A description of four types of otolith opaque zone

Analysis of the alternating opaque and translucent zones in fish otoliths is a widely used method to determine age in fish. The mechanisms underlying the annual periodicities in the formation of these zones remain unknown, although various interpretations and explanations have been presented to explain the appearance of the opaque zones. Here I review the biological and structural characteristics of the opaque zones in otoliths from a number of fish species. The results of this review suggest that the opaque zones can be classified into four different types. Type A is a dark opaque zone, displaying minute, dense crystals, and is typically formed at a young life-history stage. Type B comprises an aggregation of grooves and discontinuous crystals that are formed during growth-stagnant periods. Type C can be described as a washy black zone that has the appearance of an object that is smeared with ink; it is formed during seasons of active growth. Type D has deep grooves, appears luminous in transmitted light in etched otolith sections, and is formed during the spawning season. Types A and C otolith opaque zones are typical of younger fish and are formed during growing periods. They are complementary to Types B and D otolith opaque zones which are formed during periods of stagnant growth and/or during the spawning seasons and which are typical of older fish, which generally lack Type A zones. To ensure precise age determination in fish, it is therefore necessary to understand the structural and biological characteristics that produce these four distinct types of opaque zones.     

Age validation,growth and mortality of introduced Tilapia zillii in Crater Lake Nkuruba,Uganda

Periodicity and timing of opaque zone formation in otoliths of introduced redbelly tilapia, Tilapia zillii (Gervais), in Crater Lake Nkuruba, Uganda, were validated using marginal increment. Age and growth were assessed through readings of biannuli in thin‐sectioned sagittal otoliths. Deposition of opaque zone formation in T. zillii otoliths was bimodal (March–May and September–November), corresponding to two seasonal peaks of precipitation characteristic of this equatorial region. Ages of T. zillii ranged from 2 to 8 years, with fish gillnetted offshore having a faster growth and attaining larger size‐at‐age than fish captured inshore in minnow traps, suggesting that use of multiple gears is needed when estimating the growth of T. zillii. Total instantaneous mortality (Z), estimated using catch curve analysis, was 0.74 for gillnetted fish and 0.71 for trapped fish. These estimates were at the low end of the total mortality reported for other tilapia species. Natural mortality (M) was estimated as 0.52–0.54 by applying Rikhter and Efanov's maximum age at maturity and Hoenig's maximum age methods, respectively. Fishing mortality (F) in Lake Nkuruba was 0.17–0.22, indicating a low exploitation level in the lake.     

Age determination and growth of Pacific bluefin tuna, Thunnus orientalis, off Japan and Taiwan

Age determination of wild captured Pacific bluefin tuna, Thunnus orientalis, was conducted using sagittal otoliths of 806 specimens (47–260 cm in fork length) caught in the waters off Japan and Taiwan. Otoliths were transversely sectioned and the opaque and translucent zones were analyzed. Opaque zones mainly appeared on the otolith edge from April to July, indicating that the opaque zone is formed annually. The opaque zones formed during later life (age 10+) were more distinct than the earlier zones. The estimated ages of specimens ranged from 1 to 26 years. Parameters of the von Bertalanffy growth function were estimated to be 249.6 cm, 0.173, and −0.254 years for L_∞, k, and t₀, respectively. Growth of younger fish was rapid up to 5 years old attaining about 150 cm, and then growth rate decreased. After that, fish attained about 200 cm at 9 years old and about 225 cm (90% of L_∞) at 13 years old (50% of maximum age). This paper updates the biological information on length at age with a large size range to support stock assessment model analyses for this commercially valuable species.     

Validation of the otoliths for age determination in Barbus sclateri (Günth.)

