Stock discrimination and connectivity assessment of yellowfin seabream (Acanthopagrus latus) in northern South China Sea using otolith elemental fingerprints

Connectivity between fish stocks is fundamental to the understanding of population dynamics and the implementation of sustainable fisheries management. Otolith microchemistry is a promising tool as it can provide information on the continuous growth of otoliths and the environmental effects on otolith composition. Such elemental fingerprints can help distinguish different stocks or life history stages, identify the origins or nursery areas of fish, and assess population structure. In this study, we examined the stock discrimination and spatial connectivity of cage-cultured and wild stocks of yellowfin seabream (Acanthopagrus latus) from the coastal waters of Shantou, Yangjiang, and Zhanjiang in China southern province Guangdong during 2012–2014, based on otolith trace-elemental signatures using multivariate statistical analysis and machine learning approaches. The concentrations of 13 elements (⁷Li, ²³Na, ²⁴Mg, ⁴⁰Ca, ⁵⁵Mn, ⁵⁶Fe, ⁵⁹Co, ⁵⁹Ni, ⁶⁴Cu, ⁶⁵Zn, ⁸⁸Sr, ¹²²Sb, and ¹³⁷Ba) in the natal spot of fish otoliths, representing the embryonic and paralarval stages of fish, were analyzed using laser ablation inductively coupled plasma mass spectrometry (LA-ICP-MS). Stepwise discriminant analysis and random forests were used to distinguish the cultured and wild stocks of yellowfin seabream, and non-metric multidimensional scaling (NMDS) and cluster analysis were used to determine the spatial variation and connectivity of yellowfin seabream stocks. Overall, the cultured and wild stocks of yellowfin seabream could be identified with classification accuracy of 80.7% and 99.2% by using stepwise discriminant analysis and random forests respectively. When we compared site difference between cultured and wild stocks (site × stock interactions), the classification success was 60.4% for stepwise discriminant analysis and 85.7% for random forests. The misclassification of cultured and wild stocks within the three sites suggested the spatial connectivity between stocks and among sampling locations. Our findings suggested that the three wild stocks of yellowfin seabream from Guangdong coastal waters could be considered as one unit for management, and the difference between cultured and wild stocks was significant for yellowfin seabream from Shantou and Yangjiang, but less significant for yellowfin seabream from Zhanjiang. This study demonstrated that otolith elemental fingerprints can help improve our knowledge on the spatial connectivity, population structure, and life history of fish stocks, and random forests can be a useful tool for identifying cultured and wild stocks compared to the traditional stepwise discriminant analysis.     

Elemental signatures of Acanthochromis polyacanthus otoliths from the Great Barrier Reef have significant temporal, spatial, and between-brood variation

We evaluated the spatial and temporal scales over which otolith signatures varied in a reef fish on the Great Barrier Reef (GBR) using the non-dispersing damselfish Acanthochromis polyacanthus. We found a robust multi-element separation in otolith signatures from reef clusters in the northern and southern GBR. Variance components indicated that this spatial scale accounted for the majority of the variation in two elemental ratios (Ba/Ca and Sr/Ca) over the 2 years of the study. There was also significant variation in elemental signatures between otoliths collected over two consecutive years, as well as within a season. Individual reefs within clusters were less distinguishable based on otolith chemistry and were probably observed by differences within reefs (among sites and broods within sites). These results indicate that it may be difficult to determine the reef of origin for individual fish using otolith chemistry, while determining natal region seems a realistic goal.     

