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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
854:
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
409:
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
829:
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
690:
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,
643:
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
601:
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,
691:
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.
2017:
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".
956:
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
1737:
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".
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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.
733:
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.
1445:
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".
438:
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
875:
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.
594:
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
814:, 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.
972:
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
679:
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
399:
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.
81:
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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
980:
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.
768:. 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
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1870:
1395:"Limited interference at the early stage of infection between two recombinant novirhabdoviruses: viral hemorrhagic septicemia virus and infectious hematopoietic necrosis virus"
2202:
1947:
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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1283:
92:
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
57:. 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
2995:
2742:
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.
2501:
1153:
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1290:. United States Department of Health and Human Services, National Institutes of Health, National Library of Medicine, National Center for Biotechnology Information
610:, 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 (
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2916:
1822:
1018:
1824:
Report of the ICES Advisory Committee on Fishery Management, Advisory Committee on the Marine Environment and Advisory Committee on Ecosystems, 2006. Book 2
2888:
2323:
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1679:"Genotyping and pathogenicity of viral hemorrhagic septicemia virus from free-living turbot (Psetta maxima) in a Turkish coastal area of the Black Sea"
2383:
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784:
306:
1946:
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2263:"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"
1550:"Genetic relatedness among Japanese, American and European isolates of viral hemorrhagic septicemia virus (VHSV) based on partial G and P genes"
2075:
He M, Yan XC, Liang Y, Sun XW, Teng CB (August 2014). "Evolution of the viral hemorrhagic septicemia virus: divergence, selection and origin".
663:
This virus appears to have evolved in the Pacific Northwest of North America. It appears to have diverged into several strains ~300 years ago.
2209:. Cornell University, College of Veterinary Medicine, Department of Microbiology and Immunology, Aquatic Animal Health Program. Archived from
2579:
821:, 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.
405:
is a powerful tool to study and characterize the previously unknown viral genes. Reverse genetics system is currently available for VHSV. A
2610:
838:
VHSV can be spread from fish to fish through water transfer, as well as through contaminated eggs, and bait fish from infected waters. The
1185:"Identification of viral hemorrhagic septicemia virus isolated from Pacific cod Gadus macrocephalus in Prince William Sound, Alaska, USA"
937:(blood-forming) areas of the kidney and spleen are the initial area of infection, and should show necrosis. The gill may have thickened
2444:
1881:
434:(unique strains) of VHSV are typically grouped by genotyping. It is found that genotype groups are divided more geographically than by
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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
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1978:"An outbreak of VHSV (viral hemorrhagic septicemia virus) infection in farmed Japanese flounder Paralichthys olivaceus in Japan"
1916:
1783:"Outbreak of viral haemorrhagic septicaemia (VHS) in seawater-farmed rainbow trout in Norway caused by VHS virus Genotype III"
1344:"Identification of diverse defense mechanisms in rainbow trout red blood cells in response to halted replication of VHS virus"
3375:
3041:
3000:
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Cornell University, College of Veterinary Medicine, Department of Microbiology and Immunology, Aquatic Animal Health Program
422:
at the early stage of viral infection. This discovery unlocked the mystery of presence of NV proteins in novirhabdoviruses.
31:
2512:
846:
heron has spread the virus, but it does so mechanically; the virus is apparently inactive in the digestive tract of birds.
757:
in western New York state. This was the first case in the region outside waters contiguously connected to the Great Lakes.
372:
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′,
158:
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76:, 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.
772:. Preliminary tests later indicated VHS in specimens from Lake Winnebago. The lakes drain through the Fox River to
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2121:
938:
777:
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352:, bullet-shaped particle, about 180 nm long by 60 nm in diameter, covered with 5 to 15 nm long
333:
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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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3207:
3053:
953:
Electron microscopy can reveal the bullet-shaped rhabdovirus, but is not adequate for definitive diagnosis.
719:
to the north. The sample was not tested for VHS until 2005, after the disease was detected in Lake Ontario.
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Marine fish of the Northwest Pacific (North America), North American north Atlantic coast, Japan, and Korea
2884:
2862:
2391:
2300:
2270:
410:(NV) gene of novirhabdoviruses. Even though it has been demonstrated that the NV gene is not necessary for
3407:
3236:
3126:
3005:
2153:"Molecular characterization of the Great Lakes viral hemorrhagic septicemia virus (VHSV) isolate from USA"
3293:
3133:
3011:
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1954:
618:
1591:"Phylogenetic analysis of viral haemorrhagic septicaemia virus (VHSV) isolates from France (1971-1999)"
2701:"Great Lakes fish hatcheries could benefit from new test for deadly VHS virus | Great Lakes Echo"
393:
3511:
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3176:
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2445:"Ontario Government Acting Swiftly to Control Further Spread of New Fish Virus]. (Press release)"
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788:
153:
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is a particularly popular bait fish in the Great Lakes region, and is among the species afflicted.
142:
54:
1342:
Nombela I, Puente-Marin S, Chico V, Villena AJ, Carracedo B, Ciordia S, et al. (2018-02-09).
345:
3078:
3016:
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1498:"Nonvirion protein of novirhabdovirus suppresses apoptosis at the early stage of virus infection"
1478:
961:) and genetic sequencing and comparison for definitive confirmation and genotype classification.
