In situ experiment of ontogenetic variability in the otolith chemistry of Stegastes partitus

Otolith chemistry can be used to assess pelagic larval fish connectivity by comparing spatially variable otolith edge chemistry (corresponding to the site of collection) to otolith core chemistry (corresponding to the site of hatching). However, because the otolith’s edge and core represent different life stages, the deposition of elements may differ, thus complicating direct comparisons of edge and core chemistry to investigate connectivity. Here we present data from a field experiment in which otoliths from embryos (3 days post-fertilization) and juveniles of Stegastes partitus were collected at the same site and time, and chemically analyzed to assess whether elemental concentrations of otoliths vary ontogenetically. Separate multivariate analyses, each investigating the spatial/temporal variability in the chemistry of either embryo otoliths or the edges of juvenile otoliths, revealed significant differences, suggesting an environmental influence to the chemical signals of otoliths. A nested multivariate analysis assessing whether otolith chemistry varied with life history stage (i.e., ontogenetic variability) indicated that elemental concentrations of embryo otoliths were significantly greater than that of juvenile otolith edges. Specifically, embryo elemental concentrations of Mn, Zn, Sn, Ba, Ce, and Pb were between 2 and 163 times greater than those of the corresponding juvenile otoliths, and thus the environment was not the primary determinant of embryo otolith chemistry. Consequently, caution is warranted when interpreting environmental patterns of otolith cores, particularly when using them as a proxy for natal signatures.     

Exploring the potential of otolith microchemistry to enhance diet analysis in pinnipeds

Increasing the scope and accuracy of information about pinniped diets obtainable from non-invasive techniques is increasingly important, particularly in cases where pinniped species are threatened or endangered. This study is the first to explore the potential for using elemental analysis of the otoliths found in scat to enhance the information available for diet analyses. We investigated the effects of pinniped digestion on otolith microchemistry using laser ablation inductively coupled plasma mass spectrometry (LA-ICPMS). We compared the elements contained in the edges (adult stage) and cores (larval/juvenile stage) of otoliths from Pacific herring (Clupea pallasii), Atka mackerel (Pleurogrammus monopterygius), and Walleye pollock (Theragra chalcogramma) recovered from the scat of captive Steller sea lions (Eumetopias jubatus) to elements in a sample of pristine (undigested) otoliths. We found that digestion had a significant effect on four of the six sampled combinations of species and otolith region (herring edges and cores, mackerel edges, and pollock cores), and that Rb most frequently showed significant differences in concentration after digestion. We could significantly discriminate among species of both pristine and digested otoliths using either otolith edges or cores with the elements Ba, Rb, Sr, Y, and Mg. When compared to previously identified digested otoliths, unknown samples of the three species of digested otoliths could be discriminated with 55–100% accuracy depending on species and otolith region. When compared to a library of previously identified pristine and digested otoliths, unknown samples of digested otoliths could be discriminated to species with 65–88% accuracy. When the group of unknown digested otoliths was compared to known pristine otoliths, discrimination ranged from 45 to 65%. These results indicate that elemental analysis could be used to supplement visual identification of otoliths from scat. However, further research is required to determine whether elemental analysis of digested otoliths could be useful for prey fish population studies.     

Dietary effects on multi-element composition of European eel (<Emphasis Type="Italic">Anguilla anguilla</Emphasis>) otoliths

Otolith microchemistry is widely used as a tool to track individual migration pathways of diadromous fish under the assumption that the elemental composition of fish otoliths is directly influenced by the physicochemical properties of the surrounding water. Nevertheless, several endogenous factors are reported to affect element incorporation into fish otoliths and might lead to misinterpretations of migration studies. This study experimentally examined the influence of eight different diets on the microchemical composition of European eel (Anguilla anguilla) otoliths using laser ablation inductively coupled plasma mass spectrometry (LA-ICP-MS). Seven natural prey types and one artificial diet were fed during 8 weeks in freshwater circuits. Results show for the first time that food has no significant influence on the incorporation of Na, Sr, Ba, Mg, Mn, Cu and Y into European eel otoliths. This indicates that the incorporation of elements usually chosen for migration studies is not affected by diet and that individual feeding behaviour of A. anguilla will not lead to any misinterpretation of migration pathways.     

