Checklist of marine mammal parasites in New Zealand and Australian waters

K. Lehnert; R. Poulin; B. Presswell

doi:10.1017/S0022149X19000361

Checklist of marine mammal parasites in New Zealand and Australian waters

Published online by Cambridge University Press: 24 June 2019

K. Lehnert

R. Poulin and

B. Presswell

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K. Lehnert*: Affiliation:
Institute for Terrestrial and Aquatic Wildlife Research, University of Veterinary Medicine Hannover, Foundation, Bünteweg 2, 30559 Hannover, Germany
R. Poulin: Affiliation:
Department of Zoology, University of Otago, 340 Great King Street, PO Box 56, Dunedin 9054, New Zealand
B. Presswell: Affiliation:
Department of Zoology, University of Otago, 340 Great King Street, PO Box 56, Dunedin 9054, New Zealand
*: Author for correspondence: K. Lehnert, E-mail: kristina.lehnert@tiho-hannover.de

Article contents

Abstract
Introduction
Material and methods
Results
Discussion
References

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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.

Keywords

Metazoa protozoa cetaceans pinnipeds arthropods ecology bioindicators conservation

Type: Review Article
Information: Journal of Helminthology , Volume 93 , Issue 6 , November 2019 , pp. 649 - 676

DOI: https://doi.org/10.1017/S0022149X19000361 [Opens in a new window]
Creative Commons: This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use, distribution, and reproduction in any medium, provided the original work is properly cited.
Copyright: Copyright © Cambridge University Press 2019

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., Reference Baker, Chilvers, Constantine, DuFresne, Mattlin, Van Helden and Hitchmough2010). 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., Reference Baker, Chilvers, Constantine, DuFresne, Mattlin, Van Helden and Hitchmough2010). 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., Reference Schumann, Gales, Harcourt and Arnould2013). 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., Reference Bannister, Kemper and Warneke1996; Salgado Kent et al., Reference Salgado Kent, Jenner, Jenner, Bouchet and Rexstad2012). 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., Reference Kovacs, Aguilar and Aurioles2012) and NZ sea lion (Robertson & Chilvers, Reference Robertson and Chilvers2011). 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., Reference Kingsford, Watson and Lundquist2009).

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, Reference Measures, Samuel, Pybus and Kocan2001; Siebert et al., Reference Siebert, Wünschmann, Weiss, Frank, Benke and Frese2001). Although Australian and NZ waters are a hotspot for marine mammal species richness (Pompa et al., Reference Pompa, Ehrlich and Ceballos2011), 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., Reference Siebert, Gilles, Lucke, Ludwig, Benke, Kock and Scheidat2006; Lehnert et al., Reference Lehnert, Seibel, Hasselmeier, Wohlsein, Iversen, Nielsen and Siebert2014), reflecting habitat use (Aznar et al., Reference Aznar, Raga, Corcuera and Monzón1995), diet (Marcogliese, Reference Marcogliese2002), social behaviour and population dynamics (Balbuena and Raga, Reference Balbuena and Raga1994), but also as markers for exposure and detrimental effects of xenobiotics (Sures, Reference Sures2004; Pascual & Abollo, Reference Pascual and Abollo2005; Marcogliese & Pietrock, Reference Marcogliese and Pietrock2011). They reveal evolutionary host–parasite relationships and highlight their biogeography and phylogeny over historical timescales (Anderson, Reference Anderson1982; Leidenberger et al., Reference Leidenberger, Harding and Härkönen2007). 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, Reference Gulland and Hall2007; van Bressem et al., Reference Van Bressem, Raga and Di Guardo2009). 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., Reference Dorny, Praet, Deckers and Gabriël2009; Shamsi, Reference Shamsi2014), and can induce severe gastro-intestinal disease when ingested via undercooked fish (Mattiucci et al., Reference Mattiucci, Fazii and De Rosa2013; Bao et al., Reference Bao, Pierce, Pascual, González-Muñoz, Mattiucci, Mladineo, Cipriani, Bušelić and Strachan2017). 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., Reference Llarena-Reino, Abollo, Regueira, Rodríguez and Pascual2015). Zoonotic protozoans like Giardia and Cryptosporidium are significant enteropathogens in NZ, causing higher infection rates than in other developed countries (Britton et al., Reference Britton, Hales, Venugopal and Baker2010). 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., Reference Fayer, Dubey and Lindsay2004).

