Global spread and evolution of viral haemorrhagic septicaemia virus

Viral haemorrhagic septicaemia virus (VHSV) is a rhabdovirus that infects over 48 species of teleosts and is lethal in many. VHSV threatens marine and aquatic fisheries. VHSV was first discovered outside Europe in 1988 in fish from the Pacific coast of North America. In 1994, VHSV was discovered in Newfoundland. In 2003, VHSV was isolated from fish in Lake St. Clair (Michigan and Ontario). In this study, we used 46 nucleotide sequences for the glycoprotein gene from 12 studies and 150 nucleotide sequences for the nucleoprotein gene from nine studies. We combined phylogenetics and a geographic information system to visualize the transmission paths of VHSV lineages. We also reconstructed the spread of VHSV lineages through optimization of geographic data for viral isolates on phylogenetic trees. We demonstrate that VHSV was transmitted from the North Atlantic Ocean and/or Baltic Sea to the Atlantic coast of North America and Japan in independent events. From the Atlantic coast, the virus was transmitted independently to the Laurentian Great Lakes and the Pacific coast of Canada and the contiguous United States. From the Pacific Northwest, the virus was transmitted to Asia and Alaska in independent events. These results clarify the debate ongoing in the literature on the geographic spread of VHSV.     

Investigation of wild caught whitefish, Coregonus lavaretus (L.), for infection with viral haemorrhagic septicaemia virus (VHSV) and experimental challenge of whitefish with VHSV

Abstract One hundred and forty-eight wild whitefish, Coregonus lavaretus (L.), were caught by electrofishing and sampled for virological examination in December 1999 and 2000, during migration from the brackish water feeding grounds to the freshwater spawning grounds, where the whitefish may come into contact with farmed rainbow trout. All samples were examined on cell cultures. No viruses were isolated. Three viral haemorrhagic septicaemia virus (VHSV) isolates of different origin were tested in infection trials by immersion and intraperitoneal (IP) injection, using 1.5 g farmed whitefish: an isolate from wild caught marine fish, a farmed rainbow trout isolate with a suspected marine origin and a classical freshwater isolate. The isolates were highly pathogenic by IP injection where 99-100% of the whitefish died. Using an immersion challenge the rainbow trout isolates were moderately pathogenic with approximately 20% mortality, whereas the marine isolate was virtually non-pathogenic. At the end of the experiment it was possible to isolate VHSV from survivors infected with the marine and suspected marine isolates. Because of the low infection rate in wild whitefish in Denmark, the role of whitefish in the spread of VHSV in Denmark is probably not significant. The experimental studies, however, showed that whitefish are potential carriers of VHSV as they suffer only low mortality after infection but continue to carry virus.     

A mortality event in wrasse species (Labridae) associated with the presence of viral haemorrhagic septicaemia virus

Viral haemorrhagic septicaemia (VHS) is an infectious disease of farmed and wild fish and has an extensive host range in both freshwater and marine environments. In December 2012, a wrasse population consisting of ballan, Labrus bergylta (Ascanius), corkwing, Symphodus melops (L.), cuckoo, Labrus mixtus L., goldsinny, Ctenolabrus rupestris (L.), and rock cook, Centrolabrus exoletus (L.), held at a marine hatchery in the Shetland Isles, Scotland, experienced a mortality event. Approximately 10 000 wrasse were being held at the facility on behalf of an Atlantic salmon, Salmo salar L., aquaculture company prior to being deployed for the biological control of parasites on marine pen Atlantic salmon, aquaculture sites. Fish Health Inspectors from Marine Scotland Science initiated a diagnostic investigation, and subsequent diagnostic testing confirmed the site to be VHSV positive by qRT-PCR and virus isolation followed by ELISA. A VHSV genotype-specific qRT-PCR assay revealed that the isolates belonged to genotype III, the European marine strain of the virus. The virus genotype was further confirmed by nucleic acid sequencing of the partial nucleoprotein (N) and glycoprotein (G) genes followed by BLAST nucleotide searches. This study reports for the first time the detection of VHSV within multiple wrasse species and highlights the need for a comprehensive risk-based approach to the use of wrasse and other finfish species as biological controls within the aquaculture industry.     

