Shellfish growing areas in rural catchments are occasionally affected by elevated faecal contamination from diffuse sources and may be subject to frequent harvest closures/classification downgrades. We combined traditional risk management methods based on sanitary surveys and monitoring of Escherichia coli in seawater and shellfish with faecal source tracking, bacterial source apportionment, and hydrometeorological modelling to determine the causes of elevated E. coli concentrations contributing to harvest closures in Papanui Inlet (Aotearoa New Zealand). These multiple lines of evidence were used to inform a weight of evidence assessment of bacterial contamination in the inlet. Ruminant livestock was estimated to contribute 80% of the faecal coliform loading. Concentrations of E. coli in seawater were low (≤ 87 MPN 100 ml − 1 ) while those in tuaki ( Austrovenus stutchburyi ) occasionally exceeded the “Approved” classification limit (230 MPN 100 g − 1 ). The most frequent positive genetic markers in seawater were the seagull ( Catellicoccus marimammalium ) (54% of seawater samples), and in shellfish, the bovine and seagull markers (both 12.5% of shellfish samples). Solar radiation was negatively correlated with E. coli in tuaki. We found that the growing area is affected by faecal inputs from animal and, to a lesser extent, human (septic tank discharges) sources which elevate contamination to levels detectable in tuaki but not in seawater, particularly in the summer months. The innovative approach can enhance the management of shellfish growing areas affected by intermittent contamination and enables more targeted action to reduce pollution to improve shellfish water quality.
Stabilised organic solids derived from sewage sludge ('biosolids') are applied to land in many countries as an alternative to disposal as land ll. This study evaluates the long-term effects of biosolids applied to forestry plantations on the adjacent intertidal habitats of Moturoa / Rabbit Island (Aotearoa New Zealand). On this island, biosolids are applied to enhance soil fertility and growth of trees (Pinus radiata). However, applications may affect the quality of groundwater and surface water (e.g., increasing concentrations of nutrients and trace metals) and, via seepage into adjacent coastal areas, potentially adversely affect intertidal habitats and biota. Shoreline topography, macroalgal cover, sediment grain size, concentrations of nutrients, trace metals and faecal indicator bacteria, and benthic infaunal community characteristics were studied on three occasions (2008, 2014 and 2019) at twelve intertidal transect sites (four 'reference' and eight 'application') adjacent to forestry blocks where biosolids have been applied intermittently over a period of 24 years. No long-term adverse changes to the environment and community characteristics attributed to biosolids application were detected between reference and application sites. The study provides evidence that biosolids application can co-occur without detectable adverse effects on the nearby intertidal environment.
The frequency and intensity of cyanobacterial blooms is increasing worldwide. Multiple factors are implicated, most of which are anthropogenic. New Zealand provides a useful location to study the impacts of human settlement on lake ecosystems. The first humans (Polynesians) arrived about 750 years ago. Following their settlement, there were marked landscape modifications which intensified after European settlement about 150 years ago. The aims of this study were to reconstruct cyanobacterial communities in six lakes over the last 1000 years and explore key drivers of change. Cyanobacterial environmental DNA was extracted from sediment cores and analysed using metabarcoding and droplet digital PCR. Cyanobacteria, including potentially toxic or bloom forming species, were already present in these lakes prior to human arrival, however their overall abundance was low. Total cyanobacteria abundance and richness increased in all lakes after European settlement but was very pronounced in four lakes, where bloom-forming taxa became dominant. These shifts occurred concomitant with land-use change. The catchment of one deteriorated lake is only moderately modified, thus the introduction of non-native fish is posited as the key factor driving this change. The paleolimnological approach used in this study has enabled new insights into timing and potential causes of changes in cyanobacterial communities.
