Reproductive seasonality of the detritivorous fish <Emphasis Type="Italic">Cyphocharax abramoides</Emphasis> (Kner, 1958) (Characiformes: Curimatidae) in flooded rivers of the eastern Amazon

The Lower Anapu River, Eastern Amazon, has a discrete flood pulse with narrow flooded areas due to the reduced influence of tides. In this environment, local precipitation might act as an important factor in the input of detritus, which can serve as a trigger for fish reproductive period. Therefore, this paper aims to determine the effects of local precipitation on the reproductive ecology of an abundant detritivorous fish in flooded rivers. Between May 2012 and April 2013, 886 specimens (471 females and 415 males) were captured monthly using gill nets. Individuals were measured, weighted, sexed, and the gonadal development stage was defined macroscopically and confirmed histologically. Sex ratio, length-weight relationship (LWR), condition factor (k), Gonadosomatic Index (GSI), relative frequency of maturity stages, and size at first gonadal maturity (L₅₀) were estimated and tested in relation to rainfall periods. There was no difference between the expected sex ratio (1:1), and both males and females showed polyphasic growth. The condition factor indicated higher values at the beginning of the rainy season and lower values during the dry season for both sexes. GSI and relative frequency of mature individuals indicated a long reproduction period with gonadal investment starting within the dry season until early in the rainy season for spawning within the rains. The L₅₀ value showed that males reach sexual maturity before females. Results indicated a reproductive pattern strongly influenced by rainfall, corroborating the importance of local precipitation for lacustrine environments.     

Annual variation in energy reserves in male three-spined stickleback, Gasterosteus aculeatm L. (Pisces, Gasterosteidae)

Wet and dry weights of tissue were measured and concentrations of glycogen, lipid and protein were estimated for the liver, gonad and carcass of male sticklebacks from an annual population collected each month over one complete year. In young-of-the-year there is one period of rapid weight gain, in all three body regions (liver, carcass and gonads) but particularly of the carcass, in the autumn and a second in spring and early summer. This is accompanied by an increase in the water content of all three body regions. The gonadosomatic index also increases sharply in spring and early summer. Young sticklebacks accumulate lipid and glycogen slowly during the autumn and winter of their first year of life and more rapidly from late winter to early summer. Thus, the period of most rapid accumulation of these reserves coincides with the time when body weight and gonad maturation are also increasing sharply. Lipid and glycogen levels fall during the reproductive season in those males that breed, so that by July they are reduced to 43% and 37% (respectively) of their peak values in May. Levels of protein increase throughout the year as the fish grow, but in breeding males by July the concentration of protein in the carcass falls to 70% of pre-breeding levels. Breeding males therefore reach the end of the reproductive season with their total energy reserves severely reduced, and consequently they suffer a very high mortality. In contrast, adult males that fail to reproduce survive beyond the breeding season. They continue to gain weight and to accumulate lipid and glycogen from August to September, but these energy reserves fall (to levels comparable to those of post-breeding fish) in December, when these fish probably die. These results demonstrate that in male sticklebacks, growth and gonad maturation can be sustained in parallel with the accumulation of energy reserves, which are subsequently extensively depleted as a result of reproductive activities.     

Empirical evidence of a trade-off between reproductive effort and expectation of future reproduction in female three-spined sticklebacks

Optimal life-history models generally predict that the reproductive effort of iteroparous organisms may increase with age, as their expectation of future reproduction decreases. The population of three-spined sticklebacks (Gasterosteus aculeatus) in the Camargue (Rhone River Delta, France) is annual, all adults dying after their first breeding season. As the three-spined stickleback is a multiple spawner, we tested the hypothesis that reproductive effort may increase during the breeding season on field data. From 1987 to 1998, 653 female sticklebacks were collected in the field during the breeding seasons. The body size, body weight and weights of the liver, gonads and carcass were measured for these individuals. Only gravid females with mature eggs (176 fish) were included in the analysis. Considering the female three-spined stickleback as a capital breeder, the energetic resources available for allocation between soma and gonads were estimated by its body weight. Somatic condition decreased during the breeding season and reproductive effort (gonad weight relative to body weight) increased. These patterns did not vary significantly between years. These observed variations in reproductive effort during the breeding season can be interpreted as empirical evidence of a trade-off between reproductive effort and expectation of future reproduction.     

