WASPS THAT PROVISION THEIR NESTS WITH COCKROACHES
Family POMPILIDAE
Pompilus bracatus Bingham
Natural hosts.—Cockroaches, India (Bingham, 1900).
Pompilus sp.
Natural host.—Cockroach, Nyasaland (Lamborn in Poulton, 1926): The wasp was collected leading a nymph of the cockroach by its antenna. The cockroach was in a stupefied state, and its antennae were bitten off to about half their length.
Salius verticalis Smith
Natural hosts.—Cockroaches, India (Bingham, 1900).
Family AMPULICIDAE
The species of Ampulex do not appear to make special nests in which to lay their eggs but drag their prey to any convenient hole, or crack in the ground (Arnold, 1928). Although many species of Ampulex have been described, the prey of only a small number of species have been discovered, but the known prey are all cockroaches.
Ampulex amoena Stål
Synonymy.—Ampulex novarae Saussure [Krombein, personal communication, 1957].
Natural hosts.—Periplaneta americana and Periplaneta australasiae, both as small nymphs, Formosa (Sonan, 1924, 1927): The wasp stings a nymph about one inch long and carries it to a suitable place, such as bamboo pipes, folds of newspaper, or books (in houses), for oviposition.
Periplaneta picea, Japan (Kamo, 1957; Kohriba, 1957).
Experimental hosts.—Periplaneta picea, Japan (Kamo, 1957; Kohriba, 1957).
Kamo (1957) observed that in the field both males and females sucked juices from wounds they made in the stems of Clerodendron trichotomum Thunberg or Ilex rotunda Thunberg. Kohriba (1957), on the other hand, found both sexes sucking sap of Abies sp. and other trees from points injured by the rostrum of cicadas. Kamo (1957) observed that the female wasp grasped the cockroach by a tergum and stung it several times in the thorax. The wasp always amputated the antennae of the prey and sucked up the fluid oozing from the cut antennae. The wasp egg was placed on the mesocoxa of the cockroach. In the laboratory as many as three cockroaches, each with a wasp egg, were stored in artificial nests per day. Kohriba (1957) observed similar behavior in the laboratory and made these additional notes. The paralyzed cockroach could move its legs and was led to the nest by the wasp which seized its antennae. The egg hatched in 2 days, and after sucking up body fluid for 2 days the larva began to devour the prey. Three days later the larva spun its cocoon, and about one month after spinning a female wasp emerged.
Ampulex assimilis Kohl
Natural hosts.—Blatta lateralis, wingless females, Iraq (Hingston, 1925): Nesting sites are holes in palm trees, galleries of beetles, or tunnels in ground. The wasp first seizes a cockroach by the edge of its thorax and stings it in the thoracic region, then seizes the cockroach by an antenna and pulls and leads it to the nest. The wasp deposits her egg on the outer surface of the femur of the cockroach's midleg. The nest is closed with debris; later the cockroach recovers from the sting. The wasp larva first feeds externally, then bores into the cockroach and devours the internal organs. Pupation occurs inside the exoskeleton of the cockroach.
Ampulex canaliculata (Say)
Synonymy.—Rhinopsis caniculatus.
Natural hosts.—Ischnoptera sp., U.S.A. (Krombein, 1951).
Lobopterella dimidiatipes, Hawaii (Williams, 1928a, 1929).
Parcoblatta pensylvanica? MacNay (1954) referred to a rare sphecoid wasp in eastern Canada which provisioned its nest with nymphs and adults of P. pensylvanica. Dr. W. R. M. Mason (personal communication, 1957) wrote us that although this wasp was Ampulex canaliculata, it was not reared from the cockroach but was swept from a pine tree. There are no positive records linking A. canaliculata with P. pensylvanica.
Experimental host.—Parcoblatta virginica, females, U.S.A., Missouri (Williams, 1928a, 1929): figure 6.
