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Diseases of Penguins are a significant component of ’s Thirteen of the world’s 16 ( Spheniscidae) biodiversity with 13 species, four of which are endemic. occur in New Zealand’s maritime territories(1). Four are either rare or Disease prevalence studies have been only locally common endemics and, of the remaining species, five limited to a few species and to a few breed here and four are non-breeding visitors(1). The endemic and disease agents. The published literature is endangered yellow-eyed penguin or hoiho ( antipodes) is reviewed along with unpublished data from locally distributed on the South Island, Stewart Island, Codfish New Zealand’s diagnostic laboratories. Island, the Auckland Islands and Campbell Island. The rarest Padraig J Duignan penguin species in the world, its population has declined dramatically on the South Island(2). Fledglings go to sea in February In recent years there has been a dramatic decline in the number of and March and half die of malnutrition during their first five weeks erect-crested and rockhopper penguins. The former has decreased by at sea resulting in a peak of beach-cast mortality in May and 20% on the Antipodes since the mid 1990s and the latter by about June(3)(4)(5). The Fiordland or tawaki (Eudyptes 90% over the past 50 years on Campbell Island(8). Although the pachyrhynchus) is also an endangered endemic that has a restricted factors responsible have yet to be established, the circumpolar breeding distribution from Westland to Stewart Island. Most beach- decline of rockhoppers suggests widespread environmental changes cast mortality occurs from October to February when fledglings go (D Thompson, ESR, personal communication). Declines of yellow- to sea and breeding adults begin to moult(5). Related endemic species eyed penguins have been attributed to disturbance, loss, are the Snares crested penguin (E robustus) and the erect-crested introduced predators, periodic food shortages, and fluctuations in penguin (E sclateri). The former is locally abundant on the Snares climatic variables such as rainfall and sea surface temperatures(9)(10)(11). Islands where the only nesting sites occur. The latter breeds on In , New Zealand’s Ross Dependency has three breeding islands including the Antipodes, the Bounties, and on penguin species including the emperor ( forsteri), Adélie Disappointment Island in the Aucklands. Outside the breeding ( adeliae), and chinstrap (P antarctica)(1). Visitors and season individuals of both species disperse widely and have been vagrants to the New Zealand region include the two subspecies of E recorded regularly in . Fledgling Snares crested penguins are chrysalophus (the royal and macaroni), king (A patagonicus), and occasionally found wrecked on Southland beaches in summer(5).Low (P papua), whose closest breeding colonies are on numbers of erect-crested penguin fledglings are found beach-cast on and Heard Island. The the southeast coast of the South Island during March and April(5). (Spheniscus magellanicus) is a rare vagrant from South America(1). The native species with the most northerly breeding distribution, The threats to the various species in New Zealand territories include extending from Northland to Stewart Island and the Chatham by introduced mammals, habitat alteration and Islands, is the blue penguin or korora ( minor). At least five disturbance, competition for resources, and incidental mortality by phenotypic variants, or subspecies, occur around New Zealand; one commercial fisheries(12)(13). Infectious diseases and parasites may also is known as the white-flippered penguin. Blue penguins are common compromise the survival of individuals or relict populations and are and frequent the Hauraki Gulf, Cook Strait, Kapiti coast, Otago, the subject of this review. Foveaux Strait and the Chathams. Between1960 and1982 almost 16,000 were recorded in beach surveys for sea mortality(5). This Infectious diseases figure includes unusual mortalities (wrecks) such as the 3,729 blue The literature on infectious diseases of New Zealand’s penguins is penguins found dead on Auckland’s east coast beaches in 1974 and a patchy with few directed studies on endemic diseases. Several (5)(6) similar event in Northland in 1973-74 . Annual mortality peaks penguin species have received little attention apart from cataloguing from January to March, coinciding with fledging and the parasites. With little information on endemic disease, the risks from (5)(7) period of breeding . potentially exotic diseases are largely unknown.

