protozoan parasites nematopsis

526

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

VICTOR SPRAGUE and JOHN COUCHi

Urr iu . of Mar)~lrinil, Nnfirrnl Rrsoio.rP.c Z i ~ . ~ f i / u f r , Clrr\npcrrke niologicnl / . r i b r i w i t o i y , So lo ino i rs . 121d. 2OhS‘S and ( I . S . D e p . of Critninerrf>, Nalroiinl Oceanic nnrl : l I ~ i t o s ~ h r r i c Af/iiiiiri.\li-r,lioiz, Nufionnl Mnritie Fi$/reric,.! S f w i c e Biologirn/ I .n l ior r r lo iy , O x f o i d , Mil. 21654

SYNOPSIS. An approximately complete list of the knolvn pro- suggrstrd. Host. site of infrction and seographic location are tozoan associates (exclusive r i f a few aberrant forms like Amal- locystir) of decapods is prcsented. A few taxonomic changes arc

nientionrd. Soinc comnirnts on pathosrnicity are made.

T H I S is, principally a.taxonomic paper, intended to Pre-

deal with a few of the problems in classification and nomenclature found a t the generic and spccific levels. At Even the same time, such highly relevant matters as host and site of infection, locality and pathogenicity are mentioned.

At the higher levels of classification we followed “A revised classification of the phylum Protozoa” of the Honig- berg Committee (78) as far as we could but \\’ere unable Suh l~ l iy lun i SARCOMASTOGOPHORA Honixherg & Bala- to follow it consistently because several specialists consider different parts of that system inadequate. In classifying the Amoebae lve ha\x folloived Rovee and Jahn (20) . Commenting on their difficulties, they stated very clearly the nature of some of the basic problems: “One reason these families are difficult to deal with is the consensus of taxonoInists of protozoa ( ~ ~ ~ i ~ h ~ ~ ~ et al,, 1964) whicll

evolutionally ancient and taxonomically diverse as are the

i n 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-

capod host. I\ felt, ciliates ars pathogenic in decapods but Protozoa known to be associated with and

are llarmless coInIllensals. we ran \,.ith confidel,ce that a para-

site is 01’ is not 1,articularly pathogenic to a dc*capod host, \ce kno\,. l i t t l e i n terms of the fluctuation of tklc host poprilations,

, \ k l a t

lllutll. 1963 ( 7 8 ) .

Suprrclass SARCODINA IIcrtnig R- Lrsser, 1874 ( 7 8 ) . Class HYDRAULE.4 Jahn & Bo\cr, 1965 (85 ) .

Order HYALOPODIA Bovee & Jahn. 1966 (20 ) . Family PAKAMOEBIDAE Poche, 1913 ( 153).

Genus Pnranroeba Schaudinn. 1896 (169) .

foroit ioeb(L perniciosa Si~rague, Bpckctt & S a v e r , 1969 (190) .

teaguc nay. \.irginia and hlaryland, CSA (165, 166, 167, 18-1,

Syn . : P n r n r n o e h sp. Spraque & Bvrkrtt. 19611 ( 185) . In hrmii- lymph of hluc crah Cnllinectes supidus Rathbun, 1896. Chinco-

185, 190) ; coast of North Carolina and Georgia, USA (129) . Dr. Eugrnr E. Rovce (pers. coin.) thinks P. eilhorrfi Scliaudinn. 1896, the tWe sliec+’s, belongs t O WnuS MaYoreilfl and (in pry=) has rcjccted both gums Pnro? i roeb~ Schaudinn and family Para- nloehidar P c ~ h e . I f B0vee.s opinion is jllstificd, >,Ilich \\e see, no reason to doubt, then P. periricio\o must he transferred to another genus. Sincr ur hold no oliinion as to the correct systematic position of this anruc1)a \\e ha\? , for couvenicnce, used the old classifiration

to recognize that protozoa are at least as

metazoa. The protozoa probably should be considered at least a subkingdom (or kingdom) of organisills, containing

phyla and subphyla (Raabe, 1964). This idea \vas at length and presented by and \vas

strongly emphasized by Kozloff ( 1960) in a syrrlposial address; and kingdoms were suggested-but not adopted- by Corliss (1962) ,” In the classification of the “sporozoa” we have followed Levine (128), who has given the most recent modification. Howcver, the comment just quoted from Bovee and Jahn is applicable also to this, because the “Sporozoa” probably include at least 3 distinct phyla. M’s have followed the Honigberg Committee on classification of the ciliates, since that part, strongly influenced by the Farriily CEPHAI,OIDOPHORlD.4E Kamm. 1922 ( 9 0 ) . opinions of Corliss, has not bcen so controversial.

Suliphylum .4PICOMPLEX:\ Idmine, 1970 (128) .

Class SPOROZOEA Lruckart. 1879 ( 1 2 8 ) . Su1)class GKEC;ARINlA Dufour, 1828 ( 7 8 ) .

Order EUC;RE(:ARINIDA Liger, 1900 (78 ) .

(;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 C n r c t n o e c ~ t e r b e r i n u d u n ~ i r Ball, 1951 ( l o ) . Ill mid- and hind- of those which actually occur. Probably all decapods qut of the grapaiid crab Pachygrapius tranrrei ius (Gibbs). Ber- harbor protozoan parasites: few sPccirs are knoLtI1 hecausc muda. T h e gencric nanle Corcinoecetes Ball, 1938 ( 7 j , is a

junior synonym of Kernntopris ( 2 1 0 ) . The present species does not srrni to Iia\e any of the vsscntial characters of Nenzntopcis relatively fsbv decapods have becn examined.

is poorly understood h i t a fe\v gcneral statsmcnts may be pro\iaionaily translcrrcd. made. The one known amoeba is highly pathogenic to its L‘ephaloidophora c t t rp i lode i Dall. 1963 (13 ) . In the gut of rrab host, pathogenic, thr xanthic1 crab Carpi loder rupntzr i . Cocoanut Island, Hawaii, altho they cause some local damage and occlude passages “*.

Cephn /o idop / iorn confornt i i (Diesing, 1951 ) Lhgrr & Duboscq, 191 1 (12‘2). Syns . : Greparinu conformis Diesing, 1851 ( 9 0 ) ;

* Contribution No. 458, Natural Rrsources Institritr of thr ~ ~ ~ ~ e ~ ~ ~ t ~ ron/ormis ( ~ i e s i n g , 18 j l j LabbC, 1899 (107) ; Fren-

C m r c i n o r c u f r i c~,n!orniic (Dirsing. 11151 ) Ball, 1938 ( 7 ) . In the

The pathogenirity of protozoan infections in decapods is like ~ ~ ~ / ~ ~ l ~ i d ~ ~ ~ ~ ~ ~ ~ , ,\hicl1 grnus it is here

(;regarincs do not Seem tO tIe

University of Maryland.

Gulf Breei:?. Fla. 32561. L present address: Erlvironnll.nta{ protrrtil,n .i\sc,ncy [,a],,,ratory, rahrL co‘ i /orr tL1’ (r)icsirls ““l Id‘ser ’ Duboscq3 1907 ( ‘‘1 ;

PROTOZOA OF DECAPODS 527

cera thr gut lunirn of thr grapsid crab P c c h y g r u p s u r marmoratus Fabricius. hlcditerranean roast of France; roast of Tunisia ( 2 1 4 ) : northrast roast of Spain ( 2 1 3 ) . Tuzrt and Ormi6res (210) de- rlared the name CnrcinoecPteT Ball, 1938, a junior synonym of Nematopcir Srhncider and rrturned this spwies to genus Cepha- loidophora.

Cephaloidophorn drnchi Theodorid&, 1962 (201 ) . Syn.: Cephnloidophora sp. T u x t & OrmiEres, 1961 ( 2 1 0 ) . In the gastric cera of the xanthid crab Xantho rir~zrlosur (Risso). Banyuls. France.

Cephaloidophora d ~ o m i a e (Frenzel. 1885) VimrEs & Rubih. 1969 1213). Syn.: Gregar ina dronziae Frrmzrl. 1885 (61 ) . In gut of the crab Droniiu personoto (= d g a r i s ) (61, 214). Gulf of Naples ( 6 1 ) , Blanes. Spain ( 2 1 3 ) ; and coast of Tunisia ( 2 1 4 ) . Altho ThPodorid6s (201 ) suggested that this gregarine probably should be placed in genus Cephaloidophora, thc new combination is here attributed to Vi\ar+s and Rubih ( 2 1 3 ) . n h o first used it.

Cephaloidophora duborcqi Poisson. 1924 (154) . I n thr gut of the alphcid shrimp Athanas nitescens Lcarh. Luc-sur-Mer (Calvados) ( 154) and Roscoff ( 2 11 ) , France.

Cephaloidophora etiii (Ball. 1959) n. ctimh. Syn.: Cnrcinoe- cetes etisi Ball. 1959 ( 1 2 ) . In the rrah Etirus laer,imanus Randall. Bombay, India. Carcinoecetes Ball, 1938, is a junior synonym of Nemntopsis (210) . The dcsrription of the present species fits that of Cephuloidophora rather than that of Neniatopsis.

Cephaloidophuru foresti Th6odoridi.s. 1967 (205) . In thr gut of thc crab Xantho pores,n (Olivi) . Banyuls and Villefranche, France.

Cephaloidophora f o s w (L(.gcr & Dubosrq. 1907) Trigouboff. 1912 (208) . Syns.: A g g r e g a t u roelomica LCgrr. 1901 [ partim] (110); Frenzelina f o r s w Liger & Duljoscq, 1907 (116) . In gut of the pea crab Pinnotheres pi$un! Penn. living in M p i l u s edulis L. France (English Channel) ( 1 10).

I n gastric ceca of the xanthid crabs Xuntho granulocurpus Forest (205) , Pilumnus hirtellus rpinifer Milne-Edhards (205, 213) and P. hirtellus L. (205) . The 1st 2 hosts at Banyuls and the 3rd a t Villefranche, France (205) ; thc 2nd also at Blanes, Spain (213) .

Cephaloidophora knoepf l e r i l 'hiodoridh, 1962 (201 ) . In gastric ceca of Pinnotheres pinnotheres ( L . ) , a pea crab. ronimensal in the rlani Pinna nobilis L. Banyuls, France (201) ; Blancs, Spain ( 2 1 3 ) .

Cephaloidophora mithraxi (Ball, 1951 ) n. comb. Syn.: Car- cinoecetes mithraxi Ball. 1951 (10 ) . In the crab itlithrax forceps (Milne-Edwards) . Bermuda. The generic name Carcinoecetes Ball, 1938 ( 7 ) is a junior synonym of A'ematopsis (210) . The present species does not rt~srrnl)le NematopJis but its known characters arc consistent ui th those of genus Cephaloidophora to which genus it is here transferred.

Cephaloidophora nephropis Tuzet 8: Orrnii.res, 1961 ( 2 10) . Syn. : Porospora nephropis Liger & Duboscq, 1915 [ partiin] (126) . In thr %ut of the Norway lobster, A'ephrops norz,egicuc (126, 127, 210, 2 1 2 ) . Roscoff, France (126, 127, 210, 2 1 2 ) .

Cephuloidophora nigrofusca (Watson, 1916) Kamm, 1922 (90 ) . Syn.: Frenzelina nzerofusca Watson. 1916 ( 2 1 6 ) . In rut of thc

Cephaloidophora guinotae Thiodoridts, 1967 (205) .

, I

fiddler crabs IJca pu>nux and U . pugzlator. Cold Sprini Harbor, New York, US.4.

Cephaloidophoru oceliuta (Liger 8: Duboscq, 1907 ) Kamni. 1922 (90 ) . Syns.: Aggregata i 'ngans LCger & Duboscq, 1903 [partim] (112 , 116) ; Frenzelinu ocellata Liger 8: Duboscq. 1907 (116) . In gut lumen of the hcrmit crab Eupagurus prideauxi Leach. Banyuls, France ( 112, 201 ) ,

Cephaloidophora o h i a (Watson, 1916) Kamni. 1922 (90 ) . Syn.: Frenzelina o/iuia Watson, 1916 (216) . I n gut of 2 littoral crabs Libinia dubia Milne-Edwards ( 2 1 6 ) and Ocj 'pode cursor L. (46 ) . Cold Spring Harbor, Nc\z- York, US.4, (216) and Kalade, Lebanon (46 ) .

Cephaloidophora pinguis Ball, 1963 (13) . I n gut of the xanthid crab Carpi lodes rugntu.<. Cocoanut Island, Hawaii, USA.

Cephaloidophora poissoni 'lh(.odoridPs, 1967 (205). In the

alphrid shrimp Alpheus ruber Milnr-Edmards and A . dentipes Guerin. Banyuls. France.

Cephaloidophoro ( ? ) praemorra (Diesing, 1851) Kamm, 1922 (90); Syns. : Gregarina braemorsa Diesing,-1851 (90) ; Aggregata praemorsa (Diesins, 1851) Labb6, 1899 (107) : Frenzelina grae- morsa (Diesing-, 1851) LiSer & Duboscq, 1907 (116). In gut and vesiclrs adhrring to the ovary of the crab Cancer pagurus I,. As Kamm (90) pointed out. this nominal species is based on a niere refrrence by Redi, in 3684, to somethins which might have been a p,garinc.