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is the earliest confirmed case of VHSV within the Great Lakes region. Lake St. Clair connects to
415:
278:
85:
2618:
3485:
3439:
3119:
2863:"Viral Hemorrhagic Septicemia | National Invasive Species Information Center | USDA"
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2122:"Economic impacts of private sector aquaculture-based recreational fishing in the western USA"
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1999:
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2006 saw mass die-offs from VHS in Lake Erie, the St. Clair River, the Detroit River, and the
411:
1589:
Thiéry R, de Boisséson C, Jeffroy J, Castric J, de Kinkelin P, Benmansour A (November 2002).
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1989:
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Virus neutralisation is another important method of diagnosis, especially for carrier fish.
818:
648:
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431:
402:
1158:
Virus Taxonomy Online: Seventh Report of the International Committee on Taxonomy of Viruses
3200:
3147:
3023:
2787:"Fish Innate Immune Response to Viral Infection-An Overview of Five Major Antiviral Genes"
1781:
Dale OB, Ørpetveit I, Lyngstad TM, Kahns S, Skall HF, Olesen NJ, Dannevig BH (June 2009).
1125:. United States Department of the Interior, U.S. Fish and Wildlife Service. Archived from
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741:, which connects the Great Lakes to the Atlantic Ocean. VHS was further detected in the
84:. 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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17:
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341:(IHNV)), which causes infectious hematopoietic necrosis (IHN) disease in salmonidae.
317:
243:
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109:
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1924:
1702:
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53:. 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.
879:
754:
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723:
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373:
46:
1014:
Section requires expansion or clarification but source is not publicly accessible.
3471:
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2665:"Viral Hemorrhagic Septicemia and Spring Viremia of Carp: Threats to Aquaculture"
1677:
Nishizawa T, Savas H, Isidan H, Ustündağ C, Iwamoto H, Yoshimizu M (April 2006).
1251:"Implementation of taxon-wide non-Latinized binomial species names in the family
647:
To keep track of the distribution of different VHSV genotypes, a database called
562:
and northern France, farmed turbot in the UK and Ireland, and Greenland halibut (
418:. A new role of NV protein has been discovered and demonstrated that it inhibits
3256:
3095:
2966:
2553:
2096:
1323:(Report). Great Lakes Aquatic Nonindigenous Species Information System (GLANSIS)
906:
796:
700:
672:
630:
105:
2871:"European Union Reference Laboratory for Fish Diseases Fish Pathogens Database"
2539:"Feature Story: Viral Hemorrhagic Septicemia: A New Invader in the Great Lakes"
2062:"European Union Reference Laboratory for Fish Diseases Fish Pathogens Database"
1126:
914:
would normally have a clear membrane. The liver may be pale, mottled with red
3350:
3230:
2951:
2894:
2755:
2293:"Lake Winnebago Fish Preliminarily Test Positive for VHS]. (News release)"
1466:
730:
712:
548:
476:
377:
97:
1548:
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
3385:
3365:
3355:
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3163:
3113:
2879:
Great Lakes fish hatcheries could benefit from new test for deadly VHS virus
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69:
2891:. Lists general information and resources on Viral hemorrhagic septicemia.
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2003:
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1575:
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1474:
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1976:
Isshik T, Nishizawa T, Kobayashi T, Nagano T, Miyazaki T (November 2001).
1830:. Copenhagen: International Council for the Exploration of the Sea. 2006.
1651:
1634:
1183:
Meyers TR, Sullivan J, Emmenegger E, Follett J, Short S, Batts WN (1992).
606:. 1996 saw the first VHSV in Japan, among Japanese flounder farmed in the
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3424:
3107:
1513:
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1393:
Biacchesi S, Lamoureux A, Mérour E, Bernard J, Brémont M (October 2010).
910:
800:
484:
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369:
353:
89:
1635:"Evolution of the fish rhabdovirus viral haemorrhagic septicaemia virus"
882:("pop eye"), pale or red-dotted gills, sunken eyes, and bleeding around
687:. The isolate was named MI03GL and was sequenced for its entire genome.
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2129:
1994:
1977:
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945:. Skeletal muscle accumulates blood but does not suffer much damage.
855:
open sores that may look like the lesions from other diseases or from
3298:
3169:
2580:"New fish virus could be a little as 2 years away from Lake Michigan"
1799:
1782:
1154:"Description of Viruses: Family Rhabdoviridae, Genus Novirhabdovirus"
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958:
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disease, meaning it causes bleeding. Internally, the virus can cause
569:
533:
514:
365:
3401:
2384:"Trout from Lake Michigan likely VHS positive]. (News release)"
745:, which connects Lake Erie to Lake Ontario. It was also found in a
883:
492:
170:
29:
817:
On July 14, 2007, federal labs confirmed the presence of VHS in
652:
104:
have occurred among a wide variety of freshwater species in the
3405:
2898:
2870:
640:), as well as dozens of freshwater species in the Great Lakes.
131:
2061:
1953:(Report). Commonwealth of Australia. June 2003. Archived from
987:
30:
1633:
Einer-Jensen K, Ahrens P, Forsberg R, Lorenzen N (May 2004).
