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Diseases of Echinodermata. 111. Agents Metazoans (Annelida to Pisces)

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Diseases of Echinodermata. 111. Agents Metazoans (Annelida to Pisces) l DISEASES OF AQUATIC ORGANISMS Vol. 3: 59-83, 1987 Published October 14 Dis. aquat. Org. REVIEW Diseases of Echinodermata. 111. Agents metazoans (Annelida to Pisces) Michel Jangoux Laboratoire de Biologie marine (CP 160),Universite Libre de Bruxelles, Ave F. D. Roosevelt 50, B-1050 Bruxelles, Belgium ABSTRACT. Parasitic myzostomids mostly infest crinoids but a few are known from asteroids and ophiuroids; they are either galhcole, cysticole or endoparasitic. Although many copepods have been said to be ectoparasitic on echinoderms, this has been proven for only a few species. Some copepods are known to induce gall formation in the spines of echinoids or in the body wall of ophiuroids; others have been found to infest either the bursae or gonads of ophiuroids. Ascothoracid cirripeds were reported either as ectoparasites on cnnoids and ophiuroids or as endoparasites In the body cavlty of asteroids and spatangoid echinoids. Echinoderm castration by copepods and ascothoracids was reported several times in the literature. Parasitic crabs nlostly occur on or in echino~ds.Ectoparasitic crabs often exert major effects and may kill their host; gut-inhabiting species may produce conspicuous host deformation. Some species of carapid fishes are known to live either temporarily or permanently in the body cavity of holothuroids and asteroids Carapid infestations do not seem to seriously affect the echlnoderms except for species permanently inhabihng the host's coelom. Parasitic associations between echinoderms and polychaetes, tardigrads, barnacles, amphipods, tanaidaceans, acarians, pycnogonids and insects have been casually reported in the literature. INTRODUCTION ous species are known to live ectocommensally on echinoderms, only 3 cases of parasitism have been The present paper is the third of a series of 4 that reported with polychaetes. According to Monticelli review the dlseases of Echinodermata. It considers the (1892) the eunicid Ophryotrocha puerilis occurs In the disease agents belonging to the Annelida (Polychaeta coelomic cavity of the holothuroid Ocnus planci from and Myzostomida), Tardigrada, Crustacea (Copepoda, Naples (Italy). Ganapati & Radhakrishna (1962) noted Cirripedia and Malacostraca), Arachnida, Pycno- that 50% of the holothuroid Molpadia sp. investigated gonida, Insecta and Pisces. (Other metazoan agents harbored the small hesionid Ancistrocyllis sp. either in have been considered in Part 11; Jangoux 1987b). As the digestive tract or respiratory trees. The only case of discussed in Part I (Jangoux 1987a),I have adopted the unequivocal parasitism is that of the lumbrinereid definition of parasites proposed by Kinne (1980, p. 19) Ophiuricola cynips which forms myzostomid-like galls and used it in a vely broad sense, considering disease at the base of the arms of the deep-sea ophuroid agents (parasites sensu lato) to represent any lund of Ophioglypha tumulosa (Ludwig 1905). According to harmful associate which affects, if even slightly, the Ludwig, the galls are rather large and partly protrude echinoderm's tissues or internal fluids (Le, coelomic into the host's coelomic cavlty. and hemal fluids). Agents: Annelida, Myzostomida DISEASES CAUSED BY METAZOANS The class Myzostomida (about 110 described Agents: Annelida, Polychaeta species) occupies a peculiar place among echinoderm symblotes. They are aberrant annelids with a small Symbiotic polychaetes were reviewed by Paris (1955) flattened body several mm in length. Their most extra- and Clark (1956), both authors stating that these ordnary feature is their intimate association with polychaetes rarely affect echinoderms. While numer- echinoderms; in fact there are no known free-living O Inter-Research/Pnnted in F. R. Germany Dis. aquat. Org. 3: 59-83, 1987 Table 1. Parasitic myzostomids from echinoderms (compiled from the sources indicated). Myzostomid classificahon and species names according to Jagersten (1940). Hosts: A, asteroid; C, crinoid; 0, ophiuroid. Records of conspicuous deformations caused by unidentified myzostomids were reported on several occasions (e.g. Carpenter 1889 for Acitinometra notata). Speel & Dearborn (1983) noted that each of the 96 individuals of Promochocrinus kerguelensis they observed harbored 1 to 3 myzos- tomid cysts Host Location on/in host Remarks Geographical area Source I. Proboscidea Myzostornum beardi Perissornetra flexilis Galls on arms Only 1 worm NE Indian Ocean Graff (1887) (C) gall-' (Arafura Sea) Endoxocnnus Galls at base of arms Only 1 worm NW Paclfic Wheeler (1896) alternicirrus (C) gall-' (SPhilippines) Myzostorn um &Ietacrinusintemptus Galls on arms Indian Ocean (Bay of Wheeler (1896) cryplopodium (C) Bengal?) Myzostomurn