The use of parasites in discriminating stocks of Pacific halibut (Hippoglossus stenolepis) in the northeast Pacific

The use of parasites as indicators of the stock structure of Pacific halibut (Hippoglossus stenolepis) in the northeast Pacific was investigated by using 328 adult (>55 cm fork length) halibut from 15 composite localities ranging from northern California to the northern Bering Sea and 96 juvenile (10–55 cm) halibut from five localities ranging from the northern Queen Charlotte Islands to the Bering Sea. Counts of eight selected parasite species (the juvenile acanthocephalans Corynosoma strumosum and C. villosum, the metacestode Nybelinia surmenicola, the digenean metacercaria Otodistomum sp., and the larval nematodes Anisakis simplex, Pseudoterranova decipiens, Contracaecum sp., and Spirurid gen. sp.) that produce infections of long duration, do not multiply in the host, and that have a relatively high abundance in at least one geographic locality were subjected to discriminant function analysis. Juvenile Pacific halibut showed no separation and, even though they were not heavily infected with parasites, the analysis suggested that juveniles could be a mixed stock. Three groups of adults were identified: fish from California to the southern Queen Charlotte Islands, those from the northern Queen Charlotte Islands to the central Bering Sea, and those from the central and north-ern Bering Sea. These groups suggest that the single stock concept be more thoroughly evaluated.

Small-scale spatial and temporal variability in growth and mortality of fish larvae in the subtropical northcentral Gulf of Mexico: implications for assessing recruitment success

Extensive plankton collections were taken during seven September cruises (1990–93) along the inner continental shelf of the northcentral Gulf of Mexico (GOM). Despite the high productivity and availability of food during these cruises, significant small-scale spatial variability was found in larval growth rates for both Atlantic bumper (Chloroscombrus chrysurus, Carangidae) and vermilion snapper (Rhomboplites aurorubens, Lutjanidae). The observed variability in larval growth rates was not correlated with changes in water temperature or associated with conspicuous hydrographic features and suggested the existence of less-recognizable regions where conditions for growth vary. Cruise estimates of mortality coefficients (Z) for larval Atlantic bumper (n=32,241 larvae from six cruises) and vermilion snapper (n= 2581 larvae from four cruises) ranged from 0.20 to 0.37 and 0.19 to 0.29, respectively. Even in a subtropical climate like the GOM, where larval-stage durations may be as short as two weeks, observed variability in growth rates, particularly when combined with small changes in mortality rates, can cause order-of-magnitude differences in cumulative larval survival. To what extent the observed differences in growth rates at small spatial scales are fine-scale “noise” that ultimately is smoothed by larger-scale processes is not known. Future research is needed to further characterize the small-scale variability in growth rates of larvae, particularly with regard to microzooplankton patchiness and the temporal and spatial pattern of potential predators. Small-scale spatial variability in larval growth rates may in fact be the norm, and understanding the implications of this subtle mosaic may help us to better evaluate our ability to partition the causes of recruitment variability.

Temporal changes in population density, fecundity, and egg size of the Hawaiian spiny lobster (Panulirus marginatus) at Necker Bank, Northwestern Hawaiian Islands

Fecundity (F, number of brooded eggs) and egg size were estimated for Hawaiian spiny lobster (Panulirus marginatus) at Necker Bank, North-western Hawaiian Islands (NWHI), in June 1999, and compared with previous (1978–81, 1991) estimates. Fecundity in 1999 was best described by the power equations F = 7.995 CL 2.4017, where CL is carapace length in mm (r2=0.900), and F = 5.174 TW 2.758, where TW is tail width in mm (r2=0.889) (both n=40; P< 0.001). Based on a log-linear model ANCOVA, size-specific fecundity in 1999 was 18% greater than in 1991, which in turn was 16% greater than during 1978–81. The additional increase in size-specific fecundity observed in 1999 is interpreted as evidence for further compensatory response to decreased lobster densities and increased per capita food resources that have resulted either from natural cyclic declines in productivity, high levels of harvest by the commercial lobster trap fishery, or both.

