Population regulation of vendace (Coregonus albula) in Lake Pyhäjärvi, southwest Finland

Previous time series analysis on vendace population dynamics in Lake Pyhäjärvi, 1971–1990, revealed a 2-year cycle in year-class strength, implying powerful density-dependent regulation. Here we have extended this analysis by using multiple regression models to test whether the recruitment series is influenced by density-independent factors. We chose population size with a lag of 1 year as the density-dependent factor; the density-independent factors were the summer water temperature with a lag of 2 years (temperature sum for June, July and August, indicating the year-class strength of predators) and the temperature-derived length of the larval period of vendace. For the years 1972–1990 the coefficient of determination ( r ²) of this regression model was 0·77. We suggest that the basic mechanism producing a persistent 2-year cycle of vendace in Lake Pyhäjärvi is the asymmetrical food competition between age groups. The abundance of predators in the lake and the warming of the water after the hatching of larvae in spring together determine the final year-class strength of vendace.     

Fish stock assessment in lakes based on mass removal

The efficiency of mass removal of fishes can potentially be assessed using catch statistics collected during intensive fishing periods. The calculations are based on the general assumption that a decline in population size will produce a decline in catch per unit effort (cpue). When the removal is efficient it is possible to estimate the population size both at the beginning of the fishing period and after the removal.
Two examples are based on the winter seine net fishing of vendace ( Coregonus albula ) in Lake Pyhäjärvi, SW Finland in (1) 1983–1984 and (2) 1989–1990. The effects of the error in catch composition samples and random changes in probabilities of capture during the removal period on the final estimates of initial stock sizes are also examined.
The precision of the population estimates shown here was greatly influenced by the violation of the underlying assumption that the probability of capture is equal for all members of the target age group. In any case, particular attention should be paid to optimizing the sampling programme so that it will reveal the best information on the exploited stock with the resources in hand.     

The size of a vendace, Coregonus albula L., stock in a deep lake basin monitored by hydroacoustic methods

An acoustic estimate of the number and biomass of a vendace stock (age >2 + years) in a deep basin of Lake Karjalan Pyhäjärvi was made in August 1985. The acoustic data were collected at night during the summer stagnation. The vendace were on the lake bottom during the day and rose by midnight to the hypolimnion; they did not rise to the thermocline or water layers above it. In the research area the mean number of vendace was 1900 fish ha ⁻¹ and the mean biomass was 76 kg ha ⁻¹. Total vendace biomass was 151 in the whole 200-ha research area.     

Two-year cyclicity in recruitment of a fish population is driven by an inter-stage effect

A 25-year time series for a vendace (Coregonus albula (L.)) population was analysed with mathematical models to reveal the factors that regulate its recruitment. The fitted recruitment model was an age-structured model incorporating the compensatory effects of the spawning population and the abundance of the previous year class. Wind forcing index was also added as an example of an external source of recruitment variability. The auto-correlation analysis revealed a tendency for 2-year generation cyclicity in recruitment. The compensatory effects of both spawning biomass and previous year class abundance on recruitment had to be incorporated into the model to remove cyclicity from residuals. Wind forcing during the larval and early juvenile periods negatively affected recruitment. Re-estimating the parameters of the Cushing recruitment function and the effect of previous year-class from simulated data revealed that the time series structure and measurement errors induce strong biases in parameters exaggerating the density independent population growth rate parameter and the amount of compensation. The negative effect of previous year class was also exaggerated but less severely. Simulations with the artificially perturbed deterministic model skeleton revealed a tendency for cyclicity in recruitment. The model typically generated dampening oscillations, but the dynamics appeared as limit cycles when assuming high mortality, a low level of compensation by spawning biomass and a considerable negative effect of the previous year-class. No single ultimate mechanism for inter-stage effects causing cyclicity can be presently designated despite the rather extensive studies on vendace population dynamics. Several of the suggested mechanisms may be operating in concert.     

Unpredictability of fish recruitment: interannual variation in young–of–the–year abundance

In 1989–1998, vendace larvae Coregonus albula were sampled in Finnish lakes following a stratified random sampling design. The abundance of young–of–the–year vendace after the first growing season was estimated using catch–per–unit–effort statistics. The number or total area of nursery places hardly limited the recruitment of vendace in the study lakes. The major proportion of prerecruit mortality of vendace occurred in the larval phase and larval sampling produced significant information on young–of–the–year survival. Although larval abundances and recruitment were clearly associated only in Lake Onkamo, generally high larval abundance was needed to produce high number of recruits.     

