CONTROL OF MITES IN COCKROACH COLONIES
Fisk (1951) eliminated the mites [possibly Pimeliaphilus podapolipophagus (Baker et al., 1956)] in his cockroach colony by using a 5-percent spray and a 5-percent dust of p-chlorophenyl, p-chlorobenzene sulfonate. The exterior of the cockroach containers were sprayed with the solution and the interior, including the insects, were dusted. Within a month the mites had disappeared and the vigor of the cockroach colony improved. Piquett and Fales (1952) used flowers of sulfur and general sanitary procedures for eliminating the mites in laboratory colonies of Blatta orientalis; they cleaned the dishes every few days and applied grease around the edges of the containers to prevent new mite invasions. Qadri (1938) employed similar control measures.
Class CHILOPODA
Large centipedes which entered houses in India probably sought out cockroaches (Maxwell-Lefroy, 1909). In Puerto Rico, centipedes entered homes to which they were attracted by cockroaches (Seín, 1923). In Hawaii, centipedes preyed on insects generally but especially on cockroaches (Bryan, 1915). Sonan (1924) reported that in Formosa and Okinawa Islands a species of centipede 5 to 6 inches long comes into the houses and devours both adults and nymphs of Periplaneta americana or P. australasiae. Zimmerman (1948) found P. australasiae breeding by scores in rock piles in Hawaii accompanied by large numbers of Scolopendra and large spiders that probably preyed upon the cockroaches.
Order SCUTIGEROMORPHA
Family SCUTIGERIDAE
Scutigera coleoptrata (Linnaeus)
Synonymy.—Scutigera forceps Rafinesque [Crabill, 1952].
Common name.—House centipede.
Natural prey.—Cockroaches, U.S.A. (Felt, 1909; Back, 1947; Auerbach, 1951; Crabill, 1952; and others): This predator-prey relationship seems to be based on good circumstantial evidence (Crabill, personal communication, 1953).
Experimental prey.—Blattella germanica, newly hatched nymphs and adult female, U.S.A. (Snodgrass, 1930; Roth and Willis, unpublished data, 1953).
Periplaneta americana, U.S.A. (Roth and Willis, unpublished data, 1953).
Supella supellectilium, U.S.A. (Roth and Willis, unpublished data, 1953): See plate 31.
Our specimen caught a small American cockroach nymph that we placed in its jar. Before it had finished its meal, it caught and held two other nymphs with its legs while it continued to feed on the first. The body of this centipede reaches a maximum length of 27 mm. and it is usually found in basements, dark corners, or in spaces in the walls (Auerbach, 1951). Introduced from Europe, this species is now widespread in the United States (Crabill, 1952).
Allothereua maculata (Newport)
Synonymy.—Scutigera maculata [Crabill, personal communication, 1957].
Natural prey.—Cockroaches, Malay peninsula, Batu caves (Ridley in Annandale et al., 1913): This is a presumptive host record.
Order SCOLOPENDROMORPHA
Family SCOLOPENDRIDAE
Scolopendra cingulata (Latreille)
Experimental prey.—Cockroaches, England (Cloudsley-Thompson, 1955): After capture in France, this specimen was kept for four weeks without food. She was then fed medium-sized nymphal cockroaches of which she ate an average of about one per week throughout the summer. Adult cockroaches were attacked only after they had been disabled.
Scolopendra morsitans Linnaeus
Natural prey.—Cockroaches, Guadeloupe (Lherminier, 1837).
Experimental prey.—Cockroaches, India, Nagpur (Jangi, 1955): As soon as the centipede became aware of its prey, it rapidly embraced the cockroach within its legs and with its fangs gripped the insect's thorax. The predator continued to hold the prey with its fangs while its mouth parts prodded the victim's body. After feeding on an adult cockroach, the centipede is not inclined to kill another for 2-3 days.
Scolopendra subspinipes Leach
Natural prey.—Cockroaches, Hawaii (Williams et al., 1931): This is a common species with a body length of 6 or more inches. It is reported to be a great enemy of cockroaches.
