Checklist of marine mammal parasites in New Zealand and Australian waters

Abstract

Marine mammals are long-lived top predators with vagile lifestyles, which often inhabit remote environments. This is especially relevant in the oceanic waters around New Zealand and Australia where cetaceans and pinnipeds are considered as vulnerable and often endangered due to anthropogenic impacts on their habitat. Parasitism is ubiquitous in wildlife, and prevalence of parasitic infections as well as emerging diseases can be valuable bioindicators of the ecology and health of marine mammals. Collecting information about parasite diversity in marine mammals will provide a crucial baseline for assessing their impact on host and ecosystem ecology. New studies on marine mammals in New Zealand and Australian waters have recently added to our knowledge of parasite prevalence, life cycles and taxonomic relationships in the Australasian region, and justify a first host–parasite checklist encompassing all available data. The present checklist comprises 36 species of marine mammals, and 114 species of parasites (helminths, arthropods and protozoans). Mammal species occurring in New Zealand and Australian waters but not included in the checklist represent gaps in our knowledge. The checklist thus serves both as a guide for what information is lacking, as well as a practical resource for scientists working on the ecology and conservation of marine mammals.

Introduction

In the oceanic waters around New Zealand (NZ) and Australia, marine mammals are considered as vulnerable wildlife and often endangered due to anthropogenic impacts on their habitat. Strandings of these long-lived top predators and often pelagic species are rare and individuals are seldom available for data collection. A recent assessment has shown that the conservation status of NZ marine mammals has not improved (Baker et al., 2010). Furthermore, three endemic NZ marine mammals, i.e. NZ sea lion (Phocarctos hookeri), Hector's dolphin (Cephalorhynchus hectori hectori) (both endangered) and Maui's dolphin (Cephalorhynchus hectori maui) (nationally critical) are regarded as threatened. Thirteen taxa are considered data deficient (Baker et al., 2010). Around Australia, at least seven species are classified threatened, among them the iconic blue whale (Balaenoptera musculus), and the conservation status of 25 cetacean species is unknown due to insufficient data (Schumann et al., 2013). Australian and NZ waters include critical feeding and breeding grounds for permanent cetacean residents and visitors that migrate from summer feeding grounds in the Antarctic to the warmer waters of the Australian coast during the winter (Bannister et al., 1996; Salgado Kent et al., 2012). Recent dangers to pinnipeds in Australasian waters include exposure to marine debris and bycatch in fishing gear, which is an acute threat for the endangered Australian sea lion (Neophoca cinerea) (Kovacs et al., 2012) and NZ sea lion (Robertson & Chilvers, 2011). Continuous pressure of anthropogenic impacts such as fisheries, entanglement, vessel strike and chemical and noise pollution has prompted researchers to evaluate the effects of cumulative stress on marine mammals in Oceania and to implement conservation strategies to protect their survival (Kingsford et al., 2009).

Parasitism is ubiquitous in wildlife, and parasites in marine mammals are common. While a certain parasite load may not hamper host physiology, heavy infections can have serious pathogenic effects on host fitness (Measures, 2001; Siebert et al., 2001). Although Australian and NZ waters are a hotspot for marine mammal species richness (Pompa et al., 2011), little is known of their parasite diversity.

Parasitic infections, their prevalence and intensity as well as emerging species have proven to be valuable bioindicators of the ecology and health of marine mammals (Siebert et al., 2006; Lehnert et al., 2014), reflecting habitat use (Aznar et al., 1995), diet (Marcogliese, 2002), social behaviour and population dynamics (Balbuena and Raga, 1994), but also as markers for exposure and detrimental effects of xenobiotics (Sures, 2004; Pascual & Abollo, 2005; Marcogliese & Pietrock, 2011). They reveal evolutionary host–parasite relationships and highlight their biogeography and phylogeny over historical timescales (Anderson, 1982; Leidenberger et al., 2007). In the future, metazoan parasites, emerging infectious diseases and microparasites like viruses may be used as markers for the effects of anthropogenic stress on the health of marine mammals, as their role also as indicators for global change has become evident (Gulland & Hall, 2007; van Bressem et al., 2009). Additionally, some parasites of marine mammals have zoonotic potential, causing public health concerns and economic harm. Both the tapeworm Diphyllobothrium latum and anisakid nematodes (e.g. Anisakis spp., Pseudoterranova spp.) increasingly cause zoonotic infections in humans (Dorny et al., 2009; Shamsi, 2014), and can induce severe gastro-intestinal disease when ingested via undercooked fish (Mattiucci et al., 2013; Bao et al., 2017). These parasites are transmitted to their definitive cetacean and pinniped hosts via infective larvae within prey intermediate hosts, while infected fillets cause economic losses for the fishery industries (Llarena-Reino et al., 2015). Zoonotic protozoans like Giardia and Cryptosporidium are significant enteropathogens in NZ, causing higher infection rates than in other developed countries (Britton et al., 2010). Faeces from humans, pets and farm animals are discharged in runoff, bringing encysted parasites to coastal waters. They are filtered and concentrated by invertebrates and consumed by marine mammals, infecting a wide range of hosts, resulting in morbidity and mortality to some species (Fayer et al., 2004).

Collecting information about parasite diversity in marine mammals will provide a crucial baseline for assessing their impact on host and ecosystem ecology (Poulin et al., 2016). Study design in live marine mammals is restricted by legal as well as ethical constraints so that, since the cessation of whaling, data are collected mostly opportunistically from stranded or bycaught individuals. In the oceanic waters of the southern hemisphere, many species are seldom found stranded; therefore, few parasitological records (Berón-Vera et al., 2008; Nikolov et al., 2010) exist. Parasitology increasingly complements marine ecology to further our understanding of ecosystem dynamics (Poulin et al., 2016), but, so far, little is known about the biodiversity of Australasian marine parasites (Poulin, 2004; Stockin et al., 2009). New studies on marine mammals in NZ and Australian waters have recently added to our knowledge about parasite prevalence, life cycles (Tomo et al., 2010; Lehnert et al., 2017) and taxonomic relationships (Shamsi et al., 2012, Hernández-Orts et al., 2017), and justify a first host–parasite checklist encompassing all available data. Previous marine mammal parasite checklists include Baylis (1932), Delyamure (1955), Dailey & Brownell (1972), Raga (1994), Felix (2013) and Fraija-Fernández et al. (2016). Most include host data, but geographical data are scanty. Australian lists include that of the internal parasites of mammals by MacKerras (1958), the references of which are unfortunately disconnected from the text and, therefore, not useable, and Arundel (1978), which records parasites found in all marine mammals that are found in Australian waters. This list, however, does not differentiate between parasites found in the host species within Australian waters and those found in the same species in other parts of the world. There are no equivalent lists for the marine mammals of NZ.

A couple of useful lists are available for particular host taxa (e.g. Blair, 1981: dugong monostome digeneans; Bowie, 1984: bottle-nosed dolphin parasites), and parasite taxa (e.g. Price, 1932: trematodes in marine mammals; Leung, 1967: whale lice of cetaceans). There have also been compilations with a veterinary perspective (e.g. Duignan, 2000; Dailey, 2001; Ladds, 2009; McFarlane et al., 2009) and those looking at certain pathologies (e.g. Baylis & Daubney, 1925: lungworms of cetacean; Spratt, 2002: respiratory parasites in Australian mammals; Measures, 2001: lungworms of marine mammals).

Here, we present a host–parasite checklist collating all information about the metazoan and protozoan parasites of marine mammals in NZ and Australian waters. Although viruses (e.g. morbillivirus) and bacteria (e.g. Brucella) are relevant pathogens with zoonotic potential that can cause mortality in marine mammals (Castinel et al., 2007), we limit our list to the better-known eukaryotic parasites. Where possible, we also identify knowledge gaps and research needs, especially with regard to human interactions and zoonoses, as well as marine mammal conservation.

Material and methods

We present a list of the parasites found in pinnipeds, cetaceans and sirenians of NZ and Australia, as far as possible up to date at the time of publication. The list was assembled from primary publications found through searches on Google Scholar using all combinations of relevant keywords, plus searches of the reference lists in those publications. The parasites are presented in alphabetical order of Families under their relevant Phylum, Class and Order. Within Families, species are listed in alphabetical order. Classifications followed for helminths are: Anderson et al. (2009) (Nematoda); Caira & Jensen (2017) (Cestoda); Olson et al. (2003), Gibson et al. (2002), Jones et al. (2005) and Bray et al. (2008) (Trematoda); and Amin (2013) (Acanthocephala). For arthropods and protozoa higher taxonomy was taken from the records cited, or from searches for more up-to-date phylogenetic studies. Synonyms are taken from the references in square brackets following the entries, with corrections and updates from primary sources in some cases.

Localities of records are given as indicated in the original source. Standard abbreviations are used for Australian states: New South Wales, NSW; Queensland, QLD; South Australia, SA; Victoria, VIC; Tasmania, TAS; Western Australia, WA; and New Zealand, NZ. We have included only those records that fall within the geographical boundaries of Australia and NZ, and their subantarctic islands. Records for mainland Antarctica have not been included, because there is no geographical distinction between Australian or NZ-held territory, and those territories belonging to other countries. Although there are many records for mainland Antarctica, these are best dealt with as a separate entity.

Where there is more than one reference source, hosts and localities bear a superscript number that refers to the numbered reference. In a few cases, where references cite differing infection sites, these are also numbered with the relevant superscript number. Multiple references are listed in chronological order.

For host taxonomy we have adhered to WoRMS (2018) (Pinnipeda and Cetacea), Berta and Churchill (2012) (Pinnipeda) and Perrin (2018) (Cetacea). Host names listed are considered to be up to date as of publication. Where hosts were named differently in the original source, we have noted this in the relevant Remarks section. Hosts’ common names can be found in the host–parasite list.

The developmental stage of the parasite has been noted, where given in the original source. If such information was not given, worms are assumed to be adult if egg presence is noted or egg measurements are given.

Remarks sections contain information on intermediate hosts when available, notes on prevalence and intensity, and on pathology when available; short summaries of the latest research on disputed or complicated species; and mention of molecular data if available (i.e. from papers listed in the references and placed on GenBank) with the genetic markers used.

A list of host–parasite associations follows the parasite–host checklist.

Results

Parasite–host list

• Phylum: Acanthocephala

• Class: Palaeoacanthocephala Meyer, 1931

• Order: Polymorphida Petrochenko, 1956

• Family: Polymorphidae Meyer, 1931

Parasite name: Bolbosoma balaenae (Gmelin, 1790) Porta, 1908

• Synonyms. Bolborhynchus porrigens Porta, 1906; Bolbosoma porrigens (Rudolphi, 1814) Porta, 1908; Echinorhynchus balaenae Gmelin, 1790; E. lendix Phipps, 1774; E. mysticeti Beneden, 1870; E. porrigens Rudolphi, 1819; Sipunculus lendix Phipps, 1774 [Amin, 2013; Yamaguti, 1963].

• Hosts. Megaptera novaeangliae.

• Localities. Bondi Beach, NSW.

• Infection site.

• Stage.

• References. Johnston & Deland (1929).

• Remarks. Called B. porrigens in this paper. Host reported as probably M. nodosa (= longimana); both now synonymized with M. novaeangliae.

Parasite name: Bolbosoma capitatum (von Linstow, 1880) Porta, 1908

• Synonyms. Bolborhynchus capitatus (von Linstow, 1880) Porta, 1908; B. physeteris Gubanov, 1952 (fide Amin & Margolis, 1998); Echinorhynchus capitatus von Linstow, 1880 [Amin, 2013; Felix, 2013].

• Hosts. Pseudorca crassidens (1, 3), Globicephala melas (2).

• Localities. (1) Prime's Beach, St. Vincent Gulf, SA; (2) Macquarie Harbour, TAS; (3) Augusta WA.

• Infection site. Intestine.

• Stage.

• References. (1) Edmonds (1957b); (2) McManus et al. (1984); (3) Edmonds (1987).

• Remarks. Host reported in Edmonds (1957b) as Melon-headed whale (G. melaenas = G. melas); later corrected to false killer whale in Edmonds (1987).

Parasite name: Bolbosoma sp.

• Synonyms.

• Hosts. Tursiops truncatus.

• Localities. Otago Harbour, NZ.

• Infection site. Intestine.

• Stage. Immature.

• References. Bowie (1984).

• Remarks. Single specimen.

Parasite name: Corynosoma australe Johnston, 1937

• Synonyms. Corynosoma otariae Morino & Boero, 1960 [Amin, 2013].

• Hosts. Arctocephalus pusillus doriferus (3), Arctophoca australis forsteri (4), Hydrurga leptonyx (2), Neophocus cinerea (1, 5), Phocarctos hookeri (2, 6).

• Locality. (1) Pearson Island, SA; (2) Auckland/Campbell Island, NZ; (3) Phillip Island, VIC; (4) in captivity, NZ; (5) Port Adelaide and Dangerous Reef, SA; (6) NZ.

• Infection site. Small and large intestine.

• Stage. Adult.

• References. (1) Johnston (1937); (2) Johnston & Edmonds (1953); (3) Obendorf & Presidente (1978); (4) Cordes & O'Hara (1979); (5) Smales (1986); (6) García-Varela et al. (2013).

• Remarks. Johnston (1937) described host as A. forsteri, but corrected this to N. cinerea in Johnston & Mawson (1941). Although Johnston & Edmonds (1953) listed a number of ways in which their specimens differed from those of Johnston (1937), they nonetheless record them as C. australe. Smales (1986) agrees in her list of comparative specimens of the species. In her paper, the host is called A. pusillus. Johnston & Edmonds (1953) found immature specimens in Phalacrocorax colensoi. Molecular data available (18S, 28S, cox1) (6).

Parasite name: Corynosoma bullosum (von Linstow, 1892) Railliet & Henry, 1907

• Synonyms. Corynosoma arctocephali Zdzitowiecki, 1984; C. mirabilis Skrjabin, 1966 (fide Zdzitowiecki, 1986); C. singularis Skrjabin & Nikolski, 1971, in part; Echinorhynchus bullosus von Linstow, 1892 [The World Register of Marine Species (WoRMS); Amin, 2013].

