Chapter VIII: Section III: (p. 14). Other associations may be accidental (e.g., certain (2)

_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

=Stenophora= sp.

_Natural host._--_Blatta orientalis_, India (Bal and Rai, 1955): Organism found in the midgut of the cockroach.

Family GREGARINIDAE

=Gregarina blattarum= von 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 fastidiosa= Harrison

_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 gibbsi= Harrison

_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 illinensis= M. 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 impetuosa= Harrison

_Natural host._--_Melanosilpha capensis_, South Africa (Harrison, 1955): All specimens of this gregarine were found in the anterior mesenteron of the host.

=Gregarina légeri= Pinto

_Natural host._--_Periplaneta americana_, Brazil (Pinto, 1918, 1918a, 1919): Intestinal canal.

=Gregarina neo-brasiliensis= Al. Cunha

_Natural host._--_Periplaneta americana_, Brazil (R. de Almeida Cunha _in_ Pinto, 1919; Cunha, 1919).

=Gregarina ohioensis= Semans

_Natural host._--_Parcoblatta virginica_, U.S.A., Ohio (Semans, 1939): The protozoan was present in large numbers in the insect's midgut.

=Gregarina panchlorae= Frenzel

_Natural host._--_Panchlora exoleta_, Argentina (Frenzel, 1892): Midgut.

=Gregarina parcoblattae= Semans

_Natural hosts._--_Parcoblatta pensylvanica_ and _Parcoblatta uhleriana_, U.S.A., Ohio (Semans, 1939): Midgut.

=Gregarina rhyparobiae= J. 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 sandoni= Harrison

_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 thomasi= Semans

_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 tenax= Schneider

(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 cryptocerci= Yarwood

_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 periplanetae= Georgévitch

_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 (Ivanić, 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 kudoi= Sprague 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.

=Plistophora= sp.

_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

=Paramecium= sp.

_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 caulleryi= Weill

_Natural host._--_Periplaneta americana_, Indochina (Weill, 1929): Alimentary canal.

Order SPIROTRICHA

Family BURSARIIDAE

=Balantidium blattarum= Ghosh

_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 ovatum= Ghosh

_Natural host._--_Periplaneta americana_, India (Ghosh, 1922a; Bhatia and Gulati, 1927); Indochina, Saigon (Weill, 1929): Intestinal tract.

=Balantidium praenucleatum= Kudo 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.

=Balantidium= sp.?

_Natural host._--_Periplaneta americana_, Brazil (Magalhães, 1900): These organisms were numerous in the intestine.

Family SPIROSTOMIDAE

=Nyctotherus buissoni= Pinto

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

=Nyctotherus ovalis= Leidy

_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 (Bělǎr, 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 uichancoi= Kidder

_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 viannai= Pinto

_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 brevis= Kidder

_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 simplex= Kidder

_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).

XI. HELMINTHS

Intestinal nematodes of the family Thelastomatidae have no apparent pathological effect on their cockroach hosts. Galeb (1878) has shown experimentally that oxyurids eat the same food as the host insect and that if one starves them, by withholding food from the host, the oxyurids die and disappear. In other words, these worms are not parasites, in the sense that we use the term in this paper, but commensals. Dobrovolny and Ackert (1934) stated that "all observations seemed to indicate that the health, fertility and activity of the heavily infested cockroaches were comparable with those of the non-parasitised specimens."

