526

An Annotated List of Protozoan Parasites, Hyperparasites, and Commensals of Decapod Crustacea*

VICTOR SPRAGUE and JOHN COUCHi

Urriu. of Mar)~lrinil,Nnfirrnl Rrsoio.rP.c Zi~.~fi/ufr,Clrr\npcrrke niologicnl /.ribriwitoiy, Soloinoirs. 121d. 2OhS‘S and (I. S. Dep. of Critninerrf>, Nalroiinl Oceanic nnrl :lI~itos~hrricAf/iiiiiri.\li-r,lioiz, Nufionnl Mnritie Fi$/reric,.!Sfwice Biologirn/ I.nliorrrloiy, Oxfoid, Mil. 21654

SYNOPSIS. An approximately complete list of the knolvn pro- suggrstrd. Host. site of infrction and seographic location are tozoan associates (exclusive rif a few aberrant forms like Amal- nientionrd. Soinc comnirnts on pathosrnicity are made. locystir) of decapods is prcsented. A few taxonomic changes arc

THIS is, principally a.taxonomic paper, intended to Pre- in the in]ial,itcd organ (9) with serious results. Micro- sent In a sYstematlc manner a conlP1ete list Of sporida arc notorious pathogens and generally kill the de- Protozoa known to be associated with and capod host. I\ felt, ciliates ars pathogenic in decapods but deal with a few of the problems in classification and are llarmless coInIllensals. nomenclature found at the generic and spccific levels. At Even we ran \,.ith confidel,ce that a para- the same time, such highly relevant matters as host and site is 01’ is not 1,articularly pathogenic to a dc*capod host, site of infection, locality and pathogenicity are mentioned. \ce kno\,. little ,\klat in terms of the At the higher levels of classification we followed “A re- fluctuation of tklc host poprilations, vised classification of the phylum Protozoa” of the Honig- berg Committee (78) as far as we could but \\’ere unable Suhl~liyluni SARCOMASTOGOPHORA Honixherg & Bala- to follow it consistently because several specialists consider lllutll. 1963 (78). different parts of that system inadequate. In classifying Suprrclass SARCODINA IIcrtnig R- Lrsser, 1874 (78). the Amoebae lve ha\x folloived Rovee and Jahn (20). Class HYDRAULE.4 Jahn & Bo\cr, 1965 (85). Commenting on their difficulties, they stated very clearly Order HYALOPODIA Bovee & Jahn. 1966 (20). Family PAKAMOEBIDAE Poche, 1913 ( 153). the nature of some of the basic problems: “One reason Genus Pnranroeba Schaudinn. 1896 (169). these families are difficult to deal with is the consensus of taxonoInists of protozoa (~~~i~h~~~et al,, 1964) whicll foroitioeb(L perniciosa Si~rague, Bpckctt & Saver, 1969 (190). Syn.: Pnrnrnoeh sp. Spraque & Bvrkrtt. 19611 ( 185). In hrmii- to recognize that protozoa are at least as lymph of hluc crah Cnllinectes supidus Rathbun, 1896. Chinco- evolutionally ancient and taxonomically diverse as are the teaguc nay. \.irginia and hlaryland, CSA (165, 166, 167, 18-1, metazoa. The protozoa probably should be considered at 185, 190); coast of North Carolina and Georgia, USA (129). least a subkingdom (or kingdom) of organisills, containing Dr. Eugrnr E. Rovce (pers. coin.) thinks P. eilhorrfi Scliaudinn. phyla and subphyla (Raabe, 1964). This idea \vas 1896, the tWe sliec+’s, belongs tO WnuS MaYoreilfl and (in pry=) has rcjccted both gums Pnro?iroeb~Schaudinn and family Para- presented by and \vas at length and nloehidar Pc~he. If B0vee.s opinion is jllstificd, >,Ilich \\e see, no strongly emphasized by Kozloff ( 1960) in a syrrlposial ad- reason to doubt, then P. periricio\o must he transferred to another dress; and kingdoms were suggested-but not adopted- genus. Sincr ur hold no oliinion as to the correct systematic by Corliss (1962) ,” In the classification of the “sporozoa” position of this anruc1)a \\e ha\?, for couvenicnce, used the old we have followed Levine (128), who has given the most classifiration recent modification. Howcver, the comment just quoted Suliphylum .4PICOMPLEX:\ Idmine, 1970 (128). from Bovee and Jahn is applicable also to this, because the “Sporozoa” probably include at least 3 distinct phyla. M’s Class SPOROZOEA Lruckart. 1879 (128). Su1)class GKEC;ARINlA Dufour, 1828 (78). have followed the Honigberg Committee on classification Order EUC;RE(:ARINIDA Liger, 1900 (78). of the ciliates, since that part, strongly influenced by the Farriily CEPHAI,OIDOPHORlD.4E Kamm. 1922 (90). opinions of Corliss, has not bcen so controversial. (;enus Cephnlordophoru hiavrodiadi. 1908 (90). Altho we have listed most of the protozoa reported as Cephaloidophora berrnudeniir (Ball, 1951) n. comb. Syn.: associates of decapods, these are doubtless a small fraction Cnrctnoec~terberinudun~ir Ball, 1951 (lo). Ill mid- and hind- of those which actually occur. Probably all decapods qut of the grapaiid crab Pachygrapius tranrreiius (Gibbs). Ber- harbor protozoan parasites: few sPccirs are knoLtI1 hecausc muda. The gencric nanle Corcinoecetes Ball, 1938 (7j, is a junior synonym of Kernntopris (210). The present species does relatively fsbv decapods have becn examined. not srrni to Iia\e any of the vsscntial characters of Nenzntopcis The pathogenirity of protozoan infections in decapods is like ~~~/~~l~id~~~~~~~,,\hicl1 grnus it is here is poorly understood hit a fe\v gcneral statsmcnts may be pro\iaionaily translcrrcd. made. The one known amoeba is highly pathogenic to its L‘ephaloidophora cttrpilodei Dall. 1963 (13). In the gut of rrab host, (;regarincs do not Seem tO tIe pathogenic, thr xanthic1 crab Carpiloder rupntzri. Cocoanut Island, Hawaii, altho they cause some local damage and occlude passages “*. Cephn/oidop/iorn conforntii (Diesing, 1951 ) Lhgrr & Duboscq, 191 1 (12‘2). Syns.: Greparinu conformis Diesing, 1851 (90); * Contribution No. 458, Natural Rrsources Institritr of thr ~~~~e~~~t~ron/ormis (~iesing,18jl j LabbC, 1899 (107) ; Fren- University of Maryland. rahrL co‘i/orrtL1’(r)icsirls ““l Id‘ser ’ Duboscq3 1907 ( ‘‘1 ; L presentaddress: Erlvironnll.nta{ protrrtil,n .i\sc,ncy [,a],,,ratory, Gulf Breei:?. Fla. 32561. Cmrcinorcufri c~,n!orniic (Dirsing. 11151 ) Ball, 1938 (7). In the PROTOZOAOF DECAPODS 527 cera thr gut lunirn of thr grapsid crab Pcchygrupsur marmoratus alphrid shrimp Alpheus ruber Milnr-Edmards and A. dentipes Fabricius. hlcditerranean roast of France; roast of Tunisia (214) : Guerin. Banyuls. France. northrast roast of Spain (213). Tuzrt and Ormi6res (210) de- Cephaloidophoro ( ?) praemorra (Diesing, 1851) Kamm, 1922 rlared the name CnrcinoecPteT Ball, 1938, a junior synonym of (90); Syns. : Gregarina braemorsa Diesing,-1851 (90) ; Aggregata Nematopcir Srhncider and rrturned this spwies to genus Cepha- praemorsa (Diesins, 1851) Labb6, 1899 (107) : Frenzelina grae- loidophora. morsa (Diesing-, 1851) LiSer & Duboscq, 1907 (116). In gut Cephaloidophorn drnchi Theodorid&, 1962 (201 ). Syn.: and vesiclrs adhrring to the ovary of the crab Cancer pagurus Cephnloidophora sp. Tuxt & OrmiEres, 1961 (210). In the I,. As Kamm (90) pointed out. this nominal species is based on gastric cera of the xanthid crab Xantho rir~zrlosur (Risso). Banyuls. a niere refrrence by Redi, in 3684, to somethins which might France. have been a p,garinc. Cephaloidophora d~omiae (Frenzel. 1885) VimrEs & Rubih. Cephaloidophora sp. Syn. : Carcinoecetes sp. Ball, 1948 (8). 1969 1213). Syn.: Gregarina dronziae Frrmzrl. 1885 (61). In hlainly in the digrstive gland hut also in the gut Innirn of the gut of the crab Droniiu personoto (= dgaris) (61, 214). Gulf sand crab Emerita analoga. Southern California, USA. Since of Naples (61), Blanes. Spain (213); and coast of Tunisia (214). genus Carcinoecetes Ball, 1938 (7) is a junior synonym of Altho ThPodorid6s (201 ) suggested that this gregarine probably A'eniatopii$ Srhncider. Ifi92 (?lo), this species needs to be should be placed in genus Cephaloidophora, thc new combination reassigned. Mat knonn gresarines of crabs belong either to is here attributed to Vi\ar+s and Rubih (213). nho first used it. gcnus C~phnloidophoraor n'eniatopsis. The known characters of Cephaloidophora duborcqi Poisson. 1924 (154). In thr gut of the this species seems to he more like those of the former genus. alphcid shrimp Athanas nitescens Lcarh. Luc-sur-Mer (Calvados) ( 154) and Roscoff (211 ), France. Genus Cephalolobus Kruse, 1959 (99). Cephaloidophora etiii (Ball. 1959) n. ctimh. Syn.: Cnrcinoe- Cephnlolvbur penneus Krusr. 1959 (99). Attached to filtrr of cetes etisi Ball. 1959 (12). In the rrah Etirus laer,imanus Randall. stomach of shrimp Penaeu, artecus Ivcs and P. duorarum Burken- Bombay, India. Carcinoecetes Ball, 1938, is a junior synonym of road. Florida, USA. Type species. Kruse considered the systc- Nemntopsis (210). The dcsrription of the present species fits niatic position of the senus in dctail. He did not place it in a that of Cephuloidophora rather than that of Neniatopsis. family hut concluded. "If the life cycle of C. penaeur is proven Cephaloidophuru foresti Th6odoridi.s. 1967 (205). In thr gut to hc siniilar to that of Ani~olobur,both genera should be placed of thc crab Xantho pores,n (Olivi). Banyuls and Villefranche, in a ric'w fainily." 'l'liiodoridi~s (202). for reasons not very con- France. vincing assigned the genus to family Cephaloidophoridac, where Cephaloidophora fosw (L(.gcr & Dubosrq. 1907) Trigouboff. it is here listed for convenii.ncr. 1912 (208). Syns.: Aggregatu roelomica LCgrr. 1901 [ partim] Cephalolobur petiti Theodorid&, 1964 (202). Active stages (110); Frenzelina forsw Liger & Duljoscq, 1907 (116). In gut in stomach and cysts in hind-gut of thc penacid shrimp Solenocera of the pea crab Pinnotheres pi$un! Penn. living in Mpilus edulis nienibranacea (Risso). Banyuls, France. L. France (English Channel) ( 1 10). Cephaloidophora guinotae Thiodoridts, 1967 (205). In gastric Genus L'ardiohabitans Ball, 1959 (12) ceca of the xanthid crabs Xuntho granulocurpus Forest (205), Pilumnus hirtellus rpinifer Milne-Edhards (205, 213) and P. Curdzohnbrtans ietnai Ball, 1959 (12). In the mid-gut of the hirtellus L. (205). The 1st 2 hosts at Banyuls and the 3rd at shrimp Penaeur Tenikulcatus de Haan. Bombay, India. Type and Villefranche, France (205) ; thc 2nd also at Blanes, Spain (213). only species. Ball did not assign the genus to a family but he Cephaloidophora knoepfleri l'hiodoridh, 1962 (201 ). In gas- compared it both uith Cephaloidophora and genera in the tric ceca of Pinnotheres pinnotheres (L.), a pea crab. ronimensal Porosporidae ; it is here provisionally listed with the Cephaloido- in the rlani Pinna nobilis L. Banyuls, France (201) ; Blancs, phoridae. 'lhc validity of the genus is questionable. Spain (213). Cephaloidophora mithraxi (Ball, 1951 ) n. comb. Syn.: Car- Family URADIOPHORIDAE GrassC, 1953 (64) cinoecetes mithraxi Ball. 1951 (10). In the crab itlithrax forceps Genus C'radiofora Mrrcicr, 1912 (90). (Milne-Edwards). Bermuda. The generic name Carcinoecetes C'rcdiophorn rithnnasi Poisson, 1924 (154). In the gut of the Ball, 1938 (7) is a junior synonym of A'ematopsis (210). The alpheid shrimp Athanor nitescens Leach. Lnc-sur-Mer (Cal- present species does not rt~srrnl)le NematopJis but its known vados), France. characters arc consistent uith those of genus Cephaloidophora Uradiophora cuenoti (Mercier, 1911) Merrirr, 1912 (90). to which genus it is here transferred. Syn.: Cepholoidophora cuenoti Mercier, 1911 (90). In the gut Cephaloidophora nephropis Tuzet 8: Orrnii.res, 1961 (210). of the shrimp Atyoephya desmaresti Millet. Nancy, France. Syn. : Porospora nephropis Liger & Duboscq, 1915 [ partiin] (126). In thr %ut of the Norway lobster, A'ephrops norz,egicuc Family POROSPORIDAE Lahbi, 1899 (107). (126, 127, 210, 212). Roscoff, France (126, 127, 210, 212). Cephuloidophora nigrofusca (Watson, 1916) Kamm, 1922 (90). Gnus Porospora Schneider, 1875 (171). Syn.: Frenzelina nzerofusca Watson. 1916 (216). In rut of thc Porospora gigantea (Bencden. 1869) Schneider, 1875 (171 ). ,I fiddler crabs IJca pu>nux and U. pugzlator. Cold Sprini Harbor, Syn.: Gregarina gigantea Bcnedrn, 1869 (17). In the lobster New York, US.4. Homarus gammarus (L.) and the gastropod Trochocochlea Cephaloidophoru oceliuta (Liger 8: Duboscq, 1907 ) Kamni. mutabilis Philippi. Roscoff (171 ) and Banyuls (201 ), France. 1922 (90). Syns.: Aggregata i'ngans LCger & Duboscq, 1903 This is the type and the only spccies for which the essential [partim] (112, 116); Frenzelinu ocellata Liger 8: Duboscq. 1907 gcneric character, naked sporozoites in a molluscan host, has been (116). In gut lumen of the hcrmit crab Eupagurus prideauxi demonstrated. This demonstration was by a laboratory infection Leach. Banyuls, France ( 112, 201 ), experiment performed by Hatt (72). The same author (71) Cephaloidophora ohia (Watson, 1916) Kamni. 1922 (90). obtained partial development also in M3,tilus minmus Poli. Syn.: Frenzelina o/iuia Watson, 1916 (216). In gut of 2 littoral Porospora nephropis L6ger 8: Duboscq, 1915, emend. Tuzet & crabs Libinia dubia Milne-Edwards (216) and Ocj'pode cursor OrmiPrcs, 1961 (210). Syn. : Porospora nephropis Liger & L. (46). Cold Spring Harbor, Nc\z- York, US.4, (216) and Duboscq, I915 [partim] (126). In gut of the lobster Nephrops Kalade, Lebanon (46). norcegicus L. Molluscan host unknovn. Roscoff, France. Cephaloidophora pinguis Ball, 1963 (13). In gut of the Porospora piJae LCger & Duboscq, 1911 (123). In the spider xanthid crab Carpilodes rugntu.<. Cocoanut Island, Hawaii, USA. crabs Pisa grb~ii (123) and P. armata (213). Cette (123) and Cephaloidophora poissoni 'lh(.odoridPs, 1967 (205). In the Villefrariche-sur-Mer (209). France; northeast coast of Spain 528 PROTOZOAOF DECAPODS