Long-term temporal and spatial variation of macrobenthos in the intertidal soft-bottom flats of two small bights (Chupa Inlet, Kandalaksha Bay, White Sea)

Despite the dynamic nature of spatial pattern, the temporal variation of spatial structure of marine benthic assemblages is rarely assessed using several temporal scales. We quantified the variability of density and biomass of main benthic species in the intertidal soft-bottom flats at two bights in Chupa Inlet (Kandalaksha Bay, the White Sea). The data cover the 21-year period (1987–2008) of a long-term monitoring survey (1987–present) using a hierarchical sampling design with two temporal (year, season within a year) and three spatial scales (bights—7 km, stations within a bight—10–100 m, and replicate samples—10 s cm apart). We used nested ANOVA to test significance and variance components to compare the relative contribution of different scales of variability of density and biomass of 18 most occurring macrobenthic species. Some species demonstrated high large-scale variability, however, the majority showed high small-scale variability and residual variance. The interactive variability was at least as important as the temporal effects, indicating that the spatial pattern changes through time. The assemblages were more variable at small scales and more stable at larger scales. Potential implications for sampling design are discussed.     

Modelling habitat associations with fingernail clam (Family: Sphaeriidae) counts at multiple spatial scales using hierarchical count models

1. Macroinvertebrate count data often exhibit nested or hierarchical structure. Examples include multiple measurements along each of a set of streams, and multiple synoptic measurements from each of a set of ponds. With data exhibiting hierarchical structure, outcomes at both sampling (e.g. within stream) and aggregated (e.g. stream) scales are often of interest. Unfortunately, methods for modelling hierarchical count data have received little attention in the ecological literature. 2. We demonstrate the use of hierarchical count models using fingernail clam (Family: Sphaeriidae) count data and habitat predictors derived from sampling and aggregated spatial scales. The sampling scale corresponded to that of a standard Ponar grab (0.052 m²) and the aggregated scale to impounded and backwater regions within 38–197 km reaches of the Upper Mississippi River. Impounded and backwater regions were resampled annually for 10 years. Consequently, measurements on clams were nested within years. Counts were treated as negative binomial random variates, and means from each resampling event as random departures from the impounded and backwater region grand means. 3. Clam models were improved by the addition of covariates that varied at both the sampling and regional scales. Substrate composition varied at the sampling scale and was associated with model improvements, and reductions (for a given mean) in variance at the sampling scale. Inorganic suspended solids (ISS) levels, measured in the summer preceding sampling, also yielded model improvements and were associated with reductions in variances at the regional rather than sampling scales. ISS levels were negatively associated with mean clam counts. 4. Hierarchical models allow hierarchically structured data to be modelled without ignoring information specific to levels of the hierarchy. In addition, information at each hierarchical level may be modelled as functions of covariates that themselves vary by and within levels. As a result, hierarchical models provide researchers and resource managers with a method for modelling hierarchical data that explicitly recognises both the sampling design and the information contained in the corresponding data.     

Biological quality metrics: their variability and appropriate scale for assessing streams

The concept of spatial scale is at the research frontier in ecology, and although focus has been placed on trying to determine the role of spatial scale in structuring communities, there still is a further need to standardize which organism groups are to be used at which scale and under which circumstances in environmental assessment. This paper contributes to the understanding of the variability at different spatial scales (reach, stream, river basin) of metrics characterizing communities of different biological quality elements (macrophytes, fishes, macroinvertebrates and benthic diatoms) as defined by the Water Framework Directive. For this purpose, high-quality reaches from medium-sized lowland streams of Latvia, Ecoregion 15 (Baltic) were sampled using a nested hierarchical sampling design: (river basin → stream → reach). The variability of metrics within the different groups of biological quality elements confirmed that large-bodied organisms (macrophytes and fish) were less variable than small-bodied organisms (macroinvertebrates and benthic diatoms) at reach, stream and river basin scales. Single metrics of biological quality elements had the largest variation at the reach scale compared with stream and basin scales. There were no significant correlations between biodiversity indices of the different organism groups. The correlation between diversity indices (Shannon’s and Simpson’s) of the biological quality elements (macrophytes, fish, benthic macroinvertebrates and benthic diatoms) and a number of measured environmental variables varied among the different organism groups. Relationships between diversity indices and environmental factors were established for all groups of biological quality elements. Our results showed that metrics of macrophytes and fish could be used for assessing ecological quality at the river basin scale, whereas metrics of macroinvertebrates and benthic diatoms were most appropriate at a smaller scale.     

