Diseases of Echinodermata. Ill Agents metazoans (Annelida to Pisces)

Michel  Jangoux

The association of scyphomedusae with invertebrates has been long known in the literature; especially with hyperiids amphipods. The association of echinoderms with jellyfish is not common and rarely recorded. We reported the association of the ophiuroid Ophiocnemis marmorata with the rhizostome scyphomedusa Rhopilema hispidum collected in Vellar estuary (on the southeast coast of India). O. marmorata is supposed to be a filter feeding ophiuroid, quite common in soft bottom of shallow waters. The brittle stars possibly seek for food supply, shelter and protection through the association.

Very little information is reported for parasites of cnidarians, therefore, the present work aimed to investigate parasitic infections in one of the most widespread jellyfish in the Mediterranean Sea, Rhizostoma pulmo. The goals were to determine prevalence and intensity of parasites in R. pulmo, identify the species involved through morphological and molecular analysis, test whether infection parameters differ in different body parts and in relation to jellyfish size. 58 individuals were collected, 100% of them infected with digenean metacercariae. Intensity varied between 18.7 ± 6.7 per individual in 0–2 cm diameter jellyfish up to 505 ± 50.6 in 14 cm ones. Morphological and molecular analyses suggest that the metacercariae belonged to the family Lepocreadiidae and could be possibly assigned to the genus Clavogalea. Prevalence values of 100% suggest that R. pulmo is an important intermediate host in the life cycle of lepocreadiids in the region. Our findings also support the hypot...

l Vol. 3: 59-83, 1987 DISEASES O F AQUATIC ORGANISMS Dis. aquat. Org. Published October 14 REVIEW Diseases of Echinodermata. 111. Agents metazoans (Annelida to Pisces) Michel Jangoux Laboratoire d e Biologie marine (CP 160), Universite Libre d e Bruxelles, Ave F. D. Roosevelt 50, B-1050 Bruxelles, Belgium ABSTRACT. Parasitic myzostomids mostly infest crinoids but a few are known from asteroids a n d ophiuroids; they are either galhcole, cysticole or endoparasitic. Although many copepods have been said to b e 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 a n d ascothoracids w a s reported several times in the literature. Parasitic crabs nlostly occur on or in e c h i n o ~ d s 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 a n d polychaetes, tardigrads, barnacles, amphipods, tanaidaceans, acarians, pycnogonids a n d insects have been casually reported in the literature. INTRODUCTION The present paper is the third of a series of 4 that review the dlseases of Echinodermata. It considers the disease agents belonging to the Annelida (Polychaeta and Myzostomida), Tardigrada, Crustacea (Copepoda, Cirripedia and Malacostraca), Arachnida, Pycnogonida, Insecta and Pisces. (Other metazoan agents have been considered in Part 11; Jangoux 1987b). As discussed in Part I (Jangoux 1987a),I have adopted the definition of parasites proposed by Kinne (1980, p. 19) and used it in a vely broad sense, considering disease agents (parasites sensu lato) to represent any lund of harmful associate which affects, if even slightly, the echinoderm's tissues or internal fluids (Le, coelomic and hemal fluids). ous species are known to live ectocommensally on echinoderms, only 3 cases of parasitism have been reported with polychaetes. According to Monticelli (1892) the eunicid Ophryotrocha puerilis occurs In the coelomic cavity of the holothuroid Ocnus planci from Naples (Italy). Ganapati & Radhakrishna (1962) noted that 50% of the holothuroid Molpadia sp. investigated harbored the small hesionid Ancistrocyllis sp. either in the digestive tract or respiratory trees. The only case of unequivocal parasitism is that of the lumbrinereid Ophiuricola cynips which forms myzostomid-like galls at the base of the arms of the deep-sea ophuroid Ophioglypha tumulosa (Ludwig 1905). According to Ludwig, the galls are rather large and partly protrude into the host's coelomic cavlty. Agents: Annelida, Myzostomida DISEASES CAUSED BY METAZOANS Agents: Annelida, Polychaeta Symbiotic polychaetes were reviewed by Paris (1955) and Clark (1956), both authors stating that these polychaetes rarely affect echinoderms. While numerO Inter-Research/Pnnted in F. R. Germany The class Myzostomida (about 110 described species) occupies a peculiar place among echinoderm symblotes. They are aberrant annelids with a small flattened body several mm in length. Their most extrao r d n a r y feature is their intimate association with echinoderms; in fact there are no known free-living Dis. aquat. Org. 3: 59-83, 1987 Table 1. Parasitic myzostomids from echinoderms (compiled from the sources indicated). Myzostomid classificahon a n d species names according to Jagersten (1940). Hosts: A, asteroid; C, crinoid; 0, ophiuroid. Records of conspicuous deformations caused by unidentified myzostomids w e r e 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 myzostomid cysts Host Location on/in host Remarks Geographical area Source I. Proboscidea Myzostornum beardi Myzostorn um cryplopodium Myzostomurn deformator rllyzostornum erernita Myzostom urn pen tacrini Myzostornum ten uispin urn Myzostomum willernoes1 11. Pharyngidea Asteriomyzostornum asteriae Perissornetra flexilis (C) Endoxocnnus alternicirrus (C) &Ietacrinusintemptus (C) Endoxocnnus alternicirrus (C) Metacrinus rnoseleyl (C) Endoxocn'n us alternjcirrus (C) Pachylometra inaequabs, Perissornetra flexilis, Charitornetra basicurva, Charitometra incisa (C) Pachylometra inaequal~s,Perissorn etra flexllrs, Charitometra basicurva (C) Sclerasterias neglecta. Sclerastenasrichard~ (A) Asteriomyzostornum fish eri Cystimyzostom urn clarki Galls on arms Galls at base of arms Galls on arms Galls in pinnules Gallsat base of arms and pinnules Slight galls extending into 3 to 6 arm segments Conspicuous galls extending into 2 to 3 arms segments Wheeler (1896) Wheeler (1896) Graff (1884) Wheeler (1896) Graff (1884) Graff (1884) Hypertrophied pyloric caeca (proximalpart of the caeca) 1 to 3 worms infested asteroid-' Tosia leptoceramus (A) Coelomic cavity, loosely attached to body wall Metactinusrotundus Subcutaneous cysts on underside of arms (C) Cystimyzostorn um ijirnai Tropiometra macrodiscus, Capillaster rnultiradiatus (C) Neocoma tella pulchella, Adelometra angus tiradia (C) Horaeometra duplex, Stirernetra breviradia, Adelornetra angustiradja (C) Comanthina schlegeli (C) Metacrinus rotundus (( 2 worms ( d , Q ) Trop~calW Pacific (SE gall-'; several Phhppines, Fiji lslands galls on each and Kermadec Island) infested host Graff (1887) Q ) Tropical W Pacific (Fij~ Graff (1884 2 worms (6, gall-') and Kermadec 1887) Islands); NE Indian Ocean (Arafura Sea) Anthornetra adriani. Subcutaneous Arnphirnetra discoidea, calcified cysts on Comactinia upper side of arms meridionalis (C) Cystimyzostom urn platypus Cystimyzostornurn rubusturn Cystimyzostornurn taeniatum NE Indian Ocean (Arafura Sea) NW Paclfic (SPhilippines) Indian Ocean (Bay of Bengal?) 