Recent speciation and hybridization in Icelandic deep-sea isopods: An integrative approach using genomics and proteomics.

The crustacean marine isopod species Haploniscus bicuspis (Sars, 1877) shows circum-Icelandic distribution in a wide range of environmental conditions and along well-known geographic barriers, such as the Greenland-Iceland-Faroe (GIF) Ridge. We wanted to explore population genetics, phylogeography and cryptic speciation as well as investigate whether previously described, but unaccepted subspecies have any merit. Using the same set of specimens, we combined mitochondrial COI sequences, thousands of nuclear loci (ddRAD), and proteomic profiles, plus selected morphological characters using confocal laser scanning microscopy (CLSM). Five divergent genetic lineages were identified by COI and ddRAD, two south and three north of the GIF Ridge. Assignment of populations to the three northern lineages varied and detailed analyses revealed hybridization and gene flow between them, suggesting a single northern species with a complex phylogeographic history. No apparent hybridization was observed among lineages south of the GIF Ridge, inferring the existence of two more species. Differences in proteomic profiles between the three putative species were minimal, implying an ongoing or recent speciation process. Population differentiation was high, even among closely associated populations, and higher in mitochondrial COI than nuclear ddRAD loci. Gene flow is apparently male-biased, leading to hybrid zones and instances of complete exchange of the local nuclear genome through immigrating males. This study did not confirm the existence of subspecies defined by male characters, which probably instead refer to different male developmental stages.

Humpback whale (Megaptera novaeangliae) song unit and phrase repertoire progression on a subarctic feeding ground.

The songs of the male humpback whales have traditionally been associated with breeding activities at low latitude breeding grounds during winter. This study provides the first detailed analysis of humpback whale songs recorded in the subarctic waters of Iceland using passive acoustic recorders. Recordings were collected during three winter seasons: 2008-2009, 2009-2010, and 2011 during which singing was detected in all seasons. Peak song occurrence was during January-February in all years; this coincides with the timing of the peak breeding season of humpback whales in the Northern hemisphere. A total of 2810 song units from all years were measured and statistically divided into 14 groups, which constructed 25 phrases. The song unit repertoires included stable song unit types that occurred frequently in songs during all years while the occurrence of other song unit types varied more between years. Around 60% of the phrases were conserved between the first two study seasons, while the majority of phrases found during the last study season had not been observed before. This study indicates the importance of a subarctic feeding ground for song progression and song exchange and possibly as an opportunistic mating ground for migrating or overwintering humpback whales.

POPs in long-finned pilot whales mass stranded in Iceland as a proxy for their physiological condition.

Long-finned pilot whales (Globicephala melas) are the most frequently stranded cetaceans in the world; however, the predominant drivers of these events are poorly understood. In this study the levels of persistent organic pollutants from pilot whales stranded in North-east Iceland were quantified and compared to historical data and physical parameters to investigate whether contaminant load may have influenced the physiological state of stranded individuals, how these loads fluctuate with sex and age group, and if this is consistent with the literature. Historical comparison was also carried out to discern how pollutant contamination has changed throughout the past few decades. DDE, transnonachlor and PCB-153 were the top three pollutants respectively. The accumulation of POPs was greater on average in immature individuals than adults, whilst among adults, males had higher concentration than females. Moreover, despite an indication of decreasing POP loads throughout the years, knowledge of harmful thresholds remains exceedingly limited.

Call combination patterns in Icelandic killer whales (Orcinus orca).

Acoustic sequences have been described in a range of species and in varying complexity. Cetaceans are known to produce complex song displays but these are generally limited to mysticetes; little is known about call combinations in odontocetes. Here we investigate call combinations produced by killer whales (Orcinus orca), a highly social and vocal species. Using acoustic recordings from 22 multisensor tags, we use a first order Markov model to show that transitions between call types or subtypes were significantly different from random, with repetitions and specific call combinations occurring more often than expected by chance. The mixed call combinations were composed of two or three calls and were part of three call combination clusters. Call combinations were recorded over several years, from different individuals, and several social clusters. The most common call combination cluster consisted of six call (sub-)types. Although different combinations were generated, there were clear rules regarding which were the first and last call types produced, and combinations were highly stereotyped. Two of the three call combination clusters were produced outside of feeding contexts, but their function remains unclear and further research is required to determine possible functions and whether these combinations could be behaviour- or group-specific.

Occurrence of long-finned pilot whales (Globicephala melas) and killer whales (Orcinus orca) in Icelandic coastal waters and their interspecific interactions.

