Ovarian allometry and the use of the gonosomatic index: a case study in the Mediterranean sardine,<Emphasis Type="Italic"> Sardina pilchardus</Emphasis>

The gonosomatic or gonadosomatic index [GSI=(ovary weight/fish weight)×100] has been widely used in fisheries science and experimental reproductive studies as a simple, low-cost measure of reproductive condition. However, its properties have not been fully evaluated, and several pitfalls, such as size-dependence and changes in ovarian allometry, may invalidate its use. In the present study, we examined ovarian allometry and the appropriateness of GSI for assessing ovarian activity in the Mediterranean sardine Sardina pilchardus. The analysis was based on a large sample of histologically scored females collected over an annual cycle in the eastern Mediterranean (Aegean and Ionian Seas). First, we examined GSI dependence on fish size by comparing ovary weight–on–fish weight relationships in different stages of oocyte development. The effects of recent spawning (incidence of postovulatory follicles) and intensity of follicular atresia were also addressed. In a subsequent step of the analysis, we applied generalized linear models (GLMs) to analyze the relationship between GSI and histological stage, taking into account the parallel effects of additional factors related to geographical region, month of capture, fish length and energetic reserves (fat stage, hepatosomatic index—HSI). Ovarian growth was isometric in all stages of oocyte development and states of atresia, but altered to positive allometric at the stage of oocyte hydration. Oocyte growth and intensity of atresia significantly affected GSI. Fish length, geographical region, month of capture, fat stage and HSI did not substantially affect GSI, further strengthening the appropriateness of the index for the Mediterranean sardine. Finally, we provide first evidence, based mainly on available (but yet limited) published information for other fish species, that: (1) the pattern of ovarian allometry may not be altered by fecundity variations and (2) the pattern of ovarian allometry may reflect the pattern of oocyte growth (i.e. isometric vs. allometric ovarian growth could reflect group-synchronous vs. -asynchronous oocyte development). Alternatively, the pattern of ovarian allometry could reflect the presence or absence of size dependency in the relative fecundity (eggs per gram of body weight) and/or egg size. The isometric ovarian growth in the Mediterranean sardine could be attributed to the lack of such size-dependent effects.Communicated by O. Kinne, Oldendorf/Luhe     

Ephemeral spawning aggregations in the Mediterranean sardine, <Emphasis Type="Italic">Sardina pilchardus</Emphasis>: a comparison with other multiple-spawning clupeoids

Using previously published histological data on multiple, monthly samples of Sardina pilchardus collected in the central Aegean and Ionian Seas (September 1999–August 2000, and November 2000–February 2001), the Mediterranean sardine was treated as a case study to investigate the biological characteristics of ephemeral spawning aggregations in multiple-spawning clupeoids. Actively spawning (Day₀) females in the Mediterranean sardine, i.e., the daily class of spawners caught a few hours prior, during, or after the spawning act, were shown to separate spatially from late (Day₁₊) spawners and non-spawning females, taking with them a large proportion of conspecific males which were also in advanced spawning condition and in better somatic condition compared to the remaining population. In addition, information from 28 stocks of multiple-spawning clupeoids from a wide range of geographic locations, belonging to 14 species and 2 families (Engraulidae and Clupeidae), was reviewed and analyzed pertinent to the formation of ephemeral spawning aggregations. Results from the latter analysis indicated similar patterns of spatial segregation of Day₀ spawners in the reviewed clupeoid stocks as in the Mediterranean Sardine, which strongly suggested that the formation of ephemeral spawning aggregations is a common behavioral trait among multiple-spawning clupeoids. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Evaluation of spawning frequency in a Mediterranean sardine population (<Emphasis Type="Italic">Sardina pilchardus sardina</Emphasis>)