Application of a sensitive,specific and controlled real‐time PCR assay to surveillance indicates a low prevalence of viral haemorrhagic septicaemia virus (VHSV) in wild herring,Clupea harengus L., in Scottish waters

Surveillance data on the distribution of viral haemorrhagic septicaemia virus (VHSV) in the North Sea (UK), targeting Atlantic herring in areas with previous virus detection, were obtained from research cruises conducted during 2005. The sensitive molecular approach of real‐time RT‐PCR (qRT‐PCR) was applied alongside a newly developed endogenous positive control assay specific for herring (elongation factor 1α) to ensure integrity of template. Three hundred and five pools from 1937 individual herring were tested, and no evidence of VHSV in association with wild Atlantic herring was detected. Samples were obtained from Scottish waters where marine aquaculture is conducted. The results confirm that previous tissue culture studies have most likely not significantly underestimated the prevalence of carrier herring in this area. The significance of migratory species such as herring as a reservoir species for VHSV, with the potential to translocate virus genotypes between geographical areas, is discussed.     

Epidemiological aspects of viral haemorrhagic septicaemia virus genotype II isolated from Baltic herring, Clupea harengus membras L

This study was carried out to clarify the role of wild fish, especially Baltic herring, Clupea harengus membras L., in the epidemiology of viral haemorrhagic septicaemia virus (VHSV) in brackish water in Finland. Baltic herring with no visible signs of disease were collected from the Archipelago Sea, the Gulf of Bothnia and the eastern Gulf of Finland. In total, 7580 herring were examined by virus isolation as 758 pooled samples and 3029 wild salmonid broodfish as pooled samples during 2004-2006. VHSV was isolated from 51 pooled herring samples in bluegill fibroblast-2 cells, but not in epithelioma papulosum cyprini cells. The majority of isolations were from the coastal archipelago and from fish caught during the spawning season. Based on glycoprotein (G) gene sequences, the virus was classified as a member of genotype II of VHSV. Pairwise comparisons of the G gene regions of herring isolates revealed that all the isolates were closely related, with 98.8-100% nucleotide homology. Phylogenetic analyses revealed that they were closely related to the strains isolated previously from herring and sprat, Sprattus sprattus (L.), in Gotland and to the VHSV isolates from European river lamprey, Lampetra fluviatilis (L.), in the rivers that flow into the Bothnian Bay. The infection in Baltic herring is likely to be independent of the VHSV Id epidemic in farmed rainbow trout, Oncorhynchus mykiss (Walbaum).     

A survey of wild marine fish identifies a potential origin of an outbreak of viral haemorrhagic septicaemia in wrasse,Labridae, used as cleaner fish on marine Atlantic salmon,Salmo salar L., farms

Viral haemorrhagic septicaemia virus (VHSV) was isolated from five species of wrasse (Labridae) used as biological controls for parasitic sea lice predominantly, Lepeophtheirus salmonis (Krøyer, 1837), on marine Atlantic salmon, Salmo salar L., farms in Shetland. As part of the epidemiological investigation, 1400 wild marine fish were caught and screened in pools of 10 for VHSV using virus isolation. Eleven pools (8%) were confirmed VHSV positive from: grey gurnard, Eutrigla gurnardus L.; Atlantic herring, Clupea harengus L.; Norway pout, Trisopterus esmarkii (Nilsson); plaice, Pleuronectes platessa L.; sprat, Sprattus sprattus L. and whiting, Merlangius merlangus L. The isolation of VHSV from grey gurnard is the first documented report in this species. Nucleic acid sequencing of the partial nucleocapsid (N) and glycoprotein (G) genes was carried out for viral characterization. Sequence analysis confirmed that all wild isolates were genotype III the same as the wrasse and there was a close genetic similarity between the isolates from wild fish and wrasse on the farms. Infection from these local wild marine fish is the most likely source of VHSV isolated from wrasse on the fish farms.     

7种海洋鱼类的生物能量学模式

根据室内流水实验法，定量研究了黄、渤海生态系统7种鱼类能量收支组分，并建立了相应的生物能量学模式。7种海洋鱼类的生物能量学模式豆著不同，并可分为3类：(1)较低代谢和较高生长，如黑鲳、矛尾鳃虎鱼；(2)较高代谢和较低生长，如欧氏六线鱼和黑鲷；(3)代谢和生长均处于中等水平，如真鲷、红鳍东方纯和鲐。其差异原因可能与这7种海洋鱼类的生态习性不同相关。与淡水肉食性鱼类比较，7种海洋鱼类的代谢能明显偏高，表明海洋鱼类属于高代谢消耗、低生长效率型鱼类。     

Outbreak of viral haemorrhagic septicaemia (VHS) in lumpfish (Cyclopterus lumpus) in Iceland caused by VHS virus genotype IV

