Chapter X: Section III: (p. 14). Other associations may be accidental (e.g., certain (4)

_Synonymy._--_Entedon hagenowii_ Ratzeburg, _Blattotetrastichus hagenowii_ (Ratzeburg) [Burks, 1943]. _Tetrastichodes asthenogmus_ Waterston. G. J. Kerrich (personal communication, 1957) compared the type of _Tetrastichodes asthenogmus_ Waterston with authentically determined material of _Tetrastichus hagenowii_ and concluded that _T. asthenogmus_ is only a weakly developed specimen of _T. hagenowii_. He stated, "The longitudinal dorsal grooves of the scutellum, which are strongly developed in normal _hagenowii_, are only rather faintly developed in Waterston's type and also the second specimen, which was dissected and mounted on a series of ten microscope slides. No doubt it was this faint development that caused Waterston to describe the species in _Tetrastichodes_, a segregate that has since been recognized by Dr. Burks (Proc. U. S. Nat. Mus., 1943) as being not truly generically distinct from _Tetrastichus_."

_Natural hosts._--_Blatta orientalis_, Seychelles (Ratzeburg, 1852); India (Usman, 1949).

_Blatta_ sp., U.S.A., Louisiana (Gahan, 1914).

_Blattella germanica_ (Burks, 1943; Peck, 1951). [In personal communications, Burks and Peck cite Howard (1892) and Marlatt (1902, and the 1908 revision of 1902) as sources for this host record. However, _B. germanica_ is not mentioned specifically as a host of _T. hagenowii_ in the sources cited nor in the 1915 revision of Marlatt's 1902 paper cited by Burks (1943); see footnote 6, p. 236.]

_Neostylopyga rhombifolia_, Hawaii (Pemberton, 1941): This record is based on one parasitized oötheca. We have exposed, at three different times, groups of 10 to 20 oöthecae of _N. rhombifolia_ to many newly emerged _T. hagenowii_, but none of the eggs was parasitized (Roth and Willis, unpublished data, 1957).

_Parcoblatta_ sp., U.S.A., Ohio (Edmunds, 1953a).

_Periplaneta americana_, Africa (Crawford, 1910; Nash, 1955): Nash's record was incorrectly attributed to _Syntomosphyrum glossinae_ Wtstn., a parasite of tse-tse fly pupae (Jordan, 1956), Formosa (Takahashi, 1924; Sonan, 1924); Palestine (Bodenheimer, 1930); Puerto Rico (Seín, 1923; Plank, 1947, 1950; Wolcott, 1951); St. Croix, Virgin Islands (Beatty, 1944); Hawaii (Schmidt, 1937); U.S.A.: Missouri (Rau, 1940a); Ohio (Edmunds, 1955); Florida (parasitized oöthecae were collected near Orlando by members of the Orlando Laboratory, Entomology Research Branch, U.S. Department of Agriculture; the parasites were identified by Burks, personal communication, 1955). Fiji (Lever, 1943); India (Mani, 1936; Usman, 1949); Trinidad and Saudi Arabia (Cameron, 1955). Westwood (1839) stated that 70 parasites belonging to the genus _Eulophus_ emerged from an oötheca of _P. americana_ collected on shipboard. Burks (personal communication, 1955) stated that the wasp was probably _T. hagenowii_.

_Periplaneta australasiae_, Australia (Shaw, 1925); India (Usman, 1949); Saudi Arabia, Trinidad (Cameron, 1955); Formosa (Sonan, 1924).

_Periplaneta brunnea_, U.S.A., Florida (parasitized oöthecae were collected near Orlando, by members of the Orlando Laboratory, Entomology Research Branch, U.S. Department of Agriculture. The parasites were identified by Burks, p. c., 1955).

Cockroach eggs, Formosa (Maki, 1937); Ceylon (Waterston, 1914): Taken on an oötheca.

"Domestic cockroaches," U.S.A., Louisiana (Girault, 1917).

"Roach egg cases," Panama Canal Zone (Rau, 1933).

_Evania_ sp., Hawaii (Ashmead, 1901; Perkins, 1913); Guam (Fullaway, 1912); Fiji (Lever, 1946); Europe, Cuba, Florida (Marlatt, 1902, 1915).

_Experimental hosts._--_Blatta orientalis_, _Eurycotis floridana_, and _Periplaneta americana_, U.S.A. (Roth and Willis, 1954b): We have maintained _T. hagenowii_ for over two years through more than 30 generations on eggs of both _B. orientalis_ and _P. americana_.

_Periplaneta fuliginosa_, U.S.A., Pennsylvania (Roth and Willis, 1954b); Massachusetts (Roth and Willis, unpublished data, 1957).

Schmidt (1937) deduced that _T. hagenowii_ was a primary parasite of eggs of _P. americana_ because the parasitized oötheca was obtained from a cage covered with screen too fine to permit entry of a larger parasite, such as an evaniid. As noted above, we have reared _T. hagenowii_ for more than 30 generations on cockroach eggs, none of which was ever exposed to parasitization by an evaniid. If _T. hagenowii_ were ever hyperparasitic on _Evania_, this relationship would be accidental, the eulophid happening to oviposit into an oötheca already containing an evaniid, or vice versa.

_Adult behavior._--The male mates soon after becoming adult; he mounts the female from behind, grasps her antennae with his own antennae, and vibrates his wings during copulation. Mating is accomplished in from "several" to 20 seconds (Takahashi, 1924; Edmunds, 1955). The adults are positively phototactic and are capable of hopping for some distance (Edmunds, 1955). The females feed on material that oozes through the oviposition puncture (Roth and Willis, 1954b). Females lived 10 days (Seín, 1923). Without food, females lived 7.8 days and males 3.4 days, but when fed dilute honey females lived 12.5 days (Usman, 1949). Females lived 5-11 days (Roth and Willis, 1954b). Fed water and sugar, the wasps lived 2-6 weeks at 65°F. (Cameron, 1955). Without food, 9 females lived an average of 3.5 days and 9 males an average of 1.7 days, but when fed on raisins, 9 females lived an average of 25 days and 9 males 15 days (Edmunds, 1955). In Formosa there were six generations from April to December (Maki, 1937).

In Hawaii, Severin and Severin (1915) caught 571 _T. hagenowii_ in 10 kerosene traps that were set up to sample populations of Mediterranean fruitfly. Apparently the parasite is attracted by the odor of kerosene.

_Oviposition._--The female wasp explores the surface of the oötheca with vibrating antennae (Edmunds, 1955). She bends her abdomen ventrad and repeatedly touches the surface of the oötheca with her valvae; when she finds an acceptable oviposition site, the wasp unsheathes her ovipositor and bores through the wall of the oötheca (Roth and Willis, 1954b). The wasp deposited her eggs in 2-5 minutes (Edmunds, 1955). Wasps oviposited (pl. 34, C) into young or old eggs of _P. americana_ (Roth and Willis, 1954b). A single wasp parasitized more than one oötheca and more than one wasp oviposited into the same oötheca (Roth and Willis, 1954b; Edmunds, 1955). We found freshly laid wasp eggs in 34 empty but previously parasitized oöthecae from which the wasps had emerged (Roth and Willis, 1954b).

_Development._--In _Periplaneta americana_: Development is completed in an average of 36 days (range 29-58 days) (Maki, 1937); 29-40 days (Lever, 1943); average of 23.6 days (range 22-26 days) at 62°-85° F. (Usman, 1949); about 3 months at 60°-65° F. (Cameron, 1955); 31-60 days at 70°-80° F. (Edmunds, 1955). We found that the wasps completed development in 23-56 days at about 85° F., but the period depended on the number of wasps in the oötheca; the larger the number of wasps (up to an average of about 70 wasps per oötheca), the shorter the time required to complete development. Wasps in oöthecae containing 70 or more parasites developed in an average of about 32 days (Roth and Willis, 1954b). Wasp larvae eat the contents of the cockroach egg in which they start development, then rupture the chorion and attack adjoining eggs (Cameron, 1955; Edmunds, 1955). All eggs are consumed when the parasite density is high, but if too few larvae develop per oötheca, some cockroach eggs survive and the embryos complete development (Roth and Willis, 1954b). However, a certain number of cockroach nymphs must complete development to enable the survivors to force open the crista and emerge from the oötheca; fewer than this number of surviving nymphs will be trapped and killed as effectively as if they had been eaten by the parasite. The adult parasites emerge from one to three holes cut through the wall of the oötheca (Usman, 1949; Roth and Willis, 1954b).

_Number of offspring per female._--In _Blatta orientalis_: In the laboratory, 5 oöthecae were left with each of 25 female wasps for their entire lifespans; of the 125 oöthecae, 32 were parasitized. The average number of offspring per female was 66 (range 5-164) (Roth and Willis, 1954b). In _Periplaneta americana_: Each of 206 oöthecae was exposed to a single female wasp for 24 hours; the average number of offspring per female was 103 (range 50-139). Five oöthecae were left with each of 38 females for their entire lifespans; of the 190 oöthecae, 81 were parasitized. The average number of offspring per female was 94 (range 45-168 [from original data]) (Roth and Willis, 1954b).

