Contribution of water chemistry and fish condition to otolith chemistry: comparisons across salinity environments

This study quantified the per cent contribution of water chemistry to otolith chemistry using enriched stable isotopes of strontium (⁸⁶Sr) and barium (¹³⁷Ba). Euryhaline barramundi Lates calcarifer, were reared in marine (salinity 40), estuarine (salinity 20) and freshwater (salinity 0) under different temperature treatments. To calculate the contribution of water to Sr and Ba in otoliths, enriched isotopes in the tank water and otoliths were quantified and fitted to isotope mixing models. Fulton's K and RNA:DNA were also measured to explore the influence of fish condition on sources of element uptake. Water was the predominant source of otolith Sr (between 65 and 99%) and Ba (between 64 and 89%) in all treatments, but contributions varied with temperature (for Ba), or interactively with temperature and salinity (for Sr). Fish condition indices were affected independently by the experimental rearing conditions, as RNA:DNA differed significantly among salinity treatments and Fulton's K was significantly different between temperature treatments. Regression analyses did not detect relations between fish condition and per cent contribution values. General linear models indicated that contributions from water chemistry to otolith chemistry were primarily influenced by temperature and secondly by fish condition, with a relatively minor influence of salinity. These results further the understanding of factors that affect otolith element uptake, highlighting the necessity to consider the influence of environment and fish condition when interpreting otolith element data to reconstruct the environmental histories of fish.     

Mediterranean juvenile bluefin tuna: life history patterns

As there is a lack of information on the growth and migrations of bluefin tuna, information about them was gathered using the structural and chemical characteristics of their otoliths and mercury levels in body tissues as indicators of physiological and habitat characteristics. The otoliths of juvenile tuna caught in the Spanish Mediterranean littoral were studied. Otolith increments, assumed to be formed daily, were enumerated. Measurements by wavelength dispersive electron microprobe confirmed the presence of strontium in otolith tissue, and an inverse relationship between strontium/calcium (Sr/Ca) concentration ratio and temperature is suggested. Electron microprobe analyses combined with daily increment analyses of otoliths provided life history profiles for individual fish. Additional Sr/Ca concentration ratio data on fish supported the idea that Sr/Ca ratios can provide information on the environmental history of individual fish. Body concentrations of mercury were related to otolith analyses to suggest age structure, critical life history periods, growth environment, stock structure, food web position, and migration history. The techniques applied present an innovative approach to management-related problems, and the combination of chemical analyses with structural analyses promises to expand our knowledge of the life history of migratory fishes.     

Incorporation of strontium into otoliths of an estuarine fish

Patterns of Sr/Ca variability in fish otoliths have been widely applied as tracers of movement between freshwater and marine habitats, with the assumption that low salinity habitats correspond to lower otolith levels of Sr/Ca. On the other hand, fluvial estuaries can contain steep gradients in Sr/Ca, and in some estuaries, freshwater values of Sr/Ca can exceed marine values, which are relatively constant across marine habitats. Therefore, to interpret Sr/Ca variability in otoliths of fish that move through estuaries, information is needed about both the incorporation of strontium into otoliths and the nature of the gradient of Sr/Ca in the water. We conducted four experiments to evaluate the incorporation of strontium into fish otoliths under estuarine conditions, using white perch (Morone americana) as a model estuarine fish. One laboratory and the two field experiments tested the relationship between Sr/Ca in the otolith and that in the water. A fourth experiment investigated the effect of salinity, independently of the water chemistry (Sr was manipulated while maintaining a constant salinity and Ca level). All four experiments supported a direct relationship between Sr/Ca in the otolith and the water, across a range of estuarine salinities. Results also indicated that the incorporation of strontium into otoliths of estuarine fishes should be constant across broad gradients of Sr/Ca in estuarine waters. While the experiments supported past applications of tracing estuarine and diadromous movements with otolith Sr/Ca chronologies, we emphasize the need to understand the underlying nature of Sr/Ca gradients in estuaries, which may limit or confound reconstructions of estuarine habitat use.     

