HELMINTHS FOR WHICH COCKROACHES SERVE AS PRIMARY HOSTS
Phylum ASCHELMINTHES
Class NEMATODA
Order MERMITHOIDEA
Family MERMITHIDAE
Undetermined mermithids
Natural Hosts.—Ectobius pallidus, U.S.A., Plymouth, Massachusetts (Roth and Willis, 1957): This mermithid lies coiled in the body cavity of the host and one end may extend into the thorax. Apparently, the host is eventually killed and the worms may leave the cockroach ventrally between the thorax and abdomen (pl. [29], A) or thorax and head.
Periplaneta americana, Germany (Bode, 1936): Attacked by "Mermis" or "Gordius." It has been suggested that the name Mermis is often applied without critical identification to immature Nematoda found in insects (Buxton, 1955).
Order RHABDITOIDEA
Family DIPLOGASTERIDAE
Diplogaster sp.
Synonymy.—Lycolaimus [Goodey, 1951].
Experimental Host.—Blattella germanica, U.S.A. (Christie and Crossman, 1933).
Family STEINERNEMATIDAE
Neoaplectana sp.
Experimental hosts.—Blattella germanica, Nauphoeta cinerea, and Periplaneta americana. U.S.A. (Dutky and Hough, 1955): This nematode, found in codling moth larvae, is close to Neoaplectana chresima Steiner but apparently is a new species. Nauphoeta cinerea was very susceptible to infection; B. germanica and P. americana were less susceptible.
Order OXYUROIDEA
Family THELASTOMATIDAE
These nematodes are found in the intestinal tract of cockroaches.
Aorurus philippinensis Chitwood and Chitwood, 1934
Natural host.—Panesthia angustipennis, Philippine Islands (Chitwood and Chitwood, 1934).
Binema mirzaia (Basir, 1940) Basir, 1956
Synonymy.—Periplaneticola mirzaia Basir, 1940.
Natural host.—Periplaneta americana, India, Aligarh (Basir, 1940).
Blattelicola blattelicola Basir, 1940
Natural host.—Blattella germanica, India, Aligarh (Basir, 1940).
Blatticola blattae (Graeffe, 1860) Chitwood, 1932
Synonymy.—Oxyuris blattae Graeffe, 1860; Oxyuris blatticola Galeb, 1878; Blatticola blatticola (Galeb, 1877) Schwenck, 1926 [Chitwood, 1930, 1932].
Natural hosts.—Blattella germanica, Brazil (Pessôa and Corrêa, 1926; Schwenck, 1926); U.S.A. (Chitwood, 1930; Bozeman, 1942); Egypt? (Galeb, 1877, 1878); U.S.S.R. (Sobolev, 1937; Sondak, 1935); Czechoslovakia (Groschaft, 1956).
Ectobius lapponicus, Ectobius pallidus, Egypt? (Galeb, 1877, 1878).
Polyphaga aegyptiaca, France (Graeffe, 1860).
The life cycle has been studied by Bozeman (1942): He found never more than four worms in the large intestine of each cockroach. Embryos developed to "resting" stage in vitro. The resting stage was infective while the active stage was not. The worms seemed to have no effect on the vital activities of the host. Alicata (1934b) found that the embryo undergoes a molt before hatching.
Chitwood (1930) found 75 percent of the German cockroaches examined from houses in Washington infected. As a rule, one adult female, one or two males, and possibly two larval females are found in a single individual, apparently only in the rectum.
Sobolev (1937) found 72 percent of Blattella germanica collected in Gorkov (U.S.S.R.) infected with Blatticola blattae. The mean number of worms per host was 1.8, the maximum 5. Sondak (1935) found about 30 percent of 788 B. germanica collected in Leningrad to be infected with B. blattae. Groschaft (1956) regularly found only single worms in B. germanica, collected in a laboratory in Prague, except for two females that contained 2 and 3 worms each.
Blattophila sphaerolaima Cobb, 1920
Synonymy.—Aorurus sphaerolaima (Cobb, 1920) Travassos, 1929. Although Chitwood (1932) indicated that the taxonomic position of this nematode is questionable, Chitwood and Chitwood, 1934, apparently accepted it as a valid species in describing the variety javanica.
