proportion of females increased with length and age, suggesting that annual verification of the sex ratio is needed. The present study also revealed that the total length (L_50%) and the age (mean age‐at‐maturity; MAM_50%) at which 50% achieved first sexual maturity were higher for females than for males in the study period. The long‐term mean L_50% and MAM_50% for females were 43.9 cm and 4.3 years, respectively, and for males 34.8 cm and 3.4 years. There was also a spatial difference between calculated maturity ogives, with slightly lower L_50% (range: 1.4–8.6 cm) in the Gdańsk Basin than in the Bornholm Basin. The increasing trend in fishing mortality observed in 1993–2004 (ICES data) did not translate into a temporal trend in calculated maturity ogives. However, changes in L_50% and MAM_50% reflected recruitment variations (ICES data). The significance of these findings is discussed in the context of the environment and recruits abundance.     

Acidification affects the perch,Perca fluviatilis,populations in small lakes,of southern Finland

Synopsis Characteristics of perch were studied in 4 recently acidified lakes in southern Finland. The densities of the populations were 0–250 ha^–1 compared with 1400–3300 ha^–1 in two circumneutral reference lakes. The perch were close to extinction in one acidic lake and dominated by 7 year old fish in two other acidic lakes, suggesting decreased reproduction. Individual fecundity was not reduced by the acidity, but high mortality (close to 100%) of eggs in two acidic lakes indicated recruitment failure. In one of the acidified lakes reproduction had continued resulting in a population dominated by younger fish. However, increased egg mortality, low density, and the resulting high growth rate of perch in this lake were considered to be consequences of the acidity. These findings are the first documentation on the possible decrease of fish populations in Finland due to acid precipitation.