Spatial variability of trace elements in fish otoliths: comparison with dietary items and habitat constituents in seagrass meadows

Juvenile trumpeters Pelates sexlineatus , two main prey items (amphipods and polychaetes) and seagrass constituents (detritus, young and old leaves of Zostera capricorni ) were collected from two sites within six estuaries, to compare the spatial variability of elemental composition (Li, Mn, Sr, Cd, Ba and Pb). Average elemental values were lower in the fish otoliths except for Sr. Significant differences in elemental composition of otoliths were detected among estuaries (Mn and Pb). Spatial differences in concentrations of Cd were found in amphipods and polychaetes; the latter also showed differences for Ba and Pb. Detritus and Z. capricorni leaves showed appreciable spatial differences for most of the elements. Spatial variability between sites was generally high. There were differences in the multivariate composition of trace elements found in the otoliths of fish among estuaries. One estuary was consistently separated from the other estuaries in the ordinations. Significant correlations were detected between concentrations of Mn in otoliths and concentrations in prey, Z. capricorni and detritus. A significant relationship was also detected between Pb in otoliths and detritus. There were significant correlations between concentrations of Mn, Sr and Ba in detritus and that in otoliths. The study demonstrated that trophic transfer and habitat constituents may be considered as a potential source for some of the elemental accumulation in fish otoliths.     

Effects of handling and storage methods on the concentrations of elements in deep‐water fish otoliths

Sagittal otoliths of Coryphaenoides rupestris (roundnose grenadier), Helicolenus dactylopterus (bluemouth) and Merluccius merluccius (European hake) were collected using a variety of handling and storage treatments and their elemental composition was examined using inductively coupled plasma mass spectrometry. Some differences between element concentrations were identified between the control and treatment groups, most notably for the element Li. For H. dactylopterus and M. merluccius , Li concentrations were significantly higher in the otoliths extracted with metal forceps and stored in paper envelopes (treatment), compared to those from the same fishes that had been extracted using plastic forceps and stored in polyethylene vials (control). Lower concentrations of Ba and Cr were found in M. merluccius otoliths extracted from fish that had been stored frozen. The presence or absence of elemental concentrations above the instrumental limits of detection was noted, but no significant differences were identified between otolith pairs for any of the treatments. The differences between otolith pairs attributable to storage and handling effects are small compared to between‐area differences.     

Small-scale spatial variation in the elemental composition of otoliths of Stegastes nigricans (Pomacentridae) in French Polynesia

Solution-based inductively coupled plasma-mass spectrometry (ICP-MS) was used to determine if Stegastes nigricans collected from 15 sites in French Polynesia could be distinguished by the trace element composition of their otoliths. A total of 293 adults were collected by spearing and their otoliths were analysed. We found that elemental signatures differed significantly among sites within and between the islands of Tahiti and Moorea (p<0.001), primarily due to variation in concentrations of the elements Ba, Ca, Li, Mg, Mn, Na, Sr and Y. The otoliths of fish collected within Papeete Harbour in Tahiti had distinctive elemental signatures characterised by relatively high concentrations of Mn. Otoliths of these fish could be distinguished from others that were collected only a small distance (200 m) from the harbour. This is the first time that differences in chemical composition of otoliths have been reported at such small spatial scales and this trait may prove useful for the studies of connectivity of populations at within reef scales.     

Elemental composition of otoliths from Atlantic croaker along an estuarine pollution gradient

Elemental compositions of otoliths of young-of-the-year Atlantic croaker Micropogonias undulatus at stations along a pollution gradient in Galveston Bay, Texas, were compared with those of sediments and livers to establish the level of consonance between otolith and habitat chemistries. Both coupled and uncoupled trends were indicated, with the bulk of the results pointing to the need for additional research to test the working hypothesis that the otolith stores a temporal record of elemental chemistry of resident waters. Lack of simple correlations between elemental concentrations in sediment or liver and in otoliths indicate that other biological and geochemical factors influence otolith composition.     

