A REVIEW OF THE SPECIES OF PROTOZOAN EPIBIONTS ONCRUSTACEANS.I.PERITRICH CILIATESBYGREGORIO FERNANDEZ-LEBORANS and MARIA LUISA TATO-PORTODepartamento de Biologia Animal I(Zoologia),Facultad de Biologia,Pnta9a,Universidad Complutense,E-28040Madrid,SpainABSTRACTAn updated inventory of the peritrich(Protozoa,Ciliophora)epibiont species on crustaceans has been carried out.Data concerning268epibiont species,their taxonomic position,and the various crustacean basibionts were considered.The overview comprised in this study may be of use in further surveys of protozoan-crustacean epibiosis.RESUMENSe ha realizado un inventario actualizado de las especies de peritricos(Protozoa,Ciliophora) epibiontes en crustáceos.Se han considerado los datos concernientes a268especies epibiontes,su posición taxonómica,y los diferentes crustáceos visión general que comprende este estudio puede ser utilizada en futuras investigaciones sobre la epibiosis protozoos-crustáceos.INTRODUCTIONEpibiosis is a facultative association of two organisms:the epibiont and the basibiont(Wahl,1989).The term“epibiont”includes organisms that,during the sessile phase of their life cycle,are attached to the surface of a living substratum, while the basibiont lodges and constitutes a support for the epibiont(Threlkeld et al.,1993).Both concepts describe ecological functions(Wahl,1989).Several crustacean groups,cladocerans,copepods,cirripedes,isopods,amphi-pods,and decapods,include forms that are hosts for macroepibiont invertebrates (Ross,1983),and for protozoan microepibionts of the phylum Ciliophora:apos-tomatids,chonotrichids,suctorians,peritrichs,and heterotrichs(Corliss,1979; Small&Lynn,1985).The study of ciliate epibionts on crustaceans began in the last century.Bütschli(1887-89)made a compilation from former publications.After-wards,other authors(Keiser,1921;Kahl,1934,1935;Precht,1935;Raabe,1947; c®Koninklijke Brill NV,Leiden,2000Crustaceana73(6):643-683644G.FERNANDEZ-LEBORANS&M.L.TATO-PORTONenninger,1948)not only described epibiont species,but proposed explanations for the processes of epibiosis.A review of the protozoan epibionts found on de-capod crustaceans was carried out by Sprague&Couch(1971).Green(1974), in a study of the epibionts living on cladocerans,pays considerable attention to protozoan species.Ho&Perkins(1985)have focused on the epibionts found on copepods.In other contemporary and also earlier works,the following aspects have been taken into account:(1)speci city between ciliates and their crustacean basi-bionts(Evans et al.,1981;Batisse,1986,1992;Clamp,1991);(2)the morpholog-ical and physiological adaptations of the epibionts(D’Eliscu,1975;Batisse,1986, 1994;Fenchel,1987;Clamp,1991;Lom&De Puytorac,1994);(3)the effects pro-duced by the epibionts on the crustaceans(Herman et al.,1971;Turner et al.,1979; Kankaala&Eloranta,1987;Nagasawa,1988);(4)the possible use of epibionts for the assessment of water quality(Antipa,1977;Henebry&Ridgeway,1979;Scott &Thune,1986);(5)the implications of protozoan epibionts on cultures of vari-ous species of crustaceans(Overstreet,1973;Johnson,1977,1978;Lightner,1977, 1988;Couch,1978;Scott&Thune,1986;V ogelbein&Thune,1988;Camacho& Chinchilla,1989);and(6)the organization of the epibiont communities on plank-tonic crustaceans(Threlkeld et al.,1993).Despite the fact that there is a considerable amount of information about the protozoan epibionts on crustaceans,since the