• Hosts. Hydrurga leptonyx (2), Mirounga leonine (1–3).

• Locality. (1) Auckland/Campbell Island, NZ; (2, 3) Heard Island, Australia; (2) Macquarie Island, Australia.

• Infection site. Intestine.

• Stage. Adult.

• References. (1) Johnston & Edmonds (1953); (2) Edmonds (1955); (3) Edmonds (1957a).

• Remarks. Edmonds reported 200 (1957a), and over 100 (1955) specimens from single hosts. He states that the larval stages are found encysted in the mesenteries of Notothenia coriiceps.

Parasite name: Corynosoma cetaceum Johnston & Best, 1942

• Synonyms. Corynosoma semerme sensu Cordero, 1933; Polymorphus arctocephali Smales, 1986 (nec arctocephalus); Polymorphus (Polymorphus) cetaceum (Johnston & Best, 1942) Schmidt & Dailey, 1971 [Amin, 2013].

• Hosts. Arctocephalus pusillus doriferus (2), Delphinus delphis (1), Tursiops truncatus (1).

• Locality. (1) Port Lincoln, SA; (1) St. Vincent Gulf, SA; (2) Phillip Island, VIC.

• Infection site. Stomach and small intestine.

• Stage. Adult.

• References. (1) Johnston & Best (1942); (2) Smales (1986).

• Remarks. Described as new species in Johnston & Best (1942). Described as P. arctocephali n. sp., and host as A. pusillus in Smales (1986), synonymized by Aznar et al. (1999).

Parasite name: Coryosoma semerme (Forssell, 1904) Lühe, 1905

• Synonyms. Corynosoma gibber (Olsson, 1894) Lühe, 1911; Echinorhynchus semermis Forssell, 1904 (Amin, 2013).

• Hosts. Phocarctos hookeri.

• Locality. Auckland/Campbell Island, NZ.

• Infection site.

• Stage.

• References. Johnston & Edmonds (1953).

• Remarks. Host called Otaria hookeri in this paper.

Parasite name: Corynosoma sp.

• Synonyms.

• Hosts. Arctophoca australis forsteri (2), Cephalorhynchus hectori (3), Delphinus delphis (1), Hydrurga Leptonyx (2), Neophoca cinerea (4).

• Locality. (1) St. Vincent Gulf, SA; (2) Auckland/Campbell Island, NZ; (3) Canterbury coast, NZ; (4) Port Adelaide, SA.

• Infection site. Intestine.

• Stage.

• References. (1) Johnston & Deland (1929); (2) Johnston & Edmonds (1953); (3) McKenzie & Blair (1983); (4) Smales (1986).

• Remarks. The specimens of Johnston and Deland (1929) were later designated as C. cetaceum (Johnston & Best, 1942). Host called Otaria forsteri in Johnston and Edmonds (1953). McKenzie & Blair (1983) reported finding seven specimens in one of three dolphins.

• Phylum: Nemathelminthes

• Class: Nematoda Rudolphi, 1808

• Order: Ascaridida Sprehn, 1927

• Family: Anisakidae Railliet & Henry, 1912

Parasite name: Anisakis berlandi Mattiucci, Nascetti, Cianchi, Paggi, Arduino, Margolis, Brattey, Webb, D'Amelio, Orecchia & Bullini, 1997

• Synonyms. Anisakis simplex C, of Mattiucci et al. (1997).

• Hosts. Kogia sima.

• Locality. Coast of NSW.

• Infection site. Stomach.

• Stage. Larva.

• References. Shamsi et al. (2012).

• Remarks. Molecular data available (ITS1&2). The specimens in this paper referred to as A. simplex C following Mattiucci et al. (1997) were later assigned to A. berlandi by Mattiucci et al. (2014).

Parasite name: Anisakis brevispiculata Dollfus, 1966

• Synonyms.

• Hosts. Kogia sima.

• Locality. Coast of NSW.

• Infection site. Stomach.

• Stage. Adult.

• References. Shamsi et al. (2012).

• Remarks. Molecular data available (ITS1&2).

Parasite name: Anisakis nascettii Mattiucci, Paoletti & Webb, 2009

• Synonyms. Anisakis sp. of Pontes et al. (2005) and Iglesias et al. (2008); Anisakis sp. of Valentini et al. (2006).

• Hosts. Mesoplodon bowdoini, M. grayi, M. layardii.

• Locality. South Pacific Ocean, off NZ.

• Infection site. Stomach.

• Stage. Adult.

• References. Mattiucci et al. (2009).

• Remarks. Molecular data available (cox2). The intermediate host was confirmed molecularly as the squid, Onykia ingens by Mattiucci et al. (2009) (then Moroteuthis ingens).

Parasite name: Anisakis oceanica (Johnston & Mawson, 1951) Davey, 1971

• Synonyms. Stomachus oceanicus Johnston & Mawson, 1951.

• Hosts. Globicephala melas.

• Locality. Coast of NSW.

• Infection site. Stomach.

• Stage.

• References. Johnston & Mawson (1951).

• Remarks. Described as S. oceanicus in this paper, with the host called Globicephalus ventricosas. Davey (1971) synonymized this species with A. physeteris, but Shamsi et al. (2012) and Shamsi (2014) have kept the species separate as a sp. inq. Species is also accepted by Mattiucci et al. (2005).

Parasite name: Anisakis pegreffii Campana-Rouget & Biocca, 1955

• Synonyms.

• Hosts. Delphinus delphis, Tursiops truncatus.

• Locality. Blairgowrie and Apollo Bay, VIC.

• Infection site. Stomach.

• Stage. Adult plus immatures in D. delphis, larvae in T. truncatus.

• References. Shamsi et al. (2012).

• Remarks. Molecular data available (ITS1&2).

Parasite name: Anisakis physeteris Baylis, 1923

• Synonyms. Anisakis skrjabini Mozgovoi, 1949 [WoRMS].

• Hosts. Kogia breviceps.

• Locality. Wellington Beach, NZ.

• Infection site. Stomach.

• Stage.

• References. Hurst (1980).

• Remarks. A single host harboured 61 specimens.

Parasite name: Anisakis simplex sensu lato

• Synonyms. Anisakis catodontis Baylis, 1929; A. ivanizkii Mozgovoi, 1949; A. kogiae Johnston & Mawson, 1939; A. kuekenthalii (Cobb, 1889) Baylis, 1920; A. marina (Linnaeus, 1767) van Thiel, 1966; A. patagonica (von Linstow, 1880) Davey, 1971; A. rosmari (Baylis, 1916) Baylis, 1920; A. similis (Baird, 1853) Baylis, 1920; A. simplex (Rudolphi, 1809) Dujardin, 1845; A. tridentata Kreis, 1939; Ascaris kuekenthalii Cobb, 1889; As. similis Baird, 1853; Capsularia marina (Linnaeus, 1767) Johnston & Mawson, 1943; Stomachus similis (Baird, 1853) Davey, 1971 and many more [WoRMS; Davey 1971].

• Hosts. Arctophoca australis forsteri (11), Delphinus delphis (2), Globicephala melas (14), Hydrurga leptonyx (2, 5, 6), Kogia breviceps (1, 11), Lagenodelphis hosei (12), Lagenorhynchus obscurus (3, 7), Mirounga leonine (4, 6, 9, 10), Peponocephala electra (8), Tursiops truncatus (13).

• Locality. (1) Port Victoria, Spencer Gulf, SA; (1) Moreton Bay, QLD; (2) Sydney Harbour, NSW; (2) Port Adelaide, SA; (3) Cook Strait, NZ; (4, 6, 9) Macquarie Island, TAS; (5) Campbell Island, NZ; (6) Heard Island, Australia; (7) NZ coast; (8) Moreton Island, QLD; (8) Tweed Heads, NSW; (10) in captivity, NZ; (11) Wellington Beach, NZ; (12) Corio Bay, VIC; (13) Otago Harbour, NZ; (14) McIntyres Beach, Falmouth, and Macquarie Harbour, TAS.

• Infection site. Stomach.

• Stage. All stages are reported from larvae to adults.

• References. Johnston & Mawson ((1) 1939, (2) 1941, (3) 1942, (4) 1945, (5) 1953); (6) Mawson (1953); (7) Brundson (1956); (8) Cannon (1977); (9) Morgan et al. (1978); (10) Cordes & O'Hara (1979); (11) Hurst (1980); (12) McColl & Obendorf (1982); (13) Bowie (1984); (14) McManus et al. (1984).

• Remarks. Anisakis simplex is widely accepted to be a complex of closely related species (Nascetti et al., 1986; Mattiucci et al., 1997, 2009, 2014; D'Amelio et al., 2000; Abe et al., 2006; Abe, 2008), and the name has been attached to specimens the world over and in numerous hosts. Davey (1971) alone lists 34 different hosts from both Cetacea and Pinnipedia. While it is not possible to assign specimens reported in the literature over the years, Mattiucci et al. (2014) have made worthy inroads into resolving some of the ambiguity. As a result of their findings, it is likely that at least some of those specimens found in NZ will be their newly erected A. berlandi Mattiucci, Cipriani, Webb, Paoletti, Marcer, Bellisario, Gibson & Nascetti, 2014 (See entry for Anisakis berlandi above). Without further information, or examining each of the specimens mentioned in the literature, we here assign all references to A. simplex in the Australasian region, except those of Mattiucci et al. (2014), which have been characterized to A. simplex sensu lato. Anisakis simplex has been shown to occur as a larva in many species of fish under various names (Johnston & Mawson, 1943a, b, 1945, 1951, 1953). Brundson (1956) was able infect eels with encapsulated A. simplex from barracouta, where they re-established, demonstrating that horizontal infection is possible. This is not surprising in a system where the species complex appears to be highly generalist in its choice of host and where there exists a cascade in size of predatory fish. The larva must be able to withstand being eaten by a succession of fish before being finally taken by a cetacean, where it can mature in the stomach. Hurst (1980) completed the life cycle of A. simplex with Nyctiphanes australis (first intermediate host), large fish – for example, Thyrsites atun, Trachurus sp. (second intermediate host) – and squid or small fish – for example, hoki, anchovy, sprat (paratenic host). While host specificity does appear extremely broad when all reports are taken into account, this effect may be exaggerated by the fact that A. simplex is, in fact, many cryptic species that are yet to be fully disentangled. In the above references the species is called A. kogiae (1), A. similis (2), S. marinus (7) and S. similis (4, 6). Host species are called D. forsteri (2). In the literature cited in this section, the maximum number of worms per infection amounted to several hundred (13). The species is widely reported to cause gastric ulceration and, according to some sources, mortality (10, 11).

Parasite name: Anisakis sp.

• Synonyms.

• Hosts. Arctocephalus pusillus doriferus (1), Arctophoca australis forsteri (2), Balaenoptera acutorostrata (3), Globicephalus melas (6), Kogia breviceps (7), Mesoplodon grayi (5), M. hectori (4), Phocoena dioptrica (7).

• Locality. (1) Franklin Island Derwent Heads, TAS; (2) in captivity, NZ; (3) Pigeon Bay, Banks Peninsula, NZ; (4) Oneroa Bay, Waiheke Island Hauraki Gulf, NZ; (5) NZ coast; (6) Farewell Spit, Golden Bay, NZ; (7) Moeraki Beach, NZ; (7) Caroline Bay, Timaru, NZ.

• Infection site. Stomach and intestine. Mouth (K. breviceps).⁷

• Stage. Larva, immature, adult.

• References. (1) Johnston & Mawson (1941); (2) Cordes & O'Hara (1979); (3) Dawson & Slooten (1990); (4) Baker et al. (2001); (5) Valentini et al. (2006); (6) Beatson & O'Shea (2009); (7) Lehnert et al. (2017).

• Remarks. Molecular data available (cox2) (5). The specimens from Grey's beaked whale (Valentini et al., 2006) grouped with some from the South African coast, also unnamed. They were closest to Anisakis ziphidarum in phylogenic analyses. A lack of adult specimens limited the morphological description and proper naming of this new species. Type 1 larvae were identified from Aphanopus carbo (black scabbard fish) from Madeira and from Trachurus trachurus (Atlantic horse mackerel) from the North Atlantic. The host in Johnston & Mawson's (1941) report is called Gypsophoca tasmanica.

Parasite name: Anisakis typica (Diesing, 1860) Baylis, 1920

• Synonyms. Anisakis alexandri Hsü & Hoeppli, 1933; A. tursiopis Crusz, 1946; Ascaris typica (Diesing, 1860) Jägerskiöld, 1894; Conocephalus typicus Diesing, 1860; Peritracheilus typicus (Diesing, 1860) Jägerskiöld, 1894 [WoRMS; Li et al., 2016].

• Hosts. Peponocephala electra.

• Locality. Moreton Island, QLD; Tweed Heads, NSW.

• Infection site. Stomach.

• Stage. Adults and immatures.

• References. Cannon (1977).

• Remarks. Shamsi (2014) note that A. typica specimens in Australia appear to be genetically different from those reported in other countries. Jabbar et al. (2012) found larval stages (confirmed by DNA sequence) in a number of different fish species.

Parasite name: Anisakis ziphidarum Paggi, Nascetti, Webb, Mattiucci, Cianchi & Bullini, 1988

• Synonyms.

• Hosts. Mesoplodon bowdoini.

• Locality. South Pacific Ocean, off NZ.

• Infection site. Stomach.

• Stage. Adult.

• References. Mattiucci et al. (2009).

• Remarks. Molecular data available (cox2).

Parasite name: Contracaecum mirounga Nikolskii, 1974

• Synonyms.

• Hosts. Hydrurga leptonyx, Mirounga leonina.

• Locality. Heard Island and Macquarie Island, Australia.

• Infection site. Stomach.

• Stage.

• References. Fagerholm (1988).

• Remarks. Using the arrangement of the male caudal papillae, this study found that C. mirounga was restricted to southern hemisphere hosts, compared to C. osculatum and C. ogmorhini, which were more widespread in both hemispheres.

