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.     

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.     

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.     

Otolith geochemistry does not reflect dispersal history of clownfish larvae

Natural geochemical signatures in calcified structures are commonly employed to retrospectively estimate dispersal pathways of larval fish and invertebrates. However, the accuracy of the approach is generally untested due to the absence of individuals with known dispersal histories. We used genetic parentage analysis (genotyping) to divide 110 new recruits of the orange clownfish, Amphiprion percula, from Kimbe Island, Papua New Guinea, into two groups: “self-recruiters” spawned by parents on Kimbe Island and “immigrants” that had dispersed from distant reefs (>10 km away). Analysis of daily increments in sagittal otoliths found no significant difference in PLDs or otolith growth rates between self-recruiting and immigrant larvae. We also quantified otolith Sr/Ca and Ba/Ca ratios during the larval phase using laser ablation inductively coupled plasma mass spectrometry. Again, we found no significant differences in larval profiles of either element between self-recruits and immigrants. Our results highlight the need for caution when interpreting otolith dispersal histories based on natural geochemical tags in the absence of water chemistry data or known-origin larvae with which to test the discriminatory ability of natural tags.     

Expression of warm temperature acclimation-related protein 65-kDa (Wap65) mRNA, and physiological changes with increasing water temperature in black porgy, Acanthopagrus schlegeli

We isolated the warm temperature acclimation-related protein 65-kDa (Wap65) cDNA from the liver of black porgy and investigated the expression by increasing water temperature in black porgy, Acanthopagrus schlegeli. Black porgy Wap65 full-length cDNA consists of 1,338 nucleotides, including an open reading frame, predicted to encode a protein of 425 amino acids and showed high homology to pufferfish (79%), Medaka (73%), carp (70%), and goldfish (68%) Wap65. Increase in water temperature (20 degrees C --> 30 degrees C; 1 degrees C/day) induced the rise of Wap65 mRNA expression in liver of black porgy. Also, the levels of cortisol and glucose in plasma were significantly higher at 30 degrees C than at 20 degrees C. To determine the high water temperature stressor specificity of the induction of Wap65, black porgy were transferred from seawater (SW) to freshwater (FW) for 24 hr. Wap65 expression was not detected when the fish were transferred from SW to FW (in fish transferred from SW to FW), although the levels of cortisol and glucose in plasma were increased. These results suggest that increase in Wap65 gene is related to high water temperature stress and play important roles in high water temperature environment of black porgy.     

Limited gene flow in Uca minax (LeConte 1855) along a tidally influenced river system

For crab larvae, swimming behaviors coupled with the movement of tides suggests that larvae can normally move upstream within estuaries by avoiding ebb tides and actively swimming during flood tides (i.e., flood-tide transport [FTT]). Recently, a 1-D transport model incorporating larval behavior predicted that opposing forces of river discharge and tidal amplitude in the Pee Dee River/Winyah Bay system of South Carolina, USA, could limit dispersal within a single estuary for downstream transport as well as become a dispersal barrier to recruitment of late stage larvae to the freshwater adult habitats of Uca minax (LeConte 1855). We sequenced 394-bp of the mitochondrial cytochrome apoenzyme b for 226 adult U. minax, from four locales along a 49-km stretch of the Pee Dee River/Winyah Bay estuary, above and below the boundary of salt intrusion. Results of an analysis of molecular variance (AMOVA) and an exact test of population differentiation showed a small, but statistically significant (α=0.05) population subdivision among adults of the 4 subpopulations, as well as all subpopulations being significantly differentiated (α=0.05). This pattern fitted with model predictions, which implies that larval transport within the tidally influenced river system is limited.     

Ring formation on otoliths and scales of Pagrus pagrus: a comparative study

One annulus formed on the scales of farmed red porgy Pagrus pagrus each year during March and one opaque zone on the otoliths during June. Scales proved to be more sensitive than otoliths in recording the life history of the red porgy. A false annulus (ring of capture) was observed on the scales of the 0+ age group, but not on the otoliths. A gradual lengthening of the period of annulus formation was observed, from the younger to the older fish. The annulus formation in fish older than 2+ was less synchronized, due to the aquaculture conditions. Scales were more accurate than otoliths in annuli formation. A high percentage of missing or additional opaque zones were observed on otoliths of older fish. These irregularities were related to fish maturation and/or farming conditions. Farmed fish grew three times as fast as the wild fish.     

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 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.     

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.     

