Characterization of the complete mitochondrial genome of Desmaulus extinctorium (Littorinimorpha, Calyptraeoidea, Calyptraeidae) and molecular phylogeny of Littorinimorpha.

For the purpose of determining the placement of Calyptraeidae within the Littorinimorpha, we hereby furnish a thorough analysis of the mitochondrial genome (mitogenome) sequence of Desmaulus extinctorium. This mitogenome spans 16,605 base pairs and encompasses the entire set of 37 genes, including 13 PCGs, 22 tRNAs and two rRNAs, with an evident AT bias. Notably, tRNASer1 and tRNASer2 lack dihydrouracil (DHU) arms, resulting in an inability to form a secondary structure. Similarly, tRNAAla lacks a TΨC arm, rendering it incapable of forming a secondary structure. In contrast, the remaining tRNAs demonstrate a characteristic secondary structure reminiscent of a cloverleaf. A comparison with ancestral gastropods reveals distinct differences in three gene clusters (or genes), encompassing 15 tRNAs and eight PCGs. Notably, inversions and translocations represent the major types of rearrangements observed in D. extinctorium. Phylogenetic analysis demonstrates robust support for a monophyletic grouping of all Littorinimorpha species, with D. extinctorium representing a distinct Calyptraeoidea clade. In summary, this investigation provides the first complete mitochondrial dataset for a species of the Calyptraeidae, thus providing novel insights into the phylogenetic relationships within the Littorinimorpha.

Analysis of Genetic Diversity and Structure of Eight Populations of Nerita yoldii along the Coast of China Based on Mitochondrial COI Gene.

Nerita yoldii is a euryhaline species commonly found in the intertidal zone. To investigate the genetic diversity of 233 N. yoldii individuals from eight locations along the coast of China, we utilized the mitochondrial COI gene as a molecular marker. A total of 34 haplotypes were detected, exhibiting a mean haplotype diversity (Hd) of 0.5915 and a mean nucleotide diversity (Pi) of 0.0025, indicating high levels of genetic diversity among all populations. An analysis of molecular variance (AMOVA) indicated that the primary source of genetic variation occurs within populations. In addition, neutral tests and mismatch analyses suggested that N. yoldii populations may have experienced bottleneck events. Moderate genetic differentiation was observed between Xiapu and other populations, excluding the Taizhou population, and may be attributed to the ocean currents. Intensively studying the genetic variation and population structure of N. yoldii populations contributes to understanding the current population genetics of N. yoldii in the coastal regions of China. This not only provides a reference for the study of other organisms in the same region but also lays the foundation for the systematic evolution of the Neritidae family.

Environmental enrichment improves behaviors rather than the growth and physiology of rock bream Oplegnathus fasciatus.

Environmental enrichment has the potential to improve the welfare and post-release survival of hatchery fish stocked for conservation purposes. However, the effectiveness of environmental enrichment is partly dependent on the fish species, life stage, and specific enrichment structure used. To enhance the effectiveness of environmental enrichment, it is crucial to focus on characteristic differences in enrichment structures, such as type and level. This study investigated how differences in enrichment type and level affected physiological and behavioral aspects of the welfare of pre-release juvenile rock bream Oplegnathus fasciatus by evaluating growth performance, basal and stressed cortisol levels, antioxidant enzyme activities, and exploratory behaviors regarding anxiety and flexibility. Fish were reared for 4 weeks in different enrichment treatments: barren, low-level cover structure, high-level cover structure, low-level interference structure (LI), and high-level interference structure (HI). The results revealed that fish reared with the LI treatment showed less anxiety and greater flexibility with respect to exploratory behaviors, without oxidative damage being detected. Despite exhibiting less anxiety as well, fish reared in the HI treatment had oxidative damage, indicated by lower superoxide dismutase activity, compared to those in the barren treatment. In addition, none of these enrichment structures enhanced growth performance or mitigate chronic and acute stress responses. Overall, the low-level interference structure may be more favorable in promoting the behavioral welfare of the fish. Application of this type and level of enrichment may increase the survival of the hatchery fish after release, which is critical to stocking success.

Seasonal variations in the diversity and benthic community structure of subtidal artificial oyster reefs adjacent to the Luanhe River Estuary, Bohai Sea.

