Patterns in vendace recruitment in Lake Pyhäjärvi, south-west Finland

A vendace Coregonus albula population has long formed the basis of a flourishing fishery in Lake Pyhäjärvi, south-west Finland. Between 1971 and 1990, it exhibited variations in year-class strength generally characterized by a 2-year cycle of alternating strong and weak year classes, which implied density-dependent regulation probably due to asymmetrical food competition between age groups. The abundance of piscivorous predators in the lake and the wanning of water after the hatching of larvae in spring together determined the final year-class strength of vendace. Since 1990, year-class sizes have remained very low and the 2-year cycle has been disrupted. The continuously low levels of recruitment could not be explained by an increased larval mortality, although the decline started with 2 successive years with very high mortality of this life stage which reduced the whole spawning stock to a fraction of earlier levels, and consequently led to low larval numbers such that between 1988 and 1996 larval abundance was positively correlated with the spawning stock size. The importance of the spawning stock size to recruitment could also be seen in the whole 26-year time series as a significant correlation between year-class sizes with a 2-year lag. The observed recruitment patterns of the vendace stock in Lake Pyhäjärvi thus show the importance of spawning stock size, intraspecific competition, predator effects and abiotic environmental variation.     

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.     

Monthly variability of hydroacoustic fish stock estimates in a deep lake and its correlation to gillnet catches

Monthly (April to November) hydroacoustic surveys and parallel gillnet catches were used to determine vendace Coregonus albula abundance, biomass and population dynamics in a deep oligotrophic lake. By hydroacoustic surveys, recruitment of 0+ year vendace could clearly be detected. In contrast, gillnet catches resulted in low numbers of 0+ year fish, and similarly under-represented proportions of the oldest vendace. Consequently, the correlation between hydroacoustics and gillnets with respect to fish numbers and geometric mean fish total length was high only for the age groups 1 + to 4+ years. Annual variability in hydroacoustic estimates offish abundance and biomass was high (CV=26–29%) which reflects the seasonal population dynamics of vendace.     

Year-class fluctuations in vendace,Coregonus albula (Linnaeus): Who's got the upper hand in intraspecific competition?

The vendace Coregonus albula (L.) populations in the lakes Mjøsa and Osensjøen exhibited fluctuating year-class strength. In Mjøsa, a strong year-class emerged every third year, except for the four year period between the strong year-classes 1969 and 1973. The difference between the strong and weak year-classes decreased from the 1960s, through the 1970s to the 1980s. The Mjøsa vendace matured sexually at age 2 +, and more than ten sexually mature age-groups were present in the population. Growth ceased at maturation, and asymptotic length was 23.6 cm. In Osensjøen, one strong year-class (1969) dominated the population during the period 1976–1987. The Osensjøen vendace matured sexually at age 3, and more than 15 sexually mature age-groups were present in the population. Growth ceased at maturation, and asymptotic length was 28.4 cm. In both lakes, vendace fed on crustacean zooplankton in the epilimnion throughout summer and autumn. Our data indicate that regular year-class oscillations occur as a result of the juvenile survival being negatively correlated to the number of adults.     

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

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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.     

Hydrological attributes and rheophilic freshwater fish: stock assessment

The mathematical and statistical advances in fitting stock assessment models enabled the emergence of the paradigm of “integrated analysis”, which fits all available data to a single model of population dynamics that traditionally has total catch as the only forcing function of the system. This approach, however, allowed us to include, in a flexible way, the effect of hydrological regime as an additional forcing function. We tried to achieve this flexibility by making the annual recruitment rates and spawning biomass adjustable to the attributes of the hydrological cycle data. Our models showed that these attributes are influential in the population dynamics of Brycon hilarii of the Northern Pantanal, and their inclusion in the models allowed best partial fits (which considered fits only to the data components length- and age-compositions, CPUE of juveniles and adults) than the Base-case (without hydrological attributes). The best partial fits where obtained when the attributes “delay of floods” and “intensity of floods” were forcing the spawning biomass and the annual recruitment respectively, indicating that these characteristics of the population may be influenced by specific attributes of the water level. The use of integrated modeling contributed with the advancement of population ecology knowledge of rheophilic fish. It is recommended that freshwater fisheries management be integrated into the hydrology management.     

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.     

