Chapter IX: Section III: (p. 14). Other associations may be accidental (e.g., certain (3)

_Natural host._--_Periplaneta americana_, U.S.A. (Roth and Willis, unpublished data, 1953): Mite determined by Dr. E. W. Baker (personal communication, 1953). Mites were found in the oöthecal cavity of a female cockroach that had been isolated for her entire adult life. The mites were in a closely packed mass behind a plug of what appeared to be feces, disintegrated eggs, and dried blood; none of the mites were visible until this plug was removed. Baker (personal communication, 1953) stated that the mite is probably not parasitic and that species of the genus feed on organic matter.

=Rhizoglyphus tarsalus= Banks

_Natural host._--_Periplaneta americana_, U.S.A. (Rau, 1940a): Not normally parasitic on cockroaches, but the mites became so numerous at times they would attack living as well as dead and dying cockroaches.

Family GLYCIPHAGIDAE

=Chaetodactylus= sp.

_Synonymy._--_Trichotarsus_ sp. [Baker and Wharton, 1952].

_Natural host._--_Leucophaea maderae_, Puerto Rico (Seín, 1923): Mites found on cockroach's thorax and particularly among the folds of the wings (Seín, 1923). Mites of this genus are found infesting organic matter (Baker and Wharton, 1952).

Family PODAPOLIPODIDAE

=Locustacarus= sp.

_Natural hosts._--_Diploptera punctata_ and _Nauphoeta cinerea_, U.S.A. (Roth and Willis, unpublished data, 1954): Mite genus determined by Dr. E. W. Baker (personal communication, 1954). The mites cluster thickly on intersegmental membranes, particularly around the coxae and neck. Despite a heavy infestation, the colony of _Nauphoeta_ thrived for several years. This mite was found first on _N. cinerea_ and possibly transferred to _D. punctata_ when the latter was brought into the laboratory from Hawaii.

Family IOLINIDAE

=Iolina nana= Pritchard

_Natural hosts._--_Blaberus craniifer_ (originally from a culture at Harvard University) and _Diploptera punctata_ (originally from Hawaii), U.S.A., Pennsylvania (Roth and Willis, unpublished data, 1953; Pritchard, 1956): The mites usually attached near the wing bases of the insects. Morphologically, the species is intermediate between certain predaceous and phytophagous mites (Pritchard, 1956).

Family PTERYGOSOMIDAE

=Pimeliaphilus podapolipophagus= Trägårdh

_Common name._--Cockroach mite.

_Natural hosts._--_Parcoblatta_ sp., U.S.A. (Edmunds, 1953a).

_Periplaneta americana_, U.S.A. (Piquett and Fales, 1952).

Cockroaches. U.S.A. (Baker and Wharton, 1952).

_Experimental hosts._--_Blatta orientalis_, _Blattella germanica_, and _Periplaneta americana_, U.S.A. (Cunliffe, 1952).

Eggs of this mite (fig. 4) are usually laid indiscriminately in the rearing cages, rarely on the host. Eggs are coated with a sticky secretion which enables those laid on the host to adhere. Hatching occurs in 6-11 days at 90-95° F., and in 9-11 days at 80° F. The newly hatched larva starts to feed immediately on the cockroach. Larval stage lasts 4-6 days, rests 2-3 days, and molts. During the single nymphal instar, the mite feeds on the host and moves about for 6-7 days. The mite then rests 3-4 days before molting. Entire life cycle covers a period of 28-32 days. Adult mite lives 2-3 weeks, during which time it can produce 2-3 batches of from 1 to 20 eggs; the usual batch is about 12 eggs. The mites are unable to live on cockroach feces, cast skins, or dead cockroaches. Mites died within 4-5 days unless live cockroaches were supplied. Parasitism was proved by detecting radioactivity in mites that had fed on cockroaches which had been previously fed radioactive NaCl (Cunliffe, 1952).

The mites can destroy laboratory cultures of cockroaches (Piquett and Fales, 1952; Edmunds, 1953a). A cockroach attacked by 25 mites succumbed after about an hour, falling on its back; it died after 5 hours (Cunliffe, 1952).

When found in homes and offices, these mites are an indication of the presence of cockroaches; the mite has been twice accused of biting people (Baker et al., 1956).

RECORDS OF UNIDENTIFIED MITES

_Natural hosts._--_Aglaopteryx facies_, Puerto Rico (Seín, 1923): Four red "tick" nymphs found under wings of female.

_Blaberus craniifer_, U.S.A., Florida (Hebard, 1917): "A number of lice [mites] are present on many of these specimens [28[F][F]]."

_Blaberus discoidalis_, adventive from West Indies, taken in Scotland (Stewart, 1925): A considerable number of mites were all over the body and hind wings.

_Blatta orientalis_, Germany (Cornelius, 1853): Ex sexual organs of male.

_Blattella germanica_, U.S.A., in laboratory (Parker, 1939): Under conditions of high humidity, the cockroaches became heavily infested with mites. In cages where the infestation was heavy, an abnormally large number of females dropped their oöthecae, and the percentage of eggs hatching was low.

_Parcoblatta uhleriana_, U.S.A., North Carolina (Hatcher, 1939): Hypopi of mites were found deeply embedded in the fat body of two individuals.

Mites in the hypopial stage attach to insects by which they are dispersed. Hypopi have been found in the gill chambers of a mollusk and in the gonads of a millipede (Baker and Wharton, 1952).

_Periplaneta americana_, U.S.A., in laboratory (Fisk, 1951): The insects were sluggish and molted with difficulty. Gold Coast Colony (Macfie, 1922): Larvae of a tarsonemid mite were found in the feces.

_Pycnoscelus surinamensis_, Hawaii (Illingworth, 1915): During the summer the soil was literally swarming with young of various stages. Early in September most of the adults were dead and all were covered with mites. U.S.A., Connecticut, in laboratory (Zappe, 1918a). Hawaii, in laboratory (Schwabe, 1950): Some of the cockroaches apparently died from mite infestations.

; reproduced through the
courtesy of Dr. E. W. Baker and the National Pest Control
Association.)]

Cockroach, England? (Ealand, 1915): Cockroaches may carry the hypopial stage of the cheese mite.

CONTROL OF MITES IN COCKROACH COLONIES

Fisk (1951) eliminated the mites [possibly _Pimeliaphilus podapolipophagus_ (Baker et al., 1956)] in his cockroach colony by using a 5-percent spray and a 5-percent dust of p-chlorophenyl, p-chlorobenzene sulfonate. The exterior of the cockroach containers were sprayed with the solution and the interior, including the insects, were dusted. Within a month the mites had disappeared and the vigor of the cockroach colony improved. Piquett and Fales (1952) used flowers of sulfur and general sanitary procedures for eliminating the mites in laboratory colonies of _Blatta orientalis_; they cleaned the dishes every few days and applied grease around the edges of the containers to prevent new mite invasions. Qadri (1938) employed similar control measures.

Class CHILOPODA

Large centipedes which entered houses in India probably sought out cockroaches (Maxwell-Lefroy, 1909). In Puerto Rico, centipedes entered homes to which they were attracted by cockroaches (Seín, 1923). In Hawaii, centipedes preyed on insects generally but especially on cockroaches (Bryan, 1915). Sonan (1924) reported that in Formosa and Okinawa Islands a species of centipede 5 to 6 inches long comes into the houses and devours both adults and nymphs of _Periplaneta americana_ or _P. australasiae_. Zimmerman (1948) found _P. australasiae_ breeding by scores in rock piles in Hawaii accompanied by large numbers of _Scolopendra_ and large spiders that probably preyed upon the cockroaches.

Order SCUTIGEROMORPHA

Family SCUTIGERIDAE

=Scutigera coleoptrata= (Linnaeus)

_Synonymy._--_Scutigera forceps_ Rafinesque [Crabill, 1952].

_Common name._--House centipede.

_Natural prey._--Cockroaches, U.S.A. (Felt, 1909; Back, 1947; Auerbach, 1951; Crabill, 1952; and others): This predator-prey relationship seems to be based on good circumstantial evidence (Crabill, personal communication, 1953).

