Using productivity and susceptibility indices to assess the vulnerability of United States fish stocks to overfishing

Assessing the vulnerability of stocks to fishing practices in U.S. federal waters was recently highlighted by the National Marine Fisheries Service (NMFS), National Oceanic and Atmospheric Administration, as an important factor to consider when 1) identifying stocks that should be managed and protected under a fishery management plan; 2) grouping data-poor stocks into relevant management complexes; and 3) developing precautionary harvest control rules. To assist the regional fishery management councils in determining vulnerability, NMFS elected to use a modified version of a productivity and susceptibility analysis (PSA) because it can be based on qualitative data, has a history of use in other fisheries, and is recommended by several organizations as a reasonable approach for evaluating risk. A number of productivity and susceptibility attributes for a stock are used in a PSA and from these attributes, index scores and measures of uncertainty are computed and graphically displayed. To demonstrate the utility of the resulting vulnerability evaluation, we evaluated six U.S. fisheries targeting 162 stocks that exhibited varying degrees of productivity and susceptibility, and for which data quality varied. Overall, the PSA was capable of differentiating the vulnerability of stocks along the gradient of susceptibility and productivity indices, although fixed thresholds separating low-, moderate-, and highly vulnerable species were not observed. The PSA can be used as a flexible tool that can incorporate regional-specific information on fishery and management activity.

Diet composition and prey selection of the introduced grouper species peacock hind (Cephalopholis argus) in Hawaii

The introduced grouper species peacock hind (Cephalopholis argus), was the dominant large-body piscivore on the Main Hawaiian Island (MHI) reefs assessed by underwater visual surveys in this study. However, published data on C. argus feeding ecology are scarce, and the role of this species in Hawaiian reef ecosystems is presently not well understood. Here we provide the first comprehensive assessment of the diet composition, prey electivity (dietary importance of prey taxa compared to their availability on reefs), and size selectivity (prey sizes in the diet compared to sizes on reefs) of this important predator in the MHI. Diet consisted 97.7% of fishes and was characterized by a wide taxonomic breadth. Surprisingly, feeding was not opportunistic, as indicated by a strongly divergent electivity for different prey fishes. In addition, whereas some families of large-body species were represented in the diet exclusively by recruit-size individuals (e.g., Aulostomidae), several families of smaller-body species were also represented by juveniles or adults (e.g., Chaetodontidae). Both the strength and mechanisms of the effects of C. argus predation are therefore likely to differ among prey families. This study provides the basis for a quantitative estimate of prey consumption by C. argus, which would further increase understanding of impacts of this species on native fishes in Hawaii.

Abundance, condition, and diet of juvenile Pacific ocean perch (Sebastes alutus) in the Aleutian Islands

T he relative value of pelagic habitat for three size classes of juvenile Pacific ocean perch (Sebastes alutus) was investigated by comparing their abundance and condition in two areas of the Aleutian Islands. Diet, zooplankton biomass, and water column temperatures were examined as potential factors affecting observed differences. Juvenile Pacific ocean perch abundance and condition, and zooplankton biomass varied significantly between areas, whereas juvenile Pacific ocean perch diet varied only by size class. Observed differences in fish condition may have been due to the quantity or quality of pelagic prey items consumed. For the delineation of essential demersal fish habitat, important ecological features of the pelagic habitat must therefore be considered.

The repulsive and feeding-deterrent effects of electropositive metals on juvenile sandbar sharks (Carcharhinus plumbeus)