Scolopendra sp.
Natural prey.—Ectobius panzeri, England (Lucas, 1911, 1920): When captured, the centipede was holding a live cockroach which it had apparently just caught. The insect was held beneath its captor's body, ventral surface upward, by several of the anterior legs while the centipede fed.
Class INSECTA
We have found representatives of only 10 orders that have preyed on or parasitized cockroaches: Beetles, flies, bugs, ants, wasps, stylops, and cockroaches occurred in nature; the others resulted from feeding cockroaches to captive insects or were laboratory observations.
Order ODONATA
Family AESHNIDAE
Anax strenuus Hagen
Common name.—Giant Hawaiian dragonfly.
Experimental prey.—Cockroaches, Hawaii (Williams, 1936): The dragonfly nymph was fed with medium large cockroaches and other insects.
Order BLATTARIA
In this chapter the relations of other arthropods to cockroaches are either as parasites or as predators. Certain cockroaches have turned the tables on their adversaries and become predators themselves. This aspect of cockroach behavior is discussed in chapter XVI. Other associations of cockroaches, as commensals with other insects and as associates of other cockroaches, are discussed in chapters XV and XVII.
Order ORTHOPTERA
Family MANTIDAE
Hierodula tenuidentata (Saussure) (?) (Serville)
(Pl. 32)
Experimental prey.—Blatta orientalis, Diploptera punctata, Eurycotis floridana, Leucophaea maderae, Nauphoeta cinerea, Neostylopgya rhombifolia, and Periplaneta americana, U.S.A. (Rilling, personal communication, 1957): Mrs. Rilling wrote us that with the exception of N. rhombifolia, all the above cockroaches were readily eaten. All the mantids initially rejected N. rhombifolia after grasping and making a brief attempt to chew the cockroaches. However, if specimens of N. rhombifolia were left in the jars with the mantids, the cockroaches were usually eaten within the next 24 hours. N. rhombifolia ejects an odorous substance when seized and the mantids probably ate these insects after most of this secretion had been depleted. It is highly probable that the secretion of N. rhombifolia may deter the mantid's attack, but it should be pointed out that, with the possible exception of N. cinerea, all the other species fed to these mantids give off odorous substances when seized or disturbed. Apparently, certain naturally repellent compounds will deter this mantid, whereas others that are presumed to be repellent will not; however, the nutritional state of the mantid is undoubtedly a factor which may limit the effectiveness of certain repellent secretions against this predator.
Byrsotria fumigata, teneral males, and Periplaneta australasiae, nymphs, U.S.A. (Roth and Willis, unpublished data, 1958).
Diploptera punctata, U.S.A. (Eisner, 1958).
Mantis religiosa Linnaeus
Common name.—European mantis.
Experimental prey.—Nauphoeta cinerea, and Periplaneta americana, U.S.A. (Rilling, personal communication, 1957).
Metallyticus semiaeneus Westwood
Experimental prey.—Cockroaches, Borneo (Shelford, 1916).
Sphodromantis viridus (Forskål)
Synonymy.—Sphodromantis bioculata Burmeister [Gurney, personal communication, 1958].
Experimental prey.—Blatta orientalis, Egypt (Adair, 1923): This species of cockroach was apparently used regularly as food for the mantid in the laboratory.
Stagmomantis carolina (Johansson)
Common name.—Carolina mantis.
Experimental prey.—Blattella germanica and Periplaneta americana, U.S.A. (Breland, 1941): The mantids were fed 1-2 German cockroaches daily. One female mantid consumed 10 adult German cockroaches plus one oötheca and part of another in 2.5 hours. An adult German cockroach was consumed in an average of 8.5 minutes (range 5.5-15 minutes).
Blatta orientalis, nymphs, and Diploptera punctata, U.S.A. (Roth and Willis, unpublished data, 1953).
Tarachodes maurus (Stal)
Experimental prey.—Cockroaches, South Africa (Faure, 1940).
Tenodera aridifolia sinensis Saussure
Common name.—Chinese mantis.
