Reproduction related immunoglobulin changes in rainbow trout

Annual changes in plasma immunoglobulin (IgM) levels were investigated in three strains of rainbow trout Oncorhynchus mykiss which have different spawning periods, i.e., September–October, November–December, and January, reared under constant water temperature and natural day length. Plasma IgM levels decreased during the spawning season in all strains tested. The IgM changes became reversed in response to significant increases in plasma testosterone (T) and estradiol-17 in females and T and 11-ketotestosterone in males. Though the IgM decline showed a connection with suppressed immunocompetence, since many mature fish caught fungal diseases, no clear differences were observed in the plasma IgM levels between infected and noninfected fish during the spawning season. Incidentally, plasma IgM levels in infection prone fish were higher than in noninfection prone fish prior to the spawning season, whereas coincident differences in the plasma steroid levels were observed. Immature fish reared under lower water temperatures showed lower IgM levels. The effect of water temperature may have to be considered when analyzing the defense mechanism during the spawning season in rainbow trout.     

Frequency and distribution patterns of Saprolegnia infection in wild and hatchery-reared brown trout Salmo trutta L. and char Salvelinus alpinus (L.)

Abstract. Fungal infection of sexually mature brown trout and char was associated with a particular type of Saprolegnia exhibiting a low degree of homothallic sexuality. Hatchery-reared brown trout were more severely infected (in terms of the % area of the body covered with the fungus) than were wild fish. The fins of hatchery-reared fish were particularly prone to Saprolegnia infection regardless of sex. In wild brown trout, a sexual difference in the pattern of infection was demonstrated. The flanks of the male fish appeared to be more prone to infection when compared with the female and there was a greater susceptibility of the caudal and ventral fin of the female when compared with the male. Evidence is presented which suggests that the incidence of infection in mature male salmonid fish prior to spawning is significantly greater than in the females. This difference may not be apparent in spent fish after spawning. These findings are discussed in relation to the background concentration of fungal spores in the water, the behavioural characteristics of spawning fish and differences in epidermal structure.     

Lipid transport in female Fundulus heteroclitus during the reproductive season

Lipid transport during the reproductive cycle of seasonally spawning teleosts has been well described. However, there is little information on lipid transport in a multiple spawning teleost. We examined lipid transport during the reproductive cycle in Fundulus heteroclitus, a teleost that spawns in a semilunar fashion. Plasma levels of triglycerides more than double prior to spawning events and then drop off sharply around the time of spawning. Plasma levels of total cholesterol also changed throughout the spawning cycle, but those changes were not so pronounced. The fraction of lipid carried by very low density lipoprotein (VLDL) and low density lipoprotein (LDL) was greatest prior to spawning events, while high density lipoprotein (HDL) carried the greatest portion of lipid during the spawning period. Vitellogenin (Vtg) was detected prior to spawning events, but it never represented more than 5% of total plasma lipid, making it very unlikely that Vtg is responsible for the changes in plasma lipids that we observed. It is also unlikely that Vtg is solely responsible for transport of lipid to growing ovaries because it carries only a very small portion of plasma lipid. This is the first study we are aware of, that tracks changes in plasma lipids of a multiple spawning teleost with a great enough sampling frequency to resolve single spawning events. This revised version was published online in August 2006 with corrections to the Cover Date.     

Relationships between water temperatures and upstream migration, cold water refuge use, and spawning of adult bull trout from the Lostine River, Oregon, USA

Howell PJ, Dunham JB, Sankovich PM. Relationships between water temperatures and upstream migration, cold water refuge use, and spawning of adult bull trout from the Lostine River, Oregon, USA.
Ecology of Freshwater Fish 2010: 19: 96–106. This article is a US Government work and is in the public domain in the USA Abstract – Understanding thermal habitat use by migratory fish has been limited by difficulties in matching fish locations with water temperatures. To describe spatial and temporal patterns of thermal habitat use by migratory adult bull trout, Salvelinus confluentus, that spawn in the Lostine River, Oregon, we employed a combination of archival temperature tags, radio tags, and thermographs. We also compared temperatures of the tagged fish to ambient water temperatures to determine if the fish were using thermal refuges. The timing and temperatures at which fish moved upstream from overwintering areas to spawning locations varied considerably among individuals. The annual maximum 7‐day average daily maximum (7DADM) temperatures of tagged fish were 16–18 °C and potentially as high as 21 °C. Maximum 7DADM ambient water temperatures within the range of tagged fish during summer were 18–25 °C. However, there was no evidence of the tagged fish using localized cold water refuges. Tagged fish appeared to spawn at 7DADM temperatures of 7–14 °C. Maximum 7DADM temperatures of tagged fish and ambient temperatures at the onset of the spawning period in late August were 11–18 °C. Water temperatures in most of the upper Lostine River used for spawning and rearing appear to be largely natural since there has been little development, whereas downstream reaches used by migratory bull trout are heavily diverted for irrigation. Although the population effects of these temperatures are unknown, summer temperatures and the higher temperatures observed for spawning fish appear to be at or above the upper range of suitability reported for the species.     