Natural host.—Panesthia laevicollis [Cobb recorded the host as Panesthia brevicollis, but no such cockroach exists. Van Zwaluwenburg (1928) and Caudell (in Chitwood, 1932) believed that Cobb meant Panesthia laevicollis. According to Gurney (personal communication, 1957) Caudell's notes show that in 1933 he wrote to Dr. Chitwood and explained that he had compared Cobb's figure of the cockroach with laevicollis Saussure (figures and description) and had found them the same.] Australia, New South Wales (Cobb, 1920).
Blattophila sphaerolaima var. javanica Chitwood and Chitwood, 1934
Natural host.—Panesthia angustipennis, Philippine Islands (Chitwood and Chitwood, 1934).
Blattophila supellaima Basir, 1941
Natural host.—Supella supellectilium, India, Aligarh (Basir, 1941).
Cephalobellus brevicaudatum (Leidy, 1851) Christie, 1933
Synonymy.—Thelastoma brevicaudatum Leidy, 1851 [Christie, 1933]. Thelastoma indiana Basir, 1940 [Basir, 1949].
Natural host.—Leucophaea sp., India, Aligarh (Basir, 1940, 1949).
Cephalobellus magalhāesi (Schwenck, 1926) Basir, 1956
Synonymy.—Bulhỡesi magalhāesi Schwenck, 1926; Thelastoma magalhāesi (Schwenck, 1926) Travassos, 1929 [Basir, 1956].
Natural host.—"Barata selvagem," Brazil, São Paulo (Schwenck, 1926).
Euryconema paradisa Chitwood, 1932
Natural host.—Eurycotis floridana, U.S.A., Florida (Chitwood, 1932).
Galebia aegyptiaca (Galeb, 1878) Chitwood, 1932
Synonymy.—Oxyuris aegyptiaca Galeb, 1878; Blatticola aegyptiaca (Galeb, 1878) Schwenck, 1926 [Chitwood, 1932].
Natural hosts.—Blattella germanica, Brazil (Schwenck, 1926).
Polyphaga aegyptiaca, Egypt? (Galeb, 1878).
Hammerschmidtiella diesingi (Hammerschmidt, 1838) Chitwood, 1932
Synonymy.—Anguillula macrura Diesing, 1851; Aorurus diesingi (Hammerschmidt, 1838) Travassos, 1929; Streptostomum gracile Leidy, 1850; Oxyuris diesingi Hammerschmidt, 1838; Oxyuris blattae orientalis Hammerschmidt, 1838 [Chitwood, 1932]. Oxyuris macrura of Lankester (1865).
Natural hosts.—Blatta orientalis, Europe (Hammerschmidt 1838, 1847; Bütschli, 1871); Egypt? (Galeb, 1878); England (Lankester, 1865; Lee, 1958); U.S.A. (Leidy, 1850a); U.S.S.R. (Yakimov and Miller, 1922; Sobolev, 1937; Sondak, 1935); Brazil (Travassos, 1929); China (Chitwood, 1932); Czechoslovakia (Groschaft, 1956).
Leucophaea maderae, Brazil (Pessôa and Corrêa, 1926).
Periplaneta americana, Brazil (Magalhães, 1900; Pessôa and Corrêa, 1926). U.S.A.: Texas (Todd, 1943); Kansas (Dobrovolny, 1933; Dobrovolny and Ackert, 1934); North Carolina (Hatcher, 1939); Iowa, North Dakota, Michigan (Hoffman, 1953). China (Chitwood, 1932). India (Basir, 1940). Czechoslovakia (Groschaft, 1956). England (Lee, 1958).
Periplaneta australasiae, Brazil (Pessôa and Corrêa, 1926).
Polyphaga aegyptiaca (Linstow, 1878).
Cockroaches (Blatta orientalis, Blattella germanica, and/or Periplaneta americana), U.S.A. (McAdow, 1931).
Cockroach, Venezuela (Tejera, 1926).
According to Hammerschmidt (1847) this worm may be found throughout the intestinal canal but especially in the small intestine. It is frequently found in adults and seldom in the nymphs. There were seldom more than 5 to 10 worms in one cockroach and female worms were found more frequently than males; the male worms were found only in winter and spring while the females were present at all times of the year. Bütschli (1871) stated that all stages from those just hatching to mature males and females are found.