L'otolithe: la ≪ boîte noire ≫ des Téléostéens

Calcified structures such as otoliths have been used successfully for estimating the age of teleost fishes for many years. Otoliths record age as well as annual and/or seasonal events in the lives of fish and they are an important source of life history information. Otoliths have been found to contain characteristics that are stock specific. Elemental composition might be useful for identifying non-mixing groups of fish that may be regarded as separate stocks for fisheries management. Examination of otolith microstructure has been used to study early life history. Papers describe the use of otolith microstructure combined with chemical analysis to assess the relative significance of environmental variables: feeding frequency, photoperiod and temperature, migration from freshwater to marine water, pollution, etc. Otoliths are the first calcified structures that appear during the early development of fish and they provide a more adequate record of the growth history than scales. Otoliths represent the black box of teleost fish.     

Flexible Life History Strategies of Ninespine Sticklebacks,Genus <Emphasis Type="BoldItalic">Pungitius</Emphasis>

Synopsis We studied the life histories of the ninespine sticklebacks, Pungitius pungitius and Pungitius tymensis, collected from Japanese freshwater and brackish (sea) water habitats by examining the strontium (Sr) and calcium (Ca) concentrations in their otoliths. The Sr:Ca ratios in the otoliths changed with the salinity of the habitat regardless of identification as freshwater or brackish water type based on morphological characteristics. The ninespine sticklebacks living in a freshwater environment showed consistently low Sr:Ca ratios throughout the otolith. These samples were identified as a standard freshwater type. In contrast, all freshwater-type fishes collected from the intertidal zone showed higher otolith Sr:Ca ratios than those in the standard freshwater type, and the ratios fluctuated with the growth phase. All brackish water-type fishes collected in the intertidal zone showed the highest otolith Sr:Ca ratio throughout the otolith. In the present study, besides the two representative life history types of P. pungitius, i.e., freshwater and brackish water life history types, other sticklebacks had an anadromous life history type. These findings clearly indicate that the ninespine stickleback has a flexible migration strategy with a high degree of behavioral plasticity and an ability to utilize the full range of salinity in its life history.     

Strontium:cakium concentration ratios in otoliths of herring larvae as indicators of environmental histories

Synopsis Elemental analyses, using wave-length dispersive electron microprobe techniques on otoliths from reared Atlantic herring larvae, Clupea harengus, showed trace quantities of strontium relative to that of calcium, and an inverse relationship between Sr/Ca concentration ratios and rearing temperature. These data are consistent with those for coral aragonite, in that there appears to be an inverse temperature effect on physiological incorporation of strontium in the otolith aragonite. Our determinations of Sr/Ca concentration ratios of lab-reared herring larvae showed that the deposition of strontium relative to calcium and the rearing temperature were related, where: T (° C) = −2.955 [Sr/Ca] × 1000 ± 19.172. This principle thus makes it possible to use Sr/Ca concentration ratios in fish otoliths to delineate past temperatures experienced by an individual. Further, combining electron microprobe analyses with scanning electron microscope (SEM) examinations of daily increments in the same otolith makes it possible to reconstruct the temperature history for an individual fish on a time scale of days. An example of the application of the technique to an approximately six-month-old field-caught herring larva is given, and the limitations of the technique are discussed.     