A validated bioenergetics model for ruffe Gymnocephalus cernuus and its application to a northern lake

A bioenergetics model was constructed for ruffe Gymnocephalus cernuus , and the model was used to estimate the food consumption of a native G. cernuus stock in Pyhäjärvi, a large mesotrophic lake in south-west Finland. The parameters for the bioenergetics model were determined through laboratory experiments conducted at several temperatures between 5 and 22° C for different sized G. cernuus (from 6 to 27 g). The model was verified and validated with independent laboratory growth experiments. The validated model was applied to field data from Pyhäjärvi to estimate the food consumption of G. cernuus and its effects on the zoobenthos community during the years 2002–2004. The total annual food consumption of the G. cernuus stock ranged from 11·6 to 15·7 g m⁻² (wet mass). Chironomidae was the most abundant prey, 81% of the total consumption. The annual total production of zoobenthos in Pyhäjärvi was 15·2 g m⁻² (wet mass). Therefore, it can be concluded that G. cernuus stock is likely to exert a significant influence on the zoobenthos community in this lake, indicating a potential role of G. cernuus as an influential invasive species in many freshwater ecosystems in Europe and North America.     

Formation of year-class strength in the bream population in the shallow eutrophic Lake Tjeukemeer

Formation of year-class strength of bream Abramis brama , the most abundant fish in shallow eutrophic Lake Tjeukemeer, was studied during a 14-year period. Although the size of the spawning stock of bream was rather stable during this period, analysis of length—frequency distributions indicates that it comprised only a few year-classes. The strength of a year-class was determined at a fork length of about 11 cm, at the end of the second summer. Mortality rates of fish larger than 11 cm were low (<50% year⁻¹). As the number of fish at the end of the second summer was not correlated to the number of 0+ in the previous year, mortality rates during the second summer (ranging from 75.6 to 97.7% year⁻¹) determine the final year-class strength. These mortality rates were positively correlated to the year-class strength of 2- to 4-year-old pikeperch. The abundance of this predator, therefore, is likely to be the most important factor controlling the year-class strength of bream.     

Egg quality and female condition in vendace (Coregonus albula L.) before and during spawning

The fat content of muscle, liver and eggs of vendace (Coregonus albula L.) were measured during the maturation of ovaries in summer. The fat content of eggs was at its highest in July and that of muscle in August during the optimal feeding season. Significant differences were not found in fat content between lakes Suvasvesi and Southern-Kallavesi in central Finland. There were no essential differences in fatty acid composition between muscle, liver and eggs of Lake Suvasvesi vendace. During the last month before spawning in October the gross energy and fat contents of eggs decreased in Lake Suvasvesi vendace. The fat and vitamin C contents of muscle as well as the liver index of females decreased during this time also. In males the corresponding decrease did not occur, indicating that spawning is a greater stress for females than for males. The egg quality and condition of Lake Inarijärvi vendace were about the same as those of Lake Suvasvesi vendace. The unfavourable conditions of Lake Inarijärvi, located in northern Finland, are reflected in the size of ovaries, which is known to be smaller (gonadosomatic index < 20%) than in Lake Suvasvesi (GSI 26–27%). Vendace, then, maintain the quality of eggs at the expense of their number.     

Temperature-driven changes in early life-history stages influence the Gulf of Riga spring spawning herring (<Emphasis Type="Italic">Clupea harengus m</Emphasis>.) recruitment abundance

Processes occurring during early life-history stages influence the year-class abundance of marine fish. We found that the abundance of 1-year-old spring spawning herring is statistically significantly determined by the number of post-flexion herring larvae in the Gulf of Riga (Baltic Sea). The abundance of consecutive developmental stages of larvae: yolk-sac, pre-flexion, flexion and post-flexion strongly correlated with each other, indicating that factors which already influence the yolk-sac stage are important in determining the abundance of post-flexion herring larvae. Winter air temperature before spawning determined the timing of maximum abundance of pre-flexion herring larvae, but not their main prey: copepod nauplii, implying that different mechanisms governing major preconditions for the formation of year-class strength. The abundance of post-flexion larvae displayed a potential dome-shaped relationship with sea surface temperature experienced after hatching. We suggest that increased summer temperatures, which exceed the physiological optimum negatively, affect the survival of post-flexion herring larvae. Overall, future climate warming poses an additional risk to larval herring survival and this may lead to a reduction in those herring stock which rely on recruitment from shallow coastal areas.     

