Reproductive potential of the eastern baltic cod <Emphasis Type="Italic">Gadus morhua callarias</Emphasis> L. population

Trends in interannual variation in maturation and spawning terms of various age cohorts in the Eastern Baltic cod population in 1997–2009 were studied. Specific features in the age structure of the mature population part that were established by the end of the first decade of the 21st century were clarified. The role of cod age cohorts in the current population reproduction was considered taking into account the data on cod recruitment and fecundity.     

The potential fecundity of cod in the Baltic Sea from 1993 to 1999

Investigations on the reproduction biology of cod in the Baltic Sea (Gadus morhua L. and G. morhua callarias L.) were carried out beginning in 1993, with yearly sampling of gonads to estimate the potential absolute and relative individual fecundity in the Belt Sea (subdivision, SD: 22) and in the Arkona Sea (SD: 24). Additional samples were taken in the Bornholm Sea (SD: 25). The potential absolute fecundity of the total area of investigation is described by the regression models: F_a = 6.89 × TL^3.09 and F_a = ?40 586 + 977.7 × W. As a further result the mean relative fecundity was 978 oocytes per gram body weight. Furthermore, it was shown that there was a difference in development of the potential individual fecundity between the Belt Sea cod and the original eastern Baltic cod stock. In contrast to the stable fecundity of cod in the Belt Sea, the fecundity of the large females in the Bornholm Sea significantly increased from 1993 to 1999. This change is related to a decrease in the dry weight of oocytes in the ovaries. In the same period this cod stock dramatically decreased.     

Reproductive potential of the Eastern Baltic cod Gadus morhua callarias L. population

Trends in interannual variation in maturation and spawning terms of various age cohorts in the Eastern Baltic cod population in 1997-2009 were studied. Specific features in the age structure of the mature population part that were established by the end of the first decade of the 21st century were clarified. The role of cod age cohorts in the current population reproduction was considered taking into account the data on cod recruitment and fecundity.     

Some issues related to reproduction of Pacific cod <Emphasis Type="Italic">Gadus macrocephalus</Emphasis> in waters of the eastern coast of the northern Kuril Islands and the southern extremity of Kamchatka

The analysis of data collected in December 1996 and 1998 on the reproduction of cod Gadus macrocephalus in Pacific waters of the northern Kuril Islands and the southern part of Kamchatka was performed. It was shown that the individual absolute fecundity of the cod varies within 0.197–9.729 million eggs and the relative fecundity, in the range of 24-1386 eggs. The fecundity of 1000 mature females comprises 2179–2449 million eggs. The low individual fecundity of fish is related to pseudobranchial tumor growth. The main role in cod reproduction is played by females of two-three size groups characterized by the highest numbers of mature females. It is suggested that different fecundity within the range of Pacific cod is related to environmental conditions, in particular, water temperature in the spawning grounds during spawning rather than to the habitation latitude.     

Spatio-temporal overlap of the alien invasive ctenophore <Emphasis Type="Italic">Mnemiopsis leidyi</Emphasis> and ichthyoplankton in the Bornholm Basin (Baltic Sea)

In 2007 the alien invasive ctenophore Mnemiopsis leidyi A. Agassiz 1865 was recorded for the first time in the Bornholm Basin, an area which serves as important spawning ground for Baltic fish stocks. Since M. leidyi is capable of preying upon early life stages of fish and further might act as food competitor for fish larvae, it is of major concern to investigate the potential threat that this non-indigenous species poses to the pelagic ecosystem of the Baltic Sea. The present study investigates the temporal and spatial overlap of M. leidyi with eggs and larvae of Baltic cod (Gadus morhua L.) and sprat (Sprattus sprattus L.) in order to assess the potential impact of this new invader on two of the most important Baltic fish stocks. Results show variable inter-seasonal distribution and overlap dynamics and thus different seasonal threat-scenarios for the early life stages of cod and sprat. The spatial overlap between M. leidyi and ichthyoplankton was low for most of the period observed, and we conclude that M. leidyi presently does not have a strong impact. However, we detected situations with high overlaps, e.g. for sprat larvae and cod eggs in spring. As the population dynamics of M. leidyi in the central Baltic are not yet fully understood, a future population explosion of the alien ctenophore with possible effects on fish recruitment cannot be ruled out. Furthermore, a possible shift in peak spawning of cod to the early season, when ctenophore abundances were relatively high, might increase the impact of M. leidyi on cod.     