The postovulatory follicle method was used to assess the frequency of spawning of sardine (Sardina pilchardus sardina) in the Central Aegean Sea (eastern Mediterranean) in November–December 1999 and 2000. A field-based aging key for postovulatory follicles (POFs) was developed, and preliminary evidence is presented of a temperature influence on the degeneration of POFs. Bias in the fraction of day-0 spawners (females that had, were, or would spawn the night of sampling) with respect to sampling time was identified and attributed to spawning behavior. An appraisal of consistency in spawners' fractions and sex ratio, based on four sets of sequential trawl hauls, showed a considerably high small-scale variability, which suggested that spawning schools are structures limited in space and/or ephemeral in time. Sampling gear, i.e. commercial purse seines and research pelagic trawls, did not differ significantly with respect to spawners' fractions. The fractions of day-1 and day-2 females were the same during both years, and their combination resulted in more precise, composite estimates of spawning frequency. In both years, the Mediterranean sardine spawned every 11–12 days. The incidence of spawning increased significantly with fish size. When compared to all estimates available for other sardine genera, species and subspecies, the spawning frequency of the Mediterranean sardine was lower; this was attributed to the synergetic effect of oligotrophy and smaller body sizes in the Mediterranean.     

Length-structured analysis of the reproductive season of anchovy and common sardine off central southern Chile

The anchovy (Engraulis ringens) and common sardine (Strangomera bentincki) are two small pelagic fish which have a similar reproductive strategy off central southern Chile. The seasonal reproductive dynamics of both the species was investigated by taking into account the gonadosomatic index (GSI) and visual maturity data as a function of female size for the period 1993–1999, and also the seasonal pattern in condition factor per size classes. Larger females of common sardine (repeat spawners) have a reproductive peak earlier in the season than younger females at first maturity, while larger anchovy have a reproductive peak delayed as compared with first-time spawners. The condition factor of females exhibited an inverse cycle with the seasonal pattern of GSI, and delayed by approximately 6 months. Although larger females tend to present better and wider condition than smaller females, the condition of females seems to be delayed in larger females probably mediated by the seasonal pattern in food availability. This “energy storage strategy” in spring and summer time seems to be size-dependent and past energetic reserves could also affect the egg production and timing of reproduction. We concluded that the reproductive season of anchovy and common sardine is different and mediated by the length structure in the seasonal upwelling system off central southern Chile.     

Ovarian development and spawning in relation to the moult cycle in Northern krill, Meganyctiphanes norvegica (Crustacea: Euphausiacea), along a climatic gradient

Adaptive processes linked to reproduction were studied comparatively for three populations of Northern krill, Meganyctiphanes norvegica (M. Sars, 1857), sampled during winter and summer cruises in the Clyde Sea (W Scotland), the Kattegat (E Denmark), and the Ligurian Sea (NW Mediterranean). The aim was to investigate the functional relationship between egg production and moulting under contrasted climatic and environmental conditions. A staging system for female sexual development established for live krill was complemented by a histological study of the ovary at various developmental steps. During the reproductive season, all adult female krill were engaged in cyclical egg production. During experiments, female krill released one batch of mature oocytes in one or two spawning events. The ovary of postspawn female krill still contained developing oocytes for another egg batch. In the non-reproductive period, all female krill had a resting ovary. Ovarian structure and pattern of egg production were identical in the three populations, but seasonal timing of egg production was different. The model proposed for the Ligurian population of the annual cycle of ovarian development can be extended to the other two populations, taking into account the seasonal characteristics of each site. Random field samples were staged simultaneously for moult cycle and for sexual development. Moult stages and the seasonal variation of the intermoult period were studied for the Kattegat population using multi-year data and compared to data obtained during summer/winter cruises in the Clyde and the Ligurian Sea. At the three sites, intermoult period was shorter and temperature-dependent during the reproductive period, concurrent with the season of greatest food availability. During most of the year and the period of sexual rest, moulting activity was reduced. The relationship between spawning and the moult cycle was studied comparatively for the three populations. Eggs were released during the premoult phase of a “spawning moult cycle”, in one or two spawnings associated with apolysis and Moult Stage D1, respectively. Yolk accumulation for the next egg batch was completed during an alternating “vitellogenic moult cycle”. A model for the timing of cyclical egg production in relation to moulting, as proposed for the Kattegat, can be extended to the other populations, taking into account intermoult period variation with temperature. Temperature appeared to be the principal environmental factor controlling growth (through moulting) and egg production during the reproductive season, in connection with favourable trophic conditions. Received: 22 December 1997 / Accepted: 29 August 1998     

Sardine (Sardina pilchardus) spawning seasonality in European waters of the northeast Atlantic