works of Sprague&Couch(1971), Green(1974),and Ho&Perkins(1985),which relate to speci c crustacean groups, no further general reviews have appeared.Several new species of protozoan ciliate epibionts have recently been described(Dovgal,1985;Batisse,1992;Fernandez-Leborans&Gomez del Arco,1996;Zhadan&Mikrjukov,1996;Fernandez-Leborans et al.,1996,1997),and some of these are peritrich ciliates(Matthes& Guhl,1973;Bierhof&Roos,1977;Jankowski,1986;Dale&Blom,1987;Clamp, 1990,1991;Threlkeld&Willey,1993;Hudson&Lester,1994;Stoukal&Matis, 1994;Foissner,1996).The purpose of this work is to provide an up-to-date review of the peritrich ciliates living as epibionts on crustaceans:268species have been considered in this compilation,which may contribute data for studies of epibiosis in crustaceans.CRUSTACEAN PROTOZOAN EPIBIONTS,I.PERITRICH CILIATES645RESULTS1)Phylum CILIOPHORA Do ein,1901Class OLIGOHYMENOPHOREA De Puytorac,Batisse,Bohatier,Corliss, Deroux,Didier,Dragesco,Fryd-Versavel,Grain,Grolière,Hovasse,Iftode,Laval,Roque,Savoie&Tuffrau,1974Subclass P ERITRICHIA Calkins,1933Order S ESSILIDA Kahl,1933Family Epistylididae Kahl,1935Genus Rhabdostyla Kent,1880( g.1)R.bosminae Levander,1907.On the cladoceran Bosmina sp.R.conipes Kahl,1935.On the cladoceran Daphnia sp.Fresh water.On the cladocerans Daphnia magna,D.longispina and Scapholeberis mucronata (cf.Green,1957,1974).R.cyclopis Kahl,1935.On the copepod Cyclops sp.Fresh water.R.cylindrica Stiller,1935.On the cladoceran Leptodora ke Balaton (Hungary).On the cladoceran Leptodora kindtii.Denmark(Green,1974).R.hungarica Stiller,1931.On the cladoceran Leptodora ke Balaton (Hungary).R.globularis Stokes,1890.On the cladoceran Bosmina longirostris and on Diaphanosoma brachyurum.Germany(Nenninger,1948).R.invaginata Stokes,1886.On the ostracod Cypris sp.R.sessilis Penard,1922.On the copepod Cyclops sp.Fresh water.R.pyriformis Perty,1852(cf.Kahl,1935;on Entomostraca).On the clado-ceran Daphnia longispina(cf.Nenninger,1948).On the cladoceran Daph-nia hyalina(cf.Sommer,1950).On Daphnia pulex and Ceriodaphnia reticu-lata(cf.Hamman,1952).On Daphnia magna,D.pulex,D.cucullata,Simo-cephalus vetulus,Ceriodaphnia reticulata,and Leptodora kindtii(cf.Green, 1953).On Daphnia magna(cf.Green,1955).On Daphnia magna andD.longispina(cf.Green,1957).On Daphnia atkinsoni,D.hyalina,D.lon-gispina,D.curvirostris,D.obtusa,Ceriodaphnia laticaudata,and C.pulchel-la(cf.Green,1974).R.vernalis Stokes,1887.On the copepod Eucyclops agilis(cf.Henebry& Ridgeway,1979).1)For authors and dates of species of Crustacea mentioned herein,see separate section,below.646G.FERNANDEZ-LEBORANS&M.L.TATO-PORTOFigs.1-2.1,Rhabdostyla(R.pyriformis,after Green,1957);2,Epistylis(E.gammari,after Precht,1935).Rhabdostyla sp.Bierhof&Roos,1977.Between the spines at the end of the telson on Gammarus tigrinus.Germany.Rhabdostyla sp.Weissman et al.,1993.On the copepod Acartia hudsonica. Genus Epistylis Ehrenberg,1832( g.2)E.anastatica(Linnaeus,1767)(cf.Kent,1881).Syn.:Vorticella anastatica L.,1767.On Entomostraca and freshwater plants.On cyclopoid copepods and Daphnia pulex(cf.Green,1974).E.astaci Nenninger,1948.Fresh water.On the gills of the decapod Astacusastacus(as A. uviatilis)(Germany).On A.leptodactylus(cf.Stiller,1971).On the gills of Austropotamobius torrentium(cf.Matthes&Guhl,1973).E.bimarginata Nenninger,1948.Fresh water.On the appendages of Astacusastacus(as A. uviatilis).Germany.E.branchiophila Perty,1852.Syn.:E.formosa Nenninger,1948.On theparasitic copepod Lernaea cyprinacea,in freshwater environments of South Africa(Van As&Viljoen,1984).E.breviramosa Stiller,1931.On the antennal lament of the cladoceran Daph-nia ke Balaton(Hungary).On the copepod Cyclops sp.,Czechoslovakia (Srámek-Husek,1948).On the cladocerans Bosmina longirostris and Alona af nis(cf.Green,1974).E.cambari Kellicott,1885.On the gills of the decapod Cambarus sp.