764:
announced the likely presence of VHS in Wisconsin's inland
2611:"Great Lakes Article: VHS disease has entered Great Lakes"
791:, a popular fishing destination is in the central part of
344:
The viral cause of the disease was discovered in 1963 by
82:
European Community Reference Laboratory for Fish Diseases
80:
in Europe and is therefore being watched closely by the
68:
Historically, VHS was associated mostly with freshwater
2237:
New York State Department of Environmental Conservation
787:
confirmed the presence of VHS in the Michigan's inland
675:
region since at least 2003, resulting in some massive
1262:
International Committee on Taxonomy of Viruses (ICTV)
1229:
International Committee on Taxonomy of Viruses (ICTV)
588:
Freshwater fish in North American Great Lakes region
2354:"Fish Disease Discovered in Budd Lake, Clare County"
3455:
3414:
3338:
3307:
3087:
2932:
2785:Ortega-Villaizan MD, Chico V, Perez L (July 2022).
1871:"Viral Hemorrhagic Septicemia (VHS) Briefing Paper"
2128:. Colorado Aquaculture Association. Archived from
1082:"Disease Factsheets: Viral Hemorrhagic Septicemia"
683:in 2005, and then in an archived 2003 sample from
2851:"Fish-Killing Virus Spreading in the Great Lakes"
2887:, National Invasive Species Information Center,
2233:"Viral Hemorrhagic Septicemia (VHS) in New York"
2203:"Viral Hemorrhagic Septicemia in New York State"
1951:Department of Agriculture Fisheries and Forestry
1089:The Center for Food Security & Public Health
655:pathogens (including VHSV) and their sequences.
327:. Another related fish rhabdovirus in the genus
2476:"Michigan's Summer Fishing Turns Less Carefree"
699:An archived 2003 sample from Lake St. Clair of
2885:Species Profile - Viral Hemorrhagic Septicemia
1921:Manual of Diagnostic Tests for Aquatic animals
1917:"Chapter 2.1.5: Viral Hemorrhagic Septicaemia"
878:External signs may include darker coloration,
2910:
2722:Michigan Lakes & Streams Association, Inc
1672:
1670:
8:
2269:(Press release). 12 May 2007. Archived from
1543:
1541:
1148:
1146:
1144:
925:Histopathology (microscopic tissue analysis)
651:has been created to store data on different
2889:United States National Agricultural Library
2658:
2656:
2415:"Federal Lab Confirms VHS Caused Fish Kill"
1307:
1305:
392:(G), a unique non-virion protein (NV), and
3402:
2917:
2903:
2895:
2151:Ammayappan A, Vakharia VN (October 2009).
2116:
2114:
532:Rainbow trout in Georgia, farmed and wild
130:
119:
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2802:
2388:Wisconsin Department of Natural Resources
2297:Wisconsin Department of Natural Resources
2267:Wisconsin Department of Natural Resources
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1993:
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1854:
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1496:Ammayappan A, Vakharia VN (August 2011).
1418:
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1200:
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1067:
1037:Learn how and when to remove this message
2844:Michigan Department of Natural Resources
2358:Michigan Department of Natural Resources
1910:
1908:
1906:
1904:
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1878:Michigan Department of Natural Resources
1732:
1730:
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1626:
1312:Kipp RM, Ricciardi A (8 December 2006).
1119:"Viral Hemorrhagic Septicemia of Fishes"
568:) in Greenland. Farmed rainbow trout in
444:
368:of VHSV is composed of approximately 11-
307:negative-sense single-stranded RNA virus
1440:
1438:
1063:
339:Infectious hematopoietic necrosis virus
1683:Applied and Environmental Microbiology
1112:
1110:
1108:
1106:
3037:Photobacterium damselae ssp piscicida
2996:Hypodermal and hematopoietic necrosis
2502:"VHS Surveillance Planning Under Way"
2449:Ontario Ministry of Natural Resources
2077:Molecular Phylogenetics and Evolution
1218:"Rename two (2) species in the genus
7:
1284:"Viral hemorrhagic septicemia virus"
886:(eye sockets) and at base of fins.
3457:Viral hemorrhagic septicemia virus
2840:Viral Hemorrhagic Septicemia (VHS)
2324:"Michigan's Best Bets for Muskies"
1314:Viral Hemorrhagic Septicemia (VHS)
1216:Kurath G, Kuhn JH (18 July 2016).
795:. A major die-off of VHS-positive
296:Viral hemorrhagic septicemia virus
51:Viral hemorrhagic septicemia virus
25:
749:die-off in the landlocked inland
695:Great Lakes regional distribution
451:Prevalent host type and location
3320:Diseases and parasites in salmon
2634:"Viral Haemorrhagic Septicaemia"
2451:. 8 January 2007. Archived from
2040:10.1111/j.1365-2761.2006.00755.x
1760:10.1111/j.1365-2761.2007.00802.x
1080:Rovid-Spickler A (17 May 2007).
992:
901:Gross pathology (non-laboratory)
625:Gasterosteus aculeatus aculeatus
157:
2881:, Sarah Coefield, June 18, 2009
1703:10.1128/AEM.72.4.2373-2378.2006
1639:The Journal of General Virology
929:Preliminary diagnosis involves
3376:Neurotoxic shellfish poisoning
3001:Infectious pancreatic necrosis
2578:Boomgaard J (9 January 2007).