Very few papers have dealt with the ecology of the oxyurid parasites of cockroaches. According to Galeb (1878), usually one species of nematode is found in a single cockroach, but sometimes two species live together in the same host (e.g., in _Blatta orientalis_ and _Polyphaga aegyptiaca_) where they compete for food. Galeb claimed that _Hammerschmidtiella diesingi_ would replace _Leidynema appendiculata_; he observed that _H. diesingi_ surpassed _L. appendiculata_ in numbers and the latter became uncommon in the intestines of the cockroaches. On the other hand, Sobolev (1937) found that 48 percent of his oriental cockroaches were infected with both of the above species of nematodes. The average number of both species was 7.5, and the maximum number was 97; the mean number of _H. diesingi_ was 5.1 and the maximum 64; the mean number of _L. appendiculata_ was 2.4 and the maximum 33. More than 40 nematodes were found in each cockroach of 1.3 percent of those examined. These results apparently contradict Galeb's conclusions inasmuch as the number of each species in mixed infections was essentially the same as the number found in cockroaches infected by only one species (see pp. 195 and 197). Dobrovolny and Ackert (1934) found that 29 percent of 222 _Periplaneta americana_ contained both of the above species of nematodes; whereas 40 percent contained _L. appendiculata_ only, and 21 percent contained _H. diesingi_ only. The infestation ranged from 1 to 36 worms per cockroach with averages of 3.8 per male, 5.1 per female, and 2.7 per nymph.

The eggs of some helminths pass unharmed through the guts of cockroaches that serve as vectors of these ova and have no effect on the insect. However, helminths that are secondary parasites in cockroaches damage the insect to varying degrees depending upon the extent of the infection. Thus the larvae of _Moniliformis moniliformis_ pass through the gut wall and some may become embedded in the fat tissue (Moore, 1946). First stage larvae of _Oxyspirura mansoni_ also burrow through the midgut wall into the fat body; Sanders (1929) believed that _Pycnoscelus surinamensis_ could be killed if at one time a sufficient number of migrating larvae of _O. mansoni_ penetrated the cockroach's intestinal wall. _Gongylonema neoplasticum_ migrates through the digestive tract and encysts in the muscles of the thorax and legs of the host (Fibiger and Ditlevsen, 1914). Infective larvae of _Protospirura muricola_, after hatching from ingested eggs, pass through the cockroach's gut wall and encyst mainly in the thorax, around the crop, and at the bases of the large muscles of the prothoracic legs (Foster and Johnson, 1939). It is probably generally true that nematodes which are secondary parasites in cockroaches do some damage to the host's intestinal tract at least, and they probably also damage other organs in which they may encyst.

Cockroach tissues may react defensively to infections by parasitic nematodes. For example, encysted third-stage larvae of _Physaloptera turgida_ have been found enclosed in a thin, brown, chitinous substance that was undoubtedly deposited by the tissue of the cockroach (Alicata, 1937). Cysts of similar appearance have been found in cockroaches infected with _Physaloptera rara_, _P. maxillaris_, _P. hispida_ (Petri, 1950; Hobmaier, 1941; Schell, 1952), and _Gongylonema pulchrum_ (Schell, 1952a); in the latter species the deposit eventually completely surrounded the nematode larva which was killed and "chitinized." Apparently these pigmented cysts surround unhealthy or dead larvae and are secreted as a defensive mechanism by the host (Schell, 1952a). Oswald (1958) has reported finding similar pigmented cysts in _Blatta orientalis_ and _Periplaneta americana_ that were experimentally infected with _Rictularia coloradensis_.

Our classification of the helminths follows Hyman (1951, 1951a).

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).

HELMINTHS WHOSE EGGS HAVE BEEN CARRIED BY COCKROACHES

The use of the asterisk (*) is explained in footnote 3, page 4.

Phylum PLATYHELMINTHES

Class TREMATODA

Order DIGENEA

Family SCHISTOSOMATIDAE

* =Schistosoma haematobium= (Bilharz, 1852) Weinland, 1858

_Experimental vector._--_Periplaneta americana_, Gold Coast Colony (Macfie, 1922).

Class CESTODA

Order TAENIOIDEA

Family HYMENOLEPIDIDAE

* =Hymenolepis= sp.

_Natural vectors._--_Periplaneta americana_, Formosa (Morischita and Tsuchimochi, 1926).

_Polyphaga saussurei_, Tadzhikistan (Zmeev, 1936).

