Geographic variation among age-0 walleye pollock (Theragra chalcogramma): evidence of mesoscale variation in nursery quality?

Nurseries play an important part in the production of marine f ishes. Determining the relative importance of different nurseries in maintaining the parental population, however, can be difficult. In the western Gulf of Alaska, the Kodiak Island vicinity may be particularly well suited as a pollock nursery because of a prey-rich nearshore environment. Our objectives were 1) to examine age-0 pollock body condition, growth, and diet for evidence of a nearshore-shelf effect, and 2) to determine if variation in the potential prey field of zooplankton was associated with this effect. This was a pilot study that occurred in three bays and over the adjacent shelf off east Kodiak Island during 5−18 September 1993. Sampling occurred only during night at locations where echo sign indicated the presence of age-0 pollock. Echo sign was targeted to increase the chance of collecting fish given the limited vessel time. Fish condition was indicated by length-specific body weight. Growth rate indices were estimated for three different periods by using fish lengthage data and daily otolith increment widths: 1) from hatching date to capture, 2) 1−5 d before capture, and 3) 6−10 d before capture. Fish diet was determined from gut content analysis. Considerable variation among areas was evident in zooplankton composition, and fish condition, growth, and diet. However, relatively high prey densities, as well as fish condition and growth rates indicated that Chiniak Bay was particularly well suited as a pollock nursery. Hatching-date distributions indicated that most of the age-0 walleye pollock from bays were spawned earlier than were those from the shelf. The benefit of being reared in nearshore areas is therefore realized more by individuals that were spawned early than by individuals spawned relatively late.

Sablefish, Anoplopoma fimbria, Populations on Gulf of Alaska Seamounts

Sablefish, Anoplopoma fimbria, were tagged and released on Gulf of Alaska seamounts during 1999–2002 to determine the extent, if any, of emigration from the seamounts back to the continental slope and of movement between seamounts. Seventeen sablefish from Gulf of Alaska seamounts have been recovered on the continental slope since tagging began, verifying that seamount to slope migration occurs. Forty-two sablefish were recovered on the same seamounts where they were tagged, and none have been recaptured on seamounts other than the ones where they were released. Sablefish populations on Gulf of Alaska seamounts are made up of individuals mostly older than 5 years and are maledominant, with sex ratios varying from 4:1 up to 10:1 males to females. Males are smaller than females, but the average age of males is greater than that of females, and males have a greater range of age (4–64 yr) than females (4–48 yr). Otoliths of seamount fish frequently have an area of highly compressed annuli, known as the transition zone, where growth has suddenly and greatly slowed or even stopped. Because transition zones can be present in both younger and older seamount fish and are rare in slope fish, formation of otolith transition zones may be related to travel to the seamounts. The route sablefish use to reach the seamounts is so far unknown. One possibility is that fish enter the eastward-flowing North Pacific Current off the Aleutian Islands or western Gulf of Alaska and travel more or less passively on the current until encountering a seamount. The route from seamount back to slope would likely be the northwardflowing Alaska Current. These routes are discussed in light of tag recovery locations of slope- and seamount-tagged fish.

A Comparison of Two Stratification Schemes Used in Sampling Canadian Atlantic Cod, Gadus morhua

Sampling is a key element in the assessment of any fish stock. It is often one of the most expensive activities of the management process; thus, improved efficiency can result in significant cost savings. In most cases a two-phase sampling strategy is employed. Two commonly used versions of such stratified random schemes were simulated using a test population based on Atlantic cod, Gadus morhua. A 1 otolith per 1 cm length frequency currently used for many flatfish and some smaller gadoids and a 3 otolith per 3 cm length frequency currently used for many of the larger gadoids. No difference was detected in the age composition or mean length at age for either scheme; however, 10 percent fewer otoliths were collected in 1 for 1 sampling than 3 for 3. There was an improvement of between 30 and 60 percent in the coefficient of variation of the estimated catch numbers at age using the 1 for 1 compared with the 3 for 3 stratified sampling. For these reasons and other operational considerations, the 1 for 1 stratified random design of sampling appears to be superior.