The subantarctic islands (Aucklands, Campbell, Bounties, Antipodes, Viral infections Snares) are frequented by ten penguin species and four regularly A 1988 serological survey on a limited number of rockhopper and breed there. The yellow-eyed penguin is a solitary-nesting species yellow-eyed penguins on Campbell Island reported no antibodies while the other three species are colony-nesting crested penguins and against a panel of poultry viruses including the agents of infectious include the Snares crested and erect-crested penguins. The smallest bronchitis, reticuloendotheliosis, Newcastle disease, infectious of the group, the eastern rockhopper penguin (E chrysocome filholi), laryngotracheitis, avian encephalomyelitis, infectious bursal disease, occurs locally on New Zealand’s subantarctic islands and on Marek’s disease, and fowlpox(14). Macquarie Island in Australian territory. Two other rockhopper subspecies, E c chrysocome and E c moselyi, are vagrants. Crested Paramyxoviruses: Nine paramyxovirus isolates were recovered from penguins breed and moult ashore and then spend about four cloacal swabs collected between 1976 and 1979 from royal and king months foraging at sea over the winter(8). penguins on Macquarie and Heard Islands in Australia’s

page 5 Surveillance 28(4) 2001 subantarctic(15)(16). One isolate was indistinguishable from viruses of Orthomyxoviruses: Avian influenza virus antibodies, but not PMV-1 serotype (lentogenic Newcastle disease virus, NDV). No clinical disease, were found in Adélie penguins from Casey Station viruses were isolated from either gentoo or rockhopper penguins(15). and the in Antarctica(18)(22). King, royal and rockhopper penguins sampled at several sites had Arboviruses: Antibodies against a Flavivirus have been found antibodies against one of the isolates and royal penguins were occasionally in sera from king, royal, Adélie, and rockhopper seropositive for NDV(15). Antibodies against the same viruses were penguins(15). A new Flavivirus, Bunyavirus, and were also detected in sera from a few blue penguins sampled along the isolated from ( uriae) collected from penguins on Victorian coast of Australia(17). Macquarie Island, which suggests several possibilities for the low In Antarctica, paramyxoviruses were isolated from Adélie penguin prevalence of antibodies: transmission rates may be low, penguins cloacal swabs at several rookeries between 75°E (Davis Station) and may be a dead-end host, or disease may be fatal and few birds 170°E (the Ross Sea)(18)(19). A serological survey of seven colonies seroconvert and survive(23)(24)(25). Recently, a new arbovirus (Family Togaviridae, Alphavirus) was isolated from lice collected from elephant seals (Mirounga leonina) on Macquarie Island(26). Further characterisation is required for these novel wildlife viruses.

Unclassified RNA virus: The significance of an uncharacterised RNA virus isolated from a pool of ticks (I uriae) from rockhopper penguins on Campbell Island(14) has not been established and antibodies were not present in rockhopper or yellow- eyed penguins. It is unlikely to be associated with the species’ decline.

Birnaviruses: Infectious bursal disease virus (IBDV) occurs as avirulent strains Yellow-eyed penguins and a sealion on Enderby Island (photo Gareth W Jones) and a highly virulent strain. Even the avirulent strains can cause growth found that infection was widespread close to Casey Station in retardation and immunosuppression(27). Antibodies against IBDV (18) Australian territory and the French base of Dumont d’Urville . were detected in sera from emperor and Adélie penguins in However, no seropositive birds were detected at 13 sites near the Antarctica but there was no evidence of disease(28). Adélie chicks at a (19) Davis Station in Australian territory . It appears that titres may be colony remote from human activity were seronegative, suggesting elevated only transiently following infection and this may influence that the virus may have been introduced into Antarctica with seroprevalence data. contaminated poultry products.

Based on antigenic and biochemical characteristics, the penguin Herpesviruses: A herpesvirus caused laryngotracheitis-like paramyxoviruses are distinct from those of other avian species but infection in African black-footed penguins (Spheniscus demersus) at some isolates share epitopes with PMV-1 isolates from the northern a zoo in North America(29). The disease has not been reported for (16) hemisphere . The significance of infection for free-living birds is any free-living penguin species. unknown but the isolates were not pathogenic for chickens or blue Suspected viral mortalities: In 1972, an unusual event killed 65% penguins(17). However, infection could predispose penguins to of Adélie penguin chicks near , Antarctica(29). opportunistic infections or confer immunity against pathogenic Moribund chicks were in good body condition but were recumbent viruses such as NDV. Antibodies against the latter were detected in and would stagger and fall forward when righted. The cause was not the serum of Adélie and royal penguins(15)(18). The significance of established but it was not starvation as occurred in 1995(30). NDV for free-living birds is unknown but mortalities have occurred among Adélie penguins(20) and in a captive (21).The A mass mortality among gentoo penguins at Signey Island involved Adélies were captured in 1973 near Scott Base in the Ross Sea and several hundred birds that were in good body condition, but each shipped to the United States where several birds died from an had focal ulcers on the dorsal surface of the feet(31). No aetiology was infection characterised as velogenic neurotropic in chickens(20).It established. was believed that infection was contracted in the wild because the In 1981, an unusual number of four- to five-week-old Adélie chicks birds were maintained in quarantine. died on Petersen Island (near Casey Station). No clinical signs were

