Migration behaviour of twaite shad Alosa fallax assessed by otolith Sr:Ca and Ba:Ca profiles

Individual migration behaviour during the juvenile and adult life phase of the anadromous twaite shad Alosa fallax in the Elbe estuary was examined using otolith Sr:Ca and Ba:Ca profiles. Between hatching and the end of the first year of life, juveniles showed two migration patterns. Pattern one exhibited a single downstream migration from fresh water to the sea with no return into fresh water. In contrast, pattern two showed a first migration into the sea, then a return into fresh water and, finally, a second downstream migration into marine water. This first report of migration plasticity for A. fallax points to different exposure times to estuarine threats depending on the migration strategy. In adults, high Sr:Ca and low Ba:Ca in the majority of individuals confirmed prior reports of a primarily marine habitat use. Patterns reflecting spawning migrations were rarely observed on otoliths, possibly due to the short duration of visits to fresh water.     

Juvenile downstream migration patterns of an anadromous fish,allis shad (Alosa alosa), before and after the population collapse in the Gironde system,France

Diadromous fish have exhibited a dramatic decline since the end of the 20th century. The allis shad (Alosa alosa) population in the Gironde-Garonne-Dordogne (GGD) system, once considered as a reference in Europe, remains low despite a fishing ban in 2008. One hypothesis to explain this decline is that the downstream migration and growth dynamics of young stages have changed due to environmental modifications in the rivers and estuary. We retrospectively analysed juvenile growth and migration patterns using otoliths from adults caught in the GGD system 30 years apart during their spawning migration, in 1987 and 2016. We coupled otolith daily growth increments and laser ablation inductively-coupled plasma mass spectrometry measurements of Sr:Ca, Ba:Ca, and Mn:Ca ratios along the longest growth axis from hatching to an age of 100 days (i.e., during the juvenile stage). A back-calculation allowed us to estimate the size of juveniles at the entrance into the brackish estuary. Based on the geochemistry data, we distinguished four different zones that juveniles encountered during their downstream migration: freshwater, fluvial estuary, brackish estuary, and lower estuary. We identified three migration patterns during the first 100 days of their life: (a) Individuals that reached the lower estuary zone, (b) individuals that reached the brackish estuary zone, and (c) individuals that reached the fluvial estuary zone. On average, juveniles from the 1987 subsample stayed slightly longer in freshwater than juveniles from the 2016 subsample. In addition, juveniles from the 2016 subsample entered the brackish estuary at a smaller size. This result suggests that juveniles from the 2016 subsample might have encountered more difficult conditions during their downstream migration, which we attribute to a longer exposure to the turbid maximum zone. This assumption is supported by the microchemical analyses of the otoliths, which suggests based on wider Mn:Ca peaks that juveniles in 2010s experienced a longer period of physiological stress during their downstream migration than juveniles in 1980s. Finally, juveniles from the 2016 subsample took longer than 100 days to exit the lower estuary than we would have expected from previous studies. Adding a new marker (i.e., Ba:Ca) helped us refine the interpretation of the downstream migration for each individual.     

The Great Melting Pot. Common Sole Population Connectivity Assessed by Otolith and Water Fingerprints

Quantifying the scale and importance of individual dispersion between populations and life stages is a key challenge in marine ecology. The common sole (Solea solea), an important commercial flatfish in the North Sea, Atlantic Ocean and the Mediterranean Sea, has a marine pelagic larval stage, a benthic juvenile stage in coastal nurseries (lagoons, estuaries or shallow marine areas) and a benthic adult stage in deeper marine waters on the continental shelf. To date, the ecological connectivity among these life stages has been little assessed in the Mediterranean. Here, such an assessment is provided for the first time for the Gulf of Lions, NW Mediterranean, based on a dataset on otolith microchemistry and stable isotopic composition as indicators of the water masses inhabited by individual fish. Specifically, otolith Ba/Ca and Sr/Ca profiles, and δ¹³C and δ¹⁸O values of adults collected in four areas of the Gulf of Lions were compared with those of young-of-the-year collected in different coastal nurseries. Results showed that a high proportion of adults (>46%) were influenced by river inputs during their larval stage. Furthermore Sr/Ca ratios and the otolith length at one year of age revealed that most adults (∼70%) spent their juvenile stage in nurseries with high salinity, whereas the remainder used brackish environments. In total, data were consistent with the use of six nursery types, three with high salinity (marine areas and two types of highly saline lagoons) and three brackish (coastal areas near river mouths, and two types of brackish environments), all of which contributed to the replenishment of adult populations. These finding implicated panmixia in sole population in the Gulf of Lions and claimed for a habitat integrated management of fisheries.     