Egg data from ichthyoplankton monitoring sites in the western English Channel (1988–2003) and northern Spain (1990–2000) and macroscopic maturity data from biological samples of purse seine landings in western and southern Iberia (1980–2004) are used to describe the spawning seasonality of sardine (Sardina pilchardus) in European waters of the northeast Atlantic using generalised additive models. The fitted models reveal a double peak in spawning activity during early summer and autumn in the western Channel, a wider spring peak off northern Spain and a broad winter season in the western and southern Iberian Peninsula. At all sites, a high probability of spawning activity was observed over at least 3 months of the year, with the duration of the season increasing with both decreasing latitude and increasing fish size. Off western and southern Iberia there are indications that the spawning season has been of longer duration in recent years for all size classes (reaching in some cases 8 months of the year for large fish). These patterns are in general agreement with existing literature and theoretical expectations of sardine spawning being driven locally by the seasonal cycle of water temperature, assuming preferences for spawning at 14 –15°C and avoidance for temperatures below 12°C and above 16°C. Regional quotient plots indicated that spawning tolerance to higher temperatures increases progressively with decreasing latitude. Despite the weak evidence for geographical differences in temperature tolerance that may have some genetic origin, the degree of spatio-temporal overlap in sardine-spawning activity within Atlantic European waters is unlikely to promote any reproductive isolation in that area.     

Parental care and the cost of reproduction in a Mediterranean fish

Summary Symphodus tinca is a common near-shore Mediterranean labrid fish in which females may sometimes spawn their eggs over hundreds of square meters, or alternatively spawn into well-defined algal nests. Eggs spawned in either manner are fertilized, but widely scattered eggs receive no parental care, whereas eggs spawned into nests are usually guarded by the male until they hatch. Here, I report weight changes of individual marked fish that engaged in a variety of different reproductive behaviors during three breeding seasons. Males gained weight at 0.15% per day outside the spawning season, and added 29–78% to their overall body weight between reproductive seasons, even following substantive weight losses in a spawning season (up to 20% among nesting males). Nesting and nest-guarding males lost an average of 0.32% and 0.41% of their body weight per day in 1986 and 1987. This cost is four times greater than reproduction for nonnesting males, which registered a 0.03% daily weight gain. Actively spawning females lost 0.06% of their body weight daily during the spawning season. While long-term growth rates did not appear to be substantially affected by reproduction in either sex or by parental care in males, present work does not exclude the possibility that such long-term effects may exist.     

Analysis of the relationship between gonads and hepatopancreas in males and females of the crab Sesarma intermedia,with reference to resource use and reproduction

At the beginning of the reproductive period in May, there was an increment in both the gonad and heaptopancreas. During the peak phase in June, with high ovary production, there was a decrease in the hepatosomatic index (HSI), particularly for the Ariake females. The minimum gonadosomatic index (GSI) of 2% in August and September for the Ariake and in September and October for the Hakata populations was inversely related to the increase of HSI. A second rise in GSI was observed in November at the 1% level as against the June peak of 5% for the Ariake and 3.5% for the Hakata females, and it remained at this level until December, while the HSI decreased to the minimum. The December minima for both GSI and HSI were about the same as the initial February values. The spermathesomatic index (SSI) had a slight increase to 0.8% in August, but remained constant at 0.5% throughout the year. Likewise, GSI for males was constant at the 0.5% level, with minor fluctuations, and did not show any clear relationship with the HSI as for female GSI. Females had higher hepatosomatic indices but the soma weighed less than males throughout the year. Again in May, medium-sized females had the highest GSI followed by large and then small-sized individuals. As the HSI steadily decreased, GSI increased with advancing egg development stages for ovigerous females. The quantitative stomach analyses showed that females had higher stomach quantity indices (SQI) in May than in September, when the breeding period had ended. Contrary results were observed for males. Also, female SQI in May was higher than that for males.     