A novel viral haemorrhagic septicaemia virus (VHSV) of genotype IV was isolated from wild lumpfish (Cyclopterus lumpus), brought to a land‐based farm in Iceland, to serve as broodfish. Two groups of lumpfish juveniles, kept in tanks in the same facility, got infected. The virus isolated was identified as VHSV by ELISA and real‐time RT‐PCR. Phylogenetic analysis, based on the glycoprotein (G) gene sequences, may indicate a novel subgroup of VHSV genotype IV. In controlled laboratory exposure studies with this new isolate, there was 3% survival in the I.P. injection challenged group while there was 90% survival in the immersion group. VHSV was not re‐isolated from fish challenged by immersion. In a cohabitation trial, lumpfish infected I.P. (shedders) were placed in tanks with naïve lumpfish as well as naïve Atlantic salmon (Salmo salar L.). 10% of the lumpfish shedders and 43%–50% of the cohabiting lumpfish survived after 4 weeks. 80%–92% of the Atlantic salmon survived, but no viral RNA was detected by real‐time RT‐PCR nor VHSV was isolated from Atlantic salmon. This is the first isolation of a notifiable virus in Iceland and the first report of VHSV of genotype IV in European waters.     

A novel multiplex RT‐qPCR method based on dual‐labelled probes suitable for typing all known genotypes of viral haemorrhagic septicaemia virus

Viral haemorrhagic septicaemia (VHS) is a notifiable fish disease, whose causative agent is a rhabdovirus isolated from a wide range of fish species, not only in fresh but also in marine and brackish waters. Phylogenetic studies have identified four major genotypes, with a strong geographical relationship. In this study, we have designed and validated a new procedure – named binary multiplex RT‐qPCR (bmRT‐qPCR) – for simultaneous detection and typing of all four genotypes of VHSV by real‐time RT‐PCR based on dual‐labelled probes and composed by two multiplex systems designed for European and American/Asiatic isolates, respectively, using a combination of three different fluorophores. The specificity of the procedure was assessed by including a panel of 81 VHSV isolates covering all known genotypes and subtypes of the virus, and tissue material from experimentally infected rainbow trout, resulting in a correct detection and typing of all strains. The analytical sensitivity was evaluated in a comparative assay with titration in cell culture, observing that both methods provided similar limits of detection. The proposed method can be a powerful tool for epidemiological analysis of VHSV by genotyping unknown samples within a few hours.     

Transport of larvae and juvenile fish into central and northern Norwegian waters

The distribution of juvenile fish in Norwegian waters in spring and summer provides evidence that some of these are transported into Norwegian coastal waters from spawning grounds outside the ecosystems which comprise the central and northern Norwegian coast, the Barents Sea and the Spitsbergen area. The relevant species are capelin (Mallotus villosus Müller), haddock (Melanogrammus aeglefinus L.), herring (Clupea hareri' gus L.) and saithe (Pollachius virens L.). A substantial part of the saithe and haddock stock along the central and northern Norwegian coast is considered to be recruited from spawning grounds in the northern North Sea. A contribution of these species from the spawning grounds around the Faeroe Islands and west of Scotland may also occur in some years. Herring from the northern North Sea are found scattered off the Norwegian coast north to 70^oN. It seems likely that pelagic species such as capelin and herring are transported by the prevailing currents from Iceland.     

Using otolith microchemical analysis to investigate the importance of brackish bays for pike (Esox lucius Linnaeus, 1758) reproduction in the southern Baltic Sea

The brackish Baltic Sea is inhabited by several primary freshwater fish species. The pike (Esox lucius) is the largest of these species and acts as the top predator in the marine food web of the Baltic Sea. Prior studies have shown the importance of brackish bays as spawning grounds for the Baltic Sea pike. To identify the spawning habitat of the pike population around the island of Rügen (southern Baltic Sea)—either freshwater streams flowing into the Baltic or brackish lagoons of the Baltic—we used the ICPMS method to quantify the strontium to calcium (Sr:Ca) ratios in sagittal otoliths of 79 adult pike. In total, only six per cent (n = 5) of the analysed Baltic Sea individuals were classified as being of freshwater origin. Ninety‐four per cent (n = 74) of analysed pike were classified as having hatched in brackish water. Compared to previous studies on Baltic Sea pike, our sample shows a high proportion of pike that is of brackish origin: this highlights the adaptive capacity of this species with regard to its ability to reproduce in different salinities. Our data highlight the importance of brackish bays for pike reproduction in the southern Baltic Sea, even if pike live on the upper limit of their salinity tolerance in this area. This is an important finding considering decreases in commercial catches in the last 30 years indicating a population decline supposedly related to recruitment failures.     

