Section Iii (p. 14). Other Associations May Be Accidental (e.g., Certain

unique observations that have never again been confirmed). In the absence of contrary evidence, most associations are presumed to be benign; exceptions to this conclusion are found among the cockroaches that feed on living plants (p. 162) and those allegedly captured as prey by the carnivorous pitcher plants (Sarracenia and Nepenthes). In all the records cited below the cockroaches were stated to have been on, in, or feeding on the plant.

The plants are listed below by family according to the taxonomic arrangement of Lawrence (1951). Botanical nomenclature follows Bailey (1925), Fernald (1950), or Dr. R. A. Howard (personal communications, 1958, 1959). We take full responsibility for referring to appropriate taxa certain plants that were reported by common name only in the cited literature.

Division PTERIDOPHYTA

Family CYATHEACEAE

=Alsophila= sp.

Associate.--Pycnoscelus surinamensis, Louisiana (Anonymous, 1893): Feeding on heart of tree fern.

Family POLYPODIACEAE

=Asplenium nidus= Linnaeus

Associate.--Comptolampra liturata, Malaya (Karny, 1924): Often found between dry foliage of the beakers of this fern.

Division EMBRYOPHYTA SIPHONOGAMA

Family PINACEAE

=Pinus australis= Michaux

Associates.--Aglaopteryx gemma and Parcoblatta lata, Alabama (Hebard, 1917): The former species was common under signs on longleaf pines, and P. lata was occasional.

Parcoblatta divisa, Georgia (Rehn and Hebard, 1916): Under signs.

=Pinus caribaea= Morelet

Associates.--Eurycotis floridana, Latiblattella rehni, and Parcoblatta fulvescens, Florida (Hebard, 1917): Many records under signs on the tree trunks.

=Pinus clausa= Vasey

Associate.--Latiblattella rehni, Florida (Hebard, 1917): Under sign on tree.

=Pinus echinata= Mill.

Associates.--Parcoblatta divisa, Virginia (Rehn and Hebard, 1916): Under signs on shortleaf pine.

Parcoblatta zebra, Mississippi (Hebard, 1917): Under sign.

=Pinus sylvestris= Linnaeus

Associate.--Ectobius pallidus, England (Milton, 1899; Burr, 1899b): On Scotch fir.

=Pinus= spp.

Associates.--Plectoptera lacerna and Plectoptera vermiculata, Cuba (Rehn and Hebard, 1927).

Latiblattella rehni, Florida (Rehn and Hebard, 1905): Under signs. Cuba (Rehn and Hebard, 1927).

Family TAXODIACEAE

=Cryptomeria= sp.

Associate.--Diploptera punctata, Hawaii (Pemberton and Williams, 1938; Zimmerman, 1948).

Family CUPRESSACEAE

=Cupressus macrocarpa= Hartweg

Associate.--Diploptera punctata, Hawaii (Hebard, 1922): "The species is common and injurious in the territory infesting particularly the Monterey cypress trees ... and doing particular damage by gnawing away the bark." Similar injury has been cited by Pemberton (1934), Fullaway and Krauss (1945), and Zimmerman (1948).

=Juniperus= sp.

Associate.--Phyllodromica tartara nigrescens, Southern Uzbekistan (Bei-Bienko, 1950): Under bark.

Family PANDANACEAE

=Freycinetia= sp.

Associate.--Graptoblatta notulata and Kuchinga remota, Tahiti (Hebard, 1933).

=Pandanus= sp.

Associate.--Hololeptoblatta sp., Seychelles (Scott, 1910, 1912).

Family GRAMINEAE

=Aristida pennata= Trin.

Associate.--Phyllodromica pygmaea, U.S.S.R. (Bei-Bienko, 1950): Found in the dense turf.

=Bamboo=

Associate.--Comptolampra liturata, Malaya (Karny, 1925).

=Chloris gayana= Kunth

Associate.--Blattella vaga, Texas (Riherd, 1953): This field cockroach was rather abundant in clumps of Rhodes grass.

=Panicum purpurascens= Raddi

Synonymy.--Panicum barbinode [Hitchcock, 1936].

Associate.--Epilampra abdomen-nigrum, Puerto Rico (Seín, 1923; Wolcott, 1936): Abundant in "malojillo" meadow.

=Saccharum officinarum= Linnaeus

Associates.--Balta quadricaudata, Balta scripta, Balta torresiana, Balta verticalis, Ellipsidion simulans, and Megamareta verticalis, Australia, Queensland (Hebard, 1943): All collected by J. F. Illingworth on sugarcane.

Blattella humbertiana, Ischnoptera schenklingi, and Pycnoscelus surinamensis, Formosa (Box, 1953).

Cariblatta stenophrys, Puerto Rico (Seín, 1923; Wolcott, 1936): Between the leaves and under the leaf sheaths.

Panchlora nivea, Cuba (Rehn and Hebard, 1927): On the leaves.

Pelmatosilpha coriacea, Puerto Rico (Wolcott, 1936).

Phoraspis spp., Brazil and Guiana (Doumerc in Blanchard, 1837).

Plectoptera dorsalis, Plectoptora infulata, and Plectoptera rhabdota, Puerto Rico (Wolcott, 1950): Under the leaf sheaths.

Symploce ruficollis, Puerto Rico (Wolcott, 1950): Often found living under the leaf sheaths.

Cockroaches, Philippine Islands (Uichanco in Williams et al., 1931): Between cane leaf sheaths.

=Setaria verticillata= (Linnaeus) Beauv.

Synonymy.--Chaetochloa verticillata (Linnaeus) [Howard, personal communication, 1958].

Associate.--Diploptera punctata, Hawaii (Severin, 1911): The cockroach was caught on the barbed awns of this grass.

=Wild oats=

Associate.--Ischnoptera deropeltiformis, Missouri (Rau, 1937).

=Zea mays= Linnaeus

Associates.--Cariblatta stenophrys, Puerto Rico (Seín, 1923; Wolcott, 1936).

Ellipsidion bicolor, Australia, Queensland (Hebard, 1943).

Lophoblatta arawaka, Trinidad (Princis and Kevan, 1955).

Phoraspis sp., Brazil and Guiana (Doumerc in Blanchard, 1837).

Supella supellectilium, New Caledonia (Cohic, 1956).

Family CYPERACEAE

=Cyperus= sp.

Associate.--Maretina uahuka, Marquesas Islands, Uahuka (Hebard, 1933a).

Family PALMAE

=Acrocomia aculeata= (Jacq.) Lodd.

Associate.--Pycnoscelus surinamensis, Trinidad (Princis and Kevan, 1955): On "gru-gru" fruits.

=Cocos nucifera= Linnaeus

Associates.--Aglaopteryx gemma, Florida (Rehn and Hebard, 1912).

Cariblatta lutea minima, Florida, and Cariblatta delicatula, San Domingo (Hebard, 1916a).

Eurycotis floridana, Florida (Rehn and Hebard, 1912; Hebard, 1917).

Periplaneta australasiae, Jamaica (Rehn and Hebard, 1927).

Pycnoscelus surinamensis, Florida (Rehn and Hebard, 1912; Hebard, 1917). Jamaica (Rehn and Hebard, 1927).

=Phoenix dactylifera= Linnaeus

Associate.--Blattella germanica, California (Herms, 1926): On date palms.

=Pritchardia= sp.

Associate.--Periplaneta australasiae, Nihoa Island (Bryan, 1926). Hawaii (Zimmerman, 1948).

=Roystonea regia= O. F. Cook

Associate.--Cariblatta punctulata, San Domingo (Hebard, 1916a).

=Sabal palmetto= Lodd.

Associate.--Eurycotis floridana, Florida (Scudder, 1879).

Periplaneta australasiae, Florida (Hebard, 1917).

=Undetermined palms=

Associates.--Euthlastoblatta abortiva, Texas (Hebard, 1917).

Hormetica laevigata, Brazil (Hancock, 1926).

Panchlora antillarum, Dominican Republic (Rehn and Hebard, 1927).

Periplaneta americana, Texas (Zimmern in Gould and Deay, 1940).

Family ARACEAE

=Arum= sp.

Associate.--Latiblattella vitrea, Mexico (Hebard, 1921b): In flower shaft.

=Caladium= sp.

Associate.--Plectoptera dorsalis, Puerto Rico (Rehn and Hebard, 1927).

Family BROMELIACEAE

=Aechmaea porteoides= Britton

Associate.--Dryadoblatta scotti, Trinidad (Princis and Kevan, 1955).

=Ananas comosus= Merr.

Associates.--Pycnoscelus surinamensis, Hawaii (Illingworth, 1927, 1929): Feeding on roots of pineapple.

Blattella humbertiana, Formosa (Takahashi, 1940): Imago and grown nymphs occasionally lie concealed in the leaves.

=Catopsis fulgens= Griseb.

Associates.--Cockroaches, Costa Rica (Calvert and Calvert, 1917).

=Glomeropitcairnia erectiflora= Mez

Associate.--Dryadoblatta scotti, Trinidad (Princis and Kevan, 1955).

=Grevisia= sp.

Associate.--Notolampra antillarum, Trinidad (Princis and Kevan, 1955): One male only.

=Tillandsia fasciculata= Swartz

Associate.--Eurycotis floridana, Florida (Rehn and Hebard, 1914; Hebard, 1917).

=Tillandsia usneoides= Linnaeus

Associates.--Parcoblatta sp., Louisiana (Rainwater, 1941).

Latiblattella rehni, Florida (Blatchley, 1920): By beating.

Cockroaches, Louisiana (Rosenfeld, 1911, 1912): One mature and 39 immature blattids were collected from 8 of 12 samples of Spanish moss.

=Tillandsia uttriculata= Linnaeus

Associate.--Epilampra mona, Mona Island, West Indies (Rehn and Hebard, 1927): The type and one paratypic female of E. mona were collected in this bromeliad.

Eurycotis floridana, Florida (Blatchley, 1920).

=Tillandsia= sp.

Associates.--Aglaopteryx gemma, Texas (Hebard, 1917).

Dryadoblatta scotti, Trinidad (Scott, 1912): Found in the leaf bases.

=Undetermined bromeliads=

Associates.--Aglaopteryx diaphana, Jamaica (Hebard, 1917; Rehn and Hebard, 1927).

Anaplecta azteca and Anaplecta sp., Costa Rica (Picado, 1913).

Anaplecta mexicana, Costa Rica (Calvert and Calvert, 1917).

Audreia bromeliadarum, Panama (Caudell, 1914).

Audreia jamaicana, Jamaica (Rehn and Hebard, 1927).

Blattella sp., Costa Rica (Picado, 1913).

Buboblatta armata, Panama (Caudell, 1914): "Probably not a typical bromeliadicolous species."

Cariblatta insularis, Jamaica (Hebard, 1916a, 1917; Rehn and Hebard, 1927).

Cariblatta nebulicola, Jamaica (Rehn and Hebard, 1927); One immature male.

Dryadoblatta scotti, Trinidad (Princis and Kevan, 1955).

Epilampra conspersa, Dominica (Scott, 1912).

Epilampra maya, Panama (Hebard, 1920).

Epilampra sodalis, Panama (Caudell, 1914).

Epilampra sp. and Hormetica laevigata, Brazil (Hancock, 1926).

Eurycotis biolleyi, Costa Rica (Picado, 1913).

Ischnoptera rufa occidentalis, Mexico (Caudell, 1914).

Latiblattella chichimeca, Costa Rica (Picado, 1913).

Litopeltis biolleyi, Costa Rica (Rehn, 1928).

Litopeltis bispinosa, Panama (Caudell, 1914).

Neoblattella brunneriana, Costa Rica (Calvert and Calvert, 1917).

Neoblattella dryas, Neoblattella eurydice, Neoblattella grossbecki, and Neoblattella proserpina, Jamaica (Rehn and Hebard, 1927).

Neoblattella fratercula, Mexico (Hebard, 1921b).

Neoblattella nahua, Mexico (Caudell, 1914).

Nesomylacris relica, Jamaica (Rehn and Hebard, 1927).

Nyctibora brunnea(?), Panama (Caudell, 1914): According to Hebard (1920) Caudell's specimen was almost certainly incorrectly identified. It may have been Nyctibora noctivaga or a smaller species of the genus. Brazil (Hancock, 1926).

Nyctibora laevigata, Jamaica (Hebard, 1917; Rehn and Hebard, 1927).

Nyctibora lutzi, Puerto Rico (Rehn and Hebard, 1927): "in epiphytes with pencil-like leaves."

Pelmatosilpha rotundata, Panama (Caudell, 1914).

Pseudomops laticornis, Costa Rica (Picado, 1913).

Pycnoscelus surinamensis, Costa Rica (Picado, 1913). Mexico (Caudell, 1914). Jamaica (Rehn and Hebard, 1927).

"Rhicnoda" sp., Costa Rica (Picado, 1913). This genus is now recognized as not being in the New World fauna. Probably the specimen was a species of Epilampra or Hyporhicnoda as suggested by Gurney (personal communication, 1959) and confirmed by Rehn (p.c., 1959).

Cockroaches, Costa Rica (Calvert, 1910): Cockroaches were said to be common in bromeliads on the moist Atlantic slope.

Family LILIACEAE

=Yucca elata= Engelman

Associate.--Latiblattella lucifrons, Arizona (Ball et al., 1942).

=Easter lilies=

Associate.--Pycnoscelus surinamensis, Connecticut (Zappe, 1918).

Family MUSACEAE

=Bananas=

Cockroaches have been captured in bunches of bananas, in bracts of banana flowers, under banana leaves, and burrowing in rotten banana stalks. Although many of the species associated with bananas are indigenous to the banana-growing areas of the American Tropics, most of the specimens cited below were captured elsewhere as adventitious insects that had been imported with the fruit. It is obvious that many of these insects must have been closely associated with bananas on the plantations, where, undoubtedly, the growing plants provided attractive ecological niches. Bunting (1956) deduced, from the presence of healthy cockroaches on bananas allegedly sprayed with copper arsenate, that the insects did not feed on stems or fruit but hid among the bananas and foraged elsewhere; however, certain reports are of cockroaches actually feeding on bananas. Some of the records cited by Hebard (1917) were compiled from earlier reports not all of which we have seen. Numbers in parentheses following certain citations indicate the number of times the association had been observed. Known or suspected adventive material is so indicated.

Aglaopteryx diaphana, Jamaica (Rehn and Hebard, 1927): Found in bracts of banana blossoms. England (Bunting, 1955): Adventive, on bananas from Dominica.

Aglaopteryx vegeta, Finland (Princis, 1947): Adventive, in banana box.

Amazonina emarginata, Trinidad (Princis and Kevan, 1955): In banana bunch.

Archimandrita marmorata, Denmark (Henriksen, 1939): Adventive (2), in bananas from Jamaica(?). As Princis (1947) and Gurney (personal communication, 1959) point out, this is a Central American species, so Jamaica may be an error.

Archimandrita tessellate, Sweden (Princis, 1947): Adventive, from Honduras.

Blaberus atropos(?), Denmark (Henriksen, 1939): Adventive, from Jamaica. Princis (1947) pointed out that this species was more likely to have been Blaberus craniifer or Blaberus discoidalis, which are West Indian species, than B. atropos which is a South American species.

Blaberus boliviensis, Ecuador (Princis, 1952): In a shipment of bananas from near Puna.

Blaberus discoidalis, Puerto Rico (Rehn and Hebard, 1927): From banana ripening room. Great Britain (Pearce, 1929): Adventive. England (Bunting, 1955, 1956): Adventive, from Dominica.

Capucinella delicatula, California (Caudell, 1931): Adventive.

Cariblatta delicatula, Cuba (Rehn and Hebard, 1927).

Cariblatta hylaea, Honduras (Rehn, 1945a): Shaken from hanging dead banana leaves.

Cariblatta insularis, Finland (Frey, 1948): Adventive.

Cariblatta landalei, Jamaica (Rehn and Hebard, 1927): All specimens taken from under drying bracts of banana blossoms.

Cariblatta punctipennis and Chorisoneura barbadensis, England (Bunting, 1956): Adventive, from Dominica.

Epilampra abdomen-nigrum and Epilampra sp., England (Bunting, 1955): Adventive, from Dominica.

Epilampra maya, Massachusetts (Hebard, 1917): Adventive.

Epilampra mexicana(?), Denmark (Henriksen, 1939): Adventive (2), from Danish West Indies. Princis (1947) suggested that this should be Epilampra sp., because E. mexicana is not a West Indian species.

Eudromiella calcarata and Eurycotis bananae, U.S.S.R., Leningrad (Bei-Bienko, 1947): Adventive, from Colombia.

Euphyllodromia angustata, Sweden (Princis, 1947): Adventive.

Eurycotis caraibea, New York (Hebard, 1917): Adventive.

Eurycotis dimidiata, Washington, D. C. (Caudell, 1931): Adventive.

Eurycotis lixa, New York (Rehn, 1930): Adventive, on banana ship from Jamaica.

Graptoblatta notulata, Marquesas Islands, Uahuka (Hebard, 1933a): In banana leaves.

Holocompsa nitidula, Trinidad (Princis and Kevan, 1955): Eating banana pulp.

Hormetica laevigata, Wales (Sandemann, 1934): Adventive, in pile of banana sacks.

Hormetica ventralis, Sweden (Princis, 1947): Adventive, in local warehouse of banana company.

Hormetica spp., Europe and North America (Bei-Bienko, 1950): Adventive, introduced with bananas and other tropical fruits.

Ischnoptera rufa rufa, Puerto Rico (Wolcott, 1950): Brought into houses on bunches of bananas.

Kuchinga remota, Society Islands, Moorea (Hebard, 1933a): In dead banana leaves.

Lamproblatta albipalpus, Panama Canal Zone (Hebard, 1920): Several under decayed banana stem.

Latiblattella sp., Finland (Frey, 1948): Adventive.

Leucophaea maderae, New York (Hebard, 1917): Adventive. Dominica (Rehn and Hebard, 1927): Under banana sheaths. England (Palmer, 1928): Adventive, captured at railroad station after bananas had been unloaded. England (Bunting, 1955): Adventive, from Dominica. Trinidad (Princis and Kevan, 1955): Nymph, eating bananas in cupboard. Puerto Rico (Seín, 1923): Seín stated that bananas are the favorite food of L. maderae.

Litopeltis bispinosa, Panama Canal Zone (Hebard, 1920): From rotting banana stalks at bases of leaves.

Litopeltis musarum, Costa Rica (Rehn, 1928): Shaken from dead banana leaves.

Nauclidas nigra, England (Bunting, 1955, 1956): Adventive, from Dominica.

Nauphoeta flexivitta, Denmark (Vestergaard, 1958): Adventive.

Neoblattella carcinus, Neoblattella celeripes, and Neoblattella laodamia, England (Bunting, 1956): Adventive, from Dominica. Bunting (1955) first reported these as Neoblattella spp. and stated that they were common.

Neoblattella detersa, Jamaica (Rehn and Hebard, 1927): From under the bracts of banana blossoms. Sweden (Princis, 1947): Adventive.

Neoblattella detersa and Neoblattella tridens, Finland (Frey, 1948): Adventive.

Neoblattella fratercula, Nebraska (Hebard, 1916b): Adventive.

Neoblattella semota, Jamaica (Rehn and Hebard, 1927): From under drying bracts of banana blossoms.

Neoblattella vatia, Cuba (Rehn and Hebard, 1927).

Neoblattella sp., Finland (Princis, 1947): Adventive, from Jamaica.

Nyctibora azteca, England (Bunting, 1955): Adventive, from Dominica. Bunting reported this species as Nocticola azteca. Dr. A. B. Gurney called our attention to the fact that Nocticola is an Old World genus, presumably combined in error with the New World species azteca. The true identity of the specimen was confirmed by Dr. D. Ragge (personal communication, 1958), who examined it at the British Museum (Natural History).

Nyctibora holoserica, Canada (Walker, 1912): Adventive.

Nyctibora laevigata, Canada, Maine, Massachusetts, Pennsylvania (2) (Hebard, 1917): Adventive. Taken from banana boat Annetta at Philadelphia (Rehn and Hebard, 1927). England (Bunting, 1956): Adventive, from Dominica. Sweden, Denmark (Princis, 1947): Adventive.

Nyctibora mexicana(?), Denmark (Henriksen, 1939): Adventive (5), from Jamaica and West Indies. Princis (1947) suggested that these specimens were probably the West Indian Nyctibora noctivaga, because N. mexicana is not a West Indian insect.

Nyctibora noctivaga, Canada, Idaho, Illinois, Massachusetts, Nebraska (4), Virginia (Hebard, 1917): Adventive. Nebraska (Hauke, 1949): Adventive (2). Panama Canal Zone (Hebard, 1920): From banana stalks. England (Blair in Turner, 1930): Adventive, from Costa Rica. Washington (Hatch, 1938): Adventive. Sweden (Princis, 1947): Adventive (2). Finland (Princis, 1947): Adventive, from Jamaica.

Nyctibora obscura, Trinidad (Princis and Kevan, 1955): In banana bunch.

Nyctibora sericea, Canada (Stevenson, 1905; Walker, 1912): Adventive; Hebard (1917) synonymized Walker's specimen under N. laevigata. Isle of Wight (Meade-Waldo, 1910): Adventive, from Jamaica. England (Tulloch, 1939): Adventive, in banana crates from Brazil.

Nyctibora sp., England (Welch, 1935): Adventive, in railway truck that had carried bananas. England (Tulloch, 1939): Adventive, from Brazil.

Oxyhaloa deusta, U.S.S.R., Leningrad (Bei-Bienko, 1947): Adventive, from Colombia.

Panchlora antillarum, England (Bunting, 1955): Adventive, from Dominica.

Panchlora exoleta, Scotland (Distant, 1902): Adventive. Great Britain (Shaw, 1902): Adventive. England (Coney, 1918): Adventive. Sweden, Norway (Princis, 1947): Adventive, Norwegian specimen from Brazil. Germany (Zacher, 1917): Adventive, from Jamaica.

Panchlora nivea, Colorado, Nebraska, New Jersey, New York (2), Utah (Hebard, 1917): Adventive. Nebraska (Hauke, 1949): Adventive. Washington (Hatch, 1938): Adventive. Massachusetts (Roth and Willis, 1958): Adventive. England (Bunting, 1955): Adventive, from Dominica. U.S.S.R. (Bei-Bienko, 1947): Adventive, from Colombia. Sweden (13), Norway (3), Finland (3) (Princis, 1947): Adventive, mostly females; origin (where known) Jamaica.

Panchlora fraterna(?) and Panchlora peruana(?), Denmark (Henriksen, 1939): Adventive; origin (where known) Danish West Indies and Jamaica; Princis (1947) suggested that both species were probably Panchlora nivea.

Panchlora sagax, Puerto Rico (Wolcott, 1936).

Panchlora virescens, Canada (Walker, 1912): Adventive; this was probably P. nivea as we now know it (Gurney, personal communication, 1959).

Panchlora sp., Canada (Walker, 1912): Adventive. England (Tulloch, 1939): Adventive, from Brazil.

Pelmatosilpha coriacea, Puerto Rico (Wolcott, 1936).

Pelmatosilpha marginalis and Pelmatosilpha purpurascens, England (Bunting, 1955, 1956): Adventive, from Dominica; both species common.

Pelmatosilpha vagabunda, New Zealand (Princis, 1954): Adventive, probably from South America.

Periplaneta americana, Belgium (Schepdael, 1931): Adventive, on bananas from the American Tropics.

Periplaneta americana and Periplaneta brunnea, England (Bunting, 1955, 1956): Adventive, from Dominica.

Periplaneta americana and Periplaneta australasiae, England (Watson, 1907): Adventive; they ate ripening bananas in the tropical plant house of the Royal Botanic Gardens, Kew, where they hid in "the sheathing bases of palm, banana and pandanus leaves." Sweden (Princis, 1947): Adventive.

Periplaneta australasiae, Canada (Walker, 1912): Adventive. Denmark (Henriksen, 1939): Adventive (9); origin mostly Jamaica. England (Tulloch, 1939): Adventive, from Brazil. England (Bunting, 1955, 1956): Adventive, from Dominica; common.

Platyzosteria bifida, Nebraska (Hebard, 1917): Adventive.

Plectoptera dorsalis, Puerto Rico (Rehn and Hebard, 1927): Captured by beating banana plants.

Pycnoscelus surinamensis, Canada (Walker, 1912; Hebard, 1917): Adventive: Marquesas Islands, Nukuhiva (Hebard, 1933a): In banana leaves. England (Goodliffe, 1958): Adventive, doing considerable damage to banana plants growing in a conservatory.

Sibylloblatta panesthoides, Massachusetts (Rehn, 1937a): Adventive, from Jamaica.

Family ZINGIBERACEAE

=Renealmia= sp.

Associate.--Cariblatta orestera, Jamaica (Rehn and Hebard, 1927): The male was taken in a head of wild ginger.

Family CANNACEAE

=Canna= sp.

Associate.--Periplaneta americana, Hawaii (Zimmerman, 1948).

Family ORCHIDACEAE

=Cattleya= sp.

Associates.--Periplaneta americana, U.S.A. (Rau, 1940a).

Periplaneta australasiae, England (Lucas, 1918).

=Vanda= sp.

Associates.--Periplaneta americana, U.S.A. (Rau, 1940a).

Periplaneta australasiae, England (Lucas, 1918).

=Undetermined orchids=

Associates.--Blaberus discoidalis, Blatta orientalis, Periplaneta americana, Hawaii (Swezey, 1945).

Blatta orientalis, americana, cinerea, maderae, unidentified cockroaches, England, in bulb from Ecuador (Westwood, 1876).

Graptoblatta notulata, Hawaii (Swezey, 1945): On orchid from India.

Homalopteryx laminata and Hormetica apolinari, New York (Hebard, 1912c): In orchids shipped from Colombia.

Pelmatosilpha coriacea, Puerto Rico (Wolcott, 1936).

Periplaneta americana, Germany (Tashenberg, 1884).

Periplaneta australasiae, England (Wainwright, 1898). Pennsylvania (Skinner, 1905). Massachusetts (Morse, 1920).

Pycnoscelus surinamensis, England (Westwood, 1869). Germany (Zacher, 1920). Massachusetts (Morse, 1920). Hawaii (Swezey, 1945).

Family CASUARINACEAE

=Casuarina= sp.

Associate.--Diploptera punctata, Hawaii (Zimmerman, 1948).

Family SALICACEAE

=Populus euphratica= Oliv.

Synonymy.--Populus diversifolia Schrenk. [Howard, personal communication, 1959].

Associate.--Ectobius semenovi, Kazakhstan (Bei-Bienko, 1950).

=Populus= sp.

Associate.--Ectobius lapponicus, U.S.S.R. (Stark in Bei-Bienko, 1950): On aspen.

=Salix= sp.

Associate.--Ectobius semenovi, Kazakhstan (Bei-Bienko, 1950): On willow.

Family MYRICACEAE

=Myrica cerifera= Linnaeus

Associate.--Chorisoneura texensis, Florida (Rehn and Hebard, 1916): On bayberry. Florida (Blatchley, 1920): Beaten from foliage.

Family FAGACEAE

=Quercus alba= Linnaeus

Associate.--Parcoblatta pensylvanica, Virginia (Rehn and Hebard, 1916): Under signs on white oaks.

=Quercus rubra= Linnaeus

Associates.--Parcoblatta divisa, Virginia, and Parcoblatta pensylvanica, North Carolina (Rehn and Hebard, 1916): Under signs on red oak.

Parcoblatta lata, North Carolina (Hebard, 1917): Under sign.

=Quercus virginiana= Mill.

Associate.--Eurycotis floridana, Georgia (Rehn and Hebard, 1916): Under dead bark on live-oak tree. Georgia (Hebard, 1917): In cavity in tree.

=Quercus= spp.

Associates.--Aglaopteryx gemma, Alabama, Georgia, Florida, Louisiana, Texas (Hebard, 1917): Under signs on oaks.

Blatta orientalis, England (Donisthorpe, 1918): Under bark.

Cariblatta lutea lutea, Mississippi (Hebard, 1916a): By beating low oaks on hills.

Chorisoneura texensis, Mississippi (Hebard, 1917). Florida (Blatchley, 1920): By beating.

Ectobius pallidus, England (Milton, 1899; Burr, 1899b). Massachusetts (Flint, 1951): Under loose lichens and bark.

Parcoblatta divisa, Georgia, Louisiana, and Parcoblatta pensylvanica, Georgia (Hebard, 1917): Under signs.

Periplaneta australasiae, Florida (Rehn and Hebard, 1905): Ten specimens taken from under a tin sign.

Periplaneta brunnea, Georgia (Rehn and Hebard, 1916): Under signs.

Phyllodromica megerlei, U.S.S.R. (Bei-Bienko, 1950): By shaking oak branches.

Plectoptera lacerna, Cuba (Rehn and Hebard, 1927).

Family MORACEAE

=Cecropia= sp.

Associate.--Cariblatta hylaea, Honduras (Rehn, 1945a).

Family CHENOPODIACEAE

=Beta maritima= Linnaeus

Associate.--Ectobius panzeri, England (Lucas, 1920a).

=Beta vulgaris= var. =cicla= Linnaeus

Associate.--Ectobius pallidus, Massachusetts (Flint, 1951): Many specimens collected in the bases of Swiss chard plants.

Family LAURACEAE

=Nectandra coriacea= (Sw.) Griseb.

Synonymy.--Ocotea catesbyana Sarg. [Howard, personal communication, 1959].

Associate.--Chorisoneura texensis, Florida (Rehn and Hebard, 1912).

Family SARRACENIACEAE

Only a few records have been found of cockroaches being trapped in the pitchers of carnivorous plants of this and the following family. The insects drown in the fluid within the pitcher where they are apparently digested by proteinases secreted by the plant (Meyer and Anderson, 1939; Lloyd, 1942).

=Sarracenia flava= Linnaeus

Natural prey--Cariblatta lutea lutea, Ischnoptera deropeltiformis, Parcoblatta lata, and nymphs of Parcoblatta sp., North Carolina (Wray and Brimley, 1943): Most of the cockroaches seemed to have been trapped accidentally with the possible exception of C. lutea lutea, 11 of which were found in Sarracenia pitchers.

=Sarracenia purpurea= Linnaeus

Natural prey.--Cariblatta lutea lutea, North Carolina (Wray and Brimley, 1943).

=Sarracenia minor= Walter

Synonymy.--Sarracenia variolaris Michx. [Howard, personal communication, 1958].

Natural and experimental prey.--Periplaneta australasiae, Florida (Treat, 1876): After the insect imbibed some of the fluid in the pitcher it became docile; others became highly active and rushed wildly about before becoming quiescent. See also Treat in Scudder (1877).

Cockroaches, U.S.A. (Riley, 1875).

Family NEPENTHACEAE

=Nepenthes ampularia= Jack

Natural prey.--Cockroaches, Singapore (Dover, 1928).

=Nepenthes gracilis= Korth.

Natural prey.--Cockroaches, Singapore (Dover, 1928).

=Nepenthes= sp.

Natural prey.--Cockroach, Old World Tropics? (Hooker, 1874): The insect was apparently attracted into the pitcher, where it drowned, by a piece of cartilage placed there by Hooker.

Family CUNONIACEAE

=Weinmannia= sp.

Associates.--Aneurina viridis, Marquesas Islands, Nukuhiva and Fatuhiva (Hebard, 1933a).

Maretina uahuka, Marquesas Islands, Uahuka (Hebard, 1933a).

Family HAMAMELIDACEAE

=Liquidambar styraciflua= Linnaeus

Associate.--Parcoblatta divisa, Georgia (Rehn and Hebard, 1916): Under sign on sweet gum.

Parcoblatta zebra, Louisiana (Hebard, 1917): In decay cavity.

Family ROSACEAE

=Crataegus= sp.?

=Associates.=--Plectoptera dorsalis, Plectoptera infulata, Plectoptera rhabdota, Puerto Rico (Wolcott, 1950): In the dry flower clusters of "espino rubial."

=Rosa= sp.

Associate.--Pycnoscelus surinamensis, Connecticut (Zappe, 1918); Rhode Island and Pennsylvania (Caudell, 1925); Pennsylvania (Doucette and Smith, 1926): Feeding on canes in greenhouses.

=Rubus= spp.?

Associate.--Hololampra chavesi, Azores (Chopard, 1932): This species is exclusively dendricolous and was found only by beating the bushes on which it abounds. It was very common in hedges, particularly on brambles (ronces).

Family LEGUMINOSAE

=Acacia farnesiana= Willd.

Associate.--Diploptera punctata, Hawaii (Bridwell and Swezey, 1915; Zimmerman, 1948): Feeding on pods.

=Acacia= sp.

Associates.--Ellipsidion australe, Australia, New South Wales (Hebard, 1943).

Methana curvigera, Australia, Queensland (Pope, 1953a).

=Ceratonia siliqua= Linnaeus

Associate.--Diploptera punctata, Hawaii (Pemberton and Williams, 1938): Damaging algarroba.

=Erythrina glauca= Willd.

Associates.--Aglaopteryx absimilis, Puerto Rico (Wolcott, 1950): In abandoned cocoon.

Aglaopteryx facies, Puerto Rico (Wolcott, 1936): In empty cocoons.

=Inga laurina= Willd.

Associate.--Aglaopteryx facies, Puerto Rico (Wolcott, 1936): On trunk.

=Inga vera= Willd.

Associates.--Aglaopteryx facies, Puerto Rico (Wolcott, 1936): In larval tents.

Cariblatta stenophrys, Puerto Rico (Wolcott, 1936): On leaves.

Plectoptera dorsalis, Plectoptera infulata, and Plectoptera rhabdota, Puerto Rico (Wolcott, 1950): In "butterfly nests" in leaves.

=Mesquite=

Associate.--Nyctibora stygia, Haiti (Rehn and Hebard, 1927).

=Samanea saman= Merr.

Associate.--Aglaopteryx absimilis, Puerto Rico (Wolcott, 1950).

=Tamarindus indica= Linnaeus

Associate.--Hemiblabera brunneri, Puerto Rico (Rehn and Hebard, 1927).

Family GERANIACEAE

=Geraniums=

Associate.--Diploptera punctata, Hawaii (Zimmerman, 1948).

Family ZYGOPHYLLACEAE

=Tribulus= sp.

Associates.--Periplaneta americana and Pycnoscelus surinamensis, Johnston Island (Bryan, 1926). Zimmerman (1948) lists Tribulus as a host plant for these cockroaches.

Family RUTACEAE

=Citrus aurantifolia= Swingle

Associates.--Plectoptera dominicae and Plectoptera perscita, Dominica (Rehn and Hebard, 1927): Beaten from moss-covered lime trees.

=Citrus maxima= Merr.

Associates.--Plectoptera dorsalis, Plectoptera infulata, Plectoptera rhabdota, Puerto Rico (Wolcott, 1950).

Plectoptera rhabdota, Puerto Rico (Rehn and Hebard, 1927).

=Citrus sinensis= Osbeck

Associate.--Diploptera punctata, Hawaii (Bridwell and Swezey, 1915; Zimmerman, 1948): Feeding on oranges on tree.

=Citrus= sp.

Associates.--Diploptera punctata, Hawaii (Zimmerman, 1948).

Riatia [=Lissoblatta] fulgida, Panama, Rio Trinidad (Hebard, 1920).

Plectoptera porcellana, Puerto Rico (Sein, 1923).

=Zanthoxylum caribaeum= Lam.

Associates.--Plectoptera dorsalis, Plectoptera infulata, Plectoptera rhabdota, Puerto Rico (Wolcott, 1950): In the dry flower clusters.

Family BURSERACEAE

=Bursera simaruba= (L.) Sarg.

Associate.--Chorisoneura texensis, Florida (Rehn and Hebard, 1912; Hebard, 1917): Beaten from the lower branches of gumbo limbo.

Family EUPHORBIACEAE

=Poinsettia= sp.

Associate.--Pycnoscelus surinamensis, Connecticut (Zappe, 1918): Ate bark of greenhouse plants.

Family ANACARDIACEAE

=Mangifera indica= Linnaeus

Associate.--Diploptera punctata, Hawaii (Bridwell and Swezey, 1915; Zimmerman, 1948): Feeding on mangoes on the tree.

=Spondias mombin= Linnaeus

Associates.--Plectoptera dorsalis, Plectoptera infulata, Plectoptera rhabdota, Puerto Rico (Wolcott, 1950): Living on leaves of "jobo."

=Spondias purpurea= Linnaeus

Associate.--Eurycotis biolleyi, Costa Rica (Rehn, 1918): In the crown of dry jocoto.

Family AQUIFOLIACEAE

=Ilex cassine= Linnaeus

Associate.--Plectoptera poeyi, Florida (Rehn and Hebard, 1912, 1914; Hebard, 1917).

=Ilex coriacea= (Pursh) Chapm.

Synonymy.--Ilex lucida [Fernald, 1950].

Associate.--Cariblatta lutea lutea, Florida (Hebard, 1916a).

Family SAPINDACEAE

=Exothea paniculata= (Juss.) Radlk.

Associate.--Aglaopteryx gemma, Florida (Hebard, 1917).

Family MALVACEAE

=Gossypium= spp.

Associates.--Graptoblatta notulata, Marquesas Islands, Tahuata (Hebard, 1933a).

Periplaneta australasiae, St. Kitts, B.W.I. (Ballou, 1916).

Periplaneta fuliginosa and Plectoptera poeyi, Florida (Rainwater, 1941).

Plectoptera dorsalis, Plectoptera infulata, Plectoptera rhabdota, Puerto Rico (Wolcott, 1950).

=Hibiscus rosa-sinensis= Linnaeus

Associate.--Riatia orientis, Trinidad (Princis and Kevan, 1955).

=Hibiscus= sp.

Associates.--Amazonina emarginata, Cariblatta antiguensis, Eurycotis kevani, and Rhytidometopum dissimile, Trinidad (Princis and Kevan, 1955).

=Sida= sp.

Associate.--Periplaneta australasiae, Nihoa Island (Bryan, 1926). Hawaii (Zimmerman, 1948).

Family STERCULIACEAE

=Theobroma cacao= Linnaeus

Associate.--Ceratinoptera picta, Trinidad (Princis and Kevan, 1955).

Family BIXACEAE

=Bixa= sp.

Associate.--Notolampra antillarum, Trinidad (Princis and Kevan, 1955): Nymphs in dry fruits on "annato" tree.

Family FLACOURTIACEAE

=Xylosma suaveolens= Forst.

Associate.--Graptoblatta notulata, Marquesas Islands, Uahuka (Hebard, 1933a).

Family PASSIFLORACEAE

=Passiflora= sp.

Associate.--Aristiger [=Plumiger] histrio, Malaya (Karny, 1924).

Family CARICACEAE

=Carica papaya= Linnaeus

Associate.--Diploptera punctata, Hawaii (Bridwell and Swezey, 1915; Zimmerman, 1948): Feeding on papaya fruit on tree.

Family RHIZOPHORACEAE

=Rhizophora mangle= Linnaeus

Associate.--Aglaopteryx gemma, Florida (Hebard, 1917).

Family COMBRETACEAE

=Conocarpus erectus= Linnaeus

Associate.--Plectoptera poeyi, Florida (Rehn and Hebard, 1914): Running on leaves.

Family MYRTACEAE

=Eucalyptus= sp.

Associate.--Ellipsidion australe, Australia, New South Wales (Hebard, 1943).

=Eugenia aromatica= Baill.

Synonymy.--Syzygium aromaticum [Bailey, 1925].

Associate.--Plectoptera dorsalis, Puerto Rico (Wolcott, 1936): On flowers of "pomarrosa."

=Metrosideros collina= Gray

Associates.--Aneurina viridis, Marquesas Islands: Nukuhiva, Fatuhiva, and Tahuata (Hebard, 1933a)

Aneurina tahuata, Marquesas Islands, Tahuata (Hebard, 1933a).

Graptoblatta notulata, Marquesas Islands, Nukuhiva (Hebard, 1933a).

=Psidium guajava= Linnaeus

Associate.--Plectoptera rhabdota, Puerto Rico (Rehn and Hebard, 1927).

Family ONAGRACEAE

=Jussiaea natans= Humb. and Bonpl.

Associate.--Epilampra abdomen-nigrum, Panama (Crowell, 1946): In an aquarium the cockroach fed on leaves of this aquatic plant which had been collected in the lagoon where the insect was captured.