2 worms ( 6 , Q ) NW Pacific (SE gall-' Philipp~nes) Only 1 worms NW Paclfic (SE gall-' Philippines) 1 to 3 worms NW Pacific (SE gall-' Philippines) Galls in pinnules Cystim yzostorn um cysticolum Cystimyzostom urn m urra yi Only 1 worm gall-' Only 1 worm gall-' Mediterranean sea Tropical E Pacific (off South California) 1 worm cyst-'; up to 7 cysts host-' 2worms(d, P) cyst-'; no more than 1 cyst on each host arm N. Pacific (Japan: Sagami Sea) McClendon (1906) Cosmopolitan (Caribbean; Red Sea; Aru Islands; Eastern coast of Japan) Graff (1884). McClendon (1906),Remscheid(l916), Fishelson (1973, 1974) Hara & Okada (1921).Fishelson (1973, 1974) Graff (1884) Subcutaneous cyst on calyx (uppersurface) 1 to 2 worms cyst-' Subcutaneous calcified cysts at arm base (upper surface) Subcutaneousstalked and calcified cystson calyx (upper surface) 2worms(6,Q) Circumtropical cyst-' (Papua;Barbados) Subcutaneous cyst on calyx (upper surface) Cysts onarms Indo W Pacific (Sagami Sea, Japan; Red Sea) 2 worms ( d . Q ) Circumtropical cyst-' (Papua;Kermadec Islands; Barbados) Graff (1884) 1 worm cyst-' Graff (1887) l worm cyst-' '1 Zygornetra rnertoni (C) Juvenlle in cysts on pinnules, adults free-living Marenzeller (1895a,b ) ,Stummer-Traunfels (1903) Wheeler (1904) l worm cyst-' W Pacific (Philipp~nes: off Samboangan) NW Pacific (Japan: Sagam1 Sea) NtVInd~anOcean (Aru Islands) Hara & Okada (1921) Remscheid (1916) 61 J a n g o u x D ~ s e a s e sof E c h i n o d e r n ~ a t a :a g e n t s metazoans Table l (continued) Myzostomid Host Locat~ononlin host Gonads (genital p~nnules) Amphimetra disco~dea Coelomic cavity of arms (C) Astrocladus coniferus Encysted in gonads (0) Gorgonocephalus arc- Bursae and gonads ticus, Gorgonocephalus eucnen~js,Gorgonocephalus caput- Remarks Geographical area Sea of Japan Coman thus japonicus Source Okada (1933) (C) Protom yzostomum astrocladi Prolomyzostomum polynephris Prolomyzostornun~ sagamiense Pulvinom)~zostomurn pulvlnar Gorgonocephalus sp (0) Leptometra phalanglum, Antedon bifida (C) Only 2 indlN W Indian Ocean (Aru v ~ d u a l sknown Islands) NW Paclfic (Sagami Sea) 10 to 20 North Sea (Scandinaworms wan coast), Barents host-' Sea Bursae and gonads D~gestivetract myzostomids, nor is there any species associated with other phyla. Myzostomids n~ostlyinfest crinoids, but a few are known from asteroids and ophiuroids. Myzostomids are highly differentiated, both morphologically and ecologically. Their almost obligatory association with crinoids (they even infested now extinct crinoids; e.g. Meyer & Ausich 1983, Arendt 1985) suggests that they are a n ancient group which evolved together with clinoids. Myzostomids may be considered a unique case of 'symbiogenesis' involving a whole class of organisms ( e . g . Clark 1921, Stummer-Traunfels 1926, Prenant 1959). About 25 species of myzostomids have been recorded as echinoderm parasites (Table l ) , the remaining species being frequently referred to 'free-living'. However, these latter species are ectocommensals which generally live unattached on the echinoderm body surface. As pointed out by Stummer-Traunfels (1926), 3 types of parasitic myzostomids may be distinguished depending on whether they are gahcole, cysticole or endoparasltic. Endoparasitic species feed on the host's tissues, while cysticole and gallicole species are suspension feeders which divert the water current produced by the host's ambulacra (galls and cysts are most often located near the ambulacral grooves and always have 2 apertures allowing passage of directional watercurrents) (Prenant 1959). With rather few exceptions (e.g. Cystomyzostomum ijimai and Mesomyzostoma katoi; respectively Fishelson 1974, Okada 1933),parasitic myzostomids mostly infest bathyal echinoderms (in 200 to 1200 m water depth; Stummer-Traunfels 1926). This fact explains why these parasitoses have been so anecdotically documented. Infestation level: 10 to 20 % (Jagersten) NW Pacific (Sagam1 Sea) Mediterranean (Banyuls; Naples); NE Atlantic (Roscoff) Remscheid (1916) Fedotov (1925) Fedotov (1912. 1914. 1916), Barel & Kramers (1977) Okada (1922) Prouho (1892). Wheeler (1896). Jagersten (19401, Barel & Kramers (1977) Gallicole species belong to the genus Myzostornurn. They dig into the dermal tlssue of crinoids arms or pinnules (Fig. l A , B) a n d build more or less spacious intradermal cavities, always located under skeletal ossicles. The cavities sometimes are very complicated, with internal partitions (e.g. in Myzostomum tenuispjnum; Graff 1884). Myzostomid galls often harbor a pair of individuals. Gallicole species were termed 'deformative Arten' by Stummer-Traunfels (1926); they induce conspicuous deformation of the host's ossicles which considerably enlarge and adjust their shape to that of the myzostomid's shelter (Graff 1884, Wheeler 1896, Stummer-Traunfels 1926). Cysticole species belong to the genus Cystirnyzostomum; they behave differently in that they build remarkable stalked or unstalked cysts which are always subcutaneous (i.e. located outside the host's skeleton) and protrude into the external medium (Fig. 1C). In many cases the cyst wall is reinforced by a pavement of minute skeletal plates (Graff 1884, Stummer-Traunfels 1926). Both gallicole and cysticole myzostomids induce major host reactions affecting the crinoid skeleton either by modifying size and shape of skeletal ossicles or by inducing the formation of additional skeletal plates. Such host reactions invite further investigations. Among the 8 species of endoparasitic myzosotomids, 4 infest ophiuroid or cnnoid gonads (respectively Protomyzostomum spp. and Mesomyzostoma katol). Gonad-infesting myzostomids cause at least partial castration of their host. Fedotov (1916) reported that P. polynephns infests the gonads of its ophiuroid host by rupturing the wall of the bursae. Infestation may b e very intense (up to 119 myzostomids per host) a n d the Dis. aquat. Org. 3: 59-83, 1987 Fig. 1 Gallicole and cysticole myzostom.i.ds of crinoids. ( A ) Myzostomum deformator. Gall in the pinnules of Endoxocrinus alfernicirrus; (B) Myzostornum tenuispinum. Gall in the arms of Pachylornetra inaequalis; (C) Cystomyzostomum murrayi. Cyst on the arms of Stiremetra breviradiata. (Redrawn from Graff 1884) parasites ingest germ cells causing direct castration of the gonads. The host reacts by progressively encysting each parasite by a thick layer of dense connective tissue in which minute calcareous ossicles are seen. Fedotov suggested moreover that germ cell dedifferentiation may also occur. According to Remscheid (1916), Mesom yzostoma reichenspergiinfests gonads, while according to Prenant (1959)it affects only the arm coelom of its crinoid host a n d feeds on coelomocytes. Asteriomyzostomum asteriae (Fig. 2) is said to feed on the host's digestive contents; its occurrence supposedly increases the propensity of asteroids to shed their arms (Marenzeller 1895a, b). Pulvinomyzostomum pulvinar might be more properly classified as 'semi-parasitic'. It inhabits the crinoid's digestive tract (anterior part) living intimately attached to the host's digestive epithelium without causing injury but diverting the flow of food particles entering the crinoid's mouth (Prenant 1959, West & West 1976). Van der Land 1975, Kristensen 1980). This symbiosis has been reported only from Brittany, France (Roscoff a n d adjoining areas) where it is common. Very high infestation rates occur in various populations, viz. 45, 80 and even 100% (respectively Bare1 & Kramers 1970, Kristensen 1980, Cuenot 1912). The number of T. synaptae per holothuroid is highly variable; it ranges from 2 to 3 up to 270 individuals (respectively Cuenot 1912, Kristensen 1980). The latter author noted that infestation occurs only from May to October (eggs a n d juveniles are seen in J u n e and July). Kristensen presented photographic evidence that T. synaptae punctures the epidermal cells of L. galliennei and sucks out their contents; such behavior indicates a parasitic nature of the tardigradeholothuroid association. Another tardigrade, Actinarchus doryphorus, occurs on the body surface of the small European clypeasteroid echinoid Echinocyamus pusillus. According to Schulz (1935), A. doryphorus is a facultative parasite. Agents: Tardigrada Agents: Crustacea, Copepoda The tardigrade Tetrakentron synaptae lives attached to the body surface of the synaptid a holothuroid Leptosynapta galliennei (Cuenot 1892, 1912, There is little information regarding the relations between echinoderms and closely associated J a n g o u x . Dlseases of Ech ~ n o d e r m a t a a- g e n t s metazoans copepods. This IS paradoxical given the high numbers of copepods recorded as echinoderm symbiotes (e.g. Humes 1980a, 1986, Stock 1968b). Table 2 lists species demonstrated or presumed to be parasites. Although many ectoparasitic copepods of echinoderins have been reported in the literature, the parasitic nature of these assoclations has been proven for only 4 species, viz. Cancenlla tubulata, Scottomyzon gibberum, Astel-ocheres lilljeborgi, and Ophiopsyllus reductus (respectively Carton 1968, Rottger 1969, Rottger et al. 1972, Emson et al. 1985). A. lilljeborgi is not particularly adaptated to parasitic life. It is a motile copepod which uses its siphon-shaped mouth to feed on the skin of the asteroid Henricia sanguinolenta. S. gibberum mainly lives on or near the pedicellarial rosettes of Asterias rubens where it seems to feed more deeply in the host tissues (Fig. 3 ) . 