The effect of water temperature on gamete maturation and gamete quality in the European grayling (Thymalus thymallus) based on experimental data and on data from wild populations

To investigate the effect of water temperature on gamete maturation and gamete quality, European grayling (Thymalus thymallus) were held under different temperature regimes prior to spawning. Maturation of males and females and their gamete quality depended strongly on temperature regime. The highest percentages of maturing fish and highest gamete quality were obtained under a creek water temperature regime with natural seasonal fluctuations. In warmed creek water (3–4°C), at a constant temperature of 8–9°C, and under an abruptly increasing temperature, regime maturation rates and gamete quality were reduced. The effect was more pronounced in females than in males. The spawning dates of different wild Austrian grayling populations were also correlated with water temperature data collected during the last 10–15 years. The mean spawning date expressed as the number of days from 21 December until spawning ranged from 98 to 111 days for lower elevation populations, while it was considerably delayed for an alpine population. All populations spawned at water temperatures of 5.5–7.2°C. Regression analysis indicated that spawning date correlated with (1) the overall mean water temperature from 21 December until spawning, (2) the mean water temperature during both the last 10 days and 15 days before spawning, and (3) the difference between mean January temperature and that of the last 15 days before spawning. The regression functions indicate that an increase in water temperature from 21 December to spawning of 1°C advances spawning by 5? days, and an increase of 1°C in the last 10–15 days advances spawning by 3? days.     

Inter‐annual variation of the spawning aggregations of the white‐streaked grouper Epinephelus ongus,in relation to the lunar cycle and water temperature fluctuation

The white‐streaked grouper, Epinephelus ongus (Serranidae), which forms spawning aggregations at specific times and sites, has been traditionally targeted by local fishers in the Yaeyama Islands, Okinawa. This study examined the long‐term dynamics of the population and the spawning aggregation of E. ongus, based on the historical catch data for a 20‐yr period. The continuous declines of the catch data suggested a substantial decrease in the population. The analysis of the daily catch data, which reflect the approximate volumetric fluctuation of the spawning aggregation, demonstrated that the distinctive peaks of daily catches appeared consistently over a 20‐yr span at the last quarter moon in the spawning season, in synchrony with the lunar phase as a spawning cue of the species. Contrary to this consistency, there were inter‐annual variations of spawning aggregation formation, which formed once or twice in the two consecutive months in the lunar calendar. The present study indicated that the water temperature before the aggregation period dictated not the only timing of the first spawning aggregation, but also the frequency of spawning aggregation and that the abundance between the first and the second spawning aggregation, represented by catch data, were negatively correlated. Together with previous evidence of gonadal histology, the findings indicated that water temperatures affect the gonadal maturation in the spawning population and control spawning aggregation formation. These facts imply that most individuals join in only a single spawning aggregation in the course of the year and highlight the importance of the spawning aggregation for reproduction of E. ongus.     

Spawning site selection by Eurasian perch (Perca fluviatilis L.) in relation to temperature and wave exposure

Abstract –  The selection of spawning depth by Eurasian perch Perca fluviatilis was investigated in an experiment using artificial substrata in Lake Constance during the spawning season of 2007. The experiment compared spawning behaviour at substrata between 0.5 and 15 m depth at two sites exposed to different regimes of ship-generated wave action. The total abundance of egg ribbons did not differ significantly between the two sites, but the preferred spawning depth was deeper at the wave exposed site (5 m) compared to the sheltered site (2 m). While water temperatures could not account for the observations, differences in wave exposure may explain the different spawning depth preferences. At both sites, large egg ribbons were generally found in deeper water, and large egg ribbons occurred more frequently at the sheltered site. Because the egg ribbons of perch are likely to have a size-dependent susceptibility to hydrodynamic stress, large females may be expected to select deeper spawning locations where the effects of surface waves are considerably attenuated.     