Migration and habitat use of the tropical eels Anguilla marmorata and A. bicolor pacifica in Vietnam

The patterns of use of marine and freshwater habitats by the tropical anguillid eels Anguilla marmorata and A. bicolor pacifica were examined by analysing the otolith strontium (Sr) and calcium (Ca) concentrations of yellow (immature) and silver (mature) stage eels collected in Vietnamese waters. In A. marmorata, the change in the Sr:Ca ratios outside the high Sr:Ca core was generally divided into three patterns: (1) typical catadromous life history pattern; (2) constant residence in brackish water; and (3) habitat shifting between sea and brackish waters with no freshwater life. In A. bicolor pacifica, no eels had a general life history as freshwater residents. The eels were also divided into three patterns: (1) constant residence in sea water; (2) constantly living in brackish water; and (3) habitat shifting from brackish to sea water with no freshwater residence. The mean Sr:Ca ratio value after recruitment to coastal waters ranged from 1.73 to 5.67 × 10^?3 (mean 3.2 × 10^?3) in A. marmorata and from 2.53 to 6.32 × 10^?3 (mean 4.3 × 10^?3) in A. bicolor pacifica. The wide range of otolith Sr:Ca ratios in both species indicated that the habitat use of these tropical eels was facultative among fresh, brackish, and marine waters during their growth phases after recruitment to coastal areas. Tropical eel species may have the same behavioural plasticity as temperate anguillid species regarding whether to enter freshwater or to remain in estuarine and marine environments.     

Unravelling the life-history patterns and habitat preferences of the Japanese eel (Anguilla japonica) in the Pearl River,China

Eels have fascinated biologists for centuries due to their amazing long-distance migrations between freshwater habitats and very distant ocean spawning areas. The migratory life histories of the Japanese eel, Anguilla japonica, in the waters of south China are not very clear despite its ecological importance, and the need for fishery regulation and management. In this study, strontium (Sr) and calcium (Ca) microchemical profiles of the otoliths of silver eels were measured by X-ray electron probe microanalysis based on data collected from different habitats (including freshwater and brackish habitats), in the large subtropical Pearl River. The corresponding habitat preference characteristics were further analysed using redundancy analysis (RDA). A total of 195 Japanese eels were collected over 6 years. The collected individuals ranged from 180 to 771 mm in total length and from 8 to 612 g in body weight. Two-dimensional pictures of the Sr:Ca concentrations in otoliths revealed that the A. japonica in the Pearl River are almost entirely river eels, spending the majority of their lives in fresh water without exposure to salt water, while the catadromous migration time has delayed about 1 month in the Pearl River estuary in the past 20 years. RDA analysis further indicated that juveniles and adults preferred water with high salinity and high tide levels. Youth preferred habitats with high river fractals. Our findings contribute to a growing body of evidence showing that the eels are extremely scarce currently and conservation measures against them are imminent, including the protection of brackish and freshwater areas where they live in south China.     

Life history and migration of Sakhalin taimen, Hucho perryi, caught from Lake Akkeshi in eastern Hokkaido, Japan, as revealed by Sr:Ca ratios of otoliths

Microchemical analysis of the strontium (Sr) and calcium (Ca) ratios of otoliths was conducted to determine the life history and migration of anadromous Sakhalin taimen, Hucho perryi. In 2008 and 2009, 10 specimens were sampled from Lake Akkeshi in eastern Hokkaido, Japan. Our results indicated that some specimens migrated to brackish waters during their early life histories. Because the Sr:Ca ratios of the specimens in this study were all less than those of specimens from Sakhalin Island during a previous study, specimens from Lake Akkeshi may have migrated to brackish water, or may have remained in the ocean for only a short period.     

