鱼类耳石元素指纹研究进展

耳石是沉积在真骨鱼类内耳中的一种矿物结构,其主要成分是碳酸盐.在鱼类的生活过程中,水环境中的化学元素经过代谢稳定地沉积在耳石上,由于生活水域化学环境的差异,鱼类耳石的元素组成或含量存在一定的差异,而使耳石上的元素指纹成为一种天然标志,记录了鱼类所经历的生态环境及其变化.耳石元素指纹研究主要包括同位素成分分析、锶钙比率研究(Sr/Ca)以及一些痕量元素分析等,这些可以用于鱼类种群识别、生活史研究、生活环境重建,以及高龄鱼的年龄鉴定,在鱼类生态学和渔业资源学研究中具有广泛的应用前景.参57     

Scales of variation in otolith elemental chemistry of juvenile staghorn sculpin (<Emphasis Type="Italic">Leptocottus armatus</Emphasis>) in three Pacific Northwest estuaries

Although distinct otolith elemental signatures are often observed in fish collected from different estuaries, significant differences are also observed among sites within estuaries. Variation at these smaller spatial scales is not well quantified and has the potential to lead to inappropriate interpretations of otolith elemental data. To quantify variation at multiple scales, the otolith elemental composition (Mg:Ca, Mn:Ca, Sr:Ca, Ba:Ca, and Pb:Ca) of juvenile staghorn sculpin (Leptocottus armatus Girard, 1854) collected from five sites within three estuaries, the Columbia River (two sites) and Coos Bay (one site), Oregon, and Humboldt Bay, California (two sites), was examined. Using laser ablation-inductively coupled plasma mass spectrometry, each otolith was sampled at three zones: (1) within the primordium, which represents the egg and early larval periods; (2) at the outer edge, which represents the juvenile period just prior to collection; and (3) midway between the primordial and edge samples, which represents the late larval and early juvenile period. There were significant differences in otolith metal-to-calcium ratios at all scales examined. Using multi-element otolith signatures, fish were classified to estuary and site within estuary with relatively high levels of accuracy (av = 70–90%). The largest differences in metal-to-calcium ratios were observed between sites within estuaries (<5 km apart) and the smallest differences were observed among otolith zones. Variation in otolith chemistry may be used to provide information on probable habitat use by estuarine fish but studies must be carefully designed. Electronic supplementary material  Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Elevated manganese concentrations at the cores of clupeid otoliths: possible environmental,physiological, or structural origins

Laser ablation inductively coupled plasma mass spectrometry (LA-ICP-MS) was used to investigate the distribution of manganese (Mn) in Atlantic herring (Clupea harengus L.) otoliths. An unusual pattern of Mn concentration, which has not previously been reported in fish otoliths, was observed. Time-resolved plots (time roughly equivalent to ablation depth) of the Mn signal recorded in the ICP-MS detector during ablation of otoliths contained a distinct peak, suggesting that elevated Mn concentrations were specific to certain regions. Ablation inside and outside of the otolith core revealed that the region of high Mn concentration coincided with the ablation of the core. It was initially suggested that Mn from the gravel substrate on which herring eggs are incubated is incorporated into the otolith core, producing elevated Mn concentrations in this region. It was hypothesised that otoliths from fish whose eggs are not incubated in gravel will not contain high Mn concentrations at the core. To test this, otoliths of sprat (Sprattus sprattus), which have a pelagic embryological stage, and from reared herring, which were incubated on glass plates, were analysed using LA-ICP-MS. Peaks in the Mn signal were observed during the ablation of cores from some sprat and reared herring otoliths, indicating that incubation on a gravel substrate is not necessary for the accumulation of Mn at the otolith core and that otoliths from both pelagic and demersally spawning fish species can contain elevated Mn concentrations. The results suggest that Mn concentrations at the otolith core do not reflect the external environment of the developing embryo. Two other hypotheses are considered. A dramatic increase in Mn concentrations in the egg, associated with embryological development, may produce elevated Mn levels at the otolith core. Alternatively, the high levels of Mn observed at the otolith core may reflect spatial variation in the crystal structure of the calcium carbonate portion of the otolith. The possible influence of physiological or structural factors on the chemical composition of the otolith core may confound the environmental signal in this region of the otolith. This has important implications for the interpretation of otolith chemistry data.Communicated by J.P. Thorpe, Port Erin     

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.     

