Response of the blue mussel Mytilus edulis L. following exposure to PAHs or contaminated sediment

The polycyclic aromatic hydrocarbons fluoranthene and benzo(a)pyrene significantly reduced the feeding rate of mussels. For both compounds the tissue concentration resulting in a 50% reduction of the clearance rate (TEC₅₀) was calculated. At high tissue concentrations both aromatic compounds reduced the tolerance of mussels to aerial exposure, whereas at low tissue concentrations an improved response was noticed. The activities of the antioxidant enzymes Superoxide dismutase (SOD) and catalase were elevated only at low tissue concentrations of fluoranthene and benzo(a)pyrene. At the highest measured tissue concentrations the activity of both enzymes was reduced, possibly due to a narcotic effect. The reproductive success rate of mussels appeared to be affected negatively by the investigated hydrocarbons. The results of a pilot experiment indicate that mussels can be used also for the testing of contaminated sediments.     

Correlation of CYP1A1 induction, as measured by the P450 RGS biomarker assay, with high molecular weight PAHs in mussels deployed at various sites in San Diego Bay in 1993 and 1995

In 1993 collections of marine mussels (Mytilus galloprovincialis) were deployed 1 m from the water surface at six sites in San Diego Bay for 88 days. A similar mussel deployment was conducted in 1995, except the animals were deployed 1 m off the bottom and only for 32 days. After recovery from the sites, tissue was extracted with dichloromethane and the solvent extracts analyzed for chemical contaminant content and the ability to produce CYP1A1 induction in a transgenic cell line (TV101L cells). The cells used in the assay (P450 RGS) are stably transfected with a plasmid containing firefly luciferase linked to human CYP1A1 promoter sequences. Induction (fold increase compared to control) was determined by luminometry 16 h after application of small volumes (2–10 μl) of solvent extracts to cultured cells. Small mussels deployed in the Naval Station (NAV) in 1993 exhibited very high bioaccumulation of polycyclic aromatic hydrocarbons (PAHs; 52 μg/g) and polychlorinated biphenyls (PCBs), in addition to very strong induction of CYP1A1 measured by reporter gene system (RGS) responses. Large mussels deployed at the NAV station in 1993 and intermediate-sized animals placed at three stations within the NAV station in 1995 accumulated 13–29 μg PAH/g and exhibited relatively high RGS responses. Correlation of RGS responses for all mussel samples to the measured PAH concentrations was 0.85 (r²). When the concentrations of seven specific PAHs found in the samples are converted to benzo[a]pyrene equivalents, from previously derived toxic equivalency factors (TEFs) for this test system, and compared to measured RGS responses, the correlations are approximately 0.9. The results of these studies indicate that the RGS biomarker can be used as a screening tool for detection of CYP1A1-inducing compounds in tissues, and an estimate of potential human health or ecological risk from ingestion of contaminated organisms. Positive RGS responses can be followed by detailed chemical analyses of PAHs and coplanar PCBs using the same extract.     

Phase I and II biotransformation enzymes and polycyclic aromatic hydrocarbons in the Mediterranean mussel (Mytilus galloprovincialis,Lamarck, 1819) collected in front of an oil refinery

The aim of the present study was to investigate the responses of phase I and II biotransformation enzymes and levels of PAHs in the Mediterranean mussel (Mytilus galloprovincialis, Lamarck, 1819) collected from three sites at different distance from an oil refinery. Phase I enzyme activities as NAD(P)H-cyt c red, NADH ferry red, B(a)PMO and phase II as UDPGT, GST were measured in digestive gland while 16 PAHs (US-EPA) in whole soft tissue. An added value to the data obtained in the present study rely on the RDA analysis which showed close correlations between PAHs levels and phase I enzyme activities in mussels collected in front of the refinery. And again a significant spatial correlation between B(a)P levels and NADPH-cyt c red activities was observed using linear models. No differences among sites for B(a)PMO and phase II GST activities were observed, while the application of UDPGT as biomarkers requires further investigation.     