Changes in size and age at maturity of the northern stock of Tilefish (Lopholatilus chamaeleonticeps) after a period of overfishing

The modern fishery for Tilefish (Lopholatilus chamaeleonticeps) developed during the 1970s, offshore of southern New England, in the western North Atlantic Ocean. The population quickly became over exploited, with documented declines in catch rates and changes in demographic traits. In an earlier study, median size at maturity (L50) of males declined from 62.6 to 38.6 cm fork length (FL) and median age at maturity (A50) of males declined from 7.1 to 4.6 years between 1978 and 1982. As part of a cooperative research effort to improve the data-limited Tilefish assessment, we updated maturity parameter estimates through the use of an otolith aging method and macroscopic and microscopic evaluations of gonads. The vital rates for this species have continued to change, particularly for males. By 2008, male L50 and A50 had largely rebounded, to 54.1 cm FL and 5.9 years. Changes in female reproductive schedules were less variable among years, but the smallest L50 and youngest A50 were recorded in 2008. Tilefish are dimorphic, where the largest fish are male, and male spawning success is postulated to be socially mediated. These traits may explain the initial rapid decline and the subsequent rebound in male L50 and A50 and less dramatic effects on females. Other factors that likely contribute to the dynamics of maturity parameter estimates are the relatively short period of overfishing and the amount of time since efforts to rebuild this fishery began, as measured in numbers of generations. This study also confirms the gonochoristic sexual pattern of the northern stock, and it reveals evidence of age truncation and relatively high proportions of immature Tilefish in the recent catch.

James's rule and causes and consequences of a latitudinal cline in the demography of John's Snapper (Lutjanus johnii) in coastal waters of Australia

Demographic parameters were derived from sectioned otoliths of John’s Snapper (Lutjanus johnii) from 4 regions across 9° of latitude and 23° of longitude in northern Australia. Latitudinal variation in size and growth rates of this species greatly exceeded longitudinal variation. Populations of John’s Snapper farthest from the equator had the largest body sizes, in line with James’s rule, and the fastest growth rates, contrary to the temperature-size rule for ectotherms. A maximum age of 28.6 years, nearly 3 times previous estimates, was recorded and the largest individual was 990 mm in fork length. Females grew to a larger mean asymptotic fork length (L∞) than did males, a finding consistent with functional gonochorism. Otolith weight at age and gonad weight at length followed the same latitudinal trends seen in length at age. Length at maturity was ~72–87% of L∞ and varied by ~23% across the full latitudinal gradient, but age at first maturity was consistently in the range of 6–10 years, indicating that basic growth trajectories were similar across vastly different environments. We discuss both the need for complementary reproductive data in age-based studies and the insights gained from experiments where the concept of oxygen- and capacity-limited thermal tolerance is applied to explain the mechanistic causes of James’s rule in tropical fish species.

Age determination manual of the Alaska Fisheries Science Center Age and Growth Program

The Age and Growth Program at the Alaska Fisheries Science Center is tasked with providing age data in order to improve the basic understanding of the ecology and fisheries dynamics of Alaskan fish species. The primary focus of the Age and Growth Program is to estimate ages from otoliths and other calcified structures for age-structured modeling of commercially exploited stocks; however, the program has recently expanded its interests to include numerous studies on topics ranging from age estimate validation to the growth and life history of non-target species. Because so many applications rely upon age data and particularly upon assurances as to their accuracy and precision, the Age and Growth Program has developed this practical guide to document the age determination of key groundfish species from Alaskan waters. The main objective of this manual is to describe techniques specific to the age determination of commercially and ecologically important species studied by the Age and Growth Program. The manual also provides general background information on otolith morphology, dissection, and preparation, as well as descriptions of methods used to measure precision and accuracy of age estimates. This manual is intended not only as a reference for age readers at the AFSC and other laboratories, but also to give insight into the quality of age estimates to scientists who routinely use such data.

