Revealing the profound influence of diapause on gene expression: Insights from the annual transcriptome of the copepod Calanus finmarchicus.

Annual rhythms are observed in living organisms with numerous ecological implications. In the zooplanktonic copepod Calanus finmarchicus, such rhythms are crucial regarding its phenology, body lipid accumulation, and global carbon storage. Climate change drives annual biological rhythms out of phase with the prevailing environmental conditions with yet unknown but potentially catastrophic consequences. However, the molecular dynamics underlying phenology are still poorly described. In a rhythmic analysis of C. finmarchicus annual gene expression, results reveal that more than 90% of the transcriptome shows significant annual rhythms, with abrupt and dramatic upheaval between the active and diapause life cycle states. This work explores the implication of the circadian clock in the annual timing, which may control epigenetic mechanisms to profoundly modulate gene expression in response to calendar time. Results also suggest an increased light sensitivity during diapause that would ensure the photoperiodic entrainment of the endogenous annual clock.

Contribution of combined stressors on density and gene expression dynamics of the copepod Temora longicornis in the North Sea.

The impact of multiple environmental and anthropogenic stressors on the marine environment remains poorly understood. Therefore, we studied the contribution of environmental variables to the densities and gene expression of the dominant zooplankton species in the Belgian part of the North Sea, the calanoid copepod Temora longicornis. We observed a reduced density of copepods, which were also smaller in size, in samples taken from nearshore locations when compared to those obtained from offshore stations. To assess the factors influencing the population dynamics of this species, we applied generalised additive models. These models allowed us to quantify the relative contribution of temperature, nutrient levels, salinity, turbidity, concentrations of photosynthetic pigments, as well as chemical pollutants such as polychlorinated biphenyls and polycyclic aromatic hydrocarbons (PAHs), on copepod density. Temperature and Secchi depth, a proxy for turbidity, were the most important environmental variables predicting the densities of T. longicornis, followed by summed PAH and chlorophyll concentrations. Analysing gene expression in field-collected adults, we observed significant variation in metabolic and stress-response genes. Temperature correlated significantly with genes involved in proteolytic activities, and encoding heat shock proteins. Yet, concentrations of anthropogenic chemicals did not induce significant differences in the gene expression of genes involved in the copepod's fatty acid metabolism or well-known stress-related genes, such as glutathione transferases or cytochrome P450. Our study highlights the potential of gene expression biomonitoring and underscores the significance of a changing environment in future studies.

Highly structured populations of copepods at risk to deep-sea mining: Integration of genomic data with demogenetic and biophysical modelling.

Copepoda is the most abundant taxon in deep-sea hydrothermal vents, where hard substrate is available. Despite the increasing interest in seafloor massive sulphides exploitation, there have been no population genomic studies conducted on vent meiofauna, which are known to contribute over 50% to metazoan biodiversity at vents. To bridge this knowledge gap, restriction-site-associated DNA sequencing, specifically 2b-RADseq, was used to retrieve thousands of genome-wide single-nucleotide polymorphisms (SNPs) from abundant populations of the vent-obligate copepod Stygiopontius lauensis from the Lau Basin. SNPs were used to investigate population structure, demographic histories and genotype-environment associations at a basin scale. Genetic analyses also helped to evaluate the suitability of tailored larval dispersal models and the parameterization of life-history traits that better fit the population patterns observed in the genomic dataset for the target organism. Highly structured populations were observed on both spatial and temporal scales, with divergence of populations between the north, mid, and south of the basin estimated to have occurred after the creation of the major transform fault dividing the Australian and the Niuafo'ou tectonic plate (350 kya), with relatively recent secondary contact events (<20 kya). Larval dispersal models were able to predict the high levels of structure and the highly asymmetric northward low-level gene flow observed in the genomic data. These results differ from most studies conducted on megafauna in the region, elucidating the need to incorporate smaller size when considering site prospecting for deep-sea exploitation of seafloor massive sulphides, and the creation of area-based management tools to protect areas at risk of local extinction, should mining occur.

The fluid dynamics of barnacle feeding.