Biochemical and histological responses in mussel (Mytilus edulis) exposed to North Sea oil and to a mixture of North Sea oil and alkylphenols

Several environmental chemicals are suspected to be responsible for adverse health effects on the reproductive system in various organisms. During this work, environmentally relevant concentrations of North Sea oil were used alone or in combination with alkylphenols and additional PAH to study the effect on vitellogenin-like protein expression and gonadal development in mussels. North Sea oil (0.5 ppm) induced the expression of phospho-proteins in both sexes indicating that some compounds are oestrogen-mimics. This induction was not seen in samples dosed with the mixture but signs of toxic effects were observed in the gonads. Indeed, numerous degenerating ovarian follicles in females and foci, similar to vertebrate melanomacrophage centres, were observed in testes.     

Biochemical and histological responses in mussel (Mytilus edulis) exposed to North Sea oil and to a mixture of North Sea oil and alkylphenols

Several environmental chemicals are suspected to be responsible for adverse health effects on the reproductive system in various organisms. During this work, environmentally relevant concentrations of North Sea oil were used alone or in combination with alkylphenols and additional PAH to study the effect on vitellogenin-like protein expression and gonadal development in mussels. North Sea oil (0.5 ppm) induced the expression of phospho-proteins in both sexes indicating that some compounds are oestrogen-mimics. This induction was not seen in samples dosed with the mixture but signs of toxic effects were observed in the gonads. Indeed, numerous degenerating ovarian follicles in females and foci, similar to vertebrate melanomacrophage centres, were observed in testes.     

Anthropogenic pollution stimulates oxidative stress in soft tissues of mussel <Emphasis Type="Italic">Crenomytilus grayanus</Emphasis> (Dunker1853)

The digestive gland and gills of the mussel Crenomytilus grayanus extracted from three locations — (i) sampled from a clean and (ii) polluted site and (iii) transplanted from the nonpolluted to polluted site - were analysed for antioxidant enzymes (superoxide dismutase, catalase, glutathione reductase), total oxyradical scavenging capacity and levels of lipid peroxidation products (malondialdehyde, conjugated dienes and lipofuscin). Perturbation of redox status was found in both digestive gland and gill tissues of mussels living in the polluted area. As the activities of superoxide dismutase and catalase were 1.2–3 times higher, the total oxyradical scavenging capacity was lower by 20–35% and the levels of lipid peroxidation products were 2–7 times higher compared to mussels from the reference site. In transplanted mussels, the lipid peroxidation process in both tissues was significantly stimulated (the level of conjugated dienes was increased 1.7–2.5-fold; malondialdehyde and lipofuscin contents were increased 3.5–5-fold) and the total oxyradical scavenging capacity fell by 50–70%. In addition, the transplantation generally resulted in transient and variable responses of antioxidant enzymes for both tissues. Complex response-behaviour of the antioxidant enzymes strongly points to the necessity of employing a combined approach that takes into account activities of antioxidant enzymes and the total oxyradical scavenging capacity, as well as measurement of oxidative damage (e.g., lipid peroxidation) to evaluate the physiological health of molluscs.     

The role of oxyradicals in intracellular proteolysis and toxicity in mussels

Studies were performed on the common mussel, M. edulis L., to determine whether copper (Cu) exposure can affect the extent to which digestive cell proteins are oxidised and whether such oxidative damage is mediated by free radicals. Three age groups of mussels were exposed for 6 ^-days to environmentally realistic concentrations of Cu and then digestive gland homogenates were examined for evidence of protein carbonyl formation. Significant increases in carbonyls relative to untreated control mussels were seen for the youngest (2–4 year-old) and oldest (≥ 10 year-old) mussels only after exposure for 6 days, followed by recovery from exposure for a further 6 days. Untreated mussels also showed an age-related difference in protein oxidation, with a significantly lower concentration in the youngest animals (2–4 year olds). Copper did not affect the levels of modified tryptophan or tyrosine residues or the extent of total lipid peroxidation in digestive gland homogenate. Significant depletion of total vitamin E (a-tocopherol) was seen only in young and medium-aged mussels following exposure for 6 days. The levels of protein carbonyl groups were increased in digestive cell cytosol and lighter lysosomes but not in heavier lysosomes or digestive gland microsomes following 5 days exposure to Cu. Dihydrohodamine-123 was converted to fluorescent rhodamine-123 following sequestration into digestive cell lysosomes. The results suggest a link between the lysosomal sequestration of copper, a concomitant increase in the production of oxyradicals and the potential for intracellular oxidative damage, as well as an increased capacity for oxidative damage in older animals.     

