Daily and sub-daily otolith increments of larval and juvenile walleye pollock, Theragra chalcogramma (Pallas), as validated by alizarin complexone experiments

Walleye pollock (Theragra chalcogramma) were reared from eggs to the juvenile life stage to study daily increment formation in the sagittae otoliths, which are routinely used for age and growth analyses. The apparent deposition of sub-daily growth increments becomes problematic for determining fish age from the late larval stage throughout the juvenile (young-of-the-year) development stage. Otolith marking experiments were conducted to determine interpretation criteria to differentiate between daily and sub-daily increments. Immersion of larval and transforming walleye pollock in 25 mg/l of alizarin complexone (ALC) for 6 h once a week produced a fluorescent mark on the day of staining. Evidence of six well defined and equally spaced increments counted between the weekly ALC marks validated the deposition of daily increments. The daily increments gradually increased in width as the fish/otolith grew. The criteria for determining the presence of sub-daily increments between the daily increments were (1) weak optical definition and (2) a sudden change in incremental distance that lasted for one or two increments and were approximately <0.5 μm in width. Growth problems that occurred during the experiments were identified on otoliths as reductions in daily incremental widths and optical definition, which continued for several days. Otoliths from field-collected fish have also shown similar changes in daily increment properties during the juvenile stage, which may be an indicator of an environmental influence. The criteria for defining different increment types help to resolve our current age determination issues for late larval and early juvenile stage walleye pollock from the Gulf of Alaska.     

Semilunar spawning periodicity in brackish damsel Pomacentrus taeniometopon

ABSTRACT:   Lunar-related spawning periodicity of the brackish damsel Pomacentrus taeniometopon was investigated by changes in gonadosomatic index (GSI) and ovarian histology. During the spawning season, GSI increased toward the week of the first and last quarter moon. Concomitant with increases in GSI, oocytes laden with yolk developed from the new/full moon to the first/last quarter moon. Postovulatory follicles appeared in the ovary around and after the first/last quarter moon. These results suggest that P. taeniometopon is a semilunar spawner and repeats spawnings twice a month with lunar periodicity during the spawning season. When the fish were reared under laboratory conditions without periodicity of tide, the spawning of P. taeniometopon occurred sporadically. It is likely that the fish uses tidal stimuli in the natural habitat to entrain reproductive activities.     

Validation of daily increment formation and the effects of different temperatures and feeding regimes on short-term otolith growth in Australian smelt Retropinna semoni

Abstract –  To aid otolith interpretation of wild fish, we conducted a laboratory study using metalarval Australian smelt ( Retropinna semoni ) collected from the Murray River, to examine daily increment deposition and the effects of different temperatures and feeding regimes on otolith growth. Daily increment deposition was confirmed by comparing the number of increments from an oxytetracycline mark with the known number of days from marking. After holding fish at two temperature levels and three feeding rates, both food density and temperature were found to have a significant effect on otolith growth, with food density having the greatest influence. Overall trends in final lengths and condition of fish were well represented by recent otolith growth. The results of the experiment have implications for estimating growth histories and its relationship to various environmental conditions.     

Otolith microstructure of chum salmon <Emphasis Type="Italic">Oncorhynchus keta</Emphasis>: formation of sea entry check and daily deposition of otolith increments in seawater conditions

To apply otolith microstructure to examination of age and growth of juvenile chum salmon Oncorhynchus keta inhabiting coastal waters, formation of otolith increments was investigated for juveniles reared in a seawater aquarium and in net pens. In all otoliths examined, a distinctive check was formed at the time of sea entry of the fish. The deposition of otolith increments after the check was daily for rearing both in the aquarium (57 days) and in the net pens (26 days). Check formation associated with sea entry was also observed in otoliths of juvenile salmon collected 1 km off the coast of Shari, Hokkaido, Japan. Transmitted light observation of otoliths of those fish revealed a transition in otolith increment appearance from dark to light. Otolith Sr: Ca ratio remarkably changed from a low to a high level, coinciding with the transition in otolith appearance. It is suggested that the transition was associated with individual sea entry. This study demonstrated that the check and/or transition associated with sea entry are applicable to a benchmark for otolith increment counts of juvenile chum salmon inhabiting coastal waters.     