Reproduction and condition of female Octopus mimus (Mollusca: Cephalopoda)

The relationship between reproduction and condition was studied in a 15 mo sample of 919 maturing, mature and post-spawning female Octopus mimus (388 to 3714 g) caught in Iquique (North Chile). O. mimus is a semelparous species, with reproduction taking place all year round. Investment in reproductive tissues was, on average, 9.9% of mature female body weight, independent of season. However, somatic growth during maturation was dependent on season and varied between 26 and 63% of the whole-body growth in weight. The condition of females did not vary markedly until spawning, although seasonal variations were apparent, winter being the most unfavourable. Condition deteriorated dramatically after spawning, during parental care of the eggs. During this period, somatic-tissue depletion, mainly from the muscles, was >25% of the total body weight of mature females. The fecundity of O. mimus was probably limited by the costs associated with parental care of the eggs.     

Size at maturity,fecundity and reproductive potential of a protected population of the spiny lobster<Emphasis Type="Italic"> Palinurus elephas</Emphasis> (Fabricius, 1787) from the western Mediterranean

Protected lobster populations are expected to contribute to the replenishment of fished populations through increased egg production. We studied the reproductive biology and egg production potential of a population of the spiny lobster Palinurus elephas protected from fishing since 1990 in the Columbretes Islands Marine Reserve (western Mediterranean). An index of spawning potential was derived to compare egg production potential in the Reserve and in western Mediterranean exploited populations. Females' physiological maturity (ability to reproduce) and functional maturity (ability to mate and bear eggs) occurred at a carapace length (CL) of 76–77 mm. Males' physiological maturity occurred at a slightly larger size, 82.5 mm CL. In the Reserve, P. elephas' individual fecundity increases linearly with body size up to the females' maximum size, although maximum reproductive yield (eggs per body gram) was reached at intermediate sizes. Size-specific fecundity in the protected population was similar to that of lightly fished populations off Ireland and greater than that of western Mediterranean exploited populations. The female size class of 105–110 mm CL contributed most to egg production in the protected population and is well above the minimum landing size (MLS) for western Mediterranean fisheries. Newly mature females (below MLS) generate a very small fraction (1%) of the egg production from the Reserve. Given the pattern of exploitation in western Mediterranean fisheries, egg production potential depends more on the quantity than on the mean size and fecundity of the available females. The role of the greater availability of large males for mating in unfished populations is discussed in terms of the females' individual fecundity and mating success.     

Habitat influences on reproductive allocation and growth of the mummichog (<Emphasis Type="Italic">Fundulus heteroclitus</Emphasis>) in a coastal salt marsh

To determine whether life history differences can occur in salt marsh fishes that occupy different habitats within the same marsh, we compared reproductive allocation in female mummichog (Fundulus heteroclitus) inhabiting creeks and ponds of a coastal salt marsh in southern New Jersey, USA during the spring and summer of 2001 and 2002. Females were collected in phase with the lunar spawning cycle from four sites of each habitat type, and assessed for gonad-to-body-mass ratio and growth increment. Annual reproductive allocation, expressed as a percent of somatic mass, was estimated for each site and year from the gonadosomatic indices of individuals collected during each spawning period. Mummichogs from creeks showed little change in annual reproductive allocation from 2001 to 2002, whereas those inhabiting ponds showed a significant increase between these years. Seasonal reproductive patterns indicated that pond females cease spawning at least one lunar cycle earlier than creek fish. While ponds experienced considerably higher maximum summer temperatures than creeks as well as near-anoxic pre-dawn conditions, neither of these variables explained a significant amount of variation in annual reproductive allocation. In contrast, annual reproductive allocation of mummichogs in a pond correlated with its flood frequency in both years of study. Our results suggest that while the length of the spawning season differs in mummichogs inhabiting marsh creeks and ponds, annual reproductive allocation depends more upon the hydrodynamic conditions of the particular waterbody than its habitat type per se.     

Long-term experiments on lifespan, reproductive activity and timing of reproduction in the Arctic copepod Calanus hyperboreus