_Experimental prey._--_Blattella germanica_, newly hatched nymphs and adult female, U.S.A. (Snodgrass, 1930; Roth and Willis, unpublished data, 1953).

_Periplaneta americana_, U.S.A. (Roth and Willis, unpublished data, 1953).

_Supella supellectilium_, U.S.A. (Roth and Willis, unpublished data, 1953): See plate 31.

Our specimen caught a small American cockroach nymph that we placed in its jar. Before it had finished its meal, it caught and held two other nymphs with its legs while it continued to feed on the first. The body of this centipede reaches a maximum length of 27 mm. and it is usually found in basements, dark corners, or in spaces in the walls (Auerbach, 1951). Introduced from Europe, this species is now widespread in the United States (Crabill, 1952).

=Allothereua maculata= (Newport)

_Synonymy._--_Scutigera maculata_ [Crabill, personal communication, 1957].

_Natural prey._--Cockroaches, Malay peninsula, Batu caves (Ridley in Annandale et al., 1913): This is a presumptive host record.

Order SCOLOPENDROMORPHA

Family SCOLOPENDRIDAE

=Scolopendra cingulata= Latreille

_Experimental prey._--Cockroaches, England (Cloudsley-Thompson, 1955): After capture in France, this specimen was kept for four weeks without food. She was then fed medium-sized nymphal cockroaches of which she ate an average of about one per week throughout the summer. Adult cockroaches were attacked only after they had been disabled.

=Scolopendra morsitans= Linnaeus

_Natural prey._--Cockroaches, Guadeloupe (Lherminier, 1837).

_Experimental prey._--Cockroaches, India, Nagpur (Jangi, 1955): As soon as the centipede became aware of its prey, it rapidly embraced the cockroach within its legs and with its fangs gripped the insect's thorax. The predator continued to hold the prey with its fangs while its mouth parts prodded the victim's body. After feeding on an adult cockroach, the centipede is not inclined to kill another for 2-3 days.

=Scolopendra subspinipes= Leach

_Natural prey._--Cockroaches, Hawaii (Williams et al., 1931): This is a common species with a body length of 6 or more inches. It is reported to be a great enemy of cockroaches.

=Scolopendra= sp.

_Natural prey._--_Ectobius panzeri_, England (Lucas, 1911, 1920): When captured, the centipede was holding a live cockroach which it had apparently just caught. The insect was held beneath its captor's body, ventral surface upward, by several of the anterior legs while the centipede fed.

Class INSECTA

We have found representatives of only 10 orders that have preyed on or parasitized cockroaches: Beetles, flies, bugs, ants, wasps, stylops, and cockroaches occurred in nature; the others resulted from feeding cockroaches to captive insects or were laboratory observations.

Order ODONATA

Family AESHNIDAE

=Anax strenuus= Hagen

_Common name._--Giant Hawaiian dragonfly.

_Experimental prey._--Cockroaches, Hawaii (Williams, 1936): The dragonfly nymph was fed with medium large cockroaches and other insects.

Order BLATTARIA

In this chapter the relations of other arthropods to cockroaches are either as parasites or as predators. Certain cockroaches have turned the tables on their adversaries and become predators themselves. This aspect of cockroach behavior is discussed in chapter XVI. Other associations of cockroaches, as commensals with other insects and as associates of other cockroaches, are discussed in chapters XV and XVII.

Order ORTHOPTERA

Family MANTIDAE

=Hierodula tenuidentata= (Saussure) (?) (Serville)

(Pl. 32)

_Experimental prey._--_Blatta orientalis_, _Diploptera punctata_, _Eurycotis floridana_, _Leucophaea maderae_, _Nauphoeta cinerea_, _Neostylopgya rhombifolia_, and _Periplaneta americana_, U.S.A. (Rilling, personal communication, 1957): Mrs. Rilling wrote us that with the exception of _N. rhombifolia_, all the above cockroaches were readily eaten. All the mantids initially rejected _N. rhombifolia_ after grasping and making a brief attempt to chew the cockroaches. However, if specimens of _N. rhombifolia_ were left in the jars with the mantids, the cockroaches were usually eaten within the next 24 hours. _N. rhombifolia_ ejects an odorous substance when seized and the mantids probably ate these insects after most of this secretion had been depleted. It is highly probable that the secretion of _N. rhombifolia_ may deter the mantid's attack, but it should be pointed out that, with the possible exception of _N. cinerea_, all the other species fed to these mantids give off odorous substances when seized or disturbed. Apparently, certain naturally repellent compounds will deter this mantid, whereas others that are presumed to be repellent will not; however, the nutritional state of the mantid is undoubtedly a factor which may limit the effectiveness of certain repellent secretions against this predator.

_Byrsotria fumigata_, teneral males, and _Periplaneta australasiae_, nymphs, U.S.A. (Roth and Willis, unpublished data, 1958).

_Diploptera punctata_, U.S.A. (Eisner, 1958).

=Mantis religiosa= Linnaeus

_Common name._--European mantis.

_Experimental prey._--_Nauphoeta cinerea_, and _Periplaneta americana_, U.S.A. (Rilling, personal communication, 1957).

=Metallyticus semiaeneus= Westwood

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

=Sphodromantis viridus= (Forskål)

_Synonymy._--_Sphodromantis bioculata_ Burmeister [Gurney, personal communication, 1958].

_Experimental prey._--_Blatta orientalis_, Egypt (Adair, 1923): This species of cockroach was apparently used regularly as food for the mantid in the laboratory.

=Stagmomantis carolina= (Johansson)

_Common name._--Carolina mantis.

_Experimental prey._--_Blattella germanica_ and _Periplaneta americana_, U.S.A. (Breland, 1941): The mantids were fed 1-2 German cockroaches daily. One female mantid consumed 10 adult German cockroaches plus one oötheca and part of another in 2.5 hours. An adult German cockroach was consumed in an average of 8.5 minutes (range 5.5-15 minutes).

_Blatta orientalis_, nymphs, and _Diploptera punctata_, U.S.A. (Roth and Willis, unpublished data, 1953).

=Tarachodes maurus= (Stal)

_Experimental prey._--Cockroaches, South Africa (Faure, 1940).

=Tenodera aridifolia sinensis= Saussure

_Common name._--Chinese mantis.

_Experimental prey._--_Nauphoeta cinerea_ and _Periplaneta americana_, U.S.A. (Rilling, personal communication, 1957).

Family GRYLLACRIDIDAE

=Diestrammena apicalis= Br. v. Wattenwyl

and

=Diestrammena japanica= Blatchley

_Natural prey._--Cockroach eggs, Japan (Asano, 1937): These are questionable records. Asano found _D. apicalis_ and _D. japanica_ beneath his house near several empty cockroach oöthecae which appeared to have been eaten into. He assumed from the condition of the oöthecae and the proximity of the stone crickets that the insects had devoured the cockroach eggs.

_Experimental prey._--Eggs of _Blattella germanica_ and _Periplaneta japanica_, Japan (Asano, 1937): Seven eggs of _B. germanica_ (obtained from an oötheca being carried by a female) and eggs of _P. japanica_ (presumably in oöthecae) were fed to both species of stone crickets in the evening. The eggs were devoured by the next morning.

Order DERMAPTERA

Family FORFICULIDAE

=Undetermined earwigs=

_Experimental prey._--Cockroaches, France (Chopard, 1938): According to Chopard, Brisout de Barneville in 1848 indicated that earwigs in captivity can be fed small cockroaches.

Order HEMIPTERA

Family LYGAEIDAE

=Clerada apicicornis= Signoret

_Natural prey._--Cockroach, Hawaii (Illingworth, 1917): This predaceous bug is commonly found about buildings. Illingworth says that Kirkaldy suspected that it fed on small blattids and that Dr. Perkins saw it feeding on a dead cockroach.

Family REDUVIIDAE

=Spiniger domesticus= Pinto

_Natural prey._--_Periplaneta americana_, Brazil, Matto Grosso (Pinto, 1927, 1927a): This bug preys principally on cockroaches and was observed infesting the walls of dwellings where it preyed on _P. americana_.