Effects of mass removal and variable recruitment on nutrient excretion by a planktivorous roach stock

The effects of intensive fishing and fish recruitment on the nutrient excretion rate by a planktivorous roach Rutilus rutilus stock in Lake Vesijärvi were estimated using a bioenergetics model. The values were compared with those given by the model assuming that no mass removal occurred or that the mass removal ceased after 2 years. According to the model, during June-August 1989 phosphorus (P) excretion by the pelagic roach stock corresponded to 18% of the annual external P loading and showed a maximum of 0.4 mg m⁻² day⁻¹. As a result of mass removal, the maximum daily P excretion declined to 0.18 mg m⁻² by 1990 and to 0.10 mg m⁻² by 1992. The reduction was due to the lower number of roach, while the excretion rates of individual fish were unaffected. A 2-year mass removal would have temporarily reduced the P excretion rate, but in 1991 and 1992 the highest daily excretion rates would have been doubled compared with values after the 4-year mass removal. Assuming no fishing mortality, interannual differences were diminished and the excretion rates in 1991 and 1992 were 2.5 times higher than the values after the 4-year mass removal. The results suggest that nutrient recycling by fish stocks may be affected substantially by fisheries management. However, the effects are dependent on the timing of fishing due to the variations in the year class strength of fish. In Lake Vesijärvi the reduction in nutrient excretion was amplified coincidentally by the natural variation in recruitment of the roach stock.     

The influence of food competition and host specificity on the transmission of Triaenophorus crassus (Cestoda) and Cystidicola farionis (Nematoda) to Coregonus lavaretus and Coregonus albula (Pisces:Coregonidae) in Finland

As a superior competitor for planktonic food, vendace (Coregonus albula), when abundant, is expected to displace whitefish (Coregonus lavaretus) from feeding on plankton and to force it to rely more on benthic food. The predicted result would be a reduced abundance of the copepod-transmitted cestode Triaenophorus crassus in whitefish, but an increase in the abundance of the nematode Cystidicola farionis transmitted via benthic amphipods. We studied the occurrence of both parasites in whitefish during 1991-1996 in three interconnected areas at Lake Saimaa, Finland, where the densities of the vendace stocks varied due to natural fluctuation in year-class strengths. In accordance with our hypothesis, some indication of the effect of the density of the vendace population on abundance of C. farionis infection in whitefish was found, but not in the case of T. crassus. Only 0.2% of vendace were infected with T. crassus, while up to 100% of the whitefish in the yearly samples harboured the parasite. In further experiments we clarified which copepod species in Lake Saimaa act as first intermediate hosts of T. crassus, and verified from stomach samples the exposure of both whitefish and vendace to those species. Experimental infections indicated that the infectivity of T. crassus is lower for vendace than for whitefish. We suggest that the reason why vendace stock density does not affect T. crassus infection in whitefish is that T. crassus is transmitted in littoral areas during a short period in spring. At that time of the year copepods are abundant and available to both whitefish and vendace, but since the parasite is less infective to vendace. they do not become infected. At other times of the year, dense vendace stocks may force whitefish to shift to benthic food, which includes amphipods transmitting C. farionis. During vendace stock decline, whitefish may, however, continue to feed on plankton and avoid exposure to C. farionis.     

Effective size,census size,and genetic monitoring of the endangered razorback sucker, <Emphasis Type="Italic">Xyrauchen texanus</Emphasis>

For the past several decades the Lake Mohave population of the federally endangered razorback sucker, Xyrauchen texanus, has had no natural recruitment because of intensive predation on larvae by non-native fishes. In response to impending extirpation, a repatriation program was implemented where larval razorbacks are collected from the wild following natural spawning, reared in protective custody, and then repatriated at a much larger size. In this study, we estimated annual (N _bf) and generational female effective size (N _ef) of the spawning stock by characterizing temporal genetic changes in mtDNA among larval cohorts, and then compared these estimates to the estimated number of reproductively capable (adult) females in Lake Mohave (N _f). Razorback suckers have life history and mortality schedules that could yield values of N _ef/N _f as low as 10⁻⁵ due to match–mismatch recruitment (or the “Hedgecock effect”) that increases variance in reproductive success. Average N _bf was estimated to be 160, N _ef was 706, and the ratio N _ef/N _f was 0.29 and 0.38 for arithmetic and harmonic mean N _f, respectively. Our findings indicate that (i) larval sampling in Lake Mohave sufficiently encompasses temporal and spatial variation within annual larval cohorts as to be representative of the spawning stock, (ii) roughly 3–16% (about 8% on average) of adult females contribute genetically to larval cohorts each year, and (iii) repatriated fishes appear to be contributing genetically to larval cohorts. Simultaneous genetic and demographic monitoring offers insights that neither approach can provide alone into effects of population decline and management practices in this species.     