Family ERICACEAE

=Calluna vulgaris= Salisb.

Associates.--Ectobius lapponicus, England (Lucas, 1925): "Nymphs of varying size were beaten out of heather ... on 9 February and later."

Ectobius panzeri, England (Lucas, 1927): "numerous imagines of both sexes were swept from heather."

=Vaccinium meridionale= Sw.

Associates.--Chorisoneura formosella, Neoblattella dryas, Neoblattella proserpina, Jamaica (Rehn and Hebard, 1927).

Family SAPOTACEAE

=Sideroxylon foetidissimum= Jacq.

Associate.--Pelmatosilpha coriacea, Puerto Rico (Wolcott, 1941): Under bark.

Family APOCYNACEAE

=Vinca minor= Linnaeus

Associate.--Ectobius pallidus, Massachusetts (Willis, unpublished observation, 1958).

Family CONVOLVULACEAE

=Ipomoea tiliasea= Choisy

Associate.--Plectoptera dorsalis, Puerto Rico (Rehn and Hebard, 1927).

Family BORAGINACEAE

=Cordia dentata= Poiret

Synonymy.--Calyptracordia alba [Howard, personal communication, 1958].

Associates.--Cariblatta antiguensis, Ischnoptera rufa rufa, Supella supellectilium, Symploce ruficollis and Symploce hospes, St. Croix, Virgin Islands (Beatty, 1944): On fruits of C. dentata except S. supellectilium which was found at night on the flowers.

Family VERBENACEAE

=Citharexylum villosum= Jacq.

Associate.--Chorisoneura texensis, Florida (Rehn and Hebard, 1912).

Family SOLANACEAE

=Nicotiana= sp.

Associate.--Pycnoscelus surinamensis, Sumatra (Roeser, 1940).

=Solanum tuberosum= Linnaeus

Associate.--Pycnoscelus surinamensis, Haiti (Hoffman, 1927): Feeding on tubers in field.

Family GESNERIACEAE

=Cyrtandra= sp.

Associate.--Aneurina viridis, Marquesas Islands, Nukuhiva (Hebard, 1933a).

Family RUBIACEAE

=Canthium barbatum= (Forst.) Seem.

Associate.--Graptoblatta notulata, Marquesas Islands, Uahuka (Hebard, 1933a).

=Cinchona pubescens= Vahl.

Associate.--Periplaneta americana, Puerto Rico (Plank and Winters, 1949): In greenhouse.

=Coffea= sp.

Associate.--Plectoptera porcellana, Puerto Rico (Seín, 1923).

Family COMPOSITAE

=Goldenrod=

Associate.--Eurycotis floridana, Florida (Hebard, 1917): "Climbing about on top of goldenrod at night."

=Helianthus= sp.

Associate.--Pseudomops septentrionalis, Texas (Hebard, 1917).

=Scorzonera acanthoclada= Franch.

Associate.--Phyllodromica tartara nigrescens, Southern Uzbekistan (Bei-Bienko, 1950): On the flowers.

DAMAGE TO PLANTS BY COCKROACHES

Cockroaches characteristically feed on dead plant and animal material. Damage to living plants occurs principally in the Tropics or under subtropical conditions in greenhouses in temperate regions. Among the depredations attributed to cockroaches in text books, damage to plants is seldom emphasized. This is surprising in view of the many records cited below.

Capt. William Bligh (1792), while collecting breadfruit trees in Tahiti to take to the West Indies, wrote in his log during January 1789: "This morning, I ordered all the chests to be taken on shore, and the inside of the ship to be washed with boiling water, to kill the cockroaches. We were constantly obliged to be at great pains to keep the ship clear of vermin, on account of the plants."

Westwood (1869) stated that Pycnoscelus surinamensis was very destructive in orchid houses feeding on buds and young shoots. Later Westwood (1876) exhibited the bulb of an orchid from Ecuador which contained six species of cockroaches: Blatta orientalis, [Periplaneta?] americana, [Nauphoeta?] cinerea, [Leucophaea?] maderae, and two others unknown to him. Fullaway (1938) stated that cockroaches damage root tips, buds, and flowers of orchids. Periplaneta americana has been said to eat the root tips and blossoms of orchids (Taschenberg, 1884) and to devour the open flower petals of Cattleya orchids as well as the aerial roots and flower spikes of Vanda orchids (Rau, 1940a). Wainwright (1898) stated that Periplaneta australasiae had been observed in an orchid house in Perthshire where over a period of three years it had caused a good deal of damage. Skinner (1905) reported that P. australasiae in greenhouses in Pennsylvania showed no preference for any one plant but ate both plants and flowers of orchids, roses, and carnations. Lucas (1918) received specimens of P. australasiae which had played havoc with orchids especially Cattleya and Vanda. Morse (1920) reported that both P. australasiae and Pycnoscelus surinamensis were obnoxious in a conservatory in Massachusetts where they gnawed the tips of the aerial roots of orchids. Swezey (1945) in Hawaii stated that the following cockroaches have been reported as occasional minor pests on orchids: Blatta orientalis, Blaberus discoidalis, P. americana, and P. surinamensis; he further stated that Graptoblatta notulata had been intercepted at Honolulu on orchids from India.

Watson (1907) stated that Blatta orientalis, Periplaneta americana, and Periplaneta australasiae were injurious in the tropical plant houses at Kew: "at night they come out and run or fly about among the plants, devouring flowers and leaves like rabbits. Such plants as Eucharis, Crinum and Alpinia, when in flower, have little chance in the palm house, where the cockroaches are most abundant; they also find out the ripening bananas and soon devour them." Raffill (1910) stated that in plant houses in England B. orientalis, P. americana, and P. australasiae commonly, and Nauphoeta cinerea, Nauphoeta flexivitta, and Pycnoscelus surinamensis more rarely, are extremely destructive to plants. Flowers having a strong perfume, such as orchids, Eucharis, Crinum, and Hedychium, were often attacked while other flowers nearby were left uninjured.

Plank and Winters (1949) reported that in Puerto Rico the species of Orthoptera most injurious under greenhouse conditions was Periplaneta americana. Large nymphs destroyed 25 to 30 percent of freshly planted seed of Cinchona pubescens. In Hawaii the host plants of P. americana are blossoms of Canna and Tribulus, and the host plants of Periplaneta australasiae are Pritchardia and Sida (Zimmerman, 1948). On St. Kitts, B.W.I., young cotton plants were severely attacked by P. australasiae; this caused loss of the stand on a considerable area and necessitated replanting (Ballou, 1916). P. australasiae was reported damaging the Polystichum aristatum Presl [=Lastrea aristata variegata] in a greenhouse (Thilow and Riley, 1891). Laing (1946; British Museum [Natural History], 1951) stated that P. australasiae abounds in greenhouses and forcing pits where it may do considerable damage to the plants. Periplaneta fuliginosa is also troublesome in greenhouses because of its tendency to feed on seedlings and succulent plants (Dodge and Rickett, 1943).

Ectobius lapponicus has been observed feeding in galleries in the thick skin of young aspen in 25 percent of the trees examined (Stark in Bei-Bienko, 1950). The aquatic cockroach Epilampra abdomen nigrum fed on the leaves of Jussiaea natans in an aquarium (Crowell, 1946). Ischnoptera deropeltiformis has been taken while it was feeding on a fleshy fungus (Agaricus sp.) in dense woods in Indiana (Blatchley, 1920).

Diploptera punctata, the cypress roach or beetle roach, has been found in Hawaii feeding on ripening mangoes and papayas, oranges on the tree, and the outer covering of the pods of Acacia farnesiana (Bridwell and Swezey, 1915). Pemberton (1934) stated that D. punctata "disfigures our cypress trees by eating the bark from the young branches, often giving them a dead appearance over much of their leaf area." Fullaway and Krauss (1945) added, "This injury [to cypress] is so severe that sometimes areas of leaves die and turn brown. The Japanese cedar, ironwood, citrus and algaroba (kiawe) trees are attacked in a similar manner." Similar injury to cypress was described by Hebard (1922). In addition to girdling Cupressus, D. punctata injures Cryptomeria in the same fashion and also attacks algaroba, lime, and other plants (Pemberton and Williams, 1938). Zimmerman (1948) cited the following host plants for D. punctata in Hawaii: "Cupressus macrocarpa, Casuarina, Cryptomeria, Citrus, geraniums, Acacia farnesiana pods, mango fruits, orange fruits, papaya fruits."

In the reports of damage to plants by cockroaches, Pycnoscelus surinamensis has been implicated most often. This species is undoubtedly one of the economically most important cockroaches, being the vector of the chicken eyeworm as well as feeding on plants. In addition to the few reports of damage caused by this species that have already been mentioned, P. surinamensis has been reported to be very destructive in New Orleans to palms and ferns, attacking large alsophilas avidly, eating out the hearts (Anonymous, 1893). Zappe (1918) in Connecticut reported damage in a greenhouse to roses valued, at that time, at several hundred dollars; P. surinamensis had girdled the rose bushes, done much damage to Easter lilies, and in another greenhouse had eaten the bark from the stems of poinsettias. In Germany this species bit off the tips of the aerial roots and ate the petals of orchids (Zacher, 1920). Lucas (1923) reported damage to cucumber plants in a greenhouse in Surrey. Damage by P. surinamensis to the stems of rose bushes has been reported in Rhode Island and Pennsylvania; the canes were attacked both under and above ground (Caudell, 1925). Doucette and Smith (1926) reported a heavy infestation of P. surinamensis in a range of greenhouses in Philadelphia: "The roaches were present literally by the millions.... Although the roaches had been observed in cabinets and trash barrels for several months, it was not until the manager had occasion to go through the house one evening that he discovered that roaches were the cause of the troubles previously attributed to soil condition, watering, fungus, and other agencies.... About 30,000 to 35,000 rose plants from a total of 200,000 in the three more heavily infested houses were so badly injured by the gnawing off of the bark, young buds, and shoots of the main stems, that they were not in condition to be kept in the beds for another season."

In Haiti Pycnoscelus surinamensis damaged the tubers of growing potatoes (Hoffman, 1927). Illingworth (1927, 1929) reported that in Hawaii P. surinamensis was a minor pest of pineapples, feeding on the roots. This species was very plentiful in a propagating pit in England where it did much damage to various seeds and seedlings (Lucas, 1930). Roeser (1940) summarized some of the above-mentioned damage caused by P. surinamensis and added damage to chrysanthemums in Hawaii and tobacco in Sumatra where this cockroach destroyed 300,000 plants in a few days. Roeser was of the opinion that living plants were eaten only as a substitute when the earth became poor in food material. Zimmerman (1948) listed as host plants of P. surinamensis in Hawaii: "blossoms of Tribulus; reported feeding at roots of pineapples, and unconfirmed reports of damage to underground parts of some other plants." Goodliffe (1958) reported damage by this species to banana plants in a conservatory in northern England. Cohic (1956) implied that in New Caledonia "Racines de légumes" were attacked by P. surinamensis and that Zea mays Linnaeus was attacked by Supella supellectilium. Wolcott (1924a) reported that P. surinamensis damaged transplanted tobacco plants in Puerto Rico by eating the interior of the stalks. Tobacco planters in Cuba consider P. surinamensis injurious to the roots of tobacco plants (Bruner and Scaramuzza, 1936); this belief was confirmed in the laboratory, where adults and nymphs destroyed the roots and stems of tobacco plants two inches high and ate into the edges of the leaves. Dammerman (1929) reported that in Malaya this species often appeared in large numbers in gardens where it gnawed at the underground parts of vegetables and ornamental plants. Lever (1947) listed it as a pest on the leaves of pineapple.

Blattella vaga may occasionally damage seedlings in the laboratory (Flock, 1941a), but no damage has been reported in the field (Ball et al., 1942). Heer (1864) reported receiving a shipment of cycads from Cuba with all stages of Periplaneta americana living in holes in the branches, apparently subsisting on the starchy tissues. Goldenberg (1877) stated that sago trees provide cockroaches with their favorite nourishment. Scudder (1879) found Eurycotis floridana living in the tops of the cabbage palmetto, on which he presumed it fed. Parcoblatta americana has been observed feeding on an apple 6 feet above ground (Fulton, 1930).

X. PROTOZOA ASSOCIATED WITH COCKROACHES

The classification of the Protozoa follows that of Kudo (1954). The use of the asterisk (*) is explained in footnote 3, page 4.

=Phylum PROTOZOA=

Class MASTIGOPHORA

Order EUGLENOIDINA

Family EUGLENIDAE

=Euglena= sp.

Experimental host.--Periplaneta americana, U.S.A. (Hegner, 1929): When fed to the insects in concentrated culture, Euglena could withstand conditions in the crop up to 5 hours and were passed into the stomach in a viable state up to 6 hours. However, the majority were killed in the crop within 2 hours and very few reached the stomach alive.

Order PROTOMONADINA

Family OIKOMONADIDAE

=Oikomonas blattarum Tejera=

Natural host.--Cockroach, Venezuela (Tejera, 1926).

=Oikomonas= sp.

Natural host.--Blatta orientalis, U.S.S.R. (Yakimov and Miller, 1922): Oikomonas sp. and Monas sp. were found in the intestines of 83 percent of 124 cockroaches.

Cockroach, Venezuela (Tejera, 1926).

Family TRYPANOSOMATIDAE

=Leptomonas blaberae= Tejera

Natural host.--Blaberus sp., Venezuela (Tejera, 1926).

=Leptomonas= sp.

Natural hosts.--Parcoblatta lata, Parcoblatta pensylvanica, Parcoblatta virginica, U.S.A., Ohio (Semans, 1939, 1941): Hind intestine. Of 70 specimens examined, 86 percent harbored Leptomonas sp.

* =Herpetomonas periplanetae= Laveran and Franchini

Natural host.--Blatta orientalis, Italy, France (Laveran and Franchini, 1920, 1920a).

Family MONADIDAE

=Monas= sp.

Natural host.--Blatta orientalis, U.S.S.R. (Yakimov and Miller, 1922): Monas sp. and Oikomonas sp. were found in the intestines of 83 percent of 124 cockroaches examined.

Cockroach, Venezuela (Tejera, 1926).

Family BODONIDAE

=Bodo blattae=

Natural host.--Blatta orientalis, England (Lankester, 1865).

=Bodo= sp.

Natural host.--Blattella germanica and/or Periplaneta americana, South Africa (Porter, 1930).

=Retortamonas blattae= (Bishop)

Synonymy.--Embadomonas blattae Bishop [Wenrich, 1932].

Natural host.--Blatta orientalis, England (Bishop, 1931): Hind intestine. The organism occurred in about 40 percent of the cockroaches examined. L. G. Feo (in Wenrich, 1932) successfully cultured this protozoan (fig. 2, F).

=Retortamonas= sp.?

Natural host.--Leucophaea maderae, Philippine Islands (Hegner and Chu, 1930).

Order POLYMASTIGINA

Family CHILOMASTIGIDAE

=Chilomastix mesnili= (Wenyon)

Experimental vectors.--Blatta orientalis and Periplaneta americana, South Africa (Porter, 1918): The cockroaches were fed human excrement that contained cysts of C. mesnili. The cysts passed unharmed through the insects' digestive tract. Rats became infected with this protozoan on eating food that had been contaminated with feces from these cockroaches.

Family POLYMASTIGIDAE

=Eutrichomastix= sp.

Synonymy.--Trichomastix [Kudo, 1954].

Natural host.--Blattella germanica and/or Periplaneta americana, South Africa (Porter, 1930).

=Monocercomonoides globus= Cleveland et al.

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934): Organism occurs in practically all hosts.

=Monocercomonoides melolonthae= (Grassi)

(Fig. 2, A)

Natural host.--Platyzosteria novae seelandiae, New Zealand (Laird, 1956): Found in the intestinal tracts of the adult cockroaches, and of other species of insects.

=Monocercomonoides orthopterorum= (Parisi)

Synonymy.--Trichomonas (Trichomastix) orthopterorum Parisi; Monocercomonas orthopterorum [Bělǎr, 1916]; Trichomastic orthopterum? [Zasukhin, 1930]; Monocercomonoides orthopterorum [Travis, 1932; Cleveland et al., 1934]; Retortamonas orthopterorum [Semans, 1943].

Natural hosts.--Blatta orientalis, Italy (Parisi, 1910); U.S.S.R. (Zasukhin, 1930).

Ectobius lapponicus, Italy (Parisi, 1910).

Periplaneta americana, Philippine Islands (Hegner and Chu, 1930).

"Küchenschaben," Austria (Bělǎr, 1916).

The protozoan is found in the hind gut. Zasukhin (1930) found the organism in 85 percent of over 3,000 B. orientalis. Parisi (1910) found the flagellate present in very large numbers.

=Monocercomonoides panesthiae= Kidder

Natural host.--Panesthia angustipennis, Philippine Islands (Kidder, 1937): In hind gut.

=Tetratrichomastix blattidarum= Young

Natural hosts.--Blatta orientalis, Blattella germanica, Periplaneta americana, U.S.A. (Young, 1935): The organism when present occurs in large numbers in the posterior part of the intestine near the anus. The protozoan was successfully cultivated in a hemoglobin-saline medium.

Family OXYMONADIDAE

=Oxymonas doroaxostylus= (Cleveland et al.)

Synonymy.--Saccinobaculus doroaxostylus Cleveland et al. [Cleveland, 1950].

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934).

=Oxymonas nana= Cleveland

Synonymy.--Saccinobaculus minor Cleveland et al. [Cleveland, 1950].

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934).

Family DINENYMPHIDAE

=Saccinobaculus ambloaxostylus= Cleveland et al.

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934).

=Saccinobaculus lata= Cleveland

Natural host.--Cryptocercus punctulatus, U.S.A. (Cleveland, 1950b): There are at least two other species of Saccinobaculus in C. punctulatus that have not been described.

=Notila proteus= Cleveland

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland, 1950b).

Family TRICHOMONADIDAE

* =Trichomonas hominis= (Davaine)

Experimental vectors.--Blatta orientalis, South Africa (Porter, 1918); Italy (Mariani and Besta, 1936).

Periplaneta americana, South Africa (Porter, 1918); U.S.A. (Hegner, 1928).

=Trichomonas= sp.

Natural vector.--Cockroach, Venezuela (Tejera, 1926): Organism found in digestive tract of the cockroach.

Family HEXAMITIDAE

=Hexamita cryptocerci= Cleveland et al.

Natural hosts.--Cryptocercus punctulatus, U.S.A. (Cleveland et al., 1934).

Panesthia angustipennis, Philippine Islands (Kidder, 1937).

=Hexamita periplanetae= (Bĕlăr)

Synonymy.--Octomitus periplanetae Bĕlăr [Kudo, 1954].

Natural hosts.--Blatta orientalis, U.S.S.R. (Zasukhin, 1930): Organism is found in the hind gut. Eighty-five percent of over 3,000 B. orientalis contained this organism.

Periplaneta americana, Philippine Islands (Hegner and Chu, 1930).

"Küchenschaben," Austria (Bĕlăr, 1916).

=Hexamita= sp.?

Natural host.--Leucophaea maderae, Philippine Islands (Hegner and Chu, 1930): The flagellates were present in large numbers.

* =Giardia intestinalis= (Lambl)

Experimental vectors.--Blatta orientalis, South Africa (Porter, 1918).

Blattella germanica, Brazil (Pessôa and Corrêa, 1927).

Eurycotis floridana, U.S.A. (Young, 1937).

Leucophaea maderae, Brazil (Pessôa and Corrêa, 1927).

Periplaneta americana, South Africa (Porter, 1918); Gold Coast Colony (Macfie, 1922); Brazil (Pessôa and Corrêa, 1927); U.S.A. (Young, 1937).

Periplaneta brunnea, U.S.A. (Young, 1937).

Cockroaches, Venezuela (Tejera, 1926); Argentina (Bacigalupo, in Tejera, 1926).

* =Giardia= sp.

Natural vectors.--Cockroaches, Venezuela (Tejera, 1926).

Order HYPERMASTIGINA

Family HOLOMASTIGOTIDAE

=Leptospironympha eupora= Cleveland et al.

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian area (Cleveland et al., 1934).

=Leptospironympha rudis= Cleveland et al.

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian area (Cleveland et al., 1934).

=Leptospironympha wachula= Cleveland et al.

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian area (Cleveland et al., 1934).

=Macrospironympha xylopletha= Cleveland et al.

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian area (Cleveland et al., 1934).

Family LOPHOMONADIDAE

=Lophomonas blattarum= Stein

Natural hosts.--Blatta orientalis, Czechoslovakia (Stein, 1860); Germany (Bütschli, 1878; Schubotz, 1905; Chen, 1933); U.S.A. (Leidy, 1879a; Kudo, 1922, 1925, 1926, 1926b; McAdow, 1931); Europe (Janicki, 1908); U.S.S.R. (Yakimov and Miller, 1922; Zasukhin, 1930); Poland (Lorenc, 1939).

Blattella germanica, U.S.A., Ohio (McAdow, 1931).

Blattella germanica and/or Periplaneta americana, Egypt (DeCoursey and Otto, 1956, 1957).

Periplaneta americana, England (Schuster, 1898); Europe (Janicki, 1910); U.S.A. (Kudo, 1926b; McAdow, 1931; Hatcher, 1939; Armer, 1944); Philippine Islands (Hegner and Chu, 1930).

Periplaneta sp., Goa (Mello and Lima Ribeiro, 1924, 1925).

"Küchenschaben," Austria (Bělǎr, 1916).

The protozoan (fig. 2, E) is found in the host's colon, particularly anterior portion; encysted stages of organism are found throughout hind gut. Of 1,400 B. orientalis studied, 32 percent harbored this organism (Kudo, 1925, 1926). Yakimov and Miller (1922) found 7 percent of 124 B. orientalis infested. Zasukhin (1930) found 10 percent of over 3,000 B. orientalis infested. The flagellate does not harm the host and is never present in the host tissue; it should be considered a commensal (Kudo, 1926).

=Lophomonas striata= Bütschli

Synonymy.--Lophomonas sulcata Schuster is most probably identical with L. striata (Kudo, 1926b).

Natural hosts.--Blatta orientalis, Germany (Bütschli, 1878; Schubotz, 1905); Europe (Janicki, 1908, 1910); U.S.A. (Kudo, 1922, 1926, 1926b; McAdow, 1931); U.S.S.R. (Yakimov and Miller, 1922; Zasukhin, 1930); Poland (Lorenc, 1939).

Blattella germanica, U.S.A., Ohio (McAdow, 1931).

Blattella germanica and/or Periplaneta americana, South Africa (Porter, 1930).

Periplaneta americana, Indochina (Weill, 1929); Philippine Islands (Hegner and Chu, 1930); U.S.A. (Kudo, 1926b; McAdow, 1931; Armer, 1944).

Cockroach, Venezuela (Tejera, 1926); England or U.S.A.? (Lucas, 1928).

"Küchenschaben," Austria (Bělǎr, 1916).

Found in the host's colon, particularly the anterior portion. L. striata (fig. 2, D) was found in 29 percent of 1,400 B. orientalis and in 2 of 30 P. americana (Kudo, 1926, 1926b). Yakimov and Miller (1922) found the organism in 9.6 percent of 124 specimens of B. orientalis. Zasukhin (1930) found 8.6 percent of over 3,000 B. orientalis infested.

Grassé (1926, 1926a) identified corrugations on the surface of L. striata as a bacterial parasite which he named Fusiformis lophomonadis.

=Prolophomonas tocopola= Cleveland et al.

Natural host.--Cryptocercus punctulatus, California, Oregon, Virginia, West Virginia (Cleveland et al., 1934): Not abundant.

Family HOPLONYMPHIDAE

=Barbulanympha estaboga= Cleveland et al.

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934).

Barbulanympha coahoma (Cleveland et al., 1934) represents the diploid form of B. estaboga (Cleveland, 1953).

=Barbulanympha laurabuda= Cleveland et al.

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934): This species, B. ufalula, and Rhynchonympha tarda occur in all parts of the colon, especially in the enlarged, flexed part near the ileum.

=Barbulanympha ufalula= Cleveland et al.

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934).

=Barbulanympha wenyoni= Cleveland

Natural host.--Cryptocercus punctulatus, U.S.A., Pacific coast area (Cleveland, 1953).

=Rhynchonympha tarda= Cleveland et al.

(Fig. 3, D)

Natural host.--Cryptocercus punctulatus, U.S.A., Pacific coast area (Cleveland et al., 1934): Fairly abundant in every specimen examined from Pacific coast.

=Urinympha talea= Cleveland et al.

(Fig. 3, C)

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934): Present in fairly great numbers in every cockroach examined.

Family STAUROJOENINIDAE

=Idionympha perissa= Cleveland et al.

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian area (Cleveland et al., 1934): Present in only a few specimens.

Family TRICHONYMPHIDAE

=Trichonympha acuta= Cleveland et al.

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934).

=Trichonympha algoa= Cleveland et al.

(Fig. 3, E)

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934): Fairly abundant and present in most specimens.

=Trichonympha chula= Cleveland et al.

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934).

=Trichonympha grandis= Cleveland et al.

Natural host.--Cryptocercus punctulatus, U.S.A., Pacific coast areas (Cleveland et al., 1934): Fairly abundant in all specimens from Pacific area.

=Trichonympha lata= Cleveland et al.

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934).

=Trichonympha okolona= Cleveland et al.

(Fig. 3, E)

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934): Found in only a few specimens, never abundant.

=Trichonympha parva= Cleveland et al.

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934): This organism is smaller than any known species of Trichonympha; it is more resistant to warm weather than the other hypermastigotes.

Family EUCOMONYMPHIDAE

=Eucomonympha imla= Cleveland et al.

Natural host.--Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934): Organism (fig. 3, A) sometimes becomes attached to the intestinal wall; attached individuals were seen in 2 to 3 percent of the cockroaches examined.

=Unidentified flagellate=

Natural host.--Pycnoscelus surinamensis, Hawaii (Schwabe, 1950): A small flagellate was found in the digestive tract and malpighian tubules.

Class SARCODINA

Order MYCETOZOA

INCERTAE SEDIS

=Peltomyces periplanetae= (Léger)

Synonymy.--Peltomyces blattellae. Sprague (1940a) synonymizes Peltomyces periplanetae, with Coelosporidium periplanetae.

Natural hosts.--Blatta orientalis, France (Debaisieux, 1927).

Blattella germanica, France (Léger, 1909; Debaisieux, 1927).

The organism inhabits the malpighian tubules of cockroaches. Léger and Debaisieux concluded that their organism was a mycetozoan, but they may have erred in synonymizing Plistophora periplanetae with the organism they studied. Debaisieux found intracellular stages of Peltomyces periplanetae that have not been found in Plistophora periplanetae or Coelosporidium periplanetae.

Order AMOEBINA

Family AMOEBIDAE

=Hartmannella blattae= Ivanić

Natural host.--Blatta orientalis, Yugoslavia (Ivanić, 1937): Found in the hind gut.

Family ENDAMOEBIDAE

In the following classification we have accepted the conclusions of Kirby (1945), Kudo (1954), and others that species of Endamoeba are generically different from species of Entamoeba and that the latter genus is not a homonym of Endamoeba.

=Dobellina= sp.

Natural vectors.--Blattella germanica and/or Periplaneta americana, Egypt (DeCoursey and Otto, 1956, 1957): Thirty out of 261 cockroaches examined contained this protozoan.

=Endamoeba blattae= (Bütschli)

Synonymy.--Amoeba blattae, Entamoeba blattae, Entamoeba blattarum.

Natural hosts.--Blatta orientalis, Germany (Bütschli, 1878; Schubotz, 1905; Chen, 1933); U.S.A. (Leidy, 1879a, 1880; Kudo, 1922, 1925a, 1926a; Kirby, 1927; McAdow, 1931; Meglitsch, 1938, 1940); France (Mercier, 1907a, 1908, 1909, 1910); Europe? (Janicki, 1908, 1909); U.S.S.R. (Yakimov and Miller, 1922; Zasukhin, 1929, 1930); England (Thomson and Lucas, 1926; Lucas, 1927, 1927a, 1928); Yugoslavia (Ivanić, 1926a).

Blattella germanica and/or Periplaneta americana, South Africa (Porter, 1930); Egypt (DeCoursey and Otto, 1956, 1957): Seven out of 217 cockroaches examined harbored the protozoan.

Periplaneta americana, Philippine Islands (Hegner and Chu, 1930); U.S.A. (Morris, 1936; Armer, 1944); Gold Coast Colony (Macfie, 1922).

Periplaneta australasiae, U.S.A. (Morris, 1936).

Cockroaches, Paraguay? (Elmassian, 1909); Austria (Bělǎr, 1916); U.S.A. (Morris, 1935, 1936; Balch, 1932); Venezuela (Tejera, 1926).

The habitat of E. blattae (fig. 2, C) is the hind intestine and rectum of the cockroach. The incidence of infection varies: Kudo (1925a) found in 1,255 oriental cockroaches infections in 5 percent in March and 50 percent in the summer; Schubotz (1905) found 5 to 20 percent of the examined cockroaches to be infested; Yakimov and Miller (1922) found 4 percent of 124 oriental cockroaches infested; Zasukhin (1930) found up to 50 percent of over 3,000 B. orientalis infested; Meglitsch (1938, 1940) found almost 100 percent infection in B. orientalis kept in a crowded culture for several weeks. Chen (1933) developed two synthetic media in which E. blattae could be grown for 45 to 50 days.

Mercier (1907a) observed a fungus, Nucleophaga sp., hyperparasitic in the nucleus of Endamoeba blattae.

=Endamoeba javanica= Kidder

Natural hosts.--Panesthia angustipennis, Philippine Islands, and Panesthia spadica, Japan (Kidder, 1937): Occurred in 50 percent of P. angustipennis examined and in one of four P. spadica. The endoplasm of this amoeba contains large amounts of wood and cellulose fibers.

=Endamoeba philippinensis= Kidder

Natural host.--Panesthia angustipennis, Philippine Islands (Kidder, 1937): Occurred in about 10 percent of the Panesthia examined. The food vacuoles contained bacteria, no wood.

=Entamoeba coli= (Grassi)

Synonymy.--Endamoeba coli, Amoeba coli [Kirby, 1945].

Natural vectors.--Blaberus atropos, Venezuela (Tejera, 1926): In a lot of 60 cockroaches captured in latrines, two were found that carried apparently live cysts similar to cysts of E. coli.

Blattella germanica or Periplaneta americana, Egypt (DeCoursey and Otto, 1956, 1957): One out of 44 cockroaches collected in a village harbored E. coli.

Experimental vector.--Periplaneta americana, Gold Coast Colony (Macfie, 1922): In nine experiments cysts of E. coli were fed to the cockroaches. In seven of the experiments cysts of E. coli were found in the feces. Cysts were observed in the feces for only one to three days, and eventually disappeared completely. The cysts appeared to be unharmed. No amoebae were found.

* =Entamoeba histolytica= Schaudinn

Natural vectors.--Blatta orientalis, Blattella germanica, Periplaneta americana, Periplaneta australasiae, and/or Supella supellectilium, Peru (Schneider and Shields, 1947).

Blattella germanica and/or Periplaneta americana, Egypt (DeCoursey and Otto, 1956, 1957).

Cockroaches, Venezuela (Tejera, 1926).

Experimental vectors.--Blatta orientalis, Italian Somaliland (Mariani and Besta, 1936).

Periplaneta americana, Gold Coast Colony (Macfie, 1922); U.S.A. (Frye and Meleney, 1936).

Cockroaches, Venezuela (Tejera, 1926).

=Entamoeba pitheci= Prowazek?

Experimental vector.--Periplaneta americana, Formosa (Morischita and Tsuchimochi, 1926): Eleven of 15 cockroaches fed feces of a monkey [Macaca cyclopis (Swinhoe)] containing cysts of the amoeba voided live cysts in their own feces.

=Entamoeba thomsoni= Lucas

Synonymy.--Endamoeba thomsoni [Kudo, personal communication, 1957].

Natural hosts.--Blatta orientalis, England (Lucas, 1927a, 1928); U.S.A. (Taliaferro, 1928; McAdow, 1931); U.S.S.R. (Zasukhin, 1930); Germany (Chen, 1933).

Blattella germanica, U.S.A. (McAdow, 1931).

Periplaneta americana, England (Lucas, 1927a); U.S.A. (Smith and Barret, 1928; McAdow, 1931); Philippine Islands (Hegner and Chu, 1930).

The organism is found in the hind intestine and rectum of the cockroach. Smith and Barret (1928) developed a synthetic medium in which cultures of E. thomsoni were carried through successive transfers for 24 months.

=Entamoeba= sp.

Natural vector.--Periplaneta americana, Gold Coast Colony (Macfie, 1922): Under the heading "Entamoeba histolytica and E. coli" Macfie (p. 445) stated, "The cockroaches used in these experiments had previously been carefully examined for amoebic infections a precaution which was doubly necessary, because some of these insects at Accra had been found naturally infected."

Experimental vectors.--Periplaneta americana, Gold Coast Colony (Macfie, 1922): Entamoeba, resembling E. coli, from feces of the monkey [Erythrocebus patas patas (Schreber)] were fed to cockroaches, and on the second to fourth days thereafter apparently healthy cysts were recovered in the cockroach feces.

=Endolimax blattae= Lucas

Natural hosts.--Blatta orientalis, England (Lucas, 1927, 1927a); U.S.S.R. (Zasukhin, 1930); Germany (Chen, 1933).

Periplaneta americana, England (Lucas, 1927, 1927a); Indochina (Weill, 1929); U.S.A. (Armer, 1944).

Periplaneta australasiae, U.S.A. (Steinhaus, 1946).

Organism is found in the hind gut of the cockroach. Zasukhin (1930) found 3-percent infestation in over 3,000 B. orientalis examined.

=Endolimax nana= (Wenyon and O'Connor)?

Synonymy.--Entamoeba nana.

Natural host.--Blaberus atropos, Venezuela (Tejera, 1926): A small amoeba greatly resembling E. nana was found in the intestinal contents of the cockroach. In inoculations this amoeba was not pathogenic.

=Endolimax= sp.

Natural hosts.--Blatta orientalis, U.S.S.R. (Zasukhin, 1930): This organism was found in the hind gut of 0.3 percent of over 3,000 cockroaches examined.

Blattella germanica and/or Periplaneta americana, Egypt (DeCoursey and Otto, 1956, 1957): Seventy-four out of 261 cockroaches examined harbored this protozoan.

=Iodamoeba= sp.

Natural vectors.--Blattella garmanica and/or Periplaneta americana, Egypt (DeCoursey and Otto, 1956, 1957): Fifty-nine of 261 cockroaches examined contained this protozoan. Iodamoeba sp. was common in human feces in villages in which the cockroaches were collected.

=Undetermined species of Amoeba=

Natural host.--Panesthia angustipennis, Philippine Islands. (Kidder, 1937): Found in only one specimen.

Class SPOROZOA

Order GREGARINIDA

Family DIPLOCYSTIDAE

=Diplocystis schneideri= Kunstler

(Fig. 2, I)

Natural hosts.--Blatta orientalis, England (Woodcock, 1904; Jameson, 1920).

=Periplaneta americana=, France (Kunstler, 1884, 1887); England (Jameson, 1920); Germany (Foerster, 1939).

In body cavity of host. Cysts containing spores are ingested during cannibalistic feeding on infected cockroaches. Sporozoites penetrate the gut wall which later ruptures, freeing the gregarines into the coelom. There is no apparent pathogenic effect. Jameson (1920) found 81 percent of P. americana infested with D. schneideri.

=Diplocystis= sp.

Natural host.--Periplaneta americana, U.S.A. (Hertig, 1921): Heavy infections in body cavity.

Cockroach, India (Ray and Dasgupta, 1955): A large number of cockroaches, both adults and nymphs, collected in Calcutta were all infected.

=Diplocystis= sp.?

Natural host.--Blaberus craniifer, U.S.A. (Nutting, 1953): From 1 to 12 or more paired trophozoites or cysts may be found in the hemocoele and occasionally in the thorax.

Family STENOPHORIDAE

=Stenophora= sp.

Natural host.--Blatta orientalis, India (Bal and Rai, 1955): Organism found in the midgut of the cockroach.

Family GREGARINIDAE

=Gregarina blattarum= von Siebold

Synonymy.--Gregarina blattae orientalis; Clepsidrina blattarum.

Natural hosts.--Blatta orientalis, Germany (Siebold, 1837, 1839; Stein, 1848; Bütschli, 1881; Wolters, 1891; Marshall, 1892; Wellmer, 1910, 1911; Foerster, 1938; Schubotz, 1905); U.S.A. (Leidy, 1853a; Crawley, 1903; Watson, 1917; Kudo, 1922; McAdow, 1931; Sprague, 1940, 1941); England (Lankester, 1863); France (Schneider, 1875; Cuénot, 1901; Laveran and Franchini, 1920a); Brazil (Magalhães, 1900; Pinto, 1919); U.S.S.R. (Zasukhin, 1929, 1930).

Blattella germanica, U.S.A. (Crawley, 1903); South Africa (Fantham, 1929; Porter, 1930: these appear to be the same record).

Blattella germanica and/or Periplaneta americana, Egypt (DeCoursey and Otto, 1956, 1957).

Periplaneta americana, Brazil (Magalhães, 1900); U.S.A. (Crawley, 1903; 1907; McAdow, 1931); South Africa (Fantham, 1929; Porter, 1930: these appear to be the same record); Gold Coast Colony (Macfie, 1922).

Parcoblatta pensylvanica, U.S.A., Michigan (Ellis, 1913a).

Cockroaches, Germany (Schiffmann, 1919: probably used the oriental cockroach); Venezuela (Tejera, 1926).

Organism usually found in the intestinal tract of cockroaches where it is attached to the gut cells. Cysts are passed in the feces. Occasionally, G. blattarum (fig. 2, J) is found in the body cavity (Leidy, 1853a; Hall, 1907). Though this is considered to be one of the commonest of the Sporozoa encountered in cockroaches, DeCoursey and Otto (1956) found only 10 of 217 P. americana and B. germanica, collected in restaurants in Egypt, infested with this species. Watson (1917) found a dozen or more in one specimen of Blatta orientalis. Zasukhin (1929, 1930) found 2.6 percent of 3,000 oriental cockroaches infected with this parasite.

=Gregarina fastidiosa= Harrison

Natural host.--Aptera fusca, South Africa (Harrison, 1955): All mature females were heavily infected; in all specimens there were over 100 parasites in the gut. All nymphs were infected, the earlier instars more lightly than the later instars. Gregarines were found in all parts of the gut except the crop and gizzard.

=Gregarina gibbsi= Harrison

Natural host.--Temnopteryx phalerata, South Africa (Harrison, 1955): Although the cockroaches were found in groups, only 32 percent were infected and only 10 percent heavily. The gregarines were found in the anterior mesenteron but none in the hepatic caeca. All cysts were found in the hind gut or rectum.

=Gregarina illinensis= M. E. Watson

Natural host.--Parcoblatta pensylvanica, U.S.A., Illinois (Watson, 1915, 1916): The intestine of one cockroach was found to contain 25 of these gregarines.

=Gregarina impetuosa= Harrison

Natural host.--Melanosilpha capensis, South Africa (Harrison, 1955): All specimens of this gregarine were found in the anterior mesenteron of the host.

=Gregarina légeri= Pinto

Natural host.--Periplaneta americana, Brazil (Pinto, 1918, 1918a, 1919): Intestinal canal.

=Gregarina neo-brasiliensis= Al. Cunha

Natural host.--Periplaneta americana, Brazil (R. de Almeida Cunha in Pinto, 1919; Cunha, 1919).

=Gregarina ohioensis= Semans

Natural host.--Parcoblatta virginica, U.S.A., Ohio (Semans, 1939): The protozoan was present in large numbers in the insect's midgut.

=Gregarina panchlorae= Frenzel

Natural host.--Panchlora exoleta, Argentina (Frenzel, 1892): Midgut.

=Gregarina parcoblattae= Semans

Natural hosts.--Parcoblatta pensylvanica and Parcoblatta uhleriana, U.S.A., Ohio (Semans, 1939): Midgut.

=Gregarina rhyparobiae= J. M. Watson

Natural host.--Leucophaea maderae, Uganda (Watson, 1945): Midgut. Trophozoites could be seen in sections attached to cells of the intestinal wall (fig. 2, H).

