Chapter XIII: Section III: (p. 14). Other associations may be accidental (e.g., certain (7)

Shaw (1925) claimed that _Supella supellectilium_ tended to oust _Blattella germanica_, but Pope (1953) thought it doubtful in Queensland. Wolcott (1950) stated that "The larger and more powerful domestic cockroaches, _Periplaneta americana_ (L.), _P. australasiae_ (F.) and _P. brunnea_ Burmeister have very definitely fallen behind in Puerto Rico in competition with the little German roach." Pessôa and Corêa (1928) observed that other species of cockroaches were rare in Brazil in houses that were infested with _Leucophaea maderae_. Lederer (1952) noticed that in the reptile house of the aquarium at Frankfort am Main _Blatta orientalis_ was obviously kept down by _Blattella germanica_, even before the appearance of _P. americana_. However, _B. germanica_ was not driven out of the reptile house by _P. americana_ although the populations of each fluctuated for about 22 years after the American cockroach had settled there; both species occupied separate resting places. Lederer further observed that within four years of the introduction of _P. americana_ into the crocodile house, none of the original infestation of _B. orientalis_ could be found; a small colony of _Pycnoscelus surinamensis_ in the reptile house was apparently also driven out by _P. americana_. Chopard (1932, 1938) stated that the oriental cockroach does not exist in company with _P. americana_ which very probably destroys it. Pettit (1940) kept _B. germanica_ and _P. americana_ together in a cage for several weeks but neither species gave any indication of feeding on the other.

Froggatt (1906) stated that "It is probable that the advent of the larger and more formidable American cockroach into Australia has led to the retirement or destruction of our indigenous species" [presumably _Periplaneta australasiae_]. Tillyard (1926) noted that this statement is incorrect as neither species is native to Australia. Yet Shaw (1925) stated that in Australia "When both species live together in the same places, _australasiae_ Fabr. will probably be found gradually to displace _americana_ L." Local fluctuations in the relative abundance of these species could be a basis for such dissimilar observations. However, MacDougall (1925) observed that in the plant houses of the Royal Botanical Garden, Edinburgh, the Australian cockroach seemed to have overcome the American which had been more numerous in former years.

In conclusion, we emphasize that many of the above observations are merely tentative impressions gathered by workers who have watched many species of cockroaches in nature. Obviously, additional observations coupled with appropriate experimentation will be needed to disclose the true structure of each presumed association and to resolve apparent discrepancies. Although we are greatly indebted to the cited authors for their contributions to the known information, we anticipate that future results of cleverly designed laboratory experiments will do much to dispel the uncertainty that still surrounds our knowledge of the relations of the Blattaria to each other.

XVIII. DEFENSE OF COCKROACHES AGAINST PREDATORS

Irritating or repellent secretions provide many animals belonging
to widely unrelated groups with a more or less potent means of
defence....

It will be seen that this method of defence does not rest merely
upon a passive unpalatable attribute, but upon an active emission
of the unpalatable substance which, since it occurs when the
animal is seized or threatened by an enemy, enforces its
effectiveness. In its highest development we find different forms
whose specialized habits and modified structure enables them to
_project_ secretion at the enemy, and thus to discourage attack.

COTT (1940)

There are very few records indicating that cockroaches are unaccepted as food by other animals. Hutson (1943) found that the duck, guinea fowl, and pigeon would not normally eat _Pycnoscelus surinamensis_, and in his experiments with the chicken eye worm he had to force-feed his birds with infected cockroaches. Lederer (1952) found that insectivorous birds in the Zoological Garden, Frankfurt am Main, either refused hardened (as opposed to teneral) American cockroaches or ate them unwillingly. Carpenter (1925) reported that a monkey (_Cercopithecus_) failed to feed on cockroaches and suggested that the insects' odor made them repugnant; however, there are a number of positive records of monkeys feeding on cockroaches (see pp. 284-286).

Cockroaches may escape capture by predators through evasive behavior, concealment, protective coloration, mimicry, or secretion of malodorous materials. Nocturnal cockroaches may avoid predators that are active during the day (Crawford, 1934), but nocturnal predators are apparently quite successful in capturing cockroaches. Some cockroaches may be protected by their swiftness, others by their resemblance to vegetation (Williams, 1928). The habit of squeezing into narrow cracks may afford cockroaches some protection.