Effect of diet on otolith composition in Pomatomus saltatrix, an estuarine piscivore

To test the hypothesis that elemental composition of otoliths (sagittae) could be influenced by differences in natural prey type, young‐of‐the‐year bluefish Pomatomus saltatrix were captured immediately after their migration from oceanic waters into mid‐Atlantic Bight estuaries and fed either shrimp, Crangon septemspinosa and Palaemonetes spp. or fish Menidia menidia under similar temperature and salinity regimes in two separate 60 day experiments. Unlimited rations of fish and shrimp prey were provided in the first experiment which led to differences in bluefish growth rate between the two prey treatments; fish prey was limited in the second experiment to ensure that growth rates of bluefish in the two prey treatments were similar. Concentrations of seven elements in bluefish otoliths were determined using solution‐based inductively coupled plasma mass spectrometry (ICPMS). There was no significant effect of diet on five of the seven elements examined (Na, Mg, K, Ca and Mn). The levels of Sr and Ba in the otoliths of shrimp‐fed bluefish, however, were significantly higher than fish‐fed bluefish in both experiments. Concentrations of Ba in shrimp‐fed bluefish otoliths were double that found in fish‐fed bluefish. The results suggest that diet can explain some of the variation in otolith chemistry previously attributed to physical and chemical properties of the water.     

Inferring Processes from Spatial Patterns: The Role of Directional and Non–Directional Forces in Shaping Fish Larvae Distribution in a Freshwater Lake System

Larval dispersal is a crucial factor for fish recruitment. For fishes with relatively small-bodied larvae, drift has the potential to play a more important role than active habitat selection in determining larval dispersal; therefore, we expect small-bodied fish larvae to be poorly associated with habitat characteristics. To test this hypothesis, we used as model yellow perch (Perca flavescens), whose larvae are among the smallest among freshwater temperate fishes. Thus, we analysed the habitat association of yellow perch larvae at multiple spatial scales in a large shallow fluvial lake by explicitly modelling directional (e.g. due to water currents) and non-directional (e.g. due to aggregation) spatial patterns. This allowed us to indirectly assess the relative roles of drift (directional process) and potential habitat choice on larval dispersal. Our results give weak support to the drift hypothesis, whereas yellow perch show a strong habitat association at unexpectedly small sizes, when compared to other systems. We found consistent non-directional patterns in larvae distributions at both broad and medium spatial scales but only few significant directional components. The environmental variables alone (e.g. vegetation) generally explained a significant and biologically relevant fraction of the variation in fish larvae distribution data. These results suggest that (i) drift plays a minor role in this shallow system, (ii) larvae display spatial patterns that only partially covary with environmental variables, and (iii) larvae are associated to specific habitats. By suggesting that habitat association potentially includes an active choice component for yellow perch larvae, our results shed new light on the ecology of freshwater fish larvae and should help in building more realistic recruitment models.     

Employing a novel hybrid of GA-ANFIS model to predict distribution of whiting fish larvae and juveniles from tropical estuaries in the context of climate change

Global warming may lead to changes in nutritional composition and fish biodiversity in estuaries due to the effect of increasing temperatures on Sillago fish occurrence. Datacollected on whiting fish Sillago from two estuaries in northern Vietnam were used to employ a Genetic algorithm (GA) based Adaptive neuro fuzzy inference system (ANFIS) model for quantifying the relationships between fish distribution and environments. The present study showed that the occurrences of Sillago larvae and juveniles depended on water temperature, salinity and turbidity. The average fish CPUE predicted by GA-ANFIS was consistent with surveyed data, with the coefficient of determination (R²) being 0.63–0.65. Data obtained by applying the Hidden Markov models (HMM) indicated a rapid development of the construction area in the study site, which leads the temperature to increase. Our results suggest that as warming and land-use changes cause estuarine waters to become warmer, climate change may lead to a decline in Sillago fish occurrence numbers and abundance. The study also showed that a temperature of 23 °C is optimal for the development and growth of Sillago fish, which is shown by the higher CPUE of the Ka Long estuary than the Ba Lat estuary. These findings demonstrate applicability and reliability of GA-ANFIS and HMM models for elucidating fish larval and juvenile distribution in tropical estuaries, which will be valuable to inform management and conservation.     

COMPOSITION,ABUNDANCE AND SEASONALITY OF LARVAL FISH IN RICHARDS BAY HARBOUR,KWAZULU-NATAL,SOUTH AFRICA

Summary

The present study examined the composition, abundance and seasonality of fish larvae occurring in Richards Bay Harbour, over an 18-month period from June 1991 to November 1992. Ichthyoplankton samples were collected every six weeks at top, mid and bottom depths in the dredged channel 100 to 200 m from the harbour entrance. A total of 7 163 fish larvae, representing 105 taxa and 53 families, was collected throughout the study period. The dominant families were the Engraulidae and Gobiidae representing 50% and 37% of the total catch, respectively. The most abundant larvae were those of the thorny anchovy, Stolephorus holodon, and an unidentified goby, Gobiid 12 (32% and 30% of the total catch, respectively). Estuarine-associated taxa dominated the total catch in terms of density (61%), with 11 out of the 28 taxa being totally dependent on estuaries at some stage in their life cycle. Turbidity and temperature were significant variables accounting for 27% of the variation in larval densities of estuarine-dependent species. The main peak in larval density occurred in January 1992 (summer) with a mean larval density of 160 larvae per 100 m³. Larvae of estuarine-associated taxa were predominantly at the flexion and postflexion developmental stages. Since larval densities of certain estuarine-associated species were significantly greater in bottom samples and on flood tides, it is suggested that selective tidal stream transport is a recruitment mechanism used by these species. The present study concluded that it is important to maintain and preserve the estuarine component of Richards Bay Harbour so that the system can function as an estuarine nursery for many of the fish species depending on it at certain stages in their life cycle.     