Seasonal changes in the immune activities of common carp (Cyprinus carpio)

Various endocrinological and immunological parameters involved with the seasonal changes of immunocompetence in common carp, Cyprinus carpio reared under natural conditions were investigated. Plasma concentrations of steroid hormones, IgM levels, IgM-secreting cell (IgMSC) numbers, and water temperature were considered as major parameters. Testosterone, 11-ketotestosterone in males and estradiol-17β in females detected by RIA, increased simultaneously with gonadal maturation, but no significant correlation was found with IgM production measured by ELISA. IgM levels were very high during the spawning season, but did not fluctuate much during the other seasons even at high water temperatures. These changes of IgM are certainly related to sexual maturation. In addition, IgMSC numbers were also high during the spawning season. This positive interaction between endocrine and immune parameters could contribute to the immune systems of common carp being more effective during spawning season. Interestingly, cortisol levels increased with the water temperature and were high during the spawning season, but had no suppressive effect on the IgM levels and IgMSC numbers.     

Relationship between recruitment of Japanese sardine (Sardinops melanostictus) and environment of larval habitat in the low‐stock period (1995–2010)

Stock level of Japanese sardine (Sardinops melanostictus) was high from 1980s to early 1990s and low from late 1990s to 2000s. The warm and cold water masses in the vicinity of the Kuroshio axis from winter to early spring used to be critical for the recruitment in the high‐stock period, because most of the larvae were distributed there. However, the environmental fluctuation might not affect the recruitment in the low‐stock period. Some studies reported that spawning location and spawning season, and hence the larval habitat, differ depending on the stock level. Three points were investigated in this study: (a) how spawning location and spawning season shifted from the late 1990s, (b) confirmation of the distribution area of larvae in the recent low‐stock period and (c) whether the water temperature in the vicinity of the Kuroshio axis was still related to the recruitment in the low‐stock period. The spawning location and spawning season clearly changed after 1995. Consequently, particle tracking experiments suggested that the larvae appeared in the vicinity of the Kuroshio axis from winter to early spring decreased. Nevertheless, only the ambient temperature of larvae that appeared in the vicinity of the Kuroshio axis from winter had a significant negative correlation with an index of the recruitment in the low‐stock period. It is suggested that the warm and cold masses in the vicinity of the Kuroshio axis are critical for the recruitment regardless of the stock level.     

Stability in reproductive timing and habitat usage of Chinook salmon across six years of varying environmental conditions and abundance

The spatial and temporal distribution of spawning activity by autumn‐run Chinook salmon Oncorhynchus tshawytscha (Walbaum) was examined across multiple years. The study period included two years of extreme drought conditions when water temperatures in the spawning reach of the Stanislaus River were considered sub‐optimal for spawning and egg incubation. Despite varying levels of superimposition, redd counts and densities remained stable and positively associated with river location, indicating that superimposition may be driven by habitat preference rather than the absence of suitable spawning locations. Spawning occurred slightly later during drought years (6–10 days later compared with wetter years). This delay was attributable to deferred migration instead of deferred spawning, and the majority of redds were constructed at water temperatures exceeding the optimal temperature range. As a consequence, estimates of juvenile production during 2014 and 2015 were among the lowest on record. These findings may be related in part to the high hatchery contribution to the population, above‐average temperatures during spawning and incubation, and superimposition rates. Management recommendations include adequate cold‐water storage in the upstream reservoir, refined spawning habitat restoration techniques in the light of superimposition rates and, on a broader system scale, actions that reduce the amount of stray hatchery Chinook salmon.     