Tributyltin compounds in mussels,oysters and sediments of Chinhae Bay,Korea

To study the historical and current status of the environmental levels of organotin compounds in Chinhae Bay in southeast Korea, mussels (Mytilus edulis galloprovincialis), oysters (Crassostrea gigas) and sediment cores were analysed for tributyltin (TBT) compounds. TBT concentrations were higher in oysters (mean: 0.25 μg TBT-Sn/g dry wt) than in mussels (mean: 0.20 μg TBT-Sn/g dry wt). The major sources of TBT are in the inner bay, and the highest TBT concentrations of 0.50 μg TBT-Sn/g (mussels) and 0.74 μg TBT-Sn/g (oysters) were observed at the site adjacent to the port. Concentrations of TBT decreased sharply with distance from the inner bay toward the outer bay and there was little TBT in the outer bay. A significant correlation was observed between this distance and the tissue TBT content of oysters and mussels. The concentration and percentages of TBT in sediment cores decreased with depth as a result of debutylation. Half-lives of TBT and dibutyltin (DBT) in sediments were 6.9 and 11.6 y, respectively. The TBT concentrations in mussels were higher than those in sediments by a factor of 4–7.     

Biochemical responses in farmed mussel Perna perna transplanted to contaminated sites on Santa Catarina Island, SC, Brazil.

The effects of contaminants on the biochemical parameters of the intensively farmed mussel Perna perna, are unknown. The aim of this study was to compare biochemical responses in mussels held in clean and contaminated sites in Santa Catarina Island, Brazil. Mussels were transplanted from a farming area, Ratones Grande Island (RGI), to two contaminated sites, Itacorubi (ITAC) and Hercílio Luz Bridge (HLB). A reference group was kept at RGI. After 150 and 180 days of exposure, the digestive glands of the mussels were analyzed for catalase, glucose 6-phosphate dehydrogenase (G6PDH) and glutathione S-transferase (GST) activities. No changes were observed in the catalase activity, in both periods. Low G6PDH activity was observed in mussels transplanted for 150 days at the ITAC site. Increased GST activity was observed in mussels from ITAC and HLG sites after 180 days. These responses are probably related to the augmented discharges of domestic effluents associated with elevated rainfall index.     

Distribution of PACs in surficial sediments and bioavailability to mussels, Mytilus edulis of Halifax Harbour

Surficial sediments and blue mussels were collected around the perimeter of Halifax Harbour. Samples were analysed for the levels and fingerprint of polycyclic aromatic compounds (PACs). The objective was to explore the relationship between contaminant profiles in sediments and in mussels and to explore the possibility of deducing their sources, pattern of deposition and circulation in the harbour. Combustion derived compounds predominated in both sets of samples, with more variability observed in the level of petroleum derived hydrocarbons in mussel samples. Biota-sediment accumulation factors (BSAF, dry/dry wt.) varied throughout the harbour, ranging from means of 0.006 to 1.26, where generally fluoranthene displayed the highest BSAF. Sediments and mussels give complementary views of the geographical distribution of contaminants, they reflect the state of benthic and pelagic habitats. Results observed for Halifax Harbour are discussed from the perspective of published data.     

Histopathology of mussels (Mytilus sp.) from the Tamar estuary, UK

This study assessed numerous histological parameters as markers of health status in mussels (Mytilus spp.) collected from several locations along a contamination gradient on the River Tamar and two coastal sites on the north and south coast of Cornwall, UK. Twenty-seven health parameters were assessed including the presence of pathogens, inflammatory lesions, non-specific pathologies and reproductive condition. Logistic regression showed that reproductive condition and inflammatory lesions showed changes indicative of a contamination gradient response. Mussel populations further upstream within the Tamar were relatively less developed compared to mussels situated downstream in the estuary. Compared to coastal locations, mussels sampled within the Tamar also exhibited a higher prevalence of inflammatory lesions that increased at locations situated furthest upstream. Similarly, levels of lipofuscin within the kidney epithelium were highest in mussels sampled within the estuary and Whitsand Bay compared to Trebarwith Strand. The most prevalent pathogens observed during this survey included Rickettsia/Chlamydia-like organisms (R/CLO), gill ciliates including Ancistrum mytili, the copepod Mytilicola intestinalis, Ciliophora-like organisms and Marteilia sp. Marteilia sp. was observed in 31% of the mussels sampled from the Cremyll Ferry site. Generally, the range and prevalence of pathogens was highest from the two estuarine locations at Cremyll Ferry and Wilcove. No significant differences were observed between estuarine sampling sites and Whitsand Bay with respect to median epithelial cell height of digestive tubules, however mussels from Trebarwith Strand did exhibit a reduced median epithelial cell height compared to mussels sampled from other sampling sites. Species genotyping was also undertaken during this study in order to consider any histological differences in relation to species. Results are discussed in relation to histopathology health parameters and biological effects monitoring.     