Morphological variation of perch Perca fluviatilis in humic lakes: the effect of predator density,competition and prey abundance

Between and within‐lake variations in morphology of perch Perca fluviatilis were studied in four humic lakes in eastern Finland. Perca fluviatilis were more streamlined and smaller headed in a lake with the highest abundance of cyprinids, but lowest abundance of predators (Lake Tuopanjärvi), indicating adaptation to planktivorous feeding and low predator density. Highest bodied fish were found from a lake with the lowest cyprinid but highest predator abundance (Lake Koppelojärvi), which conversely indicates adaptation to more effective predator avoidance. Furthermore, the length of the paired fins was longest in Lake Kinnasjärvi and Lake Tuopanjärvi, where the abundance of benthic macroinvertebrates was lowest, suggesting selection for more effective benthivory. Clear morphological differences of P. fluviatilis between habitats were found only in Lake Kinnasjärvi, whereas in Lake Koppelojärvi and Lake Tuopanjärvi only the length of the paired fins differed and in Lake Harkkojärvi no differences were found. Taken together, these results suggest that inter and intrapopulation morphological differences are probably highly dependent on different biotic factors (i.e. predation risk, resource availability and competition). Spatial and temporal variations in these factors may have a great effect on body morphology of P. fluviatilis.     

Minnow predation on vendace larvae: intersection of alternative prey phenologies and size-based vulnerability

Time-intensive sampling was used to study minnow Phoxinus phoxinus density and foraging activity in the littoral area of Lake Lentua at a time of high vendace Coregonus albula larval abundance. Minnow activity and foraging during the late spring-early summer period at low temperatures was found to be mainly nocturnal and quite consistent with features reported in the literature, with the exception of feeding on vendace larvae. The absence of the latter finding from previous studies may be due to previous seasonally limited daytime samplings. The temporal or spatial scale of sampling is decisive when studying foraging on food animals with a brief period of vulnerability and seasonal phenology. However, minnows in Lake Lentua seemed to prefer Bosmina longispina whenever available and low B. longispina density during the first weeks after the break-up of ice directed minnow predation towards the vendace larvae. Estimated gross predation values showed that the minnow has the potential to become a remarkable predator on fish larvae. Slight variations in the vernal timing of the vendace hatching and larval development with respect to minnow activity, both of which are obviously temperature related, may be critical to vendace larval survival in the nearshore zone. However, the predation on the larvae is probably a strong factor only for a short period and the survival of the vendace larvae may be more related to the availability of suitably sized food resources than to predation by the minnow.     

A 27-year study of brown trout population dynamics and exploitation in Lake Songsjøen, central Norway

From 1968–1984 (period I), a brown trout Salmo trutta , population in a 70-ha oligotrophic lake in central Norway was exploited using larger mesh gill-nets selectively removing the larger fish. From 1985–1994 (period II), intermediate sized fish were removed using smaller-mesh sizes gill-nets. Fishing mortality and CPUE were correlated positively with effort and numbers of fish >3 years old for period II. The gill-net catchability was correlated negatively with spawner biomass and number of trout >3 years old. The significant positive correlation between natural mortality and stock biomass and spawning stock biomass indicated density-dependent mortality. The significant correlation between spawning stock and recruitment described by the Ricker model, indicated density-dependent recruitment of 1-year-old trout. The fishing regimes in the two periods affected the population dynamics and density differently. Selective removal of smaller fish permitted the larger fish to survive, and was beneficial in reducing fish density and maintaining stocks at low levels, consequently, achieving the expected increase in fish growth rates.     

