819:
region. As of July 13, 2007, new rules have been enacted in the
Canadian province of Ontario, and US states of Michigan, New York, Ohio, Pennsylvania, and Wisconsin, while they are currently being drafted in Illinois, Indiana, and Minnesota. Additionally, the USDA's Animal and Plant Health Inspection Service issued a federal order in the fall of 2006 barring the transfer of all live susceptible species from the eight states bordering the lower Great Lakes, as well as importing such species from the Canadian provinces of Ontario and Quebec.
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Survivors of the disease can become lifelong carriers of the virus, contaminating water with urine, sperm, and ovarian fluids. The virus has been shown to survive two freeze/thaw cycles in a conventional freezer, suggesting both live and frozen bait could be a transmission vector. In Europe, the gray
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hemorrhaging (tiny spots of blood) in internal muscle tissue, and petechial or severe hemorrhaging in internal organs and other tissues. Internal hemorrhaging can be observed as red spots inside a dead fish, particularly around the kidney, spleen, and intestines, as well as the swim bladder, which
843:
Fish that become infected experience hemorrhaging of their internal organs, skin, and muscle. Some fish show no external symptoms, but others show signs of infection that include bulging eyes, bloated abdomens, bruised-looking reddish tints to the eyes, skin, gills and fins. Some infected fish have
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free reverse genetic system for Great Lakes VHSV (Genotype IVb) was developed by a research group from the USA. This system allows the investigators to explore the functional properties of individual viral genes of VHSV in detail. This system was immediately utilized to characterize the non-virion
818:
Before the Type IVb die-offs in the Great Lakes, few states regulated transfer of live species other than for salmonids. Since 2005, new policies have been adopted concerning fish and egg transfer, use of live bait, and water transfer, aimed at curtailing the spread to new lakes and rivers in the
679:
The North
American genotype of the virus, in addition to moderate mortality to salmonid species, including various varieties of trout, is also proving virulent among a wide variety of warm-water species previously considered resistant to VHS. The Great Lakes region variant has killed lake trout,
632:
VHSV continues to be found in new geographical areas, in new species of fish. This is thought to represent both the spread of the virus into new areas, as with VHSV egg and live fish transfers from North
America to Asia, or feeding of raw marine fish to inland farmed trout in Finland, as well as
590:
In 1988, the first marine strain of VHSV, now designated type IV, was found in normal-appearing salmon returning from the
Pacific to rivers of Washington State. This strain and other marine strains were not lethal to rainbow trout. The discovery prompted further studies, and by the mid-1990s,
680:
steelhead trout, chinook salmon, yellow perch, gobies, emerald shiners, muskies, whitefish, and walleye. While the
European strain of VHSV is particularly deadly to rainbow trout, the Great Lakes region variant affects the species only mildly, as is typical with primarily marine genotypes.
2006:
Elsayed E, Faisal M, Thomas M, Whelan G, Batts W, Winton J (October 2006). "Isolation of viral haemorrhagic septicaemia virus from muskellunge, Esox masquinongy (Mitchill), in Lake St Clair, Michigan, USA reveals a new sublineage of the North
American genotype".
945:
The Manual or
Diagnostic for Aquatic Animals, 2006, is the standard reference for definitive tests. In most cases, cell culturization is recommended for surveillance, with antibody tests and reverse transcription polymerase chain reaction
1726:
Gagné N, Mackinnon AM, Boston L, Souter B, Cook-Versloot M, Griffiths S, Olivier G (April 2007). "Isolation of viral haemorrhagic septicaemia virus from mummichog, stickleback, striped bass and brown trout in eastern Canada".
1042:
gene was identified as an interferon-stimulated gene against VHSV. Whereas, viperin produces inhibitory ddhCTP (3ʹ-deoxy-3′,4ʹdidehydro-CTP), which is an elongation inhibitor. Thereby, the VHSV RNA replication is terminated.
722:
lake whitefish were infected with VHS. Initially classified as an unknown rhabdovirus, the Lake
Ontario sample was confirmed to be VHSV in 2006, while the Lake Huron sample was confirmed to be VHSV in 2007.
1434:
Ammayappan A, Kurath G, Thompson TM, Vakharia VN (August 2011). "A reverse genetics system for the Great Lakes strain of viral hemorrhagic septicemia virus: the NV gene is required for pathogenicity".
427:
species. Earlier studies used different numbering systems, but the following system has come into common usage based on genotype similarity based on sequencing of the N- and G-genes. Types I-III are
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Living fish afflicted with VHS may appear listless or limp, hang just beneath the surface, or swim very abnormally, such as constant flashing circling due to the tropism of the virus for the brain.
583:
Type I-a was the only strain known from VHSV's discovery in 1963 until the late 1988, isolated to fish farms in continental Europe, affecting primarily rainbow trout and occasionally brown trout or
803:, the second largest freshwater lake in the United States. Contamination in the lake had been expected for months by experts, since the presence of VHS was confirmed in the connected Lake Huron.
961:
Thoroughly cleaning boats, trailers, nets and other equipment when traveling between different lakes and streams also helps. The only EPA-approved disinfectant proven effective against VHS is
668:
since 2005 of many species in the affected lakes. Originally found off the
Atlantic coast of Canada, it was considered a low mortality marine strain. Its first detection in freshwater was in
388:
VHSV enters the host cell through either fusion or endocytosis. It binds to a glycoprotein, called fibronectin, in the extracellular matrix. VHSV does its binding through integrin receptors.
922:
examination, observing tissues through a microscope. Most tissue changes can be observed as minor to major necrosis (cell death) in the liver, kidneys, spleen, and skeletal muscle. The
969:
kills the VHS virus, but in concentrations that are much too caustic for ordinary use. Disinfecting stations can be found at various inland lake boat launches in the Great Lakes region.