0. reductus lives firmly attached to its ophiuroid host and feeds on surface tissues. As for endoparasitic species, Pyefinch (1940) concluded that Ophioica asyrnnietrjca found in the genital bursae of ophiuroids does not feed on the host's tissues, and Changeux (1961) reported - based on casual observations - that the intracoelomic species Allantogynus delamarei 'browses' on the holothuroid's mesothelium. Host reactions against several species of parasitic copepods have been documented. Parasitism by Scottomyzon gibberum induces the infested asteroid to produce dermal outgrowths which progressively embed the copepods (Rottger 1969). Ophiuroid hosts tend to produce a fibrous, sometimes calcified cyst around symbiotes which live in their bursae (Pyefinch 1940, Heegard 1951). However, encystment does not occur with all bursal-infesting species (Stephensen 1935a). Mortensen (1933) recorded unidentified copepods inducing gall formation in the dorsal wall of both Ophionlitrella corynephora and 0. hamata. Paterson (1958) reported that cysts, presumably produced by the host, form around the intracoelomic copepod Cucumaricola notabilis. According to him, small spherical or oval cysts contain juvenile parasites. These cysts are attached to the coelomic wall either near the circumpharyngeal calcareous ring or on the cloaca near the insertion of the respiratory trees; large amorphous cysts containing mature copepods lie detached in the coelomic cavity. Jungersen (1914) reported that the ophiuroid Asteronyx loveni produces cysts around intracoelomic copepods. These cysts are attached on the ophiuroid's coelomic wall. Other intracoelomic copepods have been noted to live often 'free in the coelomic cavity' without seemingly inducing any host reaction (e.g. intracoelomic copepods of holothuroids). However, some bursal- or coelom-infesting copepods may produce conspicuous swellings of the body wall of ophiuroids. These swellings follow the outline of the 63 parasite and do not result from a particular host reaction (Mortensen 6 Stephensen 1918, Stephensen 1933, Goudey-Perriere 1979). Copepods have been found to induce gall formation in the spines of echinothurid echinoids by Hansen (1902),Stephensen (1935bj and Stock (1968a).According to Stock, the galls consist of loose calcareous material and always occur in the middle part of the spines. Galls contain a pigmented ampula In which lives a slngle copepod. Another gallicole copepod, Pionodesmotes phormosomae, occurs inside the echinoid Hygrosoma petersi (Bonnier 1898, Koehler 1898, see also Mortensen 1935) (Fig. 4). P. phormosomae lives in conspicuous inner galls located in the oral part of the host's coelomic cavity. The spherical, calcified galls correspond morphologically to intracoelomic outgrowths of the echinoid body wall. Each gall opens on the host's outer body surface by a slit measuring 1 to 2 mm in length. The slit is protected by the spines of the echinoid. At least 1 large female copepod was found in each gall. According to Bonnier (1898) the copepod does not prey on host tissues and obtains its food mostly from the outside through the slit in the gall. Mortensen (1935) reported that empty galls progressively disappear: the slit enlarges, then the gall wall normalizes, and finally new outer appendages develop. The way in which intracoelomic copepods infest their host has been considered only for holothuroids and asteroids. According to Paterson (1958) and Changeux (19611, holothuroids are infested by larval copepods which cross the digestive wall either at the level of the esophagus or at the level of the cloaca and respiratory trees. In asteroids, Carton (1974) showed that male Botulosoma endoarrhenun~penetrate the coelom of Ech~nasterpurpureus through the respiratory papulae Female B. endoarrhenum actually settle and grow in papulae, living in a kind of integumental cavity. The effects of parasitic copepods on the biology of echinoderms have been considered practically only for Amphiurophilus amphiurae whlch parasitizes the genital bursae of a brooding ophiuroid Amphipholis squamata ( e . g Goudey-Perriere 1979, 1980). The author reported that the occurrence of the parasite does not affect the host's gonads but decreases the host's fecundity by inhibiting the development of embryos incubated in the ophiuroid genital bursae. Such inhibition could result from competition for food between parasite and embryos. Gonad destruction occurred, however, in the ophiuroids Ophiomitrella clavigera and 0. corynephora parasitized by a n unidentified copepod (Mortensen 1933). According to Emson et al. (1985), parasitic copepods on the ophluroid Ophiocomella ophiactoides may decrease the tendency of the host to undergo fission (cross-disc division). Dls aquat Org. 3. 59-83, 1987 64 Asteriomyzostom um asten'ae, a myzostomid para- Fig. 2. site of the pyloric caeca of the asteroid Sclerasterias neglecfa. E: enlarged and deformed part of pyloric caeca harboring the parasite; M: myzostomid. (Redrawn from Stummer-Traunfels 1903; sllghtly moddied) Agents: Crustacea, Cirripedia Thorac~dcirripeds (i.e. barnacles) somet~mesattach to the outer body surface of some specles of regular echinoids (Lytechninus andmesus, Tetrapygus niger, Stomopneustes variolans, Strongylocentrotus spp.) and clypeasteroids (Dendraster excentricus, Rotula orbiculus). Either of 2 situabons may occur: ( 3 ) the barnacle attaches only loosely to the host's body wall overlaping an intact cutaneous epithelium (Moore & McPherson 1963); (2) the barnacle is strongly attached to the echninoid test, the epidermis being destroyed (Giltay 1934, Dartevelle 1940, Boolootian 1958, 1964, Strachan 1970, Ganapati & Sastry 1972, Hurley 1973, Bay-Schmith & Jana 1977). Barnacles Bdlanus spp. have been not~cedon echlnoderms mainly by casual observations, except for the association between D. excentricus and Balanus concavus pacificus detailed by Booloot~an(1964).There appears to be general agreement that the activity of pedicellariae and spines would avoid settlement of barnacle cyprids on the echino~d body wall (e.g Campbell & Rainbow 1977) However, the reviewer believes that the failure of cyprids to settle results basically from the occurrence of a cutaneous epithelium which prevents larval fixation. Presumably cypr~dsettlement can occur only when the echinoderm body wall is wounded. or when the epiderm is eroded for some mechanical or pathological reasons (such as bald-sea-urchin disease; Jangoux 1987a). Barnacles of the genera Pachylasma, Scalpellum and Verruca have been noticed