Thermal regime in the hyporheic zone of two contrasting salmonid spawning streams: ecological and hydrological implications

The temperatures of stream water and hyporheic water were monitored in spawning areas of an upland stream (Girnock Burn) and a degraded lowland stream (Newmills Burn). Both streams are located in North-east Scotland and are spawning streams for Atlantic salmon, Salmo salar L., and brown trout, S. trutta L. Monitoring between the spawning and hatching of ova revealed significant differences in the thermal regimes of the two sites. Whilst temperatures in stream and hyporheic water were similar in the Girnock, notable attenuation in the Newmills Burn resulted in increasing moderation of thermal regime with hyporheic depth. This probably reflects contrasting groundwater–surface water interactions in the two systems. In the Girnock an open gravel stream bed and lack of groundwater influence resulted in stream water downwelling through the gravel. In contrast, fine bed sediments and local groundwater inputs resulted in more complex stratification of temperatures in Newmills Burn. However, even in the conditions of the Newmills Burn, temperature based salmonid embryo development models predicted that egg burial depth has an insubstantial effect on ova development.     

Plasma somatolactin levels in coho salmon (Oncorhynchus kisutch) during smoltification and sexual maturation

Somatolactin (SL) is a novel pituitary hormone recently characterized in several fish species. Structural analyses have shown that SL belongs to the growth hormone/prolactin family, and that it is a highly conserved protein. SL is synthesized by the periodic acid/Schiff-positive cells in the pars intermedia, but has an as yet unidentified function(s). We have recently developed a homologous radioimmunoassay for coho salmon SL and measured plasma levels of SL during two stages of the coho salmon life cycle, smoltification and sexual maturation. During smoltification, plasma levels of SL changed almost in parallel with plasma levels of thyroxine; levels increased as morphological indices of smoltification appeared and decreased as smoltification was completed. Following this period, SL levels remained low until the spring prior to spawning. In a separate study, plasma levels of SL were measured in sexually maturing coho salmon that remained in fresh water throughout their life cycle. During the year of sexual maturation, plasma levels of SL gradually increased from the spring onward, reaching peak levels at the time of spawning in November and December. These data are similar to those previously reported for sexually maturing coho salmon that were maintained in seawater prior to spawning (Rand-Weaver et al. 1992). Therefore, increases in plasma SL levels occurred in sexually maturing fish irrespective of whether they were maintained in fresh water or seawater. Peak levels at spawning were higher than those observed during smoltification. Possible roles for SL in metabolism and reproduction are discussed.     

Design and development of a geothermal temperature control system for broodstock management of channel catfish Ictalurus punctatus

A control system was designed to raise and maintain water temperatures within 0.03-ha earthen ponds to a range conducive for spawning (24–30 °C) channel catfish Ictalurus punctatus. Heating was done during February 2001 to April 2001, when temperatures would have otherwise prohibited spawning (<24 °C). Temperature was increased from 10 °C (ambient) by 2 °C per day, and maintained at 27 °C, by the addition of geothermally warmed water (36 °C). The control system substantially increased the controllability and precision of heating ponds compared to manual operation. Systems were designed to control sets of four ponds. In designing this control system, consideration of biological constraints was essential. Reproduction in channel catfish is most strongly influenced by temperature. Because cold fronts are common during the winter and early spring (January–March), it was essential to ensure that pond temperatures did not fall below the range for spawning. Constraints on the heating rate and temperature variability to maintain fish health and stimulate spawning behavior were considered. Components of the control system included temperature measurement devices (type-T thermocouples), a central electronic control unit, electronic switches and electrically actuated ball valves. In response to the temperature sensed by each thermocouple, the controller sent a message to close or open the valve. When the valve was opened, warm water was added to the pond to increase the average pond temperature. Hardware and algorithm design and initial system testing were the major components of this project. The final design incorporated information on relevant biological parameters and safety features including peak pond temperature, independent aeration and water pressure control mechanisms. Initial results indicate successful control of this biological system, and ongoing studies suggest similar mechanisms may be used for additional control objectives. In particular, this system could be used to vary pond temperatures to study biological responses and to cool ponds by addition of well water during summer months.     

Importance of broodstock holding temperature on fecundity and egg quality in three groups of photo‐manipulated Atlantic cod broodstock

We conducted a study to evaluate the effects of holding water temperature on the fecundity and egg quality of photo‐manipulated Atlantic cod broodstock. Adult cod Gadus morhua were distributed among several 25‐m³ tanks. Three separate photoperiods were used for each group to obtain three spawning per year (in May, July and December). While photoperiod was controlled, the temperature was not controlled and followed the ambient seasonal pattern. Due to the uncontrolled seasonal temperature used, each spawning group experienced different temperatures at a particular gonadal developmental stage. Otherwise, all groups were treated the same. Results showed that the May and July spawning groups released significantly more eggs per kg of fish (four and three times more respectively) than the December spawning group. The egg fertilization success and proportion of normal eggs were also higher in the May and July groups. Possible reason for the difference may be that the May and July groups experienced lower temperatures (4–7°C) during oocyte maturation and ovulation than the December group (7–9°C). Our results show the importance of using lower/natural temperatures during oocyte maturation and ovulation to obtain good quality eggs in photo‐manipulated cod broodstock.     