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., Reference Poulin, Blasco-Costa and Randhawa2016). 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., Reference Berón-Vera, Crespo and Raga2008; Nikolov et al., Reference Nikolov, Cappozzo, Berón-Vera, Crespo, Raga and Fernández2010) exist. Parasitology increasingly complements marine ecology to further our understanding of ecosystem dynamics (Poulin et al., Reference Poulin, Blasco-Costa and Randhawa2016), but, so far, little is known about the biodiversity of Australasian marine parasites (Poulin, Reference Poulin2004; Stockin et al., Reference Stockin, Duignan, Roe, Meynier, Alley and Fettermann2009). New studies on marine mammals in NZ and Australian waters have recently added to our knowledge about parasite prevalence, life cycles (Tomo et al., Reference Tomo, Kemper and Lavery2010; Lehnert et al., Reference Lehnert, Randhawa and Poulin2017) and taxonomic relationships (Shamsi et al., Reference Shamsi, Gasser and Beveridge2012, Hernández-Orts et al., Reference Hernández-Orts, Smales, Pinacho-Pinacho, García-Varela and Presswell2017), and justify a first host–parasite checklist encompassing all available data. Previous marine mammal parasite checklists include Baylis (Reference Baylis1932), Delyamure (Reference Delyamure1955), Dailey & Brownell (Reference Dailey, Brownell and Ridgway1972), Raga (Reference Raga, Robineau, Duguy, Robineau and Klima1994), Felix (Reference Felix2013) and Fraija-Fernández et al. (Reference Fraija-Fernández, Fernández, Raga and Aznar2016). Most include host data, but geographical data are scanty. Australian lists include that of the internal parasites of mammals by MacKerras (Reference Mackerras1958), the references of which are unfortunately disconnected from the text and, therefore, not useable, and Arundel (Reference Arundel1978), 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, Reference Blair1981: dugong monostome digeneans; Bowie, Reference Bowie1984: bottle-nosed dolphin parasites), and parasite taxa (e.g. Price, Reference Price1932: trematodes in marine mammals; Leung, Reference Leung1967: whale lice of cetaceans). There have also been compilations with a veterinary perspective (e.g. Duignan, Reference Duignan2000; Dailey, Reference Dailey, Dierauf and Gulland2001; Ladds, Reference Ladds2009; McFarlane et al., Reference McFarlane, Norman, Jones, Kerry and Riddle2009) and those looking at certain pathologies (e.g. Baylis & Daubney, Reference Baylis and Daubney1925: lungworms of cetacean; Spratt, Reference Spratt2002: respiratory parasites in Australian mammals; Measures, Reference Measures, Samuel, Pybus and Kocan2001: 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., Reference Castinel, Duignan, Pomroy, Lopez-Villalobos, Gibbs, Chilvers and Wilkinson2007), 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. (Reference Anderson, Chabaud and Willmott2009) (Nematoda); Caira & Jensen (Reference Caira and Jensen2017) (Cestoda); Olson et al. (Reference Olson, Cribb, Tkach, Bray and Littlewood2003), Gibson et al. (Reference Gibson, Jones and Bray2002), Jones et al. (Reference Jones, Bray and Gibson2005) and Bray et al. (Reference Bray, Gibson and Jones2008) (Trematoda); and Amin (Reference Amin2013) (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 (Reference Berta and Churchill2012) (Pinnipeda) and Perrin (Reference Perrin2018) (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, Reference Amin2013; Yamaguti, Reference Yamaguti1963].
Hosts. Megaptera novaeangliae.
Localities. Bondi Beach, NSW.
Infection site.
Stage.
References. Johnston & Deland (Reference Johnston and Deland1929).
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, Reference Amin2013; Felix, Reference Felix2013].
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 (Reference Edmonds1957b); (2) McManus et al. (Reference McManus, Wapstra, Guiler, Munday and Obendorf1984); (3) Edmonds (Reference Edmonds1987).
Remarks. Host reported in Edmonds (Reference Edmonds1957b) as Melon-headed whale (G. melaenas = G. melas); later corrected to false killer whale in Edmonds (Reference Edmonds1987).

Parasite name: Bolbosoma sp.

Synonyms.
Hosts. Tursiops truncatus.
Localities. Otago Harbour, NZ.
Infection site. Intestine.
Stage. Immature.
References. Bowie (Reference Bowie1984).
Remarks. Single specimen.

Parasite name: Corynosoma australe Johnston, 1937

Synonyms. Corynosoma otariae Morino & Boero, 1960 [Amin, Reference Amin2013].
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 (Reference Johnston1937); (2) Johnston & Edmonds (Reference Johnston and Edmonds1953); (3) Obendorf & Presidente (Reference Obendorf and Presidente1978); (4) Cordes & O'Hara (Reference Cordes and O'Hara1979); (5) Smales (Reference Smales1986); (6) García-Varela et al. (Reference García-Varela, Pérez-Ponce de Léon, Aznar and Nadler2013).
Remarks. Johnston (Reference Johnston1937) described host as A. forsteri, but corrected this to N. cinerea in Johnston & Mawson (Reference Johnston and Mawson1941). Although Johnston & Edmonds (Reference Johnston and Edmonds1953) listed a number of ways in which their specimens differed from those of Johnston (Reference Johnston1937), they nonetheless record them as C. australe. Smales (Reference Smales1986) agrees in her list of comparative specimens of the species. In her paper, the host is called A. pusillus. Johnston & Edmonds (Reference Johnston and Edmonds1953) 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, Reference Amin2013].
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 (Reference Johnston and Edmonds1953); (2) Edmonds (Reference Edmonds1955); (3) Edmonds (Reference Edmonds1957a).
Remarks. Edmonds reported 200 (Reference Edmonds1957a), and over 100 (Reference Edmonds1955) 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, Reference Amin2013].
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 (Reference Johnston and Best1942); (2) Smales (Reference Smales1986).
Remarks. Described as new species in Johnston & Best (Reference Johnston and Best1942). Described as P. arctocephali n. sp., and host as A. pusillus in Smales (Reference Smales1986), synonymized by Aznar et al. (Reference Aznar, Bush and Raga1999).

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

Synonyms. Corynosoma gibber (Olsson, 1894) Lühe, 1911; Echinorhynchus semermis Forssell, 1904 (Amin, Reference Amin2013).
Hosts. Phocarctos hookeri.
Locality. Auckland/Campbell Island, NZ.
Infection site.
Stage.
References. Johnston & Edmonds (Reference Johnston and Edmonds1953).
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 (Reference Johnston and Deland1929); (2) Johnston & Edmonds (Reference Johnston and Edmonds1953); (3) McKenzie & Blair (Reference McKenzie and Blair1983); (4) Smales (Reference Smales1986).
Remarks. The specimens of Johnston and Deland (Reference Johnston and Deland1929) were later designated as C. cetaceum (Johnston & Best, Reference Johnston and Best1942). Host called Otaria forsteri in Johnston and Edmonds (Reference Johnston and Edmonds1953). McKenzie & Blair (Reference McKenzie and Blair1983) 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. (Reference Mattiucci, Nascetti and Clanchi1997).
Hosts. Kogia sima.
Locality. Coast of NSW.
Infection site. Stomach.
Stage. Larva.
References. Shamsi et al. (Reference Shamsi, Gasser and Beveridge2012).
Remarks. Molecular data available (ITS1&2). The specimens in this paper referred to as A. simplex C following Mattiucci et al. (Reference Mattiucci, Nascetti and Clanchi1997) were later assigned to A. berlandi by Mattiucci et al. (Reference Mattiucci, Cipriani, Webb, Paoletti, Marcer, Bellisario, Gibson and Nascetti2014).