Family TAENIIDAE

* =Taenia saginata= Goeze, 1782

_Experimental vector._--_Periplaneta americana_, Gold Coast Colony (Macfie, 1922).

=Echinococcus granulosis= (Batsch, 1786) Rudolphi, 1805

_Synonymy._--_Taenia echinococcus_ (Zeder, 1803) [Faust, 1939].

_Experimental vector._--_Periplaneta americana_, Uruguay (Pérez Fontana, 1955): Eggs were recovered from the feces of artificially infested cockroaches under "natural" conditions.

Family Unknown

* =Undetermined tapeworm ova=

_Natural vector._--_Polyphaga saussurei_, Tadzhikistan (Zmeev, 1936).

Phylum ASCHELMINTHES

Class NEMATODA

Order OXYUROIDEA

Family OXYURIDAE

* =Enterobius vermicularis= (Linnaeus, 1758) Leach in Baird, 1853

_Natural vectors._--_Blatta orientalis_ and _Blattella germanica_, U.S.S.R. (Sondak, 1935).

Order ASCAROIDEA

Family ASCARIDAE

* =Ascaris lumbricoides= Linnaeus, 1758

_Natural vector._--_Periplaneta americana_, South Africa (Porter, 1930): The eggs may have been those of _A. suum_ Goeze, 1782.

_Experimental vectors._--_Periplaneta americana_, Gold Coast Colony (Macfie, 1922); India (Chandler, 1926).

_Periplaneta americana_, _Periplaneta australasiae_, _Neostylopyga rhombifolia_, Formosa (Morischita and Tsuchimochi, 1926).

* =Ascaris= sp.

_Natural vector._--_Blatta orientalis_, Italian Somaliland (Mariani and Besta, 1936).

_Experimental vector._--_Periplaneta americana_, Uruguay (Pérez Fontana, 1955): Eggs recovered from the insects' feces.

Order STRONGYLOIDEA

Family ANCYLOSTOMIDAE

* =Ancylostoma caninum= (Ercolani, 1859) Hall, 1913

_Experimental vector._--_Periplaneta americana_, Netherlands (Akkerman, 1933).

* =Ancylostoma ceylanicum= (Looss, 1911) Leiper, 1915

_Experimental vector._--_Periplaneta americana_, Gold Coast Colony (Macfie, 1922); Netherlands (Akkerman, 1933).

* =Ancylostoma duodenale= (Dubini, 1843) Creplin, 1845

_Natural vector._--_Periplaneta americana_, South Africa (Porter, 1929, 1930).

_Experimental vector._--_Periplaneta americana_, Gold Coast Colony (Macfie, 1922).

* =Necator americanus= (Stiles, 1902) Stiles, 1906

_Natural vector._--_Periplaneta americana_, India (Chandler, 1926).

_Experimental vector._--_Periplaneta americana_, Gold Coast Colony (Macfie, 1922).

* =Hookworm ova=

_Experimental vectors._--_Periplaneta americana_, _Periplaneta australasiae_, _Neostylopyga rhombifolia_, Formosa (Morischita and Tsuchimochi, 1926).

Family TRICHOSTRONGYLIDAE

* =Trichostrongylus= sp.

_Natural vector._--_Blatta orientalis_, Italian Somaliland (Mariani and Besta, 1936).

Order TRICHUROIDEA

Family TRICHURIDAE

* =Capillaria hepatica= (Bancroft, 1893) Travassos, 1915

_Experimental vector._--_Blatta orientalis_, Italy? (Giordano, 1950).

* =Trichuris trichiura= (Linnaeus, 1771) Stiles, 1901

_Natural vectors._--_Blatta orientalis_, Italian Somaliland (Mariani and Besta, 1936); U.S.S.R. (Sondak, 1935).

_Blattella germanica_, U.S.S.R. (Sondak, 1935).

_Periplaneta americana_, Gold Coast Colony (Macfie, 1922); Formosa (Morischita and Tsuchimochi, 1926).

