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.     

Effects of reproductive timing and hatch date on fathead minnow recruitment

Abstract –  Timing of reproduction can be an important determinant of recruitment success for fish in strongly seasonal environments, both for individuals hatching at different times over extended spawning seasons and for entire cohorts, if the spawning period is accelerated or delayed among years. To examine potential demographic consequences of delayed spawning, we staggered dates that fathead minnow ( Pimephales promelas ) were introduced into experimental ponds in Alberta, Canada, by 3 weeks and compared growth and survival of offspring from early- and late-stocked fish. By autumn, size and mass were consistently greater in early-hatched minnows, regardless of cohort density. Size differences between treatments persisted through the second summer, which likely contributed to a higher proportion of early-hatched fish reaching sexual maturity as yearlings compared with late-hatched counterparts. Because late spawning can limit growth and subsequent maturation of progeny, temporal variability in reproductive timing should be considered when assessing recruitment potential of new year-classes.     

丹江口水库太湖新银鱼不同繁殖群体的繁殖特征比较

2010年6月—2011年6月,对丹江口水库太湖新银鱼(Neosalanx taihuensis)不同繁殖群体的繁殖生物学特征进行了比较研究。结果表明:太湖新银鱼在该水库产卵时间为1—5月(春群)和9—10月(秋群);其中春季群体的体长(65.79 mm±6.05 mm)、性腺成熟指数(17.26±6.42)、繁殖力(1851粒±342粒)和成熟卵径(0.62 mm±0.05mm)均显著高于秋季群体(体长(55.04 mm±3.18 mm)、性腺成熟指数(11.72±5.01)、繁殖力(1 067粒±190粒)和成熟卵径(0.53 mm±0.04 mm))(P<0.05);但是两个群体的丰满度指数(101.43±19.71,101.87±11.85)差异并不显著(P>0.05)。春群是丹江口水库太湖新银鱼种群的主要补充者,春群管理与控制应是银鱼种群管理的重点。     

Climate‐induced vulnerability of fisheries in the Coral Triangle: Skipjack Tuna thermal spawning habitats

We investigated projected changes in sea surface temperature (SST) and the associated impacts on spawning habitat for skipjack tuna (Katsuwonus pelamis) in the Coral Triangle region (CT). A multimodel aggregate of SST CMIP5 models for the CT region, based on a comprehensive skill validation assessment, was used to identify the five best performing of 36 models tested for inclusion in a regional multimodel ensemble. Monthly 1° SST multimodel aggregate projections for the CT region under RCP8.5 show that increases in SST, as high as 2.8°C (mean value), will likely occur by the end of this century. Using these estimates of SST change, we applied three parameterizations of skipjack tuna spawning temperatures to assess the potential for change in spawning habitat within the CT region. The three spawning temperature parameterizations were as follows: (a) a square‐wave function derived from catch data with boundaries at 26 and 30°C; (b) a symmetric Gaussian function derived from the SEAPODYM models; and (c) an asymmetric Gaussian function that modifies the SEAPODYM curve in (b) to include the results of relevant physiological experiments. All three parameterizations show similar geographic patterns, with the amount of favourable spawning habitat decreasing throughout the central, equatorial CT region and increasing at higher latitudes. However, the three parameterizations show marked differences in the modelled magnitude of change, with an asymmetric Gaussian function (ASGF) showing a regionwide average of 66.1% decline in favourable spawning habitat between 2015 and 2099. These projected changes in tuna spawning habitats are likely to have important consequences on local and regional fisheries management in the CT region.     

Fishery indicators during a predictable aggregation of Atlantic sharpnose sharks Rhizoprionodon terraenovae in the southern Gulf of Mexico: An alternative to assess a heterogeneous data‐poor fishery

Small‐scale fisheries in the southern Gulf of Mexico that catch Rhizoprionodon terraenovae (Richardson) are heterogeneous and data‐poor. Fishery‐dependent monitoring was conducted from 2010 to 2017, including a target season during an aggregation of this species to estimate data‐poor fishery indicators. During the target season, the average sizes for females and males (95.3 and 89.8 cm total length, respectively) were recorded, a global male sex bias (1:1.7), the highest percentage of mature sharks for all years (>89%), the highest values of CPUE (20.1 sharks/day) and size‐selectivity higher than the size at maturity. The spawning potential ratio was over 0.6 (reference point of 0.71) in the combined (target and non‐target) and target seasons for all years, which suggests that the fishery stock is not healthy. Annual assessment of this fishery can be carried out through monitoring during the target season, where management is more feasible to implement.     

