The classification of the Protozoa follows that of Kudo (1954). The use of the asterisk (*) is explained in footnote 3, page 4.

Phylum PROTOZOA

Class MASTIGOPHORA

Order EUGLENOIDINA

Family EUGLENIDAE

Euglenasp.

Experimental host.—Periplaneta americana, U.S.A. (Hegner, 1929): When fed to the insects in concentrated culture, Euglena could withstand conditions in the crop up to 5 hours and were passed into the stomach in a viable state up to 6 hours. However, the majority were killed in the crop within 2 hours and very few reached the stomach alive.

Order PROTOMONADINA

Family OIKOMONADIDAE

Oikomonas blattarum Tejera

Natural host.—Cockroach, Venezuela (Tejera, 1926).

Oikomonassp.

Natural host.—Blatta orientalis, U.S.S.R. (Yakimov and Miller, 1922): Oikomonas sp. and Monas sp. were found in the intestines of 83 percent of 124 cockroaches.

Cockroach, Venezuela (Tejera, 1926).

Family TRYPANOSOMATIDAE

Leptomonas blaberaeTejera

Natural host.—Blaberus sp., Venezuela (Tejera, 1926).

Leptomonassp.

Natural hosts.—Parcoblatta lata, Parcoblatta pensylvanica, Parcoblatta virginica, U.S.A., Ohio (Semans, 1939, 1941): Hind intestine. Of 70 specimens examined, 86 percent harbored Leptomonas sp.

* Herpetomonas periplanetaeLaveran and Franchini

Natural host.—Blatta orientalis, Italy, France (Laveran and Franchini, 1920, 1920a).

Family MONADIDAE

Monassp.

Natural host.—Blatta orientalis, U.S.S.R. (Yakimov and Miller, 1922): Monas sp. and Oikomonas sp. were found in the intestines of 83 percent of 124 cockroaches examined.

Cockroach, Venezuela (Tejera, 1926).

Family BODONIDAE

Bodo blattae

Natural host.—Blatta orientalis, England (Lankester, 1865).

Bodosp.

Natural host.—Blattella germanica and/or Periplaneta americana, South Africa (Porter, 1930).

Retortamonas blattae(Bishop)

Synonymy.—Embadomonas blattae Bishop [Wenrich, 1932].

Natural host.—Blatta orientalis, England (Bishop, 1931): Hind intestine. The organism occurred in about 40 percent of the cockroaches examined. L. G. Feo (in Wenrich, 1932) successfully cultured this protozoan (fig. 2, F).

Retortamonassp.?

Natural host.—Leucophaea maderae, Philippine Islands (Hegner and Chu, 1930).

---

Order POLYMASTIGINA

Family CHILOMASTIGIDAE

Chilomastix mesnili(Wenyon)

Experimental vectors.—Blatta orientalis and Periplaneta americana, South Africa (Porter, 1918): The cockroaches were fed human excrement that contained cysts of C. mesnili. The cysts passed unharmed through the insects' digestive tract. Rats became infected with this protozoan on eating food that had been contaminated with feces from these cockroaches.

Family POLYMASTIGIDAE

Eutrichomastixsp.

Synonymy.—Trichomastix [Kudo, 1954].

Natural host.—Blattella germanica and/or Periplaneta americana, South Africa (Porter, 1930).

Monocercomonoides globusCleveland et al.

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934): Organism occurs in practically all hosts.

Monocercomonoides melolonthae(Grassi)

(Fig. 2, A)

Natural host.—Platyzosteria novae seelandiae, New Zealand (Laird, 1956): Found in the intestinal tracts of the adult cockroaches, and of other species of insects.

Monocercomonoides orthopterorum(Parisi)

Synonymy.—Trichomonas (Trichomastix) orthopterorum Parisi; Monocercomonas orthopterorum [Belar, 1916]; Trichomastic orthopterum? [Zasukhin, 1930]; Monocercomonoides orthopterorum [Travis, 1932; Cleveland et al., 1934]; Retortamonas orthopterorum [Semans, 1943].

Natural hosts.—Blatta orientalis, Italy (Parisi, 1910); U.S.S.R. (Zasukhin, 1930).

Ectobius lapponicus, Italy (Parisi, 1910).

Periplaneta americana, Philippine Islands (Hegner and Chu, 1930).

"KÜchenschaben," Austria (Belar, 1916).

The protozoan is found in the hind gut. Zasukhin (1930) found the organism in 85 percent of over 3,000 B. orientalis. Parisi (1910) found the flagellate present in very large numbers.

Monocercomonoides panesthiaeKidder

Natural host.—Panesthia angustipennis, Philippine Islands (Kidder, 1937): In hind gut.

Tetratrichomastix blattidarumYoung

Natural hosts.—Blatta orientalis, Blattella germanica, Periplaneta americana, U.S.A. (Young, 1935): The organism when present occurs in large numbers in the posterior part of the intestine near the anus. The protozoan was successfully cultivated in a hemoglobin-saline medium.

Family OXYMONADIDAE

Oxymonas doroaxostylus(Cleveland et al.)

Synonymy.—Saccinobaculus doroaxostylus Cleveland et al. [Cleveland, 1950].

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934).

Oxymonas nanaCleveland

Synonymy.—Saccinobaculus minor Cleveland et al. [Cleveland, 1950].

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934).