Parasite name: Contracaecum ogmorhini Johnston & Mawson, 1941

• Synonyms. Contracaecum corderoi Lent & Freitas, 1948; C. ogmohini Johnston & Mawson, 1941 (lapsus); C. osculatum of Flores-Barroeta et al., 1961 [Fagerholm & Gibson, 1987].

• Hosts. Arctocephalus pusillus doriferus (2), Hydrurga leptonyx (1).

• Locality. (1) Port Adelaide, SA; (2) coast of NZ.

• Infection site.

• Stage. Adult.

• References. (1) Johnston & Mawson (1941); (2) Mattiucci et al. (2003).

• Remarks. Molecular data available (cytb) (2). Described as new species in Johnston & Mawson (1941), but with a lapsus in the main heading of ‘ogmohini’. Has been accepted as ogmorhini ever since.

Parasite name: Contracaecum osculatum (Rudolphi, 1802) Baylis, 1920

• Synonyms. Ascaris osculata Rudolphi, 1802; C. antarcticum Johnston, 1937; C. gypsophocae Johnston & Mawson, 1941; Phocascaris hydrurgae Johnston & Mawson, 1941 [WoRMS].

• Hosts. Arctocephalus pusillus doriferus (3), Arctophoca australis forsteri (3, 5, 6), Hydruruga leptonyx (2, 3), Mirounga leonine (3, 4), Neophoca cinerea (1).

• Locality. (1) Pearson Island, SA; (2) Port Adelaide, SA; (3) Franklin Island Derwent Heads, TAS; (3, 4) Macquarie Island, TAS; (3) Heard Island, Australia; (5) in captivity, NZ; (6) Open Bay Island, Wellington, NZ.

• Infection site. Stomach and small intestine.

• Stage. Adults and immatures.

• References. (1) Johnston (1937); (2) Johnston & Mawson (1941); (3) Mawson (1953): (4) Morgan et al. (1978); (5) Cordes & O'Hara (1979); (6) Hurst (1980).

• Remarks. Described as new species, C. gypsophocae and as P. hydrurgae in Johnston & Mawson (1941). Both synonymized by Johnston & Mawson (1945). Original host recorded as Gypsophoca tasmanica. Hosts recorded as Arctocephalus forsteri (5, 6). Johnston's (1937) original description stated the host as A. forsteri, but this was corrected to N. cinerea in Johnston and Best (1942). Hurst (1980) recorded 100% prevalence for this anisakid in the NZ fur seal. They found an infection range of 5–344 in the seals examined, and the presence of C. osculatum was associated with gastric ulcers.

Parasite name: Contracaecum radiatum (von Linstow, 1907) Baylis 1920

• Synonyms. Ascaris falcigera Railliet & Henry, 1907; A. osculaia von Linstow, 1892; A. radiata von Linstow, 1907; Contracaecum falcigerum (Railliet & Henry, 1907) Baylis, 1920; Kathleena radiata Leiper & Atkinson, 1915 [WoRMS; Baylis 1937].

• Hosts. Hydrurga leptonyx, Mirounga leonina.

• Locality. Heard Island, Australia.

• Infection site.

• Stage.

• References. Mawson (1953).

• Remarks.

Parasite name: Contracaecum sp.

• Synonyms.

• Hosts. Cephalorhynchus hectori.

• Locality. Canterbury coast, NZ.

• Infection site. Stomach.

• Stage.

• References. McKenzie & Blair (1983).

• Remarks. The material reported in this paper may have been two different species of Contracaecum from three dolphins. One was found as a single immature female in one dolphin, the second species was in all three dolphins, and 15 specimens were taken in total.

Parasite name: Phocascaris sp.

• Synonyms.

• Hosts. Cephalorhynchus hectori.

• Locality. Canterbury coast, NZ.

• Infection site. Stomach.

• Stage. Immature.

• References. McKenzie & Blair (1983).

• Remarks. A single, female, specimen in one of three dolphins.

Parasite name: Pseudoterranova decipiens (Krabbe, 1878) Gibson, 1983

• Synonyms. Agamonema piscium Schneider, 1862; Ag. campbelli Chatin, 1885; Ascaris bulbosa Cobb, 1888; A. capsularia Stiles and Hasall, 1899, in part; A. capsularia Baylis, 1916, in part; A. decipiens Krabbe, 1878; A. rectangula Linstow, 1888; A. simplex von Linstow, 1888; Phocanema decipiens (Krabbe, 1878) Myers, 1959; Physaloptera guiarti Garin, 1913; Porrocaecum decipiens (Krabbe, 1878) Baylis, 1920; Por. piscium Johnston & Mawson, 1943; Por. capsularia Dogiel, 1932; Terranova decipiens (Krabbe, 1878) Mozgovoi, 1953; T. piscium (Rudolphi, 1809) Johnston and Mawson, 1943 [Johnston & Mawson, 1945; Myers, 1959].

• Hosts. Arctophoca australis forsteri (3, 5), Hydrurga leptonyx (3, 4), Mirounga leonine (2, 3), Kogia breviceps (5), Phocarctos hookeri (1, 3, 5).

• Locality. (1, 3) Campbell Island, NZ; (2, 4) Macquarie Island, TAS; (3, 5) Auckland Island, NZ; (5) Wellington Beach, NZ.

• Infection site. Stomach.

• Stage. Adult plus immatures.

• References. (1) Johnston & Mawson (1943a); (2) Johnston & Mawson (1945); (3) Johnston & Mawson (1953); (4) Mawson (1953); (5) Hurst (1980).

• Remarks. Mawson (1953) reported a prevalence of seven out of nine leopard seals from Macquarie Island. Hurst reported a range of infection between one and 122 per host animal. Hurst's (1980) data include both pinniped and cetacean hosts. Johnston & Mawson (1943a) referred to this species as Por. decipiens (Krabbe, 1878) Baylis, 1920 and (1945, 1953) T. piscium. Various fish are recorded as intermediate hosts; Thyrsites atun (5), notothenioid species (1, 3) and the flounder Rhombosolea sp. (3). Hosts variously called Arctocephalus hookeri (1), Otaria forsteri (3), O. hookeri (3). Mawson (1953) also reported this species from the Royal penguin Eudyptes schlegeli Finsch, 1876.

Parasite name: Pseudoterranova kogiae (Johnston & Mawson, 1939) Mozgovoi, 1951

• Synonyms. Porrocaecum kogiae Johnston & Mawson, 1939; Terranova kogiae (Johnston & Mawson, 1939) Johnston & Mawson, 1945 [Johnston & Mawson, 1945; Deardorff & Overstreet, 1981].

• Hosts. Kogia breviceps (1, 2).

• Locality. (1) Port Victoria, Spencer Gulf, SA and Moreton Bay, QLD; (2) Wellington Beach, NZ.

• Infection site. Stomach.

• Stage. Adults and larvae.

• References. (1) Johnston & Mawson (1939); (2) Hurst (1980).

• Remarks. Called P. kogiae in Johnston & Mawson (1939). Intermediate fish hosts include the barracouta (Thyrsites atun), subantarctic cod (Notothaenia sp.) (Hurst, 1984), hoki (Macruronus novaezelandiae) and ling (Genypterus blacodes) (Grabda & Ślósarczyk, 1981).

• Family: Ascarididae Baird, 1853

Parasite name: Paradujardinia halicoris (Owen, 1833) Travassos, 1933

• Synonyms. Ascaris dugonis Brandt, 1846; A. halichoris Owen, 1833; Dujardinia halicoris (Owen, 1833) Baylis, 1920; Dujardinascaris halicoris (Owen, 1833) Baylis, 1947 [Sprent, 1980].

• Hosts. Dugong dugon (1–4).

• Locality. (1) Wallum Creek, North Stradbroke Island, QLD; (2) Yarrabah, near Cairns, QLD; (3) north coast of QLD; (4) coast of QLD.

• Infection site. Stomach and intestine.

• Stage.

• References. (1) Dexler & Freund (1906); (2) Johnston & Mawson (1941); (3) Sprent (1980); (4) Owen et al. (2012).

• Remarks. Johnston & Mawson (1941) called their specimens Dujardinia halicoris. Owen et al. (2012) reported the presence of 5500 to 6000 worms in a single dugong, with the massive impaction possibly the cause of death of the individual.

• Order: Spirurida Chitwood, 1933

• Family: Acuariidae Railliet, Henry & Sisoff, 1912

Parasite name: Acuaria sp.

• Synonyms.

• Hosts. Cephalorhynchus hectori.

• Locality. Canterbury coast, NZ.

• Infection site. Stomach.

• Stage. Larva.

• References. McKenzie & Blair (1983).

• Remarks. A single specimen was found in one of three dolphins.

• Family: Filariidae Weinland, 1858

Parasite name: Filaria sensu lato sp.

• Synonyms.

• Hosts. Mirounga leonina.

• Locality. Heard Island, Australia.

• Infection site. Blood vessels.

• Stage. Adult.

• References. Mawson (1953).

• Remarks.

• Family: Gnathostomatidae Railliet, 1895

Parasite name: Echinocephalus overstreeti Deardorff & Ko, 1983

• Synonyms.

• Hosts. Delphinus delphis.

• Locality. St. Vincent Gulf, SA.

• Infection site. Intestine.

• Stage. Immature.

• References. Johnston & Mawson (1941).

• Remarks. This species is usually found as adults in elasmobranchs, with the larval stages in molluscs and teleosts. This finding of a larval stage in the intestine of a dolphin almost certainly represents an accidental infection from predation on a paratenic fish host. Johnston and Mawson (1941) listed this as Echinocephalus uncinatus Molin, 1858. However, Beveridge (1987) showed that all adult and larval specimens found in elasmobranchs in Australian waters belonged to E. overstreeti, and that earlier records of E. uncinatus can probably be attributed to E. overstreeti. Moravec and Justine (2006), however, questioned this decision.

• Family: Tetrameridae Travassos, 1914

Parasite name: Crassicauda boopis Baylis, 1920

• Synonyms. Crassicauda pacifica Margolis & Pike, 1955 [WoRMS].

• Hosts. Ziphius cavirostris.

• Locality. Kiritehere Beach, Purakanui Bay, Mahia Peninsula, NZ.

• Infection site. Kidneys.

• Stage. Adult.

• References. Duignan (2003).

• Remarks. Crassicauda boopis is recorded as pathogenic in this paper. The worms destroy functional renal elements and cause physical obstruction of the urinary ducts, likely causing renal failure in some cases.

Parasite name: Crassicauda grampicola Johnston & Mawson, 1941

• Synonyms.

• Hosts. Grampus griseus.

• Locality. Manley, NSW.

• Infection site. Pterygoid sinus.

• Stage. Adults and immatures.

• References. Johnston & Mawson (1941).

• Remarks.

Parasite name: Crassicauda magna Johnston & Mawson, 1939

• Synonyms. Crassicauda duguyi Dollfuss, 1966 [Jabbar et al., 2015].

• Hosts. Kogia breviceps (1, 2).

• Locality. (1, 2) Moreton Bay, QLD; (1) Port Victoria and Spencer Gulf, SA.

• Infection site. Subcutaneous tissue and connective tissue of neck.

• Stage. Adult.

• References. (1) Johnston & Mawson (1939); (2) Jabbar et al. (2015).

• Remarks. Molecular data available (18S) (2). The phylogeny in Jabbar et al. (2015) moves the genus Crassicauda from Tetrameridae to Acuariidae. This huge worm seems to be found only in fragments due to the difficulty of extracting it whole from the subcutaneous flesh.

Parasite name: Crassicauda sp.

• Synonyms.

• Hosts. Lagenorhynchus obscurus (1, 2), Ziphius cavitostris (1), Pseudorca crassidens (3), Delphinus delphis (4, 5).

• Locality. (1, 2, 4) NZ; (3) NSW; (5) Buckland's Beach, Howick, NZ.

• Infection site. (1, 3) Pterygoid sinus; (1, 2) kidneys; (1, 2) urogenital system; (1, 2) cranial sinuses; (1, 5) fascia; (3) middle ear; (4) mammary ducts.

• Stage.

• References. Duignan ((1) 2000, (2) 2003); (3) J. Boulton in Ladds (2009); (4) Stockin et al. (2009); (5) Lehnert et al. (2017).

• Remarks. Undoubtedly, based upon infection site alone, these records refer to different species of Crassicauda. Most records agree that worms of this genus embed their cephalic end into the tissues of their favoured organ, with the body of the worm projecting freely into the lumen or sinus for the release of the eggs. The species occurring in kidneys of Z. cavirostris caused ‘…massive destruction of reniculi with fibrosis and necrosis…’ (Duignan, 2000 p. 452). Twenty specimens were present in this host animal (1).

• Order: Strongylida Molin, 1861

• Family: Ancylostomatidae Looss, 1905

Parasite name: Uncinaria hamiltoni Baylis, 1933

• Synonyms.

• Hosts. Neophoca cinerea.

• Locality. Kangaroo Island, SA.

• Infection site.

• Stage.

• References. Beveridge (1980).

• Remarks. Uncinaria hamiltoni sensu Beveridge (1980) should be treated as a synonym of U. sanguinis (Professor Ian Beveridge, pers. comm.).

Parasite name: Uncinaria sanguinis Marcus, Higgins, Slapeta & Gray, 2014

• Synonyms.

• Hosts. Neophoca cinerea (1, 2).

• Locality. (1, 2) Dangerous Reef, Spencer Gulf, SA; (1) South Page Island, Backstairs Passage, SA; (1, 2) Seal Bay, Kangaroo Island, SA.

• Infection site. Intestine.

• Stage. Adults and immatures.

• References. (1) Haynes et al. (2014); (2) Marcus et al. (2014).

• Remarks. Molecular data available (cox1) (1). Haynes et al. (2014) found that all pups examined were infected and that the infection route was trans-mammary. They also found that hookworms were genetically highly variable, but female host natal site fidelity and the transmammary route of infection do not restrict hookworm gene flow between N. cinerea populations.

Parasite name: Uncinaria sp.

• Synonyms.

• Hosts. Phocarctos hookeri (1, 2, 4, 5), Arctophoca australis forsteri (3, 7), Neophoca cinerea (6, 7).