Burrowing forms such as _Pycnoscelus_ may spend much time in underground cells (Roeser, 1940). Polyphagids rapidly burrow into sand (Fausek, 1906), where they may be protected from predators. Tepper (1893) discovered that a very large Australian cockroach, _Geoscapheus robustus_, had its fore legs, especially the tibiae, adapted for digging. He observed this species in captivity and in 1894 reported that it appeared to sink into the soil without raising any considerable amount above the surface and that it did not form an unobstructed tunnel. Another large Australian cockroach, _Macropanesthia rhinocerus_, burrows about two feet below the surface of sandy soil; it also makes nests among pine roots and the nymphs rarely appear above ground (Henson _in_ Day, 1950). Tepper (1893) observed that Australian cockroaches of the genera _Epilampra_ and _Oniscosoma_ buried themselves in loose soil and dust. Baker (_in_ Rehn, 1930) observed that _Styphon bakeri_ is found in humus and rubble in the Dutch West Indies where "It is sluggish in the open, but wedges into the humus quite quickly."

_Therea nuptialis_, found in India, conceals itself at the roots of fig trees, etc. The small hairs on its elytra retain sufficient dust to conceal it, or at any rate to render it inconspicuous, when not on the wing (Annandale, _in_ Chopard, 1924c). Rehn and Hebard (1914) observed that the nymphs of _Blaberus craniifer_[13] at Key West, Fla., "were usually found half buried in loose damp earth under boards, where they remained motionless, looking much like lumps of earth (with which they were usually much dusted) until disturbed." Hebard (1917) reported of _Monastria biguttata_ from Brazil that "All of the juveniles are heavily coated with foreign particles" which adhere "to a multitude of closely placed, minute and usually curved spines, which cover the dorsal surface and marginal portions of the ventral surface."

It is apparent from the numbers of predators reported herein that many animals are not deterred by the odorous secretions of cockroaches; these secretions, because they may seem repugnant to man, are often claimed to be repellent to predators. However, Cott (1940) points out that "There are many instances in which protective devices and associated warning colours are known to be ineffectual against certain enemies. But this does not necessarily imply that they are not on the whole beneficial to the species attacked." Certain cockroach secretions may well be repellent to many predators, but as this is a purely negative aspect of the predator-prey relationship little thus far has been observed or published. Potential prey that successfully defends itself against attack is never found in a predator's stomach.

Cockroaches have a variety of glands which secrete odorous materials. Certain secretions, produced by tergal or dorsal glands in males, are involved in sexual behavior; the females feed on the secretion from these glands prior to copulating (Roth and Willis, 1954). However, other secretions which are produced by both sexes are ejected or given off when the insect is disturbed; undoubtedly these are defensive weapons that are used against predators. Very few experiments or observations are on record to show how effective these secretions may be in protecting the cockroach. Although the morphology of some of the glands has been described, relatively little is known about the chemistry of their secretions.

Many species of Australian cockroaches have been reported to emit "disgusting" odors, though the glands producing these secretions have not been described, nor is the chemistry of the compounds known. _Cosmozosteria lateralis_ exposed two orange-red spots on the abdomen while emitting a pungent odor which deterred a collector from capturing it (Shelford, 1912). Another Australian species, _Platyzosteria castanea_, when disturbed on barren ground tilts forward on the vertex and straddles out the posterior legs, supporting itself in a vertical position on the head and tarsi; in assuming this attitude it will squirt a foetid fluid as a fine spray for a distance of 6 or 7 inches (Shaw, 1914). Spencer (1892) mentions the pungent odor given off by a cockroach which had been accidentally cut in two. Rageau (1956) stated that in the New Hebrides and New Caledonia _Cutilia nitida_ emits, when disturbed, a corrosive liquid with an extremely disagreeable odor.