Reproductive biology and female parental care in the cockscomb prickleback,Anoplarchus purpurescens (Pisces: Stichaeidae)

Synopsis Reproduction and parental care in the cockscomb prickleback, a Pacific coast intertidal fish, were examined using a combination of field and laboratory observations. The sexes were dimorphic, particularly during the breeding season, and males competed with other males for access to females. Males performed lateral and spasm displays. In the wild, the breeding season extended from January to March on cobblestone beaches. Assortative mating was positive with respect to body size. Females exhibited solitary parental care of the eggs. Each female coiled around, guarded and fanned a single egg mass that likely represented her total reproductive effort for the year. The number of eggs in the mass increased linearly with female size (weight or length). Males did not remain after spawning. Aquarium observations revealed that males spawn with more than one female given the opportunity. It is not known whether this occurs in the wild. Incubation to hatching took 29 days. Upon hatching, the young swam towards the surface. Parental care did not extend beyond hatching.     

The reproductive biology of an endemic cyprinid,Zacco pachycephalus,in Taiwan

Synopsis Reproductive parameters including body weight-length relationship, condition factor, gonadosomatic index (GSI) and fecundity of an endemic cyprinid,Zacco pachycephalus, were studied from October 1991 to January 1993. Although the condition factor was similar between both sexes, GSI values of female (3.72 to 14.24%) were significantly larger than that of males (0.21 to 2.23%) during the major breeding season. The number of yolky eggs per fish ranged from 459 to 2059. Ovum size distribution, annual GSI value, and recruitment of juvenile fish indicated thatZ. pachycephalus conducted two major spawning activities in spring (February to April) and summer (June to August). The larger GSI value and number of yolky eggs in spring compared to those in summer implies thatZ. pachycephalus has evolved a reproductive style by investing more energy in spring to avoid the damage of summer freshet on its recruitment success. The mating behavior ofZ. pachycephalus is similar to that of otherZacco fishes.     

Gonad development and size‐at‐maturity of silver therapon Leiopotherapon plumbeus (Kner 1864; Teleostei:Terapontidae) in tropical volcanic lakes in south Luzon,Philippines

Gonad development of the silver therapon Leiopotherapon plumbeus in two volcanic crater  lake  habitats (Sampaloc Lake, Taal Lake) in south Luzon, Philippines was examined during the annual reproductive cycle. The minimum body size‐at‐maturity of fish in these two lake habitats was also compared. Four gonad development stages were characterized as basis for the classification of ovarian (immature, maturing, mature, spawned) and testicular maturation (immature, maturing, mature) phases. The occurrence of all development stages in individual gonads suggest an asynchronous development whereby advanced stages are recruited continuously from a pool of younger stage germ cells to result in elevated female and male GSI throughout the annual cycle due to active gonadogenesis. Together with the increasing occurrence of advanced stage oocytes and spermatozoa from March until October, the elevated GSI of fish may indicate peak gonadal growth during the onset of the dry season (December–January) for eventual spawning from the beginning (May–June) until the end of the wet season (October–November). In both lake habitats, male fish were smaller than females but, regardless of sex, the minimum size‐at‐maturity of fish in Sampaloc Lake was significantly smaller than fish in Taal Lake. Overall, asynchronous development during oogenesis and spermatogenesis allows for year‐round reproduction of silver therapon, with elevated gonad growth in the dry season in preparation for spawning during the wet season. Compared with fish in Taal Lake, a smaller size‐at‐maturity of fish in Sampaloc Lake may be a response of the wild fishery stock to long‐term high fishing mortality and degradation of the lake habitat.     