Sessile barnacles feed by sweeping their basket-like cirral fan through the water, intercepting suspended prey. A primary component of the diet of adult barnacles is copepods that are sensitive to fluid disturbances and capable of escaping. How do barnacles manage to capture copepods despite the fluid disturbances they generate? We examined this question by describing the feeding current architecture of 1 cm sized Balanus crenatus using particle image velocimetry, and by studying the trajectories of captured copepods and the escapes of evading copepods. We found that barnacles produce a feeding current that arrives both from behind and the sides of the barnacle. The flow from the sides represents quiescent corridors of low fluid deformation and uninterrupted by the beating cirral fan. Potential prey arriving from behind are likely to encounter the cirral fan and, hence, capture here is highly unlikely. Accordingly, most captured copepods arrived through the quiet corridors, while most copepods arriving from behind managed to escape. Thus, it is the unique feeding flow architecture that allows feeding on evasive prey. We used the Landau-Squire jet as a simple model of the feeding current. For the Reynolds number of our experiments, the model reproduces the main features of the feeding current, including the lateral feeding corridors. Furthermore, the model suggests that smaller barnacle specimens, operating at lower Reynolds numbers, will produce a fore-aft symmetric feeding current without the lateral corridors. This suggests an ontogenetic diet shift from non-evasive prey to inclusion of evasive prey as the barnacle grows.

Local variability of Arctic mesozooplankton biomass and production: A case summer study.

The Barents Sea is a highly productive ecosystem within the Arctic Ocean. The overall biological productivity in this region relies heavily on the secondary mesozooplankton production (MZP). Previous research has primarily focused on mesozooplankton abundance (MZA) and biomass (MZB), lacking a comprehensive analysis of the environmental factors that influence MZP in the Arctic marine environment. The primary objective of this study was to examine the key factors responsible for the spatial variability in the community structure and MZP during the summer season. Data were collected from 52 stations in the central Barents Sea, specifically during post-bloom conditions in 2015. Through cluster analysis, two distinct groups of stations were identified, differing in terms of mesozooplankton taxa abundance. Copepods were found to dominate the mesozooplankton assemblages, comprising 89% of total MZA, 83% of MZB, and 68% of MZP. The biomass stocks in the study area varied from 8 to 102 mg dry mass (DM) m-3, with an average of 44 mg DM m-3. MZP rates ranged from 0.34 to 2.33 mg DM m-3 day-1, with an average of 1.16 mg DM m-3 day-1. The highest MZB and MZP values were observed at frontal zones that separated relatively warm and cold waters. Through redundancy analysis, it was determined that the primary environmental factors affected the distribution of mesozooplankton were longitude, latitude, and sampling depth. Temperature, salinity, and chlorophyll a concentration were found to have a less significant impact. These findings emphasize the importance of oceanographic conditions as the main predictors of mesozooplankton distribution during the summer season in the Arctic marine environment. This study highlights the essential role of environmental forcing in determining the productivity of Arctic marine zooplankton. Given the ongoing climatic changes, the results of this report can serve as a valuable tool for monitoring pelagic ecosystems in the Arctic.

Integrative taxonomy of metazoan parasites of the bluntnose sixgill shark Hexanchus griseus (Bonnaterre, 1788) in the Mediterranean Sea, with the resurrection of Grillotia acanthoscolex Rees, 1944 (Cestoda: Trypanorhyncha).