Parasite name: Anisakis brevispiculata Dollfus, 1966

Synonyms.
Hosts. Kogia sima.
Locality. Coast of NSW.
Infection site. Stomach.
Stage. Adult.
References. Shamsi et al. (Reference Shamsi, Gasser and Beveridge2012).
Remarks. Molecular data available (ITS1&2).

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

Synonyms. Anisakis sp. of Pontes et al. (Reference Pontes, D'Amelio, Costa and Paggi2005) and Iglesias et al. (Reference Iglesias, D'Amelio, Ingrosso, Farjallah, Martınez-Cedeira and Garcıa-Estevez2008); Anisakis sp. of Valentini et al. (Reference Valentini, Mattiucci, Bondanelli, Webb, Mignucci-Giannone, Colom-Llavina and Nascetti2006).
Hosts. Mesoplodon bowdoini, M. grayi, M. layardii.
Locality. South Pacific Ocean, off NZ.
Infection site. Stomach.
Stage. Adult.
References. Mattiucci et al. (Reference Mattiucci, Paoletti and Webb2009).
Remarks. Molecular data available (cox2). The intermediate host was confirmed molecularly as the squid, Onykia ingens by Mattiucci et al. (Reference Mattiucci, Paoletti and Webb2009) (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 (Reference Johnston and Mawson1951).
Remarks. Described as S. oceanicus in this paper, with the host called Globicephalus ventricosas. Davey (Reference Davey1971) synonymized this species with A. physeteris, but Shamsi et al. (Reference Shamsi, Gasser and Beveridge2012) and Shamsi (Reference Shamsi2014) have kept the species separate as a sp. inq. Species is also accepted by Mattiucci et al. (Reference Mattiucci, Nascetti, Dailey, Webb, Barros, Cianchi and Bullini2005).

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. (Reference Shamsi, Gasser and Beveridge2012).
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 (Reference Hurst1980).
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 Reference Davey1971].
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) Reference Johnston and Mawson1939, (2) Reference Johnston and Mawson1941, (3) Reference Johnston and Mawson1942, (4) Reference Johnston and Mawson1945, (5) Reference Johnston and Mawson1953); (6) Mawson (Reference Mawson1953); (7) Brundson (Reference Brundson1956); (8) Cannon (Reference Cannon1977); (9) Morgan et al. (Reference Morgan, Caple, Westbury and Campbell1978); (10) Cordes & O'Hara (Reference Cordes and O'Hara1979); (11) Hurst (Reference Hurst1980); (12) McColl & Obendorf (Reference McColl and Obendorf1982); (13) Bowie (Reference Bowie1984); (14) McManus et al. (Reference McManus, Wapstra, Guiler, Munday and Obendorf1984).
Remarks. Anisakis simplex is widely accepted to be a complex of closely related species (Nascetti et al., Reference Nascetti, Paggi, Orecchia, Smith, Mattiucci and Bullini1986; Mattiucci et al., Reference Mattiucci, Nascetti and Clanchi1997, Reference Mattiucci, Paoletti and Webb2009, Reference Mattiucci, Cipriani, Webb, Paoletti, Marcer, Bellisario, Gibson and Nascetti2014; D'Amelio et al., Reference D'Amelio, Mathiopoulos, Santos, Pugachev, Webb, Picanço and Paggi2000; Abe et al., Reference Abe, Tominaga and Kimata2006; Abe, Reference Abe2008), and the name has been attached to specimens the world over and in numerous hosts. Davey (Reference Davey1971) 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. (Reference Mattiucci, Cipriani, Webb, Paoletti, Marcer, Bellisario, Gibson and Nascetti2014) 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. (Reference Mattiucci, Cipriani, Webb, Paoletti, Marcer, Bellisario, Gibson and Nascetti2014), 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, Reference Johnston and Mawson1943a, Reference Johnston and Mawsonb, Reference Johnston and Mawson1945, Reference Johnston and Mawson1951, Reference Johnston and Mawson1953). Brundson (Reference Brundson1956) 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 (Reference Hurst1980) 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 (Reference Johnston and Mawson1941); (2) Cordes & O'Hara (Reference Cordes and O'Hara1979); (3) Dawson & Slooten (Reference Dawson and Slooten1990); (4) Baker et al. (Reference Baker, Duignan, Norman and Helden2001); (5) Valentini et al. (Reference Valentini, Mattiucci, Bondanelli, Webb, Mignucci-Giannone, Colom-Llavina and Nascetti2006); (6) Beatson & O'Shea (Reference Beatson and O'Shea2009); (7) Lehnert et al. (Reference Lehnert, Randhawa and Poulin2017).
Remarks. Molecular data available (cox2) (5). The specimens from Grey's beaked whale (Valentini et al., Reference Valentini, Mattiucci, Bondanelli, Webb, Mignucci-Giannone, Colom-Llavina and Nascetti2006) 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 (Reference Johnston and Mawson1941) 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., Reference Li, Gibson and Zhang2016].
Hosts. Peponocephala electra.
Locality. Moreton Island, QLD; Tweed Heads, NSW.
Infection site. Stomach.
Stage. Adults and immatures.
References. Cannon (Reference Cannon1977).
Remarks. Shamsi (Reference Shamsi2014) note that A. typica specimens in Australia appear to be genetically different from those reported in other countries. Jabbar et al. (Reference Jabbar, Asnoussi, Norbury, Eisenbarth, Shamsi, Gasser, Lopata and Beveridge2012) 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. (Reference Mattiucci, Paoletti and Webb2009).
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 (Reference Fagerholm1988).
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., Reference Flores-Barroeta, Hidalgo-Escalante and Oiea1961 [Fagerholm & Gibson, Reference Fagerholm and Gibson1987].
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 (Reference Johnston and Mawson1941); (2) Mattiucci et al. (Reference Mattiucci, Cianchi and Nascetti2003).
Remarks. Molecular data available (cytb) (2). Described as new species in Johnston & Mawson (Reference Johnston and Mawson1941), 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 (Reference Johnston1937); (2) Johnston & Mawson (Reference Johnston and Mawson1941); (3) Mawson (Reference Mawson1953): (4) Morgan et al. (Reference Morgan, Caple, Westbury and Campbell1978); (5) Cordes & O'Hara (Reference Cordes and O'Hara1979); (6) Hurst (Reference Hurst1980).
Remarks. Described as new species, C. gypsophocae and as P. hydrurgae in Johnston & Mawson (Reference Johnston and Mawson1941). Both synonymized by Johnston & Mawson (Reference Johnston and Mawson1945). Original host recorded as Gypsophoca tasmanica. Hosts recorded as Arctocephalus forsteri (5, 6). Johnston's (Reference Johnston1937) original description stated the host as A. forsteri, but this was corrected to N. cinerea in Johnston and Best (Reference Johnston and Best1942). Hurst (Reference Hurst1980) 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 Reference Baylis1937].
Hosts. Hydrurga leptonyx, Mirounga leonina.
Locality. Heard Island, Australia.
Infection site.
Stage.
References. Mawson (Reference Mawson1953).
Remarks.