(NE ofU.S.A.).On the maxillae of the cray sh Astacus leptodactylus(fresh water) (cf.Matthes&Guhl,1973).E.crassicollis Stein,1867.On freshwater Entomostraca and on the pleopodsand gills of cray sh.On the gills of Astacus astacus(as A. uviatilis),andCRUSTACEAN PROTOZOAN EPIBIONTS,I.PERITRICH CILIATES647 the maxillae,maxillipeds,and gills of A.leptodactylus,in Europe(Matthes& Guhl,1973).E.cyprinaceae Van As&Viljoen,1984.On the parasitic copepod Lernaea cyprinacea(fresh water,South Africa).E.daphniae Fauré-Fremiet,1905.On the cladoceran Daphnia sp.On Daphnia magna(cf.Nenninger,1948).On the copepod Boeckella triarticulata(New Zealand)(Xu&Burns,1990).On the cladoceran Moina macrocopa in an urban stream.E.diaptomi Fauré-Fremiet,1905.On the copepod Diaptomus sp.E.digitalis Ehrenberg,1838.On the copepod Cyclops sp.E.epibarnimiana Van As&Viljoen,1984.On the parasitic copepod Lernaea barnimiana(fresh water,South Africa).E.fugitans Kellicott,1887.On the cladoceran Sida crystallina.North America.E.gammari Precht,1935.On the antennae of the gammarid Gammarus sp. (Kiel channel).On the proximal part of the rst antenna and,less commonly, on the second antenna of Gammarus oceanicus and G.salinus.In the Baltic Sea and areas of Norway(Fenchel,1965).On the rst antenna of Gammarus tigrinus(cf.Stiller,1971).E.halophila Stiller,1942.On the cladocerans Daphnia longispina and D.pulex (Lake Cserepeser,Hungary).E.harpacticola Kahl,1933.On harpacticoid copepods in the Kiel channel. E.helenae Green,1957.On the cladocerans Daphnia pulex,D.magna,D.ob-tusa,D.longispina,D.curvirostris,Ceriodaphnia pulchella,C.reticulata, ticaudata,Moina macrocopa,M.micrura,Chydorus sphaericus,Simo-cephalus serrulatus,and S.vetulus(cf.Green,1957,1974).On Daphnia magna(cf.Nenninger,1948).On Ceriodaphnia reticulata and Simocephalus vetulus(cf.Matthes,1950).E.humilis Kellicott,1887.On the gammarid Gammarus sp.and other Ento-mostraca.custris Imhoff,1884.On the pelagic copepod Cyclops sp.On the buccal appendages of the branchiopod Lepidurus apus(freshwater areas near Vienna, Austria)(Foissner,1996).E.magna V an As&Viljoen,1984.On the parasitic copepod Lernaea cypri-nacea(fresh water,South Africa).E.niagarae Kellicott,1883.On the body surface of cray sh(Niagara River, U.S.A.).On the antennae and body of the European cray sh Astacus lep-todactylus,on Austropotamobius torrentium,and on Orconectes limosus(as Cambarus af nis)(cf.Matthes&Guhl,1973).On the surface of the copepod648G.FERNANDEZ-LEBORANS&M.L.TATO-PORTOEucyclops serrulatus,and on the cladocerans Daphnia pulex,D.rosea,Cerio-daphnia reticulata,and Scapholeberis mucronata(lakes of Colorado,U.S.A.) (Willey&Threlkeld,1993).E.nitocrae Precht,1935.On the third pereiopod of Gammarus tigrinus(cf.Bierhof&Roos,1977).E.nympharum Engelman,1862.On cladocerans(Nenninger,1948).On Cy-clops sp.