=Triatoma arthurneivai= Lent and Martins

_Natural prey._--_Monastria_ sp., Brazil, Minas Gerais (Martins, 1941): This bug probably feeds on cockroaches of this genus, as well as on rodents.

=Undetermined reduviids=

_Natural prey._--_Arenivaga roseni_ and _Polyphaga saussurei_, Turkmen S.S.R. (Vlasov and Miram, 1937): These desert cockroaches are found in burrows of rodents and desert turtles around Ashkhabad. Reduviids are their main enemies. Vlasov (1933) found nymphs of _Reduvius christophi_ Jak. and _R. fedtschenkianus_ Osch. in similar burrows in this same area, although he did not specifically cite them as enemies of the desert cockroaches.

Family NEPIDAE

=Ranatra= sp.

_Experimental prey._--Cockroaches, U.S.A. (Hoffman, 1924).

Order NEUROPTERA

Family ASCALAPHIDAE

=Undetermined larva=

_Experimental prey._--_Blattella germanica_, Kenya Colony (Someren, 1924).

Order DIPTERA

From the few observations that have come to our attention, it seems that flies are comparatively rare parasites in cockroaches.

Family PHORIDAE

=Megaselia= sp.

_Host._--Eggs of _Parcoblatta_ sp., Ohio (Edmunds, 1952a).

Family CONOPIDAE

=Stylogaster stylata= (Fabricius)

_Hosts._--Cockroaches, Brazil (Souza Lopes, 1937): L. Travassos was quoted as having observed this species pursue cockroaches that were escaping columns of the army ant _Eciton_ sp. Souza Lopes (1937) stated that the female deposits eggs on the cuticle of the host near the end of the body; the egg is barely inserted and two recurrent hooks prevent it from falling off. Souza Lopes (1937) also observed other species of _Stylogaster_ pursue Orthoptera, but he was unable to devote proper attention to the behavior of the flies.

=Stylogaster= spp.

_Hosts._--_Chorisoneura_ sp., Brazil (Souza Lopes, 1937): An adult specimen was found in a museum collection with an egg of _Stylogaster_ attached to the posterior end of its abdomen.

Cockroaches, Panama (C.W. Rettenmeyer, personal communication, 1959): "Seven species were collected hovering over army ant swarms and a few flies were seen apparently attacking cockroaches that had been flushed by the ants."

Family LARVAEVORIDAE

=Calodexia= (?) =venteris= Curran

_Hosts._--_Periplaneta americana_, Brazil (Souza Lopes, 1937): Obtained complete evolution of the parasite in this host. This may have been an experimental host.

=Calodexia= spp.

_Hosts._--Cockroaches, Panama (Rettenmeyer, personal communication, 1959): Swarms of army ants are accompanied by about 20 species of _Calodexia_. These flies larviposit on the cockroaches, crickets, and possibly other arthropods that are flushed from cover by the ants. Larvae were found in one(?) cockroach. Larvae from an adult of _Calodexia_ were introduced experimentally into a cockroach and successfully reared.

=Undetermined tachinids=

_Hosts._--_Eurycotis floridana_, from Florida (Roth, unpublished data, 1953): Three larvae (det. by W.W. Wirth) were found in a living adult male.

_Panesthia australis_, from Australia (Roth, unpublished data, 1957): Reared from a wild-caught cockroach that was maintained in a laboratory colony.

Cockroaches, Australia (E. F. Riek, personal communication, 1955): Reared from some of the larger species.

Family MUSCIDAE

=Coenosia basalis= Stein

_Host._--Eggs of _Parcoblatta_ sp., Ohio (Edmunds, 1952a).

Family SARCOPHAGIDAE

=Sarcophaga omani= Hall

_Host._--_Arenivaga bolliana_, Texas (Wirth, personal communication, 1953): Specimens in U.S. National Museum.

=Sarcophaga lambens= Wied.

_Synonymy._--_Sarcophaga sternodontis_ (Towns.).

Hoffman (1927) claimed that approximately 40 percent of some specimens of _Pycnoscelus surinamensis_ collected in southern Haiti were parasitized by _S. lambens_. However, according to entomologists at the University of Puerto Rico Agricultural Experiment Station, Hoffman was incorrect in his observations: _S. lambens_ was never reared from a living insect and had been recovered only from dead cockroaches and other dead insects and was considered saprophytic rather than parasitic (Schwabe, 1950b).

=Sarcophaga= spp.

Sanjean (1957) reared various species of sarcophagid larvae on _Periplaneta americana_ which were freshly killed or chopped up; first instar larvae were also introduced into the body cavity of cockroaches which had their heads and legs removed. Adult sarcophagids were collected and freshly killed American cockroaches used as bait.

Order COLEOPTERA

Family CARABIDAE

=Harpalus pennsylvanicus= De Geer

_Experimental prey._--_Cryptocercus punctulatus_, U.S.A. (Cleveland et al., 1934): This beetle is often found in the galleries of _C. punctulatus_ in nature. In the laboratory it killed and devoured cockroaches as large as itself.

Family DYTISCIDAE

=Rhantus pacificus= Boisduval

_Experimental prey._--Cockroaches, disabled, Hawaii (Williams, 1936): This beetle, which is common in mountain streams, located wounded cockroaches in an aquarium by sense of smell or taste rather than sight.

Family LAMPYRIDAE

=Undetermined larva=

_Experimental prey._--_Parcoblatta virginica_, adult female (pl. 33, C), U.S.A. (Roth and Willis, unpublished data, 1953).

Family RIPIPHORIDAE[5]

=Neonephrites partiniger= Riek

_Natural host._--Cockroach (undescribed genus belonging to the Pseudomopinae), Australia Capital Territory (Riek, 1955).

=Neorhipidius neoxenus= Riek

_Natural host._--_Robshelfordia longiuscula_ or _Robshelfordia circumducta_, Australia Capital Territory (Riek, 1955).

=Paranephrites xenus= Riek

_Natural host._--_Oniscosoma granicollis_, Australia Capital Territory (Riek, 1955).

=Rhipidioides ableptus= Riek

_Natural host._--_Balta patula_, Australia, Victoria (Riek, 1955): Pupal stage lasted only 3 days.

=Rhipidioides adynatus= Riek

_Natural host._--_Escala_ sp. or an undescribed genus of Pseudomopinae, Australia, Victoria (Riek, 1955).

=Rhipidioides fuscatus= Riek

_Natural host._--_Ellipsidion affine_, Australia, New South Wales (Riek, 1955).

=Rhipidioides helenae= Riek

_Natural host._--_Robshelfordia longiuscula_ or _Robshelfordia circumducta_, Australia Capital Territory (Riek, 1955).

=Rhipidioides mollis= Riek

_Natural host._--_Robshelfordia longiuscula_ or _Robshelfordia circumducta_, Australia Capital Territory (Riek, 1955).

=Rhipidioides rubricatus= Riek

_Natural host._--_Choristima_ sp. and _Choristimodes_ sp., Australia Capital Territory (Riek, 1955).

=Riekella australis= (Riek)

_Synonymy._--_Nephrites australis_ Riek [Selander, 1957].

_Natural host._--_Cutilia_ sp., Australia Capital Territory (Riek, 1955): Two females emerged from one host.

=Riekella nitidioides= Selander

_Synonymy._--_Nephrites nitidus_ of Riek not Shuckard [Selander, 1957].

_Natural host._--_Platyzosteria_ sp., Tasmania (Riek, 1955).

=Riekella= sp.

_Synonymy._--_Nephrites_ sp. [Selander, 1957].

_Natural host._--_Platyzosteria castanea_, Australia Capital Territory (Riek, 1955).