The utility of otolith weight as a predictor of age in the emperor Lethrinus mahsena and other tropical fish species

This study examines the potential use of otolith weight as a proxy for age in the lethrinid Lethrinus mahsena from different sites in the tropical Indian Ocean: the banks of Seychelles, Mauritius and British Indian Ocean Territory (BIOT, Chagos Archipelago). The reliability of age–frequency distributions and individual ages estimated using otolith weight–age relationships was examined through comparison with those estimated through the standard method of ageing using otolith increments. Two other methods for estimating age–frequencies using age-slicing via an estimated growth curve were also examined; these used growth curves estimated by a length-based method (ELEFAN), or by fitting directly to length-at-age data (an ‘age-based’ method). Age-slicing using length-based growth parameters failed to produce reliable age–frequencies, due to inaccuracies in the growth parameter estimates. The use of age-based growth parameter estimates improved the results of age-slicing, however, age–frequencies remained significantly different from those obtained from ageing using otolith increments in two locations. The use of otolith weight–age relationships resulted in estimated age–frequency distributions that in all locations were not significantly different from those assessed through otolith increment counts. In contrast, L. mahsena otolith weight–age relationships could not be used to estimate individual ages accurately, due to the level of overlap in otolith weight between age classes. Where otolith increments are routinely used to age commercial fish species, the fact that otolith weight–age relationships could not be used to age individuals accurately may limit its application. However, where routine ageing of individuals through otolith increments is considered impractical, for instance because of its cost, the use of otolith weight–age relationships to derive catch age–frequencies represents a viable alternative approach. With this in mind, this study has also demonstrated that there is the potential to use otolith weight–age relationships for five other species caught around the Seychelles, following the validation of their otolith increments.     

Using otolith increment widths to infer spatial,temporal and gender variation in the growth of sand whiting Sillago ciliata

Abstract Two methods of otolith increment analysis were used to describe spatial, temporal and gender variation in growth of sand whiting, Sillago ciliata (Cuvier), in four south‐east Australian estuaries. Mean annual standardised otolith increment widths were used as indices of individual lifetime growth rates, while raw otolith increment widths were used to describe variation in growth throughout the life of S. ciliata. Temporal variation in growth was observed at an annual scale, while spatial variation in growth was observed between estuaries. Growth rates increased significantly with decreasing latitude and greater mean sea surface temperatures. A divergence in growth rates between sexes was detected, with females growing faster than males after sexual maturity. This study highlights how otolith increment analyses can: (1) be used to analyse temporal trends in growth from a single sample and (2) provide insight into juvenile growth when samples have an absence of undersized fish.     

Lunar-related spawning in honeycomb grouper, Epinephelus merra

Lunar-related spawning in honeycomb grouper was investigated by histological and behavioral analyses. Ovarian development and spawning of the fish were related to lunar periodicity. And also the fish migrated to out side of the reef area and spawned for a few days after the full moon.     

Age determination and growth of juveniles of the European hake,Merluccius merluccius (L., 1758), in the northern Tyrrhenian Sea (NW Mediterranean)

The aim of the present study was to provide an estimation of growth of juvenile European hake, Merluccius merluccius (L., 1758) (OSTEICHTHYES; MERLUCCIIDAE), by means of the analysis of otolith daily increments. Hake specimens were collected during trawl surveys carried out in the northern Tyrrhenian Sea (NW Mediterranean). The sagittae were removed from hakes ≤20 cm total length. Left otoliths were ground and polished to obtain thin frontal sections. Otolith microstructure was analysed under a compound green light-polarising microscope. A power curve with intercept was fitted to the length-age data to describe the growth of M. merluccius. According to the growth curve, a mean length of 18 cm was reached at the end of the first year of life. The validation of the otolith increment periodicity was performed by means of two indirect methods.     

Temporal patterns in the post-larval supply of two crustacean taxa in Rangiroa Atoll,French Polynesia

The post-larval supply of two crustacean taxa (Brachyura and Stomatopoda) was monitored using one crest net over three lunar months at Rangiroa Atoll, French Polynesia. We captured a total of 37,068 brachyuran and 12,697 stomatopod post-larvae during the study. Post-larval supply was higher during the warm season (February–April) than during the cold season (June–July) for both Brachyura (warm season: 85% of total post-larval supply) and Stomatopoda (warm season: 92%). Moreover, the pulse of the brachyuran post-larval supply occurred predominantly around the last quarter, while the pulse of stomatopods occurred predominantly around the new moon. However, for both taxa, the post-larval supply was lowest around the full moon and the first quarter. Overall, our monitoring highlighted that the post-larval supply of Brachyura and Stomatopoda was modulated by seasons and lunar phases at Rangiroa.     