_Experimental vectors._--_Periplaneta americana_, Gold Coast Colony (Macfie, 1922); India (Chandler, 1926); Uruguay (Pérez Fontana, 1955).

_Periplaneta americana_, _Periplaneta australasiae_, and _Neostylopyga rhombifolia_, Formosa (Morischita and Tsuchimochi, 1926).

XII. ARTHROPODA

The classification follows Brues et al. (1954) with the following exceptions. The Acarina are arranged according to Dr. J. H. Camin (personal communication, 1955). Family Eupelmidae of the Hymenoptera follows the classification of Peck (1951).

Class ARACHNIDA

In this class, representatives of at least four orders have utilized cockroaches as food: the whip scorpions, scorpions, spiders, and mites. Apparently none of these feed exclusively on cockroaches, but the Philippine forest scorpion _Heterometrus_ (=_Palamnaeus_) _longemanus_ seems to prefer blattids to other insects (Schultze, 1927).

Order PEDIPALPIDA

Family THELYPHONIDAE

=Mastigoproctus giganteus= (Lucas)

_Synonymy._--_Thelyphonus giganteus_ Lucas [Dr. R. E. Crabill, personal communication, 1958].

_Experimental prey._--Cockroaches, U.S.A. (Marx, 1892, 1894): Immature whip scorpion captured and fed on one or two cockroaches a week. It lived on this diet for about two years.

=Mastigoproctus= sp.

_Common name._--Whip scorpion.

_Experimental prey._--_Blattella germanica_, U.S.A., Florida [Dr. B. J. Kaston, personal communication, 1953].

Order SCORPIONIDA

Pocock (1893) noticed that a scorpion whose pectines had come in contact with a cockroach immediately turned back and ate the insect. He concluded that the scorpion detected the cockroach by means of the pectines. However, Cloudsley-Thompson (1955) has demonstrated that the main function of the pectines is probably the detection of ground vibrations. He accounted for Pocock's observation by the presence of sensory spines (presumably tactile) which project from beneath the pectines. In a house in Arizona, Lyon (1951) observed over 60 scorpions living in a kitchen cabinet that enclosed a sink. They were apparently thriving on a heavy infestation of cockroaches. Stahnke (1953) stated that he used _Periplaneta americana_ as the principal food for scorpions at the Poisonous Animals Research Laboratory of Arizona State College. Cloudsley-Thompson (1955a) cited cockroaches as one of the arthropods that scorpions feed upon.

Family BUTHIDAE

=Buthus australis= (Linnaeus)

_Synonymy._--_Androctonus australis_ [Crabill, personal communication, 1957].

_Experimental prey._--Cockroaches, England (Cloudsley-Thompson, 1955a): This African species ate at least one cockroach per week during the summer months. It can, however, survive four months' starvation and is particularly adapted to a dry climate (Cloudsley-Thompson, personal communication, 1956).

=Centruroides gracilis= (Latreille)

_Experimental prey._--_Periplaneta americana_, U.S.A. (Roth, unpublished data, 1953): Scorpion collected in Florida by Roth and identified by Dr. M. H. Muma.

=Centruroides hentzi= (Banks)

_Experimental prey._--_Periplaneta australasiae_ and _Pycnoscelus surinamensis_, U.S.A. (Muma, personal communication, 1953): This scorpion occurs in large numbers in the Florida citrus groves, together with _P. australasiae_ which is probably an important natural prey.

=Centruroides vitattus= (Say)

_Natural prey._--_Parcoblatta pensylvanica_ (?), U.S.A., Florida (Muma, personal communication, 1953). This may have been another species of this genus, possibly _P. divisa_, as _P. pensylvanica_ is not known from Florida (Rehn, personal communication, 1958).

_Experimental prey._--_Blatta orientalis_, _Blattella germanica_, _Periplaneta americana_, and _Pycnoscelus surinamensis_, U.S.A., Florida (Muma, personal communication, 1953).