Cephaloidophora sp. Syn. : Carcinoecetes sp. Ball, 1948 (8). hlainly in the digrstive gland hut also in the gut Innirn of the sand crab Emerita analoga. Southern California, USA. Since genus Carcinoecetes Ball, 1938 ( 7 ) is a junior synonym of A'eniatopii$ Srhncider. Ifi92 ( ? l o ) , this species needs to be reassigned. M a t knonn gresarines of crabs belong either to gcnus C~phnloidophora or n'eniatopsis. The known characters of this species seems to he more like those of the former genus.

Genus Cephalolobus Kruse, 1959 (99 ) .

Cephnlolvbur penneus Krusr. 1959 (99 ) . Attached to filtrr of stomach of shrimp Penaeu , ar tecus Ivcs and P . duorarum Burken- road. Florida, USA. Type species. Kruse considered the systc- niatic position of the senus in dctail. He did not place i t in a family hut concluded. "If the life cycle of C. penaeur is proven to hc siniilar to that of Ani~olobur, both genera should be placed in a ric'w fainily." 'l'liiodoridi~s ( 2 0 2 ) . for reasons not very con- vincing assigned the genus to family Cephaloidophoridac, where i t is here listed for convenii.ncr.

Cephalolobur petiti Theodorid&, 1964 ( 2 0 2 ) . Active stages in stomach and cysts in hind-gut of thc penacid shrimp Solenocera nienibranacea (Risso) . Banyuls, France.

Genus L'ardiohabitans Ball, 1959 ( 1 2 )

Curdzohnbrtans ietnai Ball, 1959 ( 1 2 ) . In the mid-gut of the shrimp Penaeur Tenikulcatus de Haan. Bombay, India. Type and only species. Ball did not assign the genus to a family but he compared it both uith Cephaloidophora and genera in the Porosporidae ; i t is here provisionally listed with the Cephaloido- phoridae. 'lhc validity of the genus is questionable.

Family URADIOPHORIDAE GrassC, 1953 (64) Genus C'radiofora Mrrcicr, 1912 (90) .

In the gut of the alpheid shrimp Athanor nitescens Leach. Lnc-sur-Mer (Cal- vados), France.

Uradiophora cuenoti (Mercier, 1911) Merrirr, 1912 (90 ) . Syn.: Cepholoidophora cuenoti Mercier, 1911 (90 ) . In the gut of the shrimp Atyoephya desmaresti Millet. Nancy, France.

C'rcdiophorn rithnnasi Poisson, 1924 (154) .

Family POROSPORIDAE Lahbi, 1899 (107) . G n u s Porospora Schneider, 1875 ( 1 71).

Porospora gigantea (Bencden. 1869) Schneider, 1875 (171 ) . Syn.: Gregarina gigantea Bcnedrn, 1869 (17 ) . In the lobster Homarus gammarus ( L . ) and the gastropod Trochocochlea mutabilis Philippi. Roscoff (171 ) and Banyuls (201 ), France. This is the type and the only spccies for which the essential gcneric character, naked sporozoites in a molluscan host, has been demonstrated. This demonstration was by a laboratory infection experiment performed by Hatt ( 7 2 ) . The same author (71) obtained partial development also in M3,tilus minmus Poli.

Porospora nephropis L6ger 8: Duboscq, 1915, emend. Tuzet & OrmiPrcs, 1961 ( 2 10). Syn. : Porospora nephropis Liger & Duboscq, I915 [partim] (126) . In gut of the lobster Nephrops norcegicus L. Molluscan host unknovn. Roscoff, France.

Porospora piJae LCger & Duboscq, 1911 (123) . In the spider crabs Pisa grb~ii (123) and P. armata (213) . Cette (123) and Villefrariche-sur-Mer (209) . France; northeast coast of Spain

528 PROTOZOA OF DECAPODS

(213) . Trhgouboff (209) observed peculiarities in this species which convinced L6ger and Duboscq (127) that it is not a Porospora. These authors did not attempt a reclassification for lack of information on final development of the cysts. Vivarks and Rubi6 concluded that it is neither a Porosfiora nor a Nematopsis (213) . I t is kept here in Porosporn provisionally because there is still no clue as to its proper generic classification.

Porospora sp. Thhodoridks & Laird, 1970 (206). In gut of the lobster Homarus americanus. St. Andrcws, New Brunswick, Canada.

Genus Nematopsis Schneider, 1892 ( 172). Nematopsis cnlappae (Ball, 1951) n. comb. Syn.: Carcino-

ecetes calappae Ball, 1951 (10 ) . In gut of the oxystomatous crab Cnlappa flammne (Herbst) , Intermediate host unknown. Ber- muda. Carcinoecetes Ball, 1938, is a junior synonym of Nema- topsis (210). Excepting rare intrarcllular stages, which could belong to another species, the known characters of this species are consistent with those of Nematopsis, to which genus this species is here provisionally assigned. I t is noteworthy that Ball’s figures show a ~iorosporid character which the author did not mention, the “muscular collar” of the protomerite.

Nematopsis doripfie Bogolepova, 1953 (18 ) . In gut of the oxystomatous crab Dorippe granulnta de Haan. Sea of Japan.

Nematopsis duorari Kruse, 1966 (100, 101). In gut of the shrimp Penneus duorarum Burkenroad. Spores in the pelecypods Aequipecten irradians. Cardita Poridana, Chione cancellata, Macrocallista nimbora. Florida, USA.

h’ematopsis goneplaxi Tuzet & OrmiPres, 1961 (210). Syn.: PorosPora petiti ThCodoridPs, 1962 (201). In gut of the crabs Coneplax angulatu (Penn.) (210, 201) and G. rhomboides (213) . Intermediate host unknown. Cette (210) and Banyuls (201) , France; northeast coast of Spain (213) .

Nemntogsis grarsei (ThCodorides, 1962) VivarPs & Rubib, 1969 (213). Syn.: Poros~ora grarsei Th6odoridPs, 1962 (201) . In gut of the oxystomatous crab Calappa granulata L. (201) . In- termediate host unknown. Banyuls, France (201) ; northeast coast of Spain (213).

Nematopsir herpertis (Ball, 1938) Tuzet & OrmiPres, 1961 (210) . Syns.: Carcinoeceter hesperus Ball, 1938 hw1: SPECIES]

( 7 ) ; Por~spora hesperus (Ball, 1938) ThCodoridb, 1961 (200) ; [ ? ] Nematopsis rp. Ball, 1951 (11 ) . In gut of the grapsid crab Pachygrap.rur crasripes Randall. Intermediate host unknown. California, USA.

Nemntopsis legeri (de Bcauchamp, 1910) Hatt, 1931 (73) . Syns. : Nematopsis mediterranen Lhger. 1905 I nomen oblitum] (113) ; Porosporn leg& de Beauchamp, 1910 (16 ) ; P. galloprovincialis LCger & Duboscq, 1925 (127, 73). In gut of the crabs Eriphia .rpinifrons (16) and E . ~ ‘ e r i u c o ~ n (214) ; spores in the mussel Myfi lus galloprovincialis and possibly other molluscs (73 ) . Southern coast of France (16) and coast of Tunisia (214) . This is one of the best known species of A’ematopsir, due largely to the work of Hatt (70, 71, 73) , who studied its entire life cycle by means of experimental infections. Previously only parts of the life cycle of a porosporid gregarine had been made known experimentally (124, 179). Tuzet and Ormikres (210) recently made a careful study of this species to confirm the generic characters.

Nematopsis marnis i (Lhger & Duhoscq, 191 1 ) Sprague, 1954 (179). Syn.: Porosporn marnisi LPger & Duboscq, 1911 (123). In gut of tlir rrabs Macropipus (= Portunus) depurator L. (123) , M . m i n d i s , Portumnus lotipes (214) and Macropipus depurator (213) . Intermediate host unknown. Cette, France (123) ; coast of Tunisia (214) ; northeast coast of Spain (213).

Nematoprir mututne (Ball, 1959) Tuzet & OrmiPres, 1961 (210) . Syn.: Cnrcinoecetes rnatutae Ball, 1959 ( 1 2 ) . I n gut of the crab Matuta lunaris (Forskil). Intermediate host unknoxn. Bombay, India.

Nematopsis miroulei (ThCodorid&s, 1964) n. comb. Syn.: Poros~ora miroulei Thdodoridcs, 1964 (202) . In gut of the

penaeid shrimp Solenocera membranncea (Risso) . Intermediate host unknown. Banyuls, France. This species is transferred to genus Nematopri.r because its vegetative stages are similar to those of confirmed species of this genus in crabs and shrimp but strikingly different from those of the single confirmed species of Porospora, known only in the lobster.

Mematopsis ostrenrum Prytherch, 1938, rmend. Sprague, 1949 (176). Syns.: Nematopric octrearum Prythrrrh, 1938 [partim] (159) , N . ostrearum Prythrrrh, 1940 [partim] (160), [non] N . legeri (de Beauchamp, 1910): Kudo (103, 104). In gut of the xanthid crabs Panopeur herbstii Milne-Edwards (159, 160), Eurypanopeus depressus (Smith) (159, 160), Neopanope texana ragi (Smith) (97 ) , Eurltium limosum (Say) (176, 180) . Spores in the oyster Crarrostrea oirginica (Gmelin). Gulf and Atlantic coasts of North America, as far north as New York (109) .

A’emotopsis panopei Ball, 1951 (10 ) . In the gut of the xanthid rrabs Panopeus occidentalis Saussure and P. h e r b ~ t i i Milne-Ed- wards. Intermediate host unknown. Bcrmuda.

Nematoprir garnpeneops i~ i (Srtna & Bhatia, 1934) n. comb. Syns. : Hirmocystis ( ? ) parnpeneopsisi Setna & Bhatia, 1934 (173) : Protomagalhensin ( 7 ) ottenunta Sctna & Bhatia, 1934 (173) . In gut of the shrimp Parapeneopsis sculptilis (Heller). Possible intermediate host unknown. Bombay, India. Grass6 (64) concluded that these 2 species described by Setna and Bhatia were crroneously assigned as to gcnera. Altho mature cysts were not seen, the known characters (especially disappearance of septa in the satellites) of both species secm to he consistent with those of the Porosporidae. Furthermore, the difference between the 2 (slight difference in shape) seems to he insignificant. Thus, they are treated here as the same spccies. The specific name of that une appearing first in the paper is rrtained. Assignment is here made to gcnus Nematopsis because the known stages are much more like corresponding stages of this genus than those of Porospora and because the former (100) hut not the latter is known to occur in shrimp. Thdodorides (202) suggested that these gregarines may belong to genus Uradiophora or maybe Cephaloidophora, hut this suggestion is untenable bccause the septa of satellites do not disappear in these genera.

Nematopsis penaeur Sprague, 1954 (179) . Syn.: Porospora penaeus (Sprague, 1954) ?’h&odoridPs, 1965 (203) . In the gut nf the shrimp Penneur ortecur Ives. Intermrdiatr host unknobn. Barataria Bay, Louisiana, USA. Sprague ( 179) evidently mistook something else for intracellular stages of this parasite, for presence of these has not been confirmed. Kruse (99) reported this species in P. aztecus, P. duorarum and P. setiferus. Hutton et al. (79) found it in these and also Sicyonia. Later, Hutton (80) reported it in P. aztecus, P. setiferus (I , .) , P. duoraruni Burkenroad, P. brasiliensis Latreil, Sicyonia typica Boeck, Trachypenaeus similis (Smith) , T . constricta ( Stinison), Xiphopeneur kregeri (Heller) and Solenocera atlantidic Burkenroad. Considering that gregarines in these hosts have not been studied in detail, that Kruse found 2 species in 1 host, and that ecologic factors related to the different shrimp are diverse, it is probable that several species of Nematopsis in these shrimp have been mistaken for one. The work of Kruse (100, 101) supports Sprague’s original generic determination.

A‘ematopsir fiortunidarum (Frenzel, 1885) Hatt, 1931 (73 ) . Syns. : Aggregata portunidnrum Frenzel, 1885 [ partim] (61, 116, 127); Nemntopsis schneideri Lkger, 1903 (111, 127, 7 3 ) ; Frenielina portunidarum (Frenzel, 1885) Ldger & Duboscq, 1907 (1 16) ; Porosjora portunidarum (Frenzel, 1885) Lhger & Du- boscq, 1911 (122) . In gut of crabs Corcinus maenas (61, 122) and C. mediterraneus (214) . Originally reported also in Macro- pipus (= Portunus) arcuatus (61) hut LCger and Duboscq (122) reserved the specific name portunidarum for the gregarine in Cnrcinus. Dcvelops experimentally in Macropipus ( 124, 125, 127). Spores in the niolluscs Cardium edule, Scrobularia plana and perhaps others (127). Naples, Italy ( 6 1 ) ; Etang PCrols, near Montpellier, France (122); coast of Tunisia (214) . ThBo- dorid& (201, 203), apparently overlooking experimental proof

PROTOZOAOF DECAPODS 529

by Liger 8: Duhoscq (124, 125) that this is a Nematopsis, has continued to call it Porospora. N . schneideri, represented by spores in Solen oagina, is nominal type species by monotypy hut, since this is regarded as a junior synonym of N . portunidarum (127, 73) , the latter is now generally accepted as the type.