Fed aquaculture is one of the fastest growing and most valuable food production industries. The efficiency with which farmed fish convert feed into biomass influences both environmental impact and economic revenue. Salmonid species, such as king salmon (Oncorhynchus tshawytscha), exhibit high levels of plasticity in vital rates such as feed intake and growth rates. Accurate estimations of individual variability in vital rates are important for production management. The use of mean trait values to evaluate feeding and growth performance can mask individual-level differences that potentially contribute to inefficiencies. Here, we apply an integral projection model (IPM) to investigate individual variation in growth performance of 1625 individually tagged king salmon fed one of three distinct rations and tracked over 276 days. To capture the observed sigmoidal growth, we compared a non-linear mixed-effects (logistic) model to a linear regression model used within the IPM framework. Ration significantly influenced several aspects of growth. Mean final body mass and mean growth rate increased with ration, however, variance in body mass and feed intake also increased significantly over time. Trends in body mass mean and variance were captured by both logistic and linear models, suggesting the linear model to be suitable for use in the IPM. Higher rations resulted in a decreasing proportion of individuals reaching the cohort’s mean size or larger by the end of the experiment. This suggests that, in our trial, feeding to satiation did not produce the desired effects of efficient and uniform growth in juvenile king salmon. While monitoring individuals through time is challenging in commercial aquaculture settings, recent technological advances combined with an IPM approach could provide new scope for tracking growth performance in experimental and farmed populations. The IPM framework also allows the exploration of other size-dependent processes affecting vital rate functions, such as competition and mortality.
Symbiodiniaceae are a diverse group of dinoflagellates, the majority of which are free-living and/or associated with a variety of protists and other invertebrate hosts. Maintenance of isolated cultures is labour-intensive and expensive, and cryopreservation provides an excellent avenue for their long-term storage. We aimed to cryopreserve 15 cultured isolates from six Symbiodiniaceae genera using dimethyl sulfoxide (DMSO) as the cryoprotectant agent (CPA). Under 15% DMSO, 10 isolates were successfully cryopreserved using either rapid freezing or controlled-rate freezing. Cultures that failed or had low survival, were subjected to (1) a reduction of CPA to 10%, or (2) increased salinity treatment before freezing. At 10% DMSO, three further isolates were successfully cryopreserved. At 15% DMSO there were high cell viabilities in Symbiodinium pilosum treated with 44 parts per thousand (ppt) and 54 ppt culture medium. An isolate of Fugacium sp. successfully cryopreserved after salinity treatments of 54 ppt and 64 ppt. Fatty acid (FA) analyses of S. pilosum after 54 ppt salinity treatment showed increased saturated FA levels, whereas Fugacium sp. had low poly-unsaturated FAs compared to normal salinity (34 ppt). Understanding the effects of salinity and roles of FAs in cryopreservation will help in developing protocols for these ecologically important taxa.
Karlodinium veneficum is a toxic benthic globally distributed dinoflagellate which has direct impacts on human health and the environment. Early and accurate detection of this harmful algal bloom-forming species could be useful for potential risks monitoring and management. In the present work, a real-time PCR targeting the internal transcribed spacer ribosomal DNA region for the specific detection and absolute quantification of K. veneficum was designed. Then, the assay conditions were adjusted and validated. The developed qPCR was highly specific for the target species and displayed no cross-reactivity with closely related dinoflagellates and/or other microalgal species commonly distributed along the Tunisian coast. Its lowest detection limit was 5 rDNA copies per reaction, which is often considered satisfying. qPCR assay enumeration accuracy was evaluated using artificially inoculated environmental samples. The comparison of the cell abundance estimates obtained by qPCR assay with the theoretical estimates showed no statistically significant difference across a range of concentrations. We suggest that the qPCR approach developed in the present study may be a valuable tool to investigate the distribution and seasonal dynamics of K. veneficum in marine environments.
Recent global trade disruptions, due to blockage of the Suez Canal and cascading effects of COVID-19, have altered the movement patterns of commercial ships and may increase worldwide invasions of marine non-indigenous species. Organisms settle on the hulls and underwater surfaces of vessels and can accumulate rapidly, especially when vessels remain stationary during lay-ups and delays. Once present, organisms can persist on vessels for long-periods (months to years), with the potential to release propagules and seed invasions as ships visit ports across the global transportation network. Shipborne propagules also may be released in increasing numbers during extended vessel residence times at port or anchor. Thus, the large scale of shipping disruptions, impacting thousands of vessels and geographic locations and still on-going for over two years, may elevate invasion rates in coastal ecosystems in the absence of policy and management efforts to prevent this outcome. Concerted international and national biosecurity actions, mobilizing existing frameworks and tools with due diligence, are urgently needed to address a critical gap and abate the associated invasion risks.