Reproductive biology of Apocryptes bato (Gobiidae) in the Payra River,southern Bangladesh

The gobioid fish Apocryptes bato (Hamilton, 1822) is a species of mudskipper found in Bangladesh, India, and Myanmar. The reproductive biology of A. bato including sex ratio, size at sexual maturity, breeding season, fecundity, and condition factor was studied in the Payra River, southern Bangladesh during May 2016 to April 2017. A total of 1,028 specimens (303 males and 725 females) were collected. The overall sex ratio was significantly different from the expected value of 1:1 (male: female = 1:2.39, χ² = 173.23, p < 0.05). Size at sexual maturity was estimated as 4.9 cm standard length. Monthly variations in gonadosomatic index indicate that the main breeding season is from May to December. There was a significant positive correlation of body length and body weight with fecundity. Fulton's condition factor varied in both sexes and was attributed to variations in GSI with maturity. Breeding season was significantly associated with rainfall, but not air temperature. Our findings should improve management and conservation of this fish in the Payra River and surrounding ecosystems.     

Geographical variation in reproductive investment across avian assemblages in Europe: effects of environmental drivers differ between altricial and precocial species

Reproductive traits provide information about the ways by which available resources are allocated during breeding. We tested for environmental drivers of large scale geographical patterns in assemblage mean clutch size, number of broods and overall reproductive investment per breeding season in European birds. We combined data about geographical distribution with published information about reproductive traits, and calculated mean trait values for avian assemblages occurring in 50 × 50 km grid cells. In total, we employed data from 499 species and 2059 assemblages. As the time available for breeding and the amount of food limit the reproductive effort, we related the geographical variation in reproductive traits to the length of breeding season, normalized difference vegetation index (NDVI) as a surrogate of resource availability, and its seasonality. Geographical patterns in traits may differ between reproductive modes, thus we performed the analyses separately for altricial Passerines (n = 203) and precocial non‐passerine species (n = 164) and controlled for the effect of taxonomy. Large clutches dominated in areas with high NDVI and, in precocial birds, also in areas with high annual seasonality and a long breeding season. High number of broods and high overall reproductive investment dominated in areas with a long breeding season, and high number of broods was found also in areas with low annual seasonality, but only in precocial species. High overall reproductive investment dominated in highly productive areas and also in areas with low annual seasonality in both groups. The increase in reproductive investment is caused mostly by an increase in the number of broods related to the length of season and partly by increase in clutch size related to NDVI. We found a negative correlation between clutch size and the number of broods in Passerines, which might suggest a trade‐off between these traits. Processes behind trait patterns differ between altricial and precocial species.     

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.     

Gonadal development and spawning cycle in the digynic hermaphrodite sharpsnout seabream Diplodus puntazzo (Sparidae) off the Canary Islands, northwest of Africa

Diplodus puntazzo is a species characterized by a rudimentary hermaphroditism, with a low level of protandry. The study found that the overall sex ratio was one male to 2.03 females, diverging significantly from 1 : 1, and that there was a ratio of one male to 0.11 transitional fish. The sex ratio varied with season: during the reproductive period the ratio was 1 : 1.86 (male : female), significantly lower than in the non‐reproductive period in which the sex ratio was approximately 1 : 2.26. Both sexes had similar size distributions; however, females were predominant in all size intervals, especially in the larger ones. No significant difference in mean total length (TL) between females (330 mm), males (319 mm) or transitionals (321 mm) was found. The smallest functional female and male analysed were 159 and 157 mm TL, and the largest were 520 and 510 mm TL, respectively. No significant difference in length at first level of maturity, attained at the second year of life, was found between males (281 mm) and females (292 mm), respectively. A protracted spawning season was identified for the species: from September to February, with a peak in November. GSI values followed the same pattern for males and females, with values of females being similar to those of males; the highest values occurred between September and December, with a maximum in November. Occurrence of transitional fish was significantly higher during the month with highest values of GSI than in the resting period. The study found that vitellogenesis did not begin synchronously for all females, and the simultaneous presence of oocytes in all stages of development indicated an asynchronous mode of ovarian development. The presence of postovulatory, together with tertiary yolk vesicle oocytes, indicated that the species is a multiple spawner.     

