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.     

Expulsion of miniature radio transmitters along with eggs of muskellunge and northern pike––a new method for locating critical spawning habitat

Identification and protection of critical spawning habitat for muskellunge Esox masquinongy and northern pike Esox lucius is important for preserving the reproductive potential of both species. In this study, we implanted miniature radio transmitters through the oviduct into the egg masses of female muskellunge and northern pike just prior to spawning. This non-surgical procedure was a novel approach for identifying spawning sites when transmitters were expelled with the eggs during egg deposition. Preliminary studies in three lakes showed that muskellunge and northern pike deposited many of the transmitters in likely spawning habitat. An inability to find eggs limited our validation of this method, but nevertheless, a relatively high proportion (70%) of northern pike larger than 690 mm (27.2 inches) expelled transmitters in a previously known spawning area in Willow Lake, Minnesota. Shoreline vegetation in that area consisted primarily of sedges Carex spp., and the adjacent water was shallow with substrate consisting of large mats of water bulrush Scirpus subterminalis. A lower proportion (50%) of muskellunge expelled transmitters in Elk Lake, Minnesota. Water depth at likely spawning sites averaged 1.1 m (3.6 feet) and vegetative cover was variable, but Chara spp. was common to most sites. In Moose Lake, Minnesota, containing sympatric populations of muskellunge and northern pike, 60% of muskellunge and 90% of pike expelled transmitters. Chara spp. beds were the predominant substrate where transmitters were expelled in Moose Lake, but the two species deposited transmitters on deepwater bars (3.7–5.2 m) in addition to shallow near-shore habitat. These results suggest more flexibility in depths used for spawning than typically reported for muskellunge and northern pike.     

Egg production of the marine,planktonic copepod,Calanus pacificus Brodsky: Laboratory observations

When acclimated to a continuous, superabundant food supply and constant temperature, Calanus pacificus Brodsky females produce eggs at a weight-specific rate ranging from 0.13 · day^?1 at 8°C to 0.21 · day^?1 at 15°C. Maximum weight-specific egg production rates do not change with seasonal changes in female body size. The relationship between egg production rate and food concentration is hyperbolic, with threshold and critical concentrations that are high relative to other species for which data are available. Food concentration and temperature influence spawning frequency (i.e., the time required for oocytes to mature) much more than the number of eggs in a single spawning event (i.e., clutch size). Clutch size is significantly related to female body size.     

Intensive fishing can mediate stronger size-dependent maternal effect in pike (Esox lucius)

In pike E. lucius L., evidence on maternal effect on reproductive output is mixed. We studied whether older and larger pike females produce eggs and larvae of higher quality (weight, starvation resistance) in three forest lakes in southern Finland. Later, the study lakes were subjected to intensive experimental pike fishing, which we assumed would increase resource availability and lead to higher maternal investment (larger egg size). Length of female pike was positively correlated with the dry weight of eggs and larvae but this relation was dependent on female age. In old females, the effect of female length on egg weight was lower or even negative. Survival analysis showed a positive effect of female length on larval survival time indicating that larvae from larger females are less vulnerable to starvation during the early stage of life. After the intensive pike fishing, the positive effect of female length on egg weight was stronger in all age classes probably due to the released resources. Based on the high quality and amount of reproductive products in large (but not very old) females, they are important for the reproduction of pike populations. This should be considered in fisheries management.     

Reproductive biology of brown trout, Salmo trutta macrostigma Dumeril 1858, in a tributary of the Ceyhan River which flows into the eastern Mediterranean Sea

The study describes some key elements of the reproductive biology, including spawning season, age at sexual maturity, fecundity and egg diameter of the native brown trout, Salmo trutta macrostigma, in a tributary of the Ceyhan River. A total of 197 brown trout (118 females and 79 males) were captured in 2000–2001 by electric fishing. In observations on monthly changes, the gonadosomatic index (GSI) and the monthly frequency distribution of egg diameter confirmed that spawning lasted from November to January. Some 27.7% of the females and 62.5% of the males attained sexual maturity in their second year. The smallest fork length (FL) of brown trout attaining sexual maturity was 17.4 cm for males and 17.8 cm for females. Mean fecundity in age groups II, III, IV and V were 360, 452, 693 and 1283 eggs per female, respectively. One 9‐year‐old female had a unique 3232 egg count. The mean fecundity of the sampled population was 554 eggs per fish, positively correlated with the FL (mm) (R = 0.8227 ) and body weight (R = 0.8130). The diameter of mature eggs in the spawning season ranged from 3.250 to 5.930 mm, with a 4.146 mm average. Mean egg diameter in age groups II, III, IV and V in the spawning season were 0.813, 3.799, 4.663 and 5.243 mm, respectively. Fecundity, egg weight and diameter were statistically different in all age groups.     