Identifying compounds responsible for the observed toxicity of the Gambierdiscus species is a critical step to ascertaining whether they contribute to ciguatera poisoning. Macroalgae samples were collected during research expeditions to Rarotonga (Cook Islands) and North Meyer Island (Kermadec Islands), from which two new Gambierdiscus species were characterized, G. cheloniae CAWD232 and G. honu CAWD242. Previous chemical and toxicological investigations of these species demonstrated that they did not produce the routinely monitored Pacific ciguatoxins nor maitotoxin-1 (MTX-1), yet were highly toxic to mice via intraperitoneal (i.p.) injection. Bioassay-guided fractionation of methanolic extracts, incorporating wet chemistry and chromatographic techniques, was used to isolate two new MTX analogs; MTX-6 from G. cheloniae CAWD232 and MTX-7 from G. honu CAWD242. Structural characterization of the new MTX analogs used a combination of analytical chemistry techniques, including LC–MS, LC–MS/MS, HR–MS, oxidative cleavage and reduction, and NMR spectroscopy. A substantial portion of the MTX-7 structure was elucidated, and (to a lesser extent) that of MTX-6. Key differences from MTX-1 included monosulfation, additional hydroxyl groups, an extra double bond, and in the case of MTX-7, an additional methyl group. To date, this is the most extensive structural characterization performed on an MTX analog since the complete structure of MTX-1 was published in 1993. MTX-7 was extremely toxic to mice via i.p. injection (LD50 of 0.235 µg/kg), although no toxicity was observed at the highest dose rate via oral administration (155.8 µg/kg). Future research is required to investigate the bioaccumulation and likely biotransformation of the MTX analogs in the marine food web.
The management of neurological disorders such as dementia associated with Alzheimer's or Parkinson's disease includes the use of cholinesterase inhibitors. These compounds can slow down the progression of these diseases and can also be used in the treatment of glaucoma and myasthenia gravis. The majority of the cholinesterase inhibitors used in the clinic are derived from natural products and our current paper describes the use of a small marine pharmacophore to develop potent and selective cholinesterase inhibitors. Fourteen small inhibitors were designed based on recent discoveries about the inhibitory potential of a range of related marine secondary metabolites. The compounds were evaluated, in kinetic enzymatic assays, for their ability to inhibit three different cholinesterase enzymes and it was shown that compounds with a high inhibitory activity towards electric eel and human recombinant acetylcholinesterase (IC50 between 20-70 μM) could be prepared. It was also shown that this compound class was particularly active against horse serum butyrylcholinesterase, with IC50 values between 0.8-16 μM, which is an order of magnitude more potent than the clinically used positive control neostigmine. The compounds were further tested for off-target toxicity against both human umbilical vein endothelial cells and bovine and human erythrocytes and were shown to display a low mammalian cellular toxicity. Overall, the study illustrates how the brominated dipeptide marine pharmacophore can be used as a versatile natural scaffold for the design of potent, and selective cholinesterase inhibitors.
The inefficient conversion of seed mussels, or 'spat' into market-ready mussels is a major problem for mussel aquaculture operations worldwide, where sustained crop losses throughout the production cycle severely disrupt production and ultimately constrain industry growth. However, despite the severity of the problem for individual mussel farm operators, few studies have sought to examine this inefficiency on an industry-wide scale to assess the broader extent of the biological and economic losses. The Greenshell™ mussel (Perna canaliculus) industry is New Zealand's largest and most valuable aquaculture industry, and yet despite its success, the industry remains extremely inefficient at converting spat into market-ready adult mussels. Consequently, this study compiled and compared historical industry-wide data on the harvesting of spat with national statistics on the production of adult mussels by the aquaculture industry. The results of the analyses show that despite substantial increases in the harvesting of wild spat over the last 10 years, the production of market-ready mussels from the industry has not increased concordantly, remaining largely static at around 90,000 t, amounting to around 1.78 billion adults a year during this period. From 2015 to 2020 on average, 344 billion spat were harvested from the wild each year, yet <1% of these were retained on Greenshell™ farms from seeding out through to final harvest. Careful evaluation of historical data reveals the efficiency of converting seeded Greenshell™ spat to harvested market-ready mussels has halved over the last 30 years, with the cause of this marked decline uncertain. These results highlight the inefficient nature of the Greenshell™ industry and show that improving production efficiency by even a small margin would result in substantial production gains for the industry.