Yakimov and Miller (1922) found H. diesingi in 50.8 percent of 124 B. orientalis collected in Petrograd. Sobolev (1937) found 96 percent of B. orientalis infected with H. diesingi with a mean number of 5.6 and maximum number of 22 in one cockroach. Groschaft (1956) found 18 in one specimen of B. orientalis. Dobrovolny and Ackert (1934) found about 50 percent of 222 P. americana infected with H. diesingi. Sondak (1935) found about 36 percent of 412 B. orientalis infected with either or both H. diesingi and Leidynema appendiculata.
Two molts occur during development of the eggs; the first takes place outside the host resulting in a resting or infective stage. After the egg in the infective stage is eaten by the host, the second molt occurs before the egg hatches. Completion of the second molt and hatching perhaps are connected with ammonia present in the digestive tract; the ammonia seems to arise from the bacteria present in the gut. There appears to be a relationship between the intestinal bacteria of the cockroach and development and hatching of nematode eggs (Todd, 1944).
At the time of oviposition the nematode eggs are in the very earliest stages of cleavage. In 36 hours a motile, tadpole-like stage is reached and in a few days the embryo becomes quiescent and nonmotile. This nonmotile stage is infective whereas the motile embryonic stage is not. Feeding experiments proved that transmission of the nematode is direct. The worm reaches sexual maturity in 20 or 30 days after being ingested by the cockroach (Dobrovolny, 1933).
The bacterium Streptomyces leidynematis Hoffman grows on the cuticle of H. diesingi (Hoffman, 1953). The bacterium apparently is only anchored to the nematode and probably obtains its food from the intestinal contents of the cockroach. See notes under Leidynema appendiculata.
Hammerschmidtiella neyrai Serrano Sánchez, 1945
Synonymy.—Hammerschmidtiella neyrae Serrano Sánchez, 1947. [According to M. B. Chitwood, personal communication, 1957, Serrano Sánchez's emendation is apparently an error.]
Natural host.—Blatta orientalis, Spain, Grenada (Serrano Sánchez, 1947): Of 2,943 specimens examined, 1,143 were parasitized by oxyurids and of these 45 percent contained H. neyrai.
Leidynema appendiculata (Leidy, 1850) Chitwood, 1932
Synonymy.—Oxyuris blattae orientalis Hammerschmidt, 1847, of Bütschli, 1871, and Oxyuris blattae-orientalis of Magalhães, 1900; Oxyuris blattae Hammerschmidt, 1847, of Galeb, 1878; Aorurus (Thelastoma) appendiculatus Leidy, 1850. [Chitwood, 1932.] Serrano Sánchez (1947) has divided this species into three geographical varieties as follows: L. appendiculata (Leidy, 1852) (Dobrovolny and Ackert, 1934) var. indiana; L. appendiculata (Leidy, 1852) (Chitwood, 1932) var. americana; L. appendiculata (Serrano Sánchez, 1947) var. hispana. However, Basir (1956) does not recognize these varieties. The Russians recognize hispana (M. B. Chitwood, personal communication, 1957).
Natural hosts.—Blaberus atropos, South America (Chitwood, 1932).
Blatta orientalis, Egypt? (Galeb, 1878); Europe (Bütschli, 1871); U.S.S.R. (Sobolev, 1937; Sondak, 1935); U.S.A., Nebraska (Todd, 1944); Spain (Serrano Sánchez, 1947): Recorded as var. hispana. Czechoslovakia (Groschaft, 1956). England (Lee, 1958a).
Blatta orientalis or Periplaneta americana, Brazil (Magalhães, 1900).
Periplaneta americana, U.S.A.: Texas (Todd, 1943); Nebraska (Todd, 1944); Kansas (Dobrovolny, 1933; Dobrovolny and Ackert, 1934); North Carolina (Hatcher, 1939); Iowa, North Dakota, Michigan (Hoffman, 1953). Czechoslovakia (Groschaft, 1956). England (Lee, 1958a).
Cockroach, Venezuela (Tejera, 1926).
Cockroaches (Blatta orientalis, Blattella germanica, and/or Periplaneta americana), U.S.A. (McAdow, 1931).
Chitwood (1932) also listed China for distribution of the worm, but we could not tell which host was involved.
The worms are found in the colon and rectum of the host. Galeb (1878) found as many as 20 individuals in a single B. orientalis. Sobolev (1937) found 52 percent of B. orientalis infected with L. appendiculata; the mean number of worms per roach was 1.5 and the maximum 2. Dobrovolny and Ackert (1934) found 69 percent of 222 P. americana infected with this species.
