< o CO "'NOlin ,SNI_NVIN0SHiltMs'^S3ldVyan ~ LlBRARIEs'^SMITHSONlAN^INSTITUTION — \ 2 ^ \ ^ lES ^ ^ 2-1 SMITHSONIAN 00 to /^w\ ^ /^^\ — Z INSTITUTION ^^ - /^rf^\ - 2 _J NVINOSHimS NQIiniliSNI -I S3iyvyan LI BR, o > ^ to to rn m Xi2ilS!i>^ ' _ to ± \. £ NviNOSHims SNi S3ibvyan libraries CO Smithsonian institution Noiir CO CO Z Z .V. 2 (A» £/) / CO z (o lES SMITHSONIAN ^ SNI r; 5 NViNOSHims V » (/) INSTITUTION </5 ^ :z ES^ SMITHS0NIAN~INSTITUTI0N Z Z I M VM LI BR z CO RAR Es'^SMITHSONIAN INSTITUTION NOIIO I 2 INSTITUTION %il ^^-1 Z a IT^LI B _J SMITHSONIAN z ^ to NOIiniliSNI~NVINOSHllWS S3ldVyan~l-'BRi CO •» SNrNVIN0SHims'"S3 ES S3l«Vyan z S3iyvyan r- libraries Z Smithsonian r^ institution z Noiin lip ^ *^ (f) NOIiniliSNI — NVINOSHillMS Z _J NOIiniliSNI NVINOSHimS — S3ldVyan LI m BR, SMITHSONIAN ^ Z INSTITUTION r Z •''- t/) NiviNOSHiiiMS NOUniliSNI ^ S3iavyan libraries 2 -1 m ^^i:_wi^ c/> Z Smithsonian — — xioiiisvtx CO Z * I f: Smithsonian Z z O NoiiniiiSNi LIBRARIES 11 ~ c/^ NoiiniiisN institution </) "^ z S3iyvaan LIBRARIES NOIiniliSNI_NVINOSHimS S3iavaan r- libraries > -I S3 dV^ 9 ^. 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A. ADVERTISEMENT The Smithsonian Miscellaneous Collections series contains, since the suspension in 1916 of the Smithsonian Contributions to Knowledge, all the publications issued directly by the Institution except the nual Report and occasional publications of a special nature. name of the series implies, its scope is As Anthe not limited, and the volumes Papers in and astrophysics have thus far issued relate to nearly every branch of science. the fields of biology, geology, anthropology, predominated. Leonard Carmichael, Secretary, Smithsonian Institution. (iii) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 141 (WHOLE VOLUME) THE BIOTIC ASSOCIATIONS OF COCKROACHES (With 37 Plates) By LOUIS M. ROTH AND EDWIN R. WILLIS Pioneering Research Division, United States Army Quartermaster Research and Engineering- Center Natick, Mass. (Publication 4422) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION DECEMBER 2, 1960 VOL. 141, PL. SMITHSONIAN MISCELLANEOUS COLLECTIONS Blahcnis craniifcr, c. X 2. i. 1 (Photograph by Jack Sahnon. Philadelphia Quartermaster Depot.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 141 (WHOLE VOLUME) THE B OTIC ASSOCIATIONS OF COCKROACHES I (With 37 Plates) By LOUIS M. ROTH AND EDWIN R. WILLIS Pioneering Research Division, United States Army Quartermaster Research and Engineering Center Natick, Mass. ^ (Publication 4422) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION DECEMBER 2, 1960 THE LORD BALTIMORE PRESS, BALTIMORE, MD., U. S. A. INC. FOREWORD People having only casual interest in insects usually express amaze- ment when they learn how much is known about this most numerous group of animals. However, while entomologists have good reason to take pride in the accomplishments of their contemporaries and predecessors, they are more likely to be appalled by how much remains to be learned. We are indeed ignorant of even the identity of fully half and probably much more than half the total number of insect species. Of those that have been described, we have reasonably complete information about the behavior and basic environmental relationships for only a comparative few. The great majority of the remainder are known only from specimens found in museum collections. Such information as we have about these species usually amounts to no more than date and locality of collection. This is true of the cockroaches, which now include approximately 3,500 described species. Conservative estimates based on partially museum and the percent of new species found in from tropical and subtropical coun4,000 species remain unnamed. Although studied collections recent acquisitions, particularly tries, indicate that at least is well known in general terms to nearly all entomologists, an almost complete void of information about all except the the group there is few domestic species and, to a progressively diminishing degree, some 400 others. Many details about the lives of even those that share man's habitations are not fully understood. This then is a rough measure of how little is With known about cockroaches. the exception of mosquitoes and a few other comparatively small groups of insects on which work has been concentrated, it is any other comparable segment of the world's insect fauna is better known. Already an estimated 800,000 kinds of insects have been described, and since this figure is generally regarded as less than half the actual total, think what this means in terms of knowledge yet to be assembled. No wonder entomology is a growing science with a promising future, but the magnitude of the task also presents a doubtful serious if obstacle to progress. Progress can continue only if the from the diversified labors of hundreds brought together and summarized in thorough and scattered literature resulting of contributors is well-organized compilations that can serve as a solid basis for future research. iii SMITHSONIAN MISCELLANEOUS COLLECTIONS iv The present work is such a compilation, for been gleaned from approximately i it VOL. I4I assembles what has ,700 sources, including correspond- ence with a large number of other workers. Original observations during some eight years of concentrated effort in U. S. Army Quartermaster research laboratories are a valuable supplement to what others have done, and with this background of experience the authors Seldom has are especially well qualified to appraise previous work. a compilation been done so thoroughly or a single large group of insects been the subject of such uninterrupted The effort. contents gives the categories of subject matter treated and the introduction discusses the value of this assembled information and offers suggestions for future study. No garded only as disagreeable pests; many longer are cockroaches respecies appear to be im- portant, actually or potentially, as carriers of disease. this Recognition of importance has grown considerably, even in the period since World War II. Consequently, anything that increases our knowledge of the basic bionomics of cockroaches will be consulted widely for factual information and for clues to new approaches. In spite of this extensive compilation, the limitations of present The information about cockroach bionomics must be kept in mind. cited observations of many clusions disagreed. But it is writers were fragmentary, or their con- fundamental to scientific inquiry that should know and attempt that what Drs. Roth and Willis have done. Fortunately, view is is readily available. maximum to evaluate the results of previous study, Sometimes, a piece of work we and their re- fails to be of value because the results are not generally accessible to later students. For this reason I am especially glad that the Smith- sonian Institution, by disseminating the results of the authors' labors, has this opportunity to exercise one of its traditional functions —that of diffusing knowledge. Throughout the period of research by Drs. Roth and Willis at I was in frequent correspondence with them, and I admire their many accomplishments. Our warmest commendations should go not only to them personally but also to those in administration who encouraged their fundamental research and who aided in the Natick, financial support of this publication. Ashley B. Gurney Entomology Research Division United States Department of Agriculture . CONTENTS Page Foreword I. II. I'l Introduction i Historical 2 Methods 4 Future work 5 Illustrations 7 Species of cockroaches III. Ecological 7 I4 relationships i6 Cave habitats I7 Cavernicolous cockroaches Cockroaches from burrows Desert habitats Desert cockroaches Aquatic habitats Amphibious cockroaches Outdoor habitats Cockroaches from outdoor habitats 23 25 27 30 31 33 35 70 Structural habitats Land-based structures Cockroaches associated with land-based structures Ti 74 82 Ships 85 Cockroaches associated with ships 87 88 Aircraft Cockroaches associated with aircraft 91 IV. Classification of the associations 96 V. Mutualism 9^ Bacteroids Cockroaches in which bacteroids have been found 99 100 Bacteria loi Protozoa VI. Viruses associated with cockroaches VII. Bacteria associated with cockroaches VIII. Fungi and yeasts Fungi associated with cockroaches IX. Higher plants associated with cockroaches by cockroaches X. Protozoa associated with cockroaches XL Helminths associated with cockroaches Helminths for which cockroaches serve as primary hosts Helminths for which cockroaches serve as intermediate hosts Helminths whose eggs have been carried by cockroaches XII. Arthropoda associated with cockroaches Arachnida Chilopoda Damage 103 104 127 129 I39 162 to plants 166 IQO 192 203 208 . 210 211 223 V SMITHSONIAN MISCELLANEOUS COLLECTIONS VI VOL. I4I Page 224 Insecta Hymenoptera 234 Host 254 255 266 268 by egg parasites Cockroach-hunting wasps Ants predaceous on cockroaches XIII. Vertebrata associated with cockroaches XIV. 234 Predators and parasites of cockroach eggs selection Pisces 268 Amphibia Reptih'a 269 272 Aves 276 Mammalia 283 Checklist of cockroaches and symbiotic associates XV. Cockroaches as commensals Hosts of commensal cockroaches Checklist of commensal cockroaches with their hosts Obscure associations XVI. Cockroaches as predators Interspecies predation Intraspecies predation 290 308 309 313 314 317 317 320 XVII. Associations among cockroaches 322 Familial associations 323 328 Gregariousness Intraspecies fighting Interspecies compatibility 334 335 Interspecies antagonism 339 XVIII. Defense of cockroaches against predators XIX. The biological control of cockroaches Invertebrates Vertebrates 341 346 347 351 Acknowledgments 352 References 354 439 Index THE BIOTIC ASSOCIATIONS OF COCKROACHES ' By Louis M. Roth and Edwin R. Willis 2 Pioneering Research Division, United States Army Quartermaster Research and Engineering Center Natick, Mass. (With 37 Plates) With most the net of us collectors the life history of an insect begins in and ends in the bottle. Hanitsch I. (1928) INTRODUCTION together much of the literature Hnking cockof certain organisms that cause disease transmission roaches with the In that paper (1957a) we concluded vertebrates. in man and other vectors of pathogenic agents, should potential being that cockroaches, Recently we brought Obviously the associanot be regarded simply as minor annoyances. diseases are of more vertebrate of agents with tions of cockroaches pathogens of lower with relations their than immediate importance cockroaches are of hand, other the On nonpathogens. animals or with their control is and importance, medical well as general economic as are unaware of their medical significance. That satisfactory may be the control of domiciliary cockroaches is far from journals in which pest-control and entomological inferred from current others found to replace to advocated new insecticides are continually of cockroaches control the approaches to inadequate. Possibly new sought by many who be Whether these lie in the direction of increased use of these insects parasites and predators for the biological control of any insect, about know we more the remains to be seen. In any event, are needed. achieving satisespecially its ecology, the greater the likelihood of 1 Part of the cost of publication of this monograph was borne by the Office of of Naval Research, Department of the Navy (through the American Institute CenEngineering and Research Quartermaster the Biological Sciences), and by ter, Department of the Army. United Fruit Present address of both authors. Central Research Laboratories, Co., Upland Road, Norwood, Mass. 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 141 (WHOLE VOL.) SMITHSONIAN MISCELLANEOUS COLLECTIONS 2 VOL. I4I factory control. In order to advance knowledge in any field of science, new research should proceed from the results of prior investigations when these exist. We hope that the observations and experiments cited herein may suggest areas for future research and exploitation. To the best of our knowledge no previous publication has brought together the vast literature on the parasites, predators, commensals, and other symbiotic associates of the Blattaria. For this reason, we have tried to assemble observations on all such known associations. Undoubtedly we have overlooked some records, as, for example, those buried in papers dealing with other phases of cockroach biology. hope that such inadvertent omissions Whatever usefulness of this compilation. We will not seriously impair the its should be a unified source of information for defects, this all who review are interested in the biotic associates of cockroaches. In addition to previously published information, this monograph and observations on the associations of first time. Although some of the observations were made by us, others were made by colleagues who have graciously made their knowledge available to us in private also contains original records cockroaches that are reported here for the communications. HISTORICAL Chopard (1938) much in his book La Biologie des Orthoptcres reviewed of the literature on cockroaches, but of the tions that exist many biotic associa- he discussed only the commensal cockroaches, gregari- ousness, and familial associations. Asano (1937), who reviewed the natural enemies of cockroaches, mentioned about 10 groups of animals Thompson (1951) in his Parasite Host was based mainly on papers abstracted or noted in that attack cockroaches. Catalogue, which Review of Applied Entomology, listed only 19 insect parasites of cockroaches. Eighteen of these were Hymenoptera which attack onl}'^ the cockroach eggs; the single Wiedemann, supposedly listed {Sarcophaga lamhens on Pycnoscelus siirinamensis) is not dipteron parasitic a parasite in this case, but deposits Cameron (1955) its eggs on the dead insects (see and predators of the cockroach 24 species of hymenopterous egg parasites, 7 species of Ampule X which hunt nymphs and adults, 17 Protozoa, 13 nematodes, p. 229). 5 bacteria, 2 mites, listed as parasites and a few other miscellaneous predators. In his protozoan parasites of the Orthoptera of the world, Semans (1943) listed about 26 species from cockroaches. Linclassified list of the stow (1878, 1889) recorded 14 species of helminths from cockroaches. (1928) listed 33 names of roundworms which Van Zwaluwenburg BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 3 are commensals or secondary parasites of cockroaches, but some of these names are synonyms. La Rivers (1949) extended this Hst with Chitwood (1932) recognized 24 species of parasites (probably commensals) of primary nematodes which are many instances of biological relagave blattids. Steinhaus (1946) 13 additional species. and tionships between cockroaches and yeasts, but the and the information is bacteria, fungi, cockroaches were not discussed as an entity scattered throughout the book. In surveying the literature on this subject more extensive list we have collected a far of animals and plants associated with cockroaches than one might have expected from an examination of any one of the previous papers on this subject. In our review of the medically important organisms associated with the Blattaria, we pointed out that in addition to found many experimental to harbor, naturally, associations cockroaches have been 4 strains of poliomyelitis virus, about 40 species of pathogenic bacteria, the eggs of 7 species of pathogenic helminths, and to serve as intermediate hosts of 12 other species of helminths pathogenic for vertebrates; cockroaches have also been found to to carry, man and on occasion, 3 species of Protozoa that are pathogenic 2 species of fungi which are sometimes found associated with pathological conditions. In addition to the above organisms of medical importance, we have compiled records of other organisms, nonpathogenic to vertebrates, which are naturally associated in some way with cockroaches. None of the following numbers can be considered absolute because some names may be synonyms. However, we believe that these figures are very close to the actual numbers of species that have been isolated because we have attempted to refer all obvious synonyms to the currently accepted name for each organism. On this basis there are about 45 species of bacteria, 40 fungi, 6 yeasts, 90 Protozoa, and 45 helminths that have been found associated naturally with cockroaches. Of the arthropods there are about 2 species of scorpions, 4 spiders, 15 mites, 4 centipedes, and 90 insects. Of vertebrates there are 4 species of fish, 16 amphibians, 12 reptiles, 20 birds, and 27 mammals. Besides these there are many records of experimental associations that have been contrived in the laboratory. Some idea of the increase in our knowledge of the biotic associations of cockroaches, during the last 70 years, may be gathered from a comparison of the above figures with those of Miall and Denny (1886) who presented "... a long list of parasites which infest the Cockroach." This list included 2 bacteria, 6 Protozoa (some of the names are synonyms), 7 nematodes (some of these names are also SMITHSONIAN MISCELLANEOUS COLLECTIONS 4 synonyms), i mite, i wasp, and as other foes of the cockroach : i VOL. I4I beetle. In addition, they mentioned monkeys, hedgehogs, polecats, cats, chameleons, and frogs. rats, birds, METHODS We have listed the organisms known to be associated with cock- roaches systematically by phylum, class, order, and family. Within each family the organisms are listed alphabetically by genus and Under each organism species. the associated cockroaches are listed as natural or experimental hosts, vectors, or prey. Identified cock- roaches are listed by the currently accepted name. Unidentified cock- roaches are indicated by the word "Cockroaches." cockroach is The name of each followed by the country in which the observation was made, the authority for the record, and with a few exceptions ^ pertinent biological information, where this is known. Question marks following the names of organisms or countries indicate tentative or questionable identifications. Records of predators capturing and feeding on cockroaches in zoos and on shipboard we consider natural, even though it is very likely that these particular predators would not normally have access to this prey in nature. Experimental prey are cockroaches that were fed to predators in the laboratory. Although these predators may have little, if any, access to these cockroaches in nature, we have included such records to indicate the relative acceptability of cockroaches as food by a wide variety of animals. Records of presumed or known cockroach associates that give no information about an associated cockroach are not included in this review, even though certain of these Evania, Podium) probably prey upon (e.g., species or parasitize of Ampulex, cockroaches exclusively. The validity of a host-parasite or predator-prey record is dependent upon the accuracy and knoMdedge of the observer. In assembling these records we have had to accept, in most instances, the identifications of species made by the original authors. However, as a result of our studies on the biology of various species of cockroaches, including 3 Names of organisms preceded by an asterisk ('^) are known or suspected pathogens of vertebrates. These records were presented with annotations in our 1957a paper on the medical and veterinary importance of cockroaches. For that reason the annotations have not been repeated herein, although the records have been included to make the listing of the biotic associates of cockroaches substantially complete. BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 5 hymenopterous parasites, we have questioned Other dubious records which have been perpetuated from one pubHcation to the next, but which apparently were not based on fact, have also been questioned or have some work on their certain records in the literature. been clarified with the aid of specialists in particular groups. Because the records cover a period of many years, the names of of the organisms as well as the names of some of the cockroaches have been changed. Although it would have been compara- many names as they appeared in the original refwould have resulted in misleading redundancy with the same organism being catalogued under several synonyms. We have attempted to list each organism by its currently accepted name. However, no attempt was made to prepare complete taxonomic synonymies the only synonyms given are those that identify the organisms by the names used by the authors of the papers cited. The synonyms under which the cockroaches may have been cited originally are listed in section II. The synonyms of associated organisms are listed with each organism. Although authorities for the name changes of the cockroaches are given, these workers are not necessarily those who were initially responsible for the synonymies. Various sections have been checked by specialists in the particular groups. Although we have accepted name changes suggested by these reviewers, we assume full tively simple to list the erences, this ; responsibility for the names. FUTURE WORK After having examined thousands of references on cockroaches, are impressed by how little is known about the biology of most we species. As a conservative estimate there are 3,500 described species of Blattaria (J. W. H. Rehn, 1951). In our literature survey we found records of biotic associations for about 400 species. Unfortunately, many of these records contain only a sentence or two of biological information. Our detailed knowledge of cockroaches is based on studies of the few domiciliary pests that man attempts to eradicate. Comparable studies of the bionomics of the less-well-known species should add much valuable information to our knowledge of this ancient group. Our understanding of most predator-prey and parasite-host rela- tionships has barely progressed beyond the taxonomic stage. effect The total of predators and parasites in limiting natural populations of cockroaches remains to be determined. It is still not known how, for example, predatory or parasitic wasps select specific cockroaches from SMITHSONIAN MISCELLANEOUS COLLECTIONS 6 among (e.g., The Secretions produced by certain cockroaches other insects. all 2-hexenal by Eurycotis floridana) will ward and identities VOL. I4I biological activities of off certain predators. most cockroach secretions are unknown, but the use of protective chemicals against predators may be widespread among how cockroaches. If so, repellents in protecting the individual or the species effective are these It is ? not known whether cockroaches are protected by apparent mimetic resemblances to other arthropods. There is no experimental proof that insect parasites can successfully attack the eggs of cockroaches that incubate by the female. their eggs while they are being carried It is conceivable that biological control of cockroaches might be achieved in limited areas such as tive to (e.g., know what structures or sewers, but effects, if It would be informa- any, organisms such as bacteria. Protozoa gregarines), intestinal nematodes, or other helminths have on cockroaches. Possibly pathogenic microorganisms can be used for biological control of cockroaches little man-made has not been thoroughly explored. this possibility ; this approach seems to have been explored. Associations of colonial cockroaches Newly hatched nymphs (e.g., may merely be truly familial or they Cryptocercus spp.) result may from gregariousness. of species that carry their oothecae until the eggs hatch cluster near the mother. This may be a response to the mother as such, a search for shelter beneath the nearest object (thigmotaxis? or negative phototaxis?), or there may well be yet another explanation. Tepper in 1893 stated that the native cockroaches of Australia are almost wholly carnivorous ; little supporting evidence for The apparent supersedure of one species of domiciliary cockroach by another may result from antagonism between different species, or it may result this claim has been brought forward since that time. from more rapid breeding and more effective utilization of available food and space; but which? Several species of cockroaches are frequently found associated with certain plants (e.g., bromeliads and bananas) ; the ecological relations in these associations remain to be determined. and other Many of the obscure associations between cockroaches insects, spiders, birds, been adequately defined. The and burrowing animals have never factors influencing cannibalism have never been thoroughly investigated experimentally. These are only a few ideas for future work that have occurred to us during the preparation of this review. as other questions that research in areas where may it is We hope that these suggestions as well occur to readers will stimulate further obviously needed. ; BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 7 ILLUSTRATIONS Unless otherwise credited, the ilkistrations were prepared from photographs taken by the authors. Except where otherwise stated, all photographs were taken of unposed living specimens. SPECIES OF COCKROACHES 11. The cockroaches referred to in this paper are listed below. name currently accepted for each species genus and species irrespective of its is taxonomic used by certain authors whose work affinities. we have quoted brackets under the respective species; the The given alphabetically by synonymy is Synonyms are given in supported by the reference citation that follows each synonym. References to illus- trations of certain species (e.g., Blaberus craniifer) that appear in the paper follow the names of the Agis orientalis Chopard Aglaopteryx absimilis Gurney Apotrogia angolensis Kirby [Acantho- diaphana (Fabricius) [Ceratinoptcra diaphana Fabricius ; Rehn and Hebard (1927)] jacics (Walker) [Aglaopteryx devia Rehn; Princis (1929). A. diaphana (Fabricius) in records from Puerto Rico only; Rehn (1932b) ; Gurney gemma Hebard ords ( 1937)1 [In Florida rec- = Ceratinoptera diaphatui R. and H.; Hebard (1917)] vegeta Rehn similis gyna deplanata Princis (i957)] Aptera Chopard (Thunberg) fusca [Apt era cingnlata (Burmeister) ; Gurney (personal communication, 1957)] Aptcroblatla perplexa Shelford Archiblatia hoevenii VoUenhoven Archimandrita marmorata (Stoll) tesscllata Rehn Arenivaga apacha (Saussure) bolliana (Saussure) erratica (Rehn) floridensls Caudell grata Hebard ypsilon Princis Allacta describers. (Saussure) [Phyllo- roseni (Brancsik) [Heterogamodes dromia obtusata Brunner; Zim- roscni; merman "Polygamia" roseni is undoubtedly an erroneous citation of Polyphaga roseni, as there is no genus Polygamia (Gurney, per- Alluauddlina (1948)] cavernicola [Alluaudella (Shelford) cavernicola Shel- ford; Chopard (1932)] Amasonina cmarginata Princis Anaplecta ascma Hebard asteca Saussure declpiens Saussure and Zehntner fallax Saussure hemiscofia Hebard Burmeister mexicana Saussure lateralis Aneurina tahuata Hebard viridis Hebard Bei-Bienko (1950). sonal communication, 1957)]. tonkazva Hebard Aristiger histrio miger (Burmeister) [Plu- (Burm.) Bruijning Hemithyrsocera (1948). histrio Burm.; Hebard (1929)] Aspiduchits boriquen J. W. H. Rehn histrio ; = Rico records Puerto [In Aspiduchus deplanatus R. and H.; Rehn, J. W. H. (ipsia)] ] ;; SMITHSONIAN MISCELLANEOUS COLLECTIONS 8 hmnbertiana (Saussure) [Blatta Phyllodromia humbertiana; humbertiana; Hebard (1929)] W. H. Rehn cavernicola J. deplanatus (Saussure) Attaphila aptera Bolivar ner) jamaicana Rehn and Hebard (Walker) (Walker) platysoma torresiana Bantua [DeroKrauss (Krauss) stigmosa Princis (i957)] atropos boliviensis Princis Saussure (1916)] giganteus (Linnaeus) Blaptica dubia Blatta W. H. i, Hebard ; Saussure [Blaberus Rehn, ; (pi. 4) He- orientalis; bard (1917)] (Shclfordella) lateralis (Walker) [Shelfordclla sure) ; tariara Princis (SausPeri- (1957). planeta tartara Saussure ; Bei- (Linnaeus) (pis. [Blatella ger5, A, B; 31, F) manica; Gurney (1952). Phyllogermanica ; Hebard dromia (1917). Ectobius germanicus; Gurney (personal communication, 1957)] landalei (Walker) Rehn and Hebard and Zehnt- lutca lutea (Saussure ner) minima Hebard (pi. 7, A, B) Rehn and Hebard orestera Rehn and Hebard punciipennis Hebard stenophrys (Walker) Rehn and Hebard instigator Rehn Hebard suave Rehn and Hebard Cariblattoides and Ceratinoptera picta Brunner barbadensis Chorisoneura Rehn and Hebard flavipennis Saussure Bienko (1950)] Blattella germanica Rehn Hebard imitans reticidosa Linnaeus [Periplaneta hylaca nebidicola 3) (1951)] orientalis ; lutca (pi. (Serville) clarazianus J. 2) [Blaberus Serville cubcnsis (Saussure and Zehntner) cuprea Hebard [Blattella delicatula (Guerin) Cariblatta delicatida Guerin punctidata (Beauvois) ; Rehn insularis craniifer Burmeister (pis. discoidalis Hebard and Hebard (1927)] [Blabera (StoU) fusca Brunner; Hebard (1917)] Blaberus nahuu (Saussure) Cariblatta antiguensis Hebard Hebard 6) (pi. Rehn Cahita borero Capucinella delicatula Hebard stigmosa Hebard jumigata (Guerin) [Temnop(Walker) (i 943)1 calymma [Latin- Caudell; (1920)] (Shelford) verticalis D) (Caudell) Byrsotria cabrerae Relin and Hebard Balta godeffroyi (Shelford) platysoma armata dia C, (pi. S, Buboblatta armata Aiidreia bromeliadarum Caudell scripta A) Bei-Bienko (1950)] vaga Hebard sexdentis Bolivar Atticola mortoni Bolivar quadricaiidata 7, (Brun- schubotzi Shelford schuppi Bolivar Hebard ; (fig. bisignata [Blattella Gurney jnngicola Wheeler teryx (Walker) lituricollis bergi Bolivar flava patula VOL. I4I and Zehntner Rehn and Hebard parishi Rehn spccilliger Hebard formosella texensis Saussure and Zehntner Ch riso n euro plocea Rehn Rehn and Hebard ( 1916) [ translucida (Saussure) ; ; ROTH & WILLIS BIOTIC ASSOCIATIONS OF COCKROACHES Choristima pallidns (Olivier) sp. Choristimodes A) sp. Chromatonotus infuscatus (Brunner) notatus (Brunner) Compsodes schwarzi (Caudell) liturata (Serville) Comptolampra [Compsolampra liturata; Comptolampra which is is the original spelling, followed by according Princis, 1959) to livens Kevan (1952) (w Roth and [Ectobius (Wesmael) cetorum (pi. (1923)] vittiventer (Costa) Bei-Bienko (Brunner) Saussure Ellipsidion ner; affine australe Saussure [Ellipsidion pel- lucidum Cyrtotria capucina (Gerstaecker) (1943)] (Brunner) Dendroblatta sohrina Rehn bicolor (Tepper) Derocalymma cruralis (Stal) [Homalodemas cruralis (Stal) simidans Hebard ; communica- [Apolyta Brun- Hebard (1943)] Hebard Gurney from photographs.] (personal Ramme ; (1951)] near sedilloti (Bolivar) (pi. 9) [Determined by Dr. A. B. Gurney [Ectobius vit- (Costa) tiventris soror (Brunner) 1957)] lampyrina Gerstaecker porcellio Gerstaecker Deropeltis autraniana Saussure (Fabricius) variegatutn conferta conspersa nigrita Saussure grisea Diploptera punctata (Eschscholtz) (pis. Burmeister (De Geer) maya Rehn mexicana Saussure niona Rehn and Hebard Eleutheroda notabilis dytiscoides (Ser- Rehn (1930)] Ectobius africanus Saussure albicinctns (Brunner) duskei Adelung lapponicus (Linnaeus) Eremoblatta subdiaphana (Scudder) Ergaiila Hgb. nicaeensis (Brisout) capensis cologamia [Ectobius perspicillaris Herbst, as used Blair Walker Walker tainana Rehn and Hebard zvheeleri Rehn sp. (fig. 7, B, C) sodalis (1948)] Dryadoblatta scotti (Shelford) [Homalopteryx scotti Shelford; ; He- Walker 3^) [Diploptera dytiscoides (Serville); Princis (19S0). io> Lucas (1920) ; azteca Saussure melanophila (Walker) Zimmerman [Ellip- sidion aurantium Saussure erythropesa Adelung ; Hebard ; bard (1943)] Epilampra abdomen-nigrum (De Geer) annandalei Shelford tion, lucidus eri- Ramme ; ; A) ville) (TurPrincis Willis, 1957)] Stephens panzeri ; semenovi Bei-Bienko Sylvester (Poda) [Ectobius sylRamme (1951)] vestris (Poda) tadzihicus Bei-Bienko ]_ Cutilia nitida sp. Ectobius ; ; 29, Gurney Cryptocercus piinctulatus Scudder relictus ton) C; (Fabri- panzeri var. nigripes Stephens Cosmozostcria lateralis (Walker) 8, cius) (pis. 7, lividus Dr. K. communication, (personal [Ectobius ( 1934)1 by (Saussure) capensis [DysSaussure Dyscologamia woUastoni Kirby Princis ( 1957)1 scarabaeoides Walker [Dyscolo- (Walker) Princis (i9S7). Parapolyphaga gamia piolosa ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 10 Chopard erecHpilis Princis ; (1950). Dyscologamia chopardi Bruijning Hanitsch; (1948). Miroblatta silpJioides Chopard; Hebard Escala sp. Euandroblatta palpalis Chopard Eublaherns posticus (Erichson) Princis ; carbo- cyanea (Burmeister) julva (Burmeister) metallica Rehn and Hebard (Fabricius) nitidula bivittata (Brulle) chavesi (Bolivar) ; B olivar ) (and hence, by inference, Eury- (Scudder) sabalianus Hebard (1917)] Rehn and Hebard improccra Rehn galeoides marginata (Schreber) punctata (Charpentier) [Aphlebia Charpentier punctata (1951)] Hololcptoblatta ; Ramme sp. Homalopteryx laminata Brunner Hoplosphoropyga babaulti Chopard Hormetica apolinari Hebard laevigata Burmeister ventralis Burmeister kevani Princis Ignabolivaria bilobata Chopard Rehn Ischnoptera deropeltiformis (Brunner) manni Rehn (pi. opaca (Brunner) panamae Hebard podoces Rehn and Hebard Geoscapheus robustus Teppcr notulata Stal; Phyllodromia rufa occidentalis Saussure [Blatta (Stal) Hebard (1929). hieroglyphica Brunner; Kirby (1904)] Gromphadorliina laevigata S. and portentosa (Schaum) ; (1917)] Euthyrrhapha nigra Chopard pacifica Coquebert Graptoblatta notulata 12A) [Temnopteryx deroBrunner Hebard peltiformis Euthlastoblatta abortiva (Caudell) (pi. rufa rufa (De Geer) Karney • schenklingi Karnyia discoidalis (Brunner) Kuchinga hemerobina (Gerstaecker) [Phyllodromia hemerobina Ger- Z. 12, A, B) staecker; Rehn (1932)] remota Hebard Gyna kazungnlana Giglio-Tos niaculipennis (Schaum) vetula Holocompsa asteca (Saussure) ; Platysosteria sabalianus Scudder lixa con- Bei-Bienko (1957) Gurney (personal communication, 1959)] jemim-eqninitm Rehn and Hebard (pi. (Walker) 11) floridana [Platyzosteria ingens Scudder cotis ; macnlata (Schreber) [Aphlebia Harz maculata Schreber Rehn (1918)] decipiens (Kirby) ( [Blatta Blattina (1950)] Hololampra caraibea (Bolivar) dimidiata cinna Haan; (Haan) zapoteca Saussure (1959)] Eurycotis bananae Bei-Bienko biollcyi Rehn [Eurycotis naria Biolley; concinna rugosa (Schulthess) Eudromiella bicolorata Hebard calcarata Bei-Bienko Euphyllodrotnia angustata (Latreille) de[Euphyllodromia litnrifera Hebard Hcbardina concinna (Haan) Hemiblabera brunneri (Saussure) Henicotyle antillarum (Brunner) Hcterogamodes krilgeri (Salfi) (1929)]. castigmata VOL. I4I Brunner; (1909b)] trisHs Hanitsch \_Gyna Shelford Lamproblatta albipalpus Hebard meridionaUs (Brunner) Latiblattella chichimeca (Saussure and Zehntner) [Blattella chichimeca S. and Z.; Hebard (1932)] BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS II rehni Hebard Hebard vitrea (B runner) Myrmeblattina longipes (Chopard) Myrmecoblatta rehni Mann sapoteca (Saussure) Naniablatta bitacniata (Stal) lucifrons wheeleri Hebard Leucophaea maderae (Fabricius) (pi. 13) [Rhyparobia maderae ; Hebard (1917). Panchlora maderae; Kirby Very (1904). probably "Blaberus" maderae is a careless reference to this species Gurney ; (personal com- (Brunner) (Saussure) Lciirolestes pallidus (Saussure) marginata Caudell [Aiidreta bispinosa ; Hebard nigra (1930)] Nauphoeta cinerca (Olivier) (pi. 14) [Nauphocta bivittata Burmeister; Zimmerman (1948)] (Walker) fiexivitta Rehn miisarum Rehn [Nauphoeta (Bolivar) (1937)] punctipennis Chopard Nelipophygtis ramsdeni Rehn ; Rehn brunneriana (Saussure) [Blaftella brunneriana; Gurney communication, (personal Lobolampra subaptera Rambur Loboptcra decipiens (Germar) thaxteri Hebard carcinus celeripes Rehn and Hebard Rehn and Hebard Lobopterella detersa (Walker) 1959)] dimidiatipes (Bolivar) [Loboptera dimidiatipes (Boli- var) Princis ; (1957a). Lobop- (Saussure) ; Hebard (1933a). Loboptera extranea Perkins; Hebard (1922). tera sakalava Princis (1957a) in erecting Lo- bopterella pointed out that only the nontypical variety of saka- lava is identical with dimidia- tipes.] Lophoblatta arawaka Hebard Macropancsthia rhinocerus Saussure Mar eta acntiventris Chopard Marctina uahuka Hebard Mcgaloblatta blabcr aides [Megaloblatta (Walker) rufipes Dohrn; Hebard (1920)] Megamareta verticalis Hebard Mclanosilpha capensis Saussure Zehntner Methana canae Pope curvigera (Walker) marginalis (Saussure) Mohichia (f) dahli Princis Monastria bigiittata (Thunberg) Miizoa madida Rehn and Hebard Neoblattella (1920)] deianira (Brunner) [PoroBrunner; Rehn nigra blatta brassae munication, 1957)] Litopeltis biolleyi Nauclidas dryas Rehn and Hebard eurydice Rehn and Hebard Hebard fratercula fratcrna (Saussure and Zehntner) Rehn and Hebard laodamia Rehn and Hebard grossbccki nahiia (Saussure) [Blattella nahua Saussure and Zehntner of Cau(1914) Hebard (1920)] Proserpina Rehn and Hebard dell ; semota Rehn and Hebard tridens Rehn and Hebard vatia Rehn and Hebard Neostylopyga rhombifolia (Stoll) (pi. [Dorylaca rhombijolia; 15) Rehn (personal communication, 1956)] and Nesomylacris cubensis Rehn and Hebard rclica Nocticola Rehn and Hebard boliz'ari Chopard caeca Bolivar decaryi Chopard simoni Bolivar sinensis Silvestri termifopliila Silvestri ; ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 12 Nothoblatta wasmanni (Bolivar) (Saussure and Zehntner) divisa Notolampra antillarmn Shelford Nyctibora astcca Saussure and Zehntner [Ischnoptera divisa S. and Z. Hebard (1917)] (Saussure and Zehntuhleriana fulvescens hrtmnea (Thunberg) laevigata (Beauvois) lutzi Rehn and Hebard mexicana Saussure ner) [Ischnoptera fulvescens S. and Z. (in part) (Brunner) lata ohscura Saussure sure) H. (not Walker tomentosa [Nyctibora Serville Burmeister latipennis Hebard ; (1917, p. 263)] chlora cubensis Saussure; Gur- Pycnosceloides ; Hebard (1921c)] angustipennis javanica B runner laeincollis (pi. 8, (Illiger) Serville C; tera borcalis fig. (pis. 6) 17, ; Hebard [?5 of Ischnoptera and H. Ischnoptera ; uhleriana fulvescens S. and Z. Hebard (1917)] (Rehn and Hebard) ; of Ischnoptera insolita R. H.; Hebard (1917)] B; [Ischnop- Brunner; Hebard (1917)] zebra Hebard Pclmatosilpha coriacea Rehn kevani Princis marginalis Brunner purpiirascens (Kirby) (Linnaeus) americana 35) [Stylopyga americana; Blatta americana L. Hebard (1917)] (pis. 19, ; (pis. 20, (pi. 21) cavcrnicola Chopard spadica (Shiraki) (in part) (Brunner) 33, brunnea Burmeister Parahormctica bilobata (Saussure) Parcoblatta amcricana (Scudder) bolliana (Saussure and Zehntner) [Kakerlac schaeffcri Rehn Hebard (1917)] desertae A; 32) Saussure insolita R. Hebard (1917)] australasiae (Fabricius) B) lobipennis Brunner caudelli 18) Saus- uhleriana vagabunda Princis Hebard (1929)] australis sure; virginica 27, 17, Periplaneta virescens (Thunberg) [Panesthia (pi. [Ischnoptera pcnnsylvanica rotundata Scudder peruana Saussure sagax Rehn and Hebard Panesthia Hebard ; (De Geer) pensylvanica [Ischnoptera Oulopteryx meliponarmn Hebard Oxyhaloa buprestoides (Saussure) deusta (Thunberg) Panchlora antillarum Saussure exoleta Burmeister fraterna Saussure and Zehntner nivea (Linnaeus) (pi. 16) [Pan- Hebard and Z.) uhleriana orientalis (Burmeister) (iQSS). S. Saussure; Hebard (1917)] (pi. (Saussure) Opisthoplatia macxdata Shiraki aporus Ischnoptera major R. and ; (1917)] notha Rehn and Hebard A) Oniscosoma granicollis (Saussure) ney [Ischnoptera coidoniana R. and H. (not Saus- sericea Burmeister stygia ; Hebard (1917)] Rehn noctivaga VOL. I4I (cfd* and ftdiginosa (Serville) ignota lata (pi. 22) Shaw (Herbst) Pcrisphaerus armadillo Serville glomeriformis (Lucas) Phaetalia pallida (Brunner) Phidon (?) dubitis Princis Phlcbonotus pollens (Serville) Pholadoblafta inusitata (Rehn) Phorticolea boliviae Caudell testacea Bolivar "Phyllodromia" treitliana Werner BIOTIC ASSOCIATIONS OF COCKROACHES Phyllodromica brevipennis (Fischer) graeca (Brunner) irinae (Bei-Bienko) maculata (Schreber) megerlei (Fieber) polita (Krauss) pygmaca (Bei-Bienko) Platysosteria analis (Saussure) [Poly- Kirby (1904)] armata Tepper bifida (Brunner) novae seelandiae [Periplaneta (pi. fortipes Walker; Shelford (1912) Pla- ; Plectoptera dorsalis (Burmeister) (Saussure) [Plectoptera ; (Linnaeus) aegyptiaca [Blafta aegyptiaca (1950). aegyptiaca (L.) ; L. Bei- ; Heterogamia Gurney (per- communication, 1957). aegyptiaca; ac- "Polygamia" cording to Gurney (p. c), there is no genus Polygamia and almost surely the reference is to Polyphaga aegyptiaca.] indica Walker [Polyphaga pellti(Redtenbacher) ; Princis (Dohrn) (Burmeis- Kirhy Gurney (personal striata ; communication, 1957)] surinamensis (Linnaeus) (pi. 24) [Leucophaea surinamensis (L.) ; cephala StoU; Kirby (1904)] Rhicnoda natatrix Shelford Rhytidometopum dissimile Princis Riatia fnlgida (Saussure) [Lissoblatta (Saussure) Gurney ; orientis Hebard (personal aethiopica [Blatta aethiopica sure; Gurney (personal munication, 1957)] ; commimica- tion, 1957)] longiuscula (Walker) [Escala longiuscula (Walker) Gurney ; (personal communication, 1957) ] Salganea morio (Burmeister) Sibylloblatta panesthoides Simblerastes jamaicanus (Walker) Rehn and Hebard Spelaeoblatta gestroi Bolivar Sphecophila polybiarum Shelford ravana Fernando tcrmitium Shelford Stclcopyga (?) sinensis Walker [Dr. Gurney (personal communicaa 1957) could not reference to this species. Walker named sinensis find tion, described species in three genera different cockroaches, and Stictolampra melanaria (Erichson) sure) (Walker) this it is of uncertain combination rep- resents.] Polysosteria limbata Burmeister Pseudoderopeltis Hebard ter) which one (1957)] saitssurei septentrionalis Pseudophoraspis nebulosa Gurney Hebard Rehn and He- Bienko cida laticornis Perty [Escala circumducta (Walker) bard (1927)] porcellana (Saussure) pygmaea (Saussure) rhabdota (Rehn and Hebard) vermiculata Rehn and Hebard sonal Hebard (1917)] Robshelfordia circumducta (Rehn and Hebard) lacerna Relin and Hebard pcrscita Rehn and Hebard infidata Polyphaga Scudder; cincta (personal communication, 1959)] scabra (Brunner) floridana (Burmeister) cincta [Thyrsocera fulgida tysosteria novae-zealandiae] poeyi 13 Hebard (1917). Blatta melano- (Saussure) castanca (Brunner) ^3) Pseudomops & WILLIS Pycnoscelus niger (Brunner) (Kirby) [Leucophaea striatus tartara (Saussure) tartara nigrcscens Bei-Bienko zosteria analis Saussure; —ROTH (SausSaus- com- buqiieti concinula (Walker) Styphon bakeri Rehn Supella hottentotta (Saussure) supcllectilium (Serville) 30, B-E; 31, A-E) 25; [Phyllo- (pis. SMITHSONIAN MISCELLANEOUS COLLECTIONS 14 dromia supcllectilium (Serv.) bilabiafa (Shelford) [Ischnop- (Shelford); cavernicola tera „, „ , ^, Fhyllodronna ntqrocincta Cno. . • Rehn and [Sytuploce Hebard; Princis Symploce breviraviis (Hanitsch) cavernicola (Fabricius) ruficollis ; Bei-Bienko (1950)] VOL. I4I , (1949a)] TartarobJatta karatavica Bei-Bienko Tcmnopteryx obliquetruncata Chov^vA phalerata (Saussure) ^ ^, j heganoptcryx straminea r-t Chopard j^,^^^ nuptialis (Gerstaecker) [Cot^, 1 pard; Hebard (1929)] curta Hanitsch flagcllata y^,^/^ Hebard hospes (Perkins) nuptialis Gerstaecker; Princis (1950)] [Symploce lita Tivia australica Princis Hebard; Hebard (1922)] jamaicana (Rehn) brunnea (Chopard) M'^^a. (Burmeister) kevani Chopard ^nacracantha Chopard parenthesis (Gerstaecker) ... , . loaromta Parenthesis staecker; Rehn (1932)] remyi (Hanitsch) remyi [Phyl- ^ f.^l^;"^ caeca Typhloblatta Ger- [Ischnoptera Hanitsch; Chopard i p^^d (1924b)] Typhloblattodcs madccassns Chopard Xestoblat fa festae (Griffmi) immacidata Hebard (1938)] III. The (Chopard) (Chopard) {Spe1. t^i r-^ j r-i laeoblatta caeca Chopard; Cho- ^ ECOLOGICAL RELATIONSHIPS ecology of extinct cockroaches From is necessarily a highly specu- and fossil same geological stratum, one might conclude that there had been intimate associations between them during prehistoric life. Heer (1864) and Goldenberg (1877) suggested that Carboniferous cockroaches fed on the plants with which they have been found as fossils. Scudder (1879) concurred with this hypothesis. However, Bolton (1911), remarking on the noticeable associations of blattoid wings with vegetable remains, suggested that the cockroaches may have been partly carnivorous, feeding on the snails Spirorbis pusilhis, which were attached to the leaves of Cordaites. Yet the proximity of fossil insects and plants in the same geological formation is hardly lative subject. the coexistence of fossil cockroaches plants in the proof of a similar association during life. In fact, Sellards (1903), Bolton (1921), and Laurentiaux (1951) have all pointed out that the may have been washed into streams by heavy rains and transported with drifting plant material to places where permanent deposits were accucockroach remains, particularly the more resistant wings, mulating. Some species of fossil cockroaches have long, well-developed ovi- positors, very unlike present-day cockroaches whose ovipositors are and nonprotruding. Brongniart (1889) and Zalesskii (1939, 1953) have suggested that certain Permian and Carboniferous cock- stuall 5 BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 1 may have inserted their eggs singly and other plants, rather than protecting the eggs with an ootheca. However, Laurentiaux (1951), although conceding the possibility of egg laying in vegetable material, suggested that oviroaches with long ovipositors into trees position into the earth is more probable because of the unbending nature of the ovipositor. Although the ecological associations of modern cockroaches should known from direct observation, actually most species are still little more than names on museum specimens, and our knowledge of them is fragmentary. All too frequently ecological observations have been only incidental to taxonomic or faunistic studies; yet the biobe well logical information that is contained in such papers is all that many of species. For this reason we have we know cited these observations in some detail, especially when they were brief; longer accounts of cockroach bionomics, of necessity, have been abstracted. Very few exclusively cockroaches. The ecological studies of insects have included native woodroaches (Parcoblatta pensylvanica, P. uhleriana, and P. wrginica) of the northern United States were included in ecological studies of the Orthoptera by Hubbell (1922), Strohecker (1937), and Cantrall (1943). Fifteen species of cockroaches were included in an ecological study of the Orthoptera of northern Florida by Friauf (1953). The original papers should be consulted for detailed descriptions of the habitats and accounts of the associated plants and other Orthoptera. In this chapter the cockroaches are grouped into those that have been found in man-made structures and those that occur in other habitats. Certain species may appear in several categories because The structural pests are divided into they live both indoors and out. cockroaches that occur in land-based structures, those on ships, and The nonstructural cockroaches are divided into those in aircraft. those that occur in quite specific habitats (caves, water, and deserts) and those that occur generally out of doors. Nests of various arthro- pods serve as microhabitats of commensal cockroaches; these latter associations are discussed on pages 310-318. In this chapter our discussion is limited to the physical environment and specific habitats of cockroaches, and only very general references are made to associated organisms. to the biota are examined in The detail in relationships of cockroaches subsequent chapters. To show the full extent of the associations, the associates, from bacteroids to vertebrates, are arranged phyletically. sifications serve within common These associate-centered clas- admirably to relate various species of cockroaches bounds, but fail to give an integrated account of the SMITHSONIAN MISCELLANEOUS COLLECTIONS l6 total biotic relationships in the ecology of each species. physically separated in this monograph, the many species of cockroach should all VOL. I4I Although associates of each be considered in appraising the ecology of that species. To assist the reader to achieve this end, we have included a checklist (p. 290) which serves as a convenient index to certain organisms associated with particular species of cockroaches. CAVE HABITATS Caves, mines, and animal burrows are somewhat similar habitats many species of cockroaches with shelter and frequently that provide with food. The been described in that microclimates of these cockroach habitats have not detail in the papers cited, but it seems rather obvious man-made caves (mines), and burrows natural caves, offer and humidities and protection from adAlthough such cavernicolous animals as relatively stable temperatures verse climatic conditions. and bats periodically leave caves to search for food, cockroaches guano and animal and plant detritus an entirely adequate dietary (Chopard, 1938). Cockroaches in mines presumably subsist on the food and feces dropped by man and mine animals (e.g., pit ponies). Food stored in their nests by burrowing animals is undoubtedly utilized by the associated cockroaches. birds find the accumulated Cavernicolous cockroaches show varying degrees of dependence on and adaptation to these specialized habitats. Some of the common domiciliary species {Blatta orientalis, Blattella germanica, and Peri- may have accompanied man into caves and remained there after he left (Chopard, 1929a, 1936, 1938). Other species, from the paucity of records noting their occurrence in caves, are undoubtedly accidental inhabitants that may never become estabplaneta americana) lished. many Besides these, however, other species of cockroaches of the latter species Although some show very pronounced morphological adaptations many others resemble their noncavernicolous relatives. have established large breeding colonies in caves. to a cave life, The possible origin of cavernicolous Orthoptera has been discussed by Chopard (1938). Cavernicolous cockroaches have been segregated into four groups according to their ability to adapt to their environment and the degree (i) Trogloxof their specialized evolution (Chopard, 1936, 1938) enes: Cockroaches that occur in caves in a sporadic fashion (the : domiciliary cockroaches and accidentals such as Ectohius and Hetero- Cockroaches found habitually in gamodes). (2) Troglophiles caves {Symploce, Periplaneta cavernicola). (3) Guanobies: Cock: . BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS I7 roaches that live in the guano of cavernicolous vertebrates (Gyna, Acanthogyna, Dyscologamia, Pycnoscelus). (4) Troglobies: Cockroaches that apparently cannot live outside of caves and which show very marked adaptive characters {Alluaudellina, Nocticola, Spelaeoblatta, Typhloblatta) . For complete discussions of these groups inshown by certain cluding descriptions of the adaptive characters genera, the original sources should be consulted. Although we know very cavernicolous cockroaches, little it is of the ethology of most of the intriguing that three of the six known two are inhabitants of termite and one holivari) (N. was found under stones and 315), species of Nocticola are cave dwellers, nests (p. cement blocks (Chopard, 1950b). In the rather extensive list of cavernicolous cockroaches only two (Arcnivaffa grata and Parcohlatta sp.) were taken from caves in North America north of Yucatan. All other records are from Africa, Asia, Central America, Europe, West and the Philippine Islands. This we find puzzHng. Packard (1888) in his extensive study of the cave fauna of North America listed no cockroaches. Dearolf (1941) found only the abovementioned Parcohlatta in one of t^'j caves in Pennsylvania. Kohls and Jellison (1948) listed no cockroaches among the arthropods from six bat caves in Texas. We would expect Periplaneta americana to inhabit mines in North America, but we have found no such records. Have cockroaches been ignored in faunal collections from North American caves, or has our cave fauna been less extensively studied than that of other parts of the world ? The two species of cockroaches found in mines (BlatteUa germanica and Periplaneta americana) are also found in caves. For this reason we have included them in the list headed Cavernicolous Cockroaches. On the other hand, the cockroaches found in animal burrows are generally different species from those found in caves, so we have grouped these together in a second list. Indies, East Indies, CAVERNICOLOUS COCKROACHES Alluaudellina cavernicola Tanganyika. —From Kulumusi caves, near Tanga. The eyes of this cockroach are reduced to a pair of slender streaks (Shelford, 1910a; Chopard, 1932a). East Africa. — Chopard ( 1936) Apotrogia angolensis Belgian Congo. acters. —A troglophile without well-marked adaptive char- Collected in moist sand on floor of a sandstone grotto in- SMITHSONIAN MISCELLANEOUS COLLECTIONS l8 habited by bats (Chopard, 1927, 1950a). Taken in VOL. I4I many caves in Bas Congo (Leleup, 1956). Apteroblatta perplexa East Africa. —Accidental inhabitant of cave (Chopard, 1936). Arenivaga grata Arizona. the guano —"A female and in a bat cave in the many nymphs were taken by Flock in Tucson Mountains" (Ball et al., 1942). Aspiduchus borinquen — Puerto Rico. In limestone cavern by thousands in grass and on (Rehn and Hebard, 1927 Rehn, J. W. H., 1951a). walls ; Aspiduchus cavernicola Puerto Rico. — In limestone cave, in caves inhabited by bats, and removed from entrance. "In this great numbers were seen on the side walls and roof" apparently seen in other caves well latter situation (Rehn,J.W.H., 1951a). Blaberus atropos Yucatan. —Found once, in Xmahit cave (Pearse, 1938). Blaberus craniif er Yucatan. — Collected within three caves, near the entrances (Pearse, 1938). Blaberus giganteus Panama. —Two males and in the second chamber of the several nymphs were taken under rocks Chilibrillo cave ; some also were on the walls (Caudell, 1924). Blatta lateralis — Turkmen S.S.R. All stages, but more often females and nymphs, were found in the middle and back part of Bakharden cavern, which was inhabited by tens of thousands of bats (Vlasov, 1929). Blatta orientalis Turkmen S.S.R. —All bat cave. This cave Bakharden was uninhabited by man but supported a variety stages found in front part of of other animals (Vlasov, 1929). BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS IQ Blattella germanica South Africa. — Numerous mine on the Witwatersrand in a gold (Porter, 1930). Tonkin. — Chopard (1929a); Colani (1952). Byrsotria fumigata Cuba. — Cueva de las Cucarachas, Province: 21 specimens, "It is La Pantana, evident . . . Baracoa, Oriente that the species is also a cave inhabitant" (Rehn and Hebard, 1927). Deropeltis erythropeza — East Africa. Found at entrance of cave; not a form according to Chopard (1936), strictly cavernico- lous Ectobius pallidus France. — Nymph in cave in Basses-Pyrenees, accidental inhabitant (Chopard, 1936). Ectobius vittiventer Italy. — In detritus at base of entrance shaft of Acquaviva cave in the Venezia Tridentina (Conci, 1951). Ectobius Italy. — Found in the sp. heap of saprophytic detritus at the base of the entrance shaft in the Acquaviva cave (Conci, 1951). Ergaula scarabaeoides —West coast (Hebard, 1929). — Found burrowing bat guano among stones Sumatra. Malaya. in to caves in Selangor at entrance (Chopard, 1919, 1929). Euthyrrhapha nigra Madagascar. —Three males and six females in guano in Antsinomy grotto (Chopard, 1949a). G3ma kazungulana — East Africa. This species is especially found in caves although it shows no special adaptive characters. It is a typical guanobe (Chopard, 1936). Gyna maculipennis Belgian Congo. —Troglophile, Lualaba (Leleup, 1956). guanophile. Found in two caves in . SMITHSONIAN MISCELLANEOUS COLLECTIONS 20 Gyna Belgian Congo. — VOL. I4I tristis In three caves in Uele (Leleup, 1956). Heterogamodes North Africa. — An kriigeri accidental inhabitant of caves (Chopard, 1938). Holocompsa zapoteca Yucatan. — Common throughout rather dry, dusty caves in southern Yucatan (Pearse, 1938). Hoplosphoropyga babaulti Stated to be a troglophile by Chopard (1938). Nocticola caeca Philippine Islands. — Bolivar (1892). Nocticola decaryi Madagascar. —A true troglobie according to Chopard ( 1945) Nocticola simoni Philippine Islands. —Bolivar (1892), Parcoblatta sp. Pennsylvania. — Found in Merkle cave, Berks County (Dearolf, 1941). Periplaneta americana IN CAVES East Africa. — Its presence in the cave at Shimoni was thought to man had sought refuge there and brought the cockbaggage or provisions (Chopard, 1936). Many present in cave at Vengurla, the floor of which was India. covered with bird guano (AbdulaH, 1942). Madagascar. Thought to have been introduced into the cave en- indicate that roaches in with — — trance by man (Chopard, 1945, 1949a). IN MINES —In a Pontewydd where they had been established for some years (Lucas, 1916). In the Pentre Pit mine where they were abundant (Lucas, 1918). Abundant in a Welch mine 2,166 feet below the surface (Lucas, 1925). This species was Great Britain. coal mine at BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 21 found quite commonly in a number of South Wales coal mines; in one deep mine a white-eyed mutant form comprised about 5 percent of the cockroach population for the preceding 11 years (Jefferson, 1958). India, western Bengal. —Very numerous human Numerous in sole food apparently was South Africa. — mines where the in coal feces (Chandler, 1926). four deep-level gold mines on the Witwatersrand. — Sumatra. Numerous males and females from Sawah Lunto " 'from a coal mine where they lived in great numbers on the faeces of miners' " (Hanitsch, 1929). Periplaneta australasiae — — Found swarming on walls of caves and in soft bird company with Symploce cavernicola (Moulton, 1912). Tonkin. Chopard (1929a); Colani (1952). Saraivak. guano in Periplaneta cavernicola —Taken Malaya. ticularly on walls of inner caverns, where they were par- abundant (Chopard, 1919). Periplaneta lata Tonkin. — Chopard suggested that linked with man (Chopard, —From a cave in Jalor Malaya. —The wingless probably sp. (Annandale Perisphaerus in a is 1929a; Colani, 1952). Periplaneta Malaya. presence in caves its females and et al., 1913). sp. nymphs mined in bats' guano cavern of the Jalor caves (Annandale, 1900). Polyphaga aegyptiaca Turkmen S.S.R. — Females found Bakharden bat in front part of cave on several occasions (Vlasov, 1929). Turkey. —At Magharadjik and Arab Dede, found in caves with various other animals (Lindberg, 1954). Polyphaga Burma. — Hsin Dawng sp. Cave, S. Shan States, i under stone in complete darkness (Chopard, 1924b). immature male ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 22 VOL. I4I Pycnoscelus niger Tonkin. —Apparently not an accidental inhabitant as nymphs were present (Chopard, 1929a; Colani, 1952). Pycnoscelus striatus Malaya. —Found burrowing Selangor, where it in bats' guano at entrance to caves in was very abundant 50 to 600 feet from entrance on walls of inner cavern (Chopard, 1919, 1929). In the absence of other evidence, the presence of P. striatus in a cave indicates that also bats also inhabit the cave (Chopard, 1929a). Pycnoscelus surinamensis — Assam. Found 300 to 400 feet from entrance of Siju cave Garo Hills (Chopard, 1924b). South Celebes. Hanitsch (1932). in the — Spelaeoblatta gestroi Burma. — Chopard teristics of dale, stated that this species adaptation to a life in shows marked charac- darkness (Bolivar, 1897; Annan- 1913; Chopard, 1919). Symploce breviramis South Celebes. —Hanitsch (1932). Symploce cavernicola — Sarawak, Borneo. Swarming on walls of caves and in soft bird guano on the cave floor (Moulton, 1912). Hanitsch (1931) noted that this species was first recorded by Shelford from a cave in Sarawak and that there is a series from a cave in the Oxford University Museum, taken by Banks in 1928. — On the walls of the inner cavern of a cave Biserat; Sumatra. — From Baso cavern, on the west coast (Hebard, 1929). Malaya. at the insects covered the walls in places (Chopard, 1919). Symploce curta South Celebes. —Hanitsch (1932). Symploce remyi Tonkin. —This seems 1929a; Colani, 1952). to be a true cavernicolous species (Chopard, BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 23 Tivia macracantha Belgian Congo. —A troglophile without well-marked adaptive char- acters (Chopard, 1950a). At Haut-Katanga, troglophile and guano- guano Antsinomy phile (Leleup, 1956). Tivia Madagascar. sp. —Last-stage nymphs captured in in grotto (Chopard, 1949a). Typhloblatta caeca Assam. India, —An eyeless species with noticeably elongated ap- pendages (Chopard, 1945). Typhloblattodes madecassus Madagascar. —Unpigmented integument and reduced eyes (Cho- pard, 1945). Xestoblatta immaculata Panama. —Found under rocks on guano-covered floor of the Chili- caves (Caudell, 1924). brillo bat Unidentified cockroaches —The walls of a cave were covered by dense groups of a pests of the mines are cockroaches, England. — "The chief Malaya. species of "Blatta" (Annandale, 1900). insect which often swarm in hot mines and those with pit pony stables ." . . (Hardy, 1941). COCKROACHES FROM THE BURROWS OF VERTEBRATES Arenivaga apacha Arizona. — In the nests of wood rats, Ncotoma sp. (Hebard, 1917). sp, (Hebard, 1917; Arenivaga boUiana Texas. — In the nests of wood rats, Ncotoma 1943a). Arenivaga erratica — The wingless females were commonly found in burrows Dipodomys spectabilis spectabilis Merriam, the kangaroo rat. The Arizona. of winged males were never found in the burrows (Vorhies and Taylor, 1922). Found most commonly in wood-rat and ground-squirrel dens in the desert regions (Ball et al., 1942). SMITHSONIAN MISCELLANEOUS COLLECTIONS 24 VOL. I4I Arenivaga floridensis — Florida. Found in a burrow of Peromyscus polionotus rhoadsi (Bangs), the white-footed mouse (Young, 1949). Arenivaga roseni — Occasionally found in burrows of Rhombomys burrows of the desert turtle, Testudo horsfieldi Gray; and frequently in burrows of the ground squirrel, Spermophilopsis leptodactylus Lichtenstein ( Vlasov, 1933 Vlasov Turkmen SS.R. opimiis Lichtenstein ; in the ; and Miram, 1937). Arenivaga tonkawa Texas. —An immature specimen was found in a prairie-dog hole (Hebard, 1943a). Cariblatta lutea Florida. Geomys — sp. It has been taken in burrows of the pocket gopher, (Hubbell and Gofif, 1940). Euthlastoblatta abortiva Texas. — In the nests of wood Texas. — In the nests of wood rats, Ncotoma sp. (Hebard, 1917). sp. (Hebard, 1917). Parcoblatta fulvescens rats, Neotoma Polyphaga aegyptiaca — Turkmen S.S.R. Nymphs and adult females were often found burrows of the sand mouse, Rhornboniys opimus (Vlasov, 1933). in Polyphaga indica Turkmen S.S.R. —This species prefers sandy soils where it can be found in burrows of Spermophilopsis leptodactylus and Pallasiomys meridionalis pennicilliger Heptner (Vlasov and Miram, 1937). Polyphaga saussurei and rodents (Zmeev, — Found burrows of Turkmen S.S.R. — Nymphs and adult females are common in burTadzhikistan. in turtles 1936). rows of Rhombomys opimus and in burrows of Testudo horsfieldi. Its principal habitat is rodent burrows in loess dust, where it is not infrequently found in the food stores of the host (Vlasov and Miram, 1937). BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 25 Pycnoscelus surinamensis Texas. — In the nests of wood rats, Neotoma sp. (Hebard, 1917). DESERT HABITATS There is relatively little ecological information about cockroaches that live in deserts, even though certain species, notably Polyphaga aegyptiaca, have long been known to inhabit arid zones. In fact, so little is known about the ecology of arid-zone insects in general that it is more a subject for research than for review (Pradhan, 1957). In their account of the cockroaches of Northern Kenya and Jubaland, Kevan and Chopard (1954) describe in some detail the vegetational areas of this arid desert or semidesert country, which averages only about 10 inches of rain per year. The other sources that are cited below contain very little more biological information than the abstracted material that Nearly all is given under each species. the Polyphaginae are said to be marked xerophiles whose distribution coincides with that of the deserts (Bei-Bienko, 1950), the exception of Arenivaga floridana, the species of Polyphaginae in the United States all occur in the Southwest, where they are (with a few exceptions) the only cockroaches that inhabit the desert With regions (Hebard, proper 1917). The Polyphaginae reach greatest diversity in the deserts of Northern Africa their and Anterior and South-Central Asia (Bei-Bienko, 1950). Some of the desert-inhabiting species have also been found under nondesert conditions. This only further exemplifies the plasticity of cockroaches in adapting to different environments. The ability of desert insects to live under what appear to be ex- tremely unfavorable conditions has been abundantly illustrated by Pradhan (1957) Uvarov (1954) has pointed out that a desert "covers . a great variety of landscapes, which provide desert animals with a wide range of habitats, some of them offering very favorable conditions Pradhan (1957) stated that many desert animals avoid extremes of desert climates by choosing suitable microclimates for the resting places, that a permanent or temporary underground diurnal for life." very common among insects in arid zones, and that many Orthoptera burrow into the soil or hide under stones where nocturnal lower. For example, the type of Parcoblatta desertae temperatures are was found under a boulder on the bare desert (Rehn and Hebard, existence is 1909). Symbiosis with burrowing animals is another solution to the prob- SMITHSONIAN MISCELLANEOUS COLLECTIONS 26 lem of existence in the desert; in fact, symbiosis is a VOL. I4I mode of life adopted by nearly half of the desert cockroaches about which we have any information. Vlasov and Miram (1937) iomid Poly phaga indica, Polyphaga saussiirei, and Arenivaga roseni in the burrows of rodents and desert turtles. In the desert regions of Arizona, females of Arenivaga erratica were found commonly in burrows of the kangaroo rat (Vorhies and Taylor, 1922) and in dens of wood rats and ground squirrels (Ball et al., 1942). Arenivaga apacha and Arenivaga holhave also been found inhabiting the nests of wood liana rats (Hebard, 1917; 1943a). Bei-Bienko (1950) has suggested that the adaptation of desert-inhabiting cockroaches to rodent burrows might enable these insects to survive in the severe climatic conditions of deserts in summer. Under desert conditions in southern Arizona, the relative humidity outside of the burrows of the kangaroo rat the day and 15 to 40 percent at night ; is i to 15 percent during but inside the burrows the rela- 30 to 50 percent, and the temperature, even during the day, is below 30° C. (Schmidt-Nielsen, 1949). Thus by living in rodent burrows during the day and going outside at night, the desert tive humidity is cockroaches could avail themselves of the most favorable microclimates obtainable. Presumably whatever food these insects eat prothem to survive under vides them with sufficient water to enable Bodenheimer (1953) has suggested that the extent of dew, which is sometimes heavy in the desert, should desert conditions. of utilization be investigated; he stated that tenebrionid beetles have been seen in the early morning eating dry [dead?] herbs that were still wet with dew. It is is a need for additional detailed informawhich we can only guess about the ecology of desert obvious that there tion without cockroaches. list we have cited only those species that were found under desert conditions. Undoubtedly, rebeen have stated to lated species that have been taken in similar localities are also desertinhabiting forms, as, for example, other species of Arenivaga that In the following were collected in Texas by Hebard (1943a). In the absence of information linking such other species with deserts, trarily relegated those forms to the section specific we have on outdoor arbi- habitats. In addition to the species listed below, desert cockroaches are said to be found in the following genera Nymphytria (Chopard, 1938). : Anisogamia, M ononychohlatta, and BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS TTJ DESERT COCKROACHES Agis orientalis Northern Kenya. —In desert-grass and thorn-bush country among acacia bush and : tered, dry tufts of grasses interspersed scatscat- (Kevan and Chopard, 1954). tered trees Arenivaga apacha U.S.A. — Inhabits desert regions of the Southwest, has been found wood in nests of (Hebard, 1917). rats Arenivaga bolliana U.S.A. of wood —On gravelly rats in hillocks, in scattered scrub, Texas. It is and in the nests a desert inhabitant in the Southwest (Hebard, 191 7; 1943a). Arenivaga erratica U.S.A. — Inhabits desert regions of the Southwest (Hebard, 1917). In Arizona it has been found in rodent burrows in the desert (Vorhies and Taylor, 1922 Ball ; et al., 1942). Arenivaga roseni Turkmen S.S.R. stages "swim" in — Predominantly found in burrows in sand; all sand and loess dust (Vlasov and Miram, 1937). Blattella vaga Arizona. —Found in small numbers on the dry desert (Flock, 1941a). Compsodes schwarzi — U.S.A. Occurs in the Southwest where it is confined to the desert and semidesert mountainous areas, rarely being found on the desert floor (Hebard, 1917). Taken in an ant nest in mountains of Arizona (Ball et al., 1942). Cyrtotria capucina Eastern Africa. — "Commonly met with under females being most frequent." debris, the apterous Thorn-bush country (Kevan and Chopard, 1954). Derocalymma lampyrina — Northern Kenya. Very abundant; both sexes under debris and thorn-bush country (Kevan and Chopard, 1954). desert-grass in SMITHSONIAN MISCELLANEOUS COLLECTIONS 28 Derocalymma — Northern Kenya. Taken and Chopard, 1954). in VOL. I4I porcellio upland grassland and bush (Kevan Deropeltis autraniana Northern Kenya. — In thorn-bush country (Kevan and Chopard, 1954). Deropeltis melanophila — Northern Kenya. "Very commonly found at the base of tufts of grass and other debris, the apterous female particularly in the latter situation" ; in upland grassland near forest ; in thorn-bush country (Kevan and Chopard, 1954). Deropeltis nigrita — Northern Kenya. Taken and Chopard, 1954). in upland grassland and bush (Kevan Eremoblatta subdiaphana —Apparently abundance in the extreme desert conditions of the southwestern United States (Hebard, 1917). Two small groups of males were observed in the midst of the sandy U.S.A. desert north of Yuma, found Ariz. in greatest ; these insects alternately flew and ran over the sand in the hot sun while headed in a southwesterly direction (Wheeler, 1911). Euandroblatta palpalis — Northern Kenya. In desert-grass and thorn-bush country (Kevan and Chopard, 1954). Heterogamodes rugosa Kenya. — "All Northern from desert grass and thorn bush (on It was stated (under discussion of Tivia fulva) that Heterogamodes females live more or less buried in the sand (Kevan and sand)." Chopard, 1954). Namablatta bitaeniata Southzvestcrn Africa. — Limited in distribution to the more arid por- tions, being peculiar to extreme desert conditions (Rehn, 1937). Nauphoeta punctipennis — Northern Kenya. In desert grass and thorn bush "probably the commonest of all the medium-sized cockroaches occurring in the area ; COCKROACHES BIOTIC ASSOCIATIONS OF under discussion, coming very freely —ROTH to light" & WILLIS 29 (Kevan and Chopard, 1954). Parcoblatta desertae U.S.A. — In the desert and semidesert mountainous areas of the on the desert floor (Hebard, 1917). Found under boulder on bare desert (Rehn and Hebard, 1909). Southwest; it is rarely found Polyphaga aegyptiaca Caucasus. —The wingless female was found buried in sand and dust (Burr, 1913). Turkmen S.S.R. —Although this species is secondarily encountered and courtyards, it is a very characteristic insect of the Trans-Caspian deserts the females are encountered fairly frequently as inhabitants of sand, where they run slowly over the surface, or dig themselves into the sand to continue their forward motion not far below the surface (Fausek, 1906). Uvarov (in Chopard, 1929b) indicated that females of this genus are found in various desert localities, in dwellings ; where vegetative debris occurs, but they are not particularly strictly attached to sandy terrain. Polyphaga indica Turkmen S.S.R. — This species prefers sandy soils where the and wingless females "swim" readily through the sand; they can also be found in the burrows of desert animals (Vlasov and Miram, 1937). nymphs, alate males, Polyphaga saussurei Turkmen S.S.R. — Its principal habitats are rodent burrows in loess dust and burrows of the desert turtle (Vlasov and Miram, 1937). Supella hottentotta Northern Kenya. —Taken grass and thorn-bush country at 1954). ". . . by dry river bed and in desertseveral stations (Kevan and Chopard, in bushes taken with Hght at night running on bark of a large acacia tree" (Rehn, 1947)- Symploce kevani — Northern Kenya. In desert grass and thorn-bush country (Kevan and Chopard, 1954). SMITHSONIAN MISCELLANEOUS COLLECTIONS 30 VOL. I4I Theganopteryx straminea Northern Kenya. —Taken at three stations in desert grass and thorn bush (Kevan and Chopard, 1954). Tivia brunnea Northern Kenya. among cover — In open sandy, riverine bush (scanty ground and doum palms) (Kevan and Chopard, acacia trees 1954). Tivia fulva Northern Kenya. — In desert grass and thorn bush; semidesert areas south of Sahara live the apterous ; distributed in females probably buried in sand (Kevan and Chopard, 1954). Tivia obscura Northern Kenya. — In desert grass and thorn bush (Kevan and Chopard, 1954). AQUATIC HABITATS The so-called aquatic or amphibious cockroaches are all members These forms are not nearly as aquatic as water beetles or aquatic Hemiptera, but in their relations to water they behave differently from nonamphibious cockroaches, which tend to avoid water except for drinking. There are apparently no special morphological characteristics that distinguish amphibious cockroaches ( Shelf ord, 1907, 1909a; Chopard, of the subfamily Epilamprinae (Chopard, 1938). 