An eco-morphological explanation of individual variability in the shape of the fish otolith: comparison of the otolith of Hoplostethus atlanticus with other species by depth

Variation in otolith shape (otolith polymorphism) in Hoplostethus atlanticus, Hoplostethus mediterraneus, Paratrichichthys trailli, Pagrus major and Trachurus murphyi , quantified using principal components analysis based on Fourier transform decompositions of outlines of otolith shape, was particularly high among the samples of H. atlanticus from the North Atlantic, New Zealand, Australia and Namibia. The scatter was uniform, however, and did not show any significant differences among regions. The implication drawn from the high variability in otolith shape of H. atlanticus was that otolith shape polymorphism was maintained by some form of balancing selection across many small local environments which may result in k-selection with consequent poor response by H. atlanticus to maximum sustainable yield harvesting strategies. The variation in otolith shape defined by the otolith morphospace of the five species that were measured, showed a decreasing trend in scatter (i.e. decreasing complexity of shape) proceeding from the species with the deepest habitat ( H. atlanticus ) to the most shallow ( T. murphyi ).     

Distinction between Mulloidichthys flavolineatus juveniles from Reunion Island and Mauritius Island (south-west Indian Ocean) based on otolith morphometrics

Sagittal otoliths of the yellowstripe goatfish Mulloidichthys flavolineatus were analysed in order to compare Reunion Island fish stocks with those of Mauritius (south-west Indian Ocean). Conventional otolith morphometric variables (area, perimeter, length and width), shape indices (form factor, roundness, circularity, rectangularity, ellipticity and eccentricity) and Fourier shape analysis were compared between three sites; two in Reunion Island and one in Mauritius. Regional and site-specific differences were found for all the conventional otolith morphometric features. Regarding the shape indices, the differences between sites were best described by form factor, roundness, circularity and rectangularity. A classification by canonical discriminant analysis indicated significant differences between the three sampling sites. The combined use of morphometric variables (size and shape) and external outlines (shape analysis through Fourier series) showed the importance of otolith shape for intraspecific discrimination.     

Latitudinal Gradient in Otolith Shape among Local Populations of Atlantic Herring (Clupea harengus L.) in Norway

Otolith shape analysis of Atlantic herring (Clupea harengus) in Norwegian waters shows significant differentiation among fjords and a latitudinal gradient along the coast where neighbouring populations are more similar to each other than to those sampled at larger distances. The otolith shape was obtained using quantitative shape analysis, the outlines were transformed with Wavelet and analysed with multivariate methods. The observed morphological differences are likely to reflect environmental differences but indicate low dispersal among the local herring populations. Otolith shape variation suggests also limited exchange between the local populations and their oceanic counterparts, which could be due to differences in spawning behaviour. Herring from the most northerly location (69°N) in Balsfjord, which is genetically more similar to Pacific herring (Clupea pallasii), differed in otolith shape from all the other populations. Our results suggest that the semi-enclosed systems, where the local populations live and breed, are efficient barriers for dispersal. Otolith shape can thus serve as a marker to identify the origin of herring along the coast of Norway.     

Ontogenetic and environmental effects on otolith shape variability in three Mediterranean European eel (Anguilla anguilla, L.) local stocks

Otolith morphology is an efficient tool for the discrimination of fish stocks, populations and species when comparative genetic data are not available. Currently, the relationship between environmental factors and otolith shape is poorly characterized for the European eel (Anguilla anguilla), a highly migratory catadromous species constituting a single, randomly mating stock. The present study analyses the differences in otolith morphology between three Mediterranean eel local stocks from different environmental contexts (i.e. two brackish lagoons and one river). The relationship between otolith shape and otolith size was studied by means of Elliptic Fourier analysis and multivariate statistics. Otolith profile was digitally acquired and Cartesian coordinates were extracted. Partial Least Square (PLS) analysis pointed to continuous allometric growth in size and shape in otoliths from all three sites. In the three environments, shape variations occurred during growth as indicated by the presence of a significant and positive relationship between otolith size and the first PLS latent vector (i.e. which bears most of the information regarding otolith outline). Differences between smaller and larger sized otoliths were investigated using PLS Discriminant Analysis (PLSDA) and cluster analysis. Results indicate that otolith shape is highly uniform at smaller than at larger sizes. These shape differences apparently overlap the initial differentiation of the small otolith outlines acquired by eels during the growing phase as elvers in the marine environment. Data were discussed considering that the physical and chemical habitat variability in brackish lagoons and river could underlie a marked change in otolith shape during the animals' growth.     