Experimental prey.—Nauphoeta cinerea and Periplaneta americana, U.S.A. (Rilling, personal communication, 1957).
Family GRYLLACRIDIDAE
Diestrammena apicalis Br. v. Wattenwyl
and
Diestrammena japanica Blatchley
Natural prey.—Cockroach eggs, Japan (Asano, 1937): These are questionable records. Asano found D. apicalis and D. japanica beneath his house near several empty cockroach oöthecae which appeared to have been eaten into. He assumed from the condition of the oöthecae and the proximity of the stone crickets that the insects had devoured the cockroach eggs.
Experimental prey.—Eggs of Blattella germanica and Periplaneta japanica, Japan (Asano, 1937): Seven eggs of B. germanica (obtained from an oötheca being carried by a female) and eggs of P. japanica (presumably in oöthecae) were fed to both species of stone crickets in the evening. The eggs were devoured by the next morning.
Order DERMAPTERA
Family FORFICULIDAE
Undetermined earwigs
Experimental prey.—Cockroaches, France (Chopard, 1938): According to Chopard, Brisout de Barneville in 1848 indicated that earwigs in captivity can be fed small cockroaches.
Order HEMIPTERA
Family LYGAEIDAE
Clerada apicicornis Signoret
Natural prey.—Cockroach, Hawaii (Illingworth, 1917): This predaceous bug is commonly found about buildings. Illingworth says that Kirkaldy suspected that it fed on small blattids and that Dr. Perkins saw it feeding on a dead cockroach.
Family REDUVIIDAE
Spiniger domesticus Pinto
Natural prey.—Periplaneta americana, Brazil, Matto Grosso (Pinto, 1927, 1927a): This bug preys principally on cockroaches and was observed infesting the walls of dwellings where it preyed on P. americana.
Triatoma arthurneivai Lent and Martins
Natural prey.—Monastria sp., Brazil, Minas Gerais (Martins, 1941): This bug probably feeds on cockroaches of this genus, as well as on rodents.
Undetermined reduviids
Natural prey.—Arenivaga roseni and Polyphaga saussurei, Turkmen S.S.R. (Vlasov and Miram, 1937): These desert cockroaches are found in burrows of rodents and desert turtles around Ashkhabad. Reduviids are their main enemies. Vlasov (1933) found nymphs of Reduvius christophi Jak. and R. fedtschenkianus Osch. in similar burrows in this same area, although he did not specifically cite them as enemies of the desert cockroaches.
Family NEPIDAE
Ranatra sp.
Experimental prey.—Cockroaches, U.S.A. (Hoffman, 1924).
Order NEUROPTERA
Family ASCALAPHIDAE
Undetermined larva
Experimental prey.—Blattella germanica, Kenya Colony (Someren, 1924).
Order DIPTERA
From the few observations that have come to our attention, it seems that flies are comparatively rare parasites in cockroaches.
Family PHORIDAE
Megaselia sp.
Host.—Eggs of Parcoblatta sp., Ohio (Edmunds, 1952a).
Family CONOPIDAE
Stylogaster stylata (Fabricius)
Hosts.—Cockroaches, Brazil (Souza Lopes, 1937): L. Travassos was quoted as having observed this species pursue cockroaches that were escaping columns of the army ant Eciton sp. Souza Lopes (1937) stated that the female deposits eggs on the cuticle of the host near the end of the body; the egg is barely inserted and two recurrent hooks prevent it from falling off. Souza Lopes (1937) also observed other species of Stylogaster pursue Orthoptera, but he was unable to devote proper attention to the behavior of the flies.
Stylogaster spp.
Hosts.—Chorisoneura sp., Brazil (Souza Lopes, 1937): An adult specimen was found in a museum collection with an egg of Stylogaster attached to the posterior end of its abdomen.
Cockroaches, Panama (C.W. Rettenmeyer, personal communication, 1959): "Seven species were collected hovering over army ant swarms and a few flies were seen apparently attacking cockroaches that had been flushed by the ants."
