Regional variation in otolith Sr:Ca ratios of African longfinned eel Anguilla mossambica and mottled eel Anguilla marmorata: a challenge to the classic tool for reconstructing migratory histories of fishes

Otolith Sr:Ca ratios of the African longfinned eel Anguilla mossambica and giant mottled eel Anguilla marmorata from nine freshwater sites in four rivers of South Africa were analysed to reconstruct their migratory life histories between freshwater and saltwater habitats. For A. mossambica, the Sr:Ca ratios in the otolith edge differed significantly among rivers and had large effect sizes, but did not differ among sites within a river. Otolith Sr:Ca ratios did not differ among rivers for A. marmorata. When rivers were pooled, the edge Sr:Ca ratios of A. mossambica were not significantly different from those of A. marmorata. According to the river-specific critical Sr:Ca ratio distinguishing freshwater from saltwater residence, most A. mossambica and A. marmorata had saltwater habitat experience after settlement in fresh water. This was primarily during their elver stage or early in the yellow eel stage. During the middle and late yellow eel stage, freshwater residency was preferred and only sporadic visits were made to saltwater habitats. The data also suggest that regional variations in otolith Sr:Ca ratios affect the critical Sr:Ca value and are a challenge for the reconstruction of migratory life histories that should be explicitly considered to avoid bias and uncertainty.     

Spatial and temporal variations in otolith chemistry and relationships with water chemistry: a useful tool to distinguish Atlantic salmon Salmo salar parr from different natal streams

Otolith elemental (Sr:Ca, Ba:Ca, Mn:Ca, Mg:Ca and Rb:Ca) and isotopic (⁸⁷Sr:⁸⁶Sr) profiles from several annual cohorts of juvenile Atlantic salmon Salmo salar were related to the physico‐chemical characteristics (chemical signatures, flow rate, temperature and conductivity) of their natal rivers over an annual hydrological cycle. Only Sr:Ca, Ba:Ca and ⁸⁷Sr:⁸⁶Sr in otoliths were determined by their respective ratios in the ambient water. Sr:Ca ratios in stream waters fluctuated strongly on a seasonal basis, but these fluctuations, mainly driven by water flow regimes, were not recorded in the otoliths. Otolith Sr:Ca ratios remained constant during freshwater residency at a given site and were exclusively related to water Sr:Ca ratios during low flow periods. While interannual differences in otolith elemental composition among rivers were observed, this variability was minor compared to geographic variability and did not limit classification of juveniles to their natal stream. Success in discriminating fish from different sites was greatest using Sr isotopes as it remained relatively constant across years at a given location.     

Migratory patterns and contribution of stocking to the population of European eel in Lithuanian waters as indicated by otolith Sr:Ca ratios

Otolith Sr:Ca ratios were examined to evaluate the contribution of the stocked eel Anguilla anguilla elvers, which have been stocked in Lithuanian waters and mixed with naturally recruited eels for several decades, to the native eel population. Stocked eels were identified by the freshwater signature (Sr:Ca ratios <2·24 × 10⁻³) on the otolith after the glass eel stage. Naturally recruited eels, that had migrated through the North and Baltic Seas, were characterized by an extended seawater and brackish-water signature (Sr:Ca ratios >3·23 × 10⁻³) after the glass eel stage. Of 108 eels analysed, 21 eels had otolith Sr:Ca ratio profiles consistent with stocking while 87 showed patterns of natural recruitment. The ages of naturally recruited eels arriving in Lithuanian fresh waters varied from 1 to 10 years, with a mean ±  s.d . age of 5·2 ± 2·1 years. Eels from the inland Lake Baluošai were all freshwater residents of stocked origin. Stocked eels, however, accounted for only 20% of the eels from the Curonian Lagoon and 2% of eels sampled in Baltic coastal waters. This finding does not support the hypothesis that the eel fishery in the Curonian Lagoon depends mostly on stocking.     

