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Diseases in This article provides a comprehensive sea review of diseases in New Zealand sea Introduction mammals. New Zealand’s marine fauna has many unique features associated with the geographical location and isolation of the country. The Padraig J. Duignan wide latitudinal range of New Zealand territories, ranging from the sub-tropical Kermadec Islands to the Ross Dependency in Antarctica, presents a wide range of habitats suitable for marine species, both main New Zealand islands but which now has breeding resident and those that migrate seasonally between polar waters concentrations only at the and Campbell Island and the tropics. Little is known about the off-shore movements of during the winter months. In the North, the Hauraki Gulf and Bay New Zealand’s sea mammals, but they are potentially at greater risk of Plenty are home to Bryde’s whales (Balaenoptera edeni) which from transmission of infectious agents than are our terrestrial fauna. occasionally become stranded in the Firth of Thames. Of most Thus, it is prudent to regard our species as part of the greater Pacific significance among the New Zealand cetacean fauna is Hector’s Basin marine province. dolphin (Cephalorhynchus hectori). It is one of the smallest and Based on stranding and sighting records since the later nineteenth rarest oceanic dolphins in the world and is an endemic species with century, New Zealand has a diverse complement of sea mammals. populations off the east and west coasts of the and a These include 38 members of the order cetacea (whales, dolphins relict population between Taranaki and the Manukau Harbour in and porpoises) and seven members of the class pinnipedia (otariidae the North Island(1). Because of its coastal distribution it is vulnerable or eared seals and phocidae or true seals). to human activities such as set-net fishing, and the Banks Peninsula marine sanctuary was established to protect Hector’s The cetaceans are broadly divided into those with baleen dolphins from this activity. (mystacoceti) and those with teeth (odontoceti). Among the former are the great baleen whales some of which are of considerable Three otariid species frequent our coasts. Most common is the New importance in New Zealand waters. These include the southern right Zealand ( forsteri). Because there are rookeries whale (Eubalaena mysticetus), which used to be common around the on or close to the mainland, weaners frequently come ashore in

page Surveillance 27(3) 2000 9 inappropriate locations and suffer the misguided attention of people only on viral and bacterial infections. Compared with North or their . The New Zealand (or Hooker’s) sea (Phocarctos America and Europe, relatively little research has been conducted on hookeri), which is our only endemic species, is encountered less health and disease among New Zealand’s sea mammals. A frequently on the main islands. It is classified as threatened because consequent serious lack of baseline data on native species became of its small population size and limited breeding sites on the apparent during an investigation of mass mortality in 1998. subantarctic islands(2). Occasional pups are born on the Otago coast The prevalence of potentially pathogenic bacteria, viruses, protozoa, and, out of their breeding season, sea are sighted quite and metazoan parasites in these species is unknown, and laboratory commonly around the southern parts of the South Island. The third findings were therefore hard to interpret. Complicating this lack of otariid to frequent our coasts is the sub- published data is a paucity of tissue and serum archives which could (A tropicalis). It does not breed in New Zealand, but strandings and be used to determine the historical prevalence of putative pathogens sightings are not uncommon. in sea mammals.