Family LARVAEVORIDAE
Calodexia (?) venteris Curran
Hosts.—Periplaneta americana, Brazil (Souza Lopes, 1937): Obtained complete evolution of the parasite in this host. This may have been an experimental host.
Calodexia spp.
Hosts.—Cockroaches, Panama (Rettenmeyer, personal communication, 1959): Swarms of army ants are accompanied by about 20 species of Calodexia. These flies larviposit on the cockroaches, crickets, and possibly other arthropods that are flushed from cover by the ants. Larvae were found in one(?) cockroach. Larvae from an adult of Calodexia were introduced experimentally into a cockroach and successfully reared.
Undetermined tachinids
Hosts.—Eurycotis floridana, from Florida (Roth, unpublished data, 1953): Three larvae (det. by W.W. Wirth) were found in a living adult male.
Panesthia australis, from Australia (Roth, unpublished data, 1957): Reared from a wild-caught cockroach that was maintained in a laboratory colony.
Cockroaches, Australia (E. F. Riek, personal communication, 1955): Reared from some of the larger species.
Family MUSCIDAE
Coenosia basalis Stein
Host.—Eggs of Parcoblatta sp., Ohio (Edmunds, 1952a).
Family SARCOPHAGIDAE
Sarcophaga omani Hall
Host.—Arenivaga bolliana, Texas (Wirth, personal communication, 1953): Specimens in U.S. National Museum.
Sarcophaga lambens Wied.
Synonymy.—Sarcophaga sternodontis (Towns.).
Hoffman (1927) claimed that approximately 40 percent of some specimens of Pycnoscelus surinamensis collected in southern Haiti were parasitized by S. lambens. However, according to entomologists at the University of Puerto Rico Agricultural Experiment Station, Hoffman was incorrect in his observations: S. lambens was never reared from a living insect and had been recovered only from dead cockroaches and other dead insects and was considered saprophytic rather than parasitic (Schwabe, 1950b).
Sarcophaga spp.
Sanjean (1957) reared various species of sarcophagid larvae on Periplaneta americana which were freshly killed or chopped up; first instar larvae were also introduced into the body cavity of cockroaches which had their heads and legs removed. Adult sarcophagids were collected and freshly killed American cockroaches used as bait.
Order COLEOPTERA
Family CARABIDAE
Harpalus pennsylvanicus De Geer
Experimental prey.—Cryptocercus punctulatus, U.S.A. (Cleveland et al., 1934): This beetle is often found in the galleries of C. punctulatus in nature. In the laboratory it killed and devoured cockroaches as large as itself.
Family DYTISCIDAE
Rhantus pacificus Boisduval
Experimental prey.—Cockroaches, disabled, Hawaii (Williams, 1936): This beetle, which is common in mountain streams, located wounded cockroaches in an aquarium by sense of smell or taste rather than sight.
Family LAMPYRIDAE
Undetermined larva
Experimental prey.—Parcoblatta virginica, adult female (pl. [33], C), U.S.A. (Roth and Willis, unpublished data, 1953).
Family RIPIPHORIDAE[5]
Neonephrites partiniger Riek
Natural host.—Cockroach (undescribed genus belonging to the Pseudomopinae), Australia Capital Territory (Riek, 1955).
Neorhipidius neoxenus Riek
Natural host.—Robshelfordia longiuscula or Robshelfordia circumducta, Australia Capital Territory (Riek, 1955).
Paranephrites xenus Riek
Natural host.—Oniscosoma granicollis, Australia Capital Territory (Riek, 1955).
Rhipidioides ableptus Riek
Natural host.—Balta patula, Australia, Victoria (Riek, 1955): Pupal stage lasted only 3 days.
Rhipidioides adynatus Riek
Natural host.—Escala sp. or an undescribed genus of Pseudomopinae, Australia, Victoria (Riek, 1955).
Rhipidioides fuscatus Riek
Natural host.—Ellipsidion affine, Australia, New South Wales (Riek, 1955).
Rhipidioides helenae Riek
Natural host.—Robshelfordia longiuscula or Robshelfordia circumducta, Australia Capital Territory (Riek, 1955).