Estrogenic responses to xenobiotics in channel catfish (Ictalwus punctatus)

Several environmentally relevant contaminants are characterized as xenoestrogens by virtue of their ability to induce responses similar to 17β-estradiol (E2). There is concern that exposure to these xenoestrogens may result in endocrine and, thus, reproductive disruption. The objective of the present research was to determine whether xenobiotics known to elicit estrogenic responses in mammals were estrogenic in channel catfish, and if these compounds were capable of altering normal physiological response to E2. Sexually immature catfish were exposed by intraperitoneal injection to E2, suspected xenoestrogens or combination doses of E2 and xenoestrogen. Appearance of vitellogenin (Vg) in serum was used as the bioindicator of estrogenicity; Vg was measured by ELISA 7 days after injection. The ED₅₀ of E2 for the appearance of Vg in blood, 0.6 mg/kg, was used as the positive control. Synthetic estrogens, ethinylestradiol, mestranol and DES were effective in inducing Vg; the antiestrogen tamoxifen inhibited response to E2 when given in a combination dose. Treatment by both methoxychlor and p-nonylphenol resulted in Vg appearance in serum; the doses required were 300 times that of E2, and the vitellogenic response was less when compared to the E2-induced response. In addition, a combination dose of methoxychlor with E2 decreased the magnitude of the response to E2. Other mammalian xenoestrogens, o,p′-DDT, chlordecone, lindane and β-HCH, were not able to produce a vitellogenic response in this study.     

Tolerance of five species of tropical marine mussels to continuous chlorination

The paper examines the relative lethal and sublethal response of five important tropical marine mussels (Perna viridis, Perna perna, Brachidontes striatulus, Brachidontes variabilis and Modiolus philippinarum) to different chlorine concentrations varying from 0.25 to 15 mg l(-1). The mussels were observed to co-exist in the cooling water circuits of a coastal power station that adopted intermittent chlorination as a fouling control technique. The five mussel species showed, in response to chlorination, 100% mortality at significantly different exposure times, indicating significant species-specific variability in chlorine tolerance. For example, at 1 mg l(-1) residual chlorine, B. variabilis and P. viridis took 288 and 816 h, respectively, to achieve 100% mortality. The time taken for 100% mortality decreased with increasing chlorine residual concentration. The effect of mussel size (= mussel age) of P. viridis, P. perna, B. striatulus and M. philippinarum on mortality was significant between 1 and 5 mg l(-1) residual chlorine, with larger mussels showing greater resistance than smaller ones. All mussel species showed progressive reduction in physiological activities when chlorine residuals were increased from 0 to 1 mg l(-1). However, species-specific differences in the relative rate of physiological activities were observed. Accordingly, relative reduction in physiological activities in response to chlorination was the lowest in P. viridis and the highest in B. variabilis. The data clearly indicate significant differences in the lethal and sublethal responses of the five mussel species to chlorination. The results, therefore, suggest that for effective fouling control, chlorine treatment against mussels has to be employed judiciously, depending on the mussel species involved.     

DNA adducts and polycyclic aromatic hydrocarbon (PAH) tissue levels in blue mussels (Mytilus spp.) from Nordic coastal sites

DNA adducts in gills and digestive gland, as well as polycyclic aromatic hydrocarbon (PAH) tissue levels were analysed in blue mussels (Mytilus spp.) from Nordic coastal areas (Iceland, Norway and Sweden) with diffuse or point sources of PAHs of various origins. Both DNA adduct and PAH tissue levels were generally low, indicating low PAH exposure to the mussels in the areas studied. DNA adducts were found to be higher in gills than in digestive gland of the mussels at all sites studied. Elevated DNA adduct levels in gills were found at 6 sites out of 18 compared to reference sites in respective coastal zones. Adduct levels ranged from 0.5 to 10 nmol adducts/mol normal nucleotides, being highest in mussels from Reykjavík harbour, Iceland (intertidal mussels), and from Fiskaatangen, Norway (subtidal mussels). Total PAH tissue levels in the mussels ranged between 40 and 11,670 ng/g dry wt., and were significantly correlated with DNA adduct levels (r(2)=0.73, p<0.001). PAH ratio values indicated that the PAHs were in most cases of pyrolytic origin, but with petrogenic input near harbours and an oil refinery.     