Family DINENYMPHIDAE

Saccinobaculus ambloaxostylusCleveland et al.

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934).

Saccinobaculus lataCleveland

Natural host.—Cryptocercus punctulatus, U.S.A. (Cleveland, 1950b): There are at least two other species of Saccinobaculus in C. punctulatus that have not been described.

Notila proteusCleveland

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland, 1950b).

Family TRICHOMONADIDAE

* Trichomonas hominis(Davaine)

Experimental vectors.—Blatta orientalis, South Africa (Porter, 1918); Italy (Mariani and Besta, 1936).

Periplaneta americana, South Africa (Porter, 1918); U.S.A. (Hegner, 1928).

Trichomonassp.

Natural vector.—Cockroach, Venezuela (Tejera, 1926): Organism found in digestive tract of the cockroach.

Family HEXAMITIDAE

Hexamita cryptocerciCleveland et al.

Natural hosts.—Cryptocercus punctulatus, U.S.A. (Cleveland et al., 1934).

Panesthia angustipennis, Philippine Islands (Kidder, 1937).

Hexamita periplanetae(Belar)

Synonymy.—Octomitus periplanetae Belar [Kudo, 1954].

Natural hosts.—Blatta orientalis, U.S.S.R. (Zasukhin, 1930): Organism is found in the hind gut. Eighty-five percent of over 3,000 B. orientalis contained this organism.

Periplaneta americana, Philippine Islands (Hegner and Chu, 1930).

"KÜchenschaben," Austria (Belar, 1916).

Hexamitasp.?

Natural host.—Leucophaea maderae, Philippine Islands (Hegner and Chu, 1930): The flagellates were present in large numbers.

* Giardia intestinalis(Lambl)

Experimental vectors.—Blatta orientalis, South Africa (Porter, 1918).

Blattella germanica, Brazil (PessÔa and CorrÊa, 1927).

Eurycotis floridana, U.S.A. (Young, 1937).

Leucophaea maderae, Brazil (PessÔa and CorrÊa, 1927).

Periplaneta americana, South Africa (Porter, 1918); Gold Coast Colony (Macfie, 1922); Brazil (PessÔa and CorrÊa, 1927); U.S.A. (Young, 1937).

Periplaneta brunnea, U.S.A. (Young, 1937).

Cockroaches, Venezuela (Tejera, 1926); Argentina (Bacigalupo, in Tejera, 1926).

* Giardiasp.

Natural vectors.—Cockroaches, Venezuela (Tejera, 1926).

Order HYPERMASTIGINA

Family HOLOMASTIGOTIDAE

Leptospironympha euporaCleveland et al.

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian area (Cleveland et al., 1934).

Leptospironympha rudisCleveland et al.

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian area (Cleveland et al., 1934).

Leptospironympha wachulaCleveland et al.

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian area (Cleveland et al., 1934).

Macrospironympha xyloplethaCleveland et al.

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian area (Cleveland et al., 1934).

Family LOPHOMONADIDAE

Lophomonas blattarumStein

Natural hosts.—Blatta orientalis, Czechoslovakia (Stein, 1860); Germany (BÜtschli, 1878; Schubotz, 1905; Chen, 1933); U.S.A. (Leidy, 1879a; Kudo, 1922, 1925, 1926, 1926b; McAdow, 1931); Europe (Janicki, 1908); U.S.S.R. (Yakimov and Miller, 1922; Zasukhin, 1930); Poland (Lorenc, 1939).

Blattella germanica, U.S.A., Ohio (McAdow, 1931).

Blattella germanica and/or Periplaneta americana, Egypt (DeCoursey and Otto, 1956, 1957).

Periplaneta americana, England (Schuster, 1898); Europe (Janicki, 1910); U.S.A. (Kudo, 1926b; McAdow, 1931; Hatcher, 1939; Armer, 1944); Philippine Islands (Hegner and Chu, 1930).

Periplaneta sp., Goa (Mello and Lima Ribeiro, 1924, 1925).

"KÜchenschaben," Austria (Belar, 1916).

The protozoan (fig. 2, E) is found in the host's colon, particularly anterior portion; encysted stages of organism are found throughout hind gut. Of 1,400 B. orientalis studied, 32 percent harbored this organism (Kudo, 1925, 1926). Yakimov and Miller (1922) found 7 percent of 124 B. orientalis infested. Zasukhin (1930) found 10 percent of over 3,000 B. orientalis infested. The flagellate does not harm the host and is never present in the host tissue; it should be considered a commensal (Kudo, 1926).

Lophomonas striataBÜtschli

Synonymy.—Lophomonas sulcata Schuster is most probably identical with L. striata (Kudo, 1926b).

Natural hosts.—Blatta orientalis, Germany (BÜtschli, 1878; Schubotz, 1905); Europe (Janicki, 1908, 1910); U.S.A. (Kudo, 1922, 1926, 1926b; McAdow, 1931); U.S.S.R. (Yakimov and Miller, 1922; Zasukhin, 1930); Poland (Lorenc, 1939).

Blattella germanica, U.S.A., Ohio (McAdow, 1931).

Blattella germanica and/or Periplaneta americana, South Africa (Porter, 1930).

Periplaneta americana, Indochina (Weill, 1929); Philippine Islands (Hegner and Chu, 1930); U.S.A. (Kudo, 1926b; McAdow, 1931; Armer, 1944).