Reproductive parameters in relation to food supply in the whiskered tern ( Chlidonias hybrida )

It is generally accepted that breeding terns are sensitive to food supply and that their reproductive effort could be substantially affected by the availability and access to resources. In this study we examined reproductive parameters in the whiskered tern Chlidonias hybrida in relation to food supply during the courtship feeding period (food brought by males to females) over a 2-year period (2004–2005). We also studied whether the condition of the nestlings [body condition index (BCI)] was related to a proxy of the reproductive investment of the adults (the clutch size) during a season of food shortage. Behavioural observations showed a decrease in the intensity of male courtship feeding between years (2004 > 2005), and a strong shift in the relative abundance of the two prey groups (invertebrates/fish; invertebrate prey dropped from 88.0 to 49.3%) brought by males. This change in food delivery rates did not result in a delay in laying, but there was a significant difference in mean clutch size between years (2.71 ± 0.49 eggs in 2004 and 2.05 ± 0.78 eggs in 2005) without any within-year variation in relation to the laying dates. The egg size (volume and length) was related to the year (2004 < 2005), suggesting a trade-off in the quantity and the quality of eggs between the two seasons. We also found no evidence that the investment in a large clutch affected nestling BCI in the course of the food shortage season. Since many pairs (about 60%) interrupted breeding during the incubation stage, we assumed that parents that succeeded in rearing nestlings in these conditions were probably ‘high-quality’ individuals. Our results therefore showed that whiskered terns are sensitive to the varying food conditions they experienced throughout the courtship period. The diversity of prey types could be a key factor in the reproductive investment of this tern species.     

Reproductive biology of Oreochromis niloticus (Perciformes: Cichlidae) at Emiliano Zapata dam, Morelos, Mexico

The reproductive season of tilapia was studied by monthly samplings at Emiliano Zapata dam, Morelos State, Mexico. From February 1999 through February 2000 a sample of 50 fish was taken from the commercial catch (castnet, 6.5 cm of mesh size). The observed sex ratio was 1:1.29 (females:males) (chi(2)=10.26; p<0.05). The tilapia reached maturity at 151.3 mm (females) and 152.0 mm (males) of total length. Rainy (August) and dry (February) seasons were determined as the breeding period. Fecundity variation was better correlated with length (r=0.7473: p<0.002) than with weight (r=0.7395; p<0.002). The fecundity ranged between 243 and 847 oocytes per fish, with egg diameter from 300 to 3 700 microm. Intensive breeding activity in August and February coincide with phytoplankton biomass increase.     

Size Dependent Male Reproductive Tactic in the Two-Spotted Goby (Gobiusculus flavescens)

Male investment in testes and sperm duct gland in the polygamous nest breeding two-spotted goby Gobiusculus flavescens (Fabricius) was investigated in relation to time in reproductive season and individual physical parameters. This small teleost fish is most likely the most abundant species found along the rocky shores of the North East Atlantic. The two-spotted goby has a single reproductive season, during which nest-caring males can raise several clutches of offspring. According to the literature the males are on average larger than the females. Here we report for the first time a population showing a reversal of this trend, with males on average being smaller than females, a difference likely caused by a large proportion of small males. Early in the breeding season these small males have typical sneaker characters, with relatively large testes and small seminal duct glands compared to the larger dominant territorial males. The presence of these two alternative male reproductive tactics is confirmed by histological studies, which shows the presence of sperm in the sperm duct glands (SDG) of smaller males, but not in the SDG of intermediate and larger males. To our knowledge, males with typical sneaker characters have not been reported in earlier studied populations of two-spotted goby. Interestingly we found that testes investment declined significantly over the course of the breeding season, and that this reduction was significantly more pronounced in small compared to the large males. Further, a significant increase in seminal duct gland (SDG) mass was observed for the smaller males over the breeding season. We propose that this indicates a possible shift in mating tactic by smaller males from a parasitic to a nest-holding tactic over the course of the breeding season. Thus, the observed size dependent plasticity in investment in SDG over time suggests that the reproductive tactic of G. flavescens is conditional, and possibly influenced by mate availability and male—male competition.     