Morphometry and composition of aragonite and vaterite otoliths of deformed laboratory reared juvenile herring from two populations

Vaterite otoliths were sampled from two reared populations (Celtic and Clyde Seas) of juvenile herring Clupea harengus. The crystallography, elemental composition and morphometry were analysed and compared with those of normal aragonite otoliths. The incidence of vaterite otoliths in the juveniles sampled (n = 601) ranged from 7·8% in the Clyde population to 13·9% in the Celtic Sea population, and was 5·5% in the small sample (n = 36) of wild adults examined. In all but one case fish had only one vaterite otolith; the corresponding otolith of the pair was completely aragonite. Although the majority of the juveniles sampled showed craniofacial deformities, there was no link between the skull or jaw malformation and the incidence of vaterite otoliths. All vaterite otoliths had an aragonite inner area, and vaterite deposition began sometime after the age of 90 days. The vaterite otoliths were larger and lighter than their corresponding aragonite partners, and were less dense as a consequence of the vaterite crystal structure. The vaterite areas of the otoliths were depleted in Sr, Na and K. Concentrations of Mn were higher in the vaterite areas. The transition between the aragonite inner areas and the vaterite areas was sharply delineated. Within a small spatial scale (20 μm³) in the vaterite areas, however, there was co‐precipitation of both vaterite and aragonite. The composition of the aragonite cores in the vaterite otoliths was the same as in the cores of the normal aragonite otoliths indicating that the composition of the aragonite cores did not seed the shift to vaterite. Vaterite is less dense than aragonite, yet the concentrations of Ca analysed with wavelength‐dispersive spectrometry (WDS) were the same between the two polymorphs, indicating that Ca concentrations measured with WDS are not a good indicator of hypermineralized zones with high mineral density. The asymmetry in density and size of the otoliths may cause disruptions of hearing and pressure sensitivity for individual fish with one vaterite otolith, however, the presence of vaterite otoliths did not seem to affect the growth of these laboratory reared juvenile herring.     

Bromine patterns in Norwegian coastal Cod otoliths—a possible marker for distinguishing stocks?

Bromine was found to accumulate in otoliths of Norwegian coastal Cod Gadus morhua that were reared under known conditions. Despite the fact that the Cod were moved from one rearing environment to another, causing marked changes in some otolith elemental concentrations, bromine appeared to accumulate continuously along certain growth axes as revealed by 2-D elemental mapping. In contrast, North Sea and Baltic Sea Cod showed little to no patterning in Br. We suggest that Br uptake in otoliths may be under physiological and genetic control, and as such, may prove useful as a stock identification tool.     

Otolith elemental fingerprints of juvenile Pacific swordfish Xiphias gladius

The trace element composition of young‐of‐the‐year (YOY) juvenile swordfish Xiphias gladius sagittal otoliths were analysed as a preliminary test of the value of otolith elemental fingerprints for determining swordfish nursery ground origins in the central Pacific Ocean. A suite of five elements (Mg, Zn, Sr, Ba and Pb) was assayed with isotope dilution ICP‐MS; all elemental concentrations were roughly comparable to otoliths of other marine fishes. Multivariate analyses of elemental fingerprints based on Ba and Sr revealed differences between sample sites, and the magnitude of the differences increased with latitudinal separation. With more comprehensive sampling of nursery grounds, it should be possible to identify origin of nursery ground for adult swordfish by analysing the YOY juvenile portion of the sagittal otolith.     

Otoliths of deep water fishes: structure,chemistry and chemically-coded life histories

• 1.1. The gross anatomy and crystalline ultrastructure of the external surfaces has been described for the sagittal otoliths of a 6 million year old fossil otolith and the following bottom-living deep-water teleosts (families in parenthesis): Coryphaenoides (Nematonurus) armatus (Macrouridae), Coryphaenoides (Chalinura) mediterraneus (Macrouridae), Celonurus globiceps (Macrouridae), Coryphaenoids (Chalinura) leptolepis (Macrouridae), Antimora rostrata (Moridae), Halosaurus pectoralis (Halosauridae), Psychrolutes obesus and Neophrynichthys angustus (Psychroultidae).
• 2.2. Sections of otoliths were scanned by proton microprobe to yield three dimensional maps of the elemental density of calcium, strontium and, in some otoliths, zinc. The same sections were examined optically for opaque and hyaline zones. For some otoliths, broken sections were examined by SEM and their internal crystalline structure described.
• 3.3. The theoretical basis for using cycles in elemental (particularly strontium) composition to interpret life history is examined. Life history information from elemental composition is compared with other information derived both from otoliths and field observations of the species involved.