Nematopsis prytherchi Sprague, 1949 ( 176). Syns.: Nema- topsir ostrearum Prytherch, 1938 [partinil (159) ; N . ortrearum Prytherch, 1940 [partim] (160) . In the stone crab Menippe mercenaria (Say) , Spores in the oyster Crarsostrea virginica (Gmclin). Louisiana. Florida and the south .4tlantic states of USA.

Nematopis raouadi Vivares, 1970 (214). In gut of the crab Porturnnus latipes. Coast of Tunisia. I t is curious that Vivares, when comparing N . raouadi \vith A‘. marnisi in the samr host. 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 ) , while finding the latter to be 7-8 p .

Nematopsis soyeri (Theodorid&, 1965) n. comb. Syn.: Poro- spora soyeri ThCodoridb, 1965 (203) . In gut of the penaeid shrimp Ari$teus nntennntul (Risso) . Intermediate host unknown. Banyuls, France. Drsportrs and Th6odoridi.s (45 ) studied the ultrastructure of the gymnospores. This sprcirs is transferred to genus A’eniutopris hrcause its vcgetative stages are similar to those of confirmed species of this genus in crabs and shrimp hut strikingly different from those of the single confirmed species of Porospora known only in the lobster.

Nematopsis sp. Krusc, 1966 (100) . In the gut of the shrimp, Penaeus duorarum Burkcnroad. Intermediate host unknown. This is the “small gymnospored specirs of Neniatopsis” which was found in shrimp with N . duorarum (the “large gymnospored species”) but which \\as not produced by spores from the pclccypods. Florida, USA.

Nematopsis sp. Sprasue, 1954 (179) . In 1932. Pearse (142) rcportcd the presence of “~rrgar ine cysts” on the walls of the rertuni of the ralico rrah Eriphici Conagrn (Fahririus) collectcd at Dry Tortugas off the coast of Florida, USA. Sprague (179) cxprcsscd the opinion that this is a h’eniutopsis, not only because the limitcd description is charactcristic of the porosporid grega- rims hut also bccause thc hest known spccics A’. legeri, also occurs in a species of Eriphia, E. rpinifrons.

Neniatopsis sp. VivarPs 8: Ruhi6, 1969 (213) . In gut of the crab E t h u w musrnrone. Northeast coast of Spain (Blanes).

Nematopsis sp. Vivarh C Rubi6. 1969 ( 2 1 3 ) . In gut of the crab Parthrnope nngulifron~. Northrast roast ( i f Spain (Blanes)

[A’enratopsis sp. ? ] “Gregarines” Cuckler & Fichter, 1939 ( 4 2 ) . In the gut of thc crab Cancer productus . Argyle Bay, San Juan Island, Washington, USA. Since this grezarine \\as found in a crab host and since as many as 3 nuclei \\ere seen in 1 com- partment (suggrsting fusion of satcllitrs), there is reason to suspect that i t is a species of Sematopsis .

Gcnus Pachyporospora TheodoridPs, 1961 ( Z O O )

Pachyporospora lamellaris (Bogolcpova, 1953 ) OrmiGrcs, 1968 (141) . Syns.: A’enmtopsis lamdlar i s Bogolepo\-a. 1953 (18) ; Tricystis @nta Poljanskij & Chejsin, 1965 (52 ) . In gut of a crab. Sea of Japan. Poljanskij and Chcjsiri (52 ) , in their revision ( 1965) of Dogicl’s ( I95 1 ) book on protozoa, reproduced Bogolepova’s (1953) sketch of this species and said it was from Dogie1 (1951). They did not say where they got the name 7’ricjds plicatu and n e ha\e been unable to find it elsewhere in the literature.

Pachyporospora laubieri ‘Ihdodoridi.s, 1961 ( 2 0 0 ) . In the gut of thr rrahs Atelecj’clus Jcptemdentntur hlontagu ( 2 0 0 ) and A . rotundatus (213) . Banyuls, Irance (200) ; northeast coast of Spain (213) . TYPE SPECIES.

Pachyporosporn retorta Orniiihcs, 1968 (141). In gut of the crabs Macropipus holcatus (F . ) (141, 214) and M . i,ernalis ( 2 1 3 ) . Shte, France ( 1 4 1 ) ; coast of l’unisia ( 2 1 4 ) ; northeast coast of Spain (213).

Family and qcnus uncertain

Gregnfina pmtrini Frcnzel. 1885 ( 6 1 ) . In gilt of the crab Macropipus (= P o , f l m u r ) nrcuatus. Naplrs. Italy. hfonocystid gregarine. certainly not stmiis Gregarina and unlike other known grcgarincs in decapods.

Gregarines, identity not spccified. Heckmann ( 74) found intestinal grrgarines in a ni~inhrr of decapods in Hawaii. Heldt (75) described. xvithout naminp. grrgarines from the shrimp Penoeus kerathurus (Forsk:il) in thy Gnlf of Tunis. ThCodoridPs ( 2 0 2 ) judgcd that thcsc last onrs helonged to 2 species, a Uradiophora and a Porospora.

Poljanskij and Chcjsin (52 ) nicritioncd “ccrtain species of Merogregarina from the alinientary tract of crahs” but we have been unable to find mention of them elsewhere.

Suhclass COCCIDIA Lcuckart, 1879 (78 ) . Order EUCOCCIDA 1,Eger & Ihboscq, 1910 (78) . Family SELENOCOCCIDIID.4E Pochr, 1913 (153) . Genus Selenococcidium LCger 8: Duhoscq, 1909 (120) .

Selenococcidium infermedium Legcr & Duboscq, 1909 ( 120). France. This is the type

The authors de- In gut of European lobster Homarus. of the gcnus and the only species in the family. scribed it only hricfly in 1909 and niurc fully in 1910 (121) .

Family AGGREGATIDAE Labhe, 1899 (107) . Griius Aggregnta Frrnzrl, 1885 ( f i l ) , emend. I.6Srr C Dul~oscq,

1906 (114).

AggrPjinta coelornica LkSer, 1901. rmend. Leger and Duboscq, 1906 ( 1 14) . Syn.: Aggregata coeloniica L6xrr. 1901 [partinil (110, 114). In gut i v a l l of the crab Pinnotheres pi;um Pcnn., \I hich lives in Mytilus edu1i.s L. Th(, original “sprcirs” rventually hecarne in part the grrgarinr, Cephaloidophora foisor. France (Errglish Channel) f 1 I O ) . Found recently hy Vixar6s and Rubi6 ( 2 1 3 ) , in the same host. on the northcast coast of Spain.

Azgregata eberfhi (I,ahht, 189.5) L6grr & Duhoscq, 1906 ( 115 ) . Syns.: Aggregata portunidnrum Fren~el . 1885 I partirnl ( 6 1 ) ; Benedenia eberthi LabhC, 1895 (105) [cited by LahbC, 1899 ( lo ; ) , altho wc can find no tnention of this name in the 1895 paperl. Schizogonic stages in the portunid crabs Macro- pipus (= Portunus) arcuatur (61 ) . M . depurntor ( 1 1 5 , 213): M . tuberculata (213) , M . 6oliz.uri (213) . ill. corrogatus (214) and probably, M . uernalis ( 2 1 4 ) . Sporogony in thr cuttlefish Sepia officinulir. Common ar.d widespread. Naples, Italy (61 ) ; Site. Francr ( 1 15) ; l‘unisia ( 2 1 1 ) ; northeast coast of Spain (213) . Since the synonymy is quite complicated, only the most significant synonyms are gken here. [See Lahbb, 1899 (107) and I,(.<er and Duboscq, 1908 (117) for more details.] Why the junior synonym eberthi \*as conserved instead of the oripinal name portunidarum \\e havr been unable to deterrriinr, altho the former is now fixed by the provisions of Art. 2 3 ( b ) of the Inter- national Code af Zoological Nomenclature (82 ) . Wr believe that this species, as Calkins ( 2 2 ) said, is the type; altho PellCrdy (143) said A . octopiana (Srhneider, 1875) is the type. Our reasoning is that the fornicr, bring the only original species in genus A g g r e g a t a , inust he type by monotypy as provided by .4rt. 68(c) (82) . I t may he pertinrwl. lro\\ever. to note that the name A. portunidarum \cas originally applied inad\,ertcntly to 2 species, 1 a gregarine I, hich later hecanie A’eriratopsii portuni- darunr. This species has heen the object of important life cycle studies by Ltger and Duhoscq ( I 14, 115, 117), Pixell-Goodrich (150) and Dobell (491, and electron microscope studies by Porchct-Hennerd and Rirhard (156, 157, 158).

Aggregata inachi Smith, 1905 (I ;+) , ernrnd. Ltger & Duboscq, 1906 (114). Schizogony stases in the gut wall of the crabs Inachus dorsettensis (174, 214) and I . communissimus (213) ; indistinpuishahle forms in I . scorpio (17-1). Stages in ~nolluscan host unknohn . Naplrs, Italy ( 1 7 4 ) ; coast of Tunisia (21-1) ; northeast coast of Spain ( 2 1 3 ) . Smith, likr earlier workers, presumed that


the cysts in thr. silt \vall hrlonged to a fireprint. xvith frcr stages in tlie linnen. altho lic sa\\- only cysts of this spc~cirs. L6gi-r and Duhoscq i 1 14, I 17) pointed out the incorrt-ctness of this viv\v.

A,cgregiitn lrnndri Pis1.1l-Goodrich. 1950 ( 152). Schizogony stages in gut wall of the shrimp I.eonder rquilln I,. Molluscan host unknov n. Naplrs. Italy. Found also in thv shrimp Solenocera inembranncea (Risso) and Acnnthephyra eximin Smith by Thfo- dorid& (204) at Banyrils. Francr.

Aggregatn xzgnnr 1,Pger S: Duboscq, 1903. emend. LGSer & Duboscq, I906 ( 1 1 I ) . Syti : A g y r e g n t n i 'ngnnr I&rr & Duhosrq. 1903 [partinil ( 1 1 2 ) . In gut wall of the rrah Eupocurur pridenuxi Leach. Ranyuls. Francr. Thr original species eventually hrcame in part the qrsarinc. Ceplrnioidopliora oc r l l a ta .

Aggrega tn sp. I.(.gcr & Dnhoscq. 1908 (117) . In codom of the crab Pachygiopriis mnrinoratuc F. Franrc ( 1 1 7 ) . Found ri~r.i~ntly in the saiiic host by \'ivar+s and Rubii, o n the northeast mast of Spain ( 2 13).

Aggrega ta sp. Srtna Rhatia, 1934 ( 1 73). Schisojiony stages i n gut \\all of thv shrimp P n r ~ i p e i i ~ ~ o f ~ i i v sculptilir (IIrl ler) . Mol- luscan host nnkno\vn. Bornhay, India.

A g g r e g n t a s p . Vi1ari.s Kr Rubih. 1969 (213) . Unidentificd species of <.iggregntn \vert% found in the crahs Coiystes cnrri- i ,e launui . rl te leci 'r lur i o t u n d u r . Xni i tho pore,rn. (;oncplrix rhoi i i - boides, Parothenope angiilifrons. and Macropodin rostrota. North- rast coast of Spain.

Aggregatu \ \ere found in tlir craln CoiciiiziT niedi trrrnnrus. Dromia perronntn. Poituiiinuc lntipes. X n i m biguttntn. Mncro- pipus z,ernali,. P i l u ~ ~ ~ n i i c rpinijer and Piiuninzir h ir te l l i i r . Coast of Tunisia.

Aggregntn sp. \'ivar+s. 1970 (214) . C..nidentificd spi

Subphylnni II1CROSPOK.Z Spra:nr, 1969 ( 188)

Class I\IICROSPORE.\ Corliss bi 1.c~ inc, 196:i (38 ) . Order hIICROSPORIDA Balliiani, 1882 ( 6 ) Family NOSEAIATIDrlE Lablit?. 1899 ( 1 0 7 )

Genus A'osenin Niigeli. 18.57 (133). rnicnd. P6ret. 1905 ( I 16).

Norema fren-elinne LCgw S: Lhlioarq. 1909 ( 1 18. 119) . Hyper- parasite of the gresarinr Cephaloidophorn confoi m i r (Dirsin:) in the crab I'nchj,grnp,zir riiormorntus (Fabricius). France.

