Marine Zooplankton Research Articles

Toxicological impacts of plastic microfibers from face masks on Artemia salina: An environmental assessment using Box-Behnken design

An increase in global plastic manufacturing and subsequent disposal has resulted in widespread increase in microplastic pollution. Particularly, the post-COVID surge in face mask usage has introduced significant volumes of synthetic plastic microfibers into the environment, posing new ecological risks. Present study examines the toxicological impacts of face mask derived microfibers on Artemia salina, a key marine zooplankton species, using the Box-Behnken design to assess the effects of microfiber dosage (0.1 mg/L- 5 mg/L), salinity (0.5 ppt–30 ppt), temperature (10 °C-45 °C), and cyst stocking density (10 cysts/L-100 cysts/L) on hatching efficiency and swimming competencies. Results demonstrated that higher microfiber dosages (2.5 mg/L-5 mg/L) significantly reduced the hatching efficiencies and swimming competencies, while temperature and cyst density also modulated these effects. Additionally, survival assays indicated a significant reduction in survival rates with increasing microfiber concentrations, attributed to the bioaccumulation, oxidative stress and developmental deformities in the organism. The study further explores the leaching of chromophoric dissolved organic matter and turbidity, revealing a direct correlation with microfiber dosage and exposure duration. These findings underscore the urgent need for mitigation strategies to address microfiber pollution and protect aquatic ecosystems. Data presented from the research provides valuable insights into the environmental impacts of plastic microfiber contamination, emphasizing the necessity for continued investigation into effective solutions for managing plastic waste.

Editorial: Ecology of marine zooplankton and micronekton in polar and sub-polar areas

Editorial: Ecology of marine zooplankton and micronekton in polar and sub-polar areas

Exploring marine zooplankton dynamics through carbon stable isotope signatures in a recently marine submarine volcano

Submarine emissions of gases from hydrothermal vents alter the surrounding chemical environment, influencing species responses to the resulting environmental gradients. The 2011 underwater eruption of the Tagoro volcano off the coast of El Hierro in the Atlantic Ocean changed the physical and chemical conditions, impacting the distribution of pelagic fauna. Post-eruptive stages from 2013 to 2018 revealed changes in both benthic and pelagic communities, with continuous eruptions further affecting local carbon cycle through shifts in nutrient concentrations and isotopic composition. The lowest δ13C values in zooplanktonic primary consumers were found in areas directly influenced by the Tagoro submarine volcano. Although the mixing model results show that phytoplankton is the primary carbon source for copepods, contributing an average of 28.3% of their carbon, magmatic CO2 and seawater dissolved inorganic carbon each account for approximately 17–18%. The isotopic signatures reveal a gradient of enrichment in copepods, reflecting the influence of inorganic nutrient and gas emissions from the volcano's centre to its periphery within the marine ecosystem. This data is useful for understanding the worldwide significance of environmental stressors such as volcanic eruptions on marine organisms.

Mechanism of barotaxis in marine zooplankton

Hydrostatic pressure is a dominant environmental cue for vertically migrating marine organisms but the physiological mechanisms of responding to pressure changes remain unclear. Here, we uncovered the cellular and circuit bases of a barokinetic response in the planktonic larva of the marine annelid Platynereis dumerilii. Increased pressure induced a rapid, graded, and adapting upward swimming response due to the faster beating of cilia in the head multiciliary band. By calcium imaging, we found that brain ciliary photoreceptors showed a graded response to pressure changes. The photoreceptors in animals mutant for ciliary opsin-1 had a smaller sensory compartment and mutant larvae showed diminished pressure responses. The ciliary photoreceptors synaptically connect to the head multiciliary band via serotonergic motoneurons. Genetic inhibition of the serotonergic cells blocked pressure-dependent increases in ciliary beating. We conclude that ciliary photoreceptors function as pressure sensors and activate ciliary beating through serotonergic signalling during barokinesis.

Mechanism of barotaxis in marine zooplankton.