(cf.Foissner&Schiffman,1974).On the branchiuran Dolops ra-narum(cf.Van As&Viljoen,1984).E.ovalis Biegel,1954.On the gnathopods of Gammarus tigrinus.On the thirdpereiopod of the gammarid Gammarus pulex,and on the spines at the end of the third uropod of Gammarus tigrinus(cf.Bierhof&Roos,1977).E.plicatilis Ehrenberg,1838.On the copepods Eucyclops agilis,Cyclopsvernalis,and C.bicuspidatus(Ashmore Lake,Illinois,U.S.A.)(Henebry& Ridgeway,1979).E.salina Stiller,1941.On the rst and second antennae,coxae,and gills of thegammarid Gammarus pulex(cf.Bierhof&Roos,1977).E.thienemanni Sommer,1951.On the gills of Gammarus tigrinus(cf.Bierhof&Roos,1977).E.zschokkei(Keiser,1921).Syn.:Opercularia zschokkei Keiser,1921.On thegnathopods of the gammarid Gammarus tigrinus and on other Entomostraca.On the cladoceran Acantholeberis curvirostris(cf.Nenninger,1948).Epistylis sp.Hutton,1964.On the decapod Penaeus duorarum(Florida,U.S.A.).Between the setae of the rst antenna of Gammarus tigrinus(cf.Bierhof& Roos,1977).Epistylis sp.Hutton,1964.On the decapod Ploeticus robustus(Daytona Beach, Florida,U.S.A.).Epistylis sp.Viljoen&Van As,1983.Two species on the thoracic appendages of a freshwater brachyuran,apparently erroneously identi ed as“Potamon sp.”(South Africa)[the genus Potamon does not occur in southern Africa].Epistylis sp.Pearse,1932.On the gills of the decapods Coenobita clypeatus, Geograpsus lividus,and Pachygrapsus transversus(Florida,U.S.A.).Epistylis sp.Hudson&Lester,1994.On the gills of the decapod Scylla serrata (Moreton Bay,Queensland,Australia).Epistylis sp.Turner et al.,1979.On the estuarine copepods Acartia tonsa andA.clausi(Escambia Bay,Florida,U.S.A.).Epistylis sp.Villarreal&Hutchings,1986.Fresh water.On the maxillipeds, pereiopods,and ventral portion of the abdomen of the decapod Cherax tenuimanus(Australia).CRUSTACEAN PROTOZOAN EPIBIONTS,I.PERITRICH CILIATES649 Family Lagenophryidae Bütschli,1889Genus Lagenophrys Stein,1852( g.3)L.aegleae Mouchet-Bennati,1932.Fresh water.On the branchial laments of the anomurans Aegla sp.,Aegla castro,and Aegla franca.Arroyo Miguelete, (Uruguay)and Parana River(Brazil).L.ampulla Stein,1851.Fresh water.On the gills of species of the genus Gammarus.L.andos(Jankowski,1986)(cf.Clamp,1991).Syn.:Circolagenophrys andos Jankowski,1986.Fresh water.On the decapod Parastacus chilensis(Chile).L.anticthos Clamp,1988.Fresh water.On the branchial laments of the decapods Parastacus pugnax,P.defossus,and P.saffordi(Chile,Brazil, Uruguay).L.aselli Plate,1886.On the branchial surface of the isopod Asellus aquaticus (Hamburg,Germany).L.awerinzewi Abonyi,1928.On the gills of the decapod Potamon uviatilis(as Telphusa uviatilis)(Africa).L.bipartita Stokes,1890.On the cladoceran Daphnia sp.(fresh water,U.S.A.).L.branchiarum Nie&Ho,1943.Fresh water.On the gills of the caridean shrimp Macrobrachium nipponense(as Palaemon nipponense)(Japan).L.callinectes Couch,1967.Marine and in estuaries.On the gills of the decapods Callinectes sapidus,C.bocourti,and C.maracaiboensis(Chesapeake Bay, Maryland,Virginia,and Gulf of Mexico).mensalis Swarczewsky,1930.Fresh water.On gammarids(Lake Baikal).L.darwini Kane,1965.On the branchial laments of the decapod Cherax quadricarinatus(stream near Darwin,Australia).L.dennisi Clamp,1987.Fresh water.On the decapods Orconectes illinoiensis, Cambarus bartonii bartonii,and C.chasmodactylus(North America).L.deserti Kane,1965.Fresh water.On the gills of the decapods Cherax tenuimanus and C.quinquecarinatus(SW rivers,Australia).L.diogenes(Jankowski,1986).Syns.:Circolagenophrys diogenes Jankowski, 