Impact of hatchery releases on the recreational fishery for Pacific threadfin (Polydactylus sexfilis) in Hawaii

The Pacific threadfin (Polydactylus sexfilis) is considered one of the premier Hawaiian food fishes but even with catch limits, seasonal closures, and size limits, catches have declined dramatically since the 1960s. It was identified as the top candidate species for stock enhancement in Hawaii, based on the decline in stocks, high market value, and importance of the fishery. In the stock enhancement program for Pacific threadfin, over 430,000 fingerlings of various sizes were implanted with coded wire tags and released in nursery habitats along the windward coast of Oahu between 1993 and 1998. Because few Pacific threadfin were present in creel surveys conducted between 1994 and 1998, Oahu fishermen were offered a $10 reward for each threadfin that was caught (for both hatchery-reared and wild fish). A total of 1882 Pacific threadfin were recovered from the reward program between March 1998 and May 1999, including 163 hatchery-reared fish, an overall contribution of 8.7% to the fishery. Hatchery-reared fish accounted for as high as 71% of returns in the release areas. Hatchery-reared fish were recovered on average 11.5 km (SD=9.8 km) from the release site, although some had moved as far away as 42 km. Average age for recovered hatchery-reared fish was 495 days; the oldest was 1021 days. Cultured Pacific threadfin juveniles survived and recruited successfully to the recreational fishery, accounting for 10% of fishermen’s catches on the windward side of Oahu. Recruitment to the fishery was highest for the 1997 release year; few juveniles from earlier releases were observed. Presence of a few large, fully developed females in the recreational fishery suggested that hatchery-reared fish can survive, grow, and reproductively contribute to the population. Implementation of an enhancement program that is focused on juveniles and perhaps large females, as part of an integrated fishery management strategy, could speed the recovery of this fish population.

Yield- and biomass-per-recruit analysis for rotational fisheries, with an application to the Atlantic sea scallop (Placopecten magellanicus)

A general model for yield-per-recruit analysis of rotational (periodic) fisheries is developed and applied to the sea scallop (Placopecten magellanicus) fishery of the northwest Atlantic. Rotational fishing slightly increases both yield- and biomass-per-recruit for sea scallops at FMAX. These quantities decline less quickly when fishing mortality is increased beyond FMAX than when fishing is at a constant rate. The improvement in biomass-per-recruit appears to be nearly independent of the selectivity pattern but increased size-at-entry can reduce or eliminate the yield-per-recruit advantage of rotation. Area closures and rotational fishing can cause difficulties with the use of standard spatially averaged fishing mortality metrics and reference points. The concept of temporally averaged fishing mortality is introduced as one that is more appropriate for sedentary resources when fishing mortality varies in time and space.

Estimating long-term growth-rate changes of southern bluefin tuna (Thunnus maccoyii) from two periods of tag-return data

Southern bluefin tuna (SBT) (Thunnus maccoyii) growth rates are estimated from tag-return data associated with two time periods, the 1960s and 1980s. The traditional von Bertalanffy growth model (VBG) and a two-phase VBG model were fitted to the data by maximum likelihood. The traditional VBG model did not provide an adequate representation of growth in SBT, and the two-phase VBG yielded a significantly better fit. The results indicated that significant change occurs in the pattern of growth in relation to a VBG curve during the juvenile stages of the SBT life cycle, which may be related to the transition from a tightly schooling fish that spends substantial time in near and surface shore waters to one that is found primarily in more offshore and deeper waters. The results suggest that more complex growth models should be considered for other tunas and for other species that show a marked change in habitat use with age. The likelihood surface for the two-phase VBG model was found to be bimodal and some implications of this are investigated. Significant and substantial differences were found in the growth for fish spawned in the 1960s and in the 1980s, such that after age four there is a difference of about one year in the expected age of a fish of similar length which persists over the size range for which meaningful recapture data are available. This difference may be a density-dependent response as a consequence of the marked reduction in the SBT population. Given the key role that estimates of growth have in most stock assessments, the results indicate that there is a need both for the regular monitoring of growth rates and for provisions for changes in growth over time (possibly related to changes in abundance) in the stock assessment models used for SBT and other species.