Appropriate diagnoses of parasites of apex marine predators are crucial to understand their biodiversity, host specificity, biogeography, and life cycles. Such diagnoses are also informative of ecological and biological characteristics of both host and environment in which the hosts and their parasites live. We here (i) investigate the parasite fauna of a bluntnose sixgill shark Hexanchus griseus (Bonnaterre, 1788) obtained from the Gulf of Naples (Tyrrhenian Sea), (ii) characterize molecularly all its metazoan parasites, and (iii) resurrect and report the main morphological features and phylogenetic position of Grillotia acanthoscolex, a cestode species previously synonymized with Grillotia adenoplusia. A rich parasite fauna represented by eight different taxa was found, including two monogeneans (Protocotyle grisea and Protocotyle taschenbergi), one digenean (Otodistomum veliporum), four cestodes (Crossobothrium dohrnii, Clistobothrium sp., G. acanthoscolex, and G. adenoplusia), and one copepod (Protodactylina pamelae). Sequencing of these samples accounts for an important molecular baseline to widen the knowledge on the parasitic fauna of bluntnose sixgill sharks worldwide and to reconstruct their correct food chains. The bluntnose sixgill shark was found to be a definitive host for all endoparasites found here, confirming that it occupies an apex trophic level in the Mediterranean Sea. The taxa composition of the trophic parasite fauna confirms that the bluntnose sixgill shark mostly feeds on teleost fish species. However, the occurrence of two phillobothrid cestodes (C. dohrnii and Clistobothrium sp.) suggests that it also feeds on squids. Finally, we emphasize the importance of using integrative taxonomic approaches in the study of parasites from definitive and intermediate hosts to elucidate biology and ecology of taxa generally understudied in the Mediterranean Sea.

Functional characterization reveals a diverse array of metazoan fatty acid biosynthesis genes.

Long-chain (≥C20 ) polyunsaturated fatty acids (LC-PUFAs) are physiologically important fatty acids for most animals, including humans. Although most LC-PUFA production occurs in aquatic primary producers such as microalgae, recent research indicates the ability of certain groups of (mainly marine) invertebrates for endogenous LC-PUFA biosynthesis and/or bioconversion from dietary precursors. The genetic pathways for and mechanisms behind LC-PUFA biosynthesis remain unknown in many invertebrates to date, especially in non-model species. However, the numerous genomic and transcriptomic resources currently available can contribute to our knowledge of the LC-PUFA biosynthetic capabilities of metazoans. Within our previously generated transcriptome of the benthic harpacticoid copepod Platychelipus littoralis, we detected expression of one methyl-end desaturase, one front-end desaturase, and seven elongases, key enzymes responsible for LC-PUFA biosynthesis. To demonstrate their functionality, we characterized eight of them using heterologous expression in yeast. The P. littoralis methyl-end desaturase has Δ15/17/19 desaturation activity, enabling biosynthesis of α-linolenic acid, eicosapentaenoic acid and docosahexaenoic acid (DHA) from 18:2 n-6, 20:4 n-6 and 22:5 n-6, respectively. Its front-end desaturase has Δ4 desaturation activity from 22:5 n-3 to DHA, implying that P. littoralis has multiple pathways to produce this physiologically important fatty acid. All studied P. littoralis elongases possess varying degrees of elongation activity for saturated and unsaturated fatty acids, producing aliphatic hydrocarbon chains with lengths of up to 30 carbons. Our investigation revealed a functionally diverse range of fatty acid biosynthesis genes in copepods, which highlights the need to scrutinize the role that primary consumers could perform in providing essential nutrients to upper trophic levels.

The recovery of crustacean zooplankton from acidification depends on lake type.

Acidification has harmed freshwater ecosystems in Northern Europe since the early 1900s. Stricter regulations aimed at decreasing acidic emissions have improved surface-water chemistry since the late 1980s but the recovery of biotic communities has not been consistent. Generally, the recovery of flora and fauna has been documented only for a few lakes or regions and large-scale assessments of long-term dynamics of biotic communities due to improved water quality are still lacking. This study investigates a large biomonitoring dataset of pelagic and littoral crustacean zooplankton (Cladocera and Copepoda) from 142 acid-sensitive lakes in Norway spanning 24 years (1997-2020). The aims were to assess the changes in zooplankton communities through time, compare patterns of changes across lake types (defined based on calcium and humic content), and identify correlations between abiotic and biological variables. Our results indicate chemical and biological recovery after acidification, as shown by a general increase in pH, acid neutralizing capacity, changes in community composition and increases in the total number of species, number of acid-sensitive species and functional richness through time. However, the zooplankton responses differ across lake types. This indicates that the concentration of calcium (or alkalinity) and total organic carbon (or humic substances) are important factors for the recovery. Therefore, assessment methods and management tools should be adapted to the diverse lake types. Long-term monitoring of freshwater ecosystems is needed to fully comprehend the recovery dynamics of biotic communities from acidification.