Parasite name: Contracaecum sp.

Synonyms.
Hosts. Cephalorhynchus hectori.
Locality. Canterbury coast, NZ.
Infection site. Stomach.
Stage.
References. McKenzie & Blair (Reference McKenzie and Blair1983).
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 (Reference McKenzie and Blair1983).
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, Reference Johnston and Mawson1945; Myers, Reference Myers1959].
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 (Reference Johnston and Mawson1943a); (2) Johnston & Mawson (Reference Johnston and Mawson1945); (3) Johnston & Mawson (Reference Johnston and Mawson1953); (4) Mawson (Reference Mawson1953); (5) Hurst (Reference Hurst1980).
Remarks. Mawson (Reference Mawson1953) 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 (Reference Hurst1980) data include both pinniped and cetacean hosts. Johnston & Mawson (Reference Johnston and Mawson1943a) 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 (Reference Mawson1953) 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, Reference Johnston and Mawson1945; Deardorff & Overstreet, Reference Deardorff and Overstreet1981].
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 (Reference Johnston and Mawson1939); (2) Hurst (Reference Hurst1980).
Remarks. Called P. kogiae in Johnston & Mawson (Reference Johnston and Mawson1939). Intermediate fish hosts include the barracouta (Thyrsites atun), subantarctic cod (Notothaenia sp.) (Hurst, 1984), hoki (Macruronus novaezelandiae) and ling (Genypterus blacodes) (Grabda & Ślósarczyk, Reference Grabda and Ślósarczyk1981).

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, Reference Sprent1980].
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 (Reference Dexler and Freund1906); (2) Johnston & Mawson (Reference Johnston and Mawson1941); (3) Sprent (Reference Sprent1980); (4) Owen et al. (Reference Owen, Gillespie and Wilkie2012).
Remarks. Johnston & Mawson (Reference Johnston and Mawson1941) called their specimens Dujardinia halicoris. Owen et al. (Reference Owen, Gillespie and Wilkie2012) 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 (Reference McKenzie and Blair1983).
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 (Reference Mawson1953).
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 (Reference Johnston and Mawson1941).
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 (Reference Johnston and Mawson1941) listed this as Echinocephalus uncinatus Molin, 1858. However, Beveridge (Reference Beveridge1987) 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 (Reference Moravec and Justine2006), 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 (Reference Duignan2003).
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 (Reference Johnston and Mawson1941).
Remarks.

Parasite name: Crassicauda magna Johnston & Mawson, 1939

Synonyms. Crassicauda duguyi Dollfuss, 1966 [Jabbar et al., Reference Jabbar, Beveridge and Bryant2015].
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 (Reference Johnston and Mawson1939); (2) Jabbar et al. (Reference Jabbar, Beveridge and Bryant2015).
Remarks. Molecular data available (18S) (2). The phylogeny in Jabbar et al. (Reference Jabbar, Beveridge and Bryant2015) 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) Reference Duignan2000, (2) Reference Duignan2003); (3) J. Boulton in Ladds (Reference Ladds2009); (4) Stockin et al. (Reference Stockin, Duignan, Roe, Meynier, Alley and Fettermann2009); (5) Lehnert et al. (Reference Lehnert, Randhawa and Poulin2017).
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, Reference Duignan2000 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 (Reference Beveridge1980).
Remarks. Uncinaria hamiltoni sensu Beveridge (Reference Beveridge1980) 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. (Reference Haynes, Marcus, Higgins, Gongora, Gray and Šlapeta2014); (2) Marcus et al. (Reference Marcus, Higgins and Gray2014).
Remarks. Molecular data available (cox1) (1). Haynes et al. (Reference Haynes, Marcus, Higgins, Gongora, Gray and Šlapeta2014) 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 (Reference Duignan and Baker1998); (2) Duignan (Reference Duignan2000); (3) Boren (Reference Boren2005); Castinel et al. ((4) Reference Castinel, Duignan, Pomroy, Lyons, Nadler, Dailey, Wilkinson and Chilvers2006, (5) Reference Castinel, Duignan, Pomroy, Lopez-Villalobos, Gibbs, Chilvers and Wilkinson2007); (6) I. Beveridge in Ladds Reference Ladds2009); (7) Ramos et al. (Reference Ramos, Lynch, Hu, Arnould, Norman and Beveridge2013).
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. (Reference Castinel, Duignan, Pomroy, Lyons, Nadler, Dailey, Wilkinson and Chilvers2006) 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. (Reference Nadler, Lyons and Pagan2013) 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. (Reference Nadler, Lyons and Pagan2013).
Remarks. Molecular data available (ITS1&2, 28S, 12S). The phylogeny of Nadler et al. (Reference Nadler, Lyons and Pagan2013) 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. (Reference Ramos, Lynch, Hu, Arnould, Norman and Beveridge2013) 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. (Reference Nadler, Lyons and Pagan2013); (2) Ramos et al. (Reference Ramos, Lynch, Hu, Arnould, Norman and Beveridge2013).
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. (Reference Ramos, Lynch, Hu, Arnould, Norman and Beveridge2013) are the same as those of Nadler et al. (Reference Nadler, Lyons and Pagan2013) 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. (Reference Nadler, Lyons and Pagan2013); (2) Ramos et al. (Reference Ramos, Lynch, Hu, Arnould, Norman and Beveridge2013).
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. (Reference Ramos, Lynch, Hu, Arnould, Norman and Beveridge2013) are equivalent to those of Nadler et al. (Reference Nadler, Lyons and Pagan2013) 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 (Reference Duignan2000).
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 (Reference Mawson1953).
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 (Reference Dailey2009); (2) Jabbar et al. (Reference Jabbar, Mohandas and Gasser2014).
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 (Reference Nicholson and Fanning1981); (2) Boren (Reference Boren2005); (3) Norman in McFarlane et al. (Reference McFarlane, Norman, Jones, Kerry and Riddle2009); (4) Stockin et al. (Reference Stockin, Duignan, Roe, Meynier, Alley and Fettermann2009).
Remarks. Nicholson & Fanning (Reference Nicholson and Fanning1981) 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. (Reference Stockin, Duignan, Roe, Meynier, Alley and Fettermann2009) 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 (Reference Johnston and Mawson1941); (2) Tomo et al. (Reference Tomo, Kemper and Lavery2010).
Remarks. Tomo et al. (Reference Tomo, Kemper and Lavery2010) 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. (Reference Lehnert, Randhawa and Poulin2017).
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 (Reference McKenzie and Blair1983); (2) Duignan (Reference Duignan2000); (3) Lehnert et al. (Reference Lehnert, Randhawa and Poulin2017).
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, Reference Baylis and Daubney1925].
Hosts. Delphinus delphis.
Locality. Coast of SA.
Infection site. Lungs.
Stage.
References. Tomo et al. (Reference Tomo, Kemper and Lavery2010).
Remarks.

