The classification follows Brues et al. (1954) with the following exceptions. The Acarina are arranged according to Dr. J. H. Camin (personal communication, 1955). Family Eupelmidae of the Hymenoptera follows the classification of Peck (1951).
Class ARACHNIDA
In this class, representatives of at least four orders have utilized cockroaches as food: the whip scorpions, scorpions, spiders, and mites. Apparently none of these feed exclusively on cockroaches, but the Philippine forest scorpion Heterometrus (=Palamnaeus) longemanus seems to prefer blattids to other insects (Schultze, 1927).
Order PEDIPALPIDA
Family THELYPHONIDAE
Mastigoproctus giganteus(Lucas)
Synonymy.—Thelyphonus giganteus Lucas [Dr. R. E. Crabill, personal communication, 1958].
Experimental prey.—Cockroaches, U.S.A. (Marx, 1892, 1894): Immature whip scorpion captured and fed on one or two cockroaches a week. It lived on this diet for about two years.
Mastigoproctussp.
Common name.—Whip scorpion.
Experimental prey.—Blattella germanica, U.S.A., Florida [Dr. B. J. Kaston, personal communication, 1953].
Order SCORPIONIDA
Pocock (1893) noticed that a scorpion whose pectines had come in contact with a cockroach immediately turned back and ate the insect. He concluded that the scorpion detected the cockroach by means of the pectines. However, Cloudsley-Thompson (1955) has demonstrated that the main function of the pectines is probably the detection of ground vibrations. He accounted for Pocock's observation by the presence of sensory spines (presumably tactile) which project from beneath the pectines. In a house in Arizona, Lyon (1951) observed over 60 scorpions living in a kitchen cabinet that enclosed a sink. They were apparently thriving on a heavy infestation of cockroaches. Stahnke (1953) stated that he used Periplaneta americana as the principal food for scorpions at the Poisonous Animals Research Laboratory of Arizona State College. Cloudsley-Thompson (1955a) cited cockroaches as one of the arthropods that scorpions feed upon.
Family BUTHIDAE
Buthus australis(Linnaeus)
Synonymy.—Androctonus australis [Crabill, personal communication, 1957].
Experimental prey.—Cockroaches, England (Cloudsley-Thompson, 1955a): This African species ate at least one cockroach per week during the summer months. It can, however, survive four months' starvation and is particularly adapted to a dry climate (Cloudsley-Thompson, personal communication, 1956).
Centruroides gracilis(Latreille)
Experimental prey.—Periplaneta americana, U.S.A. (Roth, unpublished data, 1953): Scorpion collected in Florida by Roth and identified by Dr. M. H. Muma.
Centruroides hentzi(Banks)
Experimental prey.—Periplaneta australasiae and Pycnoscelus surinamensis, U.S.A. (Muma, personal communication, 1953): This scorpion occurs in large numbers in the Florida citrus groves, together with P. australasiae which is probably an important natural prey.
Centruroides vitattus(Say)
Natural prey.—Parcoblatta pensylvanica (?), U.S.A., Florida (Muma, personal communication, 1953). This may have been another species of this genus, possibly P. divisa, as P. pensylvanica is not known from Florida (Rehn, personal communication, 1958).
Experimental prey.—Blatta orientalis, Blattella germanica, Periplaneta americana, and Pycnoscelus surinamensis, U.S.A., Florida (Muma, personal communication, 1953).
Parabuthus capensis(Hemprich and Ehrenberg)
Experimental prey.—"Common house-cockroach" (Pocock, 1893): The scorpions were collected in Cape Town, South Africa.
Family CHACTIDAE
Euscorpius germanus(Koch)
Synonymy.—Euscorpius carpathicus [Cloudsley-Thompson, 1955a].
Experimental prey.—Blattella germanica, England? (Pocock, 1893).
Periplaneta americana, nymphs, England? (Cloudsley-Thompson, personal communication, 1956): This scorpion is found naturally in southern Europe and North Africa.