Artificial oyster reefs provide important spawning and nursery grounds for a variety of fishes and large mobile crustaceans. Between July 2016 and May 2017, seasonal surveys of species composition and community structure were performed in the artificial oyster reef area and control area adjacent to the Luanhe River Estuary in China. During the survey year, 56 species belonging to 50 genera, 45 families, and 19 orders were recorded. The dominant economically important fish and mobile crustaceans were Hexagrammos otakii, Pholis fangi, Sebastes schlegelii, Charybdis japonica, and Oratosquilla oratoria. Resident fishes belonged to the Cynoglossidae, Paralichthyidae, Pleuronectidae, and Gobiidae families. Seasonally important fish species included Lateolabrax japonicus, Konosirus punctatus, Thryssa kammalensis, Hexagrammos agrammus, and Acanthopagrus schlegelii. The ranges of H' values among stations were 1.18-2.16, 0.65-1.75, 1.18-2.06, and 0.62-1.92 in spring, summer, autumn, and winter, respectively. The benthic organisms present in the community of artificial oyster reef areas can be classified into groups according to month and season. The abundance biomass curves showed that the oyster reef area in spring, autumn, and winter experienced low disturbance, whereas the community structure in summer was subject to large variations from external disturbance. We also found that as the age of the oyster reefs increased, the percentage of oysters in the low shell height group (< 40 mm) decreased. The oyster density was 324 ind/m2 for the reef created in 2016, 724 ind/m2 for the reef created in 2015, and 364 ind/m2 for the reef created in 2013. These findings can be used to develop suitable management strategies for the sustainable maintenance of artificial oyster reef ecosystems.

Comparison of Mitochondrial Genome Sequences between Two Palaemon Species of the Family Palaemonidae (Decapoda: Caridea): Gene Rearrangement and Phylogenetic Implications.

To further understand the origin and evolution of Palaemonidae (Decapoda: Caridea), we determined the mitochondrial genome sequence of Palaemon macrodactylus and Palaemon tenuidactylus. The entire mitochondrial genome sequences of these two Palaemon species encompassed 37 typical genes, including 13 protein-coding genes (PCGs), 2 ribosomal RNA genes (rRNAs), and 22 transfer RNA genes (tRNAs), and a control region (CR). The lengths of their mitochondrial genomes were 15,744 bp (P. macrodactylus) and 15,735 bp (P. tenuidactylus), respectively. We analyzed their genomic features and structural functions. In comparison with the ancestral Decapoda, these two newly sequenced Palaemon species exhibited a translocation event, where the gene order was trnK-trnD instead of trnD-trnK. Based on phylogenetic analysis constructed from 13 PCGs, the 12 families from Caridea can be divided into four major clades. Furthermore, it was revealed that Alpheidae and Palaemonidae formed sister groups, supporting the monophyly of various families within Caridea. These findings highlight the significant gene rearrangements within Palaemonidae and provide valuable evidence for the phylogenetic relationships within Caridea.

The Complete Mitochondrial Genome of Mytilisepta virgata (Mollusca: Bivalvia), Novel Gene Rearrangements, and the Phylogenetic Relationships of Mytilidae.

The circular mitochondrial genome of Mytilisepta virgata spans 14,713 bp, which contains 13 protein-coding genes (PCGs), 2 ribosomal RNA genes, and 22 transfer RNA genes. Analysis of the 13 PCGs reveals that the mitochondrial gene arrangement of Mytilisepta is relatively conserved at the genus level. The location of the atp8 gene in Mytilisepta keenae differs from that of other species. However, compared with the putative molluscan ancestral gene order, M. virgata exhibits a high level of rearrangement. We constructed phylogenetic trees based on concatenated 12 PCGs from Mytilidae. As a result, we found that M. virgata is in the same clade as other Mytilisepta spp. The result of estimated divergence times revealed that M. virgata and M. keenae diverged around the early Paleogene period, although the oldest Mytilisepta fossil was from the late or upper Eocene period. Our results provide robust statistical evidence for a sister-group relationship within Mytilida. The findings not only confirm previous results, but also provide valuable insights into the evolutionary history of Mytilidae.

Insights into the Deep Phylogeny and Novel Gene Rearrangement of Mytiloidea from Complete Mitochondrial Genome.