Learning compositional sequences with multiple time scales through a hierarchical network of spiking neurons

Sequential behaviour is often compositional and organised across multiple time scales: a set of individual elements developing on short time scales (motifs) are combined to form longer functional sequences (syntax). Such organisation leads to a natural hierarchy that can be used advantageously for learning, since the motifs and the syntax can be acquired independently. Despite mounting experimental evidence for hierarchical structures in neuroscience, models for temporal learning based on neuronal networks have mostly focused on serial methods. Here, we introduce a network model of spiking neurons with a hierarchical organisation aimed at sequence learning on multiple time scales. Using biophysically motivated neuron dynamics and local plasticity rules, the model can learn motifs and syntax independently. Furthermore, the model can relearn sequences efficiently and store multiple sequences. Compared to serial learning, the hierarchical model displays faster learning, more flexible relearning, increased capacity, and higher robustness to perturbations. The hierarchical model redistributes the variability: it achieves high motif fidelity at the cost of higher variability in the between-motif timings.     

Meta-population structure in a coral reef fish demonstrated by genetic data on patterns of migration,extinction and re-colonisation

Background  

Management strategies for coral reefs are dependant on information about the spatial population structure and connectivity of reef organisms. Genetic tools can reveal important information about population structure, however, this information is lacking for many reef species. We used a mitochondrial molecular marker to examine the population genetic structure and the potential for meta-population dynamics in a direct developing coral reef fish using 283 individuals from 15 reefs on the Great Barrier Reef, Australia. We employed a hierarchical sampling design to test genetic models of population structure at multiple geographical scales including among regions, among shelf position and reefs within regions. Predictions from island, isolation-by-distance and meta-population models, including the potential for asymmetric migration, local extinction and patterns of re-colonisation were examined.     

Spatial hierarchical approach in community ecology: a way beyond high context-dependency and low predictability in local phenomena

Patterns and functioning of communities, which are determined by a set of processes operating at a large variety of spatial and temporal scales, exhibit quite high context-dependency and low predictability at the fine spatial scales at which recent studies have concentrated. More attention to broader scale and across-scale phenomena may be useful to search for general patterns and rules in communities. In this context, it is effective to incorporate hierarchical spatial scale explicitly into the experimental and sampling design of field studies, an approach referred to here as the spatial hierarchical approach, focusing on a particular assemblage in which biological interaction and species life history are well known. The spatial hierarchical approach can provide insight into the effects of scale in operating processes and answers to a number of important questions in community ecology such as: (1) detection of patterns and processes in spatiotemporal variability in communities, including how to explain the partitioning of pattern information of species diversity at a broad scale into finer scales, and the pattern of spatial variability of community properties at the finest spatial scale; (2) evaluation of changes in patterns observed in macroecology at finer scales; (3) testing of models explaining the coexistence of competing species; and (4) detection of latitudinal patterns in spatiotemporal variability in communities and their causal processes.     

Using hierarchical sampling to understand scales of spatial variation in early coral recruitment

Ecological patterns are created by processes acting over multiple spatial and temporal scales. By combining spatially explicit sampling with variance components models, the relative importance of spatial scale to overall variability can be determined. We used a spatially structured experimental design in the Mombasa Marine National Park in Kenya to quantify variation in coral recruitment across four spatial scales (~1–1,000 m) and to generate hypotheses about processes affecting recruitment and potential sources of post-settlement mortality during early life history. For the dominant recruiting corals (Pocillopora spp.), variation in recruitment on surfaces protected from fish grazing was greatest at the largest spatial scale examined (1,000 m). We hypothesize that recruitment on protected surfaces varies mainly with larval delivery due to different lagoonal circulation and water flow between sites. Conversely, variation on surfaces exposed to fishes was greatest at the smallest spatial scale (1 m). We hypothesize that recruitment on exposed surfaces mainly reflects local differences in the scale and intensity of fish grazing, which may obscure larval delivery patterns. Spatial variation in recruitment can affect many ecological processes and factors, including growth, survival to maturity, the distribution of habitat, and variation in species interaction strengths. This study demonstrates how spatially explicit sampling, followed by variance components modeling to partition variance across scales, can help to identify potential drivers of patterns at each relevant scale.     