Reducing shark bycatch and depredation (i.e., damage caused by sharks to gear, bait, and desired fish species) in pelagic longline fisheries targeting tunas and swordfish is a priority. Electropositive metals (i.e., a mixture of the lanthanide elements lanthanum, cerium, neodymium, and praseodymium) have been shown to deter spiny dogfish (Squalus acanthias, primarily a coastal species) from attacking bait, presumably because of interactions with the electroreceptive system of this shark. We undertook to determine the possible effectiveness of electropositive metals for reducing the interactions of pelagic sharks with longline gear, using sandbar sharks (Carcharhinus plumbeus, family Carcharhinidae) as a model species. The presence of electropositive metal deterred feeding in groups of juvenile sandbar sharks and altered the swimming patterns of individuals in the absence of food motivation (these individuals generally avoided approaching electropositive metal closer than ~100 cm). The former effect was relatively short-lived however; primarily (we assume) because competition with other individuals increased feeding motivation. In field trials with bottom longline gear, electropositive metal placed within ~10 cm of the hooks reduced the catch of sandbar sharks by approximately two thirds, compared to the catch on hooks in the proximity of plastic pieces of similar dimensions. Electropositive metals therefore appear to have the potential to reduce shark interactions in pelagic longline fisheries, although the optimal mass, shape, composition, and distance to baited hooks remain to be determined.

Recalculated diet and daily ration of the shortfin mako (Isurus oxyrinchus), with a focus on quantifying predation on bluefish (Pomatomus saltatrix) in the northwest Atlantic Ocean

The diet and daily ration of the shortfin mako (Isurus oxyrinchus) in the northwest Atlantic were re-examined to determine whether fluctuations in prey abundance and availability are reflected in these two biological variables. During the summers of 2001 and 2002, stomach content data were collected from fishing tournaments along the northeast coast of the United States. These data were quantified by using four diet indices and were compared to index calculations from historical diet data collected from 1972 through 1983. Bluefish (Pomatomus saltatrix) were the predominant prey in the 1972–83 and 2001–02 diets, accounting for 92.6% of the current diet by weight and 86.9% of the historical diet by volume. From the 2001– 02 diet data, daily ration was estimated and it indicated that shortfin makos must consume roughly 4.6% of their body weight per day to fulfill energetic demands. The daily energetic requirement was broken down by using a calculated energy content for the current diet of 4909 KJ/kg. Based on the proportional energy of bluefish in the diet by weight, an average shortfin mako consumes roughly 500 kg of bluefish per year off the northeast coast of the United States. The results are discussed in relation to the potential effect of intense shortfin mako predation on bluefish abundance in the region.

Seasonal, diel, and lunar spawning periodicities and associated sound production of white seabass (Atractoscion nobilis)

Spawning periodicities of white seabass (Atractoscion nobilis) were evaluated by observing spawning behavior, by collecting eggs, and monitoring recognizable sounds produced during the release of gametes. A total of 297 spawning events were documented from 15 male and 47 female white seabass contained within the seminatural confines of a 526-m3 net pen located in Catalina Harbor, Santa Catalina Island, California. Consistent spawning occurred from March through July 2001−03, and peaked in May at a photoperiod of 14 hours. Most spawning occurred within the 2-hour period following sunset or from 19:00−20:00 hours Pacific Standard Time. White seabass spawned at every phase of the lunar cycle; but an increase in successive spawning events followed the new moon. Most spawning occurred in water temperatures from 15 to 18°C, and there was no apparent correlation with tidal cycles. Seasonal and diel spawning periods were directly correlated with increases in the rate, intensity, and variety of white seabass sounds; this correlation may indicate that sounds function to enhance reproductive success. These findings can be extended to further develop seasonal fishery regulations and to better comprehend the role of sound in the reproduction of sound-producing fishes.

Age, growth, and reproduction of dolphinfish (Coryphaena hippurus) caught off the coast of North Carolina

Age, growth, and reproductive data were obtained from dolphinfish (Coryphaena hippurus, size range: 89 to 1451 mm fork length [FL]) collected between May 2002 and May 2004 off North Carolina. Annual increments from scales (n=541) and daily increments from sagittal otoliths (n=107) were examined; estimated von Bertalanffy parameters were L∞ (asymptotic length)=1299 mm FL and k (growth coefficient)=1.08/yr. Daily growth increments reduced much of the residual error in length-at-age estimates for age-0 dolphinfish; the estimated average growth rate was 3.78 mm/day during the first six months. Size at 50% maturity was slightly smaller for female (460 mm FL) than male (475 mm FL) dolphinfish. Based on monthly length-adjusted gonad weights, peak spawning occurs from April through July off North Carolina; back-calculated hatching dates from age-0 dolphinfish and prior reproductive studies on the east coast of Florida indicate that dolphinfish spawning occurs year round off the U.S. east coast and highest levels range from January through June. No major changes in length-at-age or size-at-maturity have occurred since the early 1960s, even after substantial increases in fishery landings.

