The significance of many observed associations between cockroaches and the higher plants is still obscure. Undoubtedly many associations are ecological, but lack of adequate supporting evidence makes this conclusion somewhat tentative. The ecological aspects are covered in 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 Alsophilasp.Associate.—Pycnoscelus surinamensis, Louisiana (Anonymous, 1893): Feeding on heart of tree fern. Family POLYPODIACEAE Asplenium nidusLinnaeusAssociate.—Comptolampra liturata, Malaya (Karny, 1924): Often found between dry foliage of the beakers of this fern. Division EMBRYOPHYTA SIPHONOGAMA Family PINACEAE Pinus australisMichauxAssociates.—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 caribaeaMoreletAssociates.—Eurycotis floridana, Latiblattella rehni, and Parcoblatta fulvescens, Florida (Hebard, 1917): Many records under signs on the tree trunks. Pinus clausaVaseyAssociate.—Latiblattella rehni, Florida (Hebard, 1917): Under sign on tree. Pinus echinataMill.Associates.—Parcoblatta divisa, Virginia (Rehn and Hebard, 1916): Under signs on shortleaf pine. Parcoblatta zebra, Mississippi (Hebard, 1917): Under sign. Pinus sylvestrisLinnaeusAssociate.—Ectobius pallidus, England (Milton, 1899; Burr, 1899b): On Scotch fir. Pinusspp.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 Cryptomeriasp.Associate.—Diploptera punctata, Hawaii (Pemberton and Williams, 1938; Zimmerman, 1948). Family CUPRESSACEAE Cupressus macrocarpaHartwegAssociate.—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). Juniperussp.Associate.—Phyllodromica tartara nigrescens, Southern Uzbekistan (Bei-Bienko, 1950): Under bark. Family PANDANACEAE Freycinetiasp.Associate.—Graptoblatta notulata and Kuchinga remota, Tahiti (Hebard, 1933). Pandanussp.Associate.—Hololeptoblatta sp., Seychelles (Scott, 1910, 1912). Family GRAMINEAE Aristida pennataTrin.Associate.—Phyllodromica pygmaea, U.S.S.R. (Bei-Bienko, 1950): Found in the dense turf. BambooAssociate.—Comptolampra liturata, Malaya (Karny, 1925). Chloris gayanaKunthAssociate.—Blattella vaga, Texas (Riherd, 1953): This field cockroach was rather abundant in clumps of Rhodes grass. Panicum purpurascensRaddiSynonymy.—Panicum barbinode [Hitchcock, 1936]. Associate.—Epilampra abdomen-nigrum, Puerto Rico (SeÍn, 1923; Wolcott, 1936): Abundant in "malojillo" meadow. Saccharum officinarumLinnaeusAssociates.—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 oatsAssociate.—Ischnoptera deropeltiformis, Missouri (Rau, 1937). Zea maysLinnaeusAssociates.—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 Cyperussp.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 nuciferaLinnaeusAssociates.—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 dactyliferaLinnaeusAssociate.—Blattella germanica, California (Herms, 1926): On date palms. Pritchardiasp.Associate.—Periplaneta australasiae, Nihoa Island (Bryan, 1926). Hawaii (Zimmerman, 1948). Roystonea regiaO. F. CookAssociate.—Cariblatta punctulata, San Domingo (Hebard, 1916a). Sabal palmettoLodd.Associate.—Eurycotis floridana, Florida (Scudder, 1879). Periplaneta australasiae, Florida (Hebard, 1917). Undetermined palmsAssociates.—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 Arumsp.Associate.—Latiblattella vitrea, Mexico (Hebard, 1921b): In flower shaft. Caladiumsp.Associate.—Plectoptera dorsalis, Puerto Rico (Rehn and Hebard, 1927). Family BROMELIACEAE Aechmaea porteoidesBrittonAssociate.—Dryadoblatta scotti, Trinidad (Princis and Kevan, 1955). Ananas comosusMerr.