Distribution of mating type alleles in the wheat pathogen Mycosphaerella graminicola over spatial scales from lesions to continents

A total of 2035 Mycosphaerella graminicola strains collected from 16 geographic locations on four continents were assayed for the mating type locus. RFLP fingerprints were used to identify clones in each population. At the smallest spatial scale analyzed, both mating types were found among fungal strains sampled from different lesions of the same leaf as well as from different pycnidia in the same lesion. At larger spatial scales, the two mating types were found at equal frequencies across spatial scales ranging from several square meters to several thousand square kilometers. Though the absolute frequencies of the two mating types sometimes varied for different sampling units within the same spatial scale in the hierarchy (plots within a field, fields within a country, or different continents of the world), none of the differences were statistically significant from the null hypothesis of equal frequencies for the two mating types. The evolutionary forces likely to maintain the even distribution of the two mating types in this pathogen were discussed.     

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

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

Discrimination of French grunts (Haemulon flavolineatum, Desmarest, 1823) from mangrove and coral reef habitats using otolith microchemistry

Using French Grunts (Haemulon flavolineatum) held captive within mangrove and reef sites, we determined (a) whether otolith microchemical differences existed between mangroves and reefs separated at a biologically relevant spatial scale (0.25-7.1 km), (b) whether patterns in elemental concentrations were consistent across years, and (c) whether it was possible to identify whether a given fish occupied a mangrove as a juvenile.Three sites were established at Great Exuma, Bahamas (two reefs and one mangrove, May 2001) and at Turneffe Atoll, Belize (one reef and two mangroves, August 2001 and 2002). Using concentrations of Sr and Ba, discriminant function analysis (DFA) indicated unique spatial microchemical signatures of fish from each of the three Bahamas sites allowing an average correct classification of 77%. Using concentrations of Sr, Ba, Sn and Pb (2001), and in addition Li, Mg, Cu and Rb (2002), DFA of the three Belize sites indicated an average correct classification of 68% and 85% in 2001 and 2002, respectively. To assess temporal variability in otolith microchemistry, we compared microchemical signatures of Belize fish from 2001 to those from 2002. On average, 42% of fish from 2002 were correctly classified to their captive sites using chemical information from 2001, thus suggesting considerable temporal variability in otolith microchemistry. Finally, to identify whether a given fish occupied a mangrove during its juvenile stage, we ablated the juvenile portion of the otolith taken from reefs in Belize 2002. Results of this analysis indicated that 36% of 39 individuals taken from the reef had a signature more representative of one of the mangrove sites. Although otolith microchemistry varied temporally and our analysis was restricted to the grouping of individuals to only one of three sites, mangroves appeared to contribute to reef populations.     

Consistency of elemental and isotope‐ratio patterns across multiple scales from individual fish

Assays of elemental and stable‐isotope ratios across growth increments of scales have the potential to provide a non‐lethal alternative to otolith chemistry for identifying migration and ontogenetic trophic shifts. A central assumption when employing scales as otolith analogues is that any scale from an individual will provide equivalent information about the chemical history of that fish. This assumption was investigated with multiple scales from wild and captive euryhaline Atlantic tarpon Megalops atlanticus from the north‐west Gulf of Mexico. Elemental (Sr:Ca) and isotope‐ratio (δ¹³C and δ¹⁵N) life‐history profiles were compared among multiple scales from each fish. All three chemical proxies showed highly consistent patterns among non‐regenerated scales, while patterns in regenerated scales diverged, indicating rapid regrowth of interior scale material at the onset of regeneration. Patterns of Sr:Ca and δ¹³C covaried, supporting their use as salinity proxies, while δ¹⁵N patterns were consistent with ontogenetic diet shifts. Water samples taken from aquaria holding captive fish were used to calculate partition coefficients for a suite of elements in M. atlanticus scales for future quantification of migratory movements in the region. Together, these results support the assumption that non‐regenerated scales from individual M. atlanticus provide equivalent chemical histories, further validating their use as a viable non‐lethal alternative to otoliths.     