Nesting sites are in twigs (Krombein, 1951). The adult behavior is similar to that of A. compressa; the female wasp imbibes blood that oozes from the amputated antennae of the cockroach; the egg hatches in 2-3 days, and the development of one male was completed in 33 days (Williams, 1929).
Fig. 6. Ampulex canaliculata attacking Parcoblatta virginica. A, Female wasp stinging her prey, c. X 4.8. B, Wasp's egg attached to the coxa of the mesothoracic leg of the cockroach. C, Larva of A. canaliculata (about three-quarters grown) feeding on the internal organs of the host from the exterior, c. X 4. (Reproduced from F. X. Williams [1929], through the courtesy of Dr. F. X. Williams and F. A. Bianchi, Hawaiian Sugar Planters' Association.)
Distribution.—U.S.A.: Connecticut south to Georgia; Ohio, Wisconsin, Missouri, Kansas; in open woods (Krombein, 1951).
Ampulex compressa (Fabricius)
(Pl. 35)
Synonymy.—Guĕpe ichneumon of Réaumur [Williams, 1929]; Chlorion (Ampulex) compressum.
Natural hosts.—Periplaneta americana, New Caledonia (Lucas, 1879); India (Dutt, 1912); Reunion (Bordage, 1912).
Periplaneta australasiae, Hawaii (Swezey, 1944).
Periplaneta sp., India (Maxwell-Lefroy, 1909).
Cockroach. Mauritius (Réaumur, 1742); Burma (Bingham, 1897).
Experimental hosts.—Neostylopyga rhombifolia, Periplaneta americana, and Periplaneta australasiae, Hawaii (Williams, 1942, 1942a). Zimmerman's (1948) listings probably were taken from Williams.
Nesting sites.—Holes in walls; holes in banyan and fig trees; in houses in drawers and cartons. Behavior.—Similar to that of A. assimilis. Bordage (1912) gives a complete description of capture of prey. The female wasp cuts off part of the cockroach's antennae, legs, and wings; she sticks her egg onto the host's mesothoracic coxa. The wasp frequents houses in search of prey. Five ♀♀, 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 fasciata Jurine
Natural host.—Ectobius pallidus, France (Picard, 1911, 1919): Nesting sites are in brier or bramble stems, or in crevices in fig trees; the female possibly uses old nests of leaf-cutter bees. The feeding of the wasp larva is similar to that of other Ampulex. Adult wasp emerges by cutting open a passage through its cocoon and through the anus of the cockroach.
Ampulex ruficornis (Cameron)
Natural hosts.—Cockroaches, Oriental region (Rothney in Sharp, 1899): Nesting sites are in crevices in bark. The female grasps the cockroach by an antenna to drag it to her nest.
Ampulex sibirica Fabricius
Synonymy.—Perkins referred to this species as Ampulex sibirica. Williams (1942a), referring to Perkins's observations, mentions the species as "A. compressiventris Guérin (=A. siberica Sauss.)." Krombein (personal communication, 1956) has commented upon this synonymy as follows: Ampulex siberica Sauss. is apparently a misidentification by Saussure of sibirica Fab. Kohl (1893) in his revision of the genus Ampulex considered A. compressiventris Guérin to be the correct name for this common African species and that sibirica, described from Siberia, must be another species. However, Turner (1912) stated that he had seen Fabricius's type specimen and that it was identical with what had been called compressiventris; he considered the Siberian locality given by Fabricius as an error. Krombein suggested that Williams's use of the combination siberica Sauss. was a lapsus and that the valid name, if Turner is correct, is sibirica Fab.
Natural hosts.—Cockroaches, West Africa (Perkins in Sharp, 1899): Nesting sites are keyholes. Enters apartments in search of cockroaches. Wasp cocoon protrudes from dead body of cockroach.
Ampulex sonnerati Kohl
Synonymy.—"La mouche bleue" of Sonnerat (Kohl, 1893).
Natural host.—"Kakkerlac," Philippine Islands (Sonnerat, 1776): Nesting sites are readymade crevices. The wasp seizes the cockroach by an antenna and stings the host many times in the "abdomen." She drags the cockroach by an antenna to the nest, and, after depositing her egg, plugs the opening with moistened earth.