Fish nutrition has driven advances in the efficiency, sustainability, and product quality of aquaculture production, facilitating its expansion of aquaculture production [...]
Monitoring fish is necessary for understanding population dynamics, tracking distribution patterns and evaluating conservation efforts. Molecular techniques targeting environmental DNA (eDNA) are now considered effective methods for detecting specific species or characterising fish communities. The analysis of DNA from lake‐surface sediments (sedDNA) can provide a time‐integrated sampling approach which reduces the variability sometimes observed in water samples. However, studies of sedDNA have had varying success in detecting fish. The present study aimed to determine the most effective extraction method for recovering fish DNA from lake‐surface sediments. A literature review was undertaken to identify DNA extraction methods used previously on aquatic sediments targeting aquatic and terrestrial animals. Five methods with various modifications were tested to establish their ability to desorb extracellular DNA. Based on these results, two methods were selected and optimised, and the recovery of fish sedDNA characterised using droplet digital PCR assays targeting eel and perch (Anguilla australis, Anguilla dieffenbachii, Perca fluviatilis). A range of sediment masses (0.25–20 g) were assessed to establish the optimal amount required to accurately assess fish sedDNA. The DNA extraction methods found to be most effective at recovering extracellular DNA spiked into small sediment masses (0.25 g) were the DNeasy PowerSoil DNA Isolation Kit (QIAGEN), and the ABPS protocol which involved an initial alkaline buffer extraction followed by the PowerSoil extraction kit. For larger sediment masses (>0.25 g) the ABPS protocol or the DNeasy PowerMax Soil Kit (QIAGEN) with an additional ethanol DNA concentration step (PMET protocol) yielded the highest concentrations of target genes across a range of lake sediments. Larger sediment masses (≤20 g was tested) increased the likelihood of detection of fish in sedDNA. Optimisation of the ABPS protocol was required (65°C incubation temperature, pooling of multiple PowerSoil extractions) to overcome technical challenges related to co‐precipitation of organic content in lake‐surface sediments. This optimised ABPS protocol was called the “Lakes ABPS protocol”. We recommend the use of the Lakes ABPS protocol as it is cheaper than the PMET protocol. Additionally, after the first extraction step, the process can be automated on a DNA extraction robot, allowing for higher sample throughput. A mass of 10 g is suggested, although higher detection is achieved with more sediment, a suite of challenges, such as co‐precipitation of organic content, are encountered when the amount is increased. This study highlights the complexity of the extraction and detection of fish sedDNA from lake sediment, especially when it has a high organic content. We have optimised a DNA extraction method to overcome some of these complexities that can be applied to a wide range of lake sediments.
The objective of this study was to develop a nutrient-demand model for King salmon (Oncorhynchus tshawytscha) based on a factorial nutrient demand approach. Growth potential was defined using laboratory data to define the thermal response boundaries and benchmarked against commercial farm-data. These factors were combined to determine the size specific mass-gain of the animal across its production cycle. Algorithms of body composition across the size ranges for this species were defined based on the analysis of whole-body samples across the size range from 110 g/fish to 4786 g/fish. The utilisation efficiencies and maintenance demand for protein (46%) and energy (66%) were determined. Each of these factors were then assembled into a model that allowed the iterative estimation of macronutrient specifications of diets, based on intake being driven by the demands for energy. Demands for essential amino acids, and how these demands change with animal size, were predicted by applying the ideal protein concept to protein gain and maintenance, combined with estimates of variable utilisation efficiencies of individual amino acids. Modelled estimations of energy and protein demands were slightly higher than most of the empirical data, with the model suggesting that a higher dietary level of protein, relative to other salmonid diets, has potential benefits for King salmon.