Cockroach, Venezuela (Tejera, 1926); England or U.S.A.? (Lucas, 1928).

"KÜchenschaben," Austria (Belar, 1916).

Found in the host's colon, particularly the anterior portion. L. striata (fig. 2, D) was found in 29 percent of 1,400 B. orientalis and in 2 of 30 P. americana (Kudo, 1926, 1926b). Yakimov and Miller (1922) found the organism in 9.6 percent of 124 specimens of B. orientalis. Zasukhin (1930) found 8.6 percent of over 3,000 B. orientalis infested.

GrassÉ (1926, 1926a) identified corrugations on the surface of L. striata as a bacterial parasite which he named Fusiformis lophomonadis.

Prolophomonas tocopolaCleveland et al.

Natural host.—Cryptocercus punctulatus, California, Oregon, Virginia, West Virginia (Cleveland et al., 1934): Not abundant.

Family HOPLONYMPHIDAE

Barbulanympha estabogaCleveland et al.

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934).

Barbulanympha coahoma (Cleveland et al., 1934) represents the diploid form of B. estaboga (Cleveland, 1953).

Barbulanympha laurabudaCleveland et al.

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934): This species, B. ufalula, and Rhynchonympha tarda occur in all parts of the colon, especially in the enlarged, flexed part near the ileum.

Barbulanympha ufalulaCleveland et al.

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934).

Barbulanympha wenyoniCleveland

Natural host.—Cryptocercus punctulatus, U.S.A., Pacific coast area (Cleveland, 1953).

Rhynchonympha tardaCleveland et al.

(Fig. 3, D)

Natural host.—Cryptocercus punctulatus, U.S.A., Pacific coast area (Cleveland et al., 1934): Fairly abundant in every specimen examined from Pacific coast.

Urinympha taleaCleveland et al.

(Fig. 3, C)

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934): Present in fairly great numbers in every cockroach examined.

Family STAUROJOENINIDAE

Idionympha perissaCleveland et al.

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian area (Cleveland et al., 1934): Present in only a few specimens.

Family TRICHONYMPHIDAE

Trichonympha acutaCleveland et al.

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934).

Trichonympha algoaCleveland et al.

(Fig. 3, E)

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934): Fairly abundant and present in most specimens.

---

Trichonympha chulaCleveland et al.

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934).

Trichonympha grandisCleveland et al.

Natural host.—Cryptocercus punctulatus, U.S.A., Pacific coast areas (Cleveland et al., 1934): Fairly abundant in all specimens from Pacific area.

Trichonympha lataCleveland et al.

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934).

Trichonympha okolonaCleveland et al.

(Fig. 3, E)

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934): Found in only a few specimens, never abundant.

Trichonympha parvaCleveland et al.

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934): This organism is smaller than any known species of Trichonympha; it is more resistant to warm weather than the other hypermastigotes.

Family EUCOMONYMPHIDAE

Eucomonympha imlaCleveland et al.

Natural host.—Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934): Organism (fig. 3, A) sometimes becomes attached to the intestinal wall; attached individuals were seen in 2 to 3 percent of the cockroaches examined.

Unidentified flagellate

Natural host.—Pycnoscelus surinamensis, Hawaii (Schwabe, 1950): A small flagellate was found in the digestive tract and malpighian tubules.

Class SARCODINA

Order MYCETOZOA

INCERTAE SEDIS

Peltomyces periplanetae(LÉger)

Synonymy.—Peltomyces blattellae. Sprague (1940a) synonymizes Peltomyces periplanetae, with Coelosporidium periplanetae.

Natural hosts.—Blatta orientalis, France (Debaisieux, 1927).

Blattella germanica, France (LÉger, 1909; Debaisieux, 1927).

The organism inhabits the malpighian tubules of cockroaches. LÉger and Debaisieux concluded that their organism was a mycetozoan, but they may have erred in synonymizing Plistophora periplanetae with the organism they studied. Debaisieux found intracellular stages of Peltomyces periplanetae that have not been found in Plistophora periplanetae or Coelosporidium periplanetae.

Order AMOEBINA

Family AMOEBIDAE

Hartmannella blattaeIvanic

Natural host.—Blatta orientalis, Yugoslavia (Ivanic, 1937): Found in the hind gut.

Family ENDAMOEBIDAE

In the following classification we have accepted the conclusions of Kirby (1945), Kudo (1954), and others that species of Endamoeba are generically different from species of Entamoeba and that the latter genus is not a homonym of Endamoeba.

Dobellinasp.

Natural vectors.—Blattella germanica and/or Periplaneta americana, Egypt (DeCoursey and Otto, 1956, 1957): Thirty out of 261 cockroaches examined contained this protozoan.

Endamoeba blattae(BÜtschli)

Synonymy.—Amoeba blattae, Entamoeba blattae, Entamoeba blattarum.

Natural hosts.—Blatta orientalis, Germany (BÜtschli, 1878; Schubotz, 1905; Chen, 1933); U.S.A. (Leidy, 1879a, 1880; Kudo, 1922, 1925a, 1926a; Kirby, 1927; McAdow, 1931; Meglitsch, 1938, 1940); France (Mercier, 1907a, 1908, 1909, 1910); Europe? (Janicki, 1908, 1909); U.S.S.R. (Yakimov and Miller, 1922; Zasukhin, 1929, 1930); England (Thomson and Lucas, 1926; Lucas, 1927, 1927a, 1928); Yugoslavia (Ivanic, 1926a).