=Gregarina sandoni= Harrison

Natural host.--Melanosilpha capensis, South Africa (Harrison, 1955): This gregarine was found in the anterior and middle parts of the mesenteron and in the hepatic caeca.

=Gregarina thomasi= Semans

Natural host.--Parcoblatta pensylvanica, U.S.A., Ohio (Semans, 1939): Enteric caeca and midgut.

=Protomagalhaesia serpentula= (de Magalhães)

Synonymy.--Gregarina serpentula [Pinto, 1918a, 1919; Semans, 1943].

Natural host.--Periplaneta americana, Brazil (Magalhães, 1900): In the coelom and alimentary canal. The host of this parasite (fig. 2, K) was incorrectly cited as Blatta orientalis by Watson (1916).

=Gamocystis tenax= Schneider

(Fig. 2, L)

Natural hosts.--Ectobius lapponicus, France (Schneider, 1875); Germany (Wellmer, 1910, 1911; Foerster, 1938).

Ectobius pallidus, Germany (Foerster, 1938): In intestine.

Family ACTINOCEPHALIDAE

=Pileocephalus blaberae= (Frenzel)

Synonymy.--Gregarina blaberae [Watson, 1916].

Natural hosts.--Blaptica dubia and related forms, Argentina (Frenzel, 1892): In midgut.

=Unidentified Gregarinida=

Natural hosts.--Blaberus craniifer, U.S.A. (Roth and Willis, unpublished data, 1953): Possibly Diplocystis sp. (pl. 28, A, B).

Cryptocercus punctulatus, U.S.A. (Cleveland et al., 1934).

Leucophaea maderae, Philippine Islands (Hegner and Chu, 1930): In intestines of host. U.S.A. (Roth and Willis, unpublished data, 1958): Cysts in feces (pl. 28, C).

Gromphadorhina portentosa, U.S.A., in laboratory colony (Roth and Willis, unpublished data, 1958): In intestine of adult female.

Pycnoscelus surinamensis, Hawaii (Schwabe, 1950): A cephaline gregarine was found in the cockroach's digestive tract; it was also claimed to be present in new-born nymphs.

Order COCCIDIA

Family ADELEIDAE

=Adelina cryptocerci= Yarwood

Natural host.--Cryptocercus punctulatus, U.S.A. (Yarwood, 1937): This intracellular parasite was found in the fat body in light infestations. In heavy infections the coccidia were found in the head, antennae, mouthparts, muscles, legs, salivary glands, nerve cord, as well as fat body. Infection in freshly collected specimens was about 3 percent; when large numbers of cockroaches were kept together in culture, the rate of infection increased because the insects ate their dead companions.

Cleveland et al. (1934) mentioned a coccidium which was sometimes generally distributed through the body (head, legs, antennae, etc.) of C. punctulatus; this parasite was probably the species described by Yarwood.

Order HAPLOSPORIDIA

=Haplosporidium periplanetae= Georgévitch

Natural host.--Blatta orientalis, Yugoslavia (Georgévitch, 1953): This organism was described from the malpighian tubules of the cockroach where it apparently occurred in a mixed infection with the microsporidian Plistophora periplanetae. See synonymy under Plistophora periplanetae.

=Coelosporidium periplanetae= (Lutz and Splendore)

Synonymy.--Nosema periplanetae, Coelosporidium blattellae, Bertramia blatellae [after Semans, 1943]. Some of the observations cited under Plistophora periplanetae may pertain to C. periplanetae (see Sprague, 1940). See also Haplosporidium periplanetae.

Natural hosts.--Blatta orientalis, U.S.S.R. (Epshtein, 1911); U.S.A. (Kudo, 1922; Sprague, 1940); Yugoslavia (Ivanić, 1926).

Blattella germanica, U.S.A. (Crawley, 1905); Germany (Wellmer, 1910, 1911).

Periplaneta americana, Brazil (Lutz and Splendore, 1903).

This organism (fig. 2, B) passes its life cycle living free in the lumina of the malpighian tubules of cockroaches. The elongate trophozoite is firmly attached to the wall of the tubule as are clusters of immature spores. Mature spores are freed into the lumina of the tubules from whence they pass to the exterior. Sprague (1940) examined about 200 wild-caught B. orientalis and found them to be practically 100 percent infected.

Order MICROSPORIDIA

Family NOSEMATIDAE

=Plistophora kudoi= Sprague and Ramsey

Natural host.--Blatta orientalis, U.S.A., Illinois, West Virginia, Kentucky (Sprague and Ramsey, 1941, 1942): Found in the epithelial cells of caeca and midgut. Considerable damage is done to these cells. Seventy-five percent of 52 B. orientalis harbored the parasite.

=Plistophora periplanetae= (Lutz and Splendore)

Synonymy.--Nosema periplanetae, Pleistophora periplanetae [after Semans, 1943]. Georgévitch (1953) has pointed out that one may find in the malpighian tubules of cockroaches a mixed infection of Microsporidia, Haplosporidia, and Mycetozoa, and that some of the discrepancies in the earlier literature may be attributed to attempts to combine in one organism disparate stages belonging to different orders. See also comments under Coelosporidium periplanetae, Haplosporidium periplanetae, and Peltomyces periplanetae.

Natural hosts.--Blatta orientalis, France (Mercier, 1906a; Debaisieux, 1927); England (Perrin, 1906, 1906b); U.S.S.R. (Zhivago, 1909); Yugoslavia (Georgévitch, 1925, 1926, 1926a, 1927); Germany (Wellmer, 1910, 1911).

Blattella germanica, France (Léger, 1909; Debaisieux, 1927); U.S.S.R. (Zhivago, 1909).

Periplaneta americana, Brazil (Lutz and Splendore, 1903).

This organism lives in the lumen of the malpighian tubules of cockroaches. The cited authors appear to have been convinced that this organism was a microsporidian. Georgévitch (1927, 1953) described the polar capsule and filament characteristic of this order.

=Plistophora= sp.

Natural host.--Blatta orientalis, France (Mercier, 1908a): The organism parasitized the fat body of the cockroach. Mitoses, often abnormal, were induced in the fat cells. Infected cockroaches were easily recognizable by their distended abdomens. The fat body became chalky white and showed through the intersegmental membranes.

Porter (1930) reported finding an unidentified microsporidian in the fat bodies of Blattella germanica and Periplaneta americana collected in South Africa. It may or may not have been a species of Plistophora.

Class CILIATA

Order HOLOTRICHA

Family PARAMECIIDAE

=Paramecium= sp.

Natural associate.--Cockroaches, U.S.A., Maryland (Cleveland, 1927): Three of 30 cockroaches collected in the basement of a department store had paramecia in their stomachs but none in the rectum.

Experimental associate.--Periplaneta americana, U.S.A. (Hegner, 1929): Paramecia fed to the cockroaches were recovered from the crop at intervals from one-half to six and one-half hours. In no case were the protozoa recovered from the stomach alive.

Cockroaches, U.S.A., Maryland (Cleveland, 1927). About 200 starved cockroaches were fed a culture of Paramecium. Few, if any, of the protozoa were killed in the stomach during the first two hours, but all were killed within 5 to 6 hours after ingestion.

Family ISOTRICHIDAE

=Isotricha caulleryi= Weill

Natural host.--Periplaneta americana, Indochina (Weill, 1929): Alimentary canal.

Order SPIROTRICHA

Family BURSARIIDAE

=Balantidium blattarum= Ghosh

Natural host.--Periplaneta americana, India (Ghosh, 1922; Bhatia and Gulati, 1927); Gold Coast Colony (Macfie, 1922): Intestinal tract.

* =Balantidium coli= (Malmsten)

Experimental vector.--Cockroach, Venezuela (Tejera, 1926).

=Balantidium ovatum= Ghosh

Natural host.--Periplaneta americana, India (Ghosh, 1922a; Bhatia and Gulati, 1927); Indochina, Saigon (Weill, 1929): Intestinal tract.

=Balantidium praenucleatum= Kudo and Meglitsch

Natural host.--Blatta orientalis, U.S.A., Illinois (Kudo and Meglitsch, 1938; Meglitsch, 1940): This protozoan is found in the lumen of the anterior region of the colon in association with several other species of protozoa. Only 7.6 percent of 500 cockroaches examined contained B. praenucleatum. The largest number encountered in a single host was 59, but as a rule each host harbored a smaller number.

=Balantidium= sp.?

Natural host.--Periplaneta americana, Brazil (Magalhães, 1900): These organisms were numerous in the intestine.

Family SPIROSTOMIDAE

=Nyctotherus buissoni= Pinto

Natural host.--"Barata sylvestre," Brazil (Pinto, 1926): Organism found in the cockroach's intestine.

=Nyctotherus ovalis= Leidy

Synonymy.--Bursaria blattarum; Plagiotoma blattarum.

Natural hosts.--Blatta orientalis, U.S.A. (Leidy, 1850, 1853, 1853b, 1879a; Kudo, 1922, 1926, 1936; McAdow, 1931; Kudo and Meglitsch, 1938; Meglitsch, 1940); Germany (Stein, 1860; Schubotz, 1905; Chen, 1933); England (Lankester, 1865; Schuster, 1898; Lucas, 1927a, 1928); Spain (Zulueta, 1916); U.S.S.R. (Yakimov and Miller, 1922; Zasukhin, 1928, 1930; Ostroumov, 1929); Portugal (Lima Ribiero, 1924); Brazil (Pinto, 1926); Venezuela (Tejera, 1926).

Blattella germanica, South Africa (Porter, 1930); U.S.A. (Balch, 1932; McAdow, 1931).

Parcoblatta pensylvanica, U.S.A. (Semans, 1939, 1941).

Periplaneta americana, India (Bhatia and Gulati, 1927); Indochina (Weill, 1929); Philippine Islands (Hegner and Chu, 1930); South Africa (Porter, 1930); U.S.A. (McAdow, 1931; Hatcher, 1939; Meglitsch, 1940; Armer, 1944); Goa (Mello et al., 1934); China (Pai and Wang, 1947); Czechoslovakia (Low, 1956).

"Barata sylvestre," Brazil (Pinto, 1926).

"Küchenschaben," Austria (Bělǎr, 1916).

Nyctotherus ovalis (fig. 2, G) inhabits the hind gut of cockroaches, where it occurs almost always in the anterior half of the colon in association with other species of Protozoa (Kudo, 1936). Ninety percent of 500 B. orientalis contained N. ovalis (Kudo and Meglitsch, 1938). Yakimov and Miller (1922) found N. ovalis in 68 percent of 124 B. orientalis. Zasukhin (1930) found this organism in 63 percent of over 3,000 B. orientalis. Zasukhin (1928, 1934) found a fungus and possibly a bacterium hyperparasitic in the cytoplasm of N. ovalis. N. ovalis has been cultured outside the cockroach in several media (Lucas, 1928; Balch, 1932; Chen, 1933; Low, 1956).

=Nyctotherus uichancoi= Kidder

Natural hosts.--Panesthia angustipennis, Philippine Islands, and Panesthia spadica, Japan (Kidder, 1937): About 90 percent of all P. angustipennis harbored this ciliate in their hindguts.

=Nyctotherus viannai= Pinto

Natural host.--"Barata sylvestre," Brazil (Pinto, 1926): In the intestine of the cockroach.

Family CLEVELANDELLIDAE

Most of the Clevelandellidae are parasitized by rod-shaped or spherical bacteria-like organisms usually in clusters (Kidder, 1937).

Synonymy.--Clevelandiidae (Kidder, 1938).

Genus CLEVELANDELLA

Synonymy.--The generic name Clevelandia Kidder (1937) is preoccupied; it was therefore changed to Clevelandella by Kidder in 1938. All of the following species of Clevelandella were originally described as Clevelandia.

=Clevelandella constricta= (Kidder)

Natural hosts.--Panesthia angustipennis, Philippine Islands, and Panesthia spadica, Japan (Kidder, 1937): In the posterior end of hindgut.

=Clevelandella contorta= (Kidder)

Natural hosts.--Panesthia angustipennis, Philippine Islands, and Panesthia spadica, Japan (Kidder, 1937).

=Clevelandella elongata= (Kidder)

Natural host.--Panesthia angustipennis, Philippine Islands (Kidder, 1937).

=Clevelandella hastula= (Kidder)

Natural host.--Panesthia angustipennis, Philippine Islands (Kidder, 1937): Common in hindgut.

=Clevelandella nipponensis= (Kidder)

Natural host.--Panesthia spadica, Japan (Kidder, 1937).

=Clevelandella panesthiae= (Kidder)

Natural hosts.--Panesthia angustipennis, Philippine Islands, and Panesthia spadica, Japan (Kidder, 1937): In the hindgut. This protozoan is commonly parasitized by the microorganism Sphaerita.

=Clevelandella parapanesthiae= (Kidder)

Natural host.--Panesthia angustipennis, Philippine Islands (Kidder, 1937).

=Paraclevelandia brevis= Kidder

Natural hosts.--Panesthia angustipennis, Philippine Islands, and Panesthia spadica, Japan (Kidder, 1937): Present in 100 percent of P. angustipennis and in nearly all P. spadica.

=Paraclevelandia simplex= Kidder

Natural hosts.--Panesthia angustipennis, Philippine Islands (Kidder, 1937, 1938): Incidence of infection about 50 percent.

Panesthia spadica, Japan (Kidder, 1937).

=Unidentified ciliate=

Natural host.--Pycnoscelus surinamensis, Hawaii (Schwabe, 1950): A large ciliate was found in the digestive tract and malpighian tubules.

NEGATIVE FINDINGS

In a recent experimental study Schmidtke (1955) failed to demonstrate a host-parasite relationship between Periplaneta americana and the haemosporidian Toxoplasma gondii Nicolle and Manceaux. This protozoan is a blood parasite in a rodent in North Africa (Kudo, 1954).

XI. HELMINTHS

Intestinal nematodes of the family Thelastomatidae have no apparent pathological effect on their cockroach hosts. Galeb (1878) has shown experimentally that oxyurids eat the same food as the host insect and that if one starves them, by withholding food from the host, the oxyurids die and disappear. In other words, these worms are not parasites, in the sense that we use the term in this paper, but commensals. Dobrovolny and Ackert (1934) stated that "all observations seemed to indicate that the health, fertility and activity of the heavily infested cockroaches were comparable with those of the non-parasitised specimens."

Very few papers have dealt with the ecology of the oxyurid parasites of cockroaches. According to Galeb (1878), usually one species of nematode is found in a single cockroach, but sometimes two species live together in the same host (e.g., in Blatta orientalis and Polyphaga aegyptiaca) where they compete for food. Galeb claimed that Hammerschmidtiella diesingi would replace Leidynema appendiculata; he observed that H. diesingi surpassed L. appendiculata in numbers and the latter became uncommon in the intestines of the cockroaches. On the other hand, Sobolev (1937) found that 48 percent of his oriental cockroaches were infected with both of the above species of nematodes. The average number of both species was 7.5, and the maximum number was 97; the mean number of H. diesingi was 5.1 and the maximum 64; the mean number of L. appendiculata was 2.4 and the maximum 33. More than 40 nematodes were found in each cockroach of 1.3 percent of those examined. These results apparently contradict Galeb's conclusions inasmuch as the number of each species in mixed infections was essentially the same as the number found in cockroaches infected by only one species (see pp. 195 and 197). Dobrovolny and Ackert (1934) found that 29 percent of 222 Periplaneta americana contained both of the above species of nematodes; whereas 40 percent contained L. appendiculata only, and 21 percent contained H. diesingi only. The infestation ranged from 1 to 36 worms per cockroach with averages of 3.8 per male, 5.1 per female, and 2.7 per nymph.

The eggs of some helminths pass unharmed through the guts of cockroaches that serve as vectors of these ova and have no effect on the insect. However, helminths that are secondary parasites in cockroaches damage the insect to varying degrees depending upon the extent of the infection. Thus the larvae of Moniliformis moniliformis pass through the gut wall and some may become embedded in the fat tissue (Moore, 1946). First stage larvae of Oxyspirura mansoni also burrow through the midgut wall into the fat body; Sanders (1929) believed that Pycnoscelus surinamensis could be killed if at one time a sufficient number of migrating larvae of O. mansoni penetrated the cockroach's intestinal wall. Gongylonema neoplasticum migrates through the digestive tract and encysts in the muscles of the thorax and legs of the host (Fibiger and Ditlevsen, 1914). Infective larvae of Protospirura muricola, after hatching from ingested eggs, pass through the cockroach's gut wall and encyst mainly in the thorax, around the crop, and at the bases of the large muscles of the prothoracic legs (Foster and Johnson, 1939). It is probably generally true that nematodes which are secondary parasites in cockroaches do some damage to the host's intestinal tract at least, and they probably also damage other organs in which they may encyst.

Cockroach tissues may react defensively to infections by parasitic nematodes. For example, encysted third-stage larvae of Physaloptera turgida have been found enclosed in a thin, brown, chitinous substance that was undoubtedly deposited by the tissue of the cockroach (Alicata, 1937). Cysts of similar appearance have been found in cockroaches infected with Physaloptera rara, P. maxillaris, P. hispida (Petri, 1950; Hobmaier, 1941; Schell, 1952), and Gongylonema pulchrum (Schell, 1952a); in the latter species the deposit eventually completely surrounded the nematode larva which was killed and "chitinized." Apparently these pigmented cysts surround unhealthy or dead larvae and are secreted as a defensive mechanism by the host (Schell, 1952a). Oswald (1958) has reported finding similar pigmented cysts in Blatta orientalis and Periplaneta americana that were experimentally infected with Rictularia coloradensis.

Our classification of the helminths follows Hyman (1951, 1951a).

HELMINTHS FOR WHICH COCKROACHES SERVE AS PRIMARY HOSTS

Phylum ASCHELMINTHES

Class NEMATODA

Order MERMITHOIDEA

Family MERMITHIDAE

=Undetermined mermithids=

Natural Hosts.--Ectobius pallidus, U.S.A., Plymouth, Massachusetts (Roth and Willis, 1957): This mermithid lies coiled in the body cavity of the host and one end may extend into the thorax. Apparently, the host is eventually killed and the worms may leave the cockroach ventrally between the thorax and abdomen (pl. 29, A) or thorax and head.

Periplaneta americana, Germany (Bode, 1936): Attacked by "Mermis" or "Gordius." It has been suggested that the name Mermis is often applied without critical identification to immature Nematoda found in insects (Buxton, 1955).

Order RHABDITOIDEA

Family DIPLOGASTERIDAE

=Diplogaster= sp.

Synonymy.--Lycolaimus [Goodey, 1951].

Experimental Host.--Blattella germanica, U.S.A. (Christie and Crossman, 1933).

Family STEINERNEMATIDAE

=Neoaplectana= sp.

Experimental hosts.--Blattella germanica, Nauphoeta cinerea, and Periplaneta americana. U.S.A. (Dutky and Hough, 1955): This nematode, found in codling moth larvae, is close to Neoaplectana chresima Steiner but apparently is a new species. Nauphoeta cinerea was very susceptible to infection; B. germanica and P. americana were less susceptible.

Order OXYUROIDEA

Family THELASTOMATIDAE

These nematodes are found in the intestinal tract of cockroaches.

=Aorurus philippinensis= Chitwood and Chitwood, 1934

Natural host.--Panesthia angustipennis, Philippine Islands (Chitwood and Chitwood, 1934).

=Binema mirzaia= (Basir, 1940) Basir, 1956

Synonymy.--Periplaneticola mirzaia Basir, 1940.

Natural host.--Periplaneta americana, India, Aligarh (Basir, 1940).

=Blattelicola blattelicola= Basir, 1940

Natural host.--Blattella germanica, India, Aligarh (Basir, 1940).

=Blatticola blattae= (Graeffe, 1860) Chitwood, 1932

Synonymy.--Oxyuris blattae Graeffe, 1860; Oxyuris blatticola Galeb, 1878; Blatticola blatticola (Galeb, 1877) Schwenck, 1926 [Chitwood, 1930, 1932].

Natural hosts.--Blattella germanica, Brazil (Pessôa and Corrêa, 1926; Schwenck, 1926); U.S.A. (Chitwood, 1930; Bozeman, 1942); Egypt? (Galeb, 1877, 1878); U.S.S.R. (Sobolev, 1937; Sondak, 1935); Czechoslovakia (Groschaft, 1956).

Ectobius lapponicus, Ectobius pallidus, Egypt? (Galeb, 1877, 1878).

Polyphaga aegyptiaca, France (Graeffe, 1860).

The life cycle has been studied by Bozeman (1942): He found never more than four worms in the large intestine of each cockroach. Embryos developed to "resting" stage in vitro. The resting stage was infective while the active stage was not. The worms seemed to have no effect on the vital activities of the host. Alicata (1934b) found that the embryo undergoes a molt before hatching.

Chitwood (1930) found 75 percent of the German cockroaches examined from houses in Washington infected. As a rule, one adult female, one or two males, and possibly two larval females are found in a single individual, apparently only in the rectum.

Sobolev (1937) found 72 percent of Blattella germanica collected in Gorkov (U.S.S.R.) infected with Blatticola blattae. The mean number of worms per host was 1.8, the maximum 5. Sondak (1935) found about 30 percent of 788 B. germanica collected in Leningrad to be infected with B. blattae. Groschaft (1956) regularly found only single worms in B. germanica, collected in a laboratory in Prague, except for two females that contained 2 and 3 worms each.

=Blattophila sphaerolaima= Cobb, 1920

Synonymy.--Aorurus sphaerolaima (Cobb, 1920) Travassos, 1929. Although Chitwood (1932) indicated that the taxonomic position of this nematode is questionable, Chitwood and Chitwood, 1934, apparently accepted it as a valid species in describing the variety javanica.

Natural host.--Panesthia laevicollis [Cobb recorded the host as Panesthia brevicollis, but no such cockroach exists. Van Zwaluwenburg (1928) and Caudell (in Chitwood, 1932) believed that Cobb meant Panesthia laevicollis. According to Gurney (personal communication, 1957) Caudell's notes show that in 1933 he wrote to Dr. Chitwood and explained that he had compared Cobb's figure of the cockroach with laevicollis Saussure (figures and description) and had found them the same.] Australia, New South Wales (Cobb, 1920).

=Blattophila sphaerolaima var. javanica= Chitwood and Chitwood, 1934

Natural host.--Panesthia angustipennis, Philippine Islands (Chitwood and Chitwood, 1934).

=Blattophila supellaima= Basir, 1941

Natural host.--Supella supellectilium, India, Aligarh (Basir, 1941).

=Cephalobellus brevicaudatum= (Leidy, 1851) Christie, 1933

Synonymy.--Thelastoma brevicaudatum Leidy, 1851 [Christie, 1933]. Thelastoma indiana Basir, 1940 [Basir, 1949].

Natural host.--Leucophaea sp., India, Aligarh (Basir, 1940, 1949).

=Cephalobellus magalhāesi= (Schwenck, 1926) Basir, 1956

Synonymy.--Bulhỡesi magalhāesi Schwenck, 1926; Thelastoma magalhāesi (Schwenck, 1926) Travassos, 1929 [Basir, 1956].

Natural host.--"Barata selvagem," Brazil, São Paulo (Schwenck, 1926).

=Euryconema paradisa= Chitwood, 1932

Natural host.--Eurycotis floridana, U.S.A., Florida (Chitwood, 1932).

=Galebia aegyptiaca= (Galeb, 1878) Chitwood, 1932

Synonymy.--Oxyuris aegyptiaca Galeb, 1878; Blatticola aegyptiaca (Galeb, 1878) Schwenck, 1926 [Chitwood, 1932].

Natural hosts.--Blattella germanica, Brazil (Schwenck, 1926).

Polyphaga aegyptiaca, Egypt? (Galeb, 1878).

=Hammerschmidtiella diesingi= (Hammerschmidt, 1838) Chitwood, 1932

Synonymy.--Anguillula macrura Diesing, 1851; Aorurus diesingi (Hammerschmidt, 1838) Travassos, 1929; Streptostomum gracile Leidy, 1850; Oxyuris diesingi Hammerschmidt, 1838; Oxyuris blattae orientalis Hammerschmidt, 1838 [Chitwood, 1932]. Oxyuris macrura of Lankester (1865).

Natural hosts.--Blatta orientalis, Europe (Hammerschmidt 1838, 1847; Bütschli, 1871); Egypt? (Galeb, 1878); England (Lankester, 1865; Lee, 1958); U.S.A. (Leidy, 1850a); U.S.S.R. (Yakimov and Miller, 1922; Sobolev, 1937; Sondak, 1935); Brazil (Travassos, 1929); China (Chitwood, 1932); Czechoslovakia (Groschaft, 1956).

Leucophaea maderae, Brazil (Pessôa and Corrêa, 1926).

Periplaneta americana, Brazil (Magalhães, 1900; Pessôa and Corrêa, 1926). U.S.A.: Texas (Todd, 1943); Kansas (Dobrovolny, 1933; Dobrovolny and Ackert, 1934); North Carolina (Hatcher, 1939); Iowa, North Dakota, Michigan (Hoffman, 1953). China (Chitwood, 1932). India (Basir, 1940). Czechoslovakia (Groschaft, 1956). England (Lee, 1958).

Periplaneta australasiae, Brazil (Pessôa and Corrêa, 1926).

Polyphaga aegyptiaca (Linstow, 1878).

Cockroaches (Blatta orientalis, Blattella germanica, and/or Periplaneta americana), U.S.A. (McAdow, 1931).

Cockroach, Venezuela (Tejera, 1926).

According to Hammerschmidt (1847) this worm may be found throughout the intestinal canal but especially in the small intestine. It is frequently found in adults and seldom in the nymphs. There were seldom more than 5 to 10 worms in one cockroach and female worms were found more frequently than males; the male worms were found only in winter and spring while the females were present at all times of the year. Bütschli (1871) stated that all stages from those just hatching to mature males and females are found.

Yakimov and Miller (1922) found H. diesingi in 50.8 percent of 124 B. orientalis collected in Petrograd. Sobolev (1937) found 96 percent of B. orientalis infected with H. diesingi with a mean number of 5.6 and maximum number of 22 in one cockroach. Groschaft (1956) found 18 in one specimen of B. orientalis. Dobrovolny and Ackert (1934) found about 50 percent of 222 P. americana infected with H. diesingi. Sondak (1935) found about 36 percent of 412 B. orientalis infected with either or both H. diesingi and Leidynema appendiculata.

Two molts occur during development of the eggs; the first takes place outside the host resulting in a resting or infective stage. After the egg in the infective stage is eaten by the host, the second molt occurs before the egg hatches. Completion of the second molt and hatching perhaps are connected with ammonia present in the digestive tract; the ammonia seems to arise from the bacteria present in the gut. There appears to be a relationship between the intestinal bacteria of the cockroach and development and hatching of nematode eggs (Todd, 1944).

At the time of oviposition the nematode eggs are in the very earliest stages of cleavage. In 36 hours a motile, tadpole-like stage is reached and in a few days the embryo becomes quiescent and nonmotile. This nonmotile stage is infective whereas the motile embryonic stage is not. Feeding experiments proved that transmission of the nematode is direct. The worm reaches sexual maturity in 20 or 30 days after being ingested by the cockroach (Dobrovolny, 1933).

The bacterium Streptomyces leidynematis Hoffman grows on the cuticle of H. diesingi (Hoffman, 1953). The bacterium apparently is only anchored to the nematode and probably obtains its food from the intestinal contents of the cockroach. See notes under Leidynema appendiculata.

=Hammerschmidtiella neyrai= Serrano Sánchez, 1945

Synonymy.--Hammerschmidtiella neyrae Serrano Sánchez, 1947. [According to M. B. Chitwood, personal communication, 1957, Serrano Sánchez's emendation is apparently an error.]

Natural host.--Blatta orientalis, Spain, Grenada (Serrano Sánchez, 1947): Of 2,943 specimens examined, 1,143 were parasitized by oxyurids and of these 45 percent contained H. neyrai.

=Leidynema appendiculata= (Leidy, 1850) Chitwood, 1932

Synonymy.--Oxyuris blattae orientalis Hammerschmidt, 1847, of Bütschli, 1871, and Oxyuris blattae-orientalis of Magalhães, 1900; Oxyuris blattae Hammerschmidt, 1847, of Galeb, 1878; Aorurus (Thelastoma) appendiculatus Leidy, 1850. [Chitwood, 1932.] Serrano Sánchez (1947) has divided this species into three geographical varieties as follows: L. appendiculata (Leidy, 1852) (Dobrovolny and Ackert, 1934) var. indiana; L. appendiculata (Leidy, 1852) (Chitwood, 1932) var. americana; L. appendiculata (Serrano Sánchez, 1947) var. hispana. However, Basir (1956) does not recognize these varieties. The Russians recognize hispana (M. B. Chitwood, personal communication, 1957).

Natural hosts.--Blaberus atropos, South America (Chitwood, 1932).

Blatta orientalis, Egypt? (Galeb, 1878); Europe (Bütschli, 1871); U.S.S.R. (Sobolev, 1937; Sondak, 1935); U.S.A., Nebraska (Todd, 1944); Spain (Serrano Sánchez, 1947): Recorded as var. hispana. Czechoslovakia (Groschaft, 1956). England (Lee, 1958a).

Blatta orientalis or Periplaneta americana, Brazil (Magalhães, 1900).

Periplaneta americana, U.S.A.: Texas (Todd, 1943); Nebraska (Todd, 1944); Kansas (Dobrovolny, 1933; Dobrovolny and Ackert, 1934); North Carolina (Hatcher, 1939); Iowa, North Dakota, Michigan (Hoffman, 1953). Czechoslovakia (Groschaft, 1956). England (Lee, 1958a).

Cockroach, Venezuela (Tejera, 1926).

Cockroaches (Blatta orientalis, Blattella germanica, and/or Periplaneta americana), U.S.A. (McAdow, 1931).

Chitwood (1932) also listed China for distribution of the worm, but we could not tell which host was involved.

The worms are found in the colon and rectum of the host. Galeb (1878) found as many as 20 individuals in a single B. orientalis. Sobolev (1937) found 52 percent of B. orientalis infected with L. appendiculata; the mean number of worms per roach was 1.5 and the maximum 2. Dobrovolny and Ackert (1934) found 69 percent of 222 P. americana infected with this species.

Two molts occur within the egg during development of the larva. The first molt occurs outside the host resulting in the formation of an infective resting stage. The second molt occurs inside the cockroach (Todd, 1941, 1944).

Transmission of the nematode is direct, eggs in the resting embryonated stage being infective (Dobrovolny and Ackert, 1934).

Hoffman (1953) described a filamentous bacterium, Streptomyces leidynematis Hoffman, which grows on the cuticle of L. appendiculata in P. americana. Leidy (1853) noted the presence of simple, inarticulate, amorphous filaments, growing from nematodes infecting B. orientalis. Bütschli (1871) and Magalhães (1900) described similar filaments adhering to the surface of oxyurids from cockroaches.

=Leidynema appendiculata= (Leidy, 1850) Chitwood, 1932?

Natural host.--Eurycotis floridana, U.S.A., Massachusetts (Roth and Willis, unpublished data, 1955): Determined by Dr. G. Steiner who wrote us, "In Eurycotis floridana there were ten specimens of the nematode Leidynema appendiculata (Leidy, 1850). This cockroach is obviously a new host for this nematode. I am not sure that the nematode exactly agrees with the description as given in the literature."

=Leidynema cranifera= Chitwood, 1932

Natural hosts.--Blaberus craniifer, U.S.A., Florida (Chitwood, 1932); Massachusetts (Roth and Willis, unpublished data, 1955). Determined by Dr. G. Steiner.

Blaberus atropos?, U.S.A., Florida (Chitwood, 1932): B. craniifer has generally been recorded as B. atropos of Stoll which is a closely related but distinct South American species (Rehn and Hebard, 1927).

=Leidynema delatorrei= Chitwood, 1932

Natural host.--Leucophaea maderae, Cuba, Havana (Chitwood, 1932).

=Leidynema nocalum= Chitwood and Chitwood, 1934

Natural host.--Panesthia angustipennis, Philippine Islands (Chitwood and Chitwood, 1934).

=Leidynemella fusiformis= Cobb, 1934

Natural hosts.--Panesthia laevicollis?, Philippine Islands (Cobb in Chitwood and Chitwood, 1934).

Panesthia angustipennis, Philippine Islands (Chitwood and Chitwood, 1934).

=Leidynemella panesthiae= (Galeb, 1878) Chitwood and Chitwood, 1934

Synonymy.--Oxyuris panesthiae Galeb, 1878, in part; Thelastoma panesthiae (Galeb, 1878) Travassos, 1929. [Chitwood, 1932; Chitwood and Chitwood, 1934.]

Natural host.--Panesthia sp., New Guinea (Galeb, 1878): About 40 nematodes may be found in one insect.

=Leidynemella paracranifera= Chitwood and Chitwood, 1934

Natural host.--Panesthia angustipennis, Philippine Islands (Chitwood and Chitwood, 1934).

=Oxyuris= (?) =heterogamiae= Galeb, 1878

Synonymy.--Thelastoma heterogamiae (Galeb, 1878) Travassos, 1929. The taxonomic position of this species is questionable; it might possibly belong in Blatticola or Protrellina (Chitwood, 1932). Basir (1956) placed it in an appendix to the family Thelastomatidae.

Natural host.--Polyphaga aegyptiaca, Egypt? (Galeb, 1878).

=Protrelleta floridana= Chitwood, 1932

Natural host.--Blaberus craniifer, U.S.A., Florida (Chitwood, 1932).

=Protrellus aureus= Cobb, 1920

Synonymy.--The taxonomic position of this nematode is questionable (Chitwood, 1932).

Natural host.--Polyzosteria melanaria?, Australia, New South Wales (Cobb, 1920). [Caudell (in Chitwood, 1932) stated that this host was probably Platyzosteria analis.]

=Protrellus aurifluus= (Chitwood, 1932) Chitwood, 1933

Synonymy.--Protrellina aurifluus Chitwood, 1932.

Natural hosts.--Parcoblatta lata, U.S.A., North Carolina, Maryland (Chitwood, 1932).

Parcoblatta uhleriana, North Carolina (Hatcher, 1939).

=Protrellus australasiae= (Pessôa and Corrêa, 1926) Travassos, 1929

Synonymy.--Oxyuris australasiae Pessôa and Corrêa, 1926; Protrellina australasiae (Pessôa and Corrêa, 1926) Chitwood, 1932 [Chitwood, 1933].

Natural host.--Periplaneta australasiae, Brazil (Pessôa and Corrêa, 1926, 1927).

=Protrellus galebi= Schwenck, 1926

Synonymy.--Protrellina galebi (Schwenck, 1926) Chitwood, 1932 [Chitwood, 1933].

Natural host.--"Barata selvagem," Brazil (Schwenck, 1926).

=Protrellus künckeli= (Galeb, 1878) Travassos, 1929

Synonymy.--Oxyuris künckeli Galeb, 1878; Protrellina künckeli (Galeb, 1878) Chitwood, 1932 [Chitwood, 1933].

Natural hosts.--Periplaneta americana, Egypt? (Galeb, 1877, 1878). [Chitwood (1932) questioned the determination of this host because he failed to find this nematode in a large number of specimens from U.S.A. and China.] Brazil (Pessôa and Corrêa, 1926).

Periplaneta australasiae, Brazil (Pessôa and Corrêa, 1926).

=Protrellus manni= (Chitwood, 1932) Chitwood, 1933

Synonymy.--Protrellina manni Chitwood, 1932.

Natural host.--Aglaopteryx diaphana, Cuba (Chitwood, 1932).

=Protrellus phyllodromi= (Basir, 1942) Basir, 1956

Synonymy.--Protrellina phyllodromi Basir, 1942.

Natural host.--Blattella humbertiana, India, Aligarh (Basir, 1942): Found in the rectum.

=Protrelloides paradoxa= Chitwood, 1932

Natural host.--Eurycotis floridana, U.S.A., Florida (Chitwood, 1932).

=Schwenkiella icemi= (Schwenck, 1926) Basir, 1956

Synonymy.--Bulhõesia icemi Schwenck, 1926; Thelastoma icemi (Schwenck, 1926) Travassos, 1929; Thelastoma aligarhica Basir, 1940. [Basir, 1956.]

Natural hosts.--"Barata selvagem," Brazil, São Paulo (Schwenck, 1926).

Periplaneta americana, India, Aligarh (Basir, 1940); U.S.A., Nebraska (Todd, 1943).

Periplaneta brunnea, U.S.A., Louisiana (Todd, 1943).

=Severianoia magna= Pereira, 1935

Natural host.--"Blattidae sylvestres," Brazil (Pereira, 1935).

=Severianoia severianoi= (Schwenck, 1926) Travassos, 1929

Synonymy.--Bulhõesia severianoi Schwenck, 1926 [Travassos, 1929].

Natural hosts.--"Baratas de pau podre," Brazil (Schwenck, 1926).

Pycnoscelus surinamensis, U.S.A., Florida (Chitwood, 1932).

=Suifunema caudelli= Chitwood 1932

Natural host.--Steleopyga? sinensis, Asia: Suifu, Szchuen, China (Chitwood, 1932).

=Thelastoma pachyjuli= (Parona, 1896) Travassos, 1929

Synonymy.--Oxyuris bulhõesi de Magalhães, 1900; Bulhõesia bulhõesi (Magalhães, 1900) Schwenck, 1926 [Travassos, 1929; Chitwood, 1932]; Thelastoma bulhõesi (Magalhães, 1900) Travassos, 1929; although this last combination (from Chitwood, 1932) is not given by Basir (1956), it is implied by the synonymy that he does cite under T. pachyjuli.

Natural hosts.--Blatta orientalis, Czechoslovakia (Groschaft, 1956).

Periplaneta americana, Brazil (Magalhães, 1900); North America (Chitwood, 1932); U.S.A., North Carolina (Hatcher, 1939).

=Thelastoma palmettum= Chitwood and Chitwood, 1934

Natural host.--Panesthia angustipennis, Philippine Islands (Chitwood and Chitwood, 1934).

=Thelastoma riveroi= Chitwood, 1932

Natural host.--Periplaneta sp., Cuba (Chitwood, 1932).

=Undetermined nematodes=

Natural host.--Cutilia sp. near sedilloti, U.S.A. (hosts imported from New Zealand) (Roth, unpublished data, 1957).

Class NEMATOMORPHA

Order GORDIOIDEA

The immature stages of the following gordian worms have been found in the body cavity of cockroaches.

Family CHORDODIDAE

=Chordodes morgani= Montgomery, 1898

Synonymy.--Chordotes puerilis Montgomery, 1898 [Ward, 1918].

Natural host.--Cockroach, U.S.A. (Montgomery, 1898); Pennsylvania, Maryland, Michigan, Ohio, Florida, Iowa, Nebraska (Ward, 1918).

Family GORDIIDAE

=Gordius aquaticus= Linnaeus, 1758

Natural host.--Blatta sp., U.S.A. (Stiles and Hassall, 1894).

Leidy (1879) identified a 9-inch-long nematode which came from a cockroach (Blatta orientalis?) as probably being Gordius aquaticus. Ransom (in Pierce, 1921) states that G. aquaticus may be an accidental parasite of man. Faust (1955) summarizes the few reported cases of human parasitism. Dorier (1930) reported that the regurgitated liquid of Blatta orientalis had no effect on cysts of G. aquaticus after one hour.

=Gordius blattae orientalis= Diesing, 1851

Synonymy.--Gordius orientalis of Lankester (1865).

Natural host.--Blatta orientalis, Germany (Siebold, 1842; Linstow, 1878): Found in abdomen. Von Siebold called this "Filarien" but did not otherwise name or describe the worm.

=Gordius pilosus= (Möbius, 1855) Diesing, 1861

Synonymy.--Chordodes pilosus Möbius, 1855 [Diesing, 1861.]

Natural host.--Blaberus giganteus, Venezuela (Möbius, 1855): From the insect's abdomen.

=Gordius= sp.

Natural hosts.--Periplaneta americana, South Africa (Porter, 1930); Germany (Bode, 1936): Bode's record may have referred to a Mermis or other nematode.

Cockroaches, Venezuela (Miall and Denny, 1886; Burr, 1899a; Tejera, 1926).

=Parachordodes raphaelis= (Camerano, 1893) Camerano, 1897

Synonymy.--Gordius raphaelis Camerano, 1893 [Camerano, 1897].