1938), although Takahashi (1926) listed several characters that he considered (i) made Opisthoplatia maculafa adapted for an aquatic life: Back of body easily wetted; (2) long hairs on underside of (3) terminal abdominal spiracles open into tubes that extend rearward (4) long hairs on ventral surfaces of cerci "pro- thorax trap air ; ; tect" terminal abdominal spiracles. Annandale (1906) also suggested that the position of the posterior abdominal spiracles, at the base of tubes that project rearward from beneath the seventh tergite, are an adaptation to an aquatic life. Chopard (1938) have pointed in many However, as Shelf ord (1907) and same feature may be observed out, this terrestrial cockroaches. The legs of amphibious cockroaches are similar to those of nonaquatic species and are not modified for swimming (Shelf ord, 1909a; Takahashi, 1926). Biological observations have been made on relatively few species, but representatives of at least six genera occur in quasi-aquatic habitats. Strictly speaking, these cockroaches live on land at the edges of streams or pools and spend relatively brief periods in the water. A BIOTIC ASSOCIATIONS OF COCKROACHES few species are found in water-filled bromeliads. & WILLIS 3I The behavior of the species of cockroaches in relation to their habitats known amphibious is —ROTH discussed below. AMPHIBIOUS COCKROACHES Audreia bromeliadarum Pananta. that —These had collected insects when disturbed would in the base of the bromeliad ; dive into the water they would disappear beneath the surface and remain submerged for some considerable time (Caudell, 19 14). Dryadoblatta scotti Trinidad. —This species was taken from the leaf bases of Tillandsia between the leaves and the amphibious (Scott, 191 2). less more or to be presumed insect was "This Glomeropitcairnia erectiflora: bromeliad the Subaquatic in the larger, in common scotti] is very [D. species large and handsome waterfilled, epiphytic bromeliads of the rain forest. Within these plants it is usually to be found, often in considerable numbers, just sp. at 3,100 feet; water had collected above the surface of the water or partly immersed in it. The cockroaches will descend rapidly into the water when alarmed and probably obtain their nourishment from the accumulated organic matter in the water. Floating material is probably taken and it seems less likely that they feed below the surface. They appear to be ovoviviparous." (Princis and Kevan, 1955.) Epilampra abdomen-nigrum Puerto Rico. nymphs swim —Abundant easily The in wet "malojillo" meadows. and remain under water for long periods, as do the adults (Sein, 1923; Wolcott, 1950). swimming nymph, captured in a dipper with mosquito Panatna. —A larvae in a lagoon of the Rio Chilibre, was kept under observations in an aquarium. If disturbed, the insect dived into the water from floating vegetation and swam rapidly below the surface for a minute or two. Finally becoming quiescent, the cockroach would then cling to submerged roots; twice it remained still for 15 minutes before climbing to the surface, where it remained for five or more minutes before emerging completely (Crowell, 1946). Epilampra annandalei Lozver Burma. — One male and three nymphs were collected in the Annandale who made the following observations: "The wingless specimens were under stones in a jungle stream and Dawna Hills by ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 32 VOL. I4I behaved just as the one I obtained in Chota, Nagpur, did [Annandale, 1906]. The winged specimen was under a stone at the edge of the stream, but swam readily. It did not seem so much at home in the water, however, and apparently could not, owing to the wings, raise the tip of its abdomen above the (Shelford, 1909a.) surface." Epilampra sp. — Siamese Malay States. Wingless females rested on floating logs from which they would dive into the water upon the least disturbance they remained under water for several minutes, then surfaced beneath the shelter of the log. In the jungle all females were taken either in the water or among matted roots on the sides of the stream. Winged males were seen rising from the surface of the water (Annandale, 1900). — Sarawak. All specimens were immature; they swam and dived well, but were soon drowned if prevented from rising to the surface to breathe. "When at rest the body of the cockroach is almost entirely submerged, the tip of the abdomen alone projecting above the surface of the water the abdomen moves gently up and down and every 30-40 seconds a bubble of air issues from the prothoracic spiracle on ; each side." India. —A (Shelford, 1901, 1916.) nymphal female, found a jungle stream at Chota in Nagpur, could swim with belly or back upward. When held under water it drowned in a few minutes. The tip of the abdomen was held out of water (Annandale, 1906). Shelford (1907) has suggested that the immature stages of terrestrial species of Epilampra may well be amphibious. This is an area that could profit by more field observations. Opisthoplatia maculata — Invariably found under or between rocks near mounnymph have similar habits. tain streams. on land, and when it goes into the water Normally the cockroach lives minutes. This cockroach rarely swims, it returns to land within a few Formosa. The but when it does, wingless adult and the it maintains its body in a horizontal position just below the surface of the water. Ordinarily, it walks on the river bottom or on water-covered rocks. This insect feeds on decayed leaves and, according to Shikano, it will eat human feces. (Takahashi, 1926.) This species has a large number of long hydrophobic hairs on the ventral sides of the thorax and anterior abdominal segments. the insect submerges, air is When trapped in these hairs. The thoracic and BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 33 one pair of abdominal spiracles open into the bubble of trapped However, the replenish below), it its insect apparently does inspires air while at the surface through While the insect part of breaks it Rhicnoda natatrix (see tracheal air supply, but, like dominal spiracles and expires air into the bubble submerged, the is away and air air. not use this plastron of air to its posterior ab- under the thorax. bubble increases in volume until (Takahashi, 1926.) floats to the surface. Opisthoplatia orientalis Formosa. — Lives on or in swampy ground (Takahashi, 1924). Rhicnoda natatrix Sarawak. — Immature cockroaches were found in sodden leaves at the edge of a pool, where they rested for hours at a time. the fore part of the body was always was in the water but the tip Generally of the abdomen When disturbed the insects dived into the water and stones on the bottom. Air is inspired through the posterior abdominal spiracles, when they projected above the water surface, and expired through the thoracic spiracles. In experiments in which the insects' abdomens were held immersed in water, in air. and hid under sticks with the thorax exposed, the insects died in 6 to 12 hours or less. (Shel- ford, 1907.) Stictolampra buqueti concinula Westsumha. — Found of Melolo River. under moist The nymphs amphibious mode of life fallen leaves on gravelly shore distinguished themselves through their and were often good swimmers (Princis, 1957a). Unidentified epilamprines Brazil. —These cockroaches were found under stones at the side of a rocky stream at Ouro Preto. When disturbed they ran down under the surface of the water and hid under stones at the bottom. When thrown on the water surface, they were helpless, and to get beneath the water surface they had to walk down some object. When they had penetrated the surface film they could mens kept in jars lived domens exposed to the swim freely. Speci- several days with only a portion of their abair. (Bristowe, 1925.) OUTDOOR HABITATS This category to the is a catchall for more circumscribed all cockroaches that are not limited habitats that have been previously con- : SMITHSONIAN MISCELLANEOUS COLLECTIONS 34 Some sidered. VOL. I4I cockroaches in this section select specific microhabitats Cryptocercus spp., which Hve exclusively in rotten logs and Neohlattella dryas, N. eurydice, and A^". grossbecki in bromeliads). (e.g., ; Others are found in a wide variety of habitats deropeltiformis and Parcohlatta spp.). known But some Ischnoptera (e.g., species are so little that their actual habitats are barely suggested in the collection data. Williams (1941) made an ecological study of the rain forest. He found Orthoptera (nearly Panama fied nymphal cockroaches) floor dead leaves, in the litter of fauna of the were unidentitwigs, and other all plant products in over 90 percent of the quadrats he examined. These insects represented about 0.25 percent of the total animal population. Delamare Deboutteville (1948) made a quantitative study of the soil that had accumulated between the animal population in suspended roots of forest epiphytes of the lower Ivory Coast. He analyzed 2 from an epiphyte located 45 meters above ground Parinarium, with these results: Horizon A. main branch of on a dm.^ samples of soil — Superficial zone of large rootlets, and 4 nids, beetles. H orison B. 2 cockroaches, 4 arachZone of fine rootlets, 6 cm. deep 6 cm. deep — : — 6 cockroaches and numerous other arthropods. Horizon C. Humid zone, 8 cm. deep: 7 cockroaches and numerous other arthropods. Plants, such as Palissota, The were also living in this very original biotype. species of cockroaches listed below have been found in the In jungle, forest, and following kinds of outdoor microhabitats: woodlands they have been found dead, and fallen trees ; in in rotten wood under bark ; decay cavities in trees ; burrowing of living, in living trees, shrubs, bushes, and low herbage on vines and epiphytic ferns; under signs on trees and stumps; in piles of logs and firewood; under dead leaves and debris; in and under decaying fruit on the ground. Cockroaches have been found between the leaves and under leaf sheaths of sugarcane, corn, and other grasses under dry fibers and fronds of coconut trees in hollow stems and bases of tree-fern fronds under bracts of banana on foliage of bark ; and in bromeliads ; ; ; ; blossoms and in bunches of bananas (p. 146). Cockroaches also inhabit abandoned cocoons and larval tents, wasp nests, ant nests, termite nests, bird nests, rat nests, and burrows of other rodents (pp. 2325, 310-319). Cockroaches have been found in rock crevices and under boards and other objects on ground under seaand other debris on beaches burrowing in soil and under clods of earth; in marshes and swamps; in dumps and rubbish heaps. under rocks weed, drift, ; ; ; ; BIOTIC ASSOCIATIONS OF COCKROACHES The above list —ROTH & WILLIS 35 does not exhaust the available outdoor microhabitats that cockroaches find suitable for their continued existence, but fairly representative. Although we have no measurements tiate this conclusion, we it is to substan- suggest that the microhabitats cited above have a more constant temperature and a relatively higher humidity than is provided by the surrounding macrohabitats. We would expect insects such as cockroaches, whose water balance is dependent on a continuous supply of fluid water or moist food, to seek moist environments or to avoid situations in which their transpiration might increase. Deviations, presumably brief, from this expected behavior must occur to account for the cockroaches that are found under relatively unfavorable environmental conditions. Despite the apparent preference for cryptic habitats, some cockroaches are found in hot sunlight (Ellipsidion spp. that many ; Tepper, 1893); Rehn (1945) has stated kinds are diurnal rather than nocturnal. may cockroaches between habitats Movement of be assumed to occur; but move- ment from an unfavorable environment to a more favorable one, following a shift in water balance, has not been observed in nature however, laboratory experiments suggest that the mechanism for mediating such behavior is present in some species of cockroaches (Gunn and Cosway, 1938; Roth and Obviously, addi- Willis, 1952a). needed on the bionomics of all species. Further conclusions based on current limited knowledge can only be speculative tional research is and possibly misleading. COCKROACHES FROM OUTDOOR HABITATS (Except Amphibious, and Cavernicolous Forms) Desert, Aglaopteryx absimilis Puerto Rico. leaves of — Living in wooden fence living between abandoned cocoons of Megalopyge rotten, Samanea saman and in ; krugii on bucare trees (Wolcott, 1950). Leeward Islands. — On coconut tree (Princis and Kevan, 1955). Aglaopteryx fades Puerto Rico. —As diaphana, in dead branch 10 feet above the ground on Mona Island (Hebard, 1917). In trunks of trees under bark and very often in abandoned cocoons of the "plumilla" (Sein, On wooden fence; empty cocoons of Megalopyge on trunk of Inga laurina; in larval tents of Tetralopha scabridclla on Inga vera (Wolcott, 1936). In large numbers in nests of the gray kingbird (Wolcott, 1923). rotten, in krugii on trunks of bucare trees, Erythrina glauca; 1950)- ;; SMITHSONIAN MISCELLANEOUS COLLECTIONS 36 VOL. I4I Aglaopteryx diaphana M^est Indies. — In Cuba, under corky bark of large tree in open Jamaica, under loose bark of shade trees and in bracts of banana blossoms in bromeliads and hollow bases of dead tree-fern fronds ; (Rehn and Hebard, 1927). Aglaopteryx Florida. — On gemma Long Key, under coquina boulder under loose, dry Hebard, 1912). fibers in heavy scrub near head of standing coconut palm (Rehn and Climbing on roots of red mangrove, Rhizophora loose bark on trunk of Exothea panicidata swamp under mangle, in ; under limestone boulder in keys scrub under signs on oaks, sweet gum, and longleaf pines in southeastern and southern States (Hebard, 1917). Infrequent in the shrub growth of the Sandin dense jungle hills habitat Texas. ; ; (Friauf, 1953). — In undergrowth of pine forest river; in Tillandsia sp. (Hebard, 1917). trees ; ; under sign on oak near Usually in hiding places on only once found under a stone on ground (Hebard, 1943a). Allacta similis Hazvaii. —Common hollow stems and under bark (Swezey and in Williams, 1932). Amazonina emarginata Trinidad. — On low herbage, on hibiscus at night, and in banana bunch (Princis and Kevan, 1955). Anaplecta asema Panama. —Under dead leaves in jungle (Hebard, 1920). Anaplecta decipiens Costa Rica. — In decayed leaves (Rehn, 1906). Anaplecta fallax Costa Rica. — Under stones on borders of Surubres River (Rehn, 1906). Anaplecta hemiscotia Panama. —Under rubbish at edge of jungle and in overgrowth of heavy vines on low bushes (Hebard, 1920). Anaplecta lateralis Panama. —Under drift on edge of coral-sand beach (Hebard, 1920). • BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 37 Arenivaga bolliana Texas. — In dense jungle brush of the river plain; on gravelly under debris and leaf mold under mesquite hillocks in scattered scrub trees ; in rat's nests, bush; inhabits litter ; sp. (Hebard, 1917). In dry earth under on ground and nests of rats (Hebard, 1943a) Neotoma Arenivaga floridensis — Male on ground under leaves of cabbage palmetto Females in sand under boards and debris along lake shore (Friauf in Cantrall, 1941). Infrequent on bare soil and ground under vegetation in the longleaf-pine flatwoods habitat (Friauf, 1953). In rodent burrow (Young, 1949). Florida. (Blatchley, 1920). Arenivaga grata Texas. forest; —Under stones in upper canyon from oak-manzanita under rocks ; in pine-oak forest along dry stream bed (Hebard, 1943a). Aristiger histrio Malaya. — Lives freely on bushes and flowers of Passiflora sp. (Karny, 1924). Aspiduchus boriquen Puerto numerous Rico. — "Apparently the species in suitable locations, [as deplanatus] is locally such as caves, rock crevices and the (Rehn and Hebard, 1927). shelter of large stones." Audreia bromeliadarum Panama. — Perfectly at home in bromeliads (see p. 31) (Caudell, 1914). Audreia jamaicana Jamaica. — In bromeliads; under dead wood in dense forest (Rehn and Hebard, 1927). Balta godefEroyi Australia. —Under bark (Hebard, 1943)Balta quadricaudata Australia. — From sugarcane (Hebard, 1943). Balta scripta Australia, Queensland. 1943)- — On leaves, grass, and sugarcane (Hebard, SMITHSONIAN MISCELLANEOUS COLLECTIONS 38 VOL. I4I Balta torresiana —From Australia. under bark, from sugarcane (Hebard, leaves, 1943)Balta verticalis Australia. —In leaves, from tree, from sugarcane (Hebard, 1943). Blaberus atropus —Female Trinidad. in rotting log (Princis and Kevan, 1955). Blaberus discoidalis Jamaica. found it — Under dead coconut petioles under stones in a field in in open spot. Gundlach Cuba (Rehn and Hebard, 1927). Blaberus giganteus Trinidad. — Nymph in rotten palm tree (Princis and Kevan, 1955)- Blaberus spp. — Only — taken in the forests of the Orinoco near the Venezuela. trunks of rotten trees at night (Doumerc in Blanchard, 1837). Panama. Among dead leaves and debris on floor of rain forest (E. C. Williams, Jr., 1941). Blatta lateralis U.S.S.R. It is and found —Found among rocks at 2,000 or in cultivated areas as well as in more meters elevation. mountainous landscapes in semideserts (Bei-Bienko, 1950). Blatta orientalis Great Britain. — One female nymph under bark of the ground (Burr, 1900). tree 10 feet Swarming within a rubbish heap in above Feb- ruary (Lucas, 1912). In refuse tip under old sacks and sheets of linoleum (Hallett in Lucas, 1922). Male under bark of oak far from houses (Donisthorpe, 1918). One adult female and nymph in prone dead elm 50 yards from house (Burr, 1937). An immature male at (Buck in Gardner, 1954). Four additional records of this species outdoors aw^ay from houses (Lucas, 1920). the roots of Ballota nigra Southern Crimea. —Under stones, dead leaves, and detritus in small copses of Quercus puhescens, Carpinus orientalis, Cornus mas, Pali- urus aculeatus, and Dictamnus fraxinella; 19 specimens, apparently breeding outdoors (Adelung, 1907). BIOTIC ASSOCIATIONS OF COCKROACHES North-central U.S. —ROTH & WILLIS 39 — Observations since 1950 indicate a marked in- crease in frequency and duration of infestations outdoors; observed in bare soil, vegetation, debris, alongside foundations in sodded areas, along sidewalks, and at edge of parking areas througout the year in some urban residential areas, the yards of whole blocks of houses were ; "alive" with this species on warm summer been found under stones, leaf debris, and nights ; in winter they have near structures (Shuyler, soil 1956). Blattella germanica —Under moist leaves woods (Lucas, 1849). — Under rubbish and on date palms (Herms, 1926). dump under loose material, very numerous Connecticut. — In in Algeria. California. city (Walden, 1922). Additional infestations of dumps by this species have been reported in New York (Felt, 1926, 1928) and New Jersey (Hansens, 1949, 1950). England. Swarming within a rubbish heap in February (Lucas, — Formosa. — Lives among 1924). North-central U.S. — Reported 1912). on the ground (Takahashi, fallen leaves in soil living outdoors near buildings under basementless buildings from early summer and to late fall (Shuyler, 1956). Blattella humbertiana (Chopard — Common among decaying vegetation and on and Chatterjee, 1937). pineapple sugarcane Formosa. — Normally found trees India. in fields, fields, and grasslands where it feeds on decayed leaves and other decayed vegetable matter and dead insects. It lies concealed among and under fallen leaves and clods of earth on or close to ground and never on the upper parts of plants, except pineapple where it is found among the leaves (Takahashi, 1940). Blattella vaga Arizona. is found in —Typically an inhabitant of plant debris, and clumps of earth ; and yards, it found under stones, irrigated fields fewer numbers on the dry desert. found It is in greatest numbers around decaying dates on ground (Flock, 1941a). Texas. — Beneath duff under athel trees of Rhodes grass (Riherd, 1953). ; rather abundant in clumps SMITHSONIAN MISCELLANEOUS COLLECTIONS 40 VOL. I4I Byrsotria cabrerae Cuba. — In sea-coast woods : "The species [this and Byrsotria fumigata] are ground-dwelling, hiding under stones and other shelter" (Rehn and Hebard, 1927). Byrsotria fumigata Cuba. —Ground inhabitant dwelling, hiding under stones, etc. ; also a cave (Rehn and Hebard, 1927). Cahita borero Brazil, Matto Grosso. — Beaten from from and from undergrowth in tree foliage in dry scrub, tree foliage at edge of dry riverine tangle, a dry forest area (Rehn, 1937a). Cahita nahua Honduras. —All beaten from foliage along roads or in thickets, during rainy season (Rehn, 1937a). Cariblatta antiguensis Virgin Islands, St. Croix. —Common under heaps of rubbish (Beatty, 1944). Trinidad. — On herbage below bananas ; all stages on Hibiscus at night; in grass at dusk; on low herbage under old coconut (Princis and Kevan, 1955). Cariblatta cuprea Jamaica. — In leaves on leaf mold in hillside forest (Hebard, 1916a). Cariblatta delicatula West Indies. — In debris petioles of coconut palms, hillside forest, grass in open, Cuba. in short San Domingo. In leaves on Under dead leaf mold in Jamaica (Hebard, 1916a). Cariblatta hylaea —Found at foot and on lower slopes of first ridges of from 75 to at least 800 feet above sea level, where vegetation ranged from abandoned banana patches overgrown with Heliconia and Cecropia and interspersed with forest trees, at the foot of the hills, to primeval lowland forest (ceibas, figs, palms, etc.) on the slopes. In the banana patches C. hylaea was found on hanging dead banana and Cecropia leaves on the slopes it was found on undergrowth foliage, hanging dead leaves, and in dead leaves on ground (Rehn, 1945a). Honduras. the Sierra Pija, ; BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 4I Cariblatta imitans Panama. —Among loose on leaves leaf mold in heavy jungle (Hebard, 1916a). Cariblatta insularis Jamaica. insects — One of the most frequently encountered orthopterous bromeliads on trees in (Hebard, 1916a, 1917; Rehn and Hebard, 1927). Cariblatta jamaicensis Jamaica. — In decaying herbage (Rehn and Hebard, 1927). Cariblatta landalei Jamaica. —All specimens taken from under drying bracts of banana blossoms (Rehn and Hebard, 1927). Cariblatta lutea lutea North Carolina. —Under pine straw on ground in woods (Brimley, 1908). — Under dead oak leaves under dead needles in woods in wire grass under refuse beaten from undergrowth in pine and oak woods (Rehn and Hebard, 1916). In undergrowth of shortleaf-pine, longleaf-pine, and oak woods; in heavy scrub in damp spot of sand dune area from high bushes, Ilex coriacea "The species [ = lucida] along inland swampy area (Hebard, 1916a). is in large part terrestrial, being usually found among dead leaves and Southeastern U.S. longleaf-pine ; ; ; ; ; on the ground. Occasional specimens are, however, sometimes beaten from bushes. Individuals are decidedly active and are usually litter to be found in the greatest numbers sandy situations" (Hebard, in 1917). Florida. —Throughout winter and spring they are frequent beneath sandy locations (Blatchley, 1920). Friauf (1953) found this species under debris, fallen leaves, leaf mold, or decaying wood in these habitats: Dry, ruderal grassland (infrequent), scrub (frequent), sandhills (domi- leaves and other debris on ground, nant), xeric hammock especially in dry, (infrequent), mesic hammock (dominant), pond margin (infrequent), longleaf-pine flatwoods (frequent), bayhead (occasional), low hammock (frequent), and alluvial hammock (occasional). In the shrub stratum in these habitats: quent), sandhills (dominant), and xeric herbaceous stratum in these habitats: hammock (dominant), and hammock Scrub (fre- (infrequent). In Sandhills (dominant), mesic black-pine flatwoods (infrequent). On SMITHSONIAN MISCELLANEOUS COLLECTIONS 42 VOL. I4I soil or bare sand under vegetation in these habitats: Sandhills (dominant), pond margin (infrequent), longleaf-pine flatwoods (frequent), and slash-pine flatwoods (frequent) (Friauf, 1953). bare Cariblatta lutea minima — Series of specimens captured on Long Key under dead palm on moist ground at edges of pools of brackish water. Specimens from Key West were in dry dead grass under boards (Rehn and Hebard, 1912). Nymphs frequent under bark on decaying pine logs in pine woods occasional in leaf mold in heavy junglelike scrub (Rehn and Hebard, 1914). In water-soaked leaves in heavy red-mangrove swamp (Hebard, 1915). Under dead petioles of coconut palm on sandy soil in grapefruit grove (Hebard, 1916a). Numerous at bases of tufts of coarse grass growing just back of sea beach (Blatchley, 1920). Friauf (1953) found this species in leaf Florida. petioles of coconut ; duff, leaf mold, debris, or decaying ruderal grassland frequent), mesic wood (occasional), scrub hammock : Dry, (in- (infrequent), pond margin (occasional), longleaf-pine flatwoods (occasional), and low On bare soil or bare in these habitats (infrequent), sandhills hammock (infrequent). sand under vegetation in these habitats : Longleaf- and slash-pine flatwoods (occasional). Dominant in the spartina marsh habitat in the grass stratum and duff around clumps. Frequent in the saw-grass marsh habitat in the grass stratum and, during the dry season, in decaying vegetation on the marsh floor. pine flatwoods (occasional) Cariblatta nebulicola —Adults Jamaica. in dead leaf litter alongside the trail was bathed in in dense and other trees the forest fog much of the time (Rehn and Hebard, 1927). forest of tree ferns, Podocarpus, Cyrilla, ; Cariblatta reticulosa Jamaica. — In leaves on Moderately numerous leaf mold in hillside forest in leaf litter in (Hebard, 19163). mangrove swamp; in decaying herbage (Rehn and Hebard, 1927). Cariblatta stenophrys Puerto Rico. —Between the leaves and under the leaf sheaths of corn (Sein, 1923; Wolcott, 1936). Cariblatta spp. West — The tropical species among the fallen leaves Indies. forest, living of this genus inhabit heavy resting on the leaf mold, in ; BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS epiphytic bromeliads, and in dead agaves (Hebard, 1916a; 43 Rehn and Hebard, 1927). Cariblattoides instigator — In siftings from under (Rehn and Hebard, 1927). Cuba. trees sea grapes, other shrubs, and low Cariblattoides suave Puerto Rico. — On dry limestone hills (Rehn and Hebard, 1927). Ceratinoptera picta Trinidad. —Under bark of old cacao tree (Princis and Kevan, 1955)Chorisoneura flavipennis Costa Rica. —Under stones on borders of Surubres River (Rehn, 1906). Chorisoneura formosella Jamaica. — Swept from huckleberry trees (Vaccinium meridionale) (Rehn and Hebard, 1927). Chorisoneura parishi Panama. —From jungle undergrowth (Hebard, 1920). Chorisoneura specilliger Panama. — In grass (Hebard, 1920). Chorisoneura texensis Florida. — "The almost impenetrable jungle on Key Largo was ex- amined, and in its depths the two specimens of this species were se- gumbo limbo, other trees and among which latter are to be found such cured by beating the lower branches of the lower bushes and shrubs, forms as Ocofea catesbyana [=Nectandra coriacca] and Citharexylum villosum" (Rehn and Hebard, 1912). In nests of webworm and beaten from bushes of bayberry, Myrica cerifera, along edge of pine woods (Rehn and Hebard, 1916). Beneath dead leaves tropical in oak woods and beaten from foliage of oak and bayberry (Blatchley, 1920). Infrequent in the tall shrub stratum of the xeric hammock habitat (Friauf, 1953). —The great majority of specimens were beaten from pine woods Southeastern and southern U.S. — In undergrowth Texas. foliage of bushes (Hebard, 1943a). in SMITHSONIAN MISCELLANEOUS COLLECTIONS 44 VOL. I4I beaten from shrubbery, from bayberry bushes, from lower branches of gumbo limbo and jungle, in other trees, from lower bushes and shrubs in and from low oaks on hills. In Texas, beaten from tall weeds opening in river-plain jungle scrub (Hebard, 1917). Chorisoneura translucida — Panama. In jungle vegetation, including vines covering low bushes (Hebard, 1920). Chromatonotus infuscatus — Trinidad. Males on low herbage under old cacao tree (Princis and Kevan, 1955). Chromatonotus notatus — Trinidad. Males in orchard on low herbage at night; females under refuse and in grass (Princis and Kevan, 1955). Comptolampra Malaya. —Often liturata found between dry foliage in the beakers of the epiphytic fern, Asplenium nidus, although the species lives mainly in bamboo bushes (Karny, 1924). Cryptocercus punctulatus North Carolina. —"They were never found except in parts of the where the decayed wood was soft, punky and wet" (Rehn and Hebard, 1910). Oregon. In fir logs where sap wood was soggy (Hebard, 1917). Virginia. In decaying chestnut and pine logs taken six times in chestnut and once in pine (Hebard, 1917). In rotten logs in deep ravines of moist woods (Davis, 1926). Appalachian Mountains, U.S. In southern Virginia and eastern Tennessee, it is usually quite abundant in well-forested areas at elevations from 3,000 to 5,000 feet; "sometimes even a majority of the dead logs on a mountain side have roaches in them" (Cleveland et al., 1934). This cockroach not only lives in rotten, dead logs but also in sound logs that have been down only a few years. In Virginia it is found more often in chestnut and hemlock. "It occurs fairly often in oak, and has been found in pine, spruce, and arbor vitae. There is little evidence that they ever leave the log and enter the ground" logs [chestnut] — — ; — . (Cleveland et al., . . 1934). Cryptocercus relictus Eastern Manchuria. and in rotten — In great numbers under rotting dead wood (Bei-Bienko, 1950). fallen trees BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 45 Cutilia soror Marquesas 1933a). Hawaii. Islands. — In soil — Males under stones and dead log (Hebard, about roots of pineapple (lUingworth, 1927). Often found about roots of grasses and weeds and other debris (Williams et al., 1931). Under stones and pineapple mulching paper (Fullaway and Krauss, 1945). Wake Island. Numerous, some from rotten logs. Found in bunch grass on Ocean Island (Bryan, 1926). — Cutilia spp. Australia. —Frequent sunset and run actively woods where they leave shelter soon after on ground or ascend shrubs and trees in quest of prey (Tepper, 1893). Dendroblatta sobrina Panama. — Colony on tree trunk ; on surface of trunk of fallen tree (Hebard, 1920). Diploptera punctata Hawaii. —"Crowds of these sometimes gather in cypress insects in various stages of development trees, in suitable chinks, in old flower- head sheaths of palms, etc., and even more or less openly on leafy twigs, in bunch grass, and the species is at times locally abundant behind the older leaf bases of sugar cane" (Williams Williams also lists the et al., 1931). Cryptomeria, food plants: following as algaroba, lime trees, ripening mangoes, papayas, and oranges. ever, Bianchi (personal communication, 1954) the above are the main How- doubted that any of dietary, because the largest populations he seen "were found in the fairly dry litter had of Star Jasmine (lasminum puhescens Willd.), well removed from any of the plants mentioned by Williams." Uahuka, —Beaten out of bracken Marquesas Islands. —Under bark (Hebard, Trinidad. — Raiatea, 1927). Society (Cheesman, Islands. 1933a). Dryadoblatta scotti Very common in water-filled, epiphytic bromeliads in the rain forest (see p. 31) (Princis and Kevan, 1955). Ectobius africanus — Belgian Congo. Females growth (Rehn, 1931). in forest margin and in forest under- ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 46 VOL. I4I Ectobius albicinctus South France. —Females and young beneath stones (Blair, 1922). Ectobius duskei U.S.S.R. — In the steppe belt, it is feather-grass steppes, where it is a very characteristic found member of in associations of typically steppe vegetation, with feather grasses at the head {Stipa lessingiana and others), and on rocky slopes; it occurs frequently in cultivated fields of young crops and also in young geological strata in sections with virgin The soil. populations of this steppe cockroach average 6 to 8 individuals per square meter from the middle to the end of July. By the end of summer most individuals were observed at the bases of straw stacks with a canopy, having their south sides sheltered. This is the only species of Ectobius adapted to a purely steppe (Bei-Bienko, 1950.) biocenose. Ectobius lapponicus Southeastern Europe. 1898). U.S.S.R. — Found northern part of its — Numerous under stones on Trebovic (Burr, in wooded communities and peat bogs range) ; (in males occur predominantly on herbaceous plants and bushes, but females hide under fallen leaves, moss, etc. (Bei-Bienko, 1950). It populated about 25 percent of the aspen trees an experimental plot, feeding in galleries in the bark of young in branches; there were 25 or more individuals per tree Bei-Bienko, 1950). Germany. —Abundant in woods ; in pine Stenohothrus vagans and Tettix kraussi. (Stark in woods in company with Numerous in low aspen Numerous in deciduous and coniferous forests on and underbrush under fallen leaves and moss on oaks (Zacher, 1917). In foliage of young oak on top of mountain (Ramme, 1923). Great Britain. Under moss and dry leaves, among woodland undergrowth, and, generally, on vegetation close to the ground occasional on bushes and trees (Lucas, 1920). Nymphs in heather in February and later adults among rushes fringing pond in July (Lucas, 1925). Nymphs and males on rushy vegetation; unusually abundant on low herbage in dried-up swamp (Lucas, 1930). bushes in forest. trees ; ; — ; Ectobius nicaeensis France. — In dry woods, on bushes, and (Chopard, 1947). at the base of trees ; BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 47 Ectobius pallidus moist places —Under stones England. — Very abundant on sand dunes and July (Buxton, 191 Germany. — In deciduous and coniferous Algeria. ; that in are shaded and covered with plants (Lucas, 1849). among bracken in 4). forests; at edge of forest, from bare woods and bushes numerous under leaves in oak woods and under moss (Zacher, 1917). In forest well lighted by the sun (Ranime, 1923). Massachusetts. Under loose lichens and bark on oak trees under boxes, baskets, paper, etc., near houses on Swiss chard (Flint, 1951). ; — ; ; On roofs of houses, in shrubbery (Gurney, 1953). summers in a fairly dense, this species for several dwellings, among fallen leaves We have collected wooded area near and climbing on the erect stems and undersides of the leaves of periwinkle. Oothecae were found on the ground under leaves and debris. Ectobius panzeri England. — Abundant on among sandhills along shoreline roots of Under dead seaweed and other rubbish a few ground yards from shore on that would be washed by the sea (Lucas, among marram grass (Buxton, 1914). On 1896). Nymphs found sandhills near coast and covered with marram grass often found on heather and low herbage under old bark and rotten wood on posts in decayed stump (Lucas, 1920). Swarming on Beta maritima and grass (Burr, 1908). ; ; Very common other plants in July (Lucas, 1920a). in all stages in August, being frequently found under stones (Lucas, 1925). Common on sand dunes especially under stems of dead marram grass. Viable oothecae found buried in sand (Brown, 1952). Germany. — In beech woods and in pine woods (Zacher, 1917). Ectobius semenovi Kazakhstan. around —Along the shores of the Syr-daria living willows it is found on and and on Populus euphratica; under loose bark of dying and dead trees (Bei-Bienko, 1950). Ectobius Sylvester U.S.S.R. — In wooded steppe zones ; probably only occurs in asso- ciation with forests (Bei-Bienko, 1950). SMITHSONIAN MISCELLANEOUS COLLECTIONS 48 VOL, I4I Ectobius tadzhicus Tadshikistan. —Great numbers at the roots of Eleagnus shrubs on the banks of reservoirs and frequently under the bark of old trees (Bei-Bienko, 1950). Ectobius vittiventer South France. — One male beneath stone (Blair, 1922). EUipsidion affine Amtralia. — From leaves, from scrub (Hebard, 1943). Collected in trees (Pope, 1953a). EUipsidion australe Australia. —On eucalyptus leaves, on wattle, under bark (Hebard, 1943). Collected in trees (Pope, 1953a). EUipsidion bicolor Australia. — In corn and from tree (Hebard, 1943). EUipsidion simulans Australia. —From sugarcane (Hebard, 1943). EUipsidion spp. Australia. —All stages are diurnal moving about the shrubs and small trees in bright sunlight on hottest foliage of summer days (Tepper, 1893). Epilampra abdomen-nigrum — In dried-up drain; among grass; debris under old under old leaves (Princis and Kevan, 1955). cacao damp lowlands (Sein, 1923). Under Puerto Rico. —Abundant in Trinidad. tree; in dead leaves in wet malojillo meadow (Wolcott, 1936). Shelf ord (1907) suggested that (p. 31). the genus species of may be aquatic, which other of immature stages This species is amphibious would place them in moist situations on the shores of rivers and other bodies of water. Epilampra azteca — Panama. Very scarce, under palm trees in decaying leaf mold and litter; one found under decaying bark of a log (Hebard, 1921a). Epilampra mona Mona Island, Puerto Rico. (Ramos, 1946). — One specimen under bark of dead tree BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 49 Epilampra tainana Cuba. —Under dead leaves on stream bank (Rehn and Hebard, 1927). Epilampra wheeleri Puerto Rico. —In siftings from high-altitude primeval forest (Rehn and Hebard, 1927). Epilampra Australia. —By day the spp. insects live under bark, stones, logs, dead soil. After sunset females vegetable debris, or buried in loose dust or wander in grass or ascend low objects (Tepper, 1893). Ergaula capensis Uganda. — In open bush and short grass (Princis, 1955). Eudromiella bicolorata Panama. —Under rubbish on edge of jungle (Hebard, 1920). Euphyllodromia Colombia. — In brushwood liturifera (Princis, 1946). Eurycotis bioUeyi Costa Rica. —Numbers of individuals were found in the large bromeliads of the temperate localities (Picado, 1913). Eurycotis decipiens Trinidad. — In old, rotten coconut stump (Princis and Kevan, 1955). Eurycotis dimidiata . — "This species was recorded from under stones by Gundlach" (Rehn and Hebard, 1927). Cuba. . . in the fields Eurycotis ferrum-equinum Cuba. —Under stones in woods (Rehn and Hebard, 1927). Eurycotis floridana Florida. — Moderately common under bark of dead pine stumps and Key West it fairly swarmed under coquina boulders in the woods (Rehn and Hebard, 1905). Many specimens under palmetto leaves on ground (Caudell, 1905). In pine woods under dry bark of dead logs; on Long Key in dry fibers at the base of the heads of logs ; at ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 50 coconut palms ; "at Key West, VOL. I4I a large colony was discovered among and several were captured upon turning over coquina boulders in the dense bush" (Rehn and Hebard, 191 2). Particularly numerous in tree cavities and under bark along the edge of hammock areas (Hebard, 191 5). Abundant between basal leaves of Tillandsia utriculata; beneath loose bark of logs and stumps in and beneath decaying palmetto trunks and leaves; under rubbish (Blatchley, 1920) On ground in heavy tangle after dark in decaying log of Sabal palmetto; in bromeliads common under debris and bark in jungle; under signs on Pinus carihaea; in almost every sheltered outdoor place (Hebard, 1917). It moves about at night and hides under bark of logs and in other recesses during the day where pines are present it almost invariably hides under bark of dead logs and stumps (Rehn and Hebard, 1914). Friauf (1953) found this species boards lying on dry grass in a field, ; . ; ; wood in these habitats Sandhills hammock (dominant), mesic hammock (freand low hammock (dominant) on tree trunks in sandhills (infrequent) and mesic hammock (frequent) infrequent in in leaf duff, leaf mold, or decaying : (infrequent), xeric quent), ; habitat ; saw-grass marsh habitat in the grass stratum and, during the dry season, in decaying vegetation on floor of marsh. Under the bark of logs and beneath logs in the woodpile habitat (Friauf, 1953). Eurycotis galeoides Cuba. —Under stones in deep woods (Rehn and Hebard, 1927). Eurycotis kevani Trinidad. —Under debris, trash, and vegetable refuse (Princis and Kevan, 1955). Eurycotis opaca Cuba. —In pine and palmetto region (Rehn and Hebard, 1927). Euthlastoblatta abortiva Texas. —Under dense tangle of bushy vegetation, palms, and vines near Rio Grande in leaves and dry litter on ground on dead petiole hanging from palm tree (Hebard, 1917). Under bark of dead hackberry abundant in dead leaves, dry litter, and rats' (Neotoma sp.) nests in heavy scrub (Hebard, 1943a). ; ; ; Graptoblatta notulata — On foliage in sun or concealed among dead leaves that between the fronds of tree ferns (Cheesman, 1927). Tahiti. collect BIOTIC ASSOCIATIONS OF COCKROACHES Hawaii.— Quite —ROTH active during the day, occurring in the wetter districts ; it is & WILLIS on sugarcane, 5I etc., also a household insect (Williams et al., 1931)Hemiblabera brunneri Puerto Rico.— Under bark of tamarind tree (Rehn and Hebard, 1927). Under the bark on a fence post (Wolcott, 1950). Henicotyle antillarum Dominica. —From rotting wood and wood soil (Rehn and Hebard, 1927). Holocompsa metallica — Dominican Republic. Along railroad through jungle and swamp (Rehn and Hebard, 1927). Hololampra bivittata — Canary Islands. Found in numbers among pine needles were in the majority, adults rare (Burr, 1911). ; nymphs Hololampra chavesi Azores. —Very in the hedges, particularly in brambles. common on the ground under stones, this species is exclusively dendricolous and is only captured by beating the bushes on which it abounds (Chopard, 1932). Contrary to most species of this genus, which live Hololampra maculata —Abundant in deciduous forest in grass and under fallen lichens and between fallen needles in under leaves in pine forests stones (Zacher, 1917). under forest; edge of coniferous Germany. ; ; Hololampra marginata Macedonia. —Usually giant thistles in found crawling on the flowers and stems of thistles in June (Burr, 1923). May; common on Hololampra Caucasus. — sp. Numerous beneath dry leaves in a garden (Burr, 1913). Hololeptoblatta sp. Seychelles. —Apparently bases (Scott, 1910, 1912). only inhabits Pandanus between the leaf SMITHSONIAN MISCELLANEOUS COLLECTIONS 52 VOL, I4I Homalopteryx laminata St. Vincent. 1927). Trinidad. not — In decaying leaves in forest (Rehn and Hebard, — In forest debris and debris under old cacao trees; uncommon under dry leaves it ; when feigns death it is disturbed (Princis and Kevan, 1955). Hormetica laevigata Brazil. —From crown of palm between leaf bases (Hancock, 1926). Ignabolivaria bilobata U.S.S.R. —Under rocks and on the edges of woods in the north and in the in the lowlands mountains in the south (Bei-Bienko, 1950). Ischnoptera deropeltiformis North Carolina. — Under pine straw on ground in woods (Brimley, 1908). —Under dead oak leaves under debris garden running pine and oak woods (Rehn and Hebard, 1916). hiding "a ground-frequenting, forest-loving Indiana. — Georgia. on ground in ; ; in insect, It is beneath cover or about the edges of deep woodland, more frequently damp in places, and rarely taken beneath bark, signs, or at lights" (Blatchley, 1920). — Tw^enty to 30 males found resting on heads of wild on successive evenings (Rau, 1947). Missouri. oats Texas. — It preferred damp, open woodlands (Hebard, 1943a). — Under stone in heavy deciduous under damp, dead leaves on edges of forests; under bark of pine log; in wire grass and sphagnum bordering stream thicket; in Eastern and southeastern U.S. forest; leaf mold and rubbish about pothole in pine woods, Pinus carihaea; under debris and leaf mold in hammock; under dead oak leaves in heavy deciduous forest (Hebard, 1917). Florida. fer —"This species is distinctly geophilous damp surroundings" (Rehn and Hebard, on very wet ground and appears 1912). to pre- Under boards mold in heav>', in debris and leaf and vines (Rehn and Hebard, 1914). Adults and numerous nymphs beneath weeds, grass, and other debris washed up on beach of Lake Okeechobee (Blatchley, 1920). Friauf (1953) found this species in leaf duff, leaf mold, and/or decaying wood on ground in these habitats: Dry, ruderal grassland (infrein everglades ; junglelike areas of trees, bushes, quent), scrub (infrequent), sandhills (occasional), xeric hammock BIOTIC ASSOCIATIONS OF COCKROACHES (frequent), mesic hammock (dominant), —ROTH & WILLIS shrubby, longleaf-pine 53 flat- woods (infrequent), bayhead (dominant), and low hammock (dominant). On open bare or bare sand under vegetation in these soil (infrequent), mesic hammock (dominant), moist, ruderal grassland (infrequent), pond margin (oc- Dry, habitats: grassland ruderal casional), longleaf-pine flatwoods (infrequent), slash-pine flatwoods (infrequent), and low hammock (dominant). Infrequent in the herbaceous stratum of these habitats: Dry, ruderal grassland, moist, ruderal grassland, and longleaf-pine flatwoods. Infrequent in the shrub stratum of the dry, ruderal grassland habitat. (Friauf, 1953.) Tennessee. —Taken in traps baited with cantaloupe in a parklike stand of oak, gum, hickory, and tulip trees in a creek bottom, and in a stand of oak on a dry ridge (Walker, 1957). panamae Ischnoptera Panama. —Under rubbish at edge of jungle and under drift on edge of coral-sand beach (Hebard, 1920). Ischnoptera podoces Jamaica. forest — In dead leaf litter along side trail through mountain (Rehn and Hebard, 1927). Ischnoptera rufa rufa Virgin Islands, St. Croix. — Common under rubbish and on shrub- bery at night (Beatty, 1944). Barbados. Occasionally found in cane — fields (Tucker, 1952). Puerto Rico, under stones in cultivated area, under debris on alkalie flat. In Jamaica, under dry petioles of coconut West palm Indies. — In in grassy area; under logs, logwood on docks, and litter on In Panama, under drift on edge of coral- limestone and near beach. sand beach; under rubbish at edge of jungle (Hebard, 1916c). Jamaica. —Under limbs and leaf litter in mangrove swamp (Rehn and Hebard, 1927). Lamproblatta albipalpus Panama. —Under drift on edge of coral-sand beach. Several under decayed banana stem (Hebard, 1920). Lamproblatta meridionalis Trinidad. —Under debris (Princis and Kevan, 1955). in forest and debris under old cacao trees . SMITHSONIAN MISCELLANEOUS COLLECTIONS 54 VOL. I4I Latiblattella chichimeca — Costa Rica. Very common in the bromeliads of all Costa Rica (Picado, 1913). Latiblattella lucifrons Arizona. —"Most commonly seen feeding on pollen and dead on the flower Yucca stalks of tains" (Ball et al., elata in June in the Santa Rita insects Moun- 1942). Latiblattella rehni —Widely woods (Pinus cariand Pinus caribaea (Hebard, Pinus clausa on haea) under signs beating Spanish moss; they pine tree; bark of dead 1917). Beneath in disturbed, but hide crevices or drop to when attempt flight seldom (Blatchley, ground 1920), Florida. distributed throughout pine ; Latiblattella zapoteca — Costa Rica. Under stones on borders of Surubres River (Rehn, 1906) Leucophaea maderae (Tucker, 1952). — In cane vegetation of royal palms, guava, Dominica. — In Barbados. fields etc. ; under loose bark and banana sheaths. In Jamaica, on logwood docks (Rehn and Hebard, 1927). Litopeltis biolleyi Costa Rica. — Under bark of tree in forest ; in epiphytic bromeliads (Rehn, 1928). Litopeltis bispinosa Panama Canal Zone. —About 80 specimens from rotting banana banana stem (Hebard, stalks at bases of leaves; boring in decaying 1920). Litopeltis deianira Costa Rica. — In tree wood on ground (Rehn, stump on edge of mountain Litopeltis — forest ; in dead 1928). musarum Shaken from dead banana leaves. Footnote to specific "In relation to the liking of species of this genus for bananas (Miisa) as shelter and possibly food" (Rehn, 1928). Costa Rica. name : BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 55 Lobolampra subaptera France. — Under stones and dead leaves, always rare (Chopard, 1947)Loboptera decipiens —All stages common beneath stones 1922). Under Maltese — Quite common open country under stones 1955). Dalmatia. — On seashores under rocks and seaweed up on shore France. (Blair, stones and dead leaves (Chopard, 1947). Islands. in (Valletta, cast (Bei-Bienko, 1950). Loboptera thaxteri Argentina. — Common in rubbish and leaf litter in small woodlot (Hebard, 1932), Lobopterella dimidiatipes Hawaii. —Abundant in wet districts, both in lowlands and to a con- siderable altitude in the forests, under trash, stones, boards, etc. (Wil- liams et al., 1931). Often it is found with nymphs of Periplaneta and Krauss, 1945). australasiae (Fullaway Lophoblatta arawaka Trinidad. —On grass, maize, and cut sugarcane fodder ; under vegetable and garden refuse; under old cacao (Princis and Kevan, 1955)Macropanesthia rhinocerus Australia. — Infrequently "They burrow October. seen during dry season from quite deeply, about two feet March to below the sur- They among the face of the sandy soil in stands of cypress pine {Callitris sp.). make a nest of dead leaves, grass roots, etc., frequently The young nymphs rarely appear above ground, but lowing rain the adults burrow to the surface, especially at night. pine roots. fol. . . found in the brigalow {Acacia harpophylla) scrub about 70 miles west of Rockhampton, Queensland, and on Fraser Island off the Coast of Queensland" (Henson in Day, 1950). This species is also Megaloblatta blaberoides Panama. — Under bark on tree —Under a dense Ecuador. (Campos R., 1926). (Hebard, 1920). dead leaves around base of tree pile of ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 56 Megamareta Australia. VOL. I4I verticalis —In sugarcane (Hebard, 1943). Methana canae Australia. —Under loose bark on dead upright tree (Pope, 1953a). Methana curvigera Australia. on wattle —Under loose bark on trees where trees and logs; many specimens in strong sunlight they hid in curled-up leaves oothecae attached to underside of loose bark and leaves (Pope, 1953a). Methana marginalis Australia. —Under loose bark of trees and logs (Pope, 1953a). Moluchia (?) dahli Chile. — Collected from lichens and mosses on tree trunks (Princis, 1952). Muzoa madida — Under dead wood in dense second-growth forest; in mat of hanging dead vegetation in dense forest; under leaves Costa Rica. thick in forest (Rehn, 1930). Nauclidas nigra St. Vincent. —Under rotten fruit (Rehn and Hebard, 1927). Nelipophygus ramsdeni Cuba. —Under rotten bark (Rehn and Hebard, 1927). Neoblattella detersa — Under dried leaves of coconut palm; in dry leaves under on beach under stones on coral rock in leaf mold under dense brush on hillside; under bracts of banana blossoms (Rehn and Hebard, 1927). Jamaica. acacia on hillside ; in debris ; ; Neoblattella dryas Jamaica. —In bases of dead tree-fern fronds; numerous liads; nearly all collected and Hebard, 1927). in brome- specimens were taken in these plants (Rehn ; BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 57 Neoblattella eurydice Jamaica. — Nearly all collected specimens taken in bromeliads (Relin and Hebard, 1927). Neoblattella grossbecki Jamaica. — In epiphytic bromeliads and hollow bases of fern fronds ; nearly all dead tree- collected specimens taken in bromeliads (Rehn and Hebard, 1927). Neoblattella proseipina Jamaica. — Under mens taken in bark of huckleberry; nearly all collected speci- bromeliads (Rehn and Hebard, 1927). Neoblattella semota — Jamaica. All specimens collected from under drying bracts of banana blossoms (Rehn and Hebard, 1927). Nesomylacris cubensis Cuba. — In dry region of palmettos and pines (Rehn and Hebard, 1927). Nesomylacris relica Jamaica. —Widely distributed from sea level to 5,700 feet elevation mountain forest; among dead leaves in heavy leaf mold under dense hillside scrub; under stones and in ground litter about banana trees; under bark of tree in dense ridge-type forest; in dead agave in scrub forest (Rehn and Hebard, 1927). in bromeliads in Nocticola bolivari Ethiopia. necessarily — Always found under stones or cement blocks, but not deeply buried in the ground (Chopard, 1950b). Nyctibora laevigata Jamaica. —In cracks in dead stump of mimosa; in bromeliads (Rehn and Hebard, 1927). Nyctibora lutzi Puerto Rico. in the highest — Possibly to be found most often mountains ; in rotten tree trunks found in rotten stump with termites, ants, and beetle grubs (Wolcott, 1950). Nyctibora obscura Trinidad. —Under pile of cornstalks (Princis and Kevan, 1955). SMITHSONIAN MISCELLANEOUS COLLECTIONS 58 VOL. I4I Nyctibora stygia — Haiti. Under loose dead bark (Rehn and Hebard, 1927). of mesquite tree, Oniscosoma Australia. 52 specimens spp. —The females bury themselves in loose soil or dust (Tep- per, 1893). Opisthoplatia orientalis — Formosa. On or in swampy ground or under rotten ground (Takahashi, 1924). trees on the Panchlora antillarum — Dominican Republic. In (Rehn and Hebard, 1927). cultivated grounds, palms, fruits, etc. Panchlora nivea — Panama. As Pycnosceloides aporus, in jungle under decaying banana stem in which were boring individuals of Litopeltis bispinosa (Hebard, 1920). Texas. Lives in foliage and in the green sheaths of plants (He- — Cuba. — On bard, 1943a). cane leaves according to Gundlach this genus lives under the loose bark of trees (Rehn and Hebard, 1927). Puerto Rico. In rotting trunks of coconut palms (Sein, 1923). Most specimens have been collected from the very rotten interior of coconut palms (Wolcott, 1950). Trinidad. On corn under old log flies readily to lights ( Princis and Kevan, 1955). ; — — ; ; Panchlora sagax Dominica. —In decaying stump in banana patch and in rotting wood. In Puerto Rico, in rotten coconut palm (Rehn and Hebard, 1927). Panesthia australis Australia. trees — In burrows under the thick bark of (Shaw, 1914). In loose detritus, fallen and rotting beneath clods of earth, and in fissures at foot of cliffs along the seashore beyond direct action of the waves (Tepper, 1893). Panesthia laevicollis Australia. —Under decayed logs in coastal scrub. the soft part of the log (Froggatt, 1906). It burrows into BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 59 Parcoblatta bolliana North Carolina. —Under pine straw on ground in pine woods (Brimley, 1908). Texas. —Under dry cow dung pine woods (Hebard, —Under of old boards (Hauke, 1949). in Nebraska. 