Blattella germanica and/or Periplaneta americana, South Africa (Porter, 1930); Egypt (DeCoursey and Otto, 1956, 1957): Seven out of 217 cockroaches examined harbored the protozoan.

Periplaneta americana, Philippine Islands (Hegner and Chu, 1930); U.S.A. (Morris, 1936; Armer, 1944); Gold Coast Colony (Macfie, 1922).

Periplaneta australasiae, U.S.A. (Morris, 1936).

Cockroaches, Paraguay? (Elmassian, 1909); Austria (Belar, 1916); U.S.A. (Morris, 1935, 1936; Balch, 1932); Venezuela (Tejera, 1926).

The habitat of E. blattae (fig. 2, C) is the hind intestine and rectum of the cockroach. The incidence of infection varies: Kudo (1925a) found in 1,255 oriental cockroaches infections in 5 percent in March and 50 percent in the summer; Schubotz (1905) found 5 to 20 percent of the examined cockroaches to be infested; Yakimov and Miller (1922) found 4 percent of 124 oriental cockroaches infested; Zasukhin (1930) found up to 50 percent of over 3,000 B. orientalis infested; Meglitsch (1938, 1940) found almost 100 percent infection in B. orientalis kept in a crowded culture for several weeks. Chen (1933) developed two synthetic media in which E. blattae could be grown for 45 to 50 days.

Mercier (1907a) observed a fungus, Nucleophaga sp., hyperparasitic in the nucleus of Endamoeba blattae.

Endamoeba javanicaKidder

Natural hosts.—Panesthia angustipennis, Philippine Islands, and Panesthia spadica, Japan (Kidder, 1937): Occurred in 50 percent of P. angustipennis examined and in one of four P. spadica. The endoplasm of this amoeba contains large amounts of wood and cellulose fibers.

Endamoeba philippinensisKidder

Natural host.—Panesthia angustipennis, Philippine Islands (Kidder, 1937): Occurred in about 10 percent of the Panesthia examined. The food vacuoles contained bacteria, no wood.

Entamoeba coli(Grassi)

Synonymy.—Endamoeba coli, Amoeba coli [Kirby, 1945].

Natural vectors.—Blaberus atropos, Venezuela (Tejera, 1926): In a lot of 60 cockroaches captured in latrines, two were found that carried apparently live cysts similar to cysts of E. coli.

Blattella germanica or Periplaneta americana, Egypt (DeCoursey and Otto, 1956, 1957): One out of 44 cockroaches collected in a village harbored E. coli.

Experimental vector.—Periplaneta americana, Gold Coast Colony (Macfie, 1922): In nine experiments cysts of E. coli were fed to the cockroaches. In seven of the experiments cysts of E. coli were found in the feces. Cysts were observed in the feces for only one to three days, and eventually disappeared completely. The cysts appeared to be unharmed. No amoebae were found.

* Entamoeba histolyticaSchaudinn

Natural vectors.—Blatta orientalis, Blattella germanica, Periplaneta americana, Periplaneta australasiae, and/or Supella supellectilium, Peru (Schneider and Shields, 1947).

Blattella germanica and/or Periplaneta americana, Egypt (DeCoursey and Otto, 1956, 1957).

Cockroaches, Venezuela (Tejera, 1926).

Experimental vectors.—Blatta orientalis, Italian Somaliland (Mariani and Besta, 1936).

Periplaneta americana, Gold Coast Colony (Macfie, 1922); U.S.A. (Frye and Meleney, 1936).

Cockroaches, Venezuela (Tejera, 1926).

Entamoeba pitheciProwazek?

Experimental vector.—Periplaneta americana, Formosa (Morischita and Tsuchimochi, 1926): Eleven of 15 cockroaches fed feces of a monkey [Macaca cyclopis (Swinhoe)] containing cysts of the amoeba voided live cysts in their own feces.

Entamoeba thomsoniLucas

Synonymy.—Endamoeba thomsoni [Kudo, personal communication, 1957].

Natural hosts.—Blatta orientalis, England (Lucas, 1927a, 1928); U.S.A. (Taliaferro, 1928; McAdow, 1931); U.S.S.R. (Zasukhin, 1930); Germany (Chen, 1933).

Blattella germanica, U.S.A. (McAdow, 1931).

Periplaneta americana, England (Lucas, 1927a); U.S.A. (Smith and Barret, 1928; McAdow, 1931); Philippine Islands (Hegner and Chu, 1930).

The organism is found in the hind intestine and rectum of the cockroach. Smith and Barret (1928) developed a synthetic medium in which cultures of E. thomsoni were carried through successive transfers for 24 months.

Entamoebasp.

Natural vector.—Periplaneta americana, Gold Coast Colony (Macfie, 1922): Under the heading "Entamoeba histolytica and E. coli" Macfie (p. 445) stated, "The cockroaches used in these experiments had previously been carefully examined for amoebic infections a precaution which was doubly necessary, because some of these insects at Accra had been found naturally infected."

Experimental vectors.—Periplaneta americana, Gold Coast Colony (Macfie, 1922): Entamoeba, resembling E. coli, from feces of the monkey [Erythrocebus patas patas (Schreber)] were fed to cockroaches, and on the second to fourth days thereafter apparently healthy cysts were recovered in the cockroach feces.