• Locality. (1, 2) NZ; (3) Ohau Point, Kaikoura, NZ; (4, 5) Sandy Bay, Enderby Island, Auckland Island, NZ; (6) Adelaide Zoo, SA; (7) Kangaroo Island, Dangerous Reef, Spencer Gulf, SA.

• Infection site. Stomach, intestine.

• Stage. Adults and immatures.

• References. (1) Duignan (1998); (2) Duignan (2000); (3) Boren (2005); Castinel et al. ((4) 2006, (5) 2007); (6) I. Beveridge in Ladds 2009); (7) Ramos et al. (2013).

• Remarks. Molecular data available (ITS1&2) (7). Hookworms are the cause of haemorrhagic enteritis and are the main cause of death to pups in some seasons and localities. Sources report up to 7000 worms in a single pup. Route of infection is trans-mammary and worms develop into adults within three weeks of infection (4). Castinel et al. (2006) found a high level of size variability within adult worm populations in a single host. We suspect that most or all of the specimens reported in these papers could be referred to U. sanguinis or to the unnamed species of Nadler et al. (2013) in the following.

Parasite name: Uncinaria sp. NZSL of Nadler et al., 2013

• Synonyms.

• Hosts. Phocarctos hookeri.

• Locality. Sandy Bay Beach, Enderby Island, NZ.

• Infection site.

• Stage.

• References. Nadler et al. (2013).

• Remarks. Molecular data available (ITS1&2, 28S, 12S). The phylogeny of Nadler et al. (2013) included Uncinaria spp. from pinniped hosts worldwide and recovered seven independent evolutionary lineages or species, including the two described species (U. hamiltoni and U. lucasi) and five undescribed species, each from a different host. Uncinaria sp. NZSL was restricted to the NZ sea lion. Note, however, that Ramos et al. (2013) concluded that the three species found in Australian hosts A. p. doriferus, A. forsteri and N. cinerea were not distinguishable.

Parasite name: Uncinaria sp. AFS of Nadler et al., 2013

• Synonyms.

• Hosts. Arctocephalus pusillus doriferus (1, 2).

• Locality. (1, 2) Lady Julia Percy Island and Seal Rocks, Phillip Island, VIC; (2) Kanowna Island, VIC.

• Infection site.

• Stage.

• References. (1) Nadler et al. (2013); (2) Ramos et al. (2013).

• Remarks. Molecular data available (ITS1&2, 28S, 12S) (1, 2). Although not explicitly stated in their paper, we understand that the specimens from A. p. doriferus of Ramos et al. (2013) are the same as those of Nadler et al. (2013) from the same host.

Parasite name: Uncinaria sp. SES of Nadler et al., 2013

• Synonyms.

• Hosts. Mirounga leonine (1, 2).

• Locality. (1) Macquarie Island, TAS; (2) Possession Island, QLD and Lady Julia Percy Island, VIC.

• Infection site.

• Stage.

• References. (1) Nadler et al. (2013); (2) Ramos et al. (2013).

• Remarks. Molecular data available (ITS1&2, 28S, 12S) (1, 2). Although not explicitly stated in their paper, we understand that the specimens from M. leonina of Ramos et al. (2013) are equivalent to those of Nadler et al. (2013) from the same host.

• Family: Filaroididae Schultz, 1951

Parasite name: Parafilaroides decorus Dougherty & Herman, 1947

• Synonyms.

• Hosts. Phocarctos hookeri.

• Locality. NZ.

• Infection site. Lungs.

• Stage.

• References. Duignan (2000).

• Remarks. Duignan reported mild sub-pleural lesions caused by P. decorus.

Parasite name: Parafilaroides hydrurgae Mawson, 1953

• Synonyms.

• Hosts. Hydrurga leptonyx.

• Locality. Heard Island, Australia.

• Infection site. Lungs and bronchi.

• Stage.

• References. Mawson (1953).

• Remarks.

Parasite name: Parafilaroides normani Dailey, 2009

• Synonyms.

• Hosts. Arctocephalus pusillus doriferus (1, 2), Arctophoca australis forsteri (1).

• Locality. (1) Dog Beach, Ocean Grove, VIC; (1) Foxton Beach, Palmerston North, NZ; (2) Phillip Island, VIC.

• Infection site. Lungs.

• Stage. Adult.

• References. (1) Dailey (2009); (2) Jabbar et al. (2014).

• Remarks. Molecular data available (mitochondrial genome) (2).

Parasite name: Parafilaroides sp.

• Synonyms.

• Hosts. Arctocephalus pusillus doriferus (3), Arctophoca australis forsteri (2, 3), Delphinus delphis (4), Hydrurga leptonyx (3), Neophoca cinerea (1).

• Locality. (1) Dangerous Reef, Spencer Gulf and Seal Bay, Kangaroo Island, SA; (2) Ohau Point, Kaikoura, NZ; (3) Cape Woollamai and Ocean Grove, VIC; (3) Taranaki Bight, NZ; (3) Geelong, VIC; (4) coast of NZ.

• Infection site. Lungs.

• Stage. Adults and larvae.

• References. (1) Nicholson & Fanning (1981); (2) Boren (2005); (3) Norman in McFarlane et al. (2009); (4) Stockin et al. (2009).

• Remarks. Nicholson & Fanning (1981) stated that their specimens were probably a new species of Parafilaroides, but did not describe or name them. These authors also reported that, although the animals appeared healthy when captured, they showed signs of acute verminous pneumonia. The report of Stockin et al. (2009) seems unlikely in a dolphin host, and may represent an accidental infection or possible misidentification.

• Family: Pseudaliidae Railliet & Henry, 1909

Parasite name: Halocercus lagenorhynchi Baylis & Daubney, 1925

• Synonyms.

• Hosts. Delphinus delphis (2), Tursiops aduncus (2), Tursiops truncatus (1).

• Locality. (1) Encounter Bay, SA; (2) coast of SA.

• Infection site. Lungs.

• Stage.

• References. (1) Johnston & Mawson (1941); (2) Tomo et al. (2010).

• Remarks. Tomo et al. (2010) report ‘necrotic microabscesses, in serious acute to chronic infections, lung function would have been compromised’ in lungworm infections of dolphins, ‘moderate and heavy burdens may have contributed to animals’ deaths.’ Predominantly calves and juveniles were infected. During the 18-year study, the prevalence of lung nematode infections increased in 2005–2006 fourfold from 14% before the outbreak to 63% in short-beaked common dolphins; some cases were also recorded in Indo-Pacific bottlenose dolphins and common bottlenose dolphins.

Parasite name: Halocercus delphini Baylis & Daubney, 1925

• Synonyms.

• Hosts. Stenella coeruleoalba.

• Locality. Carter's Beach, Te Akau, NZ.

• Infection site. Lungs.

• Stage.

• References. Lehnert et al. (2017).

• Remarks.

Parasite name: Halocercus sp.

• Synonyms.

• Hosts. Cephalorhynchus hectori (1, 2), Delphinus delphis (3), Stenella coeruleoalba (3).

• Locality. (1) Canterbury coast, NZ; (2) NZ; (3) Kawaa Kawaa Bay, NZ; (3) Muriwai Beach, NZ.

• Infection site. Lung.

• Stage. Adult.

• References. (1) McKenzie & Blair (1983); (2) Duignan (2000); (3) Lehnert et al. (2017).

• Remarks.

Parasite name: Pharurus alatus (Leuckart, 1848) Stiles & Hassall, 1905

• Synonyms. Prosthecosacter alatus (Leuckart, 1848) Diesing, 1851; Pseudalius alatus (Leuckart, 1848) Diesing, 1851; Stenurus alatus (Leuckart, 1848) Yorke & Maplestone, 1926; Strongylus alatus Leuckart, 1848; Str. (Pharurus) alatus (Leuckart, 1848) Diesing, 1851; Torynurus alatus (Leuckart, 1848) Delyamure, 1952 [WoRMS; Baylis & Daubney, 1925].

• Hosts. Delphinus delphis.

• Locality. Coast of SA.

• Infection site. Lungs.

• Stage.

• References. Tomo et al. (2010).

• Remarks.

Parasite name: Skrjabinalius cryptocephalus Delyamure, 1942

• Synonyms.

• Hosts. Tursiops truncatus.

• Locality. Otago Harbour, NZ.

• Infection site. Lungs.

• Stage.

• References. Bowie (1984).

• Remarks. The anterior end of this nematode was tightly knotted, embedded in the parenchyma of the lung and surrounded by purulent fluid in a fibrous or calcified capsule (Bowie, 1984).

Parasite name: Stenurus globicephalae Baylis & Daubney, 1925

• Synonyms.

• Hosts. Globicephala melas (3, 5), Globicephala sp. (1), Grampus griseus (4), Peponocephala electra (2).

• Locality. (1) Prosperine, QLD; (2) Moreton Island and Tugun Beach, QLD; (3) McIntyres Beach, Falmouth, TAS; (4) NSW; (5) Big River, Kahurangi National Park, NZ; (5) Kaka Point, NZ.

• Infection site. Guttural pouch, ear canals, auditory sinuses.

• Stage. Adult.

• References. (1) Arnold in Cannon (1977); (2) Cannon (1977); (3) McManus et al. (1984); (4) J. Boulton in Ladds (2009); (5) Lehnert et al. (2017).

• Remarks. Molecular data available (ITS2) (5). McManus et al. (1984) reported ‘huge masses of worms: average 2300, range 1140–4200 per ear’ in the lead bulls, and their data led them to conclude that this auditory parasitism played a significant role in the mass stranding of 183 pilot whales on this occasion. However, a study on by-caught porpoises with good nutritional status has revealed high loads of Stenurus minor, with no apparent effect on echo-location or hunting ability, and no pathological changes (Faulkner et al., 1998). Another recent study on S. minor in the inner ear of harbour porpoises highlights the need of further research to assess the impact these nematodes may have on hearing (Morell et al., 2017). Host named G. malaenas in McManus et al. (1984).

Parasite name: Stenurus minor (Kuhn, 1829) Baylis & Daubney, 1925

• Synonyms. Pharurus minor (Kuhn, 1929) Cobbold, 1879; Prosthecosacter minor (Kuhn, 1829) Diesing, 1851; Pseudalius minor (Kuhn, 1829) Schneider, 1866; Stenurus inflexus Dujardin, 1845; Stenurus phocoenae Dougherty, 1943; Stenurus vagans Dougherty, 1943; Strongylus minor Kuhn, 1829 [WoRMS; Baylis & Daubney, 1925].

• Hosts. Phocoena dioptrica.

• Locality. Pipikaretu, Otago Peninsula, NZ.

• Infection site. Sinuses and tympanic cavity.

• Stage. Adult.

• References. Lehnert et al. (2017).

• Remarks. Molecular data available (ITS2).

Parasite name: Stenurus ovatus (von Linstow, 1910) Baylis & Daubney, 1925

• Synonyms. Pseudalius ovatus von Linstow, 1910 [WoRMS].

• Hosts. Delphinus delphis (3), Lagenodelphis hosei (1), Tursiops aduncus (3), T. truncatus (2, 3).

• Locality. (1) Corio Bay, VIC; (2) Otago Harbour, NZ; (3) coast of SA.

• Infection site. Lungs.

• Stage. Adults and larvae.

• References. (1) McColl & Obendorf (1982); (2) Bowie (1984); (3) Tomo et al. (2010).

• Remarks. McColl & Obendorf (1982) report the lungs of their host specimen were highly congested with worms, causing verminous pneumonia and greatly impairing respiratory function.

Parasite name: Stenurus sp.

• Synonyms.

• Hosts. Cephalorhynchus hectori.

• Locality. NZ.

• Infection site. Lungs.

• Stage.

• References. Duignan (2000).

• Remarks.

• Phylum: Platyhelminthes

• Class: Cestoda

• Order: Diphyllobothriidea Kuchta, Scholz, Brabec & Bray, 2008

• Family: Diphyllobothriidae Lühe, 1910

Parasite name: Adenocephalus pacificus Nybelin, 1931

• Synonyms. Adenocephalus septentrionalis Nybelin, 1931; Bothriocephalus sp. of Stiles & Hassall (1899); B. macrophallus von Linstow, 1905, in part; Dibothriocephalus atlanticum Delyamure & Parukhin, 1968; Diphyllobothrium glaciale (Cholodkovsky, 1915) Markowski, 1952; Dip. krotovi Delyamure, 1955; Dip. pacificum (Nybelin, 1931) Margolis, 1956; Dip. septentrionalis Nybelin, 1931 [WoRMS; Hernández-Orts et al., 2015].

• Hosts. Arctocephalus pusillus doriferus (1), Neophoca cinerea (2).

• Locality. (1) Lady Julia Percy Island, VIC; (2) Pearson Island, SA.

• Infection site. Intestine.

• Stage. Adult plus immatures.

• References. (1) Drummond (1937); (2) Johnston (1937).

• Remarks. Described in Drummond (1937) as Dip. arctocephali n. sp., and host called A. tasmanicus. Described in Johnston (1937) as Dip. arctocephalinum n. sp. and host called A. forsteri, but Johnston corrected this to N. cinerea in Johnston & Mawson (1941) and Johnston & Best (1942). Johnston (1937) described the worms as a ‘tangled mass’ in the intestine.

Parasite name: Diphyllobothrium sp.

• Synonyms.

• Hosts. Arctophoca australis forsteri, Hydrurga leptonyx.

• Locality. In captivity, NZ.

• Infection site. Intestine.

• Stage. Adult.

• References. Cordes & O'Hara (1979).

• Remarks. Cordes & O'Hara (1979) described these worms as occurring in huge masses, their scoleces embedded in a cyst in the colon wall and their strobilae free in the lumen. In the leopard seal, they described a 2–3 lb mass of worms blocking the intestine.

• Order: Phyllobothriidea Caira, Jensen, Waeschenbach, Olson & Littlewood, 2014

• Family: Phyllobothriidae Braun, 1900

Parasite name: Monorygma chamissonii (Linton, 1905) Meggitt, 1924

• Synonyms. Cysticercus delphini Rudolphi, 1819; Phyllobothrium chamissonii (Linton, 1905) Southwell & Walker, 1936; Taenia chamissonii Linton, 1905.