The adults of _Eurycotis floridana_ emit an odorous fluid when seized (Rehn and Hebard, 1905). The fluid, which may irritate sensitive skin areas, may be ejected as a spray for a distance of several inches. This secretion has been identified as 2-hexenal (Roth et al., 1956), and the ventral abdominal glands which produce it have been described (Stay, 1957). Eisner (personal communication, 1958) has found that the toad _Bufo marinus_ and the frog _Rana pipiens_ invariably spit out adults of _E. floridana_ which they have seized. The odor of 2-hexenal was strongly apparent after these attacks, and the insect was never damaged. However, the lizard _Anolis equestris_ seized and crushed _E. floridana_ before releasing its hold and dropping the insect 5 to 10 minutes later. The blue jay _Cyanocitta cristata_ readily attacked adults of _E. floridana_ and killed them but did not eat the insects until after the odor had dissipated; however, the bird carried nymphs of _E. floridana_ to its perch and ate them. Nymphs of this species do not secrete 2-hexenal (Roth et al., 1956). Recently, 2-hexenal has been tested for its antibacterial activity and has been found to be active against seven species of pathogenic bacteria (Valcurone and Baggini, 1957). _Eurycotis decipiens_ from Trinidad also ejects a fluid which may produce toxic symptoms such as vertigo and nausea (Bunting _in_ Roth and Willis, 1957a).

Large reservoirs of glands similar in appearance and position to those of _Eurycotis floridana_ are present in the adults of both sexes of _Neostylopyga rhombifolia_ and _Platyzosteria novae seelandiae_. Walker (1904) and Longstaff (_in_ Shelford, 1912) noted that the latter species had a strong odor. Roth (unpublished data, 1957) found that the secretion of _P. novae seelandiae_ when ejected is grayish or milky in color. In the reservoirs of the ventral gland of this insect the secretion is a milky liquid containing floating greenish globules. Both infrared and mass spectrographic analyses show that the secretion is a mixture containing 2-hexenal, the aldehyde that is found in _E. floridana_. Eisner (personal communication, 1958) observed that the lizard _Anolis carolinensis_ immediately released _Neostylopyga rhombifolia_ without injury, but that _Bufo marinus_, _Anolis equistris_, and _Cyanocitta cristata_ ate the insect despite the secretion; several unidentified spiders and the ant _Pogonomyrmex badius_ were not repelled by the secretion of _N. rhombifolia_.

Dorsal and ventral glands have been found in both sexes of _Blatta orientalis_ and _Periplaneta americana_ (Minchin, 1888, 1890; Kul'vets, 1898; Oettinger, 1906; Harrison, 1906; Liang, 1956). The ventral glands are found in the same general region as those of _Eurycotis_. We have also found similar ventrally located glands in both _Periplaneta australasiae_, and _P. brunnea_. The reservoirs which store the secretion of the ventral glands are smaller in _Blatta_ and _Periplaneta_ spp. than those found in _Eurycotis_, _Neostylopyga_, or _Platyzosteria_.

In _Blatta orientalis_ the dorsal glands can be everted by pressure on the abdomen; the secretion in these glands, according to Haase (1889), has the typical oriental cockroach odor. Although the dorsal glands of the oriental cockroach are usually given a defensive role (Haase, 1889, 1889a; Kul'vets, 1898; Oettinger, 1906; Konček, 1924), the functions of secretions of these nonepigamic dorsal glands and the ventral glands are still open to question. It is possible that some of the odors produced by cockroaches have functions other than defense or sex attraction. For example, Ledoux (1945) showed that the species odor is largely responsible for the gregarious behavior shown by _Blatta orientalis_ and _Blattella germanica_. The olfactory stimulus acts over a short distance only, and the source of this odor in the insect is unknown. By washing _Blattella germanica_ in warm chloroform Dusham (1918) extracted a wax which had the odor of the German cockroach. However, there is no evidence to show that cockroaches respond to the same cockroach odors that are detected by man.

Certain cockroaches have recently been found to have odorous secretions which are produced in tracheal glands. In _Diploptera punctata_ the tracheae leading to the second abdominal spiracles of nymphs and adults are modified into odoriferous glands which produce a mixture of 2-ethyl-1,4-benzoquinone; 2-methyl-1,4-benzoquinone; and _para_ benzoquinone; this material is ejected as a means of defense. The offensive odor emitted by adults and nymphs of _Leucophaea maderae_ also issues from the second abdominal spiracles (Roth and Stay, 1958).