Hydrostatic pressure is a dominant environmental cue for vertically migrating marine organisms but the physiological mechanisms of responding to pressure changes remain unclear. Here, we uncovered the cellular and circuit bases of a barokinetic response in the planktonic larva of the marine annelid Platynereis dumerilii. Increased pressure induced a rapid, graded, and adapting upward swimming response due to the faster beating of cilia in the head multiciliary band. By calcium imaging, we found that brain ciliary photoreceptors showed a graded response to pressure changes. The photoreceptors in animals mutant for ciliary opsin-1 had a smaller sensory compartment and mutant larvae showed diminished pressure responses. The ciliary photoreceptors synaptically connect to the head multiciliary band via serotonergic motoneurons. Genetic inhibition of the serotonergic cells blocked pressure-dependent increases in ciliary beating. We conclude that ciliary photoreceptors function as pressure sensors and activate ciliary beating through serotonergic signalling during barokinesis.

Body-size reductions in dacryoconarid tentaculitoids during Late Devonian warming

Body size is an essential factor in an organism’s survival, and when paired with paleoenvironmental proxies, size trends can provide insights into a lineage’s evolutionary responses to changing environmental conditions. This study explores the diversity and body-volume trends of dacryoconarid tentaculitoids, globally abundant marine zooplankton, in the Devonian of the Appalachian Basin (eastern United States), spanning the late Givetian through the middle Frasnian punctata carbon isotope excursion. Using statistical approaches to model trends, we find evidence of a gradual, within-lineage reduction in styliolinid adult body sizes starting at the Givetian-Frasnian boundary. This reduction is followed by a significant decrease in both adult and initial chamber volumes during the punctata excursion. At the Givetian-Frasnian boundary, annulated forms (nowakiids) become rare and smooth forms (styliolinids) begin to dominate the assemblage. Using pre-existing geological and geochemical data sets, we consider environmental factors, including sea level, anoxia, nutrient availability, and temperature, as potential drivers of body-size reductions. Bottom-water anoxia most likely did not influence body-size trends of this pelagic group, but frequent water-column overturning in the Frasnian or other exchange between deep and shallow water may have affected taxonomic composition, favoring styliolinids. Sea-surface temperature correlates inversely with body size, suggesting that warming beginning in the early Frasnian may have contributed to gradual, long-term size reductions. Rising temperatures through the middle Frasnian may have led to the disappearance of dacryoconarids in the northern Appalachian Basin after the excursion.

Dynamic population modeling of bacterioplankton community response to gelatinous marine zooplankton bloom collapse and its impact on marine nutrient balance

The diverse microbial community in the ocean, encompassing various metabolic types, interacts with the wide array of compounds in the dissolved organic matter (DOM) pool, thereby influencing the ocean’s biogeochemical state and, consequently, the global climate. Our understanding of the interactions between specific DOM constituents and microbial consortia remains limited, necessitating further refinement to achieve a mechanistic comprehension of the relationship between the DOM field and the microbial metabolic network. Attaining this level of understanding is crucial for accurately predicting the marine ecosystem’s response to natural and anthropogenic perturbations. To address this gap, we developed a bacterial population model based on the von Foerster equation. This model aims to describe the complex microbial-mediated degradation of gelatinous zooplankton (hereinafter ‘jellyfish’) detritus, as an important, but largely overlooked source of DOM in the ocean. By considering microbial growth and decay, as well as DOM uptake, and nutrient release, the model is able to describe the microbial community’s life cycle, and the biochemical transformations of the jellyfish-derived organic matter. We fitted the model to results of laboratory microcosm experiments conducted to simulate scenarios experienced by ambient microbiomes during decay of two different jellyfish species in the northern Adriatic Sea. By interpreting the fitted parameters, we highlight the differences in the microbial response to different jellyfish species, namely how these affect the microbial community composition and the release of nutrients. This model has been specifically designed for integration with ocean circulation models to create a comprehensive physical-biogeochemical ocean model. Such an extended model can be utilized for multi-scale simulations to assess the system’s response to jellyfish and jellyfish-derived organic matter. Given that jellyfish blooms may become more prevalent under future ocean scenarios, this modeling approach is essential for understanding their potential impact on marine ecosystems.

Net type, tow duration and day/night sampling effects on the composition of marine zooplankton derived from metabarcoding

Net type, tow duration and day/night sampling effects on the composition of marine zooplankton derived from metabarcoding

Using bioluminescence as a tool for studying diversity in marine zooplankton and dinoflagellates: an initial assessment.