Long-finned pilot whales and killer whales are widely distributed across the North Atlantic, but few studies have reported their occurrence in Icelandic coastal waters. Here, we use sightings data from research platforms and whale watching tours in six regions of Iceland from 2007 to 2020 to show that the occurrence of long-finned pilot and killer whales varied with region and season. Killer whales were regularly encountered in the south of Iceland during summer and west of Iceland during winter/spring. Long-finned pilot whales were only seen during the summer and were most often encountered in the south, west, and northwest of Iceland. Long-finned pilot whale occurrence in the south of Iceland appeared to increase during the study period but killer whale occurrence showed no noticeable changes. Long-finned pilot whales were sighted often in the areas that were also frequented by killer whales and interspecific interactions were commonly observed when both species co-occurred. Interactions appeared to be antagonistic, with killer whales often avoiding long-finned pilot whales and sometimes fleeing at high speed, similar to what has been described elsewhere in the North Atlantic. In the majority of interactions observed (68%), killer whales avoided long-finned pilot whales by moving away, but in 28% avoidance was at high speed with both species porpoising. This variability in the type of behavioural responses indicates that interactions may be more complex than previously described. We discuss regional trends in long-finned pilot whale and killer whale sightings and potential drivers of the observed interactions. Supplementary Information: The online version contains supplementary material available at 10.1007/s10211-022-00394-1.

Pollutants from shipping - new environmental challenges in the subarctic and the Arctic Ocean.

Maritime activities in the subarctic and Arctic Ocean are predicted to substantially increase in the future due to climate change and declining sea ice cover. Inevitably, the consequences will be seen in impacts on marine ecosystems in this region at many different levels, such as increased pollution load due to antifouling biocides, polycyclic aromatic hydrocarbons, metals and pharmaceuticals. Here we discuss the current situation and evaluate the effect of increased shipping on the environmental status of subarctic and Arctic waters, in relation to elevated loads of both legacy and emerging pollutants in the region. It is of high importance to evaluate the current levels of selected pollutants, which will most likely rise in near future. Furthermore, it is important to improve our understanding of the effects of these pollutants on marine organisms at high latitudes, as the pollutants may behave differently in cold environments compared to organisms at lower latitudes, due to dissimilar physiological responses and adaptations of the cold-water organisms. Integrative studies are needed to better understand the impact of pollutants on the marine fauna while monitoring programmes and research should be continued, with an increased capacity for emerging pollutants of concern.

Adding pieces to the puzzle: insights into diversity and distribution patterns of Cumacea (Crustacea: Peracarida) from the deep North Atlantic to the Arctic Ocean.

The Nordic Seas have one of the highest water-mass diversities in the world, yet large knowledge gaps exist in biodiversity structure and biogeographical distribution patterns of the deep macrobenthic fauna. This study focuses on the marine bottom-dwelling peracarid crustacean taxon Cumacea from northern waters, using a combined approach of morphological and molecular techniques to present one of the first insights into genetic variability of this taxon. In total, 947 specimens were assigned to 77 morphologically differing species, representing all seven known families from the North Atlantic. A total of 131 specimens were studied genetically (16S rRNA) and divided into 53 putative species by species delimitation methods (GMYC and ABGD). In most cases, morphological and molecular-genetic delimitation was fully congruent, highlighting the overall success and high quality of both approaches. Differences were due to eight instances resulting in either ecologically driven morphological diversification of species or morphologically cryptic species, uncovering hidden diversity. An interspecific genetic distance of at least 8% was observed with a clear barcoding gap for molecular delimitation of cumacean species. Combining these findings with data from public databases and specimens collected during different international expeditions revealed a change in the composition of taxa from a Northern Atlantic-boreal to an Arctic community. The Greenland-Iceland-Scotland-Ridge (GIS-Ridge) acts as a geographical barrier and/or predominate water masses correspond well with cumacean taxa dominance. A closer investigation on species level revealed occurrences across multiple ecoregions or patchy distributions within defined ecoregions.

Molecular evidence of the survival of subterranean amphipods (Arthropoda) during Ice Age underneath glaciers in Iceland.