Validation and efficacy of transgenerational mass marking of otoliths in viviparous fish larvae

Transgenerational mass marking of viviparous fish larvae in vivo was validated by intra‐muscular injection of elemental strontium chloride (SrCl₂) in gestating females and detection of the Sr in the otoliths of developing larvae. All otoliths of brown rockfish Sebastes auriculatus larvae produced from SrCl₂‐injected females showed enriched Sr:Ca ratios near the otolith edges, and the signatures did not appear to be affected by the anterior, centre and posterior positions of larvae within the ovary. Results from the present study indicate that transgenerational marking is a highly reliable technique for marking large numbers of extremely small viviparous fish larvae.     

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.     

Influence of hydrographic features on larval fish distribution during the south‐westerly monsoon in the waters of Taiwan,western North Pacific Ocean

The spatial distribution patterns of fish larvae and the relationships with hydrographic conditions in the waters surrounding Taiwan were studied in August 2004 when the south‐westerly monsoon prevailed. A total of 6566 fish larvae were identified, belonging to 80 families, 129 genera and 230 species. Cluster analysis revealed two station groups, one associated with the South China Sea Surface Current (SCSSC) and the other with the Kuroshio Current (KC). The SCSSC group was characterized by mostly coastal and neritic species, and the KC group was predominated by oceanic species. Larger fish larvae were generally more abundant in the neritic waters west of Taiwan where the SCSSC prevails than in the waters east of Taiwan where the KC prevails. The combination of chlorophyll a, zooplankton and nitrite best explained the relationship of larval fish distribution and environmental variables, implying that the distribution of fish larvae in summer was closely linked to the food source. The changing of monsoons potentially affects the succession of water masses and the transport and assemblage of fish larvae in this study area.     

Relative larval loss among females during dispersal of Lake Sturgeon (<Emphasis Type="Italic">Acipenser fulvescens</Emphasis>)

Mortality that occurs during larval dispersal as a consequence of environmental, maternal, and genetic effects and their interactions can affect annual recruitment in fish populations. We studied larval lake sturgeon (Acipenser fulvescens) drift for two consecutive nights to examine whether larvae from different females exposed to the same environmental conditions during dispersal differed in relative levels of mortality. We estimated proportional contributions of females to larval collections and relative larval loss among females as larvae dispersed downstream between two sampling sites based on genetically determined parentage. Larval collections were composed of unequal proportions of offspring from different females that spawned at upstream and downstream locations (~0.8 km apart). Hourly dispersal patterns of larvae produced from females spawning at both locations were similar, with the largest number of larvae observed during 22:00–23:00 h. Estimated relative larval loss did not differ significantly among females as larvae were sampled at two sites approximately 0.15 and 1.5 km from the last section downstream of spawning locations. High inter- and intra-female variation in larval contributions and relative larval loss between nights may be a common feature of lake sturgeon and other migratory fish species, and likely is a source of inter-annual and intra-annual variation in fish recruitment.     

Fish larval supply to and within a lagoonal estuary: multiple sources for Barnegat Bay,New Jersey

We evaluated spatial variation in fish larval supply to a temperate, lagoon type estuary (Barnegat Bay, New Jersey) by determining species composition, size, and stage into inlets (n = 2), thoroughfares between adjacent estuaries (n = 3), and within the estuary (n = 4) in seasonal, synoptic sampling on night time flood tides during 2010–2014. Larval supply, as sampled with identical plankton nets (1 m diameter, 1 mm mesh) was dominated by post-flexion stage individuals (most 5–10 but reaching 70+ mm) from species spawned in the Atlantic Ocean from a variety of sources (e.g., Sargasso Sea, outer and inner continental shelf) and in the bay. While abundance for individual species varied among locations and years, in general, the larval composition was similar across inlets, thoroughfares, and within the bay within the same seasons. Homogenization across locations was likely the result of the tidal exchanges between the ocean, the estuary, and the adjacent locations. These exchanges provide numerous, redundant sources of larvae to this estuarine nursery. The similarity in larval supply among inlets, thoroughfares, and within the estuary indicates that the longer term study location behind Little Egg Inlet is representative for this, and probably other, estuaries along the New Jersey shore.