Otolith Area Research Articles

Using otolith shape and morphometry to identify four Alburnus species (A. chalcoides, A. escherichii, A. mossulensis and A. tarichi ) in Turkish inland waters

Asteriscus otolith shapes as well as their morphometry and shape contours were investigated in order to identify four allopatric Alburnus species: A. chalcoides (Güldenstädt, 1772) (Ordu), A. escherichii Steindachner, 1897 (Eskişehir), A. mossulensis Heckel, 1843 (Tunceli), and A. tarichi (Güldenstädt, 1814) (Van) in Turkish inland waters. These were compared using the shape indices (form factor, roundness, circularity, ellipticity, rectangularity and aspect ratio), and the morphological characters [otolith weight (OWE), otolith length (OL), otolith width (OW), otolith perimeter (OP), and otolith area (OA)]. The overall canonical discriminant analysis (CDA) classification score was 93.8%, with the lowest score for A. escherichii (82.5%) and the highest for A. chalcoides (100%). The otolith shapes, morphology and shape contours of all sampled fish were a clear species differentiator, thereby demonstrating that the otolith shape is species-specific. The current study presents for the first time comprehensive variation information on interspecific left-right asteriscus otoliths in males and females of each Alburnus species: A. chalcoides from Ordu, A. escherichii from Eskişehir, A. mossulensis from Tunceli and A. tarichi from Van, based on a total of 307 individuals. Scanning electron microscopy (SEM) images, shape contours and other otolith characters vary within the same genus; these differences should be investigated not only in other freshwater fish species or genera but also in the same species living in different habitats. In addition, further investigation is required not only with respect to the morphometry, biometry, shape, geometry, and shape contours of the otoliths, but also regarding the genetic methods for robust identification of various sympatric and allopatric fish populations.

Effects of CO<sub>2</sub>-driven ocean acidification on early life stages of marine medaka (<i>Oryzias melastigma</i>)

Abstract. The potential effects of high CO2 and associated ocean acidification (OA) in marine fishes and other non-calcified organisms are less well understood. In this study, we investigated the responses of early life stages (ELS) of marine medaka (Oryzias melastigma) exposed to a series of experimental manipulation of CO2 levels. Results showed that CO2-driven seawater acidification (pH 7.6 and pH 7.2) had no detectable effect on hatching time, hatching rate, or heart rate of embryos. However, the deformity rate of larvae in the pH 7.2 treatment was significantly higher than that in the control treatment. There is no significant difference between the left and right otolith areas in each treatment. However, the average otolith area of larvae in the pH 7.6 treatment was significantly smaller than that in the control. Such alterations in the developmental abnormalities and otolith size of marine medaka larvae due to elevated-CO2 levels suggests that this species will be increasingly challenged by future OA. Further studies of the impacts of OA on marine fish to assess whether or not the environmental influence in one generation can affect the later life history and the phenotype of subsequent generations are needed.

The migratory history of anadromous and non-anadromous tapertail anchovy Coilia nasus in the Yangtze River Estuary revealed by the otolith Sr:Ca ratio

The migratory history of tapertail anchovy Coilia nasus in the Yangtze River Estuary, China was investigated using otolith Sr:Ca ratios and two-dimensional images of the Sr level from an X-ray electron probe microanalyzer (EPMA). The results showed that 17 of the 22 young-of-the-year (YOY) specimens had low Sr:Ca ratios (1.2–2.4 × 10−3;1.5 ± 0.3 × 10−3) at the central otolith area, indicating their riverine origin and initial freshwater residence. In addition, 11 of the 14 adult specimens had low Sr:Ca ratios (1.3–2.2 × 10−3; 1.7 ± 0.4 × 10−3) at the central otolith area but showed alternating changes between high (>4.0 × 10−3) and low (<2.5 × 10−3) values outside of this region, reflecting their riverine origin and the migration between freshwater and estuarine habitats. These 28 specimens represented the anadromous population in this region. The other 5 YOY specimens had high Sr:Ca ratios (3.6–5.9 × 10−3; 4.8 ± 0.8 × 10−3) throughout the life history. Similarly, the other 3 adult specimens had high Sr:Ca ratios (4.0–5.7 × 10−3; 4.8 ± 0.7 × 10−3) at the central otolith area but showed alternating changes between low and high values outside this region, suggesting that estuarine-origin non-anadromous individuals occurred in this region. The average of the otolith Sr:Ca ratios and Sr level mapping along the life-history transects could be used as a scalar for charting the migratory history of the tapertail anchovy in the Yangtze River Estuary: <2.0 × 10−3 for freshwater residence and 3.5–6.0 × 10−3 for estuarine residence.