Euscorpius italicus italicus(Herbst)
Experimental prey.—Cockroaches including nymphs of Periplaneta, England? (Cloudsley-Thompson, 1951): The cockroaches had to be disabled before the scorpion would feed on them. Prey is apparently detected by tactile and auditory senses, sight being poorly developed and not used. The scorpion is found in southern Europe and North Africa.
Family VEJOVIDAE
Hadrurus arizonensisEwing
Experimental prey.—Periplaneta americana, U.S.A. (Stahnke, 1949): This record is a photograph showing the scorpion eating the cockroach.
Family ISCHNURIDAE
Hormurus caudicula(Koch)
Experimental prey.—Cockroach, Australia (McKeown, 1952): This record is a photograph showing the scorpion feeding on a cockroach.
Family SCORPIONIDAE
Heterometrus longimanus(Herbst)
Synonymy.—Palamnaeus longimanus [Cloudsley-Thompson, personal communication, 1956].
Natural prey.—"Large wood cockroach," Philippine Islands (Schultze, 1927): On several occasions Schultze found fragments of wings and legs of the large wood cockroach in a scorpion cavity, under a rotten log.
Experimental prey.—Leucophaea maderae, Periplaneta americana, and other species of Blattidae, Philippine Islands (Schultze, 1927): Blattids seemed to be the favored food. This scorpion is usually found in humid, damp places in forest and jungle. Schultze describes in detail feeding behavior of the scorpion and method of capturing its prey.
Urodacus novaehollandiaePeters
Experimental prey.—Periplaneta americana, Australia (Glauert, 1946): An injured cockroach was accepted at once by the scorpion, which held the insect in its claws and tore it with the alternately moving chelicerae. The scorpion ate all the soft parts and most of the sclerotized exoskeleton.
Order ARANEIDA
Many observations of spiders feeding on cockroaches are quite general, and many observers have failed to identify either the spider or its prey. Belt (1874) stated that "the cockroaches that infest houses in the tropics ... have numerous enemies—birds, rats, scorpions, and spiders." When Belt tried to drive a cockroach toward a large cockroach-eating spider, the insect rushed away from him until it came within a foot of the spider when it would double back, never advancing nearer.
Beebe (1925) watched a giant "wood roach," which was in the grasp of a 2-inch ctenid spider, fly through the window of his British Guiana laboratory. While the spider ate the cockroach, the insect gave birth to 51 nymphs. Sonan (1924) reported that large gray spiders devour nymphs and adults of Periplaneta americana and P. australasiae in Formosa; this spider also occurs on Hiyakejima Island and Okinawa. Passmore (1936), who has produced some excellent photographs of tarantulas, stated that they destroy cockroaches. Rau (1940) stated that American and oriental cockroaches were the principal item of diet of a friend's pet tarantula for several years. Kaston (personal communication, 1953) successfully fed a tarantula with Periplaneta americana.
Bristowe (1941) found that the British species of Ectobius are readily accepted by Xysticus, Clubiona, Drassodes, Zelotes, Tarantula, and the web-builders Ciniflo and Aranea. The British domestic cockroaches were accepted by Tegenaria and Ciniflo, spiders large enough to overpower them, and were useful as food for tropical avicularids, ctenids, and sparassids in captivity.
Family THERAPHOSIDAE
Avicularia avicularia(Linnaeus) and Aviculariasp.?
Common name.—Bird-eating spider.
Natural prey.—Periplaneta americana, Trinidad (Main, 1924, 1930): The remains of the host were compressed into globular form by the spider after it had extracted the nutritive parts.
Phormictopus cancerides(Latreille)
Experimental prey.—Cockroach, West Indies (Wolcott, 1953).
Family SPARASSIDAE
Heteropoda venatoria(Linnaeus)
Synonymy.—Heteropoda regia Fabricius.