1869:Whelan GE (26 February 2007).
1361:10.12688/f1000research.12985.2
291:Oncorhynchus 2 novirhabdovirus
1:
3381:Paralytic shellfish poisoning
3361:Diarrheal shellfish poisoning
3315:Diseases and parasites in cod
3288:Tetracapsuloides bryosalmonae
3159:Epizootic ulcerative syndrome
2390:. 24 May 2007. Archived from
2360:. 17 May 2007. Archived from
2299:. 18 May 2007. Archived from
1982:Diseases of Aquatic Organisms
1787:Diseases of Aquatic Organisms
1595:Diseases of Aquatic Organisms
1554:Diseases of Aquatic Organisms
1189:Diseases of Aquatic Organisms
984:Immune reactions against VHSV
949:Virology (definitive testing)
414:, it is highly essential for
34:VHS disease in a gizzard shad
2609:Eckert M (26 January 2007).
889:Genetics researchers at the
565:Reinhardtius hippoglossoides
39:Viral hemorrhagic septicemia
3346:Amnesic shellfish poisoning
2926:Fish diseases and parasites
2715:"Michigan vs the VHS virus"
2097:10.1016/j.ympev.2014.04.002
1012:. The specific problem is:
766:Little Lake Butte des Morts
3556:
3325:Disease in ornamental fish
2500:Gustafson L (April 2007).
1008:to meet Knowledge (XXG)'s
793:Michigan's lower peninsula
3330:List of aquarium diseases
2756:10.1016/j.fsi.2019.06.015
2474:Sander L (13 July 2007).
2413:Figura D (14 July 2007).
1467:10.1007/s10126-010-9329-4
1249:Walker P (15 June 2015).
941:, and the liver may have
891:Lake Erie Research Center
807:began on April 30, 2007.
753:, the westernmost of the
284:
277:
152:
138:
129:
122:
45:) is a deadly infectious
3391:Scombroid food poisoning
2020:Journal of Fish Diseases
1740:Journal of Fish Diseases
513:Farmed rainbow trout in
447:
334:Salmonid novirhabdovirus
3416:Piscine novirhabdovirus
3276:Sphaerothecum destruens
3264:Schistocephalus solidus
3208:Lernaeocera branchialis
3054:Spring viraemia of carp
2849:Saulny S (2007-04-21).
2615:Port Huron Times-Herald
2170:10.1186/1743-422X-6-171
1091:. Iowa State University
701:Great Lakes muskellunge
270:Piscine novirhabdovirus
147:Piscine novirhabdovirus
124:Piscine novirhabdovirus
18:Piscine novirhabdovirus
3237:Nanophyetus salmincola
3127:Clinostomum marginatum
2979:Hematopoietic necrosis
2947:Nervous necrosis virus
1288:NCBI Taxonomy Database
1160:. 2000. Archived from
1117:McAllister PE (1990).
467:in continental Europe
100:. Since 2005, massive
88:. This North American
35:
3134:Dactylogyrus vastator
3012:Mycobacterium marinum
2941:Aeromonas salmonicida
2621:on 27 September 2007.
2126:CSU Research Proposal
1652:10.1099/vir.0.79820-0
825:Government regulation
783:On May 17, 2007, the
760:On May 12, 2007, the
711:to the south, and to
667:Great Lakes Type IV-b
634:), and striped bass (
619:Fundulus heteroclitus
502:Farmed rainbow trout
33:
3282:Swim bladder disease
3244:Pseudorhabdosynochus
3225:Myxobolus cerebralis
3177:Gyrodactylus salaris
3102:Amoebic gill disease
3031:Pfiesteria piscicida
2985:Heterosigma akashiwo
2744:Genes & Genomics
2731:on 18 February 2012.
2663:Groocock GH (2007).
2590:on 27 September 2007
2584:Ludington Daily News
2559:on 26 September 2007
1957:on 27 September 2007
1514:10.1128/JVI.00597-11
1447:Marine Biotechnology
1411:10.1128/JVI.00343-10
1164:on 27 September 2007
1123:Fish Disease Leaflet
1019:improve this article
976:AQUATIC). Chlorine
895:University of Toledo
154:Virus classification
3540:Fish viral diseases
3184:Henneguya zschokkei
3048:Streptococcus iniae
2552:(5). Archived from
2537:Phillips K (2007).
2364:on 14 December 2008
2334:on 27 February 2009
2089:2014MolPE..77...34H
2032:2006JFDis..29..611E
1887:on 17 February 2007
1847:on 31 October 2007.
1752:2007JFDis..30..213G
1695:2006ApEnM..72.2373N
1645:(Pt 5): 1167–1179.
1502:Journal of Virology
1459:2011MarBt..13..672A
1405:(19): 10038–10050.