Endoxocnnus Galls in pinnules 2 worms (6,Q) NW Pacific (SE Graff (1884) deformator alternicirrus (C) gall-' Philipp~nes) rllyzostornum erernita Metacrinus rnoseleyl Gallsat base of arms Only 1 worms NW Paclfic (SE Wheeler (1896) (C) and pinnules gall-' Philippines) Myzostom urn Endoxocn'n us alter- Slight galls extending 1 to 3 worms NW Pacific (SE Graff (1884) pen tacrini njcirrus (C) into 3 to 6 arm seg- gall-' Philippines) ments Myzostornum Pachylometra inae- Conspicuous galls ex- 2 worms (d , Q) Trop~calW Pacific (SE Graff (1884) ten uispin urn quabs, Perissornetra tending into 2 to 3 arms gall-'; several Phhppines, Fiji lslands flexilis, Charitornetra segments galls on each and Kermadec Island) basicurva, Charito- infested host metra incisa (C) Myzostomum Pachylometra inae- Galls in pinnules 2 worms (6,Q) Tropical W Pacific (Fij~ Graff (1884 willernoes1 qual~s,Perissorn etra gall-') and Kermadec 1887) flexllrs, Charitometra Islands); NE Indian basicurva (C) Ocean (Arafura Sea) 11. Pharyngidea Asteriomyzostornum Sclerasterias neglecta. Hypertrophied pyloric 1 to 3 worms Mediterranean sea Marenzeller asteriae Sclerastenasrichard~ caeca (proximalpart of infested (1895a,b), Stum- (A) the caeca) asteroid-' mer-Traunfels (1903) Asteriomyzostornum Tosia leptoceramus (A) Coelomic cavity, Tropical E Pacific (off Wheeler (1904) fish eri loosely attached South California) to body wall Cystimyzostom urn Metactinusrotundus Subcutaneous cysts on 1 worm cyst-'; N. Pacific (Japan: McClendon clarki (C) underside of arms up to 7 cysts Sagami Sea) (1906) host-' Cystim yzostorn um Anthornetra adriani. Subcutaneous 2worms(d, P) Cosmopolitan (Carib- Graff (1884). cysticolum Arnphirnetra discoidea, calcified cysts on cyst-'; no bean; Red Sea; Aru McClendon Comactinia upper side of arms more than 1 Islands; Eastern coast (1906),Rem- meridionalis (C) cyst on of Japan) scheid(l916), each host arm Fishelson (1973, 1974) Cystimyzostorn um Tropiometra macro- Subcutaneous cyst on 1 to 2 worms Indo W Pacific (Sagami Hara & Okada ijirnai discus, Capillaster calyx (uppersurface) cyst-' Sea, Japan; Red Sea) (1921).Fishelson rnultiradiatus (C) (1973, 1974) Neocoma tella Subcutaneous calcified 2worms(6,Q) Circumtropical Graff (1884) pulchella, Adelometra cysts at arm base cyst-' (Papua;Barbados) angus tiradia (C) (upper surface) Cystimyzostom urn Horaeometra duplex, Subcutaneousstalked 2 worms (d. Q ) Circumtropical Graff (1884) m urra yi Stirernetra breviradia, and calcified cystson cyst-' (Papua;Kermadec Adelornetra angust- calyx (uppersurface) Islands; Barbados) iradja (C) Cystimyzostom urn Comanthina Subcutaneous cyst on 1 worm cyst-' W Pacific (Philipp~nes: Graff (1887) platypus schlegeli (C) calyx (upper surface) off Samboangan) Cystimyzostornurn Metacrinus rotundus Cysts onarms l worm cyst-' NW Pacific (Japan: Hara & Okada rubusturn (( '1 Sagam1 Sea) (1921) Cystimyzostornurn Zygornetra rnertoni (C) Juvenlle in cysts on l worm cyst-' NtVInd~anOcean (Aru Remscheid taeniatum pinnules, adults Islands) (1916) free-living Jangoux D~seasesof Echinodern~ata:agents metazoans 6 1 Table l (continued) Myzostomid Host Locat~ononlin host Remarks Geographical area Source Coman thus japonicus Gonads (genital Sea of Japan Okada (1933) (C) p~nnules) Amphimetra disco~dea Coelomic cavity of Only 2 indl- NW Indian Ocean (Aru Remscheid (C) arms v~dualsknown Islands) (1916) Protom yzostomum Astrocladus coniferus Encysted in gonads NW Paclfic (Sagami Fedotov (1925) astrocladi (0) Sea) Prolomyzostomum Gorgonocephalus arc- Bursae and gonads 10 to 20 North Sea (Scandina- Fedotov (1912. polynephris ticus, Gorgonocephalus worms wan coast), Barents 1914. 1916), eucnen~js,Gorgono- host-' Sea Barel & cephalus caput- Kramers (1977) Prolomyzostornun~ Gorgonocephalus sp Bursae and gonads NW Pacific (Sagam1 Okada (1922) sagamiense (0) Sea) Pulvinom)~zostomurn Leptometra phalan- D~gestivetract Infestation Mediterranean (Ban- Prouho (1892). pulvlnar glum, Antedon level: 10 to yuls; Naples); NE At- Wheeler (1896). bifida (C) 20 % lantic (Roscoff) Jagersten (Jagersten) (19401, Barel & Kramers (1977) myzostomids, nor is there any species associated with Gallicole species belong to the genus Myzostornurn. other phyla. Myzostomids n~ostlyinfest crinoids, but a They dig into the dermal tlssue of crinoids arms or few are known from asteroids and ophiuroids. Myzos- pinnules (Fig. lA, B) and build more or less spacious tomids are highly differentiated, both morphologically intradermal cavities, always located under skeletal and ecologically. Their almost obligatory association ossicles. The cavities sometimes are very complicated, with crinoids (they even infested now extinct crinoids; with internal partitions (e.g. in Myzostomum tenuis- e.g. Meyer & Ausich 1983, Arendt 1985) suggests that pjnum; Graff 1884). Myzostomid galls often harbor a they are an ancient group
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