Dolichurus bicolor Lepeletier
Synonymy.—Schulz (1912) considered this to be Dolichurus corniculus. Berland (1925) stated that this is possibly a color variety of D. corniculus. Soyer (1947), from a study of the behavior of the wasps, believed that both D. bicolor and D. haemorrhous are varieties of D. corniculus. Krombein (personal communication, 1956) stated that D. corniculus and D. bicolor differ in characters other than color alone and that D. bicolor is considered a valid species today.
Natural host.—Cockroach, France (Benoist, 1927): The wasp was observed closing the entrance to its burrow. Its egg was attached to the coxa of the midleg of the cockroach.
Maneval (1932) stated that D. bicolor is found at the edge of dry woods along with D. corniculus and that the wasp will also accept the prey of D. corniculus if presented to it.
Dolichurus corniculus (Spinola)
Synonymy.—Dolichurus haemorrhous Costa [Schulz, 1912]. Berland (1925) listed D. haemorrhous separately but stated that it is perhaps a color variety of D. corniculus.
Natural hosts.—Blattella germanica, France (Benoist, 1927).
Ectobius lapponicus, Germany (Sickmann, 1893); Denmark (Nielsen, 1903); Sweden (Adlerz, 1903); Italy (Grandi, 1931, 1954); France (Benoist, 1927; Maneval, 1928).
Ectobius pallidus, France (Maneval, 1932; Soyer, 1947).
Ectobius panzeri, France (Soyer, 1947).
Ectobius sp., Italy (Grandi, 1954).
Hololampra punctata, Pitten (Handlirsch, 1889).
Loboptera decipiens, France (Ferton, 1894).
Cockroach, Netherlands (Bouwman, 1914).
Nesting sites.—The wasp uses already-made cavities such as rotting dead branches on ground, fissures in the earth, abandoned ant holes, chinks in stone, or the empty cocoon of the ichneumon Ophion luteus (Ferton, 1894; Maneval, 1932).
Behavior.—The prey is immobile while being dragged to the nest but recovers sufficiently from the sting so that if dug up it will run around (Ferton, 1894; Bouwman, 1914; Benoist, 1927; Grandi, 1954). The wasp cuts off about two-thirds of the cockroach's antennae prior to putting its prey in its nest (Adlerz, 1903; Bouwman, 1914; Soyer, 1947). One cockroach is placed in the nest and the wasp's egg is attached to the midcoxa (Ferton, 1894). Oviposition takes 5 to 6 minutes (Maneval, 1939). Wasp fills and seals its nest with bits of earth and stones (Ferton, 1894; Grandi, 1954). The wasp larva feeds externally and devours the entire cockroach, including its exoskeleton (Ferton, 1894).
Development.—Hatching occurs in 3 to 4 days (Ferton, 1894) or longer during cooler weather (Maneval, 1939). Larval development takes 6 days (Grandi, 1954), 8 days (Ferton, 1894), or 10 to 25 days depending on season (Maneval, 1939).
Dolichurus gilberti Turner
Natural hosts.—"Small Blattidae," India (Turner, 1917).
Dolichurus greenei Rohwer
Natural host.—Parcoblatta sp., U.S.A., Virginia (Krombein, 1951, 1955): Nesting sites are under leaf litter. The prey was a paralyzed third-instar nymph. Distribution.—Ontario. U.S.A. from Canadian border south to Florida in coastal States (Krombein, 1951).
Dolichurus ignitus Sm.
Natural hosts.—Cockroaches, Natal and Southern Rhodesia (Arnold, 1928): The wasp is "usually seen running up and down the trunks of trees searching for small cockroaches in the crevices of the bark."
Dolichurus stantoni (Ashmead)
Natural hosts.—Allacta similis, nymphs, Hawaii (Williams et al., 1931; Zimmerman, 1948).
Blattella lituricollis, usually nymphs, Philippine Islands, Hawaii (Williams, 1919).