Endolimax blattaeLucas

Natural hosts.—Blatta orientalis, England (Lucas, 1927, 1927a); U.S.S.R. (Zasukhin, 1930); Germany (Chen, 1933).

Periplaneta americana, England (Lucas, 1927, 1927a); Indochina (Weill, 1929); U.S.A. (Armer, 1944).

Periplaneta australasiae, U.S.A. (Steinhaus, 1946).

Organism is found in the hind gut of the cockroach. Zasukhin (1930) found 3-percent infestation in over 3,000 B. orientalis examined.

Endolimax nana(Wenyon and O'Connor)?

Synonymy.—Entamoeba nana.

Natural host.—Blaberus atropos, Venezuela (Tejera, 1926): A small amoeba greatly resembling E. nana was found in the intestinal contents of the cockroach. In inoculations this amoeba was not pathogenic.

Endolimaxsp.

Natural hosts.—Blatta orientalis, U.S.S.R. (Zasukhin, 1930): This organism was found in the hind gut of 0.3 percent of over 3,000 cockroaches examined.

Blattella germanica and/or Periplaneta americana, Egypt (DeCoursey and Otto, 1956, 1957): Seventy-four out of 261 cockroaches examined harbored this protozoan.

Iodamoebasp.

Natural vectors.—Blattella garmanica and/or Periplaneta americana, Egypt (DeCoursey and Otto, 1956, 1957): Fifty-nine of 261 cockroaches examined contained this protozoan. Iodamoeba sp. was common in human feces in villages in which the cockroaches were collected.

Undetermined species of Amoeba

Natural host.—Panesthia angustipennis, Philippine Islands. (Kidder, 1937): Found in only one specimen.

Class SPOROZOA

Order GREGARINIDA

Family DIPLOCYSTIDAE

Diplocystis schneideriKunstler

(Fig. 2, I)

Natural hosts.—Blatta orientalis, England (Woodcock, 1904; Jameson, 1920).

Periplaneta americana,France (Kunstler, 1884, 1887); England (Jameson, 1920); Germany (Foerster, 1939).

In body cavity of host. Cysts containing spores are ingested during cannibalistic feeding on infected cockroaches. Sporozoites penetrate the gut wall which later ruptures, freeing the gregarines into the coelom. There is no apparent pathogenic effect. Jameson (1920) found 81 percent of P. americana infested with D. schneideri.

Diplocystissp.

Natural host.—Periplaneta americana, U.S.A. (Hertig, 1921): Heavy infections in body cavity.

Cockroach, India (Ray and Dasgupta, 1955): A large number of cockroaches, both adults and nymphs, collected in Calcutta were all infected.

Diplocystissp.?

Natural host.—Blaberus craniifer, U.S.A. (Nutting, 1953): From 1 to 12 or more paired trophozoites or cysts may be found in the hemocoele and occasionally in the thorax.

Family STENOPHORIDAE

Stenophorasp.

Natural host.—Blatta orientalis, India (Bal and Rai, 1955): Organism found in the midgut of the cockroach.

Family GREGARINIDAE

Gregarina blattarumvon Siebold

Synonymy.—Gregarina blattae orientalis; Clepsidrina blattarum.

Natural hosts.—Blatta orientalis, Germany (Siebold, 1837, 1839; Stein, 1848; BÜtschli, 1881; Wolters, 1891; Marshall, 1892; Wellmer, 1910, 1911; Foerster, 1938; Schubotz, 1905); U.S.A. (Leidy, 1853a; Crawley, 1903; Watson, 1917; Kudo, 1922; McAdow, 1931; Sprague, 1940, 1941); England (Lankester, 1863); France (Schneider, 1875; CuÉnot, 1901; Laveran and Franchini, 1920a); Brazil (MagalhÃes, 1900; Pinto, 1919); U.S.S.R. (Zasukhin, 1929, 1930).

Blattella germanica, U.S.A. (Crawley, 1903); South Africa (Fantham, 1929; Porter, 1930: these appear to be the same record).

Blattella germanica and/or Periplaneta americana, Egypt (DeCoursey and Otto, 1956, 1957).

Periplaneta americana, Brazil (MagalhÃes, 1900); U.S.A. (Crawley, 1903; 1907; McAdow, 1931); South Africa (Fantham, 1929; Porter, 1930: these appear to be the same record); Gold Coast Colony (Macfie, 1922).

Parcoblatta pensylvanica, U.S.A., Michigan (Ellis, 1913a).

Cockroaches, Germany (Schiffmann, 1919: probably used the oriental cockroach); Venezuela (Tejera, 1926).

Organism usually found in the intestinal tract of cockroaches where it is attached to the gut cells. Cysts are passed in the feces. Occasionally, G. blattarum (fig. 2, J) is found in the body cavity (Leidy, 1853a; Hall, 1907). Though this is considered to be one of the commonest of the Sporozoa encountered in cockroaches, DeCoursey and Otto (1956) found only 10 of 217 P. americana and B. germanica, collected in restaurants in Egypt, infested with this species. Watson (1917) found a dozen or more in one specimen of Blatta orientalis. Zasukhin (1929, 1930) found 2.6 percent of 3,000 oriental cockroaches infected with this parasite.