Natural hosts.--Symploce parenthesis and Kuchinga hemerobina, French Equatorial Africa (Camerano, 1893, 1897).

=Undetermined gordian worms=

Natural hosts.--Eurycotis floridana, Florida (T. Eisner, personal communication, 1958): See plate 29, B.

Parahormetica bilobata, Brazil (Pessôa and Corrêa, 1929): Worm referred to as "gordiaceo."

Cockroaches, Australia (E. F. Riek, personal communication, 1953): Three undescribed gordian worms were found in undetermined cockroaches of the subfamily Blattinae.

HELMINTHS FOR WHICH COCKROACHES SERVE AS INTERMEDIATE HOSTS

The use of the asterisk (*) is explained in footnote 3, page 4.

Phylum ACANTHOCEPHALA

Order ARCHIACANTHOCEPHALA

Family OLIGACANTHORHYNCHIDAE

* =Prosthenorchis elegans= (Diesing, 1851) Travassos, 1915

Natural host.--Blattella germanica, France (Brumpt and Urbain, 1938, 1938a; Brumpt et al., 1939).

Experimental hosts.--Blaberus atropos and Leucophaea maderae, France (Brumpt and Desportes, 1938).

* =Prosthenorchis spirula= (Olfers in Rudolphi, 1819) Travassos, 1917

Natural host.--Blattella germanica, France (Brumpt and Urbain, 1938, 1938a; Brumpt et al., 1939); Netherlands (Thiel and Wiegand Bruss, 1946).

Experimental hosts.--Blattella germanica, Netherlands (Thiel and Wiegand Bruss, 1946).

Blaberus atropos and Leucophaea maderae, France (Brumpt and Desportes, 1938).

Family MONILIFORMIDAE

* =Moniliformis dubius= Meyer, 1932

Natural hosts.--Periplaneta americana, Brazil (Magalhães, 1898; Travassos, 1917); Gold Coast (Southwell, 1922); India (Pujatti, 1950); U.S.A. (Burlingame and Chandler, 1941; Moore, 1946).

Periplaneta australasiae, India (Pujatti, 1950).

Experimental hosts.--Blattella germanica, Japan (Yamaguti and Miyata, 1942).

Periplaneta americana, U.S.A. (Chandler, 1941; Moore, 1946); Japan (Yamaguti and Miyata, 1942).

* =Moniliformis kalahariensis= Meyer, 1931

Natural host.--Blattella germanica, India (Meyer, 1931, 1932).

* =Moniliformis moniliformis= (Bremser in Rudolphi, 1819) Travassos, 1915

Natural hosts.--Periplaneta americana, Argentina (Bacigalupo, 1927, 1927a, 1928); Brazil (Pessôa and Corrêa, 1929); Algeria (Seurat, 1912); Burma (Subramanian, 1927); South Africa (Porter, 1930); Madras (Sita, 1949).

Periplaneta spp., New Caledonia (Rageau, 1956).

Cockroaches, Venezuela (Tejera, 1926).

Experimental hosts.--Blaberus atropos, Blatta orientalis, Blattella germanica, Leucophaea maderae, France (Brumpt and Urbain, 1938a).

Periplaneta americana, Japan (Yamaguti and Miyata, 1942); France (Brumpt, 1949); Madras (Sita, 1949).

Phylum ASCHELMINTHES

Class NEMATODA

Order OXYUROIDEA

Family SUBULURIDAE

* =Subulura jacchi= (Diesing, 1861) Railliet and Henry, 1914

Synonymy.--Subulura jacchi (Marcel, 1857) [Dr. J. T. Lucker, personal communication, 1957].

Experimental host.--Blaberus atropos, France (Chabaud and Larivière, 1955).

Order SPIRUROIDEA

Family THELAZIIDAE

* =Oxyspirura mansoni= (Cobbold, 1879) Ransom, 1904

Natural hosts.--Pycnoscelus surinamensis, Australia (Fielding, 1926, 1927, 1928, 1928a); U.S.A. (Sanders, 1927, 1928, 1929; Shealy, 1927); Formosa (Kobayashi, 1927); Antigua (Hutson, 1938, 1943); Hawaii (Illingworth, 1931; Schwabe, 1950, 1950a, 1950b, 1951); New Caledonia (Rageau, 1956).

We have recently found (Roth and Willis, 1960) that two strains of Pycnoscelus surinamensis exist; a parthenogenetic strain (from Florida), and a bisexual strain (from Hawaii) which does not reproduce parthenogenetically. The parthenogenetic strain is undoubtedly the form that has been shown to be the host of O. mansoni in the United States and Antigua, because only this form is found in the New World. Probably the parthenogenetic strain was the form involved in most Pacific areas. However, from internal evidence in his papers, we concluded that Schwabe, in Hawaii, may well have been working with the bisexual strain and possibly also with the parthenogenetic strain; if this is true, then both parthenogenetic and bisexual strains of Pycnoscelus surinamensis may serve as intermediate hosts of the eyeworm.

Experimental hosts.--Periplaneta americana, Antigua (Hutson, 1943).

Pycnoscelus surinamensis, U.S.A. (Sanders, 1929); Australia (Fielding, 1927, 1928a); Hawaii (Schwabe, 1951).

=Rictularia coloradensis= Hall, 1916

Natural hosts.--Parcoblatta pensylvanica and Parcoblatta virginica, U.S.A., Ohio (Oswald, 1958): Of 49 wood roaches collected, one of each species contained a single larva each.

Experimental hosts.--Blatta orientalis, Blattella germanica, Parcoblatta pensylvanica, Parcoblatta virginica, Periplaneta americana, and Supella supellectilium, U.S.A. (Oswald, 1958): The larvae underwent normal development in all species of cockroaches except B. orientalis and P. americana in which cysts developed that contained a reddish-brown pigment; larvae in such cysts were dead or dying. Eggs of R. coloradensis hatched in the midgut of B. germanica and first-stage larvae entered the hindgut epithelium within 24 hours. The larvae underwent two molts within a cyst formed by tissues of the host's gut, the second molt occurring during the twelfth or thirteenth day. In Parcoblatta, cysts were found free in the body cavity as well as attached to the hindgut. In B. germanica and S. supellectilium the cysts remained attached to the hindgut. Usually over 20 cysts developed in each infected Parcoblatta; fewer than 10 per insect developed in the other species. Larvae became infective to the definitive host, the white-footed mouse [Peromyscus leucopus noveboracensis (Fischer)], as early as the tenth day.

Family SPIRURIDAE

* =Agamospirura parahormeticae= Pessôa and Corrêa, 1929

Natural host.--Parahormetica bilobata, Brazil (Pessôa and Corrêa, 1929).

* =Gongylonema ingluvicola= Ransom, 1904

Experimental host.--Blattella germanica, U.S.A. (Cram, 1935).

* =Gongylonema neoplasticum= (Fibiger and Ditlevsen, 1914) Ransom and Hall, 1916

Natural hosts.--Blatta orientalis, Netherlands (Baylis, 1925).

Blattella germanica, U.S.A. (Hitchcock and Bell, 1952).

Periplaneta americana, Denmark and St. Croix (Fibiger, 1913, 1913a; Fibiger and Ditlevsen, 1914); Netherlands (Baylis, 1925); Argentina (Bacigalupo, 1930); England (Leiper, 1926); South Africa (Porter, 1930); U.S.A. (Hitchcock and Bell, 1952); Formosa (Yokagawa, 1924, 1925, 1925a).

Periplaneta australasiae, Formosa (Yokagawa, 1924, 1925, 1925a).

Experimental hosts.--Blattella germanica, Denmark (Fibiger and Ditlevsen, 1914); U.S.A. (Hitchcock and Bell, 1952); France (Brumpt, 1949).

Blatta orientalis, Denmark (Fibiger and Ditlevsen, 1914).

Periplaneta americana, Denmark and St. Croix (Fibiger, 1913; Fibiger and Ditlevsen, 1914); U.S.A. (Hitchcock and Bell, 1952).

* =Gongylonema pulchrum= Molin, 1857

Experimental hosts.--Blattella germanica, U.S.A. (Ransom and Hall, 1915, 1916, 1917; Stiles and Baker, 1927; Schwartz and Lucker, 1931; Lucker, 1932; Alicata, 1934a, 1935); Europe (Baylis et al., 1925, 1926, 1926a; Sambon, 1926).

Parcoblatta sp., Alicata (1934, 1935).

* =Gongylonema= sp.

Natural host.--Periplaneta americana, Brazil (Magalhães, 1900); Algeria (Seurat, 1916); England? (Leiper, 1926).

* =Microtetrameres helix= Cram, 1927

Experimental host.--Blattella germanica, U.S.A. (Cram, 1934).

* =Protospirura bonnei= Ortlepp, 1924

Natural host.--Leucophaea maderae, Venezuela (Brumpt, 1931).

Experimental hosts.--Blatta orientalis, Blattella germanica, Leucophaea maderae, France (Brumpt, 1931).

* =Protospirura columbiana= Cram, 1926

Experimental host.--Blattella germanica, U.S.A. (Cram, 1926).

* =Protospirura muricola= Geodoelst, 1916

Natural host.--Leucophaea maderae, Panama (Foster and Johnson, 1938, 1939).

* =Seurocyrnea colini= (Cram, 1927) Cram, 1931

Experimental host.--Blattella germanica, U.S.A. (Cram, 1931, 1931a, 1933a).

* =Spirura gastrophila= (Müller, 1894) Seurat, 1913

Natural hosts.--Blatta orientalis, Europe? (Deslongchamps, 1824, in Seurat, 1911); Italy (Grassi, 1888); Algeria (Seurat, 1916).

Periplaneta americana, Brazil (Pessôa and Corrêa, 1929).

Cockroach, Venezuela (Tejera, 1926).

Experimental hosts.--Blatta orientalis, France (Galeb, 1878a); "Cafards," Algeria (Roger, 1906, 1907).

* =Tetrameres americana= Cram, 1927

Natural host.--Blattella germanica, U.S.A. (Cram, 1931b, personal communication, 1956); Hawaii (Alicata, 1938, 1947).

Experimental host.--Blattella germanica, U.S.A. (Cram, 1931b).

* =Tetrameres pattersoni= Cram, 1933

Experimental host.--Blattella germanica, U.S.A. (Cram, 1933).

Family PHYSALOPTERIDAE

* =Physaloptera hispida= Schell, 1950

Experimental host.--Blattella germanica, U.S.A. (Schell, 1952, 1952a).

* =Physaloptera maxillaris= Molin, 1860

Experimental host.--Blattella germanica, U.S.A. (Hobmaier, 1941).

* =Physaloptera praeputialis= von Linstow, 1889

Experimental host.--Blattella germanica, U.S.A. (Petri and Ameel, 1950).

* =Physaloptera rara= Hall and Wigdor, 1918

Experimental host.--Blattella germanica, U.S.A. (Petri and Ameel, 1950; Petri, 1950).

* =Physaloptera turgida= Rudolphi, 1819

Experimental host.--Blattella germanica, U.S.A. (Alicata, 1937; Schell, 1952).

HELMINTHS WHOSE EGGS HAVE BEEN CARRIED BY COCKROACHES

The use of the asterisk (*) is explained in footnote 3, page 4.

Phylum PLATYHELMINTHES

Class TREMATODA

Order DIGENEA

Family SCHISTOSOMATIDAE

* =Schistosoma haematobium= (Bilharz, 1852) Weinland, 1858

Experimental vector.--Periplaneta americana, Gold Coast Colony (Macfie, 1922).

Class CESTODA

Order TAENIOIDEA

Family HYMENOLEPIDIDAE

* =Hymenolepis= sp.

Natural vectors.--Periplaneta americana, Formosa (Morischita and Tsuchimochi, 1926).

Polyphaga saussurei, Tadzhikistan (Zmeev, 1936).

Family TAENIIDAE

* =Taenia saginata= Goeze, 1782

Experimental vector.--Periplaneta americana, Gold Coast Colony (Macfie, 1922).

=Echinococcus granulosis= (Batsch, 1786) Rudolphi, 1805

Synonymy.--Taenia echinococcus (Zeder, 1803) [Faust, 1939].

Experimental vector.--Periplaneta americana, Uruguay (Pérez Fontana, 1955): Eggs were recovered from the feces of artificially infested cockroaches under "natural" conditions.

Family Unknown

* =Undetermined tapeworm ova=

Natural vector.--Polyphaga saussurei, Tadzhikistan (Zmeev, 1936).

Phylum ASCHELMINTHES

Class NEMATODA

Order OXYUROIDEA

Family OXYURIDAE

* =Enterobius vermicularis= (Linnaeus, 1758) Leach in Baird, 1853

Natural vectors.--Blatta orientalis and Blattella germanica, U.S.S.R. (Sondak, 1935).

Order ASCAROIDEA

Family ASCARIDAE

* =Ascaris lumbricoides= Linnaeus, 1758

Natural vector.--Periplaneta americana, South Africa (Porter, 1930): The eggs may have been those of A. suum Goeze, 1782.

Experimental vectors.--Periplaneta americana, Gold Coast Colony (Macfie, 1922); India (Chandler, 1926).

Periplaneta americana, Periplaneta australasiae, Neostylopyga rhombifolia, Formosa (Morischita and Tsuchimochi, 1926).

* =Ascaris= sp.

Natural vector.--Blatta orientalis, Italian Somaliland (Mariani and Besta, 1936).

Experimental vector.--Periplaneta americana, Uruguay (Pérez Fontana, 1955): Eggs recovered from the insects' feces.

Order STRONGYLOIDEA

Family ANCYLOSTOMIDAE

* =Ancylostoma caninum= (Ercolani, 1859) Hall, 1913

Experimental vector.--Periplaneta americana, Netherlands (Akkerman, 1933).

* =Ancylostoma ceylanicum= (Looss, 1911) Leiper, 1915

Experimental vector.--Periplaneta americana, Gold Coast Colony (Macfie, 1922); Netherlands (Akkerman, 1933).

* =Ancylostoma duodenale= (Dubini, 1843) Creplin, 1845

Natural vector.--Periplaneta americana, South Africa (Porter, 1929, 1930).

Experimental vector.--Periplaneta americana, Gold Coast Colony (Macfie, 1922).

* =Necator americanus= (Stiles, 1902) Stiles, 1906

Natural vector.--Periplaneta americana, India (Chandler, 1926).

Experimental vector.--Periplaneta americana, Gold Coast Colony (Macfie, 1922).

* =Hookworm ova=

Experimental vectors.--Periplaneta americana, Periplaneta australasiae, Neostylopyga rhombifolia, Formosa (Morischita and Tsuchimochi, 1926).

Family TRICHOSTRONGYLIDAE

* =Trichostrongylus= sp.

Natural vector.--Blatta orientalis, Italian Somaliland (Mariani and Besta, 1936).

Order TRICHUROIDEA

Family TRICHURIDAE

* =Capillaria hepatica= (Bancroft, 1893) Travassos, 1915

Experimental vector.--Blatta orientalis, Italy? (Giordano, 1950).

* =Trichuris trichiura= (Linnaeus, 1771) Stiles, 1901

Natural vectors.--Blatta orientalis, Italian Somaliland (Mariani and Besta, 1936); U.S.S.R. (Sondak, 1935).

Blattella germanica, U.S.S.R. (Sondak, 1935).

Periplaneta americana, Gold Coast Colony (Macfie, 1922); Formosa (Morischita and Tsuchimochi, 1926).

Experimental vectors.--Periplaneta americana, Gold Coast Colony (Macfie, 1922); India (Chandler, 1926); Uruguay (Pérez Fontana, 1955).

Periplaneta americana, Periplaneta australasiae, and Neostylopyga rhombifolia, Formosa (Morischita and Tsuchimochi, 1926).

XII. ARTHROPODA

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.

=Mastigoproctus= sp.

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 arizonensis= Ewing

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 novaehollandiae= Peters

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 =Avicularia= sp.?

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 indistinctus= Pickard-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 helluo= Walckenaer

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).

=Lycosa= sp.

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 nauphoetae= Womersley

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.

=Hypoaspis= sp.

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

=Uropoda= sp.

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.

=Caloglyphus= sp.

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 noxius= A. 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 tarsalus= Banks

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

Family GLYCIPHAGIDAE

=Chaetodactylus= sp.

Synonymy.--Trichotarsus sp. [Baker and Wharton, 1952].

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

Family PODAPOLIPODIDAE

=Locustacarus= sp.

Natural hosts.--Diploptera punctata and Nauphoeta cinerea, U.S.A. (Roth and Willis, unpublished data, 1954): Mite genus determined by Dr. E. W. Baker (personal communication, 1954). The mites cluster thickly on intersegmental membranes, particularly around the coxae and neck. Despite a heavy infestation, the colony of Nauphoeta thrived for several years. This mite was found first on N. cinerea and possibly transferred to D. punctata when the latter was brought into the laboratory from Hawaii.

Family IOLINIDAE

=Iolina nana= Pritchard

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

Family PTERYGOSOMIDAE

=Pimeliaphilus podapolipophagus= Trägårdh

Common name.--Cockroach mite.

Natural hosts.--Parcoblatta sp., U.S.A. (Edmunds, 1953a).

Periplaneta americana, U.S.A. (Piquett and Fales, 1952).

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

Experimental hosts.--Blatta orientalis, Blattella germanica, and Periplaneta americana, U.S.A. (Cunliffe, 1952).

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

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

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

RECORDS OF UNIDENTIFIED MITES

Natural hosts.--Aglaopteryx facies, Puerto Rico (Seín, 1923): Four red "tick" nymphs found under wings of female.

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

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.

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

CONTROL OF MITES IN COCKROACH COLONIES

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

Class CHILOPODA

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

Order SCUTIGEROMORPHA

Family SCUTIGERIDAE

=Scutigera coleoptrata= (Linnaeus)

Synonymy.--Scutigera forceps Rafinesque [Crabill, 1952].

Common name.--House centipede.

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

Experimental prey.--Blattella germanica, newly hatched nymphs and adult female, U.S.A. (Snodgrass, 1930; Roth and Willis, unpublished data, 1953).

Periplaneta americana, U.S.A. (Roth and Willis, unpublished data, 1953).

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

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

=Allothereua maculata= (Newport)

Synonymy.--Scutigera maculata [Crabill, personal communication, 1957].

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

Order SCOLOPENDROMORPHA

Family SCOLOPENDRIDAE

=Scolopendra cingulata= Latreille

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

=Scolopendra morsitans= Linnaeus

Natural prey.--Cockroaches, Guadeloupe (Lherminier, 1837).

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

=Scolopendra subspinipes= Leach

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

=Scolopendra= sp.

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

Class INSECTA

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

Order ODONATA

Family AESHNIDAE

=Anax strenuus= Hagen

Common name.--Giant Hawaiian dragonfly.

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

Order BLATTARIA

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

Order ORTHOPTERA

Family MANTIDAE

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

(Pl. 32)

Experimental prey.--Blatta orientalis, Diploptera punctata, Eurycotis floridana, Leucophaea maderae, Nauphoeta cinerea, Neostylopgya rhombifolia, and Periplaneta americana, U.S.A. (Rilling, personal communication, 1957): Mrs. Rilling wrote us that with the exception of N. rhombifolia, all the above cockroaches were readily eaten. All the mantids initially rejected N. rhombifolia after grasping and making a brief attempt to chew the cockroaches. However, if specimens of N. rhombifolia were left in the jars with the mantids, the cockroaches were usually eaten within the next 24 hours. N. rhombifolia ejects an odorous substance when seized and the mantids probably ate these insects after most of this secretion had been depleted. It is highly probable that the secretion of N. rhombifolia may deter the mantid's attack, but it should be pointed out that, with the possible exception of N. cinerea, all the other species fed to these mantids give off odorous substances when seized or disturbed. Apparently, certain naturally repellent compounds will deter this mantid, whereas others that are presumed to be repellent will not; however, the nutritional state of the mantid is undoubtedly a factor which may limit the effectiveness of certain repellent secretions against this predator.

Byrsotria fumigata, teneral males, and Periplaneta australasiae, nymphs, U.S.A. (Roth and Willis, unpublished data, 1958).

Diploptera punctata, U.S.A. (Eisner, 1958).

=Mantis religiosa= Linnaeus

Common name.--European mantis.

Experimental prey.--Nauphoeta cinerea, and Periplaneta americana, U.S.A. (Rilling, personal communication, 1957).

=Metallyticus semiaeneus= Westwood

Experimental prey.--Cockroaches, Borneo (Shelford, 1916).

=Sphodromantis viridus= (Forskål)

Synonymy.--Sphodromantis bioculata Burmeister [Gurney, personal communication, 1958].

Experimental prey.--Blatta orientalis, Egypt (Adair, 1923): This species of cockroach was apparently used regularly as food for the mantid in the laboratory.

=Stagmomantis carolina= (Johansson)

Common name.--Carolina mantis.

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

Blatta orientalis, nymphs, and Diploptera punctata, U.S.A. (Roth and Willis, unpublished data, 1953).

=Tarachodes maurus= (Stal)

Experimental prey.--Cockroaches, South Africa (Faure, 1940).

=Tenodera aridifolia sinensis= Saussure

Common name.--Chinese mantis.

Experimental prey.--Nauphoeta cinerea and Periplaneta americana, U.S.A. (Rilling, personal communication, 1957).

Family GRYLLACRIDIDAE

=Diestrammena apicalis= Br. v. Wattenwyl

and

=Diestrammena japanica= Blatchley

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

Experimental prey.--Eggs of Blattella germanica and Periplaneta japanica, Japan (Asano, 1937): Seven eggs of B. germanica (obtained from an oötheca being carried by a female) and eggs of P. japanica (presumably in oöthecae) were fed to both species of stone crickets in the evening. The eggs were devoured by the next morning.

Order DERMAPTERA

Family FORFICULIDAE

=Undetermined earwigs=

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

Order HEMIPTERA

Family LYGAEIDAE

=Clerada apicicornis= Signoret

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

Family REDUVIIDAE

=Spiniger domesticus= Pinto

Natural prey.--Periplaneta americana, Brazil, Matto Grosso (Pinto, 1927, 1927a): This bug preys principally on cockroaches and was observed infesting the walls of dwellings where it preyed on P. americana.

=Triatoma arthurneivai= Lent and Martins

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

=Undetermined reduviids=

Natural prey.--Arenivaga roseni and Polyphaga saussurei, Turkmen S.S.R. (Vlasov and Miram, 1937): These desert cockroaches are found in burrows of rodents and desert turtles around Ashkhabad. Reduviids are their main enemies. Vlasov (1933) found nymphs of Reduvius christophi Jak. and R. fedtschenkianus Osch. in similar burrows in this same area, although he did not specifically cite them as enemies of the desert cockroaches.

Family NEPIDAE

=Ranatra= sp.

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

Order NEUROPTERA

Family ASCALAPHIDAE

=Undetermined larva=

Experimental prey.--Blattella germanica, Kenya Colony (Someren, 1924).

Order DIPTERA

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

Family PHORIDAE

=Megaselia= sp.

Host.--Eggs of Parcoblatta sp., Ohio (Edmunds, 1952a).

Family CONOPIDAE

=Stylogaster stylata= (Fabricius)

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

=Stylogaster= spp.

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

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

Family LARVAEVORIDAE

=Calodexia= (?) =venteris= Curran

Hosts.--Periplaneta americana, Brazil (Souza Lopes, 1937): Obtained complete evolution of the parasite in this host. This may have been an experimental host.

=Calodexia= spp.

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

=Undetermined tachinids=

Hosts.--Eurycotis floridana, from Florida (Roth, unpublished data, 1953): Three larvae (det. by W.W. Wirth) were found in a living adult male.

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

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

Family MUSCIDAE

=Coenosia basalis= Stein

Host.--Eggs of Parcoblatta sp., Ohio (Edmunds, 1952a).

Family SARCOPHAGIDAE

=Sarcophaga omani= Hall

Host.--Arenivaga bolliana, Texas (Wirth, personal communication, 1953): Specimens in U.S. National Museum.

=Sarcophaga lambens= Wied.

Synonymy.--Sarcophaga sternodontis (Towns.).

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

=Sarcophaga= spp.

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

Order COLEOPTERA

Family CARABIDAE

=Harpalus pennsylvanicus= De Geer

Experimental prey.--Cryptocercus punctulatus, U.S.A. (Cleveland et al., 1934): This beetle is often found in the galleries of C. punctulatus in nature. In the laboratory it killed and devoured cockroaches as large as itself.

Family DYTISCIDAE

=Rhantus pacificus= Boisduval

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

Family LAMPYRIDAE

=Undetermined larva=

Experimental prey.--Parcoblatta virginica, adult female (pl. 33, C), U.S.A. (Roth and Willis, unpublished data, 1953).

Family RIPIPHORIDAE

=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 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 as blattarum because it had been captured in the body of Ectobia livida. We presume that Abeille de Perrin was referring to Sundevall's work in which the host was given as Blattella germanica.

Periplaneta americana, on shipboard (Sundevall, 1831): One nymph only.

With the exception of the single nymph of P. americana, R. pectinicornis apparently attacks only adult females and nymphs of B. germanica. Barbier (1947) found only B. germanica parasitized, although both Blatta orientalis and Supella supellectilium were prevalent on board the ship. Primary larvae of the parasite failed to parasitize Supella.

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

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

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

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

=Ripidius scutellaris= Heller

Natural hosts.--Blattidae, Philippine Islands (Schultze, 1925).

Family DERMESTIDAE

=Dermestes ater= De Geer

Common name.--Black larder beetle.

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

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

=Dermestes= sp.

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

Order STREPSIPTERA

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

Order HYMENOPTERA

PREDATORS AND PARASITES OF COCKROACH EGGS

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

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

Family EVANIIDAE

=Acanthinevania princeps= (Westwood)

Synonymy.--Evania princeps [Dr. H. Townes, personal communication, 1956].

Natural host.--Cockroach eggs, Australia (Froggatt, 1906).

=Brachygaster minutus= (Olivier)

Synonymy.--Evania minuta Olivier [Kieffer, 1920].

Natural hosts.--Blattella germanica, Europe? (Schletterer, 1889; Kiefer, 1912; Crosskey, 1951.)

Ectobius lapponicus, Europe? (Schletterer, 1889; Kieffer, 1912; Crosskey, 1951.)

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

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

Adult wasps have been collected on Asparagus officinalis Linnaeus (Schmiedeknecht in Schletterer, 1889; Crosskey, 1951). Thompson's (1951) citation of records of B. minutus and Evania appendigaster from Blatta orientalis and Blattella germanica, and Cameron's (1957) citation of these records and one from Ectobius lapponicus, all attributed to Kadocsa (1921), are almost certainly in error. Kadocsa (1921, p. 33) listed these wasps as egg parasites of cockroaches but not necessarily in Hungary and did not name specific cockroach hosts.

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

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

=Evania appendigaster= (Linnaeus)

Synonymy.--Evania desjardinsii Bordage, Evania laevigata Latreille [Dalla Torre, 1901-1902].

Natural hosts.--Blatta, "exotic species" (Westwood, 1854, 1954a).

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

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

Cutilia soror, Hawaii (Swezey, 1929; Zimmerman, 1948).

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

Neostylopyga rhombifolia, Hawaii (Swezey, 1929).

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

Periplaneta americana or P. australasiae, Formosa (Sonan, 1924).

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

Experimental host.--Blatta orientalis, U.S.A. (Haber, 1920).

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

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

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

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

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

=Evania dimidiata= Fabricius

Synonymy.--Evania abyssinica Westwood [Schletterer, 1889].

Natural host.--Blatta orientalis, Egypt? (Alfieri, 1914).

=Evania subspinosa= Kieffer

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

=Hyptia dorsalis= of Ashmead

Synonymy.--Dr. H. Townes, (personal communication, 1956) believes that this wasp was probably either H. reticulata, H. harpyoides, or H. thoracica; it is not possible to tell which without reexamining Ashmead's specimens; these apparently have been lost.

Natural host.--Parcoblatta pensylvanica, U.S.A., Mississippi (Ashmead, 1900).

=Hyptia harpyoides= Bradley

Natural hosts.--Parcoblatta virginica, U.S.A., Ohio (Edmunds, 1952a, 1953a, 1954).

Parcoblatta pensylvanica, U.S.A. (Muesebeck, 1958).

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

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

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

=Hyptia reticulata= Say

Natural host.--Parcoblatta pensylvanica, U.S.A., Missouri (Rau, 1940).

Adult wasps have been taken on parsnip, Pastinaca sativa (Robertson, 1928).

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

=Hyptia thoracica= (Blanchard)

Natural host.--Parcoblatta pensylvanica, U.S.A., Ohio (Edmunds, 1952a, 1953a, 1954).

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

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

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

=Hyptia= sp.

Natural host.--Cariblatta delicatula, Cuba (Hebard, 1916a); Parasite identified by Ashmead.

=Hyptia= sp. (undescribed)

Natural host.--Parcoblatta sp., U.S.A., Ohio (Edmunds, 1952a).

=Prosevania punctata= (Brullé)

Synonymy.--Evania punctata Brullé [Townes, 1949].

Natural and experimental hosts.--Blatta orientalis, Istrian Peninsula (Fahringer, 1922); Algeria (Cros, 1942); U.S.A., Ohio (Edmunds, 1954).

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

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

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

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

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

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

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

=Szepligetella sericea= (Cameron)

Synonymy.--Evania sericea Cameron [Townes, 1949, personal communication, 1956]. Evania impressa Schletterer [Townes, p. c., 1956].

Natural hosts.--Cutilia soror and Neostylopyga rhombifolia, Hawaii (Swezey, 1929).

Periplaneta americana and Periplaneta australasiae, Hawaii (Swezey, 1929; Zimmerman, 1948).

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

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

=Zeuxevania splendidula= Costa

Natural hosts.--Loboptera decipiens, France (Lavagne, 1914; Genieys, 1924).

Picard (1913) believed that Z. splendidula parasitized L. decipiens and not its eggs; however, Lavagne (1914) explained the true relationship by dissecting two specimens of Z. splendidula from oöthecae of L. decipiens.

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

Family CLEONYMIDAE

=Agamerion metallica= Girault

Natural hosts.--Ellipsidion australe, Australia, Queensland (Dodd, 1917): "the parasite when ready to emerge fully occupies the whole space of the destroyed eggs."

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

Family ENCYRTIDAE

=Blatticida pulchra= Ashmead

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

=Blatticidella ashmeadi= (Girault)

Synonymy.--Blatticida ashmeadi. Blatticida Girault, 1915, is preoccupied by Blatticida Ashmead, 1904. In 1923 Gahan and Fagan renamed Blatticida Girault, Blatticidella. [Burks, p. c., 1956.]

Natural host.--Cockroach, Australia, Queensland (Girault, 1915).

=Cheiloneurus viridiscutum= (Girault)

Synonymy.--Cristatithorax Girault = Cheiloneurus Westwood [Mercet, 1921].

Natural host.--Ellipsidion australe, Australia, Queensland (Dodd, 1917).

=Comperia merceti= (Compere)

Synonymy.--Comperia merceti var. falsicornis Gomes [Peck, 1951].

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

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

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

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

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

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

=Dicarnosis alfierii= Mercet

Natural hosts.--"Phyllodromia" sp., Egypt (Mercet, 1930): According to Mercet, Dr. Alfieri claimed that this wasp parasitized one of the species of "Phyllodromia" found in Egypt, namely, Phyllodromia [= Blattella] germanica, Phyllodromia [= Supella] supellectilium and/or Phyllodromia treitliana. We do not know to which modern genus the host of this wasp belonged.

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

=Eutrichosomella blattophaga= Girault

Natural host.--Cockroach, Australia, Queensland (Girault, 1915).

Family EUPELMIDAE

=Anastatus blattidifurax= Girault

Natural host.--Cockroach, Australia, Queensland (Girault, 1915).

=Anastatus floridanus= Roth and Willis

Natural host.--Eurycotis floridana, U.S.A., Florida (Roth and Willis, 1954a).

Experimental hosts.--Blatta orientalis, Eurycotis floridana, and Periplaneta americana, U.S.A. (Roth and Willis, 1954a).

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

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

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

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

Number of parasites per oötheca.--In Blatta orientalis: One of 111 oöthecae exposed to female wasps yielded 48 parasites. In Eurycotis floridana: One oötheca parasitized in the field yielded 68 parasites; 8 oöthecae exposed to single wasps for their entire lifespan yielded an average of 50 ± 6 parasites (range 23-81); 34 oöthecae exposed to many wasps for their entire lifespan yielded an average of 198 ± 8 parasites (range 93-306). In Periplaneta americana: Nine oöthecae of 152 exposed to the wasps were found to be parasitized when dissected; 11 adults emerged from one oötheca; no parasites emerged from the other 8 oöthecae.

Sex ratio.--4 :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 tenuipes= Bolívar y Pieltain

Synonymy.--Anastatus blattidarum Ferrière. Dr. C. Ferrière (personal communication, 1957) is of the opinion that his A. blattidarum is a synonym of A. tenuipes. He stated "I have never been able to see the unique type of A. tenuipes B. y P., which is in Madrid, but the description agrees with A. blattidarum. I had not yet knowledge of Bolívar's description, when describing my species. The parasite of cockroaches eggs [Supella supellectilium] should be called A. tenuipes Bol." Mani (1938) synonymized Solindenia blattiphagus Mani with Anastatus blattidarum.

Natural hosts.--Supella supellectilium, Anglo-Egyptian Sudan (Ferrière, 1930, 1935); U.S.A., Arizona (Flock, 1941); Egypt (Alfieri, in Hafez and Afifi, 1956). Ohio (Hull and Davidson, 1958).

Periplaneta americana, India (Burks in Roth and Willis, 1954a).

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

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

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

Sex ratio.--4 :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 atriflagellum= Girault

Natural host.--Blattella germanica, Australia, Queensland (Girault, 1924).

=Eupelmus= sp.

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

=Solindenia picticornis= Cameron

Natural hosts.--Allacta similis, Hawaii (Perkins, 1906, 1913; Timberlake, 1924; Swezey, 1929; Zimmerman, 1948).

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

Family PTEROMALIDAE

=Pteromalus= sp.?

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

=Systellogaster ovivora= Gahan

Natural hosts.--Blatta orientalis, U.S.A., Illinois (Gahan, 1917).

Parcoblatta pensylvanica, Canada, Ontario (Judd, 1955).

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

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

One oötheca of P. pensylvanica yielded 14 parasites with a sex ratio of 2.5 : 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 chalybii= Ashmead

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

=Mestocharomyia oophaga= Dodd

Natural host.--Ellipsidion australe, Australia, Queensland (Dodd, 1917).

=Syntomosphyrum blattae= Burks

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

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

=Syntomosphyrum ischnopterae= (Girault)

Synonymy.--Epomphaloides ischnopterae Girault [Peck, 1951].

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

Natural hosts.--Ischnoptera sp. [probably Parcoblatta sp. (Rehn, personal communication, 1958)]. U.S.A., Maryland (Girault, 1917).

Zeuxevania splendidula Costa (an evaniid in the oöthecae of Loboptera decipiens), France (Parker, 1924; Parker and Thompson, 1928).

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

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

=Tetrastichus australasiae= Gahan

Natural host.--Periplaneta australasiae, Sumatra (Gahan, 1923).

=Tetrastichus hagenowii= (Ratzeburg)

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 periplanetae= Crawford

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).

=Tetrastichus= sp. 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).

=Tetrastichus= sp. 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 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).

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.

=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 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.

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 picea= Emery

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) wilverthi= Emery

Natural prey.--Small cockroaches, Belgian Congo (Raignier and van Boven, 1955).

=Dorylus= sp.

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 humilis= Mayr

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.

XIII. VERTEBRATA

Class PISCES

In British Guiana, Beebe (1925a) found undetermined cockroach remains in the stomachs of four species of fish belonging to three families, as follows:

Family POTAMOTRYGONTIDAE

=Potamotrygon humboldti= (Duméril)

(= Potamotrygon hystrix)

Family PIMELODIDAE

=Rhamdia sebae= Cuvier and Valenciennes

Family CHARACIDAE

=Cyrtocharax magdalenae essequibensis= (Eigenmann)

(= Cynopotamus essequibensis)

=Chalceus macrolepidotus= Cuvier and Valenciennes

The only other records of cockroaches being eaten by fish pertain to the use of cockroaches as bait. Captain William Owen (in Webster, 1834) stated that the Chinese used cockroaches as bait in their fishing excursions. At Reelfoot Lake, Tennessee, Blatta orientalis were kept in large numbers by bait dealers and were sold to fishermen who used them for catching Lepomis pallidus, a sunfish locally known as bream, blue bream, or bluegill (Rau, 1944). In Indiana, oriental cockroaches were collected at a city dump by fishermen (Gould, 1941). Peterson (1956) states that cockroaches are satisfactory bait for bluegills, crappies, channel cat, blue heads, and large mouth black bass.

Class AMPHIBIA

Order CAUDATA

Family PLETHODONTIDAE

=Plethodon glutinosus= (Green)

Natural prey.--Cryptocercus punctulatus, U.S.A. (Honigberg, 1953): Protozoa which are normally only found in C. punctulatus were present in the intestine of the salamander indicating that this cockroach had been eaten by the amphibian.

Order SALIENTIA

Family BUFONIDAE

=Bufo funereus= Bocage

Natural prey.--Cockroaches, Belgian Congo (Noble, 1924): The stomachs of 62 out of 72 specimens contained food; this included 3 cockroaches.

=Bufo ictericus= Spix

Natural prey.--Cockroaches, Brazil (Valente, 1949): Stomach contents revealed the prothorax, legs, and wings of cockroaches, and fragments of wood-cockroaches. This toad frequently feeds at night.

=Bufo marinus= (Linnaeus)

Common name.--Giant toad, marine toad, Surinam toad.

Natural prey.--Epilampra abdomen-nigrum, Trinidad (Weber, 1938): Found in the stomachs of two toads.

Diploptera punctata, Hawaii (Pemberton and Williams, 1938).

Periplaneta sp., Fiji (Lever, 1939): Many householders in Suva have seen the toad eat considerable numbers of these cockroaches.

Pycnoscelus surinamensis, Hawaii (Alicata, 1938; Illingworth, 1941).

Cockroaches, Nicaragua (Noble, 1918): Stomach contents of toads captured at street lamps in Rio Grande consisted chiefly of large cockroaches. Puerto Rico (Wolcott, 1937).

=Bufo valliceps= Wiegmann

Experimental prey.--Periplaneta americana, U.S.A. (Moore, 1946): Cockroaches containing infective acanthellas of Moniliformis dubius were fed to three toads.

Family HYLIDAE

=Hyla cinerea= (Schneider)

Common name.--Green tree frog.

Natural prey.--Ischnoptera deropeltiformis, Periplaneta americana, and undetermined cockroaches, U.S.A., Georgia (Haber, 1926): Cockroaches were found in 11 of 100 stomachs.

Family RANIDAE

=Arthroleptis variabilis= Matschie

Natural prey.--Cockroaches, Belgian Congo (Noble, 1924): Of 52 specimens examined, the stomach contents of 17 contained food, including 3 cockroaches.

=Hyperolius picturatus= Peters

Natural prey.--Cockroach, Belgian Congo (Noble, 1924): The stomachs of 12 of 56 specimens examined contained food, including one cockroach.

=Leptodactylus albilabris= (Günther)

Natural prey.--Cockroach, Puerto Rico (Schmidt, 1920): One of 25 stomachs contained a medium-sized cockroach.

=Leptodactylus pentadactylus= (Laurenti)

Common name.--"Smoky jungle frog" or "pepper frog."

Natural prey.--Cockroaches, Nicaragua (Noble, 1918): Cockroach wings were found in the stomach of a frog caught around human habitation. Brazil (Valente, 1949).

=Leptopelis calcaratus= (Boulenger)

Natural prey.--Cockroaches, Belgian Congo (Noble, 1924): The stomachs of 35 specimens were examined of which 13 contained food, including 2 cockroaches.

=Leptopelis rufus= Reichenow

Natural prey.--Cockroaches, Belgian Congo (Noble, 1924): Forty-five of 83 stomachs examined contained food, including 2 cockroaches.

=Megalixalus fornasinii= (Bianconi)

Natural prey.--Cockroaches, Belgian Congo (Noble, 1924): The stomachs of 3 of 40 specimens contained food, including 2 cockroaches.

