Timing and duration of mating and brooding periods of Atka mackerel (Pleurogrammus monopterygius) in the North Pacific Ocean

The timing and duration of the reproductive cycle of Atka mackerel (Pleurogrammus monopterygius) was validated by using observations from time-lapse video and data from archival tags, and the start, peak, and end of spawning and hatching were determined from an incubation model with aged egg samples and empirical incubation times ranging from 44 days at a water temperature of 9.85°C to 100 days at 3.89°C. From June to July, males ceased diel vertical movements, aggregated in nesting colonies, and established territories. Spawning began in late July, ended in mid-October, and peaked in early September. The male egg-brooding period that followed continued from late November to mid-January and duration was highly dependent on embryonic development as affected by ambient water temperature. Males exhibited brooding behavior for protracted periods at water depths from 23 to 117 m where average daily water temperatures ranged from 4.0° to 6.2°C. Knowledge about the timing of the reproductive cycle provides a framework for conserving Atka mackerel populations and investigating the physical and biological processes influencing recruitment.

Description of embryonic development of Atka mackerel (Pleurogrammus monopterygius)

Atka mackerel (Pleurogrammus monopterygius) is hexagrammid fish that inhabits the temperate and subarctic North Pacific Ocean and neighboring seas (Fig. 1). This highly abundant fish is a critically important prey species (Sinclair and Zeppelin, 2002; Zenger, 2004) that supports a directed commercial trawl fishery (Lowe et al., 2006). Atka mackerel is a demersal spawner and males provide parental care to eggs (Zolotov, 1993). During breeding periods, sexually mature males aggregate on the bottom at nesting sites where they establish territories (Lauth et al., in press). Sexually mature females periodically visit male nesting territories from July to October to spawn batches of demersal egg masses (McDermott and Lowe, 1997; McDermott et al., 2007). Individual nests may consist of multiple egg masses deposited by different females, and males defend nesting territories for a protracted period lasting from the time territories are being established until all eggs within the territory are completely hatched (Lauth et al., 2007). Knowledge about the timing of the reproductive cycle and the use of spawning habitat are important for understanding population structure and the dynamics of stock recruitment, which in turn are important factors in the management of Atka mackerel populations.

Validation of a method for estimating realized annual fecundity in a multiple spawner, the long-snouted seahorse (Hippocampus guttulatus), using underwater visual census

The long-snouted seahorse (Hippocampus guttulatus) (Cuvier, 1829), was used to validate the pre-dictive accuracy of three progressively realistic models for estimating the realized annual fecundity of asyn-chronous, indeterminate, multiple spawners. Underwater surveys and catch data were used to estimate the duration of the reproductive season, female spawning frequency, male brooding frequency, and batch fecun-dity. The most realistic model, a generalization of the spawning fraction method, produced unbiased estimates of male brooding frequency (mean ±standard deviation [SD]=4.2 ±1.6 broods/year). Mean batch fecundity and realized annual fecundity were 213.9 (±110.9) and 903.6 (±522.4), respectively. However, females prepared significantly more clutches than the number of broods produced by males. Thus, methods that infer spawning frequency from patterns in female egg production may lead to significant overestimates of realized annual fecundity. The spawning fraction method is broadly applicable to many taxa that exhibit parental care and can be applied nondestructively to species for which conservation is a concern.

Geographic and seasonal variations in maturation and growth of female Pacific cod (Gadus macrocephalus) in the Gulf of Alaska and Bering Sea

This study investigates the temporal stability of length- and age-at-maturity estimates for female Pacific cod (Gadus macrocephalus) in the Gulf of Alaska and eastern Bering Sea. Females reached 50% maturity (A50) at 4.4 years in the Gulf of Alaska and at 4.9 years in the eastern Bering Sea. Total body length at 50% maturity (LT50) was significantly smaller (503 mm) in the Gulf of Alaska than in the eastern Bering Sea (580 mm). The estimated length- and age-at-maturity did not differ significantly between winter and spring in either the Gulf of Alaska (1999) or Bering Sea (2003) areas. The results of this study raised the spawning biomass estimate of female Alaskan Pacific cod from 298×103 t for 2005 to 499×103 t for 2006. The increased spawning biomass estimate resulted in an increased over-fishing limit for Pacific cod.