Abstract – Otoliths were used to explore the timing of annulus formation in Barbus sclateri. Lapillus exhibited opaque and translucent zones whose combination was formed once a year. Opaque zones appeared formed from July and January, and the translucent ones from February to June. We describe a procedure to assess age by using the number of annulii, the condition of the otolith's edge and the capture date. The use of otoliths for age determination in this cyprinid endemic to the Iberian Peninsula provides a useful alternative to the traditional methods of scale reading. ^Note     

Age and changes in growth of the king weakfish Macrodon atricauda (Günther, 1880) between 1977 and 2009 in southern Brazil

The coastal demersal sciaenid Macrodon atricauda (Günther, 1880), formerly M. ancylodon (Bloch and Schneider, 1801) was sampled for ageing during four periods (1977–1979, 1984–1986, 1997–1998 and 2006–2009) in commercial fishing and scientific surveys along southern Brazil (Lat. 30°S–34°40′S). Maximum observed age was seven years, but no fish over five years old was sampled in the last period. Marginal increment analysis of thin sections validated ageing and showed that opaque and translucent bands were laid down at all ages in spring–summer and autumn–winter, respectively. Ageing M. atricauda based on sectioned otoliths is highly recommended because comparisons with readings on whole otoliths showed that ages based on whole otoliths exceeded those based on sectioned otoliths for 56.5% of the aged specimens. The growth of M. atricauda has increased in the last four decades, most noticeably in the case of adult males over two years old and females over three years old. A threefold decrease in its density and the demersal fish community as a whole are the most likely causes of the growth increase.     

Identification of hatchery-reared and naturally produced Atlantic salmon, Salmo salar L., juveniles based on examination of otoliths

. Atlantic salmon, Salmo salar L., parr and smolts from three Norwegian rivers were examined with respect to whether or not they had been released from a hatchery. The hatchery background was known for some released fish and could be ascertained for others from their aberrant body morphology and eroded fins. Fish released after one winter in a hatchery had opaque otoliths like the fish examined from two hatcheries. Naturally produced fish showed a distinct, seasonal growth pattern in the otoliths, alternating between opaque summer zones and hyaline winter zones. Fish released as fry showed an otolith pattern similar to that of naturally produced fish. A test revealed little discrepancy between two independent otolith readings even though the test reading was conducted without any information about the fish accompanying the otoliths. The results suggest that examination of otoliths may help distinguish between juveniles that are hatchery-reared and juveniles that are naturally produced in the river, provided that the hatchery-reared fish have a 1-year history in the hatchery prior to release.     

Age, growth and life history of gunnel, Pholis fangi, in the Yellow Sea

We examined the formation of annuli by marginal observations on otoliths of gunnel (Pholis fangi) in the Yellow Sea to validate the age determination method and to derive the growth equation covering from larval to adult stages. Gunnels, ranging from 46 to 173 mm in total length, were collected by a bag net fishery from the western coastal waters off Korea from November 1998 to October 1999. Marginal observations indicated that the translucent zone (annual mark) on adult otolith was formed during the winter, whereas the opaque zone was formed during the summer. However, a translucent zone was formed between May and June in juvenile otoliths. This false ring was formed when the fish transited from the inshore pelagic life of larvae to the offshore bottom life of juveniles. The observed maximum age was 58 months. Using observed length-at-monthly age, growth in length was expressed by von Bertalanffy growth curve; L_t = 144.0 (1 − e^{−0.11 (t+0.43)}). P. fangi spawned in winter recruit to inshore, and grow quickly in the nursery habitats in spring. Gunnel inhabit the bottom offshore area during the summer season, and reappear inshore thereafter.     

大黄鱼仔稚鱼不同发育阶段矢耳石形态发育和微结构特征

对人工培育大黄鱼(Larimichthys crocea)的生长发育与矢耳石形态及微结构特征进行研究,结果表明:(1)大黄鱼矢耳石上的轮纹是每日形成的,第1日轮在孵化后第2天形成,与其初次摄食相对应。(2)大黄鱼卵黄囊期和前弯曲期仔鱼的耳石形态为圆形,进入弯曲期耳石长轴迅速伸长,在后弯曲期耳石形态变为椭圆形。进入稚鱼期,矢耳石开始形成次生核。随后次生核数量逐渐增加,在孵化后47～78 d的个体中,次生核数量稳定在5～7个,耳石近似盾形。(3)根据耳石日轮宽度推算的大黄鱼稚鱼在其仔鱼期生长率(b)与第1个次生核的形成时间(tsp1)之间存在明显的线性关系,表明生长较快的个体形成次生核的时间较早,进入稚鱼期所需的时间更短。以上结论表明,大黄鱼矢耳石可以反演其早期生活史阶段的生长发育特征。     