     

The detection of elements in larval otoliths from Atlantic herring using laser ablation ICP-MS

Trace element concentrations of otoliths from larval herring Clupea harengus collected from known spawning beds in the Celtic and Irish Seas, were investigated using laser ablation ICP-MS and compared with concentrations in the larval cores of juvenile otoliths from the same populations and year class. A range of elements (Mg, Zn, Sr, Ba and Pb) was detectable in early larval otoliths (20–40 µm diameter). Larval otolith concentrations exceeded the larval core concentrations of juvenile otoliths and also the concentrations reported in the literature, for Mg, Zn, Ba and Pb, indicating that the measurement of elements in larval otoliths was severely affected by post-mortem contamination, most likely due to adherence of tissue and endolymph residue on the otolith surface. Comparison of otolith composition between larvae from two freezing treatments showed that contamination from Mg and Zn was more serious in otoliths that had remained in frozen larvae for prolonged periods. Larval populations from the two seas showed significant differences in otolith Sr concentrations, which were consistent over two sampling years. Similar differences were seen in the corresponding juvenile populations. The results show that while early larval otoliths are extremely susceptible post-mortem contamination, Sr concentrations can be reliably measured using laser ablation ICP-MS and for this element, the detection of region specific differences is possible.     

Fish otolith chemistry influenced by exposure to multiple environmental variables

There is an increasing desire for researchers to use the elemental concentrations in fish otoliths to reconstruct environmental histories of fish. These reconstructions may be plausible due to the unique incorporation of elements into discrete layers of otolith material that correspond to daily growth, and because environmental variables of temperature, salinity, and water chemistry can influence otolith chemistry. However, it is essential to establish exactly how temperature, salinity, and the ambient concentration of elements influence otolith chemistry in order to interpret environmental histories of fish. Using a controlled laboratory experiment we tested the relative and interactive effects of temperature, salinity, and ambient concentration of strontium (Sr) and barium (Ba) on the resulting concentration of Sr and Ba in otoliths of black bream Acanthopagrus butcheri (Munro 1949). Salinity and concentration, and temperature and concentration interacted to affect the elemental concentration of Sr:Ca and Ba:Ca in otoliths. Regression analysis revealed that temperature and ambient concentration contributed most to the trend in otolith chemistry for both elements. Importantly, this is the first experiment to combine three environmental variables and assess their effect on otolith chemistry. Based on these results, it should be possible to use changes in the elemental concentration in otoliths to better reconstruct previous environments of temperature, salinity, and ambient water chemistry, which is especially useful when determining occupancy in habitats such as estuaries that display variable environmental characteristics.     

Otolith chemistry of prey fish consumed by a fish predator: does digestion hinder Russian doll techniques?

The effect of digestion by a predatory fish (largemouth bass Micropterus salmoides) on stable isotopic (δ¹³C and δ¹⁸O) and trace elemental (Sr:Ca and Ba:Ca) compositions of prey fish (bluegill Lepomis macrochirus) otoliths was investigated in a laboratory experiment. Trace element and stable‐isotopic signatures of L. macrochirus otoliths were not significantly altered for up to 16 h after L. macrochirus were consumed by M. salmoides. Prey fish otoliths recovered from predator digesta can retain environmental stable isotopic and trace elemental signatures, suggesting that determination of environmental history for prey fishes by stable‐isotope and trace‐element analysis of otoliths recovered from stomachs of piscivorous fishes will be feasible.     