Life history and age at maturity of an anadromous smelt, the eulachon Thaleichthys pacificus (Richardson)

Trace element and fork length ( L _F) frequency analyses of eulachon Thaleichthys pacificus otoliths were used to determine age at maturity and repeat spawning potential, two aspects of eulachon life history that are not known but are important for successful management of this species. The L _F-frequency analysis for ocean caught and spawning eulachon was used to estimate age at maturation. Two size classes of eulachon were caught in the ocean and spawning eulachon were consistently the largest fish indicating that spawners from mid-coast of British Columbia were 3 years old. Laser-ablation inductively-coupled plasma mass spectrometry (LA-ICP-MS) was also used to reconstruct the Ba:Ca and Sr:Ca molar ratios deposited spatially into the otolith to estimate spawner age for five populations of eulachon. Age at maturation differed among populations examined. Based on the seasonal fluctuations in Ba:Ca molar ratios caused by seasonal upwelling of deep waters, it was determined that more southerly populations spawned at a younger age than the northern populations examined. Southern populations of eulachon, Columbia River, Washington, U.S.A., spawn after 2 years. Eulachon from the Fraser, Kemano and Skeena Rivers in British Columbia, Canada, generally mature after 3 years. Some Skeena River eulachon and most of the eulachon from the Copper River, Alaska, U.S.A., matured after 4 years. In contrast to the Ba:Ca molar ratios in the otolith, Sr:Ca molar ratios maintained a relatively flat profile over the life of the eulachon. The lack of a change in Sr:Ca ratios within the otolith, the single size class of spawners across all systems and the single age class within most populations strongly suggest that eulachon in the present study are semelparous.     

Spawning energetics and otolith microchemistry provide insights into the stock structure of bonga shad Ethmalosa fimbriata

The gross energy content of spawning batches and the microchemistry of sagittal otoliths in individual female bonga shad Ethmalosa fimbriata were compared between contrasting sampling sites at the Senegalese southern coast and inside the hypersaline Sine Saloum Estuary. Results show that females spawning in the estuary's middle reaches invested almost three times more energy into reproduction (115 ± 65 J g⁻¹ body mass) than their neritic counterparts (39 ± 34 J g⁻¹ body mass). Also, female otolith levels of Ba:Ca, Sr:Ca and Zn:Ca either differed significantly between study sites or could be linked to heterogeneous environmental variables. A quadratic discriminant function analysis provided evidence of segregated spawning populations of E. fimbriata in southern Senegalese waters.     

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.     

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.     

Stable Isotopic and Trace Elemental Compositions of Otoliths and the Stock Structure of Pacific Cod, Gadus macrocephalus

Synopsis We examined the stock composition and life history of Pacific cod, Gadus macrocephalus, in the northeast Pacific Ocean from sagittal otoliths collected from three marine fishing areas in Washington State, U.S.A. We analyzed both stable isotope ratios (δ¹⁸O and δ¹³C) and trace elemental concentrations (Sr, Mg, Na, Fe, Mn) for these otoliths. The combination of δ¹⁸O and δ¹³C, and correlation of δ¹⁸O vs. 1000Sr/Ca and 1000Mg/Ca showed clear separations between North Puget Sound and coastal cod, suggesting there might be two different spawning stocks in the region. The North Puget Sound cod might represent an ‘Estuary-type’ from the Strait of Georgia, whereas coastal cod might represent an ‘Ocean-type’ from the Pacific west coast. Isotopic variations from five representative otoliths also showed a two-stage life history for Pacific cod, and a critical transition period for cod in migration to the ocean or in age of sexual maturity. These chemical interpretations and conclusions appear in agreement with biological observations of Pacific cod, and are consistent with results of previous studies for other marine fish species in nearby areas.     

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.     