A Two endemic groundwater arthropod crustacean species, Crangonyx islandicus and Crymostygius thingvallensis, were recently discovered on the mid-Atlantic volcanic island of Iceland. The extent of morphological differences from closest relatives, endemism, along with the geographic isolation of Iceland and its complete coverage by glaciers 21,000 years ago, suggests that these two species have survived glaciation periods in sub-glacial refugia. Here we provide strong support for this hypothesis by an analysis of mitochondrial genetic variation within Crangonyx islandicus. Our results show that the species is divided into several distinct monophyletic groups that are found along the volcanic zone in Iceland, which have been separated by 0.5 to around 5 million years. The genetic divergence between groups reflects geographic distances between sampling sites, indicating that divergence occurred after the colonization of Iceland. The genetic patterns, as well as the dependency of genetic variation on distances from the tectonic plate boundary and altitude, points to recent expansion from several refugia within Iceland. This presents the first genetic evidence of multicellular organisms as complex as crustacean amphipods which have survived glaciations beneath an ice sheet. This survival may be explained by geothermal heat linked to volcanic activities, which may have maintained favourable habitats in fissures along the tectonic plate boundary in Iceland during glaciations.

Revision of Pleuroprion zur Strassen, 1903 (Holidoteidae) and re-evaluation of Spectrarcturus Schultz, 1981 (Arcturidae) (Crustacea, Isopoda, Valvifera).

Pleuroprion zur Strassen, 1903 is revised and restricted to its type species, Antares chuni zur Strassen, 1902, which is redescribed, using the type material from the German Deep-Sea Expedition in 1898-1899 and more recent material. Pleuroprion is transferred to Holidoteidae Wägele, 1989. Pleuroprion frigidum Hansen, 1916 is transferred to Astacilla Cordiner, 1793 (Arcturidae), while all other species previously placed in Pleuroprion, i.e. P. murdochi (Benedict, 1898), P. chlebovitschi Kussakin, 1972, P. fabulosum Gujanova, 1955, P. furcatum Kussakin, 1982, P. hystrix (Sars, 1877), P. intermedium (Richardson, 1899), P. iturupicum Kussakin Mezhov, 1979 and P. toporoki Kussakin, 1972, are transferred to Spectrarcturus Schultz, 1981 (Arcturidae). The type species of Spectrarcturus, S. multispinatus Schultz, 1981, is a subjective junior synonym of Arcturus murdochi Benedict, 1898, now S. murdochi comb. nov. Spectrarcturus murdochi (Benedict, 1898) is redescribed on the basis of its type material and new material from South Greenland.

New gnathiid isopod crustaceans (Cymothoida) from Heron Island and Wistari Reef, southern Great Barrier Reef.

Eleven species including six new species of Gnathiidae (Crustacea, Isopoda, Cymothoida) are reported from the Heron Island and Wistari Reef, Capricorn Group, southern Great Barrier Reef: Elaphognathia queenslandica sp. nov., Gnathia acrorudus sp. nov., Gnathia capricornica sp. nov., Gnathia carinodenta sp. nov., Gnathia formosa sp. nov. and Gnathia glaucostega sp. nov. Gnathia cornuta Holdich Harrison, 1980 and Gnathia grutterae Ferreira, Smit Davies, 2010 are for the first time reported from Heron Island and new records from Heron Island are provided for Gnathia biorbis Holdich Harrison, 1980, Gnathia variobranchia Holdich Harrison, 1980, and Gnathia wistari Svavarsson Bruce, 2012.

Immediate biomarker responses to benzo[a]pyrene in polluted and unpolluted populations of the blue mussel (Mytilus edulis L.) at high-latitudes.

Immediate biomarker responses of two high-latitude populations of the blue mussel Mytilus edulis to benzo[a]pyrene (B[a]P) were evaluated. Mussels collected from a clean and a polluted site in southwest Iceland were exposed to the nominal dose of 100 microg B[a]P L(-l) for 3 h, after 4 days of acclimatization in clean seawater. To test the sensitivity to the toxicant and immediate biological responses, the following biomarkers were used: DNA single strand breaks, heart rate and feeding rate. All the biomarkers revealed differences between the study sites. Irrespective of the origin of the organisms, the short time exposure to the high B[a]P concentration did not induce DNA single strand breaks or significantly affect the feeding rate. However, the heart rate results showed significantly different responses. The mussels from the polluted site (Reykjavík harbour) increased their heart rate when exposed to B[a]P, while no difference was observed between the heart rate values of the individuals from the clean site (Hvassahraun). The mussels seem to sense the pollutant they have been previously exposed to, and their acute response indicates physiological adaptation to the polluted environment. The results indicate limited sensitivity and temporal predictivity, i.e. transient measurable changes of these biomarkers, as well as showing that the background of the organisms should be considered when evaluating short-term biomarker responses to contaminants.

Spatial trends of polyfluorinated compounds in guillemot (Uria aalge) eggs from North-Western Europe.