Variation in concentrations of trace elements in otoliths and eye lenses of a temperate reef fish, Parma microlepis, as a function of depth, spatial scale, and age

 Territorial Parma microlepis (Günther) (Pomacentridae) were collected at different depths, at three sites in each of four estuaries near Sydney, Australia. Element concentrations were measured by induc- tively-coupled–plasma mass spectrometry. Significant differences in concentrations of Mn and Ba were found in the otoliths of fish sampled in different depth strata, with concentrations generally greatest in fish found in water <4 m deep. Depth-related differences varied among estuaries (e.g. 0 to 1.2 μg Ba g⁻¹ otolith). In most estuaries there was a negative linear relationship between concentrations of Mn and Ba in otoliths and actual depth. Great variation was found within an estuary among sites separated by 0.5 to 3 km. In the eye lenses, concentrations of Rb differed according to depth of capture of fish. The age of fish (1+ to 34 yr) had no influence on the concentrations of elements in otoliths or lenses. Multivariate comparisons of elemental composition (= fingerprints) detected significant differences among depth strata. Fish collected from shallow water had the clearest multivariate classification according to depth. There was a close match between our shallow strata and the average depths reached by low-salinity/high-temperature estuarine plumes. The element composition of whole otoliths and lenses represents average concentrations experienced by the fish. The temporal resolution of differences in ambient conditions is likely to be coarse in the fish (i.e. months to years). Received: 9 April 1999 / Accepted: 29 February 2000     

Nodularin concentrations in Baltic Sea zooplankton and fish during a cyanobacterial bloom

Toxic cyanobacterial blooms, dominated by Nodularia spumigena, are a recurrent phenomenon in the Baltic Sea during late summer. Nodularin, a potent hepatotoxin, has been previously observed to accumulate on different trophic levels, in zooplankton, mysid shrimps, fish as well as benthic organisms, even in waterfowl. While the largest concentrations of nodularin have been measured from the benthic organisms and the food web originating from them, the concentrations in the pelagic organisms are not negligible. The observations on concentrations in zooplankton and planktivorous fish are sporadic, however. A field study in the Gulf of Finland, northern Baltic Sea, was conducted during cyanobacterial bloom season where zooplankton (copepod Eurytemora affinis, cladoceran Pleopsis polyphemoides) and fish (herring, sprat, three-spined stickleback) samples for toxin analyses were collected from the same sampling areas, concurrently with phytoplankton community samples. N. spumigena was most abundant in the eastern Gulf of Finland. In this same sampling area, cladoceran P. polyphemoides contained more nodularin than in the other areas, suggesting that this species has a low capacity to avoid cyanobacterial exposure when the abundance of cyanobacterial filaments is high. In copepod E. affinis nodularin concentrations were high in all of the sampling areas, irrespective of the N. spumigena cell numbers. Furthermore, nodularin concentrations in herring samples were highest in the eastern Gulf of Finland. Three-spined stickleback contained the highest concentrations of nodularin of all the three fish species included in this study, probably because it prefers upper water layers where also the risk of nodularin accumulation in zooplankton is the highest. No linear relationship was found between N. spumigena abundance and nodularin concentration in zooplankton and fish, but in the eastern area where the most dense surface-floating bloom was observed, the nodularin concentrations in zooplankton were high. The maximum concentrations in zooplankton and fish samples in this study were higher than measured before, suggesting that the temporal variation of nodularin concentrations in pelagic communities can be large, and vary from negligible to potentially harmful.     