A'orema michnelir Spragiie. 1970 (191 ) . Syns.: S o i r m n supidi DeTurk. 19-10 ipartiml (47) : 1Y. si'. Spragne. 1965 (182) . In muscles of the rrah C n l l i n e c t e s snpidur Rathbun. .4tlantic and Gulf statrs of Xortti America. .A significant factor in crab mor- tality. DeTurk mistook spnri~s of this specks for sporoblasts of A'. rapidi. Spragne et al. 1 1 8 7 ) niade an clectron niicrosropr study of A'. michneli..

Xorenia nelioni Sliraqne. 1950 ( 1 7 7 ) . Syn. : Iriotll .Yorcriio puli . is: Jones ( 8 6 ) . I n riiiisclrs of prnat,id slirinip. M'idcly dis- tributcd. First rr1)orted i n Pei ioeur nztc r ic , I\ cs i n Louisiana. USA. Later found hy IVoodburn r t al. (217) , Hutton (80) and Hutton et al. ( 79 ) also in P . d u o r a i i i n i BurLenroad in Ilorida. US.4. Recently found by Sprague and Vcrnick (189) . \ \ho studird its fine atrurtnrr. i l l P. .setifeinr from Gcorgia. US.\. 1%. F. P. Champion (prrs. ~ o n i . ) f o r d a similar or identical sprcics i n t he iiiiisrIi.s o f / \ le tnpei i i ieus m o ~ ~ o c e r c o s i n thc Rrpublir of South hfrica. Kruse ( prrs. corn.) reported h'osemn sp.. which may he idrntiral \ \ i th .Y, nrlmiii. in the ahrinip Solenocera zioscni and Siphopciieu, krojrii. i n Louisiana. USA

A'osema pii!i,ic P6rzz. 1905 ( 14.5). I n rnuscles r i l t l i r crali Carcinur moen(i\. Fr;inc i'. First dtwrilied \ c r y hriefly. t h i s parasite \\as dcwribeti niorr fully in thr same year ( 1.46).

X o s ~ m a rnpid i Dc'Iiirk, 1950. cnicnd. Sprajinr. 1970 (191) Syn . : ,Vorema sirpidi l k l i i r k . 1910 lpartiinl ( 4 7 ) . In muscles of thr crab Cnlliriertei iapidur Ratliljuii. Rraufort. North (;sic>-

lina, YSA. This is diatiugiiishrd from the otlicr :+ sprcies of ,Vo.sema descrihed iii muscles of decapods by having rrlatively large spores \shich ( \ \h r i i staiiird) h a \ ? light arras at t h r 2 rnds and a dark I,and i i i t h v iiiiddlc. llcl'urk includrd in his

description also A'. michnelis Spragne. 1970 (191). the sporrs of nhich he mistook for sporohlasts of A'. ctipidi.

.Voiema sprlotremoe Guyinot 6: Poncc. 1925 (66 ) . Hyprr- parasitic in encysted metacercariae of the digrnetic trematode Microphnllus (= Spelotreina) cnrciui (198) , which is parasitic in the crah Cnrcinus inaennr. Stanier et al. rrcrntly (193) did an rlcctron microscope study of this species.

A'oserna sp. I n 1963, one of us (V.S.) found a species similar to A'. spelotremne in encysted niicrophallid metacercariae in a crab Cnl l i i i rc ter rapidus collected i n Pry Cove on the rastern shore of Chesapeakc Bay, Maryland, US:\. hl. W . Newnian ( ~irrs. cnm.) found a Xosenra, possihly thr samr spcries also, in North (:arolina. South Carolina and Georgia, USA, in 1969.

Genus Gurleya Doflcin, 1898 (51 ) . G z i r l e y z mij,nirii (Kudo, 1924) Spragne, 1970 (191). Syns.:

7'helohnnin sp. Miyairi, 1909 ( 102) : Cocconrinn inij ,uiri i Kudo. 192-4 (102). In thr niuscles of 2 sprcies of Atj'ephira. Japan.

Genus 7'helohniiin Henneguy, 1892 ( 7 7 ) . Thelohania cnnibari Sprasue, 1950 (178) . In muscles of the

crayfish Cambarus bartoni. Georgia, US.4. Thclohunia contejeani Hrnneguy. 1892 ( 7 7 ) . In muscles of

tlie crayfish A ~ t a c u s fluiiatilis ( 7 7 ) ; A . paliper (53, 168) and .4. astncus (53, 199) . France ( 7 7 ) ; Germany ( 1 6 8 ) ; Russia ( 1 7 0 ) and Finland (199) .

Thelohania rfuorarn Iverson & Manning, 1959 (83 ) . Syns.: Thelohonia hun terae Jones, 1958 [norncn nudu~n l (86) : Thelo- hnnin sp. Kruse, 1959 (99) . I n muscles of the shrimp Penaeus duorarum (83, 80, 84 ) , P. brnsiliensis (84 ) and P. nrtecur (80) . Florida, USA.

7'helohnnia giardi Hennzguy. 1892 ( 7 7 ) . In niusclrs of the shrirrip Crnngon i'ulgnris. Francr. TYPE S I T C I I . ~ h y snliscqiic~nt dtGgnation, Gurley, 1893 (Gj).

Thelohanin macrocyrtir Curley. 1893 (65 ) . In muscles of the shrimp Pnlarnioneter I nrieiis. Italy. Kudo (IO?) qncstionrd the \.alidity of this species.

7'hrlohania innenndis Perez, 1904 ( 144). In inusclrs (usually) of the rrah Cnrrinus maenas. Ptrez ( 1 4 7 ) reported 1 caw in \\ hirh an ovary was parasitized. resulting in parasitic castration. 'I'his parasite. first rqiortrd liriefly. \ \as drscrihrd inore fu l ly in 19U5 (1-16). France (Arcachon).

7'helohnnia octospora Hennrguy, 1892 ( 7 7 ) . In ninsrlrs of tlir shiinip I'niocmon rectirostris, P . rer rntur and P. elegnnr. France. England and the Black Sea (76, 77. 151).

Thelohania p a g u r i Ptrez. 1927 (148) . In hody cavity of the hcrmit crab Eiipngurur bernhordus. France.

Thelohonin prnnei Spragur, 1950 (177) . Jn gonad of the shrimp J'eiiaeu~ setiferur. Louisiana, USA. H. F. R . Champion (pers. coni.) found a siniilar or identical parasite in tlir ovary of Penaeur indicus in the Republic of South Africa.

Thelohonin petroliithir Spragur. 1970 (191 ) . Syn . : Thrluhnnia s p . Spraguc, 1950 ( 1 7 7 ) . I n niusclrs of the crah Petrolirther aimatus. Louisiana, US.A.

Thelohania sp. Sogandares-Rrrnal, 1962 ( 175). In musclrs of the crayfish Cambarclluc schujeldi.

7'helohanta sp. Thomas. ( M . M. Thomas. Pen. coni.) I n gonad and ~nusclcs ol the shrimp. Penacus ieniirulcntus d r Ilaan. South India.

C;CIIUS Pleiitophora Giirlfy, I893 (65) .

Louisiana. L'S.4.

Pleirtophorn cargoi (Sprague, 1966) Sprague, 1970 ( 191 ) . Syn.: Plistophora cargo; Spragnr. 1966 ( 1 8 3 ) . I n niusclrs of thc crah Ca l l i u rc tes rapidus Kathliun. Maryland. US.4.

PIeistophorii miJairii (Kudo. 1924) Spraguc, 1970 (191 ) . Syns.: A'osemn airoir inluin ( 7 ) Miyairi, 1909 (102) ; Plistophora miyazrii Kudo. 1924 (102) . In digestive tract of Atyephira sp. Japan.

Plei3tophoia Jogaiidaresi (Sprague, 1966) n. corrib. Syns.: Pl i~ tophora sp. Sogandares-BernaI ( 175) ; Plistophora rogandure~i

PROTOZOAOF DECAPODS 53 1

Sprague, 1966 (183) . In muscles of the crayfish Cambarellus puer Hobhs. Louisiana, USA. Since Plistophora Labbk. 1899, is a junior synonym of Pleistophora Gurley, 1893 (186, 192), this species is transferred to the latter gmus.

Pleistophora sp. Baxter, Rigdon & Hanna. 1970 ( 1 5 ) . In the niusclcs of the shrimp Penaeur ietiferus and P. aztecus. Galveston Bay, Trxas. USA. Probably identical with Plictophora sp. Kruse, 1970 (191) found in the same host.

Pleisfophora sp. Sprague, 1970 (191). In muscles of the shrimp Pnlatnionetes pugio Holthius. Georgia, USA.

Class HAPLOSPOREA Caullery, 1953 (27 ) . Ordrr H.4PI.OSPORID.4 Caullery & Mesnil, 1899 ( 2 4 ) .

Faniily H.API.OSPORIDIIDAE Caullery & Mesnil, 1905 (25 ) . Genus Minchinia Labbe, 1896 (106) .

Minchinia loui\iana Sprague. 1963 ( 1 8 1 ) . Syn.: Haplo- sporidiunz sp. Sprague, 1954 (179) . In the gut wall of the mud crab, Panopeur herbstii. Louisiana, USA.

Minchinia sp. Rosenfield, Buchanan, Chapman, 1969 (162) . In gill, niusclcs and gut tissue of the crab Eurypnnopeu , depressus. Virginia. USA.

Genus Urosporidium Caullery & Mcsnil, 1905 (26) . Urosporidium crescens DeTurk, 1940 (48) . Hyperparasitic in

encysted metacercariae of a distome identified by DeTurk as Spelotrema nicolli, now in genus Microphallur (198). Nortli Carolina ( 4 8 ) : Maryland and Virginia (191) , USA. Sprague (191) recently made observations on the structure of this species, Couch (41) studied its effect on its trematode host and Perkins (149) studied its ultrastructure.

lirosporidium sp. Sprague, 1970 (191) . Hyprrparasitc in cp.- cystcd metacercariae of Microphallus sp. in the shrimp, Palae- monetes pugio. Georgia, USA.

Subphylum CILIOPHORA Doflein, 1901 (78) .

Class CILIATEA Perty, 1852 (78) . Subclass HOLOTRICHIA Stein, 1859 (78) .

Order ASTOMATIDA Schewiakoff, 1896 (78) . Family ANOPLOPHRYIDAE C+de, 1910 ( 2 8 )

Genus Anoplophrya Stein, 1860 (136) . Anoplophiya minima Lkger & Duboscq, 1909 (120) . This

very rare species of astonie ciliate was reported from the posterior intestine and rectum of the lobster, Hornarus gamniarus. France.

Order HYMENOSTOMATIDA DPlage & Htrouard, 1896 (78). Suborder TETRAHYMENINA Faurc5-Fremiet, 1936 (78) .

Family COHNILEMBIDAE Kahl, 1933 (78). Genus Anophrys Cohn, 1866 (34 ) .

Anophrys maggii Cattaneo, 1888 (23) . Syn.: Anophrys sarco- phagu Cohn, 1866 (34) . Occurring naturally, but rarely, in hemolymph of Carcinus maenas. Western coast of France (155) . Poisson (155) stated that the parasitic A. Maggii of Cattaneo is identical witb the free-living A . sarcophaga of Cohn, and thus the narnc A . maggii should be a junior synonym of A . sarcophuga. However, Tucker (14) argued that until more is known about these forms both names should be retained. Bang ( 1 4 ) was ablr to induce fatal experimental infections of A. maggii in Cnrcinus m a m a \ , Main squinado and Macropipus depurator. The ciliates \vithin the hemolymph of these crabs were found to be eating host leukocytes. Bang (14 ) reported that the events leading to death in these Anophrys-infected crabs uere anemia and as- phyxiation folloMed by severe bacteremias.

Order APOS1'0MATID.4 Chatton 8z Lwoff, 1928 (78) . Family FOETTINGERIIDAE Chatton, 191 1 (29, 32) .

Genus Synophrya Chatton & Lwoff, 1926 (29, 32). Synophrya hypertrophica Chatton & Lwoff, 1926 (32). Found

in sills of the fo l lon in~ marine crabs along the west coast of Francr: Macropipus f = Portunur) depurator at Banyuls and Roscoff. M. puber at Roscoff and Wimereux, M. holsatus at Roscoff. M . purillus a t Roscoff and Banyuls, M . tuberculatus at Banyuls. and Cnrcinus maenas at Roscoff. Recently found by Vi\,arPs and Rubi6 (213) in Macropipus tuberculatur, M . bolir,ari, M. depurator and M . t,ernalis on the northeast coast of Spain. VivarPs (214) found the folloning decapods infected with S. hypertrophica in the Gulf of Tunis: Dorippe lanata, Carcinus mediterraneus, Portunrnur lntipes. Macropipus arcuatus, M . cor- rugatus . M . imernalis, Pilumnus Ipinifer and P. hirtellus. This apostonie causes considerable pathogenesis in gills of hosts (32 ) . Life cycle fully described and figured by Chatton and Lwoff (32) .

Genus Gj.n~nodinioides Minkiewicz, 1912 ( 130).

(r'ynznodinioides calkinsi Chatton & Lwoff, 1935 (32) . Phoronts on the gills and trophonts on the cuticle of Polaemonetes sp. Woods Hole, Massachusetts, US.4.