_Diploptera_ is capable of ejecting its quinones from either its right or left tracheal gland according to which side of the insect is attacked (pl. 36, A-B). Eisner (1958) found that the secretion repelled the ant _Pogonomyrmex badius_ (Latreille) (pl. 36, C) and the beetle _Galerita janus_ Fabricius when they attacked the cockroach. The spider _Lycosa helluo_ Walckenaer was repelled by large nymphs and adults of _D. punctata_ but young nymphs were usually eaten promptly (Eisner, 1958).

Bordas (1901, 1908) believed that the "conglobate" gland (Miall and Denny, 1886), found in males of _Periplaneta americana_ and _Blatta orientalis_, was an odoriferous gland used for defense, but Gupta (1947) has shown that in all probability this gland (the phallic gland) secretes the outermost covering of the spermatophore.

What appears to be mimicry occurs in some species of Blattaria. The nymphs of many Panchlorini and Blaberinae vaguely resemble sow bugs (Chopard, 1938). Certain members of the Perisphaerini (e.g., _Perisphaerus glomeriformis_) from the Malayan region which resemble sow bugs (Annandale, 1900; Hanitsch, 1915) can roll themselves up into a ball thus hiding their antennae and legs (Lucas, 1862). Although these cockroaches are found among dead leaves or under stones, in places in which sow bugs are also found, the benefit to either or both forms is questionable; Annandale (1900) believed that the crustacean and the cockroach, living under similar conditions, developed the same general body shape. Rolling up into a ball is nothing more than an exaggeration of a reflex common to many young cockroaches, that is, an arched position which these insects assume when they immobilize themselves in response to certain stimuli (Chopard, 1938).

There are cockroaches that resemble various Coleoptera and Hemiptera (Belt, 1874; Shelford, 1912; Hanitsch, 1915). Some look like cerambycids, lampyrids, coccinellids, pentatomids, etc. Perhaps the most striking examples are the resemblances of cockroaches in the genus _Prosoplecta_ of the Epilamprinae to beetles of the family Coccinellidae; Shelford (1912) has figured a number of species of _Prosoplecta_ together with the species of beetles which they seem to have taken for models. Williams (1928) mentioned diurnal cockroaches which by a combination of markings, shape, posture, and active flight about vegetation suggest certain wasps.

Unfortunately, practically nothing is known about the behavior of these so-called mimics and models or their relationships with predators in the field. For the most part, the examples are based on a comparison of pinned insects from museum collections (Burr, 1899); for this reason Chopard (1938) believed that not much value should be placed on superficial resemblances of this kind. However, we believe that a lack of knowledge of cockroach mimicry is not a valid reason for rejecting the idea that mimicry, if it occurs, may be of some benefit in the survival of mimetic species. Certainly Cott's (1940) voluminous compilation of the literature on adaptive coloration should make the most skeptic hesitate to conclude dogmatically that these instances of mimicry are merely accidental and meaningless.

XIX. THE BIOLOGICAL CONTROL OF COCKROACHES

In the Navy [Japanese] a seaman who has captured 300 cockroaches
will be granted one day special shore leave. They call it "shore
leave for cockroaches." The purpose is to promote extermination of
cockroaches in a warship because, on the one hand, any warship
suffers from numerous cockroaches, and, on the other hand, any
seaman likes shore leave.... The formalities for a shore leave for
cockroaches are as follows. A seaman keeps cockroaches which he
captured (mainly _B. germanica_, because _P. americana_ and _P.
australasiae_ are seldom found in Japan) in a bottle or in a bag
until the number reaches 300. Then he brings them to the deck
officer to get the confirmation that he has actually captured more
than 300 cockroaches. If the deck officer confirms it, the seaman
goes to a cabin where a petty officer reports that the deck
officer confirmed the number of cockroaches. The petty officer
signs the seaman's name, name of division, rank, and date to be on
shore leave in the log book for cockroach shore leaves. The petty
officer brings the log book again to the deck officer to get his
approval and then goes to the commander for the final approval. In
the Navy, they have another special shore leave for rats. In this
system, a seaman gets one day shore leave for one rat. The
formalities for the latter are the same as for the former, and
there is a log book for the rat shore leave in the petty officer's
quarters. The author took advantage of these systems frequently.