page 6 Surveillance 28(4) 2001 observed and no necropsies conducted. However, five of 55 adult Cause of mortality and incidental findings for birds were seropositive for influenza antibodies(18). Although penguins in New Zealand, based on records on the (13) unlikely, the virus may have been implicated. Wildlife Mortality Database () and from Hocken Diagnosis Blue penguin Yellow-eyed Bacterial and fungal diseases penguin Starvation 34 (13%) Enteric bacteria: Many bacteria have been isolated from free-living Starvation and parasitism1 15 (6%) 2 penguins but only a few are pathogens(32)(33)(34).Five Salmonella Starvation and trauma 1 (0.004%) 2 serotypes were isolated from Adélie penguins on Ross Island, Starvation and neurological signs 1 (0.004%) Trauma (35) Antarctica, but were not associated with disease . Campylobacter Dogs 35 (14%) spp, with only minor sequence difference in 16sRNA from C jejuni Mustelids (ferrets) 20 (8%) and C lari, were isolated from marine birds including apparently Shark3 8 (3%) healthy gentoo and macaroni penguins sampled on Bird Island, Vehicles 32 (13%) 1 Vandalism 10 (4%) South Georgia(36). Whether these enteric bacteria are normal flora of Traps 3 (1%) penguins is not known. It is possible that bacteria were brought to Undetermined 27 (11%) Antarctica or subantarctic islands on contaminated poultry Wrecks (multiple starvation mortalities)4 8 (3%) 5 products and disseminated through penguin rookeries by scavengers Bacterial infection 5 (2%) Plasmodium-like infection6 1 such as and giant petrels. Migratory sea birds may also act as Aspergillosis7 8 (3%) vectors of infection to penguin species that are restricted to the Drowning8 10 (4%) subantarctic or (37). Undetermined 36 (14%) 1 One blue penguin was under rehabilitation for seven days and at necropsy also Psittacosis: Antibodies against Chlamydiophila psittaci, the cause of had mycotic pneumonia (Aspergillosis). The parasite burdens included some or avian psittacosis, were found in emperor, rockhopper, royal, gentoo all of the following: Contracaecum sp nematodes in the proventriculus, Tetrabothroid cestodes in the intestine, Renicola trematodiasis of the kidney, and Adélie penguins(38)(39). There was no evidence of disease and the intestinal and renal coccidiosis. One bird had haemorrhage from a origin and significance of infection are unknown. proventricular ulcer that was probably parasite-induced. 2 One blue penguin was reported to display neurological impairment but no Pasteurellosis: Pasteurella multocida of similar serotype to that histological lesions were detected. The signs were possibly biotoxin-induced. 3 Inferred from linear lacerations in skin and shorn feathers(13). which causes avian cholera epidemics among North American 4 The eight birds in this submission were juveniles found wrecked at the same waterfowl was isolated from several septicaemic rockhopper time on the Coromandel Peninsula. penguins on Campbell Island(14). rats (Rattus norvegicus) 5 Presumptive diagnosis based on gross and histological lesions. 6 Acute multifocal necrotising hepatitis, splenitis and myocarditis associated with appear to be associated with the disease. The eradication of rats intracellular , species not determined. from Campbell Island over the winter of 2001 may reduce the 7 Aspergillosis was the only significant finding reported. prevalence of infection and establish whether it is implicated in the 8 Drowning diagnosed from the location of the carcass and postmortem finding of fluid in lungs and air sacs(13). rockhopper decline on the island.