firmly attached to the stem or cirri of several species of stalked crinolds (Clark 1921). Ascothoracida are 'naked barnacles' that parasitize octoralllan corals and echinoderms, except holothuroids. They have been found either as ectoparasites on crinoids and ophiuroids (genera Ascothorax, Parascothorax, Waginella) or as endoparasites in the coelomic cavity of asteroids and spatangoid echinoids (genera Ctenosculum, Dendrogaster, Ulophysema) (Table 3 ) . Waginella metacrinicola lives attached to the cnnoid calyx where it forms a conspicuous depression. It feeds on the 'fluids' of the host by piercing the body wall w t h its oral appendages (Okada 1926). Ophiuroid-associated ascothoracids infest the genital bursae producing marked swellings of part of the host disc (Fig. 5). According to Wagin (1946) Ascothorax ophioctenis feeds on the cells of the bursal wall and probably also on the coelomic fluid of the host. As for intracoelomic ascothoracids, Brattstrom (1947) concluded that Ulophysema oeresundense directly absorbs nutnents from the echinoid's coelomic fluid, while Wagin (1976)believed that coelomocytes form the bulk of the diet of Dendrogaster spp. (Fig. 6). Echlnoderms react against intracoelomic ascothoracids by forming a host envelope that surrounds the mantle of the parasite. According to Wagin (1946), this envelope consists of coelomocytes and covers even the mantle opening which consequently is continually cleaned and smoothed by the ascothoracid's anterior appendages; Wagin claimed that such appendage activity actually corresponds to the normal way of obtaining food by intracoelomic Dendrogaster species. However, investigations by Bresciani & Jespersen (1985) on Ulophysema oeresundense, parasitic in the coelom of Echinocardium cordatum, showed that the envelope is made of choanocyte-like cells and that consequently it should originate from host mesothelium. A conspicuous host reaction also occurs in Parascothorax sinagodoides which lives attached in the bursal cavity near the bursal slit of J a n g o u x Diseases of E c h i n o d c r ~ n a t a a g e n t s metazoans 65 T a b l e 2. Parasitic copepods from echinoderins (compiled from t h e sources indicated) Hosts: A, asteroid; C, crinoid, E, echinold; H , holothuroid; 0, ophiuroid Copepod Host Location ~nhost Geographical area Remarks Source - I. Harpacticoida Metis Actinopyga agasslzl (H) holothur~ae T ~ s b efurcata Ocnus p l a n o (H) nsbe holothuriae 11. Cyclopodia Allantogynus delamare1 Coelom~ccavlty Coelomlc c a v ~ t y Almost 100 of holothuro~dsinvestigated were infested (Monticelh) "'L Trop~calAtlantic (Bahamas) Med~terraneanSea (Naples) Edxvards (1891). Humes (1980a) Montlcelli (18921, Humes (1980a) Mediterranean Sea (Banyuls) Humes (1957) Holothuna stellatl (H) D~gestivetract (antenor part) Holoihuna tubulosa, Holothul-~a poll, (H) H o l o t h u ~ ~stellat~ a Antenor part of coelomic cavity frxed on lnesothelium (occur also in wall of pharynx, gonads, tentacular ampullae, 01Polran vesicles) H tubulosa and H stellatl. about 2 copepods holothuroid-' (82 infested/ll7 investigated); H poll is only occasronal host Mediterranean Sea Changeux (Banyuls, V~llefranche) (1958, 19611, Stock & Kleeton (1963, Humes (1980a) Bursae About 12 infested oph~uroidsIn Roscoff (Goudey-Perl-iere) NE Atlant~c(Roscoff); Sorth Sea (Bergen); NW Atlantic (IYoods Hole region) Fewkes (1887, 1888). Herouard (1906).Bocquet (1952).Masson (1965).GoudeyPerriere (1979, 1980) Rottger et a1 11972),Brun (1976) Carton (1974) Aniphlul-ophllus Amphlphohs squamata (0) amphiurae - Asterocheres hlljeborgi Henncia sangu~nolen- Free on outer body ta (AI surface Up to 24 copepods asteroid- ' NE Atlantlc (Swedlsh coast: Gullmarflord) Botulosoma endoarrhenuni Ech~nasterpurpureus (A) Respiratory papulae ( 2 copepod), coelo~nrc cavlty (d copepod) Several dozen copepods as hero id^' SW Indian Ocean (Madagascar Tulear) Calvocheres engell Hygrosoma hoplacantha (E) Galls on spine at adora1 s ~ d e Two ~nfested ech~noidswith 6 and 3 copcpods, respectively NE Indran Ocean (off Celches, Tlrnor Sea) Stock (1968a) Calvocheres globosus Calver~osoniag r a d e , Sperosoma qulncuna a l e (E) Galls on secondary splnes or on spines of adoral s ~ d e Yot more than 1 9 copepod gall (2 and 5 galls observed on 2 infested hosts) N Indo-Pacific (off Philippines, In Halmahelra Sea; off SE Japan) Hansen (19021, Stephensen (1935h),Stock (1968a) Calvocheres oblongus Hygrosoma peters1 (E) Galls on spines N Pac~flc(SE of Japan) Stephensen (1935b) Calypsanon carinaturn St~chopusnionotuberculatus (H) Internal 1 C copepod ejected from host upon collect~on N\.V Indlan Ocean (Ethiopia: Dahlak Archipelago) Stock (1968b). Humes (1980a) Calypsanon leprum Actinopyga rnauntanla, Actlnopyga miharis, Actlnopyga lecanora (H) Internal A. ~ n l l i a r1s ~ smost common host (76 copepods - 43 X and 33 9 -collected from 726 hosts) SW Indian Ocean (Madagascar NossyBe) Humes & Ho (19691, Humes (1980a) Calypsarion sentosum Bohadschia rnarn~orata Internal (H) 8 copepods (5 ( ' 3 Q ) collected from 8 hosts SLV Indian Ocean (Madagascar:NossyBe) Humes & Ho (1969),Humes (1980a) Calypana changeuxr Holothuria tubulosa, Holothuna stellati (H) Mediterranean (Banyuls) Changeux (1961),Stock & Kleeton (1963). Hurnes (l980a) Cancer~lla tubulata Amphiphol~ssquamata Attached to outer body (0) surface NE Atlantic (Brittany, North Sea) Giard (1887). Carton (1968) Esophagus and gonoduct - - Infestation rate ca 10 % (Giard) J a n g o u x : Diseases of Echinodermata. a g e n t s metazoans 67 T a b l e 2 (continued) Copepod Host Location in host Diogen ella splnicauda Internal (ejected from Holothilria mrxlcana, Actlnopyga agassrzl (H) H. niexlcana; Stock) Dlogenidium deforme Holothuna glaberrima, Holothuria arenicola, Holothuria mexicana Internal (ejected from H. glaberrima; Stock) (H) Remarks 2 c o p e p o d s ( l h , 1 9) from H n~exlcana (Stock),46 copepods ( 2 2 d , 2 4 9 ) f r o m 119 H. nlexrcana, 50 copepods (32 d . 18 Q ) from 62 A agassizi (Humes & H01 2copepods(l 6 . 1 9 ) from 1 H. glabernma (Stock);7 copepods (1 6 . 6 C)from87H. arenicola, and 5 copepods(22.3 Q ) from 24 H. mexicana Geographical area Tropical Atlantlc ( C u r a ~ a oBahamas, , Jamaica, Puerto Rico) Coelomic cavity 9copepods (5 d ,4 Q ) Tropical Atlantic from 3 H mexlcana (Bahamas, C u r a ~ a o , (Stock);9 copepods Jamaica, Puerto Rico) (4 5 5 9 ) f r o m 6 9 A agdsslzi, 10 copepods(4 8 . 6 V ) from 29 H mexlcana. and 6copepods (4 d , 2 9 ) from 11 H. grlsea (Humes & Ho) Diogenidium spin ulosum Isostichopus badionotus (H) Internal (ejected from host; Stock) 1 6 copepod from 1 holothuroid (Stock); 4 copepods (2 d . 2 9 ) from 17 holothuroids (Humesl Dlogenldlurn tectum Actinopyga agassizi (H) Internal 3 d copepods from 64 Tropical Atlantic holothuroids (Jamaica, Bahamas) D i a d e n ~ a s e t o s u n(E) ~ Esophagus 85 copepods collected from 65 echinoids Enteroynathus comatulae Antedon mediterranea. Antedon blhda (C) D ~ g e s t ~ tract, v e rarely in 11 copepods, (2 d . coelomic cavity 9 9 ) from 12A mediterranea (Changeux & Delamare Deboutteville) Enterognath us la tenpes Decametra chad~vlcki, Digestive tract Heterornetra savignyi, OUgometra serripinna (Cl Actinopyya lecanora Digestive tract Lecanunus in testinalls - (H) Lecan urius kossmannianus Actinopyga lecanora, Actinopyga rnufleri (H) Lecanurius planifron tahs Lernaeosaccus ophlacanthae Internal Stock (1968b). Humes & Ho (1970),Humes (1980a) Tropical Atlantic (Puer- Stock {1968b), Humes & H o to Rico, Barbados, (1971),Humes Bahamas) (1980a) A ctinop yga agassizi, Holothuria mexlcana, Holothuria grlsea (H) Not more than 4 copepods found Source Edwards (1891), Stock (1968b), Humes 61 Ho (1971).Humes (1980a) Tropical Atlantic (Puer- Stock (1968b). to Rico, Jamaica) Humes & Ho (1971).Humes (1980a) Humes & Ho (1971).Humes (1980a) New Caledonia Dojirl& Humes (1982) Mediterranean Sea (Banyuls,Naples);NE Atlantic (Dublin,Plymouth) Giesbrecht (1900),Grainger (1950), Changeux & De1amare.De. boutteville (1956). Stock (1959) Indian Ocean (Red Sea) Stock (1966) NW Pacific, (Philippines Bohol Island) Kossmann (1877,quoted by Stock 1968b) 14 copepods (7 d. SW Indian Ocean 7 Q ) from 1'1 ,.