Bioluminescence is light chemically produced by an organism. It is widespread across all major marine phyla and has evolved multiple times, resulting in a high diversity of spectral properties and first flash kinetic parameters (FFKP). The bioluminescence of a system is often a good proxy for planktonic biomass. The species-specific parameters of bioluminescent displays can be measured to identify species in situ and describe planktonic biodiversity. Most bioluminescent organisms will flash when mechanically stimulated i.e., when subjected to supra-threshold levels of shear stress. Here we compare first flash kinetic parameters such as flash duration, peak intensity, rise time, decay time, first-flash mechanically stimulated light and e-folding time obtained with the commercially available Underwater Bioluminescence Assessment Tool (UBAT). We provide descriptions of the first flash kinetic parameters of several species of dinoflagellates Pyrocystis fusiformis, Pyrocystis noctiluca, Pyrodinium bahamense, Lingulodinium polyedra, Alexandrium monilatum and two zooplankton (the ctenophore Mnemiopsis leidyi and the larvacean Oikopleura sp.). FFKPs are then compared and discussed using non-parametric analyses of variance (ANOVAs), hierarchical clustering and a linear discriminant analysis to assess the ability to use bioluminescence signatures for identification. Once the first flash kinetic parameters of a bioluminescent species have been described, it is possible to detect its presence using emissions collected by in situ bathyphotometers. Assessing abundance and diversity of bioluminescent species may therefore be possible.

The impact of artificial light pollution at night on the life history parameters of rotifer Brachionus plicatilis with different food experiences

Artificial light at night (ALAN) may pose threat to rotifer Brachionus plicatilis. Additionally, the food of rotifer, i.e. algal community composition, often fluctuates. Thus, we selected five wavelengths of ALAN (purple, blue, green, red, white) and a three-colored light flashing mode (3-Flash) to test their impacts on life history traits of B. plicatilis with different food experiences, including those feeding Chlorella vulgaris (RC) or Phaeocystis globosa (RP). Results indicated purple ALAN promoted RC development, white ALAN inhibited RC development, while 3-Flash and white ALAN promoted RP development. Under red and white ALAN, RP increased fecundity but decreased lifespan. High-quality food enhanced rotifer's resistance to the impact of ALAN on lifespan. ALAN and food experience interacted on B. plicatilis. The effect of blue ALAN has less negative effects on B. plicatilis, based on hierarchical cluster analysis. Such findings are helpful to evaluate the potential impact of ALAN on marine zooplankton.

The Spatial Distribution of Copepod Functional Traits in a Highly Anthropized Mediterranean Coastal Marine Region

Copepods dominate marine zooplankton in abundance and play key roles in pelagic food webs. These small crustaceans show high taxonomic and functional diversity. Although there has been considerable research on their taxonomy, only a few studies have focused on their functional traits. In this study, we analyzed the functional traits of 95 copepod species, considering their body size, trophic regime, feeding behavior, and spawning strategy. Based on samples collected during two surveys (autumn 2020 and summer 2021) located in the coastal waters of three gulfs (Gaeta, Naples, and Salerno) in the highly populated Campania region (the central Tyrrhenian Sea, NW Mediterranean), we identified nine functional groups of copepods with different characteristics. The group that comprised herbivorous copepods with feeding currents and a broadcast strategy was the most abundant in both seasons and all gulfs. This group was dominated by Acartia clausi, Centropages typicus, Temora stylifera, and the Paracalanus parvus complex. The other functional groups showed differences in their temporal and spatial distribution. Our study reports the functional diversity of copepods along the Campania coast, thus contributing to advancing our knowledge of the planktonic trophic structure in a region of considerable importance due to its marine resources and services.

Arsenic bioaccumulation and biotransformation in the marine copepod Tigriopus japonicus under chronic dietborne and waterborne exposure

Arsenic (As) can be transferred along the food chain, while little is known about the toxic effects of dietborne As on marine copepods. In this study, we investigated the short-term and long-term effects of waterborne and dietborne As exposure on the bioaccumulation and biotransformation, as well as developmental toxicity of Tigriopus japonicus. Under acute As exposure, As bioaccumulation increased and reached a plateau with increasing exposure concentration. Moreover, As accumulation at dietborne exposure was 4.3 and 5.7 times greater than that at control group for AsIII and AsV, respectively. At chronic As exposure, As accumulation increased continuously with exposure time, with a 2.8-day extension of development time and a 45% reduction in 10-d fecundity under dietborne exposure compared to control, whereas 2.3-day extension of development time and a 20% reduction in 10-d fecundity were observed under waterborne exposure. Among As species, inorganic As had the highest concentrations, but the proportion of inorganic As decreased from 89% to 63% during 4 to 21 d of exposure, suggesting the conversion of inorganic As to organic As. The organic As was dominated by arsenobetaine (AsB, 13–25%), followed by monomethylarsenic (MMA, 8–25%). These results suggest that dietborne exposure has more pronounced toxic effects on T. japonicus, but the toxicity of As could be reduced through biotransformation under chronic exposure. Therefore, the arsenic species should be considered when assessing As toxicity.