Several experiments were conducted with starved and fed females of the Arctic copepod Calanus hyperboreus (1) to investigate their lifespan, reproductive period, egg production and egg viability; (2) to study the effect of origin, i.e., Atlantic and Arctic Waters in the Greenland Sea, on the timing of reproduction; and (3) to study the effect of time of collection on the onset of reproductive activity as a first approach to study control mechanisms of the reproductive cycle. Females collected in October produced up to 1,000 eggs and had a maximum lifespan of 164 days without feeding, whereas fed females produced up to ca. 6,000 eggs and survived up to 806 days. These observations support earlier assumptions that females were multiannual-iteroparous, i.e., capable to spawn in successive years, which would be unique for calanoid copepods. In starved females, clutch size decreased significantly with each spawning event. Viable eggs were produced during most of the life time. There was no difference in the timing of reproductive activity between females from the West Spitsbergen Current and the Greenland Sea Gyre. Fed and starved females collected in May and June began to spawn circa 2 and 4 months after collection, respectively, whereas females collected in August and October started spawning at the same time, in the middle of October. This indicates initiation of reproductive activity in the field in August, coincident with the descent into deep waters. Potential cues for the untimely spawning of females collected in spring and ‘unnatural’ feeding in fall experiments are discussed. Their large size, robustness and combination of different types of diapause in their life cycle make C. hyperboreus a good model organism to study diapause control mechanisms.     

Temporal shifts in the allocation of energy in the arrow squid,<Emphasis Type="Italic"> Nototodarus gouldi</Emphasis>: sex-specific responses

Squid typically display considerable intra-specific plasticity in size and age-at-maturity in response to ambient environmental conditions, yet little is known of the mechanisms driving these variations. We examined the intra-specific variability in Nototodarus gouldi reproductive traits to determine patterns of energy allocation between somatic and reproductive processes over short temporal scales. Females caught during the cool months of May and July were larger, had slower lifetime growth, lower gonad investment, and better somatic condition than females caught during the warmer months, suggesting a trade-off between gonad investment and somatic condition in females. On the other hand, males showed a tight coupling between somatic condition and gonad investment for most months, with increases in somatic and gonad tissue occurring concurrently. In male squid, an increase in lifetime growth rate was coupled with an increase in the relative weights of somatic and reproductive structures, whereas in females, percent increase in body weight per day was correlated only with gonad development. Patterns of repro-somatic investment in mature females had implications for spawning strategies, since female squid with higher levels of gonad investment apparently released batches of eggs together as a group, regardless of body size, whereas females with low gonad investment possibly spawned their eggs independently of one another. In terms of life-history theory, male squid were able to respond rapidly to environmental fluctuations without compromising either the gonad or the soma. However, although mature females did not appear to respond as quickly to ambient conditions, female squid possibly produced two different reproductive strategies, possibly to maximise offspring survival in either a stable or a variable environment. It seems from our study that monthly variations in ambient conditions may have large effects on life-history strategies.Communicated by M.S. Johnson, Crawley     

Spatio-temporal variability in fatty acid trophic biomarkers in stomach contents and muscle of Iberian sardine (<Emphasis Type="Italic">Sardina pilchardus</Emphasis>) and its relationship with spawning

Temporal variation in the fatty acid (FA) composition of stomach contents of Iberian sardines was compared to the relative contribution to dietary carbon made by different prey types for fish from two areas off Portugal. The effect of the FA content of the diet on sardine muscle FA composition was also studied, aiming at (1) analysing if FA biomarkers can be used as a complementary technique for the study of sardine diet and (2) to relate spatial and temporal variations of prey FA content with sardine condition and reproduction. Significant spatial differences in the FA composition of sardine diet occurred with concentrations of n-3 polyunsaturated FA, namely eicosapentaenoic acid [EPA, 20:5n-3] and linolenic acid 18:3n-3, being significantly higher in the diet of sardines from the west coast, whilst the diet of sardines from the south coast was richer in monounsaturated fatty acids (MUFA), namely the carnivory biomarker oleic acid 18:1n-9. These results are in agreement with the higher contribution made by diatoms and dinoflagellates to the diet of sardines off the west coast. Spatial variation in sardine dietary FA was also detected in their muscle composition, specifically for EPA, and the eicosapentaenoic/docosahexaenoic acid and (n-3)/(n-6) ratios, which were higher in sardines from the west coast. No difference in FA composition was detected between sexes, and the seasonal variability in sardine total FA concentration was primarily related to the seasonality of spawning. Sardines accumulate high concentrations of FAs during the resting stage of reproduction when the feeding intensity is similar or lower to that observed during the spawning season. Additionally, sardines show a high selective retention of MUFA and polyunsaturated FA (PUFA) throughout the year except at the beginning of the spawning season, when these FAs are largely invested in the formation of the gonads. Therefore, temporal and regional differences of prey environments are strong enough to be reflected in fish body composition, namely on the accumulation of essential FAs, which can have a strong impact on reproduction success for this species.     