Parasite name: Skrjabinalius cryptocephalus Delyamure, 1942

Synonyms.
Hosts. Tursiops truncatus.
Locality. Otago Harbour, NZ.
Infection site. Lungs.
Stage.
References. Bowie (Reference Bowie1984).
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, Reference Bowie1984).

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 (Reference Cannon1977); (2) Cannon (Reference Cannon1977); (3) McManus et al. (Reference McManus, Wapstra, Guiler, Munday and Obendorf1984); (4) J. Boulton in Ladds (Reference Ladds2009); (5) Lehnert et al. (Reference Lehnert, Randhawa and Poulin2017).
Remarks. Molecular data available (ITS2) (5). McManus et al. (Reference McManus, Wapstra, Guiler, Munday and Obendorf1984) 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., Reference Faulkner, Measures and Whoriskey1998). 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., Reference Morell, Lehnert, IJsseldijk, Raverty, Wohlsein, Gröne and Shadwick2017). Host named G. malaenas in McManus et al. (Reference McManus, Wapstra, Guiler, Munday and Obendorf1984).

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, Reference Baylis and Daubney1925].
Hosts. Phocoena dioptrica.
Locality. Pipikaretu, Otago Peninsula, NZ.
Infection site. Sinuses and tympanic cavity.
Stage. Adult.
References. Lehnert et al. (Reference Lehnert, Randhawa and Poulin2017).
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 (Reference McColl and Obendorf1982); (2) Bowie (Reference Bowie1984); (3) Tomo et al. (Reference Tomo, Kemper and Lavery2010).
Remarks. McColl & Obendorf (Reference McColl and Obendorf1982) 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 (Reference Duignan2000).
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, Reference Delyamure1955; Dip. pacificum (Nybelin, 1931) Margolis, 1956; Dip. septentrionalis Nybelin, 1931 [WoRMS; Hernández-Orts et al., Reference Hernández-Orts, Scholz, Brabec, Kuzmina and Kuchta2015].
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 (Reference Drummond1937); (2) Johnston (Reference Johnston1937).
Remarks. Described in Drummond (Reference Drummond1937) as Dip. arctocephali n. sp., and host called A. tasmanicus. Described in Johnston (Reference Johnston1937) as Dip. arctocephalinum n. sp. and host called A. forsteri, but Johnston corrected this to N. cinerea in Johnston & Mawson (Reference Johnston and Mawson1941) and Johnston & Best (Reference Johnston and Best1942). Johnston (Reference Johnston1937) 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 (Reference Cordes and O'Hara1979).
Remarks. Cordes & O'Hara (Reference Cordes and O'Hara1979) 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 (Reference Cannon1977).
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 Reference Baylis1932; Delyamure, Reference Delyamure1955].
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 (Reference McColl and Obendorf1982); (2) Norman (Reference Norman1997); (3) Stockin et al. (Reference Stockin, Duignan, Roe, Meynier, Alley and Fettermann2009); (4) Lehnert et al. (Reference Lehnert, Randhawa and Poulin2017).
Remarks. Norman (Reference Norman1997) reported localized lymphoplasmacytic response to the cysts. These metacestodes are merocercoids of marine tapeworms infecting various marine mammals (Agustí et al., Reference Agustí, Aznar, Olson, Littlewood, Kostadinova and Raga2005; Aznar et al., Reference Aznar, Agustí, Littlewood, Raga and Olson2007); 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., Reference Aznar, Agustí, Littlewood, Raga and Olson2007; Randhawa, Reference Randhawa2011; Randhawa & Brickle, Reference Randhawa and Brickle2011). 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., Reference Klotz, Hirzmann, Bauer, Schöne, Iseringhausen, Wohlsein, Baumgärtner and Herder2018). 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, Reference Baylis1932; Delyamure, Reference Delyamure1955; Williams, Reference Williams1968].
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 (Reference Johnston and Mawson1939); (2) Morgan et al. (Reference Morgan, Caple, Westbury and Campbell1978); (3) McKenzie & Blair (Reference McKenzie and Blair1983); (4) Norman (Reference Norman1997); (5) Stockin et al. (Reference Stockin, Duignan, Roe, Meynier, Alley and Fettermann2009); (6) Lehnert et al. (Reference Lehnert, Randhawa and Poulin2017).
Remarks. Morgan et al. (Reference Morgan, Caple, Westbury and Campbell1978) 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 (Reference Delyamure1955), Dollfus (Reference Dollfus1964), Johnston & Mawson (Reference Johnston and Mawson1939) and Williams (Reference Williams1968). Molecular evidence suggests that this species belongs to the ‘Clistobothrium’ clade (Aznar et al., Reference Aznar, Agustí, Littlewood, Raga and Olson2007; Randhawa, Reference Randhawa2011; Randhawa & Brickle, Reference Randhawa and Brickle2011). 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 (Reference Bowie1984); (2) Dawson & Slooten (Reference Dawson and Slooten1990); (3) Duignan (Reference Duignan2000).
Remarks. Duignan (Reference Duignan2000) 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 (Reference Dawson and Slooten1990) 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 (Reference Cordes and O'Hara1979).
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, Reference Linton1923; 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 (Reference Krefft1871); (2) Baker et al. (Reference Baker, Duignan, Norman and Helden2001).
Remarks. Described in Krefft (Reference Krefft1871) 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 (Reference McColl and Obendorf1982); (2) Ladds (Reference Ladds2009).
Remarks. McColl & Obendorf (Reference McColl and Obendorf1982) 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 (Reference McKenzie and Blair1983); (2) Duignan (Reference Duignan2000).
Remarks. McKenzie & Blair (Reference McKenzie and Blair1983) 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 (Reference Dubois and Angel1976).
Remarks. Dubois & Angel (Reference Dubois and Angel1976) 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. (Reference Marsh, Heinsohn and Channells1984).
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. (Reference Marsh, Heinsohn and Channells1984) 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 (Reference Blair1980); (2) Olson et al. (Reference Olson, Cribb, Tkach, Bray and Littlewood2003).
Remarks. Molecular data available (18S, 28S) (2). In his redescription of I. hirudinaceus, Blair (Reference Blair1980) is adamant that it is a separate species and genus. He states that Hilmy's (Reference Hilmy1949) 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 (Reference Sey1980) 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. (Reference Olson, Cribb, Tkach, Bray and Littlewood2003).
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 (Reference Blair and Marsh1979); (2) Blanshard (Reference Blanshard, Martin and Vogelnest2001).
Remarks. Blair (Reference Blair and Marsh1979) 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. (Reference McManus, Wapstra, Guiler, Munday and Obendorf1984).
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 (Reference Blair1981).
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 (Reference Blair1981).
Remarks. Lankatrema spp. are found encapsulated in the wall of the intestine. Different species may be found in the same capsule (Blair, Reference Blair1981).