Salinity-linked growth in anguillid eels and the paradox of temperate-zone catadromy

Temperate-zone anguillid eels use both saline (marine or brackish) and fresh waters during their continental phase, but use of fresh waters is paradoxical because on average these fishes grow more rapidly in saline than in fresh waters. Based on data from anguillid eels whose habitat-residency histories had been determined by Sr:Ca otolithometry, superiority of growth rates in saline water is much greater in American eels Anguilla rostrata in north-eastern North America (mean saline:fresh growth rate ratio 2·07) than in European Anguilla anguilla , Japanese Anguilla japonica and shortfinned Anguilla australis eels (range of mean ratios 1·12–1·14). Data from A. rostrata in the Hudson Estuary, U.S.A., and Prince Edward Island, Canada, were used to test adaptive explanations of catadromous migrations. The hypothesis that lower mortality in fresh water offsets faster growth in saline water was not supported because loss (mortality + emigration ) rates did not vary between saline and fresh zones of the Hudson Estuary. Hypotheses that anguillid eels move to fresh water to escape from larger anguillid eels in saline water or to evaluate habitat quality were not supported by size and age distributions. Catadromy in temperate-zone anguillid eels increases the diversity of occupied habitats and therefore lowers fitness variance caused by environmental fluctuations. Catadromy in temperate-zone anguillid eels could be due to natural selection for maximum geometric mean fitness which is sensitive to fitness variance. Temperate-zone catadromy might also be maladaptive, at least in local areas, due to shifts over time in selective pressures or to inability of panmictic genetic systems to adapt to local conditions.     

Otolith Sr:Ca and Ba:Ca may give inconsistent indications of estuarine habitat use for American eels (<Emphasis Type="Italic">Anguilla rostrata</Emphasis>)

Temporal patterns in otolith Sr:Ca and Ba:Ca ratio values of American eels Anguilla rostrata from two sites in western Newfoundland gave insight into the use of freshwater and saline habitats. Mean Sr:Ca and Ba:Ca values at the core zone did not differ between sites, indicative of a common oceanic origin. At the otolith edge, representing continental life, both Sr:Ca and Ba:Ca values varied between sites consistent with ambient element:Ca ratio values and salinity, with typically higher Sr:Ca and lower Ba:Ca values in saline than in fresh waters. Most eels (73%) from Muddy Hole, an estuarine site, were evaluated as estuarine residents while most (70%) eels from Castors River, a freshwater site, were evaluated as freshwater residents, with the remaining eels from each site evaluated as inter-habitat migrants. An otolith element:Ca critical value appropriate for distinguishing between fresh and saline water residence is fundamental for estimating the proportion of eel residence in freshwater and their subsequent classification into habitat residence groups. Such classification is moderately robust to the critical value selected. For inter-habitat migrants, moderate otolith Sr:Ca values between the elver check and otolith edge suggestive of estuarine residence may coincide with Ba:Ca values suggestive of freshwater residence. No general critical value for separating fresh and estuarine habitats was found for otolith Ba:Ca. Otolith Ba:Ca temporal patterns may assist the use of Sr:Ca in the evaluation of historical habitat residence and inter-habitat movement but the use of otolith Ba:Ca values should be applied cautiously for American eels and perhaps of other estuarine/freshwater migratory fishes.     

Migratory patterns of anadromous white-spotted charr Salvelinus leucomaenis in Eastern Hokkaido, Japan: The solution to a mystery?

We evaluated the migratory patterns of white-spotted charr in eastern Hokkaido, Japan using: (1) field observation of fish, (2) salinity-recording archival tags, and (3) analysis of otolith Sr and Ca concentrations. Field observations suggest that the majority of anadromous white-spotted charr outmigrate to the sea in spring (April to June) and ascend the river between late summer and autumn (August to November). Salinity records from archival tags revealed a fluctuating pattern consisting of both low and high values within a month, indicating frequent short-term movements between freshwater (including non-natal rivers) and brackish/salt water habitats. Otolith Sr:Ca ratios exhibited a sharp rise at about 40–50% of the otolith radius, which was followed by several spikes consisting of both low and high values, indicating that fish migrated between freshwater and marine environments multiple times during their life history. However, the peaks in Sr:Ca ratios were generally associated with annulus zones on the otolith, indicating that the anadromous migration was generally an annual event.     

Factors influencing otolith strontium/calcium ratios in Anguilla japonica elvers

The effects of temperature and salinity on the concentration ratios of strontium (Sr) to calcium (Ca) within the sagittal otoliths of elvers of the Japanese eel, Anguilla japonica, were studied by spot analysis using a wavelength dispersive X-ray electron microprobe. A total of 340 elvers were used: 100 elvers were reared for 15 days under various salinity conditions (freshwater, one-third seawater, two-thirds seawater and pure seawater at 22 °C; 240 elvers were reared for 58 days under various water temperature conditions (12, 17, 22, 27 °C) in either freshwater or pure seawater. Otolith Sr/Ca ratios were found to be positively correlated with water salinity. On the other hand, the Sr/Ca ratios were not found to be significantly different among the various temperature groups. The above results strongly suggest that the physiological mechanism of incorporation of Sr and Ca within the otolith of an eurythermal fish, Japanese eel, does not change within this range of temperatures (12–27 °C).     