Life history of the Atlantic sharpnose shark (Rhizoprionodon terraenovae) (Richardson, 1836) off the southeastern United States

The life history of the Atlantic sharpnose shark (Rhizoprionodon terraenovae) was described from 1093 specimens collected from Virginia to northern Florida between April 1997 and March 1999. Longitudinally sectioned vertebral centra were used to age each specimen, and the periodicity of circuli deposition was verified through marginal increment analysis and focus-to-increment frequency distributions. Rhizoprionodon terraenovae reached a maximum size of 828 mm precaudal length (PCL) and a maximum age of 11+ years. Mean back-calculated lengths-at-age ranged from 445 mm PCL at age one to 785 mm PCL at age ten for females, and 448 mm PCL at age one to 747 mm PCL at age nine for males. Observed lengthat-age data (estimated to 0.1 year) yielded the following von Bertalanffy parameters estimates: L∞= 749 mm PCL (SE=4.60), K = 0.49 (SE=0.020), and t0= –0.94 (SE=0.046) for females; and L∞= 745 mm PCL (SE = 5.93), K = 0.50 (SE=0.024), and t0= –0.91 (SE = 0.052) for males. Sexual maturity was reached at age three and 611 mm PCL for females, and age three and 615 mm PCL for males. Rhizoprionodon terraenovae reproduced annually and had a gestation period of approximately 11 months. Litter size ranged from one to eight (mean=3.85) embyros, and increased with female PCL.

A general framework for integrating environmental time series into stock assessment models: model description, simulation testing, and example

We present a method to integrate environmental time series into stock assessment models and to test the significance of correlations between population processes and the environmental time series. Parameters that relate the environmental time series to population processes are included in the stock assessment model, and likelihood ratio tests are used to determine if the parameters improve the fit to the data significantly. Two approaches are considered to integrate the environmental relationship. In the environmental model, the population dynamics process (e.g. recruitment) is proportional to the environmental variable, whereas in the environmental model with process error it is proportional to the environmental variable, but the model allows an additional temporal variation (process error) constrained by a log-normal distribution. The methods are tested by using simulation analysis and compared to the traditional method of correlating model estimates with environmental variables outside the estimation procedure. In the traditional method, the estimates of recruitment were provided by a model that allowed the recruitment only to have a temporal variation constrained by a log-normal distribution. We illustrate the methods by applying them to test the statistical significance of the correlation between sea-surface temperature (SST) and recruitment to the snapper (Pagrus auratus) stock in the Hauraki Gulf–Bay of Plenty, New Zealand. Simulation analyses indicated that the integrated approach with additional process error is superior to the traditional method of correlating model estimates with environmental variables outside the estimation procedure. The results suggest that, for the snapper stock, recruitment is positively correlated with SST at the time of spawning.

Seasonal distribution, abundance, and growth of young-of-the-year Atlantic croaker (Micropogonias undulatus) in Delaware Bay and adjacent marshes

We examined the spatial and temporal distribution, abundance, and growth of young-of-the-year (YOY) Atlantic croaker (Micropogonias undulatus) in Delaware Bay, one of the northernmost estuaries in which they consistently occur along the east coast of the United States. Sampling in Delaware Bay and in tidal creeks in salt marshes adjacent to the bay with otter trawls, plankton nets and weirs, between April and November 1996–99, collected approximately 85,000 YOY. Ingress of each year class into the bay and tidal creeks consistently occurred in the fall, and the first few YOY appeared in August. Larvae as small as 2–3 mm TL were collected in September and October 1996. Epibenthic individuals <25 mm TL were present each fall and again during spring of each year, but not in 1996 when low water temperatures in January and February apparently caused widespread mortality, resulting in their absence the following spring and summer. In 1998 and 1999, a second size class of smaller YOY entered the bay and tidal creeks in June. When YOY survived the winter, there was no evidence of growth until after April. Then the YOY grew rapidly through the summer in all habitats (0.8–1.4 mm/d from May through August). In the bay, they were most abundant from June to August over mud sediments in oligohaline waters. They were present in both subtidal and intertidal creeks in the marshes where they were most abundant from April to June in the mesohaline portion of the lower bay. The larger YOY began egressing out of the marshes in late summer, and the entire year class left the tidal creeks at lengths of 100–200 mm TL by October or November when the next year class was ingressing. These patterns of seasonal distribution and abundance in Delaware Bay and the adjacent marshes are similar to those observed in more southern estuaries along the east coast; however, growth is faster—in keeping with that in other northern estuaries.