Among the true seals (phocidae) stranded or sighted around our Viruses coasts are the seal (Hydrurga leptonyx), the southern Avian influenza: The potential of viral infections to regulate marine (Mirounga leonina) and, rarely, Antarctic species such mammal populations was first recognised with the discovery that as the (Lobodon carcinophagus) and fatal avian influenza infections could be transmitted from birds to (Leptonychotes weddelli). both seals and cetaceans in the eastern North Atlantic(9),(10). Although Natural mortality no illness has been attributed to influenza A among marine mammals of the Pacific, a strain of this virus, designated A/whale/ Some understanding of the natural history of a species of interest is PO/19/76 (H1N3), was isolated from a minke whale (Balaenoptera needed when investigating causes of disease or deaths in marine acutorostrata) caught in the South Pacific during the 1975/76 mammals, and deciding what is an acceptable level of mortality is whaling season(11). A close antigenic, genetic and biological the first step. For example, as with most mammals with a high relationship was demonstrated between isolates of influenza A from reproductive rate, mortality of fur seals can exceed 50% in the first a tern and the whale(12). Close associations have also been year of life. That contrasts with the southern right whale in which a demonstrated between isolates from ducks and seals in the Atlantic, calf is produced only every third or fourth year. Thus, very few dead leading to the conclusion that introduction of avian viruses into sea calves could constitute an epidemic. mammals has occurred on several independent occasions and that For most marine mammals, we have too little understanding of these introductions were recent(12). Thus, although no investigation normal population dynamics, so it is hard to decide when unusual of influenza infection has occurred among New Zealand’s marine numbers of deaths have occurred. For some, in which population mammals, the potential for transmission exists. studies have been conducted, elevated mortality rates can be more Morbillivirus: Perhaps of greater significance for easily signalled and decisions as to whether or not to investigate population dynamics are the distemper viruses (family made. Thus, higher than normal rates of stranding, emaciation, and Paramyxoviridae, genus Morbillivirus). Recent reviews cover the death were recognised in New Zealand fur seals during the recent El extensive literature on this topic(6),(7),(13). In 1987, a field-strain of Niño/Southern Oscillation (ENSO) excursions. These events usually canine distemper virus (CDV) killed thousands of Baikal seals cause decreased availability of prey, but occasionally they result in ( sibirica) in Siberia and may also have killed thousands of blooms of toxin-producing algae that can cause paralysis. Deaths in Caspian seals (Phoca caspica) in 1997(14),(15). Since 1988, three new fur seals on the Kaikoura coast during the summer of 1998 coincided members of the genus morbillivirus have been described: phocine with blooms of Gymnodinium mikimotoi (3). However, these deaths distemper virus (PDV) first isolated from harbour seals; porpoise were not investigated. Recently, an algal toxin, domoic acid, was morbillivirus from harbour porpoises (Phocoena phocoena); and implicated in the deaths of California sea lions, dolphin morbillivirus from striped dolphins (Stenella coeruleoalba). californianus (4), suggesting that similar mortalities may occur in Each of these isolates were from that died during epidemics this country. in Europe that killed thousands of animals. The porpoise and Mass deaths in marine mammals appear to be occurring with dolphin viruses are genetically similar and may represent strains of a increased frequency worldwide. Various aetiologies have been host-adapted cetacean morbillivirus (CeMV). Reverse transcriptase implicated, including bacteria(5), viruses (6),(7), and biotoxins (4),(8). The PCR for morbillivirus P and N genes also suggest that the first such event in New Zealand occurred among New Zealand sea morbillivirus of long-finned pilot whales (Globicephala melas) is lions on the Auckland Islands in 1998, and some of the findings from another variant of CeMV(16). Based on the high sero-prevalence of the ensuing investigation are included in this review. specific antibody in both long-finned and short-finned (G macrorhynchus) pilot whales in the western North Atlantic, it was Infectious diseases hypothesised that these species may be the reservoir and vectors for This review and synthesis of unpublished observations will focus CeMV(17) (18). Similar data from the eastern North Atlantic support