Age, growth and mortality of the toadfish, Halobatrachus didactylus (Schneider, 1801) (Pisces: Batrachoididae), in the Bay of Cádiz (southwestern Spain)

Age, growth and mortality of the toadfish, Halobatrachus didactylus, were determined by examination of the whole sagittal otoliths of fish sampled in the Bay of Cádiz (southwestern Spain) from March 1999 to March 2000. A total of 844 specimens (425 males, 416 females, and 3 of indeterminate sex), ranging from 95 to 470 mm in total length were examined. Eighty-nine percent of the otoliths could be read allowing an age estimation. The opaque zone was formed between April and May coincident with the maximum reproductive peak, while the translucent zone formed mainly in summer-fall (June to December). Maximum ages for males and females were 12 and 10 years, respectively. The samples were dominated by 2- to 6-year-old specimens. Males matured at an age of approximately 2 years and females at 3 years. Fish total length and otolith radius were closely related. The von Bertalanffy growth curve was used to describe growth. The parameters were derived from back-calculated length-at-age. Significant differences in the growth parameters were found between sexes. Although the growth analysis revealed that this species is slow-growing, males reached larger sizes than females. Females appeared to experience higher natural mortality rates than males.

Sources of age determination errors for sablefish (Anoplopoma fimbria)

This study was undertaken to resolve problems in age determination of sablefish (Anoplopoma fimbria). Aging of this species has been hampered by poor agreement (averaging less than 45%) among age readers and by differences in assigned ages of as much as 15 years. Otoliths from fish that had been injected with oxytetracycline (OTC) and that had been at liberty for known durations were used to determine why age determinations were so difficult and to help determine the correct aging procedure. All fish were sampled from Oregon southwards, which represents the southern part of their range. The otoliths were examined with the aid of image processing. Some fish showed little or no growth on the otolith after eight months at liberty, whereas otoliths from other fish grew substantially. Some fish lay down two prominent hyaline zones within a single year, one in the summer and one in the winter. We classified the otoliths by morphological type and found that certain types are more likely to lay down multiple hyaline zones and other types are likely to lay down little or no zones. This finding suggests that some improvement could be achieved by detailed knowledge of the growth characteristics of the different types. This study suggests that it may not be possible to obtain reliable ages from sablefish otoliths. At the very least, more studies will be required to under-stand the growth of sablefish otoliths.

Growth, mortality, and hatchdate distributions of larval and juvenile spotted seatrout (Cynoscion nebulosus) in Florida Bay, Everglades National Park

Life history aspects of larval and, mainly, juvenile spotted seatrout (Cynoscion nebulosus) were studied in Florida Bay, Everglades National Park, Florida. Collections were made in 1994−97, although the majority of juveniles were collected in 1995. The main objective was to obtain life history data to eventually develop a spatially explicit model and provide baseline data to understand how Everglades restoration plans (i.e. increased freshwater flows) could influence spotted seatrout vital rates. Growth of larvae and juveniles (<80 mm SL) was best described by the equation loge standard length = –1.31 + 1.2162 (loge age). Growth in length of juveniles (12–80 mm SL) was best described by the equation standard length = –7.50 + 0.8417 (age). Growth in wet weight of juveniles (15–69 mm SL) was best described by the equation loge wet-weight = –4.44 + 0.0748 (age). There were no significant differences in juvenile growth in length of spotted seatrout in 1995 between three geographical subdivisions of Florida Bay: central, western, and waters adjacent to the Gulf of Mexico. We found a significant difference in wet-weight for one of six cohorts categorized by month of hatchdate in 1995, and a significant difference in length for another cohort. Juveniles (i.e. survivors) used to calculate weekly hatchdate distributions during 1995 had estimated spawning times that were cyclical and protracted, and there was no correlation between spawning and moon phase. Temperature influenced otolith increment widths during certain growth periods in 1995. There was no evidence of a relationship between otolith growth rate and temperature for the first 21 increments. For increments 22–60, otolith growth rates decreased with increasing age and the extent of the decrease depended strongly in a quadratic fashion on the temperature to which the fish was exposed. For temperatures at the lower and higher range, increment growth rates were highest. We suggest that this quadratic relationship might be influenced by an environmental factor other than temperature. There was insufficient information to obtain reliable inferences on the relationship of increment growth rate to salinity.

