Effects of maternal age and size on embryonic energy reserves, developmental timing, and fecundity in quillback rockfish (Sebastes maliger)

Maternal effects on the quality of progeny can have direct impacts on population productivity. Rockfish are viviparous and the oil globule size of larvae at parturition has been shown to have direct effects on time until starvation and growth rate. We sampled embryos and preparturition larvae opportunistically from 89 gravid quillback rockfish (Sebastes maliger) in Southeast Alaska. Because the developmental stage and sampling period were correlated with oil globule size, they were treated as covariates in an analysis of maternal age, length, and weight effects on oil globule size. Maternal factors were related to developmental timing for almost all sampling periods, indicating that older, longer, and heavier females develop embryos earlier than younger, shorter, or lighter ones. Oil globule diameter and maternal length and weight were statistically linked, but the relationships may not be biologically significant. Weight-specific fecundity did not increase with maternal size or age, suggesting that reproductive output does not increase more quickly as fish age and grow. Age or size truncation of a rockfish population, in which timing of parturition is related to age and size, could result in a shorter parturition season. This shortening of the parturition season could make the population vulnerable to fluctuating environmental conditions.

Changes in catch rates and length and age at maturity, but not growth, of an estuarine plotosid (Cnidoglanis macrocephalus) after heavy fishing

The hypothesis that heavy fishing pressure has led to changes in the biological characteristics of the estuary cobbler (Cnidoglanis macrocephalus) was tested in a large seasonally open estuary in southwestern Australia, where this species completes its life cycle and is the most valuable commercial fish species. Comparisons were made between seasonal data collected for this plotosid (eeltail catfish) in Wilson Inlet during 2005–08 and those recorded with the same fishery-independent sampling regime during 1987–89. These comparisons show that the proportions of larger and older individuals and the catch rates in the more recent period were far lower, i.e., they constituted reductions of 40% for fish ≥430 mm total length, 62% for fish ≥4 years of age, and 80% for catch rate. In addition, total mortality and fishing-induced mortality estimates increased by factors of ~2 and 2.5, respectively. The indications that the abundance and proportion of older C. macrocephalus declined between the two periods are consistent with the perception of long-term commercial fishermen and their shift toward using a smaller maximum gill net mesh to target this species. The sustained heavy fishing pressure on C. macrocephalus between 1987–89 and 2005–08 was accompanied by a marked reduction in length and age at maturity of this species. The shift in probabilistic maturation reaction norms toward smaller fish in 2005–08 and the lack of a conspicuous change in growth between the two periods indicate that the maturity changes were related to fishery-induced evolution rather than to compensatory responses to reduced fish densities.

Interactions of age-dependent mortality and selectivity functions in age-based stock assessment models

The natural mortality rate (M) of fish varies with size and age, although it is often assumed to be constant in stock assessments. Misspecification of M may bias important assessment quantities. We simulated fishery data, using an age-based population model, and then conducted stock assessments on the simulated data. Results were compared to known values. Misspecification of M had a negligible effect on the estimation of relative stock depletion; however, misspecification of M had a large effect on the estimation of parameters describing the stock recruitment relationship, age-specific selectivity, and catchability. If high M occurs in juvenile and old fish, but is misspecified in the assessment model, virgin biomass and catchability are often poorly estimated. In addition, stock recruitment relationships are often very difficult to estimate, and steepness values are commonly estimated at the upper bound (1.0) and overfishing limits tend to be biased low. Natural mortality can be estimated in assessment models if M is constant across ages or if selectivity is asymptotic. However if M is higher in old fish and selectivity is dome-shaped, M and the selectivity cannot both be adequately estimated because of strong interactions between M and selectivity.

Age, growth, and spawning season of red bream (Beryx decadactylus) off the southeastern United States

Red bream (Beryx decadactylus) is a commercially important deep-sea benthopelagic fish with a circumglobal distribution on insular and continental slopes and seamounts. In the United States, small numbers are caught incidentally in the wreckfish (Polyprion americanus) fishery which operates off the southeastern coast, but no biological information exists for the management of the U.S. red bream population. For this study, otoliths (n=163) and gonads (n=161) were collected from commercially caught red bream between 2003 and 2008 to determine life history parameters. Specimens ranged in size from 410 to 630 mm fork length and were all determined to be mature by histological examination of the gonads. Females in spawning condition were observed from June through September, and reproductively active males were found year-round. Sectioned otoliths were difficult to interpret, but maximum age estimates were much higher than the 15 years previously reported for this species from the eastern North Atlantic based on whole-otolith analysis. Estimated ages ranged from 8 to 69 years, and a minimum lifespan of 49 years was validated by using bomb radiocarbon dating. Natural mortality was estimated at 0.06/yr. This study shows that red bream are longer lived and more vulnerable to overfishing than previously assumed and should be managed carefully to prevent overexploitation.