SONAN (1924)

Little is known of the effects of predatism and parasitism on natural populations of cockroaches. Many statements in the literature are very general; yet there are a few data on egg parasites (e.g., _Tetrastichus hagenowii_) which suggest that, in the absence of parasites, populations of domestic cockroaches might be much larger than they are in certain areas. We have summarized the literature on natural control and also that on the use by man of predators and parasites in the biological control of cockroaches. However, because of the paucity of information, we have been unable to evaluate the effectiveness of biological control in reducing the numbers of pest cockroaches. This is an area that might reward further investigation.

INVERTEBRATES

_Scorpions._--In Puerto Rico, cockroaches are probably the principal food of the scorpions which live in old houses, on tree trunks, etc. (Seín, 1923). The staple diet of scorpions in Arizona is the small cockroach commonly known as the water bug (Stahnke, 1949); in the part of Arizona where he resides, Stahnke (personal communication, 1953) says that the "water-bug" is most generally _Supella supellectilium_ although _Blattella germanica_ is also found, but less abundantly.

_Spiders._--Jefferys (1760) mentioned a large spider which was protected in the Antilles and especially on Guadeloupe because it hunted down and fed on cockroaches; the spider was reputed to be common in every house. Sir Hans Sloane (1725, _in_ Cowan, 1865) reported that residents of Jamaica kept spiders in their houses to destroy cockroaches. Takahashi (1924) reported that, in the Taihoku area of Formosa, human habitations contained large numbers of spiders which caught and ate cockroaches. Smith (_in_ Marlatt, 1915) reported that Brazilians encourage large house spiders because they tend to rid the house of "other insect pests." In British Guiana tarantulas were kept in a bungalow to control _Periplaneta_ and _Pycnoscelus_ (Beebe, 1925a).

_Ants._--A Madam Merian noticed that ants cleared houses of cockroaches (Kirby and Spence, 1822). A small reddish-yellow ant, called Pucchuçiçi by Peruvian Indians, pursued and destroyed a cockroach called Chilicabra which was a pest in native huts (Tschudi, 1847). Schwabe (1950b) found swarms of ants attacking living _Pycnoscelus surinamensis_ and stated that ants are probably the chief enemy of this cockroach in Hawaii. Wallace (1891) stated that in Africa a band of driver ants may enter a house and clear it of cockroaches and other arthropods. In British Guiana, Beebe, (1925) found that several times a year army ants cleared the laboratory of all cockroaches and tarantulas.

_Wasp egg parasites._--Matsumura (1917, _in_ Asano, 1937) proposed that parasitic wasps such as _Evania_ and _Brachygaster_ be protected in Japan as the natural enemies of cockroaches. In one area in France, 20 percent of the oöthecae of _Loboptera decipiens_ were parasitized by _Zeuxevania splendidula_ (Genieys, 1924). Edmunds (1952a) found that 12 percent of 459 oöthecae of _Parcoblatta_ collected during December through April of 1950-51 in Ohio were parasitized; evaniids accounted for about 7 percent of the parasitization. Additional collection data in 1951-52 Edmunds (1953a) showed that 8.7 percent of 320 wood-cockroach oöthecae were parasitized; 2.8 percent of these parasites were evaniids; almost 13 percent of the egg capsules collected showed evidence of previous parasite emergence. Cameron (1957) reported that oöthecae of _Periplaneta americana_ collected in Saudi Arabia were 29 percent parasitized in March and 25 percent parasitized in October by _Evania appendigaster_. Sonan (1924) found 1 of 65 oöthecae of _P. americana_ and _P. australasiae_ parasitized by _E. appendigaster_ in Formosa.

Cottam (1922) stated that the increase of _Supella supellectilium_ in Khartoum was checked by a wasp egg-parasite that was later identified as _Anastatus tenuipes_ (see p. 246) (Ferrière, 1930, 1935). In this country, this wasp seemed to be effective in decreasing the numbers of _Supella_ in certain areas in Arizona (Flock, 1941).