Red snapper (Lutjanus campechanus) demographic structure in the northern Gulf of Mexico based on spatial patterns in growth rates and morphometrics

Red snapper (Lutjanus campechanus) in the United States waters of the Gulf of Mexico (GOM) has been considered a single unit stock since management of the species began in 1991. The validity of this assumption is essential to management decisions because measures of growth can differ for nonmixing populations. We examined growth rates, size-at-age, and length and weight information of red snapper collected from the recreational harvests of Alabama (n=2010), Louisiana (n=1905), and Texas (n =1277) from 1999 to 2001. Ages were obtained from 5035 otolith sections and ranged from one to 45 years. Fork length, total weight, and age-frequency distributions differed significantly among all states; Texas, however, had a much higher proportion of smaller, younger fish. All red snapper showed rapid growth until about age 10 years, after which growth slowed considerably. Von Bertalanffy growth models of both mean fork length and mean total weight-at-age predicted significantly smaller fish at age from Texas, whereas no differences were found between Alabama and Louisiana models. Texas red snapper were also shown to differ significantly from both Alabama and Louisiana red snapper in regressions of mean weight at age. Demographic variation in growth rates may indicate the existence of separate management units of red snapper in the GOM. Our data indicate that the red snapper inhabiting the waters off Texas are reaching smaller maximum sizes at a faster rate and have a consistently smaller total weight at age than those collected from Louisiana and Alabama waters. Whether these differences are environmentally induced or are the result of genetic divergence remains to be determined, but they should be considered for future management regulations.

Elemental signatures in otoliths of larval walleye pollock (Theragra chalcogramma) from the northeast Pacific Ocean

The population structure of walleye pollock (Theragra chalcogramma) in the northeastern Pacific Ocean remains unknown. We examined elemental signatures in the otoliths of larval and juvenile pollock from locations in the Bering Sea and Gulf of Alaska to determine if there were significant geographic variations in otolith composition that may be used as natural tags of population affinities. Otoliths were assayed by using both electron probe microanalysis (EPMA) and laser ablation inductively coupled plasma mass spectrometry (ICP-MS). Elements measured at the nucleus of otoliths by EPMA and laser ablation ICP-MS differed significantly among locations. However, geographic groupings identified by a multivariate statistical approach from EPMA and ICP-MS were dissimilar, indicating that the elements assayed by each technique were controlled by separate depositional processes within the endolymph. Elemental profiles across the pollock otoliths were generally consistent at distances up to 100 μm from the nucleus. At distances beyond 100 μm, profiles varied significantly but were remarkably consistent among individuals collected at each location. These data may indicate that larvae from various spawning locations are encountering water masses with differing physicochemical properties through their larval lives, and at approximately the same time. Although our results are promising, we require a better understanding of the mechanisms controlling otolith chemistry before it will be possible to reconstruct dispersal pathways of larval pollock based on probe-based analyses of otolith geochemistry. Elemental signatures in otoliths of pollock may allow for the delineation of fine-scale population structure in pollock that has yet to be consistently revealed by using population genetic approaches.

Radiometric validation of age, growth, and longevity for the blackgill rockfish (Sebastes melanostomus)

As nearshore fish populations decline, many commercial fishermen have shifted fishing effort to deeper continental slope habitats to target fishes for which biological information is limited. One such fishery that developed in the northeastern Pacific Ocean in the early 1980s was for the blackgill rockfish (Sebastes melanostomus), a deep-dwelling (300−800 m) species that congregates over rocky pinnacles, mainly from southern California to southern Oregon. Growth zone-derived age estimates from otolith thin sections were compared to ages obtained from the radioactive disequilibria of 210Pb, in relation to its parent, 226Ra, in otolith cores of blackgill rockfish. Age estimates were validated up to 41 years, and a strong pattern of agreement supported a longevity exceeding 90 years. Age and length data fitted to the von Bertalanffy growth function indicated that blackgill rockfish are slow-growing (k= 0.040 females, 0.068 males) and that females grow slower than males, but reach a greater length. Age at 50% maturity, derived from previously published length-at-maturity estimates, was 17 years for males and 21 years for females. The results of this study agree with general life history traits already recognized for many Sebastes species, such as long life, slow growth, and late age at maturation. These traits may undermine the sustainability of blackgill rockfish populations when heavy fishing pressure, such as that which occurred in the 1980s, is applied.