Borreliosis: Antibodies against Borrelia burgdorferi, the cause of Metazoan and protozoan parasites in mammals, were detected in king penguins on the Helminths, ectoparasites, and protozoa of penguins have been Crozet Archipelago, French subantarctic(40). The bacteria were also comprehensively reviewed and catalogued(45)(46)(47)(48). isolated from I uriae, a likely vector. Infection rarely results in disease in birds but it is a potential zoonosis. Ectoparasites: Fleas, ticks, mites and biting lice occur on most penguin species. The has a wide distribution on Aspergillosis: Caused by Aspergillus fumigatus, and rarely A flavus, seabirds in both hemispheres. Three other species have also been A niger or A terreus, aspergillosis is commonly diagnosed in captive recorded on blue penguins: I kohlsi, I auritulus and I eudyptidus. penguins but is rare in free-living birds. However, pulmonary Colony-nesting species have the highest parasite burdens and ticks infection has been recorded in emaciated beach-cast juvenile little are most commonly found on chicks around the face, canals, blue penguins(15)(41). Lesions have also been reported in wrecked blue cloaca and feet. Species affected include emperor, Adélie, gentoo, penguins in New Zealand (see table) and in a yellow-eyed penguin royal, rockhopper, blue and Snares crested penguins. Heavy juvenile(42). infestation may cause death of chicks and occasionally adults. Of Aspergillus spp are ubiquitous and even found in soils and greater significance may be the potential for transmission of viral nests in Antarctica(43). A serological survey of penguins from New infections. Paramyxoviruses and arboviruses were isolated from Zealand (yellow-eyed and blue), the subantarctic (yellow-eyed and I uriae from royal and king penguins on Macquarie Island(15)(23)(24). rockhopper), and Antarctica (Adélie), found an inverse relationship An uncharacterised enveloped RNA virus, morphologically but not between seroprevalence and latitude; all blue penguins from Napier antigenically similar to infectious bronchitis virus of chickens, was were seropositive but none of the Adélies(44). isolated from I uriae from Campbell Island rockhopper penguins(14).