Viability selection on early body mass and the effect of female body size on fecundity: a study on the leaf-eared mouse <Emphasis Type="Italic">Phyllotis darwini</Emphasis>

Viability selection and fecundity of size-related traits has been demonstrated to be strong in vertebrates. In small mammals, both offspring and adult size are important for viability and fecundity, respectively. We studied the role of early phenotypic selection on size attributes and female fecundity in the leaf-eared mouse (Phyllotis darwini). Our results support that larger females produce more offspring, and since the likelihood of attaining adulthood is similar for different sizes of the females, those larger females also produce more offspring that attain sexual maturity. From the offspring perspective, larger pups at birth have significantly more probability of attaining sexual maturity. However, weaning mass and growth rate did not show any differential survival. Our study suggests that early selection could be important and could prevent further episodes of selection by early culling of the distribution of sizes, and that “effective” fecundity is strongly dependent on the size of the female.     

Long-term variation in dates of mass spawning of cod Gadus morhua callarias (Gadidae) in the southeastern part of the Baltic Sea

Long-term data on the numbers of eggs of cod Gadus morhua callarias in ichthyoplankton and the seasonal variation of their distribution in the Gdansk Deep of the Baltic Sea are considered. The spring peak of spawning was observed for the years 1948–1954 and 1960–1964; the spring-summer peak for 1968–1977; and the summer peak for 1955–1959 and 1995–2001. Maximum egg numbers of cod were recorded from the late 1940s to the early 1950s; minimums were recorded in the late 1980s and the late 1990s. The decrease in the numbers of cod eggs in the ichthyoplankton, in general, coincided with a decrease in the proportion of spawners maturing at the beginning of the season of mass reproduction and a decrease in the oxygen content in the near-bottom layer of the Gdansk Deep. It is suggested that a shift of the spawning peak of cod from spring to summer is related to a decrease in the volume of near-bottom waters providing favorable conditions for the reproduction of this population.     

Phenotypic plasticity in growth and fecundity induced by strong population fluctuations affects reproductive traits of female fish

Fish are known for their high phenotypic plasticity in life‐history traits in relation to environmental variability, and this is particularly pronounced among salmonids in the Northern Hemisphere. Resource limitation leads to trade‐offs in phenotypic plasticity between life‐history traits related to the reproduction, growth, and survival of individual fish, which have consequences for the age and size distributions of populations, as well as their dynamics and productivity. We studied the effect of plasticity in growth and fecundity of vendace females on their reproductive traits using a series of long‐term incubation experiments. The wild parental fish originated from four separate populations with markedly different densities, and hence naturally induced differences in their growth and fecundity. The energy allocation to somatic tissues and eggs prior to spawning served as a proxy for total resource availability to individual females, and its effects on offspring survival and growth were analyzed. Vendace females allocated a rather constant proportion of available energy to eggs (per body mass) despite different growth patterns depending on the total resources in the different lakes; investment into eggs thus dictated the share remaining for growth. The energy allocation to eggs per mass was higher in young than in old spawners and the egg size and the relative fecundity differed between them: Young females produced more and smaller eggs and larvae than old spawners. In contrast to earlier observations of salmonids, a shortage of maternal food resources did not increase offspring size and survival. Vendace females in sparse populations with ample resources and high growth produced larger eggs and larvae. Vendace accommodate strong population fluctuations by their high plasticity in growth and fecundity, which affect their offspring size and consequently their recruitment and productivity, and account for their persistence and resilience in the face of high fishing mortality.     