757:. Preliminary tests of samples of freshwater drum collected on May 2 were positive, and the announcement came amidst a die-off of hundreds of freshwater drum there and on neighboring
2403:
1859:
1384:"Limited interference at the early stage of infection between two recombinant novirhabdoviruses: viral hemorrhagic septicemia virus and infectious hematopoietic necrosis virus"
2191:
1936:
Importation of pilchards (Sardinops sagax) for direct introduction into natural waters: Biosecurity policy review of viral haemorrhagic septicaemia virus (VHSV), Draft Report
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1272:
81:
was identified as a distinct, more marine-stable strain than the
European genotype. VHS has since been found afflicting marine fish in the northeastern Pacific Ocean, the
46:. Different strains of the virus occur in different regions, and affect different species. There are no signs that the disease affects human health. VHS is also known as
879:
20:
2731:
Im J, Kim WR, Lee HE, Kim A, Kim DH, Choi YH, et al. (January 2020). "Expression analysis of LTR-derived miR-1269a and target gene, KSR2 in Sebastes schlegelii".
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External signs are not always present, but if they are, hemorrhaging on the skin's surface can appear as anywhere from tiny red dots (petechiae) to large red patches.
322:
2490:
1142:
70:
1279:. United States Department of Health and Human Services, National Institutes of Health, National Library of Medicine, National Center for Biotechnology Information
599:, and different genotypes have appeared in different areas since then. Type IV was later found off North America's northern Atlantic coast, in Atlantic herring (
1811:
1813:
Report of the ICES Advisory Committee on Fishery Management, Advisory Committee on the Marine Environment and Advisory Committee on Ecosystems, 2006. Book 2
2312:
2342:
1668:"Genotyping and pathogenicity of viral hemorrhagic septicemia virus from free-living turbot (Psetta maxima) in a Turkish coastal area of the Black Sea"
382:
2372:
2281:
2251:
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2689:
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1935:
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2252:"Fish likely infected with deadly virus found in Little Lake Butte des Morts; Menasha Lock closed; Public asked to take steps to stop the spread"
1539:"Genetic relatedness among Japanese, American and European isolates of viral hemorrhagic septicemia virus (VHSV) based on partial G and P genes"
2064:
He M, Yan XC, Liang Y, Sun XW, Teng CB (August 2014). "Evolution of the viral hemorrhagic septicemia virus: divergence, selection and origin".
652:
This virus appears to have evolved in the Pacific Northwest of North America. It appears to have diverged into several strains ~300 years ago.
334:
2198:. Cornell University, College of Veterinary Medicine, Department of Microbiology and Immunology, Aquatic Animal Health Program. Archived from
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2905:
1007:
2568:
810:, the second of New York's Finger Lakes to test positive for the disease. The disease caused a large die-off of bass in the spring of 2007.
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is a powerful tool to study and characterize the previously unknown viral genes. Reverse genetics system is currently available for VHSV. A
2599:
827:
VHSV can be spread from fish to fish through water transfer, as well as through contaminated eggs, and bait fish from infected waters. The
2877:
1174:"Identification of viral hemorrhagic septicemia virus isolated from Pacific cod Gadus macrocephalus in Prince William Sound, Alaska, USA"
926:(blood-forming) areas of the kidney and spleen are the initial area of infection, and should show necrosis. The gill may have thickened
2433:
1870:
423:(unique strains) of VHSV are typically grouped by genotyping. It is found that genotype groups are divided more geographically than by
750:
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marine VHSV was found in eight species along the northern North America's Pacific coast, and 14 species in and around the Atlantic's
2199:
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773:
295:
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1967:"An outbreak of VHSV (viral hemorrhagic septicemia virus) infection in farmed Japanese flounder Paralichthys olivaceus in Japan"
1905:
1772:"Outbreak of viral haemorrhagic septicaemia (VHS) in seawater-farmed rainbow trout in Norway caused by VHS virus Genotype III"
1333:"Identification of diverse defense mechanisms in rainbow trout red blood cells in response to halted replication of VHS virus"
2661:
Cornell University, College of Veterinary Medicine, Department of Microbiology and Immunology, Aquatic Animal Health Program
411:
at the early stage of viral infection. This discovery unlocked the mystery of presence of NV proteins in novirhabdoviruses.
2501:
835:
heron has spread the virus, but it does so mechanically; the virus is apparently inactive in the digestive tract of birds.
746:
in western New York state. This was the first case in the region outside waters contiguously connected to the Great Lakes.
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of single stranded RNA, which contains six genes that are located along the genome in the 3′-5′ order: 3′-N-P-M-G-NV-L-5′,
2527:
1150:
65:, documented as a pathogenic disease among cultured salmonids since the 1950s. Today it is still a major concern for many
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areas, the kidney may be swollen and unusually red, the spleen may be swollen, and the digestive tract may be empty.
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There may also be a nervous form of the disease where fish are constantly flashing and showing abnormal behaviour.
761:. Preliminary tests later indicated VHS in specimens from Lake Winnebago. The lakes drain through the Fox River to
613:
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1302:
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341:, bullet-shaped particle, about 180 nm long by 60 nm in diameter, covered with 5 to 15 nm long
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discovery of existing populations, as with an apparently well established marine reservoir in the Black Sea.
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to Europe, and Type IV to North America, and Type I and type IV isolates are further subdivided, as follows:
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Electron microscopy can reveal the bullet-shaped rhabdovirus, but is not adequate for definitive diagnosis.
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to the north. The sample was not tested for VHS until 2005, after the disease was detected in Lake Ontario.
569:
Marine fish of the Northwest Pacific (North America), North American north Atlantic coast, Japan, and Korea
147:
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3178:
2891:
2873:
2851:
2380:
2289:
2259:
553:
399:(NV) gene of novirhabdoviruses. Even though it has been demonstrated that the NV gene is not necessary for
2142:"Molecular characterization of the Great Lakes viral hemorrhagic septicemia virus (VHSV) isolate from USA"
3334:
2914:
781:
754:
2350:
1943:
3523:
3313:
3052:
1580:"Phylogenetic analysis of viral haemorrhagic septicaemia virus (VHSV) isolates from France (1971-1999)"
2690:"Great Lakes fish hatcheries could benefit from new test for deadly VHS virus | Great Lakes Echo"
3500:
3318:
2434:"Ontario Government Acting Swiftly to Control Further Spread of New Fish Virus]. (Press release)"
998:
927:
766:
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is a particularly popular bait fish in the Great Lakes region, and is among the species afflicted.
1331:
Nombela I, Puente-Marin S, Chico V, Villena AJ, Carracedo B, Ciordia S, et al. (2018-02-09).