Common names.—Banana spider (Comstock, 1912); huntsman spider (Gertsch, 1949); big brown house spider (Bryan, 1915).
Natural and experimental prey.—Cockroaches, Bermuda (Verrill, 1902); Puerto Rico (Sein, 1923; Wolcott, 1924a; Petrunkevitch, 1930a); Hawaii (Bryan, 1915, Williams et al., 1931); British Guiana (Moore in Williams et al., 1931); Panama (Gertsch, 1949); New Zealand (adventive) (Parrott, 1952); England (Cloudsley-Thompson, 1953); Comstock (1912); Hawaii (Pemberton, 1917).
This (pl. 30, A) is a tropical species frequently imported into northern localities with bunches of bananas (Comstock, 1912; Cloudsley-Thompson, 1953). Adults measure 3 to 4 inches across with bodies over an inch long. They seldom leave their resting places during the day, but are active at night and search for food. The female does not spin a web (Bryan, 1915; Gertsch, 1949). The spider turns the cockroach over onto its back at the instant of seizure and holds it firmly against the substrate. The cockroach dies in 10 minutes and is gradually rolled up by the spider as it sucks out the nutriment (Moore in Williams et al., 1931). The spider does not attempt to bite when captured, but if it does, its bite is said to be painful but not dangerous (Cloudsley-Thompson, 1953). Zimmerman (1948) found scores of Periplaneta australasiae breeding in rock piles in Hawaii; also present were large numbers of these spiders and centipedes which presumably preyed upon the cockroaches.
Family THERIDIIDAE
Latrodectus indistinctusPickard-Cambridge
Common name.—Button spider.
Natural prey.—Karnyia discoidalis, South Africa, Western Cape Province (Hesse, 1942): The nest is constructed on the ground among grass stems or other vegetation. Preferred sites are slight hollows, hoof imprints, etc. Nests are roughly tubular. The remains of insects are entangled in the walls of the nest where they form dense accumulations. Predatory activities of the spider are limited to an area close to the tubular entrance to the nest and do not extend beyond the trapping strands near the entrance. Capture is dependent upon accidental contact of the insect with sticky threads surrounding the entrance. This spider apparently attacks any insect or arachnid that becomes entangled in the nest. In an examination of 40 nests, remains of 6 K. discoidalis were found.
Latrodectus mactans(Fabricius)
Common names.—Black widow, hourglass, or shoe-button spider.
Natural prey.—Cockroaches, Puerto Rico (Petrunkevitch, 1930). U.S.A., Florida, on shipboard (Anonymous, 1939): This is a presumptive host record, as the spiders were not reported as having been seen eating cockroaches; however, heavy infestations of both were found together.
Family LYCOSIDAE
Lycosa helluoWalckenaer
Experimental prey.—Young nymphs of Diploptera punctata, U.S.A. (Eisner, 1958): Larger nymphs and adults repelled the spider by ejecting a repellent secretion, which has been identified as a mixture of p-benzoquinone and its derivatives by Roth and Stay (1958).
Lycosasp.
Experimental prey.—Supella supellectilium, U.S.A. (Roth and Willis, unpublished data, 1953): The lycosid (pl. 30, B-E) was probably L. avida Walckenaer (tentatively identified by Dr. B. J. Kaston from a photograph).
Order ACARINA
Family PHYTOSEIIDAE
Blattisocius tineivorus(Oudemans)
Synonymy.—Blattisocius triodons Keegan [Baker and Wharton, 1952].
Natural host.—Blattella germanica, U.S.A. (Keegan, 1944): Three mites found on 238 cockroaches examined; others taken in debris from floor of cockroach cage (Keegan, 1944). Members of this family are predaceous (Baker and Wharton, 1952).
Family LAELAPTIDAE
Blattilaelaps nauphoetaeWomersley
Natural host.—Nauphoeta cinerea, Australia, Brisbane (Womersley, 1956).