Effects of host plant on life‐history traits in the polyphagous spider mite Tetranychus urticae

Studying antagonistic coevolution between host plants and herbivores is particularly relevant for polyphagous species that can experience a great diversity of host plants with a large range of defenses. Here, we performed experimental evolution with the polyphagous spider mite Tetranychus urticae to detect how mites can exploit host plants. We thus compared on a same host the performance of replicated populations from an ancestral one reared for hundreds of generations on cucumber plants that were shifted to either tomato or cucumber plants. We controlled for maternal effects by rearing females from all replicated populations on either tomato or cucumber leaves, crossing this factor with the host plant in a factorial design. About 24 generations after the host shift and for all individual mites, we measured the following fitness components on tomato leaf fragments: survival at all stages, acceptance of the host plant by juvenile and adult mites, longevity, and female fecundity. The host plant on which mite populations had evolved did not affect the performance of the mites, but only affected their sex ratio. Females that lived on tomato plants for circa 24 generations produced a higher proportion of daughters than did females that lived on cucumber plants. In contrast, maternal effects influenced juvenile survival, acceptance of the host plant by adult mites and female fecundity. Independently of the host plant species on which their population had evolved, females reared on the tomato maternal environment produced offspring that survived better on tomato as juveniles, but accepted less this host plant as adults and had a lower fecundity than did females reared on the cucumber maternal environment. We also found that temporal blocks affected mite dispersal and both female longevity and fecundity. Taken together, our results show that the host plant species can affect critical parameters of population dynamics, and most importantly that maternal and environmental conditions can facilitate colonization and exploitation of a novel host in the polyphagous T. urticae, by affecting dispersal behavior (host acceptance) and female fecundity.     

Lifetime fitness and age‐related female ornament signalling: evidence for survival and fecundity selection in the pied flycatcher

Ornaments displayed by females have often been denied evolutionary interest due to their frequently reduced expression relative to males, habitually attributed to a genetic correlation between the sexes. We estimated annual and lifetime reproductive success of female pied flycatchers (Ficedula hypoleuca) and applied capture–mark–recapture models to analyse annual survival rates in relation to the patterns of expression (absence/presence) of an ornament displayed by all males and a fraction of females. Overall, the likelihood of expressing the ornament increased nonlinearly with female age and was due to within‐individual variation, not to the selective appearance or disappearance of ornament‐related expression of phenotypes in the population. Accordingly, expressing the forehead patch in a given year did not influence survival probability. However, those females expressing the ornament at early ages (1–2 years old) enjoyed survival advantages throughout lifetime. Although ornamented females had higher lifetime fecundity and fledging success, their yearly reproductive performance, in terms of fledging productivity, decreased as they aged so that, late in life, ornamented females reared fewer offspring than nonexpressing females of the same age. In addition, both strategies (expressing vs. not expressing the trait) returned similar fitness payoffs in terms of recruited offspring. Our results support the hypothesis that fecundity and survival selection are involved in the displaying of this ‘male’ ornament by females.     

Oxygen and salinity characteristics of predator–prey distributional overlaps shown by predatory Baltic cod during spawning

In the distributional overlap volume of Baltic cod Gadus morhua and its prey, studied in the Bornholm Basin in the southern Baltic Sea, only a fraction of the sprat Sprattus sprattus population vertically overlapped with the Baltic cod population. Sprat occurred in the intermediate water, in the halocline and in the bottom water, while herring Clupea harengus and Baltic cod occurred exclusively in the halocline and in the bottom water. Only parts of the sprat population were hence accessible for Baltic cod, and only a fraction of the sprat had access to the Baltic cod eggs below the halocline. Baltic cod–clupeid overlap volumes appeared to be determined by salinity stratification and oxygenation of the bottom water. Hydrography time series were used to estimate average habitat volumes and overlap from July to September in 1958–1999. In the 1999 survey spawning Baltic cod had greater ratios of empty stomachs and lower average rations than non-spawning Baltic cod. The average ration for Baltic cod caught within 11· 4 m from the bottom (demersal) did not differ from the average ration of Baltic cod caught in shallower waters (pelagic), because spawning and non-spawning Baltic cod in both strata were caught at equal rates. The diet of the Baltic cod caught demersally contained more benthic invertebrates, especially Saduria entomon, but Baltic cod caught pelagically also had fresh benthic food in their stomachs, indicating vertical migration of individual fish.     