Demonstration and implications of habitat-specific chemical signatures in otoliths of juvenile summer flounder (Paralichthys dentatus Linnaeus) in North Carolina

The objective of this pilot study was to determine if otolith signatures of juvenile summer flounder Paralichthys dentatus (Linnaeus) were differentiable between beach and marsh habitat types within two North Carolina (NC), U.S.A., estuarine systems. Elemental signatures were differentiable between habitats in both systems, and in one system at smaller spatial scales (<5 km) than have been reported for any estuarine species using otolith microchemistry. These results indicate the potential for using summer flounder otolith elemental analysis to assess the nursery role of habitats within NC estuaries.     

Spatial subdivision among assemblages of Spanish mackerel, Scomberomorus commerson (Pisces: Scombridae) across northern Australia: implications for fisheries management

Aim This study investigated the use of stable δ¹³C and δ¹⁸O isotopes in the sagittal otolith carbonate of narrow‐barred Spanish mackerel, Scomberomorus commerson, as indicators of population structure across Australia. Location Samples were collected from 25 locations extending from the lower west coast of Western Australia (30°), across northern Australian waters, and to the east coast of Australia (18°) covering a coastline length of approximately 9500 km, including samples from Indonesia. Methods The stable δ¹³C and δ¹⁸O isotopes in the sagittal otolith carbonate of S. commerson were analysed using standard mass spectrometric techniques. The isotope ratios across northern Australian subregions were subjected to an agglomerative hierarchical cluster analysis to define subregions. Isotope ratios within each of the subregions were compared to assess population structure across Australia. Results Cluster analysis separated samples into four subregions: central Western Australia, north Western Australia, northern Australia and the Gulf of Carpentaria and eastern Australia. Isotope signatures for fish from a number of sampling sites from across Australia and Indonesia were significantly different, indicating population separation. No significant differences were found in otolith isotope ratios between sampling times (no temporal variation). Main conclusions Significant differences in the isotopic signatures of S. commerson demonstrate that there is unlikely to be any substantial movement of fish among these spatially discrete adult assemblages. The lack of temporal variation among otolith isotope ratios indicates that S. commerson populations do not undergo longshore spatial shifts in distribution during their life history. The temporal persistence of spatially explicit stable isotopic signatures indicates that, at these spatial scales, the population units sampled comprise functionally distinct management units or separate ‘stocks’ for many of the purposes of fisheries management. The spatial subdivision evident among populations of S. commerson across northern and western Australia indicates that it may be advantageous to consider S. commerson population dynamics and fisheries management from a metapopulation perspective (at least at the regional level).     

Scale-dependent spatial variability of coral assemblages along the Florida Reef Tract

 Coral reef communities of the western Atlantic have changed over the past two to three decades, but the magnitude and causes of this change remain controversial. Part of the problem is that small-scale patterns observed on individual reefs have been erroneously extrapolated to landscape and geographic scales. Understanding how reef coral assemblages vary through space is an essential prerequisite to devising sampling strategies to track the dynamics of coral reefs through time. In this paper we quantify variation in the cover of hard corals in spur-and-groove habitats (13–19 m depth) at spatial scales spanning five orders of magnitude along the Florida Reef Tract. A videographic sampling program was conducted to estimate variances in coral cover at the following hierarchical levels and corresponding spatial scales: (1) among transects within sites (0.01- to 0.1-km scale), (2) among sites within reefs (0.5- to 2-km scale), (3) among reefs within sectors of the reef tract (10- to 20-km scale), and (4) among sectors of the reef tract (50- to 100-km scale). Coral cover displayed low variability among transects within sites and among sites within reefs. This means that transects from a site adequately represented the variability of the spur-and-groove habitat of the reef as a whole. Variability among reefs within sectors was highly significant, compared with marginally significant variability among sectors. Estimates from an individual reef, therefore, did not adequately characterize nearby reefs, nor did those estimates sufficiently represent variability at the scale of the sector. The structure and composition of coral reef communities is probably determined by the interaction of multiple forcing functions operating on a variety of scales. Hierarchical analyses of coral assemblages from other geographic locations have detected high variability at scales different from those in the present study. A multiscale analysis should, therefore, precede any management decisions regarding large reef systems such as the Florida Reef Tract. Accepted: 19 July 1999     