• Hosts. Peponocephala electra.

• Locality. Moreton Island and Tugun Beach, QLD.

• Infection site. Stomach wall and peritoneum.

• Stage. Encysted larvae.

• References. Cannon (1977).

• Remarks. Called Phyllobothrium chamissonii in this paper. Found in cysts 20–25 mm in diameter with thick fibrous walls. The inner blastocysts (metacestode) were almond-shaped, with fat droplets in the wall giving them an orange colour.

Parasite name: Monorygma grimaldii (Moniez, 1899) Meggitt, 1924

• Synonyms. Cephalocotyleum delphini delphis Diesing, 1850; Cysticercus delphini Rudolphi, 1819; Cysticercus grimaldii Braun, 1898; Cysticercus Taenia grimaldii Moniez, 1889; Cysticerque du groupe grimaldii Baer, 1932; Dubium delphini Rudolphi, 1810; Phyllobothrium sp. Beneden, 1868; P. delphini Beneden, 1870; Taenia grimaldii Moniez, 1889; T. schamissoni Linton, 1905 [Baylis 1932; Delyamure, 1955].

• Hosts. Delphinus delphis (2, 3), Lagenodelphis hosei (1), Stenella coeruleoalba (4).

• Locality. (1) Corio Bay, VIC; (2) Barwon Heads, VIC; (3) coast of NZ; (4) Carter's Beach, Te Akau, NZ; (4) Kawera Parade, Paramon East, NZ; (4) Miruwai Beach, NZ.

• Infection site. (1) Abdominal muscle; (2, 3) peritoneal cavity; (4) urogenital area; (4) mammary gland; (4) blubber.

• Stage. Merocercoid larva.

• References. (1) McColl & Obendorf (1982); (2) Norman (1997); (3) Stockin et al. (2009); (4) Lehnert et al. (2017).

• Remarks. Norman (1997) reported localized lymphoplasmacytic response to the cysts. These metacestodes are merocercoids of marine tapeworms infecting various marine mammals (Agustí et al., 2005; Aznar et al., 2007); historically, they have been referred to as ‘Phyllobothrium delphini’ (Bosc, 1802) and ‘Monorygma grimaldii’ (Moniez, 1889), but molecular evidence suggests that they belong to the ‘Clistobothrium’ clade (Aznar et al., 2007; Randhawa, 2011; Randhawa & Brickle, 2011). Recently, they were found in subcutaneous blubber from fur seals and shown to be intermediate stages of the cestode Clistobothrium, infecting sharks (Klotz et al., 2018). The merocercoids represented the delphini-morphotype but were identified as grimaldii-genotype using 18S and 28S sequences.

Parasite name: Phyllobothrium delphini (Bosc, 1802) Gervais, 1885

• Synonyms. Cephalocotyleum delphini delphis Diesing, 1850; Cysticercus delphini Laennec, 1804; Cy. delphini Rudolphi, 1810; Cy. mysticeti Diesing, 1850; Cy. physeteris Diesing, 1863; Cysticercus sp. Bennett, 1937; Hydatis delphini Bosc, 1802; H. delphinii Bosc, 1802; Phyllbothrium sp. Beneden, 1868; Phyllobothrium sp. Carnot, 1822; Phyllobothrium sp. Rennie and Reid, 1912; P. inchoatum Leidy, 1891; P. tumidum Linton, 1922; Scolex delphini Stossich, 1897; Vermis delphini delphis Rudolphi, 1810 [Baylis, 1932; Delyamure, 1955; Williams, 1968].

• Hosts. Cephalorhynchus hectori (3), Delphinus delphis (4, 5, 6), Kogia breviceps (1), Lagenorhynchus obscurus (6), Mirounga leonina (2), Phocoena dioptrica (6), Stenella coeruleoalba (6).

• Locality. (1) Port Victoria, Spencer Gulf, SA; (2) Macquarie Island, TAS; (3) Kaikoura coast, NZ; (4) Barwon Heads, VIC; (5) coast of NZ; (6) Sandspit Estuary, Green's Point, NZ; (6) Portobello Bay, Otago Harbour and off Kaikoura, NZ; (6) Caroline Bay, Timaru, NZ; (6) Carter's Beach, Te Akau, NZ.

• Infection site. Blubber, especially around tail and genital area.

• Stage. Merocercoid larva.

• References. (1) Johnston & Mawson (1939); (2) Morgan et al. (1978); (3) McKenzie & Blair (1983); (4) Norman (1997); (5) Stockin et al. (2009); (6) Lehnert et al. (2017).

• Remarks. Morgan et al. (1978) found this cestode encysted in each of three elephant seals examined. As evidenced by the long list of synonyms, this cestode has a complicated history. Reviews of the literature can be found in Delyamure (1955), Dollfus (1964), Johnston & Mawson (1939) and Williams (1968). Molecular evidence suggests that this species belongs to the ‘Clistobothrium’ clade (Aznar et al., 2007; Randhawa, 2011; Randhawa & Brickle, 2011). See also Monorygma grimaldii.

Parasite name: Phyllobothrium sp.

• Synonyms.

• Hosts. Arctophoca australis forsteri (3), Balaenoptera acutorostrata (2), Cephalorhynchus hectori (3), Delphinus delphis (3), Lagenorhynchus obscurus (3), Phocartctos hookeri (3), Tursiops truncatus (1).

• Locality. (1) Otago Harbour, NZ; (2) Pigeon Bay, Banks Peninsula, NZ; (3) NZ.

• Infection site. Ventral blubber.

• Stage. Merocercoid larva.

• References. (1) Bowie (1984); (2) Dawson & Slooten (1990); (3) Duignan (2000).

• Remarks. Duignan (2000) found that prevalence was higher in dusky dolphins than in common or Hector's dolphins. In pinnipeds they found a higher prevalence in NZ fur seals than in NZ sea lions. They also reported granulomatous mastitis associated with the cysts that resulted in underfed pups. Dawson and Slooten (1990) reported several hundred cysts in the blubber of a dwarf minke whale.

Parasite name: Phyllobothriidae gen. sp.

• Synonyms.

• Hosts. Arctophoca australis forsteri.

• Locality. In captivity, NZ.

• Infection site. Ventral blubber.

• Stage. Merocercoid larva.

• References. Cordes & O'Hara (1979).

• Remarks.

• Order: Tetrabothriidea Baer, 1954

• Family: Tetrabothriidae Linton, 1891

Parasite name: Tetrabothrius forsteri (Krefft, 1871) Fuhrmann, 1904

• Synonyms. Prosthecotyle forsteri (Krefft, 1871) Monticelli, 1892; Taenia forsteri Krefft, 1871; Tetrabothrius dalli Yamaguti, 1952 [Linton, 1923; WoRMS].

• Hosts. Delphinus delphis (1), Mesoplodon hectori (2).

• Locality. (1) Port Jackson, NZ; (2) Oneroa Bay, Waiheke Island, NZ.

• Infection site. Stomach and intestine.

• Stage.

• References. (1) Krefft (1871); (2) Baker et al. (2001).

• Remarks. Described in Krefft (1871) as Taenia forsteri n. sp., and host recorded as D. forsteri.

Parasite name: Tetrabothrius sp.

• Synonyms.

• Hosts. Delphinus delphis (2), Lagenodelphis hosei (1).

• Locality. (1) Corio Bay, VIC; (2) unknown locality, Australia.

• Infection site. Stomach and intestine.

• Stage.

• References. (1) McColl & Obendorf (1982); (2) Ladds (2009).

• Remarks. McColl & Obendorf (1982) reported minor haemorrhaging in pancreatic ducts caused by this cestode.

• Class: Trematoda Rudolphi, 1808

• Order: Diplostomida Olson, Cribb, Tkach, Bray & Littlewood, 2003

• Family: Brauninidae Wolf, 1903

Parasite name: Braunina cordiformis Wolf, 1903

• Synonyms.

• Hosts. Cephalorhynchus hectori (1, 2).

• Locality. (1) Canterbury coast, NZ; (2) unknown locality, NZ.

• Infection site. Stomach.

• Stage.

• References. (1) McKenzie & Blair (1983); (2) Duignan (2000).

• Remarks. McKenzie & Blair (1983) found a maximum of 57 worms in one of three dolphins, attached to the stomach wall.

• Family: Cyathocotylidae Mühling, 1898

Parasite name: Mesostephanus neophocae Dubois & Angel, 1976

• Synonyms.

• Hosts. Mirounga leonina, Neophoca cinerea.

• Locality. In captivity and St. Vincent Gulf, SA.

• Infection site. Intestine.

• Stage. Adult.

• References. Dubois & Angel (1976).

• Remarks. Dubois & Angel (1976) put forward the suggestion that species of this genus are more usually parasites of phalacrocoracid birds, and that perhaps the sea lion had taken on the parasite after predating a bird.

• Family: Spirorchiidae Stunkard, 1921

Parasite name: Spirorchiidae gen. sp.

• Synonyms.

• Hosts. Dugong dugon.

• Locality. Mornington Island and Townsville, QLD.

• Infection site. Ovaries, myometrial blood vessels.

• Stage. Eggs.

• References. Marsh et al. (1984).

• Remarks. Found in 18 of 46 dugongs. Lesions were found on ovaries, containing refractile eggs 25–30 µm in diameter. Pigmented eyespots of miracidia were visible. Marsh et al. (1984) postulate that these could be the eggs of a spirorchid trematode, members of which parasitize the circulatory system of aquatic reptiles in the same habitat as dugongs.

• Order: Echinostomida La Rue, 1957

• Family: Cladorchiidae Fischoeder, 1901

Parasite name: Solenorchis travassosi Hilmy, 1949

• Synonyms. Indosolenorchis hirudinaceus Crusz, 1951; Solenorchis baeri Hilmy, 1949; S. gohari Hilmy, 1949; S. naguibmahfouzi Hilmy, 1949 [WoRMS].

• Hosts. Dugong dugon (1, 2).

• Locality. (1) Mornington Island and Townsville, QLD; (2) Lucinda, QLD.

• Infection site. Caecum and large intestine.

• Stage. Adult.

• References. (1) Blair (1980); (2) Olson et al. (2003).

• Remarks. Molecular data available (18S, 28S) (2). In his redescription of I. hirudinaceus, Blair (1980) is adamant that it is a separate species and genus. He states that Hilmy's (1949) specimens should be S. travassosi but still retains his specimens as distinct. He mentions that their size range is large and that the Australian specimens do not overlap the Indian ones in size. Sey (1980) synonymized I. hirudinaceus with S. travassossi, and now only one species, S. travassossi, is accepted from dugong.

• Family: Labicolidae Blair, 1979

Parasite name: Labicola cf. elongata

• Synonyms.

• Hosts. Dugong dugon.

• Locality. Lucinda, QLD.

• Infection site.

• Stage.

• References. Olson et al. (2003).

• Remarks. Molecular data available (18S, 28S).

Parasite name: Labicola elongata Blair, 1979

• Synonyms.

• Hosts. Dugong dugon (1, 2).

• Locality. (1) Mornington Island and Townsville, QLD; (2) unknown locality, Australia.

• Infection site. Upper lip.

• Stage. Adult.

• References. (1) Blair (1979); (2) Blanshard (2001).

• Remarks. Blair (1979) found these trematodes in all 21 specimens he observed. They were so unusual that he erected the Family Labicolidae to contain the species. Several worms together formed pus-filled abscesses along the sides of the upper lip, with pores to the outside.

• Family: Notocotylidae Lühe, 1909

Parasite name: Ogmogaster sp.

• Synonyms.

• Hosts. Caperea marginata.

• Locality. Fotheringate Bay, Flinders Island, TAS.

• Infection site. Intestine.

• Stage.

• References. McManus et al. (1984).

• Remarks.

• Family: Opisthotrematidae Poche, 1926

Parasite name: Folitrema jecoris Blair, 1981

• Synonyms.

• Hosts. Dugong dugon.

• Locality. Mornington Island, Thursday Island and Townsville, QLD.

• Infection site. Gall bladder and bile ducts.

• Stage. Adult.

• References. Blair (1981).

• Remarks.

Parasite name: Lankatrema macrocotyle Blair, 1981

• Synonyms.

• Hosts. Dugong dugon.

• Locality. Mornington Island, Thursday Island and Townsville, QLD.

• Infection site. Intestine.

• Stage. Adult.

• References. Blair (1981).

• Remarks. Lankatrema spp. are found encapsulated in the wall of the intestine. Different species may be found in the same capsule (Blair, 1981).

Parasite name: Lankatrema mannarense Crusz & Fernand, 1954

• Synonyms.

• Hosts. Dugong dugon (1, 2).

• Locality. (1) Mornington Island, QLD; (2) Townsville, QLD.

• Infection site. Stomach and intestine.

• Stage. Adult.

• References. (1) Blair (1981); (2) Olson et al. (2003).

• Remarks. Molecular data available (18S, 28S) (2). Blair (1981) admits they are poor specimens in his paper, and that the record is questionable.

Parasite name: Lankatrema microcotyle Blair, 1981

• Synonyms.

• Hosts. Dugong dugon.

• Locality. Mornington Island, Thursday Island and Townsville, QLD.

• Infection site. Intestine.

• Stage. Adult.

• References. Blair (1981).

• Remarks. Found encapsulated in wall of ileum, 2–7 in each capsule.

Parasite name: Lankatrema minutum Blair, 1981

• Synonyms.

• Hosts. Dugong dugon.

• Locality. Mornington Island and Townsville, QLD.

• Infection site. Stomach.

• Stage. Adult.

• References. Blair (1981).

• Remarks. Occurs in pairs in the wall of the stomach glands with their posterior ends towards the lumen of the gland. Their capsules are similar to those reported by Crusz & Fernand (1954) for L. mannarense. There appears to be no host response to their presence (Blair, 1981).

Parasite name: Lankatrematoides gardneri Blair, 1981

• Synonyms.

• Hosts. Dugong dugon.

• Locality. Mornington Island, Thursday Island and Townsville, QLD.

• Infection site. Pancreatic ducts.

• Stage. Adult.

• References. Blair (1981).

• Remarks.

