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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 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 , this has been proven for only a few . 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. castration by copepods and ascothoracids was reported several times in the literature. Parasitic 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 are known to live either temporarily or permanently in the of holothuroids and asteroids Carapid infestations do not seem to seriously affect the echlnoderms except for species permanently inhabihng the host's . Parasitic associations between echinoderms and , tardigrads, , 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 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 planci from and ), Tardigrada, Crustacea (Copepoda, Naples (Italy). Ganapati & Radhakrishna (1962) noted Cirripedia and ), 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- 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 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, ; 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 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 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 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) eri loosely attached South California) to body wall Cystimyzostom urn Metactinusrotundus Subcutaneous cysts on 1 worm cyst-'; N. Pacific (: 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 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 macro- Subcutaneous cyst on 1 to 2 worms Indo W Pacific (Sagami Hara & Okada ijirnai discus, calyx (upper surface) 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 (upper surface) 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 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 thusjaponicus 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) Protomyzostomum coniferus Encysted in gonads NW Paclfic (Sagami Fedotov (1925) astrocladi (0) Sea) Prolomyzostomum 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~ Gorgonocephalussp Bursae and gonads NW Pacific (Sagam1 Okada (1922) sagamiense (0) Sea) Pulvinom)~zostomurn phalan- D~gestivetract Infestation Mediterranean (Ban- Prouho (1892). pulvlnar glum, 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 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 which evolved together with pair of individuals. Gallicole species were termed clinoids. Myzostomids may be considered a unique 'deformative Arten' by Stummer-Traunfels (1926);they case of 'symbiogenesis' involving a whole class of induce conspicuous deformation of the host's ossicles organisms (e.g. Clark 1921, Stummer-Traunfels 1926, which considerably enlarge and adjust their shape to Prenant 1959). that of the myzostomid's shelter (Graff 1884, Wheeler About 25 species of myzostomids have been recorded 1896, Stummer-Traunfels 1926). Cysticole species as echinoderm parasites (Table l), the remaining belong to the genus Cystirnyzostomum; they behave species being frequently referred to 'free-living'. How- differently in that they build remarkable stalked or ever, these latter species are ectocommensals which unstalked cysts which are always subcutaneous (i.e. generally live unattached on the echinoderm body sur- located outside the host's ) and protrude into face. As pointed out by Stummer-Traunfels (1926), 3 the external medium (Fig. 1C). In many cases the cyst types of parasitic myzostomids may be distinguished wall is reinforced by a pavement of minute skeletal depending on whether they are gahcole, cysticole or plates (Graff 1884, Stummer-Traunfels 1926). Both endoparasltic. Endoparasitic species feed on the host's gallicole and cysticole myzostomids induce major host tissues, while cysticole and gallicole species are suspen- reactions affecting the crinoid skeleton either by mod- sion feeders which divert the water current produced by ifying size and shape of skeletal ossicles or by inducing the host's ambulacra (galls and cysts are most often the formation of additional skeletal plates. Such host located near the ambulacral grooves and always have 2 reactions invite further investigations. apertures allowing passage of directional water- Among the 8 species of endoparasitic myzosotomids, currents) (Prenant 1959). With rather few exceptions 4 infest ophiuroid or cnnoid gonads (respectively Pro- (e.g. Cystomyzostomum ijimai and Mesomyzostoma tomyzostomum spp. and Mesomyzostoma katol). katoi; respectively Fishelson 1974, Okada 1933),parasi- Gonad-infesting myzostomids cause at least partial tic myzostomids mostly infest bathyal echinoderms (in castration of their host. Fedotov (1916) reported that P. 200 to 1200 m water depth; Stummer-Traunfels 1926). polynephns infests the gonads of its ophiuroid host by This fact explains why these parasitoses have been so rupturing the wall of the bursae. Infestation may be anecdotically documented. very intense (up to 119 myzostomids per host) and 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 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 Van der Land 1975, Kristensen 1980). This symbiosis the gonads. The host reacts by progressively encysting has been reported only from Brittany, (Roscoff each parasite by a thick layer of dense connective and adjoining areas) where it is common. Very high tissue in which minute ossicles are seen. infestation rates occur in various populations, viz. 45, Fedotov suggested moreover that germ cell dedifferen- 80 and even 100% (respectively Bare1 & Kramers tiation may also occur. 1970, Kristensen 1980, Cuenot 1912). The number of According to Remscheid (1916), Mesom yzostoma T. synaptae per holothuroid is highly variable; it reichenspergiinfests gonads, while according to Prenant ranges from 2 to 3 up to 270 individuals (respectively (1959)it affects only the arm coelom of its crinoid host and Cuenot 1912, Kristensen 1980). The latter author feeds on coelomocytes. Asteriomyzostomum asteriae noted that infestation occurs only from May to (Fig. 2) is said to feed on the host's digestive contents; its October ( and juveniles are seen in June and occurrence supposedly increases the propensity of July). Kristensen presented photographic evidence asteroids to shed their arms (Marenzeller 1895a, b). that T. synaptae punctures the epidermal cells of L. Pulvinomyzostomum pulvinar might be more properly galliennei and sucks out their contents; such behavior classified as 'semi-parasitic'. It inhabits the crinoid's indicates a parasitic nature of the - digestive tract (anterior part) living intimately attached holothuroid association. Another tardigrade, Actinar- to the host's digestive epithelium without causing injury chus doryphorus, occurs on the body surface of the but diverting the flow of food particles entering the small European clypeasteroid echinoid Echinocyamus crinoid's mouth (Prenant 1959, West & West 1976). pusillus. According to Schulz (1935), A. doryphorus is a facultative parasite.

Agents: Tardigrada Agents: Crustacea, Copepoda The tardigrade Tetrakentron synaptae attached to the body surface of the synaptid a holo- There is little information regarding the relations thuroid Leptosynapta galliennei (Cuenot 1892, 1912, between echinoderms and closely associated Jangoux.Dlseases of Ech~nodermata- agents metazoans 6 3