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.     

Model based grouping of species across environmental gradients

We present a novel approach to the statistical analysis and prediction of multispecies data. The approach allows the simultaneous grouping and quantification of multiple species’ responses to environmental gradients. The underlying statistical model is a finite mixture model, where mixing is performed over the individual species’ responses to environmental gradients. Species with similar responses are grouped with minimal information loss. We term these groups species archetypes. Each species archetype has an associated GLM that can be used to predict distributions with appropriate measures of uncertainty. Initially, we illustrate the concept and method using artificial data and then with application to real data comprising 200 species from the Great Barrier Reef (GBR) lagoon on 13 oceanographic and geological gradients from 12°S to 24°S. The 200 species from the GBR are well represented by 15 species archetypes. The model is interpreted through maps of the probability of presence for a fine scale set of locations throughout the study area. Maps of uncertainty are also produced to provide statistical context. The presence of each species archetype was strongly influenced by oceanographic gradients, principally temperature, oxygen and salinity. The number of species in each group ranged from 4 to 34. The method has potential application to the analysis of multispecies distribution patterns and for multispecies management.     

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).     

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.     

Episodes of starvation are recorded in the otoliths of juvenile Ambassis vachelli (Chandidae), a tropical estuarine fish

Juvenile Ambassis vachelli, 40 to 50 d old, were used to test the effects of starvation and recovery-feeding on the widths of otolith increments. Three laboratory treatments (fed continuously; fed for 9d then starved for 15 d; starved for 9 d then fed for 15 d), were compared. Fish were also collected throughout the experiment from a field site to assess the natural fluctuations of increment widths. Growth, starvation and recovery were reflected in the widths of otolith increments. The width of increments deposited during starvation were similar in all fish, regardless of prior feeding history, despite the lower mortality recorded in starved fish with a history of intense feeding. Increment-width data were confounded by an ontogenetic decrease in increment widths with age. The results indicate that starvation and recovery are recorded in the increment widths of otoliths and are distinguishable from the effects of ontogenetic development. However, ontogenetic changes in the width of otolith increments confounded the interpretation of otolith-increment series. Therefore, the ontogenetic pattern of otolith increments must be known for each species before valid interpretation of otolith microstructure is possible.     

A Poisson–Gamma model for analysis of ecological non-negative continuous data

The statistical analysis of continuous data that is non-negative is a common task in quantitative ecology. An example, and our motivation, is the weight of a given fish species in a fish trawl. The analysis task is complicated by the occurrence of exactly zero observations. It makes many statistical methods for continuous data inappropriate. In this paper we propose a model that extends a Tweedie generalised linear model. The proposed model exploits the fact that a Tweedie distribution is equivalent to the distribution obtained by summing a Poisson number of gamma random variables. In the proposed model, both the number of gamma variates, and their average size, are modelled separately. The model has a composite link and has a flexible mean-variance relationship that can vary with covariates. We illustrate the model, and compare it to other models, using data from a fish trawl survey in south-east Australia.     

Otolith size-at-hatch reveals embryonic oxygen consumption in the zebrafish, <Emphasis Type="Italic">Danio rerio</Emphasis>

Otoliths have frequently been used to reconstruct growth histories in larval, juvenile and adult fish. However, there is growing evidence that otolith growth is directly determined by metabolic intensity and, consequently, only indirectly related to somatic growth. By performing measurements of oxygen consumption rate and other early life-history traits on individual eggs of zebrafish (Danio rerio), we found that oxygen consumption explained residual variance in otolith size that is not accounted for by egg size. Total oxygen consumption during the embryonic stage explained 34% of the variance in sagitta size at hatch, whereas larval size at hatch (as a proxy for growth during the embryonic period) was not significantly correlated with sagitta size. This strongly suggests that otolith growth is directly related to metabolic rate, and yields a mechanism that may explain recent observations of a link between otolith size at hatch and viability in larval fish.     