Age validation,growth and annual reproductive cycle of chub mackerel <Emphasis Type="Italic">Scomber japonicus</Emphasis> off the waters of northern Kyushu and in the East China Sea

The age and growth of chub mackerel Scomber japonicus collected from the East China Sea and the northern waters off Kyushu between June 2000 and June 2001 were determined by observing the otolith surface after dipping it in xylene. The translucent and opaque zones on the otolith surface were identified, and the number of translucent zones was counted. Monthly changes in the frequency of fish with translucent zones on the otolith margin, and in the marginal increments, indicated that the translucent zones were formed between April and June. The seasonal pattern of annulus formation on the otolith became clear by observing the otoliths of fish with known ages, and the otolith formation in wild fish was consistent with that of fish with known ages. The mean gonadosomatic index of male and female fish was high from March to May, and spawning females were observed from mid-March to mid-May. The estimated ages were 1–5 years for males and 1–6 years for females. The von Bertalanffy growth parameters did not significantly differ between male and female. The model was obtained as FL _t=406×{1−exp[−0.372×(t+1.68)]     

Does otolith macrostructure record environmental or biological events? The case of black hake (Merluccius polli and Merluccius senegalensis)

Fish age determination using otoliths requires a prior understanding of growth mark deposition patterns (translucent rings, TR) as well as their connection with internal or external events experienced by the fish. This study analysed the macrostructural seasonal ring deposition pattern observed in transversal sections of black hake otoliths. A total of 793 black hake otoliths were collected in autumn and spring 2007 from research and commercial surveys carried out in continental and shelf waters off Mauritania. Most of the Merluccius polli otoliths presented narrow and wide translucent rings (NTR and WTR, respectively) regardless of fish size, whereas Merluccius senegalensis otoliths only showed NTR. This seemed to be a sign of ontogenetic discrepancy between the two black hake species, whose otoliths confirmed the existence of significant differences in their growth patterns.The frequency distributions of the number of TR counted along the ventral radius (VR) of the otolith from the nucleus (birth date) and from the ventral edge (death date) were analysed to ascertain whether a specific endogenous event (Hypothesis A) or a precise environmental event (Hypothesis B) could restrain growth leading to the formation of TR. The general TR frequency distribution pattern was somewhat similar for both hake species, which showed marked TR at comparable distances. Within each species, TR frequency distributions of their distances from the nucleus along the otolith VR were quite similar between sexes, seasons, and fish sizes. Our results support the idea of a coincident biological event, such as first maturity, slowing down the growth process and thus provoking the formation of TR in otoliths of both species.This study also experience difficulty using the complex and highly variable macrostructural pattern of black hake otoliths to establish age interpretation criteria for these two species.     

Age and growth of the lizardfish <Emphasis Type="Italic">Saurida elongata</Emphasis> from the Tsushima/Korea Strait

We studied the age and growth of the lizardfish Saurida elongata using ground thin sections of otoliths (sagittae) from specimens collected in the Tsushima/Korea Strait between May 1999 and June 2001. A total of 695 individuals with fork lengths (FL) ranging from 189 to 478 mm were examined. The frequency of translucent zone occurrence at the outer margin of the otoliths indicated that translucent zones formed once a year between November and February. Most of the males examined were estimated to be 2–7 years old and the females, 3–9 years old. The maximum estimated age of a male specimen was 10 years and that of a female, 11 years. The estimated von Bertalanffy growth curves were FL _t  = 451{1 − exp[−0.172(t + 2.50)]} and FL _t  = 515{1 − exp[−0.151(t + 2.47)]} for males and females, respectively. At all ages, the FL at a specific age for females was greater than that for males, suggesting that females of this species grow faster than males.     