Coleolaelaps (?)sp.
Natural host.—Gromphadorhina portentosa, the hosts were imported into U.S.A. from Madagascar via Europe (Roth and Willis, unpublished data, 1958): The mites (pl. 12, C) were tentatively determined by Dr. E. W. Baker.
Hypoaspissp.
Natural host.—Panesthia australis, imported into U.S.A. from Australia (Roth and Willis, unpublished data, 1955): Cockroach determined by J. A. G. Rehn. Generic determination of mite made by Dr. R. W. Strandtmann (Camin, personal communication, 1955).
Family UROPODIDAE
Uropodasp.
Natural host.—Blattella humbertiana, Formosa (Takahashi, 1940). Nymphs of the cockroach may be destroyed (Takahashi, 1940). Uropodids frequently attach themselves to insects, especially in nymphal stages but probably are harmless (Baker and Wharton, 1952).
Family DIPLOGYNIIDAE
Undetermined diplogyniid
Natural host.—Panesthia australis, imported into U.S.A. from Australia (Roth and Willis, unpublished data, 1955): Cockroach determined by J. A. G. Rehn. According to Dr. J. H. Camin (personal communication, 1955) this is a new genus and new species in the subfamily Diplogyniinae, and is most closely related to the genus Lobogynioides. Mites of this family live as ectoparasites and commensals on beetles and possibly other insects (Baker and Wharton, 1952).
Family ANOETIDAE
Histiostoma feroniarum(Dufour)
Natural host.—Pycnoscelus surinamensis, Germany (Roeser, 1940): Though not parasitic, the mites at times became so numerous that the insects were hindered in their movement, were unable to feed, and died. The mites were introduced with soil and leaves and had originally been attached to millipedes, waterfleas, and sowbugs.
The deutonymphs, hypopial forms, or travelers are found on insects; the other stages are found in decaying organic matter (Baker and Wharton, 1952).
Family ACARIDAE
Caloglyphus spinitarsus(Hermann)
Natural host.—Pycnoscelus surinamensis, Germany (Roeser, 1940): See notes following Histiostoma feroniarum above.
Caloglyphussp.
Natural hosts.—Blattella germanica and Periplaneta americana, U.S.A. (Piquett and Fales, 1952): Mite feeds on organic matter but can reduce the vigor of a cockroach colony.
Tyrophagus lintneri(Osborne)
Common name.—Mushroom mite.
Associate.—Pycnoscelus surinamensis, U.S.A. (Roth and Willis, unpublished data, 1953): Mite determined by Dr. E. W. Baker (personal communication, 1953). Although this mite was found on the cockroach, it is a known pest in stored foods (Baker and Wharton, 1952) and probably was brought into the culture with food. Rau (1924) reported that the food of Blatta orientalis often became infested with this species, but it did not affect the health or mortality of the cockroaches in his culture.
Tyrophagus noxiusA. Z.
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 tarsalusBanks
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
Chaetodactylussp.
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
Locustacarussp.
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 nanaPritchard
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 podapolipophagusTrÄgÅrdh
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??]."
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.
Pimeliaphilus podapolipophagus. Fig. 4.—The cockroach mite, Pimeliaphilus podapolipophagus. (From Baker et al. [1956]; 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 morsitansLinnaeus
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 subspinipesLeach
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.
Scolopendrasp.
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 strenuusHagen
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 religiosaLinnaeus
Common name.—European mantis.
Experimental prey.—Nauphoeta cinerea, and Periplaneta americana, U.S.A. (Rilling, personal communication, 1957).
Metallyticus semiaeneusWestwood
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 sinensisSaussure
Common name.—Chinese mantis.
Experimental prey.—Nauphoeta cinerea and Periplaneta americana, U.S.A. (Rilling, personal communication, 1957).