3264:
3252:
3196:
3042:
2576:
1487:"Nonvirion protein of novirhabdovirus suppresses apoptosis at the early stage of virus infection"
950:) and genetic sequencing and comparison for definitive confirmation and genotype classification.
692:
is the earliest confirmed case of VHSV within the Great Lakes region. Lake St. Clair connects to
2607:
3474:
3396:
3225:
3115:
2994:
2852:"Viral Hemorrhagic Septicemia | National Invasive Species Information Center | USDA"
2111:"Economic impacts of private sector aquaculture-based recreational fishing in the western USA"
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2006 saw mass die-offs from VHS in Lake Erie, the St. Clair River, the Detroit River, and the
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3122:
3000:
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1578:
Thiéry R, de Boisséson C, Jeffroy J, Castric J, de Kinkelin P, Benmansour A (November 2002).
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Virus neutralisation is another important method of diagnosis, especially for carrier fish.
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3213:
3165:
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3019:
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2016:
1736:
1679:
1443:
1147:
Virus Taxonomy Online: Seventh Report of the International Committee on Taxonomy of Viruses
883:
793:
777:
142:
2776:"Fish Innate Immune Response to Viral Infection-An Overview of Five Major Antiviral Genes"
1770:
Dale OB, Ørpetveit I, Lyngstad TM, Kahns S, Skall HF, Olesen NJ, Dannevig BH (June 2009).
1114:. United States Department of the Interior, U.S. Fish and Wildlife Service. Archived from
8:
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131:
43:
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2020:
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730:, which connects the Great Lakes to the Atlantic Ocean. VHS was further detected in the
73:. It was first discovered in the US in 1988 among salmon returning from the Pacific in
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On May 24, 2007, preliminary tests indicated the presence of VHS in a brown trout from
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330:(IHNV)), which causes infectious hematopoietic necrosis (IHN) disease in salmonidae.
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1913:
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42:. It afflicts over 50 species of freshwater and marine fish in several parts of the
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are developing a test that will speed diagnosis from a month to a matter of hours.
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391:
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Section requires expansion or clarification but source is not publicly accessible.
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3419:
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3012:
2654:"Viral Hemorrhagic Septicemia and Spring Viremia of Carp: Threats to Aquaculture"
1666:
Nishizawa T, Savas H, Isidan H, Ustündağ C, Iwamoto H, Yoshimizu M (April 2006).
1240:"Implementation of taxon-wide non-Latinized binomial species names in the family
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715:
705:
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To keep track of the distribution of different VHSV genotypes, a database called
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and northern France, farmed turbot in the UK and Ireland, and Greenland halibut (
511:
453:
407:. A new role of NV protein has been discovered and demonstrated that it inhibits
312:
300:
244:
220:
196:
2542:
2085:
1312:(Report). Great Lakes Aquatic Nonindigenous Species Information System (GLANSIS)
3359:
3047:
2961:
2860:"European Union Reference Laboratory for Fish Diseases Fish Pathogens Database"
2528:"Feature Story: Viral Hemorrhagic Septicemia: A New Invader in the Great Lakes"
2051:"European Union Reference Laboratory for Fish Diseases Fish Pathogens Database"
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would normally have a clear membrane. The liver may be pale, mottled with red
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424:
395:
374:
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338:
208:
128:
62:
2883:
2744:
2282:"Lake Winnebago Fish Preliminarily Test Positive for VHS]. (News release)"
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3202:
2935:
1537:
Nishizawa T, Iida H, Takano R, Isshiki T, Nakajima K, Muroga K (March 2002).
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The Type IV-b strain of VHSV has been spreading among freshwater fish in the
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548:
449:
306:
232:
184:
98:
2868:
Great Lakes fish hatcheries could benefit from new test for deadly VHS virus
2158:
3239:
3183:
2880:. Lists general information and resources on Viral hemorrhagic septicemia.
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2229:
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1992:
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35:
1965:
Isshik T, Nishizawa T, Kobayashi T, Nagano T, Miyazaki T (November 2001).
1819:. Copenhagen: International Council for the Exploration of the Sea. 2006.
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1623:
1172:
Meyers TR, Sullivan J, Emmenegger E, Follett J, Short S, Batts WN (1992).
595:. 1996 saw the first VHSV in Japan, among Japanese flounder farmed in the
3245:
3084:
2955:
1502:
1399:
1382:
Biacchesi S, Lamoureux A, Mérour E, Bernard J, Brémont M (October 2010).
895:
785:
689:
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619:
94:
1624:"Evolution of the fish rhabdovirus viral haemorrhagic septicaemia virus"
871:("pop eye"), pale or red-dotted gills, sunken eyes, and bleeding around
676:. The isolate was named MI03GL and was sequenced for its entire genome.
3339:
3219:
2940:
719:
701:
537:
465:
366:
86:
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2118:
1983:
1966:
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934:. Skeletal muscle accumulates blood but does not suffer much damage.
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open sores that may look like the lesions from other diseases or from
3374:
3354:
3344:
3258:
3207:
3152:
3102:
2569:"New fish virus could be a little as 2 years away from Lake Michigan"
1788:
1771:
1143:"Description of Viruses: Family Rhabdoviridae, Genus Novirhabdovirus"
904:
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disease, meaning it causes bleeding. Internally, the virus can cause
693:
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526:
477:
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408:
172:
90:
82:
66:
58:
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3454:
3413:
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2373:"Trout from Lake Michigan likely VHS positive]. (News release)"
899:
789:
734:, which connects Lake Erie to Lake Ontario. It was also found in a
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342:
78:
1621:
2950:
2627:
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492:
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On July 14, 2007, federal labs confirmed the presence of VHS in
93:
have occurred among a wide variety of freshwater species in the
3287:
3158:
2859:
1381:
1330:
1171:
966:
962:
947:
913:
629:), as well as dozens of freshwater species in the Great Lakes.
558:
522:
503:
354:
2050:
1964:
1942:(Report). Commonwealth of Australia. June 2003. Archived from
1622:
Einer-Jensen K, Ahrens P, Forsberg R, Lorenzen N (May 2004).