\. lecan- (Madagascar:NossyBe) ora and 8 cupepods (4 6 . 4 9 ) from 205 A. m ullen Humes (1968) Actinopyga echinites, Digestive tract (postActinopyga miliaris (H) erior part) 3 copepods (2 6 . 1 9 ) from 3 A . echinites Tropical W Pacific (New Caledonia; Great Barrier Reef) Humes (l980a) Ophiacantha disjuncta (0) Only 1 individual found Antarctic seas (Palmer Archipelago) Heegard (1951) Coelomic cavity (presumably) Dis. aquat. Org. 3: 59-83, 1987 68 Table 2 (continued) Copcpod Host Locatlon in host Remarks Geographical area Source Licholhuria mandibulans Holothuria atra, Holothuria scabra. Holothuna nobihs. Holothuria fuscopunctata (H) Internal (ejected from host; Stock) 4 copepods (2 d , 2 Q ) W Indian Ocean (Red from12H.atra Sea- Eilat; Madagascar (Stock);l88 copeNossy-Be) pods(54 $, 134 ? ) from470H.scabra. 19 copepods (6 d . 13 ' 2 ) from24 H.nobilis, 139copepods(34d , 105 P ) from345 H. atra, and 1 d copepod from H.fuscopunctata (Humes&Ho) Stock (1968b), Humes & Ho (1969),Humes 1980a Ophioika appendiculata Ophiornitrella calmgera, Ophiacantha biden tata (0) Bursae, forming swelling of host's tissue 1 to 3 copepods host-' (no more than 1 bursa infested host-') N Atlantic (S of Greenland. S of Iceland. W of Hebrides) Stephensen (1935b.1940) Ophioika asymmetrica Ophlacantha imago (0) Bursae, forming sl~ght swelling of host disc Each of the 2 infested ophiuroids investlgated had apair ( d , P ) of copepods encysted in 1 of their bursae Antarctic seas Ophioika ophiacanthae Ophiacan tha severa (0) Bursae, forming swelling of host disc Only 1 Q copepod found SE Indian Ocean (Bali Sea) Stephensen (1933) Oph~o~ka tenujbrachra Oph~acanthawblpara, Ophiacantha disjuncta Bursae (presumably) 8 copepods found Antarcticseas (South Georgia; Adelaide Land) Heegaard (1951) (0) Ophiopsyllus reductus Ophiocomella ophiacto~des(0) Oraland lateral parts of arm base 40 to 65 O h of host Jamaica population parasitlzed depending on period of year. Usually l to 3 copepods ophiuroid-' Pinnodesmotes phormosomae Hygrosoma petersll (E) Intracoelomic galls l o r 2 ( d ,Q ) copepods gall-' NEAtlantic (Azores) Koehler (1898), Bonnler (1898) Scottomyzon gibberum Asteriasrubens(A) Galls in rosettes of pediceuariae Most asteroids are in- North Sea (off Helgofested; 4 to 175 land; Kattegat) copepods asterold-' Rottger (1969. see also Bare1 & Kramers (1977) Synapticola teres Polyplectana keferstein^, Synapta maculata (H) Coelomic cavity, digestive tract (posterior Part) 5 copepods (2 6 . 3 9 ) Indlan Ocean (Amboifree in coelomic cavi- na);W Pacific (Queensty (Voigt) land) Voigt (1892). Humes (1979) Stomachfolds About50°h of 0. aculeatainfested NorthSea (Gullmarfjord) Bresciani&Lutzen (1962) Atleast2copepods ( 6 ,P ) found N\VPacific(Ph~lippines: Mindanao) Mortensen & Stephensen (1918) NEIndianOcean (Arnboinal Stephensen (1933) Emson et al. (1985) 111. Monstrilloida ThespeslopOphiothrixfrag~l~s, syllusparadoxus Ophiopholisaculeata (0) IV. Incertaesedis kthrochorAstrocharisgrac~lis(O) Galls at arm base deumium appen diculosc~m Arthrochordeumium asteromorphae Codoba discoveryi Chordeum~~rm obesum Parachordeumium tetraceros ' Indentlfled Asteromorpha koehleri (0) Galls at arm base Only l copepod found Oph~urameridionalis Calcif~edcystsinside bursa Cystsw-ithincoelomic cavity (attachedcysts) 3 infestedophiuroids Antarcticsea (South were found Georgia) Almost 100%lnfesta- NorthSea (Skagerrak) tlon; no more than 1 copepod gaU-l Coelomiccavlty Only 1 copepodobsewed (0) Asteronyxloven~(O) Amphipholissquamata (0) Phormosoma uranus by Koehler (1898) Med~terranean(Villefranche) Heegard (1951) Mortensen (1912). Jungersen (1912,1914) Le Calvez (19381,Bocquet (1952) Jangoux. D ~ s e a s e sof Ech~nociermata.agents m e t ~ i ~ o a n s Fig. 3. Scottomyzon gibberum, an ectoparasitic copepod of the asteroid Asterias rubens. (A) Female on tube-foot tip. ( B & C) Copepods embedded in dermal tissue of a pedicellarial rosette. a1 & d 2 : antennae; cp: crossed pedicelloria; mxp: maxilliped; 111x2: maxilla; S : spine; s p : straight pedicellaria. (After Rottger 1969) Ophiura quadrispina (Wagin 1964, 1976). The ophiuroid first encysts the parasite with its own tissues; then both cyst and parasite grow into the bursal cavity. The cyst is gradually ejected from the cavity through the bursal slit. P. sinagodoides apparently has adapted its llfe cycle to the reaction of its host, as rejected cysts as a rule include infesting ascothoracid larvae. According to Grygier (1985b) the asteroid-associated ascothoracid Endaster hamatosculum lives in conspicuous inner galls located in the arm coelom of its host. Galls are calcified and correspond morphologically to intracoelomic outgrowths of the asteroid body wall; each gall opens on the host's outer body surface by a slit-like hole. Ascothoracids may castrate their host. Thus Ascothorax ophioctenis causes complete castration of Ophiocten sericerum, even though it does not feed on the host's gonads. Wagin (1946) found a single juvenile A. ophioctenis to Inhibit the activity of the germinal epithelium leading to a regression of the whole gonad. Similar castration occurs in spatangoids infested with large-sized Ulophysema oeresundense. According to 69 Brattstrom (1947), castration results from competition for food between gonads and ascothoracid. Casual observations of host castration have been made also by Yosii (1928b), Fisher (1930) and Korschelt (1933). Heavy infestation of Ophionotus victoriae by Ascothorax gigas (up to 7 genital bursae infested) causes occlusion of the ophiuroid mouth opening a n d reduces the development of the gonads (Grygier & Fratt 1984). The gonads do not coillpletely regress, however. The stoillach volume is reduced, but there is no demonstrable reduction in the volume of s t o n ~ a c h contents. According to Brattstrom (1947) infesting larvae of Ulophysema oeresundense usually penetrate their host through the genital apertures a n d also through the ambulacral pores. Infesting larvae thus reach either the gonadal lumen or the axial sinus. They must cross the wall of the gonad or of the axial sinus in order to reach the general coelomic cavity where they normally grow and reproduce. When mature. Ulophysema spp. most frequently attach to the apical part of the host's body wall in which they bore holes up to 1 mm in diameter. These holes are used by the larvae to escape from the host. Grygier (1981) suggested that the release of larvae from a female Dendrogaster antarcticus into the asteroid coelorn is produced by rupturing of the mantle of the female. Larvae of Dendrogaster tasmaniensis penetrate the asteroid Allostichaster polyplax through respiratory papulae; they infest the coelom and escape by crossing the asterold's stomach wall and passing to the outside through its mouth (Hickman 1959). Agents: Crustacea, Malacostraca Associations between amphipods and echinoderms were reviewed by Vader (1978).A parasitic relation has been inferred by Shoemaker (1919) for Laphystiopsis iridometra whlch l ~ v e sembedded in the calyx of the crinoid Iridometra melpomene. Typical ectoparasihsm occurs with the North Sea species Epimeria parasitica which feed on the outer tissues of the holothuroid Stichopus tremulus a n d the asteroid