=Rana catesbeiana= Shaw

Common name.--Bullfrog.

Natural prey.--Cockroaches, Puerto Rico (Derez, 1949).

=Rana mascareniensis= Duméril and Bibron

Natural prey.--Cockroaches, Belgian Congo (Noble, 1924): The stomach contents of 138 specimens were examined, 39 of which contained food, including 2 cockroaches.

=Rana pipiens= Schreber

Common name.--Leopard frog.

Experimental prey.--Periplaneta americana, U.S.A. (Moore, 1946): Cockroaches containing infective acanthellas of Moniliformis dubius were fed to two frogs.

Neostylopyga rhombifolia, U.S.A. (Dr. T. Eisner, personal communication, 1958.)

=Frogs=

Natural prey.--Blatta orientalis, U.S.A. (Rau, 1924): Frogs which escaped from a tank in the cellar consumed quantities of this cockroach.

Parcoblatta pensylvanica, U.S.A. (Frost, 1924): One adult specimen recovered from alimentary canal of a frog, probably Rana sp.

=Unidentified batrachians=

Experimental prey.--Blattella germanica, Germany, Frankfurt am Main, Zoological Garden (Lederer, 1952): These insects were preferred by all the insect eaters in the zoo.

Periplaneta americana, Germany, Frankfurt am Main, Zoological Garden (Lederer, 1952): Newly molted individuals were accepted as food, but others were usually passed by or consumed unwillingly.

Class REPTILIA

Order CHELONIA

Family EMYDIDAE

=Chrysemys picta= (Schneider)

Common name.--Painted turtle.

Natural prey.--Periplaneta australasiae, England (Lucas, 1916, 1920): The cockroach, apparently injured, fell into water in the tortoise house, Zoological Gardens, Regent's Park, and the terrapin ate it.

Order SAURIA

Family GEKKONIDAE

=Gekko gecko= (Linnaeus)

Natural prey.--Cockroaches, Philippine Islands, Laguna (Villadolid, 1934): The geckos frequent holes in trees and underside of bark which are favorable haunts of cockroaches. Stomach contents mostly Blattidae and "Locustidae."

=Hemidactylus frenatus= Duméril and Bibron

Common name.--House lizard.

Natural prey.--Cockroaches, Philippine Islands, Laguna (Villadolid, 1934): Bulk of stomach contents of 22 lizards consisted of Orthoptera, mostly cockroaches.

=Sphaerodactylus= sp.

Natural prey.--Cockroaches, British Guiana (Beebe, 1925a): The above lizard is found in houses.

=Thecadactylus= sp.

Natural prey.--Cockroaches, British Guiana (Beebe, 1925a): The above lizard is found in houses.

=Undetermined geckos=

Natural prey.--Cockroaches, Australia, Flinders River (Froggatt, 1906): The lizard lived in the walls of the hut and hunted cockroaches upon the roof at night. Arno Atoll (Usinger and La Rivers, 1953).

Family IGUANIDAE

=Anolis carolinensis= Voigt

Experimental prey.--Diploptera punctata, U.S.A. (Eisner, 1958).

=Anolis cristatellus= Duméril and Bibron

Natural prey.--Blattella sp., Cariblatta delicatula, Epilampra wheeleri, Periplaneta americana, Periplaneta australasiae, and Symploce flagellata, Puerto Rico (Wolcott, 1924): The last-named cockroach may have been S. ruficollis Rehn and Hebard, the females of which are hard to distinguish from flagellata. Rehn and Hebard (1927) stated that in all probability flagellata does not occur on the island of Puerto Rico. Wolcott (1950) stated that Symploce ruficollis [= bilabiata] serves as food for the crested lizard.

Cockroaches, Puerto Rico (Schmidt, 1920): Of 100 stomachs examined, 16 contained Orthoptera, including cockroaches. Puerto Rico (Wolcott, 1924): One hundred A. cristatellus had eaten 8 cockroaches, 4.14 percent of the total food, or 25 percent of the food for 8 lizards.

=Anolis pulchellus= Duméril and Bibron

Natural prey.--Cockroaches, Puerto Rico (Wolcott, 1924): Two small cockroaches found in 50 lizards examined.

=Anolis equestris= Merrem

Experimental prey.--Neostylopyga rhombifolia, U.S.A. (Eisner, personal communication, 1958.)

=Anolis grahami= Garman

Natural prey.--Periplaneta spp. and Blattidae, Bermuda (Simmonds, 1958): Stomachs of 176 lizards yielded 6 cockroaches.

=Anolis leachi= Duméril and Bibron

Natural prey.--Periplaneta spp. and Blattidae, Bermuda (Simmonds, 1958): Stomachs of 46 lizards yielded 31 cockroaches.

=Anolis sagrei= Cocteau

Natural and experimental prey.--Pycnoscelus surinamensis, Cuba (Darlington, 1938): This species was eaten both in captivity and in nature. The lizard ate most readily soft, immature cockroaches. Pycnoscelus surinamensis is probably a staple food of the lizard in nature, as Darlington observed wild lizards catch the nymphs.

=Anolis stratulus= Cope

Natural prey.--Aglaopteryx facies, Puerto Rico (Wolcott, 1924): One cockroach was found in 50 lizards examined.

Cockroach, Puerto Rico (Schmidt, 1920): One of 25 stomachs contained a cockroach.

=Anolis= sp.

Natural prey.--"Wood roaches," British Guiana (Beebe, 1925a): The above lizard is arboreal on foliage in low jungle.

Family SCINCIDAE

=Leiolopisma laterale= Say

Common name.--Brown skink.

Natural prey.--Woodroaches, U.S.A., Louisiana (Slater, 1949): Analysis of stomach contents of 84 adult skinks showed that nymphal and adult woodroaches comprised the majority of Orthoptera.

=Tropidophorus grayi= Günther

Common name.--Spiny lizard.

Natural prey.--Cockroaches, Philippine Islands, Laguna (Villadolid, 1934): Food of this species was mostly Blattidae.

=Unidentified skinks=

Natural prey.--Cockroaches, Arno Atoll (Usinger and La Rivers, 1953).

Family AGAMIDAE

Experimental prey?--Cockroaches, Australia (Lee and Mackerras, 1955): A general statement was made that in captivity Agamidae were observed feeding avidly on cockroaches and other insects. Three agamids studied by these workers were Amphibolurus barbatus (Gray), Physignathus lesueurii Gray, and Chlamydosaurus kingii Gray.

Family CHAMAELEONTIDAE

=Chamaeleon chamaeleon= (Linnaeus) and =Chamaeleon oustaleti= Mocquard

Experimental prey.--Cockroaches, Amsterdam (Portielje, 1914): Large cockroaches were fed to these lizards in the reptile house of Artis.

Family TEIIDAE

=Ameiva exsul= Cope

Common name.--Iguana, ground lizard.

Natural prey.--Cockroach (nymph), Epilampra wheeleri, and Periplaneta americana, Puerto Rico (Wolcott, 1924): Stomach contents of 15 lizards were analyzed. E. wheeleri formed 30 percent of the food of one lizard. The cockroach nymph formed 5 percent of the food of one lizard. One P. americana formed 20 percent of the food of one lizard; another formed 50 percent of the food of a second lizard.

Experimental prey.--Cockroach nymphs, Puerto Rico (Wolcott, 1924).

=Ameiva= sp.

Natural prey.--Cockroaches, British Guiana (Beebe, 1925a): The above lizard is terrestrial and found near clearings. The stomach contents of 18 out of 40 reptiles contained cockroach remains.

=Cnemidophorus= sp.

Natural prey.--Cockroaches, British Guiana (Beebe, 1925a): This is a terrestrial lizard found near clearings. The stomach contents of 4 out of 40 lizards contained cockroaches.

=Unidentified lizards=

Natural prey.--Cockroaches, West Indies (H., 1800).

Experimental prey.--Blatta orientalis, U.S.A. (Rau, 1924): Rau called the predator a common gray lizard.

Periplaneta americana, Germany, Frankfurt am Main, Zoological Garden (Lederer, 1952): Newly molted cockroaches were accepted as food, but others were usually passed by or consumed unwillingly.

Order SERPENTES

Family COLUBRIDAE

=Heterodon platyrhinos= Latreille

Synonymy.--Heterodon contortrix [Dr. Doris M. Cochran, personal communication, 1957].

Common name.--Hog-nosed snake.

Experimental prey.--Periplaneta americana, U.S.A. (Moore, 1946): Cockroaches containing infective acanthellas of Moniliformis dubius were fed to one snake.

=Garter Snake=

Experimental prey.--Blatta orientalis, U.S.A. (Rau, 1924).

Class AVES

The cockroach is always wrong when arguing with a chicken.

Spanish proverb (HARTNACK, 1939)

Arboreal cockroaches hidden in and under bark are much more likely to be encountered by birds than by other predators, and insectivorous birds undoubtedly consume many more cockroaches than the few records would indicate. Most of the records we have located identify the birds at least by common name. Where possible we have given the scientific names for those birds whose common names are recognizably specific. We have followed the systematic classification of Wetmore (1940).

Figuier (1869) stated that poultry and owls are very fond of cockroaches. Perkins (1913) made the general statement that some of the native birds of Hawaii are partial to the endemic Allacta similis. Asano (1937) stated that in Japan natural enemies of cockroaches may be found in the Galliformes, Strigiformes, Passeriformes, and Piciformes. Although Lederer (1952) successfully fed newly molted Periplaneta americana to insectivorous birds in the Zoological Garden, Frankfurt am Main, Blattella germanica were preferred by these birds. The following records are of specific birds feeding on cockroaches.

Order ANSERIFORMES

Family ANATIDAE

=Domestic duck=

Natural prey.--Pycnoscelus surinamensis, Australia (Fielding 1926): The ducks became infected with Manson's eye worm of which P. surinamensis is the only known intermediate host.

Cockroach, Bermuda (Jones, 1859): "All kinds of poultry feed greedily upon the cockroach; tame ducks spending entire moonlight nights in their capture."

Order GALLIFORMES

Family PHASIANIDAE

=Bambusicola thoracica= Temminck

Common name.--Kojukei.

Natural prey.--Cockroaches, Japan (Asano, 1937).

=Gallus= sp.

Common name.--Jungle fowl.

Natural prey.--Periplaneta australasiae, Hawaii (Schwartz and Schwartz, 1949).

=Phasianus calchicus karpowi= Buturlin

Common name.--Korean pheasants.

Experimental prey.--Blattella germanica and Periplaneta picea, Japan (Asano, 1937): Adults of these cockroaches were devoured at once when they were fed with the heads cut off.

=Phasianus= sp.

Common name.--Pheasant.

Natural prey.--Blattidae, unidentified (below 1 percent of the diet), Cutila soror (below 1 percent of the diet), Diploptera punctata (above 6 percent of the diet), and Pycnoscelus surinamensis (6 percent of the diet), Hawaii (Schwartz and Schwartz, 1949).

=Coturnix coturnix japonica= (Temminck and Schlegel)

Common name.--Japanese quail.

Natural prey.--Blattidae (unidentified) and Lobopterella dimidiatipes, Hawaii (Schwartz and Schwartz, 1949).

=Domestic chicken=

Natural and experimental prey.--Blaberus craniifer, U.S.A., Key West, Florida. J.A.G. Rehn in 1912 (personal communication) observed chickens feeding on nymphs of B. craniifer which had dropped to the ground from among stacked coffins in an undertaker's shack.

Blatta orientalis, U.S.A. (Rau, 1924): The chickens ate cockroaches that were caught in traps.

Hebardina concinna, Japan (Asano, 1937): Experimental feeding to white Leghorn chickens.

Periplaneta americana, Surinam (Stage, 1947): Several cockroaches ran off the floor of a house, which was being sprayed with DDT, and were eaten by chickens. Although some chickens had DDT tremors the next day, all appeared normal two days later.

Pycnoscelus surinamensis, Australia (Fielding, 1926); Formosa, experimental feeding (Kobayashi, 1927); Australia, experimental feeding (Fielding, 1927, 1928); U.S.A., Florida, experimental feeding (Sanders, 1928); Antigua (Hutson, 1943); Hawaii (Illingworth, 1931; Schwabe, 1949, 1950a, 1950b). This cockroach is the intermediate host of Oxyspirura mansoni, the chicken eye worm.

Cockroaches, Guadeloupe (Dutertre, 1654); Africa (Moiser, 1947): "Poultry" ate cockroaches which had been killed by DDT and sodium fluoride. Hawaii (Zimmerman, 1948).

=Partridge=

Natural prey.--Cockroaches, British Guiana (Beebe, 1925a): The food of two small species of leaf-colored partridges that lived on the jungle floor, consisted chiefly of cockroaches and beetles.

Family MELEAGRIDIDAE

=Meleagris gallopavo= (Linnaeus)

Common name.--Turkey.

Natural prey.--Pycnoscelus surinamensis, Antigua (Hutson, 1943): Turkeys were found heavily infected with Manson's eye worm of which P. surinamensis is the only known intermediate host. These turkeys therefore were presumed to have fed on this cockroach.

Order COLUMBIFORMES

Family COLUMBIDAE

=Streptopelia chinensis= (Scopoli)

Common name.--Chinese dove.

Natural prey.--Pycnoscelus surinamensis, Hawaii (Schwabe, 1950b).

=Pigeon=

Experimental prey.--Pycnoscelus surinamensis, Australia (Fielding, 1927); U.S.A., Florida (Sanders, 1928).

Order STRIGIFORMES

Family STRIGIDAE

=Gymnasio nudipes= (Daudin)

Common name.--Bare-legged owl.

Natural prey.--Epilampra sp., Puerto Rico (Wetmore, 1916): One specimen identified in stomach of a wild-caught owl.

Cockroaches, Puerto Rico (Wetmore, 1916): These insects were found in stomachs of five owls.

Order CORACIFORMES

Family TODIDAE

=Todus mexicanus= Lesson

Common name.--Porto Rican tody.

Natural prey.--Plectoptera poeyi?, Puerto Rico (Wetmore, 1916): The stomachs of 89 birds were examined; a single bird had eaten the above cockroach. According to Wolcott (1950) P. poeyi could be Plectoptera dorsalis, P. rhabdota, or P. infulata.

Family BUCEROTIDAE

=Tockus birostris= (Scopoli)

Synonymy.--Lophocerus birostris [Dr. H. Friedmann, personal communication, 1957].

Common name.--Common gray hornbill.

Natural prey.--Cockroaches, India, Central Provinces (D'Abreu, 1920).

Order PICIFORMES

Family PICIDAE

=Dendrocopus mahrattensis= (Latham)

Synonymy.--Liopicus mahrattensis [Friedmann, p. c. 1957].

Common name.--Yellow-fronted pied woodpecker.

Natural prey.--Cockroaches, India, Central Provinces (D'Abreu, 1920).

=Melanerpes portoricensis= (Daudin)

Common name.--Puerto Rican woodpecker.

Natural prey.--Pycnoscelus surinamensis, Puerto Rico (Wetmore, 1916): One specimen found in 59 bird stomachs examined.

Order PASSERIFORMES

Family FORMICARIIDAE

=Gymnopithys leucaspis= (Sclater)

Common name.--Bicolored antbird.

Natural prey.--Cockroaches, Panama Canal Zone (Johnson, 1954): This bird feeds on small cockroaches, and other arthropods, which are flushed from their hiding places by swarms of the army ant, Eciton burchelli.

Family ORIOLIDAE

=Icterus portoricensis= (Bryant)

Common name.--Puerto Rican oriole.

Natural prey.--Cockroaches, Puerto Rico (Wetmore, 1916): Cockroaches and oöthecae found in the birds' stomachs.

Family CORVIDAE

=Aphelocoma coerulesens= (Bosc)

Common name.--Florida jay.

Experimental prey.--Pycnoscelus surinamensis, U.S.A., Florida (Sanders, 1928).

=Cyanocitta cristata= (Linnaeus)

Common name.--Blue jay.

Experimental prey.--Diploptera punctata, U.S.A. (Eisner, 1958).

Eurycotis floridana, Neostylopyga rhombifolia, and Periplaneta americana, U.S.A. (Eisner, personal communication, 1958): E. floridana was only eaten after the odor of 2-hexenal, which was released by the insect on being attacked by the bird, had dissipated.

Family PARADISEIDAE

=Paradisea papuana= Bechstein

Experimental prey.--Cockroaches, Malaya and on shipboard (Wallace, 1869): Two adult males fed voraciously on rice, bananas, and cockroaches. Wallace collected cockroaches every night on board ship to feed the birds. "At Malta ... I got plenty of cockroaches from a bakehouse, and when I left, took with me several biscuit-tins full, as provision for the voyage home."

Family TROGLODYTIDAE

=Troglodytes aedon= Vieillot

Common name.--House wren.

Natural prey.--Cockroaches, U.S.A. (Greenewalt and Jones, 1955): The wren carried three small cockroaches to nestlings; the records probably represent incidental captures.

=Troglodytes audax= Tschudi

Common name.--Cucarachero.

Natural prey.--Cockroach (called Chilicabra by Peruvian Indians), Peru (Tschudi, 1847): The bird seized the cockroach and bit off its head then devoured the body discarding the wings.

Family LANIIDAE

=Lanius ludovicianus= Linnaeus

Common name.--Loggerhead shrike.

Experimental prey.--Pycnoscelus surinamensis, U.S.A., Florida (Sanders, 1928).

Family STURNIDAE

=Acridotheres tristis= Bonnaterre and Vieillot

Common name.--Myna, mynah.

Natural prey.--Cockroaches, Hawaii, Lanai (Illingworth, 1928): Illingworth reported that he had never seen as many cockroaches anywhere else in Hawaii. The birds followed tractors that were destroying cactus and kept close to the chain that turned over the stumps. The following species were collected: Allacta similis, Blattella germanica, Cutilia soror, Diploptera punctata, Leucophaea maderae, Periplaneta americana, Periplaneta australasiae, Pycnoscelus surinamensis. Illingworth did not state whether the birds ate all these species indiscriminately.

Pycnoscelus surinamensis, Hawaii (Williams et al., 1931; Schwabe, 1950b): In many places this species forms an important fledgling food for mynah birds.

Family VIREONIDAE

=Vireo latimeri= Baird

Common name.--Latimer's vireo.

Natural prey.--Periplaneta sp., Puerto Rico (Wetmore, 1916): Cockroaches were found in one of 43 stomachs examined.

Family ICTERIDAE

=Agelaius xanthomus= (Sclater)

Common name.--Yellow-shouldered blackbird.

Natural prey.--Cockroaches, Puerto Rico (Wetmore, 1916): Oöthecae and remains of adult cockroaches found in stomachs.

=Dolichonyx oryzivorus= (Linnaeus)

Common name.--Bobolink.

Experimental prey.--Pycnoscelus surinamensis, U.S.A., Florida (Sanders, 1928).

=Holoquiscalus brachypterus= (Cassin)

Common name.--Puerto Rican blackbird.

Natural prey.--Cockroaches, Puerto Rico (Wetmore, 1916): A few eggs (oöthecae) of cockroaches in stomachs.

=Black bird=

Experimental prey.--Pycnoscelus surinamensis, U.S.A., Florida (Sanders, 1928).

Family FRINGILLIDAE

=Passer domesticus= (Linnaeus)

Common name.--English sparrow.

Natural prey.--Pycnoscelus surinamensis, Hawaii (Illingworth, 1931; Schwabe, 1950b): Remains of this cockroach were found in the stomach of the sparrow.

=Sparrow=

Natural prey.--Periplaneta americana, England (Lucas, 1908, 1920).

Cockroaches, Japan (Asano, 1937).

=Tiaris bicolor omissa= (Jardine)

Common name.--Carib grassquit.

Natural prey.--Cockroaches, Puerto Rico (Wetmore, 1916): Animal food was found in 5 of 72 stomachs examined; one bird had eaten two cockroaches among other insects.

Class MAMMALIA

Order MARSUPIALIA

Family DIDELPHIDAE

=Monodelphis= sp.

Natural prey.--Cockroaches, British Guiana (Beebe, 1925a): The above opossum is nocturnal and arboreal but nests on the ground in grass.

Order INSECTIVORA

Family ERINACEIDAE

=Erinaceus europaeus= Linnaeus

Experimental prey.--Blattella germanica, France (Brumpt and Urbain, 1938): Two hedgehogs were fed cockroaches infested with Prosthenorchis elegans and P. spirula.

=Erinaceus= sp.

Common name.--Hedgehog.

Natural prey.--Cockroaches, England (Samouelle, 1841; Cowan, 1865).

Order CHIROPTERA

Family MOLOSSIDAE

=Molossus= sp.

Natural prey.--Cockroaches, British Guiana (Beebe, 1925a): The above bat is a common house bat of the area.

Order PRIMATES

Family LEMURIDAE

=Lemur coronatus= Gray

Natural prey.--Blattella germanica, France (Brumpt and Urbain, 1938): The monkey apparently became infested naturally with Prosthenorchis spirula for which B. germanica was the intermediate host in the monkey house.

=Lemur fulvus= E. Geoffroy

Natural prey.--Blattella germanica, France (Brumpt and Urbain, 1938): See comment under Lemur coronatus.

Family LORISIDAE

=Loris tardigradus= (Linnaeus)

Synonymy.--Lemur tardigradus [Dr. D. H. Johnson, personal communication, 1957].

Natural prey.--Cockroaches, on board ship (Cowan, 1865).

=Perodicticus potto= (P. L. S. Müller)

Common name.--Potto.

Natural prey.--Blattella germanica, France (Brumpt and Urbain, 1938): See comment under Lemur coronatus.

Experimental prey.--Blattidae, East Africa (Pitman, 1931): Both sexes of the potto ate freely of all types of cockroaches.

Family TARSIIDAE

=Tarsius= sp.

Experimental prey.--Cockroaches, Borneo (Shelford, 1916).

Family CEBIDAE

=Aotes zonalis= Goldman

Synonymy.--Aotus [Simpson, 1945].

Common name.--Canal Zone night monkey.

Natural prey.--Leucophaea maderae, Panama (Foster and Johnson, 1939): Captive monkeys became naturally infested with Protospirura muricola by eating cockroaches that contained infective larvae of the worm.

=Ateles dariensis= Goldman

Common name.--Darien black spider monkey.

Natural prey.--Leucophaea maderae, Panama (Foster and Johnson, 1939): See comment under Aotes zonalis.

=Cebus apella= (Linnaeus)

Natural prey.--Blattella germanica, France (Brumpt and Urbain, 1938): See comment under Lemur coronatus.

=Cebus capucinus= (Linnaeus)

Common name.--White-faced monkey.

Natural prey.--Leucophaea maderae, Panama (Foster and Johnson, 1939): Favorite item of food in the laboratory. See comment under Aotes zonalis.

=Saimiri sciurea= Linnaeus

Natural prey.--Blattella germanica, France (Brumpt and Urbain, 1938): See comment under Callithrix chrysoleucos.

Family CALLITHRICIDAE

=Callithrix chrysoleucos= (Natterer)

Synonymy.--Callithrix chrysolevea [Johnson, personal communication, 1957].

Natural prey.--Blattella germanica, France (Brumpt and Urbain, 1938): The monkey apparently became infested naturally with Prosthenorchis elegans for which B. germanica was the intermediate host in the monkey house.

=Callithrix jacchus= (Linnaeus)

Synonymy.--Simia jacchus.

Natural prey.--Blattella germanica, France (Brumpt and Urbain, 1938): See comment under Lemur coronatus.

Cockroaches, on board ship (Neill, 1829; also cited by Samouelle, 1841, and Cowan, 1865): "It was quite amusing to see it at its meal. When he had got hold of one of the largest cockroaches, he held it in his fore paws, and then invariably nipped the head off first; he then pulled out the viscera and cast them aside, and devoured the rest of the body, rejecting the dry elytra and wings, and also the legs of the insect, which are covered with short stiff bristles. The smaller cockroaches he eat without such fastidious nicety."

=Leontocebus oedipus= (Linnaeus)

Natural prey.--Blattella germanica, France (Brumpt and Urbain, 1938): See comment under Callithrix chrysoleucos.

=Leontocebus rosalia= (Linnaeus)

Synonymy.--Midas rosalia [Simpson, 1945].

Natural prey.--Blattella germanica, France (Brumpt and Urbain, 1938): See comments under Lemur coronatus and Callithrix chrysoleucos.

=Leontocebus ursulus= (E. Geoffroy)

Natural prey.--Blattella germanica, France (Brumpt and Urbain, 1938): See comments under Callithrix chrysoleucos.

Family CERCOPITHECIDAE

=Cercopithecus= sp.

Experimental prey.--Cockroaches, East Africa (Carpenter, 1921, 1925): The monkey rarely tasted and usually ignored cockroaches offered to it. In one experiment the monkey had to be deprived of food before it would eat the cockroach.

=Macaca mulatta= (Zimmermann)

Synonymy.--Macaca rhesus [Johnson, personal communication, 1957].

Experimental prey.--Blattella germanica, France (Brumpt and Urbain, 1938, 1938a): The macaque was fed cockroaches infested with Prosthenorchis elegans and P. spirula.

=Macaca sylvanus= (Linnaeus)

Synonymy.--Inuus sylvanus [Simpson, 1945].

Common name.--Macaque.

Natural prey.--Blattella germanica, France (Brumpt and Urbain, 1938): See comment under Lemur coronatus.

=Papio papio= (Desmarest)

Experimental prey.--Blattella germanica, France (Brumpt and Urbain, 1938, 1938a): This baboon was fed cockroaches infested with Prosthenorchis elegans and P. spirula.

Family PONGIDAE

=Pan= sp.

Common name.--Chimpanzee.

Natural prey.--Blattella germanica, Netherlands (Thiel and Wiegand Bruss, 1946): Indirect evidence for this relationship was shown by these workers who found two animals heavily infected with Prosthenorchis spirula in a zoo in Rotterdam; the intermediate host of the worm was shown to be B. germanica.

Family HOMINIDAE

=Homo sapiens= Linnaeus

Natural prey.--Oöthecae of Blatta orientalis and Neostylopyga rhombifolia, Thailand (Bristowe, 1932).

Periplaneta americana, Formosa (Takahashi, 1924).

Periplaneta americana and Periplaneta australasiae, Australia, China, and Japan (Bodenheimer, 1951).

Cockroaches, Annam and French Guinea (Brygoo, 1946).

In addition to the above records of cockroaches being used as food by man these insects have also been eaten for medicinal purposes (see Roth and Willis, 1957a).

Order EDENTATA

Family DASYPODIDAE

=Dasypus novemcinctus= Linnaeus

Synonymy.--Tatu novemcinctum [Johnson, personal communication, 1958].

Natural prey.--Ischnoptera deropeltiformis, Texas (Hebard, 1917): A specimen of this cockroach in the U. S. National Museum was taken from the stomach of the armadillo.

Order RODENTIA

Family MURIDAE

=Mus musculus= Linnaeus

Experimental prey.--Diploptera punctata, U.S.A. (Eisner, 1958).

=Rattus norvegicus= (Berkenhout)

Synonymy.--Mus decumanus; Epimys norvegicus.

Natural prey.--Leucophaea maderae, Venezuela (Brumpt, 1931): Rats infested with Protospirura bonnei presumably ate this cockroach which is the intermediate host of the worm.

Periplaneta americana, Brazil (Magalhães, 1898): Remains found in the stomachs of brown rats. Denmark (Fibiger and Ditlevsen, 1914): This cockroach was found to be the intermediate host of Gongylonema neoplasticum, a parasite of rats.

=Rattus rattus= (Linnaeus)

Natural prey.--Periplaneta americana, Denmark (Fibiger and Ditlevsen, 1914): See comment after these authors under Rattus norvegicus.

=Rattus= spp.

Natural prey.--Cockroaches, India (Maxwell-Lefroy, 1909); Burma (Subramanian, 1927).

Family CAVIIDAE

=Cavia= sp.

Experimental prey.--Blattella germanica, U.S.A. (Hobmaier, 1941): Guinea pigs were fed cockroaches infested with Physaloptera maxillaris.

Order CARNIVORA

Family CANIDAE

=Canis familiaris= Linnaeus

Experimental prey.--Blattella germanica, U.S.A. (Hobmaier, 1941): Dogs were fed cockroaches infested with Physaloptera maxillaris. U.S.A. (Petri and Ameel, 1950): Cockroaches infested with Physaloptera rara were fed to a dog.

=Canis latrans= Say

Experimental prey.--Blattella germanica, U.S.A. (Petri and Ameel, 1950): Cockroaches infested with Physaloptera rara were fed to a coyote.

=Vulpes= sp.

Experimental prey.--Blattella germanica, France (Brumpt and Urbain, 1938a): A fox was successfully infected when fed cockroaches infested with Prosthenorchis elegans and P. spirula.

Family PROCYONIDAE

=Bassariscus astutus= (Lichtenstein)

Common names.--Cacomistle, ring-tailed cat.

Natural prey.--Cockroaches, U.S.A., Arizona (Dr. H. Stahnke, personal communication, 1953): The ring-tailed cat enters dwellings located on the desert and feeds on cockroaches and other arthropods.

=Nasua narica= (Linnaeus)

Natural prey.--Blattella germanica, France (Brumpt and Urbain, 1938): The coati apparently became infested naturally with Prosthenorchis spirula for which B. germanica was the intermediate host in the laboratory.

=Nasua nasua= (Linnaeus)

Natural prey.--Blattella germanica, on board ship (Myers, 1931): This insect was eaten when other insects were absent.

Cockroach, a small outdoor species, Trinidad (Myers, 1931).

=Nasua= sp.

Natural prey.--Cockroaches, British Guiana (Beebe, 1925a).

Family MUSTELIDAE

=Meles= sp.

Experimental prey.--Blattella germanica, France (Brumpt and Urbain, 1938a): A badger was successfully infected when fed cockroaches infested with Prosthenorchis elegans and P. spirula.

Family VIVERRIDAE

=Herpestes javanicus auropunctatus= Hodgson

Natural prey.--Epilampra wheeleri, Eurycotis improcera, Panchlora nivea, Pycnoscelus surinamensis, and others unidentified to species, St. Croix and Puerto Rico (Wolcott, 1953): Based on 37 or more cockroaches obtained from stomachs of 42 mongooses collected in St. Croix (by Seaman) and 56 collected in Puerto Rico (by Pimentel).

Pimentel (personal communication, 1958) has given us the following percentage occurrence of cockroach species in the total number of mongoose stomachs that he examined in Puerto Rico: Epilampra wheeleri 1.8, Ischnoptera rufa rufa 3.6, Panchlora nivea 1.8, Periplaneta americana 1.8, and Pycnoscelus surinamensis 19.6.

=Herpestes= sp.

Natural prey.--Periplaneta americana and Periplaneta australasiae, Hawaii (Perkins, 1913): Large numbers of these cockroaches are devoured.

Cockroach, East Africa (Loveridge, 1923): Cockroach remains found in stomach of mongoose.

Family FELIDAE

=Felis catus= Linnaeus

Natural prey.--Periplaneta americana, Hawaii (Williams et al., 1931).

Cockroaches, U.S.A., Arizona (Stahnke, personal communication, 1953).

Experimental prey.--Blattella germanica, U.S.A. (Hobmaier, 1941): Cats were fed cockroaches infested with Physaloptera maxillaris. U.S.A. (Petri and Ameel, 1950): Cockroaches infested with Physaloptera rara were fed to a kitten. France (Brumpt and Urbain, 1938a): A young cat was fed cockroaches infested with Prosthenorchis elegans and P. spirula.

=Felis pardalis mearnsi= J. A. Allen

Natural prey.--Cockroaches, Panama (Dr. H. L. Sweetman, personal communication, 1958): An ocelot was seen collecting and feeding on cockroaches, possibly Blaberus sp. "The ocelot was quite efficient and seemed to relish the roaches."

XIV. CHECKLIST OF COCKROACHES AND SYMBIOTIC ASSOCIATES

Only naturally occurring associations are included in this list. Commensal cockroaches are listed on page 315. Bacteroids are not listed because they undoubtedly occur in all species. The higher plants were excluded because most of the associations may be too casual to constitute symbiosis; however, many of the plant associations were included in the chapter on ecology. The cockroaches and the associates within each category are arranged alphabetically by genus and species. Page references are to citations in the classified sections where details of the associations and/or sources of the records are given.

=Aglaopteryx facies= Mite: Undetermined, p. 220. Reptile: Anolis stratulus, p. 274.

=Aglaopteryx diaphana= Nematode: Protrellus manni, p. 200.

=Allacta similis= Insects: Dolichurus stantoni, p. 262. Solindenia picticornis, p. 247.

=Anaplecta= sp. Fungus: Herpomyces anaplectae, p. 134.

=Aptera fusca= Protozoan: Gregarina fastidiosa, p. 182.

=Arenivaga bolliana= Insect: Sarcophaga omani, p. 229.

=Arenivaga roseni= Insects: Undetermined reduviids, p. 227.

=Balta patula= Insect: Rhipidioides ableptus, p. 230.

=Blaberus atropos= Bacterium: Spirochaeta blattae, p. 125. Protozoa: Endolimax nana, p. 180. Entamoeba coli, p. 178. Nematodes: Leidynema appendiculata, p. 197. Leidynema cranifera, p. 198.

=Blaberus craniifer= Bacteria: Aerobacter aerogenes, p. 111. Alcaligenes faecalis, p. 111. Bacillus cereus, p. 120. Bacillus subtilis, p. 121. Escherichia coli var. communior, p. 113. Escherichia freundii, p. 113. Micrococcus pyogenes var. albus, p. 106. Micrococcus pyogenes var. aureus, p. 107. Proteus vulgaris, p. 114. Pseudomonas aeruginosa, p. 104. Fungi: Herpomyces tricuspidatus, p. 138. Penicillium sp., p. 131. Rhizopus nigricans, p. 133. Saccharomyces cerevisiae, p. 133. Protozoan: Diplocystis (?) sp., pp. 181, 184. Nematodes: Leidynema cranifera, p. 198. Protrelleta floridana, p. 199. Mite: Iolina nana, p. 219. Undetermined, p. 220. Bird: Chicken, p. 278.

=Blaberus discoidalis= Mite: Undetermined, p. 220.

=Blaberus giganteus= Hair worm: Gordius pilosus, p. 202.

=Blaberus= sp. Fungi: Herpomyces macropus, p. 136. Herpomyces paranensis, p. 136. Herpomyces periplanetae, p. 137. Herpomyces tricuspidatus, p. 138. Protozoan: Leptomonas blaberae, p. 167.

=Blaptica dubia= Protozoan: Pileocephalus blaberae, p. 184.

=Blatta (Shelfordella) lateralis= Bacterium: Shigella paradysenteriae, p. 119. Insect: Ampulex assimilis, p. 257.

=Blatta orientalis= Bacteria: Aerobacter aerogenes, p. 111. Alcaligenes faecalis, p. 111. Alcaligenes recti, p. 111. Arthromitus intestinalis, p. 124. B. aerobio del pseudoedema maligno, p. 125. B. alcaligenes beckeri, p. 125. B. del pseudoedema maligno, p. 125. Bacillo proteisimile, p. 126. Bacillo similcarbonchio, p. 126. Bacillo similtifo (Bacillo tifosimile), p. 126. Bacillus bütschlii, p. 120. Bacillus periplanetae, p. 121. Bacillus stellatus, p. 121. Bacillus subtilis, p. 122. Bacillus tritus, p. 122. Bacteroides uncatus, p. 119. Clostridium lentoputrescens, p. 122. Clostridium novyi or Clostridium sporogenes, p. 122. Enterococcus sp., p. 109. Escherichia coli, p. 112. Lactobacillus fermenti, p. 109. Micrococcus pyogenes var. albus, p. 107. Micrococcus pyogenes var. aureus, p. 107. Micrococcus sp., p. 108. Paracolobactrum sp., p. 113. Pasteurella pestis, p. 119. Proteus sp., p. 114. Proteus vulgaris, p. 114. Pseudomonas aeruginosa, p. 104. Pseudomonas eisenbergii, p. 105. Pseudomonas fluorescens, p. 105. Salmonella typhosa, p. 117. Sarcina alba, p. 108. Sarcina sp., pp. 108, 109. Sarcina symbiotica (host may have been B. germanica), p. 108. Sarcina ventriculi, p. 108. Serratia marcescens, p. 117. Spirillochaeta blattae, p. 127. Spirillum α, β and γ, p. 105. Spirillum sp., p. 105. Spirochaeta periplanetae, p. 125. Streptococcus faecalis, p. 109. Streptococcus liquefaciens, p. 110. Streptococcus microapoika, p. 110. Streptococcus pyogenes, p. 110. Streptococcus sp., p. 110. Treponema parvum, p. 125. Treponema stylopygae, p. 125. Vibrio sp., p. 106. Fungi: Aspergillus fumigatus, p. 130. Blastocystis hominis, p. 133. Blastocystis sp., p. 133. Candida zeylanoides, p. 129. Coccidioides periplanetae, p. 133. Herpomyces periplanetae, p. 137. Herpomyces stylopygae, p. 137. Torula gropengiesseri, p. 132. Torulopsis sp., p. 130. Protozoa: Balantidium praenucleatum, p. 187. Bodo blattae, p. 167. Coelosporidium periplanetae, p. 185. Diplocystis schneideri, p. 181. Endamoeba blattae, p. 177. Entamoeba thomsoni, p. 179. Endolimax blattae, p. 180. Endolimax sp., p. 180. Gregarina blattarum, p. 181. Haplosporidium periplanetae, p. 185. Hartmannella blattae, p. 177. Herpetomonas periplanetae, p. 167. Hexamita periplanetae, p. 171. Lophomonas blattarum, p. 172. Lophomonas striata, p. 173. Monas sp., p. 167. Monocercomonoides orthopterorum, p. 169. Nyctotherus ovalis, p. 188. Oikomonas sp., p. 166. Peltomyces periplanetae, p. 177. Plistophora kudoi, p. 185. Plistophora periplanetae, p. 186. Plistophora sp., p. 186. Retortamonas blattae, p. 167. Stenophora sp., p. 181. Tetratrichomastix blattidarum, p. 170. Helminths: Ascaris sp., p. 209. Enterobius vermicularis, p. 209. Gongylonema neoplasticum, p. 206. Gordius blattae orientalis, p. 202. Hammerschmidtiella diesingi, p. 195. Hammerschmidtiella neyrai, p. 196. Leidynema appendiculata, p. 197. Spirura gastrophila, p. 207. Thelastoma pachyjuli, p. 201. Trichostrongylus sp., p. 210. Trichuris trichiura, p. 210. Mite: Undetermined, pp. 220, 222. Insects: Dermestes ater, p. 234. Dermestes sp., p. 234. Evania appendigaster, p. 236. Evania dimidiata, p. 239. Prosevania punctata, p. 240. Systellogaster ovivora, p. 248. Tetrastichus hagenowii, p. 250. Tetrastichus sp., p. 254. Bird: Chicken, p. 278. Mammal: Homo sapiens, p. 286.

=Blatta= sp. Hair worm: Gordius aquaticus, p. 201. Insects: Evania appendigaster, p. 236. Tetrastichus hagenowii, p. 250.

=Blattella germanica= and/or =Blattella vaga= Viruses: Unspecified strains of poliomyelitis virus, p. 103.