Cutilia soror, nymphs, Hawaii (Williams et al., 1931; Zimmerman, 1948).
"Phyllodromia" sp., Philippine Islands, Hawaii (Williams, 1918; Bridwell, 1920).
Experimental hosts.—"Field cockroaches," Philippine Islands (Williams, 1944).
Nesting site.—Readymade crevices or holes in ground; porosity in lava. Behavior.—The wasp seizes the cockroach by a cercus or leg and stings it in the thorax. She (fig. 7, A) then drags the cockroach to the nest by the base of an antenna. Wasp bites off distal part of host's antennae. She deposits her egg on one of the host's midcoxae. Nest is plugged with lumps of soil. The larva eats the entire host. Development.—Eggs hatched in about a day and a half. Adults emerged about 3 weeks later. About five generations per year. (Williams, 1918, 1919; Williams et al., 1931.)
Dolichurus sp.
Natural hosts.—Cockroaches, nymphs, South Africa (Bridwell, 1917). Adult female cockroach carrying an oötheca, France (Deleurance, 1943).
Nesting site.—Plant stem, or in ground possibly an old abandoned nest of Ammophile. Behavior.—Bridwell noted that one wasp larva ate two cockroach nymphs before pupating; the adult emerged about 4 months after cocoon formation. Deleurance observed the wasp close its nest with small pebbles, balls of earth, and small dead branches. The wasp egg was placed on the femur of the midleg. The prey in the nest is alert when disturbed. Deleurance believed the wasp was a variety of D. corniculus.
Trirhogma caerulea Westwood
Natural hosts.—Periplaneta americana and Periplaneta australasiae, Formosa (Sonan, 1924): The wasp stings a nymph about one inch long and carries it to a suitable place (bamboo pipe) for oviposition.
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.)
Trirhogma sp.
Natural hosts.—Cockroaches, Oriental region (Williams, 1918, 1928): As far as is known species of this genus of wasps hunt cockroaches.
Family SPHECIDAE
Tachysphex blatticidus Williams
Natural hosts.—Chorisoneura sp., adults, Trinidad, St. Augustine (Callan, 1942): The wasps nest gregariously in sandy places. The wasp itself is parasitized by the mutillid Timulla (Timulla) eriphyla Mickel.
Cockroaches, Trinidad (Williams, 1941a; Callan, 1950).
Tachysphex coriaceus Costa
Natural hosts.—Cockroaches, Italy (Beaumont, 1954).
Tachysphex fanuiensis Cheesman
Natural hosts.—Graptoblatta notulata, Society Islands (Cheesman, 1927, 1928).
Cockroach ("except for its smaller size [it] much resembles Graptoblatta notulata."), New Caledonia (Williams, 1945).
Nesting sites.—Patches of dry soil (Cheesman, 1928); coarse sand at base of a bank (Williams, 1945). Behavior.—The female wasp pounces on the cockroach and stings it into immobility; she carries her prey in flight to the nest. Two to 13 cockroaches may be found in one nest; and one or more wasp eggs may be deposited in one nest. The egg is attached at one end to the host's thorax behind a forecoxa. Nest is sealed with dry pellets of soil. The cockroaches apparently do not recover from the wasp's sting.
Tachysphex lativalvis (Thomson)
Natural hosts.—Ectobius lapponicus, adults, Sweden (Adlerz, 1906); France (Maneval, 1932).
Ectobius pallidus, nymphs, France (Ferton, 1894, 1901; Maneval, 1932; Deleurance, 1946); Italy (Grandi, 1928).
Ectobius panzeri, Netherlands (Bouwman, 1914).
Ectobius sp., Denmark (Nielsen, 1933).