Potential effects of spawning habitat changes on the segregation of northern pike (<Emphasis Type="Italic">Esox lucius</Emphasis>) and muskellunge (<Emphasis Type="Italic">E</Emphasis>. <Emphasis Type="Italic">masquinongy</Emphasis>) in the Upper St. Lawrence River

Changes in spawning habitat of northern pike (Esox lucius) may affect their segregation from and coexistence with the closely related muskellunge (E. masquinongy). We estimated the areal coverage of robust and shallow emergent vegetation in three shared-spawning bays in the Upper St. Lawrence River from aerial photographs taken from 1948 to 2003. Robust emergent vegetation (e.g., cattail) increased in coverage by 155–241% while shallow emergents (sedges) decreased by 46–96%. The loss of sedges, an important northern pike-spawning habitat, may facilitate greater spawning overlap in offshore-submersed aquatic vegetation within bay habitats used by muskellunge. Development rates and characteristics of northern pike and muskellunge eggs and larvae were compared to better understand the implications of greater spawning overlap. Northern pike eggs developed faster than muskellunge eggs at temperatures of 4.7–19°C, and adhesive eggs and the presence of adhesive papillae were present in both species. Equations were used to predict degree-day requirements for hatching and swim-up in three habitats (shallow emergents, bay, and offshore shoal) along a temperature gradient. Northern pike required more estimated degree days to reach hatching in bay and offshore shoal habitat relative to shallow emergent habitat due to cooler temperatures. Significant spawning overlap is known to occur within bay habitats, but poor success of northern pike in deep bay habitats and overall reductions in abundance are hypothesized to currently buffer muskellunge from potential negative interactions between these species. Guest editors: J. M. Farrell, C. Skov, M. Mingelbier, T. Margenau & J. E. Cooper International Pike Symposium: Merging Knowledge of Ecology, Biology, and Management for a Circumpolar Species     

The spawning activity of cod, Gadus morhua L.

Aspects of the reproduction of reared cod, Gadus morhua L., with special emphasis on the females, were studied under laboratory conditions. The fecundity and condition factor were 2–5 and 1–5 times, respectively, that of wild cod. A total of 18 spawning females were kept in separate tanks/ chambers, each with one or two males. Seven of the 18 females were classified as stressed, based upon behaviour, irregular spawning intervals and low fertilization rates of the eggs. The reared cod were found to spawn 17–19 batches. The number of eggs liberated in each batch normally followed a smooth, dome-shaped curve with time. The fertilization rate was normally 100%. Egg size decreased from first to last batch and the egg dry weight decreased by about 20–30%. The reared cod showed the same egg diameter to dry weight relation as wild cod. Egg diameter of first batch and maternal fish length were significantly positively correlated. The mean spawning interval for the female and the mean water temperature during its spawning were negatively correlated. The reared cod spawned in both the night and the day for about 50–60 days.     

Effect of spawning number and ration on reproductive performance of the batch-spawning three-spined stickleback Gasterosteus aculeatus

An experiment quantified the effect of food ration and spawning number on the breeding season reproductive performance of batch-spawning, female three-spined sticklebacks Gasterosteus aculeatus . Individually housed females were maintained on fixed rations of 2, 4, 8 or 16% of their initial postspawning mass of live enchytraeid worms from immediately after their first spawning until they ceased spawning. Number of spawnings correlated positively with ration. Total breeding season reproductive investment (total wet mass of eggs produced as a percentage of initial postspawning female mass) ranged from 38% at the 2% ration to 147% at the 16% ration. At the lower rations, postspawning mass, batch fecundity and wet and dry masses of the batch declined over successive spawnings, with the rate of decline inversely related to ration. At the highest ration, there was no decline in batch fecundity and postspawning mass increased over successive spawnings. Mean reproductive investment per inter-spawning interval was higher at the highest ration, but at all rations declined over successive spawnings. Mean reproductive effort per inter-spawning interval (wet mass of eggs spawned as a percentage of the wet mass of food consumed over the inter-spawning interval) was inversely related to ration. At the higher rations, reproductive effort showed no trend over successive spawnings. Neither spawning number nor ration had a systematic effect on egg diameter, wet mass per egg, dry mass per egg or total lipid content of the eggs. If the rate of food intake was insufficient, although batch fecundity declined, the main adjustment was a reduction in the number of spawnings in the breeding season.     