page 7 Surveillance 28(4) 2001 Biting lice (Austrogoniodes sp) have been recorded on all species of described from blood smears from free-living Fiordland crested Antarctic and subantarctic penguins except gentoo and chinstrap penguins(53), is transmitted by blackflies (Austrosimulium spp) and is penguins(45)(46). Fleas (Parapsyllus sp) are also common in the probably not pathogenic. It was not found in blood smears from 25 subantarctic and temperate zones and have been found on blue, Snares crested penguins and one erect-crested penguin on the rockhopper, gentoo, macaroni, Magellanic, and yellow-eyed Snares(53). Trypanosoma eudyptulae was described from smears from penguins. Fleas need a suitable nest environment and spend only blue penguins in southeastern Tasmania(54). The intermediate host part of their life on the birds. Lice and fleas are of little pathogenic was not identified but ticks were proposed. Infection appears to importance but numbers may be increased on birds that are cause no disease in penguins. debilitated and not preening properly(41). They may also transmit Malaria: Avian malaria caused by Plasmodium relictum is an viruses. important cause of mortality among captive penguins in North Helminths: Cestodes, nematodes, trematodes, and acanthocephalans America and Europe(55), and also among free-living blue penguins in are common in most penguin species. Blue penguins in North Australia(56) and African black-footed penguins(57)(58). Infection in the Otago had lower than expected burdens(13). Four species of latter lasts for life and stress such as oiling or transportation may Contracaecum (gastric nematodes) have been recorded, and may be induce recrudescence(59)(60). Parasitaemia underestimates infection associated with erosion and ulceration leading to haemorrhage. but an enzyme-immunoassay detected a high seroprevalence (up to Infection rates in South Island yellow-eyed penguins were high for 100%) in yellow-eyed penguins on Banks Peninsula, Otago chicks and moderate for adults but parasite burdens were low and Peninsula, Catlins, Codfish Island, Enderby Island (the Aucklands), no pathology was observed(13)(45). Although nematodes probably do and Campbell Island(61)(62). Only 10% of yellow-eyed penguins not directly contribute to the population declines observed for sampled on Stewart Island had P r spheniscidae parasitaemia(57) and several New Zealand penguin species, they may be a factor(6)(41)(49)(50). none of 189 blood smears from 83 free-living yellow-eyed penguins on Banks Peninsula had demonstrable haemoparasites (S Cestodes of the genus Tetrabothrius occur in the intestines of blue, McDonald, personal communication). emperor, king, gentoo, Adélie, rockhopper and Magellanic penguins. Their life cycle is unknown but it may involve euphausiid Despite the apparently high level of exposure of yellow-eyed as intermediate hosts(51). Acanthocephalans are rarely penguins to Plasmodium sp, the incidence of clinical disease appears reported from marine birds but comprised 1.3% of helminths in a to be low. The cause of the deaths of 150 adult penguins on Banks sample of 20 yellow-eyed penguin chicks(49). Corynosoma sp has also Peninsula during the summer of 1989-90 was not established(42). been reported for gentoo penguins(48). Both are an unlikely cause of Avian malaria was suggested as the aetiology based on elevated significant disease among penguins. antibody titres in plasma from dead birds(61).However,the investigators found no histological lesions characteristic of malaria Several trematode species reside in the liver, gall bladder, and and proposed that acute intoxication by a marine algal neurotoxin intestines of penguins(46). Most are probably well tolerated but the was more likely, based on clinical signs, pathology and hepatic fluke Mawsonotrema eudyptulae contributes to the mortality environmental conditions (J Gill, personal communication). of juvenile blue penguins periodically wrecked in Victoria and New Zealand(5)(6)(41)(50)(52). Trematodes may exacerbate the effects of Whatever the cause of that event, a recent case on Otago Peninsula starvation in recently fledged birds. Heavy burdens of flukes were shows that avian malaria has the potential to cause mortality among noted in the kidneys of wrecked blue penguins in Northland, but yellow-eyed penguins. In February 2001, a juvenile in good body with minimal pathology(6). The species was not identified but it is condition was found dead without gross lesions. On histology, there probably of the family Echinostomatidae. was extensive acute necrosis and inflammation of the myocardium, liver and spleen associated with intracytoplasmic Plasmodium-like Coccidiosis: Oocysts were detected in the faeces of free-living parasites(63). Similar pathology was also described for a captive yellow-eyed penguins in Otago but the species was not determined Fiordland crested penguin in Australia(56). because they could not be sporulated(49). The low prevalence of infection suggests that it is not significant. Both intestinal and renal No haematozoa have been detected in blood smears from king, coccidiosis have been recorded in wrecked blue penguins in royal, and gentoo penguins on Macquarie Island, king and macaroni Victoria(41)(52). Oocysts were found in faeces of five of 48 birds but on penguins from Heard Island, and Adélie and emperor penguins histology, schizonts were detected in the intestinal mucosa of only from Antarctica(64)(51)(65). Serology confirmed the absence of infection one(41). By contrast, 64% of birds had chronic interstitial nephritis from Adélie penguins on Ross Island, Antarctica, suggesting that a and ureteritis associated with coccidia. However, the lesions were suitable vector is absent from polar latitudes(61). However, at least mild and focal and probably of little pathological significance(52). some penguins that inhabit the subantarctic islands appear to be Intestinal coccidiosis was reported for wrecked blue penguins in exposed. King and gentoo penguins on the Kerguelen and Crozet Northland but lesions were not described(6). Islands in the subantarctic had antibodies as did yellow-eyed penguins on the Aucklands and Campbell Island, but rockhoppers Haematozoa: A novel blood parasite, Leucocytozoon tawaki,

page 8 Surveillance 28(4) 2001 on Campbell Island did not(61)(62). Suitable vectors are probably Mortality in New Zealand absent from Antarctica but may be present in the subantarctic. Since 1991, diagnostic laboratories have received, and entered on Alternatively, penguins that breed on subantarctic islands may be the wildlife pathology database (Huia), 34 submissions of blue exposed to vectors if they migrate to lower latitudes outside the penguins and four of yellow-eyed penguins. Each case was re- breeding season. evaluated for this review. An additional 213 blue penguins from High seroprevalence of malaria was reported in captive blue northern Otago were examined between 1994 and 1998(13).The penguins in Napier(61) and in free-living birds from Codfish Island, diagnoses from both sources are summarised in the table. to the west of Stewart Island(62). Mortality has not been reported in New Zealand but does occur among blue penguins in Australia(56).