Two molts occur within the egg during development of the larva. The first molt occurs outside the host resulting in the formation of an infective resting stage. The second molt occurs inside the cockroach (Todd, 1941, 1944).
Transmission of the nematode is direct, eggs in the resting embryonated stage being infective (Dobrovolny and Ackert, 1934).
Hoffman (1953) described a filamentous bacterium, Streptomyces leidynematis Hoffman, which grows on the cuticle of L. appendiculata in P. americana. Leidy (1853) noted the presence of simple, inarticulate, amorphous filaments, growing from nematodes infecting B. orientalis. Bütschli (1871) and Magalhães (1900) described similar filaments adhering to the surface of oxyurids from cockroaches.
Leidynema appendiculata (Leidy, 1850) Chitwood, 1932?
Natural host.—Eurycotis floridana, U.S.A., Massachusetts (Roth and Willis, unpublished data, 1955): Determined by Dr. G. Steiner who wrote us, "In Eurycotis floridana there were ten specimens of the nematode Leidynema appendiculata (Leidy, 1850). This cockroach is obviously a new host for this nematode. I am not sure that the nematode exactly agrees with the description as given in the literature."
Leidynema cranifera Chitwood, 1932
Natural hosts.—Blaberus craniifer, U.S.A., Florida (Chitwood, 1932); Massachusetts (Roth and Willis, unpublished data, 1955). Determined by Dr. G. Steiner.
Blaberus atropos?, U.S.A., Florida (Chitwood, 1932): B. craniifer has generally been recorded as B. atropos of Stoll which is a closely related but distinct South American species (Rehn and Hebard, 1927).
Leidynema delatorrei Chitwood, 1932
Natural host.—Leucophaea maderae, Cuba, Havana (Chitwood, 1932).
Leidynema nocalum Chitwood and Chitwood, 1934
Natural host.—Panesthia angustipennis, Philippine Islands (Chitwood and Chitwood, 1934).
Leidynemella fusiformis Cobb, 1934
Natural hosts.—Panesthia laevicollis?, Philippine Islands (Cobb in Chitwood and Chitwood, 1934).
Panesthia angustipennis, Philippine Islands (Chitwood and Chitwood, 1934).
Leidynemella panesthiae (Galeb, 1878) Chitwood and Chitwood, 1934
Synonymy.—Oxyuris panesthiae Galeb, 1878, in part; Thelastoma panesthiae (Galeb, 1878) Travassos, 1929. [Chitwood, 1932; Chitwood and Chitwood, 1934.]
Natural host.—Panesthia sp., New Guinea (Galeb, 1878): About 40 nematodes may be found in one insect.
Leidynemella paracranifera Chitwood and Chitwood, 1934
Natural host.—Panesthia angustipennis, Philippine Islands (Chitwood and Chitwood, 1934).
Oxyuris (?) heterogamiae Galeb, 1878
Synonymy.—Thelastoma heterogamiae (Galeb, 1878) Travassos, 1929. The taxonomic position of this species is questionable; it might possibly belong in Blatticola or Protrellina (Chitwood, 1932). Basir (1956) placed it in an appendix to the family Thelastomatidae.
Natural host.—Polyphaga aegyptiaca, Egypt? (Galeb, 1878).
Protrelleta floridana Chitwood, 1932
Natural host.—Blaberus craniifer, U.S.A., Florida (Chitwood, 1932).
Protrellus aureus Cobb, 1920
Synonymy.—The taxonomic position of this nematode is questionable (Chitwood, 1932).
Natural host.—Polyzosteria melanaria?, Australia, New South Wales (Cobb, 1920). [Caudell (in Chitwood, 1932) stated that this host was probably Platyzosteria analis.]
Protrellus aurifluus (Chitwood, 1932) Chitwood, 1933
Synonymy.—Protrellina aurifluus Chitwood, 1932.
Natural hosts.—Parcoblatta lata, U.S.A., North Carolina, Maryland (Chitwood, 1932).
Parcoblatta uhleriana, North Carolina (Hatcher, 1939).
Protrellus australasiae (Pessôa and Corrêa, 1926) Travassos, 1929
Synonymy.—Oxyuris australasiae Pessôa and Corrêa, 1926; Protrellina australasiae (Pessôa and Corrêa, 1926) Chitwood, 1932 [Chitwood, 1933].