1986,Lagenophrys incompta Clamp,1987.Fresh water.On the gills of the decapods Orconectes illinoiensis and Cambarus diogenes(Illinois,U.S.A.).L.discoidea Kellicott,1887(cf.Clamp,1990).Syns.:Lagenophrys labiata Wallengren,1900(a junior homonym of biata Stokes,1887(cf.Clamp, 1990));L.wallengreni Abonyi,1928;Circolagenophrys entocytheris Jankow-ski,1986.Fresh water.On ostracods.On the cray sh Cambarus sp.,C.chas-modactylus,C.bartonii bartonii,and Orconectes illinoiensis(Ontario,Canada and U.S.A.).650G.FERNANDEZ-LEBORANS&M.L.TATO-PORTOFigs.3-7.3,Lagenophrys(L.eupagurus,after Clamp,1989);4,Clistolagenophrys(C.primitiva, after Swarczewsky,1930);5,Setonophrys(munis,after Clamp,1991);6,Operculigera (O.asymmetrica,after Clamp,1991);7,Usconophrys(U.aperta,after Clamp,1991).L.dungogi Kane,1965.On the branchial laments of the decapod Euastacus sp.(stream near Dungog,Australia).L.engaei Kane,1965.On the branchial laments,basal areas of the gills, branchiostegite membrane and,more rarely,on the pleopods of the decapods Engaeus victoriensis and Austroastacus hemicirratulus(Victoria,Tasmania, and Melbourne,Australia).L.eupagurus Kellicott,1893(cf.Clamp,1989).Syns.:Lagenophrys lunatus Imamura,1940;Lagenophrys articularis Nie&Ho,1943.Marine,in estu-arine areas and fresh water.On the decapods Litopenaeus setiferus(as Pe-CRUSTACEAN PROTOZOAN EPIBIONTS,I.PERITRICH CILIATES651 naeus s.)(Penaeidea,Penaeidae),on the surface of the body,Litopenaeus van-namei(as Penaeus v.),on the surface of the body,Macrobrachium nipponense (Caridea,Palaemonidae)on antennae and pleopods,Macrobrachium ohione, on the surface of the middle of the pleura,Macrobrachium rosenbergii,on the gills,Palaemon paucidens(Caridea,Palaemonidae),Palaemonetes inter-medius(Caridea,Palaemonidae),Palaemonetes kadiakensis,Palaemonetes paludosus,Palaemonetes pugio,Palaemonetes varians,on the whole body, except on the gills,Palaemonetes vulgaris,Upogebia af nis(Thalassinidea, Upogebiidae),and Pagurus longicarpus(Anomura,Paguridae),on the gills (U.S.A.,Japan,Venezuela,Thailand).L.foxi Clamp,1987.Fresh water.On the gills of the gammarids Gammarus pseudolimnaeus,G.troglophilus,G.minus,and Gammarus sp.(Missouri, U.S.A.).L.in ata Swarczewsky,1930.On the distal areas of pleopods of the gammarid Gmelinoides fasciata(Lake Baikal).L.jacobi(Kane,1969).Syn.:Stylohedra jacobi Kane,1969.On freshwater decapods in Australia.L.johnsoni Clamp,1990.Syn.:Lagenophrys labiata Stokes,1887(partim). Fresh water.On the appendages and the surface of the carapace of the gammarids Gammarus fasciatus,G.daiberi,G.tigrinus,and Crangonyx gracilis(New Jersey,Michigan,and North Carolina,U.S.A.).biata Stokes,1887(cf.Clamp,1990).Fresh water.On the appendages and on the surface of the carapace of the gammarids Gammarus fasciatus, G.daiberi,G.tigrinus,and Cangronyx gracilis(New Jersey,Michigan,and North Carolina,U.S.A.).L.leniusculus(Jankowski,1986).Syns.:Circolagenophrys leniusculus Jan-kowski,1986;L.oregonensis Clamp,1987.Fresh water.On the carapace, gills,ventral surface of the abdomen,uropods,pereiopods,and pleopods of the decapod Pacifastacus leniusculus leniusculus,and on the gills of P.leniusculus trowbridgii and P.connectens(North America).L.lenticula(Kellicott,1885)(cf.Clamp,1991).Syns.:Stylohedra lenticula Kellicott,1885;S.lenticulata Kahl,1935;Lagenophrys lenticulata(Kahl, 1935)(cf.Thomsen,1945).Fresh water.Setae of the sixth and seventh pereiopods of the gammarids