In Formosa, _Tetrastichus hagenowii_ was an important parasite of cockroach eggs (Maki, 1937). Sonan (1924) reported 30 percent parasitization of 65 oöthecae of _Periplaneta americana_ and _P. australasiae_ collected in Formosa. In Bangalore, India, the natural parasitization of randomly collected oöthecae of _P. americana_ varied from 21 percent (of 495 oöthecae), July 1947-June 1948, and 43 percent (of 288 oöthecae), July-December 1948, to 57 percent (of 178 oöthecae), July-October 1949 (Usman, 1949). Cameron (1955) obtained _T. hagenowii_ from oöthecae collected in Trinidad, B.W.I., and Saudi Arabia; some 15 percent of the oöthecae of _P. americana_ and _P. australasiae_ collected in October in Trinidad were parasitized; a later collection (March) was 34 percent parasitized; a small sample of _P. americana_ oöthecae was 65 percent parasitized. The oöthecae collected in Saudi Arabia in March were 20 percent parasitized. Plank (1947) found that the eggs of the American cockroach in Puerto Rico (probably in laboratory cultures) were so heavily parasitized by _T. hagenowii_ that he had to use _P. australasiae_ for experimental purposes; in 1950 Plank stated that more than 50 percent of American cockroach oöthecae were parasitized.

Fahringer (1922) stated that _Prosevania punctata_ could be used to eradicate cockroaches, but he did not test his hypothesis. Marlatt (1902) felt that the usefulness of _Evania appendigaster_ in biological control was impaired by _Tetrastichus_ acting as a hyperparasite (see footnote 6, p. 236). However, Wolcott (1951) stated that in Puerto Rico _E. appendigaster_ is quite abundant and is a factor of considerable importance in controlling cockroaches. Kadocsa (1921) stated that _Brachygaster minutus_ and _Evania appendigaster_ were not important in the biological control of cockroaches. These general statements are not supported by experimental evidence.

It is likely that the smaller wasp egg parasites are more effective than the evaniids in controlling cockroaches. Only one evaniid develops in a parasitized oötheca, but many individuals of the other wasps develop in one oötheca and the number of females that emerge is usually large. However, Cameron (1957) concluded that, with a parasitism rate of 25 to 29 percent and three to four generations a year, against one or less for the host, _Evania appendigaster_ in the areas where it is established is a valuable control agent.

The use of specific egg parasites to control cockroaches has not been attempted extensively. Cros (1942) liberated a species of _Tetrastichus_ (=_Eulophus_ sp.; see p. 254) in his home in Algeria to control the oriental cockroach; as far as we know, he did not report the parasite's effectiveness in reducing the cockroach population. According to Zimmerman (1948) _Comperia merceti_, when accidentally imported, practically wiped out _Supella supellectilium_ in parts of Hawaii; he claimed to have controlled the brown-banded cockroach in a store building with this parasite. In some parts of Honolulu, almost 100 percent of the oöthecae of this cockroach were parasitized (Zimmerman, 1944). We (1954b) ran some simulated field tests in which we liberated _Tetrastichus hagenowii_ in rooms artificially seeded with oöthecae; from 28 to 83 percent of American cockroach oöthecae and 56 percent of oriental cockroach oöthecae were parasitized during these tests.

_Evania appendigaster_ was introduced from Hawaii into Canton Island in 1940 against _Periplaneta americana_, and it has become established (Dumbleton, 1957). This parasite was also successfully introduced into Samoa (Dumbleton, 1957).

_Cockroach-hunting wasps._--An earnest attempt has been made to establish in Hawaii wasps that prey on cockroaches. Just how effective these wasps are in controlling cockroaches is still unknown. _Dolichurus stantoni_ was introduced from the Philippines in 1917 and spread to several of the Islands (Swezey, 1920, 1921; Williams, 1944). Bridwell (1920) stated that as a result of this introduction there was a great decrease in cockroaches of the genus "_Phyllodromia_." A number of _Podium haematogastrum_ from Brazil were liberated in Honolulu (Williams, 1925) but did not become established (Williams, 1928). The effectiveness of _Podium_ was questioned by Williams (1928) who observed that _Podium_ "destroyed innumerable Blattidae, which nonetheless swarmed in their neighborhood, and I must confess from my observations on the various cockroach-hunting wasps that the blattid more than holds its own alongside its enemy."