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.

The relative value of different estuarine nursery areas in North Carolina for transient juvenile marine fishes

Offshore winter-spawned fishes dominate the nekton of south-eastern United States estuaries. Their juveniles reside for several months in shallow, soft bottom estuarine creeks and bays called primary nursery areas. Despite similarity in many nursery characteristics, there is, between and within species, variability in the occupation of these habitats. Whether all occupied habitats are equally valuable to individuals of the same species or whether most recruiting juveniles end up in the best habitats is not known. If nursery quality varies, then factors controlling variation in pre-settlement fish distribution are important to year-class success. If nursery areas have similar values, interannual variation in distribution across nursery creeks should have less effect on population sizes or production. I used early nursery period age-specific growth and mortality rates of spot (Leiostomus xanthurus) and Atlantic croaker (Micropogonias undulatus)—two dominant estuarine fishes—to assess relative habitat quality across a wide variety of nursery conditions, assuming that fish growth and mortality rates were direct reflections of overall physical and biological conditions in the nurseries. I tested the hypothesis that habitat quality varies for these fishes by comparing growth and mortality rates and distribution patterns across a wide range of typical nursery habitats at extreme ends of two systems. Juvenile spot and Atlantic croaker were collected from 10 creeks in the Cape Fear River estuary and from 18 creeks in the Pamlico Sound system, North Carolina, during the 1987 recruitment season (mid-March–mid-June). Sampled creeks were similar in size, depth, and substrates but varied in salinities, tidal regimes, and distances from inlets. Spot was widely distributed among all the estuarine creeks, but was least abundant in the creeks in middle reaches of both systems. Atlantic croaker occurred in the greatest abundance in oligohaline creeks of both systems. Instantaneous growth rates derived from daily otolith ages were generally similar for all creeks and for both species, except that spot exhibited a short-term growth depression in the upriver Pamlico system creeks—perhaps the result of the long migration distance of this species to this area. Spot and Atlantic croaker from upriver oligohaline creeks exhibited lower mortality rates than fish from downstream polyhaline creeks. These results indicated that even though growth was similar at the ends of the estuaries, the upstream habitats provided conditions that may optimize fitness through improved survival.

Scales of spatial variation in demography of a large coral-reef fish: an exception to the typical model?

Spatial variation in demographic parameters of the red throat emperor (Lethrinus miniatus) was examined among 12 coral reefs in three geographic regions (Townsville, Mackay, and Storm Cay) spanning over 3° of latitude of the Great Barrier Reef, Australia. Estimates of demographic parameters were based on age estimates from counts of annuli in whole otoliths because there was no significant difference in age estimates between whole and sectioned otoliths. There were significant regional differences in age structures, rates of somatic and otolith growth, and total mortality. The Townsville region was characterized by the greatest proportion of older fish, the smallest maximum size, and the lowest rates of otolith growth and total mortality. In contrast the Mackay region was characterized by the highest proportion of younger fish, the largest maximum size, and the highest rates of otolith growth and total mortality. Demographic parameters for the Storm Cay region were intermediate between the other two regions. Historic differences in fishing pressure and regional differences in productivity are two alternative hypotheses given to explain the regional patterns in demographic parameters. All demographic parameters were similar among the four reefs within each region. Thus, subpopulations with relatively homogeneous demographic parameters occurred on scales of reef clusters. Previous studies, by contrast, have found substantial between-reef variation in demographic parameters within regions. Thus spatial variation in demographic parameters for L. miniatus may differ from what is assumed typical for a coral-reef fish metapopulation.