Geographical differences in early growth of walleye pollock Theragra chalcogramma,estimated by back-calculation of otolith daily growth increments

The relationship between otolith length and body length (total length) was examined in walleye pollock Theragra chalcogramma (Pallas), collected from Pacific Ocean waters adjacent to Hokkaido, Japan, and in larvae reared from fertilized eggs. A linear relationship was found between log-transformed body length and otolith length data with two inflection points at ca. 11 mm and 100 mm total length. This relationship was found to be applicable also to samples from the Japan Sea, Bering Sea and Okhotsk Sea. The early growth pattern estimated by back-calculation of otolith increments of 1-yr fish from Funka (Uchiura) Bay (Hokkaido) accorded with that obtained from size-at-age data of 0 yr fish collected in the preceding year. Differences were found in comparisons of the back-calculated early growth pattern between samples from waters adjacent to Hokkaido, the Pacific Ocean, the Japan Sea and the Okhotsk Sea. The growth curve of the Okhotsk samples was markedly different from others, showing rapid initial growth up to about 100 d after hatching and attaining a small body size in the first year.     

Planktonic larval duration of one hundred species of Pacific and Atlantic damselfishes (Pomacentridae)

The plankton larval duration for 100 species of Pacific and Atlantic damselfishes was estimated from daily growth increments on the otolith of juvenile fish collected at various localities between July 1987 and September 1988. For newly-settled fishes, larval duration was determined by counting the entire number of increments present on the otolith, while for older juveniles estimates were made by counting the number of increments between the center of the otolith and a mark corresponding to settlement. We document the development of otolith formation during the period when eggs are incubated on the reef and show that daily increments are only accreted after larvae hatch and enter the planktonic phase. The planktonic larval duration for damselfish is shorter and less variable, both between and within species, compared to other groups of reef fishes such as wrasses and surgeonfishes. Larval duration ranged from 12 to 39 d. Average duration between species ranged from 13.1 to 35.2 d. The time spent in the plankton was not significantly correlated with geographic distribution when evaluated among species, however, genera with confined regional distribution have a shorter mean larval life than do widely distributed genera. Size at settlement was positively correlated with time spent in the plankton among species, but a significant correlation between these variables was only evident within one of ten species. The low variance in planktonic larval duration within species indicates that most damselfish are unable to delay metamorphosis following competency. This inability to postpone settlement limits the potential for dispersal, especially when dispersal time between suitable habitats is greater than about 30 d.     

Population structure and life history of orange roughy (Hoplostethus atlanticus) in the SW Pacific: inferences from otolith chemistry

We examined site differences in the elemental composition of the primordium and ontogenetic variability of Sr in otoliths of fish from Australia and New Zealand and, as an out-group, the North Atlantic. Differences among sites in primordium composition are slight, but significant for all five elements assayed (Sr, Pb, Cu, Zn and Hg), but principally reflect differences between the North Atlantic and SW Pacific specimens, do not replicate for independent samples in the SW Pacific and constitute a poor “natural tag” in roughy, with <25% of fish successfully assigned to source location. However, mean Sr weight-fractions at the primordium showed similar latitudinal variation across sites in Australia, New Zealand and the Tasman Sea, indicating both spatially structured populations and a common structuring process across the region. Comparisons of ontogenetic variability of Sr in otoliths from juveniles and young adults within and between sites in the SW Pacific strongly support the hypothesis that variability in this element is site-specific and environmentally sensitive, although the environmental factors involved are not obvious. The otolith analysis confirms previous suggestions that juvenile and adult Hoplostethus atlanticus are relatively sedentary, but also indicates that the population sub-structuring by age within sites is more complex and there are likely to be more spawning areas in Australian waters than previously thought. More broadly, although single point analysis of otolith composition at the primordium resolves a population structure in roughy, alone it is not precise enough to test hypotheses about the processes causing this structure. Ontogenetic variability in Sr provides better resolution of spatial structure, even in a relatively homogenous marine environment like the deep ocean, and also provides insight into behavioural and ecological factors. Ontogenetic analyses of Sr in otoliths are expensive to obtain, require more effort in specimen preparation than single point analyses, and are difficult to compare statistically, but the increased information they yield warrants their broader consideration in marine species.     