_Number of parasites per oötheca._--In _Eurycotis floridana_: In the laboratory, 3 oöthecae that had been exposed to 20 female wasps yielded an average of 648 parasites (range 606 [from original data] to 685) (Roth and Willis, 1954b). In _Neostylopyga rhombifolia_: One oötheca yielded 73 parasites (Pemberton, 1941). In _Parcoblatta_ sp.: Two oöthecae yielded an average of 100 parasites (Edmunds, 1953a). In _Periplaneta americana_: 100 parasites per oötheca (Seín, 1923); 140 (Schmidt, 1937); 25 (Rau, 1940a); 7-38, average 33 (Usman, 1949); 71 (Wolcott, 1951); 4 oöthecae exposed to 20 female wasps yielded an average of 204 wasps (range 164 [from original data] to 261) (Roth and Willis, 1954b); average of 30-40 (Cameron, 1955); 39 oöthecae yielded an average of 93 parasites (range 12-187) (Edmunds, 1955). In _Periplaneta australasiae_: Oöthecae yielded an average of 40-50 adult parasites (Cameron, 1955); about 50 (Shaw, 1925).

_Sex ratio._--3 [F][F]:1 [M] (Usman, 1949); 4 [F][F]:1 [M] (Cameron, 1955); 2-8 [F][F]:1 [M] (Roth and Willis, 1954b); 1.2 [F][F]:1 [M] (Edmunds, 1955). Parthenogenesis exists; the unfertilized eggs produced only males (Roth and Willis, 1954b; Edmunds, 1955).

_Distribution._--Probably worldwide. Eastern and southern U.S.A.; Central and South America; Europe; Arabia; Africa; India; Formosa; Hawaii.

=Tetrastichus periplanetae= Crawford

_Natural hosts._--_Periplaneta americana_, Mozambique (Crawford, 1910); Union of South Africa (parasites reared from oöthecae collected in Durban, Natal, by the City Health Department): The parasites were identified by Burks (personal communication, 1956). Jamaica (Gowdey, 1925); Réunion Island (Bordage, 1913).

"Domestic cockroach," Puerto Rico (Wolcott, 1951).

=Tetrastichus= sp. I

_Taxonomy._--Burks (personal communication, 1956) stated that this species (specimens of which are in the U.S. National Museum) is very close to _T. hagenowii_.

_Natural hosts._--_Periplaneta americana_, Union of South Africa (parasites reared from oöthecae collected in Durban, Natal, by the City Health Department [Burks, p. c., 1956]).

_Periplaneta australasiae_, Manila, Philippine Islands (Burks, p. c., 1956).

=Tetrastichus= sp. II

_Synonymy._--Because of the war, Cros (1942) could not determine this insect specifically. He designated it provisionally and with reserve under the name _Eulophus_ sp. However, Burks (p. c., 1956) stated that the species is most certainly a _Tetrastichus_ from the description given; but, it is apparently not _T. hagenowii_ because of its brilliant steel-blue color.

_Natural host._--_Blatta orientalis_, Algeria (Cros, 1942): _Adult behavior._--Mating began as soon as wasps emerged from an oötheca. Males mated repeatedly. Adults lived up to 5 days in summer and up to 12 days in fall. There were up to four generations per year in the laboratory. _Oviposition._--Wasps oviposited into oöthecae 6, 22, 40, and 43 days old, and the parasites developed successfully. More than one female oviposited into the same oötheca. Oviposition was of long duration. _Development._--From egg to eclosion took an average of 34 days in summer (range: 30-38 days, 5 oöthecae), and an average of 67 days in fall (range: 58-73 days, 3 oöthecae). An average of 55 parasites developed per oötheca (range 21-105, 5 oöthecae); over 130 wasps emerged from a sixth oötheca. _Sex ratio._--10-20 [F][F]:1 [M].

HOST SELECTION BY EGG PARASITES

The nature of the oviposition stimulus(i) for the wasp parasites of cockroach eggs is unknown. Edmunds (1954) noted that _Prosevania punctata_ showed more interest in oöthecae that had been cemented to the substrate than in clean oöthecae that had simply been dropped. Cros (1942) experimented with two females of _P. punctata_ to see if the wasps could find oöthecae that had been buried in sand by the oriental cockroach. After prospecting the sand with their antennae, the wasps dug deep excavations with their front legs but always mistook the location of the oöthecae. Cros suggested that the wasps were misled by the odor left in the jar by the cockroaches. It is quite possible that odor helps the wasp find the host oötheca.

The extent of host selection varies among these parasites; some species will oviposit into the eggs of more than one species of cockroach, but others show some degree of host specificity. Positive selection of specific hosts by certain parasites appears in correlative data from different investigators on pages 235 to 254. There is a small body of data that shows nonacceptance of certain hosts by some of these wasps. For example, _Comperia merceti_ would not parasitize eggs of _Blatta orientalis_ or _Periplaneta americana_ in the laboratory (Lawson, 1954a). We (unpublished data, 1957) exposed a soft oötheca, recently removed from _Eurycotis floridana_, to _C. merceti_; no wasps developed; we had similar negative results with _C. merceti_ and oöthecae of _B. germanica_. We (1954b) could not induce _Tetrastichus hagenowii_ to parasitize eggs of _Blattella germanica_, _B. vaga_, or _Parcoblatta virginica_ in the laboratory. In our experiments, _T. hagenowii_ oviposited into eggs of _Supella supellectilium_, but the wasp eggs either failed to hatch, or if they hatched, the larvae died before completing development. Neither would _T. hagenowii_ parasitize eggs of _N. rhombifolia_ (Roth and Willis, unpublished data, 1957). _Anastatus tenuipes_ would not parasitize the eggs of _Latiblattella lucifrons_ Hebard, _Periplaneta americana_, _B. germanica_, or _B. vaga_ (Flock, 1941). _Anastatus floridanus_ would not oviposit into eggs of _S. supellectilium_ and only rarely into eggs of _P. americana_ or _B. orientalis_ (Roth and Willis, 1954a); in the laboratory, this wasp could not be maintained beyond one generation on the eggs of _P. americana_. Edmunds (1953b) could not induce _Prosevania punctata_ to parasitize eggs of _B. germanica_. Cros (1942) induced _P. punctata_ to oviposit into a mantid oötheca, but neither mantids nor parasite developed.

COCKROACH-HUNTING WASPS

A number of wasps of the families Ampulicidae, Sphecidae, and a very few species of Pompilidae have been found to provision their nests with nymphal or adult cockroaches. This habit of preying on cockroaches is primitive (Leclercq, 1954); Leclercq (personal communication, 1955) stated that this habit is always associated with the conservation of a number of structures considered as archaic from a purely morphological point of view.

The records of wasps of the genus _Astata_ capturing cockroaches (e.g., Sickmann, 1893; St. Fargeau _in_ Sharp, 1899) "all trace back to a questionable record by Lepeletier (1841) which probably was a misidentification of the predator" (K. V. Krombein, personal communication, 1956). Marshall (1866) suggested that the braconid _Paxylomma buccata_ Bréb., which he found frequenting cockroach runs in Pembrokeshire, was parasitic on _Ectobius nigripes_ Stephens; however, this wasp is undoubtedly parasitic on ants, probably on ant larvae (Donisthorpe and Wilkinson, 1930).

The wasps that are known to capture cockroaches, and summaries of their biology, are listed below.

WASPS THAT PROVISION THEIR NESTS WITH COCKROACHES

Family POMPILIDAE

=Pompilus bracatus= Bingham

_Natural hosts._--Cockroaches, India (Bingham, 1900).

=Pompilus= sp.

_Natural host._--Cockroach, Nyasaland (Lamborn _in_ Poulton, 1926): The wasp was collected leading a nymph of the cockroach by its antenna. The cockroach was in a stupefied state, and its antennae were bitten off to about half their length.

=Salius verticalis= Smith

_Natural hosts._--Cockroaches, India (Bingham, 1900).

Family AMPULICIDAE

The species of _Ampulex_ do not appear to make special nests in which to lay their eggs but drag their prey to any convenient hole, or crack in the ground (Arnold, 1928). Although many species of _Ampulex_ have been described, the prey of only a small number of species have been discovered, but the known prey are all cockroaches.

=Ampulex amoena= Stål

_Synonymy._--_Ampulex novarae_ Saussure [Krombein, personal communication, 1957].

_Natural hosts._--_Periplaneta americana_ and _Periplaneta australasiae_, both as small nymphs, Formosa (Sonan, 1924, 1927): The wasp stings a nymph about one inch long and carries it to a suitable place, such as bamboo pipes, folds of newspaper, or books (in houses), for oviposition.