=Blattella germanica= Bacteria: Achromobacter sp., p. 110. Aerobacter aerogenes, p. 111. Aerobacter cloacae, p. 112. Alcaligenes faecalis, p. 111. Alcaligenes viscosus, p. 111. Bacillus circulans, p. 120. Escherichia coli, p. 112. Escherichia freundii, p. 113. Micrococcus aurantiacus, p. 106. Micrococcus epidermidis, p. 106. Micrococcus pyogenes var. albus, p. 107. Micrococcus pyogenes var. aureus, p. 107. Micrococcus ureae, p. 107. Micrococcus sp., p. 108. Mycobacterium leprae, p. 123. Paracolobactrum aerogenoides, p. 113. Paracolobactrum coliforme, p. 113. Paracolobactrum sp., p. 113. Pseudomonas aeruginosa, p. 104. Sarcina symbiotica (host may have been B. orientalis), p. 108. Salmonella typhimurium, p. 116. Serratia marcescens, p. 117. Streptococcus faecalis, p. 109. Streptococcus sp., p. 110. Fungi: Aspergillus flavus, p. 130. Aspergillus tamarii, p. 130. Aspergillus sp., p. 130. Cordyceps blattae, p. 134. Herpomyces ectobiae, p. 135. Memnoniella echinata, p. 132. Saccharomyces sp., p. 133. Protozoa: Bodo sp. (host may have been P. americana), p. 167. Coelosporidium periplanetae, p. 185. Dobellina sp. (host may have been P. americana), p. 177. Endamoeba blattae (host may have been P. americana), p. 177. Endolimax sp. (host may have been P. americana), p. 180. Entamoeba coli (host may have been P. americana), p. 178. Entamoeba histolytica (host may have been P. americana), p. 179. Entamoeba thomsoni, p. 179. Eutrichomastix sp. (host may have been P. americana), p. 169. Gregarina blattarum, p. 182. Iodamoeba sp. (host may have been P. americana), p. 180. Lophomonas blattarum, p. 172. Lophomonas striata, p. 173. Nyctotherus ovalis, p. 188. Peltomyces periplanetae, p. 177. Plistophora periplanetae, p. 186. Tetratrichomastix blattidarum, p. 170. Helminths: Blattelicola blattelicola, p. 193. Blatticola blattae, p. 193. Enterobius vermicularis, p. 209. Galebia aegyptiaca, p. 195. Gongylonema neoplasticum, p. 206. Moniliformis kalahariensis, p. 203. Prosthenorchis elegans, p. 203. Prosthenorchis spirula, p. 203. Tetrameres americana, p. 207. Trichuris trichiura, p. 210. Mites: Blattisocius tineivorus, p. 216. Caloglyphus sp., p. 218. Undetermined, p. 220. Insects: Brachygaster minutus, p. 235. Eupelmus atriflagellum, p. 247. Dolichurus corniculus, p. 261. Ripidius pectinicornis, p. 232. Mammals: Callithrix chrysoleucos, p. 285. Callithrix jacchus, p. 285. Cebus apella, p. 284. Lemur coronatus, p. 283. Lemur fulvus, p. 283. Leontocebus oedipus, p. 285. Leontocebus rosalia, p. 285. Leontocebus ursulus, p. 285. Macaca sylvanus, p. 286. Nasua narica, p. 288. Nasua nasua, p. 288. Pan sp., p. 286. Perodicticus potto, p. 284. Saimiri sciurea, p. 285.

=Blattella humbertiana= Fungus: Herpomyces gracilis, p. 135. Nematode: Protrellus phyllodromi, p. 200. Mite: Uropoda sp., p. 217.

=Blattella lituricollis= Insect: Dolichurus stantoni, p. 262.

=Blattella= sp. Reptile: Anolis cristatellus, p. 273.

=Cariblatta delicatula= Insect: Hyptia sp., p. 240. Reptile: Anolis cristatellus, p. 273.

=Cariblatta lutea lutea= Plants: Sarracenia flava, p. 154. Sarracenia purpurea, p. 154.

=Chorisoneura= sp. Insects: Stylogaster sp., p. 228. Tachysphex blatticidus, p. 264.

=Choristima= sp. Insect: Rhipidioides rubricatus, p. 231.

=Choristimodes= sp. Insect: Rhipidioides rubricatus, p. 231.

=Cryptocercus punctulatus= Bacterium: Bacillus subtilis, p. 122. Protozoa: Adelina cryptocerci, p. 184. Barbulanympha estaboga, p. 173. Barbulanympha laurabuda, p. 174. Barbulanympha ufalula, p. 174. Barbulanympha wenyoni, p. 174. Eucomonympha imla, p. 176. Hexamita cryptocerci, p. 171. Idionympha perissa, p. 174. Leptospironympha eupora, p. 172. Leptospironympha rudis, p. 172. Leptospironympha wachula, p. 172. Macrospironympha xylopletha, p. 172. Monocercomonoides globus, p. 169. Notila proteus, p. 170. Oxymonas doroaxostylus, p. 170. Oxymonas nana, p. 170. Prolophomonas tocopola, p. 173. Rhynchonympha tarda, p. 174. Saccinobaculus ambloaxostylus, p. 170. Saccinobaculus lata, p. 170. Trichonympha acuta, p. 174. Trichonympha algoa, p. 174. Trichonympha chula, p. 176. Trichonympha grandis, p. 176. Trichonympha lata, p. 176. Trichonympha okolona, p. 176. Trichonympha parva, p. 176. Urinympha talea, p. 174. Undetermined gregarine, p. 184. Amphibian: Plethodon glutinosus, p. 269.

=Cutilia soror= Insects: Dolichurus stantoni, p. 262. Evania appendigaster, p. 236. Szepligetella sericea, p. 242. Bird: Phasianus sp., p. 277.

=Cutilia= sp. Nematode: Undetermined, p. 201. Insects: Riekella australis, p. 231. Undetermined strepsipteron, p. 234.

=Diploptera punctata= Bacterium: Serratia marcescens, p. 117. Fungus: Herpomyces diplopterae, p. 135. Mites: Iolina nana, p. 219. Locustacarus sp., p. 219. Insect: Nauphoeta cinerea, p. 324. Amphibian: Bufo marinus, p. 270. Bird: Phasianus sp., p. 277.

=Ectobius lapponicus= Protozoa: Gamocystis tenax, p. 184. Monocercomonoides orthopterorum, p. 169. Nematode: Blatticola blattae, p. 193. Insects: Brachygaster minutus, p. 235. Dolichurus corniculus, p. 261. Tachysphex lativalvis, p. 264.

=Ectobius pallidus= Protozoan: Gamocystis tenax, p. 184. Helminths: Blatticola blattae, p. 193. Undetermined mermithid, p. 192. Insects: Ampulex fasciata, p. 259. Aphaenogaster picea, p. 267. Dolichurus corniculus, p. 261. Lasius alienus, p. 268. Ripidius boissyi (presumptive record), p. 231. Ripidius denisi (presumptive record), p. 232. Tachysphex lativalvis, p. 264.

=Ectobius panzeri= Centipede: Scolopendra sp., p. 224. Insects: Dolichurus corniculus, p. 261. Tachysphex lativalvis, p. 264.

=Ectobius= sp. Insects: Brachygaster minutus, p. 235. Dolichurus corniculus, p. 261. Tachysphex lativalvis, p. 264.

=Ellipsidion affine= Insect: Rhipidioides fuscatus, p. 230.

=Ellipsidion australe= Insects: Agamerion metallica, p. 243. Cheiloneurus viridiscutum, p. 244. Mestocharomyia oophaga, p. 248.

=Epilampra abdomen-nigrum= Insect: Podium flavipenne, p. 265. Amphibian: Bufo marinus, p. 270.

=Epilampra conferta= Insect: Podium sp., p. 266.

=Epilampra= sp. Fungus: Herpomyces tricuspidatus, p. 138. Insect: Podium haematogastrum, p. 266. Bird: Gymnasio nudipes, p. 279.

=Epilampra wheeleri= Reptiles: Ameiva exsul, p. 275. Anolis cristatellus, p. 273. Mammal: Herpestes javanicus auropunctatus, p. 289.

=Escala= (?) sp. Insect: Rhipidioides adynatus, p. 230.

=Eurycotis floridana= Fungi: Aspergillus flavus, p. 130. Aspergillus sydowi, P. 130. Helminths: Euryconema paradisa, p. 194. Leidynema appendiculata(?), p. 198. Protrelloides paradoxa, p. 200. Undetermined gordian worm, p. 202. Insects: Anastatus floridanus, p. 245. Undetermined tachinid, p. 228.

=Eurycotis improcera= Mammal: Herpestes javanicus auropunctatus, p. 289.

=Eurycotis manni= Fungus: Herpomyces zanzibarinus, p. 138.

=Graptoblatta notulata= Insect: Tachysphex fanuiensis, p. 264.

=Gromphadorhina portentosa= Protozoan: Undetermined gregarine, p. 184. Mite: Coleolaelaps ? sp., p. 216.

=Gyna= sp.(?) Fungus: Herpomyces zanzibarinus, p. 138.

=Holocompsa fulva= Insect: Pheidole megacephala, p. 268.

=Hololampra punctata= Insect: Dolichurus corniculus, p. 261.

=Ischnoptera deropeltiformis= Plant: Sarracenia flava, p. 154 Amphibian: Hyla cinerea, p. 270. Mammal: Dasypus novemcinctus, p. 287.

=Ischnoptera rufa rufa= Fungus: Spicaria prasina, p. 130. Mammal: Herpestes javanicus auropunctatus, p. 289.

=Ischnoptera= sp. Fungus: Herpomyces arietinus, p. 134. Insects: Ampulex canaliculata, p. 257. Syntomosphyrum ischnopterae, p. 249.

=Karnyia discoidalis= Spider: Latrodectus indistinctus, p. 215.

=Kuchinga hemerobina= Hair worm: Parachordodes raphaelis, p. 202.

=Leucophaea maderae= Bacterium: Serratia marcescens, p. 117. Fungus: Herpomyces tricuspidatus, p. 138. Protozoa: Gregarina rhyparobiae, p. 183. Hexamita (?) sp., p. 171. Retortamonas (?) sp., p. 167. Undetermined gregarine, p. 184. Nematodes: Hammerschmidtiella diesingi, p. 195. Leidynema delatorrei, p. 198. Protospirura bonnei, p. 206. Protospirura muricola, p. 206. Mite: Chaetodactylus sp., p. 218. Mammals: Aotes zonalis, p. 284. Ateles dariensis, p. 284. Cebus capucinus, p. 284. Rattus norvegicus, p. 287.

=Leucophaea= sp. Nematode: Cephalobellus brevicaudatum, p. 194.

=Leurolestes pallidus= Fungus: Herpomyces leurolestis, p. 136.

=Loboptera decipiens= Insects: Dolichurus corniculus, p. 261. Zeuxevania splendidula, p. 243.

=Loboptera= sp. Fungus: Herpomyces lobopterae, p. 136.

=Lobopterella dimidiatipes= Insect: Ampulex canaliculata, p. 257. Bird: Coturnix coturnix japonica, p. 277.

=Melanosilpha capensis= Protozoa: Gregarina impetuosa, p. 183. Gregarina sandoni, p. 183.

=Monastria= sp. Insect: Triatoma arthurneivai, p. 227.

=Nauphoeta cinerea= Bacteria: Salmonella typhimurium, p. 116. Serratia marcescens, p. 117. Fungus: Herpomyces tricuspidatus, p. 138. Mites: Blattilaelaps nauphoetae, p. 216. Locustacarus sp., p. 219. Insect: Pheidole megacephala, p. 268.

=Neostylopyga rhombifolia= Bacterium: Serratia marcescens, p. 117. Insects: Evania appendigaster, p. 237. Szepligetella sericea, p. 242. Tetrastichus hagenowii, p. 250. Mammal: Homo sapiens, p. 286.

=Nyctibora obscura= Fungus: Herpomyces amazonicus, p. 134.

=Nyctibora= sp. Fungus: Herpomyces nyctoborae, p. 136.

=Nyctibora tomantosa= Fungus: Herpomyces nyctoborae, p. 136.

=Oniscosoma granicollis= Insect: Paranephrites xenus, p. 230.

=Panchlora exoleta= Protozoan: Gregarina panchlorae, p. 183.

=Panchlora nivea= Bacterium: Serratia marcescens, p. 117. Fungus: Herpomyces panchlorae, p. 136. Mammal: Herpestes javanicus auropunctatus, p. 289.

=Panesthia angustipennis= Protozoa: Clevelandella constricta, p. 189. Clevelandella contorta, p. 189. Clevelandella elongata, p. 189. Clevelandella hastula, p. 189. Clevelandella panesthiae, p. 189. Clevelandella parapanesthiae, p. 189. Endamoeba javanica, p. 178. Endamoeba philippinensis, p. 178. Hexamita cryptocerci, p. 171. Monocercomonoides panesthiae, p. 170. Nyctotherus uichancoi, p. 188. Paraclevelandia brevis, p. 190. Paraclevelandia simplex, p. 190. Undetermined amoeba, p. 181. Nematodes: Aorurus philippinensis, p. 193. Blattophila sphaerolaima var. javanica, p. 194. Leidynema nocalum, p. 198. Leidynemella fusiformis, p. 198. Leidynemella paracranifera, p. 198. Thelastoma palmettum, p. 201.

=Panesthia australis= Fungus: Metarrhizium anisopliae, p. 131. Mite: Hypoaspis sp., p. 217. Undetermined diplogyniid, p. 217. Insect: Undetermined tachinid, p. 228.

=Panesthia laevicollis= Nematode: Blattophila sphaerolaima, p. 194.

=Panesthia laevicollis= (?) Nematode: Leidynemella fusiformis, p. 198.

=Panesthia lobipennis= Fungus: Herpomyces panesthiae, p. 136.

=Panesthia spadica= Protozoa: Clevelandella constricta, p. 189. Clevelandella contorta, p. 189. Clevelandella nipponensis, p. 189. Clevelandella panesthiae, p. 189. Endamoeba javanica, p. 178. Nyctotherus uichancoi, p. 188. Paraclevelandia brevis, p. 190. Paraclevelandia simplex, p. 190.

=Panesthia= sp. Nematode: Leidynemella panesthiae, p. 198.

=Parahormetica bilobata= Helminths: Agamospirura parahormeticae, p. 205. Undetermined gordian worm, p. 202.

=Parcoblatta lata= Plant: Sarracenia flava, p. 154. Protozoan: Leptomonas sp., p. 167. Nematode: Protrellus aurifluus, p. 199.

=Parcoblatta pensylvanica= Protozoa: Gregarina blattarum, p. 182. Gregarina illinensis, p. 183. Gregarina parcoblattae, p. 183. Gregarina thomasi, p. 184. Leptomonas sp., p. 167. Nyctotherus ovalis, p. 188. Nematode: Rictularia coloradensis, p. 205. Scorpion: Centruroides vittatus (?), p. 212. Insects: Camponotus pennsylvanicus, p. 267. Hyptia dorsalis, p. 239. Hyptia harpyoides, p. 239. Hyptia reticulata, p. 240. Hyptia thoracica, p. 240. Podium carolina, p. 265. Systellogaster ovivora, p. 248. Amphibian: Rana (?) sp., p. 272.

=Parcoblatta uhleriana= Fungus: Herpomyces arietinus, p. 134. Protozoan: Gregarina parcoblattae, p. 183. Nematode: Protrellus aurifluus, p. 199. Mite: Undetermined, p. 220. Insect: Hyptia harpyoides, p. 239.

=Parcoblatta virginica= Fungus: Herpomyces arietinus, p. 134. Protozoa: Gregarina ohioensis, p. 183. Leptomonas sp., p. 167. Nematode: Rictularia coloradensis, p. 205. Insects: Hyptia harpyoides, p. 239. Podium luctuosum, p. 266.

=Parcoblatta= sp. Fungus: Herpomyces arietinus, p. 134. Plant: Sarracenia flava, p. 154. Mite: Pimeliaphilus podapolipophagus, p. 219. Insects: Coenosia basalis, p. 229. Dolichurus greenei, p. 261. Hyptia sp., p. 240. Megaselia sp., p. 227. Syntomosphyrum blattae, p. 248. Systellogaster ovivora, p. 248. Tetrastichus hagenowii, p. 250.

=Periplaneta americana= Viruses: Unspecified strain(s) of poliomyelitis virus (host may have been P. brunnea), p. 103. Bacteria: Achromobacter hyalinum, p. 110. Aerobacter aerogenes, p. 111. Aerobacter cloacae, p. 112. Aerobacter sp., p. 112. Alcaligenes faecalis, p. 111. Bacillus cereus, p. 120. Bacillus megaterium, p. 121. Bacillus subtilis, p. 122. Bacterium alkaligenes, p. 119. Clostridium sp., p. 122. Eberthella oedematiens, p. 112. Escherichia coli, p. 112. Escherichia freundii, p. 113. Escherichia intermedium, p. 113. Mycobacterium friedmannii, p. 123. Mycobacterium lacticola, p. 123. Mycobacterium leprae, p. 123. Mycobacterium phlei, p. 123. Mycobacterium piscium, p. 124. Mycobacterium sp., p. 124. Nocardia sp. (?), p. 124. Paracolobactrum aerogenoides, p. 113. Paracolobactrum coliforme, p. 113. Paracolobactrum sp., p. 113. Proteus mirabilis, p. 114. Proteus morganii, p. 114. Proteus rettgeri, p. 114. Proteus vulgaris, p. 114. Proteus sp., p. 114. Pseudomonas aeruginosa, p. 104. Pseudomonas fluorescens, p. 105. Salmonella anatis, p. 114. Salmonella morbificans, p. 115. Salmonella schottmuelleri, p. 115. Salmonella sp. (Type Bareilly), p. 115. Salmonella sp. (Type Bredeny), p. 115. Salmonella sp. (Type Kentucky), p. 115. Salmonella sp. (Type Meleagris), p. 116. Salmonella sp. (Type Newport), p. 116. Salmonella sp. (Type Oranienburg), p. 116. Salmonella sp. (Type Panama), p. 116. Salmonella sp. (Type Rubislaw), p. 116. Salmonella sp. (Type Tennessee), p. 116. Sarcina sp., p. 108. Serratia marcescens, p. 118. Shigella alkalescens, p. 118. Spirillum periplaneticum, p. 105. Streptococcus faecalis, p. 109. Streptomyces leidynematis, pp. 196, 197. Tetragenous sp., p. 127. Veillonella parvula, p. 109. Fungi: Aspergillus flavus, p. 130. Aspergillus niger, p. 130. Aspergillus sp., p. 131. Cephalosporium sp., p. 131. Herpomyces chaetophilus, p. 135. Herpomyces periplanetae, p. 137. Metarrhizium anisopliae, p. 131. Mucor guilliermondii, p. 132. Mucor sp., p. 132. Penicillium sp., p. 131. Rhizopus sp., p. 133. Syncephalastrum sp., p. 133. Torula acidophila, p. 132. Protozoa: Balantidium blattarum, p. 187. Balantidium ovatum, p. 187. Balantidium sp., p. 187. Bodo sp. (host may have been B. germanica), p. 167. Coelosporidium periplanetae, p. 185. Diplocystis schneideri, p. 181. Diplocystis sp., p. 181. Dobellina sp. (host may have been B. germanica), p. 177. Endamoeba blattae, p. 177. Endolimax blattae, p. 180. Endolimax sp. (host may have been B. germanica), p. 180. Entamoeba coli (host may have been B. germanica), p. 178. Entamoeba histolytica (host may have been B. germanica), p. 179. Entamoeba sp., p. 179. Entamoeba thomsoni, p. 179. Eutrichomastix sp. (host may have been B. germanica), p. 169. Gregarina blattarum, p. 182. Gregarina légeri, p. 183. Gregarina neo-brasiliensis, p. 183. Hexamita periplanetae, p. 171. Iodamoeba sp. (host may have been B. germanica) p. 180. Isotricha caulleryi, p. 187. Lophomonas blattarum, p. 172. Lophomonas striata, p. 173. Monocercomonoides orthopterorum, p. 169. Nyctotherus ovalis, p. 188. Plistophora periplanetae, p. 186. Protomagalhaesia serpentula, p. 184. Tetratrichomastix blattidarum, p. 170. Helminths: Ancylostoma duodenale, p. 209. Ascaris lumbricoides or Ascaris suum, p. 209. Binema mirzaia, p. 193. Gongylonema neoplasticum, p. 206. Gongylonema sp., p. 206. Gordius sp., p. 202. Hammerschmidtiella diesingi, p. 195. Hymenolepis sp., p. 208. Leidynema appendiculata, p. 197. Moniliformis dubius, p. 203. Moniliformis moniliformis, p. 204. Necator americanus, p. 210. Protrellus künckeli, p. 199. Schwenkiella icemi, p. 200. Spirura gastrophila, p. 207. Thelastoma pachyjuli, p. 201. Trichuris trichiura, p. 210. Mites: Caloglyphus sp., p. 218. Pimeliaphilus podapolipophagus, p. 219. Rhizoglyphus tarsalus, p. 218. Tyrophagus noxius, p. 218. Undetermined, p. 220. Spiders: Avicularia avicularia, p. 214. Avicularia sp., p. 214. Centipedes: Undetermined, p. 222. Insects: Ampulex amoena, p. 256. Ampulex compressa, p. 259. Anastatus tenuipes, p. 246. Calodexia (?) venteris, p. 228. Evania appendigaster, p. 237. Melittobia chalybii, p. 248. Prosevania punctata, p. 241. Ripidius pectinicornis, p. 232. Spiniger domesticus, p. 227. Szepligetella sericea, p. 242. Tetrastichus hagenowii, p. 250. Tetrastichus periplanetae, p. 253. Tetrastichus sp., p. 254. Trirhogma caerulea, p. 262. Amphibian: Hyla cinerea, p. 270. Reptiles: Ameiva exsul, p. 275. Anolis cristatellus, p. 273. Birds: Chicken, p. 278. Sparrow, p. 282. Mammals: Felis catus, p. 289. Herpestea javanicus auropunctatus, p. 289. Herpestes sp., p. 289. Homo sapiens, p. 286. Rattus norvegicus, p. 287. Rattus rattus, p. 287.

=Periplaneta australasiae= Bacteria: Mycobacterium leprae, p. 123. Serratia marcescens, p. 118. Fungus: Herpomyces periplanetae, p. 137. Plant: Sarracenia minor, p. 154. Protozoa: Endamoeba blattae, p. 177. Endolimax blattae, p. 180. Nematodes: Gongylonema neoplasticum, p. 206. Hammerschmidtiella diesingi, p. 195. Moniliformis dubius, p. 203. Protrellus australasiae, p. 199. Protrellus künckeli, p. 200. Centipedes: Undetermined, p. 222. Insects: Ampulex amoena, p. 256. Ampulex compressa, p. 259. Evania appendigaster, p. 237. Szepligetella sericea, p. 242. Tetrastichus australasiae, p. 249. Tetrastichus hagenowii, p. 250. Tetrastichus, sp., p. 254. Trirhogma caerulea, p. 262. Reptiles: Anolis cristatellus, p. 273. Chrysemys picta, p. 272. Bird: Gallus sp., p. 277. Mammals: Herpestes sp., p. 289. Homo sapiens, p. 287.

=Periplaneta brunnea= Viruses: Unspecified strain(s) of poliomyelitis virus (host may have been P. americana), p. 103. Bacterium: Serratia marcescens, p. 118. Fungus: Herpomyces periplanetae, p. 137. Nematode: Schwenkiella icemi, p. 200. Insect: Tetrastichus hagenowii, p. 251.

=Periplaneta fuliginosa= Insect: Ampulex amoena, p. 256.

=Periplaneta= sp. Bacterium: Serratia marcescens, p. 118. Fungi: Herpomyces chaetophilus, p. 135. Herpomyces periplanetae, p. 137. Protozoan: Lophomonas blattarum, p. 172. Helminths: Thelastoma riveroi, p. 201. Moniliformis moniliformis, p. 204. Insects: Ampulex compressa, p. 259. Evania subspinosa (presumptive record?), p. 239. Szepligetella sericea, p. 242. Amphibian: Bufo marinus, p. 270. Reptiles: Anolis grahami, p. 274. Anolis leachi, p. 274. Bird: Vireo latimeri, p. 281.

"=Phyllodromia=" sp. Fungus: Herpomyces phyllodromiae, p. 137. Insects: Dicarnosis alfierii, p. 245. Dolichurus stantoni, p. 262.

=Platyzosteria castanea= Insect: Riekella sp., p. 231.

=Platyzosteria novae seelandiae= Protozoan: Monocercomonoides melolonthae, p. 169.

=Platyzosteria scabra= Fungus: Herpomyces appendiculatus, p. 134.

=Platyzosteria= sp. Insect: Riekella nitidioides, p. 231.

=Plectoptera= sp. Bird: Todus mexicanus, p. 279.

=Polyphaga aegyptiaca= Nematodes: Blatticola blattae, p. 193. Galebia aegyptiaca, p. 195. Hammerschmidtiella diesingi, p. 195. Oxyuris (?) heterogamiae, p. 199.

=Polyphaga saussurei= Helminths: Hymenolepis sp., p. 208. Undetermined tapeworm ova, p. 208. Insects: Undetermined reduviids, p. 227.

=Polyzosteria melanaria= (?) or =Platyzosteria analis= Nematode: Protrellus aureus, p. 199.

=Pycnoscelus surinamensis= Bacterium: Serratia marcescens, p. 117. Fungus: Mucor sp., p. 132. Protozoa: Undetermined ciliate, p. 190. Undetermined flagellate, p. 176. Undetermined gregarine, p. 184. Nematodes: Oxyspirura mansoni, p. 204. Severianoia severianoi, p. 200. Mites: Caloglyphus spinitarsus, p. 217. Histiostoma feroniarum, p. 217. Tyrophagus lintneri, p. 218 Undetermined, p. 220. Insects: Pheidole megacephala, p. 268. Undetermined ants, p. 350. Amphibian: Bufo marinus, p. 270. Reptile: Anolis sagrei, p. 274. Birds: Acridotheres tristis, p. 281. Ducks, p. 277. Melanerpes portoricensis, p. 280. Meleagris gallopavo, p. 278. Passer domesticus, p. 282. Phasianus sp., p. 277. Streptopelia chinensis, p. 278. Mammal: Herpestes javanicus auropunctatus, p. 289.

=Robshelfordia circumducta= or =Robshelfordia longiuscula= Insects: Neorhipidius neoxenus, p. 230. Rhipidioides helenae, p. 230. Rhipidioides mollis, p. 231.

=Steleopyga= (?) =sinensis= Nematode: Suifunema caudelli, p. 200.

=Supella supellectilium= Viruses: Unspecified strain(s) of poliomyelitis virus, p. 103. Bacterium: Serratia marcescens, p. 117. Fungus: Herpomyces supellae, p. 138. Nematode: Blattophila supellaima, p. 194. Insects: Anastatus tenuipes, p. 246. Comperia merceti, p. 244.

=Symploce flagellata= or =Symploce ruficollis= Reptile: Anolis cristatellus, p. 273.

=Symploce parenthesis= Hair worm: Parachordodes raphaelis, p. 202.

=Temnopteryx phalerata= Protozoan: Gregarina gibbsi, p. 182.

=Undetermined Cockroaches= Bacteria: Arthromitus intestinalis, p. 124. Bacillus subtilis, p. 122. Clostridium perfringens, p. 122. Escherichia coli, p. 112. Fusiformis lophomonadis, p. 119. Micrococcus citreus, p. 106. Micrococcus pyogenes var. aureus, p. 107. Mycobacterium leprae, p. 123. Paracolon bacilli, p. 113. Proteus morganii, p. 114. Proteus vulgaris, p. 114. Spirillum sp., p. 105. Spirochaeta periplanetae, p. 125. Streptococcus sp., p. 110. Vibrio Types I and II Heiberg, p. 106. Fungi: Amphoromorpha blattina, p. 139. Amphoromorpha sp., p. 139. Blastocystis sp., p. 133. Cordyceps amazonica, p. 134. Herpomyces anaplectae, p. 134. Herpomyces chilensis, p. 135. Herpomyces diplopterae, p. 135. Herpomyces forficularis, p. 135. Herpomyces grenadinus, p. 136. Herpomyces macropus, p. 136. Herpomyces paranensis, p. 136. Herpomyces periplanetae, p. 137. Herpomyces platyzosteriae, p. 137. Herpomyces tricuspidatus, p. 138. Herpomyces zanzibarinus, p. 138. Metarrhizium anisopliae, p. 131. Plants: Nepenthes ampularia, p. 154. Nepenthes gracilis, p. 154. Nepenthes sp., p. 154. Protozoa: Diplocystis sp., p. 181. Endamoeba blattae, p. 177. Entamoeba histolytica, p. 179. Giardia sp., p. 172. Gregarina blattarum, p. 182. Hexamita periplanetae, p. 171. Lophomonas blattarum, p. 172. Lophomonas striata, p. 173. Monas sp., p. 167. Monocercomonoides orthopterorum, p. 169. Nyctotherus buissoni, p. 188. Nyctotherus ovalis, p. 188. Nyctotherus viannai, p. 189. Oikomonas blattarum, p. 166. Oikomonas sp., p. 166. Paramecium sp., p. 186. Trichomonas sp., p. 171. Helminths: Cephalobellus magalhãesi, p. 194. Chordodes morgani, p. 201. Gordius sp., p. 202. Hammerschmidtiella diesingi, p. 195. Leidynema appendiculata, p. 197. Moniliformis moniliformis, p. 204. Protrellus galebi, p. 199. Schwenkiella icemi, p. 200. Severianoia magna, p. 200. Severianoia severianoi, p. 200. Spirura gastrophila, p. 207. Undetermined gordian worms, p. 202. Scorpion: Heterometrus longimanus, p. 213. Spiders: Ctenid, p. 214. Heteropoda venatoria, p. 215. Latrodectus mactans, p. 216. Mite: Pimeliaphilus podapolipophagus, p. 219. Centipedes: Allothereua maculata (circumstantial evidence), p. 223. Scolopendra morsitans, p. 223. Scolopendra subspinipes, p. 224. Scutigera coleoptrata (circumstantial evidence), p. 222. Insects: Acanthinevania princeps, p. 235. Agamerion metallica, p. 243. Ampulex compressa, p. 259. Ampulex ruficornis, p. 259. Ampulex sibirica, p. 260. Ampulex sonnerati, p. 260. Anastatus blattidifurax p. 245. Anastatus tenuipes, p. 246. Blatticida pulchra, p. 243. Blatticidella ashmeadi, p. 243. Calodexia spp., p. 228. Clerada apicicornis, p. 226. Dicarnosis alfierii, p. 245. Diestrammena apicalis, p. 226. Diestrammena japonica, p. 226. Dolichurus bicolor, p. 260. Dolichurus corniculus, p. 261. Dolichurus gilberti, p. 261. Dolichurus ignitus, p. 261. Dolichurus sp., p. 262. Dorylus nigricans sjöstedi, p. 267. Dorylus sp., p. 267. Dorylus wilverthii, p. 267. Eciton burchelli, p. 268. Eupelmus sp., p. 247. Eutrichosomella blattophaga, p. 245. "Formica omnivora," p. 268. Iridomyrmex humilis, p. 268. Neonephrites partiniger, p. 230. Podium abdominale, p. 265. Podium dubium, p. 265. Podium rufipes, p. 266. Pompilus bracatus, p. 256. Pompilus sp., p. 256. Salius verticalis, p. 256. Ripidius scutellaris, p. 233. Solindenia picticornis, p. 247. Stylogaster spp., p. 228. Stylogaster stylata, p. 228. Syntomosphyrum blattae, p. 248. Systellogaster ovivora, p. 248. Tachysphex blatticidus, p. 264. Tachysphex coriaceus, p. 264. Tachysphex fanuiensis, p. 264. Tetrastichus hagenowii, p. 251. Tetrastichus periplanetae, p. 253. Trirhogma sp., p. 264. Undetermined tachinid, p. 229. Fish: Chalceus macrolepidotus, p. 269. Cyrtocharax magdalenae essequibensis, p. 269. Potamotrygon humboldti, p. 268. Rhamdia sebae, p. 268. Amphibians: Arthroleptis variabilis, p. 270. Bufo ictericus, p. 270. Bufo funereus, p. 270. Bufo marinus, p. 270. Hyla cinerea, p. 270. Hyperolius picturatus, p. 271. Leptodactylus albilabris, p. 271. Leptodactylus pentadactylus, p. 271. Leptopelis calcaratus, p. 271. Leptopelis rufus, p. 271. Megalixalus fornasinii, p. 271. Rana catesbeiana, p. 271. Rana mascareniensis, p. 271. Tree frogs, p. 351. Reptiles: Ameiva exsul, p. 275. Ameiva sp., p. 275. Anolis cristatellus, p. 273. Anolis grahami, p. 274. Anolis leachi, p. 274. Anolis pulchellus, p. 273. Anolis sp., p. 274. Anolis stratulus, p. 274. Cnemidophorus sp., p. 275. Geckos, p. 273. Gekko gecko, p. 272. Hemidactylus frenatus, p. 272. Leiolopisma laterale, p. 274. Lizards, p. 275. Skinks, p. 274. Sphaerodactylus sp., p. 273. Thecadactylus sp., p. 273. Tropidophorus grayi, p. 274. Birds: Acridotheres tristis, p. 281. Agelaius xanthomus, p. 282. Bambusicola thoracica, p. 277. Chickens, p. 278. Coturnix coturnix japonica, p. 277. Dendrocopus mahrattensis, p. 279. Ducks, p. 277. Gymnasio nudipes, p. 279. Gymnopithys leucaspis, p. 280. Holoquiscalus brachypterus, p. 282. Icterus portoricensis, p. 280. Owl, p. 351. Partridge, p. 278. Phasianus sp., p. 277. Sparrow, p. 282. Tiaris bicolor omissa, p. 282. Tockus birostris, p. 279. Troglodytes aedon, p. 281. Troglodytes audax, p. 281. Mammals: Bassariscus astutus, p. 288. Callithrix jacchus, p. 285. Erinaceus sp., p. 283. Felis catus, p. 289. Felis paradalis mearnsi, p. 290. Herpestes javanicus auropunctatus, p. 289 Herpestes sp., p. 289. Homo sapiens, p. 287. Loris tardigradus, p. 284. Molossus sp., p. 283. Monodelphis sp., p. 283. Nasua nasua, p. 288. Nasua sp., p. 289. Rattus sp., p. 287.

XV. COCKROACHES AS COMMENSALS

These particular associations may well have been accidental and due to a predilection for the same type of nesting site. But this fact in no way detracts from the interest of such records. Chance must play a very considerable part in first bringing symbiotic or commensal partners together. Once such a partnership between species has been firmly established, it is on the whole, fairly obvious, ... On the other hand, in the early stages before the relationship has become fixed as a specific habit, individual cases are generally dismissed as coincidences. It is, however, unwise to disregard such isolated observations or dismiss them lightly.

ROTHSCHILD AND CLAY (1957)

The following social insects have been found harboring cockroaches in a state of commensalism in which the cockroaches presumably benefit by acquiring food from their hosts. Benefits accruing to the hosts are not apparent. Unfortunately, biological details are not always sufficient to substantiate the suspected association. However, it seems significant that the cockroach commensals of the insects listed below have been found only in association with their hosts and, so far as we know, have never been found apart from them. Chopard (1938) has pointed out that the myrmecophilous cockroaches are all small, being only a very few millimeters long; they are apterous or subapterous; their eyes are reduced; and they are all of American origin.

HOSTS OF COMMENSAL COCKROACHES

Order ISOPTERA

Family RHINOTERMITIDAE

=Coptotermes ceylonicus= Holmgren

Commensal.--Sphecophila ravana, Ceylon (Fernando, 1957): Six females, 50 males, and nymphs of both sexes were found among decaying timber in the ground in association with a colony of this termite. The antennae of most specimens were mutilated unsymmetrically.

Family TERMITIDAE

=Macrotermes barneyi= Light

Commensal.--Nocticola sinensis, Kowloon (Silvestri, 1947): Among specimens of termites collected from a nest.

=Macrotermes bellicosus= (Smeathman)

=Macrotermes natalensis= (Haviland)

Synonymy.--Termes bellicosus [Snyder, 1949].

Commensal.--Sphecophila termitium, Kibonoto, East Africa (Shelford, 1910): Two males were collected in a termite mound.

=Macrotermes malaccensis= (Haviland)

Synonymy.--Termes malaccensis Haviland [Snyder, 1949].

Commensal.--Nocticola termitophila, Tonkin (Silvestri, 1946): The cockroach was found in the termite nest.

=Odontotermes= sp.

Commensals.--Nocticola sinensis, Kowloon (Silvestri, 1946): In termite nest.

Nocticola termitophila, Penang (Silvestri, 1946): In termite nest.

=Termes= sp.

Commensal.--Nocticola sinensis, Repulse Bay, Australia(?) (Silvestri, 1946): In a termite gallery.

=Termites=

Commensal.--Ergaula capensis [= Dyscologamia wollastoni] French Equatorial Africa, Brazzaville (Rehn, 1926; Chopard, 1949).

Order HYMENOPTERA

Family FORMICIDAE

Subfamily FORMICINAE

=Camponotus femoratus= (Fabricius)

Commensal.--Phorticolea boliviae, Bolivia, Cachuela Esperanza (Caudell, 1923): Three males collected in the joint nests of C. femoratus and Crematogaster limata.

=Camponotus maculatus= (Fabricius)?

Note.--Dr. W. L. Brown (personal communication, 1957) states that this ant is an Old World species only. So presumably Mann's record pertains to a different species.

Commensal.--Myrmecoblatta rehni, Mexico (Mann, 1914): "They were very abundant, several occurring in almost every nest, where they are no doubt very efficient scavengers."

=Camponotus rufipes= (Fabricius)

Commensals.--Atticola mortoni, Nothoblatta wasmanni, and Phorticolea testacea, Brazil, San Leopoldo (Bolívar, 1905): Found in the formicaries of C. rufipes.

=Formica rufibarbis= Fabricius

and

=Formica subcyanea= Wheeler

Commensal.--Myrmecoblatta rehni, Mexico (Mann, 1914): "They were very abundant, several occurring in almost every nest."

Subfamily MYRMICINAE

=Acromyrmex lobicornis= Emery

Commensal.--Attaphila bergi, or possibly a variety of this species, Huasán, Argentina? (Bruch, 1916).

=Acromyrmex lundi= (Guérin)

Synonymy.--Atta lundi [Brown, personal communication, 1957].

Commensal.--Attaphila bergi, Argentina and Uruguay (Bolívar, 1901): The cockroach was found in the nests of the ants sitting on the back, neck, or head of sexual individuals. It remains attached to the ant during swarming. The antennae seem always to be mutilated. Bruch (1916) stated that in La Plata A. bergi is encountered by hundreds in every nest of A. lundi.

=Acromyrmex niger= (F. Smith)

Synonymy.--Atta nigra Schupp [Brown, p.c., 1957].

Commensal.--Attaphila schuppi, Brazil, Porto Alegre (Bolívar, 1905): Found outside the nest of the ant and mixed in the columns of ants on the march.

=Acromyrmex octospinosus= (Reich)

Synonymy.--Atta octospinosa [Brown, p.c., 1957].

Commensals.--Attaphila fungicola, Panama (Wheeler, 1928): Taken in the fungus gardens of the ant.

Attaphila aptera, Esperanza, Dibulla, Colombia (Bolívar, 1905).

=Acromyrmex silvestrii= Emery

Commensal.--Attaphila bergi, or possibly a variety of this species, San Luis Province, Argentina (Bruch, 1916): According to Bruch, the behavior of this species of Attaphila is identical with the one encountered in Huasán in the nests of Acromyrmex lobicornis Emery; it differed from A. bergi in size and color.

=Atta cephalotes= (Linnaeus)

Commensal.--Attaphila fungicola, British Guiana (Wheeler, 1928): Taken in the fungus gardens of the ant.

Attaphila sp., British Guiana (Beebe, 1921): 7 of 12 queens in one nest had cockroaches hanging on them.

=Atta sexdens= (Linnaeus)

Commensal.--Attaphila sexdentis, Brazil, San Leopoldo (Bolívar, 1905): Found in nests of the ant.

=Atta texana= (Buckley)

Synonymy.--Atta fervens Say [Wheeler, 1910].

Commensal.--Attaphila fungicola, U.S.A., Texas (Wheeler, 1900, 1910): The cockroach does not feed on the fungus in the ants' nest, as Wheeler (1900) first supposed, but mounts the back of the soldiers and licks their surfaces. It is tolerated by the ants with no signs of hostility. The antennae of the cockroach are clipped short. Although Wheeler (1910) stated that this is probably accidental or unintentional, it is peculiar that Bolívar (1905) noticed the same invariable mutilation of the antennae of Attaphila bergi. Wheeler (1900) had originally suggested that the antennae were probably clipped off by the ants which are continuously trimming the fungus hyphae. Louisiana (Moser, personal communication, 1959): Numerous specimens were encountered in some nests of A. texana. This cockroach is the most closely associated inquiline in the nest and maintains very intimate terms with the ants. It is found living in the fungus cavities and tunnels.