Parasite name: Opisthotrema australe Blair, 1981

• Synonyms.

• Hosts. Dugong dugon.

• Locality. Mornington Island, Thursday Island and Townsville, QLD.

• Infection site. Eustachian tubes and middle ear.

• Stage. Adult.

• References. Blair (1981).

• Remarks.

Parasite name: Opisthotrema dujonis (Leuckart, 1874) Price, 1932

• Synonyms. Monostomum dujonis Leuckart, 1874; Opisthotrema cochleare Fischer, 1883; O. cochleariforme Travassos and Vogelsang, 1931 (lapsus) [WoRMS; Blair, 1981].

• Hosts. Dugong dugon (1–4).

• Locality. (1) Wallum Creek, North Stradbroke Island, QLD; (2) coast of QLD; (3) Mornington Island, QLD; (3) Thursday Island, QLD; (3, 4) Townsville, QLD.

• Infection site. (1, 3) Eustachian tubes; (3) middle ear; (2, 3) oesophagus.

• Stage. Adult and immature.

• References. (1) Dexler & Freund (1906); (2) Johnston (1913); (3) Blair (1981); (4) Olson et al. (2003).

• Remarks. Molecular data available (18S, 28S) (4). Dexler and Freund (1906) and Johnston (1913) record this species as O. cochleare.

Parasite name: Pulmonicola pulmonalis (von Linstow, 1904) Poche, 1926

• Synonyms. Cochleotrema indicum (Sharma & Gupta, 1971); Opisthotrema nasalis (n. n.) Purnomo, Palmieri & Budiarso in Budiarso, Palmieri, Imes, Allen & Lepes, 1979; Opisthotrema pulmonale von Linstow, 1904; Paracochleotrema indicum Sharma & Gupta, 1971; Pulmonicola indicus (Sharma & Gupta, 1971); Pulmonicola pulmonale (von Linstow, 1904) Ruiz, 1946 [WoRMS; Blair, 1981].

• Hosts. Dugong dugon (1–4).

• Locality. (1) Torres Strait, Australia; (2) Mornington Island, Thursday Island, Townsville, QLD; (3) unknown location, Australia; (4) coast of QLD.

• Infection site. (2) Nasal passages and lungs; (4) bronchi and trachea.

• Stage. Adult.

• References. (1) von Linstow (1904); (2) Blair (1981); (3) Blanshard (2001); (4) Owen et al. (2012).

• Remarks. Described as Opisthotrema pulmonale by von Linstow (1904). Called Cochleotrema indicum in Blanshard (2001) and Owen et al. (2012). Cause of chronic bronchitis, tracheitis and bronchopneumonia (3, 4).

• Family: Rhabdiopoeidae Poche, 1926

Parasite name: Faredifex clavata Blair, 1981

• Synonyms.

• Hosts. Dugong dugon.

• Locality. Mornington Island, Thursday Island and Townsville, QLD.

• Infection site. Small intestine.

• Stage. Adult.

• References. Blair (1981).

• Remarks. Found in abscesses in wall of ileum.

Parasite name: Rhabdiopoeus taylori Johnston, 1913

• Synonyms.

• Hosts. Dugong dugon (1–3).

• Locality. (1) Coast of QLD; (2) Mornington Island, Thursday Island and Townsville, QLD; (3) Lucinda, QLD.

• Infection site. Intestine.

• Stage. Adult.

• References. (1) Johnston (1913); (2) Blair (1981); (3) Olson et al. (2003).

• Remarks. Molecular data available (18S, 28S) (3). Johnston (1913) found R. taylori in both dugongs he examined, ten in one and 16 in the other.

Parasite name: Haerator caperatus Blair, 1981

• Synonyms.

• Hosts. Dugong dugon.

• Locality. Mornington Island, Thursday Island and Townsville, QLD.

• Infection site. Intestine and caecum.

• Stage. Adult.

• References. Blair (1981).

• Remarks.

Parasite name: Taprobanella bicaudata Crusz & Fernand, 1954

• Synonyms.

• Hosts. Dugong dugon (1, 2).

• Locality. (1) Mornington Island, QLD; (1) Thursday Island, QLD; (1, 2) Townsville, QLD.

• Infection site. Large intestine.

• Stage. Adult.

• References. (1) Blair (1981); (2) Olson et al. (2003).

• Remarks. Molecular data available (18S, 28S) (2).

• Order: Plagiorchiida La Rue, 1957

• Family: Brachycladiidae Odhner, 1905

Parasite name: Brachycladium delphini (Poirier, 1886) Looss, 1899

• Synonyms. Campula delphini (Poirrier, 1886) Bittner & Sprehn, 1928; Cladocoelium delphini (Poirier, 1886) Stossich, 1892; Distomum delphini Poirier, 1886; Lecithodesmus delphini (Poirier, 1886) Yamaguti, 1958 [WoRMS; Felix, 2013].

• Hosts. Delphinus delphis.

• Locality. Bethell's Beach, Auckland, NZ.

• Infection site. Liver.

• Stage. Adult.

• References. Lehnert et al. (2017).

• Remarks.

Parasite name: Brachycladium palliatum (Looss, 1885) Looss, 1899

• Synonyms. Campula palliata (Looss, 1885) Looss, 1899; Cladocoelium palliatum (Looss, 1885) Stossich, 1892; Distomum palliatum Looss, 1885; Lecithodesmus palliatus (Looss, 1885) Yamaguti, 1958 [WoRMS; Felix, 2013].

• Hosts. Delphinus delphis (1, 2).

• Locality. (1) In captivity NZ; (2) Battey's Beach, Warkworth, NZ; (2) Shakespeare Peak and Waiake Beach, Auckland, NZ.

• Infection site. Liver and bile ducts.

• Stage. Adult.

• References. (1) Cordes & O'Hara (1979); (2) Lehnert et al. (2017).

• Remarks. Cordes and O'Hara (1979) found over 100 of this species in one dolphin. The worms were associated with parasitic hepatitis in four dolphins with lesions, but no worms were found in the livers of two more. Worms and lesions blocked bile ducts, and were either contributory to, or the cause of, death.

Parasite name: Campula sp.

• Synonyms.

• Hosts. Cephalorhynchus hectori (2, 3), Lagenodelphis hosei (1).

• Locality. (1) Corio Bay VIC; (2) New Brighton Beach, Christchurch and Gillespies Point, NZ; (3) NZ.

• Infection site. (1) Pancreatic ducts; (2, 3) mesenteric lymph nodes.

• Stage. Adult and eggs.

• References. (1) McColl & Obendorf (1982); (2) Hutton et al. (1987); (3) Duignan (2000).

• Remarks. Both Duignan (2000) and Hutton et al. (1987) found these worms encapsulated within the mesenteric lymph nodes of the host. The lymph node had granulomatous lesions with inflammatory response associated with eggs/was enlarged with a tumour-like mass which contained purulent necrotic debris and a single parasitic worm. A secondary bacterial infection, Eikenella corrodens, was cultured from the lesion (2).

Parasite name: Nasitrema sp.

• Synonyms.

• Hosts. Lagenodelphis hosei.

• Locality. Corio Bay, VIC.

• Infection site. Blowhole.

• Stage. Eggs.

• References. McColl & Obendorf (1982).

• Remarks.

Parasite name: Synthesium tursionis (Marchi, 1873) Stunkard & Alvey, 1930

• Synonyms. Distomum longissimum Poirier, 1886 nec D. longissimum von Linstow, 1896; D. tursionis Marchi, 1873; Hadwenius tursionis (Marchi, 1873) Fernández, Balbuena & Raga, 1994; Orthosplanchnus tursionis (Marchi, 1873) Odhner, 1926; Synthesium tursionis (Marchi, 1873) [Price, 1932; Marigo et al., 2008].

• Hosts. Tursiops truncatus.

• Locality. Otago Harbour, NZ.

• Infection site. Small intestine.

• Stage. Adult.

• References. Bowie (1984).

• Remarks.

Parasite name: Synthesium sp.

• Synonyms.

• Hosts. Neophoca cinerea.

• Locality. St. Vincent Gulf, SA.

• Infection site. Intestine.

• Stage.

• References. Dubois & Angel (1976).

• Remarks. Genus Hadwenius in this paper. This genus was synonymized with Synthesium by Gibson (2005).

• Family: Heterophyidae Leiper, 1909

Parasite name: Galactosomum angelae Pearson, 1973

• Synonyms.

• Hosts. Neophoca cinerea.

• Locality. St. Vincent Gulf, SA.

• Infection site. Intestine.

• Stage. Adult.

• References. Dubois & Angel (1976).

• Remarks. This species was originally described by Pearson (1973) from the Caspian tern – as mentioned in the passing reference to G. angelae in Dubois and Angel (1976). There is a perpetuated error in the citation of this species and the referencing of this paper. Note that both Dailey et al. (2002) and Hernández-Orts et al. (2012) list Dubois and Angel (1976) with the title ‘Galactosomum angelae Pearson 1973 in Neophoca cinerea (Péron, 1816) the Australian sea-lion.’ There does not appear to be any paper with this title. The 1976 paper is actually entitled ‘Mesostephanus neophocae n. sp. (Strigeata: Prohemistomidae), parasite d'une otarie d'Australie, N. cinerea (Péron et Lesueur)’. In addition, the volume and page reference given in both of these works – that is, ‘82: 191–229’ – apply to a paper by Dubois and Mahon published in the Bulletin de la Société Neuchâteloise des Sciences Naturelles in 1959. The corrected reference for the Dubois and Angel (1976) paper is ‘99: 29–32’.

Parasite name: Heterophyidae ‘Gen.’ sp.

• Synonyms.

• Hosts. Delphinus delphis.

• Locality. In captivity, NZ.

• Infection site. Adipose tissue of omasum.

• Stage.

• References. Cordes & O'Hara (1979).

• Remarks.

Parasite name: Stictodora diplacantha Johnston, 1942

• Synonyms.

• Hosts. Neophoca cinerea.

• Locality. St. Vincent Gulf, SA.

• Infection site. Intestine.

• Stage.

• References. Dubois & Angel (1976).

• Remarks. This species was originally described from the pied cormorant.

In addition to the above records, ‘gastrointestinal parasites’ were reported in Arctophoca australis forsteri from Ohau Point, NZ (Boren, 2005); ‘lung worms’ in Delphinus delphis from Mooloolaba Beach, QLD (Greenland & Limpus, 2008); ‘cestodes’ in Tursiops truncatus from Otago Harbour, NZ (Bowie, 1984) and Globicephala melas from McIntyres Beach, TAS (McManus et al., 1984); and ‘trematodes’ in A. a. forsteri from Ohau Point, NZ (Boren, 2005), Lagenodelphis hosei from Corio Bay, VIC (McColl & Obendorf, 1982) and Dugong dugon from Wallum Creek, QLD (Dexler & Freund, 1906).

• Phylum: Arthropoda

• Class: Arachnida Lamarck, 1801

• Subclass: Acari Leach, 1817

• Order: Parasitiformes

• Family: Halarachnidae Oudemans, 1906

Parasite name: Halarachne miroungae Ferris, 1925

• Synonyms. Halarachne erratica Fain & Mortelmans, 1959; H. taita Eichler, 1958 [Domrow, 1962].

• Hosts. Mirounga leonine.

• Locality. Macquarie Island, TAS.

• Infection site. Respiratory tract.

• Stage. Adult, larvae.

• References. Domrow (1979).

• Remarks.

Parasite name: Orthohalarachne attenuata (Banks, 1910) Newell, 1947

• Synonyms. Halarachne attenuata Banks, 1910; H. magellanica Finnegan, 1934; H. reflexa Tubb, 1937; H. rosmari Oudemans, 1916; H. zalophi Oudemans, 1916; Orthohalarachne reflexa (Tubb, 1937) Strandtmann and Wharton, 1958 [Newell, 1947; Domrow, 1974].

• Hosts. Arctocephalus pusillus doriferus (1–4), Neophoca cinerea (4–6).

• Locality. (1) Lady Julia Percy Island, VIC; (2) Portarlington, VIC; (3) Newcastle, NSW; (4–6) Dangerous Reef, SA; (4) Seal Rocks and Geelong, VIC; (6) Kangaroo Island, SA.

• Infection site. Nasopharynx, anterior and ethmoid nasal turbinates, distal third of the trachea and in the proximal 10 cm of both bronchi.

• Stage. Adults, larva.

• References. (1) Tubb (1937); (2) Domrow (1963); (3) Seawright (1964); (4) Domrow (1974); (5) Marlow (1975); (6) Nicholson & Fanning (1981).

• Remarks. Tubb (1937) reports heavy infestations, cephalothorax and legs embedded in the mucous membrane, abdomen protruding, the locus of infection was inflamed and swollen, impossible to dislodge the mites without causing extensive damage to the mucous membrane. The host in Tubb's (1937) report is named A. tasmanicus. Seawright (1964) cites the host name as Gypsophoca tasmanica.

Parasite name: Orthohalarachne diminuata (Doetschman, 1944) Newell, 1947

• Synonyms. Halarachne diminuata Doetschman, 1944; Orthohalarachne chabaudi Gretillat, 1960; O. letalis Popp, 1961 [Newell, 1947; Domrow, 1974].

• Hosts. Arctocephalus pusillus doriferus (1), Neophoca cinerea (2).

• Locality. (1) Seal Rocks and Geelong, VIC; (2) Dangerous Reef and Kangaroo Island, SA.

• Infection site. Nasopharynx, nasal turbinates, lungs.

• Stage. Adult, larvae.

• References. (1) Domrow (1974); (2) Nicholson & Fanning (1981).

• Remarks. Domrow (1974) reports mixed infections with O. attenuata.

• Order: Trombidiformes Reuter, 1909

• Family: Demodecidae Nicolet, 1855

Parasite name: Demodex zalophi Dailey & Nutting, 1980

• Synonyms.

• Hosts. Zalophus californianus.

• Locality. Australia.

• Infection site. Skin, hair follicles.

• Stage. Adult, larvae.

• References. Dailey & Nutting (1980).

• Remarks. In Californian sea lion in captivity in Australia. First record of a demodecid in marine mammals.