Life history patterns of a livebearing fish in contrasting environments

Summary The population dynamics and energy allocations of the Gila topminnow, a small livebearing fish, were studied in two contrasting environments, a spring run of constant characteristics and a fluctuating desert wash. Topminnows grew and matured in two basic patterns. First, many fish in both areas matured the year after their birth. Second, spring fish born early in the breeding season grew rapidly, bred within five months, and died by eight months of age. Although spring fish assimilated more energy, wash fish actually expended more calories for growth and reproduction, partly because of lower maintenance costs. Reproductive effort of long-lived spring fish varied with age between 3.1 and 6.5%; whereas efforts of short-lived spring and wash fish increased steadily with age to 5.2 and 9.8%, respectively. Although spring fish produced eggs of higher energy content, females in both areas varied their investment per offspring, apparently tracking seasonal changes in the availability of food for fry. When long-lived spring fish experienced food shortage, they allocated less energy to both growth and reproduction; in contrast, wash and short-lived spring fish under similar conditions reduced only their growth allocation. The reproductive mass in spring fish appeared to be limited by food availability, incompletely filled the abdominal space, and reflected no tradeoff between fecundity and investment per offspring. Reproduction by wash fish appeared to be limited by body space and was characterized by a tradeoff between fecundity and egg size.     

Partitioning of Dry Matter and Mineral Nutrients During a Reproductive Cycle of the Mistletoe Amyema linophyllum (Fenzl.) Tieghem Parasitizing Casuarina obesa Miq.

Budgets for utilization of dry matter, nitrogen and a rangeof mineral elements were constructed for the reproductive season(September 1988–May 1989) of plants of the mistletoe Amyemalinophyllum (Loranthaceae) parasitizing the swamp she oak (Casuarinaobesa Miq.) at Gingin, W. Australia. Shed buds comprised 55%of the allotment of dry matter to reproduction, shed flowers7%, abscised perianths and styles 15%, prematurely shed fruits15%, and successfully ripened fruits a mere 12%. Commitmentto reproductive biomass peaked sharply in October, while thegenerally higher rate of allocation to vegetative biomass declinedfrom beginning to end of the 9-5 month reproductive cycle. Investmentsof dry matter in new leaves and secondary thickening of previousseason's stems were several-fold greater budget items than productionof new stems and shedding of leaves. There was no evidence ofpre-senescence retrieval of N, P, and K from leaves, and concentrationsof Ca, Mg, Na, and Cl increased steadily in dry matter withleaf age. Values for reproductive effort (RE) for dry matterand specific mineral elements were calculated as commitmentto reproduction divided by commitment to reproduction plus vegetativebiomass (? 100). Due to highly differential partitioning ofcertain elements between reproductive and vegetative parts,RE values for minerals for the reproductive season ranged widely,viz. 7% for Ca, 10% for Mn, 15% for Mg, 17% for N, 25% for Kand Zn, 26% for Cu, and 35% for P. The comparable RE for drymatter was 29%. The weight ratio for net intake of nutrientelements from the host by A. linophyllum for the reproductiveseason was 21?5:20?2:6?6:1?6:1 (N:K:Ca:Mg:P), while the meanweight ratio for the concentrations of these elements in xylem(tracheal) sap of the Casuarina host was 13?9:15?6:4?1:30:1,respectively. The data are discussed in relation to the demonstrationof a direct lumen-to-lumen tracheary continuity between hostand mistletoe in mature regions of the haustorial interface. Key words: Mistletoe, resource allocation, mineral nutrition, reproductive effort, host:parasite relationships     

Seasonal variations in plasma concentrations of sex steroid hormones and vitellogenin in wild male Japanese dace (Tribolodon hakonensis) collected from different sites of the Jinzu river basin