Is Zooplankton an Entry Point of Microplastics into the Marine Food Web?

Microplastics (MPs) overlap in size with phytoplankton and can be ingested by zooplankton, transferring them to higher trophic levels. Copepods are the most abundant metazoans among zooplankton and the main link between primary producers and higher trophic levels. Ingestion of MPs has been investigated in the laboratory, but we still know little about the ingestion of MPs by zooplankton in the natural environment. In this study, we determined the concentration and characteristics of MPs down to 10 µm in zooplankton samples, sorted calanoid copepods, and fecal pellets collected in the Kattegat/Skagerrak Sea (Denmark). We found a median concentration of 1.7 × 10-3 MPs ind-1 in the zooplankton samples, 2.9 × 10-3 MPs ind-1 in the sorted-copepods, and 3 × 10-3 MPs per fecal pellet. Most MPs in the zooplankton samples and fecal pellets were fragments smaller than 100 µm, whereas fibers dominated in the sorted copepods. Based on the collected data, we estimated a MP budget for the surface layer (0-18 m), where copepods contained only 3% of the MPs in the water, while 5% of the MPs were packed in fecal pellets. However, the number of MPs exported daily to the pycnocline via fecal pellets was estimated to be 1.4% of the total MPs in the surface layer. Our results indicate that zooplankton are an entry point of small MPs in the food web, but the number of MPs in zooplankton and their fecal pellets was low compared with the number of MPs found in the water column and the occurrence and/or ingestion of MPs reported for nekton. This suggests a low risk of MP transferring to higher trophic levels through zooplankton and a quantitatively low, but ecologically relevant, contribution of fecal pellets to the vertical exportation of MPs in the ocean.

The Behavior of Planktonic Copepods Minimizes the Entry of Microplastics in Marine Food Webs.

The entry of microplastics (MPs) into marine food webs is a major environmental concern. We investigated how the behavior of planktonic copepods influences the risk of MPs to enter marine food webs by applying a trait-based approach and by combining experiments (bottle incubations and video observations) with biogeographical analyses. We aimed to evaluate which type of feeding behavior is most risky in terms of MP ingestion and which marine geographical areas are more susceptible to MP ingestion by planktonic copepods. We used different species as models of the main foraging behaviors in planktonic copepods: feeding-current, cruising, ambush, and mixed behavior feeding. All behaviors showed a similarly low risk of MP ingestion, up to 1 order of magnitude lower than for similar-sized microalgae. We did not observe any influence of the prey type or MP size (8 and 20 µm) on MP ingestion for any of the behaviors. By mapping the global distribution of feeding behaviors, we showed that feeding-current feeding is the most common behavior, but the risk of MP ingestion remains equally low across the global ocean, independently of the predominant behavior. Overall, our results suggest a low risk of MP ingestion by planktonic copepods and therefore a minimal risk of trophic transfer of MPs via marine pelagic copepods in marine ecosystems.

Tracking Nano- and Microplastics Accumulation and Egestion in a Marine Copepod by Novel Fluorescent AIEgens: Kinetic Modeling of the Rhythm Behavior.

Nano- and microplastics (NMPs) are now prevalent in the marine environment. This study quantified the uptake and depuration kinetics of spherical polystyrene NMPs of different particle sizes (200 nm/30 µm) and functional groups (-NH2/-COOH) in a temperate calanoid copepod Calanus sinicus (C. sinicus), which exhibited rhythmic feeding patterns in natural environments. Aggregated-induced emission (AIE) fluorescent probes were employed to track and quantify the kinetics of NMPs with excellent photostability and biocompatibility. The results showed that C. sinicus consumed all NMPs types, with preference of NMPs to small size and amino group. Increased diatom concentrations also inhibited the bioaccumulation of NMPs. Influenced by rhythmic behavior, the bioaccumulation of NMPs by C. sinicus was nonstationary during the 6 h uptake phase. After 1-3 h of rapid uptake, the body burden peaked and then slowly declined. During the 3 h depuration phase, C. sinicus rapidly and efficiently removed NMPs with a mean half-life of only 0.23 h. To further quantify the body burden of C. sinicus under the influence of rhythmic feeding behavior, a biokinetic model was established, and the Markov chain Monte Carlo method was used to estimate the parameter distribution. Our results highlighted that copepods exhibited unique rhythmic feeding behavior under environmentally relevant concentrations of NMPs exposure, which may influence the bioaccumulation, trophic transfer, and environmental fate of NMPs.