Regulation of population numbers in the <Emphasis Type="Italic">Rana arvalis</Emphasis> frog according to the results of long-term observations on the same population

An analysis of the results of long-term stationary studies (1987–1996) on a Rana arvalis population showed that the mean body length and reproductive parameters of females increased with age. Significant interannual changes in body length and egg size (but not in fecundity and reproductive effort) were also accounted for by conditions of the warm season preceding the breeding season. The dependence of egg diameter and reproductive effort on the female body length weakened with age, and only the relationship between body length and fecundity was characterized by positive allometry. The survival rate of mature females depended on the premetamorphic survival rate and, therefore, was higher in earlier generations, in which the aquatic phase proceeded under more favorable conditions. The rate of reproduction (R ₀) decreased and, therefore, further population growth slowed down only after the initial abundance of the generation had reached a relatively high level. An increase in the initial abundance of the generation exerts a strong influence on R ₀ via a decrease in the survival of premetamorphic animals and, to a lesser extent, via a decrease in the body length and fecundity of tree-and four-year-old females. Changes in the annual abundance of breeding females in R. arvalis were more abrupt and less prolonged than those in the sympatric R. temporaria population. The holding capacity of the environment for the aquatic phase of the R. arvalis life cycle was higher than that for R. temporaria. Therefore, the population abundance of R. arvalis, compared to R. temporaria, was not subject to stringent negative feedback regulation dependent on the initial number of animals.     

Fishing,reproductive volume and regulation: population dynamics and exploitation of the eastern Baltic cod

The relative importance of exploitation rate and environmental variability in generating fluctuations of harvested populations is a key issue in academic ecology as well as population management. We studied how the eastern Baltic cod (Gadus morhua) is affected by fishing and environmental variation by using a newly developed single species state-space model. Survey data and auxiliary environmental data were used to estimate the model parameters. The model was then used to predict future development of the eastern Baltic cod under different fishing mortalities and abiotic conditions. Abiotic condition was represented by an index: reproductive volume which is the volume of water suitable (in terms of salinity and oxygen content) for the successful development of the early life stages of Baltic cod. The model included direct density dependence, fishing, and a lagged effect of reproductive volume. Our analysis showed that fishing rate is approximately three times more important than reproductive volume in explaining the population dynamics. Furthermore, our model suggests either under- or over-compensatory dynamics depending on the reproductive volume and long term catch levels. It follows that fishing can either reduce or increase temporal oscillations of the cod stock depending on whether the dynamics is over- or undercompensatory, respectively. The sustainable level of fishing rate is however dependent on reproductive volume. Our model predicts a dual role of fishing rate, stabilizing when reproductive volume is high and destabilizing when it is low. Exploitation rate may therefore increase or decrease the risk of the population of cod dropping below a given biomass reference point depending on the environmental conditions, which has practical implications for fisheries management.     

Age-specific survival and fecundity and their effects on the intrinsic rate of increase of Aphelinus gossypii (Hym., Aphelinidae), a parasitoid of Aphis gossypii (Hom., Aphididae)

Abstract: The age-specific survival and fecundity of female adults of the aphidophagous parasitoid, Aphelinus gossypii Timberlake (Hym., Aphelinidae), were determined at a host density of 50 Aphis gossypii Glover (Hom., Aphididae) per leaf of Ageratum houstonianum Mill each day at 25°C. The age-specific mummy production, emergence rate and sex ratio of progeny were calculated. The implication of these results in terms of potential population growth of A. gossypii and related species is discussed. The age-specific survival curve ( l _x ) of females exhibited a Type I pattern, which resulted in little difference between Σ m _x and Σ l _x   m _x . The l _x curve of males exhibited a Type II pattern, and their survival time was much shorter than that of females. Each mated female produced on average 598.9 ± 64.0 aphid mummies, and preyed on 87.9±6.2 aphids. Most of the biological performance parameters were not different significantly between mated and virgin females. The highly female-biased sex ratio of offspring produced by mated females soon after their emergence resulted in a high fecundity rate ( m _x ) during the early reproductive period. Therefore, although the sex ratio of progeny was male-biased during the entire reproductive period, the intrinsic rate of increase estimated using age-specific sex ratios was larger than that estimated using a constant sex ratio of 0.5. These results indicate that the high survival and the adaptation of producing mostly female offspring during the early reproductive period contribute much to the population increase potential of this parasitoid.     