European small pelagic fish distribution under global change scenarios

The spectre of increasing impacts on exploited fish stocks in consequence of warmer climate conditions has become a major concern over the last decades. It is now imperative to improve the way we project the effects of future climate warming on fisheries. While estimating future climate‐induced changes in fish distribution is an important contribution to sustainable resource management, the impacts on European small pelagic fish—representing over 50% of the landings in the Mediterranean and Black Sea between 2000 and 2013—are yet largely understudied. Here, we investigated potential changes in the spatial distribution of seven of the most harvested small pelagic fish species in Europe under several climate change scenarios over the 21st century. For each species, we considered eight Species Distribution Models (SDMs), five General Circulation Models (GCMs) and three emission scenarios (the IPCC Representative Concentration Pathways; RCPs). Under all scenarios, our results revealed that the environmental suitability for most of the seven species may strongly decrease in the Mediterranean and western North Sea while increasing in the Black and Baltic Seas. This potential northward range expansion of species is supported by a strong convergence among projections and a low variability between RCPs. Under the most pessimistic scenario (RCP8.5), climate‐related local extinctions were expected in the south‐eastern Mediterranean basin. Our results highlight that a multi‐SDM, multi‐GCM, multi‐RCP approach is needed to produce more robust ecological scenarios of changes in exploited fish stocks in order to better anticipate the economic and social consequences of global climate change.     

Annual and seasonal dynamics of fish in the brackish-water Matsalu Bay, Estonia

Abstract –  Matsalu Bay in the Väinameri (West-Estonian Archipelago Sea) is a relatively shallow but large bay. It is the only real delta estuary in the northern Baltic Sea. The fish fauna is strongly dominated by freshwater species such as perch Perca fluviatilis , roach Rutilus rutilus , white bream Blicca bjoerkna and rudd Scardinius erythrophthalmus . The study presents CPUE data describing annual and seasonal variations in the species composition, abundance and spatial distribution, based on 1 year of monthly samplings from August 2001 to August 2002 and routine coastal monitoring (in late July–early August) between 1994 and 2003. The bay can be divided into three rather distinct parts. The fish fauna of the inner part (salinity 0–2 p.p.t.) is different from two open parts and resembles that of freshwater lakes. In the openmost part (salinity usually 4–6 p.p.t.) some marine species, such as herring Clupea harengus membras , flounder Platichthys flesus and eelpout Zoarces viviparus may seasonally be abundant. Species composition and abundance of most species varies seasonally and interannually. Only the abundance of few species such as pike Esox lucius and pikeperch Stizostedion lucioperca was not impacted by seasonality. The most dynamic period in the bay is spring, when several species (some of them very rare in summer, such as herring and smelt Osmerus eperlanus ) enter the bay for spawning.     

Investigation of round goby viral haemorrhagic septicaemia outbreak in New York

Eleven viral haemorrhagic septicaemia virus (VHSV) genotype IVb isolates were sequenced, and their genetic variation explored to determine the source of a VHS outbreak on the eastern shore of Cayuga Lake. An active fish kill of round gobies (Neogobius melanostomus, Pallas) was intensively sampled at King Ferry, NY and nearby Long Point State Park in May 2017. Gross lesions observed on 67 moribund round gobies and two rock bass (Ambloplites rupestris, Rafinesque) included moderately haemorrhagic internal organs and erythematous areas on the head, flank, and fins. RT‐qPCR tests for VHSV were positive for all 69 fish. Viral isolation on epithelioma papulosum cyprinid cells showed cytopathic effect characteristic of VHSV for six round goby samples from King Ferry. The complete nucleotide sequence of the VHSV IVb genomes of five Cayuga Lake round goby isolates were derived on an Illumina platform along with 2017 VHSV IVb isolates from round gobies collected from the following: Lake Erie near Dunkirk, NY; the St. Lawrence River near Clayton and Cape Vincent, NY; and Lake St. Lawrence near Massena, NY. The phylogenetic tree created from these aligned sequences and four other complete VHSV IVb genomes shows Cayuga Lake isolates are closely related to the Lake Erie isolates.     

Particularities of early life stages in temperate freshwater fish species: comparisons with marine species and implications for aquaculture practices

Both egg and larvae are different between freshwater and marine fish species. Freshwater fish species have generally larger and fewer eggs than marine species. Most freshwater fish species have demersal eggs that develop stuck to various substrata, such as plants or gravels, while eggs of most marine fish species develop in the water column. These differences have consequences for both the evaluation of the quality and the incubation of eggs of freshwater fish species compared with marine species. The larvae of many freshwater fish species are larger and more developed at hatching than their marine counterparts: thus, larval feeding regimes could be different and cannibalism may emerge sooner in certain freshwater fish species. The main differences of egg and larvae between freshwater and marine species are highlighted and the possible implications for aquaculture practices are discussed.     