Morphological and mitochondrial approaches of Hatschekia sargi (Copepoda: Hatschekiidae) as a parasite of Epinephelus chlorostigma.

Little information is available until now about the copepods infecting different fish species. Therefore, this study aimed to provide light on siphonostomatoids infecting Epinephelus chlorostigma. Twenty fish specimens were taken from the Red Sea coast (Jeddah, Saudi Arabia), and ectoparasitic copepods were investigated. Light microscopy and molecular tools were used to examine copepods isolated from fish. Parasitological indexes were calculated and showed that 60% of the examined fish were infected with a mean intensity of 12 parasite/fish. Morphological examination revealed that this copepod species is characterized by all unique features of the genus Hatschekia with special reference to Hatschekia sargi. The taxonomic position of the recovered species in the Hatschekiidae family within Siphonostomatoida was confirmed using phylogenetic analysis based on partial mitochondrial cytochrome c oxidase subunit I (mt COI) gene sequences. The mt COI gene query revealed that the recovered Hatschekia species is closely related to Hatschekia maculatus (gb| JQ664005.1). This study discovers a new host for Hatschekia species isolated from Saudi Arabia and conducts the first genomic investigation of the mt COI gene.

An Experimental Test of Adaptive Introgression in Locally Adapted Populations of Splash Pool Copepods.

As species struggle to keep pace with the rapidly warming climate, adaptive introgression of beneficial alleles from closely related species or populations provides a possible avenue for rapid adaptation. We investigate the potential for adaptive introgression in the copepod, Tigriopus californicus, by hybridizing two populations with divergent heat tolerance limits. We subjected hybrids to strong heat selection for 15 generations followed by whole-genome resequencing. Utilizing a hybridize evolve and resequence (HER) technique, we can identify loci responding to heat selection via a change in allele frequency. We successfully increased the heat tolerance (measured as LT50) in selected lines, which was coupled with higher frequencies of alleles from the southern (heat tolerant) population. These repeatable changes in allele frequencies occurred on all 12 chromosomes across all independent selected lines, providing evidence that heat tolerance is polygenic. These loci contained genes with lower protein-coding sequence divergence than the genome-wide average, indicating that these loci are highly conserved between the two populations. In addition, these loci were enriched in genes that changed expression patterns between selected and control lines in response to a nonlethal heat shock. Therefore, we hypothesize that the mechanism of heat tolerance divergence is explained by differential gene expression of highly conserved genes. The HER approach offers a unique solution to identifying genetic variants contributing to polygenic traits, especially variants that might be missed through other population genomic approaches.

Costs and benefits of predator-induced defence in a toxic diatom.

Phytoplankton employ a variety of defence mechanisms against predation, including production of toxins. Domoic acid (DA) production by the diatom Pseudo-nitzschia spp. is induced by the presence of predators and is considered to provide defence benefits, but the evidence is circumstantial. We exposed eight different strains of P. seriata to chemical cues from copepods and examined the costs and the benefits of toxin production. The magnitude of the induced toxin response was highly variable among strains, while the costs in terms of growth reduction per DA cell quota were similar and the trade-off thus consistent. We found two components of the defence in induced cells: (i) a 'private good' in terms of elevated rejection of captured cells and (ii) a 'public good' facilitated by a reduction in copepod feeding activity. Induced cells were more frequently rejected by copepods and rejections were directly correlated with DA cell quota and independent of access to other food items. By contrast, the public-good effect was diminished by the presence of alternative prey suggesting that it does not play a major role in bloom formation and that its evolution is closely associated with the grazing-deterrent private good.