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 fulgensGriseb.Associates.—Cockroaches, Costa Rica (Calvert and Calvert, 1917). Glomeropitcairnia erectifloraMezAssociate.—Dryadoblatta scotti, Trinidad (Princis and Kevan, 1955). Grevisiasp.Associate.—Notolampra antillarum, Trinidad (Princis and Kevan, 1955): One male only. Tillandsia fasciculataSwartzAssociate.—Eurycotis floridana, Florida (Rehn and Hebard, 1914; Hebard, 1917). Tillandsia usneoidesLinnaeusAssociates.—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 uttriculataLinnaeusAssociate.—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). Tillandsiasp.Associates.—Aglaopteryx gemma, Texas (Hebard, 1917). Dryadoblatta scotti, Trinidad (Scott, 1912): Found in the leaf bases. Undetermined bromeliadsAssociates.—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 elataEngelmanAssociate.—Latiblattella lucifrons, Arizona (Ball et al., 1942). Easter liliesAssociate.—Pycnoscelus surinamensis, Connecticut (Zappe, 1918). Family MUSACEAE BananasCockroaches 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 Renealmiasp.Associate.—Cariblatta orestera, Jamaica (Rehn and Hebard, 1927): The male was taken in a head of wild ginger. Family CANNACEAE Cannasp.Associate.—Periplaneta americana, Hawaii (Zimmerman, 1948). Family ORCHIDACEAE Cattleyasp.Associates.—Periplaneta americana, U.S.A. (Rau, 1940a). Periplaneta australasiae, England (Lucas, 1918). Vandasp.Associates.—Periplaneta americana, U.S.A. (Rau, 1940a). Periplaneta australasiae, England (Lucas, 1918). Undetermined orchidsAssociates.—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 Casuarinasp.Associate.—Diploptera punctata, Hawaii (Zimmerman, 1948). Family SALICACEAE Populus euphraticaOliv.Synonymy.—Populus diversifolia Schrenk. [Howard, personal communication, 1959]. Associate.—Ectobius semenovi, Kazakhstan (Bei-Bienko, 1950). Populussp.Associate.—Ectobius lapponicus, U.S.S.R. (Stark in Bei-Bienko, 1950): On aspen. Salixsp.Associate.—Ectobius semenovi, Kazakhstan (Bei-Bienko, 1950): On willow. Family MYRICACEAE Myrica ceriferaLinnaeusAssociate.—Chorisoneura texensis, Florida (Rehn and Hebard, 1916): On bayberry. Florida (Blatchley, 1920): Beaten from foliage. Family FAGACEAE Quercus albaLinnaeusAssociate.—Parcoblatta pensylvanica, Virginia (Rehn and Hebard, 1916): Under signs on white oaks. Quercus rubraLinnaeusAssociates.—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 virginianaMill.Associate.—Eurycotis floridana, Georgia (Rehn and Hebard, 1916): Under dead bark on live-oak tree. Georgia (Hebard, 1917): In cavity in tree. Quercusspp.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 Cecropiasp.Associate.—Cariblatta hylaea, Honduras (Rehn, 1945a). Family CHENOPODIACEAE Beta maritimaLinnaeusAssociate.—Ectobius panzeri, England (Lucas, 1920a). Beta vulgaris var. ciclaLinnaeusAssociate.—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 flavaLinnaeusNatural 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 purpureaLinnaeusNatural prey.—Cariblatta lutea lutea, North Carolina (Wray and Brimley, 1943). Sarracenia minorWalterSynonymy.—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 ampulariaJackNatural prey.—Cockroaches, Singapore (Dover, 1928). Nepenthes gracilisKorth.Natural prey.—Cockroaches, Singapore (Dover, 1928). Nepenthessp.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 Weinmanniasp.Associates.