Natural host.—Periplaneta australasiae, Brazil (Pessôa and Corrêa, 1926, 1927).
Protrellus galebi Schwenck, 1926
Synonymy.—Protrellina galebi (Schwenck, 1926) Chitwood, 1932 [Chitwood, 1933].
Natural host.—"Barata selvagem," Brazil (Schwenck, 1926).
Protrellus künckeli (Galeb, 1878) Travassos, 1929
Synonymy.—Oxyuris künckeli Galeb, 1878; Protrellina künckeli (Galeb, 1878) Chitwood, 1932 [Chitwood, 1933].
Natural hosts.—Periplaneta americana, Egypt? (Galeb, 1877, 1878.) [Chitwood (1932) questioned the determination of this host because he failed to find this nematode in a large number of specimens from U.S.A. and China.] Brazil (Pessôa and Corrêa, 1926).
Periplaneta australasiae, Brazil (Pessôa and Corrêa, 1926).
Protrellus manni (Chitwood, 1932) Chitwood, 1933
Synonymy.—Protrellina manni Chitwood, 1932.
Natural host.—Aglaopteryx diaphana, Cuba (Chitwood, 1932).
Protrellus phyllodromi (Basir, 1942) Basir, 1956
Synonymy.—Protrellina phyllodromi Basir, 1942.
Natural host.—Blattella humbertiana, India, Aligarh (Basir, 1942): Found in the rectum.
Protrelloides paradoxa Chitwood, 1932
Natural host.—Eurycotis floridana, U.S.A., Florida (Chitwood, 1932).
Schwenkiella icemi (Schwenck, 1926) Basir, 1956
Synonymy.—Bulhõesia icemi Schwenck, 1926; Thelastoma icemi (Schwenck, 1926) Travassos, 1929; Thelastoma aligarhica Basir, 1940. [Basir, 1956.]
Natural hosts.—"Barata selvagem," Brazil, São Paulo (Schwenck, 1926).
Periplaneta americana, India, Aligarh (Basir, 1940); U.S.A., Nebraska (Todd, 1943).
Periplaneta brunnea, U.S.A., Louisiana (Todd, 1943).
Severianoia magna Pereira, 1935
Natural host.—"Blattidae sylvestres," Brazil (Pereira, 1935).
Severianoia severianoi (Schwenck, 1926) Travassos, 1929
Synonymy.—Bulhõesia severianoi Schwenck, 1926 [Travassos, 1929].
Natural hosts.—"Baratas de pau podre," Brazil (Schwenck, 1926).
Pycnoscelus surinamensis, U.S.A., Florida (Chitwood, 1932).
Suifunema caudelli Chitwood 1932
Natural host.—Steleopyga? sinensis, Asia: Suifu, Szchuen, China (Chitwood, 1932).
Thelastoma pachyjuli (Parona, 1896) Travassos, 1929
Synonymy.—Oxyuris bulhõesi de Magalhães, 1900; Bulhõesia bulhõesi (Magalhães, 1900) Schwenck, 1926 [Travassos, 1929; Chitwood, 1932]; Thelastoma bulhõesi (Magalhães, 1900) Travassos, 1929; although this last combination (from Chitwood, 1932) is not given by Basir (1956), it is implied by the synonymy that he does cite under T. pachyjuli.
Natural hosts.—Blatta orientalis, Czechoslovakia (Groschaft, 1956).
Periplaneta americana, Brazil (Magalhães, 1900); North America (Chitwood, 1932); U.S.A., North Carolina (Hatcher, 1939).
Thelastoma palmettum Chitwood and Chitwood, 1934
Natural host.—Panesthia angustipennis, Philippine Islands (Chitwood and Chitwood, 1934).
Thelastoma riveroi Chitwood, 1932
Natural host.—Periplaneta sp., Cuba (Chitwood, 1932).
Undetermined nematodes
Natural host.—Cutilia sp. near sedilloti, U.S.A. (hosts imported from New Zealand) (Roth, unpublished data, 1957).
Class NEMATOMORPHA
Order GORDIOIDEA
The immature stages of the following gordian worms have been found in the body cavity of cockroaches.
Family CHORDODIDAE
Chordodes morgani Montgomery, 1898
Synonymy.—Chordotes [sic] puerilis Montgomery, 1898 [Ward, 1918].