Identification and characterization of homeobox gene clusters in harpacticoid and calanoid copepods.

In this study, we have identified the entire complement of typical homeobox (Hox) genes (Lab, Pb, Dfd, Scr, Antp, Ubx, Abd-A, and Abd-B) in harpacticoid and calanoid copepods and compared them with the cyclopoid copepod Paracyclopina nana. The harpacticoid copepods Tigriopus japonicus and Tigriopus kingsejongensis have seven Hox genes (Lab, Dfd, Scr, Antp, Ubx, Abd-A, and Abd-B) and the Pb and Ftz genes are also present in the cyclopoid copepod P. nana. In the Hox gene cluster of the calanoid copepod Eurytemora affinis, all the Hox genes were present linearly in the genome but the Antp gene was duplicated. Of the three representative copepods, the P. nana Hox gene cluster was the most compact due to its small genome size. The Hox gene expression profile patterns in the three representative copepods were stage-specific. The Lab, Dfd, Scr, Pb, Ftz, and Hox3 genes showed a high expression in early developmental stages but Antp, Ubx, Abd-A, and Abd-B genes were mostly expressed in later developmental stages, implying that these Hox genes may be closely associated with the development of segment identity during early development.

Changes in Fecal Pellet Microbiome of the Cold-Adapted Antarctic Copepod Tigriopus kingsejongensis at Different Temperatures and Developmental Stages.

Tigriopus kingsejongensis, a copepod species reported from the King Sejong Station, Antarctica, serves as a valuable food resource in ecosystems. We cultured T. kingsejongensis at three different temperatures (2 °C, 8 °C, and 15 °C) in a laboratory to observe the changes in its fecal pellet microbiome depending on the cultivation temperatures and developmental stages. We observed that the fecal pellet microbiome of the copepod changed with temperature: a lower microbial diversity, higher abundance of the aquatic bacterium Vibrio, and lower abundance of the psychrophilic bacterium Colwellia were noted at higher temperatures. In addition, the fecal pellet microbiome of the copepod changed according to the developmental stage: a lower microbial diversity was noted in egg-attached copepods than in nauplii at 8 °C. We further analyzed three shotgun metagenomes from the fecal pellet samples of T. kingsejongensis at different temperatures and obtained 44 metagenome-assembled genomes (MAGs). We noted that MAGs of V. splendidus D contained glycosyl hydrolases (GHs) encoding chitinases and virulence factors at a higher relative abundance at 15 °C than at lower temperatures. These results indicate that increasing temperature affects the fecal pellet microbiome and the development of copepods. The findings are helpful to understand the changes in cold-adapted copepods and the effect of temperature on their growth.

Unpalatable Plastic: Efficient Taste Discrimination of Microplastics in Planktonic Copepods.

Planktonic copepods are the most abundant animals in the ocean and key players in global biochemical processes. Recent modeling suggests that zooplankton ingestion of microplastics (MPs) can disrupt the biological carbon pump and accelerate a global loss of oceanic oxygen. Here we investigate the behavioral responses and ingestion rates of a model feeding-current generating copepod when exposed to microplastics of different characteristics by small-scale video observations and bottle incubations. We found that copepods rejected 80% of the microplastics after touching them with their mouth parts, in essence exhibiting a kind of taste discrimination. High rejection rates of microplastics were independent of polymer type, shape, presence of biofilms, or sorbed pollutant (pyrene), indicating that microplastics are unpalatable for feeding-current feeding copepods and that post-capture taste discrimination is a main sensorial mechanism in the rejection of microplastics. In an ecological context, taking into account the behaviors of planktonic copepods and the concentrations of microplastics found in marine waters, our results suggest a low risk of microplastic ingestion by zooplankton and a low impact of microplastics on the vertical exportation of fecal pellets.

Chemosensory-Related Genes in Marine Copepods.