Environmental signature in the otolith elemental fingerprint of the tapertail anchovy, Coilia mystus, from the Changjiang estuary, China

The ontogenetic patterns of habitat use and the migratory history of the tapertail anchovy, Coilia mystus, collected in the Changjiang estuary around Chongming Island, China, were studied by examining the environmental signature in the otolith strontium (Sr) and calcium (Ca) fingerprints using electron probe microanalysis (EPMA). Our results suggest that the migration strategy of C. mystus is much more flexible than supposed in the literature to date. The spring spawning population of C. mystus from the studied area was found to consist of individuals with different migration histories. Although the tapertail anchovy seems to be an anadromous fish that spawns and hatches in a freshwater habitat, it can also use a freshwater environment in non‐spawning seasons. The otolith EPMA of the elemental fingerprint (Sr x‐ray maps and Sr : Ca ratios) is an environmental indicator that can be applied to the migratory ecology of other important diadromous species in China.     

Retracing migration pattern in reproductive and non-reproductive female kutum Rutilus frisii,in south Caspian Sea,using otolith microchemistry

Because trace elements of otoliths are considered a natural marker capable of recognizing the chemical composition of ambient water and fish migration history, these elements could be potentially used to analyse the movement of reproductive (R) and non-reproductive (NR) mature-sized fish. Supposedly, it is not essential for NR individuals to migrate to rivers for spawning because they do not have developed gonads. To investigate the potential differences in migration history between female R and NR kutum, Rutilus frisii, in the southwest waters of the Caspian Sea, the ratios of Sr, Ba, Mg, Na, K and P to Ca in otoliths (from the core to the edge) were examined using laser ablation–inductively coupled plasma–mass spectrometry. In NR fish, a significant increase in Sr:Ca ratio in the otoliths' growth rings, likely due to greater seawater residency, and an increase in Ba:Ca ratio in the last two rings were observed. Increased Ba:Ca ratio could be due to the movement of NR mature-sized fish to the coastal zones for foraging. Seasonal physiological factors such as gonad maturation and spawning activity are more likely to be involved in differences in the other elemental ratios (Mg, Na, K and P). These results suggest that microchemical analyses of growth rings of otolith can be used as a valuable tool for better understanding the movement pattern of different types of adult fish, which could be completed with data from other methods like tagging.     

Spatio-temporal variation in the elemental compositions of otoliths of southern bluefin tuna Thunnus maccoyii in the Indian Ocean and its ecological implication

The elements Na, Mg, Mn, Ca, Sr and Ba in otoliths of southern bluefin tuna Thunnus maccoyii , collected from their feeding ground in the central Indian Ocean and spawning ground between southern Java and north-western Australia were measured by laser-ablation inductively coupled plasma mass spectrometry (LA-ICPMS) and compared among sampling locations and developmental stages. The Na, Mg and Mn to Ca concentration ratios were significantly higher at the larval stage than at the adult stage, and the ratio reached a peak at the first inflection point of the otolith, mean ± s.d. 43·3 ± 4·9 days after hatching and decreased sharply to a low level thereafter. The temporal change of the elements:Ca ratios in the first inflection point corresponded to the life stage transition from larva to juvenile, indicating that the uptake rate of elements from ambient waters was significantly influenced by the ontogenetic change in the fish. The elemental composition at the otolith edge differed significantly in sub-adults on the feeding grounds and adults on the spawning grounds. Thus, the otolith elemental composition can be used as a biological tracer to study the time of the ontogenetic shift and to reconstruct the past migratory environmental history of T. maccoyii . In addition, the elemental composition of the otolith core of the adult was similar between feeding and spawning grounds, indicating that the fish in the Indian Ocean had the same larval origin, which is consistent with the single spawning population hypothesis.     

Habitat residency and movement patterns of Centropomus parallelus juveniles in a subtropical estuarine complex

Sixty Centropomus parallelus juveniles were collected in March 2013 in two locations (Tromomó and Guaraguaçu) inside the Paranaguá estuarine complex, southern Brazil. The habitat residency and movement patterns of the individuals were inferred from Sr:Ca ratios and age recorded in the otoliths. Data suggest that the species spawns preferentially in brackish areas mainly from October to January, and that growth rate during the early juvenile stage could be influenced by environmental salinity. Furthermore, the data also show that C. parallelus can occupy diverse salinity habitats and migrate among marine, brackish and freshwater areas within the Paranaguá estuarine complex, showing a high environmental plasticity and adaptation.     