Salinities, not diets, affect strontium/calcium ratios in otoliths of Anguilla japonica

Although otolith Strontium (Sr)/calcium (Ca) ratios have been widely used to reconstruct the past salinity environmental history of anguillid eels, factors affecting the Sr/Ca ratios in otoliths are incompletely understood. Japanese Eel (Anguilla japonica) elvers (mean length 54.7 ± 2.1 mm) were collected in the estuary during their upstream migration and reared at 5 different salinities (0, 5, 15, 25, and 35 psu) and 3 types of feeding conditions (formulated feed, tubifex, and starvation) for 30 days to evaluate the effects of salinity and diets on otolith Sr/Ca ratios. Ca and Sr concentrations in the ambient water significantly increased with salinity (SAL) as [Ca] _water = 15.50SAL − 5.56, and [Sr] _water = 0.21SAL + 0.03, respectively. Sr/Ca ratios in otoliths increased with salinity (SAL) of the rearing water as [(Sr/Ca) × 1000] _otolith = 0.091SAL + 3.790. In diets, Sr/Ca ratios were 4 times higher in tubifex than in formulated feed. However, in otoliths, ANOVA indicated that Sr/Ca ratios did not differ significantly between groups fed on tubifex or formulated feed (p = 0.118). Otolith Sr/Ca ratios were negatively correlated with fish growth rates while the growth rates differed significantly among rearing conditions with different salinities and diets. Partition coefficients of the Sr/Ca ratios from ambient water to fish tissues and otoliths significantly increased with salinity. The Sr/Ca ratios of Japanese Eel otoliths thus were positively correlated with the ambient salinity and decreased with increasing fish growth rate, but was not affected by fish diet.     

Diversity of life strategies of pikepech <Emphasis Type="Italic">Sander lucioperca</Emphasis> (L.) in the Lower Volga (by the data of microelement composition of otolithes)

Strontium (Sr) and calcium (Ca) contents and Sr/Ca ratios in otoliths of pikeperch Sander lucioperca have been studied by X-ray fluorescent microanalysis. A sample of 95 specimens from the Lower Volga region are studied. The fish were obtained in a section of the Akhtuba River at a distance of 250–279 km upstream from the Caspian Sea. Some specimens in the sample have low strontium content throughout the otolith. The other fish have increased both Sr content and Sr/Ca ratios in core zones of otolith or near its outer edge: the Sr/Ca ratio is over 6.00 × 10^–3, with a maximum at 10.02 × 10^–3. The fish with a low Sr/Ca ratio live in freshwater and demonstrate a resident life history strategy. Pikeperch that have increased ratios of Sr/Ca in outer zones of otolith demonstrate a migratory life history strategy. These fish leave the freshwater of the Volga and feed in brackish water of the Northern Caspian Sea. Some migratory fish migrate to the sea as underyearlings, stay in a brackish water for 1 year or less, and then return back to fresh water for the rest of its life. The other group migrates back and forth from freshwater to the sea several times during their life. Finally, some fish leave freshwater for 1–2 years, spending this time in the Northern Caspian Sea. The conclusion is that the pikeperch stock in the Lower Volga has a complicated composition and consists of freshwater (residents and rheodromic ones) and migratory (anadromous and semianadromous) fish.     

Use of otolith chemistry to examine patterns of diadromy in the threatened Australian grayling Prototroctes maraena

Otolith chemical analyses (proton-induced X-ray emission and laser ablation inductively coupled plasma mass spectrometry) were used to examine patterns of diadromy in Australian grayling Prototroctes maraena collected from three rivers in south-eastern Australia. Concentrations of Sr:Ca and Ba:Ca were measured in the sagittal otoliths of 25 fish and in water samples collected on two occasions from multiple sites within freshwater, estuarine and marine reaches of the rivers. The results provide evidence of marine residency during the juvenile phase, with high Sr:Ca and low Ba:Ca in the inner region of the otoliths that were consistent across all three rivers. This general pattern was apparent for all individuals examined, suggesting that diadromous migration may be an obligate aspect of the life history. The chemical signatures of the outer regions of the otoliths were distinct between the three rivers, whilst there was no difference in the inner regions of the otoliths. This suggests that juveniles from different rivers may reside in a relatively homogenous chemical environment, such as the sea, and populations in coastal Victorian rivers may share a common marine recruitment source.     