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 (Reference Blair1981); (2) Olson et al. (Reference Olson, Cribb, Tkach, Bray and Littlewood2003).
Remarks. Molecular data available (18S, 28S) (2). Blair (Reference Blair1981) 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 (Reference Blair1981).
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 (Reference Blair1981).
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 (Reference Crusz and Fernand1954) for L. mannarense. There appears to be no host response to their presence (Blair, Reference Blair1981).

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 (Reference Blair1981).
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 (Reference Blair1981).
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, Reference Blair1981].
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 (Reference Dexler and Freund1906); (2) Johnston (Reference Johnston1913); (3) Blair (Reference Blair1981); (4) Olson et al. (Reference Olson, Cribb, Tkach, Bray and Littlewood2003).
Remarks. Molecular data available (18S, 28S) (4). Dexler and Freund (Reference Dexler and Freund1906) and Johnston (Reference Johnston1913) 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, Reference Blair1981].
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 (Reference von Linstow1904); (2) Blair (Reference Blair1981); (3) Blanshard (Reference Blanshard, Martin and Vogelnest2001); (4) Owen et al. (Reference Owen, Gillespie and Wilkie2012).
Remarks. Described as Opisthotrema pulmonale by von Linstow (Reference von Linstow1904). Called Cochleotrema indicum in Blanshard (Reference Blanshard, Martin and Vogelnest2001) and Owen et al. (Reference Owen, Gillespie and Wilkie2012). 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 (Reference Blair1981).
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 (Reference Johnston1913); (2) Blair (Reference Blair1981); (3) Olson et al. (Reference Olson, Cribb, Tkach, Bray and Littlewood2003).
Remarks. Molecular data available (18S, 28S) (3). Johnston (Reference Johnston1913) 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 (Reference Blair1981).
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 (Reference Blair1981); (2) Olson et al. (Reference Olson, Cribb, Tkach, Bray and Littlewood2003).
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, Reference Felix2013].
Hosts. Delphinus delphis.
Locality. Bethell's Beach, Auckland, NZ.
Infection site. Liver.
Stage. Adult.
References. Lehnert et al. (Reference Lehnert, Randhawa and Poulin2017).
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, Reference Felix2013].
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 (Reference Cordes and O'Hara1979); (2) Lehnert et al. (Reference Lehnert, Randhawa and Poulin2017).
Remarks. Cordes and O'Hara (Reference Cordes and O'Hara1979) 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 (Reference McColl and Obendorf1982); (2) Hutton et al. (Reference Hutton, Blair, Slooten and Dawson1987); (3) Duignan (Reference Duignan2000).
Remarks. Both Duignan (Reference Duignan2000) and Hutton et al. (Reference Hutton, Blair, Slooten and Dawson1987) 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 (Reference McColl and Obendorf1982).
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, Reference Price1932; Marigo et al., Reference Marigo, Paulo Vicente, Schifino Valente, Measures and Portes Santos2008].
Hosts. Tursiops truncatus.
Locality. Otago Harbour, NZ.
Infection site. Small intestine.
Stage. Adult.
References. Bowie (Reference Bowie1984).
Remarks.

Parasite name: Synthesium sp.