Scanning proton microprobe analysis of strontium in an arctic charr,Salvelinus alpinus,otolith: implications for the interpretation of anadromy

Synopsis Scanning proton microprobe imaging has been used to map the distribution of strontium (Sr) in an arctic charr, Salvelinus alpinus, otolith. X-ray maps show that Sr variation follows an oscillatory zoned pattern which correlates with optically resolved zones (annuli). The internal region of the otolith shows optical zonation but no significant Sr content. Comparison of the optical image with the Sr X-ray map provides evidence of when the fish entered a high Sr environment. This may be interpreted as when the fish migrated from fresh water to a marine environment; this particular fish first migrated to sea in its eighth year. Micro-PIXE point analysis was used to determine the abundance of Sr in annuli along a radius from the nucleus to an outer edge; Sr levels in the internal region were about 10 ppm and variation in the outer zoned region was between 62 and 175 ppm. Detection limits for Sr are in the 1–2 ppm range. Other trace elements, such as transition elements, were found to be present in some growth regions in the 1–20 ppm range. Proton beam analysis is a non-destructive analytical technique capable of preserving the spatial integrity of trace-element data in otoliths such that element distribution may be linked to the growth structure of the otolith.     

Age at recruitment of Hawaiian freshwater gobies

Synopsis Very little is known about the dynamics of native Hawaiian stream fishes. Five species are restricted, as adults, to freshwater streams and estuaries on the major islands of the Hawaiian archipelago. This paucity of information is partly due to difficulties inherent in determination of age and subsequent determinations of life history characteristics. In the present study, we determined the age of newly recruited Hawaiian gobies,Stenogobius genivittatus andAwaous stamineus using otolith microtechniques. Internal otolith increments were enumerated using scanning electron microscopy (SEM). Increments of newly recruited juveniles were deposited on a daily basis as validated through a marking study. Results showed recruitment at an average age of 135 and 161 days for these two species, respectively, with more rapid growth following recruitment to freshwater. Chemical analyses of otolith carbonate of the Hawaiian gobies by electron microprobe for strontium and calcium concentrations provided valuable insights into a fish's past history. A combination of structural and chemical analyses makes it possible to link growth and recruitment to nutritional and environmental factors. Such information developed as a broad model would be applicable to the management of Hawaiian gobies and would greatly improve the quality of information available for these unique fish populations and other fish populations     

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.     

Geographic variation in otolith morphology among freshwater populations of Aphanius dispar (Teleostei,Cyprinodontiformes) from the southeastern Arabian Peninsula

Aphanius Nardo is a large genus of teleost fishes in the Old World, with 19 described species. Several of these species have only recently been recognized and additional species can be expected from isolated populations in remote areas. We show here that otolith morphology and statistical analyses of otolith variables can contribute to the detection of genetic differentiation in Aphanius. We studied samples of eight Aphanius dispar populations from the southeastern part of the Arabian Peninsula. Two populations originate from freshwater habitats far inland and probably have been isolated since the late Holocene some 4,000 years ago, three populations come from freshwater habitats with occasional connections to the sea, one population originates from a coastal site, and two were artificially introduced populations. The coastal population is interpreted as possessing the basic otolith type of A. dispar. The basic otolith type also occurs in fishes from the freshwater habitat which is located closest to the coast. Otoliths from the two other freshwater populations with occasional connections to the coast differ slightly from the basic type. However, the two populations from the long‐term isolated freshwater habitats far inland show distinct morphological changes. Our results are consistent with the hypotheses that i) otolith morphology is primarily genetically determined and is little influenced by physical parameters of the habitat, and ii) isolated A. dispar populations may be capable of evolving into new species within short periods of time. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.     

Age and growth of the brick soldierfish,Myripristis amaena

Otoliths (sagittae) of the coral reef fish, Myripristis amaena, the brick solderfish were examined internally by Scanning Electron Microscope methods to observe microincrements. The daily nature of increment deposition was validated through tetracycline and acetazolamide marking experiments. Utilization of multivariant mathematical models relating age to otolith size and fish size demonstrated that age could be reliably determined from body measurements and otolith weight measurements. Consequently, M. amaena grows slowly, maturing at about 6 years of age, lives at least 14 years and reaches at least 215 mm SL.     