copepods. This IS paradoxical given the high numbers parasite and do not result from a particular host reac- of copepods recorded as echinoderm symbiotes (e.g. tion (Mortensen 6 Stephensen 1918, Stephensen 1933, Humes 1980a, 1986, Stock 1968b). Table 2 lists species Goudey-Perriere 1979). demonstrated or presumed to be parasites. Although Copepods have been found to induce gall formation many ectoparasitic copepods of echinoderins have in the spines of echinothurid echinoids by Hansen been reported in the literature, the parasitic nature of (1902), Stephensen (1935bj and Stock (1968a). Accord- these assoclations has been proven for only 4 species, ing to Stock, the galls consist of loose calcareous viz. Cancenlla tubulata, Scottomyzon gibberum, material and always occur in the middle part of the Astel-ocheres lilljeborgi, and Ophiopsyllus reductus spines. Galls contain a pigmented ampula In which (respectively Carton 1968, Rottger 1969, Rottger et al. lives a slngle copepod. Another gallicole copepod, 1972, Emson et al. 1985). A. lilljeborgi is not particu- Pionodesmotes phormosomae, occurs inside the larly adaptated to parasitic . It is a motile copepod echinoid Hygrosoma petersi (Bonnier 1898, Koehler which uses its siphon-shaped mouth to feed on the skin 1898, see also Mortensen 1935) (Fig. 4). P. phor- of the asteroid sanguinolenta. S. gibberum mosomae lives in conspicuous inner galls located in the mainly lives on or near the pedicellarial rosettes of oral part of the host's coelomic cavity. The spherical, rubens where it seems to feed more deeply in calcified galls correspond morphologically to the host tissues (Fig. 3). 0. reductus lives firmly intracoelomic outgrowths of the echinoid body wall. attached to its ophiuroid host and feeds on surface Each gall opens on the host's outer body surface by a tissues. As for endoparasitic species, Pyefinch (1940) slit measuring 1 to 2 mm in length. The slit is protected concluded that Ophioica asyrnnietrjca found in the by the spines of the echinoid. At least 1 large female genital bursae of ophiuroids does not feed on the host's copepod was found in each gall. According to Bonnier tissues, and Changeux (1961) reported - based on (1898) the copepod does not prey on host tissues and casual observations - that the intracoelomic species obtains its food mostly from the outside through the slit Allantogynus delamarei 'browses' on the holothuroid's in the gall. Mortensen (1935) reported that empty galls mesothelium. progressively disappear: the slit enlarges, then the gall Host reactions against several species of parasitic wall normalizes, and finally new outer appendages copepods have been documented. Parasitism by develop. Scottomyzon gibberum induces the infested asteroid to The way in which intracoelomic copepods infest their produce dermal outgrowths which progressively host has been considered only for holothuroids and embed the copepods (Rottger 1969). Ophiuroid hosts asteroids. According to Paterson (1958) and Changeux tend to produce a fibrous, sometimes calcified cyst (19611, holothuroids are infested by larval copepods around symbiotes which live in their bursae (Pyefinch which cross the digestive wall either at the level of the 1940, Heegard 1951). However, encystment does not or at the level of the and respiratory occur with all bursal-infesting species (Stephensen trees. In asteroids, Carton (1974) showed that male 1935a). Mortensen (1933) recorded unidentified Botulosoma endoarrhenun~ penetrate the coelom of copepods inducing gall formation in the dorsal wall of Ech~nasterpurpureus through the respiratory papulae both Ophionlitrella corynephora and 0. hamata. Pater- Female B. endoarrhenum actually settle and grow in son (1958) reported that cysts, presumably produced by papulae, living in a kind of integumental cavity. the host, form around the intracoelomic copepod The effects of parasitic copepods on the of Cucumaricola notabilis. According to him, small spher- echinoderms have been considered practically only for ical or oval cysts contain juvenile parasites. These cysts Amphiurophilus amphiurae whlch parasitizes the are attached to the coelomic wall either near the cir- genital bursae of a brooding ophiuroid cumpharyngeal calcareous ring or on the cloaca near squamata (e.g Goudey-Perriere 1979, 1980). The the insertion of the respiratory trees; large amorphous author reported that the occurrence of the parasite does cysts containing mature copepods lie detached in the not affect the host's gonads but decreases the host's coelomic cavity. Jungersen (1914) reported that the fecundity by inhibiting the development of embryos ophiuroid Asteronyx loveni produces cysts around incubated in the ophiuroid genital bursae. Such inhibi- intracoelomic copepods. These cysts are attached on tion could result from competition for food between the ophiuroid's coelomic wall. Other intracoelomic parasite and embryos. Gonad destruction occurred, copepods have been noted to live often 'free in the however, in the ophiuroids Ophiomitrella clavigera and coelomic cavity' without seemingly inducing any host 0. corynephora parasitized by an unidentified copepod reaction (e.g. intracoelomic copepods of holothuroids). (Mortensen 1933). According to Emson et al. (1985), However, some bursal- or coelom-infesting copepods parasitic copepods on the ophluroid Ophiocomella may produce conspicuous swellings of the body wall of ophiactoides may decrease the tendency of the host to ophiuroids. These swellings follow the outline of the undergo (cross-disc division). 64 Dls aquat Org. 3. 59-83, 1987

Fig. 2. Asteriom yzostom um asten'ae, a myzostomid para- site of the pyloric caeca of the asteroid Sclerasterias neglec- fa. E: enlarged and deformed part of pyloric caeca harbor- ing the parasite; M: myzos- tomid. (Redrawn from Stum- mer-Traunfels 1903; sllghtly moddied)

Agents: Crustacea, Cirripedia on crinoids and ophiuroids (genera Ascothorax, Para- scothorax, Waginella) or as endoparasites in the Thorac~d cirripeds (i.e. barnacles) somet~mesattach coelomic cavity of asteroids and spatangoid echinoids to the outer body surface of some specles of regular (genera Ctenosculum, Dendrogaster, Ulophysema) echinoids (Lytechninus andmesus, niger, (Table 3). Waginella metacrinicola lives attached to the Stomopneustes variolans, spp.) and cnnoid calyx where it forms a conspicuous depression. clypeasteroids ( excentricus, Rotula It feeds on the 'fluids' of the host by piercing the body orbiculus). Either of 2 situabons may occur: (3)the wall wth its oral appendages (Okada 1926). Ophiu- attaches only loosely to the host's body wall roid-associated ascothoracids infest the genital bursae overlaping an intact cutaneous epithelium (Moore & producing marked swellings of part of the host disc McPherson 1963); (2) the barnacle is strongly attached (Fig. 5). According to Wagin (1946) Ascothorax ophioc- to the echninoid , the epidermis being destroyed tenis feeds on the cells of the bursal wall and probably (Giltay 1934, Dartevelle 1940, Boolootian 1958, 1964, also on the coelomic fluid of the host. As for Strachan 1970, Ganapati & Sastry 1972, Hurley 1973, intracoelomic ascothoracids, Brattstrom (1947) con- Bay-Schmith & Jana 1977). Barnacles Bdlanus spp. cluded that Ulophysema oeresundense directly absorbs have been not~ced on echlnoderms mainly by casual nutnents from the echinoid's coelomic fluid, while observations, except for the association between D. Wagin (1976) believed that coelomocytes form the bulk excentricus and Balanus concavus pacificus detailed by of the diet of Dendrogaster spp. (Fig. 6). Booloot~an (1964). There appears to be general agree- Echlnoderms react against intracoelomic ment that the activity of pedicellariae and spines would ascothoracids by forming a host envelope that avoid settlement of barnacle cyprids on the echino~d surrounds the mantle of the parasite. According to body wall (e.g Campbell & Rainbow 1977) However, Wagin (1946), this envelope consists of coelomocytes the reviewer believes that the failure of cyprids to settle and covers even the mantle opening which conse- results basically from the occurrence of a cutaneous quently is continually cleaned and smoothed by the epithelium which prevents larval fixation. Presumably ascothoracid's anterior appendages; Wagin claimed cypr~dsettlement can occur only when the echinoderm that such appendage activity actually corresponds to body wall is wounded. or when the epiderm is eroded the normal way of obtaining food by intracoelomic for some mechanical or pathological reasons (such as Dendrogaster species. However, investigations by bald-sea-urchin disease; Jangoux 1987a). Barnacles of Bresciani & Jespersen (1985) on Ulophysema oeresun- the genera Pachylasma, Scalpellum and Verruca have dense, parasitic in the coelom of cor- been noticed firmly attached to the stem or cirri of datum, showed that the envelope is made of choano- several species of stalked crinolds (Clark 1921). cyte-like cells and that consequently it should originate are 'naked barnacles' that parasitize from host mesothelium. A conspicuous host reaction octoralllan and echinoderms, except holo- also occurs in Parascothorax sinagodoides which lives thuroids. They have been found either as ectoparasites attached in the bursal cavity near the bursal slit of Jangoux Diseases of Echinodcr~nata agents metazoans 65