Polyfluorinated alkyl compounds (PFCs) are a group of chemicals of growing concern that have been detected in biological and abiotic samples worldwide. This study reports the concentrations of a suite of PFCs: perfluorooctyl sulfonate (PFOS), perfluorooctyl sulfonamide (PFOSA) and perfluorinated carboxylic acids (PFCAs) in guillemot (Uria aalge) eggs, collected in North-Western Europe, from Iceland, the Faroe Islands, Sweden and two locations in Norway. The highest concentrations of PFOS were found in samples from Sweden (mean 400 ng g(-1) wet weight (w.w.)), which were almost five times higher than concentrations found in Norwegian samples (mean 85 ng g(-1)w.w. from both sample sites). The concentrations found in Icelandic and Faroe samples were lowest (mean 16 and 15 ng g(-1)w.w., respectively). Only Swedish samples differed significantly from the other locations. In general, PFCAs show a different spatial trend than PFOS. Perfluorooctanoic acid (PFOA) was not detected in any sample and perfluorononanoic acid (PFNA) was only detected in samples from Sweden. The most abundant PFCA was perfluoroundecanoic acid (PFUA) with highest concentrations in samples from Sweden (mean 82 ng g(-1)w.w.), samples from the Faroe Islands had the second highest concentration (mean 57 ng g(-1)w.w.) and samples from Iceland and Norway had concentrations ranging between 18 and 30 ng g(-1)w.w. The original hypothesis was based on the idea that PFC concentrations are the highest close to more densely populated and industrialized areas and lower levels in remote areas. However, the geographic pattern is more complicated than predicted and varies among different PFCs.

Field validation, in Scotland and Iceland, of the artificial mussel for monitoring trace metals in temperate seas.

The artificial mussel (AM), a novel chemical sampling device, has been developed for monitoring dissolved trace metals in marine environments. The AM consists of Chelex-100 suspended in artificial seawater within Perspex tubing and enclosed with semi-permeable polyacrylamide gel at both ends. To validate the field performance of the AM in temperate waters, we deployed AMs alongside transplanted blue mussels Mytilus edulis in coastal environments in Scotland (Holy Loch, Loch Fyne, Loch Striven and Millport) and Iceland (Reykjavikurhöfn, Gufunes, South of thornerney, Hofsvik, Hvalfjörethur and Sandgerethi) for monitoring trace metals. While uptake patterns of Cd between the AM and M. edulis were highly comparable, discrepancies were found in the accumulation profiles of the other metals (Cu, Cr, Pb and Zn), in particular Zn. Nonetheless, the AMs gave a better resolution to accurately reveal the spatial difference in dissolved metal contamination when compared with M. edulis. AMs complement the use of mussels since AMs indicate dissolved metals in seawater, whereas uptake by mussels indicates a mixture of dissolved and particulate metals. Our results also indicated that historical metal exposure of the transplanted M. edulis could significantly confound their metal concentrations especially when the deployment period was short (i.e. <34d). This study suggested that the AM can overcome problems associated with variable biological attributes and pre-exposure history in the mussel, and provides a standardized and representative time-integrated estimate of dissolved metal concentrations in different marine environments.

Three new species of Tridentella Richardson, 1905 (Isopoda: Cymothoida: Tridentellidae) from New Caledonia.

Three new species of Tridentella Richardson, 1905 are described from the New Caledonia region. All three species are smooth bodied or largely smooth bodied, and all were collected from the shelf around New Caledonia at depths from 150-760 metres. Tridentella palmata sp. nov. is characterised by pereopod 4 having numerous large robust setae on the carpus and an exceptionally long dactylus (7.0 times as long as proximal width); Tridentella katlae sp. nov. is the only species in the genus with medially united eyes that fill the head in dorsal view; Tridentella magna sp. nov. is the largest species in the genus (36 mm) and can be identified by the head having an anterior submarginal ridge and three submarginal tubercles on the poster margin. Tridentella takedai Nunomura 2006 (off Arasaki, Hokusuka-Shi, Kyushu, Japan) is formally transferred to Alcirona Hansen, 1905, forming Alcirona takedai (Nunomura, 2006) comb. nov.

Subglacial refugia in Iceland enabled groundwater amphipods to survive glaciations.

An endemic family of subterranean amphipods (Crustacea) has recently been discovered in Iceland, in addition to a new species of a previously known amphipod family. These findings are remarkable because Iceland was covered by glaciers from about 2.6 million BP to about 10,000 BP and is isolated on the mid-Atlantic Ridge, far from the North American and European continents. We argue that there were subglacial refugia in Iceland during the Quaternary glacial period. The presence of subterranean amphipods, belonging to an old group with its present distribution mainly in North America and the Eurasian continent, indicates past contact of subterranean fresh waters of Iceland and the North Atlantic continent. The amphipods currently found in Iceland may have been present in Greenland at 40 Ma, when the precursor of Iceland drifted together with the hot spot (Iceland plume) from Greenland.