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.     

Two eggs,two different constraints: a potential explanation for the puzzling intraclutch egg size dimorphism in Eudyptes penguins

Phenotypic plasticity and phenotypic stability are major components of the adaptive evolution of organisms to environmental variation. The invariant two-egg clutch size of Eudyptes penguins has recently been proposed to be a unique example of a maladaptive phenotypic stability, while their egg mass is a plastic trait. We tested whether this phenotypic plasticity during reproduction might result from constraints imposed by migration (migratory carry-over effect) and breeding (due to the depletion of female body reserves). For the first time, we examined whether these constraints differ between eggs within clutches and between egg components (yolk and albumen). The interval between colony return and clutch initiation positively influenced the yolk mass, the albumen mass, and the subsequent total egg mass of first-laid eggs. This time interval had only a slight negative influence on the yolk mass of second-laid eggs and no influence on their albumen and subsequent total masses. For both eggs, female body mass at laying positively influenced albumen and total egg masses. Female investment into the entire clutch was not related to the time in the colony before laying but increased with female body mass. These novel results suggest that the unique intraclutch egg size dimorphism exhibited in Eudyptes penguins, with first-laid eggs being consistently smaller than second-laid eggs, might be due to a combination of constraints: a migratory carry-over effect on the first-laid egg and a body reserve depletion effect on the second-laid egg. Both these constraints might explain why the timing of reproduction, especially egg formation, is narrow in migratory capital breeders.     

Interspecific differences of parental polychlorinated biphenyl exposure on nutrient availability, egg production and brooding in two Syngnathus species

Parental nutrient contribution of two closely related, sympatric pipefish species was investigated following exposure to mid-range environmental concentrations of the polychlorinated biphenyl (PCB) mixture Aroclor 1254 (mean ± s . e . = 52·89 ± 2·12 μg l⁻¹). This study tested the hypothesis that differential nutrient supplementation during development predicts interspecific sensitivity to environmental stressors. Specifically, paternal PCB exposure was predicted to affect Syngnathus fuscus development because embryos depend on paternally derived nutrient sources in the brood pouch. Conversely, changes in Syngnathus floridae egg production were expected following maternal PCB treatment as a nurse-egg system generates the nutrient-rich pouch fluid. Lipid concentrations in adult blood plasma were significantly elevated following Aroclor 1254 treatment in S. floridae gravid females and post-brooding S. fuscus males ( P < 0·001) which suggested mobilization from body tissues in response to metabolic demands. Aroclor 1254 treatment also resulted in decreased protein concentrations in S. fuscus pouch fluid ( P < 0·05). These changes were not translated into significant exposure-related effects in the nutrient reserves of eggs and fry or fry size. Overall S. fuscus demonstrated a general sensitivity to disturbances from laboratory exposure conditions. Whereas a previous investigation of field-collected pipefishes showed comparable interspecific pouch fluid and fry nutrient concentrations and egg size, here, nutrient levels and egg diameter decreased in both laboratory-mated control and PCB-exposed S. fuscus . This study demonstrates the potential physiological effects of PCB exposure on parental nutrient allocation in syngnathids and the general sensitivity of S. fuscus to environmental disturbances.     

Predation of Northern Pike (Esox lucius L.) Eggs: A Possible Cause of Regionally Poor Recruitment in the Baltic Sea

Northern pike (Esox lucius) spawning habitat and egg mortality were studied in three spawning areas in 2001 along the Swedish coast of the Baltic proper: the Blekinge Archipelago, Kalmar Sound and coastal freshwater streams. Spawning peaked during the last week of April in streams, at temperatures ranging from 7.7 to 8.9 °C and during the first week of May in brackish waters, at temperatures ranging from 8.9 to 13.8 °C. Spawning occurred in shallow waters, at depths between 0.2–1.5 m, but generally most of the spawn was found in the shallowest areas. In streams, eggs were mainly attached to emersed vegetation, while in brackish sites pike eggs were well scattered among flooded emersed plants, submersed plants and filamentous algae. Mean egg density varied between 469–1829 m^–2 with the lowest density observed in Kalmar Sound. The calculated egg loss occurring from approximately one day after spawning to one day before hatching ranged from 41±7% in coastal streams to 67±6% in the Blekinge Archipelago and 100% in Kalmar Sound. The significant removal of eggs from spawning sites in Kalmar Sound and Blekinge was most likely due to predation from several fish species. In situ observations and stomach analyses suggested that many pike eggs in Kalmar Sound were lost to the three-spined stickleback (Gasterosteus aculeatus), a species that presently dominate the littoral small fish community. This study therefore suggests that egg predation by sticklebacks and other fishes may be a possible cause of the reported poor recruitment of coastal pike populations in the Kalmar Sound region.     