The influence of self‐fertilization and grouping on fitness attributes in the freshwater snail Physa acuta: population and individual inbreeding depression

The genetic structure, selfing rate and inbreeding depression of the hermaphroditic freshwater snail Physa acuta were jointly analysed in a population near Montpellier, France. Allozymic markers revealed moderate gene diversity (0.138), and no heterozygote deficiency. The mean outcrossing rate, estimated by using progeny arrays, was 0.9, with substantial variation among families. This also suggests that the number of fathers among outcrossed offspring of a given mother is low. Inbreeding depression was estimated over more than one generation using 83 first‐laboratory‐generation (G₁) families. The main parameters measured were parental (G₁) fecundity, offspring (G₂) survival and fecundity. Size and growth were also monitored. Parental fecundity was analysed under several conditions (isolation, pair and quadruplet outcrossing). The self‐fertilization depression, including parental fecundity, offspring survival and fecundity, was about 0.9 at the population level. The genetic data obtained in the same population indicate a value of about 0.3 using Ritland’s (1990) technique, suggesting that the depression over the whole life‐cyle might be even higher than 0.9. Grouping affected neither fecundity nor self‐fertilization depression. Substantial variation in depression for survival was detected among individuals, from no survival in some selfed families to better survival than that of outbred families in others. The overall result (outbred population structure, high outcrossing rate and high self‐fertilization depression) is consistent with what is expected in large outcrossing populations in which inbreeding depression is maintained by mutation‐selection balance.     

Size Matters: Individual Variation in Ectotherm Growth and Asymptotic Size

Body size, and, by extension, growth has impacts on physiology, survival, attainment of sexual maturity, fecundity, generation time, and population dynamics, especially in ectotherm animals that often exhibit extensive growth following attainment of sexual maturity. Frequently, growth is analyzed at the population level, providing useful population mean growth parameters but ignoring individual variation that is also of ecological and evolutionary significance. Our long-term study of Lake Erie Watersnakes, Nerodia sipedon insularum, provides data sufficient for a detailed analysis of population and individual growth. We describe population mean growth separately for males and females based on size of known age individuals (847 captures of 769 males, 748 captures of 684 females) and annual growth increments of individuals of unknown age (1,152 males, 730 females). We characterize individual variation in asymptotic size based on repeated measurements of 69 males and 71 females that were each captured in five to nine different years. The most striking result of our analyses is that asymptotic size varies dramatically among individuals, ranging from 631–820 mm snout-vent length in males and from 835–1125 mm in females. Because female fecundity increases with increasing body size, we explore the impact of individual variation in asymptotic size on lifetime reproductive success using a range of realistic estimates of annual survival. When all females commence reproduction at the same age, lifetime reproductive success is greatest for females with greater asymptotic size regardless of annual survival. But when reproduction is delayed in females with greater asymptotic size, lifetime reproductive success is greatest for females with lower asymptotic size when annual survival is low. Possible causes of individual variation in asymptotic size, including individual- and cohort-specific variation in size at birth and early growth, warrant further investigation.     

Fecundity of the European pine sawfly Neodiprion sertifer (Hymenoptera, Diprionidae) related to cyclic outbreaks: Invariance effects

At the end of ontogenesis, individual variability of the fecundity of females of Neodiprion sertifer constitutes 30–120 eggs (with rare fluctuations from 16 to 140 eggs). Average fecundity as an adaptive index in a certain biocenosis, however, depends on many factors, including the structural organization of the sawfly population. The presence of a long diapause in some individuals results in the cleavage of the offspring into groups with different dates of reactivation. At the same time, mainly heavy eonymphs, giving rise to large females, stay in the diapause. Their participation in the reproduction together with females, raised in other years under different conditions, equalize the dimensional structure of the breeding fraction of the population. During population waves, revealed during 35 years of my continuous observations, fecundity fluctuated closely to the average value with deviations constituting 19–25%. It hardly depends on the population density of the phyllophagan. The intraspecific structuredness of the population in relation to the size of females and its transformation by the moment of their emergence promotes the evolutionary stability of the degree of fecundity of the sawfly.     