(213). Trhgouboff (209) observed peculiarities in this species penaeid shrimp Solenocera membranncea (Risso). Intermediate which convinced L6ger and Duboscq (127) that it is not a host unknown. Banyuls, France. This species is transferred to Porospora. These authors did not attempt a reclassification for genus Nematopri.r because its vegetative stages are similar to those lack of information on final development of the cysts. Vivarks of confirmed species of this genus in crabs and shrimp but and Rubi6 concluded that it is neither a Porosfiora nor a strikingly different from those of the single confirmed species of Nematopsis (213). It is kept here in Porosporn provisionally Porospora, known only in the lobster. because there is still no clue as to its proper generic classification. Mematopsis ostrenrum Prytherch, 1938, rmend. Sprague, 1949 Porospora sp. Thhodoridks & Laird, 1970 (206). In gut of (176). Syns.: Nematopric octrearum Prythrrrh, 1938 [partim] the lobster Homarus americanus. St. Andrcws, New Brunswick, (159), N. ostrearum Prythrrrh, 1940 [partim] (160), [non] Canada. N. legeri (de Beauchamp, 1910): Kudo (103, 104). In gut of the xanthid crabs Panopeur herbstii Milne-Edwards (159, 160), Genus Nematopsis Schneider, 1892 ( 172). Eurypanopeus depressus (Smith) (159, 160), Neopanope texana Nematopsis cnlappae (Ball, 1951) n. comb. Syn.: Carcino- ragi (Smith) (97), Eurltium limosum (Say) (176, 180). Spores ecetes calappae Ball, 1951 (10). In gut of the oxystomatous crab in the oyster Crarrostrea oirginica (Gmelin). Gulf and Atlantic Cnlappa flammne (Herbst), Intermediate host unknown. Ber- coasts of North America, as far north as New York (109). muda. Carcinoecetes Ball, 1938, is a junior synonym of Nema- A’emotopsis panopei Ball, 1951 (10). In the gut of the xanthid topsis (210). Excepting rare intrarcllular stages, which could rrabs Panopeus occidentalis Saussure and P. herb~tiiMilne-Ed- belong to another species, the known characters of this species wards. Intermediate host unknown. Bcrmuda. are consistent with those of Nematopsis, to which genus this Nematoprir garnpeneopsi~i (Srtna & Bhatia, 1934) n. comb. species is here provisionally assigned. It is noteworthy that Ball’s Syns. : Hirmocystis ( ?) parnpeneopsisi Setna & Bhatia, 1934 figures show a ~iorosporid character which the author did not (173) : Protomagalhensin (7) ottenunta Sctna & Bhatia, 1934 mention, the “muscular collar” of the protomerite. (173). In gut of the shrimp Parapeneopsis sculptilis (Heller). Nematopsis doripfie Bogolepova, 1953 (18). In gut of the Possible intermediate host unknown. Bombay, India. Grass6 (64) oxystomatous crab Dorippe granulnta de Haan. Sea of Japan. concluded that these 2 species described by Setna and Bhatia Nematopsis duorari Kruse, 1966 (100, 101). In gut of the were crroneously assigned as to gcnera. Altho mature cysts were shrimp Penneus duorarum Burkenroad. Spores in the pelecypods not seen, the known characters (especially disappearance of septa Aequipecten irradians. Cardita Poridana, Chione cancellata, in the satellites) of both species secm to he consistent with those Macrocallista nimbora. Florida, USA. of the Porosporidae. Furthermore, the difference between the 2 h’ematopsis goneplaxi Tuzet & OrmiPres, 1961 (210). Syn.: (slight difference in shape) seems to he insignificant. Thus, they PorosPora petiti ThCodoridPs, 1962 (201). In gut of the crabs are treated here as the same spccies. The specific name of that Coneplax angulatu (Penn.) (210, 201) and G. rhomboides (213). une appearing first in the paper is rrtained. Assignment is here Intermediate host unknown. Cette (210) and Banyuls (201), made to gcnus Nematopsis because the known stages are much France; northeast coast of Spain (213). more like corresponding stages of this genus than those of Nemntogsis grarsei (ThCodorides, 1962) VivarPs & Rubib, 1969 Porospora and because the former (100) hut not the latter is (213). Syn.: Poros~oragrarsei Th6odoridPs, 1962 (201). In known to occur in shrimp. Thdodorides (202) suggested that gut of the oxystomatous crab Calappa granulata L. (201). In- these gregarines may belong to genus Uradiophora or maybe termediate host unknown. Banyuls, France (201) ; northeast coast Cephaloidophora, hut this suggestion is untenable bccause the of Spain (213). septa of satellites do not disappear in these genera. Nematopsir herpertis (Ball, 1938) Tuzet & OrmiPres, 1961 Nematopsis penaeur Sprague, 1954 (179). Syn.: Porospora (210). Syns.: Carcinoeceter hesperus Ball, 1938 hw1: SPECIES] penaeus (Sprague, 1954) ?’h&odoridPs, 1965 (203). In the gut (7); Por~sporahesperus (Ball, 1938) ThCodoridb, 1961 (200) ; nf the shrimp Penneur ortecur Ives. Intermrdiatr host unknobn. [ ?] Nematopsis rp. Ball, 1951 (11). In gut of the grapsid crab Barataria Bay, Louisiana, USA. Sprague ( 179) evidently mistook Pachygrap.rur crasripes Randall. Intermediate host unknown. something else for intracellular stages of this parasite, for presence California, USA. of these has not been confirmed. Kruse (99) reported this species Nemntopsis legeri (de Bcauchamp, 1910) Hatt, 1931 (73). in P. aztecus, P. duorarum and P. setiferus. Hutton et al. (79) Syns. : Nematopsis mediterranen Lhger. 1905 Inomen oblitum] found it in these and also Sicyonia. Later, Hutton (80) reported (113); Porosporn leg& de Beauchamp, 1910 (16); P. gallo- it in P. aztecus, P. setiferus (I,.), P. duoraruni Burkenroad, P. provincialis LCger & Duboscq, 1925 (127, 73). In gut of the brasiliensis Latreil, Sicyonia typica Boeck, Trachypenaeus similis crabs Eriphia .rpinifrons (16) and E. ~‘eriuco~n(214); spores in (Smith), T. constricta ( Stinison), Xiphopeneur kregeri (Heller) the mussel Myfilus galloprovincialis and possibly other molluscs and Solenocera atlantidic Burkenroad. Considering that gregarines (73). Southern coast of France (16) and coast of Tunisia (214). in these hosts have not been studied in detail, that Kruse found This is one of the best known species of A’ematopsir, due largely 2 species in 1 host, and that ecologic factors related to the to the work of Hatt (70, 71, 73), who studied its entire life different shrimp are diverse, it is probable that several species of cycle by means of experimental infections. Previously only parts Nematopsis in these shrimp have been mistaken for one. The of the life cycle of a porosporid gregarine had been made known work of Kruse (100, 101) supports Sprague’s original generic experimentally (124, 179). Tuzet and Ormikres (210) recently determination. made a careful study of this species to confirm the generic A‘ematopsir fiortunidarum (Frenzel, 1885) Hatt, 1931 (73). characters. Syns. : Aggregata portunidnrum Frenzel, 1885 [ partim] (61, Nematopsis marnisi (Lhger & Duhoscq, 1911) Sprague, 1954 116, 127); Nemntopsis schneideri Lkger, 1903 (111, 127, 73); (179). Syn.: Porosporn marnisi LPger & Duboscq, 1911 (123). Frenielina portunidarum (Frenzel, 1885) Ldger & Duboscq, 1907 In gut of tlir rrabs Macropipus (= Portunus) depurator L. (116) ; Porosjora portunidarum (Frenzel, 1885) Lhger & Du- (123), M. mindis, Portumnus lotipes (214) and Macropipus boscq, 1911 (122). In gut of crabs Corcinus maenas (61, 122) depurator (213). Intermediate host unknown. Cette, France and C. mediterraneus (214). Originally reported also in Macro- (123) ; coast of Tunisia (214) ; northeast coast of Spain (213). pipus (= Portunus) arcuatus (61) hut LCger and Duboscq (122) Nematoprir mututne (Ball, 1959) Tuzet & OrmiPres, 1961 reserved the specific name portunidarum for the gregarine in (210). Syn.: Cnrcinoecetes rnatutae Ball, 1959 (12). In gut Cnrcinus. Dcvelops experimentally in Macropipus ( 124, 125, of the crab Matuta lunaris (Forskil). Intermediate host unknoxn. 127). Spores in the niolluscs Cardium edule, Scrobularia plana Bombay, India. and perhaps others (127). Naples, Italy (61); Etang PCrols, Nematopsis miroulei (ThCodorid&s, 1964) n. comb. Syn.: near Montpellier, France (122); coast of Tunisia (214). ThBo- Poros~ora miroulei Thdodoridcs, 1964 (202). In gut of the dorid& (201, 203), apparently overlooking experimental proof PROTOZOAOFDECAPODS 529 by Liger 8: Duhoscq (124, 125) that this is a Nematopsis, has Family and qcnus uncertain continued to call it Porospora. N. schneideri, represented by Gregnfina pmtrini Frcnzel. 1885 (61). In gilt of the crab spores in Solen oagina, is nominal type species by monotypy hut, Macropipus (= Po,flmur) nrcuatus. Naplrs. Italy. hfonocystid since this is regarded as a junior synonym of N. portunidarum gregarine. certainly not stmiis Gregarina and unlike other known (127, 73), the latter is now generally accepted as the type. grcgarincs in decapods. Nematopsis prytherchi Sprague, 1949 ( 176). Syns.: Nema- Gregarines, identity not spccified. Heckmann ( 74) found topsir ostrearum Prytherch, 1938 [partinil (159) ; N. ortrearum intestinal grrgarines in a ni~inhrrof decapods in Hawaii. Heldt Prytherch, 1940 [partim] (160). In the stone crab Menippe (75) described. xvithout naminp. grrgarines from the shrimp mercenaria (Say), Spores in the oyster Crarsostrea virginica Penoeus kerathurus (Forsk:il) in thy Gnlf of Tunis. ThCodoridPs (Gmclin). Louisiana. Florida and the south .4tlantic states of (202) judgcd that thcsc last onrs helonged to 2 species, a USA. Uradiophora and a Porospora. Nematopis raouadi Vivares, 1970 (214). In gut of the crab Poljanskij and Chcjsin (52) nicritioncd “ccrtain species of Porturnnus latipes. Coast of Tunisia. It is curious that Vivares, Merogregarina from the alinientary tract of crahs” but we have when comparing N. raouadi \vith A‘. marnisi in the samr host. been unable to find mention of them elsewhere. found gymnospores of the formrr to he the same size (3.8-4.5 a) as that given by Leger and Duboscq (122) for the latter (4 a), Suhclass COCCIDIA Lcuckart, 1879 (78). while finding the latter to be 7-8 p. Order EUCOCCIDA 1,Eger & Ihboscq, 1910 (78). Nematopsis soyeri (Theodorid&, 1965) n. comb. Syn.: Poro- spora soyeri ThCodoridb, 1965 (203). In gut of the penaeid Family SELENOCOCCIDIID.4E Pochr, 1913 (153). shrimp Ari$teus nntennntul (Risso) . Intermediate host un- Genus Selenococcidium LCger 8: Duhoscq, 1909 (120). known. Banyuls, France. Drsportrs and Th6odoridi.s (45 ) studied Selenococcidium infermedium Legcr & Duboscq, 1909 ( 120). the ultrastructure of the gymnospores. This sprcirs is transferred In gut of European lobster Homarus. France. This is the type to genus A’eniutopris hrcause its vcgetative stages are similar to of the gcnus and the only species in the family. The authors de- those of confirmed species of this genus in crabs and shrimp hut scribed it only hricfly in 1909 and niurc fully in 1910 (121). strikingly different from those of the single confirmed species of Porospora known only in the lobster. Family AGGREGATIDAE Labhe, 1899 (107). Nematopsis sp. Krusc, 1966 (100). In the gut of the shrimp, Griius Aggregnta Frrnzrl, 1885 (fil), emend. I.6Srr C Dul~oscq, Penaeus duorarum Burkcnroad. Intermediate host unknown. This 1906 (114). is the “small gymnospored specirs of Neniatopsis” which was found in shrimp with N. duorarum (the “large gymnospored AggrPjinta coelornica LkSer, 1901. rmend. Leger and Duboscq, species”) but which \\as not produced by spores from the 1906 (114). Syn.: Aggregata coeloniica L6xrr. 1901 [partinil pclccypods. Florida, USA. (110, 114). In gut ivall of the crab Pinnotheres pi;um Pcnn., Nematopsis sp. Sprasue, 1954 (179). In 1932. Pearse (142) \I hich lives in Mytilus edu1i.s L. Th(, original “sprcirs” rventually rcportcd the presence of “~rrgarine cysts” on the walls of the hecarne in part the grrgarinr, Cephaloidophora foisor. France rertuni of the ralico rrah Eriphici Conagrn (Fahririus) collectcd (Errglish Channel) f 1 IO). Found recently hy Vixar6s and Rubi6 at Dry Tortugas off the coast of Florida, USA. Sprague (179) (213), in the same host. on the northcast coast of Spain. cxprcsscd the opinion that this is a h’eniutopsis, not only because Azgregata eberfhi (I,ahht, 189.5) L6grr & Duhoscq, 1906 the limitcd description is charactcristic of the porosporid grega- ( 115). Syns.: Aggregata portunidnrum Fren~el. 1885 I partirnl rims hut also bccause thc hest known spccics A’. legeri, also (61); Benedenia eberthi LabhC, 1895 (105) [cited by LahbC, occurs in a species of Eriphia, E. rpinifrons. 1899 (lo;), altho wc can find no tnention of this name in the Neniatopsis sp. VivarPs 8: Ruhi6, 1969 (213). In gut of the 1895 paperl. Schizogonic stages in the portunid crabs Macro- crab Ethuw musrnrone. Northeast coast of Spain (Blanes). pipus (= Portunus) arcuatur (61). M. depurntor (115, 213): Nematopsis sp. Vivarh C Rubi6. 1969 (213). In gut of the M. tuberculata (213), M. 6oliz.uri (213). ill. corrogatus (214) crab Parthrnope nngulifron~. Northrast roast (if Spain (Blanes) and probably, M. uernalis (214). Sporogony in thr cuttlefish [A’enratopsis sp. ?] “Gregarines” Cuckler & Fichter, 1939 (42). Sepia officinulir. Common ar.d widespread. Naples, Italy (61 ) ; In the gut of thc crab Cancer productus. Argyle Bay, San Juan Site. Francr (1 15) ; l‘unisia (211); northeast coast of Spain Island, Washington, USA. Since this grezarine \\as found in a (213). Since the synonymy is quite complicated, only the most crab host and since as many as 3 nuclei \\ere seen in 1 com- significant synonyms are gken here. [See Lahbb, 1899 (107) and partment (suggrsting fusion of satcllitrs), there is reason to I,(.