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.     

Migration of Sakhalin taimen (<Emphasis Type="Italic">Parahucho perryi</Emphasis>): evidence of freshwater resident life history types

Sakhalin taimen (Parahucho perryi) range from the Russian Far East mainland along the Sea of Japan coast, and Sakhalin, Kuril, and Hokkaido Islands and are considered to primarily be an anadromous species. We used otolith strontium-to-calcium ratios (Sr/Ca) to determine the chronology of migration between freshwater and saltwater and identify migratory contingents of taimen collected from the Koppi River, Russia. In addition, we examined taimen from the Sarufutsu River, Japan and Tumnin River, Russia that were captured in marine waters. Transects of otolith Sr/Ca for the Sarufutsu River fish were consistent with patterns observed in anadromous salmonids. Two fish from the Tumnin River appeared to be recent migrants to saltwater and one fish was characterized by an otolith Sr/Ca transect consistent with marine migration. Using these transects as benchmarks, all Koppi River taimen were classified as freshwater residents. These findings suggest more work is needed to assess life history variability among locations and the role of freshwater productivity in controlling migratory behavior in taimen.     

The post‐spawning movements of migratory brown trout Salmo trutta L.

The post spawning behaviour of sea trout Salmo trutta was studied over a 2 year period in the river and estuary of the River Fowey, south‐west England. Forty‐five sea trout kelts were trapped immediately after spawning in December and intraperitoneally tagged with miniature acoustic transmitters. The subsequent emigration into coastal waters was monitored using acoustic receivers deployed throughout the river catchment. The levels of gill Na⁺K⁺ATPase activity in sea trout kelts sampled at the same time as the tagged fish were within the range of 2·5 to 4·5 μmol Pi per mg protein per h indicating that the post‐spawning fish were not physiologically adapted to salt water. The tagged kelts were resident in fresh water between 4 and 70 days before entering the estuary. Sixty two per cent of the tagged kelts subsequently migrated successfully into coastal waters, with a higher success rate for male fish (75%) than females (58%). There was a significant size related difference in the run‐timing of the kelts with the larger fish moving more quickly into coastal waters after spawning than smaller fish. Seaward migration within fresh water was predominantly nocturnal and generally occurred in conjunction with increasing river discharge and rising water temperature. Migration through the estuary continued to be predominantly nocturnal and occurred during an ebbing tide. Residency within the estuary varied amongst individuals although it was invariably short, with most fish moving out into coastal waters within one to two tidal cycles. Five tagged kelts returned from the coastal zone and re‐entered fresh water during April and June. Marine residence time varied between 89 and 145 days (mean 118 days) and the minimum estimated marine survival was c. 18%. One of these sea trout was subsequently recaptured after successfully spawning in the vicinity where it had been previously tagged demonstrating a degree of spawning site fidelity.     

Variations in the migratory history of the tropical catadromous eels Anguilla bicolor bicolor and A. bicolor pacifica in south-east Asian waters

Fish movements between aquatic habitats of different salinity ranges (fresh, estuarine, marine) by the tropical catadromous eels Anguilla bicolor bicolor and A. bicolor pacifica were examined by analysing the otolith strontium and calcium concentrations of yellow (immature) and silver (mature) stage eels collected in south-east Asian (Indonesia, Malaysia and Vietnam) waters. The ratios suggest that all migratory-type eels, including freshwater, brackish water and marine residents, pass the river mouth. However, the habitat preference was different among the sites (countries). In Indonesia and Vietnam, most A. bicolor bicolor and A. bicolor pacifica were either marine or brackish water residents in this study. Alternatively, most A. bicolor bicolor were freshwater residents in Malaysia; such a typical catadromous migration pattern in these eels has not been found in previous studies. The wide range of otolith Sr:Ca in both subspecies indicates that the habitat use of these tropical eels was opportunistic among fresh, brackish and marine waters during their growth phases following recruitment to coastal areas. The geographical variability of migratory histories suggests that habitat use might be determined by the inter and intraspecific competition and environmental conditions at each site.