Trade-off between drag and catch performance when designing zooplankton trawls

The aquaculture sector is in pursuit of sustainable and cost-effective raw materials for feed, and the copepod Calanus finmarchicus is a marine zooplankton species of commercial interest because of its high abundance in northern areas. These copepods have the potential to meet the demand for vast quantities of marine raw materials. However, the lack of an energy- and catch-efficient trawl technology has limited the development of this fishery. Therefore, this study investigated the effect of two central trawl net design parameters mesh size and taper angle to identify which values that provide optimal trade-off balance between gear drag and catch efficiency. We conducted flume tank experiments using a series of plankton nets varying in mesh size (250–1000 μm), solidity ratio (0.54–0.76), and taper angle (5°– 30°) to acquire data on gear drag. The same nets were then used in fishing trials to obtain data on their catch performance. This study shows that zooplankton nets with a mesh size of approximately 500 μm and a low taper angle of about 5° provided the best trade-off between drag and catch performance.

Toxicity assays and in silico approach to assess the impacts of chlorine dioxide on survival, respiration and some biochemical markers of marine zooplankton

Chlorination is the common antifouling method in desalination and power plant water intake structures to control microbial and macrofouling growth. In this study, the impacts of chlorine dioxide on toxicity, metabolic activity and biochemical markers like glutathione S-transferase and catalase enzyme activity were tested using four zooplankton species (Centropages sp., Acartia sp., Oncaea sp., and Calanus sp.) collected from the Red Sea. The zooplankton species were treated with different concentrations (0.02, 0.05, 0.1, 0.2, and 0.5 mg L−1) of chlorine dioxide. Further, chlorite, the main decomposition product of chlorine dioxide, was used for molecular docking studies against glutathione S-transferase and catalase enzymes. The results indicated the LC50 range of 0.552–1.643 mg L−1 for the studied zooplankton species. The respiration rate of the zooplankton increased due to the chlorine dioxide treatment with a maximum of 0.562 μg O2 copepod h−1 in Acartia. The glutathione S-transferase and catalase enzyme activities showed elevated values in zooplankton treated with chlorine dioxide. Molecular docking of chlorite with enzymes involved in antioxidant defense activity, such as glutathione S-transferase and catalase enzyme showed weak interactions. Overall, this study yielded significant insights for understanding the effects of chlorine dioxide on the survival, metabolism, and biochemical composition of marine zooplankton.

Identification of Marine Zooplankton in the Maheshkhali Channel, Cox’s Bazar, Bangladesh

The purpose of the study was to identify and characterize marine zooplankton in the Maheshkhali Channel, Cox’s Bazar, Bangladesh. A total of 25 species were identified from the samples collected from the surface water during November, 2022. They were categorized into 4 major groups such as copepoda, crustaceans, ichthyoplankton and meroplankton. Copepods were the most dominant. Among the copepods, Oithona sp., Oncaea sp., Calanus sp., Subeucalanus flemingeri, Cyclops sp., Acartia sp., Labidocera sp. and Canthocalanus pauper were ubiquitous, with Oithona brevicornis, Oithona nana, Acartia tonsa, Subeucalanus sp. being moderately frequent. The presence of Centropages sp., Paracalanus aculeatus, Paracalanus sp., Pseudodiaptomas sp. and Acartia (Odonticartia) ohtsukai were rare. Shrimp larvae, crab zoea and ichthyoplankton were moderately found. Mysid shrimps were found to be highly prevalent whereas ctenophora larvae were less frequent. Copepod, naupliar larvae and copepodites were observed to be consistently present throughout the observed area. The identified species possess great significance in the marine food web. This investigation is the baseline study of marine zooplankton in the Maheshkhali Channel. Further research, therefore, is recommended to identify zooplankton species through microscopic and molecular methods in the coastal waters of Bangladesh.