the cysts in thr. silt \vall hrlonged to a fireprint. xvith frcr stages description also A'. michnelis Spragne. 1970 (191). the sporrs in tlie linnen. altho lic sa\\- only cysts of this spc~cirs. L6gi-r and of nhich he mistook for sporohlasts of A'. ctipidi. Duhoscq i 1 14, I 17) pointed out the incorrt-ctness of this viv\v. .Voiema sprlotremoe Guyinot 6: Poncc. 1925 (66). Hyprr- A,cgregiitn lrnndri Pis1.1l-Goodrich. 1950 ( 152). Schizogony parasitic in encysted metacercariae of the digrnetic trematode stages in gut wall of the shrimp I.eonder rquilln I,. Molluscan Microphnllus (= Spelotreina) cnrciui (198), which is parasitic host unknov n. Naplrs. Italy. Found also in thv shrimp Solenocera in the crah Cnrcinus inaennr. Stanier et al. rrcrntly (193) did inembranncea (Risso) and Acnnthephyra eximin Smith by Thfo- an rlcctron microscope study of this species. dorid& (204) at Banyrils. Francr. A'oserna sp. In 1963, one of us (V.S.) found a species similar Aggregatn xzgnnr 1,Pger S: Duboscq, 1903. emend. LGSer & to A'. spelotremne in encysted niicrophallid metacercariae in a Duboscq, I906 ( 1 1 I ). Syti : Agyregntn i'ngnnr I&rr & Duhosrq. crab Cnlliiircter rapidus collected in Pry Cove on the rastern 1903 [partinil (1 12). In gut wall of the rrah Eupocurur shore of Chesapeakc Bay, Maryland, US:\. hl. W. Newnian pridenuxi Leach. Ranyuls. Francr. Thr original species eventually ( ~irrs.cnm.) found a Xosenra, possihly thr samr spcries also, in hrcame in part the qrsarinc. Ceplrnioidopliora ocrllata. North (:arolina. South Carolina and Georgia, USA, in 1969. Aggregatn sp. I.(.gcr & Dnhoscq. 1908 (117). In codom of the crab Pachygiopriis mnrinoratuc F. Franrc (1 17). Found Genus Gurleya Doflcin, 1898 (51). ri~r.i~ntlyin the saiiic host by \'ivar+s and Rubii, on the northeast Gzirleyz mij,nirii (Kudo, 1924) Spragne, 1970 (191). Syns.: mast of Spain (213). 7'helohnnin sp. Miyairi, 1909 ( 102) : Cocconrinn inij,uirii Kudo. Aggregata sp. Srtna Rhatia, 1934 (1 73). Schisojiony stages 192-4 (102). In thr niuscles of 2 sprcies of Atj'ephira. Japan. in gut \\all of thv shrimp Pnr~ipeii~~of~iivsculptilir (IIrller). Mol- luscan host nnkno\vn. Bornhay, India. Genus 7'helohniiin Henneguy, 1892 (77). Aggregnta sp. Vi1ari.s Kr Rubih. 1969 (213). Unidentificd Thelohania cnnibari Sprasue, 1950 (178). In muscles of the species of <.iggregntn \vert% found in the crahs Coiystes cnrri- crayfish Cambarus bartoni. Georgia, US.4. i,elaunui. rlteleci'rlur iotundur. Xniitho pore,rn. (;oncplrix rhoiii- Thclohunia contejeani Hrnneguy. 1892 (77). In muscles of boides, Parothenope angiilifrons. and Macropodin rostrota. North- tlie crayfish A~tacusfluiiatilis (77); A. paliper (53, 168) and coast Spain. rast of .4. astncus (53, 199). France (77); Germany (168); Russia (170) Aggregntn sp. \'ivar+s. 1970 (214). C..nidentificd spi and Finland (199). Aggregatu \\ere found in tlir craln CoiciiiziT nieditrrrnnrus. Thelohania rfuorarn Iverson & Manning, 1959 (83). Syns.: Dromia perronntn. Poituiiinuc lntipes. Xnim biguttntn. Mncro- Thelohonia hunterae Jones, 1958 [norncn nudu~nl (86) : Thelo- pipus z,ernali,. Pilu~~~niicrpinijer and Piiuninzir hirtelliir. Coast hnnin sp. Kruse, 1959 (99). In muscles of the shrimp Penaeus of Tunisia. duorarum (83, 80, 84), P. brnsiliensis (84) and P. nrtecur (80). Florida, USA. Subphylnni II1CROSPOK.Z Spra:nr, 1969 ( 188) 7'helohnnia giardi Hennzguy. 1892 (77). In niusclrs of the Class I\IICROSPORE.\ Corliss bi 1.c~inc, 196:i (38). shrirrip Crnngon i'ulgnris. Francr. TYPESITCII.~ hy snliscqiic~nt dtGgnation, Gurley, 1893 (Gj). Order hIICROSPORIDA Balliiani, 1882 (6) Thelohanin macrocyrtir Curley. 1893 (65). In muscles of the Family NOSEAIATIDrlE Lablit?. 1899 ( 107) shrimp Pnlarnioneter I nrieiis. Italy. Kudo (IO?) qncstionrd the Genus A'osenin Niigeli. 18.57 (133). rnicnd. P6ret. 1905 (I 16). \.alidity of this species. Norema fren-elinne LCgw S: Lhlioarq. 1909 (118. 119). Hyper- 7'hrlohania innenndis Perez, 1904 ( 144). In inusclrs (usually) parasite of the gresarinr Cephaloidophorn confoi mir (Dirsin:) of the rrah Cnrrinus maenas. Ptrez (147) reported 1 caw in in the crab I'nchj,grnp,zir riiormorntus (Fabricius). France. \\ hirh an ovary was parasitized. resulting in parasitic castration. A'orema michnelir Spragiie. 1970 (191). Syns.: Soirmn supidi 'I'his parasite. first rqiortrd liriefly. \\as drscrihrd inore fully in 19U5 (1-16). France (Arcachon). DeTurk. 19-10 ipartiml (47) : 1Y. si'. Spragne. 1965 (182). In muscles of the rrah Cnllinectes snpidur Rathbun. .4tlantic and 7'helohnnia octospora Hennrguy, 1892 (77). In ninsrlrs of Gulf statrs of Xortti America. .A significant factor in crab mor- tlir shiinip I'niocmon rectirostris, P. rerrntur and P. elegnnr. tality. DeTurk mistook spnri~sof this specks for sporoblasts of A'. France. England and the Black Sea (76, 77. 151). rapidi. Spragne et al. 1187) niade an clectron niicrosropr study Thelohania paguri Ptrez. 1927 (148). In hody cavity of the of A'. michneli.. hcrmit crab Eiipngurur bernhordus. France. Thelohonin prnnei Spragur, 1950 (177). Jn gonad of the Xorenia nelioni Sliraqne. 1950 (177). Syn.: Iriotll .Yorcriio shrimp J'eiiaeu~ setiferur. Louisiana, USA. H. F. R. Champion puli.is: Jones (86). In riiiisclrs of prnat,id slirinip. M'idcly dis- (pers. coni.) found a siniilar or identical parasite in tlir ovary tributcd. First rr1)orted in Peiioeur nztcric, I\ cs in Louisiana. of indicus in the Republic of South Africa. USA. Later found hy IVoodburn rt al. (217), Hutton (80) and Penaeur Thelohonin petroliithir ). Syn.: Hutton et al. (79) also in P. duoraiiini BurLenroad in Ilorida. Spragur. 1970 (191 Thrluhnnia sp. Spraguc, 1950 ( 177). In niusclrs of the crah Petrolirther US.4. Recently found by Sprague and Vcrnick (189). \\ho aimatus. Louisiana, US.A. studird its fine atrurtnrr. ill P. .setifeinr from Gcorgia. US.\. 1%. Thelohania sp. Sogandares-Rrrnal, 1962 ( 175). In musclrs of F. P. Champion (prrs. ~oni.)ford a similar or identical spr- the crayfish Cambarclluc schujeldi. Louisiana. L'S.4. cics in the iiiiisrIi.s of /\letnpeiiiieus mo~~ocercosin thc Rrpublir of South hfrica. Kruse ( prrs. corn.) reported h'osemn sp.. which 7'helohanta sp. Thomas. (M. M. Thomas. Pen. coni.) In gonad may he idrntiral \\ith .Y, nrlmiii. in the ahrinip Solenocera and ~nusclcsol the shrimp. Penacus ieniirulcntus dr Ilaan. South zioscni and Siphopciieu, krojrii. in Louisiana. USA India. C;CIIUS Pleiitophora Giirlfy, I893 (65). A'osema pii!i,ic P6rzz. 1905 ( 14.5). In rnuscles ril tlir crali Carcinur moen(i\. Fr;inc i'. First dtwrilied \cry hriefly. this para- Pleirtophorn cargoi (Sprague, 1966) Sprague, 1970 ( 191 ). site \\as dcwribeti niorr fully in thr same year ( 1.46). Syn.: Plistophora cargo; Spragnr. 1966 (183). In niusclrs of Xos~marnpidi Dc'Iiirk, 1950. cnicnd. Sprajinr. 1970 (191) thc crah Calliurctes rapidus Kathliun. Maryland. US.4. Syn.: ,Vorema sirpidi lkliirk. 1910 lpartiinl (47). In muscles PIeistophorii miJairii (Kudo. 1924) Spraguc, 1970 (191 ). of thr crab Cnlliriertei iapidur Ratliljuii. Rraufort. North (;sic>- Syns.: A'osemn airoirinluin ( 7) Miyairi, 1909 (102) ; Plistophora lina, YSA. This is diatiugiiishrd from the otlicr :+ sprcies of miyazrii Kudo. 1924 (102). In digestive tract of Atyephira sp. ,Vo.sema descrihed iii muscles of decapods by having rrlatively Japan. large spores \shich (\\hrii staiiird) ha\? light arras at thr 2 Plei3tophoia Jogaiidaresi (Sprague, 1966) n. corrib. Syns.: rnds and a dark I,and iii thv iiiiddlc. llcl'urk includrd in his Pli~tophora sp. Sogandares-BernaI ( 175) ; Plistophora rogandure~i PROTOZOAOFDECAPODS 53 1