Integrating methods for determining length-at-age to improve growth estimates for two large scombrids

Fish growth is commonly estimated from length-at-age data obtained from otoliths. There are several techniques for estimating length-at-age from otoliths including 1) direct observed counts of annual increments; 2) age adjustment based on a categorization of otolith margins; 3) age adjustment based on known periods of spawning and annuli formation; 4) back-calculation to all annuli, and 5) back-calculation to the last annulus only. In this study we compared growth estimates (von Bertalanffy growth functions) obtained from the above five methods for estimating length-at-age from otoliths for two large scombrids: narrow-barred Spanish mackerel (Scomberomorus commerson) and broad-barred king mackerel (Scomberomorus semifasciatus). Likelihood ratio tests revealed that the largest differences in growth occurred between the back-calculation methods and the observed and adjusted methods for both species of mackerel. The pattern, however, was more pronounced for S. commerson than for S. semifasciatus, because of the pronounced effect of gear selectivity demonstrated for S. commerson. We propose a method of substituting length-at-age data from observed or adjusted methods with back-calculated length-at-age data to provide more appropriate estimates of population growth than those obtained with the individual methods alone, particularly when faster growing young fish are disproportionately selected for. Substitution of observed or adjusted length-at-age data with back-calculated length-at-age data provided more realistic estimates of length for younger ages than observed or adjusted methods as well as more realistic estimates of mean maximum length than those derived from backcalculation methods alone.

Age validation, growth, mortality, and demographic modeling of spotted gully shark (Triakis megalopterus) from the southeast coast of South Africa

This study documents validation of vertebral band-pair formation in spotted gully shark (Triakis megalopterus) with the use of fluorochrome injection and tagging of captive and wild sharks over a 21-year period. Growth and mortality rates of T. megalopterus were also estimated and a demographic analysis of the species was conducted. Of the 23 OTC (oxytetracycline) -marked vertebrae examined (12 from captive and 11 from wild sharks), seven vertebrae (three from captive and four from wild sharks) exhibited chelation of the OTC and fluoresced under ultraviolet light. It was concluded that a single opaque and translucent band pair was deposited annually up to at least 25 years of age, the maximum age recorded. Reader precision was assessed by using an index of average percent error calculated at 5%. No significant differences were found between male and female growth patterns (P>0.05), and von Bertalanffy growth model parameters for combined sexes were estimated to be L∞=1711.07 mm TL, k=0.11/yr and t0=–2.43 yr (n=86). Natural mortality was estimated at 0.17/yr. Age at maturity was estimated at 11 years for males and 15 years for females. Results of the demographic analysis showed that the population, in the absence of fishing mortality, was stable and not significantly different from zero and particularly sensitive to overfishing. At the current age at first capture and natural mortality rate, the fishing mortality rate required to result in negative population growth was low at F>0.004/ yr. Elasticity analysis revealed that juvenile survival was the principal factor in explaining variability in population growth rate.

Age validation of great hammerhead shark (Sphyrna mokarran), determined by bomb radiocarbon analysis

Preliminary validation of annual growth band deposition in vertebrae of great hammerhead shark (Sphyrna mokarran) was conducted by using bomb radiocarbon analysis. Adult specimens (n=2) were collected and thin sections of vertebral centra were removed for visual aging and use in radiocarbon assays. Vertebral band counts were used to estimate age, and year of formation was assigned to each growth band by subtracting estimated age from the year of capture. A total of 10 samples were extracted from growth bands and analyzed for Δ14C. Calculated Δ14C values from dated bands were compared to known-age reference chronologies, and the resulting patterns indicated annual periodicity of growth bands up to a minimum age of 42 years. Trends in Δ14C across time in individual specimens indicated that vertebral radiocarbon is conserved through time but that habitat and diet may inf luence Δ14C levels in elasmobranchs. Although the age validation reported here must be considered preliminary because of the small sample size and narrow age range of individuals sampled, it represents the first confirmation of age in S. mokarran, further illustrating the usefulness of bomb radiocarbon analysis as a tool for life history studies in elasmobranchs.