Trace elements in the otoliths and dorsal spines of albacore tuna (Thunnus alalunga, Bonnaterre, 1788): An assessment of the effectiveness of cleaning procedures at removing postmortem contamination

Trace element analysis or “elemental fingerprinting” is widely used in stock structure analyses. Postmortem contamination of bony structures can confound the results of microconstituent studies or introduce an additional source of noise to the data, thus reducing the ability of the technique to detect real variation in trace element concentrations. Despite the potential for postmortem contamination during sample preparation, the effectiveness of the procedures used to remove potential contaminants from sectioned otoliths and other calcareous structures prior to laser ablation inductively coupled plasma mass spectrometry (LA ICP-MS) has not previously been addressed. Otoliths and dorsal spine sections of albacore tuna (Thunnus alalunga) collected from the North East Atlantic Ocean and the Mediterranean Sea were deliberately contaminated prior to analysis of trace element composition using LA ICP-MS. The effectiveness of three cleaning treatments (rinsing in ultrapure water, 30% hydrogen peroxide and ultrapure 5% nitric acid) at removing this postmortem contamination were compared. Magnesium and strontium were relatively robust to postmortem effects when exposed to contamination at concentrations of 50 ppm and 200 ppm respectively. Soaking in a solution containing Mn, Cs and Ba (50 ppm) caused a marked increase in the detected concentration of each element in both structures. Translucent bands in both structures were more susceptible to contamination. Rinsing in ultrapure water or hydrogen peroxide was not effective at removing Mn, Cs and Ba contamination from either calcareous structure. Washing the otoliths and spines in nitric acid successfully removed postmortem contaminants.The removal of otoliths from tuna damages the appearance of the fish and has an adverse effect on market value. However spines are easily removed, do not affect the appearance or value of the fish and are the most commonly used structure for age determination. A weak but significant correlation was observed between Ba in opaque zones in otoliths and dorsal spines. All other spine to otolith correlations were not significant. The results do not provide support for the use of spines as an alternative to otoliths in trace elemental analyses.     

Otolith trace element and stable isotopic compositions differentiate fishes from the Middle Mississippi River, its tributaries, and floodplain lakes

Naturally occurring stable isotope and trace elemental markers in otoliths have emerged as powerful tools for determining natal origins and environmental history of fishes in a variety of marine and freshwater environments. However, few studies have examined the applicability of this technique in large river-floodplain ecosystems. This study evaluated otolith microchemistry and stable isotopic composition as tools for determining environmental history of fishes in the Middle Mississippi River, its tributaries, and floodplain lakes in Illinois and Missouri, USA. Fishes were collected from 14 sites and water samples obtained from 16 sites during summer and fall 2006 and spring 2007. Otolith and water samples were analyzed for stable oxygen isotopic composition (δ¹⁸O) and concentrations of a suite of trace elements; otoliths were also analyzed for carbon isotopic composition (δ¹³C). Tributaries, floodplain lakes, and the Mississippi and Lower Missouri Rivers possessed distinct isotopic and elemental signatures that were reflected in fish otoliths. Fish from tributaries on the Missouri and Illinois sides of the middle Mississippi River could also be distinguished from one another by their elemental and isotopic fingerprints. Linear discriminant function analysis of otolith chemical signatures indicated that fish could be classified back to their environment of capture (Mississippi River, floodplain lake, tributary on the Illinois or Missouri side of the Mississippi River, or lower Missouri River) with 71–100% accuracy. This study demonstrates the potential applicability of otolith microchemistry and stable isotope analyses to determine natal origins and describe environmental history of fishes in the Middle Mississippi River, its tributaries, and floodplain lakes. The ability to reconstruct environmental history of individual fish using naturally occurring isotopic markers in otoliths may also facilitate efforts to quantify nutrient and energy subsidies to the Mississippi River provided by fishes that emigrate from floodplain lakes or tributaries.     