Ontogenetic pattern,morphological sexual and side dimorphism in the saccular otolith of a scaleless killifish Aphanius furcatus (Teleostei: Aphaniidae)

The relationships between ontogenetic and sexual factors with otolith morphology are poorly studied in killifish genus Aphanius Nardo, 1827. We studied the ontogeny, morphological sexual and side dimorphism in the saccular otolith of a scaleless killifish Aphanius furcatus inhabiting a high‐salinity environment in Southern Iran. The results highlighted growth‐dependent variability in the otolith of A. furcatus, which might be useful to define its certain developmental stages. We underlined those characters that were consistent during fish development, which could be valuable to identify the species. Considering the ontogenetic pattern of otolith in A. furcatus, it was found that the otoliths of young individuals have largely been influenced by sexes and that it was reduced during the fish development. Morphological side dimorphism in the otolith of A. furcatus tends to be increased during fish development. The outcomes of this study are essential since they have provided new information on ontogenetic changes and the sexual and side dimorphism of Aphanius otolith.     

Migratory history of the threespine stickleback Gasterosteus aculeatus

 The migratory history of two highly divergent forms (the Japan Sea and Pacific Ocean forms) of the threespine stickleback Gasterosteus aculeatus collected from Japanese brackish water (seawater) and freshwater was studied by examining strontium (Sr) and calcium (Ca) concentrations in their otoliths using wavelength dispersive X-ray spectrometry on an electron microprobe. The Sr : Ca ratios in the otoliths changed with salinity of the habitat. The otolith Sr : Ca ratios of the freshwater resident-type samples of the Pacific Ocean form showed consistently low Sr : Ca ratios, averaging 0.85–0.96 × 10⁻³ from the core to the edge. In contrast, the otolith Sr : Ca ratios of the anadromous type of both the Japan Sea and Pacific Ocean forms fluctuated strongly along the life history transects in accordance with their migration patterns from seawater to freshwater. The higher ratios in the anadromous type, averaging 5.4 × 10⁻³, in the otolith region from the core to 200 μm, corresponded to the seagoing period, suggesting that otolith Sr : Ca ratios are affected by ambient water salinity. These findings clearly indicate that otolith Sr : Ca ratios reflect individual life histories, and that these two highly divergent forms of stickleback have a flexible migration strategy. Received: May 23, 2002 / Revised: July 29, 2002 / Accepted: August 19, 2002 Acknowledgments We are grateful to Dr. S. Mori of Gifu Keizai University, and Miss M. Yamada and Messrs. M. Kume and T. Kitamura of Hokkaido University, for their assistance in sampling. This work was supported in part by Grant-in-Aid No. 13760138 from the Ministry of Education, Culture, Sports, Science and Technology, Japan. Correspondence to:Takaomi Arai     

What otolith microchemistry and stable isotope analysis reveal and conceal about anguillid eel movements across salinity boundaries

Otolith microchemistry studies indicate that growth-phase (yellow stage) anguillid eels commonly shift at irregular intervals between fresh and saline waters, but this technique has not detected regular seasonal migrations across salinity zones. We tested the ability of otolith microchemistry and stable isotope analysis to detect migrations of American eels (Anguilla rostrata) between salinity boundaries in two small stream–estuary systems in Canada’s Bay of Fundy. Although the two methods showed concordant classifications of recent residence history, most eels caught in fresh water in spring (68.8–89.7 %) and fall (78.8–83.3 %) showed microchemical and isotopic signatures that reflected occupancy of saline waters. These eels were classified as migrants which had summered in saline waters and then migrated to freshwater wintering grounds where they retained their saline signatures. In summer, most eels (85.0–100.0 %) captured in fresh and saline water had recent microchemical and isotopic signatures matching the habitat of capture. Our results suggest that lifetime otolith microchemistry profiles are unable to detect eel wintering migrations, a failure that is likely due to winter depression of otolith accretion. Elucidation of seasonal eel movements requires cross-seasonal and cross-site sampling for the microchemistry and stable isotope methods, or tagging studies. Seasonal saline–fresh eel migrations may be more common than previously appreciated, underlining the need for conservation of both habitats, and connectivity between the two.     