Synonyms.
Hosts. Neophoca cinerea.
Locality. St. Vincent Gulf, SA.
Infection site. Intestine.
Stage.
References. Dubois & Angel (Reference Dubois and Angel1976).
Remarks. Genus Hadwenius in this paper. This genus was synonymized with Synthesium by Gibson (Reference Gibson, Jones, Bray and Gibson2005).

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 (Reference Dubois and Angel1976).
Remarks. This species was originally described by Pearson (Reference Pearson1973) from the Caspian tern – as mentioned in the passing reference to G. angelae in Dubois and Angel (Reference Dubois and Angel1976). There is a perpetuated error in the citation of this species and the referencing of this paper. Note that both Dailey et al. (Reference Dailey, Demaree and Critchfield2002) and Hernández-Orts et al. (Reference Hernández-Orts, Montero, Crespo, García, Raga and Aznar2012) list Dubois and Angel (Reference Dubois and Angel1976) with the title ‘Galactosomum angelae Pearson Reference Pearson1973 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 Reference Dubois and Mahon1959. The corrected reference for the Dubois and Angel (Reference Dubois and Angel1976) 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 (Reference Cordes and O'Hara1979).
Remarks.

Parasite name: Stictodora diplacantha Johnston, 1942

Synonyms.
Hosts. Neophoca cinerea.
Locality. St. Vincent Gulf, SA.
Infection site. Intestine.
Stage.
References. Dubois & Angel (Reference Dubois and Angel1976).
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, Reference Boren2005); ‘lung worms’ in Delphinus delphis from Mooloolaba Beach, QLD (Greenland & Limpus, Reference Greenland and Limpus2008); ‘cestodes’ in Tursiops truncatus from Otago Harbour, NZ (Bowie, Reference Bowie1984) and Globicephala melas from McIntyres Beach, TAS (McManus et al., Reference McManus, Wapstra, Guiler, Munday and Obendorf1984); and ‘trematodes’ in A. a. forsteri from Ohau Point, NZ (Boren, Reference Boren2005), Lagenodelphis hosei from Corio Bay, VIC (McColl & Obendorf, Reference McColl and Obendorf1982) and Dugong dugon from Wallum Creek, QLD (Dexler & Freund, Reference Dexler and Freund1906).

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, Reference Domrow1962].
Hosts. Mirounga leonine.
Locality. Macquarie Island, TAS.
Infection site. Respiratory tract.
Stage. Adult, larvae.
References. Domrow (Reference Domrow1979).
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, Reference Domrow1974].
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 (Reference Tubb1937); (2) Domrow (Reference Domrow1963); (3) Seawright (Reference Seawright1964); (4) Domrow (Reference Domrow1974); (5) Marlow (Reference Marlow1975); (6) Nicholson & Fanning (Reference Nicholson and Fanning1981).
Remarks. Tubb (Reference Tubb1937) 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 (Reference Tubb1937) report is named A. tasmanicus. Seawright (Reference Seawright1964) 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, Reference Domrow1974].
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 (Reference Domrow1974); (2) Nicholson & Fanning (Reference Nicholson and Fanning1981).
Remarks. Domrow (Reference Domrow1974) 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 (Reference Dailey and Nutting1980).
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, Reference Leonardi, Crespo, Raga and Fernández2009].
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 (Reference Enderlein1906); (2) Clay (Reference Clay1964); (3) Horning et al. (Reference Horning, Palma and Pilgrim1980); (4) McIntosh & Murray (Reference McIntosh and Murray2007); (5) Leonardi et al. (Reference Leonardi, Crespo, Raga and Fernández2009); (6) Marcus et al. (Reference Marcus, Higgins and Gray2015).
Remarks. Leonardi et al. (Reference Leonardi, Crespo, Raga and Fernández2009) 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 (Reference Leonardi and Palma2013) suggest that the existing records may represent a complex of cryptic species. Host is called Arctocephalus forsteri in Leonardi et al. (Reference Leonardi, Crespo, Raga and Fernández2009).

Parasite name: Antarctophthirus ogmorhini Enderlein, 1906

Synonyms. Echinophthirius setosus Rothschild, 1902 (nec E. setosus Burmeister 1838 = E. phocae Lucas, 1834) [Harrison, Reference Harrison and Johnston1937].
Hosts. Hydrurga leptonix (1, 2).
Locality. (1, 2) Macquarie Island, TAS.
Infection site. Fur/skin, body.
Stage. Adults.
References. (1) Harrison (Reference Harrison and Johnston1937); (2) Watson (Reference Watson1967).
Remarks. Leonardi et al. (Reference Leonardi, Poljak, Carlini, Galliari, Bobinac, Santos, Márquez and Negrete2014) erected Antarctophthirius carlinii for specimens from the Weddell seal (Leptonychotes weddelli (Lesson, 1826)). Antarctophthirius carlinii was, in turn, listed by Palma (Reference Palma2017) 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 (Reference Harrison and Johnston1937); (2) Murray & Nicholls (Reference Murray and Nicholls1965); (3) Watson (Reference Watson1967); (4) Lowry et al. (Reference Lowry, Horning, Poore and Ricker1978); (5) Horning et al. (Reference Horning, Palma and Pilgrim1980); (6) Palma & Horning (Reference Palma and Horning2002).
Remarks. Host named Macrorhinus leoninus in Harrison (Reference Harrison and Johnston1937). 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, Reference Pilsbry1916].
Hosts. Dugong dugon (1, 2).
Locality. (1) Sydney Harbour, Australia; (2) Magnetic Island, QLD.
Infection site. Skin, ventral surface.
Stage.
References. (1) Marlow (Reference Marlow1962); (2) Zann & Harker (Reference Zann and Harker1978).
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. (Reference Shaughnessy, Kemper, Stemmer and McKenzie2014).
Remarks. Shaughnessy et al. (Reference Shaughnessy, Kemper, Stemmer and McKenzie2014) 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 (Reference Dexler and Freund1906).
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 (Reference Dexler and Freund1906).
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 (Reference Leung1965); (2) Sedlak-Weinstein (Reference Sedlak-Weinstein1991).
Remarks. This species has been found to be highly specific in its choice of host, the humpback whale, M. novaeangliae (Carvalho et al., Reference Carvalho, Bevilaqua and Iñiguez2010), 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 (Reference Leung1965).
Remarks. In his paper, Leung (Reference Leung1965) 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, Reference Leung1967].
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 (Reference Leung1965); (2) Berzin & Vlasova (Reference Berzin and Vlasova1982); (3) Sedlak-Weinstein (Reference Sedlak-Weinstein1991).
Remarks. Leung (Reference Leung1965) names the host G. malaena. Berzin and Vlasova (Reference Berzin and Vlasova1982) 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 (Reference Sedlak-Weinstein1992b).
Remarks. Host named G. malaena in Sedlak-Weinstein (Reference Sedlak-Weinstein1992b).