• Class: Insecta Linnaeus, 1758

• Order: Phthiraptera Haeckel, 1896

• Family: Echinophthiriidae Enderlein, 1904

Parasite name: Antarctophthirus microchir (Trouessart & Neumann, 1888) Enderlein, 1906

• Synonyms. Echinophthirius microchir Trouessart & Neumann, 1888 [Leonardi, 2009].

• Hosts. Phocarctos hookeri (1–3), Neophoca cinerea (4, 6), Arctophoca australis forsteri (5).

• Locality. (1) Auckland Island, NZ; (2) Campbell Island, NZ; (3) Snares Island, NZ; (4, 6) Kangaroo Island and Dangerous Reef, SA; (4) Pages Island, SA.

• Infection site. Fur/skin, body.

• Stage. Adults.

• References. (1) Enderlein (1906); (2) Clay (1964); (3) Horning et al. (1980); (4) McIntosh & Murray (2007); (5) Leonardi et al. (2009); (6) Marcus et al. (2015).

• Remarks. Leonardi et al. (2009) report examining a group of specimens from a NZ fur seal in the Te Papa collection. Te Papa collection data show that these specimens were from Kaikoura and Open Bay Islands, NZ. This species has been found in otariids the world over and Leonardi and Palma (2013) suggest that the existing records may represent a complex of cryptic species. Host is called Arctocephalus forsteri in Leonardi et al. (2009).

Parasite name: Antarctophthirus ogmorhini Enderlein, 1906

• Synonyms. Echinophthirius setosus Rothschild, 1902 (nec E. setosus Burmeister 1838 = E. phocae Lucas, 1834) [Harrison, 1937].

• Hosts. Hydrurga leptonix (1, 2).

• Locality. (1, 2) Macquarie Island, TAS.

• Infection site. Fur/skin, body.

• Stage. Adults.

• References. (1) Harrison (1937); (2) Watson (1967).

• Remarks. Leonardi et al. (2014) erected Antarctophthirius carlinii for specimens from the Weddell seal (Leptonychotes weddelli (Lesson, 1826)). Antarctophthirius carlinii was, in turn, listed by Palma (2017) who included, as synonyms, many occurrences of A. ogmorhini in the literature.

Parasite name: Lepidophthirus macrorhini Enderlein, 1904

• Synonyms.

• Hosts. Mirounga leonina (1, 2–6).

• Locality. (1, 2–4, 6) MacQuarie Island, TAS; (5) Snares Island, NZ.

• Infection site. Fur/skin.

• Stage. Adult.

• References. (1) Harrison (1937); (2) Murray & Nicholls (1965); (3) Watson (1967); (4) Lowry et al. (1978); (5) Horning et al. (1980); (6) Palma & Horning (2002).

• Remarks. Host named Macrorhinus leoninus in Harrison (1937). A specimen in Te Papa Museum was collected from Kaikoura NZ.

• Subphylum: Crustacea

• Class: Hexanauplia Oakley, Wolfe, Lindgren & Zaharof, 2013

• Order: Sessilia Lamarck, 1818

• Family: Platylepadidae Newman & Ross, 1976

Parasite name: Platylepas hexastylos (Fabricius, 1798) Pilsbry, 1916

• Synonyms. Coronula bissexlobata Blainville, 1824; C. californiensis Chenu, 1825; Lepas hexastylos Fabricius, 1798; Platylepas bissexlobata (Blainville, 1824) Darwin, 1854; P. pulchra Gray, 1825 [Pilsbry, 1916].

• Hosts. Dugong dugon (1, 2).

• Locality. (1) Sydney Harbour, Australia; (2) Magnetic Island, QLD.

• Infection site. Skin, ventral surface.

• Stage.

• References. (1) Marlow (1962); (2) Zann & Harker (1978).

• Remarks.

• Order: Lepadiformes Buckeridge & Newman, 2006

• Family: Lepadidae Darwin, 1852

Parasite name: Lepas australis Darwin, 1851

• Synonyms.

• Hosts. Arctocephalus tropicalis.

• Locality. Adelaide, SA.

• Infection site. Body and flippers.

• Stage.

• References. Shaughnessy et al. (2014).

• Remarks. Shaughnessy et al. (2014) reported this barnacle from the pelage of the subantarctic fur seal, a vagrant in Australian waters.

• Family: Balanidae Leach, 1817

Parasite name: Balanus sp.

• Synonyms.

• Hosts. Dugong dugon.

• Locality. Wallum Creek, North Stradbroke Island, QLD.

• Infection site. Skin of back.

• Stage.

• References. Dexler & Freund (1906).

• Remarks. The authors state, ‘The dugong is the host of great numbers of parasites, both external and internal. On its back, as with whales, numerous barnacles establish themselves; a few Balanus but mostly Chelonobia [sic].’

• Family: Chelonibiidae Pilsbry, 1916

Parasite name: Chelonibia sp.

• Synonyms.

• Hosts. Dugong dugon.

• Locality. Wallum Creek, North Stradbroke Island, QLD.

• Infection site. Skin of back.

• Stage.

• References. Dexler & Freund (1906).

• Remarks. The authors report, ‘The dugong is the host of great numbers of parasites, both external and internal. On its back, as with whales, numerous barnacles establish themselves; a few Balanus but mostly Chelonobia [sic].’

• Class: Malacostraca Latreille, 1802

• Order: Amphipoda Latreille, 1816

• Family: Cyamidae Rafinesque, 1815

Parasite name: Cyamus boopis Lütken, 1870

• Synonyms. Cyamus ceti Chilton, 1883; C. elongatus Hiro, 1938; C. pacificus Lütken, 1873; C. suffuses Dall, 1872 [WoRMS].

• Hosts. Megaptera novaeangliae (1), unidentified South Australian whale (2), unidentified NZ whale (2).

• Locality. (1) Picton, NZ; (1) Carnarvon, WA; (2) unknown locality, SA, and unknown locality, NZ.

• Infection site. Skin.

• Stage. Adults, juveniles.

• References. (1) Leung (1965); (2) Sedlak-Weinstein (1991).

• Remarks. This species has been found to be highly specific in its choice of host, the humpback whale, M. novaeangliae (Carvalho et al., 2010), so it seems probable that the unidentified whales belonged to this species.

Parasite name: Cyamus catadontis Margolis, 1954

• Synonyms.

• Hosts. Physeter macrocephalus.

• Locality. Albany, WA.

• Infection site.

• Stage.

• References. Leung (1965).

• Remarks. In his paper, Leung (1965) lists a large number of cyamid specimens from all over the world, collected for a study of ectocommensal protozoans. No details are given other than the locality, date and collector. Leung names the sperm whale host as P. catodon.

Parasite name: Isocyamus delphinii (Guérin-Méneville, 1836) Gervais & Beneden, 1859

• Synonyms. Cyamus delphinii Guérin-Méneville, 1836; C. globicipitis Lütken, 1873; Isocyamus globicipitis, Hiro, 1938 [Leung, 1967].

• Hosts. Globicephala melas (1), Phocoena phocoena (2), Pseudorca crassidens (3).

• Locality. (1) Jarvis Bay, NSW; (2) Australia; (3) Little Manly and Crowdy Heads, NSW.

• Infection site. Skin.

• Stage. Adults, juveniles.

• References. (1) Leung (1965); (2) Berzin & Vlasova (1982); (3) Sedlak-Weinstein (1991).

• Remarks. Leung (1965) names the host G. malaena. Berzin and Vlasova (1982) list this species as occurring in Phocoena phocoena, which is probably a host identification error, as this porpoise does not inhabit the Australasian region. It seems probable that this host was Phocoena dioptrica.

Parasite name: Isocyamus deltobranchium Sedlak-Weinstein, 1992

• Synonyms.

• Hosts. Globicephala melas.

• Locality. Coast of TAS.

• Infection site. Skin.

• Stage. Adults.

• References. Sedlak-Weinstein (1992b).

• Remarks. Host named G. malaena in Sedlak-Weinstein (1992b).

Parasite name: Isocyamus kogiae Sedlak-Weinstein, 1992

• Synonyms.

• Hosts. Kogia breviceps.

• Locality. Moreton Island, QLD.

• Infection site. Skin.

• Stage. Adults.

• References. Sedlak-Weinstein (1992a).

• Remarks.

Parasite name: Platycyamus thompsoni (Gosse, 1855) Lütken, 1870

• Synonyms. Cyamus thompsoni Gosse, 1855 [WoRMS].

• Hosts. Mesoplodon grayi.

• Locality. Younghusband Peninsula, SA.

• Infection site. Skin.

• Stage. Adults.

• References. Sedlak-Weinstein (1991).

• Remarks. New location and new host record.

Parasite name: Scutocyamus antipodensis Lincoln & Hurley, 1980

• Synonyms.

• Hosts. Cephalorhynchus hectori (1), Phocoena dioptrica (2).

• Locality. (1) Cook Strait, NZ; (2) Caroline Bay, NZ.

• Infection site. Skin.

• Stage. Adults, juveniles.

• References. (1) Lincoln & Hurley (1980); (2) Lehnert et al. (2017).

• Remarks.

Parasite name: Syncyamus aequus Lincoln & Hurley, 1981

• Synonyms.

• Hosts. Tursiops truncatus, Stenella longirostris.

• Locality. Arafura Sea, north-west Australia.

• Infection site. Skin.

• Stage. Adults, juveniles.

• References. Sedlak-Weinstein (1991).

• Remarks. New host location in Arafura Sea for this species.

Parasite name: Syncyamus pseudorcae Bowman, 1955

• Synonyms.

• Hosts. Pseudorca crassidens.

• Locality. Crowdy Heads, NSW.

• Infection site. Skin.

• Stage. Adults, juveniles.

• References. Sedlak-Weinstein (1991).

• Remarks.

• Subphylum: Apicomplexa

• Class: Conoidasida Levine, 1988

• Order: Eucoccidiorida Léger & Duboscq, 1910

• Family: Sarcocystidae Poche, 1913

Parasite name: Cryptosporidium hominis Morgan-Ryan, Fall, Ward, Hijawi, Sulaiman, Fayer, Thompson, Olson, Lal & Xiao, 2002

• Synonyms. Cryptosporidium parvum Tyzzer, 1912 (in part).

• Hosts. Dugong dugon (1, 2).

• Locality. (1, 2) Hervey Bay, QLD.

• Infection site. Small intestine.

• Stage.

• References. (1) Hill et al. (1997); (2) Morgan et al. (2000).

• Remarks. Hill et al. (1997) do not give a species name for Cryptosporidium. Three animals were found dead, one euthanized with anorexia and lethargy, putative oocysts found. Using the same specimens, Morgan et al. (2000) provide molecular data leading to identification as C. parvum ‘human’ genotype. This genotype was later described as C. hominis by Morgan-Ryan et al. (2002).

Parasite name: Toxoplasma gondii Nicolle & Manceaux, 1908

• Synonyms.

• Hosts. Tursiops aduncus (1), Sousa chinensis (2), Dugong dugon (3), Arctophoca australis forsteri (4), Phocarctos hookeri (5).

• Locality. (1) In captivity, Australia; (2) Townsville and Gladstone Harbour, QLD; (3) south-east QLD; (4) Sydney, NSW; (5) Otago, NZ.

• Infection site. Brain (tissue cysts), heart, lungs, liver, spleen, adrenal glands (tachyzoite stages).

• Stage.

• References. (1) Jardine & Dubey (2002); (2) Bowater et al. (2003); (3) Owen et al. (2012); (4) Donahoe et al. (2014); (5) Roe et al. (2017).

• Remarks. Congenital toxoplasmosis in a stillborn late-term foetus with myocardial necrosis and nonsuppurative necrotizing encephalitis were associated with tachyzoites and tissue cysts (Jardine & Dubey, 2002); systemic toxoplasmosis, molecular data (Donahoe et al., 2014); encephalitis associated with T. gondii cysts, emaciation, ataxia in one live stranded animal (Bowater et al., 2003). Host called Arctocephalus forsteri in Donahoe et al. (2014).

• Family: Hexamitidae Kent, 1880

Parasite name: Giardia duodenalis (Lambl, 1859) Kofoid & Christiansen, 1915

• Synonyms. Cercomonas intestinalis Lambl, 1859; Giardia agilis Kunstler, 1882; G. lamblia Stiles, 1914; Hexamita duodenalis Davaine, 1875; Lamblia intestinalis Blanchard, 1888.

• Hosts. Neophoca cinerea.

• Locality. WA, SA, in captivity QLD and NSW.

• Infection site. Intestine.

• Stage.

• References. Delport et al. (2014).

• Remarks. Analysed from faecal samples, molecular data (18S).

Host–parasite list

Host taxon: Pinnipedia

• Family: Otariidae

Host name: Arctocephalus pusillus doriferus (Australian fur seal)

• Acanthocephala

  • Corynosoma australe

  • Corynosoma cetaceum

• Nematoda

  • Anisakis sp.

  • Contracaecum ogmorhini

  • Contracaecum osculatum

  • Parafilaroides normani

  • Parafilaroides sp.

  • Uncinaria sp. 2 of Nadler et al. ( 2013)

• Cestoda

  • Adenocephalus pacificus

• Arachnida

  • Orthohalarachne attenuata

  • Orthohalarachne diminuata

Host name: Arctocephalus tropicalis (subantarctic fur seal)

• Crustacea

  • Lepas australis

Host name: Arctophoca australis forsteri (NZ fur seal)

• Gastrointestinal parasites

• Acanthocephala

  • Corynosoma australe

  • Corynosoma sp.

• Nematoda

  • Anisakis simplex s.l.

  • Anisakis sp.

  • Contracaecum osculatum

  • Parafilaroides normani

  • Parafilaroides sp.

  • Pseudoterranova decipiens

  • Uncinaria sp.

• Cestoda

  • Diphyllobothrium sp.

  • Phyllobothrium sp.

• Trematoda gen. sp.

• Insecta

  • Antarctophthirus microchir

• Protozoa

  • Toxoplasma gondii

Host name: Neophoca cinerea (Australian sea lion)

• Acanthocephala

  • Corynosoma australe

  • Corynosoma sp.