=Crematogaster limata parabiotica= Forel

Commensal.--Phorticolea boliviae, Bolivia, Cachuela Esperanza (Caudell, 1923): Collected in joint nests of C. limata and Componotus femoratus.

=Solenopsis geminata= (Fabricius)

Commensal.--Myrmecoblatta wheeleri, Guatemala (Hebard, 1917a): Collected from a colony of this ant under a stone on the shores of Lake Atitlan, altitude 11,719 feet.

=Unknown host=

Cockroach.--Attaphila flava, British Honduras (Gurney, 1937): Because the known hosts of the other five species of Attaphila are ants, we presume that this species also lives in the nest of some myrmecine ant.

Subfamily PONERINAE

=Odontomachus affinis= (Guérin)

Commensal.--Myrmeblattina longipes, Brazil, Rio de Janeiro (Chopard, 1924, 1924a; Hancock, 1926): Originally described as Phileciton longipes by Chopard (1924) from the nest of an ant mistakenly identified as Eciton sp.

Family VESPIDAE

=Polybia pygmaea= Fabricius

Commensal.--Sphecophila polybiarum, French Guiana (Shelford, 1906a): Shelford stated that it was probable that the cockroaches living on the floor of the paper nest fed on small fragments of insects and spiders that were dropped by the wasp larvae feeding in the cells above.

Family MEGACHILIDAE

=Melipona nigra= Lepeletier

Commensal.--Oulopteryx meliponarum, Brazil (Hebard, 1921): According to Hebard, this cockroach is the first one to be known to inhabit the nests of bees. Nothing is known of the relationship between the cockroach and the bees. [See comment by Sonan (1924) on page 318.]

CHECKLIST OF COMMENSAL COCKROACHES WITH THEIR HOSTS

The cockroaches are arranged alphabetically by genus and species. The page references are to citations in the classified section above, where details and/or sources of the records are given.

=Attaphila aptera= Ant: Acromyrmex octospinosus, p. 313.

=Attaphila bergi= Ants: Acromyrmex lobicornis, p. 312. Acromyrmex lundi, p. 312. Acromyrmex silvestrii, p. 313.

=Attaphila flava= Host unknown, presumably an ant, p. 314.

=Attaphila fungicola= Ants: Acromyrmex octospinosus, p. 313. Atta cephalotes, p. 313. Atta texana, p. 313.

=Attaphila schuppi= Ant: Acromyrmex niger, p. 312.

=Attaphila sexdentis= Ant: Atta sexdens, p. 313.

=Attaphila= sp. Ant: Atta cephalotes, p. 313.

=Atticola mortoni= Ant: Camponotus rufipes, p. 312.

=Ergaula capensis= Termites, p. 311.

=Myrmeblattina longipes= Ant: Odontomachus affinis, p. 314.

=Myrmecoblatta rehni= Ants: Camponotus maculatus(?), p. 312. Formica rufibarbis, p. 312. Formica subcyanea, p. 312.

=Myrmecoblatta wheeleri= Ant: Solenopsis geminata, p. 314.

=Nocticola sinensis= Termites: Macrotermes barneyi, p. 311. Odontotermes sp., p. 311.

=Nocticola termitophila= Termites: Macrotermes malaccensis, p. 311. Odontotermes sp., p. 311. Termes sp., p. 311.

=Nothoblatta wasmanni= Ant: Camponotus rufipes, p. 312.

=Oulopteryx meliponarum= Bee: Melipona nigra, p. 314.

=Phorticolea boliviae= Ants: Camponotus femoratus, p. 311. Crematogaster limata parabiotica, p. 312.

=Phorticolea testacea= Ant: Camponotus rufipes, p. 312.

=Sphecophila polybiarum= Wasp: Polybia pygmaea, p. 314.

=Sphecophila ravana= Termite: Coptotermes ceylonicus, p. 310.

=Sphecophila termitium= Termite: Macrotermes bellicosus or Macrotermes natalensis, p. 311.

OBSCURE ASSOCIATIONS

Cockroaches that are sometimes found in the nests of, or in association with, other animals are not necessarily commensals. This is particularly true of cockroaches that normally are found unassociated with other animals or that merely occupy the same habitat with the other animals because of similar microclimatic requirements (see Chopard, 1924c).

McCook (1877) excavated in February a nest of Formica rufa in Pennsylvania. A hundred or more lively cockroaches occupied a part of the nest that contained few ants. Near the cockroaches McCook also found a colony of Termes flavipes. Ischnoptera deropeltiformis has been found in the company of ants, but it is probably not myrmecophilous (Donisthorpe, 1900). Mann (1911) found an "Ischnoptera" sp. (probably a species of Parcoblatta) abundant in the nests of, and tolerated by, Camponotus maccooki Forel in California. Dead and mutilated specimens of this cockroach were common in the nests of "Formicas." "Ischnoptera" sp. was also common in the nests of Veromessor andrei (Mayr) [= Stenamma andrei]. Hebard (1917) reported that W. M. Wheeler collected Eremoblatta subdiaphana in Arizona as an ant guest. Rehn (1906a; Rehn and Hebard, 1927) reported that Pholadoblatta inusitata had also been taken by Wheeler from the galleries of a jumping ant, Odontomachus clarus Roger [= O. haematodes insularis Guérin var. pallens Wheeler; Brown (personal communication, 1958)], on Andros Island, Bahamas; Rehn and Hebard (1927) stated that "This genus and species is the only blattid, which is presumably a myrmecophile, known from the West Indies." Rehn (1932a) reported Dendroblatta sobrina as taken in an ant nest in a tree in the Amazon Basin. Tivia australica was taken in an ant nest in Australia (Princis, 1954). The male of Compsodes schwarzi was taken in an ant nest in the Santa Rita Mountains of Arizona (Ball et al., 1942). A male and female of Stilpnoblatta minuta were taken in a migrating column of the ant Myrmicaria natalensis Sm. subsp. eumenoides Gerst. in Nyasaland (Princis, 1949). Princis cautioned that it is premature to derive any inference from this, possibly accidental, association. Four females of Parcoblatta desertae were taken about a nest of an ant, Ischnomyrmex sp. (Hebard, 1943a). A nymph of Parcoblatta virginica was found in a nest of Formica sp. (Hauke, 1949).

Chorisoneura texensis has been found in nests of webworm in Florida (Rehn and Hebard, 1916). Karny (1924) in Malaya found an oötheca of Aristiger histrio (sp.?) between leaves (Costus sp.) that had been stuck together by a thysanopteron, Anaphothrips sp. He pointed out that the oötheca would not adhere to leaves that were not stuck together but would fall to the ground.

Seín (in Rehn and Hebard, 1927) in Puerto Rico found Aglaopteryx facies in abandoned cocoons of Megalopyge krugii (Dewitz) and in leaves webbed together by caterpillars and in abandoned spiders nests. Wolcott (1950; and in Rehn and Hebard, 1927) also found A. facies in the empty cocoons of M. krugii and in the larval tents of Tetralopha scabridella Ragonot on Inga vera (coffee shade tree); and "Where there are no butterfly-nests, it lives in abandoned spider-nests on the leaves of other forest trees." Cotton (in Wolcott, 1950) found the type of Aglaopteryx absimilis also living in the abandoned cocoon of M. krugii on bucare trees in Puerto Rico. Wolcott (1950) reported that Plectoptera dorsalis, Plectoptera infulata, and Plectoptera rhabdota have been found living in trees between leaves or in "butterfly-nests" of Tetralopha scabridella in leaves of Inga vera, or nests of Pilocrocis secernalis (Möschler) in the leaves of Petitia domingensis in the mountains of Puerto Rico. Seín (in Rehn and Hebard, 1927) had collected P. rhabdota in the nest of larvae of T. scabridella.

Wolcott (1950) reported that Nyctibora lutzi had been found in a large rotten stump associating with "'comején' termites [Nasutitermes costalis (Holmgren)], yellow wood-ants and rhinoceros beetle grubs." Rehn and Hebard (1927) found Simblerastes jamaicanus in numbers in the debris of an abandoned termites' nest in Jamaica: "To what extent the species is dependent upon the protection of the termite or other structures remains to be determined."

In Virginia Cryptocercus punctulatus has been found living in the same galleries with Reticulitermes sp., and on the Pacific Coast it has been found occupying the same log with Termopsis sp. (Cleveland et al., 1934).

Shelford (1909) found one male and one female of Balta platysoma in a nest of a spider of the genus Phryganoporus and assumed a symbiotic association. Chopard (1924) recorded Mareta acutiventris from empty nests of spiders on Barkuda Island, India; nothing is known of the relationships, if any, between these cockroaches and spiders.

Chopard (1924c) found Margattea sp. in the nest of the ant Acropyga acutiventris Roger; he also found Margattea sp., Periplaneta sp., Polyphaga indica, and Temnopteryx obliquetruncata in deserted termite mounds in India. However, he believed that none of these species were more than accidental associates of the host insects; he considered them hygrophilous cockroaches which had found a retreat in the nests.

McClure (1936) obtained a large nest of Vespula maculata (Linnaeus) [= Vespa maculata] in March in Illinois. In it were living 65 nymphs of Parcoblatta pensylvanica, 3 spiders (Philodromus pernix Blackwall), 2 immature spiders (Drassus sp.), and 6 mites. Balduf (1936) observed four individuals of Parcoblatta pensylvanica in a nest of Vespula maculata; he suggested that they probably fed on dead bodies and organic wastes of the wasps. However, Rau (1940) has observed this cockroach devour a Polistes larva in its cell. Although we do not imply that a commensal relationship exists between Parcoblatta and the wasp, it is well to recall a statement by Rothschild and Clay (1957): "A commensal relationship is potentially even more dangerous than a merely social tie, for by nature it is more intimate. The closer the association, the more easily is the balance upset. One partner can then suddenly take a mean advantage of the other."

Cockroach nymphs may enter bees' nests where, according to Imamura (in Sonan, 1924), they do not feed on honey or pollen but presumably feed on excreta of bees or anything scattered by bees in their nest; the bees are not disturbed by the cockroaches.

Cockroaches that have been found in the burrows of vertebrates are listed on pages 23-25.

Paulian (1950) found immature cockroaches in the nests of birds (Ploceinae) in Madagascar and Ivory Coast. All nests of Fondia sp. examined in Madagascar contained many cockroaches, and Paulian believed that the blattid was a species peculiar to the nests of birds. Three nests of Ploceus sp. in Ivory Coast yielded one or two cockroaches each in association with more numerous mites, Psocoptera, Heteroptera, beetles, and lepidopterous larvae (Delamare, Deboutteville and Paulian, 1952). These last cited workers also found four cockroaches in a nest of Estrildine sp., and two in a nest of an undetermined bird, all in association with other arthropods. Moulton (1912) observed large numbers of Symploce cavernicola and Periplaneta australasiae swarming in soft bird guano on the floor of caves in Borneo. Abdulali (1942) found in India many Periplaneta americana in caves containing the edible-nest swift; there was no indication of association of the cockroaches with the birds. Danforth (in Wolcott, 1950) reported finding large numbers of Aglaopteryx facies "in the nests of the grey kingbird, in the region of the Cartagena Lagoon [Puerto Rico], 'living among the twigs.'" In Trinidad, Kevan found a male of Blaberus discoidalis in a bird's nest (Princis and Kevan, 1955).

Davis (in Rehn and Hebard, 1914a) stated that "At Punta Gorda [Florida] there was a vacant house at the end of the town frequented at night by a Nanny and Billy goat, and on warm evenings many Periplaneta australasiae would run about on the piazza floor and on the sides of the house. They were seen feeding on the excrement of the goats and were no doubt to a great degree dependent upon them." This is another example of a coprophagous insect that has taken advantage of a particular situation favorable to its survival. Similar associations exist in which many of the domiciliary cockroaches feed on the feces of man and domestic and other animals (Roth and Willis, 1957a).

XVI. COCKROACHES AS PREDATORS

INTERSPECIES PREDATION

Tepper (1893) made the broad statement that the majority of Australian and Polynesian cockroaches appear to be wholly carnivorous, eating other insects, eggs, and larvae. He stated that, because of their voracity and cannibalistic tendencies, the carnivorous species lead more or less solitary lives so that one rarely meets several in close proximity; they are never very numerous at any time because the stronger devour the weaker in the absence of other prey. Tepper stated that Australian species of Ischnoptera hunt for their prey among the foliage of shrubs, and that Australian species of Cutilia [= Drymaplaneta, Hebard (1943)] run about actively on the surface, or ascend shrubs and trees in quest of living insects and therefore are highly beneficial. Tepper (1894) also stated that Geoscapheus robustus ate earthworms, grubs, and caterpillars. Froggatt (1906) and Marlatt (1915) attributed to Tepper the statement that cockroaches, like Epilampra notabilis, which are found out-of-doors in Australia, are carnivorous and feed on caterpillars and other soft-bodied insects; but Froggatt (1907) believed that this alleged behavior needed confirmation.

A number of observations have been recorded which indicate that sometimes cockroaches may be predatory. According to Ealand (1915), nymphs of the cockroach Pseudomops cincta fed on the Argentine ant Iridomyrmex humilis. In the laboratory, Eurycotis floridana has been observed to catch and devour the wasp Anastatus floridanus which parasitizes the eggs of Eurycotis (Roth and Willis, 1954a). Parcoblatta pensylvanica was observed devouring a larva of Polistes sp. in its cell in a deserted wasps' nest (Rau, 1940). Brigham (1866) saw a cockroach kill and eat a centipede four or five inches long. Annandale (1910) described the destruction in Calcutta of termites by Periplaneta americana. During a heavy rain storm many termites flew into the dining room and were set upon by the cockroaches which seized them with their mandibles and began to gnaw their abdomens. If disturbed, the cockroaches carried the termites away in their mandibles without using their legs to seize, hold, or carry the prey. Sometimes only the abdomen, but other times the whole body with the exception of the wings, was devoured. Perhaps this observation led Allyn (Anonymous, 1937) to theorize that, first, cockroaches could eradicate termites from houses, and then the blattids in turn could be eliminated. Falls (1938) has pointed out the unfeasibility of this idea. Blattella vaga has shown some tendency to eat plant lice (Flock, 1941a). Certain small cockroaches found beneath cane leaf-sheaths, in the Philippine Islands, preyed in part upon leafhoppers (Uichanco, in Williams et al., 1931).

Takahashi (1924) stated that the American cockroach will eat the eggs of the hemipteron Cantao ocellatus (Thunberg). Cunliffe (1952) observed mite-infested cockroaches (Blatta orientalis, Blattella germanica, and/or Periplaneta americana) dislodge and eat the mite Pimeliaphilus podapolipophagus. Sonan (1924) reported that cockroaches (P. americana and P. australasiae) devoured the egg clusters and first instar larvae of Prodenia litula and the first instar larvae of Attacus atlas which were being reared in the laboratory. Lederer (1952) stated that Periplaneta americana ate reptile eggs in the aquarium at Frankfurt am Main. Pettit (1940) stated that cockroaches "are said to have destroyed a large colony of dermestids used to skeletonize carcasses at the University of Kansas."

DeFraula (1780) believed that his silent "gryllon" [obviously Blatta orientalis from his drawings; see Willemet (1784)] was the enemy of the chirping species of cricket, because after the cockroach became established in his home he no longer heard crickets chirping. Gilbert White (1905 ed.), writing in England in the late 18th century, stated that "Poda says that these [Blatta orientalis] and house crickets will not associate together; but he is mistaken in that assertion"; however, in August 1792 White noted that "Since the blattae have been so much kept under, the crickets have greatly increased in number." For several years Jolivet (1950) had observed changes in a mixed population of Blatta orientalis and Acheta domesticus in an old kitchen in France. He suggested that the cyclical fluctuations in the relative abundance of the cockroaches and crickets might be caused by reciprocal predatism with one species more susceptible than the other at certain stages. Mallis (1954) has stated that crickets prey on other insects as well as on one another. Lhéritier (1951) had also observed crickets becoming rare in bakeries in France, having been superseded everywhere by B. orientalis; however, he doubted that Jolivet's hypothesis was the correct explanation and suggested that the higher optimum temperature requirements of crickets might be the regulating factor. Lederer (1952) stated that the number of crickets decreased in the aquarium buildings at Frankfurt am Main as the population of American cockroaches increased.

Platyzosteria novae seelandiae was found under the bark of trees in New Zealand devouring bugs (Walker in Shelford, 1909b).

For years it has been believed that cockroaches feed on bedbugs (Cimex lectularius L.) and this statement has been repeated in many reference works and articles. Ealand (1915) stated that cockroaches devour bedbugs with avidity. Even today similar statements are to be found in the literature. "In the old sailing ship days, they [cockroaches] were often welcomed by crews because of the belief that they would eradicate a population of bedbugs. This belief was based on scientific fact, as cockroaches are known as predators of bedbugs" (Monro, 1951). Cockroaches will often "help rid a house of bedbugs by devouring all the little parasites they can capture" (Gaul, 1953). The basis for this belief may have originated with a statement by Webster (1834) who wrote that bedbugs disappeared aboard "H.M. Sloop Chanticleer" when cockroaches made their appearance. Newman (1855) reported the observations of a friend who claimed to have seen a cockroach seize a bedbug in an infested boardinghouse in London. In 1920 Purdy reintroduced cockroaches into a house from which they had been exterminated, in order to control the bedbugs which had become established. According to a popular account by Lillingston (1934) African natives are said to ask sailors for a cockroach or two to be used to hunt bedbugs.

In Siberia, Burr (1926, 1939) found Blattella germanica and bedbugs inhabiting the same room. Mellanby (1939) studied the populations of an animal house in which bedbugs and cockroaches occurred in large numbers; the bugs apparently were not attacked and their numbers increased greatly over a period of a few weeks (Johnson and Mellanby, 1939). Wille (1920) placed starved B. germanica with bedbugs for 20 days, but the cockroaches failed to attack the bugs. In India, captive adults and nymphs of two species of house cockroaches would not touch living bedbugs or their eggs (Cornwall, 1916). In laboratory experiments Gulati (1930) found that Periplaneta americana ate young bedbugs which had soft, blood-filled abdomens; adult bedbugs with harder exoskeletons sometimes were rejected. The maximum number of bedbugs eaten by a cockroach was 3 out of 12 during a period of 48 hours. Johnson and Mellanby (1939), also in laboratory experiments, were unable to show that bedbugs can be controlled by Blatta orientalis or that bedbugs are eaten to any extent by them. The existing evidence indicates that there is little basis for the often repeated statement that cockroaches destroy bedbugs in nature. As Lorando (1929) pointed out, assassin bugs, cockroaches, and red ants can hardly be considered as practical factors in bedbug control, though he did recommend the use of spiders.

According to Martini (1952), cockroaches prey on mosquitoes and sand flies but we have been unable to find any original sources for these statements; the only reference we have found in which cockroaches and Phlebotomus are mentioned together is a paper by Whittingham and Rook (1923); they fed ground-up cockroaches to larvae of Phlebotomus papatasii. Wharton (1951) reported that cockroaches and other predators attacked mosquitoes knocked down by insecticides and affected the number recovered.

Cockroaches will on occasion attack and bite animals other than insects. In an earlier paper (1957a) we discussed about 20 reports of cockroaches biting man. The injury is usually confined to abrasion of the callused portions of hands and feet but may result in small wounds in the softer skin of the face and neck. We failed to include the following reference in the above-mentioned paper. Sonan (1924) had his toes and breast nibbled by cockroaches on Hiyakejima Island during sleep. He had previously learned from a policeman that Periplaneta americana and P. australasiae nibbled people on that island, but he had hardly believed it before he experienced the biting himself.

INTRASPECIES PREDATION

Those who have reared cockroaches in the laboratory have undoubtedly seen cannibalism occur in the cultures. Cannibalism has been observed among the common domiciliary species of cockroaches as well as laboratory colonies of Leucophaea maderae (Scharrer, 1953), and Blaberus craniifer (Saupe, 1928). Edmunds (1957)

reported that cannibalism was common in a laboratory colony of Periplaneta brunnea and that egg capsules deposited by a female were often eaten by the other cockroaches.

Periplaneta americana occasionally ate other cockroaches and their oöthecae and also attacked members of their own species (Lederer, 1952). Griffiths and Tauber (1942) recorded the killing of male American cockroaches by females of the species: "One female was especially vicious and attacked each new male as he was introduced into the container. Most of such males had molted less than 2 days previously. Older males were more capable of defending themselves against attacks of these cannibalistic females." Even though adequate food may be present, females of Periplaneta americana may eat their own eggs (Klein, 1933). Some females may regularly eat their oöthecae as soon as they are dropped (Griffiths and Tauber, 1942). To be completely eaten an oötheca generally must be attacked before it has hardened. If a hole is eaten in one side of the capsule, the cockroach may devour the eggs and leave a portion of the oötheca. Frequently only the keel or a part of the keel is eaten and when this occurs the eggs fail to hatch and usually do not complete development because of the rapid loss of water (Roth and Willis, 1955). When adults of P. americana and P. australasiae were deprived of food, both males and females ate newly deposited eggs and, finally, the females ate the males (Sonan, 1924).

Parcoblatta virginica in laboratory cultures also may eat part of its oöthecae; in this species only the soft end of the recently deposited oötheca was eaten (Roth, unpublished data, 1957).

Cros (1942) observed oöthecae-bearing females of Blatta orientalis attack and kill males of the same species which were attempting to mate; these males were then eaten by the females. Cros also observed injured and recently molted nymphs of B. orientalis to be eaten by others of the same species.

Pettit (1940) noted that cannibalism in his culture of Blattella germanica occurred only when the insects were molting. Adult insects attacked the molting cockroaches more often than did the nymphs. However, nymphs after the fourth instar occasionally set upon other molting nymphs. First-to third-instar nymphs rarely victimized their mates. The victims were all older than third instar; the later stadia were progressively more subject to attack, and molting adults suffered the greatest mortality. No direct correlation was noted between population density and cannibalism.

German cockroaches may attack newly molted nymphs of their own kind and cause them to deflate (Gould and Deay, 1938). Lhéritier (1951) has observed the hatching nymphs of B. germanica being devoured by their congeners even before they have left the oötheca.

Nauphoeta cinerea in laboratory cultures will eat newly hatched young of the same species (Roth and Willis, 1954; Willis et al., 1958). In Hawaii, in nature, N. cinerea may kill and eat the cypress cockroach, Diploptera punctata (Illingworth, 1942; Fullaway and Krauss, 1945).

Bunting (1956) stated that species of Neoblattella are omnivorous with carnivorous and cannibalistic tendencies. An adult female Panchlora sp. was killed and eaten by Neoblattella sp. in captivity. A male, provisionally identified as N. celeripes, was killed and partly eaten by two other males of the same species.

The factors influencing the extent of cannibalism among cockroaches are not completely known. According to Wille (1920) hunger was not the cause of cannibalism in Blattella germanica. Wille claimed that the tendency toward cannibalism increased at high temperatures and decreased at low temperatures. Pettit (1940) also noted this effect. Gould and Deay (1938) stated that under crowded laboratory conditions, when there was a scarcity of food, cannibalism among Periplaneta americana was common. The injured cockroaches and those unable to molt were often eaten. Adair (1923) made similar observations. Undoubtedly, conditions of crowding, availability of food, temperature and other factors all influence cannibalism, but practically no experimental work has been done on this subject.

It is interesting, in comparison with the above positive examples of cannibalism, that both Saupe (1928) and Roeser (1940) observed no cannibalism during extensive studies with Pycnoscelus surinamensis. In fact, Roeser stated that there was never a case of cannibalism in spite of long hunger periods imposed on both nymphal and adult insects.

XVII. ASSOCIATIONS AMONG COCKROACHES

Besides preying on their own species or on other blattids, cockroaches exhibit additional symbiotic relationships among themselves. These relationships are (1) the familial associations of parent and offspring, (2) gregariousness, (3) intraspecies fighting, (4) interspecies compatibility, and (5) interspecies antagonism. There are some inconsistencies between observations made on the same species by different workers, which only further observation and experimentation will explain. Some of the reported observations are unique; this is especially true for the feral species. Because of the paucity of information, it is impossible at this time to make valid generalizations about some of these interesting relationships.

FAMILIAL ASSOCIATIONS

The females of many species of cockroaches insure varying degrees of protection to the developing young in their ways of disposing of the oötheca after it has been formed. The extent of this association between the mother and her developing progeny varies from the minimum amount of time spent by oviparous females in concealing their oöthecae, to the duration of embryogenesis in the so-called viviparous species, a period of over a month or more.

Haber (1920a) observed a female of Periplaneta americana chew a groove in a piece of pasteboard into which she attempted to deposit her oötheca. The oötheca failed to adhere to the shallow hole and fell to the floor. After several futile attempts to replace the oötheca in the hole, the female finally left the egg case on the floor of the cage and coated it with an oral secretion to which she attached bits of trash. During this operation she chased other females away when they ventured near the site. Qadri (1938) described the behavior of the female of Blatta orientalis in concealing her oötheca in a hole that she dug in sand; she deposited the egg case in the hole, coated it with saliva and sand, and then refilled the pit. Rau (1943) described in detail how females of P. americana and B. orientalis covered their oöthecae with wood dust or sand in holes they had prepared in the substrate. Both species placed a sticky oral secretion in the holes and then deposited their oöthecae therein. After coating the oöthecae with more sticky secretion, the females adjusted the oöthecae so that the keels were uppermost and then carefully concealed the oöthecae with the excavated debris. Both females spent over an hour in the act. Rau (1924) previously reported that of 90 oöthecae deposited by B. orientalis in jars containing earth and trash, 36 were placed in crevices or excavated holes, and 38 were hidden by being covered with dirt stuck to them with saliva; only 16 were left uncovered.

Edmunds (1957) described oviposition by Periplaneta brunnea. Some females spent from 30 to 40 minutes secreting from the mouth a frothy substance that was smeared on the substrate; the egg capsule was deposited in the secretion and covered with additional froth, which hardened into a very strong cement. Some females spent as long as two hours coating the capsule after it was deposited. It was not stated whether the oötheca was otherwise concealed. The female remained with her body over the oötheca for several hours and drove away other cockroaches which approached.

Sonan (1924) observed that Periplaneta americana and Periplaneta australasiae spent from 40 minutes to an hour covering their oöthecae, and that if the females were frightened away from this activity, they returned again to complete it. As well as excavating holes in the substrate in which to deposit its oöthecae, P. americana also avails itself of readymade crevices of appropriate size (Ehrlich, 1943). Species of Epilampra in Malaya were said by Annandale (1900) to deposit their oöthecae in crevices in floating logs just above the water line. However, Shelford (1906) stated that four genera (including Epilampra) of the subfamily Epilamprinae are "viviparous," in which event the females would carry their oöthecae within their bodies during embryogenesis and would not place the oöthecae in crevices in logs.

The female of Cryptocercus punctulatus was observed to make a groove in a piece of wood, then carry her oötheca 6 inches from where she had dropped it and place it in the groove; she covered the oötheca so that only a portion was visible (Cleveland in Cleveland et al., 1934). Dr. W. L. Nutting (personal communication, 1954) collected a number of oöthecae of C. punctulatus in the field and found each one almost completely sealed off with bits of wood in a deep groove in the roof of a chamber in a log. The keel of the oötheca was visible but the rest was well camouflaged. He stated that "The adult pair usually frequents the chamber at this time, while their broods of previous years occupy neighboring galleries."

Berland (1924) observed a female of Loboptera decipiens filling a hole (the abandoned nest of a hymenopteron) with earth that she carried in her mouth; he later found her oötheca behind the earthen barricade which she had erected.

In summary, the following species of oviparous cockroaches have been observed concealing their oöthecae (only those references not previously cited are given): Blatta orientalis; Cryptocercus punctulatus; Ectobius sylvester (Harz, 1956, 1957); Epilampra sp.; Eurycotis floridana (Roth and Willis, 1954a); Loboptera decipiens; Balta scripta, Methana curvigera, Methana marginalis, and Methana caneae (Pope, 1953a); Pelmatosilpha marginalis, Pelmatosilpha purpurascens, and Nauclidas nigra (Bunting, 1956); Periplaneta americana (Haber, 1919; Adair, 1923; Seín, 1923; Nigam, 1933; Gould and Deay, 1938; Rau, 1940a); Periplaneta australasiae (Girault, 1915b; Spencer, 1943; Pope, 1953); Periplaneta brunnea (Roth and Willis, unpublished data, 1958); Periplaneta fuliginosa (Gould and Deay, 1940); Periplaneta ignota (Pope, 1953); Supella supellectilium (Flock, 1941). Undoubtedly other oviparous species that drop their oöthecae long before the eggs hatch also make some attempt to conceal the oöthecae by placing them in crevices or covering them with debris.

Sometimes the oöthecae are deposited but not concealed. Hafez and Afifi (1956) reported that in Egypt Supella supellectilium attaches its oötheca to a suitable substrate with a gummy oral secretion but leaves the egg capsule otherwise exposed. We (1954) have noticed similar behavior in laboratory colonies of this species and of Blatta orientalis, as have Gould and Deay (1940). Cornelius (1853) stated that the female of B. orientalis takes care of the safety of her offspring to the extent of usually dropping her oöthecae in places which are dry and raised above the ground, although rarely one also may find some oöthecae scattered on the ground. For lack of suitable material females of Periplaneta americana sometimes did not conceal their oöthecae (Nigam, 1933). Frequently in laboratory colonies P. americana merely drops the oöthecae loosely in sand or food "in contrast to P. australasiae, which almost always went to considerable trouble to fasten their eggs securely and to conceal them with debris" (Pope, 1953). If conditions under which Nauclidas nigra is kept are not suitable, the female will drop her oötheca anywhere (Bunting, 1956). Rau (1940) stated that the female of Parcoblatta pensylvanica does not conceal her oötheca. However, Gould and Deay (1940) stated that this species deposits its oöthecae loosely behind bark. Ellipsidion affine and Ellipsidion australe attach their oöthecae to bark or the underside of leaves but apparently make no attempt to conceal them (Pope, 1953a).

The females of most of the above species have no further familial association with their offspring. The eggs hatch with no attention from the mother who is probably not even in the vicinity at that time. The young apparently do not react to the presence of the parent, as such, after hatching. This is not unexpected, as several additional oöthecae may have been deposited by these oviparous females before the eggs of the first oöthecae hatch. However, a different behavior is encountered among species that do not form a second oötheca until after the eggs of the first have hatched (see below) and in the so-called colonial species.

Shaw (1925) reported that in Australia both Panesthia australis and Panesthia laevicollis appear to live in families, and that one usually finds a pair of adults associated with from 12 to 20 nymphs in different stages of development; he continued, "it is only where the molts are very abundant that one loses sight of this familial habit." Tillyard (1926) also stated that the Australian species of Panesthia live in burrows in soil in strict family communities of a pair of adults and 10 to 20 nymphs. A related colonial species, Cryptocercus punctulatus, lives in both sound and rotten logs in colonies consisting of a pair of adults and 15 or 20 nymphs, probably representing two or three broods (Cleveland et al., 1934; Cleveland, 1948). Chopard (1938) has cited this association as an example of gregariousness, which it may well be; however, the presence of only one pair of adults in each colony suggests a more intimate relationship.

Among species of Blattella and certain other genera with similar reproductive habits the female carries her oötheca clasped in her genital cavity with the posterior portion projecting behind her. Each normal oötheca is carried for approximately the duration of embryogenesis and is not dropped until, or shortly before, hatching. We have seen (1954, fig. 65) newly hatched nymphs of Blattella vaga crawl over the body of the mother who stood quietly near the dropped oötheca; this female raised her wings and some of the nymphs crawled under them onto the dorsal surface of her abdomen. The nymphs seemed to feed on the grease covering the mother's body. The association was short-lived, however, and soon the nymphs scattered. Pettit (1940) stated that when hatching of Blattella germanica occurs in the open (on a table top), the nymphs may remain near the capsule only a few minutes. Ledoux (1945) found that newly hatched nymphs of B. germanica remained together without shelter in a single, sparse group. If the nymphs were separated by blowing on them, the group quickly reassembled, usually in the same spot. Ledoux showed that this gregarious grouping of first-instar nymphs was not necessarily a familial association by placing nymphs from two oöthecae together. In groups of 8 to 12 nymphs there was a perfect intermingling of the offspring from the two different females.

It is among the so-called viviparous cockroaches that the greatest number of observations have been made of postparturient associations between female cockroaches and their offspring. The females of these species carry their oöthecae in brood sacs within their bodies until embryogenesis has been completed. This behavior ensures protection of the young from desiccation and attack by parasites (Roth and Willis, 1955a). (See Roth and Willis, 1958a, for an analysis of oviparity and viviparity in the Blattaria.) Shelford (1906, 1916) reported that he had captured a female of Pseudophoraspis nebulosa in Borneo with numerous young nymphs clinging to the undersurface of her abdomen. He also recalled that there was in the Hope Museum (Oxford) a female of Phlebonotus pallens to which the following label was attached: "'Ceylon ... carries its young beneath its wing covers. 1878.'" Pruthi (1933) found in South India another female of P. pallens which was carrying over a dozen young nymphs on her back beneath her wings. In his paper Pruthi reproduced a photograph of this specimen with the light-colored nymphs in place on the back of the female. Hanitsch (1933) reported having seen a museum specimen from Luzon, Philippine Islands, of the apterous female of Perisphaerus glomeriformis with nymphs still clinging to her undersurface; he also reported having seen a museum specimen of a female of Ellipsidion variegatum from Australia with four young clinging to the upper side of the apex of her tegmina and six to the oötheca which projected beyond her body. Presumably this specimen was giving birth when captured. Gurney (1954; personal communication, 1958) stated that specimens of Perisphaerus sp. from Mindanao and Luzon have been found with young nymphs clinging to the middle and hind coxae. The first-instar nymph has an elongate face and specialized galeae. Karny (1925) also observed that at the slightest alarm the young of some species of Phoraspidinae creep under the dome-shaped front wings of the mother.

The newly hatched young of Leucophaea maderae have also been seen congregated under the mother on several occasions. Seín (1923) stated that after being born, the nymphs of this species gather under the mother and accompany her at night in her excursions in search of food. Pessôa and Corrêa (1928) reported that "During the first days the free larvae hide under the adult cockroach which becomes restless and active in contrast to its usual slow gait." Wolcott (1950) stated that "They are not only gregarious, but the mother broods over her young, and together they sally forth at night in search for food, until they are of such a size as to mingle with their elders."

The African mountain cockroach Aptera fusca has been observed during late summer and early winter in familial groups beneath loose bark, under stones, and in dead leaves (Skaife, 1954): "Each party consists of a number of black young ones, together with one, two or more adult females and perhaps a winged male or two. Later on they scatter and live more or less solitary lives." In Malaya Karny (1924) often found phoraspidine females between leaves surrounded by about 20 young nymphs. He stated that one also often found females of Perisphaerus armadillo surrounded by pale, yellowish-white young; similarly he had observed that Archiblatta hoevenii was found mostly in colonies made up of mothers and their young. The duration of these associations is not known.

Saupe (1928) noticed that the newly hatched nymphs of Blaberus craniifer (see footnote II, p. 322) collected together under the body of their mother and stated that this is as pronounced a case of brood care as Zacher had observed with Pycnoscelus surinamensis. Nutting (1953) stated that "A degree of maternal solicitude is exhibited by this roach [B. craniifer], for many times I have observed the female to remain motionless for an hour or more with her unpigmented brood clustered around and beneath her body." We, too, have observed similar behavior in laboratory colonies of B. craniifer and Leucophaea maderae.

Chopard (1950) noted that after hatching the young of Gromphadorhina laevigata remained grouped around the female for some time; the mother stood motionless, high on her legs, with her thorax curved up to make room for the brood which hid under her body. We (unpublished data, 1958) have seen young nymphs of Gromphadorhina portentosa also stay near their mother for some time after birth; the mother at this time produced a characteristic hissing sound when she was only slightly disturbed by the movement of our hand near her and her brood. The sound is produced as air is expelled through the second abdominal spiracle. We have seen recently hatched nymphs of Nauphoeta cinerea crawl beneath the mother, even under her wings, where they remained about an hour (Willis et al., 1958). Bunting (1956) observed a female of Blaberus discoidalis collect a mound of debris into which she inserted the tip of her abdomen; he found young in the mound later the same day. This female showed no maternal care for the young after birth. Whole families of cockroaches may be found in bromeliads in Brazil (Ohaus, 1900). Hebard (1920) observed a colony of adults and young of Dendroblatta sobrina on a tree trunk in the Panama Canal Zone.

Whether any of the above associations exemplify maternal care for the newly hatched young is questionable. The behavior of the mother, beyond placing her eggs in a suitable location, seems to be entirely passive. The first-instar nymphs are the active partners in these associations, and they may merely be seeking shelter under the nearest convenient object rather than under the mother as such. More extensive studies of some of these relationships will be needed before claims for maternal care, as suggested by Scott (1929), can be substantiated.

GREGARIOUSNESS

Casual statements that cockroaches are gregarious are often encountered in the literature. There has been some argument to the effect that large numbers of these insects seeking the same environment in a limited space would appear to be gregarious, whereas there is probably no true social tendency (Rau, 1924). Reactions of cockroaches to certain stimuli in the environment undoubtedly do result in aggregations of individuals. However, as Chopard (1938) has pointed out, it is difficult to assign the respective parts played in assembling by the attraction of the milieu and by gregarious instincts. Chopard (1938) also stated that Orthoptera with a gregarious tendency are found rarely isolated; one finds them, on the contrary nearly always collected in the same shelters, close together, as if conscious of a need for contact between themselves. He continued further that one can be tempted to attribute the assembling to taxes but that interattraction equally plays an important role; for example, if one places a large number of cockroaches in a container and offers them similar shelters composed of cardboard tubes, one finds that nearly all the individuals will assemble in one of the tubes, ignoring the others. Pettit (1940) claimed that in Blattella germanica gregariousness seemed to depend on the mutual attractiveness of body secretions as well as a thigmopositive behavior and love of warmth.

Ledoux (1945) has studied experimentally gregariousness and social interattraction in Blatta orientalis and Blattella germanica. He also found that the cockroaches tended to collect in shelters containing other cockroaches. He concluded that group formation is not the result of chance, but is a social phenomenon, and that interattraction is mainly olfactory, conditioned by (1) positive chemotaxis to odors emitted by the cockroaches themselves, (2) positive hygrotaxis, and (3) thigmotaxis. He found also that large groups are not stable and tend to break into smaller groups.

Gregariousness in the Orthoptera varies in intensity according to the species and within a species according to the age or physiological state of the insects (Chopard, 1938). This is well exemplified by several of the blattid species discussed below.

Gregarious groupings of cockroaches have been observed most frequently among the domiciliary species. A few examples will suffice. Gal'kov (1926) observed heavy infestations of undetermined cockroaches in workers' living quarters in the Ural region: "In the corners near the stove, the cockroaches covered the walls in a dense carpet." After fumigating he collected about 135,000 dead cockroaches from one barracks and about 475,000 from another. We have reviewed a few other examples of heavy infestations in our 1957(a) paper.

Periplaneta americana was observed by Gould and Deay (1938) in an old meat-packing building in Indiana. Adult cockroaches were present in large numbers between closely placed beams, but the nymphs were more common in cracks between bricks. Clusters of several hundred cockroaches were seen on the open walls of the cold, dark hide room. Gould and Deay stated "American roaches of all sizes live together in perfect harmony. Young nymphs have been noted in clusters underneath adults and crawling over the adults as they wander about in rearing jars." In the monkey house of the Hamburg zoo, P. americana spent most of the day in the cellars resting on the walls in groups of about 200 individuals (Brecher, 1929). Lederer (1952) noted that in closed, dark, heated spaces under the aquarium at Frankfurt am Main, P. americana rested in groups of 20 to 30 individuals; he stated that it was remarkable that the "herd" divided itself into groups each of which usually contained insects of the same age or stage of development. Eads (1954) found P. americana in 40 percent of 762 sewer manholes in Tyler, Tex.; 13 percent of 670 of these manholes were heavily infested with 100 or more cockroaches in each. Other heavy sewer infestations have been reviewed in our 1957(a) paper.