Growth of sea bass,Dicentrarchus labrax,larval and juvenile stages and their otoliths under quasi-steady temperature conditions

The relationship between somatic growth and growth of otoliths of sea bass larvae, postlarvae and juveniles under relatively steady temperature conditions was studied. Larvae were incubated at the constant ambient temperature of 13.5°C, whereas postlarvae and juveniles were reared at a comparatively steady temperature ranging from 18.6 to 20.4°C, with a mean of 19.67°C. The patterns of both somatic and otolith growth were found to be similar. Differentiated data on larvae length and otolith diameters indicated three periods of change in their growth rates. Since temperature was kept relatively steady during the experiment, and larvae fed ad libitum, these periods could be attributed with relative certainty to intrinsic changes which occur during stage-specific periods of growth. The third period of change in both growth rates indicates a specific phase of growth during metamorphosis. The changes in growth rates, as well as the raw time series of the growth of both larval lengths and otolith diameters, may be described by higher order polynomials with a high degree of probability levels. A non-linear relationship between body length and otolith diameters was established, indicating positive allometric growth of otoliths. It was also observed that the coefficient of allometric growth changed at the time estimated for the end of metamorphosis. Thus, a non-linear relationship and changes in the coefficients of allometry should be borne in mind when back-calculating somatic growth from the growth of otoliths.     

Interacting trophic forcing and the population dynamics of herring

Small pelagic fish occupy a central position in marine ecosystems worldwide, largely by determining the energy transfer from lower trophic levels to predators at the top of the food web, including humans. Population dynamics of small pelagic fish may therefore be regulated neither strictly bottom-up nor top-down, but rather through multiple external and internal drivers. While in many studies single drivers have been identified, potential synergies of multiple factors, as well as their relative importance in regulating population dynamics of small pelagic fish, is a largely unresolved issue. Using a statistical, age-structured modeling approach, we demonstrate the relative importance and influence of bottom-up (e.g., climate, zooplankton availability) and top-down (i.e., fishing and predation) factors on the population dynamics of Bothnian Sea herring (Clupea harengus) throughout its life cycle. Our results indicate significant bottom-up effects of zooplankton and interspecific competition from sprat (Sprattus sprattus), particularly on younger age classes of herring. Although top-down forcing through fishing and predation by grey seals (Halichoerus grypus) and Atlantic cod (Gadus morhua) also was evident, these factors were less important than resource availability and interspecific competition. Understanding key ecological processes and interactions is fundamental to ecosystem-based management practices necessary to promote sustainable exploitation of small pelagic fish.     

Interactions between cod, herring and sprat in the changing environment of the Baltic Sea: A dynamic model analysis

The interactions between cod (Gadus morhua), herring (Clupea harengus) and sprat (Sprattus sprattus) in the Central Baltic Sea were examined with a simple dynamic model, an alternative to more complicated and data-demanding multispecies and ecosystem models. The main aims of the study were to compare the effect of alternative structures on the model output and examine the control relationships in the fish assemblage under different environmental conditions. The effect of environmental conditions was modelled using a stock-recruitment equation for cod incorporating an environmental index. The model output was especially sensitive to the functional response in predation by cod on herring and sprat. The type II functional response led to a collapse of the clupeid stocks when cod was abundant, while the type III response produced more realistic stock dynamics. According to the simulations, an abundant cod stock was able to keep the sprat stock at a low level, while the herring stock was less affected and benefited from the decreased density of sprat. Simulation of different fishing scenarios indicated that reducing fishing mortality to the level currently advised by ICES would allow the recovery of the cod stock even in unfavourable environmental conditions.     

Comparison of somatic growth and otolith increment growth in laboratory-reared larvae of Pacific saury, Cololabis saira, under different temperature conditions

The relationship between somatic growth and incremental growth of otoliths of Pacific saury, Cololabis saira (Brevoort), larvae under different temperature conditions was studied in the laboratory for three age groups (0 to 9, 10 to 20 and 20 to 30 d posthatch). Larvae were incubated from hatching to 9 d at 24, 20, and 16 °C. Further, larvae initially reared at an ambient temperature of 21.7 °C were transferred to experimental temperatures of 22, 18, and 14 °C on Day 10 and reared to Day 20 and similarly from Day 20 and reared to Day 30 posthatch. Growth trajectories of larvae sampled at the end of the three experiments were back-calculated using the biological intercept method and compared to the measured values 0 and 5 d after the start of each experiment. Back-calculated knob length at the different temperatures indicated no significant difference to the measured knob lengths except for the cases at 20 °C from hatching to 9-d-old larvae and at 14 °C from 20- to 30-d-old larvae. The close correlation between somatic and otolith growth shown in this study indicated the feasibility of estimating the growth history of Pacific saury larvae using otolith readings. Received: 14 April 1999 / Accepted: 27 October 1999     