Annual fecundity, batch fecundity, and oocyte atresia of Atka mackerel (Pleurogrammus monopterygius) in Alaskan waters

Annual potential fecundity, batch fecundity, and oocyte atresia were estimated for Atka mackerel (Pleurogrammus monopterygius) collected in Alaskan waters during 1993−94. Atka mackerel were assumed to be determinate spawners on the basis of decreasing fecundity after batch spawning events. Histological examination of the ovaries indicated that oocytes in the vitellogenic stage and higher had been spawned in the current spawning season. For an average female of 40 cm, potential annual fecundity was estimated to be 41,994 eggs, average batch size (i.e., batch fecundity) was estimated to be 6689 eggs, and there were 6.13 batches per spawning season. Atresia was estimated by examining postspawning specimens and was found to be substantial. The average amount of atresia for a 40-cm fish was estimated to be 11,329 eggs, resulting in an estimated realized fecundity of only 30,664 eggs and 4.64 batches of eggs per spawning season.

Oceanic migration rates of Upper Chesapeake Bay striped bass (Morone saxatilis), determined by otolith microchemical analysis*

Oceanic incidence and spawning frequency of Chesapeake Bay striped bass (Morone saxatilis) were estimated by using microchemical analysis of strontium in otoliths. Otoliths from 40 males and 82 females sampled from Maryland’s portion of the Chesapeake Bay were analyzed for seasonal and age-specific patterns in strontium and calcium levels. The proportion of oceanic females increased from 50% to 75% between ages seven to 13; the proportion of oceanic males increased from 20% to ~50% between ages four to 13. Contrary to an earliermodel of Chesapeake Bay striped bass migration, results indicated that a substantial number of males undertook oceanic migrations. Further, we observed no mass emigration of females from three to four years of age from the Chesapeake Bay. Seasonal patterns of estuarine habitat use were consistent with annual spawning runs by striped bass of mature age classes, but with noteworthy exceptions for newly mature females. Evidence of an early oceanic presence indicated that Chesapeake Bay yearlings move into coastal regions—a pattern observed also for Hudson River striped bass. Otolith microchemical analyses revealed two types of behaviors (estuarine and oceanic) that confirm migratory behaviors recently determined for other populations of striped bass and diadromous species (e.g., American eels [Anguilla rostrata] American shad [Alosa sapidissima] and white perch [Morone Americana]).

Degeneration of postovulatory follicles in the Iberian sardine Sardina pilchardus: structural changes and factors affecting resorption

Inaccuracy in the aging of postovulatory follicles (POFs) and in estimating the effect of temperature on the resorption rate of POFs may introduce bias in the determination of the daily spawning age classes with the daily egg production method (DEPM). To explore the above two bias problems with f ield-collected European pilchard (Sardina pilchardus, known regionally as the Iberian sardine), a method was developed in which the time elapsed from spawning (POF age) was estimated from the size of POFs (i.e., from the cross-sectional area in histological sections). The potential effect of the preservative type and embedding material on POF size and the effect of ambient water temperature on POF resorption rate are taken into account with this method. A highly significant loglinear relationship was found between POF area and age; POF area shrank by approximately 50% per day. POFs were also shown to shrink faster at higher temperatures (approximately 3% per degree), but this temperature effect is unlikely to be an important source of bias in the assignment of females to daily spawning classes. The embedding material was also shown to influence the size of POFs, the latter being significantly larger in resin than in paraffin sections. In conclusion, the size of POFs provides an indirect, reliable estimation of the time elapsed from spawning and may thus be used to test both the validity of POF staging criteria for identifying daily classes of spawners and the effect of other factors (such as temperature and laboratory processing) in applications of the DEPM to S. pilchardus and other fish species.