1917). pile Parcoblatta caudelli North Carolina. Hebard, 1910). — From under the bark of dead trees (Rehn and —At night on shrubbery. In South Carolina, under sign (Hebard, 191 on a Tennessee. — In traps baited with cornmeal, cantaloupe, or Virginia. tree 7). fish in stand of oak on dry ridge, and in abandoned rocky facing slope (Walker, 1957). field on a south- Parcoblatta desertae Texas. —From mountains, arid, and semi-arid regions under small ; boulder on desert (Hebard, 1917). On ground in dry-creek bed through scrub oak, pine, and juniper forest (Hebard, 1943a). Parcoblatta divisa Eastern and southeastern U.S. —All specimens taken from under signs on red oaks and longleaf and shortleaf pines in Georgia and Virginia (Rehn and Hebard, 1916). Trapped in molasses-baited jar in oak forest in New Jersey under signs on red and white oaks, sweet gum, and other deciduous trees under signs on shortleaf and longleaf pines and pine stumps (Hebard, 1917). Widespread in south; ; eastern U.S. in habitats as diverse as dry pine lands, oak scrub, moist hammocks in northern Florida, and deep, cool ravines along Apala- chicola River (Hebard, 1943a). Parcoblatta fulvescens Eastern and southeastern U.S. —Trapped in molasses jars: in heavy, barrier-beach forest; in typical pine-barrens undergrowth; in on border of pine barheavy deciduous forest in heavy oak pine barrens with heavy, grassy undergrowth rens and on edge of swamp Found under ; in ; ; under dead leaves on edge of oak and shortleaf-pine woods under bark of pine log among dead leaves under live oaks under sign on Pinus woods. debris in dead, shortleaf-pine needles ; ; ; ; caribaea (Hebard, 1917). Georgia. —From under bark of pine log, among dead leaves under ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 60 live oaks, VOL. I4I and under leaves on edge of oak and shortleaf-pine woods (Rehn and Hebard, 1916). Florida. Very common among dead — leaves, under logs, beneath and wanders about at night in pinelands, hammock, turkey oak, and sand-scrub habitats (Hubbell and Goff, 1940). Beneath drift, cow dung, leaves, boards, bark of logs, and other debris, usually in open pine woods in sandy areas frequent at the base of thistle leaves (Blatchley, 1920). Friauf (1953) found this species in leaf duff, debris, or decaying wood in these habitats: Scrub (dominant), sand- loose bark, ; (dominant), xeric hills leaf-pine flatwoods hammock (dominant), mesic hammock, longhammock (infrequent), and (infrequent), low hammock (infrequent). In the shrub stratum in these habiScrub (dominant), sandhills (dominant), xeric hammock (dominant), and longleaf-pine flatwoods (infrequent). In the herbaceous stratum of the longleaf-pine flatwoods habitat, and under bark and beneath logs in the woodpile habitat. alluvial tats: Parcoblatta lata — Southeastern and southern U.S. Under bark of pine logs and stumps in sweet-gum logs and stumps moderately numerous under bark of dead shortleaf pines; under bark of longleaf-pine stumps; under signs on red oak and longleaf pines; in dead oak. In Texas, under bark of pine stumps (Hebard, 1917). ; ; North Carolina. —All stages under loose bark of prostrate and upright, and stumps. dead pines, both "It seems to prefer the space under the bark to be rather damp" (Brimley, 1908). Under bark of dead pine trees (Rehn and Hebard, 1910). duff and decayed wood of low ham— Infrequent Indiana. — Beneath rocks on and tops of high limestone glades where cedar abounds (Blatchley, 1920). Missouri. — In stratum of oak-hickory forest (Dowdy, 1951). Florida. mock in leaf habitat (Friauf, 1953). sides hills, in leaf Earlier, Dowdy (1947) reported finding numerous immature Pseudosp.] in soil and leaf strata of oak- mopinae [presumably Parcoblatta hickory forest. Texas. and in — Captured in molasses-baited traps in low, wet, dry woodlot on hillside oak woods (Hebard, 1943a). Parcoblatta pensylvanica oak and in chestnut forests, — Trapped in molasses-baited and on knoll with high deciduous Eastern and southeastern U.S. in jars trees. BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 6l Found in oak and pine woods, under bark of decaying chestnut log and dead chestnut stump, and under signs on trees including oaks (Hebard, 1917). North Carolina. In all stages under loose bark of upright, dead pines, when the space under the bark was dry (Brimley, 1908). Under signs on trees (white Virginia, North and South Carolina. and red oaks) under bark of dead shortleaf-pine and sweet-gum logs and stumps (Rehn and Hebard, 1916). Indiana. Beneath bark of logs and stumps empty oothecae com- — — ; — mon ley, ; beneath loose bark of logs, especially shellbark hickory (Blatch- Under 1920). Illinois. — In loose bark on logs in January (Blatchley, 1895). pine forest associes, in black oak forest on sand, in oak-hickory forest on clay, and in climax forest it evidently moved into the pine associes nightly, great numbers of oothecae were found ; under bark of pine logs, where, in October and November, hibernating nymphs were found (Strohecker, 1937). In nests of Vespula maciilata (Balduf, 1936; McClure, 1936). Missouri. —Usually in hollow trees, under loose bark, in woodpiles, and in cracks in rural buildings (Rau, 1940). — Common oak-dune and beech-maple forests, under and fallen logs, and under debris on forest floor (Hubbell, 1922). "A characteristic inhabitant of the low shrubterrestrial and probably the terrestrial-hypogeic stratum." It occurred throughout the upland forests groups were found established in and under logs 100 to 200 feet from the nearest forest (Cantrall, 1943). Michigan. in loose bark on dead trees ; Ontario. it —Very abundant in rocky, sparsely-wooded country, where occurred in rotten logs and under loose bark on rocky island in lake ; on tree trunk at night (Walker, 19 12). Parcoblatta uhleriana North Carolina. 1908). —Under pine straw on ground Under bark in woods (Brimley, (Rehn of dead trees; 92 males attracted to lights and Hebard, 1910). Virginia. —Resting on woods foliage; at night on road (Rehn and Hebard, 19 16). Eastern and southeastern U.S. Trapped in molasses-baited jars: in oak and pine woods, in heavy barrier-beach forest, in both scant and typical undergrowth on pine barrens, in heavy grassy under- — growth on pine barrens, on border of pine barrens, on edge of swamp, in heavy deciduous forest, in heavy oak woods, in upland oak and chestnut forest, in chestnut forest, in forested ravine, and on ridge : SMITHSONIAN MISCELLANEOUS COLLECTIONS 62 with heavy oak, chestnut, and maple VOL. I4I Found under damp forest. leaves on edge of forest, under bark of decayed chestnut log, inside decaying chestnut log with Cryptocercus pimctulatus, under palmetto roots, under bark of pine stump, and 1917). Tennessee. gum-oak in dry leaves under live oaks (Hebard, — In traps baited with cornmeal or cantaloupe forest in a mesic valley, and in a stand of (Walker, 1957). Indiana. Beneath cover on slopes of high wooded — essentially a forest-loving species ; maple- in oak on a dry ridge hills. "This is usually occurring beneath leaves and other debris on or along the borders of heavy hardwood timber." (Blatchley, 1920.) — In oak-hickory forest on clay and Michigan. — In oak-dune woods (Hubbell, Illinois. in climax forest (Stro- hecker, 1937). woodlands, where it Restricted to 1922). inhabited piles of moist dead leaves and rotten logs in oak -hickory forest (Cantrall, 1943). U.S.A. —This at night to species, P. uhleriana, and P. virginica were attracted honeydew secreted by aphids on Pyrus sp. (Davis, 1918). Parcoblatta virginica boards, and other — Females under loose beneath loose bark (Morse, 1920). North Carolina. —Under debris dead shortleaf-pine needles (Rehn and Hebard, 191 6). Florida. — Infrequent the shrub stratum of the scrub Nezv England. debris on ground stones, ; in habitat. in This was the only habitat of 25 studied (Friauf, 1953). Eastern and southeastern U.S. in which this species was found — Trapped in molasses-baited jars and oak woods, in pine barrens, in pine woods with heavy grass undergrowth, in oak forest, in heavily forested ravine, on rocky slope with few deciduous trees, on knoll with high deciduous trees, in lofty chestnut forest, and in heavy low chestnut and oak forest on high ridge under bark of decaying chestnut log and stump under stones in chestnut forest; under bark of pine stumps (Hebard, 191 7). Frequents borders of open woods and fields; under Indiana. debris, loose bark, and half-buried logs (Blatchley, 1920). In black-oak forest on sand, in oak-hickory forest on clay, Illinois. in pine ; ; — — (Strohecker, 1937). climax and oak-dune and beech-maple Michigan. — Common in forest in forests ; under loose bark on dead trees and fallen logs and under debris on forest floor (Hubbell, 1922). Restricted to woodlands, where it inhabited BIOTIC ASSOCIATIONS OF COCKROACHES piles of trail, —ROTH & WILLIS 63 moist dead leaves and rotten logs in oak-hickory forest (Can- 1943). T^jira^.— Captured in molasses traps in moist woods of maple, oak, and pine with much undergrowth and a heavy layer of duff in open, ; rather dry woodlot of Spanish oak and other trees and in low wet woods of willow and oak along creek (Hebard, 1943a). ; Parcoblatta zebra Indiana. —Beneath log in cypress swamp (Blatchley, 1920). — Louisiana and Mississippi. In decay cavity sign on shortleaf pine (Hebard, 1917). in sweet gum; under Parcoblatta spp. Alabama.— In the dry wall of a sweet-gum stump together with serropalpid and tenebrionid beetles (Snow, 1958). Ohio. Oothecae under loose bark of fallen trees, where as — many as 184 oothecae were found within a few feet of each other; others found under boards and in piles of firewood (Edmunds, 1952). Pelmatosilpha coriacea — Mona Island, under bark of dead trees and under (Ramos, 1946). Under bark of Sideroxylon foetidissiguava leaves (Wolcott, mum 1941). Common along the coast and in mountains, Puerto Rico. "Very much mum at home" under the loose bark of Sideroxylon foetidissi- (Wolcott, 1950). Pelmatosilpha kevani Trinidad. —Under debris in bush (Princis and Kevan, 1955). Pelmatosilpha purpurascena Dominica. — In decaying logs in forest (Rehn and Hebard, 1927). Periplaneta americana Bermuda. —Among and under decaying — Nocturnal, coming out debris, just above high-tide line (Verrill, 1902). Johnson Island. about Tribulus blossoms. at night in great numbers Under timbers on French Frigate Shoals (Bryan, 1926). United States.—AWeywdiys and yards may be overrun during the summer; adults and hundreds of nymphs found in decaying maple trees along residential street (Gould and Deay, 1938, 1940). Around SMITHSONIAN MISCELLANEOUS COLLECTIONS 64 VOL, I4I fill was burning internally (Davis, 1927), n palm trees along the gulf coast of Texas, where they around street lights at night (Zimmern in Gould and Deay, fumaroles where a railroad Common often fly 1940). Periplaneta australasiae —Very abundant under stones (Rehn, 1910). —Under bark of dead and under bases of leaves of coconut palms (Rehn and Hebard, 1927). Virgin Croix. — Common sugarcane and Bermuda. Jamaica. tree Islands, St. in in fields woodlands (Beatty, 1944). Florida. Juveniles under bark of dead logs of Pinus caribaea (Hebard, 1915). Frequently found under signs on trees near borders of towns; under bases of dead petioles of cabbage palmetto (Hebard, 1917). Beneath logs, burlap bags, and other cover in old orange — orchards (Blatchley, 1920). Marquesas 1935)- Islands. —Under coconut fronds and grass (Hebard, — Nihoa Island. Nymphs only, on Sida, Pritchardia, bunch and about camp (Bryan, 1926). grass, Periplaneta brunnea on oaks (Rehn and Hebard, 1916). —Under —Beneath bark of stump (Blatchley, 1920). Georgia. Florida. signs Periplaneta fuliginosa Southeastern and southern U.S. —"This tered out of doors, in or near towns. Over species its is range usually encounit is frequently found under signs on trees" (Hebard, 1917). Phidon (?) dubius Chile. — Collected from mosses and lichens on tree trunks (Princis, 1952). Phoraspis spp. Brazil and Guiana. —In grasslands, plantations of maize, sugarcane, and other plants on the borders of forests the cockroaches were always found between the leaves which form the branches of the ; plants (Doumerc in Blanchard, 1837). Phyllodromica brevipennis — Asia Minor and western Europe. On ground among grasses under moss and brushwood in mountain meadows (Bei-Bienko, 1950). ; BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 6$ Phyllodromica graeca pine forest mixed with deciduous USS.R., western Georgia.— In trees (Bei-Bienko, 1950). Phyllodromica irinae of "tugas" under halfoverhang the ground (Bei-Bienko, USS.R., Turan Lozuland.—Along margins fallen bushes of Salsola kali that 1950)Phyllodromica maculata Central Europe and western US.S.R.— On the edges of forests of fallen the central-European type that are lighted by the sun; under leaves; on bushes and conifers (Bei-Bienko, 1950). Phyllodromica meglerei U.S.S.R.— Among under mown fallen leaves under bushes; on oak branches; hay (Bei-Bienko, 1950). Phyllodromica polita Caucasus.—Under fallen leaves on slopes of mountains covered by forest or brushwood (Bei-Bienko, 1950). Phyllodromica pygmaea the sands of U.S.S.R.— In Un-dzhal-kum and Zhety-konur it is found in the dense turf of Aristida pennata (Bei-Bienko, 1950). Phyllodromica tartara Central Asia. fruit orchards apple worm — In lowlands and in mountains up to 2,500 meters in to trees to combat lesser ; under trap rings fastened (Bei-Bienko, 1950). Phyllodromica tartara nigrescens — Southern Uzbekistan. Under bark of Juniperus sp., under stones and on flowers of Scorsonera acanthoclada (Bei-Bienko, 1950). Platyzosteria castanea Australia. —Under loose wood or bark (Shaw, 1914)- Platyzosteria novae seelandiae Nezv Zealand. 1904). — Swarms under loose dry bark and logs (Walker, SMITHSONIAN MISCELLANEOUS COLLECTIONS d^ VOL. I4I Plectoptera dominicae Dominica. — On moss-covered lime Plectoptera are all foliage "The species of the genus and flower frequenters, generally secured trees. by beating low arborescent vegetation, or are attracted (Rehn and Hebard, 1927). light" to Plectoptera dorsalis Puerto Rico. — In caladium, flowers of Iponioea tiliasea in trees between grass, weeds, coffee, and bananas (Rehn and Hebard, 1927). ". . . ; in living leaves, or in 'butterfly-nests' of Tetralopha scabri- Inga vera, or of Pilocrocis secernalis in the leaves of 'capa bianco' (Petitia domingensis) in the mountains. Along the coast they have been found under the bracts of cotton squares or bolls, della in leaves of and under the leaf-sheaths of sugar cane, fruit, in curled-up leaves of grape- or in the dry flower clusters of 'espino rubial' (Zanthoxylum caribaeum)." These observations apply and P. rhabdota (Wolcott, 1950). also to Plectoptera infulata Plectoptera floridana — Florida. On fringe of tall bushes at edge of mangrove swamp (Rehn and Hebard, 191 4). Rehn and Hebard (1927) stated that on the Keys it frequented dry scrubby vegetation, particularly Ilex cassine. Plectoptera infulata Puerto Rico. — See Wolcott's (1950) comments under Plectoptera dorsalis above. Plectoptera lacerna Cuba. — In grasses, sedges, etc., about a waterhole ; on grass pines and oak (Rehn and Hebard, 1927). Plectoptera perscita Dominica. — On moss-covered lime trees (Rehn and Hebard, 1927). Plectoptera porcellana Cuba. —Taken on flowers of "Ji^caro" (Gundlach in Rehn and Hebard, 1927). Plectoptera Jamaica. — In pygmaea relatively dense forest foliage; in and Hebard, 1927). shrubbery (Rehn BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 67 Plectoptera rhabdota — Puerto Rico. In mixed vegetation; on grapefruit tree and guava {Psidimn guajava) on bushes and shrubs (Rehn and Hebard, 1927). On coffee trees: on Spondias; on sugarcane; in caterpillar nests of Tetralopha scabridella on Inga vera; in old cotton bolls on grapefruit (Wolcott, 1936). See also Wolcott's (1950) comments under Plec; ; toptera dorsalis. Plectoptera vermiculata Cuba. — On pine in palmetto region (Rehn and Hebard, 1927). Polyphaga aegyptiaca — Nymphal females under decaying Algeria. November (Lucas, 1849). leaves at the end of — In burrows in argillaceous cliffs along ravines. Transcaucasia. Females often covered by attached clay particles, an indication, ac- cording to Bei-Bienko, that this species is ecologically connected to compact clay soils or at least does not avoid them (Bei-Bienko, 1950). See also the section on desert habitats (p. 29). Polyphaga saussurei South-central Asia. —Occupies compact clay soils; distributed in found near dwellings, drier regions than P. aegyptiaca; frequently in yards, stables, and houses (Bei-Bienko, 1950). Polyzosteria limbata Australia. ning — Common, usually "resting among the foliage or sun- on a fence or stumps, seldom or never hiding under bark or most of the species" (Froggatt, 1906). itself logs like Poroblatta spp. Tropical America. —"The species of Poroblatta apparently live as borers in stumps and logs in a manner similar to those of Cryptoccrcus Scudder in the United States" (Gurney, 1937). Pseudomops septentrionalis — In dead-brush pile not scarce in heavy weeds, sunflowers, openings of river-plain jungle scrub (Hebard, 1917). It lives Texas. etc., in ; largely in herbage (Hebard, 1943a). Pycnoscelus surinamensis Florida. —Under stones and rubbish boulders in woods at ; very abundant under coquina 1905). "This Key West (Rehn and Hebard, ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 68 species common under is ground . . the petioles of a and other debris on the Long Key in the dry fibres at the base of coconut palm" (Rehn and Hebard, 1912). At Musa also . planks, stones, VOL. I4I found at sand (Hebard, 1915). In fallen leaves and decaying wood in xeric and mesic hammock habitats (Friauf, 1953). Hawaii. The soil swarmed with young of various stages during the Isle, found burrowing in — summer In (Illingworth, 1915). soil about roots of pineapple under feeding on pineapple roots (Illingworth, 1927, 1929). mulching paper Fakarava, Tuainoto Archipelago. Numerous among dead leaves in tree holes (Cheesman, 1927). West Indies. Under decayed stalks of sugarcane and in siftings from mangrove swamps, Cuba. Under manure, bases of leaves of coconut palm, litter, logs, and stones on coral rock and in bromeliads, Jamaica. Under wood, tiles, and boards in stable yards immature in; — — ; dividuals bored into the soil, Puerto Rico. Barbados. —Frequents cane — "Altho primarily fields Puerto Rico. among dry stones on Mona Island, (Rehn and Hebard, 1927.) (Tucker, 1952). a xerophytic species under dry cow dung at : collected Boqueron, and under boxes at Guanica, it is reasonably common in the more humid parts of Puerto Rico" (Wolcott, 1950). Common under rubbish frequently seen Virgin Islands, St. Croix. feeding on chicken feces around chicken roosts (Beatty, 1944). By feeding on chicken feces it may become the vector of the chicken eyeworm, Oxyspimra mansoni, as described in the references cited on — ; page 204. Egypt. —Large numbers were found moist (Chakour, 1942). Germany. — Under greenhouse conditions the manure in soil at the site of a pile depth to which P. was determined; 21 dug down to a depth of 8 to 10 cm., 3 dug down 10 to 12 cm., but only one dug 13 cm. below the surface. Often the tubes in the soil ended in a chamber which the cockroach might not leave for several days nymphs molted in such chambers and females bore their young there (Roeser, 1940). surinamensis penetrated the soil ; Rhytidometopum Trinidad. — Male on low herbage dissimile in orchard at night ; under sacking on Hibiscus at night (Princis and Kevan, 1955). Riatia orientis Trinidad. — Numerous specimens of both sexes at night on roadside Hibiscus rosa-sinensis or low herbage in orchard (Princis and Kevan, 1955)- . BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 69 Simblerastes jamaicanus Jamaica. —Numerous in fragmentary debris of an abandoned mite nest on ground in the dry Liguanea Plain taken under a stone in a field of short grass ; ter- a specimen was also (Rehn and Hebard, 1927) Styphon bakeri Costa Rica. ties in —Among humus and rubble in crevices and large cavi- rocks of the Tertiary limestone rim and the metamorphosed and igneous rocks of the interior of the islands (Baker in Rehn, 1930) Supella supellectilium — Virgin Islands, St. Croix. Under rubbish heaps in sugarcane straw (Beatty, 1944). "A cosmotropical species which occurs both out of doors Africa. ; — and as a household pest in many warmer parts of the world. It is apparently endemic to non- forested areas in much of Africa north of the Equator." (Kevan and Chopard, 1954.) Symploce Puerto Rico. —Under flagellata low trees on and dead leaves in —Under stones and rubbish (Illingworth, 1915). —Under rubbish and on shrubbery at hillside thicket of sea grape (Hebard, 1916c), Symploce hospes Hawaii. Virgin Islands, St. Croix. night (Beatty, 1944). Symploce jamaicana — In dead leaves under acacia and other shrubs in desert under log and rubbish in open on limestone sand near beach (Hebard, 1916c). Very common in short dry grass in roadside gutter at night, often clustered together under beach trash in stony wash of Hope River (Rehn and Hebard, 1927). Jamaica. tract; ; Symploce Virgin Islands, St. Croix. ruficollis —Under rubbish and on shrubbery at night (Beatty, 1944). Puerto Rico. In siftings from sea-grape thicket on sandy soil — (Rehn and Hebard, 1927). Often cane (Wolcott, 1950). living under leaf -sheaths of sugar- ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 70 VOL. I4I Tartaroblatta karatavica Asia, Kara-tau Mountains. — Many hundreds of individuals found only under stones on moist earth and not where ground seemed dry found on very stony slopes with sparse vegetation, often with undergrowth present (Bei-Bienko, 1950). STRUCTURAL HABITATS we In this category man habited by A nonexhaustive mess restaurants, drug stores, include or not, that list all man-made may become structures, department privies, sewers, in- of such structures would include dwellings, halls, barracks, groceries, markets, bakeries, dairies, stores, hotels, hospitals, warehouses, packing houses, animal houses, breweries, factories, whether infested with cockroaches. sewage treatment dwellings are only one of the plants, ships, aircraft, etc. many mills, incinerators, Although kinds of structures that are colonized by cockroaches, the several species that have adopted this mode of life are generally referred to as domiciliary cockroaches. This term is adequate only we remember if that these cockroaches are not restricted to domiciles but are pests in other structures as well. Associations between man and sibly started as casually as the certain species of cockroaches posshort-lived association that (1953) observed when he discovered Beebe three cockroaches in the newly couch of an orangutan. Obviously, when man came down from the trees, his fellow travelers found his cave dwellings and other abodes particularly favorable habitats. From such primitive beginbuilt nings, domiciliary cockroaches have spread into every kind of structure that man has since devised. We predict that when man develops a suitable vehicle, cockroaches will someday accompany him into space. Yet despite the apparent predilection of certain species of cockroaches for man, man is only incidental to these associations. Only the shelter and food that man unwittingly provides for these unwelcome guests attract cockroaches to him man's physical presence is unnecessary. ; Most, if not all, of the common domiciliary cockroaches apparently originated in the Tropics or sub-Tropics from whence they have spread, through normal commercial channels, into most of the in- habited world. At least eight domiciliary cockroaches originated in Blatta orientalis, Blattella germanica, LeiiAfrica (Rehn, 1945) cophaea maderae, Naiiphoeta cinerea, Oxyhaloa huprestoides, Peri: planeta americana, P. anstralasiae , and Supella supellectilium; and, perhaps, Periplaneta brunnea as well probably of Indo-Malayan origin ; ; Neostylopyga rhombifolia was Pycnoscelus surinatnensis was of BIOTIC ASSOCIATIONS OF oriental origin; COCKROACHES —ROTH & WILLIS and Leurolestes pallidus was endemic in the yi West Indies (Rehn, 1945). Princis (1954a) rejected Africa as the original home of Blatta orientalis and advanced reasons for placing its origin in Central Asia. Several domiciliary species have become well established in temperate zones and some even Bei-Bienko (1950) Usted in the Arctic. the following 10 species as sinanthropes in the Palearctic zone: Blatta lateralis, B. orientalis, Blattella germanica, Leucophaea maderae, Peri- planeta americana, P. australasiae Polyphaga saussurei, Pycnoscelus , surinamensis, and Supella siipellectiliuin. In the warmer parts of the temperate regions, as in their native Tropics, certain domiciliaryIn the species breed outdoors as well as indoors. temperate less extensions of their ranges most domiciliary species are nearly always found indoors. In regions with low winter temperatures these cockroaches do not survive in unheated structures ; but in heated buildings for example, has been able to withstand the Blattella germanica, rigorous climate of Alaska, where it has caused severe infestations (Chamberlin, 1949). The limiting factors that determine whether man-made structures will provide suitable habitats for cockroaches are favorable tempera- and food. The range of temperatures own comfort and protection fosters the rapid ture and availability of water that man provides for his Gunn (1934, 1935) has demonstrated that the preferred temperature range (zone of indifference) of Blatta orientalis is 20-29° C. The upper limit of the preincrease of cockroach populations indoors. ferred temperature of Blattella germanica and Periplaneta atnericana 33° C. (Gunn, 1935). The lower limits of temperature tolerance were not sharply defined in Gunn's work. However, less than optimum is temperatures, lethal. if they The 24-hour last for only short periods, are not necessarily mortality for P. americana that had been held for one hour at 0° C. was only 2±2 percent (Knipling and Sullivan, 1957). Gunn (1934) observed that Blatta orientalis would not settle at temperatures temperatures above 33° C. and would react violently against higher 39° C.) by running away; thus the thermotactic (e.g., behavior of cockroaches might be presumed to bring them into favorable environments within structures. Thermal death points have been Gunn and Notley (1936). determined for the above three species by It is common knowledge among those who rear cockroaches experi- mentally that, unless the water content of the food is essential in the insects' dietary. Ten is high, fluid water species of domiciliary cock- roaches have been shown to be unable to survive as long on dry food alone as they could on food and water at 36-40 percent relative SMITHSONIAN MISCELLANEOUS COLLECTIONS y2 VOL. I4I humidity (Willis and Lewis, 1957). Blatta orientalis, when in a state or normal water balance, usually spent more time in the drier part of a humidity gradient but desiccated insects tended to become hygro- ; We (Gunn and Cosway, 1938). positive ciliary species behave similarly. If water presume that other domiis available nearby, it may be presumed that partially desiccated cockroaches could locate a source through the mediation of a humidity sense. Hygroreceptors have been demonstrated on the antennae of Blattella germanica (Roth and Willis, 1952a) and suggested for Blatta Cosway, 1938). Drinking water wash on cold faucets available to cockroaches in the traps of sinks, is basins, tubs, and toilet pipes, flush tanks, ; as spillage ; bowls ; ice in flush tanks and windows ; ; as condensation around leaking pipes and in miscellaneous water-filled containers, such as pet drinking dishes, aquaria, vases drainage from (Gunn and orientalis ; empty beverage bottles ; and boxes. Soft, juicy fruits and vegetables can provide both moisture and food. There seems to be a tendency for certain and Blattella germanica) to become established in the more humid parts of structures, such as basements, around sinks, and in bathrooms. Whether this is a reaction to a species {Blatta orientalis preferred humidity or merely a fortuitous aggregation near sources of drinking water and food has never been clearly demonstrated. The rather widespread dissemination of these species into zones of low as well as high humidity suggests that detailed studies of the microclimatic conditions of structural microhabitats will be needed before meaningful conclusions can be drawn about the stratification of cock- roaches within structures according to species. In nearly is available, all food stored by some kind structures infested by cockroaches, food of either in the structure itself or nearby. man for his own This may use or the use of kept animals be crumbs, food spillage, garbage, or excreta ; ; be the it may glues and pastes on and wall paper sizing on and book covers various dried animal and plant products dead insects living plants etc. In fact, it is almost impossible, despite good housekeeping, to keep any structure used by man free of all food cartons, boxes, stamps, envelopes, labels, cloth ; ; ; ; ; suitable for cockroaches. That the requisite temperature, water, and food are provided, more is attested by the or less adequately, by a variety of structures innumerable infestations of cockroaches that develop when control measures are relaxed. Within structures the accessibility of certain harborages to cockroaches probably depends on the habits of the species and to some extent on their size. Similar types of harborages ; BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 73 may be used by the same species, although there overlapping by different species into the same kinds seems to be some ecology of domiciliary cockThe comparative of daytime shelters. in different structures roaches has not been thoroughly investigated, so any interpretation of observational data is necessarily speculative and inconclusive at this Our time. discussion on pages 324 to 343 also pertinent to this is section. LAND-BASED STRUCTURES Dwellings provide a variety of microhabitats that are acceptable to It has been stated that old houses, or houses that have cockroaches. many cracks and crevices, or have basement kitchens that are not kept clean and in good repair are particularly liable to invasion by cock- roaches (Laing, 1946; British this statement is Museum undoubtedly true, it [Nat. Hist.], 1951). Although has been our personal experience, as well as the experience of others, that new, clean, and well-planned houses and apartments are also easily and sometimes quickly invaded by cockroaches. Mallis (1954) has cited the following places that are frequently infested by cockroaches in homes. In the kitchen, cockroaches are found in and around sinks, in cupboards above and below sinks, under tables and chairs, in stoves, around breadboards, in utility cabinets, in kitchen closets, under linoleum, behind, under, and inside refrigerators and iceboxes. In living rooms cockroaches are found in furniture, studio couches, sewing machines, closets, and bookshelves behind picture frames, pennants, calendars, and other wall ornaments. In bathrooms cockroaches are found in and behind utility cabinets and toilets they may be found in wicker clothes hampers, in brooms and mops, and in door hinges. Ordinarily, cockroaches are not found in bedrooms unless they are abundant elsewhere in the dwelling. Additional harborages are cited under specific cockroaches in the list ; below. In markets DeLong (1948) found the German cockroach in bags of potatoes and onions, in crates of citrus fruits, in pads and shredded papers in banana boxes, and in cases of bottled beverages. The insects were attracted by coffee and crawled into the folds of coffee bags. They were found in cartons of canned goods in bread and baked goods in cartons of packaged cookies, cakes, and crackers. Packaged cereals were attractive, and cockroaches were sometimes found in packages of cigarettes. The insects occurred in scales (by the hundreds) and in cash registers. Rather heavy infestations were found ; ; under stainless-steel cleaning stand. The cappings that covered wooden arms on the fish insects were numerous in display cases where SMITHSONIAN MISCELLANEOUS COLLECTIONS 74 warm and they were sheltered. above produce racks, They were in electrical switch also VOL. I4I found behind mirrors boxes and conduits, and in telephone boxes, as well as generally in cracks and behind loose moldings or loose wall boards. Enclosed boxlike tables were fre- quently heavily infested. In restaurants cockroaches may be found in the following places: Crevices in wood, plaster, concrete, and metal kitchen and in the associated equipment ; ; in the bar in cupboards, ; in the lavatories, and garbage storage areas; and on the undersides of chairs and tables (Mallis, 1954). In drug stores Frings (1948) found cockroaches behind the mirror and between the sink and the cooler. Thousands were found in hollow ornamental shelf edging. The hollow bases of malted-milk dispensers and drink mixers were cockroach havens. In a hospital Frings (1948) found cockroaches in decorative trim around doorways, by the thousands in wicker laundry baskets, and in incubators for premature babies. In military hospitals we have seen cockroaches {Blattella germanica) in kitchens and dining halls in the usual hiding places mentioned above and on the undersides of stainless-steel serving tables. In department stores cockroaches have been found in food departments, beauty salons, rest rooms, dressing rooms, linen departments, and stationery departments (Anonymous, 1952). The infestation in the linen department was traced to clean towels which, when returned from the laundry, contained at least 500 cockroaches per bundle. The insects were carried into the rest rooms and beauty salon when the towels were distributed. The microhabitats of cockroaches in privies and sewers have not been studied. These habitats are particularly important in view of the demonstrated migrations of cockroaches from sewers into dwell- and the possible dissemination of pathogenic microorganisms feces to food. The reader is referred to our 1957(a) paper for a summary of the known information on cockroach dispersal from ings from sewers. COCKROACHES ASSOCIATED WITH LAND-BASED STRUCTURES Most of the cockroaches listed below are either species or they have been found one or man-made building. structures. The known more times known in domiciliary houses or other structural pests breed within the Certain other species, which have been observed only infre- quently in structures and are not known to breed there, may possibly BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 75 be incipient pests; these latter species may attain future economic importance if they establish breeding colonies within a structure. A few species have undoubtedly wandered indoors by accident. It is difficult to decide whether a particular species was an accidental invader or whether Only it was attracted indoors in response to additional information some stimulus. will provide the desired answers. Aglaopteryx ypsilon Trinidad. — Male found indoors (Princis and Kevan, 1955). AUacta Hawaii. —Found similis only indoors at Nauhi. Otherwise this is ap- parently an outdoor species (Swezey and Williams, 1932). Blaberus craniifer —Household pest (Deschapelles, 1939). Particularly abunSantiago and Havana (Rehn and Hebard, 1927). dant houses woodshed (Rehn and Hebard, 191 Florida. — Under boards Cuba. in in in 2, 1914). Blaberus discoidalis — In eating places (Campos R., 1926). —In houses (Rehn and Hebard, 1927). New Jersey. — In greenhouse (Weiss, 1917). stores Puerto Rico. — In homes (Sein, 1923). In Ecuador. Hispaniola. fruit (Wolcott, 1950). Blatta lateralis —Household S.S.R. — Males Central Asia. pest, often found in homes with clay floors (Bei-Bienko, 1950). Turkmen and females occurred in dwellings (Vlasov, 1929). Blatta orientalis This species is a cosmopolitan domiciliary pest (Hebard, 1917; Rehn, 1945). It is reported to occur particularly in basements and crawl spaces under basementless houses (Mallis, 1954). In damp basements where food may is available large colonies are not unusual, but floors off the ground (Gould and Deay, 1940). The number encountered on upper floors is seldom large, but the frequency of occurrence may reach 30 percent of the observations (Spear et al., in Shuyler, 1956). In supermarkets this species hides during the day inside concrete blocks or cracks in it also infest offices and apartments several SMITHSONIAN MISCELLANEOUS COLLECTIONS "J^i the foundation, under furniture, or behind cartons VOL. I4I it is conspicuous markets at night (De Long, 1948). In Great Britain the kitchen is preferred by this pest (and by Blattella germanica) they shelter beneath steam radiators and gas stoves, behind on the ; floors of the ; hot-water pipes, underneath furniture and floor coverings, sinks and baths ; basements and underground kitchens are especially likely to be infested (Laing, 1946; British liffe Museum may (1958) noted that B. orientalis [Nat. Hist.], 1951). Good- travel long distances to find food. Blattella germanica This species is a cosmopolitan domiciliary pest (Hebard, 191 7; Rehn, 1945). It is one of the commonest insects in homes and restaurants (Gould and Deay, 1940). It is found in kitchens, larders, bathrooms, furnace rooms, and storage rooms of bakeries, breweries, hospitals, barracks, as well as dwellings, where, during daylight, hides behind cupboards, furniture, it hanging pictures, panels and skirting boards, in cracks around drains, water pipes, electric wires, and hot-water and steam heating units (Wille, 1920). The German cockroach may be found in cracks around baseboards, pipes, conduits, sinks, and drawers behind cabinets inside switch boxes and refrigerators on under surfaces of tables, chairs, and shelves between stacks of stored goods, and in almost every place that is not readily observed (Kruse, 1948). We have also seen this species packed in electric-clock cases and loud-speaker baffles, in cash registers, and ; ; ; ; clinging to the undersurface of infestation of markets by stainless-steel this species steam tables. The has been described above. Very narrow cracks provide refuges for the German cockroach. Wille (1920) found first-instar nymphs in cracks 0.5 mm. wide and adult males and females without oothecae in cracks 1.6 mm. wide. Shuyler (1956) has observed extensions into relatively new strucby Blattella germanica in the north-central area of the tural habitats United States. A few German cockroaches are now being encountered rooms, bedrooms, clothes closets, bedroom furniture, lobbies, in living entrance halls, checkrooms, nonfood storerooms, nonfood ware- houses, and coin-vending machine repair shops. this species is behaving much like the In these situations brown-banded cockroach, Supella supellectilium. Blattella schubotzi Cameroon. — Five specimens in a house (Princis, 1955). BIOTIC ASSOCIATIONS OF COCKROACHES U.S.A. —Although it may & WILLIS 'J'] vaga Blattella seasons —ROTH mostly an outdoor species, during this is dry- temporarily enter houses in great numbers, occasionally breeding indoors in Arizona (Flock, 1941a). Two adults were col- Texas (Riherd, 1953). lected indoors in Chromatonotus notatus —A male was found indoors Trinidad. (Princis and Kevan, 1955). Cutilia soror Hawaii. —Almost as common houses as Neostylopyga rhombifolia in (Hebard, 1922). Ectobius duskei — Frequently occurs in living apartments in farming an accidental inhabitant (Bei-Bienko, 1950). U.S.S.R. ties as locali- Ectobius pallidus —A summertime pest Massachusetts. in houses along coast (Gurney, 1953; E. R. Willis, personal observation). Generally an outdoor species. Epilampra abdomen-nigrum Trinidad. —Male, indoors (Princis and Kevan, 1955). An outdoor species generally. Ergaula capensis Cameroon. —A male taken in a house (Princis, 1955). Eublaberus posticus Trinidad. — Indoors, feeding on bat feces (Princis and Kevan, 1955). Eurycotis floridana — Occasionally Florida. found in homes (Creighton, 1954; Roth and WiUis, 1954a). Euthyrrhapha pacifica — Hawaii. Found outdoors and indoors where it breeds in neglected cupboards and in rubbish (Fullaway and Krauss, 1945). Holocompsa azteca Mexico. — Household pest (Ball et al., 1942). ;; SMITHSONIAN MISCELLANEOUS COLLECTIONS 78 VOL. I4I Holocompsa cyanea Costa Rica. — One specimen in house (Rehn, 1906). Holocompsa nitidula in American Tropics (Hebard, 191 7). In Cuba (Gundlach, 1890-1891) Puerto Rico In folds of burlap bag, Florida (Rehn and Apparently domiciliary houses under chests, (Gundlach, 1887). etc., ; Hebard, 1914). Ischnoptera rufa occidentalis Panama. —Thrives about human habitations under though litter, not domiciliary (Hebard, 1920). Ischnoptera rufa rufa Jamaica. seem — In hotel. to frequent "While hardly a domiciliary form environments where man it would has considerably disturbed natural conditions, as under debris, docks, under logs and stones in cultivated areas" (Rehn and Hebard, 1927). Leucophaea maderae West "At times Indies. it habitations, warehouses, and other structures a very abundant and serious pest" (Rehn, 1945). it is Puerto Rico —In was also found in fruit stores, markets, 1923 Wolcott, 1950). Reported as a domiciliary pest In and inns (Sein, ; ward Islands (Marshall, 1878) ; Madeira (Heer, 1864) Tropics and sub-Tropics in ; Wind(Rehn, Philippine Islands (Uichanco, 1953) New York City (Anonymous, 1953; Gurney, 1953); Trinidad (Princis and Kevan, 1955). 1937) ; ; This species all is also established in coastal Brazil, Central America, the Greater Antilles, several other tropical islands, and tropical Africa, where it probably originated (Rehn, 1945). Leurolestes circumvagans Hispaniola, Grenada. —Largely domiciliary (Rehn and Hebard, 1927). Leurolestes — Florida. —Rehn and Hebard Cuba. pallidus All over island, in houses, under lockers, etc. (Rehn, 1945 Gundlach, 1890-1891). (1914). This species has been recorded from various islands in the West Indies, from Mexico, Guatemala, and Brazil (Rehn, 1945). BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 79 Methana marginalis Australia. —Reported entering houses (Pope, 1953a). Ifauphoeta cinerea — In (Mackerras and Mackerras, 1948). In dwelHngs, grain stores, and fowl-feeding pens (Pope, 1953). Sudan. Domiciliary in huts of the Shilluk natives; fairly widely Australia. hospital — Hawaii. — In feed rooms of poultry plants (Illingworth, 1942). Florida. — Established feed around Tampa (Gresham, 1952 distributed in eastern Africa (Rehn, 1945). mills in Gurney, 1953) The wide distribution of ; • originated, the to this species Orient and the from East Africa, where it was undoubtedly New World mediated by shipping (Rehn, 1945). Neoblattella Puerto Rico. — Observed sp. [as Blatta caraibea, Rehn and Hebard (1927)] in houses (Gundlach, 1887). Neostylopyga rhombifolia Indo-Malaya and Domiciliary in 1945) 1922). ; New World Philippine Islands (Uichanco, 1953) ; Tropics (Rehn, and Hawaii (Hebard, Oxyhaloa buprestoides Presumably Africa to the to some extent New World it evidently spread from (Rehn and Hebard, 1927; domiciliary, as via slave ships Rehn, 1945). Panchlora nivea Colombia. —A 1946). This is male and a female taken in a dwelling (Princis, primarily an outdoor species which is frequently taken indoors as an adventive on bananas (see p. 150 for references). Parcoblatta fulvescens Florida. — Males found in laboratory and dormitory buildings, ostensibly attracted by lights (Friauf, 1953). Parcoblatta lata Connecticut. door species. — Domiciliary pest (Moore, 1957). Generally an out- SMITHSONIAN MISCELLANEOUS COLLECTIONS 80 VOL. I4I Parcoblatta notha Arizona. — It may occasionally be a nuisance in houses (Ball et al., 1942). Parcoblatta pensylvanica — Country houses often badly infested, Indiana (Blatchley, taken in houses in wooded areas, Michigan (HubFrequently 1920). Infestation by males, females, and nymphs on fourth bell, 1922). U.S.A. floor of building, areas infested by South Dakota (Severin, 1952). Houses in wooded nymphs and occasionally by adults (Gould and Deay, 1940). Canada. Pest in summer cottages in Ontario (Walker, 1912). — Periplaneta americana This 1945). is a cosmopolitan domiciliary pest common It is (Hebard, 1917; Rehn, and in restaurants, grocery stores, bakeries, where food is prepared or stored it was trapped regularly in the basement and upper floors of store buildings, and it was also found in all heated parts of an old meat-packing plant (Gould and Deay, 1938). P. americana was numerous in latrines in Iran (Bei-Bienko, 1950) and in privies in Texas (Dow, 1955) and Georgia (Haines and Palmer, 1955). Large numbers of this species also occur in sewers adjacent to human habitations (Roth and Willis, 1957a). ; Periplaneta australasiae Generally domiciliary, but also occurs outdoors in the (Rehn and Hebard, 1927). It is West Indies a very abundant domiciliary pest in and tropical America (Rehn, 1945) Ecuador (CamPhilippine Islands Puerto Rico (Sein, 1923) 1926) (Uichanco, 1953) Australia (Pope, 1953). Also occurs as a pest in greenhouses in Pennsylvania (Thilow and tropical Africa pos ; R., ; ; ; Riley, 1891) ; France (Giard, 1900) Britain (Laing, 1946; British ; Italy (Boettger, 1930) Museum ; Great [Nat. Hist.], 1951). Periplaneta brunnea Circumtropical domiciliary pest which is apparently more nearly peculiar to the Tropics and adjacent regions than P. americana (Hebard, 1917). This species has been trapped in significant numbers in privies and dwellings in Georgia (Haines and Palmer, 1955). It is homes in San Antonio, Texas the species of Periplaneta present in (MaUis, 1954). COCKROACHES BIOTIC ASSOCIATIONS OF —ROTH & WILLIS 8l Periplaneta fuliginosa — U.S.A. Frequently encountered out of doors, but has been reported common after dark about a hotel in Alabama and was captured in a house in Louisiana was it ; also extremely abundant on the wharves at night in Jacksonville, Florida (Hebard, 1917). As a domiciliary pest it was, next to Blattella germanica, the most common cockroach inside homes in southwest Georgia, where it was also the most common cockroach in privies (Haines and Palmer, 1955)- This species has become a very common domiciliary pest in Texas (Fads, personal communication, 1955). It infested a greenhouse for five years in Indiana (Gould and Deay, 1940). Periplaneta ignota Australia. — It occurs in dwellings occasionally (Pope, 1953). Phaetalia pallida —Three specimens from three dwellings — Male indoors; male and female Colombia. Trinidad. (Princis, 1946). at light (Princis and Kevan, 1955). Plectoptera dorsalis Puerto Rico. —According night attracted by light to Gundlach (1887) it (Rehn and Hebard, 1927). enters houses at Polyphaga aegyptiaca — Common houses (Weber, 1954). —Winged male kitchen (Burr, 1913). U.S.S.R. —Listed as a sinanthrope (Bei-Bienko, 1950). Iraq. in Caucasus. in Polyphaga saussurei South-central Asia. —One of the commonest domiciliary species (Bei-Bienko, 1950). Pseudophoraspis nebulosa East Indies. —This species is sometimes difficult indoors (Karny, 1925)Pycnoscelus surinamensis A household pest in the East Indies (Karny, 1925) Tanganyika (Smith, 1955) (Uichanco, 1953) Islands ; eight records indoors (Princis and Kevan, 1955). Philippine ; ; Trinidad, It is also a green- house pest (Hebard, 1917; Zappe, 1918; Doucette and Smith, 1926; . SMITHSONIAN MISCELLANEOUS COLLECTIONS 82 VOL. I4I Saupe, 1928; Roeser, 1940). Common in or around chicken batteries in Hawaii (Schwabe, 1949). and yards Supella supellectilium (Rehn, 1945), Domiciliary wherever distributed this species is especially difficult to control because of its apparently nonselective For example, Mallis (1954) observed Texas that it was widely distributed throughout the apartment and was probably the most common cockroach seen in the bedroom its favorite harborages were beneath and behind corner braces on kitchen chairs, underneath tables, behind pictures and other objects on walls, and in shower stalls; its oothecae were commonly fastened on walls and ceilings throughout the house. Gould and Deay (1940) reported dispersal throughout dwellings. in ; that this species prefers high locations, such as shelves in closets, behind pictures, and picture molding; oothecae were found about kitchen sink, desks, tables, and other furniture, and even in bedding. Hafez and Afifi (1956) stated that the adult wanders in nearly all rooms of the house and only visits the kitchen when searching for food; it hides in cupboards, pantries, closets, bookshelves, drawers, and behind picture frames ; the nymphs normally hide in the corners of drawers, behind frames, and in similar situations. Symploce Puerto Rico. bicolor — In houses, Sardinera Beach, Mona Island (Ramos, 1946). Symploce hospes — North American Tropics. Domiciliary, but not exclusively so, and apparently widely distributed (Hebard, 1917). In Florida, as Ischnoptera rufescens, found in a greasy cupboard (Rehn and Hebard, 1914). Hawaii. — Illingworth ( 1 91 5) SHIPS Sailing ships have long been notorious for their of cockroaches, and it was by ship unwelcome hordes that at least 11 domiciliary species migrated from their centers of origin to other parts of the world (Rehn, 1945). Over 40 nondomiciliary species have been carried by ship from the American Tropics to other parts of the world in cargoes of bananas (p. 146). In addition to these, other adventive cockroaches appear from time to time in ports to which they have been carried by ships. Yet by far the most numerous cockroaches on shipboard are BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 83 few common domiciliary pests. Except most rigorously disinsectized ships, this commerce in cock- the breeding populations of a in the roaches has continued to the present day. Cockroaches undoubtedly infested the first ships that sailed the Mediterranean; of these we have no records. The earliest recognizable record of cockroaches on shipboard is Moffett's (1634) statement that when Drake captured the ship Philip, he found it overrun with cockroaches [Blattarum alatarum]. Bligh (1792) described disH.M.S. Bounty with boiling water to kill cockroaches. infesting Qiamisso (1829) reported that he had seen ships casks, or grain had been stored, that were found to be filled in which rice with Blattella germanica when opened. During a voyage from England to Van Diemen's Land, Lewis (1836) was greatly annoyed by hundreds of cockroaches flying about his cabin at night the most numerous resembled Periplaneta americana and another was similar to Ectohhis ; lapponicus. This latter was undoubtedly B. germanica, which is the only ship-infesting species that resembles the feral E. lapponicus. Lewis continued, P. americana "were in immense profusion, and had communication with every part of the ship, between the timbers or skin. The ravages they committed on everything edible were very extensive; not a biscuit but was more or less polluted by them, and amongst the cargo 300 cases of cheeses, which had holes in them to prevent their sweating, were considerably damaged, some of them being half devoured and not one without some marks of their residence," Kingsley (1870), Kellogg (1908), Gates (1912), Heiser (1936), and Bronson (1943) have all reported that cockroaches were so numerous on ships that they gnawed the skin and nails of the men on board. These are all independent observations of what may well have been a common occurrence on ships. We have discussed in detail the subject of cockroaches biting man in our 1957(a) paper. Mosely (1892) reported, "At the time that England was left the ship [H.M.S. Challenger] seemed nearly free of animals, other than men, dogs, and livestock required for food. The first cockroaches Cockapparently came on board at St. Vincent, Cape Verdes. roaches soon became plentiful on board, and showed themselves when. ever the ship was in a warm . climate. "At one period of the voyage, a number of these themselves in my cabin, and devoured parts of my all . insects established boots, nibbling ofif the margins of leather projecting beyond the seams on the upper leathers." SMITHSONIAN MISCELLANEOUS COLLECTIONS 84 VOL. I4I Edmund Freemantle (1904) recalled some of his experiences Navy. "Cockroaches in the tropics were also terrible scourges. One saw little of them in fine, dry weather, but in damp, our wet weather they seemed to come from every hidden corner Sir in the British . remedy in the 'Spartan' being caned. . . One was to make brig, the 'Lily,' was so . . — them on pain of overrun by cockroaches the boys catch that the officers' clothes smelt of them." The quotation from Sonan (1924) on page 348 describes similar conditions in the Japanese Navy. On modern cargo ships cockroaches are reported to be extremely numerous in the galley, the crew's quarters, and sometimes in the holds they dwell in hot, humid environments such as the casing around steam pipes (Monro, 1951). Williams (1931) reported that ; Blattella seen at germanica was the most important cockroach pest on ships New York. Although often numerous in the holds, the cock- They were also It was not and more than 20,000 roaches as a rule congregated in living quarters. frequently found between tarpaulins covering the hatches. unusual to kill 20,000 to 50,000 in the forecastle, have been taken from a single stateroom. Simanton (1946) inspected the S.S. William Kieth when it berthed at San Francisco from a lo-month voyage to the South Pacific. The holds were infested with thousands of Periplaneta americana, but in the crew's quarters, mess halls, and storerooms B. germanica predominated. After insecticidal treatment about 2,000 P. americana were seen in each hold and as many as 24 B. germanica in each cabin. that in Army Richardson (1947) reported transports inspected between 1943 and 1946 at New York, P. americana was found in the galleys and messes, and occasionB. gennanica ally heavy infestations were found deep in the holds ; was found in the galleys and messes Blatta ; orientalis was found only in the hold. Additional references indicating the presence of Blattella germanica on ships may be found in the account of its parasite Ripidius pectinicornis (p. 232). Although Rice (1925) and Williams (1931) cite B. germanica as the most numerous cockroach on ships, Brooke (1920) stated that the great majority of ship cockroaches were Periplaneta americana. In addition to citing cockroach infestations on ships, the following authors reported various methods for disinfecting ships: Canalis (1916), Pryor (1918), Brooke (1920), Rice (1925), Williams (1931), Simanton (1946), Richardson (1947), and Anonymous (1951, 1954)- BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 85 COCKROACHES ASSOCIATED WITH SHIPS In the following list we include some previously unpublished data on cockroaches that were recovered from ships at the Miami, Fla., Quarantine Station for the periods November 1945 through May 1946; May, June, August, and September 1950; and 17 July 1957 (Porter, personal communication, 1958). These data were lumped, without breakdown to species, under the entry Orthoptera in Porter (1958). Certain of the species listed below occur only accidentally on shipboard and will probably never establish breeding colonies on ships or become pests on shipboard or elsewhere; some were merely passengers between one land-based colony and another. Others, the truly domiciliary pests, are as likely to be pests on shipboard as they are in land-based structures. Blaberus discoidalis Hispaniola. — On board ship (Rehn and Hebard, 1927). Blatta orientalis U.S.A.— At Port of New York (Richardson, 1947)- Blattella germanica — In — Port of At sea? ships casks (Chamisso, 1829). New York (WiUiams, 1931 Richardson, 1947). San Francisco (Simanton, 1946). At Miami, 7,852 live specimens recovered from ships (Porter, personal communication, 1958). Most numerous species on ships (Rice, 1925). U.S.A. ; Epilampra maya At sea. — One male and one female found dead on S.S. Tenadores (Hebard, 1917). Epilampra Florida. — One dead specimen, sp. Miami (Porter, personal communi- cation, 1958). Ischnoptera Florida. — sp. Five live and one dead specimen, Miami (Porter, per- sonal communication, 1958). Latiblattella sp. At sea. 1917). —One female alive in hold of S.S. Tenadores (Hebard, SMITHSONIAN MISCELLANEOUS COLLECTIONS 86 VOL. I4I Leucophaea maderae Brought from West Africa (Rehn, 1945). to West Indies and Brazil by slave ships Nauphoeta cinerea Widely disseminated by sailing ships (Rehn, 1945). Neoblattella fratercula At sea. —Two females found dead on S.S. Tenadores (Hebard, 1917). Neoblattella fraterna At sea. — One male found dead in hold of S.S. Tenadores (Hebard, 1917). Neoblattella nahua At sea. — One female dead in hold of S.S. Tenadores (Hebard, 1917). Neoblattella sp. Florida. ter, —Five dead specimens recovered from ships, Miami (Por- personal communication, 1958). Neostylopyga rhombifolia Widely distributed by sailing ships (Rehn, 1945). England. Captured on a sugar vessel from Java (Lucas, 1920). — Nyctibora noctivaga — One male and one female nymph found dead on S.S. Tenadores (Hebard, 191 7). At sea. Nyctibora sp. —Two dead specimens recovered from ships (Porter, personal communication, 1958). Florida. at Miami Oxyhaloa buprestoides Spread from Africa to New World by ships (Rehn and Hebard, 1927; Rehn, 1945). Panchlora nivea —One female dead hold of S.S. Tenadores (Hebard, Florida. — Fifteen dead specimens taken from Miami (Por- At sea. in 1917)- ships, ter, personal communication, 1958). COCKROACHES BIOTIC ASSOCIATIONS OF From the —ROTH & WILLIS 87 numerous records of this species as an adventive taken it may be presumed to be a frequent traveler on on bananas (p. 150), banana boats. Periplaneta americana At sea.— Lewis (1836). Hebard (1933a) stated that this is "often a serious pest on the smaller ships sailing the South Seas." USA. San Francisco (Simanton, 1946). Port of New York (Richardson, 1947). At Miami, 62 live and 123 dead specimens — (Porter, personal communication, 1958). Periplaneta australasiae in slave ships (Rehn, by way of Africa in slave Migrated from West Africa to America 1945)Pycnoscelus surinamensis Probably in part reached the New World ships (Rehn, 1945). Supella supellectilium Reached America from West Africa by slave ship (Rehn, 1945). Xestoblatta festae At sea. — One female found dead in hold of S.S. Tenadores (Hebard, 1917). AIRCRAFT Michel (1935) stated that the development of air transportation brought the same insect dispersal problems that exist in land and water transportation; in addition, the problem of cockroach infesta- had become a very serious one, quite aside from the hygienic it had been discovered that these insects seek out the wings of airplanes, where they subsisted on the glue and dope used in airplane construction. However, Dethier (1945) found no cockroaches in dismantled or wrecked wing and tail structures of metal aircraft in central Africa. In fact, all-metal aircraft would seem to provide little in the way of food or water for stowaway cockroaches. Laird (1951, 1952, 1956a) found living specimens of Blattella germanica, Periplaneta americana, and Periplaneta australasiae in baggage compartments and/or kitchens in aircraft. Other species which have been recovered from undisclosed spaces in aircraft are listed below. Some of the cockroaches that were reported as dead may not have died from exposure during flight but may have been killed by tion point of view, because insecticide applied by inspecting personnel at the airports. . . SMITHSONIAN MISCELLANEOUS COLLECTIONS 88 VOL. I4I COCKROACHES ASSOCIATED WITH AIRCRAFT In the following list we include some previously unpublished data on cockroaches that were recovered from aircraft in Miami, Fla., International Airport from i July 1956 through 30 June 1957 (Porter, personal communication, 1958). These data were lumped under the entry Orthoptera without breakdown to species in Porter (1958). Species reported by Hughes (1949), and cited below as from southern United States, were recovered from aircraft that arrived at Brownsville, Fort Worth, Miami, New Orleans, and San Juan. There was no way of linking a specific record with any particular city. The comments we made above about species that are infrequently encountered on ships apply with equal validity to similar species found on aircraft. Anaplecta U.S.A. aft at — One sp. and 15 dead specimens recovered from 16 live Miami (Denning et al., air- 1947). Blatta orientalis U.S.A. craft at — Six live and four dead specimens recovered from six airSouthern U.S. (Hughes, et al., 1947). Miami (Denning 1949). Blattella germanica —Williams — — — Hawaii. ( 1946a) Khartoum. Whitfield ( 1940) Nezv Zealand. Laird (1951, 1952, 1956a). U.S.A. At Miami 193 live and 184 dead specimens were recovered from 141 aircraft (Denning et al., 1947). Recovered at airports in southern U.S. (Hughes, 1949). Recovered at Miami, 51 live and 24 dead specimens (Porter, personal communication, 1958). Exposed experimentally in jet aircraft (Sullivan et al., 1958). Blattella sp. — Khartoum. Whitfield ( 1940) Southern U.S. Hughes (1949). — Cariblatta spp. U.S.A. —One aircraft at 1949). live and three dead specimens recovered from three et al., 1947). Southern U.S. (Hughes, Miami (Denning BIOTIC ASSOCIATIONS OF COCKROACHES Epilampra U.S.A. —ROTH & WILLIS 89 sp. —One dead specimen, Miami (Denning et al., 1947). South- ern U.S. (Hughes, 1949). Eublaberus posticus Southern U.S. — Hughes (1949). Ischnoptera rufa rufa U.S.A. —Two dead specimens recovered Miami (Denning et al., Ischnoptera U.S.A. —Two live from two aircraft at 1947). sp. and one dead specimen recovered from three air- Miami (Denning et al., 1947). Southern U.S. (Hughes, 1949). One dead specimen, Miami (Porter, personal communication, 1958). craft, Leucophaea maderae U.S.A. —Three Miami (Denning dead specimens recovered from three aircraft 1947). Southern U.S. (Hughes, 1949). at et al., Nauphoeta cinerea U.S.A. — One live and one dead specimen, Miami (Denning et al., 1947)Neoblattella U.S.A. sp. — One dead specimen recovered at Miami (Denning et al., 1947). Panchlora nivea U.S.A. —Two dead specimens recovered from two (Denning et al., aircraft, Miami 1947). Periplaneta americana U.S.A. aircraft, — Five and three dead specimens recovered from seven Southern U.S. (Hughes, et al., 1947). and five dead, Miami (Porter, personal communi- live Miami (Denning 1949). Three live cation, 1958). Experimentally et 1958). New Zealand. exposed in jet aircraft, U.S. (Sullivan al., —Laird (1951, 1952). Periplaneta australasiae — U.S.A. Two dead specimens recovered from two (Welch, 1939). Five live and three dead from five aircraft, aircraft, Miami Miami . SMITHSONIAN MISCELLANEOUS COLLECTIONS 90 VOL. I4I (Denning et al., 1947). Southern U.S. (Hughes, 1949). Three and two dead, Miami (Porter, personal communication, 1958). New Zealand. —Laird ( live 1952 ) Periplaneta spp, U.S.A. — One Miami (Denning live live and three dead specimens from four aircraft, 1947). Southern U.S. (Hughes, 1949). One et al., and six dead, Miami (Porter, personal communication, 1958). Pycnoscelus surinamensis —Two and three dead specimens from 1947). Southern U.S. (Hughes, New Zealand. — Laird (1956a). U.S.A. live Miami (Denning five aircraft, et al., 1949). Supella supellectilium — U.S.A. Two live and one dead specimen from one aircraft, Miami (Denning et al., 1947). Southern U.S. (Hughes, 1949). Khartoum. Whitfield (1940). — Supella — Two live specimens from two Southern U.S. (Hughes, 1949). U.S.A. 1939)- sp. Symploce U.S.A. —Two ami (Denning live aircraft, Miami (Welch, sp. and one dead specimen from three aircraft, MiSouthern U.S. (Hughes, 1949). et al., 1947). Unidentified cockroaches Anglo-Egyptian Sudan. Brazil. —From —At Khartoum (Whitfield, 1940). flying boats, 62 specimens; from land planes, 45 specimens (Carneiro de Mendonga and Cerqueira, 1947). Dethier (1945). Central Africa. — —At Kisumu (Symes Whitfield, 1940). New Zealand. — Laird (1956a). and 14 dead specimens from 4 U.S.A. — Four Kenya. in live aircraft, Miami (Welch, 1939). One live and 15 dead specimens (adults?) from 16 aircraft; 8 oothecae from 7 planes; 147 live and 83 dead nymphs from 108 planes, Miami (Denning et al., 1947). Southern U.S. (Hughes, 1949). Five live and 7 dead specimens, Miami (Porter, personal communication, 1958). : : BIOTIC ASSOCIATIONS OF COCKROACHES IV. —ROTH & WILLIS 9I CLASSIFICATION OF THE ASSOCIATIONS Asano (1937) classified the natural enemies of cockroaches into two types as follows 1. 2. Enemies that feed mainly on cockroaches (certain ripiphorid beetles and certain chalcid, evaniid, and ampulicid wasps). Organisms which, in their search for food, devour cockroaches that may be ticks, centipedes, Strepsip- encountered (certain species of scorpions, spiders, tera, ants, birds, rats, and "parasitic bacteria"). Cameron (1955) arranged the associates of cockroaches groups as follows two in Group A. Parasites and predators. 1. Parasites: Hymenoptera (Evaniidae, Eulophidae, Eupelmidae, Encyrtidae, 2. Pteromalidae, Cleonymidae) and Coleoptera (Ripiphoridae). Predators: Hymenoptera (Ampulicidae), Hemiptera (Reduviidae), Coleoptera (Dermcstidae), Arachnida (Araneae, Acarina). Group B. Parasites and symbionts. 1. 2. Protozoa (including examples of both parasites and commensals). Nematoda (including both primary "parasites" and secondary parasites). 3. Bacteria (including the mutualistic bacteroids). 