Reproductive status and body condition of Atlantic bluefin tuna in the Gulf of Maine, 2000–2002

The reproductive status and body condition of 195 (≥185 cm curved fork length, CFL; assigned age 7 and above) Atlantic bluefin tuna were assessed in the Gulf of Maine during the commercial fishing season of June–October, 2000–2002. Given the distance between known spawning and feeding grounds, the prevailing paradigm for Atlantic bluefin tuna (Thunnus thynnus thynnus, L.) suggests that the most likely histological state for females arriving in the Gulf of Maine after spawning would be a resting or quiescent state with little or no perigonadal fat. Alternatively, the presence of mature or mature-inactive histological states in some females supports a more varied or individualistic model for bluefin reproduction. No relationship was found between body condition and reproductive status. Males were found in all reproductive stages, but were more likely to be in spawning condition (stages 4 and 5) or a mature-inactive state (stage 6) in June and July. Female bluefin tuna were found in stage 1 (immature or non-spawning) and stage 6 (mature-inactive). Stage 6 females were only present in June and July and smaller females (<235 cm CFL) were more likely to be in stage 6 than large females (>235 cm CFL) sampled during those same months. The presence of smaller females in stage 6 arriving at the same time as larger females in stage 1 indicates that Western Atlantic bluefin tuna may have an asynchronous reproductive schedule and may mature at a smaller size than the currently accepted paradigm suggests.     

Life-history traits of Plesionika martia (Decapoda: Caridea) from the eastern-central Mediterranean Sea

Life-history traits of Plesionika martia (Milne Edwards, 1883) were studied through data collected during six seasonal trawl surveys carried out in the Ionian Sea (eastern-central Mediterranean) between July 1997 and September 1998. P. martia was found at between 304 and 676 m depth, with the highest density in the 400-600 m range. Intraspecific, size-related depth segregation was shown. Recruitment occurred in summer at the shallowest depths. Juveniles moved to the deepest grounds as they grew. The largest female and male were 26 and 25 mm carapace length, respectively. The sex ratio was slightly in favour of females at depths >400 m. Although a seasonal spawning peak was shown, the reproduction appears to be rather prolonged throughout the year. Females with ripe gonads were found from spring to autumn. Ovigerous females with eggs in late maturity stage were found year round. Large females could spawn more than one time within their annual reproductive cycle. The size at first maturity (50% of the ovigerous females) was 15.5 mm CL. Average brood size of eggs with a well-developed embryo was 2,966ǃ,521. Iteroparity, low fecundity and large egg size patterns were observed. Brood size increased according to the carapace length. Two main annual groups were found in the field population of the Ionian Sea. Estimates of the Von Bertalanffy growth parameters are: L_X=30.5 mm, k=0.44 year^-1 in females; L_X=28.0 mm, k=0.50 year^-1 in males. A negative allometry was detected mostly in the ovigerous females. The life cycle of P. martia is discussed in the light of life-history adaptations shown in other deep-water shrimp species.     

Reproductive cycle of Zidona dufresnei (Caenogastropoda: Volutidae) from the southwestern Atlantic Ocean

Gonadal development of Zidona dufresnei (Donovan, 1823) (Caenogastropoda: Volutidae) was studied over a period of two consecutive years, through analysis of gonadal tissue. Individuals were sampled monthly at the Mar del Plata area, Argentina. Gonadosomatic index was estimated for males and oocyte size was used to estimate the stage of gonadal development in females. The reproductive season in this species in the sampled locality extended from October to March (austral spring-summer). During summer, a stage of advanced gonadal development and spawning predominated in the adult population. In autumn, gonads were generally under atresia; during wintertime (June-August), they underwent a period of recovery that lasted until the spring months, when gametes were released again. Synchronism between both sexes was evident. Marked periods of spawning were followed by resorption periods and then a growing phase; it was very clear that reproductive seasonality was linked to changes in bottom water temperature. These results suggest that Z. dufresnei gonads have a yearly cycle of gamete production, with two major activity peaks in September-October and January-February.     