Living organisms deeply rely on the acquisition of chemical signals in any aspect of their life, from searching for food, mating and defending themselves from stressors. Copepods, the most abundant and ubiquitous metazoans on Earth, possess diversified and highly specified chemoreceptive structures along their body. The detection of chemical stimuli activates specific pathways, although this process has so far been analyzed only on a relatively limited number of species. Here, in silico mining of 18 publicly available transcriptomes is performed to delve into the copepod chemosensory genes, improving current knowledge on the diversity of this multigene family and on possible physiological mechanisms involved in the detection and analysis of chemical cues. Our study identifies the presence of ionotropic receptors, chemosensory proteins and gustatory receptors in copepods belonging to the Calanoida, Cyclopoida and Harpacticoida orders. We also confirm the absence in these copepods of odorant receptors and odorant-binding proteins agreeing with their insect specificity. Copepods have evolved several mechanisms to survive in the harsh marine environment such as producing proteins to respond to external stimulii. Overall, the results of our study open new possibilities for the use of the chemosensory genes as biomarkers in chemical ecology studies on copepods and possibly also in other marine holozooplankters.

Copepod carcasses in the western Bay of Bengal and associated ecology.

Although a portion of all copepods in the natural environment are physiologically inactive or dead, there is little data on this aspect in Indian seas. Using the classical neutral red live staining technique, the dead percentage of copepods in the western Bay of Bengal (BoB) is determined for the first time in this study, which takes into account hydrography and zooplankton samples collected on April 2015 (Spring Intermonsoon [SIM]) from 6 inshore and 8 offshore locations in the western BoB. The offshore parts of the research area were infested with a persistent and extensive warm-core eddy/gyre during the sampling time, as demonstrated by satellite data of sea surface temperature, mean sea level anomaly, and surface currents. As a result, the mixed layer depth in the offshore zone was higher, whereas nutrients, chlorophyll a, and turbidity were significantly lower than inshore. Copepods dominated the zooplankton community (> 80%), with abundances ranging from 112 to 2580 No m-3 in the study area. Calanoids (particularly Paracalanidae) dominated nearshore waters (35-65%), whereas cyclopoids (notably Corycaeidae and Oncaeidae) dominated offshore (49-71%). There were 48 species of copepods identified in total, and all of them had dead individuals (carcasses). Offshore waters had a larger percentage of copepod carcasses (27-39%) than inshore seas (18-28%). The larger percentage of carcasses is possibly related to a higher top-down feeding pressure on cyclopoids in offshore waters due to extreme oligotrophy and the dominance of the microbial food web.

Spatial and temporal shift in the factors affecting the population dynamics of Calanus copepods in the North Sea.

The swap in abundance between two Calanus species in the North Sea during the 1980s constitutes a quintessential example of regime shift, with important ecosystemic and economic repercussions because these copepods constitute a major component of the diet of larval and juvenile cods. It is hypothesized that this transition was driven by gradual changes in primary productivity, the North Atlantic Oscillation (NAO) and sea surface temperatures (SST), and yet how these factors contribute to the population dynamics of these two species and the overall regime shift remains unclear. Here, we combine a highly resolved and spatially structured longitudinal dataset with population dynamics theory-based models to obtain a thorough and more detailed description of populations' responses to the regime shift observed in the North Sea. Our analyses highlight that this transition exhibits a clear spatial structure and involved a decoupling between the dynamics of Calanus finmarchicus and the NAO in western regions and between Calanus helgolandicus and SST in the eastern regions of the North Sea. Consequently, the observed switch in abundance between these species reflects the interaction between species-specific attributes, a well-defined spatial structure with a marked east-west axis and a decoupling between the ecological drivers and Calanus population dynamics following the shift. Succinctly, we suspect that higher water temperatures have favored C. helgolandicus and resulted in restrictive conditions for C. finmarchicus, eventually overshadowing the effects of NAO detected in historical records. Overall, our study illustrates how population dynamics theory can be successfully employed to disentangle the complex and multifactorial nature of a regime shift in response to gradually changing environmental conditions.