Confirmation of longevity in Sebastes diploproa (Pisces: Scorpaenidae) from 210Pb/226Ra measurements in otoliths

Fish ages are often estimated by assuming an annual frequency of the band-like, growth-zones recorded in the largest of their otoliths, the sagittae. The total number of growth-zones are normally determined either by counting external growth-zones (whole otolith technique) or by examining otolith cross-sections (otolith section technique). The two techniques do not always yield the same age, however, particularly in older specimens of certain fishes. To resolve this problem, otoliths of the splitnose rockfish Sebastes diploproa were examined morphologically and were assayed for their natural radionuclide concentrations. Four age groups of otoliths were identified based on growth-zone counting; in the first three, whole otolith and otolith section age estimates agreed, while in the fourth, the otolith section age substantially exceeded the whole otolith age. Radiometric analysis demonstrated that all otoliths were deficient in ²¹⁰Pb activity relative to ²²⁶Ra activity with the deficiency decreasing with increasing number of growth-zones. The magnitude of the ²¹⁰Pb/²²⁶Ra radioactive disequilibrium in each otolith group, when compared to the number of growth-zones and the otolith weight histories derived with the two techniques, identifies the growth-zones revealed by otolith sections as annual features. Thus when otolith section age exceeds whole otolith age (usually occurring after 20 to 25 yr of age for this species), the otolith section technique is the correct method of age determination. Estimates of longevity in the genus Sebastes near 80 yr are therefore confirmed.     

Comparison of larval duration and pre- and post-settlement growth in two species of damselfish,Chromis atripectoralis andPomacentrus coelestis (Pisces: Pomacentridae), from the Great Barrier Reef

Light traps were used to capture larval fishes, immediately before settlement, at two localities 500 km apart on the Great Barrier Reef (GBR) in December, 1987. Samples from Lizard Island, in the northern GBR, and Davies Reef, in the central GBR, were dominated by two species of damselfish:Chromis atripectralis andPomacentrus coelestis. Analysis of otoliths revealed significant differences in both size and age at settlement between the two localities forP. coelestis, but not forC. atripectoralis. Growth rates determined for pre- and post-settlementP. coelestis suggested a sigmoidal growth trajectory through the larval life, with growth slowing as fishes approached the time of settlement. Post-settlement growth rates were faster than growth prior to settlement in both species. Growth in both species was, however, similar between localities. The relationship between fish size and otolith size was complex, varying both between pre- and post-settlement fishes, and among localities. This emphasizes the need to validate the relationship between fish size and otolith size before otoliths may be used to back-calculate individual growth trajectories.Contribution No. 500 from the Australian Institute of Marine Science     

Effects of specimen handling and otolith preparation on concentration of elements in fish otoliths

The use of otolith chemistry as a tool for analysing the biology of fishes assumes that the procedures used to collect and prepare otoliths for analysis do not alter their composition. With otoliths of Nemadactylus macropterus, Hoplostethus atlanticus, and Rhombosolea tapirina, we show that this assumption is not valid for the elements that can be detected using electron-probe microanalysers (those present at concentrations greater than ≃100 parts per million): all six elements routinely measured using these techniques were affected by at least one post-mortem procedure tested. Measured concentrations of calcium and strontium were relatively insensitive to most procedures tested, whereas concentrations of sodium, potassium, sulphur and chlorine were affected substantially by many commonly used procedures. The ease with which otolith composition could be modified post-mortem suggests that apparent geographic, habitat-specific or ontogenetic differences in otolith composition should be interpreted with extreme caution because of easily induced artefacts and the problem of pseudo-replication associated with the ways otoliths are sampled and prepared for analysis. Received: 23 September 1996 / Accepted: 11 April 1998     

Effects of delayed feeding and temperature on the age of irreversible starvation and on the rates of growth and mortality of Pacific herring larvae