Rhipidioides mollis Riek
Natural host.—Robshelfordia longiuscula or Robshelfordia circumducta, Australia Capital Territory (Riek, 1955).
Rhipidioides rubricatus Riek
Natural host.—Choristima sp. and Choristimodes sp., Australia Capital Territory (Riek, 1955).
Riekella australis (Riek)
Synonymy.—Nephrites australis Riek [Selander, 1957].
Natural host.—Cutilia sp., Australia Capital Territory (Riek, 1955): Two females emerged from one host.
Riekella nitidioides Selander
Synonymy.—Nephrites nitidus of Riek not Shuckard [Selander, 1957].
Natural host.—Platyzosteria sp., Tasmania (Riek, 1955).
Riekella sp.
Synonymy.—Nephrites sp. [Selander, 1957].
Natural host.—Platyzosteria castanea, Australia Capital Territory (Riek, 1955).
Biology of Australian Ripidiini.—The Australian species of Ripidiini are parasites of apparently endemic, ground-dwelling species of cockroaches. There is some correlation between host subfamily and parasite genus: Riekella spp. [= Nephrites] have only been bred from Blattinae. Rhipidioides spp. occur only in the closely related Ectobiinae and Pseudomopinae. Neonephrites and Neorhipidius also occur in the Pseudomopinae. Paranephrites occurs in the Panchlorinae. There is some evidence that the parasitized cockroaches migrate onto trees when the larval parasite is mature, as pupae have only been found on the trunks of eucalyptus trees. In all species the larva leaves the host dorsally through an intersegmental membrane. The host continues to live for a few days after the parasite emerges. The larva attaches itself to bark on the tree trunk by a few strands of silk before pupating. The larviform, wingless female remains near the pupal skin and is sought out by the winged male. The eggs are laid in a mass around the pupal skin (Riek, 1955).
Ripidius[5] boissyi Abeille
Balduf (1935) lists Ripidius boissyi as parasitic on nymphs of Ectobius pallidus giving Abeille de Perrin (1909) as a source for this information. However, Abeille de Perrin simply presumed that R. boissyi parasitized E. pallidus because he collected this cockroach in the same habitat as the beetle. Abeille de Perrin suggested that the species of the genus Ripidius lived in the bodies of cockroaches, but there are no rearing records, as far as we know, of R. boissyi from cockroach hosts.
Ripidius denisi Chobaut
Chobaut (1919), in France, collected both R. denisi and Ectobius pallidus when beating an oak tree. Because of the known association of other species of Ripidius with cockroaches, he presumed that this beetle was parasitic on E. pallidus, a cockroach common in this beetle's habitat.
Ripidius pectinicornis Thunberg
Synonymy.—Symbius blattarum Sundevall [Leng, 1920].
Natural hosts.—Blattella germanica, on shipboard (Sundevall, 1831); Germany (Aclogue and Fowler, in Burr, 1899a); on steamship "Samui" (Stamm, 1936); on cruiser "Duguay-Trouin" (Barbier, 1947); Hawaii (Williams, 1946a): This last record was based on a specimen dissected from an adult German cockroach collected on an airplane from the South Pacific. The parasite was reported as Ripidius sp. by Williams, but Weber (1948) made the specific identification.
Ectobius pallidus? Abeille de Perrin (1909) stated that R. pectinicornis was first described by Sunders [sic] as blattarum because it had been captured in the body of Ectobia livida. We presume that Abeille de Perrin was referring to Sundevall's work in which the host was given as Blattella germanica.
Periplaneta americana, on shipboard (Sundevall, 1831): One nymph only.
With the exception of the single nymph of P. americana, R. pectinicornis apparently attacks only adult females and nymphs of B. germanica. Barbier (1947) found only B. germanica parasitized, although both Blatta orientalis and Supella supellectilium were prevalent on board the ship. Primary larvae of the parasite failed to parasitize Supella.