_Periplaneta picea_, Japan (Kamo, 1957; Kohriba, 1957).

_Experimental hosts._--_Periplaneta picea_, Japan (Kamo, 1957; Kohriba, 1957).

Kamo (1957) observed that in the field both males and females sucked juices from wounds they made in the stems of _Clerodendron_ _trichotomum_ Thunberg or _Ilex rotunda_ Thunberg. Kohriba (1957), on the other hand, found both sexes sucking sap of _Abies_ sp. and other trees from points injured by the rostrum of cicadas. Kamo (1957) observed that the female wasp grasped the cockroach by a tergum and stung it several times in the thorax. The wasp always amputated the antennae of the prey and sucked up the fluid oozing from the cut antennae. The wasp egg was placed on the mesocoxa of the cockroach. In the laboratory as many as three cockroaches, each with a wasp egg, were stored in artificial nests per day. Kohriba (1957) observed similar behavior in the laboratory and made these additional notes. The paralyzed cockroach could move its legs and was led to the nest by the wasp which seized its antennae. The egg hatched in 2 days, and after sucking up body fluid for 2 days the larva began to devour the prey. Three days later the larva spun its cocoon, and about one month after spinning a female wasp emerged.

=Ampulex assimilis= Kohl

_Natural hosts._--_Blatta lateralis_, wingless females, Iraq (Hingston, 1925): Nesting sites are holes in palm trees, galleries of beetles, or tunnels in ground. The wasp first seizes a cockroach by the edge of its thorax and stings it in the thoracic region, then seizes the cockroach by an antenna and pulls and leads it to the nest. The wasp deposits her egg on the outer surface of the femur of the cockroach's midleg. The nest is closed with debris; later the cockroach recovers from the sting. The wasp larva first feeds externally, then bores into the cockroach and devours the internal organs. Pupation occurs inside the exoskeleton of the cockroach.

=Ampulex canaliculata= (Say)

_Synonymy._--_Rhinopsis caniculatus._

_Natural hosts._--_Ischnoptera_ sp., U.S.A. (Krombein, 1951).

_Lobopterella dimidiatipes_, Hawaii (Williams, 1928a, 1929).

_Parcoblatta pensylvanica_? MacNay (1954) referred to a rare sphecoid wasp in eastern Canada which provisioned its nest with nymphs and adults of _P. pensylvanica_. Dr. W. R. M. Mason (personal communication, 1957) wrote us that although this wasp was _Ampulex canaliculata_, it was not reared from the cockroach but was swept from a pine tree. There are no positive records linking _A. canaliculata_ with _P. pensylvanica_.

_Experimental host._--_Parcoblatta virginica_, females, U.S.A., Missouri (Williams, 1928a, 1929): figure 6.

Nesting sites are in twigs (Krombein, 1951). The adult behavior is similar to that of _A. compressa_; the female wasp imbibes blood that oozes from the amputated antennae of the cockroach; the egg hatches in 2-3 days, and the development of one male was completed in 33 days (Williams, 1929).

, through the courtesy of Dr. F. X. Williams and F. A.
Bianchi, Hawaiian Sugar Planters' Association.)]

_Distribution._--U.S.A.: Connecticut south to Georgia; Ohio, Wisconsin, Missouri, Kansas; in open woods (Krombein, 1951).

=Ampulex compressa= (Fabricius)

(Pl. 35)

_Synonymy._--Guĕpe ichneumon of Réaumur [Williams, 1929]; _Chlorion_ (_Ampulex_) _compressum_.

_Natural hosts._--_Periplaneta americana_, New Caledonia (Lucas, 1879); India (Dutt, 1912); Reunion (Bordage, 1912).

_Periplaneta australasiae_, Hawaii (Swezey, 1944).

_Periplaneta_ sp., India (Maxwell-Lefroy, 1909).

Cockroach. Mauritius (Réaumur, 1742); Burma (Bingham, 1897).

_Experimental hosts._--_Neostylopyga rhombifolia_, _Periplaneta americana_, and _Periplaneta australasiae_, Hawaii (Williams, 1942, 1942a). Zimmerman's (1948) listings probably were taken from Williams.

_Nesting sites._--Holes in walls; holes in banyan and fig trees; in houses in drawers and cartons. _Behavior._--Similar to that of _A. assimilis_. Bordage (1912) gives a complete description of capture of prey. The female wasp cuts off part of the cockroach's antennae, legs, and wings; she sticks her egg onto the host's mesothoracic coxa. The wasp frequents houses in search of prey. Five [F][F], supplied with a cockroach per day, stored an average of 57±14 cockroaches; 8 [F][F] stored an average of 45±3 cockroaches; these latter wasps were not supplied with a cockroach per day throughout (mean values computed from Williams, 1942). This wasp will not attack _Nauphoeta cinerea_ (Williams, 1942a) or _Pycnoscelus surinamensis_ (Schwabe, 1950b). On one occasion, _A. compressa_ stung _Diploptera punctata_, but did not oviposit (Williams, 1942a). _Development._--Minimum 34 days, maximum 140 days (Williams, 1942). About 6 weeks (Swezey, 1944). _Longevity of adults._--13 [F][F] lived an average of 110±11 days (minimum 31, maximum 159); several [M][M] lived 2 months (Williams, 1942).

=Ampulex fasciata= Jurine

_Natural host._--_Ectobius pallidus_, France (Picard, 1911, 1919): Nesting sites are in brier or bramble stems, or in crevices in fig trees; the female possibly uses old nests of leaf-cutter bees. The feeding of the wasp larva is similar to that of other _Ampulex_. Adult wasp emerges by cutting open a passage through its cocoon and through the anus of the cockroach.

=Ampulex ruficornis= (Cameron)

_Natural hosts._--Cockroaches, Oriental region (Rothney _in_ Sharp, 1899): Nesting sites are in crevices in bark. The female grasps the cockroach by an antenna to drag it to her nest.

=Ampulex sibirica= Fabricius

_Synonymy._--Perkins referred to this species as _Ampulex sibirica_. Williams (1942a), referring to Perkins's observations, mentions the species as "_A. compressiventris_ Guérin (=_A. siberica_ Sauss.)." Krombein (personal communication, 1956) has commented upon this synonymy as follows: _Ampulex siberica_ Sauss. is apparently a misidentification by Saussure of _sibirica_ Fab. Kohl (1893) in his revision of the genus _Ampulex_ considered _A. compressiventris_ Guérin to be the correct name for this common African species and that _sibirica_, described from Siberia, must be another species. However, Turner (1912) stated that he had seen Fabricius's type specimen and that it was identical with what had been called _compressiventris_; he considered the Siberian locality given by Fabricius as an error. Krombein suggested that Williams's use of the combination _siberica_ Sauss. was a lapsus and that the valid name, if Turner is correct, is _sibirica_ Fab.

_Natural hosts._--Cockroaches, West Africa (Perkins _in_ Sharp, 1899): Nesting sites are keyholes. Enters apartments in search of cockroaches. Wasp cocoon protrudes from dead body of cockroach.

=Ampulex sonnerati= Kohl

_Synonymy._--"La mouche bleue" of Sonnerat (Kohl, 1893).

_Natural host._--"Kakkerlac," Philippine Islands (Sonnerat, 1776): Nesting sites are readymade crevices. The wasp seizes the cockroach by an antenna and stings the host many times in the "abdomen." She drags the cockroach by an antenna to the nest, and, after depositing her egg, plugs the opening with moistened earth.

=Dolichurus bicolor= Lepeletier

_Synonymy._--Schulz (1912) considered this to be _Dolichurus corniculus_. Berland (1925) stated that this is possibly a color variety of _D. corniculus_. Soyer (1947), from a study of the behavior of the wasps, believed that both _D. bicolor_ and _D. haemorrhous_ are varieties of _D. corniculus_. Krombein (personal communication, 1956) stated that _D. corniculus_ and _D. bicolor_ differ in characters other than color alone and that _D. bicolor_ is considered a valid species today.

_Natural host._--Cockroach, France (Benoist, 1927): The wasp was observed closing the entrance to its burrow. Its egg was attached to the coxa of the midleg of the cockroach.

Maneval (1932) stated that _D. bicolor_ is found at the edge of dry woods along with _D. corniculus_ and that the wasp will also accept the prey of _D. corniculus_ if presented to it.

=Dolichurus corniculus= (Spinola)

_Synonymy._--_Dolichurus haemorrhous_ Costa [Schulz, 1912]. Berland (1925) listed _D. haemorrhous_ separately but stated that it is perhaps a color variety of _D. corniculus_.

_Natural hosts._--_Blattella germanica_, France (Benoist, 1927).