Age and growth of spiny dogfish (Squalus acanthias) in the Gulf of Alaska: analysis of alternative growth models

Ten growth models were fitted to age and growth data for spiny dogfish (Squalus acanthias) in the Gulf of Alaska. Previous studies of spiny dogfish growth have all fitted the t0 formulation of the von Bertalanffy model without examination of alternative models. Among the alternatives, we present a new two-phase von Bertalanffy growth model formulation with a logistically scaled k parameter and which estimates L0. A total of 1602 dogfish were aged from opportunistic collections with longline, rod and reel, set net, and trawling gear in the eastern and central Gulf of Alaska between 2004 and 2007. Ages were estimated from the median band count of three independent readings of the second dorsal spine plus the estimated number of worn bands for worn spines. Owing to a lack of small dogfish in the samples, lengths at age of small individuals were back-calculated from a subsample of 153 dogfish with unworn spines. The von Bertalanffy, two-parameter von Bertalanffy, two-phase von Bertalanffy, Gompertz, two-parameter Gompertz, and logistic models were fitted to length-at-age data for each sex separately, both with and without back-calculated lengths at age. The two-phase von Bertalanffy growth model produced the statistically best fit for both sexes of Gulf of Alaska spiny dogfish, resulting in L∞ = 87.2 and 102.5 cm and k= 0.106 and 0.058 for males and females, respectively.

Effects of seasonal change on activity rhythms and swimming behavior of age-0 bluefish (Pomatomus saltatrix) and a description of gliding behavior

Daily and seasonal activity rhythms, swimming speed, and modes of swimming were studied in a school of spring-spawned age-0 bluefish (Pomatomus saltatrix) for nine months in a 121-kL research aquarium. Temperature was lowered from 20° to 15°C, then returned to 20°C to match the seasonal cycle. The fish grew from a mean 198 mm to 320 mm (n= 67). Bluefish swam faster and in a more organized school during day (overall mean 47 cm/s) than at night (31 cm/s). Swimming speed declined in fall as temperature declined and accelerated in spring in response to change in photoperiod. Besides powered swimming, bluefish used a gliding-upswimming mode, which has not been previously described for this species. To glide, a bluefish rolled onto its side, ceased body and tail beating, and coasted diagonally downward. Bluefish glided in all months of the study, usually in the dark, and most intensely in winter. Energy savings while the fish is gliding and upswimming may be as much as 20% of the energy used in powered swimming. Additional savings accrue from increased lift due to the hydrofoil created by the horizontal body orientation and slightly concave shape. Energy-saving swimming would be advantageous during migration and overwintering.

Age validation of Dover sole (Microstomus pacificus) by means of bomb radiocarbon

We used bomb radiocarbon (14C) in this age validation study of Dover sole (Microstomus pacificus). The otoliths of Dover sole, a commercially important fish in the North Pacific, are difficult to age and ages derived from the current break-andburn method were not previously validated. The otoliths used in this study were chosen on the basis of estimated birth year and for the ease of interpreting growth zone patterns. Otolith cores, material representing years 0 through 3, were isolated and analyzed for 14C. Additionally, a small number of otoliths with difficult-to-interpret growth patterns were analyzed for 14C to help determine age interpretation. The measured Dover sole 14C values in easier-to-interpret otoliths were compared with a 14C reference chronology for Pacific halibut (Hippoglossus stenolepis) in the North Pacific. We used an objective statistical analysis where sums of squared residuals between otolith 14C values of Dover sole and the reference chronology were examined. Our statistical analysis also included a procedure where the Dover sole 14C values were standardized to the reference chronology. These procedures allowed an evaluation of aging error. The 14C results indicated that the Dover sole age estimates from the easier-to-interpret otoliths with the break-and-burn method are accurate. This study validated Dover sole ages from 8 to 47 years.

Age-specific movement patterns of sablefish (Anoplopoma fimbria) in Alaska

Over 34,000 age 0–2 juvenile sablefish (Anoplopoma fimbria) were tagged and released in southeast Alaska waters during 1985–2005. The data set resulting from this tagging study was unusual because of its time span (20 years) and because age could be reliably inferred from release length (i.e., tagged and released fish were of known age); thus, age-specific movement patterns could be examined. The depth- and area-related recovery patterns supported the concepts that sablefish move to deeper water with age and migrate counterclockwise in the Gulf of Alaska. Availability to the fishery increased rapidly for fish of younger ages, peaked at age 5 to 6, and then gradually declined as sablefish moved deeper with age. Decreased availability with age may occur because of lower fishing effort in deep water and could have substantial implications for sablef ish stock assessments because “domeshaped” availability influences the reliability of abundance estimates. The area-related recovery pattern was not affected by year-class strength; i.e., there was no significant densitydependent relationship.