Gynznodinioides caridinae (Miyashita, 1933) Chatton & Lwoff, 1935 ( 3 2 ) . Syn.: HyaIospira caridinae Miyashita, 1933 (132) . Found on the "hairs" a t base of antennules and on the gill filaments of Xiphocuridinu comp7essa (freshwater shrimp of Japan), Leander paucidens and Macrobranchium nipponense by Miya- shita in Japan. Reported as Hyalospira caridinae from Palaemon rarianr by Debaisieux (4.4) in Europe. Debaisieux gave a fairly detailed account of the life cycle of this species on the brackish water Palaenion Lmarians.

~Ij~~rinodinioider inkytnnr hlinLicuicz. 1913 ( 1 3 0 ) . Phoronts on the gills and trophonts on the cuticles of Eufagurus bernhardus, E. prideauxii (130) and Carcinus maenas (32) . Roscoff, France (32, 1:iO). Also possibly observed by Minkiewicz, according to Chatton and Lwoff ( 3 2 ) . on the following decapod genera: Maia, Xantho, Porcellana and Galathea. Trager (207) reported this species from the gills of Pagurus longicarpus a t Woods Hole, 3lassachusetts, USA. According to Trager, the excystment of the phoronts of this species is synchronized with the molt of the host, and the conditions necessary to bring about excystment can be simulated in the laboratory,

Gymnodinioides sp. Chatton & Lwoff, 1935 (32). In the #astrovascular cavity of the anemone Sagartia parasitica found attached to Eupugurus prideauxii.

Gymnodznioides sp. Trager, 1957 (207). Phoronts on gills of Uca pugnax. Woods Hole, Massarhusetts, USA. About 80% of the crabs examined vere infested. Trager reported differences in this species and G. inkystanr to be 10 or 11 ciliary bands rather than 9 as found on the latter. Also, this undescribed species had a more pointed and twisted posterior end than G. inkystans.

Genus Phtorophrya Chatton, Lwoff & Lwoff, 1930 (30). PhtorophrTa fal lax Chatton, Lwoff & L ~ o f f , 1930 (30 ) . A

hyperparasite on the phoronts of Gymnodinioides inkystans on Carcinus maenas. Roscoff, France.

Phtorophrya iniidiosa Chatton. Luoff & Lwoff, 1Y30 (30) . Usually found as a hypcrparasite on the phoronts of Gymnodinio- ides corophii on amphipods, but in at least I instance (32) reported as a hyprrparasite of Gymnodinioides sp. from Carcinus. Membrrs c)f this grnus are characteristically hyperparasitic or phoretic on the phoronts of other apostornes (32) .

Genus Polj..ipirn Minkiewicz, 1912 (130) .

Polj'ipiru delagei Minkiewicz, 1912 (130) . O n the gills of the follouinx hosts: Eupaxurus bernhardu,, Galathea Jquamifera, and Macropodia rostrata. Roscoff, France. Found only in decapod crustaceans (32 ) . Chatton and Lwoff were unable to transmit it easily to Mrrcrupipu, si'. in the laboratory (32). These authors reported that this species is often found in company with Gymnodinioides inkystans.


Genus Phofetophryo Cliatton. 1,uofl & Luiiff. 1930 (31 )

Phoretophrjn nebaline Cliatton. I.\\off & L\\off, 1970 ( 3 1 ) . .4ltho usually found on the non-decapod ,Yebalin geflroyi. this species has also heen rrportt,d from Cnrc; f i r r r innrnni and Mocrop- ipus hohatus. RoscoIT. France.

Genus Terebrorpira Debaisieux. 1960 ( 4 4 )

Tercbrospirn lrnticiclaris (Debaisicux. 1957 ) DrhaiFieux, 1960 ( 4 4 ) . S y n . : Chnttonin lenticularis Debaisieux, 1957 ( 4 3 ) . On the cuticlr of Pnlnemon x i i n n s collrcted from the mouth ol Schclde Rivrr (brackish \ later) in Western Europe (Netherlands). Debaisieux ( 4 t ) gavr thr life cycle and host-ciliate relationship of this species in soinc detail. This specks peiietratcs the cuticle of its host shrimp and thus can cause some damage.

Grnus Wyalophysn Bradbury. 1966 ( 2 1 ) ,

Nyalophysa chattoni Bradbury, 1966 ( 2 1 ) . Phoronts found on the gills and trophonts on thc molts of scverai diffrrrnt sprcics of decapods froni the coasts of Washington and California. USA: Pagurui hirsutiusculus. P. sniriuelir. P. ,graiio~rm~nur. P. alaskeniis, Cnncer oregonenris, and Lophopanopeur belli. Ac- cording to Bradbury. the gmiis H1,alophysn brlongs to that group of apostonies phoretic on Branchyura. .4noniura, Isolwda and Cirripedia uhich develop in the exuvial fluid of thrir hosts. Othm ecologically related genera are Cymnodinioides. Poljr~pira and Phoretophrya. H . chattoni requires only 1 host for its entire life cycle.

Order CHOKOTRICHIDA Wallengren. 1895 (78 ) . Farriily CHILODOCHONIDAE Foche, 1913 ( 153) rmrnd.

Mohr: 1948 (133) . Genus Chilodochona Wallcngren, 1895 ( 2 15) .

Chilodochonn 7iiicrochilus Wallengren. 1895 (215) . On mouth parts of Macropipus dc~purator.

Chilodochona quennerstedti Wallengen. 1895 (215) . O n Ebalia turnefacta and Mncropipuc depurotor. Southern Sueden (87 ) . Molir ( 1 3 3 ) divided the Order Chonotrichida into 3 familirs and piiinted out that most chonotrichs found on decapod crustaceans are in the family Cliilodochonidae.

Southern S ~ t c d e n .

Genus Kentrochonn Koinpel, 1894 (161) Kentrochonn nebnline Rompel, 1891 (161 ) . On the dccapods

Ebalia turnefnctn and Portunus depurator. Southern Sweden. Also rcported by DoRein (50 ) from Tricstc and Rovigno. in the northern Adriatic.

Subclass PERITRICHIA Calkins, 1933 (78 ) Order PERITRICHIDA Stein. 1859 (78 ) .

Suborder SESSILINA Kahl, 1933 (78 ) . Family VORTlCELLIDAE Ehrenherg, 1838 (140) .

Genus Vorticella L., 1767 (136) , emend. Ehrenberg, 1838 (140) .

Vorticella ~ ‘ e r r z ~ c o s u Dons, 1915 (55 ) . Found on Laomedia sp. from Norway. Of many dcscribcd species of Vorticella ( 8 7 ; 137; 140), this form is the only named vorticellid habitually found on decapods. Many species have been reported from tion-dccapod crustaceans.

Genus Zouthamntuin Bory, 1826 (19)

Zoothamnium carcini Kent, 1881 (98 ) . On Carcinus maenaz from the south coast of England.

Zoothamniuin procerius Kahl, 1933 ( 8 7 ) . On legs of the crayfish Astucus fEui,iatilis from \ iciiiity of Erlangrn, Germany. Nenningcr (137) . reported a xarirty of this species, which she named Z . procerius \ar . inrius from both gamtnarids and crayfish, but until subspecies names are more generally accepted for ciliates, this form is recognized as %. procerzus.

Zoothnmnium sp. Ryder. 1883 (163). On “back and legs” of the oyster r~inini~~nsal decapod Pinnotheres ostrrum Say. Mary- land, US.4. Ryder suggested that because thr Zoothnmniuni sup- ported on its “stalks” hactrria and “\ibriones.“ it might he of henPfit as a source of food to the oyster in \\-hich its crab host lives.

Family EFISTYLIDID.4E Kahl. 1933 (87 ) . (;<%nus Epirt1,lir Ehrenherg. 1832 (58 ) .

/ipi\tTlir nrtnci Nenninser. 1948 (137) . On the gills of Artncuc Pu:,intilir. Germany.

Epi\t)’lir bimor,qinotn Kcnningrr. 1948 ( 1 3 7 ) . On legs of Astucus pui,intilir. Germany. Nenninger described this peritrich from the crayfish as a \.ariety. E. bimarginnta var. urnula. hut asain, until subspecific namrs are tnore generally accepted for ciliates this form is recognized only as a member of the species E. bimuiginata.

Epirt9’lis canibnri Krllicott, 188.5 (94 ) . O n gills of a species of Caniborur. Sorthcastvrn United States.

Epiltylir ningarne Krllicott. 1883 (92 ) . On body surface of crayfish ( ? j in freshwater. Northern United States.

I:‘pist),/ic sp. Hutton. 1964 (80 ) . On the coniiiic!rcial shrimp Penaeus duoruruni . Numerous localities in Florida, USA.

Epiil3’/i< SI). Ilutton, 1964 (110). O n the deep-uater shrimp 11~~nirnopcnaeur robuitzir, from I 7 0 fathorris off Daytona Beach, Florida.

Iipiitjlh sp. Pears?. 1932 (142) . O n gills of Coenobita cljy peactur, at Garden and Long Keys, Florida, USA. On Geogrnp- ( u s l i ; idus and Pnch~,grnprus t r n n s ~ r ~ n s in the Florida keys, L’S‘Z.

(;cmiiis Cnichciiiiiii Ehrenherg, 1832 (58 ) . Cnrchesiunz granu ln tum Kdliciitt, 1887 (95 ) . On surface of

freshuater crayfish of grnus Cnnibnrus. USA.

Farnily VAGINICOLIDAE de Fromentel, I1174 ( 6 2 ) . Genus I’aginicola Larnarck. 1816 ( 108).

Vuginicola paguri (.4ndrC. 1910) Kahl, 1933 (87 ) . S y n . : Cothuruia poguri Andri.. 1910 ( 2 ) . From cuticle of Eupngurus bernhnrdus. Roscoff, Franc?.

Genus Cothurnia, Ehrmhrrg, 1832 (58) Cothurnin nrtnci Strin. 1854 (195) . O n the branchial all-

pendages of Actacur j?u:,intilir. Europe. Thought to h e associated with mass mortalities of fresh\\-atrr crayfish in Italy as reported by Ninni (139) and by Kent (98 ) . Kent (p . 721) said, “During a fatal epideniic among the freshnater cray-fish of Lomliardy and Venetia in the year 1863, it is reported by Ninni that the dying crustacea were completely iiifrsted uith an Infusorium, referable to the genus Vaginirola. prohal)ly identical with the present species IC. astoczi. its abnorinal de\ clopmcnt on the branchial appendages apparently producing the death of the crustacean through asphyxia.”

Cothurniu currn Stein, 1851 (195) . From fresh\\ater crayfish. Europe.

Cothufnio ricboldi Stein. 1C1-1 (195) . O n thr liranchial appendages of the frcsh\\ater crayfish Astacus purintilis.

Cothurniir ;,urinbilis Kellicott. 1884 (9‘3). Occurs by thou- Europe.

sands on gills of crayfish Cambnrur afinis (93 ) and flui’iotzlir (137 j .

Family LAGENOPHRYIDAE Kahl, 1935 (89 ) . Genus Lngcnophrj’s Stein, 1852 (194) .

This large genus of loricate peritrichs (52 species at is rmiarkalh in that it has 2 2 species that are found decapod crustaceans and ha\ e strong host sprcificity.

Lagenophrys aegleae Mouchet-Bennati, 1932 ( 134) . sills of the freshnater crayfish Aeglea laetis. Uruguay.

Astncus

present) only on

O n the


Lagenophrys articularii Nie 8: Ho, 1943 ( 1 3 8 ) . On the anten- nae, legs, and s\\immerets of Polaenion nipponen\is. China.

Lagenophryr rir~~erinrrrc~i Ahonyi, 1928 ( 1 ) . O n gills of the frcshnatrr rivw crah Trlphurn fluciotilii. Africa. .2honyi gave thi, lagmophryiid its specific t iam? h s e d upon thc dc~.scription of A\\erinx\v ( 5 ) .

Lagenophrys birpinow Kaue, 1965 (91 ) . On suimnicrcts of Chernu rotundur t e to%ur . Stream nrar Nc\\castlc. N r \ v South Walrs, Australia.

Lu,qrnophryi brunchinrnm Nie & 110. 1943 (138). On the gills (only) of Palaenion 7iifiponeii,,is. China.

Lagetioflhrj,~ cnllinectes Couch. 1967 ( 4 0 ) . On both surfaces of the gill lainellac of the conitrirrcial, niarine and hrackish 1% ater crah Cnllinecter sapidur. From Chesapeake Bay, Mary-land aud Virginia: south to Gulf of Mi.xico estuarine uaters wherever blue crab is found. Durins summer months infestations are so hea\y that the gill surlaccs of the crah may be rompletely covrrcd.

Lagenophrp coniniunis Knnc, 1965 (91 ) . On the telsiins, suinrincrcts, and cuticles of the crayfish Chernx deitructor, C . nlbidui, Ijicu'tocus s u r r u t u < . Ii. nobzlis, and Engaeus sp. Aus- tralia.