Ehrlich (1943) has stated that Periplaneta americana exhibits social behavior. For instance, cockroaches of various ages inhabit a fairly large space jointly; the adults and older nymphs sense approaches with their antennae and warn and protect the young by a beating of wings and by body movements. There is complete utilization of the available living space; the imagos drive older nymphs from their resting places, and the older nymphs drive out the younger ones, until all cracks, depending on their size, are occupied by various age groups of different sizes. In his experiments Ehrlich observed that in cages with no hiding places the cockroaches would group together; when given a choice of small and large shelters, P. americana hid only under the larger ones that could shelter more insects. Finally, the cockroaches ceased to bite and fight each other when they crowded together in the face of danger.

Of Blatta orientalis Marlatt (1915) stated "This species is notably gregarious in habit, individuals living together in colonies in the most amicable way, the small ones being allowed by the larger ones to sit on them, run over them, and nestle beneath them without any resentment being shown." Haber (1919) also observed that this species is often noticed "huddled together, the younger ones crawling over, around, and beneath the older ones."

Wille (1920) observed that nymphs of Blattella germanica remained almost constantly in groups during the first and second instars, but less so during the third instar. He believed that the aggregations of young occurred because they could occupy narrow crevices where the larger insects could not penetrate. At usual room temperatures the older nymphs and adults lived completely isolated, but at certain temperatures they gathered together in large, tightly pressed groups.

Supella supellectilium is said to be gregarious (Gould and Deay, 1940). The smaller nymphs aggregate in small groups in rearing containers, but the older ones remain separate from one another (Hafez and Afifi, 1956). Leucophaea maderae is sociable and rarely found alone; in their favorite hiding places, hills of these cockroaches can be seen hanging together (Seín, 1923). Wolcott (1950) also stated that L. maderae is gregarious. Annandale (1900) observed that in the "Siamese Malay States" large colonies of Periplaneta australasiae conceal themselves in hollows of bamboo logs from which houses are built. Moulton (1912) stated that he was astonished at the large numbers of P. australasiae and Symploce cavernicola that he saw swarming on the sides of caves of Mt. Jibong, Borneo.

Rehn and Hebard (1905) stated that in Key West, Fla., Eurycotis floridana fairly swarmed under the coquina boulders in the woods, in groups of a dozen containing both young and adults; Pycnoscelus surinamensis was very abundant in the same type of habitat. Caudell (1905) also found the young of E. floridana with the mature individuals. Hebard (1917) in his discussion of Lattiblattella rehni again mentioned finding frequent colonies of E. floridana in Florida. He also found many specimens of Blaberus craniifer under boards on the ground at Key West. He found Parcoblatta lata numerous under bark of dead pine trees in Alabama. However, Dowdy (1955), in an ecological study of oak-hickory forest in Missouri, stated that "Parcoblatta [sp.] were never recorded as being gregarious, in fact they were mostly solitary. However, in some cases two were found together." Yet Blatchley (1895) stated of Parcoblatta pensylvanica that in the winter in Indiana "One cannot pull the loose bark from an old log without dislodging a colony of from ten to a hundred of the nymphs of various sizes." Males of Parcoblatta virginica were said to be often gregarious beneath loose bark and under chunks and rubbish (Blatchley, 1920).

Rehn and Hebard (1927) quoted observations made earlier by Hebard on Byrsotria fumigata in Cuba: "I found the specimens under flat stones, sometimes in colonies of 3 or 4 mature specimens and numbers of immature individuals in all stages of development." These observers also reported that Aspiduchus borinquen was found in Puerto Rico in a limestone cavern by thousands in the grass and on the walls. J. W. H. Rehn (1951a) stated that a related species, apparently Aspiduchus cavernicola, was seen in great numbers on the side walls and roof of a cave in Puerto Rico, but it was not possible to collect any of these and, we infer, confirm the species. Rehn and Hebard (1927) in their account of Simblerastes jamaicanus reported finding it in numbers in a termite nest. Pemberton and Williams (1938) stated that Diploptera punctata is of gregarious habits in Hawaii. Saupe (1928) observed a strong "herd instinct" in all age groups of Blaberus craniifer. Bunting (personal communication, 1956) stated that large nymphs and adults of Blaberus discoidalis "congregate in narrow cracks or on the underside of some low object. The younger nymphs keep in close communities of approximately the same age." Sonan (1924) stated that in Formosa(?) Salganea morio is usually found in groups of six or seven in decayed trees. Species of the genus Litopeltis may be found in small groups as they are somewhat gregarious (Rehn, 1928).

The physiological or psychological effects of gregariousness, or lack of it, are interesting aspects of the basic phenomenon. Landowski (1937) studied in Blatta orientalis the effect on development and growth of the transition from life in complete isolation to life in groups. He kept nymphs in groups of 1, 2, 4, 8, and 16 in jars of identical size and shape. Landowski found that (1) mortality increased with the size of the group and with age, as each animal occupied more of the available space. [Presumably these factors are less detrimental in nature where the group is unconfined.] He further found that (2) life in complete isolation extended the time required to produce an adult insect; and (3) the mean weight of the adult insect was, generally, in inverse proportion to the number of nymphs raised together; isolated insects usually attained the greatest adult weight.

Similarly, Griffiths and Tauber (1942a) found that isolation extended the period of nymphal development in Periplaneta americana. As most of their isolates died before reaching maturity, these workers concluded that the American cockroach does not thrive when individually isolated and that several individuals must be together for optimum development to occur. Pettit (1940, 1940a) observed that isolated nymphs of Blattella germanica take longer to mature than those reared in groups. Wallick (1954) found indications in B. germanica that there is an inverse relationship between population density and individual weight; as the population decreased the weight increased. He also noted an inverse relationship between population density and life expectancy in this species.

We (Willis et al., 1958) have confirmed the above observations that Blattella germanica, Blatta orientalis, and Periplaneta americana complete nymphal development in less time when reared in groups rather than individually. We (loc. cit.) also found that nymphs of the following additional species matured more quickly when reared in groups: Eurycotis floridana, Periplaneta fuliginosa, Supella supellectilium, Nauphoeta cinerea, and Pycnoscelus surinamensis; only a very slight decrease in the average length of the developmental period was found in grouped nymphs of Leucophaea maderae.

Wharton et al. (1954) observed that virgin adult males of Periplaneta americana that had been individually isolated upon emergence were almost wholly unresponsive to the sexually stimulating, female odor for a test period of four weeks. Similar males of comparable age that were kept in groups reacted strongly from the sixth day on. Removal of reactive males from the group inhibited the reaction in these isolates, but the response returned when the insects were regrouped. We (1952) had similarly observed that no isolated male of Blattella germanica was ever seen to give a courting response without having received some form of external stimulation. Yet when numbers of males were kept together isolated from females, on several occasions the males became active and a few individuals gave a courting response. As the sexual stimulus is received by the male of B. germanica through contact rather than odor, as in P. americana, presumably it was mutual contact between the grouped males that released the courting activity.

Cloudsley-Thompson (1953a), in his studies of diurnal rhythms in Periplaneta americana, observed a steady decline in total activity in successive 24-hour cycles: "When two cockroaches, even of different species (P. americana and P. australasiae) were kept together, this depression did not appear to set in so readily." The associates apparently kept each other active.

Isolated females of Periplaneta americana can be conditioned to run a simple maze with less time and fewer errors per trial than when paired or when a member of a group of three (Gates and Allee, 1933). There was less activity, and accordingly fewer errors per minute, among cockroaches tested as pairs and groups of three than as isolated individuals. This observation should not be contrasted with that of Cloudsley-Thompson (1953a), cited above, because the intervals during which activity was observed were quite different.

In the above account we have presumed that aggregations of some species are indications of gregariousness. However, until gregariousness has been proved experimentally for each species, we concede that reactions to environmental stimuli might be sufficient to bring about some of the observed groupings without any interaction between individuals.

In concluding this section we note that Tepper (1893) stated that carnivorous cockroaches in Australia lead more or less solitary lives, and that one rarely meets several together in close proximity. Takahashi (1940) observed that in Formosa Blattella humbertiana does not have a tendency to throng together. Rau (1947) stated that the adults of Ischnoptera deropeltiformis showed no tendency toward gregariousness, but in the laboratory newly hatched young lived close together under bark and remained together throughout the nymphal stages. We wonder whether this gregariousness was not imposed by the restricted quarters of the cage. As mentioned above, Dowdy (1955) did not find Parcoblatta sp. to be gregarious in the field.

INTRASPECIES FIGHTING

Fighting occurs among cockroaches of the same species over food or shelter or between males. Saupe (1928) observed late-instar nymphs of Blaberus craniifer attack each other and even adults. Additional records cited in the section on intraspecies predation (p. 322) imply fighting within a species. Rau (1924) saw a male of Blatta orientalis attack another male in copula and bite away a large portion of its wing. Two other males in the container had their wings badly torn overnight, presumably as a result of fighting.

Ehrlich (1943) stated that individuals of Periplaneta americana that are feeding will ward off intruders by spreading their wings and pushing with their hind legs. However, the intruder will approach again and again biting the feeder in the legs and wings. Frequently the odor of approaching food was sufficient to cause the cockroaches to fight and bite each other. Biting and fighting also occurred when individuals of this species defended their daytime hiding places. A position of attack is assumed when two antagonistic individuals of P. americana meet (Ehrlich, 1943, fig. 14). The insects raise their bodies slightly above the ground, by extending their legs, and they stretch their heads forward horizontally so that their mouth parts protrude; when the insects jump at each other, they may wound each other severely in the soft parts of the body. Fighting between sexually excited males resulted in injury to their legs, wings, cerci, and other parts of the body. Frequently an insect that could no longer defend itself was killed. Lederer (1952) also made similar but less extensive observations on fighting in this species.

Pettit (1940) quoted Woodruff as stating that nymphs of Blattella germanica, apparently healthy and perfectly normal, would do battle for no apparent cause other than a chance meeting, and that occasionally the fight was to the finish, the loser being eaten. Pettit could not substantiate such voracious attacks, although he saw nymphs engage in fights lasting about two seconds during which one would be driven off by vigorous bites on legs or cerci. Small nymphs of B. germanica tended to ignore each other, but third-and later-instar nymphs would engage in "quarrels" of short duration when two met. Pettit noted that males of B. germanica that were crowded together quickly set upon, but did not always kill, other cockroaches introduced into their cage. When he isolated a dozen males in a small cage, they became quarrelsome and three of the group were killed and partly eaten. After several days the surviving males had taken positions so that each was equidistant from his neighbors. Some of these males attacked other males and a female that were introduced, by biting their legs and cerci. Females under similar conditions were much less aggressive, although Pettit saw some females that roved about biting all large members of the group that were within easy reach.

We have frequently observed aggressive behavior between males of Nauphoeta cinerea, which resulted in torn wings. The males would wrestle with each other rolling over and over.

INTERSPECIES COMPATIBILITY

We agree in essence with Chopard (1938) who stated that it is improper to speak of associations apropos of the ecological distribution of Orthoptera. He continued that it is clearly evident that different species of Orthoptera, which are found grouped on a territory more or less narrowly limited, have no interdependence among them. Their grouping results uniquely from almost similar reactions to the different factors which characterize this limited milieu. There is neither interdependence nor interaction; the grouping is a false biocoenose, born under the action of the environment, and does not survive a modification of this milieu.

However, as there are numerous examples of mutual toleration between different species as well as examples of incompatibility, the subject has more than academic interest even if no true ecological significance. On the other hand, further study may show that certain of these associations are definitely ecological, particularly among the feral species. As might be expected, most of the following examples pertain to domiciliary cockroaches.

Dozier (1920) occasionally found Periplaneta americana with Eurycotis floridana in decaying stumps, beneath loose bark of decayed trees, and beneath corded wood. Adair (1923) stated that in his house in Egypt Periplaneta americana, Blatta orientalis, and Blattella germanica were found together in a cupboard. Sambon (1925) found B. orientalis and B. germanica side by side but not fraternizing in a home in Italy. Gould and Deay (1938) observed that apartments over stores were infested with both B. germanica and P. americana, but did not indicate whether these occupied the same microhabitat. Gould and Deay (1940) observed that in the Purdue University greenhouse Periplaneta fuliginosa was found "under benches, boxes, pots and other objects in association with the American roach." Dr. L. A. Hetrick (personal communication, 1954) wrote us that several summers before he had had a mixed infestation of cockroaches, which included Periplaneta australasiae, Periplaneta fuliginosa, and Pycnoscelus surinamensis, in his chicken shed.

Eads (personal communication, 1955), in response to our inquiry about the mixed populations of cockroaches that he had reported infesting sewers in Texas (Eads et al., 1954), stated that "Each of the ten colonies of B. orientalis found in Tyler manholes were associated with larger colonies of P. americana. True breeding colonies of B. orientalis appeared to be present since all the developmental stages were taken. The same situation existed with the P. fuliginosa and the two species of Parcoblatta. Larger colonies of P. americana were associated with the other species in each case. From our limited observations the two species always appeared to be perfectly compatible." Eads et al. (1954) had found Periplaneta fuliginosa in three manholes, Parcoblatta bolliana in one manhole and Parcoblatta pensylvanica in one manhole. We assume that the groups of each species were spacially discrete so that they were recognizable as colonies. Dr. T. A. Olson (personal communication, 1958) has observed two or more species of cockroaches in a single structure but never in mixed colonies. Each species was separated physically from the others. Olson concluded that cockroaches of different species do not mingle freely unless forced to do so by some special environmental condition. Pettit (1940) found B. germanica and P. americana similarly separated in the same building or even in the same basement laboratory.

Perkins (1899) found Lobopterella dimidiatipes generally living in company with the young of Periplaneta australasiae in Hawaii. Rehn and Hebard (1914) in Florida found P. australasiae abundant with Periplaneta americana on a quarter-boat. They also noted that the forficulid Marava [= Prolabia] arachidis (Yersin) appeared in numbers in a kitchen after dark accompanied by swarms of P. americana. These workers also found Leurolestes pallidus in a fruit store in Key West "where the species was common in a pile of old burlap bags and in cracks under the stands which it shared with one fairly large colony of Blattella germanica, occasional specimens of Holocompsa nitidula, a few specimens of Periplaneta americana, and one specimen of Supella supellectilium." They also found H. nitidula with Blaberus craniifer "between old boards in a woodshed, where nymphs were more numerous than adults."

Rehn and Hebard (1914) stated of Supella supellectilium in Florida that "The females were all taken in cupboards where Blattella germanica was found in swarms." The association in human habitations of S. supellectilium and B. germanica has been reported also by Sein (1923), Puerto Rico; Shaw (1924), Australia; Mallis (1954): "German and brown-banded roaches were often found in the same crevice."; Anonymous (1958), Texas; and Anonymous (1958a), Georgia. Gould and Deay (1940) stated that other species of cockroaches, especially B. germanica, may be found with S. supellectilium in the same part of a building. Yet Shaw (1925) stated that "when Supella supellectilium Serv. invades places already occupied by Blattella germanica L., it tends to oust the latter."

Blaberus discoidalis has been found in homes or in fruit debris in Puerto Rico in company with the more common, domiciliary species Leucophaea maderae, but never in abundance (Sein, 1923; Wolcott, 1950). Illingworth (1915) in Hawaii found Symploce hospes associated with Nauphoeta cinerea, Graptoblatta notulata, and Diploptera punctata.

Hebard (1917) found Aglaopteryx diaphana in a bromeliad on a forest tree in Jamaica together with Nyctibora laevigata and numerous Cariblatta insularis. He also found numerous Aglaopteryx gemma under signs on longleaf pines in Alabama with occasional specimens of Parcoblatta lata. In Virginia he found Parcoblatta uhleriana in a decaying chestnut log with Cryptocercus punctulalus. In Florida he found Latiblattella rehni with Eurycotis floridana and, more rarely, with Periplaneta australasiae under bark of pine trees. In Key West he found Symploce hospes in the cupboard of a hotel with swarms of Blattella germanica and a few Supella supellectilium.

Rehn and Hebard (1927) in their study of West Indian blattids reported finding Neoblattella proserpina in epiphytic bromeliads in Jamaica in company with Neoblattella eurydice and Neoblattella dryas. They also list most of the associations cited by Hebard (1917).

Ramme (1923) reported that he found in Germany four species of Ectobius (lapponicus, lucidus, pallidus, and sylvester) living together in an area about 50 m. by 200 m. Although he had stated that his specimens of E. lucidus were a distinct species in 1923, Ramme (1951) later decided that they were a form of E. sylvester, E. sylvester f. lucidus.

Dow (1955) reported trapping Blattella germanica, Periplaneta americana, and Periplaneta brunnea in houses and privies in south Texas. At our request Dr. Dow (personal communication, 1958) analyzed his records to determine whether there were indications of associations between these species, with the following results:

As stated in my published note, the roaches were at first classified to genus only. The 83 Periplaneta subsequently identified to species represented 28 different collections, 11 from houses and 17 from privies, all in Pharr, Texas. Tabulation of the data shows first that P. americana was taken only once in a house and that P. brunnea was taken only 4 times in privies. Of course this distribution greatly reduces the probability that they would be caught together, and it is not surprising that P. americana was trapped alone in the single house collection. P. brunnea, however, was trapped with P. americana 2 of the 4 times it occurred in privy collections.

To investigate the occurrence of Periplaneta with and without Blattella, an analysis has been made of 560 trap collections taken in 40 houses and 40 associated privies in Pharr, Texas, in weekly intervals (from May 14 to June 23 ). In the houses, Periplaneta and Blattella were caught in the same jar 26 times, Periplaneta alone 12 times, Blattella alone 83 times, and neither genus 159 times. In a fourfold table, the value of chi-square (14.7) is significant and indicates that the frequencies are not proportional. The number of times Periplaneta and Blattella actually occurred together (26) is, however, much larger than the expected number calculated from the row and column frequencies (14.8). In the privies, Periplaneta and Blattella were caught in the same jar 9 times, Periplaneta alone 50 times, Blattella alone 18 times, and neither genus 203 times. In a fourfold table, the value of chi-square (1.95) is not significant but the same type of disproportion is evident and the expected frequency of both genera in one trap is 5.7, lower than the actual frequency of 9. Both immature and adult roaches are included in this analysis.

The above evidence would be more satisfactory if based on more extensive data. There is also a possible objection in that the traps were operated for at least overnight, during which time one species could theoretically supplant another. Of course, it is doubtful that there is anything involved here like territory (in the ornithologists' sense). On the other hand, it is well to consider that Periplaneta and Blattella are both likely to be more abundant in the same type of favorable location and that this factor might offset in part some direct antagonism between the species.

The only known specimen of Ischnoptera podoces was captured in company with the type series of Cariblatta nebulicola, in dead leaf litter in Jamaica (Rehn and Hebard, 1927). In Florida Periplaneta australasiae was often taken in company with Pycnoscelus surinamensis and Eurycotis floridana (Blatchley, 1920).

INTERSPECIES ANTAGONISM

In contrast to the presumably amicable associations mentioned above, other observations in the literature seem to indicate that some species of cockroaches are incompatible when they attempt to occupy the same habitat niche. Marlatt (1915) stated "Rarely do two of the domestic species occur together in the same house. Often, also, of two neighboring districts one may be infested with one species, while in the other a distinct species is the commoner one. The different species are thus seemingly somewhat antagonistic, and it is even supposed that they may prey upon one another, the less numerous species being often driven out." Phelps (1924) stated "Roaches of different species are rarely found together, although roaches of the same species live together on very amicable terms."

In 1859 Darwin (1887) stated that "In Russia the small Asiatic cockroach [Blattella germanica?] has everywhere driven before it its great congener [Blatta orientalis?]." Yet in France Girard (1877) suggested that the oriental cockroach be introduced into a restaurant infested with the German cockroach as the best way to expel the latter, because the more robust species drives away cockroaches of smaller size. Wille (1920) in Germany found usually only one species of cockroach in a house. Yet when he placed B. orientalis and B. germanica together, there were no reciprocal attacks even by hungry individuals. Wille concluded that because of their greater speed, smaller size, greater number of eggs, and faster development, the German cockroaches eat the available food and so make the environment unfavorable for the oriental. However, he noted that cases may be seen in which the opposite is also possible. Laing (1946; British Museum [Natural History], 1951) observed that in the British Isles B. orientalis seems to have lost its dominant position to B. germanica in recent years; it was stated that these species are not as a rule found together and that the greater rapidity of breeding and ability to climb of B. germanica, as well as the layout of modern buildings, are some of the factors that favor the spread of B. germanica. Ledoux (1945) found that first-instar nymphs of B. germanica and fourth-instar nymphs of B. orientalis, adults of B. germanica and sixth-instar nymphs of B. orientalis, as well as adults of both species, did not form mixed groups. However, when he combined fifth-and sixth-instar nymphs of B. germanica with fourth-and fifth-instar nymphs of B. orientalis, which are all practically of equal size, sometimes he would find mixed groups, but generally the groups were distinct. Lucas (1912) stated that Burr had found B. germanica and B. orientalis swarming within a rubbish heap in England; presumably both colonies were breeding and multiplying and one species was not detrimental to the presence of the other.

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 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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1892. Ueber das Vorkommen von bakterienähnlichen Gebilden in den Geweben und Eiern verschiedener Insecten. Centralbl. Bakt. Parasitol., vol. 11, pp. 234-240.

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1936. Untersuchungen über die Symbiose von Tieren mit Pilzen und Bakterien. V. Die Bakteriensymbiose bei Blattiden und das Verhalten von Blattiden bei aseptischer Aufzucht. Arch. Mikrobiol., vol. 7, pp. 391-403. [Translated by Medical Literature Service.]

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1930. Die Schädlingsfauna a Palästinas. Monogr. Angew. Ent., Zeitschr. Angew. Ent., vol. 16. No. 10. pp. 1-439.

1951. Insects as human food. 352 pp. The Hague.

1953. Problems of animal ecology and physiology in deserts. In Desert Research. Res. Counc. of Israel, Spec. Publ. No. 2. pp. 205-229.

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1930. Untersuchungen über die Gewächshausfauna Unter-und Mittelitaliens. Zeitschr. Morph. Ökol., vol. 19, pp. 534-590.

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1892. Orthoptères. In Voyage de M. E. Simon aux Iles Philippines. Ann. Soc. Ent. France, vol. 61. pp. 29-34.

1897. Viaggio de Leonardo Fea in Birmania e regioni vicine. 78. Nouvelle espèce cavernicole de la famille des Blattaires. Ann. Mus. Civico di Storia Nat., Genova, ser. 2, vol. 18, pp. 32-36.

1901. Un nuevo ortóptero mirmecófilo Attaphila Bergi. Com. Mus. Nac. Buenos Aires, vol. 1, pp. 331-336, 1 pl.

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1911. On a collection of insect-remains from the South Wales coalfield. Quart. Journ. Geol. Soc. London, vol. 67, pp. 149-174. pls. 7-10.

1921. A monograph of the fossil insects of the British coal measures. Part I. Paleontographical Soc., London, pp. 1-80, pls. 1-4.

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1896. Sur les moeurs de l'Evania desjardinsii, Blanch. Compt. Rend. Acad. Sci., Paris, vol. 123. pp. 610-613.

1912. Notes biologique recueillies à l'Ile de la Réunion. Bull. Sci. France et Belgique, vol. 46, pp. 29-92.

1913. Notes biologiques recueillies à l'Ile de la Réunion. Chap. II-IV. Bull. Sci. France et Belgique, vol. 47, pp. 377-412.

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1901. Les glandes défensives ou odorantes des blattes. Compt. Rend. Acad. Sci., Paris, vol. 132, pp. 1352-1354.

1908. Produit de sécrétion de la glande odorante des blattes. Bull. Soc. Zool. France, vol. 33, pp. 31-32.

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1946. Richerche sul batterio simbionte della Blattella germanica (L.). Bol. Soc. Ital. Biol. Sper., vol. 22, pp. 106-107. [Translated by M. Di Paolo.]

1948. Il ciclo del batterio simbionte di Blattella germanica (L.). Mycopathologia, vol. 4, pp. 85-102, 3 pls.

1948a. Osservazioni sul batterio simbionte della Blattella germanica (L.). Monitore Zool. Ital., Florence, vol. 56, Suppl., pp. 252-253. [Translated by J. J. Buckley.]

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1914. Kakkerlakken en wespen. De Levende Natur. Tijdschr. Natuur., vol. 18, pp. 385-395. [Translated by J. H. Vanderbie.]

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1953. List of sugarcane insects. 101 pp. Commonwealth Institute of Entomology, London.

BOZEMAN, W. B., JR.

1942. An experimental investigation into the life history of Blatticola blattae, a nematode found in Blattella germanica. Trans. Kansas Acad. Sci., vol. 45, pp. 304-310.

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1929. Beitrag zur Raumorientierung der Schabe Periplaneta americana. Zeitschr. Vergl. Physiol., vol. 10, pp. 497-526. [Translated by Ruth V. Judson.]

BREED, R. S.; MURRAY, E. G. D.; and HITCHENS, A. P., EDS.

1948. Bergey's manual of determinative bacteriology. 6th ed. 1,529 pp. Baltimore.

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1941. Notes on the biology of Stagmomantis carolina (Joh.) (Orthoptera, Mantidae). Bull. Brooklyn Ent. Soc., vol. 36, pp. 170-177.

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1917. Dolichurus n. sp. Proc. Hawaiian Ent. Soc., vol. 3, p. 268.

1920. Dolichurus stantoni. Proc. Hawaiian Ent. Soc., vol. 4, pp. 277, 438.

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1915. Entomological notes. Proc. Hawaiian Ent. Soc., vol. 3, pp. 55-56.

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1866. [Cockroach attacking centipede.] Proc. Boston Soc. Nat. Hist., vol. 11, p. 100.

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1908. Notes on the Orthoptera of Raleigh, North Carolina. Ent. News, vol. 19, pp. 16-21.

BRISTOWE, W. S.

1925. Notes on the habits of insects and spiders in Brazil. Trans. Ent. Soc. London, 1924. vol. 4, pp. 475-504.

1932. Insects and other invertebrates for human consumption in Siam. Trans. Ent. Soc. London, vol. 80, pp. 387-404.

1941. The comity of spiders. Ray Soc., London, vol. 2, pp. 229-560, 3 pls.

BRITISH MUSEUM (NATURAL HISTORY).

1951. The cockroach, its life-history and how to deal with it. 5th ed. Economic Series No. 12, 26 pp. [Revision of Laing (1946).]

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1889. Les blattes de l'époque houillère. Compt. Rend. Acad. Sci., Paris, vol. 108, pp. 252-254.

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1943. The grasshopper book. 127 pp. New York.

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1920. Marine hygiene and sanitation. 409 pp. London.

BROOKS, MARION A.

1954. Certain aspects of the histochemistry and metabolic significance of the intracellular bodies (bacteroids) of cockroaches (Blattariae). Ph. D. thesis, University of Minnesota. 63 pp., 10 pls.

1957. Investigation of possible antagonism between host and symbiote. Anat. Rec., vol. 128, pp. 527-528. (Abstract.)

BROOKS, MARION A., and RICHARDS, A. G.

1954. The necessity for intracellular bacteroids (symbiotes) in growth and reproduction of cockroaches (Blattariae). Anat. Rec., vol. 120, p. 742. (Abstract.)

1955. Intracellular symbiosis in cockroaches. I. Production of aposymbiotic cockroaches. Biol. Bull., vol. 109, pp. 22-39.

1955a. Intracellular symbiosis in cockroaches. II. Mitotic division of mycetocytes. Science, vol. 122, p. 242. 1956. Intracellular symbiosis in cockroaches. III. Re-infection of aposymbiotic cockroaches with symbiotes. Journ. Exp. Zool., vol. 132, pp. 447-466.

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1952. Observations on the life-history of the cockroach Ectobius panzeri Stephens (Orth., Blattidae). Ent. Month. Mag., vol. 88, pp. 209-212.

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1916. Contribución al estudio de los hormigas de la provincia de San Luis. Rev. Mus. de la Plata, vol. 23, pp. 291-357.

BRUES, C. T., and DUNN, R. C.

1945. The effect of penicillin and certain sulfa drugs on the intracellular bacteroids of the cockroach. Science, vol. 101, pp. 336-337.

BRUES, C. T.; MELANDER, A. L.; and CARPENTER, F. M.

1954. Classification of insects. Bull. Mus. Comp. Zool., Harvard College, vol. 108, pp. 1-917.

BRUIJNING, C. F. A.

1948. Studies on Malayan Blattidae. Zool. Meded., Leiden, vol. 29, pp. 1-174.

BRUMPT, E.

1931. Némathelminthes parasites des rats sauvages (Epimys norvegicus) de Caracas. I. Protospirura bonnei. Infections expérimentales et spontanées. Formes adultes et larvaires. Ann. Parasitol. Hum. et Comp., vol. 9, pp. 344-358.

1949. Précis de parasitologie. 6th ed., vol. 1, 1,042 pp.; vol. 2, 2,120 pp. Paris.

BRUMPT, E., and DESPORTES, C.

1938. Hôtes intermédiaires expérimentaux de deux espèces d'acanthocéphales (Prosthenorchis spirula et P. elegans) parasites des lémuriens et des singes. Ann. Parasitol. Hum. et Comp., vol. 16, pp. 301-304.

BRUMPT, E., and URBAIN, A.

1938. Une curieuse épizootie vermineuse à acanthocéphales, devenue endémique à la singerie du Muséum. Mesures prophylactiques efficaces prises pour en arrêter les méfaits. Compt. Rend. Acad. Sci., Paris, vol. 206, pp. 1927-1930.

1938a. Épizootie vermineuse par acanthocéphales (Prosthenorchis) ayant sévi a la singerie du Museum de Paris. Ann. Parasitol. Hum. et Comp., vol. 16, pp. 289-300.

BRUMPT, L.; DECHAMBRE, E.; and DESPORTES, C.

1939. Prophylaxie et traitement utilisés pour combattre les acanthoceplales parasites des makis de la menagerie du Jarden des Plantes. Bull. Acad. Vét. France, vol. 12, pp. 198-202.

BRUNER, S. C., and SCARAMUZZA, L. C.

1936. Reseña de los insectos del tobaco en Cuba. Estac. Exp. Agron. Santiago de Las Vegas Prov. Habana, Circ. No. 80, 50 pp.

BRYAN, E. H., JR.

1926. Insects of Hawaii, Johnston Island and Wake Island. Bernice P. Bishop Mus. Bull., vol. 31, pp. 1-94 (especially p. 89).

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1915. Natural history of Hawaii. 596 pp. Honolulu.

BRYGOO, E. 1946. Essai de bromatologie entomologique. Les insectes comestibles. Thèse de Doctorate, Bergerac. 73 pp. Imprimerie Générale du Sud-Ouest, Bergerac.

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1912. Studien an intracellularen Symbionten. I. Die intracellularen Symbionten der Hemipteren. Arch. f. Protistenk., vol. 26, pp. 1-116, 12 pls. [Pertinent sections translated by H. L. Middleton.]

1930. Tier und Pflanze in Symbiose. 900 pp. Berlin.

1952. Historische Probleme der Endosymbiose bei Insekten. Tijdschr. Ent., vol. 95, pp. 143-165. [Pertinent section translated by K. Gingold.]

1953. Endosymbiose der Tiere mit pflanzlichen Mikroorganismen. 771 pp. Basel/Stuttgart.

BUNTING, W.

1955. Orthoptera imported into Britain with bananas from Dominica (Leeward Isles). Ent. Month. Mag., vol. 91, p. 134.

1956. Preliminary notes on some Orthoptera imported with bananas from Dominica. Ent. Month. Mag., vol. 92, pp. 284-286.

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1943. The North American parasitic wasps of the genus Tetrastichus--a contribution to biological control of insect pests. Proc. U. S. Nat. Mus., vol. 93, pp. 505-608.

1952. The North American species of Syniomosphyrum (Hymenoptera, Chalcidoidea). Proc. Ent. Soc. Washington, vol. 54, pp. 258-264.

BURLINGAME, P. L., and CHANDLER, A. C.

1941. Host-parasite relations of Moniliformis dubius (Acanthocephala) in albino rats, and the environmental nature of resistance to single and superimposed infections with this parasite. Amer. Journ. Hyg., vol. 33, sect. D, pp. 1-21.

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1898. Orthoptera collected in South-Eastern Europe. Ent. Rec. and Journ. Var., vol. 10, pp. 1-5.

1899. Mimicry in Orthoptera. Ent. Rec. and Journ. Var., vol. 11, pp. 48-50.

1899a. Parasites of Orthoptera. Ent. Rec. and Journ. Var., vol. 11, pp. 186-187.

1899b. Local Orthoptera in 1899. Ent. Rec. and Journ. Var., vol. 11, p. 333.

1900. On the British Orthoptera in the Hope Museum, Oxford. Ent. Rec. and Journ. Var., vol. 12, pp. 97-99.

1908. Orthoptera in east Kent. Ent. Rec. and Journ. Var., vol. 20, pp. 275-278.

1911. Orthoptera in the Canary Islands. Ent. Rec. and Journ. Var., vol. 23, pp. 92-95, 175-178, 193-195.

1913. Collecting Orthoptera in the Caucasus and Transcaucasus. Ent. Rec. and Journ. Var., vol. 25, pp. 12-15, 37-41.

1923. A contribution to our knowledge of the Orthoptera of Macedonia. Trans. Ent. Soc. London, 1923, pp. 110-130.

1926. In Northeast Siberia. Ent. Rec. and Journ. Var., vol. 38, pp. 97-101.

1937. Periplaneta orientalis L. in the open. Ent. Rec. and Journ. Var., vol. 49, p. 115.

1939. The insect legion. 321 pp. London.

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1871. Untersuchungen über die beiden Nematoden der Periplaneta (Blatta) orientalis L. Zeitschr. Wiss. Zool., vol. 21, pp. 252-293, 2 pls.

1878. Beiträge zur Kenntniss der Flagellaten und einiger verwandten Organismen. Zeitschr. Wiss. Zool., vol. 30, pp. 205-281, 5 pls.

1881. Kleine Beiträge zur Kenntnis der Gregarinen. Zeitschr. Wiss. Zool., vol. 35, pp. 384-409, 2 pls.

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1914. British Blattidae. Ent. Rec. and Journ. Var., vol. 26, p. 185.

1955. The natural history of tse-tse flies. London School Hyg. Trop. Med., Mem. No. 10, 816 pp.

CALLAN, E. MCC.

1942. A note on Timulla (Timulla) eriphyla Mickel (Hym., Mutillidae), a parasite of Tachysphex blatticidus F. X. Williams (Hym., Larridae), from Trinidad, B. W. I. Proc. Roy. Ent. Soc. London, ser. A, vol. 17, p. 18.

1950. Observations on tropical wasps in Trinidad. Proc. 8th Internat. Congr. Ent., Stockholm, 1948, pp. 204-206.

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1917. A year of Costa Rican natural history. 577 pp. New York.

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1910. Zoological researches in Costa Rica. Old Penn Weekly Rev., Univ. Pennsylvania, vol. 9, pp. 165-170.

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1893. Sur quelques gordiens nouveaux ou peu connus. Bull. Soc. Zool. France, vol. 18, pp. 213-216.

1897. Monografia dei Gordii. Mem. Reale Accad. Sci., Torino, ser. 2, vol. 47, pp. 339-419.

CAMERON, EWEN.

1955. On the parasites and predators of the cockroach. I.--Tetrastichus hagenowii (Ratz.). Bull. Ent. Res., vol. 46, pp. 137-147.

1957. On the parasites and predators of the cockroach. II.--Evania apppendigaster (L.). Bull. Ent. Res., vol. 48, pp. 199-209.

CAMPOS R., F.

1926. Catalogo preliminar de los blatidos (cucarachas) del Ecuador. Rev. Col. Nac. Vicente Rocafuerte, vol. 8, pp. 41-57.

CANALIS, P.

1916. Note de M. le Professeur P. Canalis, relative à quelques expériences sur l'action insecticide du gaz Clayton. Bull. Mens. Off. Internat. Hyg., Paris, vol. 8, pp. 457-463.

CANTRALL, I. J.

1941. Notes on collecting and preserving Orthoptera. Part 2 of Compendium of entomological methods. 28 pp. Ward's Natural Science Establishment, Rochester, N. Y.

1943. The ecology of the Orthoptera and Dermaptera of the George Reserve, Michigan. Misc. Publ. Mus. Zool., Univ. Michigan, No. 54, 182 pp., 10 pls.

CAO, G.

1898. Sul passagio dei microrganismi attraverso l'intestino di alcuni insetti. L'Uff. San. Riv. Igiene Med. Prat., Naples, vol. II, pp. 337-348, 385-397.

1906. Nuove osservazioni sul passaggio dei microrganismi a traverso l'intestino di alcuni insetti. Ann. Igiene Sper., vol. 16, pp. 339-368.

CARNEIRO DE MENDONÇA, F., and CERQUEIRA, N. L.

1947. Insects and other arthropods captured by the Brazilian Sanitary Service on landplanes or seaplanes arriving in Brazil between January 1942 and December 1945. Bol. Ofic. Sanataria Panamer., vol. 26, pp. 22-30.

CARPENTER, G. D. H.

1920. A naturalist on Lake Victoria with an account of sleeping sickness and the tse-tse fly. 333 pp. London.

1921. Experiments on the relative edibility of insects, with special reference to their coloration. Trans. Ent. Soc. London, 1921, pp. 1-105.

1925. A naturalist in East Africa. 187 pp., 8 pls., 1 map. Oxford, England.

CASTELLANI, A., and CHALMERS, A. J.

1919. Manual of tropical medicine. 3d ed., 2,436 pp. New York.

CAUDELL, A. N.

1905. Notes on some Florida Orthoptera. Ent. News, vol. 16, pp. 216-219.

1914. Some bromeliadicolous Blattidae from Mexico and Central America. Insecutor Inscitiae Menstruus, vol. 2, pp. 76-80.

1923. Phorticolea boliviae, a new myrmecophilous cockroach from South America. Psyche, vol. 30, pp. 28-30.

1924. Some insects from the Chilibrillo bat caves of Panama. Insecutor Inscitiae Menstruus, vol. 12, pp. 133-135.

1925. Pycnoscelus surinamensis Linnaeus (Orthoptera); on its nymphs and the damage it does to rose bushes. Proc. Ent. Soc. Washington, vol. 27, pp. 154-157.

1931. Notes on Blattidae, adventive to the United States (Orthop.). Ent. News, vol. 42, p. 204.

CHABAUD, A. G., and LARIVIÈRE, M.

1955. Cycle évolutif d'un ascaride: Subulura jacchi (Marcel 1857) parasite de primates, chez la blatte Blabera fusca. Compt. Rend. Soc. Biol., Paris, vol. 149, pp. 1416-1419.

CHAKOUR, E.

1942. Notes diverses sur Leucophaea surinamensis (L.) et Nauphoeta cinerea (Oliv.). Bull. Soc. Ent. Fouad 1^{er} Ent., vol. 26, pp. 21-23.

CHAMBERLIN, J. C.

1949. Insects of agricultural and household importance in Alaska with suggestions for their control. U. S. Dept. Agr., Alaska Agr. Exp. Sta., Circ. No. 9, 59 pp.

CHAMISSO, A. VON.

1829. Ein Zweifel und zwei Algen. Verhandl. Ges. Naturf. Freunde, Berlin, vol. 1, pp. 173-180.

CHANDLER, A. C.

1921. Notes on the occurrence of Moniliformis sp. in rats in Texas. Journ. Parasitol., vol. 7, pp. 179-183.

1922. Species of Hymenolepis as human parasites. Journ. Amer. Med. Assoc., vol. 78, pp. 636-639.

1926. Some factors affecting the propagation of hookworm infections in the Asansol Mining Settlement with special reference to the part played by cockroaches in mines. Indian Med. Gaz., vol. 61, pp. 209-212.

1941. The specific status of Moniliformis (Acanthocephala) of Texas rats, and a review of the species of this genus in the Western Hemisphere. Journ. Parasitol., vol. 27, pp. 241-244.

1949. Introduction to parasitology. 756 pp. New York.

CHARLES, VERA K.

1941. A preliminary check list of the entomogenous fungi of North America. Insect Pest Survey Bull., vol. 21, Suppl. to No. 9, pp. 707-785.

CHATTON, E., and PÉRARD, C.

1913. Schizophytes du caecum du cobaye. I. Oscillospira Guilliermondi n. g., n. sp. Compt. Rend. Soc. Biol., Paris, vol. 74, pp. 1159-1162.

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1927. A contribution towards the insect fauna of French Oceania.