1399:Journal of Virology
558:Marine fish of the
547:Marine fish of the
475:Marine fish of the
143:electron micrograph
61:, and the virus as
55:Northern Hemisphere
3017:Aquarium granuloma
2877:Great Lakes Echo:
2855:The New York Times
2480:The New York Times
2455:on 8 February 2007
2425:on 12 October 2007
2328:Michigan Sportsman
2132:on 3 February 2007
1927:on 6 February 2007
739:St. Lawrence River
416:viral pathogenesis
360:Molecular virology
348:. The virus is an
36:
3535:Novirhabdoviruses
3522:
3521:
3408:Taxon identifiers
3399:
3398:
3120:Ceratomyxa shasta
2804:10.3390/v14071546
2518:on 20 August 2007
2419:The Post-Standard
1995:10.3354/dao047087
1608:10.3354/dao052029
1567:10.3354/dao048143
1508:(16): 8393–8402.
1202:10.3354/dao012167
1047:
1046:
1039:
1010:quality standards
1001:This article may
931:histopathological
592:
591:
412:viral replication
303:
302:
16:(Redirected from
3547:
3515:
3514:
3502:
3501:
3489:
3488:
3476:
3475:
3474:
3448:
3447:
3435:
3434:
3433:
3403:
3190:Ich (freshwater)
3154:Eustrongylidosis
3141:Diphyllobothrium
3006:Koi herpes virus
2991:Hole in the head
2957:Enteric redmouth
2919:
2912:
2905:
2896:
2874:
2866:
2858:
2827:
2826:
2816:
2806:
2782:
2776:
2775:
2739:
2733:
2732:
2730:
2724:. Archived from
2719:
2711:
2705:
2704:
2697:
2691:
2690:
2688:
2686:
2680:
2674:. Archived from
2669:
2660:
2651:
2650:
2648:
2646:
2629:
2623:
2622:
2617:. Archived from
2606:
2600:
2599:
2597:
2595:
2586:. Archived from
2575:
2569:
2568:
2566:
2564:
2558:
2543:
2534:
2528:
2527:
2525:
2523:
2517:
2511:. Archived from
2506:
2497:
2491:
2490:
2488:
2486:
2471:
2465:
2464:
2462:
2460:
2441:
2435:
2434:
2432:
2430:
2421:. Archived from
2410:
2404:
2403:
2401:
2399:
2394:on 6 August 2007
2380:
2374:
2373:
2371:
2369:
2350:
2344:
2343:
2341:
2339:
2330:. Archived from
2319:
2313:
2312:
2310:
2308:
2289:
2283:
2282:
2280:
2278:
2259:
2253:
2252:
2250:
2248:
2239:. Archived from
2229:
2223:
2222:
2220:
2218:
2199:
2193:
2192:
2182:
2172:
2157:Virology Journal
2148:
2142:
2141:
2139:
2137:
2118:
2109:
2108:
2072:
2066:
2065:
2058:
2052:
2051:
2014:
2008:
2007:
1997:
1973:
1967:
1966:
1964:
1962:
1943:
1937:
1936:
1934:
1932:
1923:. Archived from
1915:Crane M (2006).
1912:
1897:
1896:
1894:
1892:
1886:
1880:. Archived from
1875:
1866:
1849:
1848:
1846:
1840:. Archived from
1829:
1819:
1813:
1812:
1802:
1800:10.3354/dao02065
1778:
1772:
1771:
1734:
1725:
1724:
1714:
1689:(4): 2373–2378.
1674:
1665:
1664:
1654:
1630:
1621:
1620:
1610:
1586:
1580:
1579:
1569:
1545:
1536:
1535:
1525:
1493:
1487:
1486:
1442:
1433:
1432:
1422:
1390:
1384:
1383:
1373:
1363:
1339:
1333:
1332:
1330:
1328:
1318:
1309:
1300:
1299:
1297:
1295:
1280:
1274:
1273:
1271:
1269:
1259:
1246:
1240:
1239:
1237:
1235:
1226:
1213:
1207:
1206:
1204:
1180:
1174:
1173:
1171:
1169:
1150:
1139:
1138:
1136:
1134:
1114:
1101:
1100:
1098:
1096:
1086:
1077:
1042:
1035:
1031:
1028:
1022:
996:
995:
988:
819:Skaneateles Lake
722:2005 samples of
649:Fishpathogens.eu
637:Morone saxatilis
628:), brown trout (
622:), stickleback (
463:and a few other
445:
403:Reverse genetics
394:virus polymerase
162:
161:
134:
120:
86:Washington state
21:
3555:
3554:
3550:
3549:
3548:
3546:
3545:
3544:
3525:
3524:
3523:
3518:
3510:
3505:
3497:
3492:
3484:
3479:
3470:
3469:
3464:
3451:
3443:
3438:
3429:
3428:
3423:
3410:
3400:
3395:
3334:
3303:
3201:Kudoa thyrsites
3148:Cymothoa exigua
3083:
3024:Novirhabdovirus
2928:
2923:
2869:
2861:
2848:
2836:
2831:
2830:
2784:
2783:
2779:
2741:
2740:
2736:
2728:
2717:
2713:
2712:
2708:
2703:. 18 June 2009.