Natural host.—Cockroach, U.S.A. (Montgomery, 1898); Pennsylvania, Maryland, Michigan, Ohio, Florida, Iowa, Nebraska (Ward, 1918).
Family GORDIIDAE
Gordius aquaticus Linnaeus, 1758
Natural host.—Blatta sp., U.S.A. (Stiles and Hassall, 1894).
Leidy (1879) identified a 9-inch-long nematode which came from a cockroach (Blatta orientalis?) as probably being Gordius aquaticus. Ransom (in Pierce, 1921) states that G. aquaticus may be an accidental parasite of man. Faust (1955) summarizes the few reported cases of human parasitism. Dorier (1930) reported that the regurgitated liquid of Blatta orientalis had no effect on cysts of G. aquaticus after one hour.
Gordius blattae orientalis Diesing, 1851
Synonymy.—Gordius orientalis of Lankester (1865).
Natural host.—Blatta orientalis, Germany (Siebold, 1842; Linstow, 1878): Found in abdomen. Von Siebold called this "Filarien" but did not otherwise name or describe the worm.
Gordius pilosus (Möbius, 1855) Diesing, 1861
Synonymy.—Chordodes pilosus Möbius, 1855 [Diesing, 1861.]
Natural host.—Blaberus giganteus, Venezuela (Möbius, 1855): From the insect's abdomen.
Gordius sp.
Natural hosts.—Periplaneta americana, South Africa (Porter, 1930); Germany (Bode, 1936): Bode's record may have referred to a Mermis or other nematode.
Cockroaches, Venezuela (Miall and Denny, 1886; Burr, 1899a; Tejera, 1926).
Parachordodes raphaelis (Camerano, 1893) Camerano, 1897
Synonymy.—Gordius raphaelis Camerano, 1893 [Camerano, 1897].
Natural hosts.—Symploce parenthesis and Kuchinga hemerobina, French Equatorial Africa (Camerano, 1893, 1897).
Undetermined gordian worms
Natural hosts.—Eurycotis floridana, Florida (T. Eisner, personal communication, 1958): See plate 29, B.
Parahormetica bilobata, Brazil (Pessôa and Corrêa, 1929): Worm referred to as "gordiaceo."
Cockroaches, Australia (E. F. Riek, personal communication, 1953): Three undescribed gordian worms were found in undetermined cockroaches of the subfamily Blattinae.
HELMINTHS[4] FOR WHICH COCKROACHES SERVE AS INTERMEDIATE HOSTS
The use of the asterisk (*) is explained in footnote [3], page [4].
Phylum ACANTHOCEPHALA
Order ARCHIACANTHOCEPHALA
Family OLIGACANTHORHYNCHIDAE
* Prosthenorchis elegans (Diesing, 1851) Travassos, 1915
Natural host.—Blattella germanica, France (Brumpt and Urbain, 1938, 1938a; Brumpt et al., 1939).
Experimental hosts.—Blaberus atropos and Leucophaea maderae, France (Brumpt and Desportes, 1938).
* Prosthenorchis spirula (Olfers in Rudolphi, 1819) Travassos, 1917
Natural host.—Blattella germanica, France (Brumpt and Urbain, 1938, 1938a; Brumpt et al., 1939); Netherlands (Thiel and Wiegand Bruss, 1946).
Experimental hosts.—Blattella germanica, Netherlands (Thiel and Wiegand Bruss, 1946).
Blaberus atropos and Leucophaea maderae, France (Brumpt and Desportes, 1938).
Family MONILIFORMIDAE
* Moniliformis dubius Meyer, 1932
Natural hosts.—Periplaneta americana, Brazil (Magalhães, 1898; Travassos, 1917); Gold Coast (Southwell, 1922); India (Pujatti, 1950); U.S.A. (Burlingame and Chandler, 1941; Moore, 1946).
Periplaneta australasiae, India (Pujatti, 1950).
Experimental hosts.—Blattella germanica, Japan (Yamaguti and Miyata, 1942).
Periplaneta americana, U.S.A. (Chandler, 1941; Moore, 1946); Japan (Yamaguti and Miyata, 1942).
* Moniliformis kalahariensis Meyer, 1931
Natural host.—Blattella germanica, India (Meyer, 1931, 1932).