Sprague, 1966 (183). In muscles of the crayfish Cambarellus puer in sills of the follonin~marine crabs along the west coast of Hobhs. Louisiana, USA. Since Plistophora Labbk. 1899, is a Francr: Macropipus f = Portunur) depurator at Banyuls and junior synonym of Pleistophora Gurley, 1893 (186, 192), this Roscoff. M. puber at Roscoff and Wimereux, M. holsatus at species is transferred to the latter gmus. Roscoff. M. purillus at Roscoff and Banyuls, M. tuberculatus at Pleistophora sp. Baxter, Rigdon & Hanna. 1970 (15). In the Banyuls. and Cnrcinus maenas at Roscoff. Recently found by niusclcs of the shrimp Penaeur ietiferus and P. aztecus. Galveston Vi\,arPs and Rubi6 (213) in Macropipus tuberculatur, M. bolir,ari, Bay, Trxas. USA. Probably identical with Plictophora sp. Kruse, M. depurator and M. t,ernalis on the northeast coast of Spain. 1970 (191) found in the same host. VivarPs (214) found the folloning decapods infected with S. Pleisfophora sp. Sprague, 1970 (191). In muscles of the shrimp hypertrophica in the Gulf of Tunis: Dorippe lanata, Carcinus Pnlatnionetes pugio Holthius. Georgia, USA. mediterraneus, Portunrnur lntipes. Macropipus arcuatus, M. cor- rugatus. M. imernalis, Pilumnus Ipinifer and P. hirtellus. This Class HAPLOSPOREA Caullery, 1953 (27). apostonie causes considerable pathogenesis in gills of hosts (32). Ordrr H.4PI.OSPORID.4 Caullery & Mesnil, 1899 (24). Life cycle fully described and figured by Chatton and Lwoff Faniily H.API.OSPORIDIIDAE Caullery & Mesnil, 1905 (25). (32). Genus Minchinia Labbe, 1896 (106). Genus Gj.n~nodinioidesMinkiewicz, 1912 ( 130). Minchinia loui\iana Sprague. 1963 (181 ). Syn.: Haplo- sporidiunz sp. Sprague, 1954 (179). In the gut wall of the mud (r'ynznodinioides calkinsi Chatton & Lwoff, 1935 (32). Phoronts crab, Panopeur herbstii. Louisiana, USA. on the gills and trophonts on the cuticle of Polaemonetes sp. Woods Minchinia sp. Rosenfield, Buchanan, Chapman, 1969 (162). Hole, Massachusetts, US.4. In gill, niusclcs and gut tissue of the crab Eurypnnopeu, de- Gynznodinioides caridinae (Miyashita, 1933) Chatton & Lwoff, pressus. Virginia. USA. 1935 (32). Syn.: HyaIospira caridinae Miyashita, 1933 (132). Found on the "hairs" at base of antennules and on the gill fila- Genus Urosporidium Caullery & Mcsnil, 1905 (26). ments of Xiphocuridinu comp7essa (freshwater shrimp of Japan), Urosporidium crescens DeTurk, 1940 (48). Hyperparasitic in Leander paucidens and Macrobranchium nipponense by Miya- encysted metacercariae of a distome identified by DeTurk as shita in Japan. Reported as Hyalospira caridinae from Palaemon Spelotrema nicolli, now in genus Microphallur (198). Nortli rarianr by Debaisieux (4.4) in Europe. Debaisieux gave a fairly Carolina (48): Maryland and Virginia (191), USA. Sprague detailed account of the life cycle of this species on the brackish (191) recently made observations on the structure of this species, water Palaenion Lmarians. Couch (41) studied its effect on its trematode host and Perkins ~Ij~~rinodinioiderinkytnnr hlinLicuicz. 1913 ( 130). Phoronts (149) studied its ultrastructure. on the gills and trophonts on the cuticles of Eufagurus bern- lirosporidium sp. Sprague, 1970 (191). Hyprrparasitc in cp.- hardus, E. prideauxii (130) and Carcinus maenas (32). Roscoff, cystcd metacercariae of Microphallus sp. in the shrimp, Palae- France (32, 1:iO). Also possibly observed by Minkiewicz, accord- monetes pugio. Georgia, USA. ing to Chatton and Lwoff (32). on the following decapod genera: Maia, Xantho, Porcellana and Galathea. Trager (207) reported this species from the gills of Pagurus longicarpus at Woods Hole, Subphylum CILIOPHORA Doflein, 1901 (78). 3lassachusetts, USA. According to Trager, the excystment of the Class CILIATEA Perty, 1852 (78). phoronts of this species is synchronized with the molt of the Subclass HOLOTRICHIA Stein, 1859 (78). host, and the conditions necessary to bring about excystment Order ASTOMATIDA Schewiakoff, 1896 (78). can be simulated in the laboratory, Family ANOPLOPHRYIDAE C+de, 1910 (28) Gymnodinioides sp. Chatton & Lwoff, 1935 (32). In the Genus Anoplophrya Stein, 1860 (136). #astrovascular cavity of the anemone Sagartia parasitica found Anoplophiya minima Lkger & Duboscq, 1909 (120). This attached to Eupugurus prideauxii. very rare species of astonie ciliate was reported from the posterior Gymnodznioides sp. Trager, 1957 (207). Phoronts on gills of intestine and rectum of the lobster, Hornarus gamniarus. France. Uca pugnax. Woods Hole, Massarhusetts, USA. About 80% of the crabs examined vere infested. Trager reported differences in Order HYMENOSTOMATIDA DPlage & Htrouard, 1896 (78). this species and G. inkystanr to be 10 or 11 ciliary bands rather Suborder TETRAHYMENINA Faurc5-Fremiet, 1936 (78). than 9 as found on the latter. Also, this undescribed species had Family COHNILEMBIDAE Kahl, 1933 (78). a more pointed and twisted posterior end than G. inkystans. Genus Anophrys Cohn, 1866 (34). Genus Phtorophrya Chatton, Lwoff & Lwoff, 1930 (30). Anophrys maggii Cattaneo, 1888 (23). Syn.: Anophrys sarco- phagu Cohn, 1866 (34). Occurring naturally, but rarely, in PhtorophrTa fallax Chatton, Lwoff & L~off,1930 (30). A hemolymph of Carcinus maenas. Western coast of France (155). hyperparasite on the phoronts of Gymnodinioides inkystans on Poisson (155) stated that the parasitic A. Maggii of Cattaneo Carcinus maenas. Roscoff, France. is identical witb the free-living A. sarcophaga of Cohn, and thus Phtorophrya iniidiosa Chatton. Luoff & Lwoff, 1Y30 (30). the narnc A. maggii should be a junior synonym of A. sarcophuga. Usually found as a hypcrparasite on the phoronts of Gymnodinio- However, Tucker (14) argued that until more is known about ides corophii on amphipods, but in at least I instance (32) re- these forms both names should be retained. Bang (14) was ablr ported as a hyprrparasite of Gymnodinioides sp. from Carcinus. to induce fatal experimental infections of A. maggii in Cnrcinus Membrrs c)f this grnus are characteristically hyperparasitic or mama\, Main squinado and Macropipus depurator. The ciliates phoretic on the phoronts of other apostornes (32). \vithin the hemolymph of these crabs were found to be eating host leukocytes. Bang (14) reported that the events leading to Genus Polj..ipirn Minkiewicz, 1912 (130). death in these Anophrys-infected crabs uere anemia and as- Polj'ipiru delagei Minkiewicz, 1912 (130). On the gills of the phyxiation folloMed by severe bacteremias. follouinx hosts: Eupaxurus bernhardu,, Galathea Jquamifera, and Macropodia rostrata. Roscoff, France. Found only in de- Order APOS1'0MATID.4 Chatton 8z Lwoff, 1928 (78). capod crustaceans (32). Chatton and Lwoff were unable to Family FOETTINGERIIDAE Chatton, 191 1 (29, 32). transmit it easily to Mrrcrupipu, si'. in the laboratory (32). These Genus Synophrya Chatton & Lwoff, 1926 (29, 32). authors reported that this species is often found in company with Synophrya hypertrophica Chatton & Lwoff, 1926 (32). Found Gymnodinioides inkystans. 532 PROTOZOAOF DECAPODS