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     

Female sticklebacks Gasterosteus aculeatus use self-reference to optimize MHC allele number during mate selection

Theoretical and empirical studies suggest that an optimal resistance to pathogens and parasites requires an optimal number of MHC alleles per individual. Here we argue that three-spined sticklebacks (Gasterosteus aculeatus) achieve this goal by applying a strategy that involves a self-referential process. According to this model, females complement their own set of alleles with a more or less diverse set of male alleles such that the combined diversity reaches an optimum. In previous experiments, we have identified allele counting as a major mate-choice strategy in large populations. The self-referential allele-counting hypothesis predicts that MHC-based mate choice favors dissimilar MHC alleles in small populations facing the risk of inbreeding. Therefore, we conducted an experiment that simulated a small effective population size with low MHC class-II diversity. Our experiments are based on the analysis of MHC class-IIB alleles that explain a major part of the overall MHC diversity in sticklebacks, as determined by mathematical modeling. The results show that females preferred males with dissimilar alleles. Our present and the previous studies (which we reanalyzed with respect to our new predictions) show that irrespective of high or low population diversity faced by female sticklebacks for their mate choice, they use information about their own and their potential mate's MHC polymorphism for optimal complementation of their own set of alleles. From combining the data of the previous and the present experiment we found that female sticklebacks try to achieve an optimum number of MHC class-IIB alleles for their offspring through mate choice. The chosen MHC diversity is close to the most frequent diversity found naturally in individual fish, which in addition have the lowest parasite burden.     

Model-based combination of spatial information for stream networks

Evolutionary improvements in Geographic Information Systems (GIS) now routinely allow the management and mapping of spatial-temporal information. In response, the development of statistical models to combine information of different types and spatial support is of vital importance to environmental science. In this paper we develop a hierarchical spatial statistical model for environmental indicators of stream and river systems in the United States Mid-Atlantic Region by combining information from separate monitoring surveys, available contextual information on hydrologic units and remote sensing information. These models are used to estimate the indicators throughout the riverine system based on information from multiple sources and aggregate scales. The analysis is based on information underlying the Landscape Atlas of the mid-Atlantic region produced by the US Environmental Monitoring and Assessment Program (EMAP). We also combine information from two overlapping separate monitoring surveys, the EMAP Stream and River Survey and the Maryland Biological Streams Survey. We present a general framework for comparative distributional analysis based on the concept of a relative spatial distribution. As an application, the spatial model is used to predict spatial distributions and relative spatial distributions for a watershed.     

A Bayesian approach to evaluating site impairment

In the United States, each state is required to list water resources that are declared to be impaired under guidelines set by the Clean Water Act. Measurements are typically collected on a number of chemical constituents and compared with a standard. If there are too many measurements exceeding the standard, then the site is declared impaired. The approach is non-statistical but similar to a Binomial test. The Binomial approach would convert the measurements to binary data then test if the proportion exceeding the standard is excessive. Both methods convert measurements to binary values hence exclude potentially important information in the data. We present a statistical approach using a Bayesian model that uses the raw data instead of the binary transformed data. The population distribution of a family of location-scale parameter models is studied under the model. Posterior distributions from the Bayesian analysis are used in the decision-making process and error probabilities for the Bayesian and the Binomial approaches are compared for a normal population.