In order to clarify the seasonal variations of plasma sex steroid hormones and vitellogenin (VTG) concentrations in the wild male Japanese dace, Tribolodon hakonensis, we measured plasma levels of testosterone (T), 11-ketotestosterone (11-KT), estradiol-17 beta (E2) and VTG, as well as spermatogenetic stages and gonadosomatic index (GSI). Wild Japanese dace were collected from different sites of the Jinzu River basin (including the Takahara River and the Itachi River). The fish from Toyama Bay were also measured the spermatogenetic stages, GSI and VTG levels. The seasonal variations of the hormone levels were discussed in the relationship with various environmental factors. In landlocked fish of the Takahara River, the plasma concentrations of T and E2 reached the highest levels in May and June. In the fish collected from the Itachi River, plasma concentrations of T, 11-KT and E2 reached the highest levels during breeding season of April and May. Sexual maturation, evaluating from GSI and the spermatogenetic stages, proceeded earlier in the fish population at Toyama Bay, and afterward it was followed in the fish population at the Takahara River, in associated with a rise of environmental water temperature at fish captured sites. In the male dace, low but detectable levels of plasma E2 were measured and there were significantly positive correlations between E2 level and the levels of GSI, VTG or T. These results suggest that E2 might be a necessary sex steroid hormone related to gonad maturation, and that circulating E2 may induce VTG production in the wild male Japanese dace.     

Relative growth and reproductive cycle of the date mussel Lithophaga lithophaga (Linnaeus, 1758) sampled from the Bizerte Bay (Northern Tunisia)

Relative growth and the reproductive cycle of the date mussel Lithophaga lithophaga were studied from September 2002 to October 2003 in the Bizerte Bay (Northern Tunisia). Relationships between shell length and shell width, shell height as well as volume-related variables (shell dry weight, soft tissues dry weight and total wet weight) showed negative allometries in both sexes except for shell width in males (isometry) and shell height in both sexes (positive allometry). The sex ratio was unbalanced, within the length range of 6–48 mm, 49.43 % of the animals were males, 24.39 % females and 26.17 % sexually undifferentiated; within the length range of 49–92 mm, the respective values were 41.19, 52.7 and 6.1 %. Histological investigations as well as analyses of the condition index and the gonadosomatic index (CI and GSI) revealed the presence of a single reproductive cycle per year. Spawning occurred at the end of August and early September and was associated with a decrease in seawater temperature and salinity. A resting phase occurred in winter, coinciding with the lowest water temperatures. Histological examinations of the gonads of a total of 130 specimens revealed only two cases of hermaphroditism. The present study constitutes a useful baseline for a sustainable management of local wild stocks of L. lithophaga.     

Experimental manipulation of tail length in female barn swallows (Hirundo rustica) affects their future reproductive success

Previous studies have shown no significant effect of experimentaltail length manipulation in female barn swallows (Hirundo rustica)at the beginning of a breeding season on reproductive successor behavior during that breeding season. In the present study,we investigate if tail length manipulation had any effect onreproductive performance the following year, the so-called long-termeffect, in contrast to the short-term effects already studied.We found that females with experimentally elongated externaltail feathers at the beginning of a breeding season producedless offspring during the breeding season the following yearthan did females with shortened or unmanipulated tails. Theseresults suggest that tail elongation caused flight deficienciesthat deteriorated the condition of females and eventually reducedreproductive success. The finding of long-term effects but nosignificant short-term effects for female tail elongation suggeststhat female barn swallows have the ability to adjust immediateparental investment. Detrimental effects of long tails in femalesin terms of decreased reproductive success might explain whyfemale tails are not as long as those of males. Finally, femalesmated to long-tailed (sexually attractive) males decreased theirreproductive success the following year more than did femalesmated to short-tailed males, possibly owing to differentialparental effort causing a deterioration of their condition.     

Quantitative studies on the reproduction of black scallop,Chlamys varia (L.) from lanveoc area (Bay of Brest)

The values of gonad index calculated by the allometric relationships of dry gonad weight over dry body weight of the black scallop, Chlamys varia (L.), gave peak values for May and September during 1976 and 1977. Histological studies on gonads from natural populations and periodic biopsies of gonads from marked animals in experimental cages showed that, in addition to intermittent spawnings between May and September, two peak spawning periods were noteworthy. These were a partial spawning in May–June and a complete spawning in September–October. The energy spent as gametes for these two spawnings has been calculated for individual males and females in each age group of the population and the cost of spawning by the whole population has been estimated. The population spent more energy for spawning in 1977 (144.49 kcal/100 m²) than in 1976 (83.70 kcal/100 m²) and the reproductive effort of an average adult was 0.04 in 1976 and 0.10 in 1977.