Table 2. Parasitic copepods from echinoderins (compiled from the sources indicated) Hosts: A, asteroid; C, crinoid, E, echinold; H, holothuroid; 0, ophiuroid

Copepod Host Location ~n- host Remarks Geographical area Source I. Harpacticoida Metis agasslzl Coelom~ccavlty Trop~calAtlantic Edxvards (1891). holothur~ae (H) (Bahamas) Humes (1980a)

T~sbefurcata Ocnus plano (H) Coelomlc cav~ty Almost 100 "'L of Med~terraneanSea Montlcelli holothuro~dsinvesti- (Naples) (18921, Humes gated were infested (1980a) (Monticelh) nsbe Holothuna stellatl (H) D~gestivetract (an- - Mediterranean Sea Humes (1957) holothuriae tenor part) (Banyuls)

11. Cyclopodia Allantogynus Holoihuna tubulosa, Antenor part of H tubulosa and H Mediterranean Sea Changeux delamare1 Holothul-~apoll, coelomic cavity frxed stellatl. about 2 (Banyuls, V~llefranche) (1958, 19611, Holothu~~astellat~ (H) on lnesothelium (occur copepods Stock & Kleeton also in wall of , holothuroid-' (82 in- (1963, Humes gonads, tentacular fested/ll7 investi- (1980a) ampullae, 01- gated); H poll is only Polran vesicles) occasronal host Aniphlul-ophllus Amphlphohs squamata Bursae About 12 infested NE Atlant~c(Roscoff); Fewkes (1887, amphiurae (0) oph~uroidsIn Roscoff Sorth Sea (Bergen); 1888). Herouard (Goudey-Perl-iere) NW Atlantic (IYoods (1906).Bocquet Hole region) (1952).Masson (1965).Goudey- Perriere (1979, 1980) Asterocheres Henncia sangu~nolen- Free on outer body Up to 24 copepods NE Atlantlc (Swedlsh Rottger et a1 hlljeborgi ta (AI surface asteroid- ' coast: Gullmarflord) 11972),Brun (1976) Botulosoma Ech~nasterpurpureus Respiratory papulae ( 2 Several dozen SW Indian Ocean Carton (1974) endoarrhenuni (A) copepod), coelo~nrc copepods as hero id^' (Madagascar Tulear) cavlty (dcopepod) Calvocheres Hygrosoma hoplacan- Galls on spine at ador- Two ~nfested NE Indran Ocean (off Stock (1968a) engell tha (E) a1 s~de ech~noidswith 6 and Celches, Tlrnor Sea) 3 copcpods, respec- tively Calvocheres Calver~osoniagrade, Galls on secondary Yot more than 1 9 N Indo-Pacific (off Hansen (19021, globosus Sperosoma qulncun- splnes or on spines of copepod gall (2 and Philippines, In Hal- Stephensen aale (E) adoral s~de 5 galls observed on 2 mahelra Sea; off SE (1935h),Stock infested hosts) Japan) (1968a) Calvocheres Hygrosoma peters1 (E) Galls on spines - N Pac~flc(SE of Japan) Stephensen oblongus (1935b)

Calypsanon St~chopusniono- Internal 1 C copepod ejected N\.V Indlan Ocean Stock (1968b). carinaturn tuberculatus (H) from host upon (Ethiopia: Dahlak Humes (1980a) collect~on Archipelago) Calypsanon Actinopyga Internal A. ~nlliar~s1s most SW Indian Ocean Humes & Ho leprum rnauntanla, Actlnopy- common host (76 (Madagascar Nossy- (19691, Humes ga miharis, Actlnopyga copepods - 43 X and Be) (1980a) lecanora (H) 33 9 -collected from 726 hosts)

Calypsarion rnarn~orata Internal 8 copepods (5 (' 3 Q) SLV Indian Ocean Humes & Ho sentosum (H) collected from 8 hosts (Madagascar: Nossy- (1969),Humes Be) (1980a) Calypana tubulosa, Esophagus and - Mediterranean Changeux changeuxr Holothuna stellati (H) gonoduct (Banyuls) (1961),Stock & Kleeton (1963). Hurnes (l980a) Cancer~lla Amphiphol~ssquamata Attached to outer body Infestation rate ca NE Atlantic (Brittany, Giard (1887). tubulata (0) surface 10 % (Giard) North Sea) Carton (1968)

Jangoux: Diseases of Echinodermata. agents metazoans 67

Table 2 (continued)

Copepod Host Location in host Remarks Geographical area Source

Diogen ella Holothilria mrxlcana, Internal (ejected from 2copepods(l h,1 9) Tropical Atlantlc Stock (1968b). splnicauda Actlnopyga agassrzl (H) H. niexlcana; Stock) from H n~exlcana (Cura~ao,Bahamas, Humes & Ho (Stock),46 copepods Jamaica, Puerto Rico) (1970),Humes (22d,24 9)from 119 (1980a) H. nlexrcana, 50 copepods (32 d. 18 Q)from 62 A agassizi (Humes & H01 Dlogenidium Holothuna glaberrima, Internal (ejected from 2copepods(l 6.1 9) Tropical Atlantic (Puer- Stock {1968b), deforme Holothuria arenicola, H. glaberrima; Stock) from 1 H. glabernma to Rico, Barbados, Humes & Ho (Stock);7 copepods Bahamas) (1971),Humes (H) (1 6.6 C)from87H. (1980a) arenicola, and 5 copepods(22.3 Q) from 24 H. mexicana

A ctinop yga agassizi, Coelomic cavity 9copepods (5 d,4 Q) Tropical Atlantic Edwards (1891), Holothuria mexlcana, from 3 H mexlcana (Bahamas, Cura~ao, Stock (1968b), Holothuria grlsea (H) (Stock);9 copepods Jamaica, Puerto Rico) Humes 61Ho (4 559)from69A (1971).Humes agdsslzi, 10 (1980a) copepods(4 8.6 V) from 29 H mexlcana. and 6copepods (4 d, 2 9)from 11 H. grlsea (Humes & Ho) Diogenidium Internal (ejected from 1 6 copepod from 1 Tropical Atlantic (Puer- Stock (1968b). spin ulosum badionotus (H) host; Stock) holothuroid (Stock); to Rico, Jamaica) Humes & Ho 4 copepods (2 d. 2 9) (1971).Humes from 17 holothuroids (1980a) (Humesl Dlogenldlurn Actinopyga agassizi (H) Internal 3 d copepods from 64 Tropical Atlantic Humes & Ho tectum holothuroids (Jamaica,Bahamas) (1971).Humes (1980a) Diaden~asetosun~(E) Esophagus 85 copepods col- New Caledonia Dojirl& Humes lected from 65 (1982) echinoids Enteroynathus . D~gest~vetract, rarely in 11 copepods, (2 d. Mediterranean Sea Giesbrecht comatulae Antedon blhda (C) coelomic cavity 9 9)from 12A (Banyuls, Naples); NE (1900),Grainger mediterranea Atlantic (Dublin, Ply- (1950), (Changeux & Dela- mouth) Changeux & De- mare Deboutteville) 1amare.De. boutteville (1956). Stock (1959) Enterognath us Decametra chad~vlcki, Digestive tract Not more than 4 Indian Ocean (Red Sea) Stock (1966) la tenpes Heterornetra savignyi, copepods found OUgometra serripinna (Cl Lecanunus Actinopyya lecanora Digestive tract - NW Pacific, (Philippi- Kossmann in testinalls (H) nes Bohol Island) (1877, quoted by Stock 1968b) Lecan urius , Internal 14 copepods (7 d. SW Indian Ocean Humes (1968) kossmannianus Actinopyga rnufleri (H) 7 Q)from 1'1 ,.\. lecan- (Madagascar: Nossy- ora and 8 cupepods Be) (4 6.4 9)from 205 A. m ullen Lecanurius , Digestive tract (post- 3 copepods (2 6.1 9) Tropical W Pacific Humes (l980a) planifron tahs (H) erior part) from 3A. echinites (New Caledonia; ) Lernaeosaccus Ophiacantha disjuncta Coelomic cavity (pre- Only 1 individual Antarctic (Palmer Heegard (1951) ophlacanthae (0) sumably) found Archipelago) 68 Dis. aquat. Org. 3: 59-83, 1987