The time periods from exhausion of the yolk to the age of irreversible starvation for Pacific herring Clupea harengus pallasi larvae were 8.5, 7.0 and 6.0 d at 6°, 8° and 10°C, respectively. These periods are within the range perviously measured for Atlantic herring larvae and other temperature zone fish species; they are long compared to the periods for tropical species. The variation in the length of this period is due almost entirely to temperature; the natural logarithm of the time period from fertilization to irreversible starvation is highly correlated (r=0.91) with the mean rearing temperature for 25 species of pelagic marine fish larvae. The rates of growth and mortality, measured for 26 experimental populations of Pacific herring larvae reared at 6°, 8° and 10°C and ten ages of delayed first feeding, decreased and increased, respectively with increasing age of first feeding and increasing temperature. These rates, adjusted for the effects of rearing conditions, were compared with the rates for natural populations of herring larvae. Growth is generally faster in the sea than in experimental enclosures. Two of the eleven estimates of natural mortality rate were high enough to indicate possible catastrophic mass starvation. This is consistent with Hjort's critical period concept of year class formation and it suggests that mass starvation occurs in 18 to 36% of the natural populations of first feeding herring larvae.     

Elemental composition of otolith growth marks in three geographically separated populations of European hake (Merluccius merluccius)

The composition of the opaque and translucent bands in the otoliths of European hake (Merluccius merluccius) from three different populations (Celtic Sea, Galicia shelf and Cretan Sea) was analysed with wavelength dispersive spectrometry (WDS) and differences in Ca, Sr and Na were compared. The translucent bands (annual marks and checks altogether) were significantly richer in Sr and Ca and significantly poorer in Na than opaque bands. Results support the idea of a more compact arrangement of crystals in translucent bands compared to opaque. Analyses did not help to discriminate between putative annual marks and checks. Clear differences in the range of Sr/Ca variation across the otolith as well as in the composition of the otolith core were observed among the three populations. As a result it was possible to discriminate the natal origin of each fish using the composition of the otolith core in Ca, Sr and Na. Nevertheless, the overall pattern across the otolith was similar among the three populations suggesting the existence of ontogenetic trends. Variation of Na/Ca ratios was largely uninformative. Based on these results, a conceptual model for hake is proposed which integrates the different sources of otolith Sr/Ca variation: ambient Sr/Ca, endogenous cues (ontogeny) and exogenous cues (environment).     

Food and feeding habits of the herring Clupea harengus and the sprat C. sprattus in inshore waters of the west coast of Scotland

The food and feeding habits of the herring Clupea harengus L. and the sprat C. sprattus (L.) as 0-group and older fish in inshore waters of the west coast of Scotland, were studied from April, 1970 to March, 1972. The two species occurred together. The diets of 0-group herring and sprats are almost identical, both qualitatively and quantitatively. Their daily feeding rhythms also coincide. It is probable that competition between these two species in their first year of life could occur if the food supply is limiting. Changes in diet between 0-group and older fish are more pronounced in herring than in sprats. The diet itself is mainly crustacean in both clupeids, with copepods contributing the major share.     

Otolith record of age, growth, and ontogeny in larval and pelagic juvenile Stephanolepishispidus (Pisces: Monacanthidae)

Juveniles of the planehead filefish Stephano-lepishispidus (Pisces: Monacanthidae) (Linnaeus, 1766) are a major component of the Sargassum spp. community, yet little is known of their ecology. In this study, the otolith record of age, growth, and ontogeny in S. hispidus was examined. Juveniles caught off Beaufort Inlet, North Carolina (USA) on 30 June 1996 were marked with alizarin complexone and reared in a flow-through, outdoor tank for up to 19 days. Examination of marked otoliths at several time intervals showed that increment formation was not significantly different than one increment per day, and thus, increment number was used to estimate age. Depth-distribution, morphology, and meristics of larvae and juveniles collected (1990–1992) between Cape Romain, South Carolina, and Cape Hatteras, North Carolina, were examined to identify the timing of the larval to juvenile transition. All indicators suggested the transition occurred between 17 and 20 days. Mean otolith increment widths exhibited a marked change at about 20 days, coinciding with the timing of the larval to juvenile transition and a change in the depth distribution from bottom to surface waters. Increment width of individual juveniles, however, did not exhibit the same pattern; only 40% conformed to the pattern identified for all fish. Thus, the record of the larval to juvenile transition is clear at the population level, but unresolved at the individual level. Received: 1 November 1999 / Accepted: 18 December 2000     