_Biology of Australian Ripidiini._--The Australian species of Ripidiini are parasites of apparently endemic, ground-dwelling species of cockroaches. There is some correlation between host subfamily and parasite genus: _Riekella_ spp. [= _Nephrites_] have only been bred from Blattinae. _Rhipidioides_ spp. occur only in the closely related Ectobiinae and Pseudomopinae. _Neonephrites_ and _Neorhipidius_ also occur in the Pseudomopinae. _Paranephrites_ occurs in the Panchlorinae. There is some evidence that the parasitized cockroaches migrate onto trees when the larval parasite is mature, as pupae have only been found on the trunks of eucalyptus trees. In all species the larva leaves the host dorsally through an intersegmental membrane. The host continues to live for a few days after the parasite emerges. The larva attaches itself to bark on the tree trunk by a few strands of silk before pupating. The larviform, wingless female remains near the pupal skin and is sought out by the winged male. The eggs are laid in a mass around the pupal skin (Riek, 1955).

=Ripidius[5] boissyi= Abeille

Balduf (1935) lists _Ripidius boissyi_ as parasitic on nymphs of _Ectobius pallidus_ giving Abeille de Perrin (1909) as a source for this information. However, Abeille de Perrin simply presumed that _R. boissyi_ parasitized _E. pallidus_ because he collected this cockroach in the same habitat as the beetle. Abeille de Perrin suggested that the species of the genus _Ripidius_ lived in the bodies of cockroaches, but there are no rearing records, as far as we know, of _R. boissyi_ from cockroach hosts.

=Ripidius denisi= Chobaut

Chobaut (1919), in France, collected both _R. denisi_ and _Ectobius pallidus_ when beating an oak tree. Because of the known association of other species of _Ripidius_ with cockroaches, he presumed that this beetle was parasitic on _E. pallidus_, a cockroach common in this beetle's habitat.

=Ripidius pectinicornis= Thunberg

_Synonymy._--_Symbius blattarum_ Sundevall [Leng, 1920].

_Natural hosts._--_Blattella germanica_, on shipboard (Sundevall, 1831); Germany (Aclogue and Fowler, _in_ Burr, 1899a); on steamship "Samui" (Stamm, 1936); on cruiser "Duguay-Trouin" (Barbier, 1947); Hawaii (Williams, 1946a): This last record was based on a specimen dissected from an adult German cockroach collected on an airplane from the South Pacific. The parasite was reported as _Ripidius_ sp. by Williams, but Weber (1948) made the specific identification.

_Ectobius pallidus_? Abeille de Perrin (1909) stated that _R. pectinicornis_ was first described by Sunders [sic] as _blattarum_ because it had been captured in the body of _Ectobia livida_. We presume that Abeille de Perrin was referring to Sundevall's work in which the host was given as _Blattella germanica_.

_Periplaneta americana_, on shipboard (Sundevall, 1831): One nymph only.

With the exception of the single nymph of _P. americana_, _R. pectinicornis_ apparently attacks only adult females and nymphs of _B. germanica_. Barbier (1947) found only _B. germanica_ parasitized, although both _Blatta orientalis_ and _Supella supellectilium_ were prevalent on board the ship. Primary larvae of the parasite failed to parasitize _Supella_.

_Adult behavior._--The winged male is relatively active compared to the apterous female; it runs around, flies well, and jumps on the female when in her vicinity. The female remains stationary and lays eggs around her by bending her long ovipositor (Sundevall, 1831). The eggs (50-100) are laid among a network of silk fibers secreted by the female. The female dies after completing oviposition (Barbier, 1947).

_Development._--The eggs hatch after 14 days, and the primary (triungulin) larvae ascend the host's legs to its body; the larvae then cut the intersegmental membrane between the metasternum and first abdominal segment of the cockroach, in order to enter the host's abdomen (Barbier, 1947). Chobaut (1892) first suggested this method of attack by the ripiphorid larva. As the parasites develop, the abdomen of the host becomes swollen. Developing larvae apparently eat the host's fat body, leaving the vital organs until the last. Parasitized female hosts were sterile and the eggs, when formed, never hatched. Development of the oötheca was also inhibited. There were usually two larval parasites per host, but three or four were found several times (Barbier, 1947). Sundevall (1831) found only one larva per cockroach except one host which, when crushed, yielded five. Stamm (1936) found three hosts infested with five larvae each. In a little over 100 cockroaches, Stamm found 10 that were parasitized.

The day before the parasite leaves the host, the cockroach shows an abrupt uneasiness and runs about, finally falling over on its back. The parasite larva emerges from the host through an opening it makes in the membrane between penultimate and last tergite. The host dies a few hours after the larva has left. The larva seeks a sheltered area and pupates within 48 hours. Adults emerge in 9 days (females) and 13 days (males) (Barbier, 1947).

_Distribution._--Adult males have been collected in light traps in Hawaii (Van Zwaluenburg, 1946), and the first female was reported by Weber (1948); the parasite is now established in the islands around Pearl Harbor (Dr. F. X. Williams, personal communication, 1953). The U. S. National Museum has specimens of _R. pectinicornis_ from England, Guatemala, Hawaii, Panama, and from Florida and Georgia in the U. S. (Dr. E. A. Chapin, personal communication, 1953). Kono (_in_ Asano, 1937) reported two species in Japan. It is noteworthy that all these records are from localities adjacent to oceans and on ships; none are from interiors of continents. The only biological data were obtained from parasites found on board ships. Sundevall (1831) believed that the parasites boarded his ship with their hosts during loading in Calcutta, since before that not any were seen on board. Barbier (1947) suggested that the parasite must be spread very easily in ports between neighboring ships by parasitized cockroaches in baskets or sacks of provisions.

=Ripidius scutellaris= Heller

_Natural hosts._--Blattidae, Philippine Islands (Schultze, 1925).

Family DERMESTIDAE

=Dermestes ater= De Geer

_Common name._--Black larder beetle.

_Natural prey._--_Blatta orientalis_, U.S.A. (Roth and Willis, unpublished data, 1953): _Dermestes ater_ is generally a scavenger, but we have seen adult beetles, which had developed in our cockroach colony, clinging to and feeding on living oriental cockroaches, eventually killing them; the beetles probably attack only the weakened or injured cockroaches in a culture. This was a natural infestation of a laboratory culture by a predator.

_Experimental prey._--_Blattella germanica_, oöthecae, U.S.A. (Roth and Willis, 1950): The beetle larvae can penetrate unhatched oöthecae of the German but not those of the American or oriental cockroaches.

=Dermestes= sp.

_Natural prey._--_Blatta orientalis_, oöthecae, U.S.A., Missouri: Rau (1924) stated that _Dermestes_ larvae often infest the egg cases of this cockroach; it is probable that Rau was referring to cockroaches in laboratory cultures.

Order STREPSIPTERA

Pierce (1909) predicted that the Blattoidea and the Grylloidea would be the only groups of the Orthoptera which would be parasitized by Strepsiptera. Essig (1926) made the statement that certain cockroaches are among the hosts of Strepsiptera. E. F. Riek (personal communication, 1952) found a strepsipteron in a late nymph of _Cutilia_ sp. from Waroona, Western Australia; he wrote us, "The female parasite is extruded between a pair of sternites towards the base of the abdomen and appears to belong to the family Halictophagidae." This is the only record that we have been able to find of a strepsipteron parasitizing cockroaches.

Order HYMENOPTERA

PREDATORS AND PARASITES OF COCKROACH EGGS

Wasps from at least six families of Hymenoptera have been recorded as developing on cockroach eggs. All the Evaniidae are presumed to be parasitic in the egg capsules of cockroaches (Clausen, 1940; Townes, 1951), although hosts for many of the described species have yet to be discovered. The presence of evaniids in dwellings indicates the presence of cockroaches (Gross, 1950). At times these wasps may become a nuisance; a family in Worthington, Ohio, complained of the evaniid wasps that they found on the windows and in other areas of their home, but they were apparently not annoyed by the oriental cockroaches in the basement (Edmunds, 1953).

The known parasites of cockroach eggs are listed below with summaries of their biology.

Family EVANIIDAE

=Acanthinevania princeps= (Westwood)

_Synonymy._--_Evania princeps_ [Dr. H. Townes, personal communication, 1956].

_Natural host._--Cockroach eggs, Australia (Froggatt, 1906).

=Brachygaster minutus= (Olivier)

_Synonymy._--_Evania minuta_ Olivier [Kieffer, 1920].