Table 2 (continued)

Copcpod Host Locatlon in host Remarks Geographical area Source

Licholhuria , Internal (ejected from 4 copepods (2 d ,2 Q) W Indian Ocean (Red Stock (1968b), mandibulans . host; Stock) from12H.atra Sea- Eilat; Madagascar Humes & Ho Holothuna nobihs. (Stock);l88 cope- Nossy-Be) (1969),Humes Holothuria fuscopunc- pods(54 $, 134 ?) 1980a tata (H) from470H.scabra. 19 copepods (6 d. 13 '2) from24 H.nobilis, 139copepods(34 d, 105 P)from345 H. atra, and 1 d copepod from H. fuscopunc- tata (Humes& Ho) Ophioika Ophiornitrella calm- Bursae, forming swel- 1 to 3 copepods N Atlantic (S of Green- Stephensen appendiculata gera, Ophiacantha ling of host's tissue host-' (no more than land. S of Iceland. W of (1935b.1940) biden tata (0) 1 bursa infested Hebrides) host-') Ophioika Ophlacantha imago (0) Bursae, forming sl~ght Each of the 2 infested Antarctic seas asymmetrica swelling of host disc ophiuroids investl- gated had apair (d, P)of copepods en- cysted in 1 of their bursae Ophioika Ophiacan tha severa (0) Bursae, forming swel- Only 1 Q copepod SE Indian Ocean (Bali Stephensen ophiacanthae ling of host disc found Sea) (1933) Oph~o~ka Oph~acanthawblpara, Bursae (presumably) 8 copepods found Antarcticseas (South Heegaard tenujbrachra Ophiacantha disjuncta Georgia; Adelaide (1951) (0) Land)

Ophiopsyllus Ophiocomella ophiac- Oraland lateral parts of 40 to 65 Ohof host Jamaica Emson et al. reductus to~des(0) arm base population para- (1985) sitlzed depending on period of year. Usual- ly l to 3 copepods ophiuroid-'

Pinnodesmotes Hygrosoma petersll (E) Intracoelomic galls l or2(d, Q) NEAtlantic (Azores) Koehler (1898), phormosomae copepods gall-' Bonnler (1898) Scottomyzon Asteriasrubens(A) Galls in rosettes of Most asteroids are in- North Sea (off Helgo- Rottger (1969. gibberum pediceuariae fested; 4 to 175 land; Kattegat) see also Bare1 & copepods asterold-' Kramers (1977) Synapticola Polyplectana kefer- Coelomic cavity, diges- 5 copepods (2 6.3 9) Indlan Ocean (Amboi- Voigt (1892). teres stein^, Synapta tive tract (posterior free in coelomic cavi- na); W Pacific (Queens- Humes (1979) maculata (H) Part) ty (Voigt) land) 111. Monstrilloida Thespeslop- Ophiothrixfrag~l~s, Stomachfolds About50°h of 0. NorthSea (Gullmar- Bresciani& Lut- syllusparadoxus Ophiopholisaculeata aculeatainfested fjord) zen (1962) (0) IV. Incertaesedis kthrochor- Astrocharisgrac~lis(O) Galls at arm base Atleast2copepods N\VPacific(Ph~lippi- Mortensen & deumium (6,P ) found nes: Mindanao) Stephensen appen - (1918) diculosc~m Arthrochor- Asteromorpha koehleri Galls at arm base Only l copepod NEIndianOcean (Arn- Stephensen deumium (0) found boinal (1933) asteromorphae Codoba Oph~urameridionalis Calcif~edcystsinside 3 infestedophiuroids Antarcticsea (South Heegard (1951) discoveryi (0) bursa were found Georgia) Chordeum~~rm Asteronyxloven~(O) Cystsw-ithincoelomic Almost 100% lnfesta- NorthSea (Skagerrak) Mortensen obesum cavity (attachedcysts) tlon; no more than 1 (1912). copepod gaU-l Jungersen (1912,1914) Parachor- Amphipholissquamata Coelomiccavlty Only 1 copepodob- Med~terranean (Ville- Le Calvez deumium (0) sewed franche) (19381, Bocquet tetraceros (1952) ' Indentlfled Phormosoma uranus by Koehler (1898) Jangoux. D~seasesof Ech~nociermata.agents met~i~oans 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 by Ascothorax gigas (up to 7 genital bursae infested) causes occlusion of the ophiuroid mouth opening and 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 ston~ach contents. According to Brattstrom (1947) infesting larvae of Ulophysema oeresundense usually penetrate their host through the genital apertures and 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 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 lar- vae from a female Dendrogaster antarcticus into the asteroid coelorn is produced by rupturing of the mantle of the female. Larvae of Dendrogaster tasmaniensis Fig. 3. Scottomyzon gibberum, an ectoparasitic copepod of the penetrate the asteroid Allostichaster polyplax through asteroid Asterias rubens. (A) Female on tube-foot tip. (B& C) respiratory papulae; they infest the coelom and escape Copepods embedded in dermal tissue of a pedicellarial by crossing the asterold's wall and passing to rosette. a1 & d2: antennae; cp: crossed pedicelloria; mxp: maxilliped; 111x2: maxilla; S: spine; sp: straight pedicellaria. the outside through its mouth (Hickman 1959). (After Rottger 1969)

Ophiura quadrispina (Wagin 1964, 1976). The Agents: Crustacea, Malacostraca ophiuroid first encysts the parasite with its own tissues; then both cyst and parasite grow into the bursal cavity. Associations between amphipods and echinoderms The cyst is gradually ejected from the cavity through were reviewed by Vader (1978).A parasitic relation has the bursal slit. P. sinagodoides apparently has adapted been inferred by Shoemaker (1919) for Laphystiopsis its llfe cycle to the reaction of its host, as rejected cysts iridometra whlch l~vesembedded in the calyx of the as a rule include infesting ascothoracid larvae. Accord- crinoid Iridometra melpomene. Typical ectoparasihsm ing to Grygier (1985b) the asteroid-associated asco- occurs with the North Sea species Epimeria parasitica thoracid Endaster hamatosculum lives in conspicuous which feed on the outer tissues of the holothuroid inner galls located in the arm coelom of its host. Galls tremulus and the asteroid pulvillus are calcified and correspond morphologically to (Vader 1978). According to Ruffo (19) the intracoelomic outgrowths of the asteroid body wall; amphipods Jassia ocia and Aristias negletus occur at each gall opens on the host's outer body surface by a Banyuls (France, Mediterranean Sea) in the coelomic slit-like hole. cavity of the holothuroid Stichopus regalis and the Ascothoracids may castrate their host. Thus crinoid Antedon mediterranea, respectively. Ascothorax ophioctenis causes complete castration of An unexpected association between the tanaidacean Ophiocten sericerum, even though it does not feed on Exspina typica and 3 species of deep-sea holothuroids the host's gonads. Wagin (1946) found a single juvenile has been reported by Thurston et al. (1987). E. typica A. ophioctenis to Inhibit the activity of the germinal should be considered a facultative parasite; it occurs epithelium leading to a regression of the whole gonad. either in the intestine or body cavity of its host. Similar castration occurs in spatangoids infested with Crabs associated with and potentially harmful to large-sized Ulophysema oeresundense. According 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 cal- cified galls. (B) Section through gall containing an ovigerous female. p. out- er pore of the gall. (A after Koehler 1898; B after Bonnier 1898)