872:
481:
159:
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announced the likely presence of VHS in Wisconsin's inland
641:
2773:
2600:"Great Lakes Article: VHS disease has entered Great Lakes"
1769:
1725:
1536:
780:, a popular fishing destination is in the central part of
2005:
333:
The viral cause of the disease was discovered in 1963 by
71:
European Community Reference Laboratory for Fish Diseases
69:
in Europe and is therefore being watched closely by the
57:
Historically, VHS was associated mostly with freshwater
2226:
New York State Department of Environmental Conservation
776:
confirmed the presence of VHS in the Michigan's inland
120:
664:
region since at least 2003, resulting in some massive
19:
1251:
International Committee on Taxonomy of Viruses (ICTV)
1218:
International Committee on Taxonomy of Viruses (ICTV)
577:
Freshwater fish in North American Great Lakes region
2343:"Fish Disease Discovered in Budd Lake, Clare County"
683:
2774:Ortega-Villaizan MD, Chico V, Perez L (July 2022).
1860:"Viral Hemorrhagic Septicemia (VHS) Briefing Paper"
889:
2139:
2117:. Colorado Aquaculture Association. Archived from
1484:
1071:"Disease Factsheets: Viral Hemorrhagic Septicemia"
672:in 2005, and then in an archived 2003 sample from
2840:"Fish-Killing Virus Spreading in the Great Lakes"
3515:
2876:, National Invasive Species Information Center,
2222:"Viral Hemorrhagic Septicemia (VHS) in New York"
2192:"Viral Hemorrhagic Septicemia in New York State"
1940:Department of Agriculture Fisheries and Forestry
1078:The Center for Food Security & Public Health
972:
937:
644:pathogens (including VHSV) and their sequences.
316:. Another related fish rhabdovirus in the genus
2913:
2465:"Michigan's Summer Fishing Turns Less Carefree"
688:An archived 2003 sample from Lake St. Clair of
2874:Species Profile - Viral Hemorrhagic Septicemia
1910:Manual of Diagnostic Tests for Aquatic animals
1906:"Chapter 2.1.5: Viral Hemorrhagic Septicaemia"
1068:
867:External signs may include darker coloration,
2899:
2711:Michigan Lakes & Streams Association, Inc
2063:
1661:
1659:
1300:
2258:(Press release). 12 May 2007. Archived from
1532:
1530:
1137:
1135:
1133:
914:Histopathology (microscopic tissue analysis)
640:has been created to store data on different
2878:United States National Agricultural Library
2730:
2647:
2645:
2404:"Federal Lab Confirms VHS Caused Fish Kill"
1296:
1294:
381:(G), a unique non-virion protein (NV), and
2906:
2892:
2140:Ammayappan A, Vakharia VN (October 2009).
2105:
2103:
1656:
1204:
1105:
521:Rainbow trout in Georgia, farmed and wild
119:
2801:
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2377:Wisconsin Department of Natural Resources
2286:Wisconsin Department of Natural Resources
2256:Wisconsin Department of Natural Resources
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1485:Ammayappan A, Vakharia VN (August 2011).
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1026:Learn how and when to remove this message
2833:Michigan Department of Natural Resources
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2347:Michigan Department of Natural Resources
1899:
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1867:Michigan Department of Natural Resources
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1719:
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1615:
1301:Kipp RM, Ricciardi A (8 December 2006).
1291:
1108:"Viral Hemorrhagic Septicemia of Fishes"
813:
655:
557:) in Greenland. Farmed rainbow trout in
357:of VHSV is composed of approximately 11-
296:negative-sense single-stranded RNA virus
18:
2614:
2100:
1429:
1427:
328:Infectious hematopoietic necrosis virus
3516:
2837:
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1672:Applied and Environmental Microbiology
1237:
1101:
1099:
1097:
1095:
1053:
3395:
3394:
3026:Photobacterium damselae ssp piscicida
2985:Hypodermal and hematopoietic necrosis
2887:
2491:"VHS Surveillance Planning Under Way"
2438:Ontario Ministry of Natural Resources
2066:Molecular Phylogenetics and Evolution
1903:
1888:
1716:
1612:
1207:"Rename two (2) species in the genus
348:
1804:
1424:
1273:"Viral hemorrhagic septicemia virus"
976:
1092:
875:(eye sockets) and at base of fins.
13:
3446:Viral hemorrhagic septicemia virus
2829:Viral Hemorrhagic Septicemia (VHS)
2313:"Michigan's Best Bets for Muskies"
1303:Viral Hemorrhagic Septicemia (VHS)
1205:Kurath G, Kuhn JH (18 July 2016).
859:
784:. A major die-off of VHS-positive
285:Viral hemorrhagic septicemia virus
40:Viral hemorrhagic septicemia virus
14:
3540:
2822:
2310:
738:die-off in the landlocked inland
684:Great Lakes regional distribution
440:Prevalent host type and location
414:
104:
3309:Diseases and parasites in salmon
2623:"Viral Haemorrhagic Septicaemia"
2440:. 8 January 2007. Archived from
2029:10.1111/j.1365-2761.2006.00755.x
1749:10.1111/j.1365-2761.2007.00802.x
1069:Rovid-Spickler A (17 May 2007).
981:
890:Gross pathology (non-laboratory)
614:Gasterosteus aculeatus aculeatus
146:
2870:, Sarah Coefield, June 18, 2009
2767:
2724:
2696:
2682:
2620:
2591:
2519:
2482:
2456:
2426:
2395:
2365:
2335:
2304:
2274:
2244:
2214:
2184:
2133:
2043:
1999:
1958:
1928:
1763:
1692:10.1128/AEM.72.4.2373-2378.2006
1628:The Journal of General Virology
1571:
1478:
1375:
918:Preliminary diagnosis involves
822:
3365:Neurotoxic shellfish poisoning
2990:Infectious pancreatic necrosis
2567:Boomgaard J (9 January 2007).
1858:Whelan GE (26 February 2007).
1350:10.12688/f1000research.12985.2
1324:
1265:
1165:
280:Oncorhynchus 2 novirhabdovirus
1:
3370:Paralytic shellfish poisoning
3350:Diarrheal shellfish poisoning
3304:Diseases and parasites in cod
3277:Tetracapsuloides bryosalmonae
3148:Epizootic ulcerative syndrome
2379:. 24 May 2007. Archived from
2349:. 17 May 2007. Archived from
2288:. 18 May 2007. Archived from
1971:Diseases of Aquatic Organisms
1776:Diseases of Aquatic Organisms
1584:Diseases of Aquatic Organisms
1543:Diseases of Aquatic Organisms
1178:Diseases of Aquatic Organisms
1046:
973:Immune reactions against VHSV
956:
938:Virology (definitive testing)
403:, it is highly essential for
23:VHS disease in a gizzard shad
2598:Eckert M (26 January 2007).