Hyalella azteca and H.curvispina(U.S.A., Canada,Mexico,and Uruguay).L.limnoria Clamp,1988.Syn.:Circolagenophrys circularis Jankowski,1986 (cf.Clamp,1991).On the isopod Limnoria lignorum.L.macrostoma Swarczewsky,1930.Fresh water.On gammarids(Lake Baikal). L.matthesi Schödel,1983.On the maxillipeds of the gammarids Gammarus pulex and Carinogammarus roeselii.652G.FERNANDEZ-LEBORANS&M.L.TATO-PORTOL.metopauliadis Corliss&Brough,1965.Fresh water.On the gills of the brachyuran Metopaulias depressus(endemic on Jamaica).L.monolistrae Stammer,1935.On the pleopods of the isopod Monolistra sp.L.nassa Stein,1852.Fresh water.On the pleopods of the gammarid Gammarus pulex.L.oblonga Swarczewsky,1930.On the antennae of the gammarid Gammarus hyacinthinus(Lake Baikal).L.orchestiae Abonyi,1928.On the amphipod Orchestia cavimana(Lake Balaton,Hungary).L.ornata Swarczewsky,1930.Fresh water.On ke Baikal.L.ovalis Swarczewsky,1930.Fresh water.On the thoracic appendages of ke Baikal.L.parva Swarczewsky,1930.On ke Baikal.L.patina Stokes,1887(cf.Clamp,1990).Syn.:Lagenophrys labiata Stokes, 1887(cf.Shomay,1955).(Corliss&Brough,1965;Clamp,1973).Fresh water.On the pereiopods and gills of the gammarids Gammarus sp.and Hyalella azteca.American continent.L.rugosa Kane,1965.Fresh water.On the gills of the decapod Geocharax falcata(Victoria,Australia).L.similis Swarczewsky,1930.On ke Baikal.L.simplex Swarczewsky,1930.On ke Baikal.L.solida Swarczewsky,1930.On ke Baikal.L.stammeri Lust,1950.On ostracods.Germany.(Lust,1950a).L.stokesi Swarczewsky,1930.On ke Baikal.L.stygia Clamp,1990.Syn.:Lagenophrys labiata Stokes,1887(cf.Jakschik, 1967).Subterranean water.On the gills of the cave-dwelling amphipod Bactrurus mucronatus(Illinois,U.S.A.).L.tattersalli Willis,1942.On European copepods.L.turneri Kane,1969.On freshwater decapods in Australia.L.vaginicola Stein,1852.Syn.:Lagenophrys obovata Stokes,1887.On the genital setae and thoracopods of the copepods Cyclops miniatus and Cantho-camptus sp.L.verecunda Felgenhauer,1982.On the decapod Palaemonetes kadiakensis (Illinois,U.S.A.).L.willisi Kane,1965.Fresh water.On the gills of the decapods Cherax destructor,C.albidus,and C.rotundus(Melbourne,New South Wales(e.g., Newcastle),and NW Australia).Genus Clistolagenophrys Clamp,1991( g.4)C.primitiva(Swarczewsky,1930)(cf.Clamp,1991).Syn.:Lagenophrys primi-tiva Swarczewsky,1930.On pereiopods and pleopods of the gammarid Pallasea cancellus(Lake Baikal).Genus Setonophrys Jankowski,1986(cf.Clamp,1991)( g.5)S.bispinosa(Kane,1965)(cf.Clamp,1991).Syn.:Lagenophrys bispinosa Kane,1965.On pereiopods of the decapod Cherax rotundus setosus.Stream near Newcastle(N.S.W.,Australia).munis(Kane,1965)(cf.Clamp,1991).Syn.:Lagenophrys communis Kane,1965.On the body surface(telson,pleopods,pereiopods,carapace...) of the decapod Cherax destructor.On the gills of the decapods C.rotundus,C.albidus,C.quadricarinatus,Euastacus armatus,and Engaeus marmoratus(Victoria,Melbourne,and Tasmania,Australia).S.lingulata(Kane,1965)(cf.Clamp,1991).Syn.:Lagenophrys lingulata Kane,1965.On the branchial laments and branchiostegite membrane of the decapods Cherax destructor, C.albidus,and C.rotundus(Victoria, Melbourne,and coastal and central areas of Australia).S.nivalis(Kane,1969)(cf.Clamp,1991).Syn.:Lagenophrys nivalis Kane, 1969.On freshwater decapods in Australia.S.occlusa(Kane,1965)(cf.Clamp,1991).Syn.:Lagenophrys occlusa Kane, 1965.On the anterior zone