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.     

Variations in reproductive potential between nearshore and offshore spawning contingents of hogfish in the eastern Gulf of Mexico

Hogfish, Lachnolaimus maximus (Walbaum), have multiple traits that confound measuring reproductive potential: they are protogynous, relatively long‐lived harem‐forming fish that spawn daily for months. Additionally, recent evidence demonstrates that size, age and timing of sex change vary on a spatial scale within the study area (West Florida shelf, USA). This study investigates the effect of this spatial variation on hogfish reproductive potential by evaluating spawning seasonality, spawning frequency and batch fecundity using an indeterminate egg production model. Offshore females were larger than nearshore females, and batch fecundity was related in a log‐linear manner to female size. Gonad histology demonstrated a more protracted reproductive period for females offshore (8 months) than nearshore (4 months). Spatial variations in size coincide with ontogeny because hogfish move offshore with growth; however, even after accounting for fish size, offshore females spawned more. In areas where male removal rates are elevated, spawning harems are disrupted; thus, greater fishing effort nearshore may further reduce the reproductive potential of these females. These nearshore and offshore spawning components of the population are not genetically distinct, but instead represent two contingent spawning strategies that likely enhance total population stability and resilience of this stock in the eastern Gulf of Mexico.     

Ocean eddies,predator pits and bluefin tuna: implications of an inferred ‘low risk–limited payoff’ reproductive scheme of a (former) archetypical top predator

The conventional view of the reproductive ecology of bluefin tunas (Thunnus thynnus, Thunnus orientalis, Thunnus maccoyii) leads to a conceptual paradox. Simple arithmetic yields an expectation for wide variation in annual reproductive success. However, the historical record does not support this prediction. A conclusion would seem to be that a much stronger than usual density dependence must be acting. It is here argued that this strong density dependence may likely occur within small strongly convergent segments of energetically forced ocean eddy structures. Success of the implied ecological scenario requires spawning schools of sufficient size to generate sufficiently copious reproductive product to circumvent resident predator pits, while exerting sufficient predatory loss on resident predators to facilitate this circumvention. This in turn implies existence of a dangerous ‘precipice’ in the form of self‐enhancing feedback loop, lurking unperceived beyond the range of historical experience, and a need for a particular degree of precaution in managing the exploitation of this iconic species.     

Selectivity matters: Rules of thumb for management of plate‐sized,sex‐changing fish in the live reef food fish trade

Effective management of fisheries depends on the selectivity of different fishing methods, control of fishing effort and the life history and mating system of the target species. For sex‐changing species, it is unclear how the truncation of age‐structure or selection of specific size or age classes (by fishing for specific markets) affects population dynamics. We specifically address the consequences of plate‐sized selectivity, whereby submature, “plate‐sized” fish are preferred in the live reef food fish trade. We use an age‐structured model to investigate the decline and recovery of populations fished with three different selectivity scenarios (asymptotic, dome‐shaped and plate‐sized) applied to two sexual systems (female‐first hermaphroditism and gonochorism). We parameterized our model with life‐history data from Brown‐marbled grouper (Epinephelus fuscoguttatus) and Napoleon fish (Cheilinus undulatus). “Plate‐sized” selectivity had the greatest negative effect on population trajectories, assuming accumulated fishing effort across ages was equal, while the relative effect of fishing on biomass was greatest with low natural mortality. Fishing such sex‐changing species before maturation decreased egg production (and the spawning potential ratio) in two ways: average individual size decreased and, assuming plasticity, females became males at a smaller size. Somatic growth rate affected biomass if selectivity was based on size at age because in slow growers, a smaller proportion of total biomass was vulnerable to fishing. We recommend fisheries avoid taking individuals near their maturation age, regardless of mating system, unless catch is tightly controlled. We also discuss the implications of fishing post‐settlement individuals on population dynamics and offer practical management recommendations.     