Seasonal changes in otolith increment width trajectories and the effect of temperature on the daily growth rate of young sardines

We studied the otolith microstructure and growth of sardine, Sardina pilchardus, in the North Aegean Sea (eastern Mediterranean Sea), using samples of larvae and juveniles that had hatched in winter (November–January) and winter–spring (February–May), respectively. The juveniles had developed during an extended period coinciding with marked pelagic ecosystem changes (from winter, mixed conditions to summer, stratified waters). To examine the relationship between environmental changes and the observed variability in their otolith increment–width trajectories (width‐at‐age), we summarized the shape of trajectories with a four‐parameter set estimated from a growth model fit to each width trajectory. The individual parameter sets were then related to the potential oceanographic conditions that fish experienced during their development, derived from a hydrodynamic–biogeochemical model (POM‐ERSEM), implemented in the sampling area. Substantial seasonal effects were demonstrated on the otolith microstructure (platykurtic versus leptokurtic trajectories in winter‐mixed versus summer‐stratified conditions), which were related to the progressive sea surface warming. In a subsequent step, in order to study the effect of oceanographic conditions on larval and juvenile daily growth rates, a GAM (Generalized Additive Model) analysis of otolith increment widths was carried out, using model‐derived oceanographic parameters and taking into account the ‘inherent otolith growth’, expressed by the explanatory variables ‘previous increment width’ and ‘Age’. Results showed a strong and positive, linear effect of temperature on the growth rate of winter‐caught larvae, whereas in juveniles, which had developed within a wide range of temperatures, an optimum temperature for growth was observed at around 24°C.     

Annual cycle of clupeiform larvae around Gran Canaria Island, Canary Islands

The distribution and abundance of fish larvae was studied along the eastern and southern shelf of Gran Canaria Island (Canary Islands) from July 2000 to June 2001. Oblique bongo hauls were carried out fortnightly during the daytime, coinciding with days of full and new moon. During February, the area was sampled every 2–5 days. About 17.3% of the ichthyoplanktonic community was composed of clupeiform larvae: 92.9% of these larvae were Sardinella aurita, whereas 4.7% and 2.4% were respectively Engraulis encrasicolus and Sardina pilchardus. Sardinella aurita larvae appeared during the whole year with two periods of maximum abundance: June to September and December to February. During the full moon their abundance was on average 38.5% (±6.8%) of their numbers during the new moon, showing a clear lunar periodicity. Engraulis encrasicolus larvae appeared from November to March, also coinciding with the new moon. Sardina pilchardus larvae only appeared during two short periods, both coinciding with filaments shed from the African coastal upwelling which reached the island. This fact confirms the transport of fish larvae from the upwelling area off northwest Africa to the Canary Islands, promoting a genetic flow among both sites.     

Validation of daily increment formation in otoliths of immature and adult Japanese anchovy <Emphasis Type="Italic">Engraulis japonicus</Emphasis>

In order to validate daily increment formation in otoliths of immature and adult Japanese anchovy Engraulis japonicus, three rearing experiments using chemical marking of otoliths were conducted on adult anchovy in summer 2004 and immature anchovy in summer 2005 and in winter 2006. In the two experiments conducted in summer, the number of otolith microincrements between alizarin complexone (ALC) marks showed that microincrements were formed daily. In the summer 2005 experiment, immature anchovy under conditions of reduced daily food rations also showed daily microincrement formation. Average increment width was 0.9 μm in adults and 1.8–3.1 μm in immature anchovy. In contrast, no clear increments were observed between ALC marks on the otoliths from the experiment in winter 2006, and scanning electron microscope (SEM) observations failed to confirm clear increment formation. We consider that low water temperatures (<13–14°C) restricted otolith growth and lowered the contrast between the discontinuous and the incremental zones of the otolith increments. For age estimation of Japanese anchovy, clear increments wider than about 1 μm in the otolith can be regarded as daily increments. However, daily age estimation of immature and adult anchovy that experience low water temperatures in winter may be difficult due to the obscurity of the increments.     

Histological observations of seasonal reproductive and lunar-related spawning cycles in the female honeycomb grouper Epinephelus merra in Okinawan waters