_Natural hosts._--_Blattella germanica_, Europe? (Schletterer, 1889; Kiefer, 1912; Crosskey, 1951.)

_Ectobius lapponicus_, Europe? (Schletterer, 1889; Kieffer, 1912; Crosskey, 1951.)

_Ectobius panzeri_ var. _nigripes_? Great Britain (Blair, 1952): This is a presumptive record. The wasp was collected at Niton and Headon Hill, Isle of Wight, an area in which this variety of _E. panzeri_ was the only species of cockroach known to occur.

_Ectobius_ sp., England (Cameron, 1955, 1957): Natural History Museum records.

Adult wasps have been collected on _Asparagus officinalis_ Linnaeus (Schmiedeknecht _in_ Schletterer, 1889; Crosskey, 1951). Thompson's (1951) citation of records of _B. minutus_ and _Evania appendigaster_ from _Blatta orientalis_ and _Blattella germanica_, and Cameron's (1957) citation of these records and one from _Ectobius lapponicus_, all attributed to Kadocsa (1921), are almost certainly in error. Kadocsa (1921, p. 33) listed these wasps as egg parasites of cockroaches but not necessarily in Hungary and did not name specific cockroach hosts.

The present writers have found no information, other than host reports, on the biology of _Brachygaster minutus_. The records of this wasp parasitizing _B. germanica_ may trace back to Schletterer, but his listing may not have been an original observation. Since the female of _B. germanica_ carries its oötheca attached to the abdomen until or just before the eggs hatch, it would seem that the female of _B. minutus_ (if the host records are valid) must oviposit into the oötheca of this species while it is still being carried by the female; this would not necessarily be true for the other hosts which drop the egg case long before the eggs hatch.

_Distribution._--Europe: Sweden, Russia, England, France, Germany, Austria, Hungary, Switzerland, Italy (Kieffer, 1920).

=Evania appendigaster= (Linnaeus)

_Synonymy._--_Evania desjardinsii_ Bordage, _Evania laevigata_ Latreille [Dalla Torre, 1901-1902].

_Natural hosts._--_Blatta_, "exotic species" (Westwood, 1854, 1954a).

_Blatta orientalis_, Europe? (Schletterer, 1886; Howard, 1888, Kieffer, 1912); Egypt? (Alfieri, 1914; Adair, 1923). [Girault (1907, 1914) erroneously attributed another record to Marlatt (1902);[see footnote 6]. See also notes under _Brachygaster minutus_ with respect to Kadocsa.]

_Blattella germanica_? (Girault 1907, 1914). [This record is obviously an error. Girault attributed the record to Marlatt (1902); see footnote 6.]

_Cutilia soror_, Hawaii (Swezey, 1929; Zimmerman, 1948).

_Leucophaea maderae_ (Schletterer, 1889; Bordage, 1896; Kieffer, 1912): These records are probably erroneous inasmuch as this cockroach incubates its eggs internally (Roth and Willis, 1954). Later, after finding that _L. maderae_ is ovoviviparous, Bordage (1913) admitted having misidentified a parasitized oötheca from some other species; he concluded that the developing eggs of this species are protected against egg parasites because they are carried within the female. Clausen (1940), in classifying the placement of parasitic wasp eggs in relation to the host, erected the category: Egg placed in the embryo while the latter is still within the parent. He stated that although this behavior was not definitely known to occur, it probably could occur. However, the records cited above do not indicate that the alleged parasitization followed this pattern.

_Neostylopyga rhombifolia_, Hawaii (Swezey, 1929).

_Periplaneta americana_, Europe (Schletterer, 1889; Bordage, 1896; Kieffer, 1912); Réunion Island (Bordage, 1913); Puerto Rico (Seín, 1923); Jamaica (Gowdey, 1925); Hawaii (Swezey, 1929); Palestine (Bodenheimer, 1930); U.S.A., Florida (Ashmead, 1900); Maryland (Piquett and Fales, 1952); Saudi Arabia, Jedda (Cameron, 1957); Canton Island and Samoa (Dumbleton, 1957).

_Periplaneta americana_ or _P. australasiae_, Formosa (Sonan, 1924).

_Periplaneta australasiae_, U.S.A., Florida (Ashmead, 1900); Hawaii (Swezey, 1929; Zimmerman, 1948). [Girault (1914) erroneously attributed another record to Marlatt (1902); see footnote 6, above.]

_Experimental host._--_Blatta orientalis_, U.S.A. (Haber, 1920).

Relatively little detailed information was known about this wasp (fig. 5), one of the earliest parasites of cockroach eggs to be discovered, until Cameron (1957) studied its biology. Arnold (Kirby and Spence, 1826) discovered that the genus _Evania_ parasitized _Blatta_, but did not know whether the wasp developed on the cockroach eggs or in the nymphs. MacLeay (Westwood, 1843) determined that _Evania_ developed within the oöthecae of cockroaches. Westwood (1854a) found the larvae, pupae, and adults of _E. appendigaster_ in egg cases of an unidentified species of cockroach found on orchids received from Calcutta.

_Adult behavior._--Adult wasps visited flowers of parsley, _Petroselium crispum_, and fennel, _Foeniculum vulgare_ (Margretti _in_ Schletterer, 1886; Crosskey, 1951). In Hawaii the adult wasps have been seen resting on leaves coated with honey dew (Williams et al., 1931); _Evania_ sp. were attracted to the honey dew secreted by a diaspine scale insect (Williams, 1931). Adults lived two to three weeks in captivity with ample food and water (Cameron, 1957).

_Oviposition._--Shelford (1912, 1916) erroneously supposed that _Evania_, by means of her cleaverlike abdomen, opened the oötheca at the crista and then deposited her egg or eggs on the eggs of the cockroach. Haber (1920) observed and described oviposition. The female wasp crawled over the surface of the oötheca, actively vibrating her antennae, and settled with the axis of her body parallel to the axis of the egg case as it lay upon its right side. Lying on her right side, the wasp extended her ovipositor and punctured the oötheca in the fifth cell on the left side; she remained in this position for about 15 minutes. Cameron (1957) described similar oviposition behavior that lasted about half an hour. Kieffer (1912) and Crosskey (1951) stated that the female deposits her eggs before the walls of the oötheca harden.

, 1951, figs. 1A and 1B.)]

_Development._--Kieffer (1912) stated that the larvae in this family eat the cockroach eggs and pupate in the oötheca without forming a cocoon. Smith (1945) stated that the larva feeds on one cockroach egg after another until all are destroyed; by that time it is full grown and it pupates within the oötheca. Cameron (1957) found that there are five larval instars and that in material from Saudi Arabia there are three or possibly four generations a year.

_Distribution._--Tropical and subtropical parts of the world as far north as New York City, and all of Europe except the northern part (Kieffer, 1920; Townes, 1949). The wide distribution of _Evania_ has been attributed to the abundance of host cockroaches on ships between the Tropics (Haldeman, 1847). Kieffer (1903) appears to have shown some correlation between the numbers of species of cockroaches found in various geographical regions and the numbers of species of evaniids found in similar regions. However, the number of blattids he listed is small.

=Evania dimidiata= Fabricius

_Synonymy._--_Evania abyssinica_ Westwood [Schletterer, 1889].

_Natural host._--_Blatta orientalis_, Egypt? (Alfieri, 1914).

=Evania subspinosa= Kieffer

_Natural host._--_Periplaneta_ sp., Fiji (Lever, 1946): Although Lever (1946) listed this species as a cockroach-egg parasite, he did not state that he actually reared it from _Periplaneta_ oöthecae.

=Hyptia dorsalis= of Ashmead

_Synonymy._--Dr. H. Townes, (personal communication, 1956) believes that this wasp was probably either _H. reticulata_, _H. harpyoides_, or _H. thoracica_; it is not possible to tell which without reexamining Ashmead's specimens; these apparently have been lost.

_Natural host._--_Parcoblatta pensylvanica_, U.S.A., Mississippi (Ashmead, 1900).

=Hyptia harpyoides= Bradley

_Natural hosts._--_Parcoblatta virginica_, U.S.A., Ohio (Edmunds, 1952a, 1953a, 1954).