La,qenophrj,r darri ini Kaur, 1965 (91 ) , On the gill filaments of C h e r u x quudricnrtnatu~. I?r&\\ater. uear Daru in , northern .lustralia.

1.ngenophrj's deserti Katie, 1965 (91 ) . In the branchial cham- hers and on gill filaments of Cherax tenuinranus and C. quinque- curinntus. Ri\ rrs in southrastcrn .4ustraha.

Lngenophrys dungogi Kaur. 1965 (91 ) . On the gill filaments of a single specimen of Euartacus si). Stream in Dungog, Australia.

Lagenuphi).\ engnei Kane, 1965 (91 ) . On brachiostegitr membranes and sill filarncnts of crayfish Engaeus rictoriensis, and on gill filanient and gill bases of crayfish Austroactncus hemi- cirratulis. Near Melbourne. ..lustralia.

Lagenophrys eupagurus Kellicwtt, 1894 (96 ) . On gills of hcrmit crab 1:'upaguru.i sp. M'ocids Hole, Massachusetts, USA.

Lagenophrys latiipinosn Kane, 1965 (91 ) . O n the flat surfaces of gills, gill bases and hranchiostegitc mcnibranes. On Cherax dertructor near Melbourne, Liustralia. O n C. dertructor and C. nlbidus from Victoria, .4ustralia. O n C. rotundu, from Nc\r South Wales, .4ustralia. Kane remzrked that the loricae of many generations of this specics arc supi~in~posed on one ariothcr on the cuticle of older hosts.

Lagenophrys lnzi'rii Kane, 196.5 (91 ) . On flat surfaces of the gills of Cherax quadricarinatuc. Near Darwin, Australia.

Lagenophrj'r lingulnto Kanc, 1965 (91 ) . Found mostly on gill filaments, occasionally on thr I~ranchiostegite membranes, On Cherax destructor and C. ailridus from Victoria, Australia, aiid on C. rotundus in central Neiv South \Vales, Australia. Type host and locality are Chernz destructor from around hlelhournc. Australia.

Lagenophiys lunutus Imaniura, 1940 (81 ) . On body cuticle of shrimp Leander pnucidens (83) and Palurnion i ,ar ians (4<!. Jaljan ( t i l ) ; Bclgiuru ( 4 I ) . One of u s (J. C . ) has found this species 011 Pnlnenionetes c'ulgnrii from Chincoteague Bay, Mary- land. Dchaisieux (44) has done an escrllent job o f \\orking out the life-cyrle of this sprcies.

Lagmophrys nietopnuliadis Corliss & Brough, 1965 (39 ) . On the gill lamr~llae of a fresti\sater crab Metopnnlini dppressus. Jamaica. l h i s was the 1st sprcics of LagenophrjBs to be reported from a brachyuran crab.

Lagenophrys occlusa Kane, 196.5 (91 ) . Restricted to the an- terior portion of the branchial chaluhcrs of Cherax deitructor, C. albidus, and C. rotundus. Victoria and Ne\v South \Vales, Australia.

Lagenophrys rugosa Kanc, 1965 191). O n the gill filaments of fresh\\ ater crayfish Georhnrax falcata. Western Victoria, .4ustralia.

Lagenophrys seticola Kane, 1965 (91 ) . Original description

hased on specimrns found on a land crayfish Engaeus fultoni lrom a burro\\. in a river bank near Melbourne. Australia. Found zttached to setae, tail fan, sjvimmerets, gill base and edge of branchiostrsites. Specimens referable to this species have also hren found on Chernx rotundus setosus, C. albidus, C . destructor and Bunitacus nobtlis from Australia.

Lagenophrys spinora Kane, 1965 (91 ) . On the cuticle, swim- merets, telson and carapace of the crayfish Cherax destructor. Victoria. .4ustralia.

Lagenophrys r d l i s i Kanr, 1965 (91 ) . O n gill filammts of fresh\*.atcr crayfish Cherax destructur, C. albidus and C. rotundus. Australia.

Subclass SPIROTRICHIA Biitschli, 1889 (78) . Order HETEROTRICHIDA Stein, 1859 (78 ) .

Suborder IfE'I'ERO'lKICHINIA Stein, 1859 ( 7 8 ) . Family FOLLICVLINIDAE Dons, 1914 (54) .

hlrmbcrs of this family belong to several different genera, all species of \\hich are loricatr. Species reported here as epibionts of decapods also may be, and usually are, found on substrates other than decapods and generally lack host specificity. FaurE- Fremict (60) and Hadii (68) reviewed the species in this family and their various substrates.

Genus Pebrilla Giard, 1888 (63 ) .

Pebrilla p a p r i Giard, 1888 (63 ) . Found on the cuticle of Pagurus bernhardus by Giard, in 1888, a t Pouliguen and at Rosruff, France. Latrr (1936), reported from cuticle of Cliba- norius misanthropus from France by Fad-Fremie t (60) , who helievrd that. tho this sprcies has fratures characteristic of members of grnus Folliculinopszs FaurE-Fremiet,l936, its generic name Pehrilla should l i e retained.

Genus Plntyfol/iculina Hadii, 1938 (67) . P/atyfu//iculina lJnguri Andrews & Reinhard, 1943 ( 3 ) . On

State of Wash- the abdomen of hermit crab Pagurus pubescens. ington, USA.

Genus Metafolliculina Dons, 1925 (56) .

Metafolliculina nndrezc,ri Hadii, 1938 (67) . On Pagurus pori- danus as reported by Andrew ( 4 ) . Texas Gulf Coast, USA. Andreus stated that tlie trsts of M . andreursi were found on the abdominal filaments, and "large holding appendages." This species \ \ as also found on oyster shells in same region by Andrews.

Genus Folliculinopris FaurE-Fremiet, 1936 ( 6 0 ) .

Follicdinopsis gunneri (Dons. 1928) FaurE-Frrmiet, 1936 (60). Syn.: Folliculina girnneri Dons, 1928 (57 ) . Found by Faure- Freniiet on thr cuticle of sacculinid parasite of Carcinus maenas and Porccllana plat),chrlri. France.

Genus Folliculina Lauiarck, 1816 (108) Folliculina r,iridis (Wright. 1858) Faxre-Fremiet, 1936 ( 6 0 ) .

Syn.: Lugotiu ziridis Wright, 1858 (218) . Found on tegument of sacculinid parasite of Carcinur maenas and Porcellana platycheles along with 1;olliculinopsis gunneri (60 ) . France.

Subclass SUCTORI.4 Haerkcl, 1866 (69 ) .

Family PODOPHRYIDAE Haeckel, 1866 (69) . Genus Podophrjsa Ehrenberg, 1835 (59 ) .

Order SCCTORIDA Claparkle 8: Lachmann, 1858 (78)

Podophrya nstaci Clapari.de 8: Lachmann, 1861 (33 ) . From the freshwater crayfish Astacus fluuiatilis. Europe.

Genus Parucineta Collin, 1912 (37) . Paracineta homari (Sand, 1899) Kahl, 1934 (88) . Syn.:

Acineta honrari Sand, 1899 (164) . Found on an unidentified

http://Clapari.de


species of lohster. Ifomaruz. and also on other decapods including Pngnruc SI) . Roscofl', Franc?.

Family .4CINETIDAE Stein, 1859 (196) . Genus Acinetn Ehrcmhcrg, 1833 ( 5 9 ) , emend. Collin. 1912 (37) .

Acinetn con\trictn Collin. 1909 (36) . On the legs of Eupagurus cuanencir. Crttr (SGtc), France.

Acinetn sp. Courh. this I)aper. On the flat surfaces of gill lamellac 111 hluc crahs. Cnllinectes snpidus. Chesapeake Bay. Maryland. USA. and Chincotrague Bay, Virginia, USA. This iindc.srrilxd species is not common and orrurs as solitary indi- viduals attarhcd near periphery of gill lamellae.

Fainily OPHRYODENDRID.4E Stein, 1867 ( 197) Gcnus Dendrosomides Collin, 1906 (35) .

Dendroroniides p q u r i Collin. 1906 ( 3 5 ) . On the "long hairs" (sctar) of tlir thoracic apprndages of Eufagurus excavatus and E. cuanen.ric. Cette (S6tc). France.

Genus Opliryodendron ClaparGdr & Lachmann, 1861 (33) . Ophr],odendron porcellnnum Kent, 1881 (98) . O n the setae

of the anomtiran Porcellnna platycheles . Jersey (English Channel Island) (98) and Roscoff, France ( 8 8 ) .

We gratefully acknoivledgr our indebtedness to Mrs. Helen Ixmg for aid in locating sonic of the literature.

REFERENCES 1 . Ahonyi, A. 1928. Ober die Epizoen der Orchestin caoinana.

Arb. B i d . Forsch . - In~ t . 2, 5-23. 2 . Andre, E. 1910. Sur quelques infusoircs marins parasites et

coniniensaux. Rer,. Suisse Zoo/. 18, 173. 3 . Andreus, E. A. 8: Rrinhard, E. G. 1943. A folliculinid

associated with a hermit crab. J . Wnrh. Acad. Sci. 33, 216-23. 4. Andrcws, E. A. 1946. Folliculinids on a hermit crab from

Trxas. l'ran\. Am. Micr. Soc. 6.5, 359-61. 5. A\verinzr\v, S. 191 2 . Beitrage zur Entwicklur.gsgeschichte

der Lagenophrys SI,. B i d . Zentralbl. P2, 714-8. 6. Balbiani, G. 1882. Sur les microsporidies ou psorospermies

dos articules. C. R. Acnd. Sci. 95, 1168-71. 7. Ball, C. H . 1938. The life history of Carcinoecetes hesperus

n. gcn.. n . sp. . a gregarine parasite of the striped shore crab, Pachygrnpsus crnecipes, \\ ith observation on related forms. Arch. Protist. !MI, 299-3 19.

8. - - 1948. .4 gregarinr from the sand-crab: Emerita rrnaloga. J . Pararit. 34, 17 LAbstr. 231.

9. - -. 1950. Some pathogenic effects of gregarines on tlirir hosts. Ibid 36( 6 / 2 ) , 36 IAbstr. 861.

10. -, 195 1, Gregarines from Bermuda marine crusta- cmns. C'nic,. Calif. Pub. Zoo/. 4 i , 351-68.

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12. . 1959. Some gregarines from crustaceans taken near Bombay, India. J . Proto-ool. 6 , 8-13.

13. 1963. Cephnloidophora carpilodei n. sp. and C. PfFquis n. sp., gregarinrs of a xanthid crab in Hawaii. Ibid 10, 3-1-7.

14, Gang, F. B. 19iO. Disease iiiechanisms in crustacean and marine arthropods. In S. F. Snirszko, ed. A Symposium on Dicenses of Fishes and Shellfishes, Spec. Publ. 5, Am. Fish. SOC., Washington, D. C., 383-404.

1970. Pleisto- phora sp. (Microsporidia: Nosematidae) : A new parasite of shrinip. J . Inz,ert. Pnth. 16, '189-91.

16. Beauchatrip. P. de 1910. Sur tine gregarine nouvelle du genre Porosfiorn. C. I ( . Acnd. Sci. 151, 997-9.

17. Bcneden. E van 1869. Sur tine nouvelle espPce de Gregarina designte sous le nom de Gregarina gigantea. Bull. Acad. Iio)'. Belg. 28, +44-56.

18. Bogolrpova, I . 1. 1953. Gregarines of Peter the Great Bay. T r o , . In l t . ZOO/. A c a d . Sci. I.IISS 13. 38-56, [In Russian].

19. Bory de Saint-Vincent. J. B. 1826. Essai d'une Classifica- tion des Animaux Microscopiques. Paris. 104 p.

20. Bovee. E. C. 8: Jahn, T. L. 1966. Mechanisms of move- ment in taxonomy of Sarcodina. 111. Orders, suborders, families, and subfamilies of the superorder Lobida. Syst. Zool. 15, 229-40.

15. Baxter, K . N., Rigdon. R. H. 8: Hanna. C.

21. Bradbury, P. C. 1966. Hj'alophysn chattoni n. g. n. sp. 1. Protorool. 13, 209-25.

22. Calkins, G. N. 1933. The Biology of the Protozoa, 2nd ed. Lea C Febiger, Philadelphia. 607 p.

23. Cattaneo, G. 1888. Su di un infusorio ciliato, parassito del sangue del Carcinur maenas. Zool . Anr. 11, 456-9.

'4. Caullery, M. C Mesnil, F. 1899. Sur le genere Aplo- sporidium (nov.) et l'ordre nouveau des Aplosporidies. Note preliminaire. C. R. Soc. B i d . Ser. 1, 11, 789-91.

25, 1905. Recherches sur les haplosporidies. Arch. Zoo l . E x p . G in . , Ser. 4, 4, 101-81.

26. ~ . 1905. Sur quelques nouvelles haplosporidies d'annelide. C. R. Soc. Biol. 58, 580-3.