Using energy dispersive x-ray fluorescence microchemistry to infer migratory life history of Atlantic sturgeon

Atlantic sturgeon migrate between ocean and freshwater habitats to spawn, and juveniles spend several years in fresh/brackish water before returning to the ocean. Because strontium/calcium (Sr/Ca) ratios are diagnostic for freshwater and marine environments, we examined the utility of energy-dispersive x-ray fluorescence (EDXRF) to quantify Sr/Ca ratios of Atlantic sturgeon pectoral fin spines. Atlantic sturgeon spines from wild adults and experimental juveniles were analyzed along a linear transect from the primordium to the outermost point. To verify the technique hatchery juvenile Atlantic sturgeon were held in experimental tanks at <0.5, 13?C15, or 33?C35?? and sampled after 5?months. Sr/Ca ratios of experimental hatchery fish increased with salinity, and Sr/Ca ratios in wild adults varied predictably along the measurement transect. However, the ratio decreased in the outermost region of the spine in mature fish collected during a return to freshwater for spawning. Therefore EDXRF is a useful tool to track individual movements of Atlantic sturgeons and other diadromous fish.     

Extrinsic factors affecting otolith chemistry – implications for interpreting migration patterns in a diadromous fish

Much has been revealed about fish migration, including diadromous behaviour, through the use of otolith chemistry. Manipulative experiments assist with unravelling information on otolith chemical composition and incorporation thereby answering specific questions on diadromous movements. In this study, a laboratory-based experiment was used to determine the relative and interactive effects of salinity and water temperature on the composition of three key elements (Sr, Ba and Mg) within the otolith of a catadromous fish, Percalates novemaculeata, endemic to south-eastern Australia. Otolith incorporation of Sr and Ba was positively related to ambient water concentration, whereas Mg incorporation was not. Sr and Ba increased and decreased significantly across salinity gradients, respectively, with minor positive effects of temperature also being detected. Salinity and temperature interacted to significantly affect the elemental concentration ratios for Ba: Ca in otoliths. Discrimination between fresh water and marine environments shows promise for interpreting P. novemaculeata residency based on these elements alone. However, deciphering finer scale movements within estuarine environments may be difficult. Our data highlights the importance of multifactorial validation experiments and suggests complementary use of multiple approaches for unravelling species-specific patterns of fish movement and habitat use.     

Diverse migratory histories of Japanese Trachidermus and Cottus species (Cottidae) as inferred from otolith microchemistry

The migratory histories of Japanese freshwater sculpins, one Trachidermus and four Cottus species, were 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 otoliths of Cottus nozawae showed consistently low Sr : Ca ratios, with an average of 3·37 × 10^?3 from the core to the edge, suggesting a freshwater resident life cycle. In contrast, the otolith Sr : Ca ratios for Trachidermus fasciatus and Cottus kazika changed along the life history transects possibly in accordance with their migration patterns from sea to fresh water. The ratios of T. fasciatus and C. kazika averaged 5·4 × 10^?3 and 5·3 × 10^?3 respectively, in the otolith region from the core to the points 450–890 μm, and changed to the lower levels, averaging 2·0 × 10^?3 and 2·7 × 10^?3, in the outer otolith region. These data suggest that both the species have a catadromous life cycle. The otoliths of Cottus hangiongensis had low Sr : Ca ratios in the two regions from the core to the points 15–30 μm and the points 415–582 μm to the edge, averaging 2·0 × 10^?3 and 1·9 × 10^?3, with significantly higher ratios in the narrow area between these regions, averaging 4·6 × 10^?3. Similar ontogenetic changes in otolith Sr : Ca ratios were found in the otoliths of Cottus amblystomopsis, suggesting their amphidromous life cycle. These findings suggest that otolith Sr : Ca ratios reflect individual life histories and that Japanese Trachidermus and Cottus species have diverse migratory histories.     

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     

Differences in the Trace Element Deposition in Otoliths Between Marine- and Freshwater-resident Japanese Eels, Anguilla japonica, as Determined by Laser Ablation ICPMS

Synopsis In order to determine whether the trace element composition in otolith of the Japanese eel Anguilla japonica could be used to determine its habitat use, we used laser ablation inductivity coupled plasma mass spectrometry (LA-ICPMS) to assay sectioned otoliths of both marine-resident (sea eels) and freshwater-resident (river eels) eels. A close linear relationship in the Sr:Ca ratios between EPMA (X-ray analysis with an electron microprobe) and LA-ICPMS analyses was found, suggesting that the latter technique could be used to separate the marine and freshwater life phases. Elemental signatures in the otolith outside the elver mark showed significant differences in Cr:Ca, Mn:Ca, and Ba:Ca ratios as well as Sr:Ca ratios between sea and river eels. These results indicate that the elemental compositions may reflect environmental variability between marine and fresh water masses. Thus, those elemental ratios determined by LA-ICPMS analysis seem to have the potential to help distinguish the habitat of the eel.     