Compensatory dynamics and the stability of phytoplankton biomass during four decades of eutrophication and oligotrophication

The link between compensatory dynamics and regime shifts is not well understood. We analyse a regime shift in phytoplankton in a large lake with respect to: (1) environmental forcing and (2) the type of dynamics (compensatory or synchronous) between phytoplankton groups. The regime shift in phytoplankton was related to gradual changes in nutrient levels, but unrelated to an almost concurrent shift in climatic conditions. The relationship between total phytoplankton biomass and phosphorus concentrations was sigmoid. Trajectories of phytoplankton biomass and community dynamics suggest that eutrophication effects can successfully be reversed when management efforts decrease nutrient loading to a level sufficiently low to overcome community resilience. The regime shift was associated with a loss of biomass compensation and compensatory dynamics among the phytoplankton groups. This suggests that the type of interactions is important for a better understanding of the existence and shape of nonlinear responses of phytoplankton biomass to environmental change.     

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.     

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.     

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.     

The role of temperature and cannibalism in interannual recruitment variation of bass in British waters

Between-year variation in bass Dicentrarchus labrax year-class strength in southern British waters is investigated. Mean spring-summer seawater temperature in the year of birth was significantly positively correlated with both the level of summer recruitment of the 0 + group to the estuarine nurseries and subsequent recruitment of III + fish to the adult population. Spectral analysis of the temperature-compensated time series showed that a statistically significant proportion of the variation not attributable to temperature was periodic at 0–33 cycles year⁻¹. Therefore, a simple, three-parameter model, combining a linear relationship between temperature and abundance and a second-order autoregressive model can be used to describe and predict variation in relative adult YCS. Bass remain for their first 3 years within their estuarine nursery areas. As I + fish were observed in Southampton water to cannibalize the 0 + group, it is suggested that strong year-classes suppress recruitment for the next 2 years even if the temperature is suitable to promote a strong year-class. Cyclic variation in recruitment caused by intraspecific interactions, particularly cannibalism, may be a feature of other marine fish that use estuaries as nursery areas.     

Possible impacts of environmental change on pelagic fish recruitment: modelling anchovy transport by advective processes in the southern Benguela

Climate change may influence pelagic fish by altering advective processes or by changing where fish choose to spawn. Using a simulation model, the effects of altered advection and spatial distribution of spawning by anchovy on recruitment off South Africa were explored. Cape anchovy Engraulis capensis spawn on the region of the Agulhas Bank, south of South Africa. Currents transport eggs and larvae to nursery areas. Transport of eggs and larvae was modelled using a flow field based on averaged Acoustic Doppler Current Profile data, Feasible Scenarious of altered advection were modelled. For modelling purposes, the ocean surrounding South Africa was divided into blocks with dimensions of a quarter-degree latitude by a quarter-degree longitude. Acoustically measured distributions of spawner biomass for the years 1986–92 were used to calculate egg production per block. In the model, batches were released from each of these blocks each day of the spawning season. The modelling study indicates that passive transport of young anchovy may account for a substantial proportion of year-class variability. Model results show that distribution of spawners influences the distribution of young of the year, as well as the number and the location of advective losses across offshore boundaries.     

Long-term variability and density dependence in Hudson River Dreissena populations

1. We used a 27-year record of Dreissena populations in the freshwater tidal Hudson River to describe interannual variation in population density, body size, and body condition; estimate long-term variation in recruitment, survivorship, and shell growth; and assess possible controls on the populations.
2. Dreissena populations in the Hudson have been highly variable, with interannual ranges of c. 100-fold in abundance and biomass, and 7-fold in mean body mass. This large interannual variation arises from both long-term trends and 2–5-year cycles.
3. Long-term trends include the 2008 appearance of the quagga mussel (Dreissena rostriformis), which still forms a small part (<10%) of the dreissenid community, and a decline in zebra mussel body size. The decline in body size was caused by a long-term decline in adult survivorship rather than a decline in rates of shell growth. We could detect no long-term trends in adult abundance or spread of Dreissena onto soft sediments in the Hudson.
4. We observed persistent, strong cycles in adult abundance and body size. These were driven by the appearance and decay of eight dominant year classes over the 27 years of our study, and were a result of temporal variation in recruitment rather than temporal variation in survivorship. The observed strongly irregular recruitment appears to arise from strong adult–larval interactions, and is consistent with previous simulation model results showing that interactions between adults and larvae can drive persistent cycling.
5. We found evidence that negative density dependence affects recruitment, somatic growth, and body condition of Dreissena in the Hudson. Warm summers may also cause high adult mortality.
6. We put our results into the context of a conceptual model of Dreissena population dynamics, and argue that neither the dynamics nor the controls of populations of these important invaders is known satisfactorily.