_Ectobius lapponicus_, Germany (Sickmann, 1893); Denmark (Nielsen, 1903); Sweden (Adlerz, 1903); Italy (Grandi, 1931, 1954); France (Benoist, 1927; Maneval, 1928).

_Ectobius pallidus_, France (Maneval, 1932; Soyer, 1947).

_Ectobius panzeri_, France (Soyer, 1947).

_Ectobius_ sp., Italy (Grandi, 1954).

_Hololampra punctata_, Pitten (Handlirsch, 1889).

_Loboptera decipiens_, France (Ferton, 1894).

Cockroach, Netherlands (Bouwman, 1914).

_Nesting sites._--The wasp uses already-made cavities such as rotting dead branches on ground, fissures in the earth, abandoned ant holes, chinks in stone, or the empty cocoon of the ichneumon _Ophion luteus_ (Ferton, 1894; Maneval, 1932).

_Behavior._--The prey is immobile while being dragged to the nest but recovers sufficiently from the sting so that if dug up it will run around (Ferton, 1894; Bouwman, 1914; Benoist, 1927; Grandi, 1954). The wasp cuts off about two-thirds of the cockroach's antennae prior to putting its prey in its nest (Adlerz, 1903; Bouwman, 1914; Soyer, 1947). One cockroach is placed in the nest and the wasp's egg is attached to the midcoxa (Ferton, 1894). Oviposition takes 5 to 6 minutes (Maneval, 1939). Wasp fills and seals its nest with bits of earth and stones (Ferton, 1894; Grandi, 1954). The wasp larva feeds externally and devours the entire cockroach, including its exoskeleton (Ferton, 1894).

_Development._--Hatching occurs in 3 to 4 days (Ferton, 1894) or longer during cooler weather (Maneval, 1939). Larval development takes 6 days (Grandi, 1954), 8 days (Ferton, 1894), or 10 to 25 days depending on season (Maneval, 1939).

=Dolichurus gilberti= Turner

_Natural hosts._--"Small Blattidae," India (Turner, 1917).

=Dolichurus greenei= Rohwer

_Natural host._--_Parcoblatta_ sp., U.S.A., Virginia (Krombein, 1951, 1955): Nesting sites are under leaf litter. The prey was a paralyzed third-instar nymph. _Distribution._--Ontario. U.S.A. from Canadian border south to Florida in coastal States (Krombein, 1951).

=Dolichurus ignitus= Sm.

_Natural hosts._--Cockroaches, Natal and Southern Rhodesia (Arnold, 1928): The wasp is "usually seen running up and down the trunks of trees searching for small cockroaches in the crevices of the bark."

=Dolichurus stantoni= (Ashmead)

_Natural hosts._--_Allacta similis_, nymphs, Hawaii (Williams et al., 1931; Zimmerman, 1948).

_Blattella lituricollis_, usually nymphs, Philippine Islands, Hawaii (Williams, 1919).

_Cutilia soror_, nymphs, Hawaii (Williams et al., 1931; Zimmerman, 1948).

"_Phyllodromia_" sp., Philippine Islands, Hawaii (Williams, 1918; Bridwell, 1920).

_Experimental hosts._--"Field cockroaches," Philippine Islands (Williams, 1944).

_Nesting site._--Readymade crevices or holes in ground; porosity in lava. _Behavior._--The wasp seizes the cockroach by a cercus or leg and stings it in the thorax. She (fig. 7, A) then drags the cockroach to the nest by the base of an antenna. Wasp bites off distal part of host's antennae. She deposits her egg on one of the host's midcoxae. Nest is plugged with lumps of soil. The larva eats the entire host. _Development._--Eggs hatched in about a day and a half. Adults emerged about 3 weeks later. About five generations per year. (Williams, 1918, 1919; Williams et al., 1931.)

=Dolichurus= sp.

_Natural hosts._--Cockroaches, nymphs, South Africa (Bridwell, 1917). Adult female cockroach carrying an oötheca, France (Deleurance, 1943).

_Nesting site._--Plant stem, or in ground possibly an old abandoned nest of _Ammophile_. _Behavior._--Bridwell noted that one wasp larva ate two cockroach nymphs before pupating; the adult emerged about 4 months after cocoon formation. Deleurance observed the wasp close its nest with small pebbles, balls of earth, and small dead branches. The wasp egg was placed on the femur of the midleg. The prey in the nest is alert when disturbed. Deleurance believed the wasp was a variety of _D. corniculus_.

=Trirhogma caerulea= Westwood

_Natural hosts._--_Periplaneta americana_ and _Periplaneta australasiae_, Formosa (Sonan, 1924): The wasp stings a nymph about one inch long and carries it to a suitable place (bamboo pipe) for oviposition.

.) B, _Podium haematogastrum_ attaching her
egg to an _Epilampra_ sp. while on the side of a termite mound that
contains the wasp's nest, c. × 1.6. C, _Epilampra_ sp. parasitized
by _P. haematogastrum_ showing the wasp's egg attached to the right
fore coxa, c. × 3.2. (B and C reproduced from Williams [1928],
through the courtesy of Dr. F. X. Williams and F. A. Bianchi.)]

=Trirhogma= sp.

_Natural hosts._--Cockroaches, Oriental region (Williams, 1918, 1928): As far as is known species of this genus of wasps hunt cockroaches.

Family SPHECIDAE

=Tachysphex blatticidus= Williams

_Natural hosts._--_Chorisoneura_ sp., adults, Trinidad, St. Augustine (Callan, 1942): The wasps nest gregariously in sandy places. The wasp itself is parasitized by the mutillid _Timulla_ (_Timulla_) _eriphyla_ Mickel.

Cockroaches, Trinidad (Williams, 1941a; Callan, 1950).

=Tachysphex coriaceus= Costa

_Natural hosts._--Cockroaches, Italy (Beaumont, 1954).

=Tachysphex fanuiensis= Cheesman

_Natural hosts._--_Graptoblatta notulata_, Society Islands (Cheesman, 1927, 1928).

Cockroach ("except for its smaller size [it] much resembles _Graptoblatta notulata_."), New Caledonia (Williams, 1945).

_Nesting sites._--Patches of dry soil (Cheesman, 1928); coarse sand at base of a bank (Williams, 1945). _Behavior._--The female wasp pounces on the cockroach and stings it into immobility; she carries her prey in flight to the nest. Two to 13 cockroaches may be found in one nest; and one or more wasp eggs may be deposited in one nest. The egg is attached at one end to the host's thorax behind a forecoxa. Nest is sealed with dry pellets of soil. The cockroaches apparently do not recover from the wasp's sting.

=Tachysphex lativalvis= (Thomson)

_Natural hosts._--_Ectobius lapponicus_, adults, Sweden (Adlerz, 1906); France (Maneval, 1932).

_Ectobius pallidus_, nymphs, France (Ferton, 1894, 1901; Maneval, 1932; Deleurance, 1946); Italy (Grandi, 1928).

_Ectobius panzeri_, Netherlands (Bouwman, 1914).

_Ectobius_ sp., Denmark (Nielsen, 1933).

Ferton (1914) stated that he had reported in 1912 that this species hunted Hemiptera, but that this observation was a lapsus. _Nesting site._--In the ground of sandy woodlot or border of dry woods; the nest is a hole 5.5 to 8 cm. long ending in a horizontal cell. Grandi (1928) stated that the entrance to the nest descended obliquely for 5 to 6 cm. and ended 4 cm. below the surface of the ground. _Behavior._--Two cockroaches, either sex, adults or nymphs, were stored in the cell (Adlerz, 1903; Grandi, 1928). The wasp laid her egg on the first prey brought, attaching it behind the front coxa. The cockroaches were not excitable and their antennae had not been injured. Grandi (1928) stated that the claws of the hind tarsi of the victims may be amputated. The hatched larva may consume one of its victims in four days leaving only the head, pronotum, tegmina, wings, and the urosternum.

=Podium abdominale= (Perty)

_Synonymy._--_Trigonopsis abdominalis_ Perty [Kohl, 1902].

_Natural hosts._--Cockroaches, nymphs, Ecuador (Williams, 1928): These wasps are apparently mainly arboreal mud daubers. The female wasp constructs a mud nest on underside of a palm leaf. Wasp egg is attached behind one of the forecoxae of the cockroach. Several cockroaches are stored in each nest. The prey is not immobilized as a result of the sting, and its antennae are left intact.

=Podium carolina= Rohwer

_Natural host._--_Parcoblatta pensylvanica_, nymphs (Rau, 1937): Nesting sites are mud nests of _Sceliphron caementarium_ (Drury). One to three cockroach nymphs are stored per nest; mud partitions are placed in tube; the nest is plugged with mud which is coated with resin. _Distribution._--U.S.A., New York to North Carolina (Murray, 1951); Florida (Krombein and Evans, 1955).