Seasonal reproductive and lunar-related spawning cycles of the female honeycomb grouper Epinephelus merra inhabiting Okinawan waters were examined by histological observation of ovaries. Gonadosomatic index (GSI) increased beginning in May and peaked in June. Histological observations revealed that many oocytes laden with yolk were present in the ovaries from May to August. From September to October, ovaries were occupied by immature oocytes. These results suggest that the reproductive season of E.   merra lasts for 4 months from May through August in Okinawan waters. When the fish were collected according to the lunar cycle, GSI increased with the approach of the full moon. Oocytes at various development stages were observed from the first quarter to the full moon. Fresh ovulatory follicles were present in the ovaries around the last quarter moon. These results suggest that E.   merra has a lunar spawning cycle and spawns between the full moon and the last quarter moon. Ovarian features of the fish collected around the last quarter moon were different among individuals; some fish had many oocytes at the tertiary yolk stage in the ovaries, while the ovaries of the others were occupied by the oocytes at the peri-nucleolus and the oil droplet stages. This observation suggests that a minor release of eggs occurs in this species before or after a major spawning lunar day. Oocytes at the migratory nucleus and the maturation stages were not observed in any ovarian samples. This may mean that maturation of oocytes is related to the spawning behavior of this species and makes rapid progress in the process of aggregation at the spawning sites.     

Natural variability of fisheries and lunar illumination: a hypothesis

Long‐term synchronous trends in small pelagic fisheries catches around the world suggest that fish populations are governed by the same global climate forcings. Recent findings regarding the population dynamics of zooplankton during the lunar cycle in sub‐tropical waters may shed light on the influence of the lunar cycle on fish spawning and mortality. Here I hypothesize that, in the short‐term, observed changes in zooplankton abundance during the lunar cycle promote periods of enhanced feeding by adult fish and lower mortality (and increased growth) in their early planktonic stages. Furthermore, a striking 9‐year periodicity in sardine and anchovy mortality was inferred in four major upwelling areas, coinciding with the long‐term variability in lunar illumination. It is suggested that both short‐ and long‐term changes in lunar illumination should be considered when modelling the effect of climate on the natural variability of fisheries.     

Variability of larval Baltic sprat (Sprattus sprattus L.) otolith growth: a modeling approach combining spatially and temporally resolved biotic and abiotic environmental key variables

Plankton sampling was conducted in the Baltic to obtain sprat larvae. Their individual drift patterns were back‐calculated using a hydrodynamic model. The modelled positions along the individual drift trajectories were subsequently used to provide insight into the environmental conditions experienced by the larvae. Autocorrelation analysis revealed that successive otolith increment widths of individual larvae were not independent. Otolith increment width was then modelled using two different generalized additive model (GAM) analyses (with and without autocorrelation), using environmental variables determined for each modelled individual larval position as explanatory variables. The results indicate that otolith growth was not only influenced by the density of potential prey but was controlled by a number of simultaneously acting environmental factors. The final model, not considering autocorrelation, explained more than 80% of the variance of otolith growth, with larval age as a factor variable showing the strongest significant impact on otolith growth. Otolith growth was further explained by statistically significant ambient environmental factors such as temperature, bottom depth, prey density and turbulence. The GAM analysis, taking autocorrelation into account, explained almost 98% of the variability, with the previous otolith increment showing the strongest significant effect. Larval age as well as ambient temperature and prey abundance also had a significant effect. An alternative approach applied individual‐based model (IBM) simulations on larval drift, feeding, growth and survival starting as exogenously feeding larvae at the back‐calculated positions. The IBM results revealed optimal growth conditions for more than 97% of the larvae, with a tendency for our IBM to slightly overestimate larval growth.     

Responses in growth rate of larval northern anchovy (Engraulis mordax) to anomalous upwelling in the northern California Current

We examined variability in growth rate during the larval stage of northern anchovy (Engraulis mordax) in response to physical and biological environmental factors in 2005 and 2006. The onset of spring upwelling was anomalously delayed by 2–3 months until mid‐July in 2005; in contrast, spring upwelling in 2006 began as a normal year in the northern California Current. Larval and early juvenile E. mordax were collected in August, September, and October off the coast of Oregon and Washington. Hatch dates ranged from May to September, with peaks in June and August in 2005 and a peak in July in 2006, based on the number of otolith daily increments. Back‐calculated body length‐at‐age in the June 2005 hatch cohort was significantly smaller than in the August 2005 cohort, which had comparable growth to the July 2006 cohort. Standardized otolith daily increment widths as a proxy for seasonal variability in somatic growth rates in 2005 were negative until late July and then changed to positive with intensification of upwelling. The standardized increment width was a positive function of biomass of chlorophyll a concentration, and neritic cold‐water and oceanic subarctic copepod species sampled biweekly off Newport, Oregon. Our results suggest that delayed upwelling in 2005 resulted in low food availability and, consequently, reduced E. mordax larval growth rate in early summer, but once upwelling began in July, high food availability enhanced larval growth rate to that typical of a normal upwelling year (e.g., 2006) in the northern California Current.     