Fucoxanthin is a carotenoid in algae with purported beneficial health-related properties including antioxidant, anti-photoaging, anti-metastasis, anti-hypertensive activity and more. These properties give fucoxanthin the potential to be used in cosmetic, dietary, and medicinal applications. This study evaluates the use of deep eutectic solvents (DESs) to extract fucoxanthin from the microalgae Tisochrysis lutea. Conductor-like Screening Model for Real Solvents (COSMO-RS) was used to screen the performance of 24 different types of DESs in the extraction of fucoxanthin based on their calculated capacities. Experimental extraction validation was then carried out using the 6 top-ranked DESs. The experimental results revealed that the extraction capacity of the thymol: dodecanoic acid DES (1.25: 1 molar ratio) for fucoxanthin was the most efficient (7.69 mg/g dry biomass weight (DW)) among the DESs explored under the screening conditions and was higher than the capacity of the conventional solvents methanol (6.29 mg/g DW) and ethanol (6.75 mg/g DW). This corresponded with COSMO-RS screening results. Then, the optimisation of extraction conditions for fucoxanthin using thymol: dodecanoic acid DES was further investigated, revealing that the highest yield of fucoxanthin (22.03 mg/g DW) was extracted at optimum experimental conditions at a temperature of 36.2 ℃, stirring time of 2.58 h, and the biomass percentage of 11.36%. Additionally, fucoxanthin showed good stability in thymol: dodecanoic acid DES over eleven days of storage. After seven extraction cycles, the final fucoxanthin concentration (13.06 mg/mL DES) resulted in a good reusability of the terpene-based food safe DES.
David Schindler and his colleagues pioneered studies in the 1970s on the role of phosphorus in stimulating cyanobacterial blooms in North American lakes. Our understanding of the nuances of phosphorus utilization by cyanobacteria has evolved since that time. We review the phosphorus utilization strategies used by cyanobacteria, such as use of organic forms, alternation between passive and active uptake, and luxury storage. While many aspects of physiological responses to phosphorus of cyanobacteria have been measured, our understanding of the critical processes that drive species diversity, adaptation and competition remains limited. We identify persistent critical knowledge gaps, particularly on the adaptation of cyanobacteria to low nutrient concentrations. We propose that traditional discipline-specific studies be adapted and expanded to encompass innovative new methodologies and take advantage of interdisciplinary opportunities among physiologists, molecular biologists, and modellers, to advance our understanding and prediction of toxic cyanobacteria, and ultimately to mitigate the occurrence of blooms.
The invasive Mediterranean fanworm Sabella spallanzanii (Gmelin, 1791) is a notifiable organism under New Zealand’s Biosecurity Act and is recognized as a marine pest of particular concern, that must be reported to the Ministry for Primary Industries (MPI), New Zealand. Since its first detection in 2008, great effort and financial resources are put into surveillance and removal of individuals to contain population growth and spread. Sensitive molecular detection techniques gain great interest and are being increasingly tested for the fanworm detection in marine high-risk sites (i.e., ports and marinas) around New Zealand. However, conventional molecular detection via PCR assays from environmental DNA (eDNA) samples requires specific laboratory resources and technical expertise. This restricts the wider applicability of this approach by biosecurity practitioners or communities willing to be engaged in biosecurity surveillance. To provide end-users with a fast, easy and highly specific way to detect S. spallanzanii directly at the site of interest, a species-specific recombinase polymerase amplification (RPA) assay was designed to be read-out with lateral flow strips (RPA-LF). The RPA generates amplification within 20 minutes at 37-39°C, with a detection limit of 10 pg of the target DNA and was matching the detection limit of digital droplet PCR (ddPCR) when performed on eDNA samples. A simplified visual protocol for non-scientist users of the assay was developed and improved through independent trials with different end-user groups. The assay applicability was verified in a final validation trial with participants without scientific background resulting in 50 percent of the participants successfully detecting S. spallanzanii. Participants rated the ease of use and performance and read-out mostly as easy-to-very easy with overall positive written feedback on its usability for citizen science applications.
Impacts of Marine Plastic Debris (MPD) on marine ecosystems are among the most critical environmental concerns of the past three decades. Virgin plastic is often cheaper to manufacture than recycled plastics, increasing rates of plastic released into the environment and thereby impacting ecosystem health and functioning. Along with other environmental effects, MPD can serve as a vector for marine hitchhikers, facilitating unwanted organisms' transport and subsequent spread. Consequently, there is a growing demand for more eco-friendly replacements of conventional plastic polymers, ideally with fit-for-purpose properties and a well-understood life cycle. We enriched polybutylene succinate (PBS) with three different concentrations of oyster shell to investigate the dynamics of biofouling formation over 18 weeks at the Nelson Marina, Aotearoa/New Zealand. Our study focused on oyster shell concentration as a determinant of fouling assemblages over time. While generally considered as a waste in the aquaculture sector, we used oyster shells as a variable of interest to investigate their potential for both, environmental and economic benefits. Using bacterial 16S and eukaryotic 18S rRNA gene metabarcoding, our results revealed that following immersion in seawater, time played a more critical role than substrate type in driving biofouling community structures over the study period. In total, 33 putative non-indigenous species (NIS) and 41 bacterial families with putative plastic-degrading capability were detected on the different substrates. Our analysis of NIS recruitment revealed a lower contribution of NIS on shell-enriched substrates than unadulterated polymers samples. In contrast, the different concentrations of oyster shells did not affect the specific recruitment of bacterial degraders. Taken together, our results suggest that bio-based polymers and composites with increased potential for biodegradability, recyclability, and aptitude for the selective recruitment of marine invertebrates might offer a sustainable alternative to conventional polymers, assisting to mitigate the numerous impacts associated with MPD.