Pair formation,pre-spawning waiting,and protandry in kokanee,<Emphasis Type="Italic"> Oncorhynchus nerka</Emphasis>

The timing of arrival to breeding areas can have profound effects on reproductive success. Under some conditions (restricted breeding seasons and mating systems characterized by a longer period of mating among males than females), the maximization of mating opportunities by males theoretically selects for the earlier arrival of males than females (a phenomenon called protandry). This study quantifies the relationship between the arrival timing and spawning success of male kokanee (non-anadromous Oncorhynchus nerka). The spawning behavior of kokanee was observed in a large pen and the spawning success of each male was estimated as the number of spawning events he participated in. A male's spawning success depended primarily on his success at pairing with and mate-guarding females, and less on participation in spawning events while unpaired. Males who paired earlier in the season had higher spawning success than males who paired later in the season because they experienced more opportunities to pair with new females. Among males who eventually paired (some males never did), arriving early was correlated with pairing early. However, selection for protandry was weak, largely because early arrival did not guarantee that a male would pair. Pre-spawning waiting by females also weakened the correlation between arrival day and pairing day. The random probability of pairing with respect to arrival day and pre-spawning waiting by females likely explains the weak selection for protandry in kokanee and the low amount of protandry observed in other sockeye salmon (anadromous O. nerka) populations.     

Spawning pattern and type of fecundity in relation to ovarian allometry in the round herring <Emphasis Type="Italic">Etrumeus teres</Emphasis>

Spawning pattern (assessed by seasonal changes in ovarian developmental stages) and type of fecundity (assessed by analysis of oocyte-size frequency distributions) of the round herring Etrumeus teres were studied in relation to ovarian growth and seasonal changes in the gonadosomatic (GSI), hepatosomatic (HSI) and liposomatic (LSI) index as well as the somatic condition of spawners (CS) in a spawning ground of southern Japan. Except for summer, mature and recently spawned ovaries occurred all year round. Oogonia and primary oocytes were present in all ovaries, and cortical alveoli stage (CA) oocytes occurred in all mature, hydrated and partially spent (PS) females (PS: females containing post-ovulatory follicles). Before hydration, a clutch of larger yolked oocytes, undergoing synchronous growth (range 0.7–1.1 mm), was present in mature ovaries which was completely separated from a more heterogeneous clutch of oogonia, primary and secondary oocytes (<0.150 mm) and oocytes in the CA stage (range 0.15–0.60 mm). As vitellogenesis progressed, the yolked clutch increased in size but the CA oocytes remained arrested. The latter entered into the secondary growth phase when hydration started in the advanced batch. Ovarian growth was isometric in all developmental stages, validating the use of GSI, which showed a consistent monthly evolution among years. Spawning stopped in summer (July and August) and peaked in winter and spring. HSI correlated positively with GSI on both a monthly mean basis (r = 0.76) and individual fish basis (liver weight explained 67–83% of the variability in ovary weight when females were grouped into 1-unit GSI intervals) suggesting a significant role of liver in vitellogenesis. LSI and CS also showed marked seasonal changes peaking from summer to middle autumn. Overall results suggest that E. teres is a multiple spawner with a group-synchronous ovarian development and indeterminate annual fecundity, with the three processes linked to an isometric growth of the ovary. We propose that such a reproductive pattern is an adaptation to produce batches of large pelagic eggs through a protracted spawning season.     

Energy storage and dynamics in bay anchovy Anchoa mitchilli

The energy of the body components and the energy costs of spawning and overwintering in the bay anchovy Anchoa mitchilli, the most abundant fish in Chesapeake Bay, were studied to determine seasonal variability during the different stages of its life cycle. Bimonthly samples were collected by trawl from April 1990 through October 1991. Fish condition and body energy levels fluctuated seasonally, and were related to anchovy size. Energy equivalents (cal g^-1 dry wt) was highest in December, before the overwintering period. The somatic weight component increased by 32 to 33% and total body weight by 26% during the spawning season, indicating that feeding not only met energy requirements of daily spawning but also provided surplus energy for growth. The overwintering loss of energy was 33 to 35% of total body calories, and was primarily derived from deposit fat in somatic and visceral tissues.