Behavioral toxicity of TDCPP in marine zooplankton: Evidence from feeding and swimming responses, molecular dynamics and metabolomics of rotifers

As new organic flame retardants, chlorinated organophosphate esters (Cl-OPEs) have high water solubility and structural similarity to organophosphate pesticides, posing risks to aquatic organisms. The potential neurotoxicity of Cl-OPEs has attracted attention, especially in marine invertebrates with a relatively simple nervous system. In this study, a marine rotifer with a cerebral ganglion, Brachionus plicatilis, was exposed to tris (1,3-dichloro-2-propyl) phosphate (TDCPP) (two environmental concentrations and one extreme level), and the changes in feeding and swimming behaviors and internal mechanism were explored. Exposure to 1.05 nM TDCPP did not change the filtration and ingestion rates of rotifers and average linear velocity. But 0.42 and 4.20 μM TDCPP inhibited these three parameters and reduced unsaturated fatty acid content, reproduction and population growth. All TDCPP test concentrations suppressed AChE activity, causing excessive accumulation of acetylcholine within rotifers, thereby disturbing the neural innervation of corona cilia. Molecular docking and molecular dynamics revealed that this inhibition was because TDCPP can bind to the catalytic active site of rotifer AChE through van der Waals forces and electrostatic interactions. TRP420 was the leading amino residue in the binding, and GLY207 contributed to a hydrogen bond. Nontargeted metabolomics using LC-MS and GC-MS identified differentially expressed metabolites in TDCPP treatments, mainly from lipid and lipid-like molecules, especially sphingolipids. TDCPP decreased ganglioside content but stimulated ceramide generation and the expression levels of 3 genes related to ceramide de novo synthesis. The mitochondrial membrane potential (MMP) and ATP content decreased, and the electron respiratory chain complex and TCA cycle were deactivated. An inhibitor of ceramide synthase, fumonisin, alleviated MMP and ATP, implying a critical role of ceramide in mitochondrial dysfunction. Thus, TDCPP exposure caused an energy supply deficit affecting ciliary movement and ultimately inhibiting rotifer behaviors. Overall, this study promotes the understanding of the neurotoxicity of Cl-OPEs in marine invertebrates.

Molecular insight into the binding properties of marine algogenic dissolved organic matter for polybrominated diphenyl ethers and their combined effect on marine zooplankton

Polybrominated diphenyl ethers (PBDEs) are widespread in marine ecosystems, despite the limits placed on several congeners, and pose a threat to marine organisms. Many coexisting factors, especially dissolved organic matter (DOM), affect the environmental behavior and ecological risk of PBDEs. Since blooms frequently occur in coastal waters, we used algogenic DOM (A-DOM) from the diatom Skeletonem costatum and examined the interaction of A-DOM with 2,2′,4,4′-tetrabromodiphenyl ether (BDE-47). Moreover, their combined effect on the rotifer Brachionus plicatilis was analyzed. During the stationary period, A-DOM had more proteins than polysaccharides, and 7 extracellular proteins were identified. A-DOM fluorescence was statically quenched by BDE-47, and amide, carbonyl, and hydroxyl groups in A-DOM were involved. Molecular docking analysis showed that all 5 selected proteins of A-DOM could spontaneously bind with BDE-47 and that hydrophobic interactions, van der Waals forces and pi-bond interactions existed. The reproductive damage, oxidative stress and inhibition of mitochondrial activity induced by BDE-47 in rotifers were relieved by A-DOM addition. Transcriptomic analysis further showed that A-DOM could activate energy metabolic pathways in rotifers and upregulate genes encoding metabolic detoxification proteins and DNA repair. Moreover, A-DOM alleviated the interference effect of BDE-47 on lysosomes, the extracellular matrix pathway and the calcium signaling system. Alcian blue staining and scanning electron microscopy showed that A-DOM aggregates were mainly stuck to the corona and cuticular surface of the rotifers; this mechanism, rather than a real increase in uptake, was the reason for enhanced bioconcentration. This study reveals the complex role of marine A-DOM in PBDEs bioavailability and enhances the knowledge related to risk assessments of PBDE-like contaminants in marine environments.