878:Genetics researchers at the
854:
647:
554:Reinhardtius hippoglossoides
28:Viral hemorrhagic septicemia
7:
3335:Amnesic shellfish poisoning
2915:Fish diseases and parasites
2704:"Michigan vs the VHS virus"
2086:10.1016/j.ympev.2014.04.002
1001:. The specific problem is:
838:
755:Little Lake Butte des Morts
10:
3545:
3314:Disease in ornamental fish
2489:Gustafson L (April 2007).
782:Michigan's lower peninsula
3444:
3403:
3327:
3319:List of aquarium diseases
3296:
3076:
2921:
2745:10.1016/j.fsi.2019.06.015
2463:Sander L (13 July 2007).
2402:Figura D (14 July 2007).
1456:10.1007/s10126-010-9329-4
1238:Walker P (15 June 2015).
930:, and the liver may have
880:Lake Erie Research Center
796:began on April 30, 2007.
742:, the westernmost of the
273:
266:
141:
127:
118:
111:
34:) is a deadly infectious
3380:Scombroid food poisoning
2009:Journal of Fish Diseases
1729:Journal of Fish Diseases
502:Farmed rainbow trout in
436:
323:Salmonid novirhabdovirus
3405:Piscine novirhabdovirus
3265:Sphaerothecum destruens
3253:Schistocephalus solidus
3197:Lernaeocera branchialis
3043:Spring viraemia of carp
2838:Saulny S (2007-04-21).
2604:Port Huron Times-Herald
2159:10.1186/1743-422X-6-171
1080:. Iowa State University
690:Great Lakes muskellunge
259:Piscine novirhabdovirus
136:Piscine novirhabdovirus
113:Piscine novirhabdovirus
3226:Nanophyetus salmincola
3116:Clinostomum marginatum
2968:Hematopoietic necrosis
2936:Nervous necrosis virus
1277:NCBI Taxonomy Database
1149:. 2000. Archived from
1106:McAllister PE (1990).
456:in continental Europe
89:. Since 2005, massive
77:. This North American
24:
3123:Dactylogyrus vastator
3001:Mycobacterium marinum
2930:Aeromonas salmonicida
2610:on 27 September 2007.
2115:CSU Research Proposal
1641:10.1099/vir.0.79820-0
814:Government regulation
772:On May 17, 2007, the
749:On May 12, 2007, the
700:to the south, and to
656:Great Lakes Type IV-b
623:), and striped bass (
608:Fundulus heteroclitus
491:Farmed rainbow trout
22:
3271:Swim bladder disease
3233:Pseudorhabdosynochus
3214:Myxobolus cerebralis
3166:Gyrodactylus salaris
3091:Amoebic gill disease
3020:Pfiesteria piscicida
2974:Heterosigma akashiwo
2733:Genes & Genomics
2720:on 18 February 2012.
2652:Groocock GH (2007).
2579:on 27 September 2007
2573:Ludington Daily News
2548:on 26 September 2007
1946:on 27 September 2007
1503:10.1128/JVI.00597-11
1436:Marine Biotechnology
1400:10.1128/JVI.00343-10
1153:on 27 September 2007
1112:Fish Disease Leaflet
1008:improve this article
997:to meet Knowledge's
965:AQUATIC). Chlorine
884:University of Toledo
143:Virus classification
3529:Fish viral diseases
3173:Henneguya zschokkei
3037:Streptococcus iniae
2541:(5). Archived from
2526:Phillips K (2007).
2353:on 14 December 2008
2323:on 27 February 2009
2078:2014MolPE..77...34H
2021:2006JFDis..29..611E
1876:on 17 February 2007
1836:on 31 October 2007.
1741:2007JFDis..30..213G
1684:2006ApEnM..72.2373N
1634:(Pt 5): 1167–1179.
1491:Journal of Virology
1448:2011MarBt..13..672A
1394:(19): 10038–10050.
1388:Journal of Virology
547:Marine fish of the
536:Marine fish of the
464:Marine fish of the
132:electron micrograph
50:, and the virus as
44:Northern Hemisphere
3006:Aquarium granuloma
2866:Great Lakes Echo:
2844:The New York Times
2469:The New York Times
2444:on 8 February 2007
2414:on 12 October 2007
2317:Michigan Sportsman
2121:on 3 February 2007
1916:on 6 February 2007
728:St. Lawrence River
405:viral pathogenesis
349:Molecular virology
337:. The virus is an
25:
3524:Novirhabdoviruses
3511:
3510:
3397:Taxon identifiers
3388:
3387:
3109:Ceratomyxa shasta
2793:10.3390/v14071546
2507:on 20 August 2007
2408:The Post-Standard
1984:10.3354/dao047087
1597:10.3354/dao052029
1556:10.3354/dao048143
1497:(16): 8393–8402.
1191:10.3354/dao012167
1036:
1035:
1028:
999:quality standards
990:This article may
920:histopathological
581:
580:
401:viral replication
292:
291:
3536:
3504:
3503:
3491:
3490:
3478:
3477:
3465:
3464:
3463:
3437:
3436:
3424:
3423:
3422:
3392:
3391:
3179:Ich (freshwater)
3143:Eustrongylidosis
3130:Diphyllobothrium
2995:Koi herpes virus
2980:Hole in the head
2946:Enteric redmouth
2908:
2901:
2894:
2885:
2884:
2863:
2855:
2847:
2816:
2815:
2805:
2795:
2771:
2765:
2764:
2728:
2722:
2721:
2719:
2713:. Archived from
2708:
2700:
2694:
2693:
2686:
2680:
2679:
2677:
2675:
2669:
2663:. Archived from
2658:
2649:
2640:
2639:
2637:
2635:
2618:
2612:
2611:
2606:. Archived from
2595:
2589:
2588:
2586:
2584:
2575:. Archived from
2564:
2558:
2557:
2555:
2553:
2547:
2532:
2523:
2517:
2516:
2514:
2512:
2506:
2500:. Archived from
2495:
2486:
2480:
2479:
2477:
2475:
2460:
2454:
2453:
2451:
2449:
2430:
2424:
2423:
2421:
2419:
2410:. Archived from
2399:
2393:
2392:
2390:
2388:
2383:on 6 August 2007
2369:
2363:
2362:
2360:
2358:
2339:
2333:
2332:
2330:
2328:
2319:. Archived from
2308:
2302:
2301:
2299:
2297:
2278:
2272:
2271:
2269:
2267:
2248:
2242:
2241:
2239:
2237:
2228:. Archived from
2218:
2212:
2211:
2209:
2207:
2188:
2182:
2181:
2171:
2161:
2146:Virology Journal
2137:
2131:
2130:
2128:
2126:
2107:
2098:
2097:
2061:
2055:
2054:
2047:
2041:
2040:
2003:
1997:
1996:
1986:
1962:
1956:
1955:
1953:
1951:
1932:
1926:
1925:
1923:
1921:
1912:. Archived from
1904:Crane M (2006).