Effect of ocean acidification on early life stages of Atlantic herring (&amp;lt;i&amp;gt;Clupea harengus&amp;lt;/i&amp;gt; L.)

Abstract. Due to atmospheric accumulation of anthropogenic CO2 the partial pressure of carbon dioxide (pCO2) in surface seawater increases and the pH decreases. This process known as ocean acidification might have severe effects on marine organisms and ecosystems. The present study addresses the effect of ocean acidification on early developmental stages, the most sensitive stages in life history, of the Atlantic herring (Clupea harengus L.). Eggs of the Atlantic herring were fertilized and incubated in artificially acidified seawater (pCO2 1260, 1859, 2626, 2903, 4635 μatm) and a control treatment (pCO2 480 μatm) until the main hatch of herring larvae occurred. The development of the embryos was monitored daily and newly hatched larvae were sampled to analyze their morphometrics, and their condition by measuring the RNA/DNA ratios. Elevated pCO2 neither affected the embryogenesis nor the hatch rate. Furthermore the results showed no linear relationship between pCO2 and total length, dry weight, yolk sac area and otolith area of the newly hatched larvae. For pCO2 and RNA/DNA ratio, however, a significant negative linear relationship was found. The RNA concentration at hatching was reduced at higher pCO2 levels, which could lead to a decreased protein biosynthesis. The results indicate that an increased pCO2 can affect the metabolism of herring embryos negatively. Accordingly, further somatic growth of the larvae could be reduced. This can have consequences for the larval fish, since smaller and slow growing individuals have a lower survival potential due to lower feeding success and increased predation mortality. The regulatory mechanisms necessary to compensate for effects of hypercapnia could therefore lead to lower larval survival. Since the recruitment of fish seems to be determined during the early life stages, future research on the factors influencing these stages are of great importance in fisheries science.

Effect of ocean acidification on otolith development in larvae of a tropical marine fish

Abstract. Calcification in many invertebrate species is predicted to decline due to ocean acidification. The potential effects of elevated CO2 and reduced carbonate saturation state on other species, such as fish, are less well understood. Fish otoliths (earbones) are composed of aragonite, and thus, might be susceptible to either the reduced availability of carbonate ions in seawater at low pH, or to changes in extracellular concentrations of bicarbonate and carbonate ions caused by acid-base regulation in fish exposed to high pCO2. We reared larvae of the clownfish Amphiprion percula from hatching to settlement at three pHNBS and pCO2 levels (control: ~pH 8.15 and 404 μatm CO2; intermediate: pH 7.8 and 1050 μatm CO2; extreme: pH 7.6 and 1721 μatm CO2) to test the possible effects of ocean acidification on otolith development. There was no effect of the intermediate treatment (pH 7.8 and 1050 μatm CO2) on otolith size, shape, symmetry between left and right otoliths, or otolith elemental chemistry, compared with controls. However, in the more extreme treatment (pH 7.6 and 1721 μatm CO2) otolith area and maximum length were larger than controls, although no other traits were significantly affected. Our results support the hypothesis that pH regulation in the otolith endolymph can lead to increased precipitation of CaCO3 in otoliths of larval fish exposed to elevated CO2, as proposed by an earlier study, however, our results also show that sensitivity varies considerably among species. Importantly, our results suggest that otolith development in clownfishes is robust to even the more pessimistic changes in ocean chemistry predicted to occur by 2100.