The extant marine mussels which belong to the Mytiloidea are widespread species inhabiting mostly coastal waters, with some distributed in the deep sea. To clarify the classification systems and phylogenetic relationships range from genus to family level within Mytiloidea, new sequence was used in a phylogenetic analysis including all the available Mytiloidea mitochondrial genomes. In this study, the complete mitochondrial genome of Vignadula atrata is 15,624 bp in length and contains 12 protein-coding genes (PCGs, atp8 is absent), two ribosomal RNA genes, and 22 transfer RNA genes. Phylogenetic analysis based on 12 PCGs showed that it has a close relationship to Bathymodiolus. The analysis of gene rearrangements in the Pteriomorphia showed that the arrangements are highly variable across species, novel gene rearrangements were found within Mytiloidea. The V. atrata mitogenome was provided in detail, with notes on the sequence and a key to the species of Vignadula. This study provides a perspective on the taxonomic histories of the marine mussels and refines the unclear relationship between the origin and evolution of species in Mytiloidea within Bivalvia.

Dual-band terahertz absorber based on square ring metamaterial structure.

In this paper, a dual-band terahertz absorber based on metamaterial structure is designed, fabricated, and measured. The metal periodic array is located on the upper surface of a silicon wafer with a metal ground plane, while the metamaterial structure is created utilizing a square metal ring with four T-shaped metal strips loaded inside of the ring. Two absorption peaks are realized at 0.715 and 1.013 THz with high Q-factors of 152.1 and 98.3, respectively, under normal TE and TM polarized incidence. A prototype of the proposed metamaterial absorber is fabricated by electron beam lithography (EBL) and electron beam evaporation (EBE) technology. Furthermore, a terahertz time-domain spectroscopy (TDS) measurement system is employed to test the absorber sample, with good measurement results obtained. This work provides a new option for the design of multi-band terahertz metamaterial absorbers.

Spoof surface plasmon polaritons based on-chip sensor for dielectric detection.

A compact millimeter-wave on-chip sensor for dielectric detection is presented using gallium arsenide technology based on spoof surface plasmon polaritons (SSPPs). The proposed structure is developed from traditional half-mode substrate integrated waveguide (HMSIW) and its dispersion characteristics is analyzed through electromagnetic simulations. Consequently, the operating frequency and bandwidth of the on-chip sensor can be easily adjusted, which provides more flexibility for the practical application of the sensor. The linear sensing for relative dielectric constant of the film materials is acquired, with thickness-insensitive property. Moreover, the low coupling to the nearby components can be achieved due to the strong field confinement characteristics of the SSPPs, which is of great significance in the application scenarios of on-chip integrated circuits for the suppression of electromagnetic interference.

[Spawning substrates of cuttlefish: Type, function, and application]. / 乌贼类产卵附着基的类型、功能和应用.

The fertilized eggs of cuttlefish are sticky eggs. Cuttlefish parents prefer to lay eggs on the attached substrates, which help increase the number of eggs and the hatching rate of fertilized eggs. If egg-attached substrates are sufficient, cuttlefish spawning will be reduced or even delayed. With the advances in the construction of marine nature reserves and research on artificial enrichment techniques, domestic and international experts have conducted research on different types and configurations of attachment substrates around cuttlefish resource enhancement. Based on the source of the substrates, we classified cuttlefish spawning substrates into two types, natural and artificial. By summarizing the differences, advantages, and disadvantages of the common economic cuttlefish spawning substrates in offshore areas worldwide, we sort out the functions of two different types of attachment bases, and discussed the practical applications of natural and artificial egg-attached substrates in spawning ground restoration and artificial enrichment. We proposed several thoughts on the future research directions of cuttlefish spawning attachment substrates, aiming to provide reasonable suggestions for cuttlefish habitat restoration, cuttlefish breeding and sustainable development of fishery resources.

A high Q-factor dual-band terahertz metamaterial absorber and its sensing characteristics.