Genus Phofetophryo Cliatton. 1,uofl & Luiiff. 1930 (31) Zoothnmnium sp. Ryder. 1883 (163). On “back and legs” of Phoretophrjn nebaline Cliatton. I.\\off & L\\off, 1970 (31 ). the oyster r~inini~~nsaldecapod Pinnotheres ostrrum Say. Mary- .4ltho usually found on the non-decapod ,Yebalin geflroyi. this land, US.4. Ryder suggested that because thr Zoothnmniuni sup- ported on its “stalks” hactrria and “\ibriones.“ it might he of species has also heen rrportt,d from Cnrc;firrr innrnni and Mocrop- ipus hohatus. RoscoIT. France. henPfit as a source of food to the oyster in \\-hich its crab host lives.

Genus Terebrorpira Debaisieux. 1960 (44) Family EFISTYLIDID.4E Kahl. 1933 (87). Tercbrospirn lrnticiclaris (Debaisicux. 1957 ) DrhaiFieux, 1960 (;<%nusEpirt1,lir Ehrenherg. 1832 (58). (44). Syn.: Chnttonin lenticularis Debaisieux, 1957 (43). On /ipi\tTlir nrtnci Nenninser. 1948 (137). On the gills of Artncuc the cuticlr of Pnlnemon xiinns collrcted from the mouth ol Pu:,intilir. Germany. Schclde Rivrr (brackish \later) in Western Europe (Netherlands). Epi\t)’lir bimor,qinotn Kcnningrr. 1948 (137). On legs of Debaisieux (4 t) gavr thr life cycle and host-ciliate relationship Astucus pui,intilir. Germany. Nenninger described this peritrich of this species in soinc detail. This specks peiietratcs the cuticle from the crayfish as a \.ariety. E. bimarginnta var. urnula. hut of its host shrimp and thus can cause some damage. asain, until subspecific namrs are tnore generally accepted for ciliates this form is recognized only as a member of the species Grnus Wyalophysn Bradbury. 1966 (21 ) , E. bimuiginata. Nyalophysa chattoni Bradbury, 1966 (21 ). Phoronts found Epirt9’lis canibnri Krllicott, 188.5 (94). On gills of a species on the gills and trophonts on thc molts of scverai diffrrrnt spr- of Caniborur. Sorthcastvrn United States. cics of decapods froni the coasts of Washington and California. Epiltylir ningarne Krllicott. 1883 (92). On body surface of USA: Pagurui hirsutiusculus. P. sniriuelir. P. ,graiio~rm~nur.P. crayfish ( ? j in freshwater. Northern United States. alaskeniis, Cnncer oregonenris, and Lophopanopeur belli. Ac- I:‘pist),/ic sp. Hutton. 1964 (80). On the coniiiic!rcial shrimp cording to Bradbury. the gmiis H1,alophysn brlongs to that group Penaeus duoruruni. Numerous localities in Florida, USA. of apostonies phoretic on Branchyura. .4noniura, Isolwda and Epiil3’/i

Genus Kentrochonn Koinpel, 1894 (161) Genus Cothurnia, Ehrmhrrg, 1832 (58) Kentrochonn nebnline Rompel, 1891 (161 ). On the dccapods Cothurnin nrtnci Strin. 1854 (195). On the branchial all- Ebalia turnefnctn and Portunus depurator. Southern Sweden. pendages of Actacur j?u:,intilir. Europe. Thought to he associated Also rcported by DoRein (50) from Tricstc and Rovigno. in the with mass mortalities of fresh\\-atrr crayfish in Italy as reported northern Adriatic. by Ninni (139) and by Kent (98). Kent (p. 721) said, “During a fatal epideniic among the freshnater cray-fish of Lomliardy and Subclass PERITRICHIA Calkins, 1933 (78) Venetia in the year 1863, it is reported by Ninni that the dying crustacea were completely iiifrsted uith an Infusorium, referable Order PERITRICHIDA Stein. 1859 (78). to the genus Vaginirola. prohal)ly identical with the present Suborder SESSILINA Kahl, 1933 (78). species IC. astoczi. its abnorinal de\ clopmcnt on the branchial Family VORTlCELLIDAE Ehrenherg, 1838 (140). appendages apparently producing the death of the crustacean Genus Vorticella L., 1767 (136), emend. Ehrenberg, 1838 (140). through asphyxia.” Vorticella ~‘errz~cosuDons, 1915 (55). Found on Laomedia sp. Cothurniu currn Stein, 1851 (195). From fresh\\ater crayfish. from Norway. Of many dcscribcd species of Vorticella (87; 137; Europe. 140), this form is the only named vorticellid habitually found Cothufnio ricboldi Stein. 1C1-1 (195). On thr liranchial ap- on decapods. Many species have been reported from tion-dccapod pendages of the frcsh\\ater crayfish Astacus purintilis. Europe. crustaceans. Cothurniir ;,urinbilis Kellicott. 1884 (9‘3). Occurs by thou- sands on gills of crayfish Cambnrur afinis (93) and Astncus Genus Zouthamntuin Bory, 1826 (19) flui’iotzlir (137 j. Zoothamnium carcini Kent, 1881 (98). On Carcinus maenaz Family LAGENOPHRYIDAE Kahl, 1935 (89). from the south coast of England. Genus Lngcnophrj’s Stein, 1852 (194). Zoothamniuin procerius Kahl, 1933 (87). On legs of the crayfish Astucus fEui,iatilis from \ iciiiity of Erlangrn, Germany. This large genus of loricate peritrichs (52 species at present) Nenningcr (137). reported a xarirty of this species, which she is rmiarkalh in that it has 22 species that are found only on named Z. procerius \ar. inrius from both gamtnarids and cray- decapod crustaceans and ha\ e strong host sprcificity. fish, but until subspecies names are more generally accepted for Lagenophrys aegleae Mouchet-Bennati, 1932 ( 134). On the ciliates, this form is recognized as %. procerzus. sills of the freshnater crayfish Aeglea laetis. Uruguay. PROTOZOA OF DECAPODS 533