     

Population studies based on relative abundance of five different life history stages of Ayu,Plecoglossus altivelis (Pisces,Plecoglossidae), in Lake Biwa

1. Comparison of relative abundance of each two successive life history stages was made for studying population dynamics of ayu, Plecoglossus altivelis, in Lake Biwa.
2. Number of eggs spawned is proportional primarily to the size of biomass of spawner population although spawning success plainly depends on presence or absence of heavy rain in the spawning period.
3. Although density of fry tends to be proportional to that of egg, yearly fluctuation of the fry mortality may be caused by changes in density of food organisms but is not in compensatory way.
4. From the fry stage to the juvenile one the rate of mortality and/or of growth seem to fluctuate in compensatory way which is supposed to be related to a shortage of food.
5. Yearly fluctuation of mortality of adult stage is caused by an artificial factor as changes of fishing intensity.
6. Necessity of studying actual events at each stage of the life history process separately in relation with environmental conditions is discussed through the result of the present analysis.

     

Representing Density Dependent Consequences of Life History Strategies in Aquatic Ecosystems: EcoSim II

EcoSim II uses results from the Ecopath procedure for trophic mass-balance analysis to define biomass dynamics models for predicting temporal change in exploited ecosystems. Key populations can be represented in further detail by using delay-difference models to account for both biomass and numbers dynamics. A major problem revealed by linking the population and biomass dynamics models is in representation of population responses to changes in food supply; simple proportional growth and reproductive responses lead to unrealistic predictions of changes in mean body size with changes in fishing mortality. EcoSim II allows users to specify life history mechanisms to avoid such unrealistic predictions: animals may translate changes in feeding rate into changes in reproductive rather than growth rates, or they may translate changes in food availability into changes in foraging time that in turn affects predation risk. These options, along with model relationships for limits on prey availability caused by predation avoidance tactics, tend to cause strong compensatory responses in modeled populations. It is likely that such compensatory responses are responsible for our inability to find obvious correlations between interacting trophic components in fisheries time-series data. But Ecosim II does not just predict strong compensatory responses: it also suggests that large piscivores may be vulnerable to delayed recruitment collapses caused by increases in prey species that are in turn competitors/predators of juvenile piscivores. Received 24 February 1999; accepted 3 August 1999.     

Ecological investigations of the zooplankton community of Balsfjorden,northern Norway: Population structure and breeding biology of the chaetognath Sagitta elegans Verrill

The population dynamics of the chaetognath Sagitta elegans Verrill has been followed in Balsfjorden in 1976 and 1977. Seasonal variation in abundance, length-frequency distribution, growth in total length, and maturity stages are presented and discussed in relation to changes in hydrography.An annual generation of S. elegans was found, with a protracted and more or less continuous breeding season from May until October during 1977. The 1976 year-class consisted of two distinct length groups, both of which participated in the 1977 spawning. This spawning gave rise to possibly four sub-populations during 1977. The variation in numbers of sub-populations produced during the spawning season in 1976 and 1977 is discussed in relation to the hydrographical conditions in Balsfjorden. From November 1976 to March 1977 the abundance of S. elegans varied between 1 and 8 ind. · m^?3. The lowest value was recorded in May (0.9 ind. · m^?3). From September to December 1977 the population abundance was ≈2 ind. · m^?3.     

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.     

On the assessment of the reproductive capacity of eastern Baltic cod <Emphasis Type="Italic">Gadus morhua callarias</Emphasis> (Gadidae) by ichthyoplankton data

Annual egg production in eastern Baltic cod Gadus morhua callarias in 1966–2000 has been assessed. A significant relationship between egg production and recruitment, progeny survival index, and 11psu-isohaline depth has been revealed. Comparison of egg production and spawning stock biomass indices has shown the time difference in their maxima and the absence of significant correlation between them. This difference is assumed to be due to an increase in the share of mature individuals that did not participate in spawning in the late 1970s–early 1980s. The determination of potential egg production, taking into account the share of individuals that spawn during the current year, yielded a significant positive relationship between its value and spawning stock biomass.