Histological observations of seasonal reproductive and lunar-related spawning cycles in the female honeycomb grouper Epinephelus merra in Okinawan waters

Seasonal reproductive and lunar-related spawning cycles of the female honeycomb grouper Epinephelus merra inhabiting Okinawan waters were examined by histological observation of ovaries. Gonadosomatic index (GSI) increased beginning in May and peaked in June. Histological observations revealed that many oocytes laden with yolk were present in the ovaries from May to August. From September to October, ovaries were occupied by immature oocytes. These results suggest that the reproductive season of E.   merra lasts for 4 months from May through August in Okinawan waters. When the fish were collected according to the lunar cycle, GSI increased with the approach of the full moon. Oocytes at various development stages were observed from the first quarter to the full moon. Fresh ovulatory follicles were present in the ovaries around the last quarter moon. These results suggest that E.   merra has a lunar spawning cycle and spawns between the full moon and the last quarter moon. Ovarian features of the fish collected around the last quarter moon were different among individuals; some fish had many oocytes at the tertiary yolk stage in the ovaries, while the ovaries of the others were occupied by the oocytes at the peri-nucleolus and the oil droplet stages. This observation suggests that a minor release of eggs occurs in this species before or after a major spawning lunar day. Oocytes at the migratory nucleus and the maturation stages were not observed in any ovarian samples. This may mean that maturation of oocytes is related to the spawning behavior of this species and makes rapid progress in the process of aggregation at the spawning sites.     

西南大西洋阿根廷滑柔鱼夏季产卵种群繁殖生物学的初步研究

根据我国鱿钓船于2012年12月—2013年3月渔汛期间,在西南大西洋公海海域作业时随机采集的阿根廷滑柔鱼夏季产卵种群样本(雌性148尾,雄性99尾),初步研究了其种群繁殖生物学。结果表明,渔汛期间阿根廷滑柔鱼夏季产卵种群雌性胴长172~252 mm,雄性胴长164~248 mm。随着月份推移,雌性占比逐月递减,从12月份的84.21%递减至次年3月份的27.03%;雄性个体占比则逐月递增,从12月份的15.79%递增至次年3月份的72.97%。雌性样本超过30%的个体性腺发育成熟并且部分处于繁殖期,雄性个体超过60%的个体性腺发育成熟并且部分处于繁殖期。雌雄个体的胴长和体质量均在性腺功能发育成熟时达到最大值,但是雌性胴长大于雄性,体质量却小于雄性。性腺指数、缠卵腺指数和精荚复合体指数随着个体生长逐渐增长。雌性个体性腺指数为0.001 2~0.31,平均值0.074;缠卵腺指数为0.003 1~0.55,平均值0.29;两者随着个体生长逐渐增长,在性腺功能成熟时达到最大值。雄性个体性腺指数为0.007 5~0.084,平均值0.038;精荚复合体指数为0.000 79~0.035,平均值0.012;两指数在繁殖期时达到最大值。     

Reproductive resilience or sweepstakes recruitment? Assessing drivers of lifetime reproductive success in exploited marine fish

Understanding the processes that drive reproductive success in marine fish stocks is critical to effective fisheries management. These processes can be difficult to investigate, especially in age-structured populations, because they occur at transgenerational scales. Reproductive success is often attributed to a small portion of the adult population (<0.01%) and thought to be driven primarily by random external factors, consistent with the concept of sweepstake reproductive success (SRS). A competing concept, the reproductive resilience paradigm, posits that fishes have evolved complex spawner-recruit systems to achieve lifetime reproductive success and maintain population stability within highly variable environments. Here, we examine these two concepts. First, we analyse the popular sport fish red drum (Sciaenops ocellatus, Sciaenidae), drawing on genetic and reproductive data to estimate a plausible range for the ratio of effective population size ($N_e$) to adult abundance ($N_A$) and to infer variance in lifetime reproductive success ($V_k^{*}$). Then, we synthesize available data and infer $V_k^{*}$ for two other fishes that have $N_e/N_A$ ratios reportedly >0.10, the southern bluefin tuna (Thunnus maccoyii, Scombridae) and the silver seabream (Chrysophrys auratus, Sparidae). Although commonly regarded as an SRS species, red drum did not meet the SRS criterion. Overdispersed $V_k^{*}$ values were inferred for all three species, with those for red drum and silver seabream being dependent upon population-closure assumptions. Results are presented within the conceptual framework of reproductive resilience, considering the roles of random extrinsic forces versus evolved traits to achieve lifetime reproductive success and population stability in high and variable mortality environments.     