Pinnotheridae (genera , Ophisthopus, chilensis) are generally said to feed on host fecal pellets Pinnaxodes, Pinnixa, Pinnotheres), Parthenopidae (e.g. Miyake 1939, Fenucci 1967). Trophic relations (genera , Zebrida), and Portunidae (genus between E. pentagonus and the echinoid Lissocarcinus) (Table 4). Feeding habits of associated calamaris were studied carefully by Castro (1971) (see species of Ophisthopus, Pinnixa, Pinnotheres and also Suzukl& Takeda 1974). Males and small immature Lissocarcinus have not been reported. Presumably females inhabit the peristomeal region where they feed these forms do not feed on host tissues, nor cause any on epithelia1 tissue and , damage being in other detrimental effects, except to slightly wound the equilibrium with the regenerating capacities of the wall of the respiratory trees or of the cloaca (Tao 1930, host. Large mature females live in the and Jones & Mahadevan 1965). Species inhab~tingthe post- ingest material from fecal pellets and aggregates of erior part of the echinoid digestive tract (Echinoecus pigmented coelomocytes that migrate across the rectal convictor, E. pentagonus, E. ra thbunae and Pinnaxodes wall. Jangoux: Diseases of Echinodermata. agents metazoans 7 1

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) and from ophiuroids by Bartsch (1982; Ophiurolepis inornata)

Ascothorac~d Host Location Remarks Geographical area Source on/in host

Ascothorax Amphiclra belgrcar, An,- Bursae 1 ascothoracicl host-' Southern Atlantic [off Heegard (19511 bulbos~rs phicli-a mrcroplax (0) South Georgia) Ascothorax Oph~notusvictoriae (0) Bursae Up to 27 parasites South Sandwlch Is- Wagin (1968),Gry- 9lPs host-' (Grygier & Fratl) lands gier (1983~).Gry- gier & Fratt (1984) Ascothorax niicroplax (0) Bursae - South Georgia Gryg~er(1983~) mortensenl Ascothoras Ophiocten sericeum (0) Bursae Infestation rate: 0.7 to Barents Sea; Kara Sea; Djakonov (1914), oph~octenrs 3.4 '10 (Wagin) North Atlantic (around Stephensen Iceland and Faroe Is- (1935a). Wagin lands) (1946) Ascothorax Ophiosphalma ar- Bursae NE Atlantic Gryg~er(1983~) pilocauda tus migerun~(0) Ctenosculum evermanni (A) Coelomic c-avity Tropical 1VPacif~c Heath (1910),Wa- ha waiiense' (Hawail) ren (1981) Dendrogaster con- Coeloi~~i(cav~ty lnfestat~onlrlvel ca 2 '% Antarctic seas (off Ross Fisher (1930),Gry- antarcticus splcuus (A) Island) yier (1981) Dendrogaster Dlpsacaster sladeni, Coelom~ccavity 1 to 3 ascothoracids Southern Indlan Le Roi (1905, 1907), arborescens host-' (infestation not Ocean (off Cape Okada (1925) calan~aria(A) uncommon) (Okada) Town); Misaki (Japan) Dendrogasler Hlppastena cabfornrca Coelornlc cavity 1 observation NE Pacific (off F~sher(191 l), Gry- arbusculus (A) California) gier (1982) Dendrogasler groenlandi- Coelomic cavity - N Pacific (Bering Sea) Flsher (1930), Kor- arctica ca (A) schelt (1933), Wa- gin (1950),Grygier (1986) Dendrogasler Anasterias minuta (A) Coelomic cavity 5 asteroids infested SW Atlantic Grygier & Salvat argentinensrs (326 investigated) (1984) Dendrogaster Coelom~ccavity 3 infested H. Barents Sea (White Kn~powitsch(1891) astericola 'A) sangujnolenta (500 in- Sea) vestigated) Dendrogdster Asterjna burtoni (A) Coeloni~ccavity 1 to l0 ascothoracids Red Sea (Gulf of Achituv (1971) astennap asteroid- Aqaba) Dendrogaster scoparius Coelom~ccavlty 5 infested asteroids Misaki (Japan) Yosii 11928b) astropectlnls (A) (250 investigated) Dendrogaster Eremicaster tenebrarius, Coelorn~ccavity - Bering Sea Wagin (19571, bldd- beringensis Hyphalaster inermis (A) sen (19611, G~~gier (1985a) Dendrogasler Crossasterpapposus (A) Coelom~ccavity 3 Infested asteroids (13 NE Pacific Wagin (1950) dichotomus investigated) Dendorgaster obscurus (A) Coelornic cavity - Bering Sea Wagln (1950) dogie11 Dendrogaster Leptasteriaspolaris (A) Coelornic cavity Bering Sea; Gulf of St Wagin (1950). Gry- elegans Lawrence gier (1986) Dendorgaster Pedicellaster magister Coelomic cavity NE Paclhc (off Fisher (1928),Gry- fish eri megalabis (A) California) gier (1982) Dendrogaster Leptasterias fisher; (A) Coelomic cavity Berlng Sea Wagin (1950) iwanowi Dendrogaster Leptastenas fjshen' (A) Coelomic cavity Bering Sea Wagin (1950)

Dendrogaster luzonicus, Coelorn~ccavlty W. Pacific (Philippines, Le Roi (1905, 1907). Iudwigi Certonardoa seniire- Japan Sea; Tasmania) Yosii (1928b), gularis, be]- Kenny (1959) cheri (A) Dendrogaster Crossasterpapposus, Coelornic cavity Barents Sea; N Pacific Korschelt (1933). murmanensls endeca (A) (Okhotsk Sea) Wagin (1950) Dendrogaster Coscinastenas Coelonlic cavity - Misaki (Japan) Yosii (1928b) okadai calamaria (A) 72 Dis. aquat. Org. 3: 59-83, 1987

Table 3 (continued)

1Ascothoracrd Host Location on/in host Remarks Geographical area Source

Dendrogaster Leptaster~aspolans(A) Coelomic cavity Bering Sea Wagin (1950) onen talis Dendrogaster Poraniops~sinflata(A) Coelomic cavity NE Pacific (off Grygier (1982) punctata California)