Stock discrimination of spottedtail goby (Synechogobius ommaturus) in the Yellow Sea by analysis of otolith shape

Otolith shape is species specific and is an ideal marker of fish population affiliation. In this study, otolith shape of spottedtail goby Synechogobius ommaturus is used to identify stocks in different spawning locations in the Yellow Sea. The main objectives of this study are to explore the potential existence of local stocks of spottedtail goby in the Yellow Sea by analysis of otolith shape, and to investigate ambient impacts on otolith shape. Spottedtail goby was sampled in five locations in the Yellow Sea in 2007 and 2008. Otoliths are described using variables correlated to size (otolith area, perimeter, length, width, and weight) and shape (rectangularity, circularity, and 20 Fourier harmonics). Only standardized otolith variables are used so that the effect of otolith size on the shape variables could be eliminated. There is no significant difference among variables of sex, year, and side (left and right). However, the otolith shapes of the spring stocks and the autumn stocks differ significantly. Otolith shape differences are greater among locations than between years. Correct classification rate of spottedtail goby with the otolith shape at different sampling locations range from 29.7%–77.4%.

Use of otolith shape for the identification of trumpeter sillago (Sillago maculata) and silver sillago (Sillago sihama)

Otolith shape is species-specific in fish. In this study, 161 samples of Trumpeter sillago Sillago maculata and 164 of Silver sillago Sillago sihama were collected from Beibu Gulf in July 2009. The main objective of this study was to use morphological variables and Fourier harmonics to describe otolith characteristics and use discriminant analyses to separate the two species. Otoliths were measured by traditional one and two dimensional measures (otolith length, width, diameters, area, perimeter, rectangularity and circularity), as well as by Fourier analysis to capture the finer regions of the otolith. Analysis of covariance (ANCOVA) showed that there was significant correlation between morphological variables (diameter, perimeter, otolith length, otolith width, otolith area, density, harmonics 3, harmonics 4, harmonics 5, harmonics 13, harmonics 15, and harmonics 16) and body length. To minimize size effects on the morphological variables between species, only fish with a body length between 90 and 140 mm were included in the data analysis and the variables which had significant relation with body length were transformed using the residual. The result showed that the accuracy of discriminant analysis was 97.8% and 100% for Trumpeter sillago and Silver sillago, respectively. This high accuracy indicated that otolith shape was described accurately by morphological variables and Fourier harmonics, and discriminant analysis was an effective way to identify and separate the two species.

Replenishment success linked to fluctuating asymmetry in larval fish

Fluctuating asymmetry (FA), defined as random deviations from perfect symmetry, has become a popular tool with which to examine the effects of stress during the development of bilaterally symmetrical organisms. Recent studies have suggested that FA in otoliths may serve as an indicator of stress in fish larvae. We examined the relationship between otolith asymmetry and temporal patterns in the occurrence of late-stage larvae to a tropical reef (i.e. replenishment) for the Caribbean lizardfish, Saurida suspicio (family Synodontidae). Late-stage larvae were collected in light traps over a period of 18 consecutive lunar months in the San Blas Archipelago, Panama. Asymmetry within otolith pairs was calculated from 24 variables: area, perimeter, longest and shortest axis of the otolith and 20 shape descriptors (Fourier harmonics). Otolith asymmetry was correlated strongly with fluctuations in lunar light trap catches. Two measured variables, otolith area and one of the 20 shape descriptors, accounted for 60% of the variability in lunar replenishment of S. suspicio. Individuals from small replenishment pulses exhibited higher levels of asymmetry compared to larvae from large pulses. When dry and wet seasons were analysed separately, otolith asymmetry explained a surprising 70 and 97% of the variation, respectively. Although the generality of these results remain to be tested among other populations and species, otolith asymmetry may be an important indicator, and potentially a predictor, of larval quality and replenishment success.

Eco-morphological patterns of the sagitta of Antarctic fish

Morphology and morphometry of the sagittae otolith were studied in pelagic and mesopelagic fish. The shape, margins and rostrum of four groups of otoliths from several species were analyzed: group 1 (pelagic fish associated with the under ice cover N = 42), group 2 (pelagic fish associated with water offshore N = 9), group 3 (mesopelagic fish associated with extensive vertical migration N = 57) and group 4 (mesopelagic fish associated with short vertical migration N = 54). E (maximum width of the sagitta /maximum length of the sagitta %), R (rostrum length (RL)/maximum length of the sagitta %) and S (sulcus area (SS)/otolith area (OS) %) indexes were calculated for each species. Sagittae of pelagic groups (1 and 2) showed the smallest sagitta dimensions in relation to the total length of the fish, in this group the sagitta shape is variable. Sagittae of mesopelagic fish (groups 3 and 4) showed variable shape and edges. The shape in group 4 was polygonal and these species have more width than length. Statistical analysis showed significant differences in the E, R and S indexes. These results were compared with other 19 species, belonging to six families, taken from a publisher-edited literature. E, R and S-values could be used to characterize the sagittae of the Antarctic fish and could be considered as a useful tool for fish ecology studies.