of the branchial cavity of the decapods Cherax destructor,C.albidus,and C.rotundus(Victoria and New South Wales, Australia).S.seticola(Kane,1965)(cf.Clamp,1991).Syn.:Lagenophrys seticola Kane, 1965.On the setae of the decapods Engaeus fultoni and Geocharax falcata (Victoria,Melbourne,and Templestowe,Australia).S.spinosa(Kane,1965)(cf.Clamp,1991).Syn.:Lagenophrys spinosa Kane, 1965.On the pleopods,carapace,and telson of the decapod Cherax destructor (Victoria,Melbourne,and Heathcote,Australia).S.tricorniculata Clamp,1991.On the pleopods of the decapod Geocharax falcata(Victoria,Grampian Mountains,and Wannon River,Australia). Genus Operculigera Kane,1969( g.6)O.asymmetrica Clamp,1991.On the base of the gills of the freshwater decapods Parastacus pugnax and Samastacus spinifrons(Concepción and Talcahuano,Chile).O.insolita Clamp,1991.On the base of the gills of the freshwater decapod Parastacus pugnax(Concepción,Talcahuano,Malleco,and Puren,Chile).O.montanea Kane,1969.On the freshwater decapod Colubotelson sp.(Aus-tralia).O.obstipa Clamp,1991.Pleopods of the isopod Metaphreatoicus australis (New South Wales,Australia).O.parastacis Jankowski,1986.On the base of the gills of the decapod Parastacus nicoleti(Isla Teja,Valdivia,Chile).O.seticola Clamp,1991.On the setae at the base of gills of the decapod Parastacus pugnax(Concepción,Chile).O.striata Jankowski,1986.On the decapod Parastacus chilensis.Chile.O.taura Clamp,1991.On the branchial laments of the freshwater decapod Parastacus pugnax(Concepción,Malleco,and Puren,Chile).O.velata Jankowski,1986.On the gills of the anomuran Aegla laevis.Chile.O.zeenahensis Kane,1969.On freshwater decapods in Australia.Family Usconophryidae Clamp,1991Genus Usconophrys Jankowski,1985(cf.Clamp,1991)( g.7)U.aperta(Plate,1889)(cf.Clamp,1991).Syns.:Lagenophrys aperta Plate, 1889;Usconophrys dauricus Jankowski,1986.On the gills and pleopods of the isopod Asellus aquaticus(Marburg and Hessen,Germany;North Carolina, U.S.A.;Brittany,Finisterre,Plougarneau,Pont-Menou,and Douron River, France).U.rotunda(Precht,1935)(cf.Clamp,1991).Syn.:Lagenophrys rotunda Precht,1935.On ostracods.Germany.Family Operculariidae Fauré-Fremiet,1979(in Corliss,1979)Genus Opercularia Stein,1854( g.8)O.allensi Stokes,1887.Syn.:O.ramosa Stokes,1887.On several living and inert substrata.On the body of the cray sh Astacus leptodactylus(cf.Matthes &Guhl,1973).O.asellicola Kahl,1935.On the isopod Asellus sp.Germany.O.coarctata Claparède&Lachmann,1858.On crabs(Buck,1961).O.crustaceorum Biegel,1954.On the gills of the cray sh Astacus astacus(asA. uviatilis).On the maxillae,maxillipeds,and pleopods of Austropotamo-bius torrentium(cf.Matthes&Guhl,1973).O.cylindrata Wrzesniowski,1807.On the copepod Cyclops sp.O.gammari Fauré-Fremiet,1905.Pereiopods of the gammarid amphipod Gammarus sp.O.lichtensteini Stein,1868.On various crabs and molluscs.O.nutans Ehrenberg,1838.Syn.:O.microstoma Stein,1854.On Entomostraca.On the cladoceran Alona af nis(cf.Matthes,1950).On the maxillipeds of the European cray sh Astacus leptodactylus(cf.Matthes&Guhl,1973).O.protecta Penard,1922.On the setae of pereiopods of the gammarid amphi-pod Gammarus pulex.O.reichelei Matthes&Guhl,1973.Found exclusively on the maxillipeds of the cray sh Astacus leptodactylus.O.stenostoma Stein,1868.On the isopod Asellus aquaticus.Genus Orbopercularia Lust,1950(cf.Lust,1950b)( g.9)O.astacicola(Matthes,1950)(cf.Matthes&Guhl,1973).Syn.:Opercularia