Breeding phenology,variation in reproductive effort and offspring size in a tropical population of the woodlouse Porcellionides pruinosus

Summary Most species of woodlice in temperate habitats have discrete breeding seasons. It is hypothesised that breeding synchronises with favourable environmental conditions to maximise offspring growth and survivorship (Willows 1984). We measured the breeding phenology of a species introduced to a tropical environment, primarily to consider the assumption that life histories in the tropics will differ fundamentally from those in temperate habitats. In addition to breeding phenology we considered variation in reproductive effort between individual females and the division of this effort between the size and number of young.A continuous breeding phenology was observed in a synanthropic population of Porcellionides pruinosus within the tropics. Reproductive effort varied between months, showed a weak relationship with female size and was independent of female fecundity. Female sizefecundity relationships varied between samples and when the proportion of reproductive females was high size-fecundity slopes were steeper than at other times. Mean offspring size varied between months and there was a wide range in offspring size within broods. Offspring size was not related to female body mass, reproductive effort or fecundity; consequently brood mass increased linearly with an increase in fecundity. Increased reproductive effort goes into more rather than larger offspring.We propose that the continuous breeding in this population was the result of the constant presence of an environmental cue to reproduction evolved in temperate habitats. Continuous breeding is not necessarily equivocal to high individual reproductive success even though overall population growth may be rapid. However, variation in reproductive effort suggests that individuals respond to current environmental conditions on short time scales.     

Effects of food availability on reproductive output,offspring quality and reproductive efficiency in the apple snail <Emphasis Type="Italic">Pomacea canaliculata</Emphasis>

Phenotypic plasticity in life history traits favors the establishment of invaders and may magnify their ecological impacts. Pomacea canaliculata, the only freshwater snail listed among the 100 worst invaders worldwide, is able to complete its life cycle within a wide range of conditions, a capacity attributed to its life history plasticity. Using snails from their native range in Argentina we investigated the changes in fecundity, egg mass traits, offspring quality, and efficiency of food conversion into eggs in response to different levels of food availability throughout different life stages. Pre-maturity mortality was not affected by chronic reductions of up to 80% in food availability. Females fed ad libitum demonstrated no significant reproductive output differences when mated with males raised at different food availability levels. For females, the number and total weight of eggs and the size of egg masses decreased at high levels of food deprivation. Their efficiency of conversion into eggs of the food ingested during the reproductive period increased with deprivation, as did the survival time of their offspring. In contrast, the egg mass laying rate and the individual egg weight did not differ under different food availability regimes. Reductions in food availability have been suggested as a control method but our results indicate that fecundity would be lessened only at deprivation levels higher than 50% and would be partially compensated by an increase in hatchling survival.     

Does Breeding Ornamentation Signal Genetic Quality in Arctic charr, <Emphasis Type="Italic">Salvelinus alpinus</Emphasis>?

Genetic theories of sexual selection predict that most ornamental secondary sexual traits provide reliable indication of the genetic quality of their bearers. Accordingly, also the offspring of mates with elaborate mating display should perform better than those of less conspicuous counterparts. In this study, we used Arctic charr (Salvelinus alpinus) as a model species to investigate whether the variation in a carotenoid-based red breeding coloration (a sexually dichromatic trait) in different sexes would reflect differences in individual genetic variability, one measure of individual quality, and/or indirectly, be manifested in variation in the offspring’s early viability and growth. We created maternal half-sibling families by artificially fertilizing the eggs with milt from bright- and pale-coloured males and then held the resulting progenies under identical hatchery conditions. The expression of red coloration among parental fish was not associated with their genetic diversity estimates in either sex nor did offspring sired by bright males consistently differ in terms of embryo survival or endogenous growth efficiency from offspring sired by pale males. By contrast, maternal effects were notably strong and, additionally, the degree of female coloration was negatively linked to their reproductive potential. The more intensely coloured females had a smaller relative fecundity and they also produced offspring of lower viability, implying a significant trade-off in resource allocation between ornamentation and offspring. Our results indicate that the red breeding ornamentation of Arctic charr is likely to be informative rather among females than males when the reproductive quality is predicted on grounds of the number of offspring produced. Nevertheless, this study does not support the direct selection hypothesis in explaining the evolution of female ornamentation, but rather suggests that the less intense coloration of female charr compared to males may reflect inter-sexual differences in the trade-off between natural and sexual selection.