• Nematoda

  • Contracaecum osculatum

  • Parafilaroides sp.

  • Uncinaria hamiltoni

  • Uncinaria sanguinis

  • Uncinaria sp.

• Cestoda

  • Adenocephalus pacificus

• Trematoda

  • Galactosomum angelae

  • Mesostephanus neophocae

  • Stictodora diplacantha

  • Synthesium sp.

• Arachnida

  • Orthohalarachne attenuata

  • Orthohalarachne diminuata

• Insecta

  • Antarctophthirus microchir

• Protozoa

  • Giardia duodenalis

Host name: Phocarctos hookeri (NZ sea lion)

• Acanthocephala

  • Corynosoma australe

  • Corynosoma semerme

• Nematoda

  • Anisakis simplex s.l.

  • Pseudoterranova decipiens

  • Parafilaroides decorus

  • Uncinaria sp.

  • Uncinaria sp. 1 of Nadler et al. (2013)

• Cestoda

  • Phyllobothrium sp.

• Insecta

  • Antarctophthirus michrochir

• Protozoa

  • Toxoplasma gondii

Host name: Zalophus californianus (Californian sea lion) captive animal

• Arachnida

  • Demodex zalophi

• Family: Phocidae

Host name: Hydrurga leptonyx (Leopard seal)

• Acanthocephala

  • Corynosoma australe

  • Corynosoma bullosum

  • Corynosoma sp.

• Nematoda

  • Anisakis simplex s.l.

  • Contracaecum mirounga

  • Contracaecum ogmorhini

  • Contracaecum osculatum

  • Contracaecum radiatum

  • Parafilaroides hydrurgae

  • Parafilaroides sp.

  • Pseudoterranova decipiens

• Cestoda

  • Diphyllobothrium sp.

• Insecta

  • Antarctophthirus ogmorhini

Host name: Mirounga leonina (Southern elephant seal)

• Acanthocephala

  • Corynosoma bullosum

• Nematoda

  • Anisakis simplex s.l.

  • Contracaecum mirounga

  • Contracaecum osculatum

  • Contracaecum radiatum

  • Filaria s.l. sp.

  • Pseudoterranova decipiens

  • Uncinaria sp. 3 of Nadler et al. (2013)

• Cestoda

  • Phyllobothrium delphini

• Trematoda

  • Mesostephanus neophocae

• Arachnida

  • Halarachne miroungae

• Insecta

  • Lepidophthirus macrorhini

Host taxon: Cetacea

• Family: Balaenopteridae

Host name: Balaenoptera acutorostrata (Minke whale)

• Nematoda

  • Anisakis sp.

• Cestoda

  • Phyllobothrium sp.

Host name: Megaptera novaeangliae (Humpback whale)

• Acanthocephala

  • Bolbosoma balaenae

• Crustacea

  • Cyamus boopis

• Family: Neobalaenidae

Host name: Caperea marginata (pygmy right whale)

• Trematoda

  • Ogmogaster sp.

• Family: Delphinidae

Host name: Cephalorhynchus hectori (Hector's dolphin)

• Acanthocephala

  • Corynosoma sp.

• Nematoda

  • Acuaria sp.

  • Contracaecum sp.

  • Halocercus sp.

  • Phocascaris sp.

  • Stenurus sp.

• Cestoda

  • Phyllobothrium delphini

  • Phyllobothrium sp.

• Trematoda

  • Braunina cordiformis

  • Campula sp.

• Crustacea

  • Scutocyamus antipodensis

Host name: Delphinus delphis (short-beaked common dolphin)

• Acanthocephala

  • Corynosoma cetaceum

  • Corynosoma sp.

• Nematoda

  • lung worms

  • Anisakis pegreffii

  • Anisakis simplex s.l.

  • Crassicauda sp.

  • Echinocephalus overstreeti

  • Halocercus lagenorhynchi

  • Halocercus sp.

  • Parafilaroides sp.

  • Pharurus alatus

  • Stenurus ovatus

• Cestoda

  • Monorygma grimaldii

  • Phyllobothrium delphini

  • Phyllobothrium sp.

  • Tetrabothrius forsteri

  • Tetrabothrius sp.

• Trematoda

  • Brachycladium delphini

  • Brachycladium palliatum

  • Heterophyidae gen. sp.

Host name: Globicephala melas (long-finned pilot whale)

• Acanthocephala

  • Bolbosoma capitatum

• Nematoda

  • Anisakis oceanica

  • Anisakis simplex s.l.

  • Anisakis sp.

  • Stenurus globicephalae

• Crustacea

  • Isocyamus delphinii

  • Isocyamus deltobranchium

Host name: Globicephala sp.

• Nematoda

  • Stenurus globicephalae

Host name: Grampus griseus (Risso's dolphin)

• Nematoda

  • Crassicauda grampicola

  • Stenurus globicephalae

Host name: Lagenodelphis hosei (Fraser's dolphin)

• Nematoda

  • Anisakis simplex s.l.

  • Stenurus ovatus

• Cestoda

  • Monorygma grimaldii

  • Tetrabothrius sp.

• Trematoda

  • Campula sp.

  • Nasitrema sp.

Host name: Lagenorhynchus obscurus (dusky dolphin)

• Nematoda

  • Anisakis simplex s.l.

  • Crassicauda sp.

• Cestoda

  • Phyllobothrium delphini

  • Phyllobothrium sp.

Host name: Peponocephala electra (melon-headed whale)

• Nematoda

  • Anisakis simplex s.l.

  • Anisakis typica

  • Stenurus globicephalae

• Cestoda

  • Monorygma chamissonii

Host name: Pseudorca crassidens (false killer whale)

• Acanthocephala

  • Bolbosoma capitatum

• Nematoda

  • Crassicauda sp.

• Crustacea

  • Isocyamus delphinii

  • Syncyamus pseudorcae

Host name: Sousa chinensis (Indo-Pacific humpbacked dolphin)

• Protozoa

  • Toxoplasma gondii

Host name: Stenella coeruleoalba (striped dolphin)

• Nematoda

  • Halocercus sp.

  • Halocercus delphinii

• Cestoda

  • Monorygma grimaldii

  • Phyllobothrium delphini

Host name: Stenella longirostris (spinner dolphin)

• Crustacea

  • Syncyamus aequus

Host name: Tursiops aduncus (Indo-Pacific bottlenose dolphin)

• Nematoda

  • Halocercus lagenorhynchi

  • Stenurus ovatus

• Protozoa

  • Toxoplasma gondii

Host name: Tursiops truncatus (common bottlenose dolphin)

• Acanthocephala

  • Bolbosoma sp.

  • Corynosoma cetaceum

• Nematoda

  • Anisakis pegreffii

  • Anisakis simplex s.l.

  • Halocercus lagenorhynchi

  • Skrjabinalius cryptocephalus

  • Stenurus ovatus

• Cestoda

  • Phyllobothrium sp.

• Trematoda

  • Synthesium tursionis

• Crustacea

  • Syncyamus aequus

• Family: Kogiidae

Host name: Kogia breviceps (pygmy sperm whale)

• Nematoda

  • Anisakis physeteris

  • Anisakis simplex s.l.

  • Anisakis sp.

  • Crassicauda magna

  • Pseudoterranova decipiens

  • Pseudoterranova kogiae

• Cestoda

  • Phyllobothrium delphini

• Crustacea

  • Isocyamus kogiae

Host name: Kogia sima (dwarf sperm whale)

• Nematoda

  • Anisakis berlandi

  • Anisakis brevispiculata

• Family: Phocoenidae

Host name: Phocoena dioptrica (spectacled porpoise)

• Nematoda

  • Anisakis sp.

  • Stenurus minor

• Cestoda

  • Phyllobothrium delphini

• Crustacea

  • Scutocyamus antipodensis

Host name: Phocoena (harbour porpoise)

• Crustacea

  • Isocyamus delphinii

• Family: Physeteridae

Host name: Physeter macrocephalus (sperm whale)

• Crustacea

  • Cyamus catadontis

• Family: Ziphiidae

Host name: Mesoplodon bowdoini (Andrew's beaked whale)

• Nematoda

  • Anisakis nascetti

  • Anisakis ziphidarum

Host name: Mesoplodon grayi (Gray's beaked whale)

• Nematoda

  • Anisakis nascetti

  • Anisakis sp.

• Crustacea

  • Platycyamus thompsoni

Host name: Mesoplodon hectori (Hector's beaked whale)

• Nematoda

  • Anisakis sp.

• Cestoda

  • Tetrabothrius forsteri

Host name: Mesoplodon layardii (strap-toothed whale)

• Nematoda

  • Anisakis nascetti

Host name: Ziphius cavirostris (Cuvier's beaked whale)

• Nematoda

  • Crassicauda boopis

  • Crassicauda sp.

Host taxon: Sirenia

• Family: Sirenidae

Host name: Dugong dugon (dugong)

• Nematoda

  • Paradujardinia halicoris

• Trematoda

  • Faredifex clavata

  • Folitrema jecoris

  • Haerator caperatus

  • Labicola cf. elongata

  • Labicola elongata

  • Lankatrema macrocotyle

  • Lankatrema mannarense

  • Lankatrema microcotyle

  • Lankatrema minutum

  • Lankatrematoides gardneri

  • Opisthotrema australe

  • Opisthotrema dujonis

  • Pulmonicola pulmonalis

  • Rhabdiopoeus taylori

  • Solenorchis travassosi

  • Spirorchidae gen. sp.

  • Taprobanella bicaudata

• Crustacea

  • Platylepas hexastylos

  • Balanus sp.

  • Chelonibia sp.

• Protozoa

  • Cryptosporidium hominis

  • Toxplasma gondii

Discussion

In this first host–parasite checklist, information about metazoan and protozoan parasites of marine mammals in NZ and Australian waters was collated. From 51 species of cetacean known from Australian and NZ waters, only 27 species have recorded parasites. From 11 species of pinnipeds known from Australian and NZ waters, eight have recorded parasites. The absence of records certainly does not signify that the remaining hosts are parasite free. There is still a lot left to learn. However, checklists such as this one remain valuable tools to ecologists and can help to further our understanding of parasite diversity and be a practical resource for scientists.

Nematodes were the most diverse group reported, with 30 different species determined, and many more records where identification was restricted to the genus or family level. Anisakid stomach nematodes (14 species) represented the family most often found in a wide range of host species, reflecting their generalist nature, followed by pseudaliid lungworms (seven species), specific to the respiratory tract of odontocetes. Trematodes (22 species), mostly from the gastro-intestinal tract, were found in sirenians (15 species), cetaceans (six species) and pinnipeds (six species). Six species of acanthocephalans were identified from pinniped (n = 6) and cetacean (n = 3) hosts. Adult cestodes (five species) were recorded in the intestinal tract of three cetacean and two pinniped species. Cestode larvae within the subcutaneous blubber and the peritoneum were recorded from multiple cetacean (n = 9) and pinniped (n = 3) species. Thirteen different ectoparasitic crustacean species were found on cetaceans and sirenians. Other arthropod ecto- (Insecta, three species) and endoparasites (Arachnida, four species) were recorded from pinniped species. Three protozoan species were encountered in several marine mammal species including sirenians, pinnipeds and cetaceans within studies dating from 1997 to 2017. This may reflect, on the one hand, the relatively recent occurrence of some of these pathogens in the marine environment due to human activities and, on the other hand, new survey and diagnostic techniques (Lasek-Nesselquist et al., 2010), as well as growing awareness of these emerging zoonotic agents in the research community.

Thirty-four parasite records were from dugongs and 25 species were found in common dolphins, while leopard seals with 17 parasite species were the pinnipeds most often infected. This reflects the opportunistic nature of sampling these animals. The findings indicate that, for example, sperm whales with one crustacean ectoparasite record or elusive beaked whales with 2–3 species records are seldom encountered hosts.

Many old records have complicated histories with multiple synonyms, which need updating. Also, the advent of molecular tools has identified several species complexes (Mattiucci et al., 2014; Klotz et al., 2018), for which retrospective analyses would be useful. New techniques will likely also continue to improve the identification of parasite fragments and minute larvae (Jabbar et al., 2015), as well as provide insights on the phylogeny of parasite species (Hernández-Orts et al., 2017).

The protozoans Toxoplasma, Giardia and Cryptosporidium are emerging parasites in marine mammals. In the future, the combination of prevalence surveys with molecular techniques will probably identify further protozoans and provide knowledge on host specificity and transmission pathways (Fayer et al., 2004; Applebee et al., 2005; Grilo et al., 2018).

Foodborne parasites like cestodes (Diphyllobothrium sp.) and anisakids (Anisakis sp., Pseudoterranova sp.) are abundant in marine mammals in Australasia and pathogenic for humans when infective larvae are accidentally consumed with undercooked fish (Yamasaki & Kuramochi, 2009; Shamsi & Butcher, 2011). Human health concerns as well as implications for fisheries and seafood control underline the importance of better understanding the epidemiology of relevant species (Cipriani et al., 2016).

Opportunistic parasite surveys of dead or live stranded cetaceans, in cooperation with established stranding networks (e.g. that of NZ's Department of Conservation) and systematic, minimally invasive studies to monitor live and free ranging pinnipeds (e.g. analysing faeces from pinniped colonies; Presswell & Lagrue, 2016), would enable a better overview of prevalence, intensity of infections and emerging parasite species. The importance of well-maintained and curated museum collections cannot be underestimated in their contribution towards our ongoing knowledge of parasite biodiversity (Lehnert et al., 2017). It remains unclear how contaminant exposure causing immune suppression or cumulative effects of human-induced pressures (shipping, fisheries, global change) make marine mammals more susceptible to infectious disease in Australasian waters (Van Bressem et al., 2009). Studies combining ecotoxicological analyses with systematic monitoring of parasite prevalence and impact (Lehnert et al., 2018) would help to elucidate these relationships in the future.

Parasites should be an integral part of biodiversity and conservation research in their marine mammal hosts (Aznar et al., 2010; Poulin et al., 2016). Ultimately, this research can inform managers and may guide species, habitat and population assessment and conservation, as well as encourage further investigations into the biodiversity and ecology of marine mammal parasites.