Family GRYLLACRIDIDAE
Diestrammena apicalisBr. v. Wattenwyl
and
Diestrammena japanicaBlatchley
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 apicicornisSignoret
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 domesticusPinto
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 arthurneivaiLent 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
Ranatrasp.
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
Megaseliasp.
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.
Stylogasterspp.
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 (?) venterisCurran
Hosts.—Periplaneta americana, Brazil (Souza Lopes, 1937): Obtained complete evolution of the parasite in this host. This may have been an experimental host.
Calodexiaspp.
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 basalisStein
Host.—Eggs of Parcoblatta sp., Ohio (Edmunds, 1952a).
Family SARCOPHAGIDAE
Sarcophaga omaniHall
Host.—Arenivaga bolliana, Texas (Wirth, personal communication, 1953): Specimens in U.S. National Museum.
Sarcophaga lambensWied.
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).
Sarcophagaspp.
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 pennsylvanicusDe 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 pacificusBoisduval
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 partinigerRiek
Natural host.—Cockroach (undescribed genus belonging to the Pseudomopinae), Australia Capital Territory (Riek, 1955).
Neorhipidius neoxenusRiek
Natural host.—Robshelfordia longiuscula or Robshelfordia circumducta, Australia Capital Territory (Riek, 1955).
Paranephrites xenusRiek
Natural host.—Oniscosoma granicollis, Australia Capital Territory (Riek, 1955).
Rhipidioides ableptusRiek
Natural host.—Balta patula, Australia, Victoria (Riek, 1955): Pupal stage lasted only 3 days.
Rhipidioides adynatusRiek
Natural host.—Escala sp. or an undescribed genus of Pseudomopinae, Australia, Victoria (Riek, 1955).
Rhipidioides fuscatusRiek
Natural host.—Ellipsidion affine, Australia, New South Wales (Riek, 1955).
Rhipidioides helenaeRiek
Natural host.—Robshelfordia longiuscula or Robshelfordia circumducta, Australia Capital Territory (Riek, 1955).
Rhipidioides mollisRiek
Natural host.—Robshelfordia longiuscula or Robshelfordia circumducta, Australia Capital Territory (Riek, 1955).
Rhipidioides rubricatusRiek
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 nitidioidesSelander
Synonymy.—Nephrites nitidus of Riek not Shuckard [Selander, 1957].
Natural host.—Platyzosteria sp., Tasmania (Riek, 1955).
Riekellasp.
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] boissyiAbeille
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 denisiChobaut
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 pectinicornisThunberg
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 scutellarisHeller
Natural hosts.—Blattidae, Philippine Islands (Schultze, 1925).
Family DERMESTIDAE
Dermestes aterDe 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.
Dermestessp.
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);[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.
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 dimidiataFabricius
Synonymy.—Evania abyssinica Westwood [Schletterer, 1889].
Natural host.—Blatta orientalis, Egypt? (Alfieri, 1914).
Evania subspinosaKieffer
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 dorsalisof 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 harpyoidesBradley
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 reticulataSay
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.)
Hyptiasp.
Natural host.—Cariblatta delicatula, Cuba (Hebard, 1916a); Parasite identified by Ashmead.
Hyptiasp. (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 splendidulaCosta
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 metallicaGirault
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 pulchraAshmead
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 alfieriiMercet
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 blattophagaGirault
Natural host.—Cockroach, Australia, Queensland (Girault, 1915).
Family EUPELMIDAE[7]
Anastatus blattidifuraxGirault
Natural host.—Cockroach, Australia, Queensland (Girault, 1915).
Anastatus floridanusRoth 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 ??:1 ? from ovipositions by isolated females. In the one oÖtheca collected in the field, the ratio was 21.6 ??:1 ?. Parthenogenesis exists; the unfertilized eggs produced only males.
Anastatus tenuipesBolÍvar 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 ??:1 ? (FerriÈre, 1935); average of 6 ??:1 ? (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 atriflagellumGirault
Natural host.—Blattella germanica, Australia, Queensland (Girault, 1924).