4. Algae [Arthromitis (= Hygrocrocis) intestinalis; see p. 124]. Asano's arrangement, although essentially true, is limited; Camsystem is divided into arthropods (group A) and lower forms (group B), but does not include higher animals. Both attempts eron's need amplification at classification ; this we have endeavored to do below. In classifying the biotic associates of cockroaches, mediately confronted with a problem in semantics. parasitism, predatism, and symbiosis have shades of meaning by different authors. merely by accepting as authoritative all we were imThe concepts been used with various The problem specific definitions, is not solved however apt they seem to be, because, unfortunately, these concepts are not mu- For example, among the entomophagous insects, as Sweetman (1936) has pointed out, there can be no definite line of the two intergrade, separation between parasitism and predatism distinct. In Andrewartha and quite fact, only the extremes being relationships calling generalized these both cateBirch (1954) by divided natural populations of These authors gories predatism. associated organisms into nonpredators and predators. Although this tually exclusive. : simplifies their presentation of the general principles of ecology, for our purpose more narrowly defined terms have proved useful. In the main we have followed Sweetman (1936) and Allee (1949) in arriving at the following definitions: et al. ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 92 Symbiosis is the living together in tion of organisms of different species more or ; it VOL. I4I less intimate associa- includes virtually all rela- and other organisms, such as parasitism, predatism, commensalism, and mutuahsm. Alice et al. (1949) apparently do not include predatism in symbiosis. Mutualism is symbiosis in which both members benefit by the association. The smaller partner has commonly been called a symtionships between cockroaches biont or symbiote by authors. Commensalism includes associations in which neither party ap- partner may live on the surplus and transport may be involved. Parasitism is the state of symbiosis in which one of the members feeds upon the other during the whole of either the immature or mature feeding stage; the host is harmed in some way and may be pears to benefit or be harmed. food or wastes of the other ; One shelter killed. Predatism is an association upon, or stores as food for its in which one member attacks and feeds progeny, one or more other organisms immature or mature feeding period on the prey. This category includes a few invertebrates and all the vertebrates that capture, kill, and feed on cockroaches. This association may be divided into interspecies predatism, in which the predator preys upon a different species, and intraspecies predatism (cannibalism) in which the predator preys upon its own species. Although we have attempted to adhere to these definitions throughout this discussion, we realize that in doing so we may have tended to oversimplify complex relations. Some questionable interpretations stem from insufficient knowledge of the basic relationships between cockroaches and their associates. Only further study will clarify these relationships. Some of the problems are discussed below. the predator spends less than the Probably many of the so-called parasites Protozoa (e.g.. like Nyctotherus, and intestinal nematodes of the family Thelastomatidae), which do not invade the host's tissues and seem to have no effect on the activity and vitality of the host, are commensals. Although we consider these forms to be commensals, we realize that they might actually affect the host in some way even though this has not been shown. It is possible that Rothschild and Clay's (1957) statement about bird parasites may well apply to the apparently harmless organ- isms found in the cockroach. These authors wrote, "It cannot be too strongly emphasized that the effect of host is detrimental. If we all find that a bird types of parasites on the seems convenienced by the presence of Protozoa or cuckoo in the nest, we can little, if worms nevertheless assume that it or at all, lice, would be in- or a better COCKROACHES BIOTIC ASSOCIATIONS OF off without them . . Small . —ROTH such as lack of effects degree. gauge. difficult to Potentially all Nevertheless, it is parasites are harmful." 93 vitality, loss of eating are dirt voice, excessive blinking, or perverted habits like extremely & WILLIS only a question of should also be It pointed out that some workers would consider certain of our com- mensals of cockroaches to be parasites. that "A truly successful parasite of equilibrium with its is host, so that Thus Faust (1955) stated one which has developed a state no detectable damage is produced In a suitable host the which endangers the health or life of the host. parasite may obtain food and shelter without any evidence of trauma or toxicity. The damage produced may be so slight that repair and functional readjustment keep pace with the injury." Faust's suc- cessful parasite would be indistinguishable from a commensal, but there is undeniably a difference between an organism causing slight, and undetectable, damage to a host and one causing none. Certain of we the organisms list as commensals may eventually be shown to be parasites. Certain organisms which live in cockroaches appear to have no effect on the vitality of the host invaded. Gregarines may even though the tissues of the host are penetrate the intestinal wall of the cockroach without seeming to injure the host. Fungi of the genus Herpomyces invade the cuticle of cockroaches producing pathological changes yet consider the insects' behavior is apparently unaffected (see p. 129). ; We these organisms to be parasites because the host's tissues are invaded and, as far as we know, no benefit to the host results. In the literature certain insects have been considered to be either parasites or predators or both. Among these are the ensign wasps (Evaniidae), whose larvae feed on the eggs of cockroaches within the ootheca, and the ampulicid wasps, which capture, paralyze, store in their nests (as food for their larvae) cockroaches. splendidiila is nymphs and and adults of Clausen (1940) claimed that the evaniid Zeuxevania a true egg parasite when it destroys the first egg in a cockroach ootheca; but after the wasp larva molts and proceeds to devour the other eggs, he considered it to be a predator. Clausen's definition of an entomophagous parasite is an insect that in its larval stage develops either internally or externally upon a single host which is eventually killed and their food is ; with few exceptions the adults are free-living usually different from that of the larvae. atory insect, by Clausen's definition, is A pred- principally free-living in the larval as well as adult stage, kills the host immediately and requires a number of victims to reach maturity by direct attack, ; the predator is SMITHSONIAN MISCELLANEOUS COLLECTIONS 94 VOL. I4I of greater size than the prey, and the food sources of the adults and immature stages are frequently the same. It is apparent, as Clausen and other writers have pointed out, that there are instances of a particular species showing characteristics which fit both the definitions for predator and parasite. Thus, among the evaniids one wasp larva destroys all the eggs in an ootheca, but in spite of this the larva has more of the characteristics of a parasite than of a predator the adult wasp does not utilize the same food as the larva (adults have been taken on flowers and on honey dew from ; scale insects). It is questionable whether the evaniid larva kills the cockroach egg outright. The wasp larva, being restricted to the inside of the ootheca, is not free-living. Probably the only criterion by which the evaniid could be judged to be a predator, by Clausen's definition, is more than that a single egg devoured by the maturing wasp larva. (Encyrtidae, Eulophidae, Eupelmi- is Among the other wasp parasites dae) of cockroach eggs When many individuals develop in a single ootheca. a hundred or more wasps emerge from an ootheca which con- tained less than 20 eggs, it is obvious that a single cockroach egg sup- it is possible that one particular wasp more than one cockroach egg before be- ported more than one wasp, yet larva may have coming an fed upon adult. We consider in cockroach oothecae to entomophagous wasps that develop all be parasites rather than predators. On the other hand, even though Anastatus floridanus, A. tenuipes, and Tetrastichus hagenowii are egg parasites as larvae, the adult females are, in a sense, predators when they sometimes eat part of the cockroach egg that oozes through the oviposition puncture (Roth and Willis, 1954a, Ampulex 1954b). Williams (1929) has seen the female of from the cut ends of canalicidata imbibe blood that oozed the cockroach's antennae after she had clipped the prey to her nest. Yet despite this evident them off before leading predatism on the part of the adult, the larva feeds as a parasite on the stored cockroaches Sweetman's (1936) (though not Clausen's 1940) which is "that form of symbiosis in which one symbiont lives in or on the host organism and feeds at its expense during the whole of either the immature or mature feeding stage." The ampulicid larva, as the evaniid, is not free-living and does not kill the host immediately by direct attack, even though it may require more than one victim to reach maturity. Thus, within one individual both parasitic and predatory behavior are operant during different in accordance with definition of parasitism, stages of With its life history. the above discussion in mind we have summarized below the Only a few examples are various biotic associations of cockroaches. BIOTIC ASSOCIATIONS OF COCKROACHES given for each section, but sumably would be all classified in the —ROTH & WILLIS 95 organisms with similar habits pre- same categories. Associations in which cockroaches serve as hosts, vectors, or prey for Class A. other organisms. Animals and plants that normally develop on or in the cockroach; in general, these organisms depend entirely upon the cockroach for survival. Type I. Obligate associates. only Group Mutuals (symbiotes or symbionts of authors). (bacteroids which are found in the fat body of all cockroaches that have been examined; p. 96), (b) Bacteria (wood-digesting forms in Panesthia, and possibly certain I. (a) Bacteria-like organisms bacteria in the intestines, of other cockroaches; p. 100). (c) Protozoa Group 2. (several genera and species found in Cryptocercus; lOl). p. Commensals. (a) Protozoa {Nyctotheriis, Herpetomonas, Lophomonas, etc.; (b) Nematodes (Thelastomatidae; Group 3. p. 172). p. 201). Parasites. (a) Fungi (Laboulbeniales ; p. 134). (b) Protozoa (gregarines, Plistophora, etc.; (c) p. 193). p. 181). Helminths. worms; (7) Primary parasites (mermithids and gordian {2) Secondary parasites {Gongylonema neoplasticuni, Oxyiiris inansoni, Moniliformis spp. p. 206). ; (d) Arthropods. (/) Mites (Pimeliaphilus podapolipophagiis ; (2) Insects p. Type II. (larvae of ripiphorids, 219). p. evaniids, and ampulicids; 231). Facultative associates. Animals and plants that prey on cockroaches or are incidentally or accidentally picked up by the cockroach, but which can survive or propagate readily on some other host or prey. Steinhaus (1946) emphasized the importance of the environment in determining the type of microbial flora associated with the cockroach which may is exposed to filth and a Because many of these organisms survive passage through or on the cockroach, the blattid may act as a vector of these animals and plants. carry one type of flora in an area which different type in other areas. Group I. Commensals. (a) Viruses (strains of poliomyelitis virus; p. 103). (b) Bacteria (Enterobacteriaceae, Pseudomonadaceae, Micrococcaceae, etc.; p. III). (c) Fungi (Aspergillus; p. 130). (lodamoeba, Dobellina, and cysts of Entamoeba and Entamoeba histolytica; p. 179). (d) Protozoa (e) Helminths (cysts of various helminths parasitic p. (f) 208). Arthropods. (i) Mites (Tyrophagus lintneri; p. 218). in colt vertebrates; SMITHSONIAN MISCELLANEOUS COLLECTIONS 96 Group 2. Parasites. (a) Bacteria {Serratia marcescens ; (b) Helminths {Protospirura spp. (c) VOL. I4I p. p. ; 117). 206). Arthropods. (j) Mites (Lociistacarns sp. p. 219). (^) Insects (Melittobia chalybii; p. 248). ; Group 3. Predators, active. (a) Arthropods. (i) Spiders (p. 214). {2) Scorpions (p. 212). (j) Centipedes (p. 222). (^) Mites {Rhizoglyphiis tarsaliis; p. 218). (5) Insects (dermestids, reduviids, and on occasion adult females of Tetrastichiis, Anastatns, Ampulex; p. 234). (b) Vertebrates. (j) Amphibia (p. 269). {2) Reptilia (p. 272). {3) Group Class B. 4. Aves (p. 276). {4) Mammalia (p. 283). Predators, passive: Pitcher plants (p. 154). Associations in which cockroaches serve as commensals or predators. Group I. Commensal cockroaches. (a) Associates of social insects (^Attaphila spp., etc.; p. 315). (b) Obscure associates (p. 316). Group 2. Predatory cockroaches. (a) Interspecies predators (p. 319). Class C. (b) Intraspecies predators (p. 322). Associations of cockroaches with other cockroaches. Group I. Intraspecies associations. (a) Familial associations (p. 325). (b) Other conspecific associations (aggregations and fighting) (p. 336). Group 2. Interspecies associations. (a) Compatible associations (p. 337). (b) Antagonistic associations (p. 329). Class D. Ecological associations of cockroaches with higher plants. Group Group I. 2. Benign associations (p. 139). Associations detrimental to plants (p. 162). V. MUTUALISM BACTEROIDS Blochmann (1887, 1888) discovered intracellular particles (the bacteroids or symbiotes of authors) that resembled bacteria in the body of males and females of Blatta orientalis and Blattella germanica (pi. 26), in the ova of these insects, and in their embryos. Bacteroids have since been found in at least 25 species and 19 genera of cockroaches. Presumably such microorganisms are universally fat distributed throughout the Blattaria. General reviews of the bacteroids BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 97 of cockroaches and other insects have been published by Glaser (1930b), Schwartz (1935), Steinhaus (1946, 1949), Buchner (1952, Brooks (1954), and Richards and Brooks (1958). The 1953) » reader is referred to these papers, and those of authors cited in the Hst at the end of this section, for discussions of the morphology of the bacteroids, their distribution within the host, and attempts to them culture It in vitro. has long been assumed, without proof, that cockroaches and their bacteroids form a mutually may As beneficial association. possible to cultivate bacteroids apart be assumed that the host is from it has not been their cockroach hosts, it essential to the continued existence of the microorganism, which also derives from the association other Experiments to show that the host also from the association have centered around rendering cock- obvious benefits as well. benefits roaches bacteroid free. tion, heat or cold, Starvation, parasites, electromagnetic radia- or chemicals have eliminate the bacteroids. Of been used in attempts to all these, only chemical treatment has pro- vided a satisfactory technique. Few chemicals other than antibiotics have proved to be useful in the elimination or reduction of bacteroids. Yetwin (1932) injected He observed decreases in the bacteroids of the fat body only following various dilutions of 22 compounds into Blattella germanica. injection of methylene blue, but did not pursue this lead further, Gier (m Steinhaus, 1946) observed reduction in the numbers of bacteroids after cockroaches were injected with crystal violet, hexyl- Bode (1936) reported resorcinol, or metaphen. irritants such as lithium salts that injection of or quinine hydrochloride had no ap- parent effect on the symbiotes. Brooks (1957) reared Blattella germanica on diets containing dif- ferent concentrations of inorganic ions. diet the cockroaches normal bacteroids ; On a manganese-deficient grew poorly and some of their progeny lacked about 10 percent of the aposymbiotic generation grew and reproduced on a diet fortified with yeast. Varying the con- centrations of other salts in the diets gave results in which the progeny were either aposymbiotic or the the mycetocytes were abundant on ; all fat body was abnormal but these cockroaches soon died even fortified diets. The administration of certain antibiotic drugs has produced cock- roaches very nearly free of bacteroids. Brues and Dunn (1945) found had no effect on the bacteroids, large doses reduced the number of bacteroids in Blaherus that although sulfa drugs penicillin in craniifer, SMITHSONIAN MISCELLANEOUS COLLECTIONS 98 VOL. I4I but the cockroaches died within a few days. Death was attributed to lack of bacteroids rather than effect of the drug; this is perhaps an unwarranted conclusion in view of the survival of aposymbiotic cock- roaches that have been produced by other drugs (Brooks, 1954; Brooks and Richards, 1955). Glaser (1946) found that the bacteroids of Periplaneta americana could be adversely affected or destroyed by sulfathiazole, or sodium or calcium penicillin. Fraenkel (1952) questioned the conclusions of Brues and Dunn (1945) and of Glaser (1946) because of the high mortality in their experiments, and he suggested that the described phenomena "were due rather to direct toxic effects on the host than to loss of the symbionts." Noland (in Brooks, 1954; Brooks and Richards, 1955) confirmed Glaser's results with penicillin and extended sulfa treatments to include Blattella germanica. Every female whose bacteroids were reduced to the vanishing point resorbed her ovaries and was incapable of reproduction. Brooks (1954; Brooks and Richards, 1955) found that adminisfrom the tration of several antibiotics did not eliminate the bacteroids body oi B. germanica unless the dose was so high that it caused Frank (1955, 1956) was able to eliminate bacteroids from Blatta orientalis by injecting or feeding chlortetracycline, oxytetracycline, or penicillin survival of treated insects was not good and reproduction was poor the aposymbiotic individuals were smaller than normal. As Richards and Brooks (1958) have pointed out, it is uncertain how much of this difference was the result of loss of bacteroids and how much the effect of the drug. It is obvious that in all these experiments the action of the drugs on the bacteroids was accompanied by equivocal side effects which confused interpretation of the results. The effect on the cockroach of a loss of bacteroids cannot be separated from a possible toxic effect of the drug. Fortunately Brooks (1954; Brooks and Richards, 1955) obtained completely aposymbiotic offspring from Blattella germanica that had been reared on aureomycin. These bacteroid-free nymphs were practically incapable of growth on a natural diet that was adequate for nymphs with symbiotes. However, the addition of large amounts of dried yeast to the diet enabled aposymbiotic nymphs to mature in two to three times the period required by normal nymphs. Final proof of the function of the bacteroids was obtained by reestablishing them fat excessive mortality. ; ; in aposymbiotic cockroaches. The insects that received implants of body of B. germanica showed a slow, steady gain in weight over the controls (Brooks, 1954; Brooks and Richards, 1956). Obviously, the intracellular symbiotes subserve the normal nutrition of the cockroach. Whether the bacteroids produce only vitaminlike sub- normal fat BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 99 by Keller (1950), or function in some other way Brooks (1954) concluded that the amount of vitamin-containing food required for increased growth by aposymbiotic cockroaches is much greater than the known vitamin requirements hence the factor (s) needed is unknown and present in low stances, as suggested to be determined. is still ; concentration, or synthesis is it serves as a precursor of a second factor (s) whose aided by the bacteroids. Brooks (in Richards and Brooks, 1958) has since found that the bacteroids of Blattella germanica "can supply the insect with B vitamins, amino acids and some larger protein fragment." all Gier (1947) stated that the symbiotes of cockroaches are generically the same. However, as the symbiotes are presumed to have been associated with cockroaches for over 300,000,000 years (Buchner, may be assumed to have developed them inseparably to their respective hosts. Ries (1932) transplanted symbiote-containing fat body from Blatta orientalis into the mealworm and larva of Ephestia kuhniella, and from Blattella germanica and Stegobium paniceum {=Sitodrepa panicea) into B. orientalis. The implants did not become established in the new host, although most of the transplantations were successful in that the hosts survived and the implants remained intact for some time before they were encapsulated by host tissue. Brooks ( 1954 Brooks and Richards, 1956) transplanted fat body of Periplaneta americana and B. orientalis into aposymbiotic B. germanica. The growth of the cockroaches injected with foreign tissue was not different from that of aposymbiotic controls. Sections of host insects did not contain mycetocytes and no bacteroids were found. Haller (1955a) injected bacteroids or implanted mycetocytes of B. germanica into gryllids, acridids, and locustids. These implants and innoculations were rapidly destroyed by the hosts. But as Richards and Brooks (1958) have pointed out, none of these experiments provide information about the specificity 1952) they specific differences that link ; of the bacteroids themselves. COCKROACHES IN WHICH BACTEROIDS HAVE BEEN FOUND Bantua stigmosa. Fraenkel (1921). Blaberus craniijer. Brooks (1954); Brues and Diinn (1945); Hoover (1945). Blaberus giganteiis. Blochmann (1892). Brooks (1954) Blatta orientalis. Blochmann (1887, 1888, 1892) Bode (1936) Frank (1955, 1956); Buchner (1912) Cuenot (1896); Fraenkel (1921) ; ; Gier (1936, 1947) Heymons (1930) ; (1895) ; ; ; ; ; Glaser (1920) ; Gropengiesser (1925) Javelly (1914) ; Keller (1950) ; ; Gubler (1948) ; Hoover (1945) Hovasse Koch (1949) Mencl (1907)?; Mer- Hollande and Favre (1931) ; ; ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 100 cier (1906a, 1907, 1907b, 1907c) (1946) ; Wolf Ries ; (1932) ; Ronzoni VOL. I4I (1949) Tacchini ; (1924, 1924a). BlaUeUa germanica. Blochmann (1946, 1948, 1948a) ; (1887, 1888, 1892); Brooks (1954) ; Bode (1936); Borghese Brooks and Richards (1954, 1955, 1955a, 1956); Fraenkel (1921); Gier (1936, 1947) Glaser (1920, 1930a); Gropengiesser (1925); Haller (1955, 1955a); Heymons (1892, 1895); Hoover (1945); Koch (1949); Lwoff (1923); Milovidov (1928); Neukomm (1927, ; 1927a, 1932) ; Perez Silva (1954, i9S4a) ; Ries (1932) ; Rizki (1954) ; Ronzoni Wollman (1926) Yetwin (1932, 1953). (1949) ; Tacchini (1946) Cryptocercus pnnctulatiis. Gier (1936, 1947); Hoover (1945). Derocalymma cruralis. Fraenkel (1921). ; ; Heymons (1892); Cuenot (1896); Koch Heymons (1892, 1895); Cuenot (1896); Ectohius lapponicus. Ectobius pallidus. (1949). Fraenkel (1921). Eurycotis floridana. Gier (1936, 1947). Loboptcra dccipiens. Buchner (1930). Epilaiupra grisca. Nauphoeta cincrea. Fraenkel (1921). Nyctibora noctivaga. Gier (1947). Parcoblatta lata. Gier (1936, 1947). Parcoblatta pensylvanica. Brooks (1954) Gier (1936, 1947). Parcoblatta uhleriana. Gier (1936, 1947) Hoover (1945). Parcoblatta znrghiica. Glaser (1920) Gier (1947). ; ; ; Baudisch (1956); Bode (1936); Cuenot (1896); Gier (1936, 1937, 1947) Glaser (1920, 1930, 1946) Gubler (1948) Hertig (1921) Hoover (1945) Ketchel and Williams (1953). Pcriplaneta americana. ; ; ; ; ; Periplaiieta australasiae . Gier (1936, 1947) ; Koch (1949). Platyzosteria armata. Fraenkel (1921). Polyphaga aegyptiaca. Fraenkel (1921), Pscndoderopeltis aethiopica. Fraenkel (1921). Pycnoscelus siirinamcnsts. Bode (1936) Supclla supellectilium. ; Koch (1949). Brooks (1954). BACTERIA Evidence showing that intestinal bacteria contribute to the nutrition is meager. Cleveland et al. (1934) isolated a bacterial organism from the foregut of the wood-feeding cockroach Panesthia of cockroaches angustipennis. The bacterium digested cellulose rapidly in vitro and these workers believe that this cockroach and other related wood- feeding species are dependent on symbiotic bacteria for the digestion of their food. Mencl (1907) described cell nuclei in "symbiotic," not closely de- abundance in the digestive tract (presumably Blatta orientalis). Unfortunately, he was more concerned about the morphology of the bacteria than the stated mutualistic relationship, so nothing is known fined types of bacilli that he found in of the KiJchenschabe, Periplaneta of their physiology. BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS lOI The growth rates of Periplaneta aniericana and Blattella gcrmanica were retarded when the insects were reared aseptically, which suggests that microorganisms normally found certain necessary dietary constituents Noland supply in the digestive tract (Gier, 1947a; House, 1949). (1949) suggested that microorganisms in the digestive germanica synthesized riboflavin since the nymphs reared et al. tract of B. on a low riboflavin diet accumulated more of the vitamin than could have been ingested in the diet. However, Metcalf and Patton (1942) found little or no bacterial synthesis of riboflavin in P. americana. Noland and Baumann (1951) suggested that methionine, one of the amino acids essential for rapid growth of B. germanica, was synthesized by intestinal microorganisms in the insects. PROTOZOA It probable that with few exceptions protozoa found in the is How- digestive tract are not necessary for survival of the cockroach. ever, very importance, few experiments have been performed if to determine the any, of these microorganisms to the host. Cleveland (1925) removed the protozoa from the cockroach (possibly Periplaneta americana) by oxygenation at 3.5 atmospheres. The ciliates Nyctotherus and Balantidium, flagellates Lophomonas and Polymastix, the amoeba Endamoeba hlattae, and three unidentified pro- tozoa were killed by this treatment, yet the insects lived normally after defaunation. Armer (1944) studied the effects of high-carbohydrate, high-fat, and high-protein diets, as well as starvation, on the intestinal protozoa {Nyctotherus ovalis, Endamoeba blattae, Endolimax blattae, Lophomonas striata, and Lophomonas blattarum) in Periplaneta americana. Starvation of the host lowered the incidence or eliminated most of the protozoa, but a high-carbohydrate diet maintained them at a relatively high level. Lophomonas blattarum was eliminated by a high- protein diet, and practically eliminated by a high-fat diet. was eliminated from some hosts Lophomonas were kept on high- fat and high-protein diets. Endamoeba blattae showed a decrease in infection rate when the cockroaches were maintained on high-fat and high-protein diets. The effects of diets on Endolimax blattae were not striata that uniform. has been shown by Cleveland (1930, 1948) and Cleveland et al. (1931, 1934) that the wood-feeding cockroach Cryptocercus punctuIt latus depends protozoa upon utilize as certain intestinal protozoa food the wood ingested by for survival ; this cockroach. these The SMITHSONIAN MISCELLANEOUS COLLECTIONS 102 wood VOL. I4I broken down into compounds the cockroach can utiHze by the is protozoa which elaborate a cellulase and possibly a cellobiase (Trager, 1932), Only molting nymphs of Cryptocercus can pass the protozoa on to the newly hatched young, so that molting and hatching must happen concurrently each year or the young die. The sexual cycles in species of protozoa in the genera Tricho- nympha, Saccinohaculus, Oxymonas, Monocercomonoides, Hexamita, Encomonympha, Leptospironympha, Urinynipha, Rhynchonympha, Macrospironympha, and Barhulanympha (fig. 3, B) are induced by hormones produced by Cryptocercus only during its molting period (Cleveland, 1931, 1947, 1947a, 1949- 1956a). Perhaps the prothoracic may be responsible for initiation of the gland hormone of the host (Cleveland and Nutting, 1955). The protozoan be used as indicators of the onset of molting in flagellate sexual cycles sexual cycles may Cryptocercus; thus different species of protozoa begin their sexual from 35 days before to 2 days after molting of the cockroach (Cleveland and Nutting, 1954). Hollande (1952) and Grasse (1952) have reviewed the roles and the evolution of the flagellates in Cryptocycles cercus and in termites. The protozoa of cockroaches and termites are clues to the relation- ship between these similarities two groups of between Endamoeba insects. hlattae Kirby (1927) pointed out of Periplaneta and the amoebae of the termite Mirotermes, suggesting that these protozoans were probably derived from an amoeba in an ancestor common to both blattid and termite. Kirby (1932, in Kidder, 1937) found a species of Nyctotherus in Amitermes that resembles Nyctotherus ovalis from domestic cockroaches. The belief that the termites and cockroaches had a common origin is also strengthened by the similarities between the hypermastigotes of both Cryptocercus and termites (Cleveland et al, 1934). The cockroaches Cryptocercus and Panesthia both feed on wood, but the protozoa found in Panesthia resemble more closely the species in domestic cockroaches than those in Cryptocercus. The ClevelanNyctotherus and have However, the separation of the Clevelandellidae from Nyctotherus must have taken place at a dellidae ( from Panesthia ) are probably evolved from closely related to common ancestors. later date than the divergence of their hosts, otherwise representatives of that family would probably also be found in Periplaneta and Blatta (Kidder, 1937). The protozoa of Cryptocercus can be transferred from one indi- vidual to another (Nutting and Cleveland, 1954). They can also be BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS IO3 transferred to the termite Zootermopsis where they survive only until the host molts the reverse ; also true, Zootermopsis Protozoa can is (Nutting and survive in Cryptocercus until the cockroach molts Cleveland, 1954a). VI. VIRUSES ASSOCIATED WITH COCKROACHES Annotations on some of the following observations in Roth and Willis (1957a). Use of note 3, page asterisk may be found explained in foot- is 4. POLIOMYELITIS VIRUSES Lansing Experimental vectors. — strain germanica, Blattella U.S.A. (Hurlbutt, 1949, 1950). Periplaneta americana, U.S.A. (Hsiang et Brunhilde type, Minnesota and —Periplaneta Experimental vectors. and Syverton, 1951 Syverton ; et al., vectors. : Mahoney strains americana, U.S.A. (Fischer virus —Periplaneta (Findlay and Howard, 1951) 1952). 1952). Columbia SK Experimental al., americana, Great Britain? Results with Blattella germanica were negative. Four Natural vectors. — unspecified strains Blattella germanica and/or Blattella vaga, Peri- planeta americana and/or Periplaneta brunnea, and Supella supellectilium, and U.S.A. (Syverton et al., 1952; Dow, 1955; Dow in Roth Willis, 1957a). OTHER VIRUSES * Coxsackie viruses Experimental vectors. —Periplaneta americana, U.S.A. (Fischer Recently Fischer and Syverton (1957) single meal of Coxsackie virus to Perifeeding after a found that and Syverton, 1951a, 1957) : planeta americana, the gastrointestinal tracts of the insects, which were removed at 5-day intervals up to 20 days, contained sufficient virus to paralyze and moved kill test mice. 5 days after the insects had Cockroach salivary glands, rewhich fed, contained the virus caused paralysis and death in test mice mice were also infected by virus obtained from salivary glands removed from the insects 10 and ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 104 VOL. I4I 20 days after the cockroaches had fed once on the virus. The virus was also isolated from the cockroaches' feces and rarely from the fat bodies and reproductive organs. Fischer and Syverton concluded that it is possible that cockroaches could acquire the virus, by feeding on mammalian excreta, maintain it for a period of time, and transmit by contamination of food. The virus could also be transmitted through the feces of wild mice if the mice happened to feed on virusit infected cockroaches. * Mouse encephalomyelitis virus Experimental vectors. —Periplaneta anicricana, U.S.A. (Syverton and Fischer, 1950). * Yellow-fever virus Experimental vectors. lay — Blattclla germanica, Great Britain? (Find- and MacCallum, 1939). VII. BACTERIA ASSOCIATED WITH COCKROACHES Breed et al. (1948). Synonymy most cases was taken from the same source. Names of bacteria preceded by the symbol f are either not listed by Breed et al. or are stated by them to be insufficiently characterized for definite classification. Use of asterisk is explained in footnote 3, page 4. Classification of the bacteria follows in Phylum SCHIZOPHYTA SCHIZOMYCETES Class EUBACTERIALES PSEUDOMONADACEAE Order Family * Pseudomonas aeruginosa Natural vectors. —Blab ems (Scliroeter) craniifer, Migula U.S.A. (Wedberg et al., 1949). Blatta orientalis, U.S.A. (Olson and Rueger, 1950). Blattella germanica, U.S.A. (Olson and Rueger, 1950; Janssen and Wedberg, 1952). Periplaneta americana, U.S.A. (Bitter and Williams, 1949, 1949a; Olson and Rueger, 1950). Experimental vectors. Blattella germanica, U.S.A. (Herms and — Nelson, 1913). Cockroaches, U.S.A. (Longfellow, 1913). BIOTIC ASSOCIATIONS OF COCKROACHES * Pseudomonas & WILLIS IO5 Migula eisenbergii Natural and experimental vectors. —ROTH —Blatta (Cao, orientalis, Italy 1906). * Pseudomonas Migula fluorescens Natural and experimental vectors. —Blatta (Cao, orientalis, Italy and Reitano, 1932). Periplaneta americana, U.S.A. (Gier, 1947) 1898, 1906; Spinelli pathogenic to the cockroach when The organism was : injected. t Spirillum periplaneticum Kunstler and Gineste Habitat. —Periplaneta americana, 1906) : From France? (Kunstler and Gineste, intestinal tract. t Spirillum a, Habitat. —Blatta orientalis, /3, and 7 Dobell England (Dobell, 191 1, 1912) From : hind gut. Spirillum Habitat. —Blatta orientalis, sp. U.S.S.R. (Zasukhin, 1930) From : intestinal tract. Cockroaches, Venezuela (Tejera, 1926) : From Vibrio comma (Schroeter) Winslow Experimental vectors. and Reitano, 1932). Blattella germanica. —Blatta Orient orientalis, Italy (Toda, 1923) ; digestive tract. et al. (Cao, 1898; SpinelH Germany (Jettmar, 1927). Periplaneta americana, Philippine Islands (Barber, 1914) NetherFormosa (Morischita and Tsuchimochi, ; lands (Akkerman, 1933) ; 1926). Periplaneta australasiae, Formosa (Morischita and Tsuchimochi, 1926). * Vibrio metschnikovii Gamaleia Experimental vectors. — Blatta orientalis, Italy (Cao, 1898, 1906). Vibrio tyrogenus (Fliigge) Holland Synonymy. — Vibrio of Deneke. Experimental vectors. Blatta orientalis, Italy (Cao, 1898) organism passed through the intestinal tract unchanged. — : The SMITHSONIAN MISCELLANEOUS COLLECTIONS I06 t Vibrio Types — and I II VOL. I4I Heiberg Water. Natural vectors. Cockroaches, India (Pasricha et al., 1938) The vibrios were found in 16 or 17 percent of 94 cockroaches and reHabitat. — sembled Vibrio reactions : comma morphology and in their their main biochemical however, serum-agglutination reactions differed. ; Vibrio Habitat. —Blatta orientalis, Family U.S.A. (Leidy, 1853) Habitat. —B. —Water. : From intestine. RHIZOBIACEAE Chromobacterium violaceum Synonymy. sp. (Schroeter) Bergonzini violaceus. — Experimental vectors. Cockroach, U.S.A. (Longfellow, 1913) Recovered from outer part of body and intestinal tract. Family : MICROCOCCACEAE Micrococcus aurantiacus (Schroeter) Cohn — Natural vectors. Blattella germanica, Wed- U.S.A. (Janssen and berg, 1952). * Micrococcus Natural vectors. —Cockroaches, —Blatta Experimental vectors. — Migula U.S.A. (Longfellow, 1913). orientalis, Italy * Micrococcus epidermidis Natural vectors. citrous (Cao, 1898). (Winslow and Winslow) Hucker Blattella germanica, U.S.A. (Janssen and Wed- berg, 1952). t Micrococcus nigrofaciens Northrup — Source. Diseased June beetle larvae. Experimental infection. Periplaneta americana, U.S.A. (NorthThree of four adults were infected by feeding them rup, 1914) bread saturated with a broth culture of the Micrococcus. After 11 — : tarsi of the cockroaches became infected, and the hind legs and broke off. Antennae and setae also were affected and micrococci were recovered from the feces. days the split * Micrococcus Natural 1949). pyogenes var. albus (Rosenbach) Schroeter vectors. —Blaberus craniifer, U.S.A. (Wedberg et al., BIOTIC ASSOCIATIONS OF COCKROACHES & WILLIS 1940; Tauber and (Tauber, Blatta orientalis, U.S.A. —ROTH IO7 Griffiths, 1942). Blattella germanica, U.S.A. (Herms and Nelson, 1913; Herms, 1939; Janssen and Wedberg, 1952). — Experimental vectors. Blatta orientalis, Italy (Cao, 1898) U.S.A. (Tauber and Griffiths, 1942). Micrococcus pyogenes var. albus ( = Staphylococcus albus) and an unidentified short rod form were found by Tauber (1940) in the hemolymph of B. orientalis. These microorganisms were never found ; together in the same insect and caused loss of appetite, sluggishness, irregular respiratory movements, and paralysis in the cockroach ; in the final stages of the disease the legs were folded under the body, the head was tucked beneath the forelegs, the whole insect became arched and maintained this position until death. In some cockroaches infected with the rod pathogen, conjunctival folds, particularly those between the dorsal abdominal and the sclerites, thoracic legs ruptured liberating thick white meta- joints of the hemolymph with filled Tauber suggested that the infection might be spread by contact, especially to newly molted individuals or by actual ingestion of the bacteria by the cockroaches feeding on dead or dying indibacteria. viduals. All the roaches died after successful inoculation with the Micrococcus. The bacterial infection was associated with high total hemocyte counts and high percentages of mitotically dividing hemolymph cells (Tauber, 1940) these responses of the insect were in; mechanism whereby the number of hemocytes increases resulting in an increase in the number of phagocytes for combating the bacteria (Tauber and Griffiths, 1942). terpreted as a * Micrococcus pyogenes var. Natural vectors. —Blaberus (Rosenbach) aureus U.S.A. craniifer, Zopf (Wedberg et al., 1949). Blatta orientalis, Italy (Cao, 1906). U.S.A. (Herms, 1939). Cockroaches, U.S.A. (Longfellow, 1913). Blattella germanica, Cohn Micrococcus ureae Habitat. — Stale Natural vectors. berg, 1952) : urine and — From soil containing urine. Blattella germanica, intestinal tract and Micrococcus U.S.A. (Janssen and Wed- feces. spp. These organisms were obtained from pus or were designated as staphylococci [i.e., pathogenic micrococci (Blair in Dubos, 1948)]. SMITHSONIAN MISCELLANEOUS COLLECTIONS I08 Natural vectors. —Blatta (Spinelli Italy orientalis, VOL, I4I and Reitano, 1932); Germany (Jettmar, 1935). Germany Blattella germanica, — (Jettmar, 1935). Experimental vectors. Blattella germanica, on shipboard (MorGermany (VoUbrechtshausen, 1953). rell, 191 1 ) ; Micrococcus Natural vectors. sp. —Periplaneta americana? England (Shrewsbury and Barson, 1948) : {"Blatella americana"), From intestinal tract. fSarcina alba Zimmermann — Habitat. Water. Natural and experimental vectors. 1898, 1906) From : —Blatta orientalis, Italy (Cao intestinal contents. Sarcina aurantiaca Flugge Habitat. —Air and water. —Blatta Experimental vectors. orientalis, Italy (Cao, 1906) Intes- : tinal contents. Sarcina lutea Schroeter Habitat. —Air, soil, water, skin surfaces. Experimental vectors. —Blatta orientalis, Italy (Cao, 1906) : From intestinal contents. t Sarcina Habitat. symbiotica —Oothecae of Blatta Pribram orientalis Germany (Gropengiesser, 1925) Sarcina" ; Pribram (1933) named and/or Blattella germanica, was described It : as "eine gelbe the organism. Sarcina ventriculi Goodsir — Garden Natural vectors. —Blatta Habitat. From soil, dust, mud ; from a diseased stomach. Poland (Nicewicz et al., 1946) isolated orientalis, : intestinal tract. Sarcina sp. —Periplaneta americana, U.S,A, Natural and experimental vectors. Organism not pathogenic to the cockroach when (Gier, 1947) : in- jected. fSarcina Synonymy. Clerc. Natural vectors. 1908) : sp. —Sarcina "blanche" of Sartory and —Blatta France (Sartory Isolated from orientalis, intestinal tract. and Clerc, BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS IO9 * t Sarcina sp. Synonymy. —Sarcina alba "patogena" of Cao. —Blatta (Cao, 1898, 1906). Natural vectors. orientalis, Italy * t Sarcina sp. Synonymy. —Sarcina "bianca" and "gialla" of Cao. —Blatta (Cao, 1898). Experimental vectors. orientalis, Italy NEISSERIACEAE Family Neisseria meningitidis (Albrecht and Ghon) Holland Experimental vectors. — Cockroaches, U.S.A. (Longfellow, 1913). Veillonella parvula (Veillon and Zuber) Prevot Natural vectors. —Periplaneta americana, U.S.A. Family LACTOBACTERIACEAE * Diplococcus Experimental vectors. (Hatcher, 1939). pneumoniae Weichselbaum —Blatta orientalis, Italy (Cao, 1898). Cockroaches, U.S.A. (Longfellow, 191 3). t Enterococcus Natural vectors. 1932) : From —Blatta orientalis, sp. Italy (Spinelli and Reitano, intestinal tract. Lactobacillus fermenti Beijerinck Habitat. —Fermenting plant or animal products. —Blatta Poland (Nicewicz Natural vectors. From orientalis, et al., 1946) : intestinal canal. t Pneumococcus Type Experimental vectors. — I, No. 1231 Blattella germanica, Germany ( Vollbrechts- hausen, 1953). * Streptococcus faecalis Natural vectors. —Blatta Andrewes and Horder orientalis, Poland (Nicewicz et al., 1946). U.S.A. (Steinhaus, 1941). Periplaneta americanaf {"Blatella americana"), England (Shrewsbury and Barson, 1948). Cockroaches [presumably any or all of the above three species], Egypt (El-Kholy and Gohar, 1945). Blattella germanica, no SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I * Streptococcus liquefaciens Sternberg emend. Orla-Jensen Natural vectors. —Blatta orientalis, Poland (Nicewicz et al., 1946). t Streptococcus microapoika Cooper, Keller, and Johnson Natural vectors. —Blatta orientalis, Poland (Nicewicz * t Streptococcus non-hemolyticus II Natural vectors. —Periplaneta ainericana? et al., 1946). Holman {"Blatella americana"), England (Shrewsbury and Barson, 1948). Streptococcus pyogenes Rosenbach — Natural vectors. Blatta orientalis, Italy (Cao, 1906). Experimental vectors. Cockroaches, U.S.A. (Longfellow, 1913). — Streptococcus sp. Experimental vectors. —Blatta Streptococcus Experimental vectors. —Blatta (pyogenic group) sp. —Blatta Germany orientalis, orientalis, Germany (Jettmar, 1935). {viridans group) Streptococcus Natural vectors. Germany orientalis, (Jettmar, 1935). spp. Germany (Jettmar, 1935). U.S.A. (Janssen and Wedberg, 1952). Cockroaches, U.S.A. (Longfellow, 1913). Experimental vectors. Blatta orientalis, Italy (Cao, 1898). Blattella germanica, (Jettmar, 1935) ; — Family CORYNEBACTERIACEAE Corynebacterium diphtheriae Experimental vectors. —Blatta Lehmann and Neumann (Flugge) orientalis, Italy (Cao, 1898). Cockroaches, U.S.A. (Longfellow, 1913). Family ACHROMOBACTERIACEAE t Achromobacter hyalinum (Jordan) Bergey et al. — Natural vectors. Periplaneta americana, U.S.A. (Hatcher, 1939) Isolated from feces. Achromobacter Natural vectors. berg, 1952). — : sp. Blattella germanica, U.S.A. (Janssen and Wed- BIOTIC ASSOCIATIONS OF COCKROACHES Alcaligenes faecalis Synonymy. —Bacillus calis; Castellani —ROTH & WILLIS III and Chalmers faecalis alkaligenes; Bacillus alcaligenes fae- B. alcaligenes faecalis. Habitat. — Intestinal canal of man. Has been isolated from feces, abscesses related to intestinal canal, and occasionally in the blood- stream. However, Natural vectors. it is generally considered nonpathogenic. —Blaherus : U.S.A. craniifer, from feces. Blatta orientalis, Poland (Nicewicz 1949) (Wedberg et al., Isolated et al., 1946) from Isolated : intestinal tract. Blattella germanica, intestinal tract and U.S.A. (Janssen and Wedberg, 1952) From : feces, Periplaneta americana, U.S.A. (Bitter and Williams, 1949) lated from Iso- : intestinal tract. Cockroaches [presumably Blatta orientalis, Blattella germanica and/or Periplaneta americana], Egypt (El-Kholy and Gohar, 1945) From suspensions of macerated whole insects. : Alcaligenes recti Synonymy. —B. alcaligenes Habitat. — —Blatta Natural (Ford) Bergey et al. recti. Intestinal canal. orientalis, vectors. Isolated from Poland (Nicewicz et al., 1946) : intestinal tract. Alcaligenes viscosus (Weldin and Levine) Weldin Habitat. —Water, dairy — Natural vectors. berg, 1952) : utensils ; produces ropiness in milk. Blattella germanica, Isolated Family from intestine and Wed- U.S.A. (Janssen and feces. ENTEROBACTERIACEAE Aerobacter aerogenes (Kruse) aerogenes. Synonymy. —Bacillus Habitat. — Grains, —Blaberus Natural Beijerinck lactis plants, intestinal tract of vectors. craniifer, from feces. Blatta orientalis, Poland (Nicewicz 1949) : man and U.S.A. other animals. (Wedberg et al., Isolated et al., 1946) : Isolated from Isolated from intestinal tract. Blattella germanica, on shipboard (Morrell, 191 1) : feces, Periplaneta americana, U.S.A, (Bitter and Williams, 1949) lated from intestinal tract. : Iso- SMITHSONIAN MISCELLANEOUS COLLECTIONS 112 Cockroaches [presumably Blatta orientalis, VOL. I4I Blattella germanica and/or Periplaneta americana], Egypt (El-Kholy and Gohar, 1945) Isolated from outer surface of body, intestinal tract, and suspensions : of macerated whole insects. (Jordan) Aerobacter cloacae —Bacillus — Sewage, — Natural Synonymy. Habitat. Bergey et al. cloacae. soil, water, human and other animal feces. on shipboard (Morrell, From feces. U.S.A. ( Janssen and Wedberg, 1952; Steinhaus, 191 1 ) From intestinal tract, feces, and ootheca. 1941 ) Periplaneta americana, U.S.A. (Bitter and Williams, 1949) From Blattella vectors. germanica, : : : intestinal tract. Aerobacter Natural vectors. liams, 1949) : sp. —periplaneta americana, U.S.A. from Isolated (Bitter and Wil- intestines. tEberthella oedematiens Assis Habitat. — Intestinal canal. Natural vectors. liams, 1949) : —Periplaneta americana, U.S.A. (Bitter and Wil- Intestinal tract. * Escherichia coli (Migula) Castellani and Chalmers Natural vectors. —Blatta orientalis, Italy (Cao 1898, 1906; Spinelli and Reitano, 1932) France (Sartory and Clerc, 1908) Europe (Jettmar, 1935) Poland (Nicewicz et al., 1946). Blattella germanica, U.S.A. (Steinhaus, 1941). Periplaneta americana, U.S.A. (Bitter and Williams, 1949, 1949a). Cockroaches [presumably one or all of the above three species], Egypt (El-Kholy and Gohar, 1945). Cockroaches, U.S.A. (Longfellow, 1913). Experimental vectors. Blattella germanica, Germany ( Vollbrechts; ; ; — hausen, 1953) : When injected anally or orally the bacteria invaded the intestinal cells and in heavy infections killed the cockroaches. Escherichia coli var. acidilactici Synonymy. —Bacillus Source. — Diseased nun moth —Blatta Experimental (Topley and Wilson) Yale acidi lactici. larvae. vectors. Organism pathogenic orientalis, to cockroach when Europe (Filatoff, injected but not when 1904) fed. : BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS II3 Escherichia coli var. communior (Topley and Wilson) Yale (Wedberg Natural vectors.— Blaherus craniifer, U.S.A. 1949) : From — (Braak) Yale Soil, water, intestinal canal of Natural vectors.— Blaherus craniifer, 1949) : al., feces. Escherichia freundii Habitat. et From man and other animals. U.S.A. (Wedberg et al., feces. Blattella germanica, intestinal canal and U.S.A. (Janssen and Wedberg, 1952) : From : From feces. Periplaneta americana, U.S.A. (Bitter and Williams, 1949) intestinal tract. Escherichia intermedium (Werkman and Gillen) Vaughn and Levine Hahitat.—SoW, water, intestinal canal of man and other animals. Natural vectors. Periplaneta americana, U.S.A. (Bitter and Wil- — liams, 1949) From : intestinal canal. Klebsiella pneumoniae (Schroeter) Trevisan Experimental vectors. * —Blatta orientalis, Italy (Cao, 1898). Paracolobactrum aerogenoides Borman, Stuart and Wheeler Natural vectors.— Blattella germanica, U.S.A. (Janssen and Wed- berg, 1952). Periplaneta americana, U.S.A. (Bitter and Williams, 1949). Paracolobactrum coliforme Borman, Stuart and Wheeler Natural vectors. — Blattella germanica, U.S.A. (Janssen and Wed- berg, 1952). Periplaneta americana, U.S.A. (Bitter and Williams, 1949)- Paracolobactrum Natural vectors.—Blatta orientalis, spp. U.S.A. (Olson and Rueger, 1950)- U.S.A. (Olson and Rueger, 1950). Periplaneta americana, U.S.A. (Bitter and Williams, 1949; Olson Blattella germanica, and Rueger, 1950). Paracolon — bacilli Natural vectors. Cockroaches [presumably Blatta orientalis, Blattella germanica and/or Periplaneta americana], Egypt (El-Kholy and Gohar, 1945)- SMITHSONIAN MISCELLANEOUS COLLECTIONS 114 * Proteus mirabilis VOL. I4I Hauser Natural vectors. —Periplaneta americana, U.S.A. (Bitter and Wil- liams, 1949, 1949a). Proteus morganii (Winslow Natural vectors. Rauss et al.) —Periplaneta americana, U.S.A. (Bitter and Wil- liams, 1949, 1949a). Cockroaches [presumably Blatta orientalis, Blattella germanica, and/or Periplaneta americana], Egypt (El-Kholy and Gohar, 1945). (Hadley * Proteus rettgeri Rustigian and Stuart et al.) Natural vectors. —Periplaneta americana, U.S.A. (Bitter and Wil- liams, 1949). Proteus Natural vectors. — Blaherus vulgaris Hauser U.S.A. craniifer, (Wedberg et al., 1949). Blatta orientalis, Italy (Spinelli and Reitano, 1932). Periplaneta americana, U.S.A. (Bitter and Williams, 1949, 1949a). Cockroaches, U.S.A. (Longfellow, 1913). Proteus Natural vectors. —Blatta spp. orientalis, (Olson and Rueger, U.S.A. 1950). Periplaneta americana, U.S.A. (Bitter and Williams, 1949; Olson and Rueger, 1950). Salmonella anatis (Rettger and Natural vectors. —Periplaneta Bergey Scoville) americana, U.S.A. et al. (Eads et al., 1954). Salmonella Experimental vectors. choleraesuis —Polyphaga (Smith) Weldin saussurei, (Zmeev U.S.S.R. in Pavlovskii, 1948). Salmonella enteritidis (Gaertner) Castellani and Chalmers Experimental vectors. —Blatta orientalis, U.S.S.R. (Rozengolts and ludina in Pavlovskii, 1948). Blattella germanica, (Rozengolts and U.S.A. (Olson and Rueger, 1950) ; U.S.S.R. ludina in Pavlovskii, 1948). Polyphaga saussurei, U.S.S.R. (Zmeev in Pavlovskii, 1948). 5 BIOTIC ASSOCIATIONS OF * Salmonella COCKROACHES morbificans (Migula) —Periplaneta americana, and Mackerras, 1948, 1949). —Nauphoeta Experimental Natural vectors. & WILLIS II Haupt Australia (Mackerras cinerea, Periplaneta australa- vectors. siae, —ROTH Periplaneta ignota, and Supella supellectilium, Australia (Mac- kerras and Pope, 1948). * Salmonella paratyphi Experimental vectors. (Kayser) —Polyphaga and Chalmers Castellani saussiirei, U.S.S.R. (Zmeev in Pavlovskii, 1948). Salmonella schottmuelleri (Winslow Natural vectors. et al.) Bergey —Periplaneta americana, U.S.A. et al. (Bitter and Wil- liams, 1949, 1949a). Experimental vectors. Periplaneta americana, Gold Coast Colony (Macfie, 1922). Polyphaga saussurei, U.S.S.R. (Zmeev * Salmonella Experimental vectors. siae, in Pavlovskii, 1948). (Type Adelaide) sp. —Nauphoeta cinerea, Periplaneta aiistrala- Periplaneta ignota, and Supella supellectilium, Australia (Mac- kerras and Pope, 1948). * Natural vectors. Salmonella (Type Bareilly) sp. —Periplaneta americana, U.S.A. (Eads et al., 1954). * Salmonella sp. Natural vectors. (Type Bredeny) —Periplaneta americana, U.S.A. (Bitter and Wil- liams, 1949, 1949a). * Salmonella sp. (Type Derby) —Nauphoeta Experimental vectors. cinerea, Periplaneta australaand Supella supellectilium, Australia (Mackerras and Pope, siae, 1948). * Salmonella sp. Natural vectors. —Periplaneta (Type Kentucky) americana, U.S.A. (Eads et al., 1954). Salmonella Experimental vectors. and Pope, 1948). erras sp. (Type Kottbus) —Periplaneta australasiae, Australia (Mack- SMITHSONIAN MISCELLANEOUS COLLECTIONS Il6 (Type Meleagris) * Salmonella sp. Natural VOL. I4I vectors. —Periplaneta americana, U.S.A. (Eads et al., 1954). (Type Montevideo) * Salmonella sp. Experimental vectors. —Periplaneta americana, U.S.A. (Jung and Shaffer, 1952). (Type Newport) * Salmonella sp. —Periplaneta Natural vectors. americana, U.S.A. (Eads et al., 1954). * Salmonella sp. (Type Oranienburg) Natural vectors. —Periplaneta americana, U.S.A. liams, 1949, 1949a; Eads 1954). —Blatta Experimental (Bitter and Wil- et al., orientalis, Blattella vectors. germanica, and Periplaneta americana, U.S.A. (Olson and Riieger, 1950). * Natural vectors. Salmonella (Type Panama) sp. —Periplaneta americana, U.S.A. (Eads et al., (Eads et al., (Eads et al., 1954). Salmonella (Type Rubislaw) sp. —Periplaneta americana, * Salmonella sp. (Type Tennessee) Natural vectors. U.S.A. 1954). Natural vectors. —Periplaneta americana, U.S.A. 1954). * Salmonella typhimurium Natural vectors. — (Loeffler) Blattella germanica, Castellani and Chalmers Belgium (Graffar and Mer- tens, 1950). Nauphoeta cinerea, Australia Experimental vectors. (Mackerras and Mackerras, 1948). craniifer, U.S.A. (Wedberg et al., —Blaberus ^ 1949)Blatta orientalis, U.S.S.R. (Rozengolts and ludina in Pavlovskii, _ 1948). Belgium (Graffar and Mertens, 1950) U.S.A. (Olson and Rueger, 1950; Janssen and Wedberg, 1952; Beck and Coffee, 1943) U.S.S.R. (Rozengolts and ludina in Pavlovskii, 1948). Nauphoeta cinerea, Australia (Mackerras and Pope, 1948). Periplaneta americana, U.S.A. (Beck and Coffee, 1943; Jung and Blattella germanica, ; Shaffer, 1952). ; BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH II7 & WILLIS Periplaneta australasiae and Supella supellectilmm, Australia (Mackerras and Pope, 1948). Polyphaga saussitrei, U.S.S.R. (Zmeev in Pavlovskii, 1948). * Salmonella typhosa (Zopf ) White Natural vectors. — Blatta Experimental vectors. — Blaberus orientalis, Italy (Antonelli, 1930, 1943). craniifer, U.S.A. (Wedberg et 1949). U.S.A. (McBlatta orientalis, Italy (Spinelli and Reitano, 1932) Burney and Davis, 1930) U.S.S.R. (Rozengolts and ludina in Pav- al., ; ; lovskii, 1948). Blattella germanica, many (Jettmar, 1927) U.S.A. (Janssen and Wedberg^ 1952) GerU.S.S.R. (Rozengolts and ludina in Pav; ; lovskii, 1948). Periplaneta americana, Gold Coast Colony (Macfie, 1922) ; Nether- (Akkerman, 1933) Formosa (Morischita and Tsuchimochi, 1926) U.S.A. (Olson in Roth and WilHs, 1957a). Periplaneta australasiae, Formosa (Morischita and Tsuchimochi, lands ; ; 1926). Polyphaga saiissurei, U.S.S.R. (Zmeev in Pavlovskii, 1948). Cockroaches [presumably Blatta orientalis, Blattella germanica, and/or Periplaneta americana], Egypt (El-Kholy and Gohar, 1945). Serratia marcescens Bizio Synonymy. —Bacillus prodigiosus. Bacterium prodigiosum. milk, foods, and various Habitat. —Water, —Blatta Poland (Nicewicz Natural insects. soil, hosts. From orientalis, et al., 1946) : and Reitano, 1932). Blattella germanica, Canada (Heimpel and West, 1959). Diploptera punctata, Nauphoeta cinerea, Neostylopyga rhomhifolia, Panchlora nivea, Pycnoscelus surinamensis, and Supella supellectilium, The organism U.S.A. (Roth and Willis, unpublished data, 1958) vi^as isolated and identified by Dr. Hillel Levinson, Quartermaster bacteriologist, from dead specimens found in our laboratory colonies w^hich showed the red coloration characteristic of insects that have died with infections of 6". marcescens (pi. 16, A, B). Leucophaea maderae, U.S.A. (Levinson, personal communication, The organism was isolated from the hemolymph of living 1958) insects while attempting to determine the cause of unexplained morintestinal tract. Italy (Spinelli : : tality in our laboratory colony of this insect. Leucophaea maderae or Periplaneta americana, Philippine Islands From hemolymph. (Barber, 1912) : SMITHSONIAN MISCELLANEOUS COLLECTIONS Il8 VOL. I4I Periplaneta americana, U.S.A. (Gier, 1947; Steinhaus, 1959). Periplaneta australasiae and Periplaneta brunnea (Roth and Willis, In laboratory colonies. Isolated from susunpublished data [1954] ) pensions of ground insects. In 1954 we received a culture of Peri: planeta brunnea from the Department of Public Health, University of off rapidly and the normally pigmented parts of the body became red. Dr. Hillel Levinson, Quartermaster bacteriologist, cultured Serratia marcescens from sev- Minnesota. These insects began to die lightly eral moribund individuals. The Department of Public Health of Min- nesota had at times in the past cultured S. marcescens but had dis- carded the cultures and was unaware that it might be surviving in the cockroach colonies (Richards, personal communication, 1954). Isosp., U.S.A. (Olson in Roth and Willis, 1957a) from an undetermined species of Periplaneta, received in a shipment from the South, a strain of 5". marcescens which was toxic Periplaneta : lated to mice when administered Experimental hosts. intraperitoneally. —Blaberus craniifer, U.S.A. (Wedberg et al., When fed in small numbers, 6". marcescens increased to such 1949) an extent that the insect's extremities and upper halves of their bodies : turned deep red. tically The insects died after this color appeared and prac- pure cultures of Serratia were recovered from the reddened areas. Blatta orientalis, Italy (Cao, 1898) tents. : from Isolated intestinal con- Passed unchanged through the gut. Blattella germanica, Canada (Heimpel and West, 1959) mally pathogenic per os; LD50 by injection, is : Not nor- approximately 38,000 bacteria per insect. Periplaneta americana, U.S.A. (Gier, 1947) the cockroach when : Organism toxic to injected. Cockroaches, U.S.A. (Longfellow, 1913) : Isolated from legs and viscera after feeding experiments. * Shigella alkalescens (Andrewes) Weldin Natural vectors. —Periplaneta americana, U.S.A. (Bitter and Wil- liams, 1949, 1949a). Shigella dysenteriae (Shiga) Experimental vectors. —Blatta Castellani and Chalmers orientalis, Italy (Spinelli and Rei- tano, 1932). Periplaneta americana, Formosa (Morischita and Tsuchimochi, 1926). Polyphaga saussurei, U.S.S.R. (Zmeev in Pavlovskii, 1948). BIOTIC ASSOCIATIONS OF COCKROACHES * Shigella paradysenteriae —ROTH (Collins) & WILLIS IIQ Weldin —Blatta Tadzhikistan (Zmeev, 1940). Experimental vectors. —Periplaneta americana, Gold Coast Colony Natural vectors. lateralis, (Macfie, 1922). Polyphaga saussurei, U.S.S.R. (Zmeev in Pavlovskii, 1948). Cockroaches, Venezuela (Tejera, 1926). PARVOBACTERIACEAE Family Bacteroides uncatus Habitat. — Probably —Blatta intestinal canal of Natural vectors. From Eggerth and Gagnon orientalis, mammals from human feces. ; Poland (Nicewicz et al., 1946) : intestinal canal. Brucella abortus (Schmidt and Weis) Meyer and Shaw Experimental vectors. and Huddleson, 1941). —Periplaneta (Ruhland americana, U.S.A. Fusifonnis lophomonadis Grasse Habitat. — Surface of a flagellate (Lophomonas (Breed in the intestine of cockroaches et al., striata) which lives 1948; Grasse 1926, 1926a). Malleomyces mallei (Zopf ) * Experimental vectors. Experimental vectors ; —Blatta (Cao, 1898). Germany orientalis, (Kiister, 1902, Italy (Cao, 1906). (Lehmann and Neumann) Holland * Pasteurella pestis Natural vectors. —Blatta — Blatta orientalis, Experimental vectors. many orientalis, Italy (Lehmann and Neumann) Rosenbusch and Merchant * Pasteurella multocida 1903) —Blatta Pribram Hongkong (Hunter, orientalis, Italy 1906). (Cao, 1898) ; Ger- (Kiister, 1903). Blattella germanica, Germany (Jettmar, 1927). Leucophaea maderae and Periplaneta americana, Philippine Islands (Barber, 1912). Family BACTERIACEAE t Bacterium alkaligenes Natural vectors. Nyberg —Periplaneta americana? England (Shrewsbury and Barson, 1948) : {"Blatella americana"), From intestinal tract. SMITHSONIAN MISCELLANEOUS COLLECTIONS 120 delendae-muscae Roubaud and Descazeaux t Bacterium Source. —Diseased fly larvae. Experimental infection. cazeaux, 1923) : VOL. I4I — Cockroach, Organism pathogenic France (Roubaud and Desto cockroach when injected. Bacterium haemophosphoreum Pfeiffer and Stammer Habitat. — Diseased larvae of Mamestra — oleracea. Experimental infection. Blatta orientalis and Blattella germanica, Germany (Pfeiffer and Stammer, 193 1) Organism pathogenic, when injected, to eight B. orientalis and two B. germanica. : Coccobacillus cajae Picard and Blanc Experimental 1913) : host. —Blatta orientalis, The organism was pathogenic Experimental vectors. Germany injected. Cohen emend. Koch anthracis —Blatta orientalis, Italy (Cao, 1898, 1906) ; (Kiister, 1903). t Bacillus butschlii Habitat. from when BACILLACEAE Family * Bacillus France (Picard and Blanc, to B. orientalis —Blatta Germany (Schaudinn, 1902) orientalis, intestinal tract. Schaudinn Isolated : Three percent of the cockroaches from Berlin bakeries were infected. Bacillus cereus Frankland and Frankland —Bacillus vectors. —Blabcnis Synonymy. Habitat. — Natural albolactis. Soil, dust, milk, plants. craniifcr, (Wedberg U.S.A. From feces. 1949) Periplaneta americana, U.S.A. (Hatcher, 1939) Experimental host. Periplaneta americana, et al., : — : In feces. U.S.A. (Babers, The cockroaches died within 96 hours after being injected 1938) with io~^ ml. of a 24-hour broth culture. : Bacillus circulans Jordan Habitat. — Soil, water, dust. Natural vectors. berg, 1952) : — From Blattella germanica, intestine and feces. U.S.A. (Janssen and Wed- BIOTIC ASSOCIATIONS OF COCKROACHES t Bacillus flacheriae — —ROTH & WILLIS 121 (Hoffman) Source. Diseased nun moth larvae. (Filatoff, Experimental infection. Blatta orientalis, Europe The organism was not pathogenic when fed to the cockroach, 1904) but killed the insects when injected into the body cavity; after the insects died Filatoff reisolated this pathogen together with another — : from the cadavers. He succeeded in culturing the new microorganism and found it to be pathogenic when injected into, but not when fed to, the cockroaches. The diseased insects became sluggish, failed to eat or drink, turned over on their backs, their extremities became totally paralyzed, and they finally died. bacillus Bacillus Habitat. — Soil, water, megaterium De Bary decomposing materials. —Periplaneta americanaf England (Shrewsbury and Barson, 1948) From —Blatta Experimental {"Blatella americana"), Natural vectors. : intestinal tract. orientalis, Italy vectors. ganism recovered, apparently unchanged, from t Bacillus monachae (Cao, 1898) : Or- intestinal contents. (von Tubeuf) Eckstein Synonymy. —Bacterium monache. Source. — Diseased larvae of nun moth, Lymantria monacha. —Blatta Europe Experimental 1904) (Filatoff, orientalis, infection. Organism pathogenic to the cockroach when injected but not when : fed. t Bacillus periplanetae Tichomiroff Habitat. —Blatta Filatoff, 1904) : orientalis, The U.S.S.R. ? (Tichomiroff, i87o[?], in infected insects suffered from a diarrhea and the liquid feces were yellow-brown. t Bacillus Natural infection. — Blatta Organism observed regularly stellatus Hollande orientalis, in France the intestine (Hollande, 1934) : (especially rectum). Extensive description given. t Bacillus radiciformis —Blatta orientalis, Italy (Cao 1898) OrExperimental vectors. ganism recovered, apparently unchanged, from intestinal contents. Bacillus Natural vectors. 1949). subtilis —Blaberus : Cohn emend. Prazmowski craniifer, U.S.A. (Wedberg et al., SMITHSONIAN MISCELLANEOUS COLLECTIONS 122 Blatta orientalis, Italy 1932) ; VOL. I4I (Cao, 1898, 1906; Spinelli and Reitano, Poland (Nicewicz et al., France (Sartory and Clerc, 1908) ; 1946). Cryptocercus punctulatus, v. S.A. (Hatcher, 1939). Periplaneta americana? {"Blatella americana") , England (Shrews- bury and Barson, 1948). Cockroaches, U.S.A. (Longfellow, 1913). Experimental vectors. Blatta orientalis, Italy (Cao, 1898, 1906). — "Bacillus subtilis group" Natural infection. lated —Blatta orientalis, Italy (Ronzoni, 1949) : Iso- from oothecae. t Bacillus Habitat. —Isolated from —Blatta Natural vectors. Isolated from tritus Batchelor (man?). Poland (Nicewicz feces orientalis, et al., 1946) : 1946) : intestinal tract. * Clostridium Experimental vectors. —Blatta f eseri Trevisan orientalis, lisXy (Cao, 1898). Clostridium lentoputrescens Hartsell and Rettger Habitat. — Soil, intestinal tract of Natural vectors. Isolated from —Blatta man. Poland (Nicewicz orientalis, et al., intestinal tract. * Clostridium novyi Clostridium sporogenes Natural and experimental (Migula) Bergey et al. or (Metchnikoff) Bergey et vectors. —Blatta al. orientalis, Italy (Cao, 1898). * Clostridium Natural vectors. tella perfringens — Cockroaches ( Veillon and Zuber) [presumably Blatta Holland orientalis, Blat- germanica, and/or Periplaneta americana], Egypt (El-Kholy and Gohar, 1945). Clostridium tetani (Fliigge) Holland Experimental vectors. —Blatta orientalis, Italy (Cao, 1898). * Clostridium spp. Natural vectors. —Periplaneta americana? England (Shrewsbury and Barson, 1948). {"Blatella americana"), BIOTIC ASSOCIATIONS OF COCKROACHES — ROTH Order ACTINOMYCETALES Family MYCOF.ACTERIACEAE * & WILLIS I23 Mycobacterium avium Chester Experimental vectors. —Blatta orientalis, U.S.S.R. (Ekzenipliar- skaia in Pavlovskii, 1948). Mycobacterium friedmannii Holland — Parasitic and possibly sparingly distributed in Natural vectors. — Pcriplaneta americana, U.SA., Texas (Micks, Habitat. in turtles soils. in Roth and Willis, 1957a) testinal tracts of * Mycobacterium Natural vectors. * : Organism isolated lacticola in- Lehmann and Neumann? —Pcriplaneta americana, U.S.A. (Leibovitz, 1951). Mycobacterium leprae (Armauer-Hansen) Natural vectors. Kenya (Moiser, from batches of cockroaches collected at random. — Blattella germanica, Lehmann and Neumann Southern Rhodesia and Anonymous, 1946). Periplaneta americana and Pcriplaneta australasiae, Formosa (Ari1945, 1946, 1946a; zumi, 1934, 1934a). Cockroaches, Venezuela (Tejera, 1926); Belgian Congo (Radna, 1939)- Experimental vectors. — Blatta orientalis, Europe (Paldrock in Nyasaland (Lamborn, 1940). Blattella germanica, Europe (Paldrock in Klingmiiller, 1930) Southern Rhodesia and Keyna (Moiser, 1945, 1946, 1946a, 1947; KlingmiJller, 1930); ; Anonymous, 1946). Nauphoeta cinerea, Nyasaland (Lamborn, 1940). Periplaneta americana. Gold Coast Colony (Macfie, 1922) mosa (Arizumi, Periplaneta australasiae, Formosa (Arizumi, Cockroaches, Belgian Congo (Radna, 1939) 1926). * ; For- 1934, 1934a). 