Biological characteristics and mortality of western butterfish (Pentapodus vitta), an abundant bycatch species of prawn trawling and recreational fishing in a large subtropical embayment

The western butterfish (Pentapodus vitta) is numerous in the bycatch of prawn trawling and recreational fishing in Shark Bay, Western Australia. We have thus determined crucial aspects of its biological characteristics and the potential impact of fishing on its abundance within this large subtropical marine embayment. Although both sexes attained a maximum age of 8 years, males grow more rapidly and to a larger size. Maturity is attained at the end of the first year of life and spawning occurs between October and January. The use of a Bayesian approach to combine independent estimates for total mortality, Z, and natural mortality, M, yielded slightly higher point estimates for Z than M. This result indicates that P. vitta is lightly impacted by fishing. It is relevant that, potentially, the individuals can spawn twice before recruitment into the fishery and that 73% of recreationally caught individuals are returned live to the water.

Interdecadal change in growth of sablefish (Anoplopoma fimbria) in the northeast Pacific Ocean

Errors in growth estimates can affect drastically the spawner-perrecruit threshold used to recommend quotas for commercial fish catches. Growth parameters for sablefish (Anoplopoma fimbria) in Alaska have not been updated for stock assessment purposes for more than 20 years, although aging of sablefish has continued. In this study, length-stratified data (1981–93 data from the annual longline survey conducted cooperatively by the Fisheries Agency of Japan and the Alaska Fisheries Science Center of the National Marine Fisheries Service) were updated and corrected for discovered sampling bias. In addition, more recent, randomly collected samples (1996–2004 data from the annual longline survey conducted by the Alaska Fisheries Science Center) were analyzed and new length-at-age and weight-at-age parameters were estimated. Results were similar between this analysis with length-at-age data from 1981 to 2004 and analysis with updated longline survey data through 2010; therefore, we used our initial results from analysis done with data through 2004. We found that, because of a stratified sampling scheme, growth estimates of sablefish were overestimated with the older data (1981–93), and growth parameters used in the Alaskan sablefish assessment model were, thus, too large. In addition, a comparison of the bias-corrected 1981–93 data and the 1996–2004 data showed that, in more recent years, sablefish grew larger and growth differed among regions. The updated growth information improves the fit of the data to the sablefish stock assessment model with biologically reasonable results. These findings indicate that when the updated growth data (1996–2004) are used in the existing sablefish assessment model, estimates of fishing mortality increase slightly and estimates of female spawning biomass decrease slightly. This study provides evidence of the importance of periodically revisiting biological parameter estimates, especially as data accumulate, because the addition of more recent data often will be more biologically realistic. In addition, it exemplifies the importance of correcting biases from sampling that may contribute to erroneous parameter estimates.

Abundance of adult horseshoe crabs (Limulus polylphemus) in Delaware Bay estimated from a bay-wide mark-recapture study

Estimates of the abundance of American horseshoe crabs (Limulus polyphemus) are important to determine egg production and to manage populations for the energetic needs of shorebirds that feed on horseshoe crab eggs. In 2003, over 17,500 horseshoe crabs were tagged and released throughout Delaware Bay, and recaptured crabs came from spawning surveys that were conducted during peak spawning. We used two release cohorts to test for a temporary effect of tagging on spawning behavior and we adjusted the number of releases according to relocation rates from a telemetry study. The abundance estimate was 20 million horseshoe crabs (90 % confidence interval: 13−28 million), of which 6.25 million (90% CI: 4.0−8.8 million) were females. The combined harvest rate for Delaware, New Jersey, Virginia, and Maryland in 2003 was 4% (90% CI: 3−6%) of the abundance estimate. Over-wintering of adults in Delaware Bay could explain, in part, differences in estimates from ocean-trawl surveys. Based on fecundity of 88,000 eggs per female, egg production was 5.5×1011 (90% CI: 3.5×1011, 7.7×1011), but egg availability for shorebirds also depended on overlap between horseshoe crab and shorebird migrations, density-dependent bioturbation, and wave-mediated vertical transport.