The highs and lows of herring: A meta‐analysis of patterns and factors in herring collapse and recovery

Pacific and Atlantic herring populations (genus Clupea) commonly experience episodic collapse and recovery. Recovery time durations are of great importance for the sustainability of fisheries and ecosystems. We collated information from 64 herring populations to characterize herring fluctuations and determine the time scales at low biomass and at high and low recruitment, and use generalized linear models and Random Survival Forests to identify the most important bottom‐up, top‐down and intrinsic factors influencing recovery times. Compared to non‐forage fish taxa, herring decline to lower minima, recover to higher maxima and show larger changes in biomass, implying herring are more prone to booms and busts than non‐forage fish species. Large year classes are more common in herring, but occur infrequently and are uncorrelated among regionally grouped stocks, implying local drivers of high recruitment. Management differs between Pacific and Atlantic herring fisheries, where at similarly low biomass, Pacific fisheries tend to be closed while Atlantic fisheries remain open. This difference had no apparent effect on herring recovery times, which averaged 11 years, although most stocks with longer recovery periods had not yet recovered at the end of the observation period. Biomass recovery is best explained by median recruitment and variability in sea surface height anomalies and sea surface temperatures—higher variability leads to shorter recovery times. In addition, the duration of recruitment failure is closely linked with low biomass. While recovery times rely on the nature of the relationship between spawning biomass and recruitment, they are still largely governed by complex and uncertain processes.     

Wind‐induced variability in coastal larval retention areas: a case study on Western Baltic spring‐spawning herring

The investigation of larval dispersal and retention, their variability and dependence on wind conditions, has become a major topic in fisheries research owing to potential effects on stock recruitment and stock structuring. The present study quantifies the wind‐induced variability of larval retention of herring in a highly productive coastal lagoon of the Western Baltic Sea. This lagoon, the Greifswalder Bodden, represents the main spawning area of Western Baltic Spring‐Spawning Herring, a stock that has recently undergone a continuous decline in recruitment. The study tests whether this decline was related to changes in larval retention, more precisely to changes in wind conditions, the main forcing of the lagoon's circulation. To answer this, a model approach was applied. Larvae were tracked as Lagrangian drifters under constant and variable wind conditions, examining the main drift patterns and reconstructing the incidents during the period of recruitment decline. For the latter, weekly cohorts of virtual larvae were released in the lagoon over the entire spawning period (April–June; >16 weeks). The fraction of retained larvae per cohort was related to observed larval abundances. On this basis, a new retention index was defined to evaluate the annual larval retention. The results presented cannot explain the observed recruitment decline but characterize the lagoon as an important larval retention area by virtue of unsteady wind conditions that prevent a steady outflow of larvae.     

Modelling potential spawning habitat of sardine (Sardina pilchardus) and anchovy (Engraulis encrasicolus) in the Bay of Biscay

Large amplitude variations in recruitment of small pelagic fish result from interactions between a fluctuating environment and population dynamics processes such as spawning. The spatial extent and location of spawning, which is critical to the fate of eggs and larvae, can vary strongly from year to year, as a result of changing population structure and environmental conditions. Spawning habitat can be divided into ‘potential spawning habitat’, defined as habitat where the hydrographic conditions are suitable for spawning, ‘realized spawning habitat’, defined as habitat where spawning actually occurs, and ‘successful spawning habitat’, defined as habitat from where successful recruitment has resulted. Using biological data collected during the period 2000–2004, as well as hydrographic data, we investigate the role of environmental parameters in controlling the potential spawning habitat of anchovy and sardine in the Bay of Biscay. Anchovy potential spawning habitat appears to be primarily related to bottom temperature followed by surface temperature and mixed‐layer depth, whilst surface and bottom salinity appear to play a lesser role. The possible influence of hydrographic factors on the spawning habitat of sardine seems less clear than for anchovy. Modelled relationships between anchovy and sardine spawning are used to predict potential spawning habitat from hydrodynamical simulations. The results show that the seasonal patterns in spawning are well reproduced by the model, indicating that hydrographic changes may explain a large fraction of spawning spatial dynamics. Such models may prove useful in the context of forecasting potential impacts of future environmental changes on sardine and anchovy reproductive strategy in the north‐east Atlantic.     