1934, 1934a). ; Venezuela (Tejera, Mycobacterium lepraemurium Marchoux and Sorel Experimental vectors. —Cockroaches, Belgian Congo (Radna, 1939)* Mycobacterium phlei Lehmann and Neumann Natural vectors. Micks in —Periplaneta americana, U.S.A. Roth and Willis, 1957a). (Leibovitz, 1951; ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 124 * Natural vectors. Experimental Mycobacterium piscium Bergey et VOL. I4I al. —Periplaneta americana, U.S.A. (Leibovitz, 1951). —Blatta U.S.S.R. (Ekzempliar- vectors. orientalis, skaia in Pavlovskii, 1948). * Mycobacterium tuberculosis Experimental vectors. many (Kiister, 1903) —Blatta (Cao, 1898) Ger(Ekzempliarskaia in Pavlovskii, orientalis, Italy U.S.S.R. ; Lehmann and Neumann (Schroeter) ; 1948). Blattella germanica, on shipboard (Morrell, 1911). Periplaneta americana, Gold Coast Colony (Macfie, 1922). Cockroaches, Venezuela (Tejera, 1926) * Mycobacterium Natural vectors. Micks in ; U.S.A. (Read, 1933). spp. —Periplaneta americana, U.S.A. Roth and (Leibovitz, 195 1 Willis, 1957a). ACTINOMYCETACEAE Family *Nocardia sp.? Natural vectors. —Periplaneta americana, U.S.A. (Leibovitz, 1951). STREPTOMYCETACEAE Family Streptomyces leidynematis Hoffman Habitat. — Surface of the nematodes Hammerschmidtiella dicsingi and Leidynema appendiculata in Periplaneta americana, U.S.A. (Hoffman, 1952, 1953) Eighteen percent of 192 nematodes found in 52 adult cockroaches were infected with the bacterium. : Order CARYOPHANALES Family ARTHROMITACEAE Arthromitus intestinalis Synonymy. —Hygrocrocis Habitat. —Blatta (Valentin) Peshkoff intestinalis. Europe (Valentin, 1836; Robin, 1847, Isolated from intestinal tract. The organ1853 Peshkoff, 1940) ism appears as fragments in fecal masses or as fibers adhering to the orientalis, ; mucous membrane : of the large intestine (Robin, 1853). Cockroach, France? (Chatton and Perard, 1913). ? BIOTIC ASSOCIATIONS OF COCKROACHES Order Family —ROTH & WILLIS I25 SPIROCHAETALES SPIROCHAETACEAE fSpirochaeta blattae Tejera Habitat. from —Blahcrus atropos, Venezuela (Tejera, 1926) : Isolated intestinal tract. * t Spirochaeta periplanetae Habitat. —Blatta orientaHs, Laveran and Franchini France (Laveran and Franchini, 1920a). Cockroaches, Venezuela (Tejera, 1926) "Spirochaeta blatarum Laveran et : Tejera reported finding Franchini" which may have been a lapsus. TREPONEMATACEAE Family t Habitat. —Blatta Zasukhin (1930) Treponema parvum Dobell orientalis, From : t England (Dobell, 1912) ; U.S.S.R. ; U.S.S.R.? intestinal tract. Treponema stylopygae Dobell —Spirochaeta stylopygae Zuelzer. —Blatta England (Dobell, Synonymy. Habitat. orientalis, Zasukhin (1930) From : 1912) intestinal tract. Unidentified spirochaetes Habitat. —Blatta orientalis, U.S.S.R. (Yakimov and Miller, 1922) Spirochaetes and spirilla were found in the intestines of 70 percent of 124 specimens collected in Petrograd. : Periplaneta amcricana, Gold Coast Colony (Macfie, 1922). ADDITIONAL BACTERIA WHOSE TAXONOMIC POSITION IS UNKNOWN *"B. aerobio del pseudoedema maligno" of Cao Natural vectors. —Blatta Natural vectors. —Blatta orientalis, Italy (Cao, 1906). B. alcaligenes beckeri Isolated from orientalis, Poland (Nicewicz intestinal tract. *"B. del pseudoedema maligno" Natural vectors. —Blatta orientalis, Italy of Cao (Cao, 1906). et al., 1946) : SMITHSONIAN MISCELLANEOUS COLLECTIONS 126 * "Bacillo proteisimile" of Natural and experimental vectors. VOL. I4I Cao —Blatta orientalis, (Cao, Italy 1898, 1906). * "Bacillo barbone del bufali" of del Experimenal vectors. — Blatta orientalis, Italy * "Bacillo similcarbonchio" of Natural and experimental vectors. — Blatta Cao (Cao, 1898). Cao (Cao, orientalis, Italy 1898, 1906). * "Bacillo similtifo" or "Bacillo tifosimile" of vectors. — Blatta Natural and experimental Cao (Cao, orientalis, Italy 1898, 1906). "Bacillus" —Blatta Germany (Heinecke, 1956) hemolymph of infected cockroaches. It can be spread by mouth and through wound infection. The animals died with symptoms of paralysis in 85-90 days. The organism has Natural infection. orientalis, : Disease organism found in the been isolated and is in the culture collection of the Institute for Microbiology and Experimental Therapy, Jena, under the numbers SO 896, Strain A SG ; 897, Strain Experimental infection. 1956) : — B SG 898, Strain C. ; Blattclla germanica, Germany (Heinecke, Infected animals died in 26-30 days. Periplaneta americana was unaffected even by heavy inoculations of the pathogen, "Bacterium" Source. — (I) Diseased silkworm larvae. (II) Diseased Ocneria dispar larvae and blood of Blatta orientalis. Experimental infection. toff, 1904) : — (I) (II) Blatta orientalis, Organism pathogenic when Europe (Fila- injected, nonpathogenic when ingested. (I) Cockroach, U.S.A. (Glaser, 1925) : Organism pathogenic to cockroach when injected. "Coccobacillus" Natural infection. —Blatta orientalis, France (Hollande, 1934) : Organism described morphologically. "Colon bacilli" — Natural vectors. Cockroaches [presumably Blatta orientalis, Blatgermanica, and/or Periplaneta americana], Egypt (El-Kholy tclla BIOTIC ASSOCIATIONS OF COCKROACHES and Gohar, 1945) From : —ROTH & WILLIS \2'] the outer surface, intestinal tract, and suspensions of macerated insects. "Diplococci" Natural From vectors. — Blatta Germany orientalis, (Jettmar, 1935) : intestinal tract. Germany Blattella germanica, (Jettmar, 1935) : From outer surface of body. "Diphtheroid Source. I and II" —Periplaneta americana. — and experimental infections. Periplaneta americana, U.S.A. (Gier, 1947) Pathogenicity to the cockroach variable when organism injected. Natural : "Gram — positive rods" Source. Feces of Blattella germanica. Experimental vector. Blattella germanica, Germany (Vollbrechts- — hausen, 1953) mouth or anus. : Nonpathogenic to the insect "Silkworm disease when injected into the bacillus" Cockroaches that were inoculated with living cultures succumbed in a few days (Glaser, 1925). Habitat. — Blatta t Spirillochaeta Hollande blattae France (Hollande, 1934; Hollande and Organism found in hind intestine. This spirillum orientalis, Hollande, 1946) stated to be related in external morphology to Spirillum peri: was planeticum Kunstler and Gineste, but it was believed that S. blattae should be in the Spirochaetaceae rather than the Spirillaccae. "Spirochaetoid bacteria" Habitat. —Blatta orientalis, France (Hollande, 1934) : Two kinds described but not named. fTetragenous Natural and experimental U.S.A. (Gier, 1947) organism injected. : infections. far the greatest —Periplaneta americana, Pathogenicity to the cockroach variable VHI. FUNGI By sp. number when AND YEASTS of fungi known to be associated with cockroaches belong to the Laboulbeniaceae, genus Herpomyces, the species of which are restricted to parasitizing cockroaches (Thaxter, SMITHSONIAN MISCELLANEOUS COLLECTIONS 128 VOL. I4I 1908). Most Species are hyaline, small and inconspicuous (Thaxter, 1931) and are usually, but not exclusively, found on the insects' antennae. Species of Herpomyces are highly, but not completely, host specific (Richards and Smith, 1954). While attached to the host, these fungi appear like minute dark-colored, yellow, or white (e.g., H. arietinus) bristles or bushy hairs (pi. 27, A). Richards and Smith (1955, 1955a) have studied the life history of Herpomyces stylopygae on the oriental cockroach. The plants grow only on living cockroaches, and the infection is disseminated by contact. The mature plants are found mostly on the antennae (pi. 27, B), either on setae or on hard or soft cuticle. Spores are ejected from perithecia singly or in groups of 2 to 4 spores, although groups as large as 12 spores have been found. The presence of single, paired. Herpomyces stylopygae Speg. — Fig. I. Diagram illustrating the relationship between a mature plant of Herpomyces stylopygae and the integument of Blatta orientalis. (Reproduced from Richards and Smith [1956], through the courtesy of Dr. A. G. Richards.) or multiple spore groups on the surface of the host was correlated with the presence of single, paired, or multiple plants on infected cockroaches. Development from spore about two weeks. The to mature perithecia takes plant obtains nutriment from the host by means of a tubular haustorium that extends through the cockroach's and expands into a large bulb in the underlying epidermal cells (fig. i). Infections on nymphs are lost when the nymph moults, but infections on adults persist throughout life. However, nymphs which have lost the fungus upon moulting are readily reinfected. cuticle BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS I29 Collart's (1947) statement that nymphs are never infected with Hcrponiyces is not true. Richards and Smith (1956) concluded that there is no evidence Herpomyces infections because heavily infected of pathogenicity in cockroaches appear fully active in laboratory colonies ; they can run same speed as uninfected cockroaches they reproduce normally and do not appear to die prematurely. These workers stated that the infections cause a dermatitis which is neither pathogenic nor debilitant. So far as we know there are no comparative data on longevity and reproductive performance of fungus-infected versus normal cockroaches. However, Gunn and Cos way (1938) have shown that the presence of these fungi (identified as Stigmatomyces sp. see p. 138) on the antennae seemed to interfere with the humidity reactions of Blatta orientalis. Although Richards and Smith (1956) admit that humidity receptors and other sense organs on the antennae may be at the ; ; destroyed by the fungus, they state that "insects possess such a large number of sensilla that the result may well be sensory physiologist than to the insect." Yet more distressing to the seems to us that the loss of sense organs from fungal infection and concomitant shortening of the antennae (pi. 27, A) might be considerably more of a handicap it to free-living cockroaches than those in laboratory colonies. Bode (1936) studied the flora of Periplaneta americana and cultured Aspergillaceae and Mucorinae from the insect's body surface and intestinal contents americana. Griffiths To ; he also found nonsporulating yeasts in P. prevent fungal growth on oothecae of P. americana, and Tauber (1942a) autoclaved their rearing containers and dipped the oothecae in 70-percent alcohol for 10 seconds. Mercier (1906) isolated and cultured a pathogenic yeastlike parasite which had invaded the fat body and blood of Blatta orientalis. The abdomens of the infected insects became swollen, distended, and soft. McShan (unpublished MS., 1953) consistently isolated Saccharomycetes from the feces of Periplaneta americana. FUNGI ASSOCIATED WITH COCKROACHES The use of the asterisk (*) is explained in footnote 3, page 4. Phylum THALLOPHYTA Class FUNGI IMPERFECTI MONILIALES PSEUDOSACCHAROMYCETACEAE Order Family Candida zeylanoides (Castellani) Langeron and Guerra Natural host. — Ootheca of Blatta orientalis, Italy (Ronzoni, 1949). SMITHSONIAN MISCELLANEOUS COLLECTIONS 130 Torulopsis Natural host. —Ootheca of Blatta sp. orientalis, Italy Spicaria prasina (Maublanc) 1950) : A (Ronzoni, 1949). MONILIACEAE Family Natural VOL. I4I Sawada —Ischnoptera rufa rufa, Puerto Rico (Wolcott, dead specimen of this cockroach was found stuck to a leaf liost. and covered with this fungus. Aspergillus Natural hosts. — Oothecae flavus Link of BlatfeUa gcrmanica and Eiirycotis floridana, U.S.A., Pennsylvania (Roth and Willis, unpublished data, On outer surface. Determination by Miss Mary Downing. 1952) : Oothecae of Pcriplaneta americana, U.S.A., Pennsylvania (Roth Inside oothecae. Determination and Willis, unpublished data, 1952) : by Miss Mary Downing. * Aspergillus f umigatus Fresenius Natural vector. 1908) : From —Blatta orientalis, * Aspergillus Natural vector. in Roth and France (Sartory and niger van Tieghem —Periplaneta americana, From 1957a) — Blatta Willis, Experimental vector. U.S.A., Texas Aspergillus Natural host. orientalis, Italy (Cao, 1898) : Organ- insects. sydowi (Bainier and Sartory) Thorn and Church — Ootheca of Eiirycotis floridana, U.S.A., Pennsyl- vania (Roth and Willis, unpublished data, 1952) Determination by On : outer surface. Mary Downing. ]\Iiss Aspergillus Natural host. (McShan feces. : ism passed unchanged through the gut of the — Oothecae of tamarii Kita Blattella germanica, U.S.A., Pennsyl- vania (Roth and Willis, unpublished data, 1952) face. Clerc, intestine. Determination by Miss : On exterior sur- Mary Downing. Aspergillus sp.? — Natural and e.xperimental vector. Blaitella germanica, on shipIsolated from feces. Experimentally Morrell board (Morrell, 1911) also showed that the spores of the fungus could be recovered from : feces of cockroaches that had fed on them. BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS I3I Aspergillus sp. — Natural vector. Periplaneta americana, England (Bunting, 1956) The fungus was isolated mostly from imperfectly excreted feces. Beauveria bassiana (Balsamo) : Vuillemin — Experimental host. Blattella germanica and Periplaneta americana, U.S.A. (Dresner, 1949, 1950) The nymphs of American cockroaches became infected when they (i) were injected with a i -percent suspen: sion of spores, (2) ate rat pellets sprayed with the spore suspension, or (3) were dusted with the fungus spores. The symptoms of the fungus infection were paralysis followed by death some of the ; infected insects liquefied, others dried up after the appearance of a subcuticular blackening. Cephalosporium sp. Natural vector. — Periplaneta americana, U.S.A., Texas (McShan, unpublished MS., 1953) From feces of cockroaches collected in the basement of a grain elevator at the docks in Galveston. : * Experimental vector. ism retained its Geotrichum candidum Link —Blatta orientalis, Italy Penicillium — Blaberiis Natural vector. From (Cao, 1898) Organ- sp. craniifer, U.S.A. (Wedberg et al., feces. 1949) Periplaneta americana, England (Bunting, 1956) imperfectly excreted feces. : : pathogenicity after passing through the insect's gut. Metarrhizium anisopliae (Metschnikoff) — Mostly from : Sorokin Natural liosfs. Blattidae, Seymour (1929) Charles (1941). Panestkia australis, U.S.A., Massachusetts (Roth and Willis, un; published data, 1957) Growing on adult specimens that were found dead in a laboratory colony. Determination by Miss Dorothy Fennell. : Periplaneta americana, England genitalia of females where it (Bunting, 1956) Growing on prevented oothecal formation. : Cockroach, Puerto Rico (Johnston, 191 5) From a "small roach" Rio Piedras (no data). : in the pathological collection at ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 132 DEMATIACEAE Family Memnoniella echinata Natural host. — Ootheca VOL. I4I Galloway (Rivolta) of Blattella germanica, U.S.A., Pennsyl- vania (Roth and Willis, unpublished data, 1952) On material that had oozed from a damaged ootheca. Determination by Miss Mary : Downing. Torula acidophila Natural host. 1948) : The this yeast Owen and Mobley —Periplaneta americana, U.S.A. digestive tract of this cockroach which was transmitted superimposed a foreign is (Owen and Mobley, the normal habitat of by the to sirup insects. The yeast suggestive of malic acid, upon the taste, original flavor of the sirup. Torula Natural host. Lodder, 1934) is —Blatta from Lodder Germany (Gropengiesser, 1925 orientalis, Isolated : gropengiesseri body and oothecae. Gier (1947) fat of the opinion that the so-called yeasts that supposedly the bacteroids may 1925) in the fat body (Mercier, actually represent poorly fixed 1907b; and may displace Gropengiesser, insufficiently stained bacteroids. Torula rosea Preuss Experimental host. —Blaherus craniifer, U.S.A. (Wedberg et al., Upon repeated feeding of massive doses of this yeast to the 1949) cockroach, these workers were able to isolate the organism from the : feces up to six days thereafter. There was no evidence was pathogenic for B. craniifer. Class that T. rosea PHYCOMYCETES Order Family MUCORALES MUCORACEAE Mucor guilliermondii Nadson and Filippov Natural host. Filippov, 1925 —Periplaneta americana, Filippov, 1926) ; : U.S.S.R. Isolated and cultured Mucor (Nadson and from intestine. sp. — Ootheca of Periplaneta americana, U.S.A., Pennsyl- Natural host. vania (Roth and Willis, unpublished data, 1952) Inside ootheca. : Mary Downing. Pycnoscelus surinamensis, Germany (Bode, 1936) Determination by Miss fat body which it had stained red. : Isolated from BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS I33 Rhizopus nigricans Ehrenberg Natural 1949) : vector. — Blaberus From (Wedberg U.S.A. craniifer, et al., feces. Rhizopus sp. — Natural vector. Periplaneta americana, U.S.A., Texas (McShan, unpublished MS., 1953) From feces. : Syncephalastrum Natural vector. unpubUshed MS., 1953) From : (McShan, feces. ENTOMOPHTHORALES Order Family BLASTOCYSTIDACEAE Blastocystis Natural sp. —Periplaneta americana, U.S.A., Texas vector. — Blatta Brumpt hominis orientalis, U.S.S.R. (Zasukhin, 1930) : In hind gut in 40 percent of over 3,000 cockroaches. Blastocystis sp. Natural vectors. 1922) : Found —Blatta orientalis, U.S.S.R. (Yakimov and Miller, 29 percent of 124 B. in the intestinal contents of ori- entalis. Cockroaches, Venezuela (Tejera, 1926). The placement of the following fungus is Avrech Coccidioides periplanetae Natural host. —Blatta orientalis, problematic. Germany (Avrech, 1931) : Found lumen of midgut and caeca. The whole upper part of the epithelium was filled with sporangia and spores. in cells lining the Class Order Family ASCOMYCETES ENDOMYCETALES SACCHAROMYCETACEAE Saccharomyces cerevisiae Hansen Natural 1949) : vector. — Blaberus craniifer, U.S.A. (Wedberg et al., In feces. Saccharomyces Natural vector. berg, 1952) : — Blattella germanica, Found sp. U.S.A. (Janssen and Wed- consistently in alimentary tract of B. germanica fed sucrose solutions. SMITHSONIAN MISCELLANEOUS COLLECTIONS 134 Order Family HYPOCREALES HYPOCREACEAE Hennings Cordyceps amazonica Natural host. VOL. I4I — Cockroaches, British Honduras (Mains, 1940), Cordyceps blattae Fetch Natural host. — Ceylon (Petch, 1924) Blattella germanica, : Col- Hakgala twice. A slight covering of brown mycelium overran the insect and fastened it to the underside of a living leaf. lected at Order Family LABOULBENIALES LABOULBENIACEAE Herpomyces amazonicus Thaxter Natural host. On —Nyctibora ohsciira, Brazil, Natal (Thaxter, 1931) : antennae. Herpomyces anaplectae Thaxter Natural hosts. —Anaplecta sp., Venezuela, Caracas (Thaxter, 1905, On antennae. 1908) Trinidad (Thaxter, 1931) Cockroach, Sumatra (Thaxter, 1931). : ; Herpomyces appendiculatus Thaxter Natural host. 1 931): On —Platysostcria scahra, Australia, N.S.W. (Thaxter, antennae. Herpomyces arietinus Thaxter Natural hosts. —Ischnoptera sp., U.S.A., Georgia (Thaxter, 1908). Parcoblatta uhleriana, U.S.A., Massachusetts (Roth, unpublished data, 1957) The nymphs were in a culture of Parcoblatta virginica which was infected with this fungus it is possible that these P. uhleriana became infected by contact with P. virginica. Fungus identified by Dr. R. K. Benjamin. : ; Parcoblatta virginica, U.S.A., Massachusetts (Roth, unpublished Fungus determined by Dr. R. K. Benjamin. Fungus found on antennae, palpi, legs, body surface (pi. 27, A). data, 1957) : Parcoblatta 1908) : On sp., U.S.A., Kentucky, Massachusetts (Thaxter, 1902, antennae. It is likely that Thaxter's host records (certainly those assigned to Temnopteryx and possibly those assigned to Ischnoptera) were species of Parcoblatta. Hebard (1917) has shown that all the species BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH referred to Ischnoptera in the United States, except /. & WILLIS I35 deropcllifonais, now belong in the genus Parcohlatta. All species originally referred to the genus Temnopteryx in the United States are now synonyniized with species of Parcohlatta. Herpomyces chaetophilus Thaxter Natural hosts. Periplaneta On —Periplaneta americana, Brazil (Thaxter, 1931). Zanzibar and Mauritius (Thaxter, 1902, 1908) sp., : spines of legs, antennae, and cerci. Herpomyces Natural host. — Cockroach, chilensis Thaxter Chile (Thaxter, 1918) : On antennae. Herpomyces diplopterae Thaxter Natural hosts. 1902, 1908) : —Diploptera On antennae. punctata, Ascension Island (Thaxter, This species also was infected experi- mentally (Richards and Smith, 1954). Cockroach, Fiji (Thaxter, 1931). Herpomyces ectobiae Thaxter — Natural hosts. ter, 1902, 1908) Blattella germanica, U.S.A., Massachusetts Burma, Tenasserim (Spegazzini, 191 5) ; ; (Thax- Argentina, Buenos Aires (Spegazzini, 1917) On antennae. U.S.A., Minnesota Scattered over entire body, wings. (Richards and Smith, 1955) On tibial spines. Chile and Philippine IsFrance? (Picard, 1913) : : : lands (Thaxter, 1931). "Ectohia" spp., Zanzibar and Saint Kitts, B.W.I. (Thaxter, 1902, Possibly on species that are now in the genus Blattella rather 1908) : than in the genus Ectobius as it is known today, because Thaxter also used the synonym Ectobia germanica for the German cockroach, Blattella germanica. Experimental hosts. — Blattella germanica and Blattella vaga, U.S.A. (Richards and Smith, 1954). Herpomyces Natural hosts. 1908, 193 1 ) : forficularis — Cockroaches, Mauritius? and On Natural host. : Fiji (Thaxter, 1902, antennae. Herpomyces — (Thaxter, 1931) Thaxter Blattella On gracilis Thaxter humhertiana, Philippine Islands, Luzon antennae. SMITHSONIAN MISCELLANEOUS COLLECTIONS 136 VOL. I4I Herpomyces grenadinus Thaxter — Natural host. Cockroach, Grenada, B.W.I, (Thaxter, 1931) antennae of a "brown wingless blattid." Herpomyces Natural Jiost. —Leurolestes (Thaxter, 1931) : On leurolestis : On Thaxter Guiana and Trinidad pallidus, British antennae. Herpomyces lobopterae Thaxter Natural host. —Lohoptera sp., Argentina (Thaxter, 1931) On : an- tennae. Herpomyces macropus Spegazzini —Blahcrus sp. ?, Argentina (Spegazzini, 1917). Cockroaches, Peru, Puerto Rico, Ecuador, and Haiti (Spegazzini, Natural host. 1915, 1917) : Material previously assigned by Spegazzini (1915) to also placed by him in this new species. Hovv^ever, H. paranensis was Thaxter (1931) believed that H. macropus may be synonymous with H. paranensis, but he provisionally retained H. macropus because he had not seen Spegazzini's material. Herpomyces nyctoborae Thaxter Natural hosts. 1905, 1908) : — Nyctibora On antennae. Texas, and the specimen tomentosa, U.S.A., Texas This cockroach may have is (Thaxter, not established in been misidentified (Gurney, per- sonal communication, 1958). Nyctibora sp., Argentina (Spegazzini, 1917) : On antennae. Herpomyces panchlorae Thaxter Natural hosts. —Panchlora nivea, Trinidad (Thaxter, 1931): On antennae. Herpomyces panesthiae Thaxter Natural host. —Panesthia lobipennis, Ceylon (Thaxter, 1915) : On antennae. Herpomyces paranensis Thaxter Natural hosts. — Blaberus sp.? Brazil (Thaxter, 1902, 1908): On On an- antennae. Blaberus sp., Brazil and Argentina (Spegazzini, 1917) tennae. Cockroaches, Trinidad and Argentina (Thaxter, 1931). : BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS I37 Herpomyces periplanetae Thaxter Natural hosts. —Blaherus sp. ?, Argentina (Spegazzini, 1917). (Thaxter, 1902, 1908) Blatta orientalis, U.S.A., Massachusetts ; France? (Picard, 1913). Periplaneta americana, Bermuda and U.S.A., Massachusetts Plains of Biajar, Italian Somaliland, and (Thaxter, 1902, 1908) Locality? (Spegazzini, 191 5) ; ; Argentina (Spegazzini, 1915, 1917). Periplaneta australasiae, Bermuda (Thaxter, 1902, 1908). Periplaneta hrunnea, Brazil (Thaxter, 1931). Periplaneta sp., Mexico, West Indies, Panama, Brazil, Africa, South Seas, and China (Thaxter, 1902, 1908). Cockroaches, Belgium (Collart, 1947). Grenada, Trinidad, B.W.I., and TanAdditional locality records : gier (Thaxter, 1931). The fungus was found growing on spines, tegmina, integument, and antennae, at times abundantly. Experimental hosts. All the following data are from Richards and Smith (1954): Blatta orientalis: A few plants matured. Neostylopyga rhomhifolia: Some development but no mature — plants. Periplaneta americana: on sity equal to that Periplaneta australasiae : Periplaneta hrunnea: equal to that on Fungus developed prolifically with a den- original host. Some development Fungus developed but no mature plants. prolifically with a density original host. Herpomyces phyllodromiae Thaxter Natural host. 1908) : On —"Phyllodromia" sp., Abyssinia (Thaxter, 1905, (Thaxter, 1905, antennae. Herpomyces platyzosteriae Thaxter Natural host. — "Eurycotis floridana," Mexico On antennal setae. 1908) Since this cockroach is not found in Mexico (J. A. G. Rehn, personal communication, 1957), E. floridana is undoubtedly not the host : for this fungus. W. B. Brown (personal communication, searched the cockroach collection at the Museum Zoology but was unable to find Thaxter's insect for 1957) of Comparative reidentification. Herpomyces stylopygae Spegazzini Natural hosts. —Blatta orientalis, Argentina (Spegazzini, 1917) U.S.A. (Richards and Smith, 1955a). ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 138 Experimental Jiosts. —Neostylopyga rhomhifoUa VOL. I4I (Richards and Smith, 1954) A few plants matured. Pycnoscelns surinamciisis (Richards and Smith, 1954) velopment but no mature plants. : : Some de- The fungus (fig. i) is found on antennae (pi. 27, B, C), palpi, and femurs. Thaxter (1931) believed H. stylo pygae to be synonymous with H. periplanctae. However, Richards and Smith (1954) concluded that H. stylopygae would not grow on P. americana under their laboratory conditions although H. periplanetac would grow on B. orientalis. This indicated a strain or species difference between the two fungi. Gunn and Cosway (1938) reported a species of Stigmatomyces on the antennae of B. orientalis; this fungus was probably H. stylopygae (Richards and Smith, 1956). cerci, Herpomyces supellae (Thaxter) Natural host. On —Supclla Trinidad (Thaxter, 1931J: supellectiliiim, antennal spines. Herpomyces tricuspidatus Thaxter hosts. — Blahcrus Natural craniifer, U.S.A., Key West (Richards and Smith, 1955). Blaberus sp. and Epilampra? sp., Panama (Thaxter, 1902, 1908). Epilampra sp., Saint Kitts, B.W.I., and Haiti (Thaxter, 1902, 1908). Leucophaea maderae, Fernando Po (Spegazzini, 1915). Nauphoeta cinerea, Brazil (Thaxter, 1931). Philippine Islands, MinCockroaches, China? (Thaxter, 1902) danao (Thaxter, 1931). Experimental hosts. Blaberus craniifer, U.S.A. (Richards and ; — Smith, 1955). Infections on the antennae. Richards and Smith (1954) were unable to secure experimental infections in L. maderae with H. tricuspidatus. Experiments with N. cinerea showed some development but no mature plants although identification of the growing fungus was uncertain because of simultaneous exposure H. tricuspidatus. to H. ectohiae, II. stylopygae, and Herpomyces zanzibarinus Thaxter Natural hosts. —Eurycotis manni, Brazil (Thaxter, 1931): antennae. Gyna sp. ?, Isle of Nias (Spegazzini, 1915) On antennae. On antennae. Cockroach, Zanzibar (Thaxter, 1902) : : On BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS I39 INCERTAE SEDIS According to Dr. R. K. Benjamin (personal communication, 1957) and Dr. E. G. Simmons (personal communication, 1957), the phylogenetic position of the following genus is uncertain. Amphoromorpha Natural hosts. On — Cockroaches, blattina Thaxter Grenada, B.W.I. (Thaxter, 1920) the axis of the antennae of a dark wingless and a pale : winged blattid. Amphoromorpha Natural host. sp. —Cockroach, Grenada, B.W.I. (Thaxter, 1920) : On antennal setae. IX. The significance of and the higher plants HIGHER PLANTS many observed associations between cockroaches obscure. Undoubtedly is still many associations are ecological, but lack of adequate supporting evidence makes this conclusion somewhat tentative. The ecological aspects are covered 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 in Section III (p. 14). ; 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 Family PTERIDOPHYTA CYATHEACEAE Alsophila Pycnoscelus surinamensis, Feeding on heart of tree fern. Associate. 1893) : — sp. Louisiana (Anonymous, SMITHSONIAN MISCELLANEOUS COLLECTIONS 140 VOL. I4I POLYPODIACEAE Family Asplenium nidus Linnaeus Associate. — Comptolampra liturata, Malaya (Karny, 1924) found between dry foliage of the beakers of Division Often : this fern. EMBRYOPHYTA SIPHONOGAMA Family PINACEAE Pinus australis Michaux Associates. —Aglaopteryx gemma and Parcohlatta lata, Alabama (Hebard, 191 7) The former species was common under signs on longleaf pines, and P. lata was occasional. : Parcohlatta divisa, Georgia (Rehn and Hebard, 1916) Under : signs. Pinus caribaea Morelet Associates. —Eurycotis hlatta fnlvescens, signs on the floridana, Florida Latihlattella rehni, (Hebard, 1917) : Many and Parco- records under tree trunks. Pinus clausa Vasey Associate. sign on —Latihlattella rehni, Florida (Hebard, 1917) Under : tree. Pinus echinata Mill. Associates. — Parcohlatta Virginia divisa, (Rehn Under signs on shortleaf pine. 1916) Parcohlatta sehra,'M.\ss\s,s\'p^i (Hebard, 1917) and Hebard, : : Undersign. Pinus sylvestris Linnaeus Associate. —Ectohius 1899b) : On Scotch pallidus, England (Milton, 1899; Burr, fir. Pinus spp. Associates. —Plectoptcra laccrna and Plectoptera vermiculata, Cuba (Rehn and Hebard, 1927). Florida (Rehn and Hebard, 1905) Cuba (Rehn and Hebard, 1927). Latihlattella rehni, signs. Family liams, 1938; Under TAXODIACEAE Cryptomeria Associate. —Diploptera : punctata, Zimmerman, 1948). sp. Hawaii (Pemberton and Wil- BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS I4I CUPRESSACEAE Family Cupressus macrocarpa Hartweg Associate. species is —Diploptera common and Hawaii (Hebard, 1922) punctata, "The : injurious in the territory infesting particularly and doing particular damage by Monterey cypress trees gnawing away the bark." Similar injury has been cited by Pemberton (1934), FuUaway and Krauss (i945)» and Zimmerman (1948). the . . . Juniperus Associate. sp. —Phyllodromica tartara nigrescens, Southern Uzbekistan (Bei-Bienko, 1950) Under : Family bark. PANDANACEAE Freycinetia Associate. — Graptoblatta sp. notulata and Kuchinga remota, Tahiti (Hebard, 1933). Pandanus Associate. —Hololeptoblatta sp., sp. Seychelles (Scott, 1910, 1912). GRAMINEAE Family Aristida pennata Trin. Associate. —Phyllodromica pygmaea, Found IJ .S.S.R. (Bei-Bienko, 1950) : dense turf. in the Bamboo Associate. — Comptolampra liturata, Chloris Malaya (Karny, 1925). gayana Kunth — This Blattella vaga, Texas (Riherd, 1953) Associate. roach was rather abundant in clumps of Rhodes grass. : field cock- Panicum purpurascens Raddi —Panicum barbinodc 1936]. —Epilampra abdonien-nigrum, Puerto Rico Synonymy. Associate. Wolcott, 1936) [H^itchcock, : Abundant in "malojillo" (Sein, 1923; meadow. Saccharum officinarum Linnaeus Associates. —Balta qttadricaudata, Balta scripta, Balta torresiana, Balta verticalis, Ellipsidion simulans, and Megamareta verticalis, AusAll collected by J. F. Illingworth tralia, Queensland (Hebard, 1943) : on sugarcane. SMITHSONIAN MISCELLANEOUS COLLECTIONS 142 VOL. I4I Ischnoptera schenklingi, and Pycnoscelus Blattella hiimhertiana, Surinam ensis, Formosa (Box, 1953). Cariblatta stenophrys, Puerto Rico (Sein, 1923; Wolcott, 1936): Between the leaves and under the leaf sheaths. On the leaves. Panchlora nivea, Cuba (Rehn and Hebard, 1927) : Pelmatosilpha coriacca, Puerto Rico (Wolcott, 1936). Phoraspis spp., Brazil and Guiana (Doumerc in Blanchard, 1837). Plectoptera dorsalis, Plcctoptora infulata, and Plectoptera rhabdota, Under the leaf sheaths, Puerto Rico (Wolcott, 1950) Symploce ruficollis, Puerto Rico (Wolcott, 1950) : Often found : living under the Cockroaches, 1931) : leaf sheaths. (Uichanco in Williams Islands Philippine Between cane et al., leaf sheaths. Setaria verticillata (Linnaeus) — Chaetochloa Associate. — Diploptera punctata, Synonymy. verticillata Beauv. [Howard, per- (Linnaeus) sonal communication, 1958]. Hawaii (Severin, 1911) : The cockroach was caught on the barbed awns of this grass. Wiia oats Associate. —Ischnoptera deropeltiforrnis, Missouri (Rau, 1937). Zea mays Linnaeus Associates. cott, — Cariblatta stenophrys, Puerto Rico (Sein, 1923; Wol- 1936). Ellipsidion bicolor, Australia, Queensland (Hebard, 1943). Lo phoblatf a arawaka, TnmdRd Phoraspis sp., (Princis and Kevan, 1955). Brazil and Guiana Supella supellectilium. New (Doumerc CYPERACEAE Family Cyperus Associate. —Maretina in Blanchard, 1837). Caledonia (Cohic, 1956). sp. uahuka, Marquesas Islands, Uahuka (He- bard, 1933a). Family PALMAE Acrocomia aculeata Associate. —Pycnoscelus Kevan, 1955) : On (Jacq.) surinamensis, "gru-gru" fruits. Lodd. Trinidad (Princis and BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS I43 Cocos nucifera Linnaeus Associates. —Aglaopteryx gemma, Florida (Rehn and Hebard, 1912). Cariblatta lutea minima, Florida, and Carihlatta delicatula, San Domingo (Hebard, 1916a). Eurycotis floridana, Florida (Rehn and Hebard, 1912; Hebard, 1917). Periplaneta australasiae, Jamaica (Rehn and Hebard, 1927). Pycnoscehis surinamensis, Florida (Rehn and Hebard, 191 2; bard, 191 7). Jamaica He- (Rehn and Hebard, 1927). Phoenix dactylifera Linnaeus Associate. — Dlattella 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 Family (Zimmern — Latiblattella Gould and Deay, 1940). ARACEAE Arum Associate. in vitrea, sp. Mexico (Hebard, 1921b) : In flower shaft. Caladium Associate. — Plcctoptera 1927). dorsalis, sp. Puerto Rico (Rehn and Hebard, SMITHSONIAN MISCELLANEOUS COLLECTIONS 144 Family VOL. I4I BROMELIACEAE Aechmaea Associate. —Dryadohlatta porteoides scotti, Britton Trinidad and Kevan, (Princis 1955)- Ananas comosus Merr. Associates. 1927, 1929) —Pycnoscclus Hawaii surinamcnsis, (Illingworth, Feeding on roots of pineapple. : Formosa (Takahashi, 1940) Blattella humhertiana, grown nymphs occasionally lie Imago and : concealed in the leaves. Catopsis fulgens Griseb. Associates. — Cockroaches, Costa Rica (Calvert and Calvert, 1917). Glomeropitcairnia erectiflora Associate. —Dryadohlatta scotti, Mez Trinidad (Princis and Kevan, 1955)Grevisia sp. Associate. 195s) • —Notolampra antiUariim, Trinidad One male —Eurycotis Swartz fasciculata Tillandsia Associate. (Princis and Kevan, only. floridana, Florida (Rehn and Hebard, 1914; Hebard, 1917). usneoides Tillandsia Associates. —Parcohlatta sp., Linnaeus Louisiana (Rainwater, 1941). Latiblattella rehni, Florida (Blatchley, 1920) By : 39 immature moss. blattids were collected Tillandsia Associate. uttriculata —Epilampra mona, Mona Hebard, 1927) : The type and one beating. One mature and 1, 1912) from 8 of 12 samples of Spanish Cockroaches, Louisiana (Rosenfeld, 191 : Linnaeus West Indies (Rehn and paratypic female of E. mona were Island, collected in this bromeliad. Eurycotis floridana, Florida (Blatchley, 1920). Tillandsia sp. Associates. —Aglaopteryx gemma, Texas Dryadohlatta bases. scotti, (Hebard, 1917). Trinidad (Scott, 191 2) : Found in the leaf BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS I45 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 hromcliadariim, 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." Carihlatta insniaris, Jamaica (Plebard, 1916a, 1917; Rehn and Hebard, 1927). Carihlatta nehiilicola, Jamaica (Rehn and Hel)ard, 1927) : One immature male. scotti, Trinidad (Princis and Kevan, 1955). Epilampra conspersa, Dominica (Scott, 1912). Epilampra niaya, Panama (Hebard, 1920). Epilampra sodalis, Panama (Caudell, 1914). Epilampra sp. and Hormetica laevigata, Brazil (Hancock, 1926). Dryadoblatta Eurycotis hiolleyi, Costa Rica (Picado, 1913). Ischnopterariifa occidentalis, Mexico (Caudell, 1914). Latiblattella chichimeca, Costa Rica (Picado, 1913). Litopeltis hiolleyi, Costa Rica (Rehn, 1928). Litopeltis bispinosa, Panama (Caudell, 1914). Neohlattella brunneriana, Costa Rica (Calvert and Calvert, 1917). Neoblattella dryas, Neohlattella eurydice, Neohlattella grosshecki, and Neohlattella proserpina, Jamaica (Rehn and Hebard, 1927). Neohlattella fratercula, Mexico (Hebard, 1921b). Neohlattella nahua, Mexico (Caudell, 1914). Nesomylacris relica, Jamaica (Rehn and Hebard, 1927). According to Nyctihora hrunneaC^), Panama (Caudell, 1914) Hebard (1920) Caudell's specimen was almost certainly incorrectly : identified. It may have been Nyctihora noctivaga or a smaller species of the genus. Brazil (Hancock, 1926). Nyctihora laevigata, Jamaica (Hebard, 1917; Rehn and Hebard, 1927). Nyctihora lutzi, Puerto Rico (Rehn and Hebard, 1927) : "in epi- phytes with pencil-like leaves." Panama (Caudell, 1914). Costa Rica (Picado, 1913). Pycnoscelus surinamensis, Costa Rica (Picado, 1913). Pelmatosilpha rotundata, Pseudomops laticornis, (Caudell, 1914). Jamaica (Rehn and Hebard, 1927). Mexico SMITHSONIAN MISCELLANEOUS COLLECTIONS 146 "Rhicnoda" sp., Costa Rica (Picado, 1913). recognized as not being in the New World VOL. I4I This genus is now Probably the fauna. specimen was a species of Epilampra or Hyporhicnoda as suggested by Gurney (personal communication, 1959) and confirmed by Rehn (p.c, 1959). Cockroaches were said Cockroaches, Costa Rica (Calvert, 1910) common in bromeliads on the moist Atlantic slope. : to be Family LILIACEAE Yucca elata Engelman Associate.— Latiblattella lucifrons, Easter Associate. Arizona (Ball et al., lilies —Pycnoscehis surinamensis, Connecticut Family 1942). (Zappe, 1918). MUSACEAE Bananas Cockroaches have been captured in bunches of bananas, in bracts of banana flowers, under banana leaves, and burrowing in rotten Although many of the species associated with bananas 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 banana stalks. are indigenous to the banana-growing areas of the 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 by Hebard (191 7) 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 obcited served. Known or suspected adventive material is so indicated. Found Aglaopteryx diaphana, Jamaica (Rehn and Hebard, 1927) Adventive, in bracts of banana blossoms. England (Bunting, 1955) on bananas from Dominica. Aglaopteryx vegeta, Finland (Princis, 1947) Adventive, in banana : : : box. Amasonina emarginata, Trinidad banana bunch. (Princis and Kevan, 1955) : In BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS I47 AdvenArchimandrita marmorata, Denmark (Henriksen, 1939) bananas from Jamaica ( ?). As Princis (1947) and Gurney (personal communication, 1959) point out, this is a Central American : tive (2), in Jamaica may be an error. Archimandrita tcssellata, Sweden species, so (Princis, 1947) Adventive, : from Honduras. Blaherus atropos{?), Denmark (Henriksen, 1939): Adventive, from Jamaica. Princis (1947) pointed out that this species was more have been Blaherus craniifer or Blaherus discoidalis, which are West Indian species, than B. atropos which is a South American likely to species. Blaherus holiviensis, Ecuador (Princis, 1952) In a shipment of : bananas from near Puna. Blaherus discoidalis, Puerto Rico (Rehn and Hebard, 1927) From Adventive. banana ripening room. Great Britain (Pearce, 1929) England (Bunting, 1955, 1956) Adventive, from Dominica. : : : Capucinella delicatida, California (Caudell, 1931) Adventive. : Cuba (Rehn and Hebard, 1927). Honduras (Rehn, 1945a) Shaken from hanging Carihlatta delicatula, Carihlatta hylaea, : dead banana leaves. Carihlatta insularis, Finland (Frey, 1948) Carihlatta landalei, Jamaica mens taken from under drying Carihlatta punctipennis (Bunting, 1956) : Adventive. (Rehn and Hebard, 1927) : All speci- bracts of banana blossoms. and Chorisoneura harhadensis, England Adventive, from Dominica. : Epilampra abdomen-nigrum and Epilampra sp., England (Bunting, Adventive, from Dominica. 1955) Epilampra may a, Massachusetts (Hebard, 191 7) 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 hananae, U.S.S.R., Leningrad Adventive, from Colombia. (Bei-Bienko, 1947) : Euphyllodromia angustata, Sweden (Princis, 1947) Adventive. Eurycotis caraihea, New York (Hebard, 1917) Adventive. AdEurycotis dimidiata, Washington, D. C. (Caudell, 1931) : : : ventive. Eurycotis ship lixa, New York (Rehn, 1930) : Adventive, on banana from Jamaica. Graptohlatta 1933a) : notidata, In banana leaves. Marquesas Islands, Uahuka (Hebard, SMITHSONIAN MISCELLANEOUS COLLECTIONS 148 VOL. I4I Holoconipsa nitidula, Trinidad (Princis and Kevan, 1955) banana pulp. Hormetica laevigata, Wales (Sandemann, 1934) Eating : Adventive, in pile : of banana sacks. Adventive, in local Hormetica ventralis, Sweden (Princis, 1947) warehouse of banana company. Hormetica spp., Europe and North America (Bei-Bienko, 1950) Adventive, introduced with bananas and other tropical fruits. Brought into Ischnoptera rufa rufa, Puerto Rico (Wolcott, 1950) : : : houses on bunches of bananas. Kuchinga Moorea (Hebard, 1933a) retnota. Society Islands, : In dead banana leaves. Lamproblatta alhipalpus, Panama Canal Zone (Hebard, 1920) Several under decayed banana stem. La^i/^/ai^^//a sp., Finland (Frey, 1948) : : Adventive. Leucophaea maderae, New York (Hebard, 1917) Adventive. Under banana sheaths. EngDominica (Rehn and Hebard, 1927) : : Adventive, captured at railroad station after land (Palmer, 1928) Adventive, bananas had been unloaded. England (Bunting, 1955) Nymph, from Dominica. Trinidad (Princis and Kevan, 1955) : : : eating bananas in cupboard. Puerto Rico (Sein, 1923) Sein stated : that bananas are the favorite food of L. maderae. Litopeltis bispinosa, rotting banana Panama Canal Zone (Hebard, 1920) Litopeltis musarimi, Costa Rica (Rehn, 1928) banana : From stalks at bases of leaves. : Shaken from dead leaves. Nauclidas nigra, England (Bunting, 1955, 1956) : Adventive, from Dominica. Nauphoeta flexivitta, Denmark (Vestergaard, 1958) : Adventive. and Neoblattella Adventive, from Dominica. laodamia, England (Bunting, 1956) Bunting (1955) first reported these as Neoblattella spp, and stated that they were common. Neoblattella carcinus, Neoblattella celeripes, : From (Rehn and Hebard, 1927) (Princis, Adof banana blossoms. Sweden 1947) Neoblattella detersa, Jamaica under the bracts : : ventive. Neoblattella 1948) : detersa and Neoblattella tridens, Finland (Frey, Adventive. Neoblattella fratercula, Nebraska (Hebard, 1916b) Neoblattella semota, Jamaica : Adventive. (Rehn and Hebard, 1927) under drying bracts of banana blossoms. : From BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS Neoblattella vatia, Cuba (Rehn and Hebard, 1927). Neohlattella Finland sp., (Princis, 1947) I49 from Adventive, : Jamaica. Adventive, from (Bunting, 1955) asteca. Dr. A. B. Nocticola as species this reported Bunting Dominica. Nocticola is an Old World that fact the attention to our called Gurney World species New with the error in combined presumably genus, confirmed by Dr. D. was specimen the of identity true The azteca. examined it at the who communication, (personal 1958), Ragge Nyctibora asteca, England British Museum : (Natural History). Adventive. Nyctibora holoserica, Canada (Walker, 1912) Pennsylvania Massachusetts, Maine, Nyctibora laevigata, Canada, : Adventive. Taken from banana boat Annetta (2) (Hebard, 1917) at Philadelphia (Rehn and Hebard, 1927). England (Bunting, 1956) : : from Dominica. Adventive, Sweden, Denmark (Princis, 1947) : Adventive. Nyctibora 7nexicana{F), Denm3.r\<. (Henriksen, 1939): Adventive (5), from Jamaica and West Indies. Princis (1947) suggested that these specimens were probably the West Indian Nyctibora noctivaga, because A^. mexicana is not a West Indian insect. Nyctibora noctivaga, Canada, Idaho, Illinois, Massachusetts, Nebraska (4), Virginia (Hebard, 1917) Adventive. Nebraska (Hauke, : From Adventive (2). Panama Canal Zone (Hebard, 1920) 1949) Adventive, from banana stalks. England (Blair in Turner, 1930) Adventive. Sweden Costa Rica. Washington (Hatch, 1938) : : : : (Princis, ventive, 1947): Adventive (2). Finland (Princis, Ad- 1947): from Jamaica. Nyctibora obscura, Trinidad banana bunch. (Princis and Kevan, 1955) : In AdNyctibora sericea, Canada (Stevenson, 1905; Walker, 1912) Hebard (1917) synonymized Walker's specimen under A^. : ventive; laevigata. Jamaica. from Adventive, from Isle of Wight (Meade-Waldo, 1910) Adventive, England (Tulloch, 1939) : in : Nyctibora sp., England (Welch, 1935) Adventive, in railway England (Tulloch, 1939) Adven: truck that had carried bananas. tive, banana crates Brazil. from : Brazil. Oxyhaloa deusta, U.S.S.R., Leningrad (Bei-Bienko, 1947) from Colombia. : Ad- ventive, Panchlora Dominica. antillaritm, England (Bunting, 1955) : Adventive, from SMITHSONIAN MISCELLANEOUS COLLECTIONS 150 VOL. I4I Adventive. Great Panchlora exolcta, Scotland (Distant, 1902) AdvenAdventive. England (Coney, 1918) : Britain (Shaw, 1902) : : Sweden, Norway (Princis, 1947) Adventive, Norwegian specimen from Brazil. Germany (Zacher, 1917) Adventive, from Ja- tive. : : maica. Panchlora nivea, Colorado, Nebraska, New Jersey, New York (2), Utah (Hebard, 191 7) Adventive. Nebraska (Hauke, 1949) AdAdventive. Massachusetts Washington (Hatch, 1938) ventive. Adventive. England (Bunting, 1955) (Roth and Willis, 1958) AdvenAdventive, from Dominica. U.S.S.R. (Bei-Bienko, 1947) tive, from Colombia. Sweden (13), Norway (3), Finland (3) (PrinAdventive, mostly females; origin (where known) Jacis, 1947): : : : : : : maica. Panchlora fraterna( ?) and Panchlora peruana{ ?), Denmark (HenAdventive; origin (where known) Danish West Indies and Jamaica; Princis (1947) suggested that both species were riksen, 1939) : probably Panchlora nivea. Panchlora sagax, Puerto Rico (Wolcott, 1936). Adventive; this was Panchlora virescens, Canada (Walker, 1912) probably P. nivea as we now know it (Gurney, personal communica: tion, 1959). Panchlora loch, 1939) : Canada (Walker, 1912) Adventive, from Brazil. sp., : Adventive. England (Tul- Pelmatosilpha coriacea, Puerto Rico (Wolcott, 1936). Pelmatosilpha marginalis and Pelmatosilpha purpurascens, England (Bunting, 1955, 1956) Adventive, from Dominica; both species : common. New Pelmatosilpha vagabunda, tive, Zealand (Princis, 1954) : Adven- probably from South America. Adventive, Periplaneta americana, Belgium (Schepdael, 1931) on bananas from the American Tropics. Periplaneta americana and Periplaneta hrnnnea, England (BunAdventive, from Dominica. ting, 1955, 1956) : : Periplaneta americana and Periplaneta anstralasiae, England (Watson, 1907): they ate ripening bananas in the tropical Adventive; Kew, where they hid in Sweden plant house of the Royal Botanic Gardens, "the sheathing bases of palm, banana and pandanus leaves." (Princis, 1947) Adventive. : Periplaneta australasiae, Canada (Walker, 1912) mark (Henriksen, 1939) : England (Tulloch, 1939) : ting, 1955, 1956) : Adventive (9) ; : Adventive. Den- origin mostly Jamaica. Adventive, from Brazil. England (Bun- Adventive, from Dominica; common. BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH Platyzosteria bifida, Nebraska (Hebard, 1917) Plectoptera dorsalis, : & WILLIS I5I Adventive. Puerto Rico (Rehn and Hebard, 1927) : Cap- tured by beating banana plants. Pycnoscelus surinamensis, Canada (Walker, 1912; Hebard, 1917) Adventive: Marquesas Islands, Nukubiva (Hebard, 1933a): In Adventive, doing conbanana leaves. England (Goodliffe, 1958) conservatory. in a growing plants damage banana siderable to AdvenSihyllohlatta panesthoides, Massachusetts (Rehn, 1937a) : : : tive, from Jamaica. Family ZINGIBERACEAE Renealmia Associate. — Cariblatta The male was taken in a Jamaica (Rehn and Hebard, 1927) head of wild ginger. Family sp. —Periplaneta americana, Hawaii (Zimmerman, Family : CANNACEAE Canna Associate. sp. orestera, 1948). ORCHIDACEAE Cattleya sp. Associates.—Periplaneta americana, \].S. A. (Rau, 1940a). Periplaneta australasiae, England (Lucas, 1918). Vanda sp. Associates. —Periplaneta americana, \].S.K. (Rau, 1940a). Periplaneta australasiae, England (Lucas, 1918). Undetermined orchids Associates. —Blaherus discoidalis, Blatta oricntalis, Periplaneta americana, Hawaii (Swezey, 1945)Blatta orientalis, americana, cincrea, madcrac, unidentified cockroaches, England, in bulb from Ecuador (Westwood, 1876). Graptohlatta notulata, Hawaii (Swezey, 1945) : On orchid from India. Homalopteryx laminata and Hormetica apolinari, In orchids shipped from Colombia, ard, 1912c) New York (Heb- : Pelmatosilpha coriacea, Puerto Rico (Wolcott, 1936). Periplaneta americana, Germany (Tashenberg, 1884). Periplaneta australasiae, England (Wainwright, 1898). vania (Skinner, 1905). Massachusetts (Morse, 1920). Pennsyl- SMITHSONIAN MISCELLANEOUS COLLECTIONS 152 VOL. I4I Pycnoscelus surinamensis, England (Westwood, 1869). Germany Massachusetts (Morse, 1920). Hawaii (Swezey, (Zacher, 1920). 1945)- Family CASUARINACEAE Casuarina sp. Associate. — Diploptcra punctata, Hawaii (Zimmerman, 1948). SALICACEAE Family Populus euphratica Oliv. —Populus Schrenk. [Howard, communication, 1959]. Associate. —Ectohius semenovi, Kazakhstan (Bei-Bienko, Synonymy. diversifolia Populus Associate. 1950) : On —Ectohius 1950). sp. lapponicus, U.S.S.R. (Stark in Bei-Bienko, aspen. Salix Associate. —Ectohius On personal sp. semenovi, Kazakhstan (Bei-Bienko, 1950) : willow. Family MYRICACEAE Myrica cerifera Linnaeus Associate. — Chorisonenra 1916) : On texensis, Florida bayberry. Florida (Blatchley, (Rehn and Hebard, Beaten from 1920) : foliage. Family FAGACEAE Quercus alba Linnaeus Associate. —Parcohlatta pensylvanica, Virginia 191 6) : Under (Rehn and Flebard, signs on white oaks. Quercus rubra Linnaeus Associates. — Parcohlatta vanica, divisa, Virginia, and Parcohlatta pensylUnder signs on North Carolina (Rehn and Hebard, 19 16) : red oak. Parcohlatta lata. North Carolina (Hebard, 1917) : Under sign. Quercus virginiana Mill. Associate. —Eurycotis floridana, Under dead bark on live-oak 1916) In cavity in tree. : Georgia tree. (Rehn and Hebard, Georgia (Hebard, 1917) : BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS I53 Quercus spp. Associates. —Aglaopteryx gemma, Alabama, Georgia, Florida, Lou- Texas (Hebard, 19 17) Under signs on oaks. Under bark. Blatta orientalis, England (Donisthorpe, 1918) isiana, : : Cariblatta lutea lutea, Mississippi (Hebard, 1916a) low oaks on By : beating hills. Chorisoneura (Blatchley, 1920) : By (Hebard, Mississippi texensis, Florida 1917). beating. Massa- Ectobius pallidus, England (Milton, 1899; Burr, 1899b). chusetts (Flint, 195 1 ) Parcohlatta divisa, Under : loose lichens and bark. Georgia, Louisiana, and Parcoblatta pensyl- Under signs. Florida (Rehn and Hebard, 1905) vanica, Georgia (Hebard, 191 7) Periplaneta australasiae, specimens taken from under a : : Ten tin sign. Periplaneta brunnea, Georgia : Under By shaking (Rehn and Hebard, 1916) signs. Phyllodromica megerlei, U.S.S.R. (Bei-Bienko, 1950) : oak branches. Plectoptera lacerna, Cuba (Rehn and Hebard, 1927). Family MORACEAE Cecropia Associate. — Cariblatta Family hylaea, sp. Honduras (Rehn, 1945a). CHENOPODIACEAE Beta maritima Linnaeus Associate. —Ectobius panseri,'E.ng\2ind (Lucas, 1920a). Beta vulgaris var. Associate. —Ectobius pallidus, cicla Linnaeus 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. 1912). — Chorisoneura texensis, Florida (Rehn and Hebard, SMITHSONIAN MISCELLANEOUS COLLECTIONS 154 Family VOL. I4I SARRACENIACEAE Only a few records have been found of cockroaches being trapped 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 in the pitchers of carnivorous plants of this Anderson, 1939; Lloyd, 1942). Sarracenia flava Linnaeus Natural prey— Cariblatta lutea Parcoblatta lata, lutca, Ischnoptera deropeltiformis, and nymphs of Parcoblatta (Wray and sp.. North Carolina Brimley, 1943) Most of the cockroaches seemed to have been trapped accidentally with the possible exception of C. lutea lutea, II of : which were found in Sarracenia pitchers. Sarracenia purpurea — Cariblatta lutea Natural prey. lutea, Linnaeus North Carolina (Wray and Brimley, 1943). Sarracenia minor Walter Synonymy. —Sarracenia variolaris Michx. [Howard, personal com—Periplancia Florida munication, 1958]. Natural and experimental prey. (Treat, 1876) australasiac, After the insect imbibed some of the : fluid in the 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). pitcher it ; Family NEPENTHACEAE Nepenthes ampularia Jack Natural prey. — Cockroaches, Singapore (Dover, 1928). Nepenthes Natural prey. gracilis Kortli. — Cockroaches, Singapore Nepenthes Natural prey. The insect — Cockroach, was apparently by a piece of (Dover, 1928). sp. Old World Tropics? (Hooker, 1874) where it drowned, : attracted into the pitcher, cartilage placed there by Hooker. BIOTIC ASSOCIATIONS OF COCKROACHES & WILLIS 1 55 CUNONIACEAE Family Weinmannia Associates. —Aneurina —ROTH sp. Marquesas Islands, Nukuhiva and viridis, Fatuhiva (Hebard, 1933a). Maretina uahuka, Marquesas Islands, Uahuka (Hebard, 1933a). Family HAMAMELIDACEAE Liquidambar styraciflua Linnaeus Associate. Under —Parcohlatta sign on sweet divisa, Georgia (Rehn and Hebard, 1916) : gum. Parcoblatta zebra, Louisiana (Hebard, 1917) : In decay cavity. ROSACEAE Family Crataegus sp.? Associates. —Plectoptera dorsalis, Plectoptera infulata, Plectoptcra rhahdota, Puerto Rico (Wolcott, 1950) : In the dry flower clusters of "espino rubial." Rosa sp. Associate. —Pycnoscelus surinamensis, Connecticut (Zappe, 1918) ; Pennsylvania and Pennsylvania (Caudell, 1925) Feeding on canes in greenhouses. (Doucette and Smith, 1926) Rhode Island ; : Rubus Associate. species is —Hololampra spp.? This Azores (Chopard, 1932) and was found only by beating the was very common in hedges, particu- chavesi, : exclusively dendricolous bushes on which it abounds. It larly on brambles (ronces). Family LEGUMINOSAE Acacia farnesiana Willd. Associate. 191 5 ; —Diploptera Zimmerman, 1948) punctata, Hawaii : Acacia Associates. —Ellipsidion (Bridwell and Swezey, Feeding on pods. australe, sp. Australia, New South Wales (Hebard, 1943). Methana curmgera, Australia, Queensland (Pope, 1953a). SMITHSONIAN MISCELLANEOUS COLLECTIONS 156 VOL. I4I Ceratonia siliqua Linnaeus Associate. —Diploptera punctata, Damaging algarroba. liams, 1938) : Hawaii (Pemberton and Wil- Erythrina glauca Willd. Associates. —Aglaopteryx absimilis, Puerto Rico (Wolcott, 1950) : In abandoned cocoon. Aglaopteryx fades, Puerto Rico (Wolcott, 1936) In empty co- : coons. Inga laurina Willd. Associate. —Aglaopteryx fades, Puerto Rico (Wolcott, 1936) : On trunk. Inga vera Willd. Associates. —Aglaopteryx fades, Puerto Rico (Wolcott, 1936) : In larval tents. Cariblatta stenophrys, Puerto Rico (Wolcott, 1936) : On leaves. Plectoptera dorsalis, Plectoptera infulata, and Plectoptera rhahdota, 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 Family (Zimmerman, 1948). ZYGOPHYLLACEAE Tribulus sp. Associates. — Periplanefa americana and Pycnoscelus surinamensis, Johnston Island (Bryan, 1926). Zimmerman (1948) as a host plant for these cockroaches. lists Tribulus BIOTIC ASSOCIATIONS OF COCKROACHES Family —ROTH & WILLIS I57 RUTACEAE Citrus aurantifolia Swingle Associates. —Plectoptera dominicae and Plectoptera perscita, Do- minica (Rehn and Hebard, 1927) : Beaten from moss-covered lime trees. maxima Merr. Citrus Associates. —Plectoptera dorsalis, Plectoptera infulata, Plectoptera rhahdota, Puerto Rico (Wolcott, 1950). Plectoptera rhahdota, Puerto Rico (Rehn and Hebard, 1927). Citrus sinensis Osbeck Associate. — Diploptera punctata, 1915; Zimmerman, 1948) Hawaii (Bridwell and Swezey, 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 rhahdota, Puerto Rico (Wolcott, 1950) Family : In the dry flower clusters. BURSERACEAE Bursera simaruba (L.) Sarg. — Chorisoneura texensis, Florida (Rehn and Hebard, Hebard, 1917) Beaten from the lower branches of gumbo Associate. 1912; : limbo. EUPHORBIACEAE Family Poinsettia sp. Associate. —Pycnoscelus surinamensis, Connecticut Ate bark of greenhouse (Zappe, 1918) : plants. Family ANACARDIACEAE Mangifera indica Linnaeus Associate. 191 5 ; —Diploptera Zimmerman, 1948) punctata, Hawaii : (Bridwell and Swezey, Feeding on mangoes on the tree. SMITHSONIAN MISCELLANEOUS COLLECTIONS 158 VOL. I4I Spondias mombin Linnaeus Associates. —Plectoptera dorsalis, Plectoptera infulata, Plectoptera rhabdota, Puerto Rico (Wolcott, 1950) : Living on leaves of "jobo." Spondias purpurea Linnaeus Associate. — Eiirycofis crown of dry biolleyi, Costa Rica (Rehn, 1918) In the : jocoto. AQUIFOLIACEAE Family Ilex cassine Linnaeus Associate. —Plectoptera poeyi, Florida (Rehn and Hebard, 1912, 1914; Hebard, 1917). Ilex coriacea Synonymy.— Ilex Associate. (Pursh) Qiapm. lucida [Fernald, 1950]. — Cariblatta lutea lutea, Florida (Hebard, 1916a). SAPINDACEAE Family Exothea paniculata (Juss.) Radlk. Associate. —Aglaopteryx gemma, Family Florida (Hebard, 1917). MALVACEAE Gossypium Associates. — Graptohlatta spp. 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 Associate. —Riatia orientis, Trinidad (Princis and Kevan, 1955). Hibiscus Associates. cotis kevani, —Amazonina sp. emarginata, Cariblatta antiguensis, Eury- and Rhytidometopnm Kevan, 1955). Linnaeus dissimile, Trinidad (Princis and BIOTIC ASSOCIATIONS OF COCKROACHES Sida Associate. —Periplaneta — ROTH & WILLIS 1 59 sp. australasiae, Nihoa Island (Bryan, 1926). Hawaii (Zimmerman, 1948). STERCULIACEAE Family Theobroma cacao Linnaeus Associate. — Ceratinoptera picta, Trinidad (Princis and Kevan, 1955)- Family BIXACEAE Bixa Associate. 1955) : sp. —N otolampra antillarum, Trinidad Nymphs in dry fruits on "annato" Family (Princis and Kevan, tree. FLACOURTIACEAE Xylosma suaveolens Associate. — Graptohlatta notulata, Forst. Marquesas Islands, Uahuka (Hebard, 1933a). PASSIFLORACEAE Family Passiflora sp. Associate. —Arisfiger [=Plnmiger] histrio, Malaya (Karny, 1924). Family CARICACEAE Carica papaya Linnaeus Associate. 191 5 ; — Diploptera Zimmerman, 1948) Family punctata, Hawaii : (Bridwell and Swezey, Feeding on papaya fruit on tree. RHIZOPHORACEAE Rhizophora mangle Linnaeus Associate. —Aglaopteryx Family gemma, Florida (Hebard, 1917). COMBRETACEAE Conocarpus erectus Linnaeus Associate. Running on —Plectoptera poeyi, Florida leaves. (Rehn and Hebard, 1914) : SMITHSONIAN MISCELLANEOUS COLLECTIONS l6o MYRTACEAE Family Eucalyptus Associate. —Ellipsidion VOL. I4I australe, sp. New Australia, Wales South (Hebard, 1943). Eugenia aromatica Baill. —Syzygium aromaticuni Puerto Associate. —Plectoptera Synonymy. [Bailey, 1925]. Rico dorsalis, On (Wolcott, 1936) : flowers of "pomarrosa." Metrosideros coUina Gray Associates. —Anenrina viridis, Marquesas Islands Nukuhiva, : Fatuhiva, and Tahuata (Hebard, 1933a). Aneurina tahuata, Marquesas Islands, Tahuata (Hebard, 1933a). Graptohlatta notulata, Marquesas Islands, Nukuhiva (Elebard, 1933a). Psidium guajava Linnaeus Associate. —Plectoptera rhahdota, Puerto Rico (Rehn and Hebard, 1927). Family ONAGRACEAE Jussiaea natans Associate. Humb. and Bonpl. —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 Family where the insect was captured. ERICACEAE Calluna vulgaris Salisb. Associates. "Nymphs —Ectobius lapponicus, England of varying size were beaten out of February and (Lucas, heather 1925) : ... on 9 later." Ectobius panseri, England (Lucas, 1927) both sexes were swept from heather." : "numerous imagines of Vaccinium meridionale Sw. Associates. — Chorisoneura blattella proserpina, formosella, Neoblattella dryas, Neo- Jamaica (Rehn and Hebard, 1927). Family SAPOTACEAE Sideroxylon foetidissimum Jacq. Associate. Under bark. —Pelmatosilpha coria^ea, Puerto Rico (Wolcott, 1941) : BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS l6l APOCYNACEAE Family Vinca minor Linnaeus Associate. — Ectobius (Willis, unpublished pallidus, Massachusetts observation, 1958). Family CONVOLVULACEAE Ipomoea Associate. —Plectoptera tiliasea dorsalis, Choisy Puerto Rico (Rehn and Hebard, 1927). BORAGINACEAE Family Cordia dentata Poiret Synonymy. — Calyptracordia alba [Howard, personal communica1958]. Associates. — Cariblatta antiguensis, Ischnoptera rufa rufa, Supella tion, supellectilium, Symploce ruficollis and Symploce hospes, St. Croix, Virgin Islands (Beatty, 1944) On fruits of C. dentata except supellectilium which was found at night on the flowers. : .S. VERBENACEAE Family Citharexylum villosum Jacq. Associate. — Chorisoneura texensis, Florida (Rehn and Hebard, 1912). Family SOLANACEAE Nicotiana Associate. —Pycnoscelus sp. surinamensis, Sumatra (Roeser, 1940). Solanum tuberosum Linnaeus — Associate. Pycnoscelus surinamensis, Haiti Feeding on tubers in field. Family —Aneurina 1927) : GESNERIACEAE Cyrtandra Associate. (Hoffman, viridis, sp. Marquesas Islands, Nukuhiva (He- bard, 1933a). Family RUBIACEAE Canthium barbatum (Forst.) Seem. Associate. — Graptoblatta (Hebard, 1933a). notidata, Marquesas Islands, Uahuka SMITHSONIAN MISCELLANEOUS COLLECTIONS l62 VOL. I4I Cinchona pubescens Vahl. Associate. —Periplaneta americana, ters, 1949) Puerto Rico (Plank and Win- In greenhouse. : Coffea sp. Associate. —Plectoptera porcellana, Family Puerto Rico (Sein, 1923). COMPOSITAE Goldenrod Associate. —Eurycotis floridana, Florida (Hebard, 1917) : "Climb- ing about on top of goldenrod at night." Helianthus Associate. —Pseudomops sp. 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 maDamage to living plants occurs principally in the Tropics or terial. under subtropical conditions in greenhouses in temperate regions. Among the depredations attributed to cockroaches in text books, damage to plants the many is seldom emphasized. This is surprising in view of 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, and the I ordered inside of the ship to be cockroaches. We were all the chests to be taken on shore, washed with boiling water, to kill the 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 BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 163 orchids as well as the aerial roots and flower spikes of Vanda orchids (Rau, 1940a). Wainwright (1898) stated that Periplaneta austra- 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 lasiae had been observed in 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 Graptohlatta notulata had been inter- cepted at Honolulu on orchids from India. Watson (1907) stated that Blatta orientalis, Periplaneta americana, 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 and Periplaneta australasiae : 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. 