Dispersion of vendace eggs and larvae around potential nursery areas reveals their reproductive strategy

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Feeding and fate of wild larval razorback sucker

The razorback sucker (Xyrauchen texanus) is disappearing throughout its native range in the Colorado River basin of western North America. The largest remaining wild population in Lake Mohave, Arizona-Nevada, has shown no recruitment since the 1950s. Although annual spawning is successful and larvae are seasonally abundant, no juveniles have been collected in recent decades. To evaluate the potential role of food availability in determining fate of larvae, fish and zooplankton samples were taken in 1985 from the reservoir and an adjacent, isolated backwater in which larvae were naturally produced. Food availability and primary dietary constituents were similar in both habitats. Reservoir larvae selectedBosmina spp. (Cladocera) and apparently avoided Copepoda, while larvae from the backwater selectedBosmina, but avoided Rotifera. Larvae from both places showed evidence of selection for certain sizes of zooplankters, but preferred sizes differed between habitats. These differences were neither attributable to larval size nor zooplankton community structure. Nutritional factors such as type, number, or size of available foods do not explain disappearance of larval razorback suckers from Lake Mohave, since larvae survive to far greater ages and size in the backwater. Predation by introduced fishes appears a significant cause of larval mortality.     

Krill (Euphausia superba) density, proportional and absolute recruitment and biomass in the Elephant Island region (Antarctic Peninsula) during the period 1977 to 1997

The Elephant Island region (Antarctic Peninsula) was selected as a long-term monitoring site to describe the interannual variability of important krill stock parameters. The analysis reviewed and updated krill density and proportional recruitment indices. Krill absolute recruitment and biomass from net sampling surveys are introduced as additional indices from this time series. New survey results from the past two seasons indicate a very successful 1994/1995 year-class and slightly below average proportional recruitment of the 1995/1996 krill year-class. Absolute recruitment of the 1995/1996 year-class was high compared to preceding years, because total stock size was relatively high in 1996/1997. After a period of low krill density and biomass in the area for almost a decade, krill density and biomass have increased. Caution is expressed as to whether this observed increase represents a long-term recovery of the stock. Received: 15 October 1997 / Accepted: 5 January 1998     

Influence of climate variability on whitefish (<Emphasis Type="Italic">Coregonus lavaretus</Emphasis>) year-class strength in a deep,warm monomictic lake

The year-class strength (YCS) of Blaufelchen (Coregonus lavaretus) in deep Upper Lake Constance was analysed for a 52-year period, from 1947 to 1998. Despite strong anthropogenic influences on the species' population dynamics due to cultural eutrophication and oligotrophication, intense fishing, and large-scale stocking, the influence of climate variability associated with the North Atlantic Oscillation (NAO) is apparent in the data set. This influence is significant although large-scale stocking of cold-bred larvae was performed to avoid a mismatch of larvae with their food. The importance of stocking on YCS, however, is unclear and was only detectable when analysing a subset of the data. In addition to climate variability a yet unidentified factor related to zooplankton suitability as food for fish larvae, and density-dependent mortality probably related to cannibalism do significantly influence YCS. The NAO seemed to influence YCS twofold, through temperature effects on egg development time and on larval growth rate. The first of these two mechanisms is related to the NAO via a time lag of 1 year due to the specific mixing dynamics of warm monomictic Lake Constance. Hence, a warm winter in the year before spawning results in earlier hatching of larvae, that is, hatching is decoupled from the actual meteorological conditions. This should make the larvae very prone to mismatch the dynamics of their food. However, we found no evidence for such a mismatch in this 52-year data set.     

Individual-based modelling of recruitment variability and biomass production of bay anchovy in mid-Chesapeake Bay

Production of bay anchovy Anchoa mitchilli is highest in the larval and juvenile stages. The interplay between vital rates, stage durations, prey resources, and anchovy abundance ultimately determines the relative magnitude of recruitment (which in the model varies by about three-fold) and of stage-specific production. Changes in adult seasonal spawning patterns that increase the number of larval survivors result in only a slight increase in overall production due to density-dependent decreases in growth rates of later life stages. Bay anchovy in the mid-Chesapeake Bay may reach a compensatory threshold during late summer-autumn as fish growth is affected by competition for food resources. Density dependence in the population is evident in the relationships between spawner-recruit, size-recruit, and production of larval or juvenile to young-of-the-year life stages. Density-dependent growth acts differentially upon the early life stage that exceeds the compensatory threshold in any given year, due either to environmental variability or population size, or both. This could explain partially the relatively low recruitment variability observed for this anchovy.