Parasite name: Isocyamus kogiae Sedlak-Weinstein, 1992

Synonyms.
Hosts. Kogia breviceps.
Locality. Moreton Island, QLD.
Infection site. Skin.
Stage. Adults.
References. Sedlak-Weinstein (Reference Sedlak-Weinstein1992a).
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 (Reference Sedlak-Weinstein1991).
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 (Reference Lincoln and Hurley1980); (2) Lehnert et al. (Reference Lehnert, Randhawa and Poulin2017).
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 (Reference Sedlak-Weinstein1991).
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 (Reference Sedlak-Weinstein1991).
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. (Reference Hill, Fraser and Prior1997); (2) Morgan et al. (Reference Morgan, Xiao, Hill, O'Donoghue, Limor, Lal and Thompson2000).
Remarks. Hill et al. (Reference Hill, Fraser and Prior1997) 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. (Reference Morgan, Xiao, Hill, O'Donoghue, Limor, Lal and Thompson2000) provide molecular data leading to identification as C. parvum ‘human’ genotype. This genotype was later described as C. hominis by Morgan-Ryan et al. (Reference Morgan-Ryan, Fall and Ward2002).

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 (Reference Jardine and Dubey2002); (2) Bowater et al. (Reference Bowater, Norton, Johnson, Hill, O'Donoghue and Prior2003); (3) Owen et al. (Reference Owen, Gillespie and Wilkie2012); (4) Donahoe et al. (Reference Donahoe, Rose and Šlapeta2014); (5) Roe et al. (Reference Roe, Michael, Fyfe, Burrows, Hunter and Howe2017).
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, Reference Jardine and Dubey2002); systemic toxoplasmosis, molecular data (Donahoe et al., Reference Donahoe, Rose and Šlapeta2014); encephalitis associated with T. gondii cysts, emaciation, ataxia in one live stranded animal (Bowater et al., Reference Bowater, Norton, Johnson, Hill, O'Donoghue and Prior2003). Host called Arctocephalus forsteri in Donahoe et al. (Reference Donahoe, Rose and Šlapeta2014).

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. (Reference Delport, Asher, Beaumont, Webster, Harcourt and Power2014).
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. ( Reference Nadler, Lyons and Pagan2013)
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. (Reference Nadler, Lyons and Pagan2013)
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. (Reference Nadler, Lyons and Pagan2013)
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., Reference Lasek-Nesselquist, Welch and Sogin2010), 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., Reference Mattiucci, Cipriani, Webb, Paoletti, Marcer, Bellisario, Gibson and Nascetti2014; Klotz et al., Reference Klotz, Hirzmann, Bauer, Schöne, Iseringhausen, Wohlsein, Baumgärtner and Herder2018), 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., Reference Jabbar, Beveridge and Bryant2015), as well as provide insights on the phylogeny of parasite species (Hernández-Orts et al., Reference Hernández-Orts, Smales, Pinacho-Pinacho, García-Varela and Presswell2017).

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., Reference Fayer, Dubey and Lindsay2004; Applebee et al., Reference Applebee, Thompson and Olson2005; Grilo et al., Reference Grilo, Gomes, Wohlsein, de Carvalho, Siebert and Lehnert2018).

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, Reference Yamasaki and Kuramochi2009; Shamsi & Butcher, Reference Shamsi and Butcher2011). 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., Reference Cipriani, Acerra, Bellisario, Sbaraglia, Cheleschi, Nascetti and Mattiucci2016).

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, Reference Presswell and Lagrue2016), 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., Reference Lehnert, Randhawa and Poulin2017). 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., Reference Van Bressem, Raga and Di Guardo2009). Studies combining ecotoxicological analyses with systematic monitoring of parasite prevalence and impact (Lehnert et al., Reference Lehnert, Desforges, Das, Siebert, Fossi and Panti2018) 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., Reference Aznar, Fernández, Balbuena and Murray2010; Poulin et al., Reference Poulin, Blasco-Costa and Randhawa2016). 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.

Author ORCIDs

K. Lehnert, 0000-0001-8938-3340

Acknowledgements

Our grateful thanks to Ian Beveridge and one anonymous reviewer who took great trouble to read this work carefully and give detailed and helpful comments. We would like to thank the University of Otago Library for helping to find many of the more obscure references. We also made much use of the Biodiversity Heritage Library (www.biodiversitylibrary.org) whom we thank for their continued commitment to making older texts available.

Financial support

Kristina Lehnert was partly funded by the Volkswagen Foundation under the ‘Research in Museums’ (AZ 89911) programme. Bronwen Presswell was indirectly funded by a grant from the Marsden Fund (Royal Society of New Zealand) to Robert Poulin. Grant No: UOO1718.

Conflict of interest

None.

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Article contents

Checklist of marine mammal parasites in New Zealand and Australian waters

Abstract

Keywords

Introduction

Material and methods

Results

Parasite–host list

Host–parasite list

Host taxon: Pinnipedia

Host taxon: Cetacea

Host taxon: Sirenia

Discussion

Author ORCIDs

Acknowledgements

Financial support

Conflict of interest

References

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