Lagenophrys articularii Nie 8: Ho, 1943 (138). On the anten- hased on specimrns found on a land crayfish Engaeus fultoni nae, legs, and s\\immerets of Polaenion nipponen\is. China. lrom a burro\\. in a river bank near Melbourne. Australia. Found Lagenophryr rir~~erinrrrc~iAhonyi, 1928 (1). On gills of the zttached to setae, tail fan, sjvimmerets, gill base and edge of frcshnatrr rivw crah Trlphurn fluciotilii. Africa. .2honyi gave branchiostrsites. Specimens referable to this species have also thi, lagmophryiid its specific tiam? hsed upon thc dc~.scription hren found on Chernx rotundus setosus, C. albidus, C. destructor of A\\erinx\v (5). and Bunitacus nobtlis from Australia. Lagenophrys birpinow Kaue, 1965 (91). On suimnicrcts of Lagenophrys spinora Kane, 1965 (91). On the cuticle, swim- Chernu rotundur teto%ur. Stream nrar Nc\\castlc. Nr\v South merets, telson and carapace of the crayfish Cherax destructor. Walrs, Australia. Victoria. .4ustralia. Lu,qrnophryi brunchinrnm Nie & 110. 1943 (138). On the gills Lagenophrys rdlisi Kanr, 1965 (91). On gill filammts of (only) of Palaenion 7iifiponeii,,is. China. fresh\*.atcr crayfish Cherax destructur, C. albidus and C. rotundus. Lagetioflhrj,~cnllinectes Couch. 1967 (40). On both surfaces Australia. of the gill lainellac of the conitrirrcial, niarine and hrackish 1% ater crah Cnllinecter sapidur. From Chesapeake Bay, Mary-land Subclass SPIROTRICHIA Biitschli, 1889 (78). aud Virginia: south to Gulf of Mi.xico estuarine uaters wherever Order HETEROTRICHIDA Stein, 1859 (78). blue crab is found. Durins summer months infestations are so Suborder IfE'I'ERO'lKICHINIA Stein, 1859 (78). hea\y that the gill surlaccs of the crah may be rompletely Family FOLLICVLINIDAE Dons, 1914 (54). covrrcd. hlrmbcrs of this family belong to several different genera, all Lagenophrp coniniunis Knnc, 1965 (91). On the telsiins, species of \\hich are loricatr. Species reported here as epibionts suinrincrcts, and cuticles of the crayfish Chernx deitructor, C. of decapods also may be, and usually are, found on substrates nlbidui, Ijicu'tocus surrutu<. Ii. nobzlis, and Engaeus sp. Aus- other than decapods and generally lack host specificity. FaurE- tralia. Fremict (60) and Hadii (68) reviewed the species in this family La,qenophrj,r darri ini Kaur, 1965 (91 ), On the gill filaments and their various substrates. of Cherux quudricnrtnatu~. I?r&\\ater. uear Daru in, northern .lustralia. Genus Pebrilla Giard, 1888 (63). 1.ngenophrj's deserti Katie, 1965 (91 ). In the branchial cham- hers and on gill filaments of Cherax tenuinranus and C. quinque- Pebrilla papri Giard, 1888 (63). Found on the cuticle of curinntus. Ri\ rrs in southrastcrn .4ustraha. Pagurus bernhardus by Giard, in 1888, at Pouliguen and at Lngenophrys dungogi Kaur. 1965 (91). On the gill filaments Rosruff, France. Latrr (1936), reported from cuticle of Cliba- of a single specimen of Euartacus si). Stream in Dungog, norius misanthropus from France by Fad-Fremiet (60), who Australia. helievrd that. tho this sprcies has fratures characteristic of Lagenuphi).\ engnei Kane, 1965 (91). On brachiostegitr members of grnus Folliculinopszs FaurE-Fremiet,l936, its generic membranes and sill filarncnts of crayfish Engaeus rictoriensis, and name Pehrilla should lie retained. on gill filanient and gill bases of crayfish Austroactncus hemi- cirratulis. Near Melbourne. ..lustralia. Genus Plntyfol/iculina Hadii, 1938 (67). Lagenophrys eupagurus Kellicwtt, 1894 (96). On gills of P/atyfu//iculina lJnguri Andrews & Reinhard, 1943 (3). On hcrmit crab 1:'upaguru.i sp. M'ocids Hole, Massachusetts, USA. the abdomen of hermit crab Pagurus pubescens. State of Wash- Lagenophrys latiipinosn Kane, 1965 (91). On the flat surfaces ington, USA. of gills, gill bases and hranchiostegitc mcnibranes. On Cherax dertructor near Melbourne, Liustralia. On C. dertructor and C. Genus Metafolliculina Dons, 1925 (56). nlbidus from Victoria, .4ustralia. On C. rotundu, from Nc\r Metafolliculina nndrezc,ri Hadii, 1938 (67). On Pagurus pori- South Wales, .4ustralia. Kane remzrked that the loricae of many danus as reported by Andrew (4). Texas Gulf Coast, USA. generations of this specics arc supi~in~posedon one ariothcr on Andreus stated that tlie trsts of M. andreursi were found on the the cuticle of older hosts. abdominal filaments, and "large holding appendages." This spe- Lagenophrys lnzi'rii Kane, 196.5 (91). On flat surfaces of the cies \\as also found on oyster shells in same region by Andrews. gills of Cherax quadricarinatuc. Near Darwin, Australia. Lagenophrj'r lingulnto Kanc, 1965 (91 ). Found mostly on gill Genus Folliculinopris FaurE-Fremiet, 1936 (60). filaments, occasionally on thr I~ranchiostegite membranes, On Cherax destructor and C. ailridus from Victoria, Australia, aiid on Follicdinopsis gunneri (Dons. 1928) FaurE-Frrmiet, 1936 (60). C. rotundus in central Neiv South \Vales, Australia. Type host Syn.: Folliculina girnneri Dons, 1928 (57). Found by Faure- and locality are Chernz destructor from around hlelhournc. Freniiet on thr cuticle of sacculinid parasite of Carcinus maenas Australia. and Porccllana plat),chrlri. France. Lagenophiys lunutus Imaniura, 1940 (81 ). On body cuticle of shrimp Leander pnucidens (83) and Palurnion i,arians (4arthropods. In S. F. Snirszko, ed. A Symposium on Acad. Sci. 260, 1761-2. Dicenses of Fishes and Shellfishes, Spec. Publ. 5, Am. Fish. SOC., 46. Desportes, I. 1968. Sur la prCsence de Cephaloidophora Washington, D. C., 383-404. olim'a (Watson ) (Eugregarina, Cephaloidophoridae) chez le crabe 15. Baxter, K. N., Rigdon. R. H. 8: Hanna. C. 1970. Pleisto- Ocypode cursor L. (Crustacea Decapoda) au Liban. Vie Milieu 19( phora sp. (Microsporidia: Nosematidae) : A new parasite of I-A), 201-2. shrinip. J. Inz,ert. Pnth. 16, '189-91. 47. DeTurk, W. E. 1940. The parasites and commensals of 16. Beauchatrip. P. de 1910. Sur tine gregarine nouvelle du some crabs of Beaufort, North Carolina. Thesis, Duke University. genre Porosfiorn. C. I(. Acnd. Sci. 151, 997-9. Unpublished. 17. Bcneden. E van 1869. Sur tine nouvelle espPce de 48. __ , 1940. The occurrence and development of a hyper- Gregarina designte sous le nom de Gregarina gigantea. Bull. parasite, Urosporidium crescens (Sporozoa, Haplosporidia) which Acad. Iio)'. Belg. 28, +44-56. infests the metacercariae of Spelotrema nicolli, parasitic in Cal- 18. Bogolrpova, I. 1. 1953. Gregarines of Peter the Great Bay. linectes sapidus. J. Elisha Mitchell Sci. Soc. 56, 231-2. Tro,. Inlt. ZOO/.Acad. Sci. I.IISS 13. 38-56, [In Russian]. 49. Dobell, C. 1925. The life-history and chromosome cycle 19. Bory de Saint-Vincent. J. B. 1826. Essai d'une Classifica- of Aggreyata eberthi (Protozoa, Sporozoa, Coccidia). Parasitology 1: 1.12Ii tion des Animaux Microscopiques. Paris. 104 p. - ., . .I - . 20. Bovee. E. C. 8: Jahn, T. L. 1966. Mechanisms of move- 50. Doflein, F. 1897. Studien zur Naturgeschichte der Proto- ment in taxonomy of Sarcodina. 111. Orders, suborders, families, zoen. I. Kentrochona nebaliae Rompel. Zool. Jahrb. Anat. 10, and subfamilies of the superorder Lobida. Syst. Zool. 15, 229-40. 619-41. PROTOZOAOF DECAPODS 535

51. 1898. Studien zur Naturgeschichte der Protozoen. 82. International Coinmission on Zoological Nomenrlature. 111. Uher Myxosporidien. Ibid 11, 281-350. 1961. International Code of Zoological Aomenclature Adopted 52. Dogiel, V. .4. 1965. General Protozoology, 2nd ed. LRev. by the XV International Congres of Zoology. Intern. Trust J. I. Poljanskij & E. M. Chejsinl, Oxford, 747 p. Zool. Nomen., London. London, 176 p. 53. Dollfus, R. P. 1935. Thelohanose de I’ecrivisse (Astacu.s 83. Iversen, E. S. & Manning, R. B. 1959. A new micro- pnllipes Lereboullet j, Rirhelieu ( Indre-et-Loire) , BuIl. Soc. sporidian parasitc from thr pink shrimp (Penaeui duortirum) . Centr. d’Aquires 42( 10-12), 119-22. Trans. Am. Fish. Sac. 88, 130-2. 54. Dons, C. 1914. Folliculina - Studien IV. Vorliiufige Bemer- 84. Iversen, E. S. & Van Meter, N. N. 1964. A record of the kungen uher die Systematik der Folliculiniden nebst Beschreibung niicrosporidian, I’helohania duorara, parasitizing the shrimp. neuer norwegischer Arten. Tromsyl Mus. Aarsh. 37, 59-92. Penaeus brasiliensis. Bull. Mar. Sci. Gulf Carib. 14, 549-53. 55. 1918. Neue marine ciliaten und siictorien. Ibid 85. Jahn, T. L. & Bovee, E. C. 1965. Mechanisms of move- 38 & 39(1915/16), 75-1QO. ment in taxonomy of Sarcodina. 1. As a basis for a new major 56. 1925. M:.rrfolliculina nordgordi n. gen., n. sp. dichotomy into two classes, Autotractea and Hydraulca. Ani. Kgl. Norsk. Videnskab. 5elsk. Skr., 1924, 1, 1-13. Midl. Nat. 73, 30-40. 57. 1928(1927). Neue und wenig bckannte Protozoen. 86. Jones, E. E. 1958. Microsporidiosis in shrimp. Ass. South- Ibid 7, 1-17. east. Biol. Bull. 5, 10. 58. Ehrenbrrg, C. G. 1832. Ubcr die Entwicklung und Lebens- 87. Kahl, A. 1933. Ciliata lihera et rctorommcrisalia. In G. dauer der Infusionsthiere: nehst frrneren Beitrigen zu einer CrimDe & E. Waeler (ed.). Die Tierzuelt der Nord-und Ostee. Vegleichung ihrer organischen Systrme. Abh. Akad. Wiss. Berlin Lief. ‘23 (Teil II,-ct), ‘29-ii6. Leipzig. 1831, 1-154. 88. 1934. Suctoria. Ibid 26 (Td 11. e;), 184-226. 59. 1835. Dritter Beitragen zur Erkrnntniss der gros- 89. __ . 1935. I. Wiinoerthiere oder Ciliata (Infusoria). ser Organisation in der Richtung des Kleinsten Raumes. Abh. In F. Dahl, ed. Die Tierzvcli Deutschlands, G. Fisher, Jena,~i, Akad. Wiss. Berlin, 1833, 145.336. Peritricha and Chonotricha, 65 1-799. 60. Fauri-Frcmiet, E. 1936. La famille des Folliculir..dae (In- 90. Kamm, M. W. 1922. Studies on grcaarines. 11. Synopsis fusoria Heterotricha). Miui. Muc. Hict. Nat. Belg. (Ser. 2), 3, of the polycystid gregarines of the world, excluding thosc from 1129-75. the Myriopoda, Orthoptera and Coleoptera. Ill. Biol. Monogr. 61. Frenzel, J. 1885. Ueber cinige in Seethieren lebende Greg- 7, 1-104. arinen. Arch. Mikroskop. Anat. 24, 545-88. 91. Kane, J. R. 1965. The genus Lagenophrys Stein, 1852 62. Fromental, E. G. de 1874. Etudes sur les microroaires ou (Ciliata, Prritricha) on Australian Parastaridae. J. Protorool. 12, infuioires proprement dits comprenaut de nouoelles recherches 109-22. sur leur orgunisation leur classification et la description des 92. Kellicott. D. S. 1883. Efiistylis niagarue sp. 11. on cray- erp2ce.s nouzxlles ou jeu connues. Paris, viii + 364 pp. fish. Trans. Am. Micr. Soc. S, 110. 63. Giard, A. 1888. Fragments biologiques. XIII. Sur les 93. ~- . 1884. Observations on infusoria, with descriptions genres Folliculina et Pebrilla. Bull. Sci. Fr. Belg. 19, 310. of ne\\- species. [bid 6, 110-24. 64. Grassi, P. P. 1953. Classe des Gregarinomorphes. In P. 94. 1885. Epirtglis cumbari sp. n. on Canzbarus. Ibid P. (kassi, ed. Traitd de Zoologic, Masson, Paris, I(?). 550-600. i, 43. 65. Gurley, R. R. 1893. The Myxosporidia, or psorosperms of 95. 1887. Notice of some fresh-water infusoria. With fishes, and the epidemics produced by them. Bull. 0’. S. Fish. remarks on collecting and preservinl: these delicate animals. Conzm. 1892, 190-205. Microscope i, 225-33. 66. Guyhot, E., Naville, A. & Ponce, K. 1925. Deux micro- 96. __. 1894. On certain marine infusoria observed at sporidies parasites de trCmatodes. Rev. Suisre Zool. 31, 399-422. Woods Hole, Massachusetts. 2nd Ann. Rep. Ohio State Acad. 67. Hadii, J. 1938. Beitrag zur Kenntnis der adriatischen Sci. 10-1 I lAhstr.1. Follikuliniden. ( Inf. Heterotricha ) . I. Sul~fanl. Eufolliculininae. 97. Kenk, V. C. 1967. A new crab host of the grcgarine Nema- Acta Adriat. 2, 1. fopsis ostrearuni. Proc. Nut. Shellfiish. Ass. 6.5, 87-8. 68. 1951. Stiidien iiber Follikuliniden. Dela Raar. Pri- 98. Kent, W. S. 1881-1882. A Manual of the Infusoria rod. Med. Vedr. Slou. Akad. ZnGn. linnzet., Ljubljana 1-390. 2, 4, 473-913; 3, 51. Bogue, London. 69. Haeckel, E. H. P. .4. 1866. Generelle Morphologie der Organismen 11. Allgemeine Eiitzi,icklungegeichichte der Organis- 99. Kruse, D. N. 1959. Parasites of the commercial shrimps, men. G. Reimer, Berlin. Penaeus aztecus Ives. P. duorarunz Burkenroad and P. retiferus 70. Hatt, P. 1927. Pororpoia legeri de Beauchaxnp (= P. (Linnaeus). l’ulane Stud. Zool. 7, 123-44. galloproi8incialis Leger et Dubosrq) et les premiers stades de son 100. 1966. Life cycle studies on Neniatopsis duorari evolution chez I’Eriphia. Arch. Zool. Exp. Gin. N & R 67, 8-11. 11. sp. (Gregarinia: Porosporidae), A parasite of the pink shrimp 71. 1927. Le dCbut de I’ivolution des Porosfiora chez (Penoeus duorarunz) and pelecypod molluscs. Diss. Abstr. 27B, ICs mollusques. [bid 6i, 2-7. 2919-B. 72. __~. 1928. L’Pvolution de la gregarine du Homard 101. 1966. Liie cyclc studies on Nematopsis duoiari 11. (Porospora gigantea E. V. Bened.) chez les mollusques. C. R. si’. (Gregarinia: Porosporidae), A parasite of thr pink shrimp Soc. Biol. 98, 647-9. (Penaeur duorarum) and pelccypod molluscs. Ph.D. ‘l‘hesis, Florida 73. - - 1951. L’ivolution des porosporides chez les mol- State. Univ., Mirrofilrus, Inc., Ann Arbor, Michigan, i-x\- -+ lusques. Arch. Zool. Exp. Gdn. 72, 341-415. 17.4 pp. 74. Heckmann, R. 1961. Gregarines of Hauaiian marine crusta- 102. Kudo, R. R. 192-1. .4 biolosic and taxonomic study of the ceans. J. Protozool. 8 (Suppl.), 17 IAbstr. 611. Microsporidia. Ill. Bid. Monogr. 9, 1-268. 75. Heldt, J. H. 1954. La gresarine de la crevette caramote 103. 1939. Protozoology, 2nd id. Thomas, Spring- (P,enaeuc kerathurus ForskAl). Bull. Soc. Sci. Nut. TuniJie 6, field, Ill. xi + 689 pp. 103-1 12. 104. 1946. Protozoology, 3rd cd. Thomas, Spring- 76. Henneguy, F. & Th&lohan, P. 1892. Sur un sporozoaire field, 111. xiii + 778 pp. parasite de muscles des crustaces dtcapodes. C. R. Acad. Sci. 105. LabbC, A. 1895. Sur le noyau et la division nucEaire 114, 1552-5. chez les Benedenia. C. R. Acad. Sci. 120, 381-3. 77. 1892. Myxosporidies parasites des muscles chez 106. __ . 1896. Recherches zoologiques, cytologiqucs et bio- quelques rrustacCs dtcapodes. Ann. Microgr. 4, 617-41. logiqucs sur Ics coccidies. Arch. Zool. Exp. Gin. Ser. 3, 4, 533- 78. Honiberg, B. M. (Chairman). 1964. A revised classifica- 5:54, 608. tion of the phylum Protozoa. J. Protozool. 11, 7-20. 107. 1899. Sporozoa. In 0. Biitschli, ed. Dar Tier- 79. Hutton, R. F., Sogandares-Bernal, F., Eldred, B., Ingle, reich. 5, Friedliinder, Berlin. xii + 180 pp. R. M. & Woodburn, K. D. 1959. Investigations on the parasites 108. Lamarck, J. B. 1816. Histoire A’aturelle des Anirnaux and diseases of saltuatcr shrimps (Penaeidae) of sports and com- sans Vertibres. (11). Paris. mercial importance to Florida. (Preliminary report). Fla. Slate 109. Landau, H. & Galtsoff. P. S. 1951. Distribution of Nema- Bd. Conserc. Tech. Ser. 26, 1-38. topsis infection on thr oyster grounds of the Chcsapeakc Bay and 80. Hutton, R. F. 1964. A serond list of parasites from marine in other \\-aters of the Atlantic and Gulf Statrs. Texas J. Sci. I, and coastal animals of Florida. Trans. Am. Micr. Sac. 83, 439-47. 115.30. 81. Imamura, T. 1940. ‘luo species of Lagenophrys from Sap- 110. Ltger, I,. 1901. Sur iine noiiwlle Sregarinc parasite drs poro. Annot. Zool. Jup. 19, 267-70. Pinnotheres des moules. C. li. Acad. Sci. 132, 1343-6. 556 PROTOZOAOF DECAPODS