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, australasiae 1891). Laing (1946; British stated that P. australasiae Museum abounds in [Natural History], 1951) greenhouses and forcing pits SMITHSONIAN MISCELLANEOUS COLLECTIONS 164 where inosa it may do damage considerable to the plants. Periplaneta fulig- also troublesome in greenhouses because of is VOL. I41 feed on seedlings and succulent plants (Dodge and its tendency to Rickett, 1943). Ectobiits lapponicus has been observed feeding in galleries in the thick skin of young aspen in Bei-Bienko, 1950). in The 25 percent of the trees examined (Stark aquatic cockroach Epilampra abdomen- nigrum fed on the leaves of well, 1946) . an aquarium (Croit was dense woods in Indiana Jitssiaea natans in Ischnoptera deropeltiformis has been taken while feeding on a fleshy fungus {Agaricus sp.) in (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." FuUaway 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 un- doubtedly one of the economically most important cockroaches, being the vector of the chicken eyeworm as well as feeding on plants. In damage caused by this species that have addition to the few reports of already been mentioned, P. surinamensis has been reported to be very destructive in New Orleans to palms and ferns, attacking large al- (Anonymous, 1893). Zappe (1918) in Connecticut reported damage in a greenhouse to roses P. surinamensis had valued, at that time, at several hundred dollars 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. surinamensophilas avidly, eating out the hearts ; BIOTIC ASSOCIATIONS OF sis to COCKROACHES — ROTH Rhode the stems of rose bushes has been reported in Pennsylvania ; & WILLIS 165 Island and the canes were attacked both under and above ground (Caudell, 1925). Doucette and Smith (1926) reported a heavy in- festation 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, About 30,000 to 35,000 rose plants from a 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 and other agencies . . . total 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). lUingworth (1927, 1929) reported that in Hawaii P. surinamensis was a minor pest of pineapples, feeding on 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 bethe roots. came poor in food material. Zimmerman (1948) of P. surinamensis in Hawaii listed as host plants "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 legumes" were attacked by P. surinamensis and that Zea mays Linnaeus was attacked by Supella supeUectiliimi. Wolcott (1924a) reported that P. surinamensis damaged transplanted tobacco plants in Puerto Rico by eating the interior of the Cuba consider P. surinamensis stalks. Tobacco planters in injurious to the roots of tobacco plants this belief was confirmed in the (Bruner and Scaramuzza, 1936) 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. ; laboratory, SMITHSONIAN MISCELLANEOUS COLLECTIONS l66 Blattella may vaga VOL. I4I damage seedlings in the laboratory damage has been reported in the field (Ball occasionally (Flock, 1941a), but no 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. Golden- et al., 1942), berg (1877) stated that sago trees provide cockroaches with their favorite nourishment. Scudder (1879) found Eurycotis floridana living in the tops of the cabbage palmetto, fed. feet on which he presumed it Parcoblatta americana has been observed feeding on an apple 6 above ground (Fulton, 1930). X. PROTOZOA ASSOCIATED WITH COCKROACHES The classification of The use of the asterisk the Protozoa follows that of (*) is Phylum Class explained in footnote 3, Kudo page (1954). 4. PROTOZOA MASTIGOPHORA EUGLENOIDINA Family EUGLENIDAE Order Euglena Experimental host. sp. —Periplaneta americana, U.S.A. (Hegner, When fed to the insects in concentrated culture, Euglena 1929) could withstand conditions in the crop up to 5 hours and were passed : stomach were killed into the in a viable state jority in the stomach up to 6 hours. However, the ma- crop within 2 hours and very few reached the alive. Order PROTOMONADINA Family OIKOMONADIDAE Oikomonas blattarum Tejera Natural host. — Cockroach, Venezuela (Tejera, 1926). Oikomonas Natural host. 1922) : —Blatta Oikomonas sp. orientalis, and Monas sp. U.S.S.R. (Yakimov and Miller, sp. were found 83 percent of 124 cockroaches. Cockroach, Venezuela (Tejera, 1926). in the intestines of COCKROACHES BIOTIC ASSOCIATIONS OF Family —ROTH & WILLIS 167 TRYPANOSOMATIDAE Leptomonas blaberae Tejera Natural host. — Blaherus sp., Venezuela (Tejera, 1926). Leptomonas Natural hosts. —Parcohlatta lata, sp. Parcohlatta pensylvanica, Parco- Ohio (Semans, 1939, 1941) Hind intestine. Of 70 specimens examined, 86 percent harbored Leptomonas sp. blatta virginica, U.S.A., : *Herpetomonas periplanetae Laveran and Franchini Natural host. chini, 1920, —Blatta orientalis, Italy, France (Laveran and Fran- 1920a). MONADIDAE Family Monas —Blatta Natural host. orientalis, sp. U.S.S.R. (Yakimov and Miller, Monas sp. and Oikomonas sp. were found 1922) percent of 124 cockroaches examined. 83 : in the intestines of Cockroach, Venezuela (Tejera, 1926). Family BODONIDAE Bodo blattae Natural host. —Blatta orientalis, Bodo Natural host. — Blattella England (Lankester, 1865). sp. germanica and/or Periplaneta americana, South Africa (Porter, 1930). Retortamonas blattae (Bishop) Synonymy. —Emhadomonas —Blatta blattae Bishop [Wenrich, 1932]. England (Bishop, 1931): Hind intestine. The organism occurred in about 40 percent of the cockroaches examined. L. G. Feo {in Wenrich, 1932) successfully culNatural host. orientalis, tured this protozoan (fig. 2, F). Retortamonas sp.? Natural host. and Chu, 1930). —Leucophaea maderae, Philippine Islands (Hegner ; SMITHSONIAN MISCELLANEOUS COLLECTIONS i68 — VOL. I4I Fig. 2. Representative Protozoa associated with cockroaches. A, Monocercomonoidcs mrlolonthae, X 3094 (after Grasse). B, Coelosporidiutn periplanetae, 1310 (after Sprague) trophozoite with spores and chromatoid bodies. C, Endamocba blattae, X 273 (after Kudo) trophozoite. D, Lophomanas striata, X 330 (after Kudo). E, Lophomonas blattartim, X 660 (after Kudo). F, Retortamonas blattae, X 3004 (after Wenrich). G, Nyctotherus ovalis, I75_ (after Kudo). H, Gregarina rhyparobiae, c. 52; mature trophozoite attached to intestinal wall of Leucophaea maderae. (Redrawn from J. M. Watson [1945].) I, Diplocystis srhneideri, c. X 14.4 (after Kunstler). J, GrtKiarina blaftarnm, c. 57 (after Kudo). K, Protomagalhaesia serpentula. X 36 (after Pinto). L, Gamocysti.'! tena.r, magnification not known (after Schneider). (All figures except redrawn from Kudo [1954! after sources X ; X X X H indicated.) COCKROACHES BIOTIC ASSOCIATIONS OF —ROTH Order POLYMASTIGINA Family CHILOMASTIGIDAE & WILLIS 169 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 germanica and/or Blattella host. Periplaneta americana, South Africa (Porter, 1930). Monocercomonoides globus Cleveland Natural host. et al. — Cryptocercus pimctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et 1934) al., Organism occurs : in practically all hosts, Monocercomonoides melolonthae (Grassi) (Fig. Natural host. (Laird, 1956) —Platysosteria Found : in the 2, A) novae seelandiae, New Zealand intestinal tracts of the adult cock- roaches, and of other species of insects. Monocercomonoides orthopterorum Synonymy. — Trichomonas (Parisi) {Trichomastix) orthopterorum Parisi; Monocercomonas orthopterorum [Belar, 1916] ; Trichomastic or- thopterum? [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). Ectohius lapponicus, Italy (Parisi, 1910). Periplaneta americana, Philippine Islands (Hegner and Chu, 1930). "Kiichenschaben," Austria (Belar, 191 6). The protozoan is found ; in the hind gut. Zasukhin (1930) found SMITHSONIAN MISCELLANEOUS COLLECTIONS 170 the organism in 85 percent of over 3,000 B. orientalis. found the flagellate present in very large numbers. Monocercomonoides panesthiae Natural host. der, 1937) : —Panesthia angustipennis, Natural hosts. —Blatta saline Kidder Philippine Islands (Kid- Young orientalis, Blattella gcrmanica, Periplancta (Young, 1935) The organism when present large numbers in the posterior part of the intestine near the americana, U.S.A. occurs in Parisi (1910) In hind gut. Tetratrichomastix blattidarum : The protozoan was anus. VOL. I4I successfully cultivated in a hemoglobin- medium. Family OXYMONADIDAE Oxymonas doroaxostylus (Cleveland Synonymy. land, 1950]. et al.) —Saccinohaculus doroaxostylus Cleveland [Cleve— Cryptocercus punctulatus, U.S.A., Appalachian and et al. Natural host. Pacific coast areas (Cleveland et 1934). al., Oxymonas nana Cleveland [Cleveland, Synonymy. — Saccinohaculus minor Cleveland 1950]— Cryptocercus punctulatus, U.S.A., Appalachian and Natural et al. host. Pacific coast areas (Cleveland et Natural host. 1934). DINENYMPHIDAE Family Saccinobaculus al., ambloaxostylus Pacific coast areas (Cleveland et al., Natural : et al. 1934). Saccinobaculus lata 1950b) Cleveland — Cryptocercus punctulatus, U.S.A., Appalachian and host. — Cryptocercus There are at least Cleveland punctulatus, two other U.S.A. (Cleveland, species of Saccinohaculus 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). BIOTIC ASSOCIATIONS OF COCKROACHES & WILLIS I7I TRICHOMONADIDAE Family * —ROTH Trichomonas hominis (Davaine) — Experimental vectors. Blatta orientalis, South Africa Italy (Mariani and Besta, 1936). 1918) Periplaneta americana, South Africa (Porter, 1918) (Porter, ; U.S.A. ; (Hegner, 1928). Trichomonas Natural vector. found sp. —Cockroach, Venezuela (Tejera, 1926) : Organism in digestive tract of the cockroach. HEXAMITIDAE Family Hexamita Natural hosts. cryptocerci — Cryptocercus Cleveland et al. punctulatus, U.S.A. (Cleveland et 1934). Panesthia angustipennis, Philippine Islands (Kidder, 1937). al., Hexamita periplanetae (Belar) Synonymy. — Octomitus periplanetae Belar —Blatta U.S.S.R. Natural hosts. ganism is found [Kudo, 1954]. (Zasukhin, 1930) Orhind gut. Eighty-five percent of over 3,000 B. orientalis, in the : orientalis contained this organism. Periplaneta americana, Philippine Islands (Hegner and Chu, 1930). "Kiichenschaben," Austria (Belar, 1916). Hexamita Natural host. and Chu, 1930) —Leucophaea The : sp.? maderae, Philippine Islands (Hegner flagellates were present * Giardia intestinalis Experimental vectors. — Blatta in large numbers. (Lamb!) orientalis, South Africa (Porter, 1918)- (Pessoa and Corrca, 1927). Eurycotis floridana, U.S.A. (Young, 1937). Leucophaea maderae, Brazil (Pessoa and Correa, 1927). Gold Coast Periplaneta americana, South Africa (Porter, 1918) Blattella gcrnianica, Brazil ; Colony (Macfie, 1922) (Young, 1937). ; Brazil (Pessoa and Correa, 1927) ; U.S.A. Periplaneta brunnea, U.S.A. (Young, 1937). Argentina (Bacigalupo, Cockroaches, Venezuela (Tejera, 1926) ; in Tejera, 1926). SMITHSONIAN MISCELLANEOUS COLLECTIONS 172 Giardia Natural vectors. sp. —Cockroaches, Venezuela Order Family (Tejera, 1926). HYPERMASTIGINA HOLOMASTIGOTIDAE Leptospironympha eupora Cleveland Natural host. (Cleveland —Cryptocercus et al., Natural host. et punctulatus, U.S.A., Appalachian area et al. — Cryptocercus punctulatus, U.S.A., Appalachian area 1934). al., Leptospironympha wachula Cleveland Natural host. (Cleveland al. 1934). Leptospironympha rudis Qeveland (Cleveland et VOL. I4I et al. — Cryptocercus punctulatus, U.S.A., Appalachian area et al., 1934). Macrospironympha xylopletha Cleveland Natural host. — Cryptocercus fornia (Cleveland et al., Family et al. punctulatus, U.S.A., northern Cali- 1934). LOPHOMONADIDAE Lophomonas blattarum Stein Natural hosts. Germany —Blatta orientalis, Czechoslovakia (Stein, i860) ; 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). 1878; (Biitschli, Schubotz, 1905; ; ; Blattella germanica, U.S.A., Ohio (McAdow, 1931). germanica and/or Periplaneta americana, Egypt Coursey and Otto, 1956, 1957). Blattella Periplaneta americana, England nicki, 1910) ; (Schuster, 1898) U.S.A. (Kudo, 1926b; McAdow, ; (De- Europe (Ja- 1931 ; Hatcher, Philippine Islands (Hegner and Chu, 1930). 1939; Armer, 1944) Periplaneta sp., Goa (Mello and Lima Ribeiro, 1924, 1925). ; "Kiichenschaben," Austria (Belar, 1916). The protozoan (fig. 2, E) is found in the host's colon, particularly anterior portion; encysted stages of organism are found throughout Of 1,400 B. orientalis studied, 32 percent harbored this organism (Kudo, 1925, 1926). Yakimov and Miller (1922) found 7 hind gut. BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS I73 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 never present in the host tissue; is it should be con- sidered a commensal (Kudo, 1926). Lophomonas striata Butschli most probably Synonymy. — Lophomonas sulcata Schuster (Kudo, 1926b), with L. —Blatta Germany (Butschli, Natural is identi- striata cal orientalis, hosts. 1878; U.S.A. (Kudo, Europe (Janicki, 1908, 1910) Schubotz, 1905) U.S.S.R. (Yakimov and 1922, 1926, 1926b; McAdow, 1931) ; ; ; Zasukhin, 1930); Poland (Lorenc, 1939). Blattella germanica, U.S.A., Ohio (McAdow, 1931). Blattella germanica and/or Periplaneta americana, South Africa Miller, 1922; (Porter, 1930). Philippine IsPeriplaneta americana, Indochina (Weill, 1929) U.S.A. (Kudo, 1926b; McAdow, ; lands (Hegner and Chu, 1930); 1 931; Armer, 1944). Cockroach, Venezuela (Tejera, 1926) England ; or U.S.A.? (Lucas, 1928). "Kiichenschaben," Austria (Belar, 1916). Found in the host's colon, particularly the anterior portion. L. 29 percent of 1,400 B. orientalis and 30 P. americana (Kudo, 1926, 1926b), Yakimov and Miller (1922) found the organism in 9.6 percent of 124 specimens of B. Zasukhin (1930) found 8.6 percent of over 3,000 B. orientalis. striata (fig. 2, D) was found in in 2 of orientalis infested. Grasse (1926, 1926a) identified corrugations on the surface of L. which he named Fttsiformis lopho- striata as a bacterial parasite monadis. Prolophomonas tocopola Natural ginia, host. Cleveland et — Cryptoccrcns punctulatus, California, Oregon, Vir- West Virginia (Cleveland Family et al., 1934) : Not abundant. HOPLONYMPHIDAE Barbulanympha estaboga Cleveland Natural host. al. et al. — 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). SMITHSONIAN MISCELLANEOUS COLLECTIONS 174 Barbulanympha laurabuda Cleveland Natural host. — Cryptocercus et VOL. I4I al. Appalachian and punctulatiis, U.S.A., This species, B. uf alula, Pacific coast areas (Cleveland et al., and Rhynchonympha tarda occur in all parts of the colon, especially in 1934) : the enlarged, flexed part near the ileum. Barbulanympha ufalula Natural host. — Cryptocercus Cleveland et al. punctulatiis, U.S.A., Pacific coast areas (Cleveland et al., Appalachian and 1934). Barbulanympha wenyoni Cleveland Natural host. — Cryptocercus punctulatus, U.S.A., Pacific coast area (Cleveland, 1953). Rhynchonympha tarda Cleveland (Fig. 3, Natural host. (Cleveland et amined from 1934) al., : Pacific coast. Cleveland talea 3, et Natural host. C) al., 1934) cockroach examined. in every Present in fairly great : STAUROJOENINIDAE Family Idionympha perissa Cleveland host. et al. — Cryptocercus punctulatus, U.S.A., Appalachian area et al., 1934) Family Present in only a few specimens. : TRICHONYMPHIDAE Trichonympha acuta Cleveland Natural al. — Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et Natural Pacific coast area Fairly abundant in every specimen ex- (Fig. (Cleveland al. — Cryptocercus punctulatus, U.S.A., Urinympha numbers et D) host. et al. — Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., 1934). Trichonympha algoa Cleveland (Fig. Natural host. 3, et al. E) — Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et present in most specimens. al., 1934) : Fairly abundant and —Protozoa from the gut of the wood-feeding cockroach Cryptocercus A, Eiiconwnympha itnla, female above, male below, c. X 375. (From Cleveland [iQSOcl.) B, Barbulanympha sp. (From Cleveland [1953].) C, f/nnympha talea, c. X 712. (From Cleveland [iQSia].) D, Rhynchonympha tarda, c. X4S0. (From Cleveland [1952].) E, Trichonympha okolona or T. algoa, c. X 390. (From Cleveland [1949].) (All drawings reproduced through the courtesy of Dr. L. R. Cleveland.) Fig. 3. piinctitlatus. 175 SMITHSONIAN MISCELLANEOUS COLLECTIONS 176 Trichonympha chula Cleveland Natural host. et VOL, I4I al. — Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et Trichonympha Natural host. grandis — Cryptocercus areas (Cleveland et al., 1934) 1934). al., Cleveland et al. U.S.A., punctulatus, Fairly abundant in : all Pacific coast specimens from Pacific area. Trichonympha lata Cleveland Natural host. et al. — Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et 1934). al., Trichonympha okolona Cleveland (Fig. Natural host. 3, et al. E) — 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 Natural host. et al. — Cryptocercus punctulatus, U.S.A., Appalachian and This organism is smaller al., 1934) than any knov^^n species of Trichonympha; it is more resistant to Pacific coast areas (Cleveland et warm : weather than the other hypermastigotes. Family EUCOMONYMPHIDAE Eucomonympha imla Natural host. Cleveland et al. — Cryptocercus punctulatus, U.S.A., Appalachian and Pacific coast areas (Cleveland et al., sometimes becomes attached the to 1934) : Organism intestinal wall ; (fig. 3, attached A) in- dividuals were seen in 2 to 3 percent of the cockroaches examined. Unidentified flagellate Natural host. —Pycnoscelus surinamensis, Hawaii (Schwabe, A small flagellate was found in the digestive tract and mal1950) pighian tubules. : Class Order SARCODINA MYCETOZOA INCERTAE SEDIS Peltomyces periplanetae (Leger) Synonymy. —Peltomyces hlattellae. Sprague (1940a) synonymizes Peltomyces periplanetae, with Coelosporidium, periplanetae. BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH I77 & WILLIS —Blatta orientalis, France (Debaisieux, 1927). France (Leger, 1909; Debaisieux, 1927). The organism inhabits the malpighian tubules of cockroaches. Leger 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 Natural hosts. Blattella germanica, Plistophora periplanetae or Coelosporidium periplanetae. Order AMOEBINA Family AMOEBIDAE Hartmannella blattae Ivanic Natural in the host. — Blatta orientalis, Yugoslavia (Ivanic, 1937) : Found 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 Natural vectors. — Blattella sp. germanica and/or Periplaneta ameri- cana, Egypt (DeCoursey and Otto, 1956, 1957) cockroaches examined contained this protozoan. Endamoeba Synonymy. tarum. —Amoeba —Blatta blattae blattae, : Thirty out of 261 (Butschli) Entamoeba blattae, Entamoeba blat- orientalis, Germany (Biitschli, 1878; SchuU.S.A. (Leidy, 1879a, 1880; Kudo, 1922, Chen, 1933) Meglitsch, 1938, 1925a, 1926a; Kirby, 1927; McAdow, 1931 (Mercier, Europe? (JaFrance 1907a, 1908, 1940); 1909, 1910) nicki, 1908, 1909) U.S.S.R. ( Yakimov and Miller, 1922 Zasukhin, Natural hosts. botz, 1905; ; ; ; ; ; 1929, 1930); England (Thomson and Lucas, 1926; Lucas, 1927, Yugoslavia (Ivanic, 1926a). germanica and/or Periplaneta americana. South Africa Egypt (DeCoursey and Otto, 1956, 1957) Seven (Porter, 1930) out of 217 cockroaches examined harbored the protozoan. 1927a, 1928) ; Blattella : ; Periplaneta 1930) ; americana, Philippine U.S.A. (Morris, 1936; (Macfie, 1922). Islands Armer, 1944) ; (Hegner and Chu, Gold Coast Colony SMITHSONIAN MISCELLANEOUS COLLECTIONS 178 Periplaneta australasiae, U.S.A. (Morris, 1936). Cockroaches, Paraguay ? (Elmassian, 1909) Austria ; 1916) ; U.S.A. (Morris, Balch, 1936; 1935, VOL. I4I (Belar, 1932); Venezuela (Tejera, 1926). The habitat of E. hlattae (fig. 2, The of the cockroach. C) is the hind intestine and rectum Kudo (1925a) percent in March incidence of infection varies: found in 1,255 oriental cockroaches infections in 5 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 parasitic in the nucleus of fungus, Endamoeba Nucleophaga sp., hyper- blattae. Endamoeba javanica Kidder Natural hosts. —Panesthia angustipennis, Philippine Islands, and Occurred in 50 percent of Panesthia spadica, Japan (Kidder, 1937) P. angustipennis examined and in one of four P. spadica. The en: doplasm of this amoeba contains large amounts of wood and cellulose fibers. Endamoeba Natural host. der, 1937) The food : philippinensis Kidder —Panesthia angustipennis, Occurred Synonymy. —Endamoeba —Blaberns coli, Natural vectors. lot examined. vacuoles contained bacteria, no wood. Entamoeba a Philippine Islands (Kid- in about 10 percent of the Panesthia coli (Grassi) Amoeba coli [Kirby, 1945]. atropos, Venezuela (Tejera, 1926) : In of 60 cockroaches captured in latrines, two were found that carried apparently live cysts similar to cysts of E. coli. germanica or Periplaneta americana, Egypt (DeCoursey One out of 44 cockroaches collected in a and Otto, 1956, 1957) Blattella : village harbored E. coli. Experimental vector. (Macfie, 1922) : —Periplaneta americana, Gold Coast Colony In nine experiments cysts oi E. coli were fed to the BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 1/9 cockroaches. In seven of the experiments cysts of E. coli were found Cysts were observed in the feces for only one to three in the feces. days, and eventually disappeared completely. The cysts appeared to No amoebae were found. be unharmed. Entamoeba Natural vectors. —Blatta histolytica Schaudinn orientalis, Blattella germanica, Periplaneta americana, Periplaneta australasiae , and/or Sitpella 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). U.S.A. Periplaneta americana. Gold Coast Colony (Macfie, 1922) — ; (Frye and Meleney, 1936). Cockroaches, Venezuela (Tejera, 1926). Entamoeba Experimental pitheci vector. —Periplaneta Prowazek? americana, Formosa (Mori- 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. schita : Entamoeba thomsoni Lucas Synonymy. tion, 1957]. —Endamoeba —Blatta Natural hosts. thomsoni England (Lucas, 1927a, 1928) orientalis, U.S.A. (Taliaferro, 1928; [Kudo, personal communica- McAdow, 1931); U.S.S.R. 1930); Germany (Chen, 1933). Blattella germanica, U.S.A. (McAdow, 1931). Periplaneta americana, England (Lucas, 1927a) and Barret, 1928; McAdow, 1931) ; ; ; (Zasukhin, U.S.A. (Smith 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 fie, 1922) : Under the heading "Entamoeba histolytica (Macand E. coli" SMITHSONIAN MISCELLANEOUS COLLECTIONS l80 Macfie (p. 445) stated, "The cockroaches used in these VOL. I4I 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 Entamoeba, resembling E. coli, from feces of the monkey [Erythrocebus patas patas (Schreber)] were fed to cock(Macfie, 1922) roaches, 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) ; U.S.A. (Armer, 1944). Periplaneta australasiae, U.S.A. (Steinhaus, 1946). Organism is found in the hind gut of the cockroach. (Weill, 1929) ; Indochina ; (1930) found 3-percent infestation in Zasukhin over 3,000 B. orientalis ex- amined. Endolimax nana (Wenyon and O'Connor) ? Synonymy. —Entamoeba nana. —Blaberus atropos, Natural Venezuela (Tejera, 1926) A nana was found in the intestinal contents of the cockroach. In inoculations this amoeba was not pathohost. small amoeba : greatly resembling E. genic. Endolimax —Blatta Natural hosts. organism was found sp. U.S.S.R. (Zasukhin, 1930) This hind gut of 0.3 percent of over 3,000 cock- orientalis, in the : roaches examined. Blattella germanica and/or Periplaneta americana, Egypt (DeCoursey and Otto, 1956, 1957) Seventy-four out of 261 cockroaches examined harbored this protozoan. : lodamoeba sp. — Blattella garmanica and/or Periplaneta ameriFifty-nine of 261 Egypt (DeCoursey and Otto, 1956, 1957) cockroaches examined contained this protozoan. lodamoeba sp. was common in human feces in villages in which the cockroaches were Natural vectors. cana, collected. : BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS l8l Undetermined species of Amoeba Natural host. der, 1937) —Panesthia angustipennis, Philippine Found : in only Islands. (Kid- (Woodcock, 1904; one specimen. SPOROZOA Class Order GREGARINIDA Family DIPLOCYSTIDAE Diplocystis schneideri Kunstler (Fig. Natural hosts. —Blatta I) 2, orientalis, England Jameson, 1920). Periplaneta americana, France (Kunstler, 1884, 1887) England ; Germany (Foerster, 1939). (Jameson, 1920) 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 There coelom. is no apparent pathogenic effect. Jameson (1920) found 81 percent of P. americana infested with D. schneideri. Diplocystis sp. Natural host. Heavy —Periplaneta infections in body americana, U.S.A. (Hertig, 1921): cavity. A large number of Cockroach, India (Ray and Dasgupta, 1955) collected in Calcutta were all cockroaches, both adults and nymphs, : infected. Diplocystis Natural host. —Blaherus craniifer, sp. ? U.S.A. (Nutting, 1953) more paired trophozoites or cysts may be found mocoele and occasionally in the thorax. I to 12 or : From in the he- STENOPHORIDAE Family Stenophora sp. —Blatta orientalis, India (Bal and Rai, 1955) Natural host. ganism found in the midgut of the cockroach. Family : Or- GREGARINIDAE Gregarina blattarum von Siebold Synonymy. — Gregarina —Blatta Natural hosts. hlattae orientalis; orientalis, Germany Clepsidrina blattarum. (Siebold, 1837, 1839; SMITHSONIAN MISCELLANEOUS COLLECTIONS l82 VOL. I4I Marshall, 1892; WellWolters, 1891 Stein, 1848; Butschli, 1881 Foerster, 1938; Schubotz, 1905); U.S.A. (Leidy, mer, 1910, 1911 McAdow, Kudo, 1922 Watson, 1917 Crawley, 1903 1853a France (Lankester, England Sprague, 1863); 1940, 1941) 1931; Laveran and Franchini, 1920a); (Schneider, 1875; Cuenot, 1901 ; ; ; ; ; ; ; I ; Brazil (Magalhaes, 1900; Pinto, 1919) ; U.S.S.R. (Zasukhin, 1929, 1930)- South Africa (FanU.S.A. (Crawley, 1903) these appear to be the same record). Blattella gennanica and/or Periplaneta americana, Egypt (De- Blattella germanica, tham, 1929 ; ; Porter, 1930 : Coursey and Otto, 1956, 1957). Periplaneta americana, Brazil (Magalhaes, 1900) McAdow, ; U.S.A. (Craw- South Africa (Fantham, 1929; 1931) 1903; Gold Coast Porter, 1930: these appear to be the same record) Colony (Macfie, 1922), 1907; ley, ; ; Parcohlatta pensylvanica, U.S.A., Michigan (Ellis, 1913a). Germany Cockroaches, (Schififmann, 1919: probably used the ori- Venezuela (Tejera, 1926). Organism usually found in the intestinal tract of cockroaches where Cysts are passed in the feces. Occait is attached to the gut cells. sionally, G. bhittarmn (fig. 2, J) is found in the body cavity (Leidy, 1853a; Hall, 1907). Though this is considered to be one of the ental cockroach) ; 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. Wat- son (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 — All Natural host. Aptera fusca, South Africa (Harrison, 1955) 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 — Temnopteryx phalerata, South Africa (Harrison, 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 Natural host. 1955) : cysts w^ere found in the hind gut or rectum. BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 183 Gregarina illinensis M. E. Watson Natural host. 1 91 6) —Parcohlatta pensylvanica, U.S.A., The intestine of 191 of these gregarines. 5, : (Watson, Illinois one cockroach was found to contain 25 Gregarina impetuosa Harrison Natural host. —Melanosilpha (Harrison, South Africa capensis, All specimens of this gregarine were found in the anterior 1955) mesenteron of the host. : Pinto Gregarina l^geri Natural 1918a, 1919) host. : —Periplaneta americana, Brazil (Pinto, 1918, Intestinal canal. Gregarina neo-brasiliensis Al. Cunha Natural host. Cunha —Periplaneta americana, Natural : (R. Almeida de Semans Gregarina ohioensis 1939) midgut. Brazil 1919; Cunha, 1919). in Pinto, host. —Parcohlatta The protozoan was U.S.A., virginica, Ohio (Semans, present in large numbers in the insect's Gregarina panchlorae Frenzel Natural host. —Panchlora exoleta, Argentina (Frenzel, 1892) : Midgut. Gregarina parcoblattae Semans Natural hosts. riana, U.S.A., —Parcohlatta pensylvanica and Parcohlatta uhle- Ohio (Semans, 1939) : Gregarina rhyparobiae Midgut. J. M. Watson — Natural host. Lciicophaca 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. • : ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 184 VOL. I4I Gregarina thomasi Semans Natural host. 1939) : —Parcohlatta pensylvanica, U.S.A., Ohio (Semans, Enteric caeca and midgut. Protomagalhaesia serpentula Synonymy. 1943]- (de Magalhaes) — Gregarina serpentula [Pinto, 1918a, 1919; Semans, —Periplaneta americana, Brazil (Magalhaes, 1900) Natural host. The In the coelom and alimentary canal. K) was host of this parasite (fig. 2, Watson (1916). incorrectly cited as Blatta orientalis by Gamocystis tenax Schneider (Fig. Natural hosts. —Ectobius Germany (Wellmer, L) 2, lapponicus, France (Schneider, 191 o, 1911; Ectobius pallidus, Germany (Foerster, 1938) Family Natural — Gregarina blaberae —Blaptica dubia hosts. (Frenzel, 1892) : : In intestine. ACTINOCEPHALIDAE Pileocephalus blaberae Synonymy. 1875) Foerster, 1938). (Frenzel) [Watson, 1916]. and related forms, Argentina In midgut. Unidentified Gregarinida Natural hosts. —Blaberus craniifer, U.S.A. (Roth and Willis, un- published data, 1953) Possibly Diplocystis sp. (pi. 28, A, B). Cryptocercus punctulatus, U.S.A. (Cleveland et al., 1934). : Leucophaea maderae, Philippine Islands (Hegner and Chu, 1930) U.S.A. (Roth and Willis, unpublished data, : In intestines of host. 1958) : Cysts in feces (pi. 28, C). Gromphadorhina portentosa, U.S.A., in laboratory colony (Roth In intestine of adult female. and Willis, unpublished data, 1958) 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 Family COCCIDIA ADELEIDAE Adelina cryptocerci Yarwood Natural 1937) • host. — Cryptocercus punctulatus, This intracellular parasite was found U.S.A. in the fat (Yarwood, body in light BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 185 In heavy infections the coccidia were found in the head, infestations. antennae, mouthparts, muscles, legs, salivary glands, nerve cord, as Infection in freshly collected specimens well as fat body. 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 (1934) mentioned a coccidium which was some- et al. times generally distributed through the body (head, legs, antennae, etc.) of C. punctiilatus ; this parasite was probably the species de- scribed by Yarwood. Order HAPLOSPORIDIA Haplosporidium periplanetae Georgevitch — Natural host. Blatta orientalis, Yugoslavia (Georgevitch, 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 Plisto- phora periplanetae. Coelosporidium periplanetae Synonymy. (Lutz and Splendore) —Noscma periplanetae, Coelosporidium hlattellae, Bcr- tramia hlatellae [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, — 191 1) ; U.S.A. (Kudo, 1922; Sprague, 1940) Yugoslavia (Ivanic, 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. MICROSPORIDIA Family NOSEMATIDAE Order Plistophora kudoi Sprague and Natural host. —Blatta orientalis, U.S.A., Ramsey West Virginia, Found in the epithe- Illinois, Kentucky (Sprague and Ramsey, 1941, 1942) : SMITHSONIAN MISCELLANEOUS COLLECTIONS l86 lial cells cells. of caeca and midgut. Considerable damage is VOL. I4I done to these Seventy-five percent of 52 B. orientaUs harbored the parasite. Plistophora periplanetae (Lutz and Splendore) Synonymy. —Nosema periplanetae, Pleistophora periplanetae [after Semans, 1943]. Georgevitch (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 dis- may crepancies in the earlier literature 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 orientaUs, France (Mercier, 1906a; Debaisieux, 1927) England (Perrin, 1906, 1906b) U.S.S.R. (ZhiYugoslavia (Georgevitch, 1925, 1926, 1926a, 1927) vago, 1909) Germany (Wellmer, 1910, 1911). — ; ; ; ; Blattella gcrmanica, France (Leger, 1909; Debaisieux, U.S.S.R. (Zhivago, 1909). Periplaneta americana, Brazil (Lutz and Splendore, 1903). This organism roaches. The lives in the 1927); lumen of the malpighian tubules of cock- cited authors appear to have been convinced that this organism was a microsporidian. Georgevitch (1927, 1953) described the polar capsule and filament characteristic of this order. Plistophora sp. — Natural host. Blatta orientaUs, 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 gertnanica and Periplaneta americana collected in South Africa. may It or may not have been a species of Plistophora. CILIATA HOLOTRICHA Class Order Family PARAMECIIDAE Paramecium Natural associate. 1927) : — Cockroaches, Three of 30 cockroaches sp. U.S.A., Maryland collected in the (Cleveland, basement of a de- COCKROACHES BIOTIC ASSOCIATIONS OF —ROTH & WILLIS partment store had paramecia in their stomachs but none rectum. Experimental associate. —Periplaneta americana, 187 in the U.S.A. (Hegner, Paramecia fed to the cockroaches were recovered from the 1929) 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. SPIROTRICHA Family BURSARIIDAE Order Balantidium blattarum Ghosh Natural host. —Periplaneta Bhatia and Gulati, 1927) ; americana, India (Ghosh, 1922; InGold Coast Colony (Macfie, 1922) : testinal tract. Balantidium Experimental vector. (Malmsten) coli — 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. litsch, 1938 ; —Blatta orientalis, Meglitsch, 1940) : U.S.A., Illinois This protozoan is (Kudo and Meg- found in the lumen of the anterior region of the colon in association with several other species of protozoa. Only y.6 percent of 500 cockroaches contained B. praenucleatum. single host was The largest examined number encountered in a 59, but as a rule each host harbored a smaller number. Balantidium — sp. ? Natural host. Periplaneta americana, Brazil (Magalhaes, 1900) These organisms were numerous in the intestine. : SMITHSONIAN MISCELLANEOUS COLLECTIONS l88 VOL. I4I SPIROSTOMIDAE Family Nyctotherus buissoni Pinto —"Barata sylvestre," Brazil (Pinto, 1926) Natural host. ism found in the cockroach's intestine. Nyctotherus Synonymy. Organ- : Leidy ovalis —Bursaria hlattarum; Plagiotoma hlattarum. —Blatta U.S.A. (Leidy, 1850, 1853, 1853b, Natural hosts. orientalis, Kudo and Meg1879a; Kudo, 1922, 1926, 1936; McAdow, 1931 litsch, 1938; Meghtsch, 1940); Germany (Stein, i860; Schubotz, ; 1905; Chen, 1933); England (Lankester, 1865; Schuster, 1898; U.S.S.R. (Yakimov Lucas, 1927a, 1928) Spain (Zulueta, 1916) and Miller, 1922 Zasukhin, 1928, 1930 Ostroumov, 1929) Portu; ; ; gal (Lima ; Ribiero, 1924) ; ; Brazil (Pinto, 1926) Venezuela (Te- ; jera, 1926). Blattella germanica. South Africa (Porter, 1930) ; U.S.A. (Balch, 1932; McAdow, 1931). Parcohlatta pensyhanica, U.S.A. (Semans, 1939, 1941). Indochina Periplaneta americana, India (Bhatiaand Gulati, 1927) ; (Weill, 1929) ; Philippine Islands (Hegner and Chu, 1930) ; South Hatcher, 1939; U.S.A. (McAdow, 193 1 Meglitsch, 1940; Armer, 1944)) Goa (Mello et al., 1934) China (Pai and Wang, 1947); Czechoslovakia (Lom, 1956). Africa (Porter, 1930); ; ; ; "Barata sylvestre," Brazil (Pinto, 1926). "Kiichenschaben," Austria (Belar, 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 (Kudo, 1936). Ninety (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 association with other species of Protozoa percent of 500 B. orientalis contained N. ovalis (Lucas, 1928; Balch, 1932; Chen, 1933; Lom, 1956). Nyctotherus uichancoi Kidder Natural hosts. —Panesthia angustipennis, Philippine Islands, and Panesthia spadica, Japan (Kidder, 1937) About 90 percent of angustipennis harbored this ciliate in their hindguts. : all P. BIOTIC ASSOCIATIONS OF COCKROACHES Nyctotherus viannai Natural host. — "Barata —ROTH & WILLIS 189 Pinto sylvestre," Brazil (Pinto, 1926) : In the in- testine of the cockroach. Family Most of CLEVELANDELLIDAE the Clevelandellidae are parasitized by rod-shaped or spherical bacteria-like organisms usually in clusters (Kidder, 1937). Synonymy. — Clevelandiidae Genus Synonymy. occupied 1938. ; it (Kidder, 1938). CLEVELANDELLA —The generic name Clevelandia Kidder was therefore changed to Clevelandella (1937) is preby Kidder in All of the following species of Clevelandella were originally described as Clevelandia. Clevelandella constricta Natural hosts. —Panesthia (Kidder) angustipennis, Philippine Islands, and Panesthia spadica, Japan (Kidder, 1937) In the posterior end of : hindgut. Clevelandella contorta Natural hosts. —Panesthia (Kidder) angustipennis, Philippine Islands, and Panesthia spadica, Japan (Kidder, 1937). Clevelandella elongata Natural host. (Kidder) —Panesthia angustipennis, Philippine Islands (Kid- der, 1937). Clevelandella hastula Natural host. der, 1937) : (Kidder) —Panesthia angustipennis, Common Philippine Islands (Kid- in hindgut, Clevelandella nipponensis (Kidder) Natural host. —Panesthia spadica, Japan (Kidder, 1937). Clevelandella panesthiae Natural hosts. —Panesthia (Kidder) angustipennis, Philippine Islands, and In the hindgut. This Panesthia spadica, Japan (Kidder, 1937) protozoan is commonly parasitized by the microorganism Sphaerita. : Clevelandella parapanesthiae (Kidder) Natural host. der, 1937). —Panesthia angustipennis, Philippine Islands (Kid- SMITHSONIAN MISCELLANEOUS COLLECTIONS igO Paraclevelandia Natural hosts. —Panesthia VOL. I4I Kidder brevis 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 Natural 1950) : host. —Pycnoscelus A large ciliate was ciliate surinamensis, found Hawaii (Schwabe, and malpighian in the digestive tract tubules. NEGATIVE FINDINGS In a recent experimental study Schmidtke (1955) failed to demonbetween Periplaneta americana and strate a host-parasite relationship the haemosporidian protozoan is Toxoplasma gondii Nicolle and Manceaux. This a blood parasite in a rodent in North Africa (Kudo, 1954). XI. HELMINTHS nematodes of the family Thelastomatidae have no apefifect on their cockroach hosts. Caleb (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, Intestinal parent pathological the oxyurids die and disappear. parasites, in the sense that we In other words, these worms are not use the term in this paper, but commen- 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-parasals. sitised specimens." Very few papers have dealt with the ecology of the oxyurid para- of cockroaches. According to Caleb (1878), usually one species of nematode is found in a single cockroach, but sometimes two species sites live together in the same host (e.g., in Blatta orientalis and Polyphaga aegyptiaca) where they compete for food. Caleb claimed that Ham- would replace Leidynema appendiculata; he surpassed L. appendiculata in numbers and that diesingi observed H. uncommon in the intestines of the cockroaches. On the latter became the other hand, Sobolev (1937) found that 48 percent of his oriental merschmidtiella diesingi BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS I9I 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 Caleb's conclusions inasmuch as the number of each species in mixed infections was essentially the same as the number found in cock- roaches 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 i 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 ex; tent of the infection. Thus the larvae of Moniliformis moniliformis may become embedded in the fat (Moore, 1946). First stage larvae of Oxyspirura mansoni also burrow through the midgut wall into the fat body; Sanders (1929) believed that Pycnoscehis siirinamensis could be killed if at one time a sufficient number of migrating larvae of O. mansoni penetrated the pass through the gut wall and some tissue cockroach's intestinal wall. Gongylonema neoplasticum migrates through the digestive tract and encysts in the muscles of the thorax and Ditlevsen, 1914). Infective larvae of Protospirnra 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 cockand legs of the host (Fibiger roaches infected with Physaloptera vara, P. maxillaris, P. hispida (Petri, 1950; Hobmaier, 1941 ; Schell, 1952), and Gongylonema SMITHSONIAN MISCELLANEOUS COLLECTIONS 192 pulchrum (Schell, 1952a) VOL. I4I in the latter species the deposit eventually ; completely surrounded the nematode larva which was killed and "chitinized." Apparently these pigmented cysts surround unhealthy mechanism by the host Oswald (1958) has reported finding similar pigBlatta orientaUs and Periplaneta americana that were or dead larvae and are secreted as a defensive (Schell, 1952a). mented cysts in experimentally infected with Rictiilaria coloradensis. Our classification of the helminths follows Hyman (1951, 1951a). HELMINTHS FOR WHICH COCKROACHES SERVE AS PRIMARY HOSTS ASCHELMINTHES Phylum NEMATODA Class Order MERMITHOIDEA MERMITHIDAE Family Undetermined mermithids Natural hosts. setts (Roth and —Ectobius : is the body extend into the thorax. Ap- This mermithid cavity of the host and one end parently, the host Plymouth, Massachu- pallidus, U.S.A., Willis, 1957) may eventually killed and the lies coiled in worms may cockroach ventrally between the thorax and abdomen leave the A) (pi. 29, or thorax and head. Periplaneta is Germany americana, "Mermis" or "Gordius." It (Bode, 1936) Attacked by has been suggested that the name Mermis : often applied without critical identification to immature found in insects Nematoda (Buxton, 1955). Order Family RHABDITOIDEA DIPLOGASTERIDAE Diplogaster sp. Synonymy. —Lycolaimus [Goodey, 1951]. — Experimental germanica, Blattella host. U.S.A. (Christie and Grossman, 1933). Family STEINERNEMATIDAE Neoaplectana — Nauphoeta cinerea, and This (Dutky and Hough, 1955) codling moth larvae, is close to Neoaplectana Experimental hosts. Blattella germanica, Periplaneta americana. U.S.A. nematode, found in sp. : COCKROACHES BIOTIC ASSOCIATIONS OF chresima Steiner but apparently was very were a is susceptible to infection ; new — ROTH & WILLIS I93 Nauphoeta cinerea species. B. germanica and P. americana less susceptible. Order Family OXYUROIDEA THELASTOMATIDAE These nematodes are found in the intestinal tract of cockroaches. Aorurus philippinensis Chitwood and Chitwood, 1934 Natural host. —Panesthia angustipennis, wood and Chitwood, Binema mirzaia Synonymy. Natural (Basir, —Periplaneticola host. Philippine Islands (Chit- 1934). —Periplaneta Basir, 1940) mirzaia Basir, americana, 1956 1940. Aligarh India, (Basir, 1940). Blattelicola blattelicola Natural host. — 1940 Blattella germanica, India, Aligarh (Basir, 1940). Blatticola Wattae (Graeffe, i860) Chitwood, 1932 Synonymy. — Oxyuris hlattae Graeffe, Caleb, 1878; Blatticola blatticola [Chitwood, 1930, 1932]. Natural hosts. Basir, — i860 (Caleb, ; 1877) Blattella germanica, Brazil Oxyuris blatticola Schwenck, 1926 (Pessoa and Correa, 1930; Bozeman, 1942); Egypt? (Caleb, 1877, 1878); U.S.S.R. (Sobolev, 1937; Sondak, 1935) Czechoslovakia (Groschaft, 1956). Ectobius lapponicus, Ectobius pallidus, Egypt? (Caleb, 1877, 1926; Schwenck, 1926); U.S.A. (Chitwood, ; 1878). Polyphaga aegyptiaca, France (Craeffe, i860). The life cycle has been studied by Bozeman (1942) never more than four Embryos developed worms effect on the He found The resting stage was The worms seemed to have to "resting" stage in vitro. infective while the active stage no : in the large intestine of each cockroach. was not. vital activities of the host. Alicata (1934b) found that 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 the in a single individual, apparently only in the rectum. SMITHSONIAN MISCELLANEOUS COLLECTIONS 194 in VOL. I4I Sobolev (1937) found 72 percent of Blattella germanica collected Gorkov (U.S.S.R.) infected with Blatticola hlattae. The mean 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. hlattae. 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. number 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 Natural host. it as a valid species in describing the variety javanica. —Panesthia laevicollis [Cobb recorded the host as Van Zwaluwen- Panesthia hrevicollis, but no such cockroach exists. burg (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. Natural host. Chitwood and Qiitwood, javanica —Panesthia angustipennis, 1934 Philippine Islands (Chit- wood 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, 185 1 [Christie, Thelastoma indiana Basir, 1940 [Basir, 1949]. Natural host. Leucophaca sp., India, Aligarh (Basir, 1940, 1949). 1933]- Cephalobellus magalhaesi Synonymy. —Bulhoesi — (Schwenck, 1926) Basir, 1956 magalhaesi Schwenck, 1926; Thelastoma magalhaesi (Schwenck, 1926) Travassos, 1929 [Basir, 1956]. Natural host. "Barata selvagem," Brazil, Sao Paulo (Schwenck, 1926). Euryconema paradisa Chitwood, 1932 Natural host. 1932). —Eurycotis floridana, U.S.A., Florida (Chitwood, BIOTIC ASSOCIATIONS OF COCKROACHES Galebia aegyptiaca (Caleb, 1878) — ROTH & WILLIS I95 Chitwood, 1932 Synonymy. — Oxyuris aegyptiaca Galeb, 1878 Blatticola aegyptiaca (Caleb, 1878) Schvvenck, 1926 [Chitwood, 1932]. — germanica, Brazil (Schwenck, 1926). Natural ; Blattella hosts. Polyphaga aegyptiaca, Egypt? (Caleb, 1878). Hammerschmidtiella diesingi (Hammerschmidt, 1838) Chitwood, 1932 Synonyrny. —Anguillula macriira Diesing, 185 1 ; Aoruriis 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, England BiJtschli, (Caleb, Egypt? 1847; 1878); 1871) (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 (Cro- schaft, 1956). Leucophaea maderae, Brazil (Pessoa and Correa, 1926). Periplaneta americana, Brazil (Magalhaes, 1900; Pessoa and Cor- 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 (Pessoa and Correa, 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 rea, 1926). : ; ; ; ; throughout the intestinal canal but especially in the small is intestine. It frequently found in adults and seldom in the nymphs. There were 5 to 10 worms in one cockroach and female worms 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. Biitschli (1871) stated that all stages from those just hatching to mature males and females are found. seldom more than w^ere ; 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 (1956) found 18 in of 22 in one cockroach. one specimen of B. orientalis. Groschaft Dobrovolny and SMITHSONIAN MISCELLANEOUS COLLECTIONS 196 VOL. I4I 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 place outside the host resulting in After the tgg in the infective stage molt occurs before the egg hatches. a is resting or ; the first infective takes stage. eaten by the host, the second 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 stages of cleavage. nematode eggs are in the very earliest In 36 hours a motile, tadpole-like stage is reached 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). and in a The bacterium Streptoniyces Jeidynematis Hofifman grows on the H. diesingi (Hoffman, 1953). The bacterium apparently is cuticle of only anchored to the nematode and probably obtains intestinal contents of the cockroach. its food from the See notes under Leidynema ap- pendiculata. Hammerschmidtiella neyrai Serrano Sanchez, 1945 Synonymy. —Hammerschmidtiella neyrae Serrano Sanchez, 1947. [According to M. B, Chitwood, personal communication, 1957, Serrano Sanchez's emendation Natural host. —Blatta is apparently an error.] orientalis, Spain, Grenada (Serrano Sanchez, Of 2,943 specimens examined, 1,143 were parasitized by 1947) oxyurids and of these 45 percent contained H. neyrai. : Leidynema appendiculata (Leidy, 1850) Chitwood, 1932 Synonymy. — Oxyuris hlattae orientalis Hammerschmidt, 1847, of and Oxyuris blattae-orientalis of Magalhaes, 1900) Oxyuris hlattae Hammerschmidt, 1847, of Galeb, 1878 Aorurus Biitschli, 1871, ; ; (Thelastoma) appendiculatus Leidy, 1850. [Chitwood, 1932.] Ser- rano Sanchez (1947) has divided this species into three geographical varieties as follows: L. appendiculata (Leidy, 1852) (Dobrovolny BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS I97 var. Indiana; L. appendiculata (Leidy, 1852) (Chitwood, 1932) var. americana; L. appendiculata (Serrano Sanchez, 1947) var. hispana. However, Basir (1956) does not recognize and Ackert, 1934) these varieties. The Russians recognize hispana (M. B. Chitwood, personal communication, 1957). Natural hosts. —Blahcnis atropos, South America 1932). Blatta orientalis, Egypt? (Caleb, 1878) U.S.S.R. (Sobolev, 1937; Sondak, 1935) Spain (Serrano Sanchez, 1947) 1944) ; ; ; : Europe (Chitwood, (Biitschli, 1871) U.S.A., Nebraska (Todd, ; Recorded as var. hispana. Czechoslovakia (Groschaft, 1956). England (Lee, 1958a). Blatta orientalis or Periplaneta americana, Brazil (Magalhaes, 1900). Texas (Todd, 1943); Nebraska Kansas (Dobrovolny, 1933; Dobrovolny and Ackert, 1934) North Carolina (Hatcher, 1939) Iowa, North Dakota, Michigan (Hoffman, 1953). Czechoslovakia (Groschaft, 1956). EngPeriplaneta americana, U.S.A.: (Todd, 1944) ; ; ; land (Lee, 1958a). Cockroach, Venezuela (Tejera, 1926). Cockroaches (Blatta orientalis, Blattella germanica, and/or Peri- 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. Caleb planeta americana) , (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 The molts occur within the tgg during development of the larva. first molt occurs outside the host resulting infective resting stage. in the The second molt occurs formation of an 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 Leidy (1853) noted the presence of simple, inarticugrowing from nematodes infecting B. oriBiitschli (1871) and Magalhaes (1900) described similar entalis. filaments adhering to the surface of oxyurids from cockroaches. in P. americana. late, amorphous filaments, SMITHSONIAN MISCELLANEOUS COLLECTIONS 198 VOL. I4I Leidynema appendiculata (Leidy, 1850) Chitwood, 1932? —Eurycotis (Roth 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 Natural host. and floridana, U.S.A., Massachusetts Willis, unpublished data, 1955) : literature." Leidynema cranifera Chitwood, 1932 Natural hosts. —Blaherus craniifer, U.S.A., Florida (Chitwood, Massachusetts (Roth and Willis, unpublished data, 1955). 1932) Determined by Dr. G. Steiner. ; Blaherus atroposf, U.S.A., Florida (Chitvv^ood, 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 Natural host. delatorrei Chitwood, 1932 —Leucophaea maderae, Cuba, Havana (Chitwood, 1932). Leidynema nocalum Chitwood and Chitwood, 1934 Natural host. —Panesthia angustipennis, Philippine Islands (Chit- wood and Chitwood, 1934). Leidynemella fusiformis Cobb, 1934 Natural hosts. —Panesthia laevicollisf, Philippine Islands (Cobb in Chitwood and Chitwood, 1934). Panesthia angustipennis^ Philippine Islands (Chitwood and Chit- wood, 1934). Leidynemella panesthiae (Caleb, 1878) Chitwood and Chitwood, 1934 Synonymy. — O.vyuris panesthiae Galeb, 1878, panesthiae (Galeb, 1878) Travassos, 1929. wood and Chitwood, Natural host. in part; Thelastoma [Chitwood, 1932; Chit- 1934.] —Panesthia 40 nematodes may be found sp.. in New one Guinea (Galeb, 1878) : About insect. Leidynemella paracranifera Chitwood and Chitwood, 1934 Natural host. —Panesthia wood and Chitwood, 1934). angustipennis, Philippine Islands (Chit- BIOTIC ASSOCIATIONS OF COCKROACHES Oxyuris Synonymy. 1929. — Thelastoma The taxonomic heterogamiae (?) —ROTH Caleb, & WILLIS I99 1878 heterogamiae (Caleb, 1878) Travassos, position of this species is questionable; it might possibly belong in Blattkola or Protrellina (Chitwood, 1932). Basir (1956) placed it in an appendix to the family Thelastomatidae. Natural host. —Polyphaga acgyptiaca, Egypt? Protrelleta Chitwood, floridana Natural host.—Blaberiis craniifcr, (Caleb, 1878). 1932 (Chitwood, U.S.A., Florida 1932). Protrellus aureus Cobb, 1920 —The taxonomic position of able (Chitwood, 1932). —Polyzosteria melanariaf, Natural nematode this Synonymy. Protrellus aurifluus South {in Chitwood, 1932) stated that this analis.] (Chitwood, 1932) Chitwood, Snyonymy. —Protrellina aurifluus Chitwood, Natural hosts.—Parcoblatta question- New Australia, host. Wales (Cobb, 1920). [Caudell host was probably Platyzosteria is lata, I933 1932. U.S.A., North Carolina, Maryland (Chitwood, 1932). Parcoblatta uhleriana, North Carolina (Hatcher, 1939). Protrellus australasiae (Pessoa and Coriea, 1926) Travassos, 1929 Synonymy. — Oxyuris australasiae trellina Pessoa and Correa, 1926; Proaustralasiae (Pessoa and Correa, 1926) Chitwood, 1932 [Chit- wood, 1933]. Natural host. —Periplaneta n (Pessoa and Correa, ^ australasiae, Brazil - , 1926, 1927). Protrellus galebi Schwenck, 1926 Synonymy. —Protrellina galebi (Schwenck, 1926) Chitwood, [Chitwood, 1933]. —"Barata selvagem," Brazil (Schwenck, 1926). Natural 1932 host. Protrellus kiinckeli (Caleb, 1878) Travassos, 1929 Caleb, 1878; Protrellina Synonymy. — Oxyuris (Caleb, 1878) Chitwood, 1932 [Chitwood, 1933]. —Periplaneta americana, Egypt? (Caleb, Natural kiinckeli hosts. 1878. [Chitwood (1932) questioned the determination of kiinckeli 1877, this host SMITHSONIAN MISCELLANEOUS COLLECTIONS 200 VOL. I4I because he failed to find this nematode in a large number of specimens from U.S.A. and China.] Brazil (Pessoa and Correa, 1926). Periplaneta australasiae, Brazil (Pessoa and Correa, 1926). Protrellus manni (Chit wood, 1932) Chitwood, 1933 Synonymy. —Protrellma manni Chitwood, 1932. —Aglaopteryx diaphana, Cuba (Chitwood, Natural host. Protrellus phyllodromi (Basir, 1942) Synonymy. —Protrellina phyllodromi — Natural humhertiana, Basir, 1932). 1956 Basir, 1942. Blattella host. 1942) : Found Protrelloides Natural Aligarh India, (Basir, in the rectum. host. paradoxa Chitwood, 1932 —Enrycotis floridana, (Chitwood, U.S.A., Florida 1932). Schwenkiella icemi (Schwenck, 1926) Basir, 1956 — Synonymy. Bulhoesia icemi Schwenck, 1926; Thelastoma icemi (Schwenck, 1926) Travassos, 1929; Thelastoma aligarhica Basir, 1940. [Basir, 1956.] Natural hosts. —"Barata selvagem," Brazil, Sao Paulo (Schwenck, 1926). Periplaneta americana, India, Aligarh (Basir, 1940) ; U.S.A., Nebraska (Todd, 1943). Periplaneta hrunnea, U.S.A., Louisiana (Todd, 1943). Severianoia Natural host. —"Blattidae magna Pereira, sylvestres," 1935 Brazil (Pereira, 1935). Severianoia severianoi (Schwenck, 1926) Travassos, 1929 Synonymy. 1929]. —Bulhoesia severianoi Schwenck, —"Baratas de pau podre," Brazil Natural hosts. 1926 [Travassos, (Schwenck, 1926). Pycnoscelus surinamensis, U.S.A., Florida (Chitwood, 1932). Suifunema Natural host. caudelli — Steleopygaf Chitwood 1932 sinensis, Asia : Suif u, Szchuen, China (Chitwood, 1932). Thelastoma pachyjuli (Parona, 1896) Travassos, 1929 Synonymy. — Oxyuris hulhoesi de Magalhaes, hoesi (Magalhaes, 1900) 1900 Bulhoesia hulSchwenck, 1926 [Travassos, 1929; Chit; BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 201 Thelastoma hulhoesi (Magalhaes, 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 wood, 1932] ; under T. pachyjuli. Natural hosts. —Blatta orientalis, Czechoslovakia (Groschaft, 1956). Periplaneta americana, Brazil (Magalhaes, 1900) North America (Chitwood, 1932); U.S.A., North Carolina (Hatcher, 1939). ; Thelastoma palmettum Chitwood and Chitwood, 1934 —Panesthia angustipennis, Natural host. Philippine Islands (Chit- wood and Chitwood, 1934). Thelastoma Natural host. —Periplaneta riveroi Chitwood, 1932 sp., Cuba (Chitwood, 1932). Undetermined nematodes Natural host. from New — Cutilia sp. Class sedilloti, U.S.A. (hosts imported NEMATOMORPHA Order The immature found near Zealand) (Roth, unpublished data, 1957). in the GORDIOIDEA stages of the following gordian worms have been body cavity of cockroaches. Family CHORDODIDAE Chordodes morgan! Montgomery, Synonymy. — Chordotes [sic] puerilis 1898 Montgomery, 1898 [Ward, 1918]. Natural host. —Cockroach, U.S.A. (Montgomery, 1898) PennsylNebraska (Ward, ; vania, Maryland, Michigan, Ohio, Florida, Iowa, 1918). Family Gordius — GORDIIDAE aquaticus Linnaeus, 1758 Blatta sp., U.S.A. (Stiles and Hassall, 1894). host. Leidy (1879) identified a 9-inch-long nematode which came from a cockroach {Blatta orientalis?) as probably being Gordius aquaticus. Natural 202 Ransom SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I may be an acci- (in Pierce, 1921) states that G. aquaticus Faust (1955) summarizes the few reported cases of human parasitism. Dorier (1930) reported that the redental parasite of man. gurgitated liquid of Blatta orientalis had no effect on cysts of G. aquaticus after one hour. Gordius blattae Synonynny. — Gordius —Blatta Natural host. 1878) : Found in 1851 Germsiny (Siebold, 1842; Linstow, Von abdomen. Siebold called this "Filarien" but describe the Gordius pilosus Synonymy. Diesing, orientalis of Lankester (1865). orientalis, name or did not otherwise orientalis (Mobius, worm. 1855) Diesing, 1861 — Chordodes pilosus Mobius, 1855 [Diesing, 1861]. —Blaberus giganteus, Venezuela (Mobius, 1855) Natural host. From : the insect's abdomen. Gordius Natural hosts. —Periplaneta Germany (Bode, 1936) 1930) a Mermis or other nematode. ; Cockroaches, Venezuela sp. South Africa (Porter, may have referred to americana, : Bode's record (Miall and Denny, 1886; Burr, 1899a; Tejera, 1926). Parachordodes raphaelis Synonymy. (Camerano, 1893) Camerano, 1897 — Gordius raphaelis 1893 [Camerano, 1897]. —Symploce parenthesis and Kuchinga hemerohina, C^LVCitrdino, Natural hosts. French Equatorial Africa (Camerano, 1893, 1897). Undetermined gordian worms Natural hosts. —Eurycotis floridana, communication, 1958) : Florida (T. Eisner, personal See plate 29, B. Parahormetica hilohata, Brazil (Pessoa and Correa, 1929) : Worm referred to as "gordiaceo." Cockroaches, 1953) : Australia (E. F. Riek, personal communication, Three undescribed gordian worms were found mined cockroaches of the subfamily Blattinae. in undeter- BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 203 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 Natural host. 1938, 1938a; — (Diesing, Blattella germanica, Brumpt Experimental elegans hosts. 1851) Travassos, 1915 France (Brumpt and Urbain, et al., — 1939). Blaherus atropos and Leucophaea maderae, France (Brumpt and Desportes, 1938). in Rudolphi, 1819) Travassos, 1917 Prosthenorchis spirula (Olfers Natural host. — France (Brumpt and Urbain, Netherlands (Thiel and Wiegand Blattella germanica, 1938, 1938a; Brumpt Bruss, 1946). Experimental hosts. et al, — 1939) ; Blattella germanica, Netherlands (Thiel and Wiegand Bruss, 1946). Blaherus atropos and Leucophaea maderae, France (Brumpt and Desportes, 1938). Family MONILIFORMIDAE * Moniliformis dubius Meyer, 1932 Natural hosts. —Periplaneta americana, Brazil (Magalhaes, 1898; India (Pujatti, Gold Coast (Southwell, 1922) Travassos, 1917) Moore, and Chandler, (Burlingame U.S.A. 1941 1946). 1950) ; ; ; ; Periplaneta australasiae , India (Pujatti, 1950). Experimental hosts. Blattella germanica, Japan — Miyata, 1942). Periplaneta americana, U.S.A. (Chandler, 1941 ; (Yamaguti and Moore, 1946) ; Japan (Yamaguti and Miyata, 1942). Moniliformis kalahariensis Meyer, 1931 Natural host. * — Blattella germanica, India The following helminths also have been stated in the literature to pass their intermediate stages in cockroaches: [Blanchard 1870) [Baer (1891)]; (1956)]; (Meyer, 1931, 1932). Inennicapsifer Spirocerca Hymenolepis diminuta (Rudolphi, 1819) madagascariensis (Davaine in Grenet, sangninolcnta (Rudolphi, 1819) [Seurat (1913)]. These doubtful records are discussed in Roth and Willis (i957a). SMITHSONIAN MISCELLANEOUS COLLECTIONS 204 * Moniliformis moniliformis Natural hosts. 1927, VOL. I4I (Bremser in Rudolphi, 1819) Travassos, 1915 — Periplaneta americana, Argentina (Bacigalupo, 1928); Brazil (Pessoa and Correa, 1929); Algeria Burma (Subramanian, 1927) South Africa 1912) 1927a, (Seurat, ; (Porter, 1930) ; ; Madras (Sita, 1949). New Caledonia (Rageau, 1956). Cockroaches, Venezuela (Tejera, 1^26). Experimental hosts. Blaberus atropos, Blatta orientalis, Blattella Periplaneta spp.. — Leucophaea maderae, germanica, (Brumpt and Urbain, France 1938a). Periplaneta americana, France (Brumpt, 1949) Phylum ; (Yamaguti and Miyata, Japan Madras 1942) ; (Sita, 1949). ASCHELMINTHES Class NEMATODA Order OXYUROIDEA Family SUBULURIDAE * Subulura jacchi (Diesing, 1861) Railliet and Henry, 1914 —Suhidura jacchi (Marcel, 1857) [Dr. T. Lucker, personal communication, 1957]. —Blaberus atropos, France (Chabaud and Experimental Synonymy. J. host. Lariviere, 1955). SPIRUROIDEA Order Family * Oxyspirura Natural hosts. 1926, Shealy, 1927) ; mansoni (Cobbold, 1879) Ransom, 1904 —Pycnoscelus 1928, 1927, THELAZIIDAE 1928a); Australia surinamensis, U.S.A. (Sanders, 1927, (Fielding, 1928, 1929; Antigua (Hutson, Formosa (Kobayashi, 1927) Schwabe, 1950, 1950a, Hawaii (Illingworth, 1931 ; 1938, 1943) 1950b, New Caledonia (Rageau, 1956). 951) have recently found (Roth and Willis, i960) that two strains We 1 ; ; ; of Pycnoscelus surinamensis exist; a parthenogenetic strain (from Florida), and a bisexual strain (from Hawaii) which does not re- produce parthenogenetically. The parthenogenetic strain shown is undoubt- 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 edly the form that has been to be the host of O. BIOTIC ASSOCIATIONS OF COCKROACHES papers, we concluded & WILLIS 205 Schwabe, in Hawaii, may well have been and possibly also with the parthenotrue, then both parthenogenetic and bisexual that working with the bisexual genetic strain; —ROTH if this is strain strains of Pycnoscelus surinamensis of the eyeworm. Experimental hosts. —Periplaneta may serve as intermediate hosts americana, Antigua (Hutson, 1943)- U.S.A. (Sanders, 1929) Hawaii (Schwabe, 1951). Pycnoscelus surinamensis, (Fielding, 1927, 1928a) ; Rictularia coloradensis Hall, Australia ; 1916 — Natural hosts. Parcoblatta pensylvanica and Parcoblatfa virginica, U.S.A., Ohio (Oswald, 1958) Of 49 M^ood roaches collected, one of : each species contained a single larva each. Experimental hosts. —Blatta orientalis, Blattella germanica, Parco- blatta pensylvanica, Parcoblatta virginica, Periplaneta americana, and The larvae underSupella supellectilium, U.S.A. (Oswald, 1958) went normal development in all species of cockroaches except B. : orientalis and P. americana a reddish-brown pigment ; in which cysts developed that contained 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. larvae underwent two molts within a cyst formed by The 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 cysts remained attached to the hindgut. 6*. supellectilium the Usually over 20 cysts de- veloped in each infected Parcoblatta; fewer than 10 per insect devel- oped 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 Pessoa and Correa, Natural host. —Parahormetica bilobata, Brazil 1929 (Pessoa and Correa, 1929). * Gongylonema ingluvicola Ransom, 1904 Experimental host. — Blattella germanica, U.S.A. (Cram, 1935). ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 206 * Gongylonema neoplasticum (Fibiger and VOL. I4I Ransom and Ditlevsen, 1914) Hall, 1916 Natural hosts. —Blatta orientalis, Netherlands (Baylis, 1925). U.S.A. (Hitchcock and Bell, 1952). Periplaneta americana, Denmark and St. Croix (Fibiger, 1913, Netherlands (Baylis, 1925) 1913a; Fibiger and Ditlevsen, 1914) England (Leiper, 1926) South Argentina (Bacigalupo, 1930) ForAfrica (Porter, 1930) U.S.A. (Hitchcock and Bell, 1952) mosa (Yokagawa, 1924, 1925, 1925a). Periplaneta australasiae, Formosa (Yokagawa, 1924, 1925, 1925a). Experimental hosts. Blattella germanica, Denmark (Fibiger and France Ditlevsen, 1914) U.S.A. (Hitchcock and Bell, 1952) Blattella germanica, ; ; ; ; ; ; — ; ; (Brumpt, 1949). Denmark (Fibiger and Ditlevsen, 1914). Denmark and St. Croix (Fibiger, 1913 Blatta orientalis, Periplaneta americana, Fibiger and Ditlevsen, 1914) * ; U.S.A. (Hitchcock and Bell, 1952). Gongylonema pulchrum MoHn, 1857 Experimental hosts. — Blattella germanica, U.S.A. (Ransom and 1917; Stiles and Baker, 1927; Schwartz and Europe (Baylis Lucker, 1931 Lucker, 1932; Alicata, 1934a, 1935) et al., 1925, 1926, 1926a; Sambon, 1926). Parcoblatta sp., Alicata (1934, 1935). Hall, 1915, 1916, ; ; * Natural host. Gongylonema sp. —Periplaneta americana, Algeria (Seurat, 1916) ; * Microtetrameres helix Experimental host. — ; Cram, 1927 Blattella germanica, * Protospirura bonnei — Brazil (Magalhaes, 1900) England? (Leiper, 1926). U.S.A. (Cram, 1934). Ortlepp, 1924 Natural host. Leucophaca maderae, Venezuela (Brumpt, 1931). Experimental hosts. Blatta orientalis, Blattella germanica, Leucophaca maderae, France (Brumpt, 1931). — * Protospirura columbiana Cram, 1926 Experimental host. — Blattella germanica, U.S.A. (Cram, 1926). * Protospirura muricola Geodoelst, 1916 Natural host. 19.-^8, 1939). —Leucophaca maderae, Panama (Foster and Johnson, BIOTIC ASSOCIATIONS OF COCKROACHES *Seurocyrnea Experimental host. — colini —ROTH & WILLIS 20"] (Cram, 1927) Cram, 1931 Blattella germanica, U.S.A. (Cram, 1931, 1931a, 1933a). Spirura Natural hosts. in Seurat, 1911) gastrophila (Miiller, 1894) Seurat, 1913 —Blatta orientalis, Europe? (Deslongchamps, 1824, Italy (Grassi, 1888); Algeria (Seurat, 1916). ; Periplaneta americana, Brazil (Pessoa and Correa, 1929). Cockroach, Venezuela (Tejera, 1926). — Experimental hosts. Blatta orientalis, France "Cafards," Algeria (Roger, 1906, 1907). * Natural host. — (Caleb, 1878a) ; Tetrameres americana Cram, 1927 Blattella germanica, U.S.A. (Cram, 1931b, per- Hawaii (Alicata, 1938, 1947)sonal communication, 1956) Experimental host. Blattella germanica, U.S.A. (Cram, 1931b). ; — Tetrameres Experimental host. — pattersoni Cram, PHYSALOPTERIDAE * Physaloptera hispida Experimental host. U.S.A. (Cram, 1933). Blattella germanica, Family — 1933 Schell, 1950 U.S.A. Blattella germanica, (Schell, 1952, 1952a). Physaloptera maxillaris Molin, i860 Experimental host. — Blattella germanica, U.S.A. (Hobmaier, 1941). * Physaloptera praeputialis Experimental host. — von Linstow, 1889 Blattella germanica, U.S.A. (Petri and Ameel, 1950). * Physaloptera rata Experimental 1950 ; host. — Hall and Wigdor, 1918 Blattella germanica, U.S.A. (Petri and Ameel, Petri, 1950) Physaloptera turgida Rudolphi, 1819 Experimental host. Schell, 1952). — Blattella germanica, U.S.A. (Alicata, 1937; SMITHSONIAN MISCELLANEOUS COLLECTIONS 208 VOL. I4I HELMINTHS WHOSE EGGS HAVE BEEN CARRIED BY COCKROACHES The use of the asterisk (*) Phylum is 3, page 4. PLATYHELMINTHES Class TREMATODA Order Family * explained in footnote DIGENEA SCHISTOSOMATIDAE Schistosoma haematobium (Bilharz, 1852) Weinland, 1858 Experimental vector. —Periplaneta americana. Gold Coast Colony (Macfie, 1922). Class CESTODA TAENIOIDEA HYMENOLEPIDIDAE Order Family * Hymenolepis sp. Natural vectors. —Periplaneta americana, Formosa (Morischita and Tsuchimochi, 1926). Polyphaga saussurei, Tadzhikistan (Zmeev, 1936). Family Taenia Experimental vector. TAENIIDAE saginata Goeze, 1782 —Periplaneta americana, Gold Coast Colony (Macfie, 1922). Echinococcus granulosis (Batsch, 1786) Rudolphi, 1805 Synonymy. — Taenia echinococcus (Zeder, 1803) [Faust, 1939]. —Periplaneta americana, Uruguay (Perez Experimental vector. Eggs were recovered from the feces of Fontana, 1955) infested cockroaches under "natural" conditions. : artificially Family Unknown Undetermined tapeworm ova Natural vector. —Polyphaga saussurei, Tadzhikistan (Zmeev, 1936). BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH Class NEMATODA Order OXYUROIDEA Family OXYURIDAE *Enterobius vermicularis (Linnaeus, 1758) Leach vectors. 