astacicola Matthes,1950.Maxillipeds and pleopods of the cray sh Aus-tropotamobius torrentium.Genus Propyxidium Corliss,1979( g.10)P.aselli Penard,1922.On the isopod Asellus sp.P.asymmetrica Matthes&Guhl,1973.On the European cray sh Astacus astacus(as A. uviatilis).P.bosminae Kahl,1935.On the cladoceran Bosmina sp.P.canthocampti Penard,1922.On the pereiopods of the harpacticoid copepod Canthocamptus sp.Fresh water.P.cothurnioide Kent,1880.On the ostracod Cypris sp.P.hebes Kellicott,1888.On the pereiopods of the isopod Asellus aquaticus.P.henneguyi(Fauré-Fremiet,1905)(cf.Kahl,1935).Syn.:Opercularia hen-neguyi Fauré-Fremiet,1905.On the rst abdominal segment of the copepod Cyclops sp.Genus Ballodora Dogiel&Furssenko,1921( g.11)B.dimorpha Dogiel&Furssenko,1921.On Porcellio sp.and other terrestrialisopods.Genus Nuechterleinella Matthes,1990( g.12)N.corneliae Matthes,1990.On the ostracod Cypria ophthalmica.Genus Bezedniella Stoukal&Matis,1994( g.13)B.prima Stoukal&Matis,1994.Fresh water.On the ostracod Cypria sp.(Slovakia).Figs.8-14.8,Opercularia(O.nutans,after Foissner et al.,1992);9,Orbopercularia(O.astacicola, after Matthes&Guhl,1973);10,Propyxidium(P.canthocampti,after Penard,1922);11,Ballodora (B.dimorpha,after Dogiel&Furssenko,1921);12,Nuechterleinella(N.corneliae,after Matthes, 1990);13,Bezedniella(B.prima,after Stoukal&Matis,1994);14,Rovinjella(R.spheromae,afterMatthes,1972).Family Rovinjellidae Matthes,1972Genus Rovinjella Matthes,1972( g.14)R.spheromae Matthes,1972.On the marine isopod Sphaeroma serratum. Family Scyphidiidae Kahl,1933Genus Scyphidia Dujardin,1841( g.15)Scyphidia sp.Henebry&Ridgeway,1979.On the cladocerans Scapholeberis kingi,Alona costata,and Pleuroxus denticulatus(Ashmore Lake,Illinois, U.S.A.).Family Vaginicolidae De Fromentel,1874Genus Platycola Kent,1881( g.16)P.baikalica(Swarczewsky,1930).Syn.:Vaginicola baicalica Swarczewsky, 1930.Fresh water.On the gills of the gammarids Brandtia lata,Pallasea grubei,and Echinogammarus fuscus(Lake Baikal).P.callistoma Hadzi,1940.Fresh water.On the cave-dwelling isopod Microlis-tra spinosissima(former Yugoslavia).P.circularis Dons,1940.Marine.On the uropods of the isopod Limnoria sp.P.decumbens(Ehrenberg,1830).Syns.:Vaginicola decumbens Ehrenberg, 1830;Platycola ampulla De Fromentel,1874;P.regularis De Fromentel, 1874;P.striata De Fromentel,1874;P.truncata De Fromentel,1874;P.longicollis Kent,1882;P.intermedia Kahl,1935;P.re exa Kahl,1935;P.amphora Swarcezwsky,1930;P.amphoroides Sommer,1951.Fresh water.On several vegetable and animal substrata.On the gills of the gammarid Brachiuropus sp.(Lake Baikal)(Swarczewsky,1930).geniformis Hadzi,1940.Fresh water.On the cave-dwelling isopod Micro-listra spinosissima(former Yugoslavia).P.pala Swarczewsky,1930.Syn.:Vaginicola pala Swarczewsky,1930.On the gills of the gammarid Palicarinus puzyllii(as Parapallesa pazill)(Lake Baikal).Genus Cothurnia Ehrenberg,1831(cf.Claparède&Lachmann,1858)( g.17)C.angusta Kahl,1933.Brackish or fresh water.On ostracods(Kiel,Germany).C.anomala Stiller,1951.Fresh water.On the amphipod Corophium curvispi-num(Lake Balaton,Hungary).C.antarctica(Daday,1911)(cf.Warren&Paynter,1991).Syn.:Cothurniopsisantarctica Daday,1911.Marine.Epibiont on the ostracod Philomedes lae-vipes(Antarctic areas).C.astaci Stein,1854.Fresh water.On the pleopods and gills of cray sh.On the maxillae,maxillipeds,and pleopods of the cray sh Astacus astacus。