Discrimination of red porgy Pagrus pagrus (Sparidae) potential stocks in the south-western Atlantic by otolith shape analysis

Otolith shape analysis is a powerful method for fish stock identification. We compared the otolith shape of Pagrus pagrus (Linnaeus 1758) along with its distribution in four south-western Atlantic regions where it is commercially fished: Rio de Janeiro, Rio Grande do Sul in southern Brazil, the Argentine-Uruguayan Common Fishing Zone (UA) and the Argentinian Exclusive Fishing Zone (AR). Otolith shapes were compared by Elliptical Fourier and Wavelet coefficients among specimens in a size range with similar otoliths, morphometric parameters and ages. Four potential stocks were identified: one in the AR, a second along the UA which included specimens from southern Brazil with well-marked opaque bands in its otoliths (MRS), the third in southern Brazil with faint or absent opaque bands in its otoliths (FRS) and the fourth along Rio de Janeiro. The difference in the otolith shape among regions followed differences reported using other stock identification techniques. The similarity between otoliths from UA and MRS (ANOVA-like, P > 0.01) can be explained by seasonal short-range migrations. Otoliths shape differences between MRS and FRS (ANOVA-like, P < 0.01) suggest that P. pagrus does not form a homogeneous group in southern Brazil.     

Otolith shape: a population marker for Atlantic herring Clupea harengus

Otolith shape variation of seven Atlantic herring Clupea harengus populations from Canada, the Faroe Islands, Iceland, Ireland, Norway and Scotland, U.K., covering a large area of the species' distribution, was studied in order to see if otolith shape can be used to discriminate between populations. The otolith shape was obtained using quantitative shape analysis, transformed with Wavelet and analysed with multivariate methods. Significant differences were detected among the seven populations, which could be traced to three morphological structures in the otoliths. The differentiation in otolith shape between populations was not only correlated with their spawning time, indicating a strong environmental effect, but could also be due to differing life‐history strategies. A model based on the shape differences discriminates with 94% accuracy between Icelandic summer spawners and Norwegian spring spawners, which are known to mix at feeding grounds. This study shows that otolith shape could become an accurate marker for C. harengus population discrimination.     

基于形态分析探究鳞头犬牙南极鱼各生活史阶段耳石形态变化

鳞头犬牙南极鱼为南极最为重要的底层鱼类,具有重要的生态作用和极高的商业价值,鱼类耳石形态随着生活史过程的推移有所差异,故耳石形态分析可用于推断鳞头犬牙南极鱼的生活史过程。本研究利用采集自南极罗斯海、阿蒙森海、威德尔海及拉扎列夫海120尾分属4个生活史阶段的鳞头犬牙南极鱼耳石,结合传统测量分析与椭圆傅里叶分析两种形态学研究方法,对其各生活史阶段耳石形态的差异进行研究。结果表明: 鳞头犬牙南极鱼各生活史阶段耳石形态存在差异,随着生活史过程的推移,耳石的变化趋势为整体上由轮廓平滑、复杂度较低向轮廓曲折、高度复杂化转变,且纵轴方向增长速度低于横轴方向,翼叶等特征部位变化显著。针对耳石形态的线性判别分析结果显示,椭圆傅里叶分析判别值较高,为85.4%,而传统测量分析判别值较低,为71.9%,表明相较于传统测量分析,椭圆傅里叶分析更具优势。     