2699:
2698:
2694:
2684:
2682:
2678:
2667:
2662:
2661:
2654:
2644:
2642:
2631:
2630:
2626:
2608:
2607:
2603:
2593:
2591:
2577:
2576:
2572:
2562:
2560:
2556:
2541:
2536:
2535:
2531:
2521:
2519:
2515:
2504:
2499:
2498:
2494:
2484:
2482:
2473:
2472:
2468:
2458:
2456:
2443:
2442:
2438:
2428:
2426:
2412:
2411:
2407:
2397:
2395:
2382:
2381:
2377:
2367:
2365:
2352:
2351:
2347:
2337:
2335:
2321:
2320:
2316:
2306:
2304:
2303:on 25 June 2007
2291:
2290:
2286:
2276:
2274:
2273:on 24 June 2007
2261:
2260:
2256:
2246:
2244:
2231:
2230:
2226:
2216:
2214:
2213:on 10 June 2007
2201:
2200:
2196:
2150:
2149:
2145:
2135:
2133:
2120:
2119:
2112:
2074:
2073:
2069:
2060:
2059:
2055:
2026:(10): 611–619.
2016:
2015:
2011:
1975:
1974:
1970:
1960:
1958:
1945:
1944:
1940:
1930:
1928:
1914:
1913:
1900:
1890:
1888:
1884:
1873:
1868:
1867:
1852:
1844:
1838:
1827:
1821:
1820:
1816:
1780:
1779:
1775:
1736:
1735:
1728:
1676:
1675:
1668:
1632:
1631:
1624:
1588:
1587:
1583:
1547:
1546:
1539:
1495:
1494:
1490:
1444:
1443:
1436:
1392:
1391:
1387:
1341:
1340:
1336:
1326:
1324:
1316:
1311:
1310:
1303:
1293:
1291:
1282:
1281:
1277:
1267:
1265:
1257:
1248:
1247:
1243:
1233:
1231:
1224:
1220:Novirhabdovirus
1215:
1214:
1210:
1182:
1181:
1177:
1167:
1165:
1152:
1151:
1142:
1132:
1130:
1129:on 15 June 2007
1116:
1115:
1104:
1094:
1092:
1084:
1079:
1078:
1065:
1060:
1043:
1032:
1026:
1023:
1016:
997:
993:
986:
970:
951:
943:pyknotic nuclei
927:
903:
873:
871:Field diagnosis
868:
852:
836:
827:
727:freshwater drum
717:St. Clair River
697:
669:
661:
613:Clupea harengus
608:Seto Inland Sea
523:Gulf of Bothnia
465:freshwater fish
428:
362:
329:Novirhabdovirus
324:Novirhabdovirus
312:Mononegavirales
273:
257:Novirhabdovirus
233:Mononegavirales
209:Negarnaviricota
156:
118:
28:
27:Disease of fish
23:
22:
15:
12:
11:
5:
3553:
3551:
3543:
3542:
3537:
3527:
3526:
3520:
3519:
3517:
3516:
3503:
3490:
3477:
3461:
3459:
3453:
3452:
3450:
3449:
3436:
3420:
3418:
3412:
3411:
3406:
3397:
3396:
3394:
3393:
3388:
3383:
3378:
3373:
3371:Marine viruses
3368:
3363:
3358:
3353:
3348:
3342:
3340:
3339:Related topics
3336:
3335:
3333:
3332:
3327:
3322:
3317:
3311:
3309:
3305:
3304:
3302:
3301:
3296:
3291:
3284:
3279:
3272:
3267:
3260:
3253:
3248:
3240:
3233:
3228:
3221:
3216:
3211:
3204:
3197:
3192:
3187:
3180:
3173:
3166:
3161:
3156:
3151:
3144:
3137:
3130:
3123:
3116:
3111:
3104:
3099:
3091:
3089:
3085:
3084:
3082:
3081:
3076:
3071:
3066:
3061:
3059:Taura syndrome
3056:
3051:
3044:
3039:
3034:
3027:
3020:
3008:
3003:
2998:
2993:
2988:
2981:
2976:
2973:Flavobacterium
2969:
2964:
2959:
2954:
2949:
2944:
2936:
2934:
2930:
2929:
2924:
2922:
2921:
2914:
2907:
2899:
2893:
2892:
2882:
2875:
2867:
2859:
2846:
2835:
2834:External links
2832:
2829:
2828:
2777:
2734:
2706:
2692:
2681:on 7 June 2007
2652:
2624:
2601:
2570:
2529:
2492:
2466:
2436:
2405:
2375:
2345:
2314:
2284:
2254:
2243:on 6 July 2007
2224:
2194:
2143:
2110:
2067:
2053:
2009:
1968:
1938:
1898:
1850:
1836:
1814:
1773:
1746:(4): 213–223.
1726:
1666:
1622:
1581:
1560:(2): 143–148.
1537:
1488:
1453:(4): 672–683.