Patterns of connectivity among populations of a coral reef fish

Knowledge of the patterns and scale of connectivity among populations is essential for the effective management of species, but our understanding is still poor for marine species. We used otolith microchemistry of newly settled bicolor damselfish (Stegastes partitus) in the Mesoamerican Reef System (MRS), Western Caribbean, to investigate patterns of connectivity among populations over 2 years. First, we assessed spatial and temporal variability in trace elemental concentrations from the otolith edge to make a ‘chemical map’ of potential source reef(s) in the region. Significant otolith chemical differences were detected at three spatial scales (within-atoll, between-atolls, and region-wide), such that individuals were classified to locations with moderate (52 % jackknife classification) to high (99 %) accuracy. Most sites at Turneffe Atoll, Belize showed significant temporal variability in otolith concentrations on the scale of 1–2 months. Using a maximum likelihood approach, we estimated the natal source of larvae recruiting to reefs across the MRS by comparing ‘natal’ chemical signatures from the otolith of recruits to the ‘chemical map’ of potential source reef(s). Our results indicated that populations at both Turneffe Atoll and Banco Chinchorro supply a substantial amount of individuals to their own reefs (i.e., self-recruitment) and thus emphasize that marine conservation and management in the MRS region would benefit from localized management efforts as well as international cooperation.     

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.     

Identifying migratory contingents of fish by combining otolith Sr:Ca with temporal collections of ambient Sr:Ca concentrations

Ambient strontium:calcium (Sr:Ca) concentrations were determined at the temporal scales of months, weeks and days, in summer and winter at two estuarine sites, and experimentally derived correlations between ambient and otolith Sr:Ca were used to estimate the otolith Sr:Ca concentrations of 'resident' black bream Acanthopagrus butcheri . Wild black bream were collected in summer and winter at the end of the temporal water sampling, and their otolith Sr:Ca concentrations were examined. Wild fish were classified as 'resident' if their otolith Sr:Ca matched the predicted concentrations of resident fish, and 'migrant' if this did not occur. In winter, all fish were classified as resident. In summer, all fish were classified as migrants, with fish spending an average of only 16·8% (estuary 1) and 61·1% (estuary 2) of their time at each estuarine location.     

Unexpected fine-scale population structure in a broadcast-spawning Antarctic marine mollusc

Several recent empirical studies have challenged the prevailing dogma that broadcast-spawning species exhibit little or no population genetic structure by documenting genetic discontinuities associated with large-scale oceanographic features. However, relatively few studies have explored patterns of genetic differentiation over fine spatial scales. Consequently, we used a hierarchical sampling design to investigate the basis of a weak but significant genetic difference previously reported between Antarctic limpets (Nacella concinna) sampled from Adelaide and Galindez Islands near the base of the Antarctic Peninsula. Three sites within Ryder Bay, Adelaide Island (Rothera Point, Leonie and Anchorage Islands) were each sub-sampled three times, yielding a total of 405 samples that were genotyped at 155 informative Amplified Fragment Length Polymorphisms (AFLPs). Contrary to our initial expectations, limpets from Anchorage Island were found to be subtly, but significantly distinct from those sampled from the other sites. This suggests that local processes may play an important role in generating fine-scale population structure even in species with excellent dispersal capabilities, and highlights the importance of sampling at multiple spatial scales in population genetic surveys.