=Podium dubium= Taschenberg

_Natural hosts._--Epilamprine cockroaches, Brazil (Williams, 1928): Burrows, lenticular in cross section, are found on shaded trails. The wasp's habits are similar to those of _P. flavipenne_ and _P. haematogastrum_.

=Podium flavipenne= Lepeletier

_Natural host._--_Epilampra abdomen-nigrum_, British Guiana (Williams, 1928): _Nesting site._--Burrows, about 2 inches deep and lenticular in cross section, are dug in the ground in well-drained, partly sheltered areas; also old _Podium_ nests are used. _Behavior._--The wasp stings the cockroach to helplessness and flies with it back to her nest where the host may recover from the sting; one or more cockroaches are stored per nest; the egg is deposited behind the forecoxa while the cockroach is still outside the burrow. The nest is sealed with mud. The larva feeds on most of the cockroach and leaves only some heavily sclerotized portions in the cell. In 153 nests examined, there was an average of 2.2 ± 0.08 [standard error computed from cited data] cockroaches per cell; four nests contained five cockroaches apiece. Of the 331 cockroaches in the nests, only 6 percent were adults. _Development._--Egg hatches in about 2 days; larva feeds about 4 days and pupates about 2 weeks later; adult emerges about 10-12 days later.

=Podium haematogastrum= Spinola

_Natural host._--_Epilampra_ sp., Brazil, Pará (Williams, 1928): The female wasp (fig. 7, B) burrows into the surface of termite mounds, in banks, and in level ground. This wasp's behavior is similar to that of _P. flavipenne_. There was an average of 1.6 cockroaches (fig. 7, C) per cell in 74 nests examined. Of the 121 cockroaches collected, 28 percent were adults. Under artificial conditions, the life cycle varied from about a month to 45 days or more.

=Podium luctuosum= Smith

_Natural host._--_Parcoblatta virginica_, female, U.S.A., New York (Pate, 1949).

_Distribution._--U.S.A.: New York to Texas (Murray, 1951).

=Podium rufipes= Fabricius

_Natural hosts._--"Wood roaches," British Guiana (Howes, 1917, 1919); Brazil (Williams, 1928): Nesting sites were clay column nests on houses, sides of stumps, or forest trees; banks; termite mound. Variable numbers of cockroaches were placed in the nests with one wasp egg attached behind forecoxa of the last host. The egg hatches in 2 days, the larva pupates about 2 weeks later, and the adult emerges 24 days later.

=Podium= sp.

_Natural host._--_Epilampra conferta_, Brazil (Poulton, 1917): The burrow contained several cockroaches of the same species.

ANTS PREDACEOUS ON COCKROACHES

A large roach endeavored to escape by crossing the main front of
the army. The creature made several powerful jumps, but each time
it touched the ground ... its legs were grasped by the fearless
ants.... In the end it fell ... and was instantly torn to bits and
carried to the rear.... Another ant with the body of a wood roach
was assisted by a worker who held the carrier's abdomen high in the
air out of the way of her burden, all the way to the nest.

HOWES (1919)

Family FORMICIDAE

From the known entomophagous habits of the lower ants (Wheeler, 1928), we wonder that there are not more records of ants feeding on cockroaches, because this act must occur frequently. Kirby and Spence (1822) stated that R. Kittoe had observed in Antigua that ants which nested in the roofs would seize a cockroach by the legs so it could not move, kill it, and carry it up to their nest. Hotchkiss (1874) observed ants kill cockroaches on shipboard. Cockroaches attracted to sugar in the pantry were killed and carried off by the ants. The destruction of cockroaches by army ants has been recorded by Bates (1863), Wallace (1891), Beebe (1917, 1919), Howes (1919), and others. Dead and mutilated specimens of _Ischnoptera_ sp. [undoubtedly _Parcoblatta americana_ (Gurney, personal communication, 1958)] are common in the nests of species of _Formica_ in California (Mann, 1911).

=Aphaenogaster picea= Emery

_Natural prey._--_Ectobius pallidus_, U.S.A., Massachusetts (Roth and Willis, 1957).

=Camponotus pennsylvanicus= (De Geer)

_Common name._--Carpenter ant.

_Natural prey._--_Parcoblatta pensylvanica_, U.S.A. (Rau, 1940): The ants entered traps set up to capture the cockroach and carried off about a dozen adults of both sexes.

=Dorylus (Anomma) nigricans= subsp. =sjöstedi Emery=

_Natural prey._--Small cockroach, Belgian Congo (Raignier and van Boven, 1955).

=Dorylus (Anomma) wilverthi= Emery

_Natural prey._--Small cockroaches, Belgian Congo (Raignier and van Boven, 1955).

=Dorylus= sp.

_Common name._--"Safari ant."

_Natural prey._--Cockroaches, Africa, Lake Victoria (Carpenter, 1920): When the "Safari ants" were hunting, many species of cockroaches were driven from hiding among dead leaves in the forest. The cockroaches rushed about but easily fell prey to the ants which tore them to bits.

=Eciton burchelli= (Westwood)

_Common name._--Army ant.

_Natural prey._--Cockroaches, Panama Canal Zone (Johnson, 1954; Schneirla, 1956).

=Formica omnivora=

_Synonymy_.--The identity of this form is unknown. There are no species of _Formica_ on Ceylon. There was another _Formica omnivora_ described from tropical America, whose identity is also unknown (W. L. Brown, personal communication, 1956).

_Natural prey._--Cockroaches, Ceylon (Kirby and Spence, 1822).

=Iridomyrmex humilis= Mayr

_Common name._--Argentine ant.

_Natural prey._--Cockroaches, injured individuals only (Ealand, 1915).

=Lasius alienus= (Förster)

_Natural prey._--_Ectobius pallidus_, U.S.A., Massachusetts (Roth and Willis, 1957).

=Pheidole megacephala= (Fabricius)

_Common name._--Big-headed ant.

_Natural prey._--_Holocompsa fulva_, Hawaii (Illingworth, 1916).

_Nauphoeta cinerea_ and _Pycnoscelus surinamensis_, Hawaii (Illingworth, 1914, 1942): The ants followed and killed _N. cinerea_ and _P. surinamensis_ as they burrowed in moist soil and attacked and destroyed _N. cinerea_ in breeding cages.

XIII. VERTEBRATA

Class PISCES

In British Guiana, Beebe (1925a) found undetermined cockroach remains in the stomachs of four species of fish belonging to three families, as follows:

Family POTAMOTRYGONTIDAE

=Potamotrygon humboldti= (Duméril)

(= _Potamotrygon hystrix_)

Family PIMELODIDAE

=Rhamdia sebae= Cuvier and Valenciennes

Family CHARACIDAE

=Cyrtocharax magdalenae essequibensis= (Eigenmann)

(= _Cynopotamus essequibensis_)

=Chalceus macrolepidotus= Cuvier and Valenciennes

The only other records of cockroaches being eaten by fish pertain to the use of cockroaches as bait.[8] Captain William Owen (_in_ Webster, 1834) stated that the Chinese used cockroaches as bait in their fishing excursions. At Reelfoot Lake, Tennessee, _Blatta orientalis_ were kept in large numbers by bait dealers and were sold to fishermen who used them for catching _Lepomis pallidus_, a sunfish locally known as bream, blue bream, or bluegill (Rau, 1944). In Indiana, oriental cockroaches were collected at a city dump by fishermen (Gould, 1941). Peterson (1956) states that cockroaches are satisfactory bait for bluegills, crappies, channel cat, blue heads, and large mouth black bass.

Class AMPHIBIA[9]

Order CAUDATA

Family PLETHODONTIDAE

=Plethodon glutinosus= (Green)

_Natural prey._--_Cryptocercus punctulatus_, U.S.A. (Honigberg, 1953): Protozoa which are normally only found in _C. punctulatus_ were present in the intestine of the salamander indicating that this cockroach had been eaten by the amphibian.

Order SALIENTIA

Family BUFONIDAE

=Bufo funereus= Bocage

_Natural prey._--Cockroaches, Belgian Congo (Noble, 1924): The stomachs of 62 out of 72 specimens contained food; this included 3 cockroaches.

=Bufo ictericus= Spix

_Natural prey._--Cockroaches, Brazil (Valente, 1949): Stomach contents revealed the prothorax, legs, and wings of cockroaches, and fragments of wood-cockroaches. This toad frequently feeds at night.

=Bufo marinus= (Linnaeus)

_Common name._--Giant toad, marine toad, Surinam toad.

_Natural prey._--_Epilampra abdomen-nigrum_, Trinidad (Weber, 1938): Found in the stomachs of two toads.

_Diploptera punctata_, Hawaii (Pemberton and Williams, 1938).

_Periplaneta_ sp., Fiji (Lever, 1939): Many householders in Suva have seen the toad eat considerable numbers of these cockroaches.

_Pycnoscelus surinamensis_, Hawaii (Alicata, 1938; Illingworth, 1941).