1Dendrogaster Leptasterias hsheri (A) Coelomic cavity Bering Sea Wagin (1950) rarnosus Den drogaster Ctenodiscus crispatus, Coelomic cavity NW Pacific (Okhotsk Wagin (1950) rimskykor- Hippastena leiopelta Sea) sakowi (A) Dendrogaster Sidonaster vaneyi (A) Coelomic cavity 2 individuals In a Phihppine Seas Flsher (1919). sagittaria single asteroid Grygier (1985a) Den drogaster AUoshchasterpolyplax Coelomic cavity 1 ascothoracid as- Around Tasman~a Hickman (1959) tasmaniae (A) teroid-' (26 infested/ 11 6 investigated) Endaster Zeroaster carinatus (A) Intracoelomic galls 1 to 2 ascothoracids Ph~lippinesSea Grygier (1985b) hamatosculum asterold-' Parascothorax Ophlura quadrispina In cysts In bursae 1 to 9 ascothoracids Okhotsk Sea Wagin (1964) synagodoides (0) oph~uroid-' Ulophyserna Echinocardium cor- Coelomic cavity; more 1 to 9 (mostly 1) asco- North Sea (Scandina- Brattstrom oeresundense datum, Echlnocard~um rarely gonads, am- thoracids echinoid-' vian coast) (1936. 1938, flavescens, Brissopsis bulacral ampullae or (all stations together: 1946, 1947), lyrifera (E) axial sinus 814 infested/42 874 Bresciani & Jes- investigated) (Bratt- persen (1985) strom 1947);infesta- tion rate: 20 to 25 % (Brescian~& Jesper- sen) Ulophysema jeffreys~(E) Coelom~ccavity 1 or 2 ascothoricids N Atlantic (between Brattstrom pourtalesiae echinoid-' Norway. Spitzberg, (1937) Greenland and the Faroes) Waginella Metacrinusrotundus On stalk Sea of Japan Okada (1926, metacrin~cola (0) 19381, Gqfg~er (1983b) Waginella Metacrinus acutus, On cirri ; Sea Grygier (1983b) axotrernata Metacnnus angulatus, Metacrinus clngulatus (C) 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 & Gut-inhabiting species may produce deformations Takeda (1974) an obvious parasitic relation occurs that are sometimes very conspicuous. Verrill (1867) between the parthenopid Zebrida adamsi and reported that Pinnotheres chilensis considerably dis- several echinoid species (Table 4). the crab feeds on tends the rectal wall of the echinoid Coenocentrotus appendages, skin and peripheral dermal tissue of gibbosus to form a membranous cyst. The cyst attaches echinoids, producing conspicuous naked test areas. aborally to the echinoid body wall and extends near the Similar parasitic behavior also occurs with some mouth into the coelomic cavity. The anal area of the species of Dissodactylus living on clypeasteroid and host is depressed and distorded, and the echinoid test is spatangoid echinoids (Dexter 1977, Telford 1982, Bell & usually swollen on the side over the cyst (Fenucci Stancyk 1983). Telford (1982) reported that, depending 1967). P. chilensis parasitizes both C. gibbosus and on the species studied, the associated Dissodactylus Loxech~nusalbus. According to Mortensen (1943b) the take 50 to 100% of their diet from host tissues. Accord- crab does not induce major test deformation when ing to him differences in feeding habits can be attri- infesting L. albus. Similar but less conspicuous deforma- buted partly to the structure of host spines, viz,larger tions are caused also by Echinoecus convictor, E. pen- test areas are denuded if the spines of the echinoid host tagonusand E. rathbunae (respectively Bouvier & Seurat are more porous (Fig. 7). 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 senceun~ (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. bs bursa1 slit, bw ophiuroid body-wall, f , m dwarf male; t thoracic limbs (Redrawn from Wagin 1946: slightly modified) swellings of anal tube and periproct of their host. spine actlvity which decreased the ability of clypeas- According to Suzuki & Takeda (1974) and Dexter terolds to feed and to move about in aquaria, resulting (1977), infestations by ectoparasitic crabs often exert in a substantial increase in mortality. 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). Agents: Arachnida; Pycnogonida; Insecta This produces an obvious decrease in movements and spine activities of infested regular echinoids (Suzuki & The only arachnid reported to be an echlnoderm Takeda 1974). Dexter (1977) observed a reduction in 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 Dip- sacaster 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 infestation levels - from 3.6 to 83.7% - depending on (Viets 1939). Three specimens were collected in the host species. He presented evidence that A. echinoid material dredged by the Albatross off North ophiurophilus has the potential to feed on the host's Moluccas, Indonesia. Northing is known about host- tube feet. The occurrence of pycnogonids on echinoids associate relations. and asteroids has also been reported casually (Stock Pycnogonid-echinoderm associations have been 1981). noticed by several authors who generally suggested an An unexpected association between the marine ectoparasitic relation. Associations with holothuroids trichopter Philanisus plebeius and the asteroids were observed by Prell (1910; Pycnogonum littorale/ exigua and Patiriella regularis was reported frondosa) and Ohshima (1927; Lecy- by Anderson et al. (1976) and by Winterbourn & Ander- throrhynchus hilgendorfi/Holothuria lubrica). Accord- son (1980). Caddis fly eggs were found in the asteroid ing to Ohshima, a single host may carry up to 30 coelomic cavity. Singly or in small clumps, eggs occur pycnogonids which supposedly absorb the blood or loose within the coelom close to the peristome or 'body juice' of the holothuroid. An obvious pycno- enveloped in stomach folds. The authors presumed that gonid-ophiuroid association occurs at Aldabra (Indian eggs are deposited through the respiratory papulae Ocean) with 3 species of and the sea- and that newly hatched intracoelomic larvae leave the spider, Anoplodactylus ophiurophilus (Sloan 1979, host either via the same route or through the stomach Sloan et al. 1979, Stock 1979). Sloan reported various 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 Host Locat~onon/in host Remarks Geographical area Source