Gregarina fastidiosaHarrison

Natural host.—Aptera fusca, South Africa (Harrison, 1955): All mature females were heavily infected; in all specimens there were over 100 parasites in the gut. All nymphs were infected, the earlier instars more lightly than the later instars. Gregarines were found in all parts of the gut except the crop and gizzard.

Gregarina gibbsiHarrison

Natural host.—Temnopteryx phalerata, South Africa (Harrison, 1955): Although the cockroaches were found in groups, only 32 percent were infected and only 10 percent heavily. The gregarines were found in the anterior mesenteron but none in the hepatic caeca. All cysts were found in the hind gut or rectum.

Gregarina illinensisM. E. Watson

Natural host.—Parcoblatta pensylvanica, U.S.A., Illinois (Watson, 1915, 1916): The intestine of one cockroach was found to contain 25 of these gregarines.

Gregarina impetuosaHarrison

Natural host.—Melanosilpha capensis, South Africa (Harrison, 1955): All specimens of this gregarine were found in the anterior mesenteron of the host.

Gregarina lÉgeriPinto

Natural host.—Periplaneta americana, Brazil (Pinto, 1918, 1918a, 1919): Intestinal canal.

Gregarina neo-brasiliensisAl. Cunha

Natural host.—Periplaneta americana, Brazil (R. de Almeida Cunha in Pinto, 1919; Cunha, 1919).

Gregarina ohioensisSemans

Natural host.—Parcoblatta virginica, U.S.A., Ohio (Semans, 1939): The protozoan was present in large numbers in the insect's midgut.

Gregarina panchloraeFrenzel

Natural host.—Panchlora exoleta, Argentina (Frenzel, 1892): Midgut.

Gregarina parcoblattaeSemans

Natural hosts.—Parcoblatta pensylvanica and Parcoblatta uhleriana, U.S.A., Ohio (Semans, 1939): Midgut.

Gregarina rhyparobiaeJ. M. Watson

Natural host.—Leucophaea maderae, Uganda (Watson, 1945): Midgut. Trophozoites could be seen in sections attached to cells of the intestinal wall (fig. 2, H).

Gregarina sandoniHarrison

Natural host.—Melanosilpha capensis, South Africa (Harrison, 1955): This gregarine was found in the anterior and middle parts of the mesenteron and in the hepatic caeca.

Gregarina thomasiSemans

Natural host.—Parcoblatta pensylvanica, U.S.A., Ohio (Semans, 1939): Enteric caeca and midgut.

Protomagalhaesia serpentula(de MagalhÃes)

Synonymy.—Gregarina serpentula [Pinto, 1918a, 1919; Semans, 1943].

Natural host.—Periplaneta americana, Brazil (MagalhÃes, 1900): In the coelom and alimentary canal. The host of this parasite (fig. 2, K) was incorrectly cited as Blatta orientalis by Watson (1916).

Gamocystis tenaxSchneider

(Fig. 2, L)

Natural hosts.—Ectobius lapponicus, France (Schneider, 1875); Germany (Wellmer, 1910, 1911; Foerster, 1938).

Ectobius pallidus, Germany (Foerster, 1938): In intestine.

Family ACTINOCEPHALIDAE

Pileocephalus blaberae(Frenzel)

Synonymy.—Gregarina blaberae [Watson, 1916].

Natural hosts.—Blaptica dubia and related forms, Argentina (Frenzel, 1892): In midgut.

Unidentified Gregarinida

Natural hosts.—Blaberus craniifer, U.S.A. (Roth and Willis, unpublished data, 1953): Possibly Diplocystis sp. (pl. 28, A, B).

Cryptocercus punctulatus, U.S.A. (Cleveland et al., 1934).

Leucophaea maderae, Philippine Islands (Hegner and Chu, 1930): In intestines of host. U.S.A. (Roth and Willis, unpublished data, 1958): Cysts in feces (pl. 28, C).

Gromphadorhina portentosa, U.S.A., in laboratory colony (Roth and Willis, unpublished data, 1958): In intestine of adult female.

Pycnoscelus surinamensis, Hawaii (Schwabe, 1950): A cephaline gregarine was found in the cockroach's digestive tract; it was also claimed to be present in new-born nymphs.

Order COCCIDIA

Family ADELEIDAE

Adelina cryptocerciYarwood

Natural host.—Cryptocercus punctulatus, U.S.A. (Yarwood, 1937): This intracellular parasite was found in the fat body in light infestations. In heavy infections the coccidia were found in the head, antennae, mouthparts, muscles, legs, salivary glands, nerve cord, as well as fat body. Infection in freshly collected specimens was about 3 percent; when large numbers of cockroaches were kept together in culture, the rate of infection increased because the insects ate their dead companions.

Cleveland et al. (1934) mentioned a coccidium which was sometimes generally distributed through the body (head, legs, antennae, etc.) of C. punctulatus; this parasite was probably the species described by Yarwood.

Order HAPLOSPORIDIA

Haplosporidium periplanetaeGeorgÉvitch

Natural host.—Blatta orientalis, Yugoslavia (GeorgÉvitch, 1953): This organism was described from the malpighian tubules of the cockroach where it apparently occurred in a mixed infection with the microsporidian Plistophora periplanetae. See synonymy under Plistophora periplanetae.