Cockroaches, Nicaragua (Noble, 1918): Stomach contents of toads captured at street lamps in Rio Grande consisted chiefly of large cockroaches. Puerto Rico (Wolcott, 1937).

=Bufo valliceps= Wiegmann

_Experimental prey._--_Periplaneta americana_, U.S.A. (Moore, 1946): Cockroaches containing infective acanthellas of _Moniliformis dubius_ were fed to three toads.

Family HYLIDAE

=Hyla cinerea= (Schneider)

_Common name._--Green tree frog.

_Natural prey._--_Ischnoptera deropeltiformis_, _Periplaneta americana_, and undetermined cockroaches, U.S.A., Georgia (Haber, 1926): Cockroaches were found in 11 of 100 stomachs.

Family RANIDAE

=Arthroleptis variabilis= Matschie

_Natural prey._--Cockroaches, Belgian Congo (Noble, 1924): Of 52 specimens examined, the stomach contents of 17 contained food, including 3 cockroaches.

=Hyperolius picturatus= Peters

_Natural prey._--Cockroach, Belgian Congo (Noble, 1924): The stomachs of 12 of 56 specimens examined contained food, including one cockroach.

=Leptodactylus albilabris= (Günther)

_Natural prey._--Cockroach, Puerto Rico (Schmidt, 1920): One of 25 stomachs contained a medium-sized cockroach.

=Leptodactylus pentadactylus= (Laurenti)

_Common name._--"Smoky jungle frog" or "pepper frog."

_Natural prey._--Cockroaches, Nicaragua (Noble, 1918): Cockroach wings were found in the stomach of a frog caught around human habitation. Brazil (Valente, 1949).

=Leptopelis calcaratus= (Boulenger)

_Natural prey._--Cockroaches, Belgian Congo (Noble, 1924): The stomachs of 35 specimens were examined of which 13 contained food, including 2 cockroaches.

=Leptopelis rufus= Reichenow

_Natural prey._--Cockroaches, Belgian Congo (Noble, 1924): Forty-five of 83 stomachs examined contained food, including 2 cockroaches.

=Megalixalus fornasinii= (Bianconi)

_Natural prey._--Cockroaches, Belgian Congo (Noble, 1924): The stomachs of 3 of 40 specimens contained food, including 2 cockroaches.

=Rana catesbeiana= Shaw

_Common name._--Bullfrog.

_Natural prey._--Cockroaches, Puerto Rico (Derez, 1949).

=Rana mascareniensis= Duméril and Bibron

_Natural prey._--Cockroaches, Belgian Congo (Noble, 1924): The stomach contents of 138 specimens were examined, 39 of which contained food, including 2 cockroaches.

=Rana pipiens= Schreber

_Common name._--Leopard frog.

_Experimental prey._--_Periplaneta americana_, U.S.A. (Moore, 1946): Cockroaches containing infective acanthellas of _Moniliformis dubius_ were fed to two frogs.

_Neostylopyga rhombifolia_, U.S.A. (Dr. T. Eisner, personal communication, 1958.)

=Frogs=

_Natural prey._--_Blatta orientalis_, U.S.A. (Rau, 1924): Frogs which escaped from a tank in the cellar consumed quantities of this cockroach.

_Parcoblatta pensylvanica_, U.S.A. (Frost, 1924): One adult specimen recovered from alimentary canal of a frog, probably _Rana_ sp.

=Unidentified batrachians=

_Experimental prey._--_Blattella germanica_, Germany, Frankfurt am Main, Zoological Garden (Lederer, 1952): These insects were preferred by all the insect eaters in the zoo.

_Periplaneta americana_, Germany, Frankfurt am Main, Zoological Garden (Lederer, 1952): Newly molted individuals were accepted as food, but others were usually passed by or consumed unwillingly.

Class REPTILIA[10]

Order CHELONIA

Family EMYDIDAE

=Chrysemys picta= (Schneider)

_Common name._--Painted turtle.

_Natural prey._--_Periplaneta australasiae_, England (Lucas, 1916, 1920): The cockroach, apparently injured, fell into water in the tortoise house, Zoological Gardens, Regent's Park, and the terrapin ate it.

Order SAURIA

Family GEKKONIDAE

=Gekko gecko= (Linnaeus)

_Natural prey._--Cockroaches, Philippine Islands, Laguna (Villadolid, 1934): The geckos frequent holes in trees and underside of bark which are favorable haunts of cockroaches. Stomach contents mostly Blattidae and "Locustidae."

=Hemidactylus frenatus= Duméril and Bibron

_Common name._--House lizard.

_Natural prey._--Cockroaches, Philippine Islands, Laguna (Villadolid, 1934): Bulk of stomach contents of 22 lizards consisted of Orthoptera, mostly cockroaches.

=Sphaerodactylus= sp.

_Natural prey._--Cockroaches, British Guiana (Beebe, 1925a): The above lizard is found in houses.

=Thecadactylus= sp.

_Natural prey._--Cockroaches, British Guiana (Beebe, 1925a): The above lizard is found in houses.

=Undetermined geckos=

_Natural prey._--Cockroaches, Australia, Flinders River (Froggatt, 1906): The lizard lived in the walls of the hut and hunted cockroaches upon the roof at night. Arno Atoll (Usinger and La Rivers, 1953).

Family IGUANIDAE

=Anolis carolinensis= Voigt

_Experimental prey._--_Diploptera punctata_, U.S.A. (Eisner, 1958).

=Anolis cristatellus= Duméril and Bibron

_Natural prey._--_Blattella_ sp., _Cariblatta delicatula_, _Epilampra wheeleri_, _Periplaneta americana_, _Periplaneta australasiae_, and _Symploce flagellata_, Puerto Rico (Wolcott, 1924): The last-named cockroach may have been _S. ruficollis_ Rehn and Hebard, the females of which are hard to distinguish from _flagellata_. Rehn and Hebard (1927) stated that in all probability _flagellata_ does not occur on the island of Puerto Rico. Wolcott (1950) stated that _Symploce ruficollis_ [= _bilabiata_] serves as food for the crested lizard.

Cockroaches, Puerto Rico (Schmidt, 1920): Of 100 stomachs examined, 16 contained Orthoptera, including cockroaches. Puerto Rico (Wolcott, 1924): One hundred _A. cristatellus_ had eaten 8 cockroaches, 4.14 percent of the total food, or 25 percent of the food for 8 lizards.

=Anolis pulchellus= Duméril and Bibron

_Natural prey._--Cockroaches, Puerto Rico (Wolcott, 1924): Two small cockroaches found in 50 lizards examined.

=Anolis equestris= Merrem

_Experimental prey._--_Neostylopyga rhombifolia_, U.S.A. (Eisner, personal communication, 1958.)

=Anolis grahami= Garman

_Natural prey._--_Periplaneta_ spp. and Blattidae, Bermuda (Simmonds, 1958): Stomachs of 176 lizards yielded 6 cockroaches.

=Anolis leachi= Duméril and Bibron

_Natural prey._--_Periplaneta_ spp. and Blattidae, Bermuda (Simmonds, 1958): Stomachs of 46 lizards yielded 31 cockroaches.

=Anolis sagrei= Cocteau

_Natural and experimental prey._--_Pycnoscelus surinamensis_, Cuba (Darlington, 1938): This species was eaten both in captivity and in nature. The lizard ate most readily soft, immature cockroaches. _Pycnoscelus surinamensis_ is probably a staple food of the lizard in nature, as Darlington observed wild lizards catch the nymphs.

=Anolis stratulus= Cope

_Natural prey._--_Aglaopteryx facies_, Puerto Rico (Wolcott, 1924): One cockroach was found in 50 lizards examined.

Cockroach, Puerto Rico (Schmidt, 1920): One of 25 stomachs contained a cockroach.

=Anolis= sp.

_Natural prey._--"Wood roaches," British Guiana (Beebe, 1925a): The above lizard is arboreal on foliage in low jungle.

Family SCINCIDAE

=Leiolopisma laterale= Say

_Common name._--Brown skink.

_Natural prey._--Woodroaches, U.S.A., Louisiana (Slater, 1949): Analysis of stomach contents of 84 adult skinks showed that nymphal and adult woodroaches comprised the majority of Orthoptera.

=Tropidophorus grayi= Günther

_Common name._--Spiny lizard.

_Natural prey._--Cockroaches, Philippine Islands, Laguna (Villadolid, 1934): Food of this species was mostly Blattidae.

=Unidentified skinks=

_Natural prey._--Cockroaches, Arno Atoll (Usinger and La Rivers, 1953).

Family AGAMIDAE

_Experimental prey?_--Cockroaches, Australia (Lee and Mackerras, 1955): A general statement was made that in captivity Agamidae were observed feeding avidly on cockroaches and other insects. Three agamids studied by these workers were _Amphibolurus barbatus_ (Gray), _Physignathus lesueurii_ Gray, and _Chlamydosaurus kingii_ Gray.