Ferton (1914) stated that he had reported in 1912 that this species hunted Hemiptera, but that this observation was a lapsus. Nesting site.—In the ground of sandy woodlot or border of dry woods; the nest is a hole 5.5 to 8 cm. long ending in a horizontal cell. Grandi (1928) stated that the entrance to the nest descended obliquely for 5 to 6 cm. and ended 4 cm. below the surface of the ground. Behavior.—Two cockroaches, either sex, adults or nymphs, were stored in the cell (Adlerz, 1903; Grandi, 1928). The wasp laid her egg on the first prey brought, attaching it behind the front coxa. The cockroaches were not excitable and their antennae had not been injured. Grandi (1928) stated that the claws of the hind tarsi of the victims may be amputated. The hatched larva may consume one of its victims in four days leaving only the head, pronotum, tegmina, wings, and the urosternum.
Podium abdominale (Perty)
Synonymy.—Trigonopsis abdominalis Perty [Kohl, 1902].
Natural hosts.—Cockroaches, nymphs, Ecuador (Williams, 1928): These wasps are apparently mainly arboreal mud daubers. The female wasp constructs a mud nest on underside of a palm leaf. Wasp egg is attached behind one of the forecoxae of the cockroach. Several cockroaches are stored in each nest. The prey is not immobilized as a result of the sting, and its antennae are left intact.
Podium carolina Rohwer
Natural host.—Parcoblatta pensylvanica, nymphs (Rau, 1937): Nesting sites are mud nests of Sceliphron caementarium (Drury). One to three cockroach nymphs are stored per nest; mud partitions are placed in tube; the nest is plugged with mud which is coated with resin. Distribution.—U.S.A., New York to North Carolina (Murray, 1951); Florida (Krombein and Evans, 1955).
Podium dubium Taschenberg
Natural hosts.—Epilamprine cockroaches, Brazil (Williams, 1928): Burrows, lenticular in cross section, are found on shaded trails. The wasp's habits are similar to those of P. flavipenne and P. haematogastrum.
Podium flavipenne Lepeletier
Natural host.—Epilampra abdomen-nigrum, British Guiana (Williams, 1928): Nesting site.—Burrows, about 2 inches deep and lenticular in cross section, are dug in the ground in well-drained, partly sheltered areas; also old Podium nests are used. Behavior.—The wasp stings the cockroach to helplessness and flies with it back to her nest where the host may recover from the sting; one or more cockroaches are stored per nest; the egg is deposited behind the forecoxa while the cockroach is still outside the burrow. The nest is sealed with mud. The larva feeds on most of the cockroach and leaves only some heavily sclerotized portions in the cell. In 153 nests examined, there was an average of 2.2 ± 0.08 [standard error computed from cited data] cockroaches per cell; four nests contained five cockroaches apiece. Of the 331 cockroaches in the nests, only 6 percent were adults. Development.—Egg hatches in about 2 days; larva feeds about 4 days and pupates about 2 weeks later; adult emerges about 10-12 days later.
Podium haematogastrum Spinola
Natural host.—Epilampra sp., Brazil, Pará (Williams, 1928): The female wasp (fig. 7, B) burrows into the surface of termite mounds, in banks, and in level ground. This wasp's behavior is similar to that of P. flavipenne. There was an average of 1.6 cockroaches (fig. 7, C) per cell in 74 nests examined. Of the 121 cockroaches collected, 28 percent were adults. Under artificial conditions, the life cycle varied from about a month to 45 days or more.
Podium luctuosum Smith
Natural host.—Parcoblatta virginica, female, U.S.A., New York (Pate, 1949).
Distribution.—U.S.A.: New York to Texas (Murray, 1951).
Podium rufipes Fabricius
Natural hosts.—"Wood roaches," British Guiana (Howes, 1917, 1919); Brazil (Williams, 1928): Nesting sites were clay column nests on houses, sides of stumps, or forest trees; banks; termite mound. Variable numbers of cockroaches were placed in the nests with one wasp egg attached behind forecoxa of the last host. The egg hatches in 2 days, the larva pupates about 2 weeks later, and the adult emerges 24 days later.
Podium sp.
Natural host.—Epilampra conferta, Brazil (Poulton, 1917): The burrow contained several cockroaches of the same species.