Incubation success and habitat selection of shore‐spawning kokanee Oncorhynchus nerka: effects of water‐level regulation and habitat characteristics

Changes to water‐level regimes have been known to restructure fish assemblages and interfere with the population dynamics of both littoral and pelagic species. The effect of altered water‐level regimes on shore‐spawning kokanee Oncorhynchus nerka incubation success was evaluated using a comprehensive in situ study in Lake Pend Oreille, ID, USA. Survival was not related to substrate size composition or depth, indicating that shore‐spawning kokanee do not currently receive a substrate‐mediated survival benefit from higher winter water levels. Substrate composition also did not differ among isobaths in the nearshore area. On average, the odds of an egg surviving to the preemergent stage were more than three times greater for sites in downwelling areas than those lacking downwelling. This study revealed that shoreline spawning habitat is not as limited as previously thought. Downwelling areas appear to contribute substantially to shore‐spawning kokanee recruitment. This research illustrates the value of rigorous in situ studies both for testing potential mechanisms underlying population trends and providing insight into spawning habitat selection.     

White sturgeon spawning and discharge augmentation

Abstract Observed spawning events of endangered Kootenai River white sturgeon, Acipenser transmontanus (Richardson), during 1994–2002 were examined to evaluate the effects of discharge from Libby Dam on spawning. Discharge from Libby Dam was manipulated in most years to provide enhanced flows (hereafter, augmented discharge) for white sturgeon spawning. The annual onset of spawning appeared independent of augmented discharge because initial spawning occurred prior to augmented discharge in 4 of 6 years. Spawning began 2–11 days after the highest river discharge in 4 of 9 years. Linear regression analysis indicated the onset of spawning was positively related to mean daily discharge from the dam but not mean daily temperature, 1‐day change in discharge (Δ discharge) or 3‐day Δ discharge. Logistic regression analysis suggested the probability of a spawning event was influenced by Julian date, mean daily water temperature, mean daily discharge and the 7‐day Δ discharge, but the predictive indices were small. Minor fluctuations in temperature and discharge characteristics had little additional predictive benefit for spawning after the river reached spawning thresholds. The highest probability of spawning (0.48) was for the temperature interval 9.5–9.9 °C, and 93% of the estimated spawning events occurred above 8 °C. Sixty percent of the estimated spawning events occurred at discharges ≥600 m³ s^?1, which comprised 45% of the range of discharge values.     

金乌贼新型产卵附着基的实验研究

2015年6—7月和2016年6—7月在青岛金沙滩水产有限公司室内产卵池中开展了不同材料、不同颜色、不同水深金乌贼(Sepia esculenta)产卵附着基附卵效果对比实验,以及柽柳附卵器对附卵效果的影响实验。实验用产卵附着基为0.33 m×0.33 m×0.33 m的十字形可折叠钢丝框架,上覆黑白2种聚乙烯网衣材料;每个产卵池中金乌贼亲体数量约35只,雌雄比约为1∶1,胴长(139.2±1.1)mm,体重(435.9±8.5)g。同时选用等比例放大的1.0 m×1.0 m×1.0 m钢筋框架与不同颜色网衣制作的十字形可折叠附着基,于2015年6—7月在青岛斋堂岛附近海域(35?36?6?N,119?54?8?E)和杨家洼湾前附近海域(35?36'6?N,119?50'50?E),以及2016年6—7月在青岛薛家岛近海(35?57?24?N,120?16?22?E)和杨家洼湾前附近海域(35?35?45?N,119?50?38?E)开展了海区附卵验证实验。结果表明:室内同一水体环境中柽柳附卵效果最佳,其次是十字形附着基,海参框附卵效果最差;白色十字形附着基的附卵效果显著优于黑色(P0.05);柽柳附卵器对附卵效果有积极影响,加装柽柳附卵器的十字形附着基附卵效果明显提高;产卵池底层摆放的十字形附着基的附卵效果显著优于表层悬挂的十字形附着基(P0.01)。海区附卵验证结果表明:2015年和2016年海区投放的十字形附着基附卵效果良好,最高附卵率分别为82.14%和56.72%;海区投放的黑色和白色十字形附着基的附卵效果差异不显著(P0.05);海底悬挂的十字形附着基的附卵效果显著优于近底层悬挂的十字形附着基(P0.05)。综合分析认为,在金乌贼产卵繁殖季节,选择其重要产卵场和主要增殖放流海区水深15~20 m的缓流区,底层设置十字形可折叠产卵附着基,可有效进行资源增殖与产卵场修复。     