—Aneurina viridis, Marquesas Islands, Nukuhiva and Fatuhiva (Hebard, 1933a). Maretina uahuka, Marquesas Islands, Uahuka (Hebard, 1933a). Family HAMAMELIDACEAE Liquidambar styracifluaLinnaeusAssociate.—Parcoblatta divisa, Georgia (Rehn and Hebard, 1916): Under sign on sweet gum. Parcoblatta zebra, Louisiana (Hebard, 1917): In decay cavity. Family ROSACEAE Crataegussp.?Associates.—Plectoptera dorsalis, Plectoptera infulata, Plectoptera rhabdota, Puerto Rico (Wolcott, 1950): In the dry flower clusters of "espino rubial." Rosasp.Associate.—Pycnoscelus surinamensis, Connecticut (Zappe, 1918); Rhode Island and Pennsylvania (Caudell, 1925); Pennsylvania (Doucette and Smith, 1926): Feeding on canes in greenhouses. Rubusspp.?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 farnesianaWilld.Associate.—Diploptera punctata, Hawaii (Bridwell and Swezey, 1915; Zimmerman, 1948): Feeding on pods. Acaciasp.Associates.—Ellipsidion australe, Australia, New South Wales (Hebard, 1943). Methana curvigera, Australia, Queensland (Pope, 1953a). Ceratonia siliquaLinnaeusAssociate.—Diploptera punctata, Hawaii (Pemberton and Williams, 1938): Damaging algarroba. Erythrina glaucaWilld.Associates.—Aglaopteryx absimilis, Puerto Rico (Wolcott, 1950): In abandoned cocoon. Aglaopteryx facies, Puerto Rico (Wolcott, 1936): In empty cocoons. Inga laurinaWilld.Associate.—Aglaopteryx facies, Puerto Rico (Wolcott, 1936): On trunk. Inga veraWilld.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. MesquiteAssociate.—Nyctibora stygia, Haiti (Rehn and Hebard, 1927). Samanea samanMerr.Associate.—Aglaopteryx absimilis, Puerto Rico (Wolcott, 1950). Tamarindus indicaLinnaeusAssociate.—Hemiblabera brunneri, Puerto Rico (Rehn and Hebard, 1927). Family GERANIACEAE GeraniumsAssociate.—Diploptera punctata, Hawaii (Zimmerman, 1948). Family ZYGOPHYLLACEAE Tribulussp.Associates.—Periplaneta americana and Pycnoscelus surinamensis, Johnston Island (Bryan, 1926). Zimmerman (1948) lists Tribulus as a host plant for these cockroaches. Family RUTACEAE Citrus aurantifoliaSwingleAssociates.—Plectoptera dominicae and Plectoptera perscita, Dominica (Rehn and Hebard, 1927): Beaten from moss-covered lime trees. Citrus maximaMerr.Associates.—Plectoptera dorsalis, Plectoptera infulata, Plectoptera rhabdota, Puerto Rico (Wolcott, 1950). Plectoptera rhabdota, Puerto Rico (Rehn and Hebard, 1927). Citrus sinensisOsbeckAssociate.—Diploptera punctata, Hawaii (Bridwell and Swezey, 1915; Zimmerman, 1948): Feeding on oranges on tree. Citrussp.Associates.—Diploptera punctata, Hawaii (Zimmerman, 1948). Riatia [= Lissoblatta] fulgida, Panama, Rio Trinidad (Hebard, 1920). Plectoptera porcellana, Puerto Rico (Sein, 1923). Zanthoxylum caribaeumLam.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 Poinsettiasp.Associate.—Pycnoscelus surinamensis, Connecticut (Zappe, 1918): Ate bark of greenhouse plants. Family ANACARDIACEAE Mangifera indicaLinnaeusAssociate.—Diploptera punctata, Hawaii (Bridwell and Swezey, 1915; Zimmerman, 1948): Feeding on mangoes on the tree. Spondias mombinLinnaeusAssociates.—Plectoptera dorsalis, Plectoptera infulata, Plectoptera rhabdota, Puerto Rico (Wolcott, 1950): Living on leaves of "jobo." Spondias purpureaLinnaeusAssociate.—Eurycotis biolleyi, Costa Rica (Rehn, 1918): In the crown of dry jocoto. Family AQUIFOLIACEAE Ilex cassineLinnaeusAssociate.—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 Gossypiumspp.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-sinensisLinnaeusAssociate.—Riatia orientis, Trinidad (Princis and Kevan, 1955). Hibiscussp.Associates.