Otolith microchemistry of tropical diadromous fishes: spatial and migratory dynamics

Otolith microchemistry was applied to quantify migratory variation and the proportion of native Caribbean stream fishes that undergo full or partial marine migration. Strontium and barium water chemistry in four Puerto Rico, U.S.A., rivers was clearly related to a salinity gradient; however, variation in water barium, and thus fish otoliths, was also dependent on river basin. Strontium was the most accurate index of longitudinal migration in tropical diadromous fish otoliths. Among the four species examined, bigmouth sleeper Gobiomorus dormitor, mountain mullet Agonostomus monticola, sirajo goby Sicydium spp. and river goby Awaous banana, most individuals were fully amphidromous, but 9–12% were semi‐amphidromous as recruits, having never experienced marine or estuarine conditions in early life stages and showing no evidence of marine elemental signatures in their otolith core. Populations of one species, G. dormitor, may have contained a small contingent of semi‐amphidromous adults, migratory individuals that periodically occupied marine or estuarine habitats (4%); however, adult migratory elemental signatures may have been confounded with those related to diet and physiology. These findings indicate the plasticity of migratory strategies of tropical diadromous fishes, which may be more variable than simple categorization might suggest.     

Interactive effects of ontogeny, food ration and temperature on elemental incorporation in otoliths of a coral reef fish

The potential for environmental and physiological modification of elemental incorporation in otoliths is significant and must be validated before otoliths can be used reliably to estimate water parameters over the life history of a fish. We experimentally manipulated temperature and diet quantity for juvenile, sub-adult, and adult Acanthochromis polyacanthus, a tropical damselfish of the SW Pacific. Significant interactive effects between life history stage, temperature and food quantity were observed for otolith Ba/Ca, while significant interactions between stage and food were observed for Sr/Ca. Specific growth rates were negatively correlated with D_Ba and D_Sr for juveniles and sub-adults. These interactions indicated elemental incorporation dynamics varied depending on the life history stage, suggesting variation in effects of stage-specific metabolism or reproductive status. Our results highlight complex responses of elemental incorporation to both endogenous and exogenous factors. Interpretations of life history transects across otoliths must account for these effects to avoid confounding environmental variability with ontogenetic changes in physiology.     

An assessment of otoliths,dorsal spines and scales to age the long‐finned gurnard,Lepidotrigla argus,Ogilby, 1910 (Family: Triglidae)

Sagittal otoliths, dorsal spines and scales were critically assessed as structures to potentially determine the age of the long‐finned gurnard, Lepidotrigla argus. Counts were made of opaque growth increments and a readability score was assigned to each structure. Comparisons of growth increment counts were made between structures and between readings. All three structures showed some degree of readability and quantifiable growth increments, but this varied within fishes and between structures. Initial results showed that whole otoliths were more suitable to determine age estimates than dorsal spines and scales. Scales were considered unsuitable due to between reading ageing bias, variation in age estimates between structures, low precision and poor readability for this species. Dorsal spines showed evidence of loss of growth increments due to hollowing of the vascular core, which resulted in underestimation of older individuals in comparison to whole otoliths. Further analysis showed that growth increment counts from whole otoliths were lower for older individuals in comparison to sectioned otoliths. It is suggested that this is because of decreased clarity of growth increments towards the outer margin of whole otoliths in older individuals; this problem was not present with sectioned otoliths. It was concluded that sectioned otoliths were a more suitable structure from which to estimate age of L. argus than were whole otoliths, dorsal spines and/or scales.     

Movements of diadromous fish in large unregulated tropical rivers inferred from geochemical tracers

Patterns of migration and habitat use in diadromous fishes can be highly variable among individuals. Most investigations into diadromous movement patterns have been restricted to populations in regulated rivers, and little information exists for those in unregulated catchments. We quantified movements of migratory barramundi Lates calcarifer (Bloch) in two large unregulated rivers in northern Australia using both elemental (Sr/Ba) and isotope ((87)Sr/(86)Sr) ratios in aragonitic ear stones, or otoliths. Chemical life history profiles indicated significant individual variation in habitat use, particularly among chemically distinct freshwater habitats within a catchment. A global zoning algorithm was used to quantify distinct changes in chemical signatures across profiles. This algorithm identified between 2 and 6 distinct chemical habitats in individual profiles, indicating variable movement among habitats. Profiles of (87)Sr/(86)Sr ratios were notably distinct among individuals, with highly radiogenic values recorded in some otoliths. This variation suggested that fish made full use of habitats across the entire catchment basin. Our results show that unrestricted movement among freshwater habitats is an important component of diadromous life histories for populations in unregulated systems.     