Fluctuating asymmetry in the otoliths of larval Atlantic menhaden Brevoortia tyrannus (Latrobe) – a condition indicator?

Fluctuating asymmetry (FA) was described for otolith length, width and area for two groups of larval and juvenile Atlantic menhaden Brevoortia tyrannus. The larvae and juveniles (age range: 20–120 days; standard length, LS, range: 14–35 mm) were separated into the two distinct groups a priori on the basis of their growth indices estimated from residuals of a LS‐at‐age model; slow growers had residuals &lt;8% of the average size at age and fast growers had residuals &gt;8%. Some indication supporting the fluctuating asymmetry methodology was found when unsigned normalized FA2 index was used for otolith area. That conclusion was not, however, supported by the other method used in this work, i.e. between‐group ANCOVA analysis of individual unsigned and not normalized FA1 indices plotted against trait size. These inconsistent results are most probably a consequence of error introduced during FA index normalization (accounting for trait size effect on absolute FA). Although it is concluded here that FA analysis of sagittal otoliths was not effective in describing differences in the natural variability of growth and condition of larval Atlantic menhaden, the possible species‐ and otolith type‐related differences should be considered in the future research on FA.

Variation in otolith macrostructure of Japanese flounder ( Paralichthys olivaceus): A method to discriminate between wild and released fish

The main objective of this study was to develop a method to discriminate between wild and hatchery-produced Japanese flounder, Paralichthys olivaceus, based on variations in otolith macrostructure. Otoliths of wild flounder were more elliptical than those of hatchery-produced fish, whereas otolith area and marginal coarseness showed no clear differences. Otolith morphometry did not vary significantly with water temperature or feeding conditions in rearing experiments. Reduced ellipticity in the otoliths of hatchery-produced fish could be caused by biotic and abiotic conditions after release. Throughout the study, it was found that otoliths of Japanese flounder reared at 15 and 20 °C regimes showed opaque zones regardless of feeding condition, while otolith of fish reared at 25 °C had translucent zones. The potential of thermal marks and secondary zones as a new mass-marking system is presented.

Comparison of otolith growth and morphology with somatic growth and age in young‐of‐the‐year bluefin tuna

Otolith morphological characteristics were studied using image analysis techniques and the relationships between otolith growth and somatic growth and age, as estimated from counting daily otolith increments, were examined in young‐of‐the‐year (YOY) bluefin tuna Thunnus thynnus ranging in fork length (LF) from 8·5 to 55·5 cm. Whole otolith length, width, area and perimeter, and three shape indexes, circularity, E value and rectangularity, were extracted for each pair of sagittae. Since no statistical significant differences between left and right otolith morphometrics were found, only one otolith from each fish was used for correlations. Statistically significant relationships were observed between otoliths measurements and fish somatic growth when a linear regression was applied after logarithmic transformation of all variables tested. Among the variables, otolith length was the one that showed the highest correlation with LF, followed by otolith area and perimeter, whereas otolith rectangularity exhibited the lowest correlation. Statistically significant relationships were also observed between the otolith variables tested and the age of the fish, which ranged from 20 to 129 days. The ages estimated using otolith mass were very close to those assessed using daily increment counts (bias ranged from 1 to 24 days). Therefore, otolith mass could represent a valuable criterion for age estimation in YOY bluefin tuna that is objective, economic and easy to perform compared to daily increment counting method.

Otolith shape and temporal stability of spawning groups of Icelandic cod (Gadus morhua L.)