2O9 ASCHELMINTHES Phylum Natural & WILLIS —Blatta and orientalis in Baird, 1853 germanica, Blattella U.S.S.R. (Sondak, 1935). Order ASCAROIDEA Family ASCARIDAE * Ascaris lumbricoides Linnaeus, 1758 Natural vector. —Periplaneta americana, South Africa (Porter, those of A. suum Goeze, 1782. —Periplaneta americana, Gold Coast Colony The eggs may have been 1930) Experimental vectors. : (Macfie, 1922) Periplaneta rhomhifolia, India (Chandler, 1926). ; americana, Periplaneta australasiae , Neostylopyga Formosa (Morischita and Tsuchimochi, 1926). * Ascaris sp. Natural vector. Besta, 1936). —Blatta —Periplaneta orientalis, Italian Experimental vector. Fontana, 1955) Order Family (Perez insects' feces. STRONGYLOIDEA ANCYLOSTOMIDAE *Ancylostoma caninum (Ercolani, 1859) Experimental Uruguay americana, Eggs recovered from the : Somaliland (Mariani and vector. —Periplaneta Hall, 1913 americana, Netherlands (Ak- kerman, 1933). *Ancylostoma ceylanicum Experimental vector. (Macfie, 1922); * 1911) Leiper, —Periplaneta americana. 1915 Gold Coast Colony Netherlands (Akkerman, 1933). Ancylostoma duodenale Natural vector. 1929, 1930). (Looss, —Periplaneta (Dubini, 1843) americana. Creplin, 1845 South Africa (Porter, SMITHSONIAN MISCELLANEOUS COLLECTIONS 210 Experimental vector. —Periplaneta americana, VOL. I4I Gold Coast Colony (Macfie, 1922). *Necator americanus Natural vector. 1902) (Stiles, Stiles 1906 —Periplaneta americana, India (Chandler, 1926). —Periplaneta americana, Gold Coast Colony Experimental vector. (Macfie, 1922). * Hookworm ova Experimental vectors. —Periplaneta americana, Periplaneta australasiae, Neostylopyga rhombifolia, Formosa (Morischita and Tsuchi- mochi, 1926). TRICHOSTRONGYLIDAE Family * Trichostrongylus sp. Natural vector. —Blatta orientalis, Italian (Mariani Somaliland and Besta, 1936). Order TRICHUROIDEA TRICHURIDAE Family * Capillaria hepatica (Bancroft, 1893) Travassos, 1915 Experimental vector. —Blatta * Trichuris trichiura Natural vectors. and Besta, 1936) —Blatta orientalis, Italy? (Linnaeus, orientalis, 1771) Italian (Giordano, 1950). Stiles, 1901 (Mariani Somaliland 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 (Perez Fontana, 1955)Periplaneta americana, Periplaneta australasiae, and Neostylopyga rhombifolia, Formosa (Morischita and Tsuchimochi, 1926). XII. The classification follows exceptions. The Acarina ARTHROPODA Brues et al. (1954) with the following are arranged according to Dr. J. H. Camin (personal communication, 1955). Family Eupelmidae of the optera follows the classification of Peck (1951). Hymen- BIOTIC ASSOCIATIONS OF Class In this mites. —ROTH & WILLIS of at least four orders have utilized the whip scorpions, scorpions, spiders, : 211 ARACHNIDA class, representatives cockroaches as food COCKROACHES and Apparently none of these feed exclusively on cockroaches, but the Philippine forest scorpion Heterometrus {=Palamnaeus) longe- manits seems to prefer blattids to other insects (Schultze, 1927). PEDIPALPIDA THELYPHONIDAE Order Family Mastigoproctus giganteus (Lucas) — Thelyphonus giganteus Lucas Synonymy. [Dr. R. E. Crabill, per- sonal 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 Common name. —Whip — Experimental prey. B. J. sp. scorpion. Blattella germanica, U.S.A., Florida [Dr. 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 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 pectines. 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. Qoudsley-Thompson (1955a) cited cockroaches as one of the arthropods that scorpions feed upon. Family Buthus Synonymy. tion, 1957], —Androctonus BUTHIDAE australis (Linnaeus) australis [Crabill, personal communica- SMITHSONIAN MISCELLANEOUS COLLECTIONS 212 VOL. I4I — Experimental prey. Cockroaches, England (Cloudsley-Thompson, This African species ate at least one cockroach per week during the summer months. It can, however, survive four months' 1955a) : starvation and is particularly adapted to a dry climate (CloudsleyThompson, personal communication, 1956). Centruroides gracilis Experimental prey. —Periplaneta published data, 1953) identified by Dr. M. PL (Latreille) americana, U.S.A. Muma. Centruroides hentzi Experimental prey. (Roth, un- Scorpion collected in Florida by Roth and : —Periplaneta (Banks) australasiae and Pycnosceliis (Muma, surinamensis, U.S.A. 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 Natural prey. —Parcohlatta (Say) pensylvanica Florida U.S.A., (?), personal communication, 1953). This may have been another species of this genus, possibly P. divisa, as P. pensylvanica is not (Muma, known from Florida (Rehn, personal communication, 1958). Experimental prey. —Blatta orientalis, Blattella germanica, Peri- planeta americana, and Pycnoscelus surinamensis, U.S.A., Florida (Muma, personal communication, 1953). Parabuthus capensis (Hemprich and Ehrenberg) Experimental prey. —"Common house-cockroach" The scorpions were collected in Cape Family (Pocock, 1893) : Town, South Africa. CHACTIDAE Euscorpius germanus (Koch) Synonymy. — Euscorpius 1955a]. Experimental prey. — carpathicus Blattella germanica, [Cloudsley-Thompson, England? (Pocock, 1893)Periplaneta americana, nymphs, England? (Cloudsley-Thompson, personal communication, 1956) : This scorpion southern Europe and North Africa. is found naturally in BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 2I3 Euscorpius italicus italicus (Herbst) — Experimental prey. Cockroaches including nymphs of Pcriplaneta, 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. 1949) : This record —Periplaneta U.S.A. americana, (Stahnke, a photograph showing the scorpion eating the is cockroach. Family ISCHNURIDAE Hormurus caudicula (Koch) Experimental prey. This record is — Cockroach, (McKeown, 1952) Australia : a photograph showing the scorpion feeding on a cock- roach. Family SCORPIONIDAE Heterometrus longimanus (Herbst) —Palamnaeus prey. — "Large Synonymy. longimanus [Cloudsley-Thompson, per- sonal communication, 1956]. Natural (Schultze, 1927) : On wood wings and legs of the large wood cockroach a rotten log. Philippine cockroach," Islands found fragments of a scorpion cavity, under several occasions Schultze in — 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 novaehoUandiae Peters — Experimental prey. Periplaneta americana, Australia (Glauert, An injured cockroach was accepted at once by the scorpion, 1946) : which held the moving insect in its claws chelicerae. The sclerotized exoskeleton. scorpion ate and tore all it with the alternately the soft parts and most of the ? SMITHSONIAN MISCELLANEOUS COLLECTIONS 214 ARANEIDA Order Many general, VOL. I4I observations of spiders feeding on cockroaches are quite and many observers have failed to identify either the spider (1874) stated that "the cockroaches that infest houses in the tropics .... have numerous enemies birds, rats, or prey. its Belt — 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 nymphs. Sonan (1924) reported that large gray spiders devour nymphs and adults of Periplaneta americana and P. austrabirth to 51 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. Ran (1940) stated that American and oriental cockroaches were the lasiae in ; 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 Ecfobiiis are by Xysticus, Cluhiona, Drassodes, Zelotes, Tarantula, readily accepted 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 Common and Avicularia — Bird-eating —Periplaneta americana, name. Natural prey. (Linnaeus) sp. spider. Trinidad (Main, 1924, The remains of the host were compressed into globular form 1930) by the spider after it had extracted the nutritive parts. : Phormictopus cancerides (Latreille) Experimental prey. — Cockroach, West Indies (Wolcott, 1953). BIOTIC ASSOCIATIONS OF Family COCKROACHES —ROTH & WILLIS 21$ SPARASSIDAE (Linnaeus) Heteropoda venatoria —Heteropoda regia Fabricius. — Banana spider (Conistock, spider (Gertsch, 1949) big brown house spider Natural and experimental prey. — Cockroaches, Synonymy. Common riames. ; 191 2) ; huntsman (Bryan, 1915). Bermuda (Verrill, Puerto Rico (Sein, 1923; Wolcott, 1924a; Petrunkevitch, Hawaii (Bryan, 1915. Williams et al., 1931) British Guiana (Moore in Williams et al., 1931) Panama (Gertsch, 1949) 1902); 1930a) ; ; ; New Zealand (adventive) (Parrott, Thompson, 1953); Comstock (1912) This (pi. 30, A) is ; England (CloudsleyHawaii (Pemberton, 1917). 1952) ; ; a tropical species frequently imported into localities with bunches of bananas (Comstock, 191 2 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 fe- northern ; male 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 when captured, but if it does, its bite bite spider does not attempt to 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 THERIDHDAE Latrodectus indistinctus Pickard-Cambridge Common — Button —Karnyia name. Natural prey. spider. discoidalis, South Africa, Western Cape The nest is constructed on the ground Province (Hesse, 1942) 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 and do not extend beyond the Capture is dependent upon ac- close to the tubular entrance to the nest trapping strands near the entrance. cidental contact of the insect with sticky threads surrounding the entrance. This spider apparently attacks any insect or arachnid that be- SMITHSONIAN MISCELLANEOUS COLLECTIONS 2l6 comes entangled in the nest. In an examination of 40 6 K. discoidalis were found. Latrodectus mactans Common VOL. I4I remains of nests, (Fabricius) —Black widow, hourglass, or shoe-button — Cockroaches, Puerto Rico (Petrunkevitch, names. Natural prey. spider. 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. LYCOSIDAE Family 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 /?-benzoquinone and its derivatives by Roth and Stay (1958). : Lycosa Experimental prey. —Supella Willis, unpublished data, 1953) sp. supellectilium, : The U.S.A. lycosid (pi. 30, (Roth and B-E) was prob- ably L. avida Walckenaer (tentatively identified by Dr. B. J. Kaston from a photograph). Order ACARINA PHYTOSEIIDAE Family Blattisocius tineivorus Synonyrny. —Blattisocius 1952]. — Natural triodons (Oudemans) Keegan [Baker and Wharton, Blattella germanica, U.S.A. (Keegan, 1944) host. 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 Natural host. sley, —Nauphoeta Womersley cinerea, Australia, Brisbane (Womer- 1956). Coleolaelaps (?) sp. Natural host. — Gromphadorhina portentosa, the hosts were im- ported into U.S.A. from Madagascar via Europe (Roth and Willis, BIOTIC ASSOCIATIONS OF COCKROACHES The mites unpublished data, 1958) termined by Dr. E. W. Baker. : Hypoaspis Natural host. —Pancsthia —ROTH (pi. 12, & WILLIS C) were 217 tentatively de- sp. australis, imported into U.S.A. from Cockroach Australia (Roth and Willis, unpublished data, 1955) made by determination of mite determined by J. A. G. Rehn. Generic : W. Strandtmann Dr. R. (Gamin, personal communication, 1955). UROPODIDAE Family Uropoda Natural host. Nymphs — sp. Formosa (Takahashi, 1940). be destroyed (Takahashi, 1940). Urop- Blattella hmnbertiana, of the cockroach may odids frequently attach themselves to insects, especially in nymphal stages but probably are harmless (Baker Family and Wharton, 1952). DIPLOGYNIIDAE Undetermined diplogyniid —Pancsthia australis, imported into U.S.A. from Gockroach (Roth and Willis, unpublished data, 1955) 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, Natural host. Australia : 1952)- Family ANOETIDAE Histiostoma feroniarum (Dufour) Natural host. —Pycnoscelus Though not surinamensis , Germany (Roeser, 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). 1940) : parasitic, the mites at times ; Family ACARIDAE Caloglyphus spinitarsus (Hermann) Natural 1940) : host. —Pycnoscelus surinamensis, Germany See notes following Histiostoma feroniarum above. (Roeser, SMITHSONIAN MISCELLANEOUS COLLECTIONS 2l8 Caloglyphus — VOL. I4I sp. germanica and Periplaneta americana, Natural hosts. Fales, and Mite feeds on organic matter but U.S.A. (Piquett 1952) of cockroach can reduce the vigor a colony. Blattella : Tyrophagus Common Associate. lintneri (Osborne) — name. Mushroom mite. Pycnoscelus surinamcnsis, U.S.A. (Roth and WilHs, — Mite determined by Dr. E. W. Baker (perunpublished data, 1953) sonal communication, 1953). Although this mite was found on the : cockroach, a it is 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 orientaUs often became infested with this species, but it did not affect the health or mortality of the cockroaches in his culture. Tyrophagus noxius A. Natural host. —Periplaneta Z. americana, U.S.A. (Roth and Willis, Mite determined by Dr. E. W. Baker (perunpublished data, 1953) sonal communication, 1953). Mites were found in the oothecal cavity : of a female cockroach that had been isolated for her entire adult The life. mites were in a closely packed mass behind a plug of what ap- peared 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. Not numerous (Rau, 1940a) : normally parasitic on cockroaches, but the mites became so would attack living as well as dead and dying cock- at times they roaches. Family GLYCIPHAGIDAE Chaetodactylus Synonymy. — Trichotarsus —Leucophaea sp. sp. [Baker and Wharton, 1952]. maderae, Puerto Rico (Sein, 1923) Mites found on cockroach's thorax and particularly among the folds of the wings (Sein, 1923). Mites of this genus are found infesting Natural host. organic matter (Baker and Wharton, 1952). : BIOTIC ASSOCIATIONS OF COCKROACHES Locustacarus Natural hosts. & WILLIS 2ig PODAPOLIPODIDAE Family (Roth and by Dr. E. —ROTH sp. —Diploptera punctata and Nauphoeta cinerea, U.S.A. Willis, unpublished data, 1954) W. Baker : Mite genus determined (personal communication, 1954). The mites 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. cluster thickly lOLINIDAE Family lolina Natural hosts. —Blaberus Harvard University) and nana Pritchard craniifer from a culture at (originally from (originally Diploptera punctata Hawaii), U.S.A., Pennsylvania (Roth and Willis, unpublished data, 1953 Pritchard, 1956) ; bases of the insects. mites usually attached near the wing The : Morphologically, the species is intermediate be- tween certain predaceous and phytophagous mites (Pritchard, 1956). Family PTERYGOSOMIDAE Pimeliaphilus podapolipophagus Tragardh Common —Cockroach —Parcoblatta name. Natural hosts. mite. 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. (CunliflFe, 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 secreon the host to adhere. Hatching occurs The newly 1 days at 80° F. 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 tion which enables those laid days at 90-95° F., and in 9-1 in 6-1 1 days. The mite then rests 3-4 days before molting. Entire life cycle Adult mite lives 2-3 weeks, during which time it can produce 2-3 batches of from i 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 covers a period of 28-32 days. ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 220 VOL. I4I Parasitism was proved by dehad fed on cockroaches which had unless live cockroaches were supplied. tecting radioactivity in mites that 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 the presence of cockroaches people (Baker et al., ; offices, these mites are an indication of the mite has been twice accused of biting 1956). RECORDS OF UNIDENTIFIED MITES Natural hosts. Four red —Aglaoptcryx fades, Puerto Rico nymphs found under wings of "tick" (Sein, 1923) : female. Blaberus craniifer, U.S.A., Florida (Hebard, 1917) "A number of [mites] are present on many of these specimens [28 ?$]." : lice Blaberus discoidalis, adventive from West Indies, taken in Scotland (Stewart, 1925) body and hind : A considerable number of mites were all over the Vv^ings. 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 was low. their oothecae, and the percentage of eggs hatching Parcohlatta uhleriana, U.S.A., North Carolina (Hatcher, 1939) Hypopi of mites were found deeply embedded in the fat body of two : individuals. by which they are dischambers of a mollusk the gonads of a millipede (Baker and Wharton, 1952). Mites persed. and in in the hypopial stage attach to insects Hypopi have been found in the gill Periplaneta americana, U.S.A., in laboratory (Fisk, 195 1) : The insects were sluggish and molted with difficulty. Gold Coast Colony (Macfie, 1922) Larvae of a tarsonemid mite were found in the feces. : Pycnoscelus Surinam ensis, Hawaii (Illingworth, 191 5) During the summer the soil was literally swarming with young of various stages. : Early in September most of the adults were dead and with mites. U.S.A., Connecticut, (Schwabe, 1950) from mite infestations. waii, in laboratory ently died in laboratory : Some all were covered (Zappe, 1918a). Ha- of the cockroaches appar- — The cockroach mite, Pimeliaphihis podapolipophagus. (From Baker reproduced through the courtesy of Dr. E. W. Baker and the National Pest Control Association.) Fig. 4. et al. [1956] ; 222 SMITHSONIAN MISCELLANEOUS COLLECTIONS Cockroach, England? (Ealand, 191 5) : Cockroaches may VOL. I4I carry the hypopial stage of the cheese mite. CONTROL OF MITES IN COCKROACH COLONIES Fisk (1951) eliminated the mites [possibly Piw^/ia/'////M.y podapoli- pophagns (Baker et al., 1956)] in his cockroach colony by using a 5percent spray and a s-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 new mite invasions. Oadri (1938) employed similar control prevent 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 (Sein, 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 —Scutigera forceps —House centipede. prey. — Cockroaches, U.S.A. Synonymy. Common (Linnaeus) 'R.z.^nesc^nt [Crabill, 1952]. name. Natural (Felt, 1909; Back, 1947; Auerbach, 195 1; Crabill, 1952; and others): This predator-prey relationship seems to be based on good circumstantial evidence (Crabill, personal communication, 1953). Experimental prey. — BlatteUa gernianica, newly hatched nymphs BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS and adult female, U.S.A. (Snodgrass, 1930; Roth and 223 Willis, un- published 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 nymph a small American cockroach that we caught and held Before it had finished its its jar. 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). meal, placed in it AUothereua maculata (Newport) Synonymy. —Scutigera maculata 1957]- Natural prey. m Annandale [Crabill, personal —Cockroaches, Malay peninsula, Batu caves et al., 191 3) : This Family SCOLOPENDRIDAE Scolopendra cingulata Experimental prey. 1955) (Ridley a presumptive host record. is SCOLOPENDROMORPHA Order '• communication, Latreille — Cockroaches, England After capture in France, this (Cloudsley-Thompson, 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 —Cockroaches, India, Experimental prey. As soon (Lherminier, 1837). Nagpur (Jangi, 1955) : became aware of its prey, it rapidly embraced the cockroach within its legs and with its fangs gripped the as the centipede insect's thorax. fangs while its The predator continued mouth parts prodded on an adult cockroach, the centipede 2-3 days. to hold the prey with its the victim's body. After feeding is not inclined to kill another for SMITHSONIAN MISCELLANEOUS COLLECTIONS 224 VOL. I4I Scolopendra subspinipes Leach Natural prey. is a common —Cockroaches, Hawaii species with a (Williams 1931) et al., body length of 6 or more inches. : This It is re- ported to be a great enemy of cockroaches. Scolopendra Natural When prey. —Eciobius sp. England (Lucas, 191 1, 1920) was holding a live cockroach which it caught. The insect was held beneath its captor's panzeri, : captured, the centipede had apparently just body, ventral surface upward, by several of the anterior legs while the centipede fed. INSECTA Class 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. : ; ODONATA Order AESHNIDAE Family 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 associations of cockroaches, as is associates of other cockroaches, are discussed in XVI. Other and as chapters XV and discussed in chapter commensals with other insects XVII. Order ORTHOPTERA Family MANTIDAE Hierodula tenuidentata (Saussure) (?) (Serville) (PI. 32) — Experimental prey. Blatta orientalis, Diploptera punctata, Eurycotis floridana, Leucophaea maderae, Nauphoeta cinerea, Neostylo- BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 225 pgya rhomhifolia, and Periplaneta amcricana, 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. rhomhifolia after grasping and making a brief attempt to chew the cockroaches. However, if specimens of A'', rhomhifolia were left in the jars with the mantids, the cockroaches were usually eaten within the next 24 hours. A^. rhomhifolia 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 the mantid's attack, but it exception oi N. cinerea, A^. rhomhifolia may deter should be pointed out that, with the possible all the other species fed to these mantids give odorous substances when seized or disturbed. Apparently, certain naturally repellent compounds will deter this mantid, whereas others off that are presumed state of the to be repellent will not; mantid is however, the nutritional undoubtedly a factor which may limit the ef- fectiveness of certain repellent secretions against this predator. Byrsotria fumigata, teneral males, and Periplaneta aiistralasiae, nymphs, U.S.A. (Roth and Willis, unpublished data, 1958). Diploptera punctata, U.S.A. (Eisner, 1958). Mantis religiosa Linnaeus Common name. —European mantis. —Naiiphoeta Experimental prey. cinerea, and Periplaneta ameri- cana, U.S.A. (Rilling, personal communication, 1957). Metallyticus semiaeneus — Cockroaches, Experimental prey. Borneo (Shelford, 1916). Sphodromantis viridus —Sphodromantis sonal communication, 1958]. —Blatta Experimental Synonymy. prey. species of cockroach mantid Westwood bioculata (Forskal) Burmeister [Gurney, per- orientalis, Egypt (Adair, 1923) This was apparently used regularly as food for the : in the laboratory. Stagmomantis Carolina (Johansson) Common name. — Carolina mantis. — germanica Experimental prey. Blattclla cana, U.S.A. (Breland, 1941) cockroaches daily. One : and Periplaneta amcri- The mantids were fed 1-2 female mantid consumed 10 adult German German cockroaches plus one ootheca and part of another in 2.5 hours. An SMITHSONIAN MISCELLANEOUS COLLECTIONS 226 adult German cockroach was consumed in VOL. I4I 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 Tenodera — aridifolia sinensis (Faure, 1940). 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 — Cockroach eggs, Japan (Asano, 1937) These are Asano found D. apicalis and D. japanica beneath his house near several empty cockroach oothecae which appeared to have been eaten into. He assumed from the condition of the Natural prey. questionable : records. oothecae 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 (ob- — : from an ootheca being carried by a female) and eggs of P. japanica (presumably in oothecae) were fed to both species of stone crickets in the evening. The eggs w^ere devoured by the next morning. tained 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. daceous bug is — Cockroach, Hawaii commonly found about (Illingworth, 191 7) : This pre- buildings. Illingworth says that BIOTIC ASSOCIATIONS OF COCKROACHES Kirkaldy suspected that saw it it on small fed —ROTH blattids & WILLIS 22/ and that Dr. Perkins feeding on a dead cockroach. REDUVIIDAE Family Pinto Spiniger domesticus —Periplaneta americana, Brazil, Matto Grosso This bug preys principally on cockroaches and was observed infesting the walls of dwellings where it preyed on P. Natural prey. (Pinto, 1927, 1927a) : americana. Triatoma arthurneivai Lent and Martins Natural prey. 1941) : —Monastria Brazil, sp., Minas Gerais This bug probably feeds on cockroaches of (Martins, this genus, as well as on rodents. Undetermined reduviids Natural prey. —Arenivaga roseni and Polyphaga saussurei, Turk- These desert cockroaches S.S.R. (Vlasov and Miram, 1937) are found in burrows of rodents and desert turtles around Ashkhabad. Reduviids are their main enemies. Vlasov (1933) found nymphs of men : Reduvms burrows christophi Jak. and R. fedtschenkianus Osch. in similar same area, although he did not specifically cite them as in this enemies of the desert cockroaches. Family NEPIDAE Kanatra Experimental prey. sp. — Cockroaches, U.S.A. (Hoffman, 1924). Order NEUROPTERA Family ASCALAPHIDAE Undetermined larva Experimental prey. — germanica, Blattella Kenya Colony (So- meren, 1924). Order From the that flies few observations DIPTERA that have come to our attention, are comparatively rare parasites in cockroaches. Family PHORIDAE Megaselia Host. —Eggs of Parcoblatta sp., sp. Ohio (Edmunds, 1952a). it seems SMITHSONIAN MISCELLANEOUS COLLECTIONS 228 Family VOL. I4I CONOPIDAE Stylogaster sty lata (Fabricius) — Cockroaches, Brazil (Souza Lopes, 1937) L. Travassos was quoted as having observed this species pursue cockroaches that Hosts. : 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 Qgg 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 specimen was found in a sp., Brazil (Souza Lopes, 1937) museum attached to the posterior end of Cockroaches, Panama (C. collection with its W. : An adult an egg of Stylogaster abdomen. Rettenmeyer, personal communica- "Seven species were collected hovering over army ant flies were seen apparently attacking cockroaches that had been flushed by the ants." tion, 1959) : swarms and a few Family LARVAEVORIDAE Calodexia (?) venteris Cur ran Hosts. —Periplaneta amerkana, Brazil (Souza Lopes, 1937) tained complete evolution of the parasite in this host. This : Ob- may have been an experimental host. Calodexia spp. Hosts. — Cockroaches, Panama (Rettenmeyer, personal communica- 1959) Swarms of army ants are accompanied by about 20 species of Calodexia. These flies larviposit on the cockroaches, crickets, and tion, : from cover by the ants. Larvae were found in one(?) cockroach. Larvae from an adult of Calodexia were introduced experimentally into a cockroach and suc- possibly other arthropods that are flushed cessfully reared. Undetermined tachinids Hosts. —Eurycotis data, 1953) : fioridana, Three larvae (det. from Florida (Roth, unpublished by W. W. Wirth) were found in a living adult male. Panesthia anstralis, from Australia (Roth, unpublished data, Reared from a wild-caught cockroach that was maintained in 1957) a laboratory colony. : COCKROACHES BIOTIC ASSOCIATIONS OF Cockroaches, 1955) : (E. Australia Family personal Riek, F. Reared from some of the larger —ROTH & WILLIS 229 communication, species. MUSCIDAE Coenosia basalis Stein Host. — Eggs of Parcoblatta sp., Ohio (Edmunds, 1952a). SARCOPHAGIDAE Family Sarcophaga omani Hall Host. tion, —Arenivaga 1953) holliana, Specimens : Texas (Wirth, personal communica- U.S. National Museum. in Sarcophaga lambens Wied. — Synonymy. Sarcophaga sternodontis (Towns.). Hoffman (1927) claimed that approximately 40 percent of some specimens of Pycnoscelus stirinamensis collected in southern Haiti were parasitized by .9. at the University of Hoffman was lambens. However, according to entomologists Puerto Rico Agricultural Experiment Station, 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 were also introduced ; first into the body cavity of cockroaches which had their heads and legs removed. Adult sarcophagids were instar larvae collected at freshly killed American cockroaches used as Order bait. COLEOPTERA Family CARABIDAE Harpalus pennsylvanicus De Geer Experimental prey. et al., 1934) : — Cryptocercus punctulatus This beetle latus in nature. is In the laboratory as large as itself. , U.S.A. (Cleveland often found in the galleries of C. punctuit killed and devoured cockroaches SMITHSONIAN MISCELLANEOUS COLLECTIONS 230 Family Rhantus VOL. I4I DYTISCIDAE pacificus Boisduval — Experimental prey. Cockroaches, disabled, Hawaii (Williams, This beetle, which is common in mountain streams, located 1936) : wounded cockroaches in an aquarium by sense of smell or taste rather than sight. Family LAMPYRIDAE Undetermined larva Experimental prey. —Parcoblatta virginica, adult female (pi. 33, to the C), U.S.A. (Roth and Willis, unpublished data, 1953). Family RIPIPHORIDAE Neonephrites partiniger ^ Riek — Natural host. Cockroach (undescribed genus belonging Pseudomopinae), Australia Capital Territory (Riek, 1955). Neorhipidius neoxenus Riek Natural host. —Robshelfordia longiuscitla or Roh shelf ordia circum- ducta, 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 Pupal stage lasted only 3 days. (Riek, 1955) : Rhipidioides adynatus Riek Natural host. —Escala sp. or an undescribed genus of Pseudo- mopinae, Australia, Victoria (Riek, 1955). Rhipidioides Natural host. —Ellipsidion fuscatus affine, Riek Australia, New South Wales (Riek, 1955). Rhipidioides helenae Riek Natural host. —Robshelfordia longiuscula or Robshelfordia circum- ducta, Australia Capital Territory (Riek, 1955). s Barber (1939) has shown that the correct spelling of the name of the type genus is Ripidius and not Rhipidius, as it is frequently written, and that, conse- quently, the family should be Ripiphoridae and not Rhipiphoridae. COCKROACHES BIOTIC ASSOCIATIONS OF —ROTH & WILLIS 23 Rhipidioides mollis Riek Natural host. —Rohshelfordia longiuscula or Rohshelfordia circum- ducta, Australia Capital Territory (Riek, 1955). Rhipidioides rubricatus Riek Natural liost. — Choristinia and Choristimodes sp. sp., Australia Capital Territory (Riek, 1955). Riekella australis (Riek) Synonymy. —Nephrites — Cutilia Natural host. 1955) • Two australis Riek [Selander, 1957]. Australia sp., Capital Territory (Riek, females emerged from one host. Riekella nitidioides Selander Synonymy. —Nephrites nitidus 1957]. —Platyzosleria Natural host. of Riek not Shuckard sp., [Selander, Tasmania (Riek, 1955). Riekella sp. Synonymy. Natural —Nephrites [Selander, 1957]. —Platysosferia castanea, Australia sp. host. (Riek, 1955). Capital Territory — Biology of Australian Ripidiini. The Australian species of Ripiof apparently endemic, ground-dwelHng species of diini are parasites cockroaches. There parasite genus Blattinae. : is some Riekella spp. correlation between host subfamily [ = Nephrites] and have only been bred from Rhipidioides spp. occur only in the closely related Ecto- 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 biinae 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 trees pupating. The larviform, vringless female remains near the pupal skin and is sought out by the winged male. The eggs are around the pupal skin (Riek, 1955). Ripidius = laid in a mass boissyi Abeille Balduf (1935) lists Ripidius boissyi as parasitic on nymphs of Ectobius pallidus giving Abeille de Perrin (1909) as a source for this SMITHSONIAN MISCELLANEOUS COLLECTIONS 232 information. VOL. I4I However, Abeille de Perrin simply presumed that R. boissyi parasitized E. pallidiis because he collected this cockroach in the same Abeille de Perrin suggested that the habitat as the beetle. species of the genus Ripidiiis lived in the bodies of cockroaches, but there are no rearing records, as far as v^e know, oi 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 — gcrmanica, on Natural Blattella hosts. [Leng, 1920]. shipboard (Sundevall, Germany (Aclogue and Fowler, in Burr, 1899a) on steam"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 identifi1831) ; ; ship ; : ; cation. ( 1909) stated that R. pectinidescribed by Sunders [sic] as blattarum because it had Ectobius pallidus f Abeille de Perrin cornis was first 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, nymph on shipboard (Sundevall, 1831) : One only. With the exception of the single nymph of P. americana, R. pectini- and nymphs of B. gerBarbier (1947) found only B. gernmnica parasitized, although both Blatta orien talis and Supella supcllcctilium were prevalent on board the ship. Primary larvae of the parasite failed to parasitize cornis apparently attacks only adult females manica. Supella. Adult behavior. —The winged male compared to 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 apterous female The eggs (50-100) ; it runs around, are laid among is relatively active flies well, a network of silk fibers secreted by BIOTIC ASSOCIATIONS OF The female the female. 1947)- Development. —The COCKROACHES —ROTH & WILLIS dies after completing oviposition 233 (Barbier, eggs hatch after 14 days, and the primary its body; the larvae then (triungulin) larvae ascend the host's legs to cut the intersegmental membrane between abdominal segment of the cockroach, the metasternum and first in order to enter the host's ab- domen (Barbier, 1947). Chobaut (1892) first suggested this method of attack by the ripiphorid larva. As the parasites develop, the abdo- men of the host becomes swollen. Developing larvae apparently eat the host's fat body, leaving the vital organs until the last. Parasitized and the eggs, when formed, never hatched. Development of the ootheca 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 female hosts were sterile cockroach except one host which, when crushed, yielded five. Stamm (1936) found three hosts infested with five larvae each. In a over 100 cockroaches, Stamm found 10 that were parasitized. The day before little the parasite leaves the host, the cockroach shows an finally falling over on its back. The 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 abrupt uneasiness and runs about, parasite larva emerges 13 days (males) (Barbier, 1947). —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 Distribution. ; ; (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 in ports between neighboring ships by parasitized cock- very easily roaches in baskets or sacks of provisions. Ripidius Natural hosts. — Blattidae, Scutellaria Heller Philippine Islands (Schultze, 1925). SMITHSONIAN MISCFXLANEOUS COLLECTIONS 234 VOL. I4I DERMESTIDAE Family Dermestes ater —Black larder —Blatta Common name. De Geer beetle. U.S.A. (Roth and Willis, unpubDermestes ater is generally a scavenger, but vs^e have seen adult beetles, which had developed in our cockroach colony, clinging to and feeding on living oriental cockroaches, eventually killthe beetles probably attack only the v^^eakened or injured ing them cockroaches in a culture. This was a natural infestation of a laboratory culture by a predator. Experimental prey. Blattella germanica, oothecae, U.S.A. (Roth and Willis, 1950) The beetle larvae can penetrate unhatched oothecae of the German but not those of the American or oriental cockroaches. Natural prey. orientalis, lished data, 1953) : ; — : Dermestes Natural prey. —Blatta sp. orientalis, oothecae, U.S.A., Missouri : Rau (1924) stated that Dermestes larvae often infest the tgg 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 ; extruded between a pair of sternites towards the base of the abdomen and appears to belong to the family Halictofemale parasite is phagidae." 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 re- corded as developing on cockroach eggs. All the Evaniidae are pre- sumed 1940; egg capsules of cockroaches (Clausen, Townes, 1951), although hosts for many of the described to be parasitic in the species have yet to be discovered. The presence of evaniids in dwell- ings indicates the presence of cockroaches (Gross, 1950). At times COCKROACHES—ROTH & WILLIS BIOTIC ASSOCIATIONS OF may become these wasps a nuisance ; 235 a family in Worthington, Ohio, complained of the evaniid wasps that they found on the windows and not annoyed in other areas of their home, but they were apparently by the oriental cockroaches in the basement (Edmunds, 1953). The known parasites of cockroach eggs are listed below with summaries of their biology. EVANIIDAE Family Acanthinevania princeps (Westwood) Synonymy. —Evania princeps [Dr. H. Townes, personal communi- cation, 1956]. Natural host.— Cockroach eggs, Australia (Froggatt, 1906). Brachygaster minutus Synonymy. —Evania minuta Olivier — germanica, Natural Blattella hosts. (Olivier) [Kieflfer, 1920]. Europe? (Schletterer, 1889; Kiefer, 1912; Crosskey, 1951.) Ectobius lapponicus, Europe? (Schletterer, 1889; Crosskey, 195 1.) Ectobius panseri var. nigripesf Great Britain (Blair, 1952) is 1912; Kieflfer, : This Niton and Headon a presumptive record. The wasp was collected at Wight, an area in which this variety of E. panseri was the Hill, Isle of only species of cockroach Ectobius Museum sp., known to occur. England (Cameron, 1955, 1957) : Natural History records. Adult wasps have been collected on Asparagus officinalis Linnaeus (Schmiedeknechtm Schletterer, 1889; Crosskey, 1951). Thompson's appendigaster (1951) citation of records of B. minutus and Evania from Blatta orientalis and Blattella germanica, and Cameron's (1957) and one from Ectobius lapponicus, all attrib(1921), are almost certainly in error. Kadocsa but (1921, p. 33) listed these wasps as egg parasites of cockroaches not necessarily in Hungary and did not name specific cockroach hosts. citation of these records uted to Kadocsa The present writers have found no information, other than host re- 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 ports, B. germanica carries its before the eggs hatch, it ootheca attached to the abdomen until or just would seem that the female of B. minutus (if the host records are valid) species while it is still must oviposit into the ootheca of this being carried by the female ; this would not : SMITHSONIAN MISCELLANEOUS COLLECTIONS 236 VOL. I4I 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 (Kiefifer, 1920). : Evania appendigaster (Linnaeus) Synonymy. —Evania — dcsjardinsii Bordage, Evania laevigata La- [Dalla Torre, 1901-1902]. treille Natural hosts. (Westwood, 1854, 1954a). Europe? (Schletterer, 1886; Howard, 1888, Egypt? (Alfieri, 1914; Adair, 1923). [Girault Blatta, "exotic species" Blatta orientalis, Kieffer, ( 1912) ; 1907, 1914) erroneously attributed another record to Marlatt ( 1902) ; See also notes under Brachygaster minutus with respect to Kadocsa.] see footnote 6. Blattella germanicaf (Girault 1907, 1914). ously an error. [This record is Girault attributed the record to Marlatt (1902) obvi; see footnote 6.] Cutilia soror, Hawaii (Swezey, 1929; Zimmerman, 1948). Leucophaea maderae (Schletterer, 1889; Bordage, 1896; Kiefifer, These records are probably erroneous inasmuch as this cock1 91 2) roach incubates its eggs internally (Roth and Willis, 1954). Later, after finding that L. maderae is ovoviviparous, Bordage (1913) admitted having misidentified a parasitized ootheca from some other : 6 Page II of Marlatt (1902) has been cited erroneously so many times in support of host records for T. hagcnowii and Evania appendigaster that we are quoting the pertinent section below. In the section preceding the quoted material Marlatt discusses the American, Australian, oriental, and German cockroaches. is nothing in the paper to connect any of these cockroaches specifically There with the parasites mentioned below NATURAL ENEMIES AND PARASITES "In Europe the egg capsules of the cockroach are often parasitized by an ichneumon fly (Evania appendigaster). This insect has become widely distributed over the world following its host insect, and has been redescribed under a great many dififerent names. It was found in Cuba as early as 1829, and has been several times collected in the United States. Unfortunately, its usefulness as a means of keeping the roach in check by destroying the egg capsules is greatly impaired by the occurrence of another ichneumon fly {Entedon hagenowi), which is parasitic upon the has been brought over with its first. This host parasite. is also a European species which egg capsule parasite of If the true the roach could have been introduced into this country without this secondary would doubtless have been very much greater. The secondary parasite, however, seems to have been introduced early, and has been found in Cuba and Florida, and probably occurs as widely as its host and prevents the latter from multiplying very greatly." parasite, its usefulness BIOTIC ASSOCIATIONS OF COCKROACHES species ; —ROTH & WILLIS 237 this species are pro- he concluded that the developing eggs of tected 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 while the latter within the parent. is still known behavior was not definitely However, the records cited : Egg He placed in the embryo stated that although this to occur, it probably could occur. above do not indicate that the alleged parasitization followed this pattern. Neostylopyga rhombifolia, Hawaii (Swezey, 1929). Periplaneta americana, Europe ( Schletterer, 1889; Bordage, 1896; Kiefifer, 1912) Reunion Island (Bordage, 1913) Puerto Rico (Sein, ; ; PalHawaii (Swezey, 1929) Jamaica (Gowdey, 1925) (Ashmead, estine (Bodenheimer, 1930) Florida U.S.A., 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). 1923) ; ; ; ; ; ; ; Periplaneta australasiae, U.S.A., Florida (Ashmead, 1900) Hawaii (Swezey, 1929; Zimmerman, 1948). [Girault (1914) errone; ously attributed another record to Marlatt (1902) above.] Experimental Relatively (%• host. little —Blatta orientalis, detailed information ; see footnote 6, U.S.A. (Haber, 1920). was known about this wasp 5)> one of the earliest parasites of cockroach eggs to be dis- covered, until Cameron (1957) and Spence, 1826) studied its biology. Arnold (Kirby discovered that the genus Evania parasitized know whether the wasp developed on the cockroach MacLeay (West wood, 1843) determined that Evania developed within the oothecae of cockroaches. Westwood Blatta, but did not eggs or in the nymphs. (1854a) found the larvae, pupae, and adults of E. appcndigastcr in egg cases of an unidentified species of cockroach found on orchids received from Calcutta. Adult behavior. —Adult wasps visited flowers of parsley, Pctro- selium crispum, and fennel, Foeniculum vulgarc (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) ; 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). Evania sp. were attracted to the — Shelford (1912, 1916) erroneously supposed that Oviposition. Evania, by means of her cleaverlike abdomen, opened the ootheca at the crista and then deposited her egg or eggs on the eggs of the cockroach. Haber (1920) observed and described oviposition. The female — SMITHSONIAN MISCELLANEOUS COLLECTIONS 238 VOL. I4I wasp crawled over the surface of the ootheca, 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 ootheca in the fifth cell on the minutes. left side; she remained in this position for about 15 similar oviposition behavior that Cameron (1957) described lasted about half an hour. Kieffer (191 2) and Crosskey (1951) Fig. 5. Evania appendigaster. Left, dorsal view, X 8. Right, side view, X 5. (Reproduced with permission. British Museum [Natural History], 1951, figs. I A and iB.) stated that the female deposits her eggs before the walls of the ootheca harden. Development. — Kieffer eat the cockroach eggs (1912) stated that the larvae in this family and pupate in the ootheca without forming a Smith (1945) stated that the larva feeds on one cockroach tgg after another until all are destroyed by that time it is full grown and it pupates within the ootheca. Cameron (1957) found that there cocoon. ; are five larval instars and that in material from Saudi Arabia there are three or possibly four generations a year. BIOTIC ASSOCIATIONS OF COCKROACHES Distribution. —Tropical and New York —ROTH & WILLIS 239 subtropical parts of the world as far of Europe except the northern part 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 north as (Kieffer, 1920 City, and all ; of blattids he listed small. is Evania dimidiata Fabricius Synonymy. —Evania abyssinica —Blatta Natural host. orientalis, Westwood Egypt? [Schletterer, 1889]. (Alfieri, 1914). Evania subspinosa Kieffer —Periplaneta Natural host. Lever (1946) listed sp., this species as a state that he actually reared it Although Fiji (Lever, 1946) cockroach-egg parasite, he did not : from Periplaneta oothecae. Hyptia dorsalis of Ashmead Synonymy. — Dr. lieves that this H. Townes, (personal communication, 1956) bewasp 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 (Ash- — ; mead, 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. May : Ootheca collected by de17, 1956; wasp emerged June 12, 1956 (pi. 33, B) termined by Dr. H. Townes. The keel region of the ootheca of P. uhleriana (pi. 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. L. Roth, ; Development. —The last instar larva overwinters inside the cock- roach ootheca (Edmunds, 1954). Five oothecae yielded one parasite each (Edmunds, 1953a). — Canada, Ontario. U.S.A. Distribution. New Hampshire and Minnesota to South Carolina, Mississippi, Texas, and Kansas. Upper and Lower Austral Zones (Townes, 1951). : SMITHSONIAN MISCELLANEOUS COLLECTIONS 240 VOL, I4I 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. Mexico. — U.S.A. Upper Austral Pennsylvania to Florida and Louisiana. : to Tropical Zones (Townes, 1951). Hyptia thoracica (Blanchard) —Parcoblatta pensylvanica, U.S.A., Ohio Natural host. (Edmunds, 1952a, 1953a, 1954). Adult behavior. Copulation — was rapid, lasting only a few seconds. Blooms of Asmorrhisa longistylis were placed in a cage with adult wasps. The insects were attracted to and fed on the flowers (Edmunds, 1954). Development. larva. — Entire contents of ootheca are eaten by the single Last instar larva overwinters inside the o5theca. Emergence in Ohio was around the middle of June. The emergence hole made by genus was about 2 mm. in diameter. The hole was made at the this Adult took about 65 minutes emerge from the time its mandibles first broke through the oothecal wall. (Edmunds, 1954.) Connecticut to WisconCanada, Ontario, U.S.A. Distribution. sin, south to Florida and Texas. Upper Austral to Tropical Zones. (Townes, 1951.) top side of the ootheca near one end. to — : Hyptia Natural host. site identified — Cariblatta sp. delicatula, Cuba (Hebard, 1916a) ; Para- by Ashmead. Hyptia Natural host. —Parcoblatta sp. sp., (undescribed) U.S.A., Ohio (Edmunds, 1952a). Prosevania punctata (Brulle) Synonymy. —Evania punctata Brulle [Townes, —Blatta Natural and experimental hosts. sula (Fahringer, 1922) ; 1949]. orientalis, Istrian Algeria (Cros, 1942) ; Penin- U.S.A., Ohio (Ed- munds, 1954). Blattellagermanicaf Europe? (Girault 1907, 1914) Europe (Fahr1922). [The records on this host are extremely doubtful. ; inger, Girault erroneously cited Marlatt (1902) as the source of this record ; see footnote 6, page 236. Fahringer, however, claimed that he obtained BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 24I seven female parasites from oothecae of Blattella germanica. He placed female parasites witli adults of B. germanica in a glass cage. As soon as oothecae could be seen between folds of a woolen rag, he re- moved sites all the larger cockroaches and held the oothecae until the para- emerged. Fahringer may have been dealing with a different species of cockroach, because placing oothecae in crevices (or between folds of rag) a habit foreign to B. germanica, the female of which is usually carries her ootheca until hatching or until about a day before. Edmunds (1953b) could not induce this wasp to parasitize eggs of B. germanica. ] Periplancta americana, Istrian Peninsula (Fahringer, 1922) estine (Bodenheimer, 1930) 1954)- Adult behavior. ; —The wasps about a great deal and fly ; Pal- U.S.A., Ohio (Edmunds, 1952, 1953b, (pi. 33, A) are very active; they walk They short distances. 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) main- tained adults 17 days without food. unidentified flowers in the laboratory. Edmunds (1954) He fed adults on also maintained them for 20 days after capture on a 5-percent honey solution. Oviposition. —A female P. punctata selected oothecae of P. ameri- cana for oviposition and ignored those of B. orientalis and Parcoblatta pensylvanica in the same cage. Oviposition was accomplished as described for Evania appendigaster. One ootheca was turned over onto by the wasp before she oviposited. (Edmunds, 1952.) Although there seemed to be a "preferred" position for oviposition, it its right side was not obligatory. The usual position was for the female to face the from the opposite side or, keel of the ootheca, but she also oviposited rarely, directly down into the side of the ootheca. The average time spent by females in 10 ovipositions was 29 minutes (range 16-62 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 oothecae 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. minutes) . 242 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I Edmunds (1954) placed females of Periplaneta americana that were carrying oothecae, into cages with Prosevania; some of the female wasps showed considerable interest in the attached oothecae, but he observed oviposition only into egg cases that had been dropped by the cockroaches. Cros (1942) described an interesting reaction that he called "intoward Prosevania. A wasp was placed in a jar in which a cockroach had just deposited its stinctive hostility" of the oriental cockroach The wasp tried to oviposit into the egg case but was upset and pursued by the cockroach. The cockroach placed herself over the ootheca, standing high on her legs, and remained there motionless. The wasp then approached from the rear, slipped under the cockroach, ootheca. and, unnoticed by the cockroach, climbed on the ootheca 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 Peri- days (Edmunds, 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 planeta americana: 1952). Only one Three wasps developed in 127 parasite develops in each ootheca. mm. (Cros, 1942.) Partheno- genesis exists; the unfertilized eggs produced only males (Edmunds, ootheca through a hole 4 1954). Distribution. in diameter. —Eastern U.S.A., from New York and Ohio south to Georgia (Townes, 1949). Europe, Syria, Palestine (Kieffer, 1920). Szepligetella sericea Synonymy. —Evania sericea (Cameron) Cameron [Townes, 1949, personal communication, 1956]. Evania impressa Schletterer [Townes, 1956]. Natural hosts. — Cutilia soror p. c, and Neostylopyga rhombifolia, Ha- waii (Swezey, 1929). Periplaneta americana and Periplaneta (Swezey, 1929; Zimmerman, 1948). Periplaneta sp., Fiji australasiae, Hawaii (Lever, 1943, 1946). Adults are sometimes found resting on leaves covered with honey dew (Williams et al., 1931). BIOTIC ASSOCIATIONS OF COCKROACHES Zeuzevania Natural hosts. splendidula —Loboptera —ROTH 243 Costa France decipiens, & WILLIS (Lavagne, 1914; Genieys, 1924). Picard (1913) believed that Z. splendidula parasitized L. decipiens its eggs; however, Lavagne (1914) explained the true re- and not lationship by dissecting two specimens of Z. splendidula from oothecae of L. decipiens. The following information position. —Wasp ^gg wall hardens. is Some is taken from Genieys (1924) Ovistill-soft o5theca before the : introduced into the oothecae had four oviposition scars but never contained more than two parasite eggs. mences development development, but it in July or — Larva comOnly one larva completes the ootheca. The wasp passes Development. August. eats all the eggs in 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 oothecae 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 "the parasite australe, Australia, when ready 191 7) space of the destroyed eggs." : Cockroach, Australia, New to emerge Queensland (Dodd, whole fully occupies the 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 ootheca 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, 19 15, 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). ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 244 VOL. I4I Cheiloneurus viridiscutum (Girault) Synonymy. — Cristatithorax [Mercet, 1921]. —Ellipsidion Natural host. Girault = Cheiloneurus Westwood Queensland (Dodd, australe, Australia, 1917). Comperia merceti (Compere) Synonymy. — Comperia 1951]. — Natural Blattella hosts. (Gomes, 1941) merceti germanica, In the English : falsicornis var. Brazil, summary of Gomes [Peck, Federal Distrito his paper, Gomes states How- that C. merceti var, falsicornis v^as reared from B. germanica. body of the paper, he states that the supposed origin of the parasite was the ootheca of B. germanica. Burks (personal comever, in the munication, 1956) does not believe that this wasp parasitizes the eggs (unpublished data, 1957) exposed six oothecae of B. germanica. We In order to retard water loss the of B. germanica to C. merceti. oothecae were removed from the females by cutting the insects in two so that each ootheca remained attached to the posterior pari of the abdomen. No wasps developed in these oothecae. Supella supcllectilium, U.S.A., Kansas (Lawson, 1954a) Hawaii ; (Zimmerman, 1944; Compere, 1946; Keck, 1951). Adult behavior. Males and nonovipositing females showed a flealike 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 (pi. 34, with the sheath of her ovipositor ; it B) selected a site on an ootheca was uncertain whether there was a definite preference for oviposition sites. Wasp tended to choose a She preferred to oviposit would place eggs in oothecae old, although she into eggs about 2 weeks less than a week old and in embryos in the green band stage. There were 1-50 oviposition punctures per ootheca. (Lawson, 1954a.) Development. If enough wasp larvae were present, they ate all eggs in an ootheca. Occasionally wasps developed in one end of an nearly horizontal position for oviposition. — ootheca while cockroaches developed in the other when this occurred, the cockroach nymphs always emerged last. The developmental pe; riod was 30-41 days per ootheca. The at room temperature. There were 5-25 single exit hole in the ootheca varied parasites from 0.6 to 0.9 mm. in diameter. (Lawson, 1954a.) U.S.A. New Jersey south to Florida, west to IlliDistribution. nois, Kansas, and Arizona. West Indies Central and South America Hawaii. (Burks, personal communication, 1956.) — : ; BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS Dicarnosis alfierii Natural hosts.—"Phyllodromia" sp., 245 Mercet Egypt (Mercet, 1930) : Ac- parasitized one cording to Mercet, Dr. Alfieri claimed that this wasp Phyllonamely, Egypt, in found 'Thyllodromia" of the species of dromia lectilium [=Blattella] Phyllodromia gernianica, and/or Phyllodromia treitliana. We [ = Supella] supel- do not know to which wasp belonged. Cockroach, Egypt? (Mercet m Compere, 1938.) modern genus the host of this Eutrichosomella blattophaga Girault Natural host.— Cockrosich, Australia, Queensland (Girault, 1915). Family EUPELMIDAE Anastatus blattidifurax Girault Natural host.— Cockroach, Australia, Queensland (Girault, 191 5). 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 WilHs, 1954a). Adult behavior.— Female wasps are sexually receptive almost im- Mating takes 3-4 seconds. Males females may also females several fertilize may and mate repeatedly 80° wasps lived 2-4 female the F. about At mate more than once. days, males one day. mediately on leaving the ootheca. ; first probes the ootheca with her acceptable spot she then drills an sheathed ovipositor until she finds ovipositor. One female oviher through the wall of the ootheca with were more usual. We have periods posited for 5 hours, but briefer Oviposition.—The female wasp ; ovipositing simultaneously into an ootheca that of Eurycotis floridana. One female was seen to feed on material ovimay A) (pi. wasp The 34, from the oviposition puncture. seen six or more females oozed being carried by and which dropped been the female, as well as in oothecae that have parasitized. successfully have hard walls. Eggs 36 days old were posit into the ootheca of E. floridana while it is still — Development. In Eurycotis floridana: In the laboratory, development was completed in 34-36 days at about 85° F. This time was ootheca. regulated to some extent by the number of parasites in the TBrues et al. (1954) include this family in the Encyrtidae. SMITHSONIAN MISCELLANEOUS COLLECTIONS 246 There is VOL. I4I evidence that larvae eat unhatched wasp eggs or other larvae. In 34 oothecae 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 oothecae 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 ootheca oothecae was two holes. ; the average number in 42 — Number of parasites per ootheca. In Blatta orientalis: One of III oothecae exposed to female wasps yielded 48 parasites. In EuryOne cotis floridana: sites ; ootheca parasitized in the field yielded 68 para- 8 oothecae exposed to single wasps for their entire lifespan yielded an average of 50 ±6 parasites (range 23-81); 34 oothecae exposed to many wasps for their entire lifespan yielded an average of (range 93-306). In Periplaneta americana: Nine 1 98 ±8 parasites oothecae of 152 exposed to the wasps were found to be parasitized when dissected 1 1 adults emerged from one ootheca no parasites emerged from the other 8 oothecae. Sex ratio. 4 $$: i J* from ovipositions by isolated females. In the one ootheca collected in the field, the ratio was 21.6 $$ i cf Parthenogenesis exists the unfertilized eggs produced only males. ; ; — : ; Anastatus Synonymy. tenuipes Bolivar —Anastatus hlattidarum sonal communication, 1957) is a synonym of A. tenuipes. is y Pieltain Ferriere. Dr. C. Ferriere (per- of the opinion that his A. hlattidarum He stated "I have never been able to see scription agrees with y P., which is in Madrid, but the deA. hlattidarum. I had not yet knowledge of Bolivar's description, when the unique type of A. tenuipes B. describing my The species. parasite of cockroaches eggs [Supella supellectilium'] should be called A. tenuipes Bol." Mani (1938) synonymized Solindenia blattiphagus Mani with Anastatus hlattidarum. Natural hosts. Supella supellectilium, Anglo-Egyptian Sudan (Fer- — Egypt (Alfieri, U.S.A., Arizona (Flock, 1941) 1956). Ohio (Hull and Davidson, 1958). Periplaneta americana, India (Burks in Roth and Willis, 1954a). riere, 1930, in 1935) Hafez and ; ; Afifi, India (Mani, 1936). Cockroach, Hawaii (Weber, 1951) The following is based on parasites that developed on eggs of ; — Adult behavior. Wasp may (Flock, 1941) be seen running rapidly on walls in buildings infested with the cockroach host. The wasp rarely flies but hops proficiently; when dis- Supella supellectilium : BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 247 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 ootheca by feeling with turbed, — her antennae. Flock stated, without citing experimental evidence, that chief factor determining was apparently the the age of the egg case The wasp took 15-45 minutes to oviposit. Three females ovia single female reposited simultaneously into a single ootheca choice. ; — Development. a constant temperature of peatedly oviposited into one ootheca at intervals. Completed in an average of 32.6 days at 82° F. Number 16) of parasites per ootheca. (Flock, 1941) Sex ratio. —4 ; —Average about 10.7 (range 4- 15 (Ferriere, 1935). $5:i J* 1935); average of 6 ?? i J* the unfertilized eggs pro- (Ferriere, : Parthenogenesis occurs; (Flock, 1941). duced only males (Flock, 1941). Distribution. U.S.A. Maryland, south to Florida, west to Illinois, Arizona. Guatemala Hawaii India Egypt Sudan. Kansas, and (Burks, personal communication, 1956). — : ; ; ; ; Eupelmus atriflagellum Girault Natural host. — Blattella germanica, Australia, Queensland (Girault, 1924). Eupelmus Natural host. —"Tree sp. cockroach," U.S.A., (Howard, Florida 1892). Solindenia picticomis Natural hosts. —Allacta similis, Cameron Hawaii (Perkins, 1906, 1913 Timberlake, 1924; Swezey, 1929; Zimmerman, 1948). Other species of cockroaches, Hawaii (Perkins, 1913). PTEROMALIDAE Family Pteromalus Natural Sells, host. sp. ? —Leucophaea maderaef, Jamaica 1842). [This host is (Westwood, 1839; undoubtedly an error. Sells stated that the ootheca which contained 96 unidentified chalcids had 16 dentations at the edge the description fits the ootheca 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 logical Society in 1838 Mr. Entomo- Sells exhibited 94 specimens of a small SMITHSONIAN MISCELLANEOUS COLLECTIONS 248 Pteromaliis (apparently identified by VOL. I4I Westwood) obtained from one cockroach ootheca. This same record of Sells was published posthu- mously in 1842, although in this paper he identified the host ootheca as Cameron (1955) lists a European record of from Periplaneta americana citing Girault (1914) as the source of the record. Girault's record was apparently taken from Westwood's footnote mentioned above.] "Blaherus" maderae. Pteromalus sp. Systellogaster ovivora Natural hosts. —Blatta orientalis, Gahan U.S.A., Illinois (Gahan, 1917). Parcohlatta pensylvanica, Canada, Ontario (Judd, 1955). Parcohlatta sp., U.S.A., Ohio (Edmunds, igS^a., 1953a). Maryland (Gahan, 1917). ootheca of P. pensylvanica yielded 14 parasites with a sex ratio of 2.5 55: I (^ (Judd, 1955). The average number of parasites in II oothecae of Parcohlatta sp. collected in 1950-51 was 27 wasps (Ed"Blattid," U.S.A., One munds, 1952a, 1953a). The adults made two to three emergence holes in the ootheca (Edmunds, 1953a; Judd, 1955). EULOPHIDAE Family Melittobia chalybii Natural host. M. —Periplaneta Ashmead americana, U.S.A., Missouri (Rau, and Hymenoptera (Peck, 1951). This is the only record from cockroach eggs. Burks (personal communication, 1956) stated that this species 1940a) : chalybii any insect is normally a parasite of Coleoptera which it is exposed and can be a serious pest in 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). will attack to insect cultures of practically ; Mestocharomyia oophaga Dodd Natural host. —Ellipsidion aiistrale, Australia, Queensland (Dodd, 1917). Syntomosphyrum blattae Burks —Parcohlatta Natural hosts. sp., U.S.A., Ohio (Burks, 1952; Edmunds, 1952a, 1953a) Ten oothecae yielded an average of 92 wasps (Edmunds, 1952a). Five oothecae, collected a year later, yielded an average of 74 wasps adults sometimes made two to three exit holes in the ootheca (Edmunds, 1953a). : ; Cockroach, U.S.A., West Virginia (Burks, 1952). BIOTIC ASSOCIATIONS OF COCKROACHES Syntomosphyrum ischnopterae Synonymy. —Epomphaloides —ROTH & WILLIS 249 (GIrault) 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 dehe stated that the species is posited in the U.S. National Mueseum apparently Syntomosphyrum ischnopterae. In view of the experi; mental work by Parker and Thompson (see below), this wasp may prove to be a hyperparasite on evaniids in cockroach oothecae rather than a primary parasite on cockroach eggs. (See Zeuxevania splendidula, p. 243.) Natural hosts. —Ischnoptera sp. [probably Parcohlatta sp. (Rehn, personal communication, 1958)]. U.S.A., Maryland (Girault, 1917). Zeuxevania splendidula Costa (an evaniid in the oothecae of Loboptera France decipiens), (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 oothecae that were parasitized by Zeuxevania, never into normal, nonparasitized oothecae. Oviposition. : — — Oviposition occurred two days after mating. The female wasp the ootheca with her antennae, selected a site, and bored stroked into the She inserted the ovipositor deeply and 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 hyperparasites in two oothecae. Sex ratio. 