Improvements To The Method Of Fourier Shape Analysis As Applied In Morphometric Studies

Fourier outline shape analysis is a powerful tool for the morphometric study of two-dimensional form in organisms lacking many biologically homologous landmarks. Several improvements to the method are described herein; these modifications are incorporated into the new computer programs H angle , H match and H curve . First, automated tracing of outlines using image capture software, although desirable, results in high frequency pixel 'noise' that can corrupt the Fourier analysis. Program H angle eliminates this noise using optional and variable levels of outline smoothing. Secondly, a widely used Fourier technique, elliptic Fourier analysis (EFA, Kuhl and Giardina 1982), yields coefficients that are not computationally independent of each other, a condition that hampers and compromises statistical analysis. In addition, EFA increasingly downweights successively more detailed features of the outline. Program H angle solves both of these problems. Lastly, Fourier methods in general are sensitive to the placement of the starting position of the digitized trace. This problem is acute when the organisms under study have no unambiguously defined, homologous point on the outline from which to start the trace. Program H angle allows the user to normalize for starting position using various properties of individual outlines. Alternatively, H match takes a new approach and can be used to normalize using properties of the entire population under study. key words : Fourier shape analysis, morphometric studies, new computer programs, foraminiferal outlines.     

Sex-linked variations in the sagittal otolith biometry of Nemipterus randalli (Russell, 1986) from the eastern Mediterranean Sea

Few studies have been conducted on the sagittal otolith shape and morphometry of Nemipterus randalli, and none of these studies has examined the effect of sexual dimorphism on the otolith morphology of this species, therefore this study aimed to contribute to knowledge about the otolith morphology of N. randalli, an invasive fish species for the Mediterranean Sea. For this purpose, a total of 132 samples (51 female and 81 male) were obtained from İskenderun Bay with the help of commercial fishermen in November 2018. Relationships between otolith measurements and fish size were determined. Shape indices and elliptic Fourier coefficients were calculated. Significant differences were detected between males and females in all analysis. The sexes were separated from each other using both shape indices and elliptic Fourier coefficients. However, shape analysis was more effective in distinguishing sexes than traditional morphometric analysis. Asymmetry in the otolith morphology of sexes has been attributed to differences in the growth and sexual maturity of male and female fish. The results of this study indicated that sexual dimorphism in Nemipterus randalli was also reflected in the otolith morphology.     

Otolith shape analysis for three <Emphasis Type="Italic">Sicyopterus</Emphasis> (Teleostei: Gobioidei: Sicydiinae) species from New Caledonia and Vanuatu

Otolith shape analysis has been used in a number of studies as an inexpensive and powerful method for categorising fish in individual stocks. Elliptical Fourier analysis was used on three different amphidromous Sicyopterus species. Sicyopterus lagocephalus is a widespread species while the other two have a limited distribution area, Sicyopterus aiensis being endemic to Vanuatu, and Sicyopterus sarasini to New Caledonia. Both endemics live in sympatry with the widespread species. The otolith shape of all fish sampled was a clear species differentiator, thereby demonstrating that otolith shape is species-specific. At an intraspecific level there are different river populations within samples from Vanuatu, indicating a western group and an eastern “central” group.These results are congruent both for the endemic species, S. aiensis and for the cosmopolitan species. Finally, we found that, for S. lagocephalus, the cosmopolitan species, New Caledonian samples are close to western Vanuatu samples, the latter two being well differentiated from the eastern “central” Vanuatu samples. The explanation for these results may lay either in the influence of environmental factors on the otolith shape, or in the influence of common early life history thus reflecting genetic factors, or a combination of both.     