Assessment of egg quality and realised fecundity of whiting Merlangius merlangus L. in captivity

Potential predictors of egg quality were assessed in whiting Merlangius merlangus L. permitted to spawn in a tank from which eggs were collected. These included fertilisation rate, the proportion of viable buoyant eggs, egg diameter, and egg wet and dry weights; all were influenced by temporal effects and were negatively correlated with days from start of spawning. The spawning period was protracted, from February to June. Mean daily egg production per female was 2.74 ± 2.43 g and 2338 ± 2075 eggs, equivalent to 14.6 ± 13.1 g kg^?1 day^?1 female^?1. Egg diameter was 1.21 ± 0.04 mm, egg wet weight 1.20 ± 0.21 mg, dry weight 0.10 ± 0.02 mg, and mean fertilisation rate and hatching rates were 76.8 and 73.3%, respectively. The incubation period ranged from 72 to 80 degree days and was dependent on temperature (x) and described by the equation y = 25.92 e^?0.1219x. Realised fecundity was also assessed to determine if this gave a more accurate measure of reproductive potential, and this was compared with potential fecundity estimated from predictive regressions on fish length from fisheries data. Realised egg production of 20 females of 185 g mean weight and 256 mm fork length was 4 444 360 (95% CL 4 093 961–4 743 018), similar to predicted seasonal egg production based on gravimetric fecundity measurements of wild caught fish.     

Natural spawning,egg and fry production of milkfish,Chanos chanos (Forsskal), broodstock reared in concrete tanks1

Gonadal maturation, rematuration, and spawning of nine- (1982 stock) and ten-year-old (1981 stock) milkfish reared in 150 and 200 m³ concrete tanks were observed in 1990 and 1991. From 23 September to 9 November 1990, the 1981 stock spawned 15 times with an average daily collection of 61,000 ± 21,000 eggs. From 4 larval rearing trials, the mean overall survival rate (from eggs to 21 -day-old fry) was 9.68 ± 4.3%. From 25 Aril to 31 October 1991, the stock spawned a totaf of 108 times. Mean daily egg collection increasef to 305,000 ± 32,000. Successful rearing trials conducted during the latter part of the spawning season had a mean overall survival rate of 17 ± 3% (n = 17) indicating that mass fry production techniques can be improved. Spawning of milkfish broodstock in concrete tanks provides an alternative to other existing methods of seed production.     

Relationship Between Condition,Aggression, Signaling,Courtship, and Egg Laying in the Field Cricket,Gryllus assimilis

Sexual selection theory suggests males in good condition should be more successful than males in poor condition when competing with rivals for territories and mates. Understanding how condition influences the interplay between aggression, mate attraction, and courtship displays could help explain why variation is maintained in traits that confer fitness. Using laboratory‐reared Jamaican field crickets, Gryllus assimilis, we found that fine‐scale temporal components of mate attraction signals were positively correlated with body condition (residual body mass) and body size; signaling effort was positively correlated with both body condition and fine‐scale temporal signaling components; aggression was positively correlated with signaling effort; number of eggs laid was positively correlated with female body size, male body condition and aggression. Together our correlative study suggests that variation in body condition and size may drive some of the variation in cricket mate attraction signaling and aggression. Given condition and body size are influenced by foraging ability, nutrient availability and the organism’s ability to uptake and retain these essential nutrients could explain some of the persistent variation in fitness conferring traits.     