* Moniliformis moniliformis (Bremser in Rudolphi, 1819) Travassos, 1915
Natural hosts.—Periplaneta americana, Argentina (Bacigalupo, 1927, 1927a, 1928); Brazil (Pessôa and Corrêa, 1929); Algeria (Seurat, 1912); Burma (Subramanian, 1927); South Africa (Porter, 1930); Madras (Sita, 1949).
Periplaneta spp., New Caledonia (Rageau, 1956).
Cockroaches, Venezuela (Tejera, 1926).
Experimental hosts.—Blaberus atropos, Blatta orientalis, Blattella germanica, Leucophaea maderae, France (Brumpt and Urbain, 1938a).
Periplaneta americana, Japan (Yamaguti and Miyata, 1942); France (Brumpt, 1949); Madras (Sita, 1949).
Phylum ASCHELMINTHES
Class NEMATODA
Order OXYUROIDEA
Family SUBULURIDAE
* Subulura jacchi (Diesing, 1861) Railliet and Henry, 1914
Synonymy.—Subulura jacchi (Marcel, 1857) [Dr. J. T. Lucker, personal communication, 1957].
Experimental host.—Blaberus atropos, France (Chabaud and Larivière, 1955).
Order SPIRUROIDEA
Family THELAZIIDAE
* Oxyspirura mansoni (Cobbold, 1879) Ransom, 1904
Natural hosts.—Pycnoscelus surinamensis, Australia (Fielding, 1926, 1927, 1928, 1928a); U.S.A. (Sanders, 1927, 1928, 1929; Shealy, 1927); Formosa (Kobayashi, 1927); Antigua (Hutson, 1938, 1943); Hawaii (Illingworth, 1931; Schwabe, 1950, 1950a, 1950b, 1951); New Caledonia (Rageau, 1956).
We have recently found (Roth and Willis, 1960) that two strains of Pycnoscelus surinamensis exist; a parthenogenetic strain (from Florida), and a bisexual strain (from Hawaii) which does not reproduce parthenogenetically. The parthenogenetic strain is undoubtedly the form that has been shown to be the host of O. mansoni in the United States and Antigua, because only this form is found in the New World. Probably the parthenogenetic strain was the form involved in most Pacific areas. However, from internal evidence in his papers, we concluded that Schwabe, in Hawaii, may well have been working with the bisexual strain and possibly also with the parthenogenetic strain; if this is true, then both parthenogenetic and bisexual strains of Pycnoscelus surinamensis may serve as intermediate hosts of the eyeworm.
Experimental hosts.—Periplaneta americana, Antigua (Hutson, 1943).
Pycnoscelus surinamensis, U.S.A. (Sanders, 1929); Australia (Fielding, 1927, 1928a); Hawaii (Schwabe, 1951).
Rictularia coloradensis Hall, 1916
Natural hosts.—Parcoblatta pensylvanica and Parcoblatta virginica, U.S.A., Ohio (Oswald, 1958): Of 49 wood roaches collected, one of each species contained a single larva each.
Experimental hosts.—Blatta orientalis, Blattella germanica, Parcoblatta pensylvanica, Parcoblatta virginica, Periplaneta americana, and Supella supellectilium, U.S.A. (Oswald, 1958): The larvae underwent normal development in all species of cockroaches except B. orientalis and P. americana in which cysts developed that contained a reddish-brown pigment; larvae in such cysts were dead or dying. Eggs of R. coloradensis hatched in the midgut of B. germanica and first-stage larvae entered the hindgut epithelium within 24 hours. The larvae underwent two molts within a cyst formed by tissues of the host's gut, the second molt occurring during the twelfth or thirteenth day. In Parcoblatta, cysts were found free in the body cavity as well as attached to the hindgut. In B. germanica and S. supellectilium the cysts remained attached to the hindgut. Usually over 20 cysts developed in each infected Parcoblatta; fewer than 10 per insect developed in the other species. Larvae became infective to the definitive host, the white-footed mouse [Peromyscus leucopus noveboracensis (Fischer)], as early as the tenth day.
Family SPIRURIDAE
* Agamospirura parahormeticae Pessôa and Corrêa, 1929
Natural host.—Parahormetica bilobata, Brazil (Pessôa and Corrêa, 1929).
* Gongylonema ingluvicola Ransom, 1904
Experimental host.—Blattella germanica, U.S.A. (Cram, 1935).