Age- and length-at-maturity of female arrowtooth flounder (Atheresthes stomias) in the Gulf of Alaska

Arrowtooth flounder (Atheresthes stomias) has had the highest abundance of any groundfish species in the Gulf of Alaska since the 1970s (Matarese et al., 2003; Turnock et al., 2005; Blood et al., 2007); however, commercial catches have been restricted because Pacific halibut (Hippoglossus stenolepis) are caught as bycatch in the fishery. Arrowtooth flounder plays a key role in the ecosystem because it is a dominant organism within the food web, both as an apex predator of fish and invertebrates, as well as an important prey for walleye pollock (Theragra chalcogramma; Aydin et al., 2002). Walleye pollock is the dominant groundfish in the Bering Sea, a principal groundfish in the Gulf of Alaska, and the primary prey for marine mammals. The distribution of arrowtooth flounder extends from Cape Navarin and the eastern Sea of Okhotsk in Russia, across the Bering Sea, Aleutian Islands, Gulf of Alaska, and south to the coast of central California (Shuntov, 1964; Britt and Martin, 2001; Chetvergov, 2001; Weinberg et al., 2002; Zenger, 2004). Because of the importance of arrowtooth flounder in the marine ecosystem of A laska, a maturity study of this species was undertaken to determine age-at-maturity, which is essential for age-based stock management models. Before these results, management has had to rely upon a length-at-maturity-based estimate (Zimmermann, 1997) to manage stocks in the Gulf of Alaska (GOA), Bering Sea, and Aleutian Islands. The central GOA was selected as the location for this maturity study Age- and length-at-maturity of female arrowtooth flounder (Atheresthes stomias) in the Gulf of Alaska because it contains approximately 70% of the total Gulf of Alaska arrowtooth flounder biomass (1.9×106 t, age 3 and older)— the highest percentage in the world (Shuntov, 1964; Britt and Martin, 2001; Weinberg et al., 2002; Wilderbuer and Nichol, 2006).

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.

Incorporating fishery observer data into an integrated catch-at-age and multiyear tagging model for estimating mortality rates and abundance

Tagging experiments are a useful tool in fisheries for estimating mortality rates and abundance of fish. Unfortunately, nonreporting of recovered tags is a common problem in commercial fisheries which, if unaccounted for, can render these estimates meaningless. Observers are often employed to monitor a portion of the catches as a means of estimating reporting rates. In our study, observer data were incorporated into an integrated model for multiyear tagging and catch data to provide joint estimates of mortality rates (natural and f ishing), abundance, and reporting rates. Simulations were used to explore model performance under a range of scenarios (e.g., different parameter values, parameter constraints, and numbers of release and recapture years). Overall, results indicated that all parameters can be estimated with reasonable accuracy, but that fishing mortality, reporting rates, and abundance can be estimated with much higher precision than natural mortality. An example of how the model can be applied to provide guidance on experimental design for a large-scale tagging study is presented. Such guidance can contribute to the successful and cost-effective management of tagging programs for commercial fisheries.

Age and growth of swordfish (Xiphias gladius) caught by the Hawaii-based pelagic longline fishery

We verified the age and growth of swordfish (Xiphias gla-dius) by comparing ages determined from annuli in fin ray sections with daily growth increments in otoliths. Growth of swordfish of exploitable sizes is described on the basis of annuli present in cross sections of the second ray of the first anal fins of 1292 specimens (60−260 cm eye-to-fork length, EFL) caught in the region of the Hawaii-based pelagic longline fishery. The position of the initial fin ray annulus of swordfish was verified for the first time with the use of scanning electron micrographs of presumed daily growth increments present in the otoliths of juveniles. Fish growth through age 7 was validated by marginal increment analysis. Faster growth of females was confirmed, and the standard von Bertalanffy growth model was identified as the most parsimonious for describing growth in length for fish greater than 60 cm EFL. The observed growth of three fish, a year-old in size when first caught and then recaptured from 364 to1490 days later, is consistent with modeled growth for fish of this size range. Our novel approach to verifying age and growth should increase confidence in conducting an age-structured stock assessment for swordfish in the North Pacific Ocean.