_Parcoblatta pensylvanica_, U.S.A. (Muesebeck, 1958).

_Parcoblatta uhleriana_, U.S.A., Natick, Mass.: Oötheca collected by L. Roth, May 17, 1956; wasp emerged June 12, 1956 (pl. 33, B); determined by Dr. H. Townes. The keel region of the oötheca of _P. uhleriana_ (pl. 18, B) is different from that of any other species of _Parcoblatta_ (Hebard, 1917; Lawson, 1954) so there can be no doubt as to the species of cockroach parasitized by this wasp.

_Development._--The last instar larva overwinters inside the cockroach oötheca (Edmunds, 1954). Five oöthecae yielded one parasite each (Edmunds, 1953a).

_Distribution._--Canada, Ontario. U.S.A.: New Hampshire and Minnesota to South Carolina, Mississippi, Texas, and Kansas. Upper and Lower Austral Zones (Townes, 1951).

=Hyptia reticulata= Say

_Natural host._--_Parcoblatta pensylvanica_, U.S.A., Missouri (Rau, 1940).

Adult wasps have been taken on parsnip, _Pastinaca sativa_ (Robertson, 1928).

_Distribution._--U.S.A.: Pennsylvania to Florida and Louisiana. Mexico. Upper Austral to Tropical Zones (Townes, 1951).

=Hyptia thoracica= (Blanchard)

_Natural host._--_Parcoblatta pensylvanica_, U.S.A., Ohio (Edmunds, 1952a, 1953a, 1954).

_Adult behavior._--Copulation was rapid, lasting only a few seconds. Blooms of _Asmorrhiza longistylis_ were placed in a cage with adult wasps. The insects were attracted to and fed on the flowers (Edmunds, 1954).

_Development._--Entire contents of oötheca are eaten by the single larva. Last instar larva overwinters inside the oötheca. Emergence in Ohio was around the middle of June. The emergence hole made by this genus was about 2 mm. in diameter. The hole was made at the top side of the oötheca near one end. Adult took about 65 minutes to emerge from the time its mandibles first broke through the oöthecal wall. (Edmunds, 1954.)

_Distribution._--Canada, Ontario, U.S.A.: Connecticut to Wisconsin, south to Florida and Texas. Upper Austral to Tropical Zones. (Townes, 1951.)

=Hyptia= sp.

_Natural host._--_Cariblatta delicatula_, Cuba (Hebard, 1916a); Parasite identified by Ashmead.

=Hyptia= sp. (undescribed)

_Natural host._--_Parcoblatta_ sp., U.S.A., Ohio (Edmunds, 1952a).

=Prosevania punctata= (Brullé)

_Synonymy._--_Evania punctata_ Brullé [Townes, 1949].

_Natural and experimental hosts._--_Blatta orientalis_, Istrian Peninsula (Fahringer, 1922); Algeria (Cros, 1942); U.S.A., Ohio (Edmunds, 1954).

_Blattella germanica_? Europe? (Girault 1907, 1914); Europe (Fahringer, 1922). [The records on this host are extremely doubtful. Girault erroneously cited Marlatt (1902) as the source of this record; see footnote 6, page 236. Fahringer, however, claimed that he obtained seven female parasites from oöthecae of _Blattella germanica_. He placed female parasites with adults of _B. germanica_ in a glass cage. As soon as oöthecae could be seen between folds of a woolen rag, he removed all the larger cockroaches and held the oöthecae until the parasites emerged. Fahringer may have been dealing with a different species of cockroach, because placing oöthecae in crevices (or between folds of rag) is a habit foreign to _B. germanica_, the female of which usually carries her oötheca until hatching or until about a day before. Edmunds (1953b) could not induce this wasp to parasitize eggs of _B. germanica_.]

_Periplaneta americana_, Istrian Peninsula (Fahringer, 1922); Palestine (Bodenheimer, 1930); U.S.A., Ohio (Edmunds, 1952, 1953b, 1954).

_Adult behavior._--The wasps (pl. 33, A) are very active; they walk about a great deal and fly short distances. They are often found in abundance in buildings infested with the larger domiciliary cockroaches where they may reproduce for many generations without leaving the premises. Specimens have also been collected outdoors. (Edmunds, 1953, 1954.) As the adult walks about, the laterally compressed abdomen moves up and down like a waving flag; because of this behavior, these insects are commonly known as ensign-flies. Cros (1942) maintained adults 17 days without food. Edmunds (1954) fed adults on unidentified flowers in the laboratory. He also maintained them for 20 days after capture on a 5-percent honey solution.

_Oviposition._--A female _P. punctata_ selected oöthecae of _P. americana_ for oviposition and ignored those of _B. orientalis_ and _Parcoblatta pensylvanica_ in the same cage. Oviposition was accomplished as described for _Evania appendigaster_. One oötheca was turned over onto its right side by the wasp before she oviposited. (Edmunds, 1952.) Although there seemed to be a "preferred" position for oviposition, it was not obligatory. The usual position was for the female to face the keel of the oötheca, but she also oviposited from the opposite side or, rarely, directly down into the side of the oötheca. The average time spent by females in 10 ovipositions was 29 minutes (range 16-62 minutes). The wasp apparently could not determine whether the eggs had been previously parasitized. The wasp laid her egg between the cockroach eggs rather than in them and she oviposited into oöthecae that had just been dropped and those two weeks old or older. On three occasions nymphal cockroaches emerged within a few hours after the wasp had oviposited. (Edmunds, 1954.) Apparently, for successful parasitization the wasp must oviposit before the cockroaches have reached the final stages of preemergence development. Edmunds (1954) placed females of _Periplaneta americana_ that were carrying oöthecae, into cages with _Prosevania_; some of the female wasps showed considerable interest in the attached oöthecae, but he observed oviposition only into egg cases that had been dropped by the cockroaches.

Cros (1942) described an interesting reaction that he called "instinctive hostility" of the oriental cockroach toward _Prosevania_. A wasp was placed in a jar in which a cockroach had just deposited its oötheca. The wasp tried to oviposit into the egg case but was upset and pursued by the cockroach. The cockroach placed herself over the oötheca, standing high on her legs, and remained there motionless. The wasp then approached from the rear, slipped under the cockroach, and, unnoticed by the cockroach, climbed on the oötheca and oviposited successfully.

_Development._--In _Blatta orientalis_: The developmental period was completed in 40-57 days in summer and fall (Cros, 1942). Time from oviposition to emergence of adult varied from 45-177 days; three parthenotes from an oviposition by an unfertilized female wasp developed in 45-53 days (Edmunds, 1954). In _Blattella germanica_: Almost 4 weeks spent in development (Fahringer, 1922). In _Periplaneta americana_: Three wasps developed in 127 days (Edmunds, 1952). Only one parasite develops in each oötheca. There were three generations a year in Ohio. (Edmunds, 1954.) In Algeria there were two to three generations per year. The adult emerged from the oötheca through a hole 4 mm. in diameter. (Cros, 1942.) Parthenogenesis exists; the unfertilized eggs produced only males (Edmunds, 1954).

_Distribution._--Eastern U.S.A., from New York and Ohio south to Georgia (Townes, 1949). Europe, Syria, Palestine (Kieffer, 1920).

=Szepligetella sericea= (Cameron)

_Synonymy._--_Evania sericea_ Cameron [Townes, 1949, personal communication, 1956]. _Evania impressa_ Schletterer [Townes, p. c., 1956].

_Natural hosts._--_Cutilia soror_ and _Neostylopyga rhombifolia_, Hawaii (Swezey, 1929).

_Periplaneta americana_ and _Periplaneta australasiae_, Hawaii (Swezey, 1929; Zimmerman, 1948).

_Periplaneta_ sp., Fiji (Lever, 1943, 1946).

Adults are sometimes found resting on leaves covered with honey dew (Williams et al., 1931).

=Zeuxevania splendidula= Costa

_Natural hosts._--_Loboptera decipiens_, France (Lavagne, 1914; Genieys, 1924).