Porania pulvillus (Vader 1978). According to Ruffo ( 1 9 ) the amphipods Jassia ocia and Aristias negletus occur at Banyuls (France, Mediterranean Sea) in the coelomic cavity of the holothuroid Stichopus regalis a n d the crinoid Antedon mediterranea, respectively. An unexpected association between the tanaidacean Exspina typica a n d 3 species of deep-sea holothuroids has been reported by Thurston et al. (1987). E. typica should be considered a facultative parasite; it occurs either in the intestine or body cavity of its host. Crabs associated with and potentially harmful to echinoderms belong to 3 distinct families: 70 Dis. aquat. Org. 3: 59-83, 1987 Fig. 4. Pionodesmotes phormosomae, a gallicole copepode parasite of the echinoid Hygrosorna petersi (A) Inner view of the oral part of a dissected echinoid showing the location of calcified galls. (B) Section through gall containing an ovigerous female. p . outer pore of the gall. (A after Koehler 1898; B after Bonnier 1898) Pinnotheridae (genera Dissodactylus, Ophisthopus, Pinnaxodes, Pinnixa, Pinnotheres), Parthenopidae (genera Echinoecus, Zebrida), and Portunidae (genus Lissocarcinus) (Table 4 ) . Feeding habits of associated species of Ophisthopus, Pinnixa, Pinnotheres and Lissocarcinus have not been reported. Presumably these forms do not feed on host tissues, nor cause any other detrimental effects, except to slightly wound the wall of the respiratory trees or of the cloaca (Tao 1930, Jones & Mahadevan 1965).Species inhab~tingthe posterior part of the echinoid digestive tract (Echinoecus convictor, E. pentagonus, E. ra thbunae and Pinnaxodes chilensis) are generally said to feed on host fecal pellets (e.g. Miyake 1939, Fenucci 1967). Trophic relations between E. pentagonus and the echinoid Echinothrix calamaris were studied carefully by Castro (1971) (see also Suzukl& Takeda 1974). Males and small immature females inhabit the peristomeal region where they feed on epithelia1 tissue and tube feet, damage being in equilibrium with the regenerating capacities of the host. Large mature females live in the rectum and ingest material from fecal pellets and aggregates of pigmented coelomocytes that migrate across the rectal wall. 71 Jangoux: Diseases of Echinodermata. agents metazoans Table 3. Parasitic ascothoracid cirripeds (compiled from the sources indicated). Ascothoracid species names according to Wagin (1976) and Grygier (1981 to 1983). Hosts: A, asteroids; C. crinoids, E, echinoids; 0, ophiuroids. Unidentified species of ascothoracids were reported from asteroids by Fisher (1940; Diplasterias meridionalis) a n d from ophiuroids by Bartsch (1982; Ophiurolepis inornata) Ascothorac~d Host Location on/in host Remarks Ascothorax bulbos~rs Ascothorax 9lPs Amphiclra belgrcar, An,- Bursae phicli-a mrcroplax (0) Oph~notusvictoriae (0) Bursae Ascothorax mortensenl Ascothoras oph~octenrs Amphiura niicroplax (0) Bursae Ascothorax pilocauda tus Ctenosculum ha waiiense' Dendrogaster antarcticus Dendrogaster arborescens Ophiosphalma armigerun~(0) Brisinga evermanni (A) Bursae Acodontaster consplcuus (A) Dlpsacaster sladeni, Coscinasterias calan~aria( A ) Hlppastena cabfornrca Coeloi~~i( cav~ty lnfestat~onlrlvel ca 2 '% Dendrogasler arbusculus Dendrogasler arctica Ophiocten sericeum (0) Bursae Coelornlc cavity Coelomic cavity Dendrogasler argentinensrs Dendrogaster astericola Anasterias minuta (A) Coelomic cavity Henricia sanguinolenta 'A) Coelom~ccavity Dendrogdster astennap Dendrogaster astropectlnls Dendrogaster beringensis Asterjna burtoni (A) Coeloni~ccavity Astropecten scoparius Coelom~ccavlty Dendrogaster Iudwigi Dendrogaster murmanensls Dendrogaster okadai Southern Atlantic [off South Georgia) Up to 27 parasites South Sandwlch Ishost-' (Grygier & Fratl) lands Infestation rate: 0.7 to 3.4 '10 (Wagin) 1 to 3 ascothoracids host-' (infestation not uncommon) (Okada) 1 observation (A) Leptasterias groenlandica (A) Dendrogasler dichotomus Dendorgaster dogie11 Dendrogaster elegans Dendorgaster fish eri Dendrogaster iwanowi Dendrogaster 1 ascothoracicl host-' Coelomic c-avity Coelom~ccavity (A) Eremicaster tenebrarius, Coelorn~ccavity Hyphalaster inermis (A) Crossasterpapposus (A) Coelom~ccavity Geographical area - 5 asteroids infested (326 investigated) 3 infested H. sangujnolenta (500 investigated) 1 to l 0 ascothoracids asteroid5 infested asteroids (250 investigated) 3 Infested asteroids (13 investigated) - Source Heegard (19511 South Georgia Wagin (1968),Grygier ( 1 9 8 3 ~ )Gry. gier & Fratt (1984) Gryg~er( 1 9 8 3 ~ ) Barents Sea; Kara Sea; North Atlantic (around Iceland and Faroe Islands) NE Atlantic Djakonov (1914), Stephensen (1935a).Wagin (1946) Gryg~er( 1 9 8 3 ~ ) Tropical 1V Pacif~c (Hawail) Antarctic seas (off Ross Island) Southern Indlan Ocean (off Cape Town); Misaki (Japan) NE Pacific (off California) N Pacific (Bering Sea) Heath (1910),Waren (1981) Fisher (1930),Gryyier (1981) Le Roi (1905, 1907), Okada (1925) SW Atlantic Barents Sea (White Sea) F~sher(191l), Grygier (1982) Flsher (1930),Korschelt (1933),Wagin (1950),Grygier (1986) Grygier & Salvat (1984) Kn~powitsch(1891) Red Sea (Gulf of Aqaba) Misaki (Japan) Achituv (1971) Bering Sea NE Pacific Wagin (19571,blddsen (19611, G ~ ~ g i e r (1985a) Wagin (1950) Bering Sea Wagln (1950) Wagin (1950). Grygier (1986) Fisher (1928),Grygier ( 1982) Wagin (1950) Wagin (1950) Yosii 11928b) Pteraster obscurus (A) Coelornic cavity Leptasteriaspolaris (A) Coelornic cavity Pedicellaster magister megalabis (A) Leptasterias fisher; (A) Coelomic cavity Coelomic cavity Bering Sea; Gulf of St Lawrence NE Paclhc (off California) Berlng Sea Leptastenas fjshen' (A) Coelomic cavity Bering Sea Echinaster luzonicus, Certonardoa seniiregularis, Nepanthia be]cheri (A) Crossasterpapposus, Solaster endeca (A) Coscinastenas calamaria (A) Coelorn~ccavlty W. Pacific (Philippines, Le Roi (1905, 1907). Japan Sea; Tasmania) Yosii (1928b), Kenny (1959) Coelornic cavity Barents Sea; N Pacific (Okhotsk Sea) Misaki (Japan) Coelonlic cavity - Korschelt (1933). Wagin (1950) Yosii (1928b) Dis. aquat. Org. 3: 59-83, 1987 72 Table 3 (continued) 1 1 Ascothoracrd Dendrogaster onen talis Dendrogaster punctata Dendrogaster rarnosus Den drogaster rimskykorsakowi Dendrogaster sagittaria Den drogaster tasmaniae Endaster hamatosculum Parascothorax synagodoides Ulophyserna oeresundense Host Location on/in host Remarks Geographical area Source Leptaster~aspolans( A ) Coelomic cavity Bering Sea Wagin (1950) Poraniops~sinflata(A) Coelomic cavity Grygier (1982) Leptasterias hsheri (A) Coelomic cavity NE Pacific (off California) Bering Sea Ctenodiscus crispatus, Hippastena leiopelta Coelomic cavity Wagin (1950) NW Pacific (Okhotsk Sea) Wagin (1950) Phihppine Seas Around Tasman~a Flsher (1919). Grygier (1985a) Hickman (1959) Ph~lippinesSea Grygier (1985b) Okhotsk Sea Wagin (1964) North Sea (Scandinavian coast) Brattstrom (1936. 