The record of daily growth in otoliths of Atlantic silversides, Menidia menidia, from field and laboratory

Otoliths were removed from field-collected silversides of age less than 3 months. Otolith diameter was highly correlated with total length of the fish. Daily growth ring counts for this species are known to be a function of age rather than size, so widths for the daily growth rings provide a record of daily increases in length of the fish. Measurement of ring widths showed that weekly specific growth rate was greater than 70% at age 1 week, but declined to about 30% at age 1 month and about 15% at age 2 months. A laboratory experiment in which temperature was changed on a weekly basis demonstrated that environmental variables can affect the width of rings. Nevertheless, the growth rate of field-collected fish, as calculated from otolith ring widths, was more highly correlated with size of fish, as measured by otolith radius, than with the environmental variables of temperature, salinity and plankton abundance. Back-calculation of growth rates from otolith ring widths of five fish collected at the end of the growing season yielded the same age-growth curves as were obtained from 203 fish collected biweekly during the season.     

Application of otolith microchemistry to estimate the migratory history of Japanese eel Anguilla japonica on the Sanriku Coast of Japan

The age and migratory history of the Japanese eel, Anguilla japonica Temminck & Schlegel, collected in Miyako Bay along the Sanriku coast of Japan, was examined using the otolith microstructure and analysis of strontium (Sr) and calcium (Ca) concentrations conducted with wavelength dispersive X‐ray spectrometry by an electron microprobe. The line analysis of Sr : Ca ratios along the life history transect of each otolith showed a peak (ca. 15–17 × 10^?3) which corresponded with the period of their leptocephalus and early glass eel stages in the ocean. The mean Sr : Ca ratios from the elver mark to the otolith edge indicated that there were eels with several general categories of migratory history, including sea eels that never entered freshwater (average Sr : Ca ratios, ≥6.0 × 10^?3), and others that entered freshwater for brief periods but returned to the estuary or bay. This evidence of the occurrence of sea eels in this northern area indicates that Japanese eels of the Sanriku coast do not necessarily migrate into freshwater rivers during recruitment as do glass eels at the beginning of their growth phase; even those that do enter freshwater may later return to the marine environment. Thus, anguillid eel migrations into freshwater are clearly not an obligatory migratory pathway, but rather a facultative catadromy with seawater or estuarine residents as an ecophenotype.     

Development of a species-specific fractionation equation for Arctic charr (<Emphasis Type="Italic">Salvelinus alpinus</Emphasis> (L.)): an experimental approach

A species-specific fractionation equation for Arctic charr (Salvelinus alpinus (L.)) was developed experimentally for use in ecological studies of temperature-driven phenologies for the species. Juvenile Arctic charr were reared in controlled conditions at different temperatures (2–14°C), with three replicates of each temperature. Otoliths from the fish and water samples from the chambers were analysed for oxygen isotope composition and used to estimate temperature-dependent fractionation equations relating the isotopic ratio to rearing temperature. A linear and a second order polynomial relationship were estimated and validated using comparable Arctic charr data from another study. Temperatures predicted using the polynomial equation were not significantly different from recorded experimental temperatures, whereas with the linear equation there were significant differences between the predicted and measured temperatures. The polynomial equation also showed the least bias as measured by mean predictive error. Statistical comparisons of the polynomial fractionation equation to a similarly estimated equation for brook charr (Salvelinus fontinalis (Mitchill)) indicated significant differences. Results imply the need for species-specific fractionation equations, even for closely related fish. Results further suggest the polynomial form of the fractionation equation will facilitate more accurate characterisation of water temperatures suitable for use in ecological studies of temperature-driven phenologies of Arctic charr.     