Coelosporidium periplanetae(Lutz and Splendore)

Synonymy.—Nosema periplanetae, Coelosporidium blattellae, Bertramia blatellae [after Semans, 1943]. Some of the observations cited under Plistophora periplanetae may pertain to C. periplanetae (see Sprague, 1940). See also Haplosporidium periplanetae.

Natural hosts.—Blatta orientalis, U.S.S.R. (Epshtein, 1911); U.S.A. (Kudo, 1922; Sprague, 1940); Yugoslavia (Ivanic, 1926).

Blattella germanica, U.S.A. (Crawley, 1905); Germany (Wellmer, 1910, 1911).

Periplaneta americana, Brazil (Lutz and Splendore, 1903).

This organism (fig. 2, B) passes its life cycle living free in the lumina of the malpighian tubules of cockroaches. The elongate trophozoite is firmly attached to the wall of the tubule as are clusters of immature spores. Mature spores are freed into the lumina of the tubules from whence they pass to the exterior. Sprague (1940) examined about 200 wild-caught B. orientalis and found them to be practically 100 percent infected.

Order MICROSPORIDIA

Family NOSEMATIDAE

Plistophora kudoiSprague and Ramsey

Natural host.—Blatta orientalis, U.S.A., Illinois, West Virginia, Kentucky (Sprague and Ramsey, 1941, 1942): Found in the epithelial cells of caeca and midgut. Considerable damage is done to these cells. Seventy-five percent of 52 B. orientalis harbored the parasite.

Plistophora periplanetae(Lutz and Splendore)

Synonymy.—Nosema periplanetae, Pleistophora periplanetae [after Semans, 1943]. GeorgÉvitch (1953) has pointed out that one may find in the malpighian tubules of cockroaches a mixed infection of Microsporidia, Haplosporidia, and Mycetozoa, and that some of the discrepancies in the earlier literature may be attributed to attempts to combine in one organism disparate stages belonging to different orders. See also comments under Coelosporidium periplanetae, Haplosporidium periplanetae, and Peltomyces periplanetae.

Natural hosts.—Blatta orientalis, France (Mercier, 1906a; Debaisieux, 1927); England (Perrin, 1906, 1906b); U.S.S.R. (Zhivago, 1909); Yugoslavia (GeorgÉvitch, 1925, 1926, 1926a, 1927); Germany (Wellmer, 1910, 1911).

Blattella germanica, France (LÉger, 1909; Debaisieux, 1927); U.S.S.R. (Zhivago, 1909).

Periplaneta americana, Brazil (Lutz and Splendore, 1903).

This organism lives in the lumen of the malpighian tubules of cockroaches. The cited authors appear to have been convinced that this organism was a microsporidian. GeorgÉvitch (1927, 1953) described the polar capsule and filament characteristic of this order.

Plistophorasp.

Natural host.—Blatta orientalis, France (Mercier, 1908a): The organism parasitized the fat body of the cockroach. Mitoses, often abnormal, were induced in the fat cells. Infected cockroaches were easily recognizable by their distended abdomens. The fat body became chalky white and showed through the intersegmental membranes.

Porter (1930) reported finding an unidentified microsporidian in the fat bodies of Blattella germanica and Periplaneta americana collected in South Africa. It may or may not have been a species of Plistophora.

Class CILIATA

Order HOLOTRICHA

Family PARAMECIIDAE

Parameciumsp.

Natural associate.—Cockroaches, U.S.A., Maryland (Cleveland, 1927): Three of 30 cockroaches collected in the basement of a department store had paramecia in their stomachs but none in the rectum.

Experimental associate.—Periplaneta americana, U.S.A. (Hegner, 1929): Paramecia fed to the cockroaches were recovered from the crop at intervals from one-half to six and one-half hours. In no case were the protozoa recovered from the stomach alive.

Cockroaches, U.S.A., Maryland (Cleveland, 1927). About 200 starved cockroaches were fed a culture of Paramecium. Few, if any, of the protozoa were killed in the stomach during the first two hours, but all were killed within 5 to 6 hours after ingestion.

Family ISOTRICHIDAE

Isotricha caulleryiWeill

Natural host.—Periplaneta americana, Indochina (Weill, 1929): Alimentary canal.

Order SPIROTRICHA

Family BURSARIIDAE

Balantidium blattarumGhosh

Natural host.—Periplaneta americana, India (Ghosh, 1922; Bhatia and Gulati, 1927); Gold Coast Colony (Macfie, 1922): Intestinal tract.

* Balantidium coli(Malmsten)

Experimental vector.—Cockroach, Venezuela (Tejera, 1926).

Balantidium ovatumGhosh

Natural host.—Periplaneta americana, India (Ghosh, 1922a; Bhatia and Gulati, 1927); Indochina, Saigon (Weill, 1929): Intestinal tract.

Balantidium praenucleatumKudo and Meglitsch

Natural host.—Blatta orientalis, U.S.A., Illinois (Kudo and Meglitsch, 1938; Meglitsch, 1940): This protozoan is found in the lumen of the anterior region of the colon in association with several other species of protozoa. Only 7.6 percent of 500 cockroaches examined contained B. praenucleatum. The largest number encountered in a single host was 59, but as a rule each host harbored a smaller number.

Balantidiumsp.?

Natural host.—Periplaneta americana, Brazil (MagalhÃes, 1900): These organisms were numerous in the intestine.