Effect of seasonality on oxidative stress responses and metal accumulation in soft tissues of Aulacomya atra,a mussel from the South Atlantic Patagonian coast

This study investigated the effects of pollution and its interaction with temperature on the oxidative status of the ribbed mussel Aulacomya atra in the southern Atlantic Patagonian coast. Animals were collected from four sites with different degree and type of human activity impact, during the summer and winter of 2011. Seawater chromium, copper, manganese, nickel and zinc concentrations were measured, as well as metal accumulation, lipid peroxidation, protein oxidation, reduced glutathione levels, and enzymatic activities of superoxide dismutase and glutathione-S-transferase in gills and digestive glands.Metal bioaccumulation and oxidative stress responses in both tissues were generally higher in mussels from harbor areas. Water temperature had a remarkable effect on gill SOD activity and protein oxidation during winter in mussels from all locations.Methodologically, we conclude that measuring both metal bioaccumulation and oxidative stress responses allowed for a more accurate assessment of the biological effects of metal present in seawater.     

Oestrogenic markers and reduced population fertile egg production in a sand goby partial life-cycle test

Understanding the ecological significance of oestrogenic biomarker responses, seen in field surveys of some UK estuarine fish, requires a suitable model to link biochemical and reproductive effects. The sand goby (Pomatoschistus minutus) is a common inshore fish that has been successfully bred in laboratory aquaria. Juvenile sand goby were continuously exposed to 17beta-oestradiol (E2) during gonadal maturation, and their subsequent reproductive success investigated. Exposure endpoints included hepatic vitellogenin (VTG) mRNA expression and maturation (male nuptial colouration, sperm duct gland- and gonado-somatic indices (SDGSI, GSI)). Reproductive endpoints included spawning behaviour, brood size, fertility and population-level fertile egg production. Measured E2 exposure concentrations were <5 (controls), 17, 71 and 530 ngl(-1). Male maturation decreased with increasing E2 concentration. VTG mRNA expression increased with E2 concentration, but not with duration of exposure. Reproductive endpoints showed progressive impairment with increasing E2 concentration. For most endpoints, the lowest observed effect concentration (LOEC) was 71 ngl(-1), and EC50s were in the range 87-165 ngl(-1). Pre-breeding male colouration, VTG expression and SDGSI were good predictors of subsequent population-level fertile egg production, but GSI was not. The sensitivity of oestrogenic markers in sand goby are comparable to those used in other species, and feral fish populations exhibiting such oestrogenic responses at molecular and organ levels are likely to have reduced fertile egg production.     

Molecular investigations into pollutant impact on roundnose grenadier (C. rupestris) and transplanted common mussel (M. edulis) in Skagerrak, the North Sea

The use of a suite of different biochemical markers in the deep sea fish Roundnose grenadier and caged common mussels, and analyses of contaminant levels in the mussels, indicate that Skagerrak are more polluted than waters around the Faroe Islands. In Roundnose, induction of ethoxyresorufin-O-deethylase (EROD) is consistent with increased benzo(a)pyrene hydroxylase (BPH) in mussels and elevated levels of polynuclear aromatic hydrocarbons (PAHs) in mussels and sediment from Skagerrak. This evidence of a large-scale environmental impact of pollutants should lead to further efforts to decrease transport into, and deposition of, waste compounds in the Skagerrak area.     