In this paper, a dual-band metamaterial absorber in the terahertz frequencies is proposed and its refractive index sensing characteristics is analyzed. The metamaterial structure is designed using a square metal ring with four T-shaped strips loaded outside of the ring, where the metal periodic array is on top of a silicon wafer backed with a metal ground plane. The resonant frequencies of the absorber are at 0.89 and 1.36 THz, whose absorption rates are both over 99% under normal TE and TM polarized incidences. The full widths at half maximum of them are 4.4 and 11.2 GHz, respectively, resulting in high quality factors (Q-factors) for these two frequency bands. The absorption rate of the absorber remains stable as the incident and polarized angles are changed. Several proposed metamaterial absorbers are experimentally fabricated and electron beam lithography (EBL) technology is employed. Good measurement results of the dual-band absorption performance are obtained using a terahertz time-domain spectroscopy system based on photoconductive antennas. Furthermore, the metamaterial absorber also shows sensing properties for analytes with different refractive indices or thicknesses. This work provides a new choice for the design of high-Q dual-band terahertz metamaterial absorbers and their application to refractive index sensing.

Novel gene re-arrangement in the mitochondrial genome of Pisidiaserratifrons (Anomura, Galatheoidea, Porcellanidae) and phylogenetic associations in Anomura.

To improve the taxonomy and systematics of Porcellanidae within the evolution of Anomura, we describe the complete mitochondrial genomes (mitogenomes) sequence of Pisidiaserratifrons, which is 15,344 bp in size, contains the entire set of 37 genes and has an AT-rich region. Compared with the pancrustacean ground pattern, at least five gene clusters (or genes) are significantly different with the typical genes, involving eleven tRNA genes and four PCGs and the tandem duplication/random loss and recombination models were used to explain the observed large-scale gene re-arrangements. The phylogenetic results showed that all Porcellanidae species clustered together as a group with well nodal support. Most Anomura superfamilies were found to be monophyletic, except Paguroidea. Divergence time estimation implies that the age of Anomura is over 225 MYA, dating back to at least the late Triassic. Most of the extant superfamilies and families arose during the late Cretaceous to early Tertiary. In general, the results obtained in this study will contribute to a better understanding of gene re-arrangements in Porcellanidae mitogenomes and provide new insights into the phylogeny of Anomura.

The Complete Mitochondrial Genome of Pilumnopeus Makianus (Brachyura: Pilumnidae), Novel Gene Rearrangements, and Phylogenetic Relationships of Brachyura.

Pilumnopeus makianus is a crab that belongs to Pilumnidae, Brachyura. Although many recent studies have focused on the phylogeny of Brachyura, the internal relationships in this clade are far from settled. In this study, the complete mitogenome of P. makianus was sequenced and annotated for the first time. The length of the mitogenome is 15,863 bp, and includes 13 protein-coding genes (PCGs), 22 transfer RNA genes (tRNA), and 2 ribosomal RNA genes (rRNA). The mitogenome exhibits a high AT content (72.26%), with a negative AT-skew (-0.01) and a GC-skew (-0.256). In the mitogenome of P. makianus, all the tRNA genes are folded into the typical cloverleaf secondary structure, except trnS1 (TCT). A comparison with the ancestors of Brachyura reveals that gene rearrangement occurred in P. makianus. In addition, phylogenetic analyses based on thirteen PCGs indicated that P. makianus, Pilumnus vespertilio, and Echinoecus nipponicus clustered into a well-supported clade that supports the monophyly of the family Pilumnidae. These findings enabled a better understanding of phylogenetic relationships within Brachyura.

Dissolved metal assessment in surface seawater: A spatial-seasonal evaluation in the Zhejiang coastal waters, the East China Sea.

The spatial-seasonal distributions and variations, correlations with environmental variables and the pollution degrees of dissolved metals in the Zhejiang coastal seawater were investigated. The concentrations of six dissolved metals (i.e. Cu, Pb, Zn, Cd, As and Hg) were in the ranges of 0.10-6.40 (1.6 ± 0.8), 0.16-3.60 (1.2 ± 0.7), 2.50-24.0 (8.5 ± 4.8), 0.011-0.180 (0.07 ± 0.03), 0.85-4.20 (2.1 ± 0.8) and 0.001-0.110 (0.06 ± 0.02) µg/L, throughout the four seasons, respectively. Significant differences in all the dissolved metals were found among seasons, whereas no significant differences were found among stations. The average concentrations of metals were in the following order: Zn > As>Cu > Pb > Cd > Hg. Single metal contamination factor was in the following order: Pb > Hg > Zn > Cu > As>Cd. The pollution level of dissolved metals in the Zhejiang coastal waters (ZCW) was low. Most of the dissolved metals were correlated to temperature, indicating seasonal differences. The redundancy analysis (RDA) indicated that depth, temperature, nitrate and phosphate could best explain the variance pattern of dissolved metals in the ZCW.