1901:
1886:
1885:
1883:
1881:
1875:
1869:. Archived from
1864:
1855:
1838:
1837:
1835:
1829:. Archived from
1818:
1808:
1802:
1801:
1791:
1789:10.3354/dao02065
1767:
1761:
1760:
1723:
1714:
1713:
1703:
1678:(4): 2373–2378.
1663:
1654:
1653:
1643:
1619:
1610:
1609:
1599:
1575:
1569:
1568:
1558:
1534:
1525:
1524:
1514:
1482:
1476:
1475:
1431:
1422:
1421:
1411:
1379:
1373:
1372:
1362:
1352:
1328:
1322:
1321:
1319:
1317:
1307:
1298:
1289:
1288:
1286:
1284:
1269:
1263:
1262:
1260:
1258:
1248:
1235:
1229:
1228:
1226:
1224:
1215:
1202:
1196:
1195:
1193:
1169:
1163:
1162:
1160:
1158:
1139:
1128:
1127:
1125:
1123:
1103:
1090:
1089:
1087:
1085:
1075:
1066:
1031:
1024:
1020:
1017:
1011:
985:
984:
977:
808:Skaneateles Lake
711:2005 samples of
638:Fishpathogens.eu
626:Morone saxatilis
617:), brown trout (
611:), stickleback (
452:and a few other
434:
433:
392:Reverse genetics
383:virus polymerase
151:
150:
123:
109:
108:
75:Washington state
3544:
3543:
3539:
3538:
3537:
3535:
3534:
3533:
3514:
3513:
3512:
3507:
3499:
3494:
3486:
3481:
3473:
3468:
3459:
3458:
3453:
3440:
3432:
3427:
3418:
3417:
3412:
3399:
3389:
3384:
3323:
3292:
3190:Kudoa thyrsites
3137:Cymothoa exigua
3072:
3013:Novirhabdovirus
2917:
2912:
2858:
2850:
2825:
2820:
2819:
2772:
2768:
2729:
2725:
2717:
2706:
2702:
2701:
2697:
2692:. 18 June 2009.
2688:
2687:
2683:
2673:
2671:
2667:
2656:
2650:
2643:
2633:
2631:
2619:
2615:
2596:
2592:
2582:
2580:
2565:
2561:
2551:
2549:
2545:
2530:
2524:
2520:
2510:
2508:
2504:
2493:
2487:
2483:
2473:
2471:
2461:
2457:
2447:
2445:
2432:
2431:
2427:
2417:
2415:
2400:
2396:
2386:
2384:
2371:
2370:
2366:
2356:
2354:
2341:
2340:
2336:
2326:
2324:
2309:
2305:
2295:
2293:
2292:on 25 June 2007
2280:
2279:
2275:
2265:
2263:
2262:on 24 June 2007
2250:
2249:
2245:
2235:
2233:
2220:
2219:
2215:
2205:
2203:
2202:on 10 June 2007
2190:
2189:
2185:
2138:
2134:
2124:
2122:
2109:
2108:
2101:
2062:
2058:
2049:
2048:
2044:
2015:(10): 611–619.
2004:
2000:
1963:
1959:
1949:
1947:
1934:
1933:
1929:
1919:
1917:
1902:
1889:
1879:
1877:
1873:
1862:
1856:
1841:
1833:
1827:
1816:
1810:
1809:
1805:
1768:
1764:
1724:
1717:
1664:
1657:
1620:
1613:
1576:
1572:
1535:
1528:
1483:
1479:
1432:
1425:
1380:
1376:
1329:
1325:
1315:
1313:
1305:
1299:
1292:
1282:
1280:
1271:
1270:
1266:
1256:
1254:
1246:
1236:
1232:
1222:
1220:
1213:
1209:Novirhabdovirus
1203:
1199:
1170:
1166:
1156:
1154:
1141:
1140:
1131:
1121:
1119:
1118:on 15 June 2007
1104:
1093:
1083:
1081:
1073:
1067:
1054:
1049:
1032:
1021:
1015:
1012:
1005:
986:
982:
975:
959:
940:
932:pyknotic nuclei
916:
892:
862:
860:Field diagnosis
857:
841:
825:
816:
716:freshwater drum
706:St. Clair River
686:
658:
650:
602:Clupea harengus
597:Seto Inland Sea
512:Gulf of Bothnia
454:freshwater fish
417:
351:
318:Novirhabdovirus
313:Novirhabdovirus
301:Mononegavirales
262:
246:Novirhabdovirus
222:Mononegavirales
198:Negarnaviricota
145:
107:
17:
16:Disease of fish
12:
11:
5:
3542:
3532:
3531:
3526:
3509:
3508:
3506:
3505:
3492:
3479:
3466:
3450:
3448:
3442:
3441:
3439:
3438:
3425:
3409:
3407:
3401:
3400:
3386:
3385:
3383:
3382:
3377:
3372:
3367:
3362:
3360:Marine viruses
3357:
3352:
3347:
3342:
3337:
3331:
3329:
3328:Related topics
3325:
3324:
3322:
3321:
3316:
3311:
3306:
3300:
3298:
3294:
3293:
3291:
3290:
3285:
3280:
3273:
3268:
3261:
3256:
3249:
3242:
3237:
3229:
3222:
3217:
3210:
3205:
3200:
3193:
3186:
3181:
3176:
3169:
3162:
3155:
3150:
3145:
3140:
3133:
3126:
3119:
3112:
3105:
3100:
3093:
3088:
3080:
3078:
3074:
3073:
3071:
3070:
3065:
3060:
3055:
3050:
3048:Taura syndrome
3045:
3040:
3033:
3028:
3023:
3016:
3009:
2997:
2992:
2987:
2982:
2977:
2970:
2965:
2962:Flavobacterium
2958:
2953:
2948:
2943:
2938:
2933:
2925:
2923:
2919:
2918:
2911:
2910:
2903:
2896:
2888:
2882:
2881:
2871:
2864:
2856:
2848:
2835:
2824:
2823:External links
2821:
2818:
2817:
2766:
2723:
2695:
2681:
2670:on 7 June 2007
2641:
2613:
2590:
2559:
2518:
2481:
2455:
2425:
2394:
2364:
2334:
2303:
2273:
2243:
2232:on 6 July 2007
2213:
2183:
2132:
2099:
2056:
2042:
1998:
1957:
1927:
1887:
1839:
1825:
1803:
1762:
1735:(4): 213–223.