To enable environmental management actions to be more effectively prioritized, cumulative effects between multiple stressors need to be accounted for in risk-assessment frameworks. Ecological risk and uncertainty are generally high when multiple stressors occur. In the face of high uncertainty, transparent communication is essential to inform decision-making. The impact of stressor interactions on risk and uncertainty was assessed using generalized linear models for additive and multiplicative effect of six anthropogenic stressors on the abundance of estuarine macrofauna across New Zealand. Models that accounted for multiplicative stressor interactions demonstrated that non-additive effects dominated, had increased explanatory power (6 to 73 % relative increase between models), and thereby reduced the risk of unexpected ecological responses to stress. Secondly, 3D-plots provide important insights in the direction, magnitude and gradients of change, and aid transparency and communication of complex stressor effects. Notably, small changes in a stressor can cause a disproportionally steep gradient of change for a synergistic effect where the tolerance to stressors are lost, and would invoke precautionary management. 3D-plots were able to clearly identify directional shifts where the nature of the interaction changed from antagonistic to synergistic along increasing stressor gradients. For example, increased nitrogen load and exposure caused a shift from positive to negative effect on the abundance of a deposit-feeding polychaete (Magelona). Assessments relying on model coefficient estimates, which provide one effect term, could not capture the complexities observed in 3D-plots and are at risk of mis-identifying interaction types. Finally, visualising model uncertainty demonstrated that although error terms were higher for multiplicative models, they better captured the uncertainty caused by data availability. Together, the steep gradients of change identified in 3D-plots and the higher uncertainty in model predictions in multiplicative models urges more conservative limits to be set for management that account for risk and uncertainty in multiple stressor effects.
Nutrient enrichment of lakes from anthropogenic activities is a significant and increasing issue globally, impairing the health, biodiversity and service provisioning from lakes, with impacts on cultural, recreational, economic and aesthetic values. Internal nutrient loads from lakebed sediment releases are a primary cause of lake eutrophication and have necessitated geoengineering methods to mitigate releases and speed up recovery from eutrophication. Our objective in this review was to evaluate the use of oxygen nanobubbles as a geoengineering technology to remediate low oxygen conditions at the lake sediment/water interface, as a precursor to alleviating eutrophication linked to high internal nutrient loads. Oxygen nanobubbles (NBs) are bubbles < 1000 nm formed at the interface of solid surfaces and aqueous solutions. These bubbles have higher density than water, persist for longer and facilitate greater oxygen solubility than larger bubbles. Methods have been developed to enable NB formation at the surface of carrier materials, which are then used in conjunction with modified local soils (MLSs), to ‘floc, lock and oxygenate’ to strip nutrients from the water column, locking them in lakebed sediments and oxygenating the sediments to prevent re-release of nutrients. Most studies of NBs for lake restoration have thus far only demonstrated their potential for this purpose, using short-term, small-scale core incubations conducted mainly in laboratory settings. Work is required to (1) address scalability, including procurement and cost, (2) extend laboratory incubation studies to large outdoor enclosures and pond/lake trials, (3) examine longevity of the effects in the natural environment, including potential for MLSs to smother benthos and/or have toxic effects, and (4) extend to a range of lake environments and MLS types. Legal, cultural and social acceptance of the technology is another prerequisite of applications in the natural environment and requires individualised analysis. Until these issues are addressed in a systematic way that addresses scalability and recommends suitable carrier materials and MLSs, NBs may continue to remain largely untried as a geoengineering method to address lake eutrophication.
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