Arctic Rabies – a Review
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Acta vet. scand. 2004, 45, 1-9. Arctic Rabies – A Review By Torill Mørk1 and Pål Prestrud2 1National Veterinary Institute, Regional laboratory, and 2Norwegian Polar Institute, Polar Environmental Centre, N-9296 Tromsø, Norway. Mørk T, Prestrud P: Arctic rabies – a review. Acta vet. scand. 2004, 45, 1-9. – Rabies seems to persist throughout most arctic regions, and the northern parts of Nor- way, Sweden and Finland, is the only part of the Arctic where rabies has not been diag- nosed in recent time. The arctic fox is the main host, and the same arctic virus variant seems to infect the arctic fox throughout the range of this species. The epidemiology of rabies seems to have certain common characteristics in arctic regions, but main ques- tions such as the maintenance and spread of the disease remains largely unknown. The virus has spread and initiated new epidemics also in other species such as the red fox and the racoon dog. Large land areas and cold climate complicate the control of the dis- ease, but experimental oral vaccination of arctic foxes has been successful. This article summarises the current knowledge and the typical characteristics of arctic rabies in- cluding its distribution and epidemiology. Arctic fox rabies; epidemiology, review. Introduction The history of rabies in the Arctic before 1945 nent potential source of infection of sub-arctic is sparsely known. The folklore of the Canadian areas (Anonymous 1990). inuits indicates that these people knew of a ra- bies-like disease that was transmitted from arc- The arctic virus variant tic foxes to dogs and people (Singleton 1969), Rabies virus belongs to the family Rhabdoviri- and in Greenland epidemics among sledge dogs dae and the genus Lyssavirus. The genotype 1 have been described for almost 150 years consists of the classic rabies virus, while the (Lassen 1962). Rabies virus was first identified other known genotypes consist mainly of in arctic regions by Williams (1949). Epidemics viruses circulating among fruit and insect eat- among sledge dogs, initiated by transmission ing bats. By monoclonal antibody technique from rabies-infected arctic foxes, have caused and genome analysis, different antigenic vari- severe problems in several arctic areas, some- ants circulating in different main hosts within times reducing the number of sledge dogs dras- certain geographical areas have been identified tically (Sikes 1968, Holck 1989, Crandell (Webster et al. 1986, Nadin-Davis 1998, Bour- 1991). After the onset of dog vaccination, ra- hy et al. 1999). This supports the theory that ra- bies has remained mainly as a disease of the bies virus adapts to different main hosts for per- arctic fox in these regions. There are several ex- sistence in the host population (Wandeler et al. amples of southward spread of rabies virus 1994). from arctic regions (Tabel et al. 1974, Webster Characterisation with monoclonal antibodies et al. 1986, Johnston & Fong 1992, Selimov et has indicated that a specific virus variant react- al. 1990) and arctic rabies represents a perma- ing with the monoclonal antibody (Mab) P-41, Acta vet. scand. vol. 45 no. 1-2, 2004 2 T. Mørk & P. Prestrud circulates in arctic areas (Schneider et al. in arctic foxes and similar epidemics among 1985), and the arctic fox is regarded as the main sledge dogs was noted in 1916 (Elton 1931). host of this virus variant (Johnston & Fong Today, the arctic strain virus persists among 1992). The arctic strain has also been isolated arctic foxes in the north and until recently from other animal species, such as red fox caused several epidemics among red foxes and (Vulpes vulpes), racoon dog (Nyctereutes pro- skunks in Ontario and Qubeck in the southeast cyonoides) and skunk (Mephitis mephitis) in (Webster et al. 1986, MacInnes et al. 2001). sub-arctic areas (Webster et al. 1986, Westerling In Greenland, epidemics of rabies-like disease 1989, Selimov et al. 1990, Nyberg et al. 1992). among sledge dogs were reported as far back as However, there are examples of virus isolates 1859, but the virus was first identified among from far more southern areas, not likely to be dogs and arctic foxes in 1959. There have been the arctic strain, which have reacted positively several large epidemics among sledge dogs in with Mab P-41, indicating that a more complete Greenland prior to 1960, when vaccination of monoclonal antibody panel is necessary for dogs was initiated (Holck 1989, Lassen 1962). identification of the arctic strain (Selimov et al. Rabies virus is still considered as endemic 1994). The reverse transcriptase-polymerase among the arctic fox population. chain reaction (RT-PCR) methodology has fa- In Arctic Russia, rabies virus was first identi- cilitated genetic characterisation of rabies virus fied during a major research study from 1954 to isolates and has been used both as a