111. __. 1903. Sporozoaire parasite des moules et autres 142. Pearse, A. S. 1932. Observations on the parasites and lamellibranches comestibles. Ibid 137. 1003-6. commensals found associated with crustaceans and fishes at Dry 112. Leger, L. & Duboscq, 0. 1903. Aggregata ragans n. sp. Tortugas. Pap. Tortugas Lab., Carnegie Inst., Washington, D. C.. gregarine gymnosporCe parasite des Pagures. Arch. Zool. Exp. 28, 103-15. Gin. 31. 147-51. 143. Pellbrdy, L. P. 1965. Coccidia and Coccidiosis. Pub. 113. Lkger, L. 1905. Un nouveau Nematopsis parasite des House Hungarian Acad. Sci., Budapest. 657 pp. moules de la MkditerranCe. Paris Bull. Mem. Franc. Auanc. Sci. 144 Pkrez, C. 1904. Sur une mirrosporidie parasite du Carcirrus 9, 331. maenas. C. R. Soc. Lid. 67, 214-5. 114. Leger, L. & Duboscq, 0. 1906. Sur l’evolution des grega- 145. 1905. Sur une nouvelle gluq?idee parasite du rines gymnosporees des crustac6s. C. R. Acad. Sci. 142, 1225-7. Carcinus maenac. Ibid 58, 146-8. 115. __ . 1906. L’evolution d’une Aggregata de la seiche 146. 1905. Microsporidies parasites des crabes d’Ar- chrz le Portunus dejurator Leach. C. R. SOC.Bid. 58, 1001-3. cachon. SOL. Sci. Arcachon Trau. Lab. 8, 15-36. 116. __. 1907. L’Cvolution des Frenzelina (n.g.), gregt- 147. 1906. Sur un cas d’envahissement de l’ovaire par rims intcstinalrs des crustacfs d6capodes. C. R. Acad. Scz. lla, Thelohonia maenadis. C. R. SOL. Biol. GO, 1091-2. 771.4 .. 148. 1927. Notes sur les epicarides et les rhizocephales 117. __. 1908. Evolution schizogonique d’dggregata eber- des cBtes de France. I. Sur I”‘Eupagurus bernhardus” et sur thi. Arch. Protist. 12. 44-108. quelques-uns de ses parasites. Bull. Soc. Zool. Fr. 52, 99-104. 118. 1909. Sur une microsporidie parasite d’une 149. Pcrkins, F. 0. 1971. Sporulation in the trematode hyper- gregarine. C. R. Acad. Sci. 148, 733-4. parasite Urosporidiurn crescen.r DeTurk, 1940 (Haplosporida: 119. 1909. Etudes sur la sexualit6 chez les gregarines. Hap1osporidiidac)-An electron microscope study. J. Parasit. 57, Arch. Protist. 17, 19-134. 9-23. 120. 1909. Protozoaires parasites de I’intestin du 150. Pixcll-Goodrich, H. L. M. 1914. The sporogony and system- homard. C. R. Acad. Sci. 148, 363-5. atic position of the Aggregatidac. Quart. J. Micr. Sci. 60, 121. __. 1910. Selenococcidium intermedium Leg. et Dub. _-”159-74 , .. et la systernatiques des sporozoaires. Arch. Zool. Exp. Gin. Ser. 151. 1920. The spore of Thelohania. Arch. Zool. Exp. 5, 5, 187-238. Gdn. N & R. 59. 17-9. 122. 191 1. Deux gregarines de crustacbs. Porospora 152. 1950. Aggregata leandri n. sp. Quart. J. Micr. portunidarum Frcnz. et Cephaloidophora maculata n. sp. Arch. Sci. 91, 465-7. Zool. Exp. Gin., N & R 6, 59-60. 153. Poche, F. 1913. Das System der Protozoa. Arch. Protist. 123. 1911. Deux nouvelles espkes de gregarines ap- 30, 125-310. partenant au genre Porospora. Ann. Uniu. Grenoble 23, 401-4. 154. Poisson, R. 1924. Sur quelqucs grkgarines, parasites de 124. __. 1913. Le cycle evoliitif de Porosfora portuni- Crustarts, obserctes a Luc-sur-Mer (Calvados). Bull. Soc. Zool. durum Frcnzel. C. R. Acad. Sci. 156, 1932-4. Fr. 4!). 238-84. 125. 1913. Sur les premiers stades du dkveloppement 155. 1930. Observations sur Anophrys sarcophaga des gregarines du genre Porospora (= Nematopsis). C. R. Soc. Cohn. Bull. Bid. Fr. Belg. 64, 289-331. Bid. 75, 95-8. 156. Porchrt-Hrnnere, E. & Richard, A. 1970. Structure finc 126. 1915. Porospora nephrogis n. sp. Ibid is, 368-71. dcs microganietes d’Aggregatn eberthi Labbe. Protistologica 6, 127. __ . 1925. Les porosporides et lcur 6volution. Trau. 71-81. Stat. Zool. Wimereux 9, 126-39. 157. 1970. Structure finc du sporoblaste immature uni- 128. Levine, N. D. 1970. Taxonomy of the sporozoa. I. Parasit. nurlee d‘Aggregatu eberthi LabbC. (Sporozoaire Coccidioinorphc) , X(4/11/1), 208 [Abstr. 3781. C. R. Acad. Sci. 269, 1681-3. 129. Lunz, G. R. 1968. In “Hearing before the subcommittee 158. -- -. 1970. Ultrastructurc dvs stadrs vfgktatifs d’Ag- on fisheries and wildlife conservation of the Committee on gregata eberthi LabbC: le trophozoite ct IC schizonte. Z. Zell- Merchant Marine and Fisheries-House of Representatives.” forsch. 103, 179-9 1. Comrn. Fish. Res. Legislation H. R. 188808. Ser. No. 90-49, 99- 159. Prytherch, H. F. 1938. Life cycle of a sporozoan parasitc 104, U. S. Printing Office, Washington, D. C. of the oyster. Science 88, 451-2. 130. Minkiewicz, R. 1912. Ciliata chromatophora, nouvel ordrc 160. 1940. The life cycle and morphology of Nema- d’infusoires B morphologic et reproduction bizarres. C. R. Acad. topsis ostrearum, sp. nov., a gregarine parasite of the mud crah Sci. 155, 513-5. and oystcr. J. Morphol. 66, 39-64. 131. ___ . 1914. Etudes sur les infusoires syndesmogames, i 161. Rompel, J. 1894. Kentrochona nebaliae n. g., n. sp. ein gamontes et gamltes. Bull. Int. Acad. Cracocie (Acad. Polon. neues Infusor aus der Familic der Spirochoninen, zugleich cin Sci.) (B) 1913, 742-9. Beitrag zur Lehre von der Kernteilung und den1 Centrosoma. Z. 132. Miyashita, Y. 1933. Studies on a freshwater foettingeriid W’iJs. Zool. 59, 618-35. ciliate, Hyalospira caridinae n.g., n. sp. lap. J. ZooE. 4, 439-60. 162. Roscnficld, A,, Buchanan, L. 8: Chapman, G. B. 1969. 133. Mnhr, J. L. 1948. Trichochona lecythoides: a new genus Comparison of the fine structure of spores of three species of and species of marine chonotrichous ciliate from California, with Minchinia (Haplosporidia, Haplosporidiidae). 1. Parasit. 5.5, a consideration of the composition of the Order Chonotricha 921-41. Wallengren, 1895. Allan Hancock Found. Pub. Occas. Pap. 6, 163. Ryder, J. A. 1883. The protozoan parasites of the oyster. 21 PP. Science 1, 567-8. 134. Mouchet-Bennati, S. 1932. Sur la branchie d‘deglca 164. Sand, R. 1899. Etude monographique sur le groupe dcs laeuzs (Latreille) et son parasite Lagrnophrys aeglea nov. sp. infusoires tmtaculifrrs. Ann. SOL. Belg. Micr. 24, 57- 189. C. R. Soc. Biol. 109, 148-50. 165. Sauycr, T. K. 1969. Preliminary study on the epizootio- 135. Nigeli. 1857. Ubcr die neue Krankheit der Seidenraupe logy and host-parasite relationship of Paramoeba sp. in the blue und verwandte Organismen. Bot. Zeit. l5(44), 760. crab, Callinectes sapidus. Proc. Nat. Shellfish. Ass. 69, 60-4. 136. Neave, S. A,, ed. 1939-40. Nomenclator Zoologicus, Vols. 166. 1969. Blue crab studied by Maryland biologists. I - IV. Zool. SOC. London, Regent’s Park, London. Chesapeake Bay Affairs Comm Fish. News 2(6), 2. 137. Ncnninger, U. 1948. Die Peritrichen der Unigehung von 167. Sauyrr. T. K., Cox, R. & Higginbottom, M. 1970. Hemo- Erlangen mit brsonderer Beriicksichtigung ihrer Wirtsspezifitiit. cyte values in hralthy blue crabs, Callinectes sapidus, and crabs Zool. Jahrb. Syst. 77, 168-265. infected wtih thr amoeba, Pmon~oebnperniciosa. J. Inuert. Path. 138. Nie, D. & Ha, Y. 1943. Notes on some epizoic infusoria 15, 440-6. from the freshjvater shrimp, Palaemon nipponensis. Sinensia 14, 168. Schiiperclaus. W. 1954. Fischkrankheiten, 3 Auflagc. 143-9. .4kadc*mie-Verlag, Berlin. xii + 708 pp. 139. Ninni, A. P. 1864. Sulla mortaliti dei gamberi (Astacus 169. Schaudinn, F. 1896. Der Zeugungskreis von Paramoeba fluz’iatilis) ne1 Veneto c pi“ particolarmente nella prnvincia eilhardi 11. g.. n. sp. Sitzung~ler. Kgl. Preusr. Aknd. d. Wirc. trevigiana. Atti Imp. Roy Irrstituto Veneto 10 (3). Berlin, 25-35, (Reprinted in Srhaudinn’s Arbeiten, 191 1, 31-41.) 140. Noland, L. E. & Finley, H. E. 1931. Studies on the taxon- 170. Srhereschensky, H. 1926. Mikrosporidien als Erreger omy of the genus L’orticella. Trans. Am. Micr. SOL. 50, XI. riner Muskelerkrankung der Flusskrrbse. Zoo/. Anr. 65, 69-74. 141. Ormihs, R. 1968. Pachyporospora retorta n. sp. gregarine 171. Schnrider, .4. 1875. Contribution a I’historic des greg-arincs parasite de Macropipus holsatus F. crustac6-dbcapode. Ann. Para- des invcrtrhres de Paris et de Roscoff. Arch. Zool. Exp. Gin. 1, sit. Hum. Comp. 13, 413-9. 493-604. PROTOZOAOF DECAPODS 537