=Parabuthus capensis= (Hemprich and Ehrenberg)

_Experimental prey._--"Common house-cockroach" (Pocock, 1893): The scorpions were collected in Cape Town, South Africa.

Family CHACTIDAE

=Euscorpius germanus= (Koch)

_Synonymy._--_Euscorpius carpathicus_ [Cloudsley-Thompson, 1955a].

_Experimental prey._--_Blattella germanica_, England? (Pocock, 1893).

_Periplaneta americana_, nymphs, England? (Cloudsley-Thompson, personal communication, 1956): This scorpion is found naturally in southern Europe and North Africa.

=Euscorpius italicus italicus= (Herbst)

_Experimental prey._--Cockroaches including nymphs of _Periplaneta_, England? (Cloudsley-Thompson, 1951): The cockroaches had to be disabled before the scorpion would feed on them. Prey is apparently detected by tactile and auditory senses, sight being poorly developed and not used. The scorpion is found in southern Europe and North Africa.

Family VEJOVIDAE

=Hadrurus arizonensis= Ewing

_Experimental prey._--_Periplaneta americana_, U.S.A. (Stahnke, 1949): This record is a photograph showing the scorpion eating the cockroach.

Family ISCHNURIDAE

=Hormurus caudicula= (Koch)

_Experimental prey._--Cockroach, Australia (McKeown, 1952): This record is a photograph showing the scorpion feeding on a cockroach.

Family SCORPIONIDAE

=Heterometrus longimanus= (Herbst)

_Synonymy._--_Palamnaeus longimanus_ [Cloudsley-Thompson, personal communication, 1956].

_Natural prey._--"Large wood cockroach," Philippine Islands (Schultze, 1927): On several occasions Schultze found fragments of wings and legs of the large wood cockroach in a scorpion cavity, under a rotten log.

_Experimental prey._--_Leucophaea maderae_, _Periplaneta americana_, and other species of Blattidae, Philippine Islands (Schultze, 1927): Blattids seemed to be the favored food. This scorpion is usually found in humid, damp places in forest and jungle. Schultze describes in detail feeding behavior of the scorpion and method of capturing its prey.

=Urodacus novaehollandiae= Peters

_Experimental prey._--_Periplaneta americana_, Australia (Glauert, 1946): An injured cockroach was accepted at once by the scorpion, which held the insect in its claws and tore it with the alternately moving chelicerae. The scorpion ate all the soft parts and most of the sclerotized exoskeleton.

Order ARANEIDA

Many observations of spiders feeding on cockroaches are quite general, and many observers have failed to identify either the spider or its prey. Belt (1874) stated that "the cockroaches that infest houses in the tropics ... have numerous enemies--birds, rats, scorpions, and spiders." When Belt tried to drive a cockroach toward a large cockroach-eating spider, the insect rushed away from him until it came within a foot of the spider when it would double back, never advancing nearer.

Beebe (1925) watched a giant "wood roach," which was in the grasp of a 2-inch ctenid spider, fly through the window of his British Guiana laboratory. While the spider ate the cockroach, the insect gave birth to 51 nymphs. Sonan (1924) reported that large gray spiders devour nymphs and adults of _Periplaneta americana_ and _P. australasiae_ in Formosa; this spider also occurs on Hiyakejima Island and Okinawa. Passmore (1936), who has produced some excellent photographs of tarantulas, stated that they destroy cockroaches. Rau (1940) stated that American and oriental cockroaches were the principal item of diet of a friend's pet tarantula for several years. Kaston (personal communication, 1953) successfully fed a tarantula with _Periplaneta americana_.

Bristowe (1941) found that the British species of _Ectobius_ are readily accepted by _Xysticus_, _Clubiona_, _Drassodes_, _Zelotes_, _Tarantula_, and the web-builders _Ciniflo_ and _Aranea_. The British domestic cockroaches were accepted by _Tegenaria_ and _Ciniflo_, spiders large enough to overpower them, and were useful as food for tropical avicularids, ctenids, and sparassids in captivity.