Geographic influences on and the accuracy and precision of age estimates for the red bass, Lutjanus bohar (Forsskal 1775): A large tropical reef fish

The red bass is a large tropical reef fish (Lutjanidae, tropical snappers) that is harvested to varying extents throughout a widespread Indo-Pacific distribution. The aims of this study were to investigate the accuracy and precision of age estimates from transverse sections of sagittal otoliths and to assess effects on these of the geographic area of collection and otolith preparation method. Two independent validation studies suggested an approximately annual formation of annuli in otoliths, predominantly for otoliths with 4–10 annuli but also for one otolith with 29 annuli. Otolith sections produced exceptionally high annulus counts: up to 56 annuli for samples from the Great Barrier Reef (GBR), Australia; and up to 55 annuli from the Seychelles, indicating a high longevity for this species. The precision of otolith readings from the GBR (index of average percent error, IAPE = 3.21 ± 0.26 S.E.) was within commonly accepted bounds for age estimation (IAPE up to 5%) but precision of readings from the Seychelles was significantly lower (IAPE = 9.18 ± 0.47 S.E.) and outside of this “acceptable” range. Age-based biological parameters for red bass from the Seychelles should thus be applied with greater caution than those for red bass from the GBR. If basic demographic properties are assumed relatively constant across the wide geographic range sampled, however, results from the GBR could be used as more reliable preliminary data for precautionary management strategies in the Seychelles and elsewhere.     

Somatic growth and otolith microstructure of larval and juvenile pacific cod <Emphasis Type="Italic">Gadus macrocephalus</Emphasis>

ABSTRACT:   Microstructures of sagittae and lapilli were examined in relation to somatic growth for reared larvae and juveniles of Pacific cod. The Laird–Gompertz model was fitted to the daily age and somatic growth relationship. Growth increments were deposited on a daily basis in both kinds of otoliths, with a check formed at hatching. Two subsequent checks and an accessory primordia (AP) occurred in the sagittae. The lapillus was adequate for increment width measurement through the early life stages. Sagittal and frontal plane of sagitta was adequate for measurement in the pre-AP and post-AP formation stages, respectively. The shift of desirable plane was caused by changes in otolith and increment shapes with AP formation. Back-calculated total lengths using the biological intercept method did not significantly differ with certain body lengths ( P  > 0.05), suggesting validity of back-calculation in this species. Using the back-calculated total length, morphological and ecologic changes that seemed to affect checks and AP formations are discussed.     

Age and maturation of jack mackerel Trachurus japonicus in the East China Sea

Growth and reproductive characteristics of jack mackerel Trachurus japonicus collected in the East China Sea were determined based on otolith readings and gonad histology, respectively. Translucent and opaque zones on sectioned otoliths were identified and opaque rings counted. Von Bertalanffy growth parameters did not significantly differ between males and females, and the combined growth curve was: L _t  = 401{1 ? exp[?0.275 (t + 1.149)]} (0.8 < t < 6.9), where L _t is fork length (mm) at age t. The calculated lengths at age 1 in our study were larger than those reported 50 years ago from the East China Sea. The spawning period was evaluated to be from December to June, but primarily from February to May, based on the monthly changes in the gonadosomatic index and histological observations. The minimum size and age at first maturity were smaller and younger, respectively, than those reported in previous studies.     