diagnostic 1956 in the Nenets Autonomous Okrug in and as a characterisation tool to identify differ- northwest of Russia. The study confirmed that ent strains persisting in certain host species the rabies-like disease known from arctic foxes within geographically defined regions. In and dogs in arctic regions of Russia, was caused Canada, 5 different variants of the arctic strain, by rabies virus (Kantorovich 1964). The arctic obtained from different geographical areas, virus strain has later been found in several arc- have been identified by RT-PCR (Nadin-Davis tic regions of Russia and also in several differ- 1998). ent animal species south of the arctic region (Selimov et al. 1990, 1994). From the Kola Distribution Peninsula, which borders Finland and Norway, Rabies virus is endemic throughout most parts arctic rabies was reported in the late 1980s of the Arctic, and several epidemics have been (Westerling 1989, Selimov 1990). reported during the last 40-50 years (Raush In the high-arctic Svalbard islands of Norway, 1958, Kantorovich 1964, Crandell 1975, Ritter rabies was detected for the first time in 1980 1981, Holck 1989). In Alaska, rabies-like dis- when there was an outbreak in the arctic fox ease was first reported in 1887 and rabies virus population. Rabies was also diagnosed in 3 was identified during an epidemic in 1945-47 reindeers and 1 seal (Ødegaard & Krogsrud (Williams 1949). Today, rabies is regarded as 1981). Only few cases were reported from endemic among the red fox and the arctic fox in 1980-1992 and during the last years there have northern and western areas of Alaska, with a been speculations whether the rabies virus had cyclic occurrence of epidemics about every 3-4 died out naturally. However, in an ongoing re- years (Ritter 1981, Follmann 1990). search project, rabies was diagnosed recently in In arctic Canada, rabies-like disease was re- one fox from 1998 and one from 1999 (Mørk & ported from the North West Territory already in Fuglei, unpublished data). 1867 and the relationship between this disease The mainlands of Norway and Sweden have Acta vet. scand. vol. 45 no. 1-2, 2004 Arctic rabies 3 both been regarded as rabies free countries dur- RNA was demonstrated among 13% of the ing the last 150 years. Reports of rabies in Swe- hyena population. Despite this high frequency den date back to 1886 (Wierup & Engvall of exposure, there were no cases of symp- 1990), while there are no reports of rabies in the tomatic rabies or decreased survival among a mainland of Norway. group of hyenas monitored for 9 to 13 years Finland was declared free of rabies in 1936. (East et al. 2001). There are reports of experi- Since then there have been several epidemics, mental rabies cases, where dogs have recovered all close to the Russian border. In 1988 there from the disease and have been secreting virus was an outbreak in southern Finland, and the in the saliva for a longer period after recovery main species involved was the racoon dog, a (Fekadu et al. 1981, Fekadu 1983). Apparently species established in Finland about 1970-80. healthy dogs have been found to secret rabies The epidemic was caused by the arctic virus virus in their saliva during a period, and re- strain and was successfully stopped by bait vac- mained clinically normal during several years cination (Westerling 1989, Nyberg et al. 1992). after the first virus isolation (Nanavati 1973, Except for one imported case in a horse in Fekadu 1975). Serological surveys of wild ani- 2003, no cases of rabies have been reported mal populations have been limited, but rabies from Finland since 1989. antibodies have also been found in arctic foxes in Alaska, indicating that some foxes survive Clinical appearance virus exposure (Ballard et al. 2001). The incubation period in experimentally in- fected arctic foxes has been reported to vary Zoonotic aspects from 8 days to 6 months (Konovalov et al. 1965, There have been few human cases of rabies in Rausch 1972). The clinical course is usually the arctic regions, and it has therefore been short and foxes may die within a day or 2 after claimed that the arctic virus strain is less the onset of symptoms. Initially, the arctic fox pathogenic to man (Johnston & Fong 1992). looses its natural timidity. It may enter villages However, there might be other explanations. An or human settlements and there are examples of important aspect is that people rarely become foxes having followed dog teams. In the exita- infected from foxes, thus the most probable ex- tive phase, the fox becomes aggressive and may posure is through dogs. There are several re- snap and bite, and sometimes runs in circles. ports indicating that most dogs infected with Excessive drivelling and foaming are also typi- arctic rabies develop "dumb rabies", which re- cal symptoms. In the following phase, the ani- duces the risk of human infection (Kantorovich mal becomes paralytic and eventually dies.