—Amazonina emarginata, Cariblatta antiguensis, Eurycotis kevani, and Rhytidometopum dissimile, Trinidad (Princis and Kevan, 1955). Sidasp.Associate.—Periplaneta australasiae, Nihoa Island (Bryan, 1926). Hawaii (Zimmerman, 1948). Family STERCULIACEAE Theobroma cacaoLinnaeusAssociate.—Ceratinoptera picta, Trinidad (Princis and Kevan, 1955). Family BIXACEAE Bixasp.Associate.—Notolampra antillarum, Trinidad (Princis and Kevan, 1955): Nymphs in dry fruits on "annato" tree. Family FLACOURTIACEAE Xylosma suaveolensForst.Associate.—Graptoblatta notulata, Marquesas Islands, Uahuka (Hebard, 1933a). Family PASSIFLORACEAE Passiflorasp.Associate.—Aristiger [= Plumiger] histrio, Malaya (Karny, 1924). Family CARICACEAE Carica papayaLinnaeusAssociate.—Diploptera punctata, Hawaii (Bridwell and Swezey, 1915; Zimmerman, 1948): Feeding on papaya fruit on tree. Family RHIZOPHORACEAE Rhizophora mangleLinnaeusAssociate.—Aglaopteryx gemma, Florida (Hebard, 1917). Family COMBRETACEAE Conocarpus erectusLinnaeusAssociate.—Plectoptera poeyi, Florida (Rehn and Hebard, 1914): Running on leaves. Family MYRTACEAE Eucalyptussp.Associate.—Ellipsidion australe, Australia, New South Wales (Hebard, 1943). Eugenia aromaticaBaill.Synonymy.—Syzygium aromaticum [Bailey, 1925]. Associate.—Plectoptera dorsalis, Puerto Rico (Wolcott, 1936): On flowers of "pomarrosa." Metrosideros collinaGrayAssociates.—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 guajavaLinnaeusAssociate.—Plectoptera rhabdota, Puerto Rico (Rehn and Hebard, 1927). Family ONAGRACEAE Jussiaea natansHumb. 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 vulgarisSalisb.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 meridionaleSw.Associates.—Chorisoneura formosella, Neoblattella dryas, Neoblattella proserpina, Jamaica (Rehn and Hebard, 1927). Family SAPOTACEAE Sideroxylon foetidissimumJacq.Associate.—Pelmatosilpha coriacea, Puerto Rico (Wolcott, 1941): Under bark. Family APOCYNACEAE Vinca minorLinnaeusAssociate.—Ectobius pallidus, Massachusetts (Willis, unpublished observation, 1958). Family CONVOLVULACEAE Ipomoea tiliaseaChoisyAssociate.—Plectoptera dorsalis, Puerto Rico (Rehn and Hebard, 1927). Family BORAGINACEAE Cordia dentataPoiretSynonymy.—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 villosumJacq.Associate.—Chorisoneura texensis, Florida (Rehn and Hebard, 1912). Family SOLANACEAE Nicotianasp.Associate.—Pycnoscelus surinamensis, Sumatra (Roeser, 1940). Solanum tuberosumLinnaeusAssociate.—Pycnoscelus surinamensis, Haiti (Hoffman, 1927): Feeding on tubers in field. Family GESNERIACEAE Cyrtandrasp.Associate.—Aneurina viridis, Marquesas Islands, Nukuhiva (Hebard, 1933a). Family RUBIACEAE Canthium barbatum(Forst.) Seem.Associate.—Graptoblatta notulata, Marquesas Islands, Uahuka (Hebard, 1933a). Cinchona pubescensVahl.Associate.—Periplaneta americana, Puerto Rico (Plank and Winters, 1949): In greenhouse. Coffeasp.Associate.—Plectoptera porcellana, Puerto Rico (SeÍn, 1923). Family COMPOSITAE GoldenrodAssociate.—Eurycotis floridana, Florida (Hebard, 1917): "Climbing about on top of goldenrod at night." Helianthussp.Associate.—Pseudomops septentrionalis, Texas (Hebard, 1917). Scorzonera acanthocladaFranch.Associate.—Phyllodromica tartara nigrescens, Southern Uzbekistan (Bei-Bienko, 1950): On the flowers. 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 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 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 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). |