元江水系越鳠年龄鉴定材料的比较

分析比较了元江越鳠(Mystus pluriradiatus)不同年龄鉴定材料的鉴定结果。结果显示:微耳石磨片、星耳石和脊椎骨材料为鉴定元江越鳠年龄的可利用材料,矢耳石细小且脆不适合用于越鳠的年龄鉴定。微耳石磨片和星耳石的年龄判别能力最高,均达到96.0%,脊椎骨的年龄判别能力为93.3%;最大年龄读数出现在微耳石磨片和脊椎骨上,读数为5+龄;微耳石磨片的同一鉴定者两次独立年龄读数的吻合率最高为94.5%、脊椎骨次之为91.7%、星耳石最低为88.1%;两种年龄材料读数之间的吻合率比较中,微耳石磨片与星耳石的吻合率较高为90.9%,两者的年龄读数在低龄组时吻合较好,微耳石磨片与脊椎骨的吻合率为89.3%,3+龄以上时二者的年龄读数吻合率高。结果表明,鉴定低龄个体选择星耳石材料(n+龄≤3+龄),高龄个体用微耳石磨片进行辅助鉴定的年龄鉴定方法对元江越鳠进行年龄鉴定较为准确。     

Mass-processing method for aging otoliths of walleye pollock <Emphasis Type="Italic">Theragra chalcogramma</Emphasis> using black resin

ABSTRACT:   The reliability of a mass-processing aging method for walleye pollock Theragra chalcogramma was examined. The method is a half-otolith method, in which 10–20 otoliths are embedded simultaneously in black resin and the mass-processed half plane section is viewed under reflected light (black-resin method). Seasonal periodicity of otolith growth was confirmed by the monthly observations of the percent of translucent zone at the otolith edge, and interannual variation in age distribution detected the progression of dominant year-classes. These results support the assumption that the translucent zone is formed once a year. The black-resin method was compared to the break–burn method, which is the standard technique for this species. The estimated ages of these methods were not biased throughout the age range examined. The coefficient of variation in the black-resin method (4.9–5.7%) was similar to those of the break–burn method (3.5–5.5%). These results suggest that the black-resin method is reliable as an aging method for walleye pollock.     

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.     

Validation of otolith daily increments for larval and juvenile Japanese halfbeak Hyporhamphus sajori

Daily ring formation was verified by examining the growth of the marginal increment on sagittal otoliths of larval and juvenile Japanese halfbeak Hyporhamphus sajori. The relationship between age (x) and number of increments (y) is y = 1.0x + 2.0. The first daily increment was formed during the 2 days before hatching. The relationship between the number of rings deposited after the alizarin complexone (ALC) mark (x) and the number of increments (y) is given by: y = 1.0x − 0.2. The index of completion of the marginal increment was 99 ± 4.1% (mean ± S.D.) at 04:00, and from 24 ± 4.4% at 08:00 it increased with time of day and reached 98 ± 3.6% until the next day at 04:00. Growth of the incremental zone started a few hours after sunrise prior to which the discontinuous zone seemed to be formed. Light rhythms tend to be one of the most important factors for the formation of the marginal increment on otoliths. Based on the relationships between time of day and the marginal increment on otoliths, it would be possible to estimate the predation time for specimens retrieved from stomach contents, and also clearer analysis of the growth history immediately prior to the sampling time.     

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.     

基于微卫星标记的图们江大麻哈鱼亲子鉴定技术研究

放流标记技术研究是支撑增殖放流效果评估的重要内容。该研究用质量浓度为100 mg·L⁻¹的茜素络合物 (ALC) 对约2月龄的青鱼 (Mylopharyngodon piceus) 幼鱼进行48 h浸泡,随后移入正常水体中恢复续养20 d。对其3种耳石在可见光和不同的激发光下进行比较分析,评价了茜素络合物在不同耳石上的沉积情况与标记效果。结果表明,通过在可见光和不同的激发光下进行观察,其3种耳石上均可见明显的标记区,标记率达100%。在对标记鱼恢复续养20 d后发现,耳石上的荧光标记区强度仍很高,表现出了较好的标记效果。此外,相关标记的最佳检测光源为蓝色激发光,绿色激发光次之。微耳石是用于青鱼幼鱼耳石荧光标记观察的最适类型,同时也是用于时滞检测的最适类型,其次是星耳石。此外,将鱼体浸泡入茜素络合物溶液中至其在耳石上形成标记沉积,以及将鱼体移出浸泡溶液至其耳石上标记区域消失,均存在1 d时滞。