27. Caullery, M. 1953. Appendice aux sporozoaires. Classe des haplosporidies (Haplosporidia Caullery et Mesnil 1899). In P. P. Grass&, ed. Trait t de Zoologie, Masson, Paris 1 ( 2 ) , 922-34.

28. C&pPde, C. 1910. Recherches sur les infusoires astomes. Arch. Zool . E x p . GQn. 3, 341.

29. Chatton, E. C Lwoff, A. 1926. Les Synophrya, infusoires parasites internes des crabes. Leur evolution A le mue. Leur place parmi les Foettingeriidae. C. R. Acad. Sci . 183, 1131-4.

30. Chatton, E., Lwoff, A. C Lwoff, M. 1930. Les Phtorophrya n. g., ciliks Foettingeriidae hyperparasites des Gyrnnodinioides, Foettingeriidae parasites des crustaces. Ibid 190, 1080-2.

31. 1930. Phoretophrya nebaliae n. g., n. sp., et I'interpretation du cycle evolutif des cilies Foettingeriidae. Ibid 190, 1154-6.

32. Chatton, E. & Lwoff, A. 1935. Les ciliCs apostomes I. Arch. Zool . E x p . G i n . 57, 1-453.

33. Claparl.de, E. 8: Lachmann, J. 1861. etudes sur les infusoires et ICs rhizopodes. Metn. Inst. N a t . Genevois. 7 , 1-291.

34. Cohn, F. 1866. h-cue lnfusorien im Seeaquarium. Z . Wiss. Zoo/. 16, 253.

35 . Collin, R. 1906. Note preliniinaire sur un Arinetien nouvcau: DendroJoniides paguri, n. g. n. sp. Arch. Zool. Exp . Gin . 7 ( 4 ) .

36. 1909. Diagnoses pdliminaires d'Acinetiens nou- veaux OLI mal cunnus. C. R. Acad. Sci. 194,

37. 1912. etude monographiques sur les acinetiens. 11. Morphologie, physiologie, systematique. Arch. Zool. E x p . G t n . 51, 1-457.

38. Corliss, J. 0. & Levine, N. D. 1963. Establishment of the Mirroaporidea as a new class in the protozoan subphylum Cnido- spora. 1. Protorool. 10 (Suppl.), 26-7.

39. Corliss, J. 0. 8: Brough, I . 1965. A new species of Lagenophrys (Ciliata: Peritrichida) from the Jamaican crab Metopaulias depressus. Trans. A m . Micr. Soc. W, 73-80.

40. Couch, J. A. 1967. .4 new species of Lagenophrys (Cili- atea: Peritrichida: Lagenopliryidae) from a marine crab, Calli- nectes sapidus. Ibid 86, 204-1 1.

41. ( I n ms.) Pathogenic effects of Urosporidium infections in microphallid metacercariae. A histological and histo- chemical study.

42. Cuckler, A. C. 8: Fichter, E. 1939. Gregarines from the crab, Cancer productus. J . Parasit. 2S, 184.

43. Debaisieux, P. 1957. Foettingeride A migration trans- cuticulaire. Bull. Acad. Roy. Belg. ( 0 1 . Sciences) 43, 98-105.

44, 1960. Ciliates apostomes parasites de Palaemon. Cellule 60, 333-52.

45. Desportes, I. C ThOodoridPs, J. 1965. L'ultrastructure de la gymnospores des Porospora (Eugregarina, Porosporidae). C. R . Acad. Sci. 260, 1761-2.

46. Desportes, I. 1968. Sur la prCsence de Cephaloidophora olim'a (Watson ) (Eugregarina, Cephaloidophoridae) chez le crabe Ocypode cursor L. (Crustacea Decapoda) au Liban. Vie Milieu 19( I-A), 201-2.

47. DeTurk, W. E. 1940. The parasites and commensals of some crabs of Beaufort, North Carolina. Thesis, Duke University. Unpublished.

48. __ , 1940. The occurrence and development of a hyperparasite, Urosporidium crescens (Sporozoa, Haplosporidia) which infests the metacercariae of Spelotrema nicolli, parasitic in Cal- linectes sapidus. J . Elisha Mitchell Sci . Soc. 56, 231-2.

49. Dobell, C. 1925. The life-history and chromosome cycle of Aggreyata eberthi (Protozoa, Sporozoa, Coccidia). Parasitology 1: 1.12Ii - . , . . I - .

50. Doflein, F. 1897. Studien zur Naturgeschichte der Proto- zoen. I. Kentrochona nebaliae Rompel. Zool. Jahrb. Anat. 10, 619-41.


51. 1898. Studien zur Naturgeschichte der Protozoen. 111. Uher Myxosporidien. Ibid 11, 281-350.

52. Dogiel, V. .4. 1965. General Protozoology, 2nd ed. LRev. J. I. Poljanskij & E. M. Chejsinl, Oxford, 747 p.

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54. Dons, C. 1914. Folliculina - Studien IV. Vorliiufige Bemer- kungen uher die Systematik der Folliculiniden nebst Beschreibung neuer norwegischer Arten. Tromsyl Mus. Aarsh. 37, 59-92.

55. 1918. Neue marine ciliaten und siictorien. Ibid 38 & 39(1915/16), 75-1QO.

56. 1925. M:.rrfolliculina nordgordi n. gen., n. sp. Kgl. Norsk. Videnskab. 5elsk . Skr., 1924, 1, 1-13.

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61. Frenzel, J. 1885. Ueber cinige in Seethieren lebende Greg- arinen. Arch. Mikroskop. Anat. 24, 545-88.

62. Fromental, E. G. de 1874. Etudes sur les microroaires ou infuioires proprement dits comprenaut de nouoelles recherches sur leur orgunisation leur classification et la description des erp2ce.s nouzxlles ou j e u connues. Paris, viii + 364 pp.

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76. Henneguy, F. & Th&lohan, P. 1892. Sur un sporozoaire parasite de muscles des crustaces dtcapodes. C . R . Acad. Sci. 114, 1552-5.

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81. Imamura, T. 1940. ‘luo species o f Lagenophrys from Sap- poro. Annot. Zool. J u p . 19, 267-70.

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85. Jahn, T. L. & Bovee, E. C. 1965. Mechanisms of move- ment in taxonomy of Sarcodina. 1. As a basis for a new major dichotomy into two classes, Autotractea and Hydraulca. Ani. Midl. Nat. 73, 30-40.

86. Jones, E. E. 1958. Microsporidiosis in shrimp. Ass. South- east. Biol. Bull. 5, 10.

87. Kahl, A. 1933. Ciliata lihera e t rctorommcrisalia. In G. CrimDe & E. Waeler (ed.) . Die Tierzuelt der Nord-und Ostee. Lief. ‘23 (Teil II,-ct), ‘29-ii6. Leipzig.

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90. Kamm, M. W. 1922. Studies on grcaarines. 11. Synopsis of the polycystid gregarines of the world, excluding thosc from the Myriopoda, Orthoptera and Coleoptera. Il l . Biol. Monogr. 7 , 1-104.

91. Kane, J. R. 1965. The genus Lagenophrys Stein, 1852 (Ciliata, Prritricha) on Australian Parastaridae. J . Protorool. 12, 109-22.

92. Kellicott. D. S. 1883. Efiistylis niagarue sp. 11. on crayfish. Trans. A m . M i c r . Soc. S, 110.

93. ~- . 1884. Observations on infusoria, with descriptions of ne\\- species. [bid 6, 110-24.

94. 1885. Epirtglis cumbari sp. n. on Canzbarus. Ibid i , 43.

95. 1887. Notice of some fresh-water infusoria. With remarks on collecting and preservinl: these delicate animals. Microscope i , 225-33.

96. __ . 1894. On certain marine infusoria observed at Woods Hole, Massachusetts. 2nd Ann. Rep. Ohio State Acad. Sci. 10-1 I lAhstr.1.

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98. Kent, W. S. 1881-1882. A Manual of the Infusoria 2 , 473-913; 3, 51. Bogue, London.

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100. 1966. Life cycle studies on Neniatopsis duorari 11. sp. (Gregarinia: Porosporidae), A parasite of the pink shrimp (Penoeus duorarunz) and pelecypod molluscs. Diss. Abstr. 27B, 2919-B.

1966. Liie cyclc studies on Nematopsis duoiari 11. si’. (Gregarinia: Porosporidae), A parasite of thr pink shrimp (Penaeur duorarum) and pelccypod molluscs. Ph.D. ‘l‘hesis, Florida State. Univ., Mirrofilrus, Inc., Ann Arbor, Michigan, i-x\- -+

102. Kudo, R. R. 192-1. .4 biolosic and taxonomic study of the Microsporidia. I l l . B i d . Monogr. 9, 1-268.

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104. 1946. Protozoology, 3rd cd. Thomas, Spring- field, 111. xiii + 778 pp.

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107. 1899. Sporozoa. In 0. Biitschli, ed. Dar Tier- reich. 5, Friedliinder, Berlin. xii + 180 pp.

108. Lamarck, J. B. 1816. Histoire A’aturelle des Anirnaux sans Vertibres. ( 1 1 ) . Paris.

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110. Ltger, I,. 1901. Sur iine noiiwlle Sregarinc parasite drs Pinnotheres des moules. C. li. Acad. Sci. 132, 1343-6.

101.

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111. __ . 1903. Sporozoaire parasite des moules et autres lamellibranches comestibles. Ibid 137. 1003-6.

112. Leger, L. & Duboscq, 0. 1903. Aggregata ragans n. sp. gregarine gymnosporCe parasite des Pagures. Arch. Zool. Exp . Gin. 31. 147-51.

113. Lkger, L. 1905. Un nouveau Nematopsis parasite des moules de la MkditerranCe. Paris Bull. Mem. Franc. Auanc. Sci. 9, 331.

114. Leger, L. & Duboscq, 0. 1906. Sur l’evolution des gregarines gymnosporees des crustac6s. C. R . Acad. Sci. 142, 1225-7.

115. __ . 1906. L’evolution d’une Aggregata de la seiche chrz le Portunus dejurator Leach. C. R . SOC. B i d . 58, 1001-3.

116. __ . 1907. L’Cvolution des Frenzelina (n.g.) , gregt- rims intcstinalrs des crustacfs d6capodes. C. R . Acad. Scz. l la , 771.4 ..

117. __ . 1908. Evolution schizogonique d’dggregata eberthi. Arch. Protist. 12. 44-108.

118. 1909. Sur une microsporidie parasite d’une gregarine. C . R . Acad. Sci. 148, 733-4.

119. 1909. Etudes sur la sexualit6 chez les gregarines. Arch. Protist. 17, 19-134.

120. 1909. Protozoaires parasites de I’intestin du homard. C. R . Acad. Sci. 148, 363-5.

121. __ . 1910. Selenococcidium intermedium Leg. et Dub. et la systernatiques des sporozoaires. Arch. Zool. E x p . Gin. Ser. 5, 5, 187-238.

122. 191 1. Deux gregarines de crustacbs. Porospora portunidarum Frcnz. et Cephaloidophora maculata n. sp. Arch. Zool . E x p . Gin., N & R 6, 59-60.

123. 1911. Deux nouvelles espkes de gregarines ap- partenant au genre Porospora. Ann. Uniu. Grenoble 23, 401-4.

124. __ . 1913. Le cycle evoliitif de Porosfora portuni- durum Frcnzel. C. R . Acad. Sci. 156, 1932-4.

125. 1913. Sur les premiers stades du dkveloppement des gregarines du genre Porospora (= Nematopsis). C. R . Soc. Bid . 75, 95-8.

126. 1915. Porospora nephrogis n. sp. Ibid is , 368-71. 127. __ . 1925. Les porosporides et lcur 6volution. Trau.

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130. Minkiewicz, R. 1912. Ciliata chromatophora, nouvel ordrc d’infusoires B morphologic et reproduction bizarres. C. R . Acad. Sci. 155, 513-5.

131. ___ . 1914. Etudes sur les infusoires syndesmogames, i gamontes et gamltes. Bull. Int. Acad. Cracocie (Acad. Polon. S c i . ) (B) 1913, 742-9.

132. Miyashita, Y. 1933. Studies on a freshwater foettingeriid ciliate, Hyalospira caridinae n.g., n. sp. lap. J . ZooE. 4, 439-60.

133. Mnhr, J. L. 1948. Trichochona lecythoides: a new genus and species of marine chonotrichous ciliate from California, with a consideration of the composition of the Order Chonotricha Wallengren, 1895. Allan Hancock Found. Pub. Occas. Pap. 6, 21 PP.

134. Mouchet-Bennati, S. 1932. Sur la branchie d‘deglca laeuzs (Latreille) et son parasite Lagrnophrys aeglea nov. sp. C. R . Soc. Biol. 109, 148-50.

135. Nigeli. 1857. Ubcr die neue Krankheit der Seidenraupe und verwandte Organismen. Bot. Zeit. l 5 ( 4 4 ) , 760.

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137. Ncnninger, U. 1948. Die Peritrichen der Unigehung von Erlangen mit brsonderer Beriicksichtigung ihrer Wirtsspezifitiit. Zool. Jahrb. Syst. 7 7 , 168-265.