The use of otolith shape to determine stock structure of <Emphasis Type="Italic">Engraulis encrasicolus</Emphasis> along the Tunisian coast

Engraulis encrasicolus is of great economic importance in the Mediterranean. However, little is known about its stock structure. Otolith shape analysis has been successfully used for fish stock identification. In this study, the stock structure of anchovy caught off the open sea and the coastal area of the Gulf of Tunis, lagoon of Bizerte and Lake of Ichkeul were investigated using otolith shape. Otolith shape was determined by Fourier analysis and then compared among specimens sampled from different areas with forward stepwise canonical discriminant analysis. Significant differences in otolith shape between the open sea and inshore anchovy groups were detected. Otolith shape of anchovy collected in the Lake of Ichkeul was distinct from the other groups. This finding suggests a clear discreteness of the open sea and the continental groups. The data highlighted the potential for using otolith shape analysis for anchovy stock identification, as well as the role of oceanographic features in determining stock separation. These findings will have major implications for anchovy fisheries management in Tunisia. By using a precautionary approach and considering the three areas as separate stocks, fisheries management strategies should be adjusted to achieve optimum sustainable production from each stock and to avoid decreases in genetic variety.     

Sagittal otolith morphogenesis asymmetry in marine fishes

This study investigated and compared asymmetry in sagittal otolith shape and length between left and right inner ears in four roundfish and four flatfish species of commercial interest. For each species, the effects of ontogenetic changes (individual age and total body length), sexual dimorphism (individual sex) and the otolith's location on the right or left side of the head, on the shape and length of paired otoliths (between 143 and 702 pairs according to species) were evaluated. Ontogenetic changes in otolith shape and length were observed for all species. Sexual dimorphism, either in otolith shape and length or in their ontogenetic changes, was detected for half of the species, be they round or flat. Significant directional asymmetry in otolith shape and length was detected in one roundfish species each, but its inconsistency across species and its small average amplitude (6·17% for shape and 1·99% for length) suggested that it has barely any biological relevance. Significant directional asymmetry in otolith shape and length was found for all flatfish species except otolith length for one species. Its average amplitude varied between 2·06 and 17·50% for shape and between 0·00 and 11·83% for length and increased significantly throughout ontogeny for two species, one dextral and one sinistral. The longer (length) and rounder otolith (shape) appeared to be always on the blind side whatever the species. These results suggest differential biomineralization between the blind and ocular inner ears in flatfish species that could result from perturbations of the proximal‐distal gradient of otolith precursors in the endolymph and the otolith position relative to the geometry of the saccular epithelium due to body morphology asymmetry and lateralized behaviour. The fact that asymmetry never exceeded 18% even at the individual level suggests an evolutionary canalization of otolith shape symmetry to avoid negative effects on fish hearing and balance. Technically, asymmetry should be accounted for in future studies based on otolith shape.     

Stock discrimination of the Japanese Spanish mackerel (Scomberomorus niphonius) based on the otolith shape analysis in the Yellow Sea and Bohai Sea

Geographic variability in sagittal otoliths shape of the Japanese Spanish mackerel (Scomberomorus niphonius) was studied to test whether stock discrimination might be possible using this method. It was hypothesized that S. niphonius would show little difference in otolith shape, likely as a result of mixing between seas. A total of 164 sagittal otoliths were collected from commercial fishing vessels between April and June 2010 in three spawning grounds, that of Qingdao in the middle Yellow Sea, Lvsi in the southern Yellow Sea, Huanghua in the the Bohai Sea. To minimize the potential effect of the fish size among the three fishing grounds, a narrow size range (400–550 mm fork length) was selected. The discriminant function analysis conducted with standardized otolith shape indices and Fourier harmonics produced classification success rates ranging from 57.0 to 88.2% and worked well in the separation between the Bohai Sea group and the Yellow Sea group. No significant differences were detected among the three groups for the Lvsi spawning grounds. The otolith variables showed a distinct gradual variation tendency with the movement of fish schools from south to north. The results suggest that stock from the Bohai Sea could be managed as a separate entity from those found in the Yellow Sea.