AbstractDuring the past 5–10 years, studies exploring small-scale structure of cod populations have provided accumulating evidence for the existence of local populations. In Iceland, all cod have been thought to belong to a single management unit. Recent studies on genetic and life history variation have, however, indicated the existence of local populations. The main objective of this study is to explore the potential existence of local populations by use of otolith shape to discriminate between spawning groups of Icelandic cod. Otoliths were sampled from mature and spawning cod at spawning locations around Iceland in 2002 and 2003, and otolith shape was described using variables correlated with size (otolith area, length, width, perimeter, and weight) as well as shape (rectangularity, circularity, and 10 Fourier coefficients). Only standardized otolith variables were used so as to remove the effect of otolith size on the shape variables. Cod were on average larger and older south of Iceland, where ambient temperature was higher than northwest, north, and east of Iceland. Otolith shape effectively discriminated between cod north and south of Iceland, and it was also possible to discriminate among cod spawning below and above 125 m at spawning locations south of Iceland. Recent genetic and tagging studies have indicated differences between cod at these two depths. Correct classification of groups of cod at the different spawning locations ranged between 0% and 44%. Incorrectly classified cod were in most cases classified to adjacent spawning locations, and a high percentage of cod south of Iceland was classified to other southern locations and cod north of Iceland to other northern locations. The temporal stability of otolith shape was studied at seven spawning locations in two consecutive years. Otolith shape differences were greater between locations than among years within a location. The spawning groups are therefore likely to have remained separate during much of their lifetime.

Fluctuating asymmetry in fish otoliths and heterozygosity in stressful estuarine environments (West Africa)

Fluctuating asymmetry (FA) is assumed to reflect the developmental instability caused by environmental or genetic stress. Fish otoliths represent a very good tool for investigating the consequence of different effects on FA. Otolith FA analysis, coupled with genetic analysis, has been undertaken on two common West African estuarine species, Ethmalosa fimbriata (EFI) and Sarotherodon melanotheron (SME), in two neighbouring estuaries, in order to highlight the impact of salinity on developmental stability. The Gambia estuary has a normal functioning and the Saloum estuary is inverse (saltier waters in the upper river), reaching extremely high salinities (&gt;100 psu) and constituting severe environmental stress. Five sub-populations of EFI and six of SME were studied along a salinity gradient. The differences between right and left otoliths were estimated with image processing by measuring five dimensions (area, perimeter, diameter, rostrum and posterior radii). Analyses of genetic differentiation at three EPIC and one anonymous nuclear gene loci for EFI and six polymorphic enzymatic loci for SME were carried out to measure the level of heterozygosity. Absolute FA in all otolith traits examined was unaffected by gender but increased significantly with fish size. Size-corrected absolute FA did not show any significant difference among sites differing largely in salinity, although a higher asymmetry in otolith area was recorded in the saltiest site. These findings suggest that otolith asymmetry is a poor indicator of osmotic stress. The individual heterozygosity level did not seem to have an effect on otolith FA for either species, even though a slight correlation appeared with otolith area or perimeter. Otolith FA cannot be considered to be a useful indicator for estimating changes linked with environmental or genetic stress in these estuaries.

Ethoxyresorufin O-deethylase (EROD) activity and fluctuating asymmetry (FA) in Zosterisessor ophiocephalus (Teleostei, Gobiidae) as indicators of environmental stress in the Venice lagoon

The impact of environmental stress on the grassgoby, Zosterisessor ophiocephalus, wasstudied at both biochemical and morphologicallevels, by comparing three sites in the Venicelagoon (the Northern, Central and Southernbasins) which differ in terms of the degree ofanthropogenic disturbance, in particular withregard to organic sediment contamination. At abiochemical level, the hepatic ethoxyresorufinO-deethylase (EROD) enzyme activity wasmeasured as a biomarker of the degree oforganic pollution (in particular regardingcontamination by PCBs and Dioxins). At amorphological level, the amount of fluctuatingasymmetry (FA) with regard to four bilateraltraits (postorbital length; otolith area;minimum otolith diameter; maximum and minimumotolith radii) was measured as a long termindicator of developmental stress. The ERODactivity was higher in grass gobies from theCentral basin, whereas the amount of FA washigher in the gobies collected from theNorthern basin. Both measurements indicatedthat the Southern basin was affected by lowlevels of environmental stress (low ERODinduction and a lower amount of FA). Theresults were discussed in the light of both theenvironmental differences among the threebasins and the different levels of biologicalorganisation at which the two indicators act.