Diets of and trophic relationships among dominant marine nekton within the northern California Current ecosystem

In this study we analyzed the diets of 26 nekton species collected from two years (2000 and 2002) off Oregon and northern California to describe dominant nekton trophic groups of the northern California Current (NCC) pelagic ecosystem. We also examined interannual variation in the diets of three nekton species. Cluster analysis of predator diets resulted in nekton trophic groups based on the consumption of copepods, euphausiids, brachyuran larvae, larval juvenile fishes, and adult nekton. However, many fish within trophic groups consumed prey from multiple trophic levels—euphausiids being the most widely consumed. Comparison of diets between years showed that most variation occurred with changes in the contribution of euphausiids and brachyuran larvae to nekton diets. The importance of euphausiids and other crustacean prey to nekton indicates that omnivory is an important characteristic of the NCC food web; however it may change during periods of lower or higher upwelling and ecosystem production.

Abundance, distribution, and habitat of leatherback turtles (Dermochelys coriacea) off California, 1990−2003

Leatherback turtles (Dermochelys coriacea) are regularly seen off the U.S. West Coast, where they forage on jellyfish (Scyphomedusae) during summer and fall. Aerial line-transect surveys were conducted in neritic waters (<92 m depth) off central and northern California during 1990−2003, providing the first foraging population estimates for Pacific leatherback turtles. Males and females of about 1.1 to 2.1 m length were observed. Estimated abundance was linked to the Northern Oscillation Index and ranged from 12 (coefficient of variation [CV] =0.75) in 1995 to 379 (CV= 0.23) in 1990, averaging 178 (CV= 0.15). Greatest densities were found off central California, where oceanographic retention areas or upwelling shadows created favorable habitat for leatherback turtle prey. Results from independent telemetry studies have linked leatherback turtles off the U.S. West Coast to one of the two largest remaining Pacific breeding populations, at Jamursba Medi, Indonesia. Nearshore waters off California thus represent an important foraging region for the critically endangered Pacific leatherback turtle.

Ecotypic variation and predatory behavior among killer whales (Orcinus orca) off the eastern Aleutian Islands, Alaska

From 2001 to 2004 in the eastern Aleutian Islands, Alaska, killer whales (Orcinus orca) were encountered 250 times during 421 days of surveys that covered a total of 22,491 miles. Three killer whale groups (resident, transient, and offshore) were identified acoustically and genetically. Resident killer whales were found 12 times more frequently than transient killer whales, and offshore killer whales were encountered only once. A minimum of 901 photographically identified resident whales used the region during our study. A total of 165 mammal-eating transient killer whales were identified, and the majority (70%) were encountered during spring (May and June). The diet of transient killer whales in spring was primarily gray whales (Eschrichtius robustus), and in summer primarily northern fur seals (Callorhinus ursinus). Steller sea lions (Eumetopias jubatus) did not appear to be a preferred prey or major prey item during spring and summer. The majority of killer whales in the eastern Aleutian Islands are the resident ecotype, which does not consume marine mammals.

Precision estimates and suggested sample sizes for length-frequency data

For most fisheries applications, the shape of a length-frequency distribution is much more important than its mean length or variance. This makes it difficult to evaluate at which point a sample size is adequate. By estimating the coefficient of variation of the counts in each length class and taking a weighted mean of these, a measure of precision was obtained that takes the precision in all length classes into account. The precision estimates were closely associated with the ratio of the sample size to the number of size classes in each sample. As a rule-of-thumb, a minimum sample size of 10 times the number of length classes in the sample is suggested because the precision deteriorates rapidly for smaller sample sizes. In absence of such a rule-of-thumb, samplers have previously under-estimated the required sample size for samples with large fish, while over-sampling small fish of the same 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.