[Growth heterogeneity of Larimichthys polyactis in spring and autumn along the coast of Zhejiang, China]. / 浙江沿岸春秋季小黄鱼生长异质性.

In order to understand the growth dynamics of Larimichthys polyactis in spring and autumn along the coast of Zhejiang Province, we used the survey data of bottom trawlers in spring (April) and autumn (November) from 2014 to 2019 along the coast of Zhejiang Province to examine the heterogeneity of L. polyactis growth by constructing a generalized linear model (GLM) and nine linear mixed effect models (LMEM). The results showed that body length of L. polyactis ranged from 15 mm to 210 mm, with an average of 124.12 mm. The dominant body length group ranged from 110 mm to 140 mm. The body weight ranged from 0.04 g to 156.2 g, with an average of 33.28 g. The dominant body weight group ranged from 30 to 50 g. According to the AIC minimum criterion, the LMEM model with the random effects of seasons and water zones on growth parameters a and b was the best. The results of cross validation also showed that the prediction of this model was the best. In the optimal model, the fixed value of growth parameter a was 0.61×10-4, which ranged in (0.32-1.91)×10-4 after adding season and water random effect. The fixed value of parameter b was 2.73, and ranged from 2.49 to 2.86 after adding the random effects of season and water area, indicating that small yellow croaker was under negative allometric growth. Season and water zone had significant effects on the relationship between body length and body weight of small yellow croaker. From a seasonal perspective, the growth rate of small yellow croaker in spring was higher than that in autumn. From water zone perspective, the closer the offshore distance, the faster the growth rate of small yellow croaker.

Insights into the Deep Phylogeny and Novel Divergence Time Estimation of Patellogastropoda from Complete Mitogenomes.

To further understand the origin and evolution of Patellogastropoda, we determined the mitochondrial genome sequence of Cellana toreuma, and compared its mitogenome characteristics with the other four limpets of Nacellidae. The ratio of Ka and Ks indicated that these Nacellidae species were suffering a purifying selection, with exception of the atp6 gene. The gene sequence is basically consistent among families, while there are great differences among Lottidae species. According to the mitogenome sequences of selected gastropod species, we reconstructed a new phylogenetic tree with two methods. The data complement the mitogenome database of limpets and is a favorable research tool for the phylogenetic analysis of Gastropoda. It is found that there is a long-branch attraction (LBA) artefact in the family Lottiidae of Patellogastropoda. Therefore, the Patellogastropoda was separated by Heterobranchia, and Lottiidae is located at the root of the whole phylogenetic tree. Furthermore, we constructed the divergence time tree according to the Bayesian method and discussed the internal historical dynamics, and divergence differences among the main lineages of 12 Patellogastropoda under an uncorrelated relaxed molecular clock. In turn, we made a more comprehensive discussion on the divergence time of limpets at the molecular level.

Sequence comparison of the mitochondrial genomes of five brackish water species of the family Neritidae: Phylogenetic implications and divergence time estimation.

Neritids are ancient gastropod species which can live in marine, brackish water, and freshwater environments. In this study, we sequenced and annotated the mitochondrial genomes of five brackish water neritids (i.e., Clithon corona, Clithon lentiginosum, Clithon squarrosum, Neritina iris, and Septaria lineata). The mitogenomes ranged from 15,618 to 15,975 bp, and all contain 13 protein-coding genes (PCGs), 22 tRNA genes, and two rRNA genes, with a closed ring structure. We calculated the Ka/Ks values of all 13 PCGs of Neritidae species, all ratios are less than 1, under purification selection. Phylogenetic analysis of the 13 PCGs showed that Neritimorpha is a sister group with Vetigastropoda and Caenogastopoda, genus Clithon is a sister group with Neritina and Septaria. Estimation of divergence time for all species of Neritidae showed that the main differentiation of Neritidae occurred in Cenozoic period (65 Mya), C. corona and C. lentiginosum were differentiated in the Cenozoic Neogene, the other three species diverged in the Cenozoic Paleogene. These results will help to better understand the evolutionary position of Neritidae and provide reference for further phylogenetic research on Neritidae species.

Transcriptome analysis of the ink sac and brain tissues from Sepiella inermis: A resource for discovering genes related to the inking of cephalopods.