Hatching date, nursery grounds, and early growth of juvenile walleye pollock (Theragra chalcogramma) off northern Japan*

Walleye pollock (Theragra chalcogramma) is widely distributed in the North Pacific Ocean and plays an important role in coastal subarctic ecosystems. The Japanese Pacific population of this species is one of the most important demersal fishes for commercial fisheries in northern Japan. The population is distributed along the Pacific coast of Hokkaido and the Tohoku area (Fig. 1), which is the southern limit of distribution of the species in the western North Pacific. In Funka Bay, the main spawning ground for this population, pollock spawn from December to March (Kendall and Nakatani, 1992). Planktonic eggs and larvae are transported into the bay, where juveniles usually remain until late July when they reach 60−85 mm in total length (Hayashi et al., 1968; Nakatani and Maeda, 1987). These juvenile pollock then migrate from Funka Bay eastward to the Doto area off southeastern Hokkaido (Honda et al., 2004). Many studies on eggs, larvae, and juveniles of the species have been conducted in or near Funka Bay, but little information is available on the ecology of the early life stages in the Tohoku area. Hashimoto and Ishito (1991) suggested that eggs are transported from Funka Bay southward to the Tohoku area by the coastal branch of the Oyashio Current, but there has been no study to verify this hypothesis.

The fishery for California market squid (Loligo opalescens) (Cephalopoda: Myopsida), from 1981 through 2003

The California market squid (Loligo opalescens) has been harvested since the 1860s and it has become the largest fishery in California in terms of tonnage and dollars since 1993. The fishery began in Monterey Bay and then shifted to southern California, where effort has increased steadily since 1983. The California Department of Fish and Game (CDFG) collects information on landings of squid, including tonnage, location, and date of capture. We compared landings data gathered by CDFG with sea surface temperature (SST), upwelling index (UI), the southern oscillation index (SOI), and their respective anomalies. We found that the squid fishery in Monterey Bay expends twice the effort of that in southern California. Squid landings decreased substantially following large El Niño events in 1982−83 and 1997−98, but not following the smaller El Niño events of 1987 and 1992. Spectral analysis revealed autocorrelation at annual and 4.5-year intervals (similar to the time period between El Niño cycles). But this analysis did not reveal any fortnightly or monthly spawning peaks, thus squid spawning did not correlate with tides. A paralarvae density index (PDI) for February correlated well with catch per unit of effort (CPUE) for the following November recruitment of adults to the spawning grounds. This stock– recruitment analysis was significant for 2000−03 (CPUE=8.42+0.41PDI, adjusted coefficient of determination, r2=0.978, P=0.0074). Surveys of squid paralarvae explained 97.8% of the variance for catches of adult squid nine months later. The regression of CPUE on PDI could be used to manage the fishery. Catch limits for the fishery could be set on the basis of paralarvae abundance surveyed nine months earlier.

Variability in supply and cross-shelf transport of pink shrimp (Farfantepenaeus duorarum) postlarvae into western Florida Bay