Why are so many boreal freshwater fishes anadromous? Confronting ‘conventional wisdom’

Suggestions that anadromy predominates in northern high latitudes because boreal marine seas have high primary productivity are questioned. Instead, post‐Pleistocene invasion of far northern fresh waters, which became inhabitable as Pleistocene temperatures ameliorated, via coastal seas, is suggested as a primary cause for northern distributions of widely anadromous fish families. Also, cold temperature preferences may be implicated in determining the ranges of these families, as well as the inhospitability of far northern fresh waters and the lack of ecosystem space to support the huge populations of boreal anadromous fishes. Moreover, there are significant numbers of tropical anadromous fish species.     

Migration of eels tagged in the Baltic Sea and Lake Mälaren—in the context of the stocking question

Eels (Anguilla spp.) are in decline worldwide, and the signs of reduced recruitment have been observed in continental Europe since the early 1970s. To aid recovery of the European eel, stocking is used by many European countries as a management option. In this study, tagging experiments were conducted to follow eel migration from Lake Mälaren and four sites along the Swedish east coast in the Baltic Sea. The recaptured tagged eels were retrieved from fishermen, allowing for the opportunity to investigate their origin (brackish water, stocked in freshwater or a mix in between) by otolith microchemistry and to assess for morphological differences after tagging. Several changes took place; for example, eye index increased while weight and condition decreased with migrated distance and time until recapture. In Lake Mälaren, the majority of tagged eels did not migrate out of the outlets, irrespective of their origin. Most of them were caught in the opposite direction and continued to be caught in the lake 1–3 years after tagging, with significant weight losses. Overall, overwintering is suggested to be an inferior option, but it is uncertain whether this is a natural behaviour or a result of translocation and restocking. For coastal eels, origin had no effect on migratory behaviour; a majority of the tagged eels migrated towards the outlet of the Baltic Sea. Interestingly, a minority of the recaptured eels originated from stocked fish. Instead, recaptures were dominated by natural immigrants that had spent most of their lives in brackish waters.     

Elemental fingerprinting in otoliths reveals natal homing of anadromous Baltic Sea pike (Esox lucius L.)

We examined the element pattern in the otoliths of a migratory fish species that inhabit the coastal areas in the brackish of the Baltic Sea. The northern pike (Esox lucius) show migratory behaviour, spawning in streams and rivers and foraging in the sea. We examined spawning migration in four nearby streams in the south‐west part of the Baltic. Otolith analysis by microPIXE revealed unique elemental patterns (Sr, Zn, Br, Co and Mn) for the juveniles in each of the different streams. The strontium signal in the otolith of the juveniles was used as an indicator of freshwater origin and the time spent in the stream. Adult pike in their migrating spawning phase were caught in each of the streams. The elemental composition in otoliths in their freshwater phase (using juvenile pike in the streams as references) was determined. A principal component analysis showed that the elemental fingerprint during the freshwater phase several years back in time was similar for the adult fish and for juveniles inhabiting the stream today. The results indicated natal homing of the adults to a specific stream, a conclusion that was strengthened by the fact that marked fish returned to spawn over consecutive years. Anadromous pike in the Baltic Sea may thus be divided in subpopulations. The results of the study may have implications for fishery management, as pike in the Baltic Sea cannot be seen as homogenous population.     

Structure and stability in exploited marine fish communities: quantifying critical transitions

Correlations between time series of the abundance of predator and prey fish species in heavily exploited western North Atlantic marine fisheries vary temporally but are generally positive in southern, warmer waters and negative in northern, colder ones. The correlations provide an index of trophic structure and dynamics. We construct a framework to quantify critical thresholds between states in which the predator–prey correlations are positive or negative. We do so by developing a quantitative model of the distribution of the correlations between predator (15 species) and prey (8 species) functional groups based on the annual predator depletion rates and bottom temperatures (or alternatively species richness). The model accounts for 58% of the variance of the correlations with a root mean square error of 0.3. This index of trophic structure indicates that warmer, species-rich, southern fish populations resist transformation from positive to negative predator–prey correlations at exploitation rates that can be double those in the colder, relatively species-poor, northern areas. The model can be used to set limits for exploitation rates that preserve the functional relationships between predator–prey groups in emerging fisheries, and to assess the potential for and measures required to achieve recovery of degraded fish communities.