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.

Dioxins and PCBs in Greenland shark (Somniosus microcephalus) from the North-East Atlantic.

The Greenland shark (Somniosus microcephalus) is a high-trophic fish species present in Arctic waters. The present study aimed to determine concentrations of polychlorinated dibenzo-p-dioxins and furans (PCDDs/Fs), dioxin-like (DL) PCBs and six non-dioxin-like (NDL) PCBs in muscle and liver from 10 female Greenland sharks collected between 2001 and 2003 from Icelandic waters. The mean total concentrations of PCDDs/Fs were 13 and 530pg/g fat for muscle and liver, respectively, and show enrichment in the liver. Concentrations of DL-non-ortho PCBs were also higher in liver compared to muscle with mean concentrations of 7.8 and 0.36ng/g fat, respectively. No enrichment in the liver was found for DL-mono-ortho- and NDL-PCBs. No correlation was found between the size range studied and total concentrations of the compounds analyzed. Total WHO-TEQs (PCDDs/Fs and DL-PCBs) ranged between 7.1-70 and 54-1500pg/g fat in muscle and liver, respectively.

Distribution of haploniscids (Isopoda, Asellota, Haploniscidae) in Icelandic waters, with description of Haploniscus astraphes n. sp. from the Iceland basin and the Southeast Atlantic Ocean.

Ten species of Haploniscidae Hansen, 1916 were sampled in Icelandic waters during expeditions in the framework of the BIOICE project. Nine of these were known from the North Atlantic Ocean, i.e. Haploniscus aduncus Lincoln, 1985, H. ampliatus Lincoln, 1985, Haploniscus angustus Lincoln, 1985, H. bicuspis (Sars, 1877), H. foresti Chardy, 1974, H. hamatus Lincoln, 1985, H. spinifer Hansen, 1916, Antennuloniscus simplex Lincoln, 1985 and Chauliodoniscus armadilloides (Hansen, 1916). All but H. bicuspis and H. angustus were restricted to the Atlantic Ocean south of the Greenland-Iceland-Faeroe Ridge (GIF Ridge), while H. bicuspis occurred at considerable depth ranges both north and south of the GIF Ridge. A new species, Haploniscus astraphes n. sp., is described based on material from the Denmark Strait, North Atlantic and the Guinea Basin, South Atlantic. H. astraphes n. sp. belongs to a group of Haploniscus species closely related to the genus Antennuloniscus and shares several characters with species from that genus, especially the spine row on pleopod 1, the stout sensory seta on the carpus of pereopod 7 and characters of the antennae. H. astraphes n. sp. is characterized by a rectangular body shape, the straight frontal margin of the head and the strongly convex posterior margin of the pleotelson.

Integrated chemical and biological assessment of contaminant impacts in selected European coastal and offshore marine areas.

This paper reports a full assessment of results from ICON, an international workshop on marine integrated contaminant monitoring, encompassing different matrices (sediment, fish, mussels, gastropods), areas (Iceland, North Sea, Baltic, Wadden Sea, Seine estuary and the western Mediterranean) and endpoints (chemical analyses, biological effects). ICON has demonstrated the use of a framework for integrated contaminant assessment on European coastal and offshore areas. The assessment showed that chemical contamination did not always correspond with biological effects, indicating that both are required. The framework can be used to develop assessments for EU directives. If a 95% target were to be used as a regional indicator of MSFD GES, Iceland and offshore North Sea would achieve the target using the ICON dataset, but inshore North Sea, Baltic and Spanish Mediterranean regions would fail.

How can we quantify impacts of contaminants in marine ecosystems? The ICON project.

An international workshop on marine integrated contaminant monitoring (ICON) was organised to test a framework on integrated environmental assessment and simultaneously assess the status of selected European marine areas. Biota and sediment were sampled in selected estuarine, inshore and offshore locations encompassing marine habitats from Iceland to the Spanish Mediterranean. The outcome of the ICON project is reported in this special issue as method-oriented papers addressing chemical analyses, PAH metabolites, oxidative stress, biotransformation, lysosomal membrane stability, genotoxicity, disease in fish, and sediment assessment, as well as papers assessing specific areas. This paper provides a background and introduction to the ICON project, by reviewing how effects of contaminants on marine organisms can be monitored and by describing strategies that have been employed to monitor and assess such effects. Through the ICON project we have demonstrated the use of an integrating framework and gleaned more knowledge than ever before in any single field campaign about the impacts contaminants may have in European marine areas.