172. - -. 1892. Signalement d’un nouveau sporozoaire. 195. -854. Die Infusionsthiere auf ihre Entwicklungr- Tabl. Zool. 2, 209-10. ge.rchichte untz sucht. Englemann, Leipzig. 265 pp. 173. Setna, S. B. & Bhatia, B. L. 1934. On some gregarines from 196. 1859. Der Organismus der Infusionsthiere nach the prawn, Parapeneopsis sculptilis (Heller). Parasitology 26, eigenen Porschunger in Systematircher Reihenfolge bearbeitet. “.14.61 .”. I. Leipzig. 206 pp. 174. Smith, G. 1905. Note on a gregarine (Aggregata inachi, 197. __ . 1867. Der Organismur der Infusionsthiere nach n. sp.) which may cause the parasitic castration of its host eigenen Forschunger in systematisrher Reihenfolge bearbeitet. (Inachus dorsettensis). Mitt. Zool. Stat. Neapel. 17, 406-10. 11. Leipzig. 355 pp. 175. Sogandares-Bernal, F. 1962. Presumable microsporidiosis 198. Stunkard, H. N. 1957. The morphology and life-history in the dwarf crayfishes Cambarellus puer Hobbs and C. shufeldti of the digenetic trematode, Microphallus similis (Jagerskiold, (Faxon) in Louisiana. J. Parasit. 48, 493. 1900) Baer, 1943. Biol. Bull. 112, 254-66. 176. Sprague, V. 1949. Species of Nematopsis in Ostrea uir- ginica. Ibid 35(6/2), 42 [Abstr. 1081. 199. Sumari, 0. 8: Westman, K. 1970. The crayfish parasite 177. 1950. Notes on three microsporidian parasites of Thelohania contejeani Henneguy (Sporozoa, Microsporidia) found dccapod Crustacea of Louisiana coastal waters. Occ. Pap. Mar. in Finland. Ann. Zool. Fenn. 7, 193-4. Lab. La. St. Uniu. 5, 1-8. 200. Theodorid&, J. 1961. Sur la distinction entre les griga- 178. 1950. Thelohania cambari n. sp., a microsporidian rines des familles des Cephaloidophoridae et des Porosporidae parasite of North American crayfish. J. Parasit. 36, 46. parasites de crustaces decapod&. C. R. Acad. Sci. 252, 3640-2. 179. __ . 1954. Protozoa. In P. S. Galtsoff, ed. Gulf of 201. 1962. Gregarines d’inverthbres marins de la region Mexico. Its Origin, Waters and Marine Life, U. S. Fish Wildl. de Banyuls I. Eugrkgarines parasites de crustaces decapodes. Vie Serv. Fish. Bull. 55(89), 243-56. Milieu 13, 95-122. 180. Sprague, V. & Orr, P. F., Jr. 1955. Nematopsis ostrearum 202. - 1964. Gregarines d’invertebrks marins de la region and N. prytherchi (Eugregarinina: Porosporidae) with special de Banyuls 11. Deux nouvelles eugrkgarines parasites de Solenocera reference to the host-parasite relations. J. Parasit. 41, 89-1 04. membranacea (Risso, 1816) (Decapoda Penaiedae) et remarques 181. Sprague, V. 1963. Minchinia louisiana n. sp. (Haplo- sur les gregarines de Penaeidae. Vol. Jubil. Prof. G. Petit, Vie sporidia, Haplosporidiidae) , a parasite of Panopeus herbstii. J. Milieu 17 (Suppl.), 115-27. Protozoal. 10, 267-74. 203. ___ . 1965. Gregarines d’invertCbrCs marins de la region 182. __ . 1965. Nosema sp. (Microsporida, Nosematidae) de Banyuls 111. Porospora soyeri n. sp. (Eugregarina, Poro- in the musculature of the crab Callinectes sapidus. Ibid 12, 66-70. sporidae) parasite de Aristeus antennotus (Risso, 1816) (De- 183. __ . 1966. Two new species of Plistophora (Micro- ca:ioda, Penaeidae). Vie Milieu 16, 537-46. sporida, Nosematidae) in decapods, with particular reference to 204. ___ . 1965. Parasitisme de decapodes Natantia de one in the blue crab. Ibid 13, 196-9. Banyuls par Aggregata leandri Pixell-Goodrich, 1950 (Coccidia 184. Sprague, V. & Beckett, R. L. 1966. A disease of blue Aggregatidae). Ibid 16, 229-30. crabs (Callinectes sapidus) in Maryland and Virginia. J. Inuert. 205.---. 1967. Gregarines d’invertCbres marins de la region Path. 8, 287-9. de Banyuls IV. Quelques eugregarines du genre Cephaloidophora 185. ~ , 1958. The nature of the etioloeical- aeent- of parasites de crustaces amphipodes et decapodes. Ibid 18, 265-79. “Gray Crab” discasc. Ibid 11. 503. 206. TheodoridPs, J. & Laird, M. 1970. Quelques eugregarines 186. Sprague, V. & Vernick, S. H. 1968. Observations on the parasites d’invertbbres marins de St. Andrews (Nouveau-Bruns- spores of Pleistophora gigantea (Thdohan, 1895) Swellengrebel, wick). Can. J. Zool. 48, 1013-6. 1911, a microsporidan parasite of the fish Crenilabrus melops. 207. Trager, W. 1957. Excystation of apostome ciliates in re- 1. Pro~ozool.15, 662-5. lation to molting of their crustacran hosts. Bid. Bull. 112, 132-6. 187. Sprague, V., Vernick, S. H. & Lloyd, B. J., Jr. 1968. The 208. Trhgouboff, G. 1912. Sur les gregarines des balanes. Arch. fine structure of Nosema sp. Sprague, 1965 (Microsporida, Nose- Zool. Exp. Gin. N & R Ser. 5, 10, 53-61. matidae) with particular reference to stages in sporogony. J. 209. -- -. 1916. Sur quelques protistes parasites rencontres Inuert. Path. 12, 105-17. B Villefranche-sur-Mer. Ibid 55: 35-47. 188. Sprague, V. 1969. Need for drastic revision of subphylum 210. Tuzet, 0. & OrmiPres, R. 1961. Sur quelques gregarines .4moebogena. In A. A. Strelkov, K. M. Sukhanova & I. B. Raikov, parasites de CrustacCs Decapod&. Ann. Sci. Nat. Zool. Ser. 12, 3, cds. Proeress in Protozoolo~~~--SrdI, Intern. Coner. Protorool.. 773-83. Nauka, cening-rad. 372. 211. 1962. Gregarines de Roscoff. Cahiers Bid. Mar. 189. Sprague, V. & Vernick, S. H. 1969. Light and electron 289-306. microscolle observation on Nosema nelsoni Smaeue. 1950 (Micro- 3, sporida, Nosematidae) with particular referenceto’its Golgi com- 212. 1962. A propos du moyau protomkritique des plex. J. Protorool. 16, 264-71. grCgarines. Ibid 3, 209-1 1. 190. Sprague, V., Beckett, R. L. & Sawyer, T. K. 1969. A new 213. VivarPs, C. P. & Rubi6, M. 1969. Protozoa parasites de species of Paramoeba (Amoebida, Paramoebidae) parasitic in the Crustacea Decapoda Brachyura de la cBte nord-est de l’Espagne. crab Callinectes sapidus. J. Inuert. Path. 14, 167-74. Publ. Inst. Biol. Apl. 4, 111-29. 191. Sprague, V. 1970. Some protozoan parasites and hyper- 214. Vivark, C. P. 1970. Parasites de crustaces decapodes parasites in marine decapod Crustacea. In S. F. Snieszko, ed. brachyoures du Golfe et du Lac de Tunis. Bull. Inst. Ocdanog. A Symposium on Diseases of Fishes and Shellfishes, Spec. Puhl. PSche, SalammbB 1(4), 181-203. 5, Am. Fish. Soc., Washington, D. C. 416-30.. 215. Wallengrcn. H. 1895. Idem. 11. Slagtet Heliochona Plate 192. ___ , 1971. Plistophora Labbe, 1899, IS a junior synonym S13gtc.t Chilodochona n. g. SIiiStet Hemispeira Fabre-Domerque. of Pleistophora Gurley, 1893, and should be discarded. J. Inuert. Lunds Uniu. Arsskr. 31, 1-72. Path. 17, 1-2. 216. Watson, M. E. 1916. Three new gregarines from marine 193. Stanier, J. E., Woodhouse, M. A. & Griffin, R. L. 1968. Crustacea. J. Pnrosit. 2. 129-36. The fine strurture of spore of a Nosema spelotremae, a micro- 217. Woodhurn. K. D., Elder, B., Clark, E., Hutton, R. F. & sporidian parasite of a Sfielotrema metacercaria encysted in the Ingle, R. M. 1957. The live bait shrimp industry of the west crab Carcinus maenas. J. Invert. Path. 12, 73-82. coast of Florida (Cedar Key to Naples). Flu. St. Bd. Conserv. 194. Stein, F. 1852. Neue Beitrgge zur Kenntnis der Entwick- Tech. Ser. 21, 33 pp. lungs-geschirhte und des fcinerer Baues der Infusionsthiere. Z. 218. Wright. S. 1858. Description of new Protozoa. Edinburgh IVirs. Zool. 3, 475-509. New Philos. J. 7, 276-81.