Estimating species and size composition of rockfishes to verify targets in acoustic surveys of untrawlable areas

Rockfish (Sebastes spp.) biomass is difficult to assess with standard bottom trawl or acoustic surveys because of their propensity to aggregate near the seafloor in highrelief areas that are inaccessible to sampling by trawling. We compared the ability of a remotely operated vehicle (ROV), a modified bottom trawl, and a stereo drop camera system (SDC) to identify rockfish species and estimate their size composition. The ability to discriminate species was highest for the bottom trawl and lowest for the SDC. Mean lengths and size distributions varied among the gear types, although a larger number of length measurements could be collected with the bottom trawl and SDC than with the ROV. Dusky (S. variabilis), harlequin (S. variegatus), and northern rockfish (S. polyspinis), and Pacific ocean perch (S. alutus) were the species observed in greatest abundance. Only dusky and northern rockfish regularly occurred in trawlable areas, whereas these two species and many more occurred in untrawlable areas. The SDC was able to resolve the height of fish off the seafloor, and some of the rockfish species were observed only near the seafloor in the acoustic dead zone. This finding is important, in that fish found exclusively in the acoustic dead zone cannot be assessed acoustically. For these species, methods such as bottom trawls, long-lines, or optical surveys using line transect or area swept methods will be the only adequate means to estimate the abundance of these fishes. Our results suggest that the selection of appropriate methods for verifying targets will depend on the habitat types and species complexes to be examined.

Prevalence, intensity, and effect of a nematode (Philometra saltatrix) in the ovaries of bluefish (Pomatomus saltatrix)

Examination of 203 adult bluefish (Pomatomus saltatrix) from Long Island, New York, in 2002 and 2003 and 66 from the Outer Banks, North Carolina, in 2003 revealed the presence of dracunculoid nematodes (Philometra saltatrix) in the ovaries of female fish. Percent prevalence reached 88% in July and then decreased after the peak of the spawning season. Bluefish contained up to 100 parasites per fish. Infection was associated with a range of disorders, including hemorrhage, inf lammation, edema, prenecrotic and necrotic changes, and follicular atresia, that may prevent proper development of oocytes and probably affect bluefish fecundity. Historical occurrences, life cycle, and geographical distribution of this nematode remain largely unknown, but may play important roles in recruitment processes of bluefish.

Population structure and variance effective size of red snapper (Lutjanus campechanus) in the northern Gulf of Mexico*

We assayed allelic variation at 19 nuclear-encoded microsatellites among 1622 Gulf red snapper (Lutjanus campechanus) sampled from the 1995 and 1997 cohorts at each of three offshore localities in the northern Gulf of Mexico (Gulf). Localities represented western, central, and eastern subregions within the northern Gulf. Number of alleles per microsatellite per sample ranged from four to 23, and gene diversity ranged from 0.170 to 0.917. Tests of conformity to Hardy-Weinberg equilibrium expectations and of genotypic equilibrium between pairs of micro-satellites were generally nonsignificant following Bonferroni correction. Significant genic or genotypic heterogeneity (or both) among samples was detected at four microsatellites and over all microsatellites. Levels of divergence among samples were low (FST ≤0.001). Pairwise exact tests revealed that six of seven “significant” comparisons involved temporal rather than spatial heterogeneity. Contemporaneous or variance effective size (NeV) was estimated from the temporal variance in allele frequencies by using a maximum-likelihood method. Estimates of NeV ranged between 1098 and >75,000 and differed significantly among localities; the NeV estimate for the sample from the northcentral Gulf was >60 times as large as the estimates for the other two localities. The differences in variance effective size could ref lect differences in number of individuals successfully reproducing, differences in patterns and intensity of immigration, or both, and are consistent with the hypothesis, supported by life-history data, that different “demographic stocks” of red snapper are found in the northern Gulf. Estimates of NeV for red snapper in the northern Gulf were at least three orders of magnitude lower than current estimates of census size (N). The ratio of effective to census size (Ne/N) is far below that expected in an ideal population and may reflect high variance in individual reproductive success, high temporal and spatial variance in productivity among subregions or a combination of the two.