1434:
1385:
1334:
1301:
1275:
1241:
1208:
1175:
1140:
1102:
1062:
1061:
1059:
1056:
1045:
1044:
1000:
998:
991:
985:
982:
969:
966:
950:
947:
926:
923:
902:
899:
872:
869:
867:
864:
851:
848:
840:emerald shiner
835:
832:
826:
823:
770:Lake Winnebago
696:
693:
685:Lake St. Clair
668:
665:
660:
657:
590:
589:
586:
582:
581:
578:
574:
573:
556:
552:
551:
545:
541:
540:
530:
526:
525:
511:
507:
506:
500:
496:
495:
473:
469:
468:
457:
453:
452:
449:
427:
426:Virus subtypes
424:
407:vaccinia virus
386:matrix protein
382:phosphoprotein
361:
358:
301:
300:
299:
298:
293:
288:
282:
281:
275:
274:
267:
265:
261:
260:
253:
249:
248:
241:
237:
236:
229:
225:
224:
221:Monjiviricetes
217:
213:
212:
205:
201:
200:
193:
189:
188:
181:
174:
173:
168:
164:
163:
150:
149:
140:Negative stain
136:
135:
127:
126:
117:
116:Virus taxonomy
114:
74:western Europe
59:Egtved disease
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
3552:
3541:
3538:
3536:
3533:
3532:
3530:
3513:
3508:
3504:
3500:
3495:
3491:
3487:
3482:
3478:
3473:
3467:
3463:
3462:
3460:
3458:
3454:
3446:
3441:
3437:
3432:
3426:
3422:
3421:
3419:
3417:
3413:
3409:
3404:
3392:
3389:
3387:
3384:
3382:
3379:
3377:
3374:
3372:
3369:
3367:
3364:
3362:
3359:
3357:
3354:
3352:
3349:
3347:
3344:
3343:
3341:
3337:
3331:
3328:
3326:
3323:
3321:
3318:
3316:
3313:
3312:
3310:
3306:
3300:
3297:
3295:
3292:
3290:
3289:
3285:
3283:
3280:
3278:
3277:
3273:
3271:
3268:
3266:
3265:
3261:
3259:
3258:
3254:
3252:
3249:
3246:
3245:
3241:
3239:
3238:
3234:
3232:
3229:
3227:
3226:
3222:
3220:
3217:
3215:
3214:Microsporidia
3212:
3210:
3209:
3205:
3203:
3202:
3198:
3196:
3193:
3191:
3188:
3186:
3185:
3181:
3179:
3178:
3174:
3172:
3171:
3167:
3165:
3162:
3160:
3157:
3155:
3152:
3150:
3149:
3145:
3143:
3142:
3138:
3136:
3135:
3131:
3129:
3128:
3124:
3122:
3121:
3117:
3115:
3112:
3110:
3109:
3105:
3103:
3100:
3098:
3097:
3093:
3092:
3090:
3086:
3080:
3077:
3075:
3072:
3070:
3067:
3065:
3062:
3060:
3057:
3055:
3052:
3050:
3049:
3045:
3043:
3042:Salmon anemia
3040:
3038:
3035:
3033:
3032:
3028:
3026:
3025:
3021:
3018:
3014:
3013:
3009:
3007:
3004:
3002:
2999:
2997:
2994:
2992:
2989:
2987:
2986:
2982:
2980:
2977:
2975:
2974:
2970:
2968:
2965:
2963:
2960:
2958:
2955:
2953:
2950:
2948:
2945:
2943:
2942:
2938:
2937:
2935:
2931:
2927:
2920:
2915:
2913:
2908:
2906:
2901:
2900:
2897:
2890:
2886:
2883:
2880:
2876:
2872:
2868:
2864:
2860:
2856:
2852:
2847:
2845:
2841:
2838:
2837:
2833:
2824:
2820:
2815:
2810:
2805:
2800:
2796:
2792:
2788:
2781:
2778:
2773:
2769:
2765:
2761:
2757:
2753:
2749:
2745:
2738:
2735:
2727:
2723:
2716:
2710:
2707:
2702:
2696:
2693:
2677:
2673:
2666:
2659:
2657:
2653:
2641:
2640:
2635:
2632:Woo P (ed.).
2628:
2625:
2620:
2616:
2612:
2605:
2602:
2589:
2585:
2581:
2574:
2571:
2555:
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1253:Rhabdoviridae
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774:Lake Michigan
771:
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762:Wisconsin DNR
758:
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748:
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743:Niagara River
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318:Rhabdoviridae
314:
313:
309:of the order
308:
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2747:
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2076:
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2019:
2012:
1988:(2): 87–99.
1985:
1981:
1971:
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1337:
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1320:
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1278:
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1157:
1131:. Retrieved
1127:the original
1122:
1093:. Retrieved
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1024:
1017:Please help
1013:
1002:
971:
963:
955:
952:
928:
920:
904:
888:
880:exophthalmia
877:
874:
861:
853:
844:
837:
834:Transmission
828:
816:
809:
785:Michigan DNR
782:
759:
755:Finger Lakes
751:Conesus Lake
736:
724:Lake Ontario
721:
715:through the
707:through the
698:
689:
681:Lake Ontario
670:
662:
646:
642:
635:
631:Salmo trutta
629:
623:
617:
611:
600:
593:
585: IV-b
577: IV-a
563:
429:
401:
398:
390:glycoprotein
380:-associated
374:nucleocapsid
363:
346:M. H. Jenson
343:
338:
332:
328:
322:
321:, and genus
316:
310:
304:
295:
290:
287:Egtved virus
269:
268:
256:
244:
232:
220:
208:
196:
184:
177:
167:(unranked):
146:
123:
67:
63:Egtved virus
62:
58:
50:
47:fish disease
42:
38:
37:
3308:Fish groups
3257:Saprolegnia
3251:Salmon lice
3096:Abergasilus
2967:Fish dropsy
2797:(7): 1546.
1264:. p. 7
1021:if you can.