Reconstructing migratory patterns of fish based on environmental influences on otolith chemistry

The analysis of elements in calcifiedstructures of fish (e.g., otoliths) todiscriminate among fish stocks and determineconnectivity between populations is becomingwidespread in fisheries research. Recently, theconcentrations of elements in otoliths arebeing analysed on finer scales that allow thedetermination of a continuous record of otolithchemistry over a fish's entire life history.These elemental concentrations can potentiallybe used to reconstruct migration patterns,based upon the influence that water chemistry,temperature, and salinity have on otolithchemistry. In doing so, assumptions are madeabout how environmental and biological factorsinfluence the concentration of elements in fishotoliths. However, there have been fewexperiments that have tested crucialassumptions regarding what influences elementaluptake and incorporation into fish otoliths.Specifically, knowledge regarding interactionsamong environmental variables, such as theambient concentration of elements in water,temperature, and salinity, and how they mayaffect otolith chemistry, is limited.Similarly, our understanding of the rate atwhich elements are incorporated into otolithsand the implications this may have forinterpretations is lacking. This reviewdiscusses methods of determining movement offish, the development of otolith research, andsome physiological aspects of otoliths (e.g.,pathways of elemental uptake). The types ofanalysis techniques that will lead to reliableand accurate migratory reconstructions areoutlined. The effects that have on otolith chemistry arereviewed with the specific aim of highlightingareas lacking environmentalvariables in experimental data. Theinfluences of the rate of elementalincorporation and ontogeny on otolith chemistryare also addressed. Finally, future researchdirections are suggested that will fill thegaps in our current knowledge of otolithchemistry. Hypotheses that need to be tested inorder to reconstruct the migratory histories offish are outlined, in a bid to clarify thedirection that research should take beforecomplex reconstructions are attempted.     

Morphometry and composition of red drum otoliths: Changes associated with temperature,somatic growth rate,and age

• 1.1. Size and composition of sagittal otoliths from red drum, Sciaenops ocellatus (Sciaenidae), reared at various constant temperatures were compared with otoliths from wild-caught fish.
• 2.2. Uncoupling of otolith growth and somatic growth in laboratory-reared fish was evident in otolith length, area, volume, weight, density, and organic fraction.
• 3.3. Fish grown at low temperatures had significantly smaller and less dense otoliths having a greater organic content than fish of the same size grown at higher temperatures.
• 4.4. Changes in inorganic elements were poorly related to temperature in laboratory-reared fish.
• 5.5. The effect of temperature on otolith elemental composition was small relative to the effects of age and its associated physiological changes.

     

Patterns of increment width and strontium: calcium ratios in otoliths of juvenile rock blackfish, Girella elevata (M.)

Otoliths of juvenile Girella elevata (M.) were examined to obtain information about the environmental conditions experienced during early life. Patterns of increment deposition and elemental ratios in otoliths were compared in wild fish. A tetracycline experiment indicated that increments were deposited daily in juveniles. Although different patterns in the spacing of increments were found among juveniles collected at different locations and times, the widest increments were always found in the first 40 increments. Strontium: calcium (Sr: Ca) ratios increased with age in the otoliths of most wild G. elevata .
The patterns of increment width and Sr: Ca ratios were not related and, therefore, were probably not under the same relative control by environmental or physiological factors. Although the number of increments can be used to age juvenile G. elevata , the utility of increment widths and Sr: Ca ratios as environmental predictors in this species is questionable without experimental validation.