Temporal variation in the antioxidant defence system and lipid peroxidation in the gills and mantle of hydrothermal vent mussel Bathymodiolus azoricus

Hydrothermal vent mussels are exposed continually to toxic compounds, including high metal concentrations and other substances like dissolved sulphide, methane and natural radioactivity. Fluctuations in these parameters appear to be common because of the characteristic instability of the hydrothermal environment. Temporal variation in the antioxidant enzymes (superoxide dismutase (SOD), catalase (CAT), total glutathione peroxidases (Total GPx), selenium dependent glutathione peroxidases (Se-GPx)), metallothioneins and lipid peroxidation (LPO) in the gills and mantle of the mussel Bathymodiolus azoricus from Menez–Gwen hydrothermal vent site was evaluated and related to the accumulated metal concentrations (Ag, Cu, Cd, Fe, Mn and Zn) in the tissues. Maximum antioxidant enzyme activities in the gills were detected in the beginning of summer, followed by a gradual decrease throughout the following months. One year after, the levels of antioxidant enzyme activities were similar to those reported one year before. LPO in this tissue exhibited a similar temporal variation trend. A different pattern of temporal variation in antioxidant enzyme activities was observed in the mantle, with a gradual increase from summer to the end of autumn (November). LPO in the mantle exhibited an almost reverse trend of temporal variation to that of antioxidant enzyme activities in this tissue. Antioxidant defences in the gills of B. azoricus were significantly enhanced with increasing concentrations of Ag, Cu and Mn, while negative relationships between antioxidant enzymes and Cd, Cu, Mn and Zn concentrations in the mantle were observed, suggesting different pathways of reactive oxygen species (ROS) production and that these tissues responded differently to the metal accumulation. However, temporal variation in biomarkers of defence and damage were in general similar to coastal bivalve species and can be associated with temporal variations of the physiological status due to reproduction. These variations might also be linked to the highly unstable nature of the hydrothermal environment.     

Concentrations of unsubstituted polynuclear aromatic hydrocarbons in bay mussels (Mytilus edulis) from Oregon, USA

Concentrations of fifteen unsubstituted polynuclear aromatic hydrocarbons (PNAH) were measured in Mytilus edulis from two sites in Yaquina Bay, Oregon, USA, during 1979–1980. There were significant differences in PNAH levels between the two populations. The average total concentration in mussels inhabiting the more industrialized bayfront was 986·2 μg/kg compared with 273·9 μg/kg in mussels from a more remote site across the bay. Substantial differences were found in the concentrations of different PNAH in M. edulis examined during this study. The smaller more water soluble, compounds were concentrated to one or two orders of magnitude above the larger, less soluble PNAH.     

Life-history traits of the symbiotic scale-worm Branchipolynoe seepensis and its relationships with host mussels of the genus Bathymodiolus from hydrothermal vents

Associations between scale‐worms and giant mussels are common constituents of hydrothermal vent and cold seep ecosystems, but very little is known about their nature and ecology. Here, we analyze the ecological characteristics of the associations between Branchipolynoe seepensis, an obligate symbiotic polychaete, and their host mytilid mussels Bathymodiolus puteoserpentis and B. azoricus inhabiting hydrothermal vent fields on the Mid‐Atlantic Ridge. Infested mussels generally harbored a single symbiont (<4% had two to six worms). Infestation rate varied from 7.2% to 76.5%, increasing with mussel size, and was significantly lower for B. puteoserpentis. Symbiont density at Lucky Strike ranged between 1071 and 1191 individuals m⁻². Female symbiont size was always positively correlated with host size, while only males and juveniles from small mussels showed the same trend. This suggested a relatively long‐lasting host/symbiont association for females and short‐lasting association with successive reproductive migrations for adult males. The sex ratio of symbionts was always biased in favor of females. Males were smaller and more slender than females and had one mode in their size distributions, whereas females typically had three or more modes, suggestive of a longer life span in females. Between 59.1% and 72.2% of mussels had damaged soft tissues with substantially higher incidence of trauma in infested ones, suggesting that symbionts may cause trauma. The symbionts also induce tunnel‐like structures among the ctenidia, indicating fidelity to a particular location inside the host. Based on our data, together with the fact that infested mussels became relatively wider than non‐infested ones, this association is considered parasitic (likely kleptoparasitic). Our data, together with those from previous studies, allowed us to define the main life‐history traits of B. seepensis: (i) the relationship with their host is parasitic, (ii) the association begins at the smallest mytilid size classes, (iii) there is sexual dimorphism in body size, (iv) sex ratio deviates from 1:1 in favor of females, (v) fertilization occurs through temporal pairing and pseudocopulation, (vi) sperm are stored by females, (vii) eggs are large (likely lecithotrophic or with direct development), (viii) females have a longer life span than males, (ix) adult males may be semalparous, undertaking reproductive migrations followed by a short period of pairing and then death, and (x) females have a semi‐continuous iteroparous reproductive cycle.