page 10 Surveillance 27(3) 2000 this hypothesis(19). A clear parallel exists between this situation and Calicivirus: Since 1972, 20 calicivirus serotypes (family that of PDV among phocid seals of the North Atlantic. For that Caliciviridae) have been isolated from at least five and one virus, serological evidence suggests that harp seals (Phoca cetacean species(34). All of these isolates were made from species groenlandica) are the most probable reservoir host(20),(21),(22). found on the Pacific coast of North America and in the western Arctic(35). Based on serological surveys, several other whale species Serological and pathological surveys of marine mammals in the in the North Pacific appear to have been infected by these Canadian Arctic and along the Atlantic seaboard of North America serotypes(35),(36). This pattern of infection is probably a research have shown that many pinniped and cetacean species have endemic artefact rather than a true indication of a worldwide distribution morbillivirus infections(17),(18),(22),(23),(24),(26). Other species, such as of the viruses. bottlenose dolphins (Tursiops truncatus) and harbour seals, with small fragmented populations, are more vulnerable to Infection in sea mammals is generally benign and is associated with epidemics(25),(26),(27). Phocine distemper appears not to have reached mild vesicular skin disease, though infection might also result the phocid populations along the North American Pacific coast(26),(28). occasionally in premature parturition, neonatal death, encephalitis, However, there is growing evidence that CeMV is present in short- and pneumonia(37),(38). However, the ‘marine caliciviruses’ are beaked common dolphins, Delphinus delphis, in the north-east classified on the basis of sequence homology and pathology into a Pacific and in several dolphin species from the south-west single genogroup which includes vesicular exanthema of swine virus Pacific(29),(30). One case of distemper encephalitis was also reported in (VESV). The latter was associated with costly outbreaks of swine a Pacific white-sided dolphin (Lagenorhynchus obliquidens) stranded disease in the US between 1932 and 1956, and infection probably in Japan(31). originated with the feeding of sea lion offal to pigs(37). There is accumulating evidence of repeated movement of calicivirus Unpublished data for New Zealand indicate a very high serotypes between marine and terrestrial animals and vice versa(34). seroprevalence of CeMV among long-finned pilot whales (Duignan, Furthermore, these serotypes appear to readily cross zoological taxa unpublished). This is consistent with data on this species from the between mammals, birds, reptiles and amphibians(34),(39). The North Atlantic and suggests that pilot whales may play a similar calicivirus status of New Zealand marine mammals is unknown, but reservoir role for the virus in the Pacific(17),(19). Limited data also show these viruses appear to be highly contagious and have no apparent that bottlenose dolphins, short-beaked common dolphins, and dusky host restrictions, so transmission to New Zealand marine mammals dolphins (Lagenorhynchus obscurus) have also been exposed to from avian or other sources is highly probable(39). infection (Duignan unpublished). Although epidemiological studies in New Zealand are still preliminary, it is of concern that the few Poxvirus: Hector’s dolphins accidentally killed in fishing nets had Hector’s dolphins that have been tested were sero-negative. Given skin lesions characteristic of the unclassified poxvirus (family that this species is already threatened with extinction, a CeMV Poxviridae) of cetaceans (Duignan unpublished, Fig. 1). Dolphin pox epidemic in New Zealand waters could pose a serious risk to is primarily a skin disease of young animals that rarely compromises the population. the health of the but severe generalised infections can occur (7),(40),(41). The implications for Hector’s dolphins are not yet known New Zealand fur seals and sea lions have also been exposed to a but investigations are in progress. PDV-like virus as determined serologically but there are too few data to determine the role of morbillivirus in the natural mortality of Orthopox and parapox infections have been reported in (42),(43),(44) either species (Duignan, unpublished). The prevalence of antibody overseas . The status of New Zealand marine mammals and the magnitude of titres among adult female sea lions sampled is unknown. shortly after the peak of the 1998 Auckland Islands mass mortality Herpesvirus: Both fatal encephalitis and benign skin disease caused were certainly too low to implicate the virus in the epidemic(32). by herpesvirus or herpesvirus-like infections have been described Indeed, the sea lions were not expected to be seropositive for two reasons: 1) PDV has not been previously reported in any otariid species worldwide(6); 2), the sea lion population appeared to be too small to maintain an endemic infection. Morbilliviruses generally require populations of several hundred thousand susceptible individuals in order for endemic infection to persist, but there are, at most, only 15,000 sea lions on the Auckland Islands(2) (33). However, it is tempting to speculate that the more numerous fur seals may be a reservoir for sympatric sea lions, and the occurrence of New Zealand fur seals in Australia further suggests that this species could act as a vector between the two countries. More data are required to test this hypothesis. Figure 1: Unclassified poxvirus lesion of Hector’s dolphins.

page Surveillance 27(3) 2000 11 overseas among several species of cetaceans(45),(46),(47),(48). Herpesvirus Pseudomonas sp., Gardnerella vaginalis, and Bordatella infection also appears to be ubiquitous among pinnipeds in Europe, bronchiseptica(67). However, lesions are not always associated with North America and Antarctica(49),(50),(51). The status of New Zealand these findings. Similar bacteria are regularly isolated from marine marine mammals is unknown. mammals in the United States and Europe(68),(69).

Papovavirus: Genital and cutaneous warts associated with Salmonella: During the investigation of the 1998 Auckland Islands papovavirus have been described overseas in several cetacean sea lion mass mortality, four distinct Salmonella serotypes were species(7),(52),(53). The status of New Zealand marine mammals isolated: S Cerro, S Derby, S Newport and S enteritidis phage types is unknown. 4 and 8(70). S Newport is an uncommon serotype in New Zealand. Rabies: Rabies has been identified in a (Phoca hispida) Between 1985 and 1997 there were only four cases confirmed, three from Svalbard, Norway. An unclassified rhabdovirus was isolated from humans and one from an overseas environmental source from a white beaked dolphin (L albirostris) that stranded in the (C. Nicol, ESR Communicable Disease Group, Porirua, pers. comm.). Netherlands (54),(55). Neutralizing antibodies against the latter virus Once case was reported in a pig in 1999 (ESR Communicable Disease were subsequently found in several cetacean and pinniped species Group). However, this serotype was also the most common of nine (68) from the North Sea and Mediterranean (55). The status of New serotypes isolated from marine pinnipeds stranded in California , Zealand marine mammals is unknown. suggesting that it may have a marine reservoir.