The common Chinese cuttlefish (Sepiella inermis) is an important cephalopod with nutritional and commercial value. Intensive inking stimulated by swilling seawater in transfer containers threatens the survival of cephalopods during transportation. However, the molecular basis for the inking behavior of S. inermis remains unclear. In the present study, transcriptome analysis was performed on ink sac and brain tissues from S. inermis under two different conditions, i.e. the control group (with individuals immersed in static seawater) and the experimental group (with individuals immersed in swilling seawater) to determine the global gene expression differences. The individuals from the experimental group ejected ink in response to the swilling of seawater. 330,699 unigenes were obtained from twelve transcriptome libraries via the Illumina Hiseq X platform, and the differentially expressed genes in the ink sac and brain tissues were identified respectively. Multiple upregulated genes in the ink sac were involved in cation transporter activity. Besides, an autocrine/paracrine factor wnt10b like and two important transcription factors (homeobox 1 and Hes-1-b-like) were also significantly upregulated in the ink sac. Moreover, a neuronal nitric oxide synthase (nNOS) was significantly downregulated in the brain. The findings from this study provide an important transcriptomic resource for discovering critical genes related to inking behavior of S. inermis, providing a basis for developing potential methods for protecting S. inermis from intensive inking.

Determination of Phthalates in Marine Sediments Using Ultrasonic Extraction Followed by Dispersive Solid-Phase Extraction and Gas Chromatography-Mass Spectrometry.

A simple, rapid and novel method has been developed and validated for determination of 16 phthalates in marine sediment samples by gas chromatography coupled to mass spectrometry. Freeze dried samples were first ultrasonic extraction by n-hexane:methylene chloride (1:1, v/v) and n-hexane:ethyl acetate (1:1, v/v) and followed by dispersive solid-phase extraction cleanup. The linearity of this method ranged from 1 to 1,000 µg/L, with regression coefficients ranging between 0.9993 and 0.9999. The limits of detection were in ng/g level, ranging between 0.1 and 0.25 ng/g (dry weight). The concentration of the total phthalates in marine sediment samples from Waters of Dongji (Zhoushan, China), Yueqing Bay (Wenzhou, China) and Coastal Waters of Yuhuan (Taizhou, China) ranged from 235.4 to 608.7 µg/kg with diisobutyl phthalate, dibutyl phthalate and di(2-ethylhexyl) phthalate being the major species, which constitutes of 94.6 and 98.1% of the total phthalates. The recoveries of spiked 16 phthalates at different concentration levels in sediment sample 3 of Waters of Dongji (Zhoushan, China) and sediment sample 3 of Yueqing bay (Wenzhou, China) were in the range of 78-117% and 83-114%, respectively, with relative standard deviations of 2.4-6.8% and 3.4-7.5% (n = 5), respectively. The performance of the proposed method was also compared with traditional Soxhlet extraction and column chromatography cleanup on the same genuine sediment samples and comparable efficiencies were obtained. It is concluded that this method can be successfully applied for the determination of phthalates in different marine sediment samples.

Spatial distribution, partitioning, and ecological risk of short chain chlorinated paraffins in seawater and sediment from East China Sea.

Short chain chlorinated paraffins (SCCPs) have attracted increasing attention due to their potential risks to the ecosystem and human health. However, there is still a lack of systematic research on their environmental fate in the coastal marine environment. In this study, we collected paired seawater (n = 40) and surface sediment samples (n = 40) from East China Sea, and investigated their spatial distribution, partitioning behaviors and ecological risks. The total SCCP concentrations (∑SCCPs) in seawater and sediment samples were in the range of 12.2-430 ng/L and 89.6-351 ng/g (dry weight), respectively. C10-11 SCCPs and Cl5-7 SCCPs were the predominant homologues in all of the samples. This study first calculated the field-based logKOC values for ∑SCCPs in the marine environment, which ranged from 5.0 to 6.5 (mean 5.5). The logKOC values of SCCP homologues (range 3.6-8.0, mean 5.6) showed a weak correlation with their logKOW (R = 0.214, p < 0.05). A preliminary risk assessment indicated that SCCPs at current levels posed no significant ecological risk. Overall, this study contributes to the further understanding of environmental behaviors of SCCPs in the marine environment.