Identifying sea-run brown trout, Salmo trutta, using Sr : Ca ratios of otolith

 The migratory history of the brown trout, Salmo trutta, collected from Japanese rivers, was examined in terms of strontium (Sr) and calcium (Ca) uptake in the otolith, by means of wavelength dispersive X-ray spectrometry on an electron microprobe. Sea-run (anadromous) and freshwater-resident (nonanadromous) types of S. trutta were found to occur sympatrically. Otolith Sr concentration or Sr : Ca ratios of anadromous S. trutta fluctuated strongly along the life history transect in accordance with the migration (habitat) pattern from sea to freshwater. In contrast, the Sr concentration or the Sr : Ca ratios of nonanadromous fish remained at consistently low levels throughout the otolith. The higher ratios in anadromous S. trutta, in the otolith region from the core to 1500 μm, corresponded to the initial seagoing period, probably reflecting the ambient salinity or the seawater–freshwater gradient in Sr concentration. The findings clearly indicated that otolith Sr : Ca ratios reflected individual life histories, enabling the sea-run S. trutta to be distinguished from the freshwater-resident brown trout. Received: March 18, 2002 / Revised: May 15, 2002 / Accepted: June 5, 2002 Acknowledgments The authors are grateful to Messrs. T. Ikeda, S. Kudo, Y. Miyakoshi, M. Nagata, K. Shimoda, T. Takami, K. Takeuchi, and M. Ueda for their assistance in collecting specimens. This work was supported in part by Grant-in-Aid No. 13760138 from the Ministry of Education, Culture, Sports, Science and Technology, Japan. Correspondence to:Takaomi Arai     

Migratory history of the spotted flagtail, <Emphasis Type="Italic">Kuhlia marginata</Emphasis>

The spotted flagtail, Kuhlia marginata (Kuhliidae) is commonly found in streams throughout the Ryukyu Archipelago, including Okinawa Island, Japan. Although it has been suggested that they spawn at sea, little is known about their migratory history. The aim of the study was to clarify their migration history based on otolith microchemistry analysis, size composition and the gonadal development of fish collected from the Genka River on Okinawa Island. All strontium/calcium (Sr/Ca) ratios of otoliths remained high around their cores, and then dropped rapidly at some distance from the core. The estimated standard length corresponding to decreases correlated with the size of new recruits collected in the river (ca. 20 mm in standard length: SL). Multiple increases, the sign for migrations into the high saline area, were recognized intermittently around the otolith margin. As the estimated SL at these increases in the Sr/Ca ratio occurred above the minimum maturation size (female: 95.5 mm SL; male: 83.5 mm SL), such multiple increases in Sr/Ca ratios were caused by spawning migration. Therefore, we conclude that the catadromous pattern of K. marginata is as follows; this species grows in the sea during the early life stage until ca. 20 mm SL and then grow in the freshwater area until maturation, before they migrate again to the sea for spawning. Sr/Ca ratio profiles also suggested that a large number of males did not return to the freshwater area after spawning, whereas females might spawn several times during their lives around the inshore area.     

The use of otolith shape analysis for ageing juvenile red snapper, Lutjanus campechanus

Morphological changes in otolith shape with age, of young (<age 3) red snapper were examined through shape analysis and tested as an objective method for age determination. Otoliths from two collections of juvenile fish (hatchery and wild) were used in the study. First, shape analysis was applied to a series of known-age otoliths from hatchery-reared age 0, 1 and 2 fish. Multidimensional scaling and non-parametric analysis of similarities showed significant shape differences among the three age classes of fish. Discriminant function analysis and cross-validation classification showed 65.6% correct age classification based on shape variables alone, and 86.7% correct age classification with inclusion of otolith weight in the discriminant function (n?=?90). Subsequently, the method was applied to otoliths from a series of age 0, 1 and 2 wild caught red snapper. Otoliths from wild fish showed a similar age classification success rate of 68.9% based on shape variables alone and 86.7% correct age classification with the inclusion of otolith weight in the discriminant function (n?=?90). Ageing of juvenile red snapper through otolith increment counts has been difficult in past studies and this study provides an alternative, objective method of otolith shape analysis for ageing young fish of this species.