Picard (1913) believed that _Z. splendidula_ parasitized _L. decipiens_ and not its eggs; however, Lavagne (1914) explained the true relationship by dissecting two specimens of _Z. splendidula_ from oöthecae of _L. decipiens_.

The following information is taken from Genieys (1924): _Oviposition._--Wasp egg is introduced into the still-soft oötheca before the wall hardens. Some oöthecae had four oviposition scars but never contained more than two parasite eggs. _Development._--Larva commences development in July or August. Only one larva completes development, but it eats all the eggs in the oötheca. The wasp passes the winter as a last instar larva and pupates in the spring; the adult emerges during the spring or in June. _Hyperparasitism._--About 10 percent of the oöthecae of _Loboptera decipiens_ that were parasitized by _Z. splendidula_ were also hyperparasitized by an eulophid (see _Syntomosphyrum ischnopterae_, p. 249).

Family CLEONYMIDAE

=Agamerion metallica= Girault

_Natural hosts._--_Ellipsidion australe_, Australia, Queensland (Dodd, 1917): "the parasite when ready to emerge fully occupies the whole space of the destroyed eggs."

Cockroach, Australia, New South Wales (Dr. B. D. Girault, 1915a).

Family ENCYRTIDAE

=Blatticida pulchra= Ashmead

_Natural host._--Cockroach eggs on orange leaves, Australia, New South Wales (Gahan and Peck, 1946). According to Dr. A. B. Gurney the oötheca associated with the type specimens of the wasps in the United States National Museum is possibly _Balta_ sp. (Burks, personal communication, 1956).

=Blatticidella ashmeadi= (Girault)

_Synonymy._--_Blatticida ashmeadi._ _Blatticida_ Girault, 1915, is preoccupied by _Blatticida_ Ashmead, 1904. In 1923 Gahan and Fagan renamed _Blatticida_ Girault, _Blatticidella_. [Burks, p. c., 1956.]

_Natural host._--Cockroach, Australia, Queensland (Girault, 1915).

=Cheiloneurus viridiscutum= (Girault)

_Synonymy._--_Cristatithorax_ Girault = _Cheiloneurus_ Westwood [Mercet, 1921].

_Natural host._--_Ellipsidion australe_, Australia, Queensland (Dodd, 1917).

=Comperia merceti= (Compere)

_Synonymy._--_Comperia merceti_ var. _falsicornis_ Gomes [Peck, 1951].

_Natural hosts._--_Blattella germanica_, Brazil, Distrito Federal (Gomes, 1941): In the English summary of his paper, Gomes states that _C. merceti_ var. _falsicornis_ was reared from _B. germanica_. However, in the body of the paper, he states that the _supposed_ origin of the parasite was the oötheca of _B. germanica_. Burks (personal communication, 1956) does not believe that this wasp parasitizes the eggs of _B. germanica_. We (unpublished data, 1957) exposed six oöthecae of _B. germanica_ to _C. merceti_. In order to retard water loss the oöthecae were removed from the females by cutting the insects in two so that each oötheca remained attached to the posterior part of the abdomen. No wasps developed in these oöthecae.

_Supella supellectilium_, U.S.A., Kansas (Lawson, 1954a); Hawaii (Zimmerman, 1944; Compere, 1946; Keck, 1951).

_Adult behavior._--Males and nonovipositing females showed a flea-like jumping tendency. Adults were attracted to light and were found near windows. Both sexes pursued an erratic course in walking and continually touched the surface with their antennae. (Lawson, 1954a.)

_Oviposition._--The wasp (pl. 34, B) selected a site on an oötheca with the sheath of her ovipositor; it was uncertain whether there was a definite preference for oviposition sites. Wasp tended to choose a nearly horizontal position for oviposition. She preferred to oviposit into eggs about 2 weeks old, although she would place eggs in oöthecae less than a week old and in embryos in the green band stage. There were 1-50 oviposition punctures per oötheca. (Lawson, 1954a.)

_Development._--If enough wasp larvae were present, they ate all eggs in an oötheca. Occasionally wasps developed in one end of an oötheca while cockroaches developed in the other; when this occurred, the cockroach nymphs always emerged last. The developmental period was 30-41 days at room temperature. There were 5-25 parasites per oötheca. The single exit hole in the oötheca varied from 0.6 to 0.9 mm. in diameter. (Lawson, 1954a.)

_Distribution._--U.S.A.: New Jersey south to Florida, west to Illinois, Kansas, and Arizona. West Indies; Central and South America; Hawaii. (Burks, personal communication, 1956.)

=Dicarnosis alfierii= Mercet

_Natural hosts._--"_Phyllodromia_" sp., Egypt (Mercet, 1930): According to Mercet, Dr. Alfieri claimed that this wasp parasitized one of the species of "_Phyllodromia_" found in Egypt, namely, _Phyllodromia_ [= _Blattella_] _germanica_, _Phyllodromia_ [= _Supella_] _supellectilium_ and/or _Phyllodromia treitliana_. We do not know to which modern genus the host of this wasp belonged.

Cockroach, Egypt? (Mercet in Compere, 1938.)

=Eutrichosomella blattophaga= Girault

_Natural host._--Cockroach, Australia, Queensland (Girault, 1915).

Family EUPELMIDAE[7]

=Anastatus blattidifurax= Girault

_Natural host._--Cockroach, Australia, Queensland (Girault, 1915).

=Anastatus floridanus= Roth and Willis

_Natural host._--_Eurycotis floridana_, U.S.A., Florida (Roth and Willis, 1954a).

_Experimental hosts._--_Blatta orientalis_, _Eurycotis floridana_, and _Periplaneta americana_, U.S.A. (Roth and Willis, 1954a).

_Adult behavior._--Female wasps are sexually receptive almost immediately on leaving the oötheca. Mating takes 3-4 seconds. Males mate repeatedly and may fertilize several females; females may also mate more than once. At about 80° F. the female wasps lived 2-4 days, males one day.

_Oviposition._--The female wasp first probes the oötheca with her sheathed ovipositor until she finds an acceptable spot; she then drills through the wall of the oötheca with her ovipositor. One female oviposited for 5 hours, but briefer periods were more usual. We have seen six or more females ovipositing simultaneously into an oötheca of _Eurycotis floridana_. One female was seen to feed on material that oozed from the oviposition puncture. The wasp (pl. 34, A) may oviposit into the oötheca of _E. floridana_ while it is still being carried by the female, as well as in oöthecae that have been dropped and which have hard walls. Eggs 36 days old were successfully parasitized.

_Development._--In _Eurycotis floridana_: In the laboratory, development was completed in 34-36 days at about 85° F. This time was regulated to some extent by the number of parasites in the oötheca.

There is evidence that larvae eat unhatched wasp eggs or other larvae. In 34 oöthecae exposed to many female wasps, the maximum number of parasites to emerge was 306; yet an average of 601 wasp larvae were dissected from four oöthecae that had each been exposed to 50 female wasps one week earlier. The larvae usually eat all the host eggs. Cockroach eggs that were not eaten by the wasp larvae sometimes developed but usually failed to hatch. Adult wasps made one to six emergence holes in the oötheca; the average number in 42 oöthecae was two holes.

_Number of parasites per oötheca._--In _Blatta orientalis_: One of 111 oöthecae exposed to female wasps yielded 48 parasites. In _Eurycotis floridana_: One oötheca parasitized in the field yielded 68 parasites; 8 oöthecae exposed to single wasps for their entire lifespan yielded an average of 50 ± 6 parasites (range 23-81); 34 oöthecae exposed to many wasps for their entire lifespan yielded an average of 198 ± 8 parasites (range 93-306). In _Periplaneta americana_: Nine oöthecae of 152 exposed to the wasps were found to be parasitized when dissected; 11 adults emerged from one oötheca; no parasites emerged from the other 8 oöthecae.

_Sex ratio._--4 [F][F]:1 [M] from ovipositions by isolated females. In the one oötheca collected in the field, the ratio was 21.6 [F][F]:1 [M]. Parthenogenesis exists; the unfertilized eggs produced only males.