D~ssodactylus rosaceus Outer body surface lnfestatiorl level 25 to Tropical W Atlantic Telford (1982) caln~anr (E) (oral surface) 59 'X, (Florida, Jama~ca) Dlssodactyl~ls Mellita sexiesperforata Outer body surface Inlestation level 30 to Tropical W Atlantlc Telford (1978, cr~nitichelis (E) 60 'Vo (Barbados) 1982) D~ssoddctylus Encope emarginata (E) Outer body surface - Tropical W Atlantic Rathbun (1918) encopei (Florida to N Brazil) Dissodactylus Mellita Outer body surface Tropical E Pacific Rioja (1944) glass ell^ longifissa (E) (Mexican coast) D~ssodactylus Mellitalongifissa, En- Outer body surface Tropical E Pacific Glassell (1935) lockingtoni cope mlcropora, En- (proximal porhon of (Northern Mexico) cope grandis, Encope pasterior interam- californica (E) bulacral lunule) Dissodactylus Mellita quinquiesper- Outer body surface 13 crabs collected NW Atlantic Rathbun (1918). mellitae forata, Ech~narachnius from 50 M. qu~nqujes- (Massachusetts to Pearse (1947), parma (E) perforata (Pearse);al- Florida) Johnson (l952), most 100 9b inlesta- Gray et al. tion (Johnson) (1968),Telford (l982) Dissodactylus Encope stokesi, En- Outer body surface 1 to 2 crabs host-'; in- Tropical E Pacific Dexter (1977) nitidus cope n~icropora(E) (near lunule or margi- fest mostly large (Panama) nal slits) echinoids (diamete~ > 30 mm) Dissodactylus , Outer body surface Infestation level 79 to Tropical W Atlantic Telford (1978, pn'mit~vus Plagiobnssusgrandis (mostly on oral surface) 100 YO (Barbados,Jamaica) 1982) (E) D~ssodactylus i~&talongifjssa (E) Outer body surface - Tropical L: Pdciflc Rioja (1944) smlthi (Mexican coast) Dissodactylus Encope stokesi, En- Outer body surface More otten on E mic- Trollicdl E Pacific Dexter (1977) xan tusl cope micropora (E) (near lunule or margi- ropora (i'dn~mdj nal slits) Echinoecus Echinothrix Internal swelling of - Tropical Indo-West Bouvler & Seu- convictor IEI anal tube Pacific (Gambier Is- rar (1905).Mor- 1dnd; Mauritius) tcnsen (1940) Echlnoecus Pseudocentrotus de- Outer body surface 10 crabs collected on Tropical Indo-W Castro (1971, pentagonus pressus, Anthocidaris (near peristome) 48 P. depressus Pacific (Andaman. 1978). Suzuki & crassispina, Echino- (Suzuk~& Takeda) Hawaii, Japan: coast of Takcda (1974). ihrix calamaris, Uchida) Sastry (1977, Echlnothru. diadema 1981) (El Ech~neocus Echlnothrix calamaris, Internal swelling of the NW Pacific (Ogasa- Miyake (1939) rathbunae Phyllacanthus dubius anal tubes wara Islands) (El Lissocarcin us (H) Mainly respiratory 11 holothuroids in NW Pacific (Philippi Trott & Garth orbicularus trees, also cloaca fested (61 investi- nes: Puerto Galera) (1970) gated) Llssocarcin us Holothuna sp. (H) Cloaca - N Indian Ocean (An- Chopra (1931) ornatus daman Sea) Ophisthoptrs Parastlchop 11s califor- Cloaca Ocfur also in other in- NE Pacific (California) Rathbun (1918) transversus n~cus(H) (e.y , bivalve molluscs) Pinnaxodes , Partly embedded in 1 to 2 crabs SE Pacific. (Ecuador to \/errill (18671, chilensis' Coenocen trotus rectum wall echinoid-'. Infesta- Patagonia) Ortmann (1894), gibbosus (E) tion generally very Meissner (1896). high (90 to 100 "/.) Clark (l910), (Verrill, Clark. Rath- Rathbun (1918). bun, Schwabe) Schwdbe (1936), Mortensen (1943b), Fenucci (1967) Pinniva iMolpadja arenicola (H) Cloaca - NE Pacific (California) Rathbun (1918) barnharti Pinnixa timida Caudina chilensis (H) Cloaca Infestation level 75 % NW Pacific (Japan: Tao (1930) Mutsu Bay) Pinnotheres Holothuria scabra (H) Cloaca and respiratory 10 holothuroids In- N Indian Ocean (In Chopra (1931), decanus trees; rarely in the fested (25 investi- dian coast) Jones & coelomic cavity gated) Mahadevan (1965). Adithiya (1969) 76 Dis. aquat. Org. 3. 59-83. 1987

Table 4 (continued)

Reptant Host Location on/in host Remarks Geographical area Source

Pinnotheres Unidentified Respiratory trees - NW Indian Ocean (An- Chopra (1931) setnai holothuroid daman Sea) Pinnolheres Achnopyga mauritia- Cloaca NW Indian Ocean (An- Doflein (1904). villosissirnus na, Actlnopyga lecan- daman Sea; west coast Chopra (1931) ora (H) of Sumatra) Zebnda adams~ Diadema selosum, Outer body surface As a rule 1 crab NW Pacific (Japan: Rathbun (1910). Trjpneust gratilla, (near ambitus) echinoid-' (11 crabs Suruga Bay; coast of Mortensen pileoius, collected) (Suzukl & Thailand) (1943a),Suzuki Toxopneustes elegans, Takeda) & Takeda ijimai, (1974),Daniel & Salmacis sphaeroides Krishnan (1978) (E) ' Often identified Fabia chilensis

eggs are found in the coelom during most of the year, 1940, Arnold 1956, Trott 1970, Van Meter & Ache 1974, the infestation level being about 10%. Smith et al. 1981, Trott 1981). species associate with holothuroids and asteroids are listed in Table 5. Agents: Pisces Most authors report that the stomach contents of Associations between echinoderms and carapid spp. from echinoderm coelomic cavities do not fishes () are of particular interest as they show remains of host tissue (e.g. Trott 1970, Smith et al. concern - at least in a few cases - typical parasitic 1981). Carapus spp. are predators. Basically, they use relations between an host and a the host's coelom as shelter (Fig. 8). They leave it in associate. The systematics, general biology, and ecol- order to catch prey, feeding primarily on . ogy of pearlfishes are documented satisfactorily, and Casual observations of feeding by Carapus on the some species have been studied in detail (e.g. Zankert host's gonad have been reported however (Hipeau-

I i Fig. 7 Dissodactylus crinitichelis, an ectoparasitic crab of the clypeastroid echinoid Mellita sexiesperforata. The crab is seen beside the devastated area where it had been feeding. (After Telford 1982) Jangoux: Diseases of Echinodern~ata: agents metazoans 77

Table 5. Fishes endosymbiot~c in echinoderms (compiled from the sources indicated). Hosts. A, asteroid; H, holothuroid

Fish Host Location ~nhost Remarks Geographical area Source

Carspils actrs Holothuna tubulosa, Mostly coelomic cav~ty; 29 infested Mediterranean Sea Emery (1880). Stichopus regahs (H) sometimes respiratory holothuro~ds (89 in- (mostly western part) Zanker (1940), Ar- trees or cloaca vestigated) (Arnold) nold (1953, 1956). Gustato (1977) Carapus Act~nopygaagasslzi Coelomlc cavity: some- 1 to 10 fishes W Atlantic from Brciz~l Linton (1907), bermudensis (usual host); also As- times respiratory trees holothurold ' (mostly to Nerth to Bermuda Parker (1926), An- tlchopus rnultif~dus, 1); infestation level cona-Lopez , may reach 50 % (1956). Smith & Holothuria lentlginosa, (Smith et al.) Tyler (1969). Trott Holothuria princeps, (1970), Dawson Isost~chopus (1971), Koster & badionotus, sp. Caycedo (1979). (H) Smith et al. (1981) Carapus Unidentified Internal Tropical W Atlantic Putnam (1874) dubius holothuroid (Caribbeans) Carapus Mostly coelomic cavlty, Infestation level 16 to Tropical Indo-W Bedford (l899), home1 and Bohadschia argus also respiratory trees 88 % according to Pacific Fisher (1907). (usual host); also in Ac- period of year Sivickis & Doman- tionpyga mauritiana, (Smith) tay (1928), Muker- Holothuria atra, ji (1932), Smith Thelonota ananas, (1964). Hipeau- Stichopus tropicalis (H) Jacquotte (19671, Trott (1970. 1981), Trott & Trott (1972) Carapus schmideliana Coelornic cavity of as- 1 to 2 flshes as- Tropical Indo-W Putnam (1874), mourlanl' and ('ulcita novae- terolds; coelomic cavity teroidC1 Infestation Pacific Simpson & Brown guineae (usual hosts); and respiratory trees of level may reach (1910), Mortensen also Acanthasterplan- holothuroids about 100 % (Mor- (1923), Mukerji ci,Chonaster tensen, Trott 1970) (1932).Strasburg granulosus, hotoreas- A single B. argus (1961).Smith ter linckl, Thromidia contained 15 fishes (1964), Hipeau- seychellensis2 (A) (Meyer-Rochow Jacquotte (1967), Bohadschia argus (H) 1977) Trott (1970), Trott & Trott (1972). Cheney (1973). Jangoux (1974), Meyer-Rochow (1977, 1979) Bohadschia argus (usu- Coelornic cavity Infestation level may Tropical lndo-W Doleschall(1861). a1 host); also reach 30 % (Trott & Pacific Arnold (1956). Holothuria atra, Trottl Strasburg (1961). Holothuria scabra, Smith (1964),Trott Stichopus chloronotus, (1970), Trott & Thelonola ananas (H) Trott (1972), Culcita novaeguinea, Cheney (1973) Acanthasterplanci (A) Jordanicus Holothuria monacaria Intestine (1)of Infestation common NW Pacific (Japan: Tanaka (1908), sagamianus (H) holothuroid; coelomic in H. monacaria Sagami, Misaki) Yosii (1928a) Certonardoa semire- cavity of asteroid (Tanaka) gularis (A) ' Identified Carapus homei by many authors Previously identified fisheri [Jangoux 1974)