The variability in the supply of pink shrimp (Farfantepenaeus duorarum) postlarvae and the transport mechanisms of planktonic stages were investigated with field data and simulations of transport. Postlarvae entering the nursery grounds of Florida Bay were collected for three consecutive years at channels that connect the Bay with the Gulf of Mexico, and in channels of the Middle Florida Keys that connect the southeastern margin of the Bay with the Atlantic Ocean. The influx of postlarvae in the Middle Florida Keys was low in magnitude and varied seasonally and among years. In contrast, the greater postlarval influx occurred at the northwestern border of the Bay, where there was a strong seasonal pattern with peaks in influx from July through September each year. Planktonic stages need to travel up to 150 km eastward between spawning grounds (northeast of Dry Tortugas) and nursery grounds (western Florida Bay) in about 30 days, the estimated time of planktonic development for this species. A Lagrangian trajectory model was developed to estimate the drift of planktonic stages across the SW Florida shelf. The model simulated the maximal distance traveled by planktonic stages under various assumptions of behavior. Simulation results indicated that larvae traveling with the instantaneous current and exhibiting a diel behavior travel up to 65 km and 75% of the larvae travel only 30 km. However, the eastward distance traveled increased substantially when a larval response to tides was added to the behavioral variable (distance increased to 200 km and 85% of larvae traveled 150 km). The question is, when during larval development, and where on the shallow SW Florida shelf, does the tidal response become incorporated into the behavior of pink shrimp.

Biology and assessment of the painted sweetlips (Diagramma pictum (Thunberg, 1792)) and the spangled emperor (Lethrinus nebulosus (Forsskål, 1775)) in the southern Arabian Gulf

The population biology and status of the painted sweeplips (Diagramma pictum) and spangled emperor (Lethrinus nebulosus) in the southern Arabian Gulf were established by using a combination of size-frequency, biological, and size-at-age data. Transverse sections of sagittal otoliths were characterized by alternating translucent and opaque bands that were validated as annuli. Comparisons of growth characteristics showed that there were no significant differences (P>0.05) between sexes. There were well defined peaks in the reproductive cycle, spawning occurred from April to May for both species, and the mean size at which females attained sexual maturity was 31.8 cm fork length (LF) for D. pictum and 27.6 cm (LF) for L. nebulosus. The mean sizes at first capture (21.1 cm LF for D. pictum and 26.4 cm LF for L. nebulosus) were smaller than the sizes for both at first sexual maturity and those at which yield per recruit would be maximized. The range of fishing-induced mortality rates for D. pictum (0.37−0.62/yr) was substantially greater than the target (Fopt=0.07/yr) and limit (Flimit=0.09/ yr) estimates. The range of fishing-induced mortality rates for L. nebulosus (0.15/yr to 0.57/yr) was also in excess of biological reference points (Fopt=0.10/yr and Flimit=0.13/yr). In addition to growth overfishing, the stocks were considered to be recruitment overfished because the biomass per recruit was less than 20% of the unexploited levels for both species. The results of the study are important to fisheries management authorities in the region because they indicate that both a reduction in fishing effort and mesh-size regulations are required for the demersal trap fishery.

A catch-free stock assessment model with application to goliath grouper (Epinephelus itajara) off southern Florida

Many modern stock assessment methods provide the machinery for determining the status of a stock in relation to certain reference points and for estimating how quickly a stock can be rebuilt. However, these methods typically require catch data, which are not always available. We introduce a model-based framework for estimating reference points, stock status, and recovery times in situations where catch data and other measures of absolute abundance are unavailable. The specif ic estimator developed is essentially an age-structured production model recast in terms relative to pre-exploitation levels. A Bayesian estimation scheme is adopted to allow the incorporation of pertinent auxiliary information such as might be obtained from meta-analyses of similar stocks or anecdotal observations. The approach is applied to the population of goliath grouper (Epinephelus itajara) off southern Florida, for which there are three indices of relative abundance but no reliable catch data. The results confirm anecdotal accounts of a marked decline in abundance during the 1980s followed by a substantial increase after the harvest of goliath grouper was banned in 1990. The ban appears to have reduced fishing pressure to between 10% and 50% of the levels observed during the 1980s. Nevertheless, the predicted fishing mortality rate under the ban appears to remain substantial, perhaps owing to illegal harvest and depth-related release mortality. As a result, the base model predicts that there is less than a 40% chance that the spawning biomass will recover to a level that would produce a 50% spawning potential ratio.