Age validation, growth, mortality, and additional population parameters of the goldband snapper (Pristipomoides multidens) off the Kimberley coast of northwestern Australia

Goldband snapper (Pristipomoides multidens) collected from commercial trap and line fishermen off the Kimberley coast of northwestern Australia were aged by examination of sectioned otoliths (sagittae).A total of 3833 P. multidens, 80–701 mm fork length (98–805 mm total length), were examined from commercial catches from 1995 to 1999. The oldest fish was estimated to be age 30+ years. Validation of age estimates was achieved with marginal increment analysis. The opaque and translucent zones were each formed once per year and are considered valid annual growth increments (the translucent zone was formed once per year between January and May). A strong link between water temperature and translucent zone formation was evident in P. multidens. The von Bertalanffy growth function was used to describe growth from length-at-age data derived from sectioned otoliths.

An approach to estimating rockfish biomass based on larval production, with application to Sebastes jordani

An assessment of the total biomass of shortbelly rockfish (Sebastes jordani) off the central California coast is presented that is based on a spatially extensive but temporally restricted ichthyoplankton survey conducted during the 1991 spawning season. Contemporaneous samples of adults were obtained by trawl sampling in the study region. Daily larval production (7.56 × 1010 larvae/d) and the larval mortality rate (Z=0.11/d) during the cruise were estimated from a larval “catch curve,” wherein the logarithm of total age-specific larval abundance was regressed against larval age. For this analysis, larval age compositions at each of the 150 sample sites were determined by examination of otolith microstructure from subsampled larvae (n=2203), which were weighted by the polygonal Sette-Ahlstrom area surrounding each station. Female population weight-specific fecundity was estimated through a life table analysis that incorporated sex-specific differences in adult growth rate, female maturity, fecundity, and natural mortality (M). The resulting statistic (102.17 larvae/g) was insensitive to errors in estimating M and to the pattern of recruitment. Together, the two analyses indicated that a total biomass equal to 1366 metric tons (t)/d of age-1+ shortbelly rockfish (sexes combined) was needed to account for the observed level of spawning output during the cruise. Given the long-term seasonal distribution of spawning activity in the study area, as elucidated from a retrospective examination of California Cooperative Oceanic Fisheries Investigation (CalCOFI) ichthyoplankton samples from 1952 to 1984, the “daily” total biomass was expanded to an annual total of 67,392 t. An attempt to account for all sources of error in the derivation of this estimate was made by application of the delta-method, which yielded a coefficient of variation of 19%. The relatively high precision of this larval production method, and the rapidity with which an absolute biomass estimate can be obtained, establishes that, for some species of rockfish (Sebastes spp.), it is an attractive alternative to traditional age-structured stock assessments.

Prey consumption of Steller sea lions (Eumetopias jubatus) off Alaska: How much prey do they require?

The effects of seasonal and regional differences in diet composition on the food requirements of Steller sea lions (Eumetopias jubatus) were estimated by using a bioenergetic model. The model considered differences in the energy density of the prey, and differences in digestive efficiency and the heat increment of feeding of different diets. The model predicted that Steller sea lions in southeast Alaska required 45–60% more food per day in early spring (March) than after the breeding season in late summer (August) because of seasonal changes in the energy density of the diets (along with seasonal changes in energy requirements). The southeast Alaska population, at 23,000 (±1660 SD) animals (all ages), consumed an estimated 140,000 (±27,800) t of prey in 1998. In contrast, we estimated that the 51,000 (±3680) animals making up the western Alaska population in the Gulf of Alaska and Aleutian Islands consumed just over twice this amount (303,000 [±57,500] t). In terms of biomass removed in 1998 from Alaskan waters, we estimated that Steller sea lions accounted for about 5% of the natural mortality of gadids (pollock and cod) and up to 75% of the natural mortality of hexagrammids (adult Atka mackerel). These two groups of species were consumed in higher amounts than any other. The predicted average daily food requirement per individual ranged from 16 (±2.8) to 20 (±3.6) kg (all ages combined). Per capita food requirements differed by as much as 24% between regions of Alaska depending on the relative amounts of low–energy-density prey (e.g. gadids) versus high–energy-density prey (e.g. forage fish and salmon) consumed. Estimated requirements were highest in regions where Steller sea lions consumed higher proportions of low–energy-density prey and experienced the highest rates of population decline