Hepnavirus: Hepatitis B virus (family Hepadnaviridae) was : Histopathologically, the 1998 sea identified in a captive Pacific white-sided dolphin with chronic lion mass mortality event was characterised by an acute septicaemia (71) hepatitis(56). The status of New Zealand marine mammals causing necrotizing vasculitis and a haemorrhagic pneumonia . A is unknown. similar disease was detected later in 1998 in a fur seal stranded in Otago(71). Microaerophilic gram negative bacteria isolated from both Adenovirus: Apparently non-pathogenic adenoviruses have been the diseased sea lions and the fur seal possessed the culture isolated from rectal swabs from whales in the Arctic, Gulf of St. characteristics, staining properties and morphological features of a Lawrence and the Antarctic (57),(58),(59). The relationship of these Campylobacter. Genetic characterisation of isolates from both isolates to each other or to adenoviruses from terrestrial mammals is species, based on 16s RNA, showed that some isolates had high not known. The status of New Zealand marine mammals sequence homology with C mucosalis while others had sequence is unknown. homology with Acinetobacter sp. Further investigation into the role Picornavirus: Picornaviruses of unknown significance have been of these isolates in the 1998 sea lion mass mortality is underway. isolated from the intestines of both California grey whales Campylobacter sp. infection has not been associated with disease (Eschrichtius robustus) and European harbour seals(60),(61). The status among marine mammals in other countries. Neither have the of New Zealand marine mammals is unknown. histopathological lesions in the New Zealand sea lions and fur seals been reported in any other species of marine mammal. However, Bacteria Campylobacter sp. have been isolated from several sea bird species There are numerous reports on the isolation of bacteria from marine and from Antarctic fur seals (Arctocephalus gazella) sampled on Bird mammals but the association between the bacteria and disease is Island, South Georgia, so perhaps clinical disease may yet be often tenuous. In New Zealand, the bacterial diseases of particular discovered in other species(72). interest include tuberculosis, salmonellosis and campylobacteriosis. Brucella: Brucellosis has not been diagnosed in New Zealand marine Mycobacteria: Recently, mycobacteria have been isolated from a fur mammals. Brucella sp have been associated with abortion among seal that stranded with granulomatous pneumonia, and from others captive bottlenose dolphins in the US, and with abscesses in seals, incidentally caught in fishing gear, and with lesions in peripheral porpoises and dolphins in the UK(73),(74),(75). Based on serology, there (62) lymph nodes . Genetic characterisation of these isolates, and others is now evidence that infection is widespread in the North Atlantic, from Australian and South American pinnipeds, confirmed that they the Arctic and the Antarctic (76),(77),(78),(79). Genetic and antigenic belong to the M tuberculosis complex and were related to both characterisation of the isolates from bottlenose dolphins indicate (62),(63),(64),(65),66) M tuberculosis and M bovis . However, genetic and that the dolphin organism is a new species now called Brucella biochemical comparison of isolates from pinnipeds worldwide delphini(73). The zoonotic potential of this particular isolate is suggest that the strain infecting New Zealand fur seals is unique unknown but a similar isolate from marine mammals in the UK has (Cousins, pers. comm). Further work is required to determine the caused clinical brucellosis in laboratory worker(80). epidemiology of infection in sea mammals and to compare the Characterization based on genetics, culture characteristics, marine isolates with those from terrestrial mammals. morphology, biochemistry, phage typing, and metabolic profiles, has Frequent bacterial isolates from stranded marine mammals been conducted on isolates from several northern hemisphere in New Zealand include Gemella haemolysans, haemolytic and species (75),(81),(82). The marine mammal isolates appear to be distinct non-haemolytic coliforms, Bacillus sp., Oligella urealytica, from the six known Brucella sp. from terrestrial mammals and it was

page 12 Surveillance 27(3) 2000 suggested that they be assigned to a new species called B maris, with Miller (eds). Zoo and Wildlife Medicine IV. Pp 497-500. W.B. Saunders, three distinct biovars(75). The relationship between the isolates from Philadelphia, 1999. various species and regions has yet to be established. The (7) van Bressem MF, van Waerebeek K, Raga JA. A review of virus infections of epidemiology of infection is likewise unknown but transmission is cetaceans and the potential impact of morbilliviruses, poxviruses and probably through infected body fluids, as in terrestrial mammals. papillomaviruses on host population dynamics. Diseases of Aquatic Organisms One intriguing finding has been the immunohistochemical 38, 53-65, 1999. identification of bacteria with characteristics of B abortus in the uteri (8) Geraci JR, Anderson DM, Timperi RJ, St. Aubin DJ, Early GA. Et al. Humpback of harbour seal lungworms(83). 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