Adult behavior.—The winged male is relatively active compared to the apterous female; it runs around, flies well, and jumps on the female when in her vicinity. The female remains stationary and lays eggs around her by bending her long ovipositor (Sundevall, 1831). The eggs (50-100) are laid among a network of silk fibers secreted by the female. The female dies after completing oviposition (Barbier, 1947).
Development.—The eggs hatch after 14 days, and the primary (triungulin) larvae ascend the host's legs to its body; the larvae then cut the intersegmental membrane between the metasternum and first abdominal segment of the cockroach, in order to enter the host's abdomen (Barbier, 1947). Chobaut (1892) first suggested this method of attack by the ripiphorid larva. As the parasites develop, the abdomen of the host becomes swollen. Developing larvae apparently eat the host's fat body, leaving the vital organs until the last. Parasitized female hosts were sterile and the eggs, when formed, never hatched. Development of the oötheca was also inhibited. There were usually two larval parasites per host, but three or four were found several times (Barbier, 1947). Sundevall (1831) found only one larva per cockroach except one host which, when crushed, yielded five. Stamm (1936) found three hosts infested with five larvae each. In a little over 100 cockroaches, Stamm found 10 that were parasitized.
The day before the parasite leaves the host, the cockroach shows an abrupt uneasiness and runs about, finally falling over on its back. The parasite larva emerges from the host through an opening it makes in the membrane between penultimate and last tergite. The host dies a few hours after the larva has left. The larva seeks a sheltered area and pupates within 48 hours. Adults emerge in 9 days (females) and 13 days (males) (Barbier, 1947).
Distribution.—Adult males have been collected in light traps in Hawaii (Van Zwaluenburg, 1946), and the first female was reported by Weber (1948); the parasite is now established in the islands around Pearl Harbor (Dr. F. X. Williams, personal communication, 1953). The U. S. National Museum has specimens of R. pectinicornis from England, Guatemala, Hawaii, Panama, and from Florida and Georgia in the U. S. (Dr. E. A. Chapin, personal communication, 1953). Kono (in Asano, 1937) reported two species in Japan. It is noteworthy that all these records are from localities adjacent to oceans and on ships; none are from interiors of continents. The only biological data were obtained from parasites found on board ships. Sundevall (1831) believed that the parasites boarded his ship with their hosts during loading in Calcutta, since before that not any were seen on board. Barbier (1947) suggested that the parasite must be spread very easily in ports between neighboring ships by parasitized cockroaches in baskets or sacks of provisions.
Ripidius scutellaris Heller
Natural hosts.—Blattidae, Philippine Islands (Schultze, 1925).
Family DERMESTIDAE
Dermestes ater De Geer
Common name.—Black larder beetle.
Natural prey.—Blatta orientalis, U.S.A. (Roth and Willis, unpublished data, 1953): Dermestes ater is generally a scavenger, but we have seen adult beetles, which had developed in our cockroach colony, clinging to and feeding on living oriental cockroaches, eventually killing them; the beetles probably attack only the weakened or injured cockroaches in a culture. This was a natural infestation of a laboratory culture by a predator.
Experimental prey.—Blattella germanica, oöthecae, U.S.A. (Roth and Willis, 1950): The beetle larvae can penetrate unhatched oöthecae of the German but not those of the American or oriental cockroaches.
Dermestes sp.
Natural prey.—Blatta orientalis, oöthecae, U.S.A., Missouri: Rau (1924) stated that Dermestes larvae often infest the egg cases of this cockroach; it is probable that Rau was referring to cockroaches in laboratory cultures.
Order STREPSIPTERA
Pierce (1909) predicted that the Blattoidea and the Grylloidea would be the only groups of the Orthoptera which would be parasitized by Strepsiptera. Essig (1926) made the statement that certain cockroaches are among the hosts of Strepsiptera. E. F. Riek (personal communication, 1952) found a strepsipteron in a late nymph of Cutilia sp. from Waroona, Western Australia; he wrote us, "The female parasite is extruded between a pair of sternites towards the base of the abdomen and appears to belong to the family Halictophagidae." This is the only record that we have been able to find of a strepsipteron parasitizing cockroaches.
Order HYMENOPTERA