* Gongylonema neoplasticum (Fibiger and Ditlevsen, 1914) Ransom and Hall, 1916
Natural hosts.—Blatta orientalis, Netherlands (Baylis, 1925).
Blattella germanica, U.S.A. (Hitchcock and Bell, 1952).
Periplaneta americana, Denmark and St. Croix (Fibiger, 1913, 1913a; Fibiger and Ditlevsen, 1914); Netherlands (Baylis, 1925); Argentina (Bacigalupo, 1930); England (Leiper, 1926); South Africa (Porter, 1930); U.S.A. (Hitchcock and Bell, 1952); Formosa (Yokagawa, 1924, 1925, 1925a).
Periplaneta australasiae, Formosa (Yokagawa, 1924, 1925, 1925a).
Experimental hosts.—Blattella germanica, Denmark (Fibiger and Ditlevsen, 1914); U.S.A. (Hitchcock and Bell, 1952); France (Brumpt, 1949).
Blatta orientalis, Denmark (Fibiger and Ditlevsen, 1914).
Periplaneta americana, Denmark and St. Croix (Fibiger, 1913; Fibiger and Ditlevsen, 1914); U.S.A. (Hitchcock and Bell, 1952).
* Gongylonema pulchrum Molin, 1857
Experimental hosts.—Blattella germanica, U.S.A. (Ransom and Hall, 1915, 1916, 1917; Stiles and Baker, 1927; Schwartz and Lucker, 1931; Lucker, 1932; Alicata, 1934a, 1935); Europe (Baylis et al., 1925, 1926, 1926a; Sambon, 1926).
Parcoblatta sp., Alicata (1934, 1935).
* Gongylonema sp.
Natural host.—Periplaneta americana, Brazil (Magalhães, 1900); Algeria (Seurat, 1916); England? (Leiper, 1926).
* Microtetrameres helix Cram, 1927
Experimental host.—Blattella germanica, U.S.A. (Cram, 1934).
* Protospirura bonnei Ortlepp, 1924
Natural host.—Leucophaea maderae, Venezuela (Brumpt, 1931).
Experimental hosts.—Blatta orientalis, Blattella germanica, Leucophaea maderae, France (Brumpt, 1931).
* Protospirura columbiana Cram, 1926
Experimental host.—Blattella germanica, U.S.A. (Cram, 1926).
* Protospirura muricola Geodoelst, 1916
Natural host.—Leucophaea maderae, Panama (Foster and Johnson, 1938, 1939).
* Seurocyrnea colini (Cram, 1927) Cram, 1931
Experimental host.—Blattella germanica, U.S.A. (Cram, 1931, 1931a, 1933a).
* Spirura gastrophila (Müller, 1894) Seurat, 1913
Natural hosts.—Blatta orientalis, Europe? (Deslongchamps, 1824, in Seurat, 1911); Italy (Grassi, 1888); Algeria (Seurat, 1916).
Periplaneta americana, Brazil (Pessôa and Corrêa, 1929).
Cockroach, Venezuela (Tejera, 1926).
Experimental hosts.—Blatta orientalis, France (Galeb, 1878a); "Cafards," Algeria (Roger, 1906, 1907).
* Tetrameres americana Cram, 1927
Natural host.—Blattella germanica, U.S.A. (Cram, 1931b, personal communication, 1956); Hawaii (Alicata, 1938, 1947).
Experimental host.—Blattella germanica, U.S.A. (Cram, 1931b).
* Tetrameres pattersoni Cram, 1933
Experimental host.—Blattella germanica, U.S.A. (Cram, 1933).
Family PHYSALOPTERIDAE
* Physaloptera hispida Schell, 1950
Experimental host.—Blattella germanica, U.S.A. (Schell, 1952, 1952a).
* Physaloptera maxillaris Molin, 1860
Experimental host.—Blattella germanica, U.S.A. (Hobmaier, 1941).
* Physaloptera praeputialis von Linstow, 1889
Experimental host.—Blattella germanica, U.S.A. (Petri and Ameel, 1950).
* Physaloptera rara Hall and Wigdor, 1918
Experimental host.—Blattella germanica, U.S.A. (Petri and Ameel, 1950; Petri, 1950).
* Physaloptera turgida Rudolphi, 1819
Experimental host.—Blattella germanica, U.S.A. (Alicata, 1937; Schell, 1952).