东海条石鲷仔鱼耳石日轮与生长的关系

2009年4月15日-5月5日在浙江省舟山水产研究所条石鲷(Oplegnathus fasciatus)人工繁殖期间,逐日选择胚胎和仔鱼样本,连续解剖观察发育后期胚胎和前期仔鱼,光镜观察仔鱼矢耳石和微耳石的形态、日轮数,测定其直径,研究条石鲷的耳石日轮和生长。结果表明,受精后约26h,条石鲷胚胎听囊内出现1对矢耳石和1对微耳石;仔鱼孵出第2天形成第1个轮纹,之后每天形成1轮,孵化后天数(N)和矢耳石日轮数(D)的关系为N=D+1;在第8天左右,矢耳石上出现第2条明显的标记轮,为初次摄食轮。仔鱼耳石长径(rs,μm)与鱼体体长(L,mm)呈线性相关,其关系式为rs=18.146L-44.436;矢耳石长径rs与微耳石长径rl之间存在线性相关,其关系式为rs=0.6125rl+1.9882。根据结果确认,矢耳石轮纹可作为条石鲷仔鱼日龄的判别依据。     

Growth of juvenile chub mackerel Scomber japonicus in the western North Pacific Ocean: with application and validation of otolith daily increment formation

The growth of juvenile chub mackerel Scomber japonicus collected in the western North Pacific Ocean in 2007 and 2009 was examined based on the evidence of otolith daily increment formation in captive specimens. There was a significant difference in the relationship between known age and number of increments in the frontal and sagittal planes. Repeated markings on the otolith using Alizarin complexone and the coefficient of variation in number of increments suggest that the increments in the frontal plane of the otolith are more suitable for age estimation than those in the long and short axes of the sagittal plane. The increments in the frontal plane formed daily, and the first ring was usually deposited 3 days after hatch. Age of wild juveniles ranged from 24 to 211 days after hatch based on the frontal plane method. The estimated hatching periods of specimens ranged from February to June, but the April-hatched specimens were collected throughout the sampling periods of 2007 and 2009. The Gompertz growth model showed a difference in growth pattern in specimens between 2007 and 2009. The juveniles in 2009 appeared to grow more quickly than those in 2007 until summer, but thereafter the 2009 specimens seemed to grow more slowly.     

Otolith development and daily increment formation in laboratory-reared larval and juvenile black-spot tuskfish <Emphasis Type="Italic">Choerodon schoenleinii</Emphasis>

Microstructures of lapilli were examined for reared larvae and juveniles of black-spot tuskfish Choerodon schoenleinii. Lapilli of larvae at 1 day after hatching have one diffuse and obscure ring with an otolith radius of 4.3 ± 0.50 μm (mean ± SD, N = 8). The slope and intercept of the regression between the number of days after hatching and increment counts did not differ significantly from one and zero, respectively, indicating that lapillus increments were formed on a daily basis after hatching. There was an ontogenetic shift in the relative values of somatic and otolith growth, which corresponded to the transition from pelagic larvae to settlement stage. Simultaneously, the daily increment width reached the maximum value. These findings suggest that age at maximum value of increment width can be used as an indicator of the planktonic larval duration while settlement mark is not found. Since ontogenetic shift in the relationship between otolith radius and body size was observed, back-calculation of somatic growth in black-spot tuskfish using the otolith radius during the early life stages should be analyzed with caution, and the method requires further validation.     

Validation of otolith increment daily periodicity in captive juvenile sablefish (Anoplopoma fimbria) experimentally immersed in strontium chloride (SrCl2)

We used a chemical marking experiment to validate the daily periodicity of otolith increment deposition in juvenile sablefish, Anoplopoma fimbria. The sagittal otoliths of 26 live juvenile sablefish were marked twice by immersion in saltwater baths containing elevated levels of strontium chloride (SrCl₂) during June–August of their first year of life (age 0). The number of otolith increments detected between strontium bands from three readings (median 15–20) was compared to the number of days between marking events (15–17 days). Median discrepancies between otolith increment counts and days between strontium bands were small (mean 1.1, S.E. 0.81, n = 20) and suggest that otolith increment counts from age 0 juvenile sablefish may provide a useful proxy for daily age. However, median discrepancies in June (mean 4.7, S.E. 0.65, n = 9 otoliths) were significantly (^*P ≤ 0.05) higher than those in July (mean −1.8, S.E. 0.40, n = 6) and August (mean −1.8, S.E. 0.66, n = 5). Increment banding patterns were more difficult to identify in otoliths marked in June (mean sablefish size 84.9 mm FL) because of changes in the structure of the sagittae associated with the formation of accessory primordia.