907:hemorrhagic
673:Great Lakes
555: III
529: I-e
510: I-d
499: I-c
472: I-b
456: I-a
106:Great Lakes
3529:Categories
3351:Brevetoxin
3231:Myxosporea
3079:Yellowhead
3074:White spot
2952:Columnaris
2645:11 October
2546:Fish Lines
2207:Highlights
1195:: 167–75.
1058:References
1027:March 2022
968:Prevention
905:VHSV is a
731:Lake Huron
713:Lake Huron
549:Baltic Sea
544: II
477:Baltic Sea
430:Different
378:polymerase
337:(formerly
305:VHSV is a
108:region of
98:Baltic Sea
96:, and the
78:fish farms
49:caused by
3472:Q18965285
3431:Q29004930
3386:Saxitoxin
3366:Fish kill
3356:Ciguatera
3270:Sea louse
3219:Monogenea
3114:Carp lice
3088:Parasites
2933:Pathogens
2772:195760294
2322:Barta J.
2083:: 34–40.
916:hyperemic
911:petechial
866:Diagnosis
859:attacks.
801:bluegills
789:Budd Lake
778:Green Bay
705:Lake Erie
659:Evolution
604:North Sea
538:Black Sea
489:North Sea
481:Skagerrak
420:apoptosis
354:peplomers
350:enveloped
315:, family
264:Species:
192:Kingdom:
185:Riboviria
94:North Sea
70:salmonids
3486:11461179
3466:Wikidata
3425:Wikidata
3108:Anisakis
2823:35891526
2764:31721105
2189:19852863
2105:24727199
2048:17026670
2004:11775799
1809:19694169
1768:17394523
1721:16597932
1661:15105533
1617:12517003
1576:12005236
1532:21653667
1483:24597813
1475:20936318
1429:20631140
1380:29527292
1354:: 1958.
1003:require
939:lamellae
850:Symptoms
677:die-offs
485:Kattegat
440:enzootic
432:isolates
279:Synonyms
240:Family:
204:Phylum:
102:die-offs
90:genotype
2962:Fin rot
2814:9317989
2791:Viruses
2685:18 July
2639:WikiVet
2594:13 July
2563:16 July
2522:16 July
2485:13 July
2459:15 July
2429:15 July
2398:13 July
2368:15 July
2338:15 July
2307:13 July
2277:13 July
2247:14 July
2217:14 July
2180:2771013
2163:: 171.
2136:12 July
2085:Bibcode
2028:Bibcode
1961:16 July
1931:16 July
1891:13 July
1748:Bibcode
1712:1449023
1691:Bibcode
1523:3147959
1455:Bibcode
1420:2937753
1371:5820608
1327:16 July
1294:14 July
1268:15 June
1234:15 June
1168:12 July
1133:12 July
1095:12 July
1051:viperin
1005:cleanup
893:at the
857:lamprey
797:muskies
747:walleye
536:in the
519:Finland
504:Denmark
459:Farmed
252:Genus:
228:Order:
216:Class:
3512:816302
3299:Xenoma
3294:Velvet
3170:Glugea
3164:Flukes
2821:
2811:
2770:
2762:
2187:
2177:
2103:
2046:
2002:
1834:
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1530:
1520:
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1473:
1427:
1417:
1378:
1368:
978:bleach
974:Virkon
959:RT-PCR
884:orbits
803:, and
570:Norway
534:turbot
515:Norway
366:genome
3507:WoRMS
3499:11287
3481:IRMNG
3445:4J8KZ
2768:S2CID
2729:(PDF)
2718:(PDF)
2679:(PDF)
2668:(PDF)
2557:(PDF)
2542:(PDF)
2516:(PDF)
2505:(PDF)
1885:(PDF)
1874:(PDF)
1845:(PDF)
1828:(PDF)
1479:S2CID
1317:(PDF)
1258:(PDF)
1225:(PDF)
1085:(PDF)
1049:Fish
493:Japan
448:Type
396:(L).
384:(P),
178:Realm
171:Virus
3494:NCBI
3247:spp.
2819:PMID
2760:PMID
2687:2007
2647:2011
2596:2007
2565:2007
2524:2007
2487:2007
2461:2007
2431:2007
2400:2007
2370:2007
2340:2007
2309:2007
2279:2007
2249:2007
2219:2007
2185:PMID
2138:2007
2101:PMID
2044:PMID
2000:PMID
1963:2007
1933:2007
1893:2007
1832:ISBN
1805:PMID
1764:PMID
1717:PMID
1657:PMID
1613:PMID
1572:PMID
1528:PMID
1471:PMID
1425:PMID
1376:PMID
1329:2007
1296:2007
1270:2019
1236:2019
1170:2007
1135:2007
1097:2007
729:and
653:fish
596:pike
436:host
364:The
3440:CoL
3069:VHS
3064:UDN
2809:PMC
2799:doi
2752:doi
2175:PMC
2165:doi
2093:doi
2036:doi
1990:doi
1795:doi
1756:doi
1707:PMC
1699:doi
1647:doi
1603:doi
1562:doi
1518:PMC
1510:doi
1463:doi
1415:PMC
1407:doi
1366:PMC
1356:doi
1197:doi
776:'s
331:is
145:of
72:in
43:VHS
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