Family CHAMAELEONTIDAE

=Chamaeleon chamaeleon= (Linnaeus) and =Chamaeleon oustaleti= Mocquard

_Experimental prey._--Cockroaches, Amsterdam (Portielje, 1914): Large cockroaches were fed to these lizards in the reptile house of Artis.

Family TEIIDAE

=Ameiva exsul= Cope

_Common name._--Iguana, ground lizard.

_Natural prey._--Cockroach (nymph), _Epilampra wheeleri_, and _Periplaneta americana_, Puerto Rico (Wolcott, 1924): Stomach contents of 15 lizards were analyzed. _E. wheeleri_ formed 30 percent of the food of one lizard. The cockroach nymph formed 5 percent of the food of one lizard. One _P. americana_ formed 20 percent of the food of one lizard; another formed 50 percent of the food of a second lizard.

_Experimental prey._--Cockroach nymphs, Puerto Rico (Wolcott, 1924).

=Ameiva= sp.

_Natural prey._--Cockroaches, British Guiana (Beebe, 1925a): The above lizard is terrestrial and found near clearings. The stomach contents of 18 out of 40 reptiles contained cockroach remains.

=Cnemidophorus= sp.

_Natural prey._--Cockroaches, British Guiana (Beebe, 1925a): This is a terrestrial lizard found near clearings. The stomach contents of 4 out of 40 lizards contained cockroaches.

=Unidentified lizards=

_Natural prey._--Cockroaches, West Indies (H., 1800).

_Experimental prey._--_Blatta orientalis_, U.S.A. (Rau, 1924): Rau called the predator a common gray lizard.

_Periplaneta americana_, Germany, Frankfurt am Main, Zoological Garden (Lederer, 1952): Newly molted cockroaches were accepted as food, but others were usually passed by or consumed unwillingly.

Order SERPENTES

Family COLUBRIDAE

=Heterodon platyrhinos= Latreille

_Synonymy._--_Heterodon contortrix_ [Dr. Doris M. Cochran, personal communication, 1957].

_Common name._--Hog-nosed snake.

_Experimental prey._--_Periplaneta americana_, U.S.A. (Moore, 1946): Cockroaches containing infective acanthellas of _Moniliformis dubius_ were fed to one snake.

=Garter Snake=

_Experimental prey._--_Blatta orientalis_, U.S.A. (Rau, 1924).

Class AVES

The cockroach is always wrong when arguing with a chicken.

_Spanish proverb_ (HARTNACK, 1939)

Arboreal cockroaches hidden in and under bark are much more likely to be encountered by birds than by other predators, and insectivorous birds undoubtedly consume many more cockroaches than the few records would indicate. Most of the records we have located identify the birds at least by common name. Where possible we have given the scientific names for those birds whose common names are recognizably specific. We have followed the systematic classification of Wetmore (1940).

Figuier (1869) stated that poultry and owls are very fond of cockroaches. Perkins (1913) made the general statement that some of the native birds of Hawaii are partial to the endemic _Allacta similis_. Asano (1937) stated that in Japan natural enemies of cockroaches may be found in the Galliformes, Strigiformes, Passeriformes, and Piciformes. Although Lederer (1952) successfully fed newly molted _Periplaneta americana_ to insectivorous birds in the Zoological Garden, Frankfurt am Main, _Blattella germanica_ were preferred by these birds. The following records are of specific birds feeding on cockroaches.

Order ANSERIFORMES

Family ANATIDAE

=Domestic duck=

_Natural prey._--_Pycnoscelus surinamensis_, Australia (Fielding 1926): The ducks became infected with Manson's eye worm of which _P. surinamensis_ is the only known intermediate host.

Cockroach, Bermuda (Jones, 1859): "All kinds of poultry feed greedily upon the cockroach; tame ducks spending entire moonlight nights in their capture."

Order GALLIFORMES

Family PHASIANIDAE

=Bambusicola thoracica= Temminck

_Common name._--Kojukei.

_Natural prey._--Cockroaches, Japan (Asano, 1937).

=Gallus= sp.

_Common name._--Jungle fowl.

_Natural prey._--_Periplaneta australasiae_, Hawaii (Schwartz and Schwartz, 1949).

=Phasianus calchicus karpowi= Buturlin

_Common name._--Korean pheasants.

_Experimental prey._--_Blattella germanica_ and _Periplaneta picea_, Japan (Asano, 1937): Adults of these cockroaches were devoured at once when they were fed with the heads cut off.

=Phasianus= sp.

_Common name._--Pheasant.

_Natural prey._--Blattidae, unidentified (below 1 percent of the diet), _Cutila soror_ (below 1 percent of the diet), _Diploptera punctata_ (above 6 percent of the diet), and _Pycnoscelus surinamensis_ (6 percent of the diet), Hawaii (Schwartz and Schwartz, 1949).

=Coturnix coturnix japonica= (Temminck and Schlegel)

_Common name._--Japanese quail.

_Natural prey._--Blattidae (unidentified) and _Lobopterella dimidiatipes_, Hawaii (Schwartz and Schwartz, 1949).

=Domestic chicken=

_Natural and experimental prey._--_Blaberus craniifer_, U.S.A., Key West, Florida. J.A.G. Rehn in 1912 (personal communication) observed chickens feeding on nymphs of _B. craniifer_ which had dropped to the ground from among stacked coffins in an undertaker's shack.

_Blatta orientalis_, U.S.A. (Rau, 1924): The chickens ate cockroaches that were caught in traps.

_Hebardina concinna_, Japan (Asano, 1937): Experimental feeding to white Leghorn chickens.

_Periplaneta americana_, Surinam (Stage, 1947): Several cockroaches ran off the floor of a house, which was being sprayed with DDT, and were eaten by chickens. Although some chickens had DDT tremors the next day, all appeared normal two days later.

_Pycnoscelus surinamensis_, Australia (Fielding, 1926); Formosa, experimental feeding (Kobayashi, 1927); Australia, experimental feeding (Fielding, 1927, 1928); U.S.A., Florida, experimental feeding (Sanders, 1928); Antigua (Hutson, 1943); Hawaii (Illingworth, 1931; Schwabe, 1949, 1950a, 1950b). This cockroach is the intermediate host of _Oxyspirura mansoni_, the chicken eye worm.

Cockroaches, Guadeloupe (Dutertre, 1654); Africa (Moiser, 1947): "Poultry" ate cockroaches which had been killed by DDT and sodium fluoride. Hawaii (Zimmerman, 1948).

=Partridge=

_Natural prey._--Cockroaches, British Guiana (Beebe, 1925a): The food of two small species of leaf-colored partridges that lived on the jungle floor, consisted chiefly of cockroaches and beetles.

Family MELEAGRIDIDAE

=Meleagris gallopavo= (Linnaeus)

_Common name._--Turkey.

_Natural prey._--_Pycnoscelus surinamensis_, Antigua (Hutson, 1943): Turkeys were found heavily infected with Manson's eye worm of which _P. surinamensis_ is the only known intermediate host. These turkeys therefore were presumed to have fed on this cockroach.

Order COLUMBIFORMES

Family COLUMBIDAE

=Streptopelia chinensis= (Scopoli)

_Common name._--Chinese dove.

_Natural prey._--_Pycnoscelus surinamensis_, Hawaii (Schwabe, 1950b).

=Pigeon=

_Experimental prey._--_Pycnoscelus surinamensis_, Australia (Fielding, 1927); U.S.A., Florida (Sanders, 1928).

Order STRIGIFORMES

Family STRIGIDAE

=Gymnasio nudipes= (Daudin)

_Common name._--Bare-legged owl.

_Natural prey._--_Epilampra_ sp., Puerto Rico (Wetmore, 1916): One specimen identified in stomach of a wild-caught owl.

Cockroaches, Puerto Rico (Wetmore, 1916): These insects were found in stomachs of five owls.

Order CORACIFORMES

Family TODIDAE

=Todus mexicanus= Lesson

_Common name._--Porto Rican tody.

_Natural prey._--_Plectoptera poeyi?_, Puerto Rico (Wetmore, 1916): The stomachs of 89 birds were examined; a single bird had eaten the above cockroach. According to Wolcott (1950) _P. poeyi_ could be _Plectoptera dorsalis_, _P. rhabdota_, or _P. infulata_.

Family BUCEROTIDAE

=Tockus birostris= (Scopoli)

_Synonymy._--_Lophocerus birostris_ [Dr. H. Friedmann, personal communication, 1957].

_Common name._--Common gray hornbill.

_Natural prey._--Cockroaches, India, Central Provinces (D'Abreu, 1920).