洞庭湖中是否存在溯河洄游型刀鲚

利用电子探针微区分析(EPMA)技术, 分析了2018年8月采自湖南省岳阳市洞庭湖畔水产市场2尾刀鲚(Coilia nasus)干制个体的矢耳石中锶和钙微化学特征。定量线分析结果显示, 2尾样本的耳石锶钙比值(Sr/Ca×1000)波动显著, 从耳石核心到边缘可分别出现对应淡水生境的锶钙比值<3的低值区, 对应于半咸水生境的3—7高值区和再次为对应于淡水生境<3的低值区。面分布分析结果更直观显示两尾刀鲚个体的耳石从核心到边缘对应于不同盐度生境的锶含量变化图谱, 即从耳石核心到边缘可分别出现对应于淡水生境的蓝色区域, 对应长江口外半咸水生境的黄绿色区域和再次对应于淡水生境的蓝色区域。这种淡水-半咸水-淡水的生境转换显示出了上述个体的典型溯河洄游生活史“履历”。研究显示目前距长江河口约1400 km的洞庭湖中仍可能存在溯河洄游型刀鲚个体。     

基于耳石微化学的乌苏里白鲑生境履历分析

为了分析乌苏里白鲑(Coregonus ussuriensis Berg)的洄游生态类型及生境履历, 采用X射线电子探针微区分析技术(EPMA)研究了2013年12月、2014年1月黑龙江干流2尾和松花江干流2尾乌苏里白鲑矢耳石的锶(Sr)、钙(Ca)元素微化学特征。定量线分析结果显示, 乌苏里白鲑耳石锶钙比值(Sr/Ca×103)波动显著, 具有对应淡水生活的低值区(小于5), 而且还有对应咸水生活的高值区(大于10), 显示了乌苏里白鲑的溯河洄游履历; 耳石元素面分析结果也显示乌苏里白鲑的淡水-咸水洄游的特征。结合耳石年轮特征分析, 乌苏里白鲑具有年度(季节)洄游特征, 部分群体有规律在淡水-咸水中季节洄游, 不同个体在淡水、咸水之间停留时间存在较大差异。研究表明, 乌苏里白鲑伴随着季节性洄游特征, 具有江海或者淡水-河口之间洄游的履历。     

Factors influencing Sr/Ca ratios in otoliths of Girella elevata: an experimental investigation

The utility of Sr/Ca ratios in otoliths as indicators of thermal history in fish was investigated for juvenile Girella elevata . There was no direct relationship between ratios of Sr/Ca and temperature of the water, as has been assumed in many previous studies. Sr/Ca ratios did not decrease when water temperature was elevated from 19 to 28° C. Elevation of ambient Sr levels in the sea water caused a significant increase in Sr/Ca ratios in otoliths, and there was a detectable increase in Sr/Ca ratios in otoliths of some individuals fed an Sr-enriched diet. Multiple factors may influence natural Sr/Ca ratios in otoliths of juvenile G. elevata and their interactions make it difficult to interpret the chronology of conditions experienced by an individual fish during early life in the pelagic or benthic environment.     

Timing and rate of sexual maturation of European eel in brackish and freshwater environments

Maturation rates in three European eel Anguilla anguilla populations increased from September to October and were slightly lower in fresh water than in brackish waters. Average and variance of total length at maturation were larger in females than males.     

Migration flexibility between freshwater and marine habitats of the pond smelt Hypomesus nipponensis

The life histories of the pond smelt Hypomesus nipponensis collected from Japanese fresh waters and brackish (sea) waters were studied 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. The pond smelt living in a freshwater environment showed consistently low Sr:Ca ratios throughout the otolith, averaging 1·2–1·3 × 10⁻³. These samples were identified as a standard freshwater type. In contrast, fish collected from the intertidal zone showed higher otolith Sr:Ca ratios than those in the standard freshwater type, and the ratios fluctuated along the growth phase. In addition to the two representative life‐history types of H. nipponensis , i. e . freshwater and anadromous life‐history types, other pond smelts were found to have an estuarine resident life history‐type with no freshwater phase, indicating that the pond smelt has a flexible migration strategy with a high degree of behavioural plasticity and an ability to utilize the full range of salinity in its life history.