5 5? i cf (from 3 oothecae). U.S.A., District of Columbia, Maryland (Burks, Distribution. ootheca with her ovipositor. oviposited for 10-30 minutes. — — — ^ 1952). Tetrastichus australasiae Gahan Natural host. —Periplaneta australasiae, Tetrastichus hagenowii Synonymy. enowii Sumatra (Gahan, 1923). (Ratzeburg) —Entedon hagenowii Ratzeburg, Blattotetrastichus hag- (Ratzeburg) [Burks, 1943]. Tetrastichodes asthenogmus compared J. Kerrich (personal communication, 1957) the type of Tetrastichodes asthenogmus Waterston with authentically determined material of Tetrastichus hagcnozvii and concluded that T. Waterston. G. asthenogmus is only a weakly developed specimen of T. hagenozcni. SMITHSONIAN MISCELLANEOUS COLLECTIONS 250 He Stated, "The VOL. I4I 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 it was and mounted on a series of ten microscope slides. No doubt development that caused Waterston to describe the this faint species in Tetrastichodes, a segregate that has since been recognized by Dr. Burks (Proc. U. nerically distinct Natural hosts. S. Nat. Mus., 1943) as being not truly ge- from Tetrastichus." —Blatta orientaHs, 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. hagenozvii in the sources cited nor in the 1915 revision of Marlatt's 1902 paper cited by Burks (1943) see footnote 6, p. 236.] Neostylopyga rhomhifolia, Hawaii (Pemberton, 1941) This record is based on one parasitized ootheca. We have exposed, at three ; : different times, groups of 10 to 20 oothecae of A^. rhomhifolia to many newly emerged T. hagenozvii, 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 tsetse fly pupae (Jordan, 1956), Formosa (Takahashi, 1924; Sonan, 1924); Palestine (Bodenheimer, 1930); St. Puerto Rico (Sein, 1923; Plank, 1947, 1950; Wolcott, 1951) Hawaii (Schmidt, 1937) Croix, Virgin Islands (Beatty, 1944) ; ; ; U.S.A.: Missouri (Ran, 1940a); Ohio (Edmunds, 1955); Florida (parasitized oothecae 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 com- ; Usman, Trinidad and Saudi Arabia (Cameron, 1955). Westwood (1839) stated that 70 parasites belonging to the genus Eiilophns munication, 1955). Fiji (Lever, 1943) 1949) ; India (Mani, 1936; emerged from an ootheca of P. americana Burks (personal communication, 1955) stated collected that the on shipboard. wasp was prob- ably T. hagenowii. India (Usman, Periplaneta ausfralasiae, Austrnlia (Shaw, 1925) Formosa (Sonan, Saudi Arabia, Trinidad (Cameron, 1955) ; 1949) 1924). ; ; BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 25I Periplaneta brunnea, U.S.A., Florida (parasitized oothecae were 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, collected near Orlando, ; 1 91 4) Taken on an : ootheca. "Domestic cockroaches," U.S.A., Louisiana (Girault, 191 7). "Roach egg cases," Panama Canal Zone (Rau, 1933). Evania sp., Hawaii (Ashmead, 1901 Perkins, 1913) Guam ; (Fullaway, 1912) Fiji ; (Marlatt, 1902, 1915). Experimental hosts. (Lever, —Blatla 1946) orientalis, ; Europe, Cuba, Florida ; 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 1954b) : (Roth and Willis, 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 ootheca was obtained from a cage covered with screen too fine to permit entry of a larger 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 acciparasite, such as dental, the eulophid happening to oviposit into an ootheca already con- taining 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 (Sein, 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 5-1 1 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). ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 252 VOL. I4I In Hawaii, Severin and Severin (191 5) caught 571 T. hagenozvii in 10 kerosene traps that were set up to sample populations of Mediter- ranean fruitfly. Apparently the parasite is attracted by the odor of kerosene. Oviposition. —The female wasp explores the surface of the ootheca with vibrating antennae (Edmunds, 1955). She bends her abdomen ventrad and repeatedly touches the surface of the ootheca with her valvae ; when she finds an acceptable oviposition site, the wasp un- sheathes her ovipositor and bores through the wall of the ootheca (Roth and Willis, 1954b). The wasp deposited her eggs in 2-5 minutes (Edmunds, 1955). Wasps oviposited (pi. 34, C) into young or old eggs of P. americana (Roth and Willis, 1954b). A single wasp more than one ootheca and more than one wasp oviposited same ootheca (Roth and Willis, 1954b; Edmunds, 1955). We found freshly laid wasp eggs in 34 empty but previously parasitized oothecae from which the wasps had emerged (Roth and Willis, parasitized into the 1954b). Development. —In Periplaneta americana: Development com- is 2)^ 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 6o°-65° F. (Cameron, pleted in an average of ; ; ; We found that 31-60 days at 70° -80° F. (Edmunds, 1955). the wasps completed development in 23-56 days at about 85° F., but 1955) ; number of wasps in the ootheca the number of wasps (up to an average of about 70 wasps per the period depended on the larger the ; ootheca), the shorter the time required to complete development. Wasps in oothecae 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 ootheca, 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 ootheca 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 ootheca (Usman, 1949; Roth and ; Willis, 1954b). Number — In Blatta orientalis: In the of offspring per female. were left with each of 25 female wasps for their laboratory, 5 oothecae BIOTIC ASSOCIATIONS OF COCKROACHES of the lifespans; entire —ROTH & WILLIS 253 125 oothecae, 32 were parasitized. The average number of offspring per female was 66 (range 5-164) (Roth and WilHs, 1954b). In Periplaneta americana: Each of 206 oothecae the average was exposed to a single female wasp for 24 hours number of offspring per female was 103 (range 50-139). Five oothecae were left with each of 38 females for their entire lifespans ; of the 190 oothecae, 81 were parasitized. The average number of off- spring per female was 94 (range 45-168 [from original data]) (Roth and Willis, 1954b). Number of parasites per ootheca. laboratory, 3 oothecae that — In Eurycotis floridana: had been exposed to In the 20 female wasps yielded an average of 648 parasites (range 606 [from original data] 685) (Roth and Willis, 1954b). In Neostylopyga rhomhifolia: ootheca yielded "jt, parasites (Pemberton, 1941). In Parcohlatta to One sp. Two : oothecae yielded an average of 100 parasites (Edmunds, 1953a). In Periplaneta americana: 100 parasites per ootheca (Sein, 140 (Schmidt, 1937) 7-38, average 33 25 (Rau, 1940a) (Usman, 1949) 71 (Wolcott, 1951) 4 oothecae exposed to 20 female wasps yielded an average of 204 wasps (range 164 [from 1923) ; ; ; ; ; original data] to 261) (Roth and Willis, 1954b) ; average of 30-40 (Cameron, 1955) 39 oothecae yielded an average of 93 parasites (range 12-187) (Edmunds, 1955). In Periplaneta australasiae : Oothecae yielded an average of 40-50 adult parasites (Cameron, 1955) ; ; about 50 (Shaw, 1925). Sex 1955) ratio. ; — 3 2-8 ??: J* (Usman, 1949); 4 $$:i ^ (Cameron, (Roth and Wilhs, 1954b) 1.2 ??: i (^ (Ed- $$:i I c^ ; munds, 1955). Parthenogenesis exists; the unfertilized eggs produced only males (Roth and WilHs, 1954b; Edmunds, 1955). Distribution. — Probably worldwide. Central and South America mosa ; Europe ; Eastern and southern U.S.A. Arabia ; Africa ; India ; For- Hawaii. ; Tetrastichus periplanetae Crawford Natural hosts. 1910) ; —Periplaneta americana, Union of South Africa collected in Mozambique (Crawford, (parasites reared from oothecae Durban, Natal, by the City Health Department) were : The by Burks (personal communication, 1956). Jamaica (Gowdey, 1925) Reunion Island (Bordage, 1913). "Domestic cockroach," Puerto Rico (Wolcott, 1951). parasites identified ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 254 Tetrastichus Taxonomy. sp. VOL. I4I I — Burks (personal communication, 1956) stated that species (specimens of which are in the U.S. National this Museum) is very close to T. hagenowii. Natural hosts. —Periplancta Union of South Africa aniericana, from oothecae City Health Department [Burks, (parasites reared collected in Durban, Natal, by the c, 1956]). p. Periplaneta australasiae, Manila, Philippine Islands (Burks, p. c, 1956). Tetrastichus Synonymy. sp. II — Because of the war, Cros this insect specifically. He designated it (1942) could not determine 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 steel-blue color. Natural host. havior. apparently not T. hagenozvii because of —Blatta — Mating began orientalis, its brilliant Adult beemerged from an ootheca. up to 5 days in summer and up Algeria (Cros, 1942) : as soon as wasps Males mated repeatedly. Adults lived There were up to four generations per year to 12 days in fall. laboratory, Oviposition. —Wasps in the oviposited into oothecae 6, 22, 40, and 43 days old, and the parasites developed successfully. More than one female oviposited into the same ootheca. Oviposition was of long duration. Development. From tgg to eclosion took an average of 34 days in summer (range: 30-38 days, 5 oothecae), and an average of — 6y days in fall (range: 58-73 days, 3 oothecae). An average of 55 over 130 10-20 55 i c^. parasites developed per ootheca (range 21-105, 5 oothecae) wasps emerged from a sixth ootheca. Sex ratio. — ; : HOST SELECTION BY EGG PARASITES The nature of the oviposition stimulus (i) for the wasp parasites of is unknown. Edmunds (1954) noted that Prosevania punctata showed more interest in oothecae that had been cemented to cockroach eggs the substrate than in clean oothecae that had simply been dropped. Cros (1942) experimented with two females of P. punctata to see if 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 oothecae. Cros suggested that the wasps were the wasps could find oothecae that misled by the odor left in the jar by the cockroaches. possible that odor helps the wasp find the host ootheca. It is quite BIOTIC ASSOCIATIONS OF COCKROACHES The extent of host selection varies species will oviposit into the —ROTH among .^t WILLIS these parasites ; 255 some eggs of more than one species of cock- show some degree of host specificity. Positive selecby certain parasites appears in correlative data on pages 235 to 254. There is a small investigators different from of certain hosts by some of nonacceptance shows that body of data would not parasitize merceti Comperia example, For these wasps. roach, but others tion of specific hosts eggs of Blatta orientalis or Periplaneta americana in the laboratory (unpublished data, 1957) exposed a soft (Lawson, 1954a). from Eurycotis fioridana, to C. merceti; no recently removed ootheca, We wasps developed ; similar negative results with C. merceti we had and We (1954b) could not induce Tetrastichus germanica, B. vaga, or ParcoBlattella of eggs parasitize hagenozvii to experiments, T. hagenozvii In our laboratory. in the hlatta virginica ellect ilium, but the wasp eggs either sup Supella of oviposited into eggs oothecae of B. germanica. failed to hatch, or if they hatched, the larvae died before completing Neither would T. hagenozvii parasitize eggs of N. (Roth and Willis, unpublished data, 1957). Anastatus rhomhifolia tenuipes would not parasitize the eggs of Latihlattella lucifrons Hebard, Periplaneta americana, B. germanica, or B. vaga (Flock, 1941). development. Anastatus floridanus would not oviposit into eggs ol S. supellectilium and only rarely into eggs of P. americana or B. orientalis (Roth and in the laboratory, this wasp could not be maintained Willis, 1954a) ; 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. /'Mwc/ato to oviposit into a mantid ootheca, 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 Leclercq (personal comis primitive (Leclercq, 1954) munication, 1955) stated that this habit is always associated with the conservation of a number of structures considered as archaic from a cockroaches ; purely morphological point of view. The (e.g., records of wasps of the genus Astata capturing cockroaches Sickmann, 1893 ; St. Fargeau in Sharp, 1899) "all trace back was a personal comKrombein, of the predator" (K. V. to a questionable record by Lepeletier (1841) which probably misidentification (1866) suggested that the braconid Paxylomma huccata Breb., which he found frequenting cockroach munication, 1956). Marshall ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 256 VOL. I4I runs in Pembrokeshire, was parasitic on Ectohius nigripes Stephens however, this wasp undoubtedly parasitic on ants, probably on ant is larvae (Donisthorpe and Wilkinson, 1930). The wasps known that are 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 Pompilus sp. — Cockroach, Nyasaland (Lamborn in Poulton, host. The wasp was collected leading a nymph of the cockroach by antenna. The cockroach was in a stupefied state, and its antennae Natural 1926) its (Bingham, 1900). were : bitten of¥ to about half their length. Salius verticalis Smith Natural hosts. — Cockroaches, India Family The which species of AMPULICIDAE Ampulex do to lay their eggs but (Bingham, 1900). make not appear to special nests in drag their prey to any convenient hole, or crack in the ground (Arnold, 1928). Although many species of Am- pulex have been described, the prey of only a small number of species have been discovered, but the known prey are Ampulex amoena Synonymy. all cockroaches. Stal —Ampulex novarae Saussure [Krombein, personal com—Periplaneta americana and Periplaneta munication, 1957]. Natural hosts. aiistralasiae, The wasp both as small nymphs, Formosa (Sonan, 1924, 1927) 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, Japzn Experimental hosts. riba, (Kamo, 1957; Kohriba, 1957). picea, Japan (Kamo, 1957; Koh- —Periplaneta 1957). Kamo (1957) observed that in the sucked juices from wounds they made field both males and females in the stems of Clerodendron BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 257 trichotommn 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. many In the laboratory as with a wasp egg, were stored in as three cockroaches, each artificial Kohriba nests per day. (1957) observed similar behavior in the laboratory and made these additional notes. The paralyzed cockroach could move its legs and was by the wasp which seized led to the nest its The egg antennae. hatched in 2 days, and after sucking up body fluid for 2 days the Three days larva began to devour the prey. cocoon, and about one month Ampulex Natural hosts. 1925) : Nesting —Blatta The wasp tunnels in ground. it by an antenna and its Kohl assimilis lateralis, spun wasp emerged. wingless females, Iraq (Kingston, are holes in palm trees, galleries of beetles, or sites thorax and stings later the larva after spinning a female first seizes a cockroach by the edge of in the thoracic region, then seizes the pulls and leads it to the nest. its cockroach The wasp deposits her egg on the outer surface of the femur of the cockroach's midleg. The nest sting. is closed with debris The wasp larva first later the ; cockroach recovers from the feeds externally, then bores into the cock- roach and devours the internal organs. Pupation occurs inside the exoskeleton of the cockroach. Ampulex canaliculata (Say) Synonymy. —Rhino caniculatus. —Ischnoptera U.S.A. Natural psis (Krombein, 1951). sp., Lohoptcrella dimidiatipcs, Hawaii (Williams, 1928a, 1929). Parcohlatta pensylvanicaf MacNay (1954) referred to a rare hosts. 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 sphecoid swept from a pine tree. There are no positive records linking A. canaliculata with P. pensylvanica. Experimental host. —Parcohlatta souri (Williams, 1928a, 1929) : virginica, figure 6. females, U.S.A., Alis- — 258 Nesting SMITHSONIAN MISCELLANEOUS COLLECTIONS sites are in twigs (Krombein, 1951). similar to that oi A. compressa; is that oozes The VOL. I4I adult behavior the female w^asp imbibes blood 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), 6. Ampulex canaliculata attacking Parcoblatta virginica. A, Female stinging her prey, c. X4. 8. B, Wasp's c^g attached to the coxa of the mesothoracic leg of the cockroach. C, Larva of A. canaliculata (about threequarters grown) feeding on the internal organs of the host from the exterior, c. X4. (Reproduced from F. X. WilHams [1929], through the courtesy of Dr. F. X. Williams and F. A. Bianchi, Hawaiian Sugar Planters' Association.) Fig. wasp Distribution. —U.S.A.: consin, Missouri, Kansas; Connecticut south to Georgia; Ohio, Wisin open woods (Krombein, 1951). Ampulex compressa (Fabricius) (PI. 35) Synonymy. —Guepe ichneumon of Reaumur Chlorion (Ampulex) compressum. [Williams, 1929] ; BIOTIC ASSOCIATIONS OF COCKROACHES Natural hosts. —Periplaneta americana, —ROTH New & WILLIS Caledonia Reunion (Bordage, 1912). 1879) Periplaneta australasiae, Hawaii (Swezey, 1944). Periplaneta sp., India (Maxwell-Lefroy, 1909). Burma Mauritius (Reaumur, 1742) Cockroach. India (Dutt, 1912) ; (Lucas, ; ; 1897). 259 (Bingham, — Experimental hosts. Neostylopyga rhonihifolia, Perplaneta ameriand Periplaneta australasiae, Hawaii (Williams, 1942, 1942a). Zimmerman's (1948) listings probably were taken from Williams. in Nesting sites. Holes in walls holes in banyan and fig trees Similar to that of A. ashouses in drawers and cartons. Behavior. cana, — — Bordage (1912) gives a complete description of capture of similis. prey. ; ; The female wasp cuts off part of the cockroach's antennae, legs, and wings she sticks her &gg onto the host's mesothoracic coxa. The wasp frequents houses in search of prey. Five $?, supplied with a cockroach per day, stored an average of 57 ±14 cockroaches; 8 5$ these latter wasps were not stored an average of 45 ± 3 cockroaches supplied with a cockroach per day throughout (mean values computed from Williams, 1942). This wasp will not attack Nauphoeta cinerea (Williams, 19423.) or Pycnoscehts surituimensis (Schwabe, 1950b). ; ; On one occasion, A. compressa stung Diploptera punctata, but did not Minimum 34 days, maxioviposit (Williams, 1942a). Development. weeks (Swezey, 1944)About 6 (Williams, mum 140 days 1942). average of iio±ii days (minan lived Longevity of adults. 13 $$ — — imum 31, maximum 159) ; several J'J* lived 2 months (Williams, 1942). Ampulex fasciata Jurine —Ectobins pallidus, France (Picard, 191 1, 1919) Natural host. 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 Ampulex. Adult wasp emerges by cutting open a passage through its cocoon and through the the wasp larva is similar to that of other anus of the cockroach, Ampulex ruficornis (Cameron) — Natural hosts. Cockroaches, Oriental region (Rothney in Sharp, Nesting sites are in crevices in bark. The female grasps the 1899) 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 SMITHSONIAN MISCELLANEOUS COLLECTIONS 26o species as (=A. "A. compressiventris Guerin siberica VOL. I4I 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 : Ampulex considered A. compressiventris Guerin to be name for this common African species and that sibirica, of the genus the correct described from Siberia, must be another species. However, Turner (191 2) 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, Natural hosts. 1899) : Nesting Turner is correct, is sibirica Fab. West Africa (Perkins sites are keyholes. Wasp cockroaches. if —Cockroaches, in Sharp, Enters apartments in search of cocoon protrudes from dead body of cockroach. Ampulex sonnerati Kohl Synonymy. — "La mouche bleue" of Sonnerat (Kohl, 1893). —"Kakkerlac," Philippine Islands (Sonnerat, 1776) Natural host. Nesting : The wasp seizes the cockroach many times in the "abdomen." She are ready-made crevices. sites by an antenna and stings the host 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 cornic- Berland (1925) stated that this is possibly a color variety of D. corniculus. Soyer (1947), from a study of the behavior of the wasps, uliis. believed that both D. bicolor corniculus. Krombein and D. haemorrhous are varieties of D. (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 Synonymy. (Spinola) —Dolichurus haemorrhous Costa [Schulz, 1912]. land (1925) listed D. haemorrhous separately but stated that haps a color variety of D. corniculus. it is Berper- BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 261 — Natural hosts. Blattella germanica, France (Benoist, 1927). Denmark Ectobius lapponicus, Germany (Sickmann, 1893) ; (Nielsen, 1903) Sweden (Adlerz, 1903) ; Italy ; (Grandi, 1931, 1954); France (Benoist, 1927; Maneval, 1928). Ectobms pallidus, France (Maneval, 1932; Soyer, 1947). Ectobius panseri, France (Soyer, 1947). Ectobius sp., Italy (Grandi, 1954). Pitten (Handlirsch, 1889). H ololampra punctata, Loboptera dccipiens, 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 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 but recovers sufficiently from the sting so that (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 Natural hosts. —"Small Blattidae," India Dolichurus greenei Natural host. 1955) • —Parcoblatta Nesting third-instar Turner sites nymph. (Turner, 1917). Rohwer U.S.A., Virginia (Krombein, 1951, are under leaf litter. The prey was a paralyzed sp., Distribution. — Ontario. U.S.A. from Canadian border south to Florida in coastal States (Krombein, 1951). Dolichurus ignitus Sm. Natural hosts. nold, 1928) : — Cockroaches, Natal and Southern Rhodesia (Ar- The wasp is "usually seen running up and down the , SMITHSONIAN MISCELLANEOUS COLLECTIONS 262 VOL. I4I trunks of trees searching for small cockroaches in the crevices of the bark." Dolichurus stantoni (Ashmead) Natural 1931 Jiosts. —Allacta nymphs, Hawaii (Williams similis, et al., Zimmerman, 1948). ; Blattella litiiricollis, usually nymphs, Philippine Islands, Hawaii (Williams, 19 19). Cutilia soror, nymphs, Hawaii (Williams et al., 1931 ; Zimmerman, 1948). "Phyllodromia" sp., Bridwell, 1920). Experimental hosts. Philippine Islands, Hawaii (Williams, 1918; —"Field cockroaches," Philippine Islands (Williams, 1944). Nesting lava. site. —Readymade crevices or holes ground porosity —The wasp the cockroach by a cercus or in Behavior. ; in leg seizes in the thorax. She (fig. 7, A) then drags the cockroach 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.) and stings it to the nest — Dolichurus Natural hosts. —Cockroaches, sp. nymphs. South Africa (Bridwell, 1917). Adult female cockroach carrying an ootheca, France (Deleurance, 1943). Nesting site. — Plant stem, or ground possibly an old abandoned Behavior. — Bridwell noted one wasp larva in nest of Ammophile. that two cockroach nymphs before pupating; the adult emerged about months after cocoon formation. Deleurance observed the wasp close 4 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 ate variety of D. corniculus. Trirhogma caerulea Westwood Natural hosts. —Periplaneta americana and Periplaneta australasiae Formosa (Sonan, 1924) long and carries it : The wasp to a suitable place stings a nymph (bamboo pipe) about one inch for oviposition. BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 263 nymph Fig. 7.— Cockroach-hunting wasps. A, DoUchurus stanfoni leading a Wilhams of BlatteUa litnricollis to her nest, c. X4. (Reproduced from F. X. an Epilampra sp. [1919].) B, Podium haematogastrwn attaching her egg to^ while on the side of a termite mound that contains the wasp's nest, c.^X 1.6. C, Epilampra sp. parasitized by P. haematogastriim showing the wasps egg at32. (B and C reproduced from Williams tached to the right fore coxa, c. F. A. Bianchi.) [1928], through the courtesy of Dr. F. X. Williams and X SMITHSONIAN MISCELLANEOUS COLLECTIONS 264 Trirhogma Natural hosts. 1928) : As — Cockroaches, far as is VOL. I4I sp. Oriental region (Williams, knovv'n species of this genus of 1918, wasps hunt cock- roaches. Family SPHECIDAE Tachysphex blatticidus Williams Natural hosts. (Callan, 1942) wasp — Chorisoneura The wasps : itself is parasitized sp., adults, Trinidad, St. Augustine nest gregariously in sandy places. The by the mutillid Timulla (Timulla) eriphyla Mickel. Cockroaches, Trinidad (Williams, 1941a; Callan, 1950). Tachysphex coriaceus Costa Natural hosts. — Cockroaches, Natural hosts. — Graptoblatta notulata. Society Islands Italy (Beaumont, 1954). Tachysphex fanuiensis Cheesman (Cheesman, 1927, 1928). Cockroach ("except for Nesting sites. — its smaller size [it] much resembles Grap- New Caledonia (Williams, 1945). Patches of dry soil (Cheesman, 1928) toblatta notulata."), ; coarse sand — bank (Williams, 1945). BeJiavior. 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. at base of a ; ; Tachysphex Natural hosts. —Ectobius lativalvis (Thomson) lapponicus, adults, Sweden France (Maneval, 1932). 1906) Ectobius pallidus, nymphs, France (Ferton, 1894, 1901 1932; Deleurance, 1946); Italy (Grandi, 1928). (Adlerz, ; ; Maneval, Ectobius panseri, 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 — Grandi is a hole 5.5 to 8 cm. long ending in a horizontal cell. (1928) stated that the entrance to the nest descended obliquely for 5 nest BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 265 6 cm. and ended 4 cm. below the surface of the ground. Behavior. either sex, adults or nymphs, were stored in the cell (Adlerz, 1903; Grandi, 1928). The wasp laid her egg on the to —Two cockroaches, first prey brought, attaching it The cockroaches behind the front coxa. were not excitable and their antennae had not been injured. Grandi (1928) stated that the claws of the hind tarsi of the victims may amputated. The hatched larva may consume one of its be victims in four days leaving only the head, pronotum, tegmina, wings, and the urosternum. Podium abdominale (Perty) Synonymy. — Trigonopsis ahdominalis Perty [Kohl, 1902]. — Cockroaches, nymphs, Ecuador (WilHams, 1928) Natural hosts. : 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 cock- The prey roaches are stored in each nest. sult of the sting, and its antennae are Podium Natural host. Nesting One —Parcohlatta mud are sites to three cockroach are placed in tube resin. 195 1 ) Distribution. ; not immobilized as a re- Carolina Rohwer pensylvanica, nymphs (Rau, 1937) : nests of Sceliphron caementarium (Drury). nymphs the nest ; is left intact. is mud partitions mud which is coated with are stored per nest plugged with —U.S.A., New York to ; North Carolina (Murray, 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. haemaiogastrum. Podium flavipenne Lepeletier Natural —Epilampra abdomen-nigriim, British Guiana (Wil— Burrows, about 2 inches deep and are dug the ground well-drained, partly cross areas also old Podium nests are used. Behavior. —The host. liams, 1928) ticular in sheltered wasp : : Nesting len- site. in section, in ; stings the cockroach to helplessness nest where the host may roaches are stored per nest while the cockroach is still and flies recover from the sting ; the egg is with ; it back to her one or more cock- deposited behind the forecoxa outside the burrow. The nest is sealed with mud. The larva feeds on most of the cockroach and leaves only some SMITHSONIAN MISCELLANEOUS COLLECTIONS 266 VOL. I4I cell. In 153 nests examined, there [standard error computed from cited was an average of 2.2 ±0.08 heavily sclerotized portions in the cockroaches per data] apiece. Of adults. Development. cell four nests contained ; five cockroaches the 331 cockroaches in the nests, only 6 percent were —Egg hatches in about 2 days about 4 days and pupates about 2 weeks later 10-12 days later. ; ; feeds larva adult emerges about Podium haematogastrum Spinola Natural host. —Epilampra The female wasp mounds, in banks, (fig. and B) 7, Para (Williams, 1928) There was an average of is similar 1.6 cockroaches 74 nests examined. Of the 121 cockroaches lected, 28 percent were adults. Under artificial conditions, the C) per (fig. 7, : burrows into the surface of termite ground. This wasp's behavior in level to that of P. flavipenne. Brazil, sp., col- cell in cycle varied from about a month Natural —Parcoblatta life to 45 days or more. Podium luctuosum Smith host. (Pate, 1949). Distribution. —U.S.A.: virginica, New York Podium Natural hosts. 1919) ; —"Wood mound. Variable numbers with one wasp egg attached hatches in 2 days, the larva emerges 24 days Texas (Murray, 1951). (Howes, 1917, were clay column stumps, or forest trees banks termite of cockroaches were placed in the nests behind forecoxa of the last host. The egg pupates about 2 weeks later, and the adult roaches," British Guiana : Nesting sites ; ; later. Podium Natural host. New York rufipes Fabricius Brazil (Williams, 1928) nests on houses, sides of to female, U.S.A., sp. —Epilampra conferta, Brazil (Poulton, 1917) burrow contained several cockroaches of the same The : 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 and was instantly torn to bits and carried to the In the end it fell Another ant with the body of a wood roach was assisted by a rear 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) BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 267 FORMICIDAE Family From the known entomophagous habits of the lower ants (Wheeler, we wonder that there are not more records of ants feeding on 1928), Kirby and this act must occur frequently. 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 cockroaches, because to sugar in the pantry were killed destruction of cockroaches by and carried army by the off ants. The ants has been recorded by Bates (1863), Wallace (1891), Beebe (1917, 1919), Howes (1919), and Dead and mutilated specimens of Ischnoptera sp. [un- others. doubtedly Parcohlatta 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 The —Carpenter —Parcohlatta name. Natural prey. ant. U.S.A. pensylvanica, (Rau, 1940) : ants entered traps set up to capture the cockroach and carried off about a dozen adults of both sexes. Dorylus (Anomma) nigricans subsp. sjostedi Emery Natural prey. — Small cockroach, Belgian Congo (Raignier and van Boven, 1955). Dorylus Natural prey. — Small (Anomma) wilverthi Emery cockroaches, Belgian Congo (Raignier and van Boven, 1955). Dorylus Common Natural sp. — "Safari ant." prey. — Cockroaches, Africa, name. Lake Victoria (Carpenter, When the "Safari ants" were hunting, many species of cock1920) roaches were driven from hiding among dead leaves in the forest. The : cockroaches rushed about but easily them to bits. fell prey to the ants which tore : SMITHSONIAN MISCELLANEOUS COLLECTIONS 268 VOL. I4I 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). Iridomynnex humilis Mayr Common —Argentine —Cockroaches, name. Natural prey. ant. injured individuals only (Ealand, 1915)Lasius alienus Natural prey. (Forster) —Ectobius pallidus,\J.S.A., Massachusetts (Roth and Willis, 1957). Pheidole megacephala (Fabricius) Common —Big-headed —Holocompsa name. ant. Hawaii (Illingworth, 1916). Nauphoeta cinerea and Pycnoscelus surinamensis, Hawaii (Illingworth, 1914, 1942) The ants followed and killed A^. cinerea and P. surinamensis as they burrowed in moist soil and attacked and deNatural prey. fulva, : stroyed A'^. cinerea in breeding cages. VERTEBRATA Class PISCES XHI. 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 (Dumeril) (= Potamotrygon Family hystrix) PIMELODIDAE Shamdia sebae Cuvier and Valenciennes BIOTIC ASSOCIATIONS OF COCKROACHES Family —ROTH & WILLIS 269 CHARACIDAE Cyrtocharax magdalenae essequibensis (Eigenmann) (z= Cynopotamiis essequibensis) Chalceus macrolepidotus Cuvier and Valenciennes The only other records of cockroaches being eaten by fish pertain Captain to the use of cockroaches as bait.® WiUiam Owen (in Web- 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 ster, who used them for catching Lepomis pallidus, a sunfish locally cockroaches were collected at a city known In Indiana, oriental as bream, blue bream, or bluegill (Rau, 1944). dump by fishermen (Gould, Peterson (1956) states that cockroaches are satisfactory bait for bluegills, crappies, channel cat, blue heads, and large mouth black 1941). bass. AMPHIBIA^ Order CAUDATA Class Family PLETHODONTIDAE Plethodon glutinosus (Green) Natural 1953) : prey. — Cryptocercus punctulatus, U.S.A. (Honigberg, Protozoa which are normally only found in C. punctulatus were present in the intestine of the salamander indicating that cockroach had been eaten by the amphibian. ^archy (Marquis, 1931) was living in a dream world when he typed: is always something to be thankful "there for you would not think that a cockroach had much ground for optimism but as the fishing season grow more and more opens up i cheerful at the thought that nobody ever got the notion of using cockroaches for bait" s Classification of Amphibia and Reptilia follows Hegner (1936). this SMITHSONIAN MISCELLANEOUS COLLECTIONS 270 Order Family VOL. I4I SALIENTIA 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 Spix ictericus — Cockroaches, Natural prey. 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 —Giant —Epilampra Surinam toad. ahdomen-nigrum, Trinidad (Weber, Found in the stomachs of two toads. 1938) Diploptera punctata, Hawaii (Pemberton and Williams, 1938). name. Natural prey. toad, marine toad, : 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 cock: roaches. Puerto Rico (Wolcott, 1937). Bufo valliceps Wiegmann Experimental prey. —Periplaneta americana, U.S.A. (Moore, Cockroaches containing infective acanthellas of Moniliformis 1946) dubius were fed to three toads. : Family HYLIDAE Hyla cinerea (Schneider) frog. — Green Natural prey. —Ischnoptera dcropeltiformis, Periplaneta americana, Common name. tree and undetermined cockroaches, U.S.A., Georgia Cockroaches were found in 11 of 100 stomachs. Family (Haber, 1926) : RANIDAE Arthroleptis variabilis Alatschie —Cockroaches, Belgian Congo (Noble, 1924) Of 52 specimens examined, the stomach contents of 17 contained food, Natural prey. including 3 cockroaches. : BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 2/1 Hyperolius picturatus Peters — Cockroach, Belgian Congo (Noble, 1924) The stomachs of 12 of 56 specimens examined contained food, including one cockroach. Natural prey. : Leptodactylus albilabris (Giinther) — Natural prey. Cockroach, Puerto Rico (Schmidt, 1920) 25 stomachs contained a medium-sized cockroach. One : of Leptodactylus pentadactylus (Laurenti) Common Natural —"Smoky jungle frog" or "pepper frog." prey. — Cockroaches, Nicaragua (Noble, 1918) name. Cock- : roach wings were found in the stomach of a frog caught around hu- man Brazil (Valente, 1949). habitation. Leptopelis calcaratus (Boulenger) — Cockroaches, Belgian Congo (Noble, 1924) The stomachs of 35 specimens were examined of which 13 contained food, including 2 cockroaches. Natural prey. : Leptopelis rufus Reichenow Natural prey. five — Cockroaches, Belgian Congo (Noble, 1924) Fortyof 83 stomachs examined contained food, including 2 cockroaches. : Megalixalus fornasinii (Bianconi) — Cockroaches, Belgian Congo (Noble, 1924) The stomachs of 3 of 40 specimens contained food, including 2 cock- Natural prey. : roaches. Rana catesbeiana Shaw Common — Bullfrog. — Cockroaches, Puerto Rico name. Natural prey. (Derez, 1949). Rana mascareniensis Dumeril and Bibron — The Natural prey. Cockroaches, Belgian Congo (Noble, 1924) stomach contents of 138 specimens were examined, 39 of which con: tained food, including 2 cockroaches. Rana pipiens Schreber — Common name. Leopard frog. Experimental prey. Periplaneta — (Moore, americana, U.S.A. Cockroaches containing infective acanthellas of Moniliformis 1946) dubius were fed to two frogs. : SMITHSONIAN MISCELLANEOUS COLLECTIONS 272 VOL. I4I Neostylopyga rhomhifolia, U.S.A. (Dr. T. Eisner, personal communication, 1958.) Frogs Natural prey. —Blatta orientalis, which escaped from a tank U.S.A. in the cellar (Rau, consumed 1924) Frogs : quantities of this cockroach. Parcohlatta pensylvanica, U.S.A. (Frost, 1924) men One : recovered from alimentary canal of a frog, probably adult speci- Rana sp. Unidentified batrachians Experimental prey. — Blattella germanica, Main, Zoological Garden (Lederer, 1952) ferred by all : Germany, Frankfurt am These insects v^ere pre- the insect eaters in the zoo. am Periplaneta americana, Germany, Frankfurt Garden (Lederer, 1952) : Newly molted Main, Zoological were accepted individuals as food, but others were usually passed by or consumed unwillingly. REPTILIA^ Class CHELONIA Family EMYDIDAE Order Chrysemys picta (Schneider) Common 1920) : — Painted —Periplaneta name. Natural prey. The turtle. England (Lucas, 1916, australasiae, cockroach, apparently injured, fell toise house, Zoological Gardens, Regent's Park, into water in the tor- and the terrapin ate it. SAURIA GEKKONIDAE Order Family Gekko gecko (Linnaeus) Natural prey. dolid, — Cockroaches, The geckos Philippine Islands, Laguna (Villa- frequent holes in trees and underside of 1934) bark which are favorable haunts of cockroaches. : Stomach contents mostly Blattidae and "Locustidae." Hemidactylus frenatus Dumeril and Bibron Common dolid, —House —Cockroaches, name. Natural prey. lizard. Philippine Islands, Bulk of stomach contents of 22 1934) Orthoptera, mostly cockroaches. : Laguna (Villa- lizards consisted of BIOTIC ASSOCIATIONS OF COCKROACHES Sphaerodactylus Natural prey. above lizard is —ROTH 273 sp. — Cockroaches, British Guiana found & WILLIS (Beebe, 1925a) : The : The in houses. Thecadactylus sp. — Natural prey. Cockroaches, British Guiana (Beebe, 1925a) is found in houses. above hzard Undetermined geckos Natural prey. 1906) : The — Cockroaches, AustraHa, Flinders River (Froggatt, and hunted cockroaches lizard lived in the walls of the hut upon the roof at night. Arno Atoll (Usinger and Family La Rivers, 1953). IGUANIDAE Anolis carolinensis Voigt Experimental prey. — Diploptera punctata, U.S.A. (Eisner, 1958), Anolis cristatellus Dumeril and Bibron Natural prey. — Blattella Cariblatta sp., delicatula, wheeleri, Periplaneta americana, Periplaneta australasiae, The last-named cockRehn and Hebard, the females of from flagellata. Rehn and Hebard ploce flagellata, Puerto Rico (Wolcott, 1924) roach may have been which are hard to 6*. Epilampra and Sym- : ruficollis distinguish (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. Of 100 stomachs exCockroaches, Puerto Rico (Schmidt, 1920) amined, 16 contained Orthoptera, including cockroaches. Puerto Rico : (Wolcott, 1924) : One hundred A. cristatellus had eaten 8 cock- roaches, 4.14 percent of the total food, or 25 percent of the food for 8 lizards. Anolis pulchellus Dumeril and Bibron Natural prey. — Cockroaches, Puerto Rico (Wolcott, 1924) : Two small cockroaches found in 50 lizards examined. Anolis equestris Experimental prey. — Neostylopyga personal communication, 1958.) Merrem rhombifolia, U.S.A. (Eisner, SMITHSONIAN MISCELLANEOUS COLLECTIONS 274 Anolis grahami VOL. I4I Garman — Natural prey. Pej'iplaneta spp. and Blattidae, Bermuda (SimStomachs of 176 lizards yielded 6 cockroaches. monds, 1958) : Anolis leachi Dumeril and Bibron prey. —Periplaneta Natural monds, 1958) and spp. Blattidae, Bermuda (Sim- Stomachs of 46 lizards yielded 31 cockroaches. : Anolis sagrei Cocteau — Natural and experimental prey. Pycnoscelus surinamensis, Cuba This species was eaten both in captivity and in (Darlington, 1938) nature. The : lizard most readily ate Pycnoscelus surinamensis is soft, immature cockroaches. probably a staple food of the lizard in nature, as Darlington observed wild lizards catch the nymphs. Anolis stratulus Cope Natural prey. — Aglaopteryx fades, Puerto Rico (Wolcott, 1924) examined. One cockroach was found in 50 lizards One of 25 stomachs Cockroach, Puerto Rico (Schmidt, 1920) : : contained a cockroach. Anolis Natural prey. — "Wood The above lizard is sp. roaches," British Guiana (Beebe, 1925a) : arboreal on foliage in low jungle. Family SCINCIDAE Leiolopisma laterale Say Common — Brown skink. —Woodroaches, U.S.A., name. Louisiana (Slater, 1949) : Analysis of stomach contents of 84 adult skinks showed that nymphal and adult woodroaches comprised the majority of Orthoptera. Natural prey. Tropidophorus grayi Gitnther Common dolid, 1934) — Spiny —Cockroaches, name. Natural prey. : Food of lizard. Philippine Islands, this species was mostly Laguna (Villa- Blattidae. Unidentified skinks Natural prey. 1953)- —Cockroaches, Arno Atoll (Usinger and La Rivers, BIOTIC ASSOCIATIONS OF COCKROACHES Family —ROTH — Cockroaches, Australia 1955) -^ general statement 2/5 AGAMIDAE Experimental prey? • & WILLIS was made (Lee and Mackerras, that in captivity Agamidae were observed feeding avidly on cockroaches and other insects. Three agamids studied by these workers were Amphibolurus harhatus (Gray), Physignathiis lesueurii Gray, and Chlamydosaurus kingii Gray. Family CHAMAELEONTIDAE Chamaeleon chamaeleon (Linnaeus) and Chamaeleon oustaleti Mocquard Experimental prey. — Cockroaches, Large cockroaches were fed Amsterdam (Portielje, 1914) to these lizards in the reptile : house of Artis. Common Family TEIIDAE Ameiva exsul — Iguana, ground — Cockroach (nymph), Epilampra wheeleri, and Peri- name. Natural prey. Cope lizard. Stomach contents planeta americana, Puerto Rico (Wolcott, 1924) 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. — The Natural prey. Cockroaches, British Guiana (Beebe, 1925a) above lizard is terrestrial and found near clearings. The stomach contents of 18 out of 40 reptiles contained cockroach remains. : Cnemidophorus Natural prey. is —Cockroaches, British sp. Guiana (Beebe, 1925a) This The stomach contents of : a terrestrial lizard found near clearings. 4 out of 40 lizards contained cockroaches. Unidentified Natural prey. lizards — Cockroaches, West Indies (H., 1800). — Blatta U.S.A. (Rau, 1924) Experimental prey. called the predator a orientalis, common gray lizard. : Rau SMITHSONIAN MISCELLANEOUS COLLECTIONS 276 VOL. I4I Periplaneta americana, Germany, Frankfurt am Main, Zoological Newly molted cockroaches were accepted Garden (Lederer, 1952) as food, but others were usually passed by or consumed unwillingly. : Order Family SERPENTES COLUBRIDAE Heterodon platyrhinos Latreille Synonymy. —Heterodon sonal communication, Common name. Experimental contortrix [Dr. Doris M. Cochran, per- 1957]. — Hog-nosed snake. prey. —Periplaneta americana, U.S.A. (Moore, Cockroaches containing infective acanthellas of Moniliformis 1946) dubius were fed to one snake. : Garter Snake Experimental prey. —Blatta orientalis, Class The cockroach is U.S.A. (Rau, 1924). AVES always wrong when arguing with a chicken. Spanish proverb (Hartnack, 1939) Arboreal cockroaches hidden in and under bark are likely to sectivorous birds undoubtedly the few records would scientific common name. Where possible we have names for those birds whose common names are recognizably specific. Wetmore consume many more cockroaches than Most of the records we have located indicate. identify the birds at least by given the of much more be encountered by birds than by other predators, and in- We have followed the systematic classification (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) may stated that in Japan natural enemies of cockroaches be found in the Galli formes, Strigi formes. Passed formes, and Piciformes. Although Lederer (1952) successfully fed newly molted Periplaneta americana to insectivorous birds in the Zoological Garden, Frankfurt birds. The roaches. am Main, Blattella germanica were preferred by these following records are of specific birds feeding on cock- BIOTIC ASSOCIATIONS OF Order COCKROACHES —ROTH & WILLIS 277 ANSERIFORMES ANATIDAE Family Domestic duck Natural 1926) : prey. —Pycnoscelus The ducks became surinamensis, Australia infected with Hansen's eye (Fielding worm of which P. surinamensis is the only known intermediate host. "All kinds of poultry feed Cockroach, Bermuda (Jones, 1859) greedily upon the cockroach; tame ducks spending entire moonlight : nights in their capture." GALLIFORMES Order Family PHASIANIDAE Bambusicola thoracica Temminck Common — Kojukei. — Cockroaches, Japan name. Natural prey. (Asano, 1937). Gallus sp. Common Jungle fowl. ——Pcriplaneta name. Natural prey. australasiae, Hawaii (Schwartz and Schwartz, 1949). Phasianus calchicus karpowi Buturlin Common name. —Korean pheasants. — germanica Blattella and Periplaneta picea, Japan (Asano, 1937) Adults of these cockroaches were devoured at once when they were fed with the heads cut off. Experimental prey. : Phasianus sp. — Pheasant. — unidentified (below Cojnmon name. Natural prey. Blattidae, Cutila soror (below i i percent of the diet), percent of the diet), Diploptera punctata (above 6 percent of the diet), and Pycnoscelus surinamensis (6 percent of the Hawaii (Schwartz and Schwartz, 1949). diet), Coturnix coturnix japonica Common tipes, —Japanese — Blattidae (unidentified) name. Natural prey. (Temminck and Schlegel) quail. and Lohopterella dimidiaHawaii (Schwartz and Schwartz, 1949). SMITHSONIAN MISCELLANEOUS COLLECTIONS 278 VOL. I4I 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 oi B. craniifer which had dropped to the ground from among stacked coffins in an undertaker's shack. The chickens ate cockBlatta oricntalis, U.S.A. (Rau, 1924) roaches that were caught in traps. Hebardina concinna, Japan (Asano, 1937) Experimental feeding to white Leghorn chickens. : : Surinam (Stage, 1947) Several cockwas being sprayed with DDT, and were eaten by chickens. Although some chickens had DDT tremors the next day, all appeared normal two days later. Formosa, Pynoscelus surinamensis, Australia (Fielding, 1926) Periplaneta americana, roaches ran ofif : the floor of a house, which ; (Kobayashi, 1927) Australia, experimental feeding (Fielding, 1927, 1928) U.S.A., Florida, experimental feed- experimental feeding ; ; Hawaii (Illing(Sanders, 1928) Antigua (Hutson, 1943) worth, 1931; Schwabe, 1949, 1950a, 1950b). This cockroach is the ing ; ; intermediate host of Oxyspirura mansoni, the chicken eye 1947) : Guadeloupe worm. Africa 1654) "Poultry" ate cockroaches which had been killed by Cockroaches, (Dutertre, ; fluoride. Hawaii (Zimmerman, 1948). Natural prey. — Cockroaches, British Guiana sodium (Moiser, DDT and Partridge (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, Turkeys were found heavily infected with Hanson's eye 1943) worm of which P. surinamensis is the only known intermediate host. These turkeys therefore were presumed to have fed on this cockroach. : COLUMBIFORMES Family COLUMBIDAE Order Streptopelia chinensis —Chinese dove. —Pycnoscelus surinamensis, Common name. Natural prey. 1950b). (Scopoli) Hawaii (Schwabe, BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 279 Pigeon Experimental prey. ing, 1927) ; —Pycnoscelus surinamensis, Australia (Field- U.S.A., Florida (Sanders, 1928). Order STRIGIFORMES Family STRIGIDAE Gymnasio nudipes (Daudin) Common —Bare-legged owl. —Epilampra Puerto name. Rico (Wetmore, 1916) sp., stomach of a wild-caught owl. Cockroaches, Puerto Rico (Wetmore, 191 6) These insects were found in stomachs of five owls. Natural prey. One specimen : identified in : CORACIFORMES Order Family TODIDAE Todus mexicanus Lesson Common — Porto Rican tody. —Plectoptera poeyif, Puerto Rico name. Natural prey. The stomachs of 89 birds were examined above cockroach. According to ; (Wetmore, 1916) : a single bird had eaten the Wolcott (1950) P. poeyi could be Plectoptera dorsalis, P. rhabdota, or P. infulata. Family Tockus Synonymy. —Lophoceriis BUCEROTIDAE birostris birostris (Scopoli) [Dr. H. Friedmann, personal communication, 1957]. Common name. Common gray hornbill. Natural prey. — —Cockroaches, India, Central Provinces (D'Abreu, 1920). Order PICIFORMES Family PICIDAE Dendrocopos mahrattensis (Latham) Synonymy. —Liopicus maJirattensis [Friedmann, 1957]. Common name. — Yellow-fronted pied woodpecker. Natural prey. — Cockroaches, Central Provinces (D'Abreu, p. c. India, 1920). : SMITHSONIAN MISCELLANEOUS COLLECTIONS 280 (Daudin) Melanerpes portoricensis — Puerto Rican woodpecker. Natural prey. —Pycnosceliis surinamensis, Puerto Rico Common 1916) : VOL. I4I name. One specimen found (Wetmore, 59 bird stomachs examined. in PASSERIFORMES Order FORMICARIIDAE Family Gymnopithys leucaspis (Sclater) Common —Bicolored —Cockroaches, Panama Canal Zone (Johnson, 1954) name. Natural prey. antbird. : 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 Icterus Common ORIOLIDAE (Bryant) portoricensis — Puerto Rican prey. — Cockroaches, Puerto name. oriole. Natural Cockroaches and oothecae found (Wetmore, Rico 1916) in the birds' stomachs. CORVIDAE Family Aphelocoma coerulesens (Bosc) Common name. —Florida —Pycnoscclus jay. Experimental prey. 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 rhomhifolia, and Periplaneta E. americana, U.S.A. (Eisner, personal communication, 1958) — : floridana was only eaten after the odor of 2-hexenal, leased by the insect on being attacked by the bird, Family which was had re- dissipated. PARADISEIDAE Paradisea papuana Bechstein Experimental prey. lace, 1869) : Two — Cockroaches, Malaya and on shipboard (Wal- 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 provision for the voyage home." me several biscuit-tins full, as COCKROACHES BIOTIC ASSOCIATIONS OF Family —ROTH & WILLIS 281 TROGLODYTIDAE Troglodytes aedon Vieillot — House wren. — Cockroaches, Common name. Natural prey. 1955) The wren : U.S.A. (Greenewalt and Jones, carried three small cockroaches to nestlings; the records probably represent incidental captures. Troglodytes audax Tschudi Common —Cucarachero. — Cockroach (called Chilicabra by Peruvian Indians), name. Natural prey. Peru (Tschudi, 1847) : The bird seized the cockroach and bit ofif its head then devoured the body discarding the wings. LANIIDAE Family Lanius ludovicianus Linnaeus Common name. — Loggerhead shrike. prey. —Pycnoscelus Experimental surinamensis , U.S.A., Florida (Sanders, 1928). Family STURNIDAE Acridotheres tristis Bonnaterre and Vieillot Common — Myna, mynah. — Cockroaches, Hawaii, name. Natural prey. Lanai (Illingworth, 1928) many cockroaches any: lUingworth reported that he had never seen as where else in Hawaii. The birds followed tractors that were destroy- ing cactus and kept close to the chain that turned over the stumps. The following species were collected : Allacta smiilis, Blattella ger- manica, Cutilia soror, Diploptera punctata, Leucophaea maderae, Periplaneta americana, Pcriplaneta australasiae , Pycnoscelus surinamensis. Illingworth did not state whether the birds ate all these species in- discriminately. Pycnoscelus surinamensis, Hawaii (Williams 1950b) : food for many places mynah birds. In this species Family Schwabe, et al., 1931 forms an important fledgling ; VIREONIDAE Vireo latimeri Baird Common name. —Latimer's —Periplaneta Natural prey. vireo. sp., Puerto Rico (Wetmore, 1916) Cockroaches were found in one of 43 stomachs examined. : SMITHSONIAN MISCELLANEOUS COLLECTIONS 282 Family VOL. I4I ICTERIDAE Agelaius xanthomus (Sclater) Common —Yellow-shouldered blackbird. — name. Natural prey. Cockroaches, Puerto Rico (Wetmore, 191 6) thecae and remains of adult cockroaches found in stomachs. Dolichonyx oryzivorus Common — Bobolink. prey. —Pycnoscelus : 06- (Linnaeus) name. Experimental surinamensis, U.S.A., Florida (Sanders, 1928). Holoquiscalus brachypterus (Cassin) Common — Puerto Rican blackbird. — Cockroaches, Puerto Rico name. Natural prey. few eggs (oothecae) of cockroaches (Wetmore, 1916) : A in stomachs. Black bird prey. —Pycnoscelus Experimental surinamensis, U.S.A., Florida (Sanders, 1928). Family FRINGILLIDAE Passer domesticus (Linnaeus) Common —English sparrow. —Pycnoscelus surinamensis, name. Natural prey. Schwabe, 1950b) Remains of 193 the stomach of the sparrow. 1 : ; this Hawaii (Illingworth, cockroach were found in Sparrow Natural prey. —Periplaneta amcricana, England (Lucas, 1908, 1920). Cockroaches, Japan (Asano, 1937). Tiaris bicolor omissa Common (Jardine) — Carib —Cockroaches, Puerto Rico name. grassquit. (Wetmore, 1916) Anione bird had was found in of stomachs examined mal food 72 5 cockroaches anions: other insects. two eaten Natural prey. : ; COCKROACHES BIOTIC ASSOCIATIONS OF —ROTH & WILLIS 283 MAMMALIA" Class MARSUPIALIA Order DIDELPHIDAE Family Monodelphis sp. — Natural prey. Cockroaches, British Guiana (Beebe, 1925a) The above opossum is nocturnal and arboreal but nests on the ground in : grass. INSECTIVORA Family ERINACEIDAE Order Erinaceus europaeus Linnaeus Experimental prey. — Blattella germanica, France (Brumpt and Urbain, 1938) Two hedgehogs were fed cockroaches infested with Prosthenorchis elcgans and P. spirilla. : Erinaceus —Hedgehog. —Cockroaches, sp. Common name. Natural prey. England (Samouelle, 1841 ; Cowan, 1865). CHIROPTERA Order Family MOLOSSIDAE Molossus Natural prey. above bat is a sp. — Cockroaches, British Guiana common house Order Family (Beebe, 1925a) : The bat of the area. PRIMATES LEMURIDAE Lemur coronatus Gray Natural prey. 1938) : — Blattella germanica, France (Brumpt and Urbain, infested naturally with Pros- The monkey apparently became thenorchis spirula for which B. germanica was the intermediate host in the monkey house. Lemur fulvus E. Geoffrey Natural prey. — 1938) : Blattella germanica, See comment under 1° Classification of Mammalia Lemur France (Brumpt and Urbain, coronatus. follows Simpson (1945). SMITHSONIAN MISCELLANEOUS COLLECTIONS 284 Family VOL. I4I LORISIDAE Loris tardigradus (Linnaeus) Synonymy. —Lemur tardigradus [Dr. D. H. Johnson, personal com- munication, 1957]. Natural prey. Cockroaches, on board ship (Cowan, 1865). — Perodicticus potto (P. L. S. Miiller) Common — Potto. — name. Natural prey. Blattella germanica, France (Brumpt and Urbain, See comment under Lemur Experimental prey. Blattidae, East Africa (Pitman, 1931) 1938) coronatus. : — sexes of the potto ate freely of Family all Both TARSI IDAE Tarsius Experimental prey. : types of cockroaches. sp. —Cockroaches, Family Borneo (Shelford, 1916). CEBIDAE Aotes zonalis Goldman Synonymy. — Aotus [Simpson, 1945]. Common name. — Canal Zone night monkey. Natural prey. —Leucophaea madcrac, Panama son, 1939) (Foster and John- Captive monkeys became naturally infested with Proto- : spirura muricola by eating cockroaches that contained infective larvae of the worm. Ateles dariensis Goldman Common son, 1939) —Darien black spider monkey. —Leucophaea maderac, Panama name. Natural prey. (Foster and John- See comment under Aotes zonalis. : Cebus apella (Linnaeus) Natural prey. — 1938) : Blattella germanica, See comment under Lemur France (Brumpt and Urbain, coronatus. Cebus capucinus (Linnaeus) — — Common name. White- faced monkey. Natural prey. Leucophaea maderae, Panama (Foster and JohnFavorite item of food in the laboratory. See comment son, 1939) : under Aotes zonalis. BIOTIC ASSOCIATIONS OF COCKROACHES —ROTPI & WILLIS 285 Saimiri sciurea Linnaeus Natural prey. — 1938) : Blattella germanica, France (Brumpt and Urbain, See comment under Callithrix chrysoleucos. Family CALLITHRICIDAE Callithrix chrysoleucos (Natterer) Synonymy. — Callithrix chrysolcvea [Johnson, personal communica- tion, 1957]. Natural prey. 1938) : — Blattella germanica, France (Brumpt and Urbain, The monkey apparently became infested naturally with Pros- thenorchis elegans for which B. germanica was the intermediate host monkey in the house. (Linnaeus) Callithrix jacchus — Synonymy. Simla jacch us. Natural prey. Blattella germanica, France (Brumpt and Urbain, See comment under Lemur coronatus. 1938) — : Cockroaches, on board ship (Neill, 1829; also cited by Samouelle, "It was quite amusing to see it at its meal. 84 1, and Cowan, 1865) 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 1 : When ; 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 cockpulled out the viscera and cast roaches he eat[s] without such fastidious nicety." Leontocebus oedipus : — Blattella germanica, France (Brumpt and Urbain, See comment under Callithrix chrysoleucos. Natural prey. 1938) (Linnaeus) Leontocebus rosalia —Midas Natural prey. — Synonymy. rosalia (Linnaeus) [Simpson, 1945]. France (Brumpt and Urbain, Blattella germanica, 1938) : See comments under Lemur coronatus and Callithrix chrys- oleucos. Leontocebus ursulus (E. Geoffrey) Natural prey. 1938) : — Blattella germanica, France (Brumpt and Urbain, See comments under Callithrix chrysoleucos. SMITHSONIAN MISCELLANEOUS COLLECTIONS 286 Family VOL. I4I CERCOPITHECIDAE Cercopithecus sp. — Cockroaches, East Africa (Carpenter, 1921, and usually ignored cockroaches ofifered to it. In one experiment the monkey had to be deprived of food before it would eat the cockroach. Experimental prey. 1925) : The monkey rarely tasted Macaca mulatta (Zimmermann) —Macaca rhesus 1957]Experimental prey. — Synonymy. [Johnson, personal communication, Blattella germanica, bain, 1938, 1938a) : The macaque was France (Brumpt and Ur- fed cockroaches infested with Prosthenorchis elcgans and P. spirula. Macaca sylvanus (Linnaeus) Synonymy. —Inuus sylvanus [Simpson, 1945]. Common name. — Macaque. germanica, France (Brumpt and Urbain, Natural prey. — Blattella 1938) : See comment under Lemur coronatus. Papio papio (Desmarest) prey. — Experimental germanica, France (Brumpt and This baboon was fed cockroaches infested Blattella Urbain, 1938, 1938a) with Prosthenorchis elegans and P. spirula. : Family PONGIDAE Pan Common sp. — Chimpanzee. — germanica, Netherlands name. Blattella (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. Natural prey. : ; Family Homo Natural prey. —Oothecae HOMINIDAE sapiens Linnaeus of Blatta orient alis and Neostylopyga rhomhifolia, Thailand (Bristowe, 1932). Periplaneta americana, Formosa (Takahashi, 1924). BIOTIC ASSOCIATIONS OF COCKROACHES Periplaneta americana —ROTH and Periplaneta & WILLIS australasiae, 28/ Australia, China, and Japan (Bodenheimer, 1951). 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 Cockroaches, Roth and Willis, 1957a). EDENTATA Order Family DASYPODIDAE Dasypus novemcinctus Linnaeus Synonymy. — Tatu novemcinctum [Johnson, personal communica1958]. Natural prey. —Ischnoptera deropeltiformis, Texas (Hebard, tion, A specimen of this cockroach in the U. 191 7) was taken from the stomach of the armadillo. : S. National Museum RODENTIA Family MURIDAE Order 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, ; 193 1) Rats infested with Protospirura bonnei presumably ate this cock- roach which is the intermediate host of the worm. Periplaneta americana, Brazil (Magalhaes, 1898) stomachs of brown in the rats. Denmark : : Remains found (Fibiger and Ditlevsen, This cockroach was found to be the intermediate host of 1914) Gongylonema neoplasticum, a parasite of rats. : Rattus rattus (Linnaeus) Natural prey. Ditlevsen, —Periplaneta 191 4) : americana, Denmark (Fibiger and See comment after these authors under Rattus norvegicus. Rattus spp. Natural prey. — Cockroaches, Burma (Subramanian, 1927). India (Maxwell-Lefroy, 1909) ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 288 Family CAVIIDAE Cavia sp. prey. — (Hobmaier, Blattella germanica, U.S.A. Guinea pigs were fed cockroaches infested with Physaloptera Experimental 1941) VOL. I4I : maxillaris. Order CARNIVORA CANIDAE Family Canis familiaris Linnaeus — U.S.A. (Hobmaier, Blattella germanica, Experimental Physaloptera maxilinfested with cockroaches fed Dogs were 1941) infested with Cockroaches Ameel, laris. U.S.A. (Petri and 1950) prey. : : Physaloptera rara were fed to a dog. Canis latrans Say Blattella germanica, U.S.A. (Petri and Ameel, Cockroaches infested with Physaloptera rara were fed to a Experimental prey. 1950) : — coyote. Vulpes prey. — Experimental Urbain, 1938a) : A sp. (Brumpt and when fed cock- Blattella germanica, France fox w^as successfully infected roaches 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, The ring-tailed cat enters dwellings personal communication, 1953) located on the desert and feeds on cockroaches and other arthropods. — : Nasua narica (Linnaeus) Natural prey. — France (Brumpt and Urbain, The coati apparently became infested naturally with Pros1938) thenorchis spirula for which B. germanica was the intermediate host Blattella germanica, : 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). : BIOTIC ASSOCIATIONS OF COCKROACHES Nasua Natural prey. — Cockroaches, Family bain, 1938a) A : — & WILLIS 289 sp. British Guiana (Beebe, 1925a). MUSTELIDAE Meles Experimental prey. —ROTH sp. Blattella gcrmanica, France (Brumpt and Ur- badger was successfully infected when fed cock- roaches infested v/ith Prosthenorchis elegans and P. spirula. Family VIVERRIDAE Herpestes javanicus auropunctatus Hodgson — Natural prey. Epilampra ivheeleri, Eurycotis improcera, Panchlora nivea, Pycnoscelus stmnamcnsis, 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 of mongoose stomachs that he examined in Puerto Rico Panchlora nivea and Pycnoscelus surinamensis 19.6. zvheeleri 1.8, Ischnoptera rufa rufa 3.6, planeta americana 1.8, Herpestes Natural prey. : 1.8, Peri- sp. — Periplaneta americana and Periplancta Hawaii (Perkins, 191 3) number Epilampra : australasiae, Large numbers of these cockroaches are devoured. Cockroach, East Africa (Loveridge, 1923) found in stomach of mongoose. Family Felis Natural prey. —Periplaneta : Cockroach remains FELIDAE catus Linnaeus americana, Hawaii (Williams et al., 1931)Cockroaches, U.S.A., Arizona (Stahnke, personal communication, 1953)- Experimental 1941) : prey. — Blattella germanica, U.S.A. (Petri and Ameel, 1950) Physaloptera rara were fed to a kitten. laris. U.S.A. (Hobmaier, Cats were fed cockroaches infested with Physaloptera maxil: Cockroaches infested with France (Brumpt and Ur- : SMITHSONIAN MISCELLANEOUS COLLECTIONS 290 bain, 1938a) : A young cat VOL, I4I was fed cockroaches infested with Pros- thenorchis elegans and P. spirula. Felis pardalis mearnsi J. — Cockroaches, A. Allen Panama (Dr. H. L. Sweetman, perwas seen collecting and feeding on cockroaches, possibly Blaherus sp. "The ocelot was quite efficient and seemed to relish the roaches." Natural prey. sonal communication, 1958) An : ocelot CHECKLIST OF COCKROACHES AND SYMBIOTIC ASSOCIATES XIV. Only naturally occurring associations are included in this on page 315. mensal cockroaches listed because they undoubtedly occur are listed constitute symbiosis The may be in all species. were excluded because most of the associations list. Com- Bacteroids are not higher plants too casual to many of the plant associations were inThe cockroaches and the associates ecology. on however, ; cluded in the chapter within each category are arranged alphabetically by genus and species. Page references are to citations in the classified sections where of the associations and/or sources of the records are given. Aglaopteryx facies Mite Undetermined, p. 220. Anolis stratulus, p. 274. : Reptile: Aglaopteryx diaphana Nematode: Protrellus manni, 200. p. Allacta similis Insects Dolichurus stantoni, Solvidenia picticornis, p. 262. Anaplecta Fungus : Herpomyces sp. anaplectae, p. 134- Aptera fusca Protozoan: Gregarina fastidiosa, 182. p. Arenivaga boUiana Insect : Sarcophaga omani, p. 229. p. 247. details BIOTIC ASSOCIATIONS OF COCKROACHES Arenivaga Insects —ROTH & WILLIS 29I roseni Undetermined reduviids, : p. 227. Balta patula Insect Rhipidtoides ablcptus, : 230. p. Blaberus atropos Bacterium Spirochaeta blattae, Protozoa Endoliinax nana, p. 180. p. 125. : Nematodes Leidynema appendictdata, 197. p. Entamoeba colt, p. 178. Leidyncma cranijera, p. 198. Blaberus craniifer Bacteria Aerobacter aerogenes, Alcaligenes faecalis, Bacillus cereus, in. in. Escherichia jreundii, p. p. Micrococcus p. 120. colt var. 113. var. albus, p. 106. Micrococcus pyogenes var. aureus, Bacillus subtilis, p. 121. Escherichia p. pyogenes communior, p. 107. Proteus vulgaris, p. 113. p. 114. Pseudomonas aeruginosa, p. 104. Fungi Herpomyces Penicillium Protozoan: trictispidatus, p. 138. sp., p. 131. Diplocystis (?) Rhicopus nigricans, p. 133. Saccharomyces cerevisiae,p. 133. sp., pp. 181, 184. Nematodes Leidynema Undetermined, Bird Chicken, : Protrelleta floridana, p. 199. cranijera, p. 198. Mite: lolina nana, p. 219. p. 220. p. 278. Blaberus discoidalis Undetermined, Mite: p. 220. Blaberus giganteus Hair worm : Gordius pilosiis, p. 202. Blaberus sp. Fungi Herpomyces macropus, p. 136. Herpomyces paranensis, p. 136. Protozoan Leptomonas blaberae, : p. 167. Herpomyces Herpomyces periplanetae, p. 137. tricuspidatus, p. 138. : : : SMITHSONIAN MISCELLANEOUS COLLECTIONS 292 VOL. I4I Blaptica dubia Protozoan : Pileocephalus blaberae, 184. p. Blatta Bacterium: lateralis 119. p. Insect (Shelfordella) Shigella paradysenteriae, Ampulex : assimilis, p. 257. Blatta orientalis Bacteria Alcaligcnes p. p. recti, p. Arthromitus Micrococcus sp., p. 108. Paracolobactrum sp., p. in. in. in. Aerobaclcr aerogenes, Alcaligenes faecalis, intestinalis, p. Pasteurella pestis, 124. B. aerobio del pseudoedema maligno, p. 125. B. alcaligenes beckcri, 125. p. B. del pseudoedema maligno, p. 125. Proteus sp., p. 114. Proteus vulgaris, p. 113. 119. p. 114. Pseudomonas aeruginosa, Pseudomonas eisenbergii, Pseudomonas fiuorescens, Bacillo proteisimile, p. 126. Salmonella typhosa, Bacillo similcarbonchio, p. 126. Sarcina alba, Bacillo similtifo (Bacillo tifosimile), Sarcina p. 104. p. 105. p. 105. p. 117. p. 108. pp. 108, 109. sp., Sarcina symbiotica (host may have 108. Bacillus periplanetae, p. 121. been B. germanica), p. Sarcina ventriculi, p. 108. Bacillus stellatus, Serratia marcescens, p. 126. Bacillus biitschlii, 120. p. p. 121. S pirillochaeta Bacillus subtilis, p. 122. Bacillus tritus, p. 122. Bacleroides uncatus, iig. p. Clostridium lentoputrescens, novyi Clostridium sporogenes, Enterococcns Escherichia p. 122. coli, p. sp., p. 105. 105. 7, p. Spirochaeta periplanetae, Streptococcus faecalis, Streptococcus pyogenes, 112. p. pyogenes Streptococcus 109. var. albiis, 107. sp., p. 125. p. p. 109. no. no. no. p. Streptococcus microapoika, 109. Micrococcus pyogenes var. aureus, p. a, Spirillum Streptococcus liqtiefaciens, p. 122. sp., p. Lactobacillus fermenti, Micrococcus p. Clostridium or and Spirillum /3 117. p. blattae, p. 127. p. p. no. Treponema parvum, p. Treponema stylopygae, 125. p. 125. I'ibrio sp., p. 106. 107. Fungi Aspergillus fumigatus, Blasfocystis hominis, Blastocystis sp., p. 133. Torula gropengiesseri, 133. Candida seylanoides, Tondopsis p. 129. Coccidioides periplanetae, sp., p. p. 132. 130. p. 133. Protozoa Balantidium praenucleatmn, Bodo Herpomyccs periplanetae, p. 137. Herpomyces stylopygae, p. 137. p. 130. p. p. 187. blattae, p. 167. Coelosporidium periplanetae, p. 185. Diplocystis schneideri, p. 181. Endamoeba blattae, p. 177. Entamoeba thomsoni, p. 179. BIOTIC ASSOCIATIONS OF COCKROACHES Endolimax Endolimax Nyctotherns blattae, p. 180. sp., p. Oikomonas 180