Order PICIFORMES

Family PICIDAE

=Dendrocopus mahrattensis= (Latham)

_Synonymy._--_Liopicus mahrattensis_ [Friedmann, p. c. 1957].

_Common name._--Yellow-fronted pied woodpecker.

_Natural prey._--Cockroaches, India, Central Provinces (D'Abreu, 1920).

=Melanerpes portoricensis= (Daudin)

_Common name._--Puerto Rican woodpecker.

_Natural prey._--_Pycnoscelus surinamensis_, Puerto Rico (Wetmore, 1916): One specimen found in 59 bird stomachs examined.

Order PASSERIFORMES

Family FORMICARIIDAE

=Gymnopithys leucaspis= (Sclater)

_Common name._--Bicolored antbird.

_Natural prey._--Cockroaches, Panama Canal Zone (Johnson, 1954): This bird feeds on small cockroaches, and other arthropods, which are flushed from their hiding places by swarms of the army ant, _Eciton burchelli_.

Family ORIOLIDAE

=Icterus portoricensis= (Bryant)

_Common name._--Puerto Rican oriole.

_Natural prey._--Cockroaches, Puerto Rico (Wetmore, 1916): Cockroaches and oöthecae found in the birds' stomachs.

Family CORVIDAE

=Aphelocoma coerulesens= (Bosc)

_Common name._--Florida jay.

_Experimental prey._--_Pycnoscelus surinamensis_, U.S.A., Florida (Sanders, 1928).

=Cyanocitta cristata= (Linnaeus)

_Common name._--Blue jay.

_Experimental prey._--_Diploptera punctata_, U.S.A. (Eisner, 1958).

_Eurycotis floridana_, _Neostylopyga rhombifolia_, and _Periplaneta americana_, U.S.A. (Eisner, personal communication, 1958): _E. floridana_ was only eaten after the odor of 2-hexenal, which was released by the insect on being attacked by the bird, had dissipated.

Family PARADISEIDAE

=Paradisea papuana= Bechstein

_Experimental prey._--Cockroaches, Malaya and on shipboard (Wallace, 1869): Two adult males fed voraciously on rice, bananas, and cockroaches. Wallace collected cockroaches every night on board ship to feed the birds. "At Malta ... I got plenty of cockroaches from a bakehouse, and when I left, took with me several biscuit-tins full, as provision for the voyage home."

Family TROGLODYTIDAE

=Troglodytes aedon= Vieillot

_Common name._--House wren.

_Natural prey._--Cockroaches, U.S.A. (Greenewalt and Jones, 1955): The wren carried three small cockroaches to nestlings; the records probably represent incidental captures.

=Troglodytes audax= Tschudi

_Common name._--Cucarachero.

_Natural prey._--Cockroach (called Chilicabra by Peruvian Indians), Peru (Tschudi, 1847): The bird seized the cockroach and bit off its head then devoured the body discarding the wings.

Family LANIIDAE

=Lanius ludovicianus= Linnaeus

_Common name._--Loggerhead shrike.

_Experimental prey._--_Pycnoscelus surinamensis_, U.S.A., Florida (Sanders, 1928).

Family STURNIDAE

=Acridotheres tristis= Bonnaterre and Vieillot

_Common name._--Myna, mynah.

_Natural prey._--Cockroaches, Hawaii, Lanai (Illingworth, 1928): Illingworth reported that he had never seen as many cockroaches anywhere else in Hawaii. The birds followed tractors that were destroying cactus and kept close to the chain that turned over the stumps. The following species were collected: _Allacta similis_, _Blattella germanica_, _Cutilia soror_, _Diploptera punctata_, _Leucophaea maderae_, _Periplaneta americana_, _Periplaneta australasiae_, _Pycnoscelus surinamensis_. Illingworth did not state whether the birds ate all these species indiscriminately.

_Pycnoscelus surinamensis_, Hawaii (Williams et al., 1931; Schwabe, 1950b): In many places this species forms an important fledgling food for mynah birds.

Family VIREONIDAE

=Vireo latimeri= Baird

_Common name._--Latimer's vireo.

_Natural prey._--_Periplaneta_ sp., Puerto Rico (Wetmore, 1916): Cockroaches were found in one of 43 stomachs examined.

Family ICTERIDAE

=Agelaius xanthomus= (Sclater)

_Common name._--Yellow-shouldered blackbird.

_Natural prey._--Cockroaches, Puerto Rico (Wetmore, 1916): Oöthecae and remains of adult cockroaches found in stomachs.

=Dolichonyx oryzivorus= (Linnaeus)

_Common name._--Bobolink.

_Experimental prey._--_Pycnoscelus surinamensis_, U.S.A., Florida (Sanders, 1928).

=Holoquiscalus brachypterus= (Cassin)

_Common name._--Puerto Rican blackbird.

_Natural prey._--Cockroaches, Puerto Rico (Wetmore, 1916): A few eggs (oöthecae) of cockroaches in stomachs.

=Black bird=

_Experimental prey._--_Pycnoscelus surinamensis_, U.S.A., Florida (Sanders, 1928).

Family FRINGILLIDAE

=Passer domesticus= (Linnaeus)

_Common name._--English sparrow.

_Natural prey._--_Pycnoscelus surinamensis_, Hawaii (Illingworth, 1931; Schwabe, 1950b): Remains of this cockroach were found in the stomach of the sparrow.

=Sparrow=

_Natural prey._--_Periplaneta americana_, England (Lucas, 1908, 1920).

Cockroaches, Japan (Asano, 1937).

=Tiaris bicolor omissa= (Jardine)

_Common name._--Carib grassquit.

_Natural prey._--Cockroaches, Puerto Rico (Wetmore, 1916): Animal food was found in 5 of 72 stomachs examined; one bird had eaten two cockroaches among other insects.

Class MAMMALIA[11]

Order MARSUPIALIA

Family DIDELPHIDAE

=Monodelphis= sp.

_Natural prey._--Cockroaches, British Guiana (Beebe, 1925a): The above opossum is nocturnal and arboreal but nests on the ground in grass.

Order INSECTIVORA

Family ERINACEIDAE

=Erinaceus europaeus= Linnaeus

_Experimental prey._--_Blattella germanica_, France (Brumpt and Urbain, 1938): Two hedgehogs were fed cockroaches infested with _Prosthenorchis elegans_ and _P. spirula_.

=Erinaceus= sp.

_Common name._--Hedgehog.

_Natural prey._--Cockroaches, England (Samouelle, 1841; Cowan, 1865).

Order CHIROPTERA

Family MOLOSSIDAE

=Molossus= sp.

_Natural prey._--Cockroaches, British Guiana (Beebe, 1925a): The above bat is a common house bat of the area.

Order PRIMATES

Family LEMURIDAE

=Lemur coronatus= Gray

_Natural prey._--_Blattella germanica_, France (Brumpt and Urbain, 1938): The monkey apparently became infested naturally with _Prosthenorchis spirula_ for which _B. germanica_ was the intermediate host in the monkey house.

=Lemur fulvus= E. Geoffroy

_Natural prey._--_Blattella germanica_, France (Brumpt and Urbain, 1938): See comment under _Lemur coronatus_.

Family LORISIDAE

=Loris tardigradus= (Linnaeus)

_Synonymy._--_Lemur tardigradus_ [Dr. D. H. Johnson, personal communication, 1957].

_Natural prey._--Cockroaches, on board ship (Cowan, 1865).

=Perodicticus potto= (P. L. S. Müller)

_Common name._--Potto.

_Natural prey._--_Blattella germanica_, France (Brumpt and Urbain, 1938): See comment under _Lemur coronatus_.

_Experimental prey._--Blattidae, East Africa (Pitman, 1931): Both sexes of the potto ate freely of all types of cockroaches.

Family TARSIIDAE

=Tarsius= sp.

_Experimental prey._--Cockroaches, Borneo (Shelford, 1916).

Family CEBIDAE

=Aotes zonalis= Goldman

_Synonymy._--_Aotus_ [Simpson, 1945].

_Common name._--Canal Zone night monkey.

_Natural prey._--_Leucophaea maderae_, Panama (Foster and Johnson, 1939): Captive monkeys became naturally infested with _Protospirura muricola_ by eating cockroaches that contained infective larvae of the worm.

=Ateles dariensis= Goldman

_Common name._--Darien black spider monkey.

_Natural prey._--_Leucophaea maderae_, Panama (Foster and Johnson, 1939): See comment under _Aotes zonalis_.

=Cebus apella= (Linnaeus)

_Natural prey._--_Blattella germanica_, France (Brumpt and Urbain, 1938): See comment under _Lemur coronatus_.

=Cebus capucinus= (Linnaeus)

_Common name._--White-faced monkey.

_Natural prey._--_Leucophaea maderae_, Panama (Foster and Johnson, 1939): Favorite item of food in the laboratory. See comment under _Aotes zonalis_.

=Saimiri sciurea= Linnaeus