Otolith regularities

The masses and the area sizes of the otoliths for the utriculus, sacculus and lagena of 15 species of the Black Sea fish are analyzed. Morphometrical otolith regularities are derived and their functional and ecomorphological explanations are suggested. The otolith regularities are summarized in four otolith rules: (1) the masses of the otoliths gradually increase with the fish growth. (2) The mass ratio of the sacculus and utriculus or the sacculus and lagena otoliths does not change with the fish growth. (3) The ratio between the otolith area s and the otolith mass m is described by the exponential equation s= αm 2/3. (4) The ratio between the otolith and macula sizes does not change with fish growth. Mathematical modeling of the otolith displacement responses to the acoustic and the instant force stimuli is performed. Based on the modeling the functional and ecomorphological explanations of the otolith regularities are suggested: (1) the greater the otolith mass, the higher the acoustic sensitivity at low frequencies and the sharper the frequency–response curve at its maximum. (2) The separation between maxima of the frequency–response curves for the saccular and lagenar otoliths remains virtually constant with the fish growth. (3) The bottom and littoral fish have better auditory capabilities than the pelagic fish. (4) The sensitivity to vestibular stimuli for greater otoliths is higher but the response is slower. The corresponding acceleration resolution for greater otoliths is higher and the range of accelerations in which the otolith organ can operate is narrower. (5) The relative vestibular sensitivities of the utriculus, sacculus and lagena otolith organs remain constant with fish growth. (6) The otolith organs of the bottom and littoral fish are tuned to different accelerations and possess different functional properties. The otolith organs of pelagic fish are adapted to a limited range of accelerations and are less sensitive to low accelerations as compared to the bottom and littoral fish.

Stock Discrimination Using Otolith Shape Analysis

Otolith shape has long been known to be species specific, but recent reports have pointed to its value as an indicator of stock identity. To test this hypothesis, all three pairs of otoliths were sampled from 2349 Atlantic cod (Gadus morhua) collected on spawning grounds throughout the northwest Atlantic. Otolith shape was determined with Fourier analysis and combined with measures of otolith area and perimeter. There were highly significant differences in otolith shape among most of the cod samples, but shape also differed among ages, sexes, and year-classes. The sagittal otoliths (largest pair) provided slightly better stock discrimination than did the lapillar or asteriscal otoliths. The first discriminant function was highly correlated with both fish and otolith growth rate, indicating that stock discrimination improved as the difference in stock-specific growth rate increased. Reevaluation of published studies on other species indicated that growth rate contributes more variation to regional differences in otolith shape than does stock origin. Differences in otolith shape among ages, sexes, and year-classes were also attributable to growth rate differences. To the extent that growth rates vary more between than within stocks, otolith shape analysis can provide an easily determined measure of stock identity.

Changes in otolith area: sensory area ratio with body size and depth

The ratio between the sensory areas of the sulcus acusticus and the sagittal otolith (S : O ratio) in the species Merluccius capensis and M. paradoxus was analyzed using a digital image processing system. Sagittal growth in both species is negatively allometric with respect to total fish length but is more negative in the deeper-dwelling species, Merluccius paradoxus. In contrast, the sulcus acusticus undergoes a relative increase in size, that is, the S : O ratio increases with fish length. There was a clear relationship between the increase in the S : 0 ratio and depth.

Growth determination of tropical marine fishes by means of otolith interpretation and length frequency analysis

Age and growth of four tropical marine fish species: Lethrinus choerorynchus, Lutjanus kasmira, Nemipterus furcosus and Scomberomorus brasiliensis, were determined by means of otolith interpretation and length-frequency analysis. The results showed the presence of annual rings in the studied species. These rings were consistently interpreted and their annual periodicity tested by means of various methods. The annuli are composed of distinct daily increments, of different thickness depending of the otolith area. Growth determination by means of length-frequency analysis was problematic in some species due to the lack of clear modal classes (in the length distributions). The joint analysis of the age and the length-frequency data improved the estimation of growth parameters, and allowed the selection of results that were biologically acceptable.