Functional responses of juvenile herring and sprat in relation to different prey types

The relationship between particulate-feeding rates and prey concentrations (functional response) of juvenile herring and sprat (5–9 cm total length) was investigated in controlled feeding experiments monitored by an underwater camera system. A special tank system was developed allowing the regulation and quantification of low prey concentrations (1–160 L^?1). Non-evasive Artemia nauplii was used as prey to estimate the maximum biting rate of both predators. In contrast, Acartia tonsa with a high escape ability was used as a realistic prey type. Herring and sprat showed a type II functional response for both prey types. Nonlinear mixed effects model revealed no significant difference between the functional responses of both predators, except that herring showed significantly higher biting rates than sprat at A. tonsa concentrations below ~40 L^?1. For both predators feeding rates were significantly higher with Artemia nauplii than with A. tonsa. Video analysis indicated that sprat, unlike herring, is an obligate particulate-feeder.     

Age determination of orange roughy,Hoplostethus atlanticus (Pisces: Trachichthyidae) using210Pb:226Ra disequilibria

Natural levels of²¹⁰Pb:²²⁶Ra in otoliths of orange roughy,Hoplostethus atlanticus, from south-east Australian waters, were measured to determine fish ages radiometrically. Up to maturity, radiometric age estimates were consistent with a single constant otolith growth rate. Radiometric ages for juveniles were comparable with, but greater than, those obtained in a recent, validated New Zealand study which employed counts of annuli on the surface of otoliths. Beyond maturity, radiometric ages were obtained by modelling with an otolith growth rate set at 45% of the juvenile rate. Radiometric ageing confirms that orange roughy is very slow-growing, with an age at maturity (32 cm standard length, SL) of ~ 32 yr, and is very long-lived, with fish 38 to 40 cm being 77 to 149 yr old. These results have important implications for the management of the fishery.     

Chemistry of tuna otoliths: assessment of base composition and postmortem handling effects

Protocols used to collect and prepare otoliths for chemical analysis may result in either contamination or loss of elements, thus biasing population studies in unknown ways. We evaluated precision and bias associated with collection and cleaning procedures for three Atlantic tuna species: Atlantic bluefin tuna (Thunnus thynnus), yellowfin tuna (T. albacares), and blackfin tuna (T. atlanticus). Elemental concentrations were measured using solution-based inductively coupled plasma mass spectrometry and atomic absorption spectrophotometry. Seven elements were present above detection limits in all samples (Na, Mg, K, Ca, Mn, Sr, Ba). Mean concentrations of all seven elements were statistically indistinguishable in fresh pairs of otoliths of T. thynnus (mean error 5%, range 2-8%) and T. albacares (mean error 5%, range 3-7%); no indication of a left versus right otolith effect was observed. Otolith elemental concentrations were size dependent and significant inverse relationships were observed for Mg, Na, and K. Deliberate contamination of previously cleaned samples using a 10-ppm solution of a mixture of elements demonstrated that otoliths easily acquire surface contamination. Recleaning contaminated otoliths restored elemental concentrations to levels similar to control otoliths. Precision of paired comparisons between cleaned otoliths and those exposed to contamination and recleaned was high (mean error 6%). The effects of storage at two temperatures (7 days at -20°C, 3 days at 1°C) were investigated. For K, Ca, Sr, Mn, and Ba, variation between control (removed immediately) and treatment otoliths (in situ freezing or chilling) was similar to variation observed within fresh otolith pairs (mean error: fresh vs frozen 5%, fresh vs iced 5%). Statistically significant but small (<10%) postmortem storage effects were observed for Na and Mg. Estimates of error indexed to natural ranges in otolith chemistry of T. thynnus and T. albacares from different geographic regions in the Atlantic and Pacific showed that error values of several elements (Mg, Mn, Ba, Na, K) accounted for a small proportion of the natural range, suggesting levels of precision achieved in this study are suitable for the purpose of stock delineation.