Fatty acid profiles reveal dietary variability of a large calanoid copepod Limnocalanus macrurus in the northern Baltic Sea

Eutrophication, climate-induced warming, and salinity fluctuations are altering the fatty acid profiles and the availability of essential polyunsaturated fatty acids (PUFAs) in marine zooplankton communities. Limnocalanus macrurus Sars G.O., 1863 is a large calanoid copepod inhabiting the low-salinity areas in the Baltic Sea, where it is a major source of docosahexaenoic acid (DHA) and eicosapentaenoic acid (EPA) to commercially important fish. L. macrurus is sensitive to warming, eutrophication and hypoxia. As an opportunistic feeder, it is capable of dietary shifts, which affects its fatty acid profiles. Although much studied in boreal lakes, there are only a few studies on the fatty acid profiles of the Baltic Sea populations. This study aimed to compare the fatty acid profiles of L. macrurus in three basins of the Baltic Sea, in relation to the community fatty acids and environmental variables. We collected samples of L. macrurus and filtered plankton community for gas chromatographic fatty acid analyses in August 2021 on R/V Aranda. The nutritional quality of L. macrurus to consumers was lower in the Gulf of Finland (GoF) compared to the Gulf of Bothnia, indicated by the low levels of DHA and EPA, as well as the low n-3/n-6 ratio of PUFAs. The lower ratio of 18:1n-7 to 18:1n-9 implied higher degree of omnivory in GoF. In contrast, a diatom marker 16:1n-7 had high proportion in the Bothnian Bay. High temperatures in GoF may have restricted feeding in the upper water column, possibly forcing a shift towards cyanobacteria or seston-based diet, as interpreted from a high proportion of 18:2n-6 and 18:3n-3. We conclude that the ability of L. macrurus to utilize multiple food sources increases its resilience to environmental change, while the consequences on the nutritional quality may have further cascading effects on the food webs.

Research trend on marine zooplankton in Indonesian Waters: A systematic review

Research on marine zooplankton in Indonesian waters was conducted in the 19th century. However, there is limited information regarding evaluating the research trends and notable scientists involved in this area. This study aims to know the research trend on marine zooplankton research in Indonesian waters. This study analyzed publications that were included in the Scopus database from 1900 to 2021. A systematic review was undertaken using a three-stage procedure including identification, screening, and final following PRISMA chart. The final list was analysed in terms of topics of research, keywoards, location, time of publication, and authorship.. We found that the research trend was changed. During the early stages of the study, there was a prevalent focus on studying both taxonomy copepods and parasite Copepods. However, in recent times, there has been a shift towards favoring the plankton ecology. Despite being a hotspot for marine biodiversity, Indonesia only had a few taxonomists, therefore it was overlooked when marine zooplankton was discovered in its seas. We also found that most of the new finding species come from copepods whereas the other taxa of zooplankton remain undescribed or misidentification. Additionally, young scientists pay less attention to the study of taxonomy. To address these problems, priority is given to intensive training for early-career scientists. Comprehensive approaches using morphological traits and genetic tools will solve this issue.

MetaZooGene Atlas and Database: Reference Sequences for Marine Ecosystems.

The MetaZooGene Atlas and Database (MZGdb; https://metazoogene.org/mzgdb/ ) is an open-access data and metadata portal synchronized with the NCBI GenBank and BOLD data repositories. The MZGdb includes sequences for genes used for the classification and identification of marine organisms based on DNA barcoding and metabarcoding. The focus of the MZGdb is biodiversity of marine ecosystems, including phytoplankton and microbes, zooplankton and invertebrates, fish, and other marine vertebrates (pinnipeds, cetaceans, and sea turtles). DNA sequences currently included are mitochondrial cytochrome oxidase I (COI), 12S, and 16S rRNA, and nuclear 18S and 28S rRNA. The MZGdb provides data and mapping tools for assembling and downloading compilations of reference sequence data that are specific to selected genes, taxonomic groups, and/or ocean regions. An additional feature of the MZGdb is the Atlas which summarizes data coverage and proportional completeness based on statistics of species with available sequences versus species commonly found in each ocean region.This chapter is a collaborative effort of the Scientific Committee for Ocean Research (SCOR) Working Group WG157: MetaZooGene: Toward a new global view of marine zooplankton biodiversity based on DNA metabarcoding and reference DNA sequence databases ( https://metazoogene.org ).