Family SPIROSTOMIDAE

Nyctotherus buissoniPinto

Natural host.—"Barata sylvestre," Brazil (Pinto, 1926): Organism found in the cockroach's intestine.

Nyctotherus ovalisLeidy

Synonymy.—Bursaria blattarum; Plagiotoma blattarum.

Natural hosts.—Blatta orientalis, U.S.A. (Leidy, 1850, 1853, 1853b, 1879a; Kudo, 1922, 1926, 1936; McAdow, 1931; Kudo and Meglitsch, 1938; Meglitsch, 1940); Germany (Stein, 1860; Schubotz, 1905; Chen, 1933); England (Lankester, 1865; Schuster, 1898; Lucas, 1927a, 1928); Spain (Zulueta, 1916); U.S.S.R. (Yakimov and Miller, 1922; Zasukhin, 1928, 1930; Ostroumov, 1929); Portugal (Lima Ribiero, 1924); Brazil (Pinto, 1926); Venezuela (Tejera, 1926).

Blattella germanica, South Africa (Porter, 1930); U.S.A. (Balch, 1932; McAdow, 1931).

Parcoblatta pensylvanica, U.S.A. (Semans, 1939, 1941).

Periplaneta americana, India (Bhatia and Gulati, 1927); Indochina (Weill, 1929); Philippine Islands (Hegner and Chu, 1930); South Africa (Porter, 1930); U.S.A. (McAdow, 1931; Hatcher, 1939; Meglitsch, 1940; Armer, 1944); Goa (Mello et al., 1934); China (Pai and Wang, 1947); Czechoslovakia (Low, 1956).

"Barata sylvestre," Brazil (Pinto, 1926).

"KÜchenschaben," Austria (Belar, 1916).

Nyctotherus ovalis (fig. 2, G) inhabits the hind gut of cockroaches, where it occurs almost always in the anterior half of the colon in association with other species of Protozoa (Kudo, 1936). Ninety percent of 500 B. orientalis contained N. ovalis (Kudo and Meglitsch, 1938). Yakimov and Miller (1922) found N. ovalis in 68 percent of 124 B. orientalis. Zasukhin (1930) found this organism in 63 percent of over 3,000 B. orientalis. Zasukhin (1928, 1934) found a fungus and possibly a bacterium hyperparasitic in the cytoplasm of N. ovalis. N. ovalis has been cultured outside the cockroach in several media (Lucas, 1928; Balch, 1932; Chen, 1933; Low, 1956).

Nyctotherus uichancoiKidder

Natural hosts.—Panesthia angustipennis, Philippine Islands, and Panesthia spadica, Japan (Kidder, 1937): About 90 percent of all P. angustipennis harbored this ciliate in their hindguts.

Nyctotherus viannaiPinto

Natural host.—"Barata sylvestre," Brazil (Pinto, 1926): In the intestine of the cockroach.

Family CLEVELANDELLIDAE

Most of the Clevelandellidae are parasitized by rod-shaped or spherical bacteria-like organisms usually in clusters (Kidder, 1937).

Synonymy.—Clevelandiidae (Kidder, 1938).

Genus CLEVELANDELLA

Synonymy.—The generic name Clevelandia Kidder (1937) is preoccupied; it was therefore changed to Clevelandella by Kidder in 1938. All of the following species of Clevelandella were originally described as Clevelandia.

Clevelandella constricta(Kidder)

Natural hosts.—Panesthia angustipennis, Philippine Islands, and Panesthia spadica, Japan (Kidder, 1937): In the posterior end of hindgut.

Clevelandella contorta(Kidder)

Natural hosts.—Panesthia angustipennis, Philippine Islands, and Panesthia spadica, Japan (Kidder, 1937).

Clevelandella elongata(Kidder)

Natural host.—Panesthia angustipennis, Philippine Islands (Kidder, 1937).

Clevelandella hastula(Kidder)

Natural host.—Panesthia angustipennis, Philippine Islands (Kidder, 1937): Common in hindgut.

Clevelandella nipponensis(Kidder)

Natural host.—Panesthia spadica, Japan (Kidder, 1937).

Clevelandella panesthiae(Kidder)

Natural hosts.—Panesthia angustipennis, Philippine Islands, and Panesthia spadica, Japan (Kidder, 1937): In the hindgut. This protozoan is commonly parasitized by the microorganism Sphaerita.

Clevelandella parapanesthiae(Kidder)

Natural host.—Panesthia angustipennis, Philippine Islands (Kidder, 1937).

Paraclevelandia brevisKidder

Natural hosts.—Panesthia angustipennis, Philippine Islands, and Panesthia spadica, Japan (Kidder, 1937): Present in 100 percent of P. angustipennis and in nearly all P. spadica.

Paraclevelandia simplexKidder

Natural hosts.—Panesthia angustipennis, Philippine Islands (Kidder, 1937, 1938): Incidence of infection about 50 percent.

Panesthia spadica, Japan (Kidder, 1937).

Unidentified ciliate

Natural host.—Pycnoscelus surinamensis, Hawaii (Schwabe, 1950): A large ciliate was found in the digestive tract and malpighian tubules.

NEGATIVE FINDINGS

In a recent experimental study Schmidtke (1955) failed to demonstrate a host-parasite relationship between Periplaneta americana and the haemosporidian Toxoplasma gondii Nicolle and Manceaux. This protozoan is a blood parasite in a rodent in North Africa (Kudo, 1954).

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