Family THERAPHOSIDAE

=Avicularia avicularia= (Linnaeus) and =Avicularia= sp.?

_Common name._--Bird-eating spider.

_Natural prey._--_Periplaneta americana_, Trinidad (Main, 1924, 1930): The remains of the host were compressed into globular form by the spider after it had extracted the nutritive parts.

=Phormictopus cancerides= (Latreille)

_Experimental prey._--Cockroach, West Indies (Wolcott, 1953).

Family SPARASSIDAE

=Heteropoda venatoria= (Linnaeus)

_Synonymy._--_Heteropoda regia_ Fabricius.

_Common names._--Banana spider (Comstock, 1912); huntsman spider (Gertsch, 1949); big brown house spider (Bryan, 1915).

_Natural and experimental prey._--Cockroaches, Bermuda (Verrill, 1902); Puerto Rico (Sein, 1923; Wolcott, 1924a; Petrunkevitch, 1930a); Hawaii (Bryan, 1915, Williams et al., 1931); British Guiana (Moore _in_ Williams et al., 1931); Panama (Gertsch, 1949); New Zealand (adventive) (Parrott, 1952); England (Cloudsley-Thompson, 1953); Comstock (1912); Hawaii (Pemberton, 1917).

This (pl. 30, A) is a tropical species frequently imported into northern localities with bunches of bananas (Comstock, 1912; Cloudsley-Thompson, 1953). Adults measure 3 to 4 inches across with bodies over an inch long. They seldom leave their resting places during the day, but are active at night and search for food. The female does not spin a web (Bryan, 1915; Gertsch, 1949). The spider turns the cockroach over onto its back at the instant of seizure and holds it firmly against the substrate. The cockroach dies in 10 minutes and is gradually rolled up by the spider as it sucks out the nutriment (Moore _in_ Williams et al., 1931). The spider does not attempt to bite when captured, but if it does, its bite is said to be painful but not dangerous (Cloudsley-Thompson, 1953). Zimmerman (1948) found scores of _Periplaneta australasiae_ breeding in rock piles in Hawaii; also present were large numbers of these spiders and centipedes which presumably preyed upon the cockroaches.

Family THERIDIIDAE

=Latrodectus indistinctus= Pickard-Cambridge

_Common name._--Button spider.

_Natural prey._--_Karnyia discoidalis_, South Africa, Western Cape Province (Hesse, 1942): The nest is constructed on the ground among grass stems or other vegetation. Preferred sites are slight hollows, hoof imprints, etc. Nests are roughly tubular. The remains of insects are entangled in the walls of the nest where they form dense accumulations. Predatory activities of the spider are limited to an area close to the tubular entrance to the nest and do not extend beyond the trapping strands near the entrance. Capture is dependent upon accidental contact of the insect with sticky threads surrounding the entrance. This spider apparently attacks any insect or arachnid that becomes entangled in the nest. In an examination of 40 nests, remains of 6 _K. discoidalis_ were found.

=Latrodectus mactans= (Fabricius)

_Common names._--Black widow, hourglass, or shoe-button spider.

_Natural prey._--Cockroaches, Puerto Rico (Petrunkevitch, 1930). U.S.A., Florida, on shipboard (Anonymous, 1939): This is a presumptive host record, as the spiders were not reported as having been seen eating cockroaches; however, heavy infestations of both were found together.

Family LYCOSIDAE

=Lycosa helluo= Walckenaer

_Experimental prey._--Young nymphs of _Diploptera punctata_, U.S.A. (Eisner, 1958): Larger nymphs and adults repelled the spider by ejecting a repellent secretion, which has been identified as a mixture of _p_-benzoquinone and its derivatives by Roth and Stay (1958).

=Lycosa= sp.