138. Nie, D. & Ha, Y. 1943. Notes on some epizoic infusoria from the freshjvater shrimp, Palaemon nipponensis. Sinensia 14, 143-9.

139. Ninni, A. P. 1864. Sulla mortaliti dei gamberi (Astacus fluz’iatilis) ne1 Veneto c pi“ particolarmente nella prnvincia trevigiana. Atti Imp. R o y Irrstituto Veneto 10 ( 3 ) .

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141. O r m i h s , R. 1968. Pachyporospora retorta n. sp. gregarine parasite de Macropipus holsatus F. crustac6-dbcapode. Ann. Para- sit. Hum. Comp. 13, 413-9.

142. Pearse, A. S. 1932. Observations on the parasites and commensals found associated with crustaceans and fishes at Dry Tortugas. Pap. Tortugas Lab., Carnegie Inst., Washington, D. C.. 28, 103-15.

143. Pellbrdy, L. P. 1965. Coccidia and Coccidiosis. Pub. House Hungarian Acad. Sci., Budapest. 657 pp.

144 Pkrez, C. 1904. Sur une mirrosporidie parasite du Carcirrus maenas. C. R . Soc. L i d . 67, 214-5.

145. 1905. Sur une nouvelle gluq?idee parasite du Carcinus maenac. Ibid 58, 146-8.

146. 1905. Microsporidies parasites des crabes d’Ar- cachon. SOL. Sci. Arcachon Trau. Lab. 8, 15-36.

147. 1906. Sur un cas d’envahissement de l’ovaire par Thelohonia maenadis. C. R . SOL. Biol. GO, 1091-2.

148. 1927. Notes sur les epicarides et les rhizocephales des cBtes de France. I . Sur I”‘Eupagurus bernhardus” et sur quelques-uns de ses parasites. Bull. Soc. Zool. Fr. 52, 99-104.

149. Pcrkins, F. 0. 1971. Sporulation in the trematode hyperparasite Urosporidiurn crescen.r DeTurk, 1940 (Haplosporida: Hap1osporidiidac)-An electron microscope study. J . Parasit. 57,

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155. 1930. Observations sur Anophrys sarcophaga Cohn. Bull. Bid . Fr. Belg. 64, 289-331.

156. Porchrt-Hrnnere, E. & Richard, A. 1970. Structure finc dcs microganietes d’Aggregatn eberthi Labbe. Protistologica 6 , 71-81.

157. 1970. Structure finc du sporoblaste immature uni- nurlee d‘Aggregatu eberthi LabbC. (Sporozoaire Coccidioinorphc) , C . R . Acad. Sci. 269, 1681-3.

158. -- -. 1970. Ultrastructurc dvs stadrs vfgktatifs d’Ag- gregata eberthi LabbC: le trophozoite ct IC schizonte. Z. Zell- forsch. 103, 179-9 1.

159. Prytherch, H. F. 1938. Life cycle of a sporozoan parasitc of the oyster. Science 88, 451-2.

160. 1940. The life cycle and morphology of Nema- topsis ostrearum, sp. nov., a gregarine parasite of the mud crah and oystcr. J . Morphol. 66, 39-64.

161. Rompel, J. 1894. Kentrochona nebaliae n. g., n. sp. ein neues Infusor aus der Familic der Spirochoninen, zugleich cin Beitrag zur Lehre von der Kernteilung und den1 Centrosoma. Z. W’iJs. Zool. 59, 618-35.

162. Roscnficld, A,, Buchanan, L. 8: Chapman, G . B. 1969. Comparison of the fine structure of spores of three species of Minchinia (Haplosporidia, Haplosporidiidae). 1. Parasit. 5.5, 921-41.

163. Ryder, J. A. 1883. The protozoan parasites of the oyster. Science 1, 567-8.

164. Sand, R. 1899. Etude monographique sur le groupe dcs infusoires tmtaculifrrs. Ann. SOL. Belg. Micr. 24, 57- 189.

165. Sauycr, T. K . 1969. Preliminary study on the epizootio- logy and host-parasite relationship of Paramoeba sp. in the blue crab, Callinectes sapidus. Proc. N a t . Shellfish. Ass. 69, 60-4.

166. 1969. Blue crab studied by Maryland biologists. Chesapeake Bay Affairs C o m m Fish. N e w s 2 ( 6 ) , 2.

167. Sauyrr. T. K., Cox, R. & Higginbottom, M. 1970. Hemo- cyte values in hralthy blue crabs, Callinectes sapidus, and crabs infected wtih thr amoeba, Pmon~oebn perniciosa. J . Inuert. Path. 15, 440-6.

168. Schiiperclaus. W. 1954. Fischkrankheiten, 3 Auflagc. .4kadc*mie-Verlag, Berlin. xii + 708 pp.

169. Schaudinn, F. 1896. Der Zeugungskreis von Paramoeba eilhardi 11. g.. n. sp. S i t z u n g ~ l e r . Kgl. Preusr. Aknd. d . Wirc. Berlin, 25-35, (Reprinted in Srhaudinn’s Arbeiten, 191 1, 31-41.)

170. Srhereschensky, H. 1926. Mikrosporidien als Erreger riner Muskelerkrankung der Flusskrrbse. Zoo/. Anr. 65, 69-74.

171. Schnrider, .4. 1875. Contribution a I’historic des greg-arincs des invcrtrhres de Paris et de Roscoff. Arch. Zool. E x p . Gin. 1, 493-604.


172. - -. 1892. Signalement d’un nouveau sporozoaire. Tabl . Zool. 2, 209-10.

173. Setna, S. B. & Bhatia, B. L. 1934. O n some gregarines from the prawn, Parapeneopsis sculptilis (Heller) . Parasitology 26, 14.61 “. .”.

174. Smith, G. 1905. Note on a gregarine (Aggregata inachi, n. sp.) which may cause the parasitic castration of its host ( Inachus dorsettensis). Mit t . Zool. Stat. Neapel. 17, 406-10.

175. Sogandares-Bernal, F. 1962. Presumable microsporidiosis in the dwarf crayfishes Cambarellus puer Hobbs and C. shufeldti (Faxon) in Louisiana. J . Parasit. 48, 493.

176. Sprague, V. 1949. Species of Nematopsis in Ostrea uir- ginica. Ibid 35(6/2) , 42 [Abstr. 1081.

177. 1950. Notes on three microsporidian parasites of dccapod Crustacea of Louisiana coastal waters. Occ. Pap. Mar. Lab. La. St. Uniu. 5, 1-8.

178. 1950. Thelohania cambari n. sp., a microsporidian parasite of North American crayfish. J . Parasit. 36, 46.

179. __ . 1954. Protozoa. In P. S. Galtsoff, ed. Gulf of Mexico. Its Origin, Waters and Marine Life, U. S . Fish Wildl. Serv. Fish. Bull. 55(89) , 243-56.

180. Sprague, V. & Orr, P. F., Jr. 1955. Nematopsis ostrearum and N . prytherchi (Eugregarinina: Porosporidae) with special reference to the host-parasite relations. J . Parasit. 41, 89-1 04.

181. Sprague, V. 1963. Minchinia louisiana n. sp. (Haplo- sporidia, Haplosporidiidae) , a parasite of Panopeus herbstii. J . Protozoal. 10, 267-74.

182. __ . 1965. Nosema sp. (Microsporida, Nosematidae) in the musculature of the crab Callinectes sapidus. Ibid 12, 66-70.

183. __ . 1966. Two new species of Plistophora (Micro- sporida, Nosematidae) in decapods, with particular reference to one in the blue crab. Ibid 13, 196-9.

184. Sprague, V. & Beckett, R. L. 1966. A disease of blue crabs (Callinectes sapidus) in Maryland and Virginia. J . Inuert. Path. 8 , 287-9.

185. ~ , 1958. The nature of the etioloeical aeent of - “Gray Crab” discasc. Ibid 11. 503.

186. Sprague, V. & Vernick, S. H . 1968. Observations on the spores of Pleistophora gigantea (Thdohan, 1895) Swellengrebel, 1911, a microsporidan parasite of the fish Crenilabrus melops.

187. Sprague, V., Vernick, S. H. & Lloyd, B. J., Jr. 1968. The fine structure of Nosema sp. Sprague, 1965 (Microsporida, Nose- matidae) with particular reference to stages in sporogony. J . I n u e r t . Path. 12, 105-17.

188. Sprague, V. 1969. Need for drastic revision of subphylum .4moebogena. In A. A. Strelkov, K . M. Sukhanova & I. B. Raikov, cds. Proeress in Protozoolo~~~--Srd Intern. Coner. Protorool..

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190. Sprague, V., Beckett, R. L. & Sawyer, T. K. 1969. A new species of Paramoeba (Amoebida, Paramoebidae) parasitic in the crab Callinectes sapidus. J . Inuert. Path. 14, 167-74.

191. Sprague, V. 1970. Some protozoan parasites and hyperparasites in marine decapod Crustacea. In S. F. Snieszko, ed. A Symposium on Diseases of Fishes and Shellfishes, Spec. Puhl. 5, Am. Fish. Soc., Washington, D. C. 416-30..

192. ___ , 1971. Plistophora Labbe, 1899, IS a junior synonym of Pleistophora Gurley, 1893, and should be discarded. J . Inuert. Path. 17, 1-2.

193. Stanier, J. E., Woodhouse, M. A. & Griffin, R . L. 1968. The fine strurture of spore of a Nosema spelotremae, a microsporidian parasite of a Sfielotrema metacercaria encysted in the crab Carcinus maenas. J . Invert. Path. 12, 73-82.

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1964. Gregarines d’invertebrks marins de la region de Banyuls 11. Deux nouvelles eugrkgarines parasites de Solenocera membranacea (Risso, 1816) (Decapoda Penaiedae) et remarques sur les gregarines de Penaeidae. Vol. Jubil. Prof. G . Petit, Vie Milieu 17 (Suppl.), 115-27.

203. ___ . 1965. Gregarines d’invertCbrCs marins de la region de Banyuls 111. Porospora soyeri n. sp. (Eugregarina, Poro- sporidae) parasite de Aristeus antennotus (Risso, 1816) (De- ca:ioda, Penaeidae). Vie Milieu 16, 537-46.

204. ___ . 1965. Parasitisme de decapodes Natantia de Banyuls par Aggregata leandri Pixell-Goodrich, 1950 (Coccidia Aggregatidae). Ibid 16, 229-30.

205.---. 1967. Gregarines d’invertCbres marins de la region de Banyuls IV. Quelques eugregarines du genre Cephaloidophora parasites de crustaces amphipodes et decapodes. Ibid 18, 265-79.

206. TheodoridPs, J. & Laird, M. 1970. Quelques eugregarines parasites d’invertbbres marins de St. Andrews (Nouveau-Bruns- wick). Can. J . Zool . 48, 1013-6.

207. Trager, W. 1957. Excystation of apostome ciliates in re- lation to molting of their crustacran hosts. B i d . Bull. 112, 132-6.

208. Trhgouboff, G. 1912. Sur les gregarines des balanes. Arch. Zool. E x p . Gin. N & R Ser. 5, 10, 53-61.

209. -- -. 1916. Sur quelques protistes parasites rencontres B Villefranche-sur-Mer. Ibid 55: 35-47.

210. Tuzet, 0. & OrmiPres, R. 1961. Sur quelques gregarines parasites de CrustacCs Decapod&. Ann. Sci. N a t . Zool. Ser. 12, 3,

211. 1962. Gregarines de Roscoff. Cahiers B i d . Mar. 3, 289-306.

2 1 2 . 1962. A propos du moyau protomkritique des grCgarines. Ibid 3, 209-1 1.

213. VivarPs, C. P. & Rubi6, M. 1969. Protozoa parasites de Crustacea Decapoda Brachyura de la cBte nord-est de l’Espagne. Publ. Inst. Biol. Apl. 4, 111-29.

214. Vivark, C . P. 1970. Parasites de crustaces decapodes brachyoures du Golfe et du Lac de Tunis. Bull. Inst. Ocdanog. PSche, SalammbB 1 ( 4 ) , 181-203.

215. Wallengrcn. H. 1895. Idem. 11. Slagtet Heliochona Plate S13gtc.t Chilodochona n. g. SIiiStet Hemispeira Fabre-Domerque. Lunds Uniu. Arsskr. 31, 1-72.

216. Watson, M. E. 1916. Three new gregarines from marine Crustacea. J . Pnrosit. 2. 129-36.

217. Woodhurn. K. D., Elder, B., Clark, E., Hutton, R. F. & Ingle, R. M . 1957. The live bait shrimp industry of the west coast of Florida (Cedar Key to Naples). Flu. St. Bd. Conserv. Tech. Ser. 21, 33 pp.

218. Wright. S. 1858. Description of new Protozoa. Edinburgh New Philos. J . 7 , 276-81.

201.

202. -

773-83.

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