1938, 1946, 1947), Bresciani & Jespersen (1985) Brattstrom (1937) (A) Sidonaster vaneyi (A) Coelomic cavity AUoshchasterpolyplax (A) Coelomic cavity Zeroaster carinatus (A) Intracoelomic galls Ophlura quadrispina In cysts In bursae (0) Echinocardium cordatum, Echlnocard~um flavescens, Brissopsis lyrifera (E) Coelomic cavity; more rarely gonads, ambulacral ampullae or axial sinus 2 individuals In a single asteroid 1 ascothoracid asteroid-' (26 infested/ 116 investigated) 1 to 2 ascothoracids asterold-' 1 to 9 ascothoracids oph~uroid-' 1 to 9 (mostly 1) ascothoracids echinoid-' (all stations together: 814 infested/42 874 investigated) (Brattstrom 1947);infestation rate: 20 to 25 % (Brescian~& Jespersen) 1 or 2 ascothoricids echinoid-' Ulophysema pourtalesiae Pourtalesia jeffreys~(E) Coelom~ccavity Waginella metacrin~cola Metacrinusrotundus (0) On stalk N Atlantic (between Norway. Spitzberg, Greenland and the Faroes) Sea of Japan Waginella axotrernata Metacrinus acutus, Metacnnus angulatus, Metacrinus clngulatus (C) On cirri Indonesia; China Sea Okada (1926, 19381, G q f g ~ e r (1983b) Grygier (1983b) Waren (1981) showed that Ctenosculum hawaiiense (Heath 1910) is better interpreted as an ascothoracid than as a mollusc; Grygier (1983a) confirmed this op~nionfrom d~rectexaminailon of C. hawailense According to Mortensen (1943a) and Suzuki & Takeda (1974) an obvious parasitic relation occurs between the parthenopid crab Zebrida adamsi and several echinoid species (Table 4 ) . the crab feeds on appendages, skin and peripheral dermal tissue of echinoids, producing conspicuous naked test areas. Similar parasitic behavior also occurs with some species of Dissodactylus living on clypeasteroid and spatangoid echinoids (Dexter 1977, Telford 1982, Bell & Stancyk 1983).Telford (1982) reported that, depending on the species studied, the associated Dissodactylus take 50 to 100% of their diet from host tissues. According to him differences in feeding habits can be attributed partly to the structure of host spines, viz,larger test areas are denuded if the spines of the echinoid host are more porous (Fig. 7). Gut-inhabiting species may produce deformations that are sometimes very conspicuous. Verrill (1867) reported that Pinnotheres chilensis considerably distends the rectal wall of the echinoid Coenocentrotus gibbosus to form a membranous cyst. The cyst attaches aborally to the echinoid body wall and extends near the mouth into the coelomic cavity. The anal area of the host is depressed and distorded, and the echinoid test is usually swollen on the side over the cyst (Fenucci 1967). P. chilensis parasitizes both C. gibbosus and Loxech~nusalbus. According to Mortensen (1943b) the crab does not induce major test deformation when infesting L. albus. Similar but less conspicuous deformations are caused also by Echinoecus convictor, E. pentagonusand E. rathbunae (respectively Bouvier & Seurat 1905, Castro 1971, Miyake 1939),all of them producing Jangoux Diseases of Echinodermata agents ~netazodns Fig. 5. Ascothorax ophioctenis, an ascothoracid parasite of the genital bursae of the ophiuroid Ophlocten s e n c e u n ~(A) Aboral view of disc of an infested ophiuroid. (B) Dissected genital bursa containing a pair of A. ophloctenls (C) Diagramatic drawing of parasites in situ. a : antennule; br: brood pouch. b s bursa1 slit, bw ophiuroid body-wall, f furca, m dwarf male; t thoracic limbs (Redrawn from Wagin 1946: slightly modified) swellings of anal tube and periproct of their host. According to Suzuki & Takeda (1974) and Dexter (1977), infestations by ectoparasitic crabs often exert major effects and may h11 echinoids. The crabs select a particular site on the host's body surface and clear away the spines of the region (see also Telford 1982). This produces a n obvious decrease in movements and spine activities of infested regular echinoids (Suzuki & Takeda 1974). Dexter (1977) observed a reduction in spine actlvity which decreased the ability of clypeasterolds to feed a n d to move about in aquaria, resulting in a substantial increase in mortality. Agents: Arachnida; Pycnogonida; Insecta The only arachnid reported to be an echlnoderm parasite is the acarian Enterohalacarus mlnutipalpus Dls. aquat. Org 3: 59-83. 1987 Fig. 6. Dendrogaster arborescens, an ascothoracid parasite of the coelomic cavity of the asteroid Dipsacaster sladeni, rnb: main branch; rnp: middle piece; pb. primary branch; sl: distal slit. (After Le Roi 1907) found in the gut of the echinoid Plesiodiadema indicum (Viets 1939). Three specimens were collected in echinoid material dredged by the Albatross off North Moluccas, Indonesia. Northing is known about hostassociate relations. Pycnogonid-echinoderm associations have been noticed by several authors who generally suggested a n ectoparasitic relation. Associations with holothuroids were observed by Prell (1910; Pycnogonum littorale/ Cucumaria frondosa) and Ohshima (1927; Lecythrorhynchus hilgendorfi/Holoth uria lubrica). According to Ohshima, a single host may carry up to 30 pycnogonids which supposedly absorb the blood or 'body juice' of the holothuroid. An obvious pycnogonid-ophiuroid association occurs a t Aldabra (Indian Ocean) with 3 species of Ophioderma a n d the seaspider, Anoplodactylus ophiurophilus (Sloan 1979, Sloan et al. 1979, Stock 1979). Sloan reported various infestation levels - from 3.6 to 83.7% - depending on the host species. He presented evidence that A. ophiurophilus has the potential to feed on the host's tube feet. The occurrence of pycnogonids on echinoids and asteroids has also been reported casually (Stock 1981). An unexpected association between the marine trichopter Philanisus plebeius and the asteroids Patiriella exigua and Patiriella regularis was reported by Anderson et al. (1976) and by Winterbourn & Anderson (1980). Caddis fly eggs were found in the asteroid coelomic cavity. Singly or in small clumps, eggs occur loose within the coelom close to the peristome or enveloped in stomach folds. The authors presumed that eggs are deposited through the respiratory papulae and that newly hatched intracoelomic larvae leave the host either via the same route or through the stomach wall. According to Winterbourn & Anderson (1980), Jangoux. Dlseases of Echinodermata: agents metazoans Table 4. Parasitic decapods (Reptantia) of echinoderms (compiled from the sources indicated). Hosts: E, echinoid; H, holothuroi Reptant D~ssodactylus caln~anr Dlssodactyl~ls cr~nitichelis D~ssoddctylus encopei Dissodactylus glass ell^ D~ssodactylus lockingtoni Dissodactylus mellitae Host Clypeaster rosaceus (E) Locat~onon/in host Outer body surface (oral surface) Outer body surface Mellita sexiesperforata (E) Encope emarginata (E) Outer body surface Mellita longifissa (E) Mellitalongifissa, Encope mlcropora, Encope grandis, Encope californica (E) Mellita quinquiesperforata, Ech~narachnius parma (E) Remarks lnfestatiorl level 25 to 59 ' X , Inlestation level 30 to 60 'Vo - Outer body surface Outer body surface (proximal porhon of pasterior interambulacral lunule) Outer body surface Encope stokesi, Encope n~icropora(E) Dissodactylus pn'mit~vus Meoma ventricosa, Plagiobnssusgrandis D~ssodactylus smlthi Dissodactylus xan tusl i ~ & t a longifjssa (E) Outer body surface Encope stokesi, Encope micropora (E) More otten on E micropora Echinoecus convictor Echinothrix diadema IEI Outer body surface (near lunule or marginal slits) Internal swelling of anal tube Echlnoecus pentagonus Pseudocentrotus depressus, Anthocidaris crassispina, Echinoihrix calamaris, Echlnothru. diadema (El Echlnothrix calamaris, Phyllacanthus dubius (El Bohadschia argus (H) Outer body surface (near peristome) 10 crabs collected on 48 P. depressus ( S u z u k ~& Takeda) Ech~neocus rathbunae Lissocarcin us orbicularus Outer body surface (near lunule or marginal slits) 1 to 2 crabs host-'; infest mostly large echinoids (diamete~ > 30 mm) Outer body surface Infestation level 79 to (mostly on oral surface) 100 YO - - Internal swelling of the anal tubes Mainly respiratory trees, also cloaca Tropical W Atlantic (Florida, J a m a ~ c a ) Tropical W Atlantlc (Barbados) Tropical W Atlantic (Florida to N Brazil) Tropical E Pacific (Mexican coast) Tropical E Pacific (Northern Mexico) 13 crabs collected NW Atlantic from 50 M . qu~nqujes- (Massachusetts to perforata (Pearse);al- Florida) most 100 9b inlestation (Johnson) Dissodactylus nitidus (E) Geographical area 11 holothuroids in fested (61 investigated) Tropical E Pacific (Panama) Cloaca Parastlchop 11scaliforn ~ c u s(H) Cloaca Pinnaxodes chilensis' Loxechinus albus, Coenocen trotus gibbosus (E) Partly embedded in rectum wall Pinniva barnharti Pinnixa timida iMolpadja arenicola (H) Cloaca Caudina chilensis (H) Cloaca Infestation level 75 % Pinnotheres decanus Holothuria scabra (H) Cloaca and respiratory trees; rarely in the coelomic cavity 10 holothuroids Infested (25 investigated) - Telford (1978, 1982) Rathbun (1918) Rioja (1944) Glassell (1935) Rathbun (1918). Pearse (1947), Johnson (l952), Gray et al. (1968),Telford ( l 982) Dexter (1977) Telford (1978, 1982) Tropical L: Pdciflc (Mexican coast) Trollicdl E Pacific Rioja (1944) Dexter (1977) (i'dn~mdj Tropical Indo-West Pacific (Gambier Is1dnd; Mauritius) Tropical Indo-W Pacific (Andaman. Hawaii, Japan: coast of Uchida) Bouvler & Seurar (1905).Mortcnsen (1940) Castro (1971, 1978).Suzuki & Takcda (1974). Sastry (1977, 1981) NW Pacific (Ogasawara Islands) Miyake (1939) NW Pacific (Philippi nes: Puerto Galera) Trott & Garth (1970) N Indian Ocean (Andaman Sea) Ocfur also in other in- NE Pacific (California) vertebrates (e.y , bivalve molluscs) SE Pacific. (Ecuador to 1 to 2 crabs echinoid-'. InfestaPatagonia) tion generally very high (90 to 100 "/.) (Verrill, Clark. Rathbun, Schwabe) Holothuna sp. (H) Telford (1982) Tropical W Atlantic (Barbados,Jamaica) - Llssocarcin us ornatus Ophisthoptrs transversus Source NE Pacific (California) NW Pacific (Japan: Mutsu Bay) N Indian Ocean (In dian coast) Chopra (1931) Rathbun (1918) \/errill (18671, Ortmann (1894), Meissner (1896). Clark (l910), Rathbun (1918). Schwdbe (1936), Mortensen (1943b),Fenucci (1967) Rathbun (1918) Tao (1930) Chopra (1931), Jones & Mahadevan (1965). Adithiya (1969) Dis. aquat. Org. 3. 59-83. 1987 76 Table 4 (continued) Reptant Host Pinnotheres setnai Pinnolheres villosissirnus Unidentified holothuroid Achnopyga mauritiana, Actlnopyga lecanora (H) Zebnda a d a m s ~ Diadema selosum, T r j p n e u s t gratilla, Toxopneustes pileoius, Toxopneustes elegans, Asthenosoma ijimai, Salmacis sphaeroides (E) Location on/in host Respiratory trees Remarks - Cloaca Outer body surface (near ambitus) As a rule 1 crab echinoid-' (11 crabs collected) (Suzukl & Takeda) Geographical area NW Indian Ocean (Andaman Sea) NW Indian Ocean (Andaman Sea; west coast of Sumatra) NW Pacific (Japan: Suruga Bay; coast of Thailand) Source Chopra (1931) Doflein (1904). Chopra (1931) Rathbun (1910). Mortensen (1943a),Suzuki & Takeda (1974),Daniel & Krishnan (1978) ' Often identified Fabia chilensis eggs are found in the coelom during most of the year, the infestation level being about 10%. Agents: Pisces Associations between echinoderms a n d carapid fishes (pearlfishes) are of particular interest as they concern - at least in a few cases - typical parasitic relations between an invertebrate host a n d a vertebrate associate. The systematics, general biology, and ecology of pearlfishes are documented satisfactorily, and some species have been studied in detail (e.g. Zankert 1940, Arnold 1956, Trott 1970, Van Meter & Ache 1974, Smith et al. 1981, Trott 1981).Pearlfish species associate with holothuroids and asteroids are listed in Table 5. Most authors report that the stomach contents of Carapus spp. from echinoderm coelomic cavities do not show remains of host tissue (e.g. Trott 1970, Smith et al. 1981). Carapus spp. are predators. Basically, they use the host's coelom as shelter (Fig. 8). They leave it in order to catch prey, feeding primarily on crustaceans. Casual observations of feeding by Carapus on the host's gonad have been reported however (Hipeau- i I Fig. 7 Dissodactylus crinitichelis, a n ectoparasitic crab of the clypeastroid echinoid Mellita sexiesperforata. The crab is seen beside the devastated area where it had been feeding. (After Telford 1982) 77 Jangoux: Diseases of Echinodern~ata:agents metazoans Table 5. Fishes endosymbiot~cin echinoderms (compiled from the sources indicated). Hosts. A , asteroid; H, holothuroid Fish Carspils actrs Carapus bermudensis Carapus dubius Carapus home1 Carapus mourlanl' Jordanicus sagamianus Host Location ~nhost Remarks Mostly coelomic cav~ty; 29 infested holothuro~ds(89 insometimes respiratory trees or cloaca vestigated) (Arnold) Geographical area Source Mediterranean Sea (mostly western part) Emery (1880). Zanker (1940),Arnold (1953, 1956). Gustato (1977) Act~nopygaagasslzi Coelomlc cavity: some- 1 to 10 fishes W Atlantic from Brciz~l Linton (1907), (usual host); also Astimes respiratory trees holothurold ' (mostly to Nerth to Bermuda Parker (1926),Antlchopus rnultif~dus, 1); infestation level cona-Lopez Holothuria glaberrima, may reach 50 % (1956).Smith & (Smith et al.) Holothuria lentlginosa, Tyler (1969).Trott Holothuria princeps, (1970),Dawson (1971),Koster & Isost~chopus Caycedo (1979). badionotus, Thyone sp. Smith et al. (1981) (H) Tropical W Atlantic Putnam (1874) Unidentified Internal (Caribbeans) holothuroid Bedford (l899), Stichopus chloronotus Mostly coelomic cavlty, Infestation level 16 to Tropical Indo-W Pacific Fisher (1907). 88 % according to and Bohadschia argus also respiratory trees period of year Sivickis & Doman(usual host); also in Actay (1928),Muker(Smith) tionpyga mauritiana, ji (1932),Smith Holothuria atra, (1964).HipeauThelonota ananas, Jacquotte (19671, Stichopus tropicalis (H) Trott (1970. 1981), Trott & Trott (1972) Putnam (1874), Coelornic cavity of as1 to 2 flshes asTropical Indo-W Culcita schmideliana Simpson & Brown terolds; coelomic cavity teroidC1 Infestation Pacific and ('ulcita novae(1910),Mortensen and respiratory trees of level may reach guineae (usual hosts); (1923),Mukerji about 100 % (Moralso Acanthasterplan- holothuroids tensen, Trott 1970) (1932).Strasburg ci,Chonaster A single B. argus (1961).Smith granulosus, hotoreascontained 15 fishes ter linckl, Thromidia (1964),Hipeau(Meyer-Rochow seychellensis2 ( A ) Jacquotte (1967), 1977) Trott (1970),Trott Bohadschia argus (H) & Trott (1972). Cheney (1973). Jangoux (1974), Meyer-Rochow (1977, 1979) Infestation level may Tropical lndo-W Bohadschia argus (usu- Coelornic cavity Doleschall(1861). Pacific reach 30 % (Trott & Arnold (1956). a1 host); also Trottl Holothuria atra, Strasburg (1961). Holothuria scabra, Smith (1964),Trott Stichopus chloronotus, (1970),Trott & Trott (1972), Thelonola ananas (H) Cheney (1973) Culcita novaeguinea, Acanthasterplanci (A) NW Pacific (Japan: Infestation common Tanaka (1908), Holothuria monacaria Intestine ( 1 ) of in H. monacaria Sagami, Misaki) Yosii (1928a) (H) holothuroid; coelomic (Tanaka) Certonardoa semirecavity of asteroid Holothuna tubulosa, Stichopus regahs (H) gularis (A) ' Identified Carapus homei by many authors Previously identified Mithrodia fisheri [Jangoux 1974) Jacquotte 1967), and some authors suggested that juvenlle individuals could depend on their host for food (Jangoux 1974, Meyer-Rochow 1979). In contrast, Encheliophis spp, are considered true parasites which stay permanently in the echinoderm coelom feeding on host viscera (Strasburg 1961, Smith 1964). Effects of pearlfish infestations are said to be practically non-existent except for Enclieliophis spp. which 78 Dis. aquat. Org. 3: 59-83, 1987 Fig. 8. Carapus acus penetrating the holothuroid Holothuria tubulosa. (A) Recognition. (B) Twisting. (C) Penetration. (After Emery 1880) presumably cause castration by consuming the host's gonads. Carapus spp. cause only slight injury by piercing the host's digestive wall when entering or leaving the coelomic cavity. At low infestation levels the effect on the host should be negligible as pearlfishes do not always infest the same host, but shelter in the nearest suitable host found. 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