=Anastatus tenuipes= Bolívar y Pieltain

_Synonymy._--_Anastatus blattidarum_ Ferrière. Dr. C. Ferrière (personal communication, 1957) is of the opinion that his _A. blattidarum_ is a synonym of _A. tenuipes_. He stated "I have never been able to see the unique type of _A. tenuipes_ B. y P., which is in Madrid, but the description agrees with _A. blattidarum_. I had not yet knowledge of Bolívar's description, when describing my species. The parasite of cockroaches eggs [_Supella supellectilium_] should be called _A. tenuipes_ Bol." Mani (1938) synonymized _Solindenia blattiphagus_ Mani with _Anastatus blattidarum_.

_Natural hosts._--_Supella supellectilium_, Anglo-Egyptian Sudan (Ferrière, 1930, 1935); U.S.A., Arizona (Flock, 1941); Egypt (Alfieri, _in_ Hafez and Afifi, 1956). Ohio (Hull and Davidson, 1958).

_Periplaneta americana_, India (Burks _in_ Roth and Willis, 1954a).

Cockroach, Hawaii (Weber, 1951); India (Mani, 1936).

The following is based on parasites that developed on eggs of _Supella supellectilium_ (Flock, 1941): _Adult behavior._--Wasp may be seen running rapidly on walls in buildings infested with the cockroach host. The wasp rarely flies but hops proficiently; when disturbed it can hop from several inches to several feet. The female licks up the drop of fluid that oozes from the oviposition puncture. Females die in a few days, but if fed honey and water may live two weeks. _Oviposition._--The female selects an oötheca by feeling with her antennae. Flock stated, without citing experimental evidence, that the age of the egg case was apparently the chief factor determining choice. The wasp took 15-45 minutes to oviposit. Three females oviposited simultaneously into a single oötheca; a single female repeatedly oviposited into one oötheca at intervals. _Development._--Completed in an average of 32.6 days at a constant temperature of 82° F.

_Number of parasites per oötheca._--Average about 10.7 (range 4-16) (Flock, 1941); 15 (Ferrière, 1935).

_Sex ratio._--4 [F][F]:1 [M] (Ferrière, 1935); average of 6 [F][F]:1 [M] (Flock, 1941). Parthenogenesis occurs; the unfertilized eggs produced only males (Flock, 1941).

_Distribution._--U.S.A.: Maryland, south to Florida, west to Illinois, Kansas, and Arizona. Guatemala; Hawaii; India; Egypt; Sudan. (Burks, personal communication, 1956).

=Eupelmus atriflagellum= Girault

_Natural host._--_Blattella germanica_, Australia, Queensland (Girault, 1924).

=Eupelmus= sp.

_Natural host._--"Tree cockroach," U.S.A., Florida (Howard, 1892).

=Solindenia picticornis= Cameron

_Natural hosts._--_Allacta similis_, Hawaii (Perkins, 1906, 1913; Timberlake, 1924; Swezey, 1929; Zimmerman, 1948).

Other species of cockroaches, Hawaii (Perkins, 1913).

Family PTEROMALIDAE

=Pteromalus= sp.?

_Natural host._--_Leucophaea maderae_?, Jamaica (Westwood, 1839; Sells, 1842). [This host is undoubtedly an error. Sells stated that the oötheca which contained 96 unidentified chalcids had 16 dentations at the edge; the description fits the oötheca of an oviparous cockroach and not that of _L. maderae_ (see Roth and Willis, 1954). Westwood (1839, footnote p. 423) stated that at the meeting of the Entomological Society in 1838 Mr. Sells exhibited 94 specimens of a small _Pteromalus_ (apparently identified by Westwood) obtained from one cockroach oötheca. This same record of Sells was published posthumously in 1842, although in this paper he identified the host oötheca as "_Blaberus_" _maderae_. Cameron (1955) lists a European record of _Pteromalus_ sp. from _Periplaneta americana_ citing Girault (1914) as the source of the record. Girault's record was apparently taken from Westwood's footnote mentioned above.]

=Systellogaster ovivora= Gahan

_Natural hosts._--_Blatta orientalis_, U.S.A., Illinois (Gahan, 1917).

_Parcoblatta pensylvanica_, Canada, Ontario (Judd, 1955).

_Parcoblatta_ sp., U.S.A., Ohio (Edmunds, 1952a, 1953a).

"Blattid," U.S.A., Maryland (Gahan, 1917).

One oötheca of _P. pensylvanica_ yielded 14 parasites with a sex ratio of 2.5 [F][F]: 1[M] (Judd, 1955). The average number of parasites in 11 oöthecae of _Parcoblatta_ sp. collected in 1950-51 was 27 wasps (Edmunds, 1952a, 1953a). The adults made two to three emergence holes in the oötheca (Edmunds, 1953a; Judd, 1955).

Family EULOPHIDAE

=Melittobia chalybii= Ashmead

_Natural host._--_Periplaneta americana_, U.S.A., Missouri (Rau, 1940a): _M. chalybii_ is normally a parasite of Coleoptera and Hymenoptera (Peck, 1951). This is the only record from cockroach eggs. Burks (personal communication, 1956) stated that this species will attack any insect to which it is exposed and can be a serious pest in insect cultures of practically any insect order. In nature it seems to prefer the nests of aculeate Hymenoptera; Rau suggested that the parasites were probably brought into his laboratory with mud nests of _Sceliphron caementarium_ (Drury).

=Mestocharomyia oophaga= Dodd

_Natural host._--_Ellipsidion australe_, Australia, Queensland (Dodd, 1917).

=Syntomosphyrum blattae= Burks

_Natural hosts._--_Parcoblatta_ sp., U.S.A., Ohio (Burks, 1952; Edmunds, 1952a, 1953a): Ten oöthecae yielded an average of 92 wasps (Edmunds, 1952a). Five oöthecae, collected a year later, yielded an average of 74 wasps; adults sometimes made two to three exit holes in the oötheca (Edmunds, 1953a).

Cockroach, U.S.A., West Virginia (Burks, 1952).

=Syntomosphyrum ischnopterae= (Girault)

_Synonymy._--_Epomphaloides ischnopterae_ Girault [Peck, 1951].

Parker and Thompson (1928) called their hyperparasite _Tetrastichus_ sp. However, Dr. B. D. Burks (personal communication, 1955) has examined the teneral specimens which Parker and Thompson deposited in the U.S. National Museum; he stated that the species is apparently _Syntomosphyrum ischnopterae_. In view of the experimental work by Parker and Thompson (see below), this wasp may prove to be a hyperparasite on evaniids in cockroach oöthecae rather than a primary parasite on cockroach eggs. (See _Zeuxevania splendidula_, p. 243.)

_Natural hosts._--_Ischnoptera_ sp. [probably _Parcoblatta_ sp. (Rehn, personal communication, 1958)]. U.S.A., Maryland (Girault, 1917).

_Zeuxevania splendidula_ Costa (an evaniid in the oöthecae of _Loboptera decipiens_), France (Parker, 1924; Parker and Thompson, 1928).

The following information is from Parker and Thompson (1928): _Adult behavior._--Courtship and mating were accomplished as soon as adults emerged, and in a manner similar to that in other chalcids. The females oviposited only into oöthecae that were parasitized by _Zeuxevania_, never into normal, nonparasitized oöthecae. _Oviposition._--Oviposition occurred two days after mating. The female wasp stroked the oötheca with her antennae, selected a site, and bored into the oötheca with her ovipositor. She inserted the ovipositor deeply and oviposited for 10-30 minutes. The eggs were deposited randomly on the evaniid larva, some upright and others lying down. _Development._--Eggs of the hyperparasite hatched within 3 days and the larvae commenced feeding on the host larva. There were 30 and 50 hyper-parasites in two oöthecae. _Sex ratio._--5 [F][F]:1 [M] (from 3 oöthecae).

_Distribution._--U.S.A., District of Columbia, Maryland (Burks, 1952).

=Tetrastichus australasiae= Gahan

_Natural host._--_Periplaneta australasiae_, Sumatra (Gahan, 1923).

=Tetrastichus hagenowii= (Ratzeburg)