Introductions of _Ampulex_ have proved more successful. _Ampulex canaliculata_ was introduced into Hawaii from the United States (Williams, 1928a, 1929). Williams also introduced _A. compressa_ into Hawaii in 1940, and the species was reared in large numbers for distribution (Pemberton, 1942). _A. compressa_ has since become established on most of the Islands (Pemberton, 1945a, 1947; Williams, 1946; Van Zwaluwenburg, 1950). The thousand of _A. compressa_ now found in the Hawaiian Islands are all descendants of three wasps captured in Noumea, New Caledonia (Williams, 1944). According to Williams (1941), the number of cockroaches was noticeably reduced at the University of Hawaii poultry farm, where some _A. compressa_ were released. Pemberton (1953) believed that this wasp has become sufficiently abundant to be of definite value. Simmonds (1941) recommended importing _A. compressa_ into Fiji for cockroach control. _A. compressa_ was introduced from Hawaii into Guam in 1954 against _Periplaneta americana_ and into the Cook Islands in 1955 against _Periplaneta_ spp.; it is not yet known whether the parasite became established in either place (Dumbleton, 1957).

VERTEBRATES

... on conserve avec soin les crapauds dans les maisons, et que les
dames les tolèrent, même sous leurs robes, en raison de leurs
continuels services, car ils se promènent sans cesse à la recherche
des Kakerlacs.

GIRARD (1877)

_Toads._--_Bufo marinus_ was first introduced into Puerto Rico from Barbados in 1920 to reduce several major insect pests including cockroaches (Leonard, 1933). It was introduced from Puerto Rico into Hawaii by C. E. Pemberton in 1932 where it rapidly became established; it has since been distributed throughout the Pacific area. _B. marinus_ is one of the world's largest toads; it attains a body length (exclusive of the hind legs) of 7 to 9 inches (Oliver, 1949) and has been kept alive for more than 11 years in captivity (Pemberton, 1945). Alicata (1938) placed giant toads in a fenced area in Hawaii containing an infestation of _Pycnoscelus surinamensis_; after 24 hours the toads were dissected and each was found to have eaten from 11 to 25 cockroaches. Illingworth (1941) found that 40 to 90 percent of 53 stools of this toad in Hawaii contained remains of _P. surinamensis_. Alicata (1947) recommended the maintenance of _B. marinus_ in poultry yards to reduce the population of _P. surinamensis_, the vector of the chicken eye worm.

Toads have also been recommended for controlling cockroaches in houses (Meech, 1889; Sweetman, 1936). Girard (1877) cited a note in a French newspaper which stated that toads were kept in houses in Cuba to control the American cockroach.

_Tree frogs._--Tree frogs enclosed in a room overnight were said to effectively clear it of cockroaches (Marlatt, 1915); on sugar plantations in Australia, these amphibians were encouraged in houses and kept as pets because they hunted and devoured large brown cockroaches (Froggatt, 1906).

_Birds._--In Guadeloupe, Dutertre (1654) claimed that all the fowls of the country were fond of small cockroaches and lived on practically nothing else. In Hawaii (Zimmerman, 1948) and in the Lesser Antilles (Ballou, 1912) cockroaches are eaten by poultry whenever the birds can find them. In Puerto Rico, Wetmore (1916) stated that owls kept in houses feed extensively on cockroaches; the stomach of one owl which had been kept in a native house was filled entirely with cockroaches. In British Guiana, Beebe (1925) found that cockroaches were eaten by 27 species of birds.

_Reptiles._--H. (1800) claimed that two lizards cleared his house of the "true brown cockroach" and suggested that lizards be used for cockroach control because the reptiles are docile and harmless. On Arno Atoll geckos and night-feeding skinks eat large numbers of cockroaches (Usinger and La Rivers, 1953). According to Wolcott (1924) the number of cockroaches eaten by lizards is surprisingly large considering the nocturnal habits of these insects. Beebe (1925a) kept geckos in a bungalow to help control _Periplaneta_ and _Pycnoscelus_.

_Mammalia._--Cowan (1865) stated that in England hedgehogs were often kept domesticated in kitchens to destroy cockroaches. This writer also stated that a lemur was kept on board ship to destroy cockroaches.

Large numbers of the American and Australian cockroaches were eaten by the mongoose in Hawaii (Perkins, 1913).

ACKNOWLEDGMENTS

We would have been unable to complete this review without the help of many people who have generously given us their time and the benefit of their special experience. We are exceedingly grateful to these individuals for they have contributed much to whatever merit this work possesses; we alone are responsible for the deficiencies and inaccuracies that remain in the text.