The influence of maternal length and age on the size and weight of the eggs and the relative fecundity of the haddock, Melanogrammus aeglefinus, in British waters

The diameter and dry weight of the eggs of haddock, Melanogrammus aeglefinus , are positively correlated with fish length. The correlations are largely due to the fact that many of the smaller fish that were sampled were 2 years old, and the eggs of these young haddock, which can be regarded as precocious spawners, are significantly smaller and lighter than those of older fish. The relative fecundity of 2–year–old haddock (274 eggs g⁻¹) is also lower than that of the other age classes (493 eggs g⁻¹); this has important implications for the estimation of egg production from female spawning stock biomass. It is pointed out that in some years precocious spawners represent a large component of the North Sea haddock spawning stock. When the annual egg production of this stock is calculated, it may be appropriate to apply a weighting factor to the number of eggs contributed by the 2–year–olds, on the assumption that the small eggs of these fish produce larvae that are less viable than those of older haddock.     

The size composition of the pelagic eggs of walleye pollack <Emphasis Type="Italic">Theragra chalcogramma</Emphasis> in spawning areas of the northern Sea of Okhotsk

The data on the size composition of the pollack developing eggs of walleye pollack collected in the northern Sea of Okhotsk in 2001 during mass spawning are given. The eggs caught in western Kamchatka and in Shelikhov Bay had the greatest diameter; they were somewhat less in the northern Okhotsk Sea area; and the smallest eggs were found in eastern Sakhalin. The clinal variability of the size composition of walleye pollack eggs was observed in the northern Sea of Okhotsk, and in the western and eastern parts it had a local pattern. Parameters such as spawning female size composition, temperature, and salinity, as well their influence on the size composition of eggs, were considered.Original Russian Text Copyright © 2004 by Biologiya Morya, Ovsyannikov.     

Lake sturgeon spawning on artificial habitat in the St Lawrence River

In 1996, lake sturgeon (Acipenser fulvescens) spawning was documented for the third consecutive year on an artificially placed gravel bed in the St Lawrence River. Two distinct spawning periods were observed in 1996. Spawning initially commenced on 17 June, when water temperature reached 15°C. A second spawning event was documented from 28 June to 1 July (16°C). Sturgeon egg densities were monitored in three transects on egg trays, on the gravel surface, and within interstitial spaces in the gravel. Counts of developing eggs in the gravel bed during both spawning periods were used to estimate a total of 275 000 eggs on the study area (0.075 ha). Average egg density was highest in the transect with the highest water velocities. Lake sturgeon fry were first observed in the gravel on 24 June (15.5°C), and first emergence from the gravel was documented on 28 June. Hatching following the second spawning event commenced on 3 July. Based on assessment of average embryo viability (61.6%) and egg‐to‐emergent fry survival (17.6%) an estimate of about 171 000 sturgeon eggs hatched, producing over 49 000 emergent fry. Current velocity, substrate particle size, depth of substrate, and maintenance of sediment‐free interstitial spaces are important considerations in planning future spawning habitat enhancement projects.     

Transmission experiments with pike fry (Esox Indus L.) rhabdovirus

Experimental infection of fertilized pike eggs with 'red-disease' virus produced 100% mortality in the fry. This mortality was associated with a disease that had previously been described as hydrocephalus internus, indicating that 'red-disease' and hydrocephalus are different manifestations of the same disease. The name pike fry rhabdovirus disease (PFRD) is suggested for the disease complex, and the name pike fry rhabdovirus (PFR) for the causative agent. Exposure of PFR to a Wescodyne * solution containing 25 ppm of iodine resulted in an inactivation of at least 99–99% of viral activity within 30 sec. Experimental egg transmission of PFR could be interrupted by disinfecting the eggs in a Wescodyne solution, suggesting that the virus was located on the egg surface. Conclusive evidence of a naturally occurring vertical transmission in pike culture is still lacking because, using FHM cells as a detection system, PFR could not be found in spawners and their sexual products. The susceptibility of pike fry to PFR rapidly decreases at increasing age.     

The relationship between fecundity and fertility in the mouthbrooding cichlid fish Tilapia leucosticta

The development of fry in the mouth of the female mouthbrooder Tilapia leucosticta Trewavas is described. The egg production by fish of various lengths is plotted and a curvilinear regression for standard and total length fitted. The number of eggs produced is approximately equal tothe square of the total length in centimetres of the parent fish. The brood numbers of 460 femalesare plotted against length, and linear regressions for mean brood number and mean fry number fitted. The relationship between the expressions for numbers of fry brooded and egg production gives anestimate of brooding efficiency that decreases with increasing parent length. Individual female fish show signs of reripening and the timing of this process is estimated from gonad state and from the length of fry brooded in the mouth. The relationship between fecundity (egg production) and fertility (fry brooded) is considered and the effect of maturation size, growth rate and spawning frequency assessed.