Jacquotte 1967), and some authors suggested that stay permanently in the echinoderm coelom feeding on juvenlle individuals could depend on their host for food host viscera (Strasburg 1961, Smith 1964). (Jangoux 1974, Meyer-Rochow 1979). In contrast, Effects of pearlfish infestations are said to be practi- Encheliophis spp, are considered true parasites which cally non-existent except for Enclieliophis spp. which 78 Dis. aquat. Org. 3: 59-83, 1987

Anderson, D. T., Fletcher, M. J., Lawson-Kerr, C. (1976). A marine caddis fly, Philanius plebeius, ovipositing in a Patiriella exigua. Search 7: 483-484 Arendt, Y A. (1985) Biotic relations of crinoids Paleontol. Zh. USSR 1982(2): 69-76. (Russian; English summary) Arnold, D. C. (1953). Observations on Carapus acus (Briinnich), (Jugulares. Carapidae). Pubbl. Staz. zool. Napoli 24: 152-166 Arnold, D. C. (1956). A systematic revision of the fishes of the teleost family Carapidae (Percornorphi, Blennioidea), with descriptions of two new species. Bull. Br. Mus. nat. Hist. 4: 247-307 Barel, C. D., Kramers, P. G. (1970). Notes on associates of echinoderms from Plymouth and the coast of Brittany. Proc. K. ned. Akad. Wet. (C)73: 159-170 Barel, C. D., Kramers, P. G. (1977).A survey of the echinoderm associates of the north-east Atlantic area. 2001. Verh., Leiden 156. 1-159 Rartsrh, I. (1982). Ophiuroidea (Echinodermata) from the Patagonian shelf. Mitt. zool. Mus. Hamb 79: 211-250 Bay-Schmith, E., Jana, C. (1977). Balanus tintinabulum con- cinnus Darwin, 1854, epibionte poco comun del erizo negro, Tetrapygus niger (Molina. 1782). Boln Soc. Biol. Concepcion 51. 59-61 Bedford, F. P. (1899). Holothurians. A. Willey's 2001. Results (London) 2: 141-150 Bell, J. L., Stancyk, S. E. (1983). Population dynamics and reproduction of Dissodactylus rnellitae (Brachyura: innotheridae) on its host, Mellita quinquiesper- Fig. 8. Carapus acus penetrating the holothuroid Holothuria forata (Echinodermata). Mar. Ecol. Prog. Ser. 13: 141-149 tubulosa. (A) Recognition. (B) Twisting. (C) Penetration. (After Bocquet, T (1952). Copepodes semi-parasites et parasites des Emery 1880) echinodermes de la region de Roscoff. Bull. Soc. zool. Fr. presumably cause castration by consuming the host's 77: 495-504 Bonnier, J. (1898). Note sur le Pionodesrnotes phormosomae, gonads. Carapus spp. cause only slight injury by pier- copepode parasite du Phormosorna ulanus. Res. Camp. cing the host's digestive wall when entering or leaving scient. Albert I Monaco 12 (Appendice): 61-66 the coelomic cavity. At low infestation levels the effect Boolootian, R. A. (1958). Notes on an unexpected association on the host should be negligible as pearlfishes do not between a common barnacle and an echinoid. Bull. Sth. always infest the same host, but shelter in the nearest Calif. Acad. Sci. 57: 91-92 Boolootian, R. A. (1964). The occurrence of Balanus concavus suitable host found. However, in a higher infested on the test of . Bull. Sth. Calif. echinoderm population (high infestation levels were, Acad. Sci. 63: 185-191 for example, recorded by Mortensen [l9231 and Trott Bouvier, E. L., Seurat, G. (1905). Eumedon convictor crabe [l9701 for Carapus mourlani inhabiting Culcita novae- commensal d'un oursin. C.r. hebd. Seanc. Acad. Sci., Paris 140: 629-631 guineae) chances increase that a given host is infested Brattstrom, H. (1936). Ulophysema oeresundense n. gen. et sp. regularly; hence repeated loss of coelomic fluid and eine neue Art der Ordnung Cirripedia Ascothoracida. Ark. successive wound repairs have to be considered. 2001. 28A(23): 1-10 Brattstrom, H. (1937). On the genus Ulophysema Brattstrom with description of a new species from east Greenland. Acknowledgements. I thank Drs M. J. Grygier, Kinne and 0. Meddr Gronland 118(7): 1-23 J. M. Lawrence for information and crit~cism;Dr C Massin for Brattstrom, H. (1938). Ascothoracica, eine fur die norwegische helping with literature research; N. Biot, Dr G. Coppois, M. Fauna neue Ordnung parasitischer Crustaceen. Det. Kong. Doize and M. Klinkert for assisting in the preparation of the Norske Vidensk. Selsk. Forh. 11(6): 22-25 manuscript and illustrations. Brattstrom, H. (1946). Observations on (For- bes) in the Gullmar Fjord. Ark. Zool. 37A(18): 1-25 Brattstrom, H. (1947). On the ecology of the ascothoracid LITERATURE CITED Ulophysema oeresundense Brattstrom. Lunds Univ. Arsskr. (N.F. 2) 43(7): 1-75 Achituv, Y. (1971). Dendrogaster asterinae n. sp., an Brescianl, J., Jespersen, A. (1985). The ultrastructure of the ascothoracid (Cirripedia) parasite of the starfish integument of Ulophyserna oeresundense Brattstrom, 1936 burtoni of the Gulf of Eilat. Crustaceana 21: 14 (Crustacea, Ascothoracida]. J. crust. Biol. 5: 146-159 Adithiya, L. (1969). Some notes on the of Holothuria. Bresciani, J., Liitzen, J. (1962). Parasitic copepods from the Loris 11: 385-388 west coast of Sweden includ~ngsome new or little known Ancona-Lopez, A. A. (1956). Occorrencia de Carapus Raf. specjes. Vidensk Meddr dansk naturh. Foren. 124: (= Fierasfer Oken) no Brasil. Pap. Avuls. Dpto Zool. 367408 (S. Paulo) 12: 389-397 Brun, E. (1976). Ecology and taxonomic position of Henricia Jangoux: Diseases of Echino dermata: agents metazoans 79

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Editonal responslblllty Managing Edltor; accepted for pnntlng on June 3, 1987