Dr. A. B. Gurney, Entomology Research Division, U. S. Department of Agriculture, and J. A. G. Rehn, Academy of Natural Sciences of Philadelphia, have given us much help and advice throughout the preparation of this monograph. Both have patiently answered our many queries, and Mr. Rehn allowed us free access to his large collection of cockroach literature. We are especially pleased to thank them for their many favors.

Many persons have determined at our request the identity of specific organisms. These individuals are cited in the text and to them we extend our thanks. We thank our colleagues, cited in the text, who have made their unpublished observations available to us. We also thank the individuals and organizations for the use of photographs and/or drawings for which they are credited in the accompanying legends.

We thank the following individuals for supplying us with living specimens of the species indicated: T. Campbell, Commonwealth Scientific and Industrial Research Organization, Canberra, New South Wales (_Panesthia australis_); Dr. L. R. Cleveland, Harvard University (_Cryptocercus punctulatus_); Dr. N. T. Davis, University of Connecticut (_Byrsotria fumigata_); Dr. F. Englemann, Albert Einstein Medical School (_Gromphadorhina portentosa_); Dr. F. A. Lawson, Kansas State College (_Comperia merceti_); Dr. K. D. Roeder, Tufts University (_Hierodula tenuidentata_ (?)); Dr. E. O. Wilson, Harvard University (_Ischnoptera deropeltiformis_).

We thank the following individuals for checking the taxonomy of the following organisms or for reading the indicated sections of the manuscript: _Fungi._--Dr. R. K. Benjamin, University of California; Dr. E. G. Simmons, Quartermaster Research Laboratories. _Protozoa._--Dr. R. R. Kudo, Professor Emeritus, University of Illinois. _Helminths._--Mrs. May Belle Chitwood and Dr. J. T. Lucker, U. S. Department of Agriculture. _Centipedes and whipscorpions._--Dr. R. E. Crabill, Jr., U. S. National Museum. _Scorpions._--Dr. J. L. Cloudsley-Thompson, University of London; Dr. R. E. Crabill, Jr. _Spiders._--Dr. B. J. Kaston, University of Connecticut; Dr. R. E. Crabill, Jr. _Mites._--Dr. J. H. Camin, Chicago Academy of Sciences; Dr. E. W. Baker, U. S. National Museum. _Cockroaches._--Dr. A. B. Gurney and J. A. G. Rehn. _Ants._--Dr. W. L. Brown, Jr., Museum of Comparative Zoology, Harvard College. _Hymenopterous parasites of cockroach eggs._--Dr. B. D. Burks, U. S. National Museum; Dr. H. K. Townes, University of Michigan. _Cockroach-hunting wasps._--Dr. K. V. Krombein, U. S. National Museum. _Lepidoptera._--Dr. J. F. G. Clarke, U. S. National Museum; Dr. E. L. Todd, U. S. Department of Agriculture. _Miscellaneous insects._--Dr. R. S. Beal, Dr. A. B. Gurney, C. W. Sabrosky, Dr. R. I. Sailer, and J. T. Spilman, U. S. National Museum. _Fishes._--Dr. L. P. Schultz, U. S. National Museum. _Amphibians and reptiles._--Dr. Doris M. Cochran, U. S. National Museum. _Birds._--Dr. Herbert Friedmann, U. S. National Museum. _Mammals._--Dr. D. H. Johnson, U. S. National Museum.

We thank the following individuals for reading the entire manuscript: J. A. G. Rehn; Dr. A. B. Gurney; Maj. Gordon Field, U. S. Army; and Dr. H. L. Sweetman, University of Massachusetts. The monograph has profited by the friendly criticism of these entomologists.

We thank Dr. R. A. Howard, Harvard University, for checking lists of plant names; Mrs. Maria E. W. Torok, formerly of the Quartermaster Technical Library, for assistance in obtaining obscure literature; Miss Louise Bercaw, U. S. Department of Agriculture Library, for identifying the journal containing the paper by Vlasov and Miram; the individuals who translated foreign language articles, for which they are credited in the bibliography; and Miss G. Lillian Fede, Quartermaster Research Laboratories, for typing the manuscript.

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