Other protozoa: A blue penguin that died of toxoplasmosis while under rehabilitation in Tasmania had been fed on uncooked meat and held in a household with several cats(66).A tick-borne piroplasmid, presumed to be Babesia sp, has also been found in blue penguins in Australia where it causes mild regenerative anaemia in juveniles (Cunningham cited by Clark and Kerry(46)). Non-infectious causes of mortality Starvation Juvenile blue penguins appear susceptible to periodic mass mortalities that may reflect fluctuations in prey availability. Rockhopper penguins may also be susceptible to fluctuations in prey availability that are influenced by the El Niño-Southern Oscillation(67). Blue penguin wrecks have been recorded in New Zealand since the early 1940s(68). During two large wrecks in Northland in July-August 1973 and April-May 1974, there was a hundredfold increase in the number of beach-cast birds(6). Most were immature females in poor body condition with an empty gastrointestinal tract, depleted fat reserves, muscle atrophy, and moderate parasite burdens. It was concluded that death resulted Conclusions from starvation exacerbated by parasitism and bad weather, as was Most of the disease surveillance on penguins has occurred in the proposed for a wreck of mixed-age blue penguins on the Victorian Antarctic and subantarctic. The focus has been on potentially exotic coast between December 1977 and June 1978(41).A wreck ofblue diseases that may have been introduced through human traffic to penguins in South Australia and Victoria in March-April 1986 also these once remote regions. The risk has been minimised by a involved emaciated and parasitised juvenile birds that washed protocol on Environmental Protection (Madrid, 1998) as part of the ashore following southerly storms(50). Antarctic Treaty, 1991. This banned the importation of live poultry During the summer of 1989-90 a catastrophic event claimed at least and other birds into the Antarctic and requires the inspection of half of the adult breeding population of yellow-eyed penguins on dressed poultry for Newcastle disease and tuberculosis. However, (42) the Otago Peninsula . Moribund birds were neurologically the international regulations are less strict than national quarantine impaired and dead birds were in good body condition with no measures and there are no protocols to control outbreaks of disease evidence of infectious disease, suggesting that marine algal should they occur in Antarctica(74). In New Zealand, the Department biotoxins may have been implicated (J Gill, personal of Conservation bans the transportation of poultry or poultry communication). Over the past two decades marine dinoflagellate products, apart from , to the subantarctic islands. Tourist vessels blooms have claimed a variety of marine vertebrates worldwide do carry such products but are not permitted to discharge organic (69) (70) (71) ranging from and sea birds , to , sirenians , and waste in the vicinity of islands. However, the risks posed by tourist (72) cetaceans . Blooms of neurotoxin-producing diatoms occur and fishing vessel traffic are largely unknown. frequently in New Zealand waters and have the potential to cause Clearly more surveillance is required for penguins in New Zealand neurological signs and death in penguins eating prey that has territories. Of the four endemic species, two are endangered accumulated the toxin(73). Laboratory assays to detect these biotoxins (yellow-eyed and Fiordland) and a third (erect-crested) is in are now available in New Zealand. decline. There have been no studies on disease or mortality factors among Snares crested penguins, Fiordland crested penguins, or

page 9 Surveillance 28(4) 2001 erect-crested penguins, apart from scattered records of parasite rockhopper penguins (Eudyptes chrysocome) from Campbell Island, New Zealand. Journal of Wildlife Diseases 26, 283-5, 1990. occurrence. Studies on yellow-eyed penguins have also been limited (15) Morgan IR, Westbury HA, Caple IW, Campbell J. A survey of virus infection in to serology for avian malaria and aspergillosis, and opportunistic subantarctic penguins on Macquarie Island, . Australian sampling of wrecked birds. The alarming decline of rockhopper Veterinary Journal 57, 333-5, 1981. (16) Alexander DJ, Manvell RJ, Collins MS, Brockman SJ, Westbury HA, et al. penguins on Campbell Island prompted a limited disease Characterization of paramyxoviruses isolated from penguins in Antarctica and investigation in the mid 1980s(14). No firm conclusions could be sub-Antarctica during 1976-1979. Archives of Virology 109, 135-43, 1989. drawn other than the suggestion that avian cholera was probably (17) Morgan IR, Westbury HA, Campbell J. Viral infections of little blue penguins (Eudyptula minor) along the southern coast of Australia. Journal of Wildlife not implicated. A novel enveloped RNA virus was isolated but not Diseases 21, 193-8, 1985. fully characterised and its pathogenicity not established. Research is (18) Morgan IR, Westbury HA. Virological studies of Adélie penguins (Pygoscelis adeliae) in Antarctica. Avian Diseases 25, 1019-26, 1981. continuing into the associated environmental factors but disease is (19) Morgan IR, Westbury HA. Studies of viruses in penguins in the Vestfold Hills. not included (D Thompson, personal communication). Hydrobiologia 165, 263-9, 1988. (20) Pierson GP, Pflow CJ. Newcastle disease surveillance in the United States. 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