Eupelmussp.
Natural host.—"Tree cockroach," U.S.A., Florida (Howard, 1892).
Solindenia picticornisCameron
Natural hosts.—Allacta similis, Hawaii (Perkins, 1906, 1913; Timberlake, 1924; Swezey, 1929; Zimmerman, 1948).
Other species of cockroaches, Hawaii (Perkins, 1913).
Family PTEROMALIDAE
Pteromalussp.?
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 ovivoraGahan
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 ??: 1? (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 chalybiiAshmead
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 oophagaDodd
Natural host.—Ellipsidion australe, Australia, Queensland (Dodd, 1917).
Syntomosphyrum blattaeBurks
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 ??:1 ? (from 3 oÖthecae).
Distribution.—U.S.A., District of Columbia, Maryland (Burks, 1952).
Tetrastichus australasiaeGahan
Natural host.—Periplaneta australasiae, Sumatra (Gahan, 1923).
Tetrastichus hagenowii(Ratzeburg)
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 ??:1 ? (Usman, 1949); 4 ??:1 ? (Cameron, 1955); 2-8 ??:1 ? (Roth and Willis, 1954b); 1.2 ??:1 ? (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 periplanetaeCrawford
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).
Tetrastichussp. 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).
Tetrastichussp. 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 ??:1 ?.
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 bracatusBingham
Natural hosts.—Cockroaches, India (Bingham, 1900).
Pompilussp.
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 verticalisSmith
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 amoenaStÅl
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 assimilisKohl
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).
Distribution.—U.S.A.: Connecticut south to Georgia; Ohio, Wisconsin, Missouri, Kansas; in open woods (Krombein, 1951).
Ampulex compressa(Fabricius)
(Pl. 35)
Synonymy.—Guepe 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 ??, supplied with a cockroach per day, stored an average of 57±14 cockroaches; 8 ?? 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 ?? lived an average of 110±11 days (minimum 31, maximum 159); several ?? lived 2 months (Williams, 1942).
Ampulex fasciataJurine
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 sibiricaFabricius
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 sonneratiKohl
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 bicolorLepeletier
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 gilbertiTurner
Natural hosts.—"Small Blattidae," India (Turner, 1917).
Dolichurus greeneiRohwer
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 ignitusSm.
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.)
Dolichurussp.
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 caeruleaWestwood
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.
Cockroach-hunting wasps. Fig. 7.—Cockroach-hunting wasps. A, Dolichurus stantoni leading a nymph of Blattella lituricollis to her nest, c. X 4. (Reproduced from F. N. Williams [1919].) 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. X 1.6. C, Epilampra sp. parasitized by P. haematogastrum showing the wasp's egg attached to the right fore coxa, c. X 3.2. (B and C reproduced from Williams [1928], through the courtesy of Dr. F. X. Williams and F. A. Bianchi.)
Trirhogmasp.
Natural hosts.—Cockroaches, Oriental region (Williams, 1918, 1928): As far as is known species of this genus of wasps hunt cockroaches.
Family SPHECIDAE
Tachysphex blatticidusWilliams
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 coriaceusCosta
Natural hosts.—Cockroaches, Italy (Beaumont, 1954).
Tachysphex fanuiensisCheesman
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 carolinaRohwer
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 dubiumTaschenberg
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 flavipenneLepeletier
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 haematogastrumSpinola
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 luctuosumSmith
Natural host.—Parcoblatta virginica, female, U.S.A., New York (Pate, 1949).
Distribution.—U.S.A.: New York to Texas (Murray, 1951).
Podium rufipesFabricius
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.
Podiumsp.
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 piceaEmery
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) wilverthiEmery
Natural prey.—Small cockroaches, Belgian Congo (Raignier and van Boven, 1955).
Dorylussp.
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 humilisMayr
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.