_Experimental prey._--_Supella supellectilium_, U.S.A. (Roth and Willis, unpublished data, 1953): The lycosid (pl. 30, B-E) was probably _L. avida_ Walckenaer (tentatively identified by Dr. B. J. Kaston from a photograph).

Order ACARINA

Family PHYTOSEIIDAE

=Blattisocius tineivorus= (Oudemans)

_Synonymy._--_Blattisocius triodons_ Keegan [Baker and Wharton, 1952].

_Natural host._--_Blattella germanica_, U.S.A. (Keegan, 1944): Three mites found on 238 cockroaches examined; others taken in debris from floor of cockroach cage (Keegan, 1944). Members of this family are predaceous (Baker and Wharton, 1952).

Family LAELAPTIDAE

=Blattilaelaps nauphoetae= Womersley

_Natural host._--_Nauphoeta cinerea_, Australia, Brisbane (Womersley, 1956).

=Coleolaelaps= (?) sp.

_Natural host._--_Gromphadorhina portentosa_, the hosts were imported into U.S.A. from Madagascar via Europe (Roth and Willis, unpublished data, 1958): The mites (pl. 12, C) were tentatively determined by Dr. E. W. Baker.

=Hypoaspis= sp.

_Natural host._--_Panesthia australis_, imported into U.S.A. from Australia (Roth and Willis, unpublished data, 1955): Cockroach determined by J. A. G. Rehn. Generic determination of mite made by Dr. R. W. Strandtmann (Camin, personal communication, 1955).

Family UROPODIDAE

=Uropoda= sp.

_Natural host._--_Blattella humbertiana_, Formosa (Takahashi, 1940). Nymphs of the cockroach may be destroyed (Takahashi, 1940). Uropodids frequently attach themselves to insects, especially in nymphal stages but probably are harmless (Baker and Wharton, 1952).

Family DIPLOGYNIIDAE

=Undetermined diplogyniid=

_Natural host._--_Panesthia australis_, imported into U.S.A. from Australia (Roth and Willis, unpublished data, 1955): Cockroach determined by J. A. G. Rehn. According to Dr. J. H. Camin (personal communication, 1955) this is a new genus and new species in the subfamily Diplogyniinae, and is most closely related to the genus _Lobogynioides_. Mites of this family live as ectoparasites and commensals on beetles and possibly other insects (Baker and Wharton, 1952).

Family ANOETIDAE

=Histiostoma feroniarum= (Dufour)

_Natural host._--_Pycnoscelus surinamensis_, Germany (Roeser, 1940): Though not parasitic, the mites at times became so numerous that the insects were hindered in their movement, were unable to feed, and died. The mites were introduced with soil and leaves and had originally been attached to millipedes, waterfleas, and sowbugs.

The deutonymphs, hypopial forms, or travelers are found on insects; the other stages are found in decaying organic matter (Baker and Wharton, 1952).

Family ACARIDAE

=Caloglyphus spinitarsus= (Hermann)

_Natural host._--_Pycnoscelus surinamensis_, Germany (Roeser, 1940): See notes following _Histiostoma feroniarum_ above.

=Caloglyphus= sp.

_Natural hosts._--_Blattella germanica_ and _Periplaneta americana_, U.S.A. (Piquett and Fales, 1952): Mite feeds on organic matter but can reduce the vigor of a cockroach colony.

=Tyrophagus lintneri= (Osborne)

_Common name._--Mushroom mite.

_Associate._--_Pycnoscelus surinamensis_, U.S.A. (Roth and Willis, unpublished data, 1953): Mite determined by Dr. E. W. Baker (personal communication, 1953). Although this mite was found on the cockroach, it is a known pest in stored foods (Baker and Wharton, 1952) and probably was brought into the culture with food. Rau (1924) reported that the food of _Blatta orientalis_ often became infested with this species, but it did not affect the health or mortality of the cockroaches in his culture.

=Tyrophagus noxius= A. Z.