A basic experiment of coral culture using sexual reproduction in the open sea

Coral larvae, produced from a mass spawning event, were successfully settled on special stone settlement sticks and raised in situ for eventual transport to other reefs. The test area, Sekisei lagoon, Okinawa Prefecture, Japan, is located close to the warm Kuroshio current and is surmised to be the source from which major Japanese corals are derived. A total of 131 settlement sticks, with small holes in their sides to increase protection from grazing (4 mm in diameter and approximately 5 mm deep), were deployed in the lagoon the day before the coral’s mass spawning. After 3 months, 61 sticks were recovered containing 71 corals, mostly in the holes. After 1 year, three corals were confirmed to be growing well and extending outside the holes of the three sticks out of 70 sticks left in the water at the lagoon site. They survived two potentially lethal conditions, that is, high water temperatures with associated extensive coral bleaching and continuous grazing pressure from predators. This procedure is applicable for large-scale coral transplantation, not only in Japan but also in other tropical countries.     

Reproductive characteristics of the fishery important temperate demersal berycid Centroberyx gerrardi indicate greater reproductive output in regions of upwelling

The reproductive biology of Centroberyx gerrardi (Günther) was investigated across ~2,000 km of its southern Australian distribution, encompassing different jurisdictions and varying environmental features. Greater gonad mass and prevalence of spawning fish, along with lower ratios of lengths at maturity:maximum lengths and ages at maturity:maximum ages, were identified at the western‐most (Capes) and eastern‐most (Great Australian Bight; GAB) regions. Across the study region, spawning peaks in summer/autumn, when water temperatures are warmest. Regional differences in potential “reproductive output,” while not consistent with the eastward decline in mean monthly water temperature, may instead be related to summer upwelling in the Capes and GAB, driving greater oceanic productivity prior to peak spawning, supporting larval survival. In autumn, the prevailing southward and eastward flowing, downwelling Leeuwin Current (LC) strengthens, providing a dispersal mechanism along the west and south Australian coasts, but limiting upwelling effects. Predicted changes in environmental conditions and their potential impacts on C. gerrardi are discussed, in particular how these factors may affect recruitment to stocks and fisheries, requiring a better understanding of source‐sink relationships for this species. As environmental changes occur, management strategies to sustain fish resources must adapt to spatially variable and changing reproductive output and be collaborative across jurisdictions.     

Skipped spawning in female iteroparous fishes

It is often assumed that iteroparous fishes spawn annually once reaching sexual maturity, but this is not always the case. This paper reviews available information on skipped spawning in female teleost fishes. All instances of non‐annual spawning are described as one of three types (retaining, reabsorbing, resting), depending on where in the normal spawning cycle development has been interrupted. Retaining ripe eggs is caused by conditions experienced during the spawning season (fish density, mate availability, pollution), whereas failure to start vitellogenesis (resting) or the breakdown of all oocytes that enter into vitellogenesis (reabsorbing) is caused by factors experienced prior to the spawning season (primarily temperature and poor nutrition). It is speculated that the relative shortage of data on non‐annual spawning may be because of difficulties in identifying non‐reproductive individuals. In an attempt to rectify this situation, the criteria needed to identify females undergoing the three forms of spawning omission are presented in terms of external appearance of gonads, gonad indices, and histological analysis. The energy saved by not spawning in a poor year may lead to increased survival and the probability of spawning in subsequent years. As the cumulative number of progeny gained by surviving to spawn in multiple subsequent years outweighs the number of progeny lost by not spawning in a given single year, occasional omission of spawning may constitute an adaptive trait in long‐lived iteroparous fishes.