不同交配,越冬方式对河蟹育苗的影响

不同交配、越冬方式对河蟹育苗产量有着显著的影响，三年的生产实践表明：秋季室外土池交配的抱卵率、抱卵量明显高于水泥交配；秋季交配优于春季交配；室外土池自然越冬抱卵卵蟹的胚胎发育优于室外塑料大棚保温越冬的抱卵蟹。适宜的秋，春交配有利于河蟹多苗育苗，延长生长期，增加产量。     

Solar and lunar influences on the spawning activity of Pacific bluefin tuna (Thunnus orientalis) in the south‐western North Pacific spawning ground

Pacific bluefin tuna (Thunnus orientalis) migrate to the south‐western North Pacific spawning grounds (area off Taiwan and the Ryukyu Islands) from mid‐April to early July. Reproductive traits of the species in this area have been studied for some time; however, temporal changes in spawning activity during this season are not well understood. In this study, the spawning fraction (proportion of females with post‐ovulatory follicles to total mature females) in relation to solar calendar date and lunar day was investigated using a generalized linear model. Sampling was conducted almost every day throughout the fishing season at Ishigaki Fishing Port (24°21′N, 124°09′E), southern Japan between 2012 and 2014, and a total of 212 ovaries were histologically observed. Spawning fraction continued to increase from the beginning to the end of the season. This indicates that the peak season of occurrence in the area and the peak of spawning activity did not co‐occur. Spawning fraction also changed with the lunar day and significantly increased around the new moon. Eggs and early larvae hatched around the new moon may have reduced the predation risk during night‐time. These findings improve the current knowledge of reproductive biology and resource evaluation of Pacific bluefin tuna and indicate that spawning activity is not constant, and has two or three peaks in each season.     

Temperature conditions in the Argentine chub mackerel (Scomber japonicus) fishing ground: implications for fishery management

We describe the seasonal migrations of Argentine chub mackerel ( Scomber japonicus ) into the fishing ground in relation to sea surface temperature conditions and analyse the temperature evolution during the fishing season on the basis of 1955–97 time-series. The upper temperature limit for the presence of schools was around 19°C. At higher values, most schools leave coastal waters (< 50 m) for the shelf. An inverse relationship was observed between catches per unit of effort (CPUE) of the inshore fishery and sea surface temperature. The highest standardized CPUE values corresponded to temperatures lower than 19°C. Above this temperature, the probability of obtaining a standardized CPUE higher than 1 was extremely low, and the standard deviations were minimal. The economic benefit from catches was also analysed through the different months of the fishing season. It was found that the mean benefit corresponding to temperatures lower than 19°C exceeded approximately 15 times that obtained at higher temperatures. A simple fishing strategy was proposed to the commercial fleet in order to improve the economic benefit of the catches. The migratory pattern followed by mackerel during the spawning period was also analysed in relationship to other environmental factors like food availability. Zooplankton composition, size structure and abundance in coastal and shelf waters are described, and their implications for feeding ecology of larvae and adults of the species are discussed.     

Fish spawning aggregations: where well‐placed management actions can yield big benefits for fisheries and conservation

Marine ecosystem management has traditionally been divided between fisheries management and biodiversity conservation approaches, and the merging of these disparate agendas has proven difficult. Here, we offer a pathway that can unite fishers, scientists, resource managers and conservationists towards a single vision for some areas of the ocean where small investments in management can offer disproportionately large benefits to fisheries and biodiversity conservation. Specifically, we provide a series of evidenced‐based arguments that support an urgent need to recognize fish spawning aggregations (FSAs) as a focal point for fisheries management and conservation on a global scale, with a particular emphasis placed on the protection of multispecies FSA sites. We illustrate that these sites serve as productivity hotspots – small areas of the ocean that are dictated by the interactions between physical forces and geomorphology, attract multiple species to reproduce in large numbers and support food web dynamics, ecosystem health and robust fisheries. FSAs are comparable in vulnerability, importance and magnificence to breeding aggregations of seabirds, sea turtles and whales yet they receive insufficient attention and are declining worldwide. Numerous case‐studies confirm that protected aggregations do recover to benefit fisheries through increases in fish biomass, catch rates and larval recruitment at fished sites. The small size and spatio‐temporal predictability of FSAs allow monitoring, assessment and enforcement to be scaled down while benefits of protection scale up to entire populations. Fishers intuitively understand the linkages between protecting FSAs and healthy fisheries and thus tend to support their protection.