1715:
1655:
1611:
1570:
1549:(2): 143–148.
1526:
1477:
1442:(4): 672–683.
1423:
1374:
1323:
1290:
1264:
1230:
1197:
1164:
1129:
1091:
1051:
1050:
1048:
1045:
1034:
1033:
989:
987:
980:
974:
971:
958:
955:
939:
936:
915:
912:
891:
888:
861:
858:
856:
853:
840:
837:
829:emerald shiner
824:
821:
815:
812:
759:Lake Winnebago
685:
682:
674:Lake St. Clair
657:
654:
649:
646:
579:
578:
575:
571:
570:
567:
563:
562:
545:
541:
540:
534:
530:
529:
519:
515:
514:
500:
496:
495:
489:
485:
484:
462:
458:
457:
446:
442:
441:
438:
416:
415:Virus subtypes
413:
396:vaccinia virus
375:matrix protein
371:phosphoprotein
350:
347:
290:
289:
288:
287:
282:
277:
271:
270:
264:
263:
256:
254:
250:
249:
242:
238:
237:
230:
226:
225:
218:
214:
213:
210:Monjiviricetes
206:
202:
201:
194:
190:
189:
182:
178:
177:
170:
163:
162:
157:
153:
152:
139:
138:
129:Negative stain
125:
124:
116:
115:
106:
105:Virus taxonomy
103:
63:western Europe
48:Egtved disease
15:
9:
6:
4:
3:
2:
3541:
3530:
3527:
3525:
3522:
3521:
3519:
3502:
3497:
3493:
3489:
3484:
3480:
3476:
3471:
3467:
3462:
3456:
3452:
3451:
3449:
3447:
3443:
3435:
3430:
3426:
3421:
3415:
3411:
3410:
3408:
3406:
3402:
3398:
3393:
3381:
3378:
3376:
3373:
3371:
3368:
3366:
3363:
3361:
3358:
3356:
3353:
3351:
3348:
3346:
3343:
3341:
3338:
3336:
3333:
3332:
3330:
3326:
3320:
3317:
3315:
3312:
3310:
3307:
3305:
3302:
3301:
3299:
3295:
3289:
3286:
3284:
3281:
3279:
3278:
3274:
3272:
3269:
3267:
3266:
3262:
3260:
3257:
3255:
3254:
3250:
3248:
3247:
3243:
3241:
3238:
3235:
3234:
3230:
3228:
3227:
3223:
3221:
3218:
3216:
3215:
3211:
3209:
3206:
3204:
3203:Microsporidia
3201:
3199:
3198:
3194:
3192:
3191:
3187:
3185:
3182:
3180:
3177:
3175:
3174:
3170:
3168:
3167:
3163:
3161:
3160:
3156:
3154:
3151:
3149:
3146:
3144:
3141:
3139:
3138:
3134:
3132:
3131:
3127:
3125:
3124:
3120:
3118:
3117:
3113:
3111:
3110:
3106:
3104:
3101:
3099:
3098:
3094:
3092:
3089:
3087:
3086:
3082:
3081:
3079:
3075:
3069:
3066:
3064:
3061:
3059:
3056:
3054:
3051:
3049:
3046:
3044:
3041:
3039:
3038:
3034:
3032:
3031:Salmon anemia
3029:
3027:
3024:
3022:
3021:
3017:
3015:
3014:
3010:
3007:
3003:
3002:
2998:
2996:
2993:
2991:
2988:
2986:
2983:
2981:
2978:
2976:
2975:
2971:
2969:
2966:
2964:
2963:
2959:
2957:
2954:
2952:
2949:
2947:
2944:
2942:
2939:
2937:
2934:
2932:
2931:
2927:
2926:
2924:
2920:
2916:
2909:
2904:
2902:
2897:
2895:
2890:
2889:
2886:
2879:
2875:
2872:
2869:
2865:
2861:
2857:
2853:
2849:
2845:
2841:
2836:
2834:
2830:
2827:
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1310:VHS Factsheet
1304:
1297:
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1274:
1268:
1252:
1245:
1243:
1242:Rhabdoviridae
1234:
1219:
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1187:
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801:Lake Michigan
797:
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794:black crappie
791:
787:
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775:
770:
768:
764:
763:Lake Michigan
760:
756:
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751:Wisconsin DNR
747:
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741:
737:
733:
732:Niagara River
729:
724:
721:
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695:
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450:rainbow trout
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377:(M), surface
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340:
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307:Rhabdoviridae
303:
302:
298:of the order
297:
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3184:Ich (marine)
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2736:
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2715:the original
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2665:the original
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2012:
2008:
2001:
1977:(2): 87–99.
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1970:
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1944:the original
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1878:. Retrieved
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1732:
1728:
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1326:
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1309:
1281:. Retrieved
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1267:
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1200:
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1146:
1120:. Retrieved
1116:the original
1111:
1082:. Retrieved
1077:
1037:
1022:
1013:
1006:Please help
1002:
991:
960:
952:
944:
941:
917:
909:
893:
877:
869:exophthalmia
866:
863:
850:
842:
833:
826:
823:Transmission
817:
805:
798:
774:Michigan DNR
771:
748:
744:Finger Lakes
740:Conesus Lake
725:
713:Lake Ontario
710:
704:through the
696:through the
687:
678:
670:Lake Ontario
659:
651:
635:
631:
624:
620:Salmo trutta
618:
612:
606:
600:
589:
582:
574: IV-b
566: IV-a
552:
418:
390:
387:
379:glycoprotein
369:-associated
363:nucleocapsid
352:
335:M. H. Jenson
332:
327:
321:
317:
311:
310:, and genus
305:
299:
293:
284:
279:
276:Egtved virus
258:
257:
245:
233:
221:
209:
197:
185:
173:
166:
156:(unranked):
135:
112:
56:
52:Egtved virus
51:
47:
39:
36:fish disease
31:
27:
26:
3297:Fish groups
3246:Saprolegnia
3240:Salmon lice
3085:Abergasilus
2956:Fish dropsy
2786:(7): 1546.
1253:. p. 7
1010:if you can.
896:hemorrhagic
662:Great Lakes
544: III
518: I-e
499: I-d
488: I-c
461: I-b
445: I-a
95:Great Lakes
3518:Categories
3340:Brevetoxin
3220:Myxosporea
3068:Yellowhead
3063:White spot
2941:Columnaris
2634:11 October
2535:Fish Lines
2196:Highlights
1184:: 167–75.
1047:References
1016:March 2022
957:Prevention
894:VHSV is a
720:Lake Huron
702:Lake Huron
538:Baltic Sea
533: II
466:Baltic Sea
419:Different
367:polymerase
326:(formerly
294:VHSV is a
97:region of
87:Baltic Sea
85:, and the
67:fish farms
38:caused by
3461:Q18965285
3420:Q29004930
3375:Saxitoxin
3355:Fish kill
3345:Ciguatera
3259:Sea louse
3208:Monogenea
3103:Carp lice
3077:Parasites
2922:Pathogens
2761:195760294
2311:Barta J.
2072:: 34–40.
905:hyperemic
900:petechial
855:Diagnosis
848:attacks.
790:bluegills
778:Budd Lake
767:Green Bay
694:Lake Erie
648:Evolution
593:North Sea
527:Black Sea
478:North Sea
470:Skagerrak
409:apoptosis
343:peplomers
339:enveloped
304:, family
253:Species:
181:Kingdom:
174:Riboviria
83:North Sea
59:salmonids
3475:11461179
3455:Wikidata
3414:Wikidata
3097:Anisakis
2812:35891526
2753:31721105
2178:19852863
2094:24727199
2037:17026670
1993:11775799
1798:19694169
1757:17394523
1710:16597932
1650:15105533
1606:12517003
1565:12005236
1521:21653667
1472:24597813
1464:20936318
1418:20631140
1369:29527292
1343:: 1958.
992:require
928:lamellae
839:Symptoms
666:die-offs
474:Kattegat
429:enzootic
421:isolates
268:Synonyms
229:Family:
193:Phylum:
91:die-offs
79:genotype
2951:Fin rot
2803:9317989
2780:Viruses
2674:18 July
2628:WikiVet
2583:13 July
2552:16 July
2511:16 July
2474:13 July
2448:15 July
2418:15 July
2387:13 July
2357:15 July
2327:15 July
2296:13 July
2266:13 July
2236:14 July
2206:14 July
2169:2771013
2152:: 171.
2125:12 July
2074:Bibcode
2017:Bibcode
1950:16 July
1920:16 July
1880:13 July
1737:Bibcode
1701:1449023
1680:Bibcode
1512:3147959
1444:Bibcode
1409:2937753
1360:5820608
1316:16 July
1283:14 July
1257:15 June
1223:15 June
1157:12 July
1122:12 July
1084:12 July
1040:viperin
994:cleanup
882:at the
846:lamprey
786:muskies
736:walleye
525:in the
508:Finland
493:Denmark
448:Farmed
241:Genus:
217:Order:
205:Class:
3501:816302
3288:Xenoma
3283:Velvet
3159:Glugea
3153:Flukes
2810:
2800:
2759:
2751:
2176:
2166:
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2035:
1991:
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1462:
1416:
1406:
1367:
1357:
967:bleach
963:Virkon
948:RT-PCR
873:orbits
792:, and
559:Norway
523:turbot
504:Norway
355:genome
3496:WoRMS
3488:11287
3470:IRMNG
3434:4J8KZ
2757:S2CID
2718:(PDF)
2707:(PDF)
2668:(PDF)
2657:(PDF)
2546:(PDF)
2531:(PDF)
2505:(PDF)
2494:(PDF)
1874:(PDF)
1863:(PDF)
1834:(PDF)
1817:(PDF)
1468:S2CID
1306:(PDF)
1247:(PDF)
1214:(PDF)
1074:(PDF)
1038:Fish
482:Japan
437:Type
385:(L).
373:(P),
167:Realm
160:Virus
3483:NCBI
3236:spp.
2808:PMID
2749:PMID
2676:2007
2636:2011
2585:2007
2554:2007
2513:2007
2476:2007
2450:2007
2420:2007
2389:2007
2359:2007
2329:2007
2298:2007
2268:2007
2238:2007
2208:2007
2174:PMID
2127:2007
2090:PMID
2033:PMID
1989:PMID
1952:2007
1922:2007
1882:2007
1821:ISBN
1794:PMID
1753:PMID
1706:PMID
1646:PMID
1602:PMID
1561:PMID
1517:PMID
1460:PMID
1414:PMID
1365:PMID
1318:2007
1285:2007
1259:2019
1225:2019
1159:2007
1124:2007
1086:2007
718:and
642:fish
585:pike
425:host
353:The
3429:CoL
3058:VHS
3053:UDN
2798:PMC
2788:doi
2741:doi
2164:PMC
2154:doi
2082:doi
2025:doi
1979:doi
1784:doi
1745:doi
1696:PMC
1688:doi
1636:doi
1592:doi
1551:doi
1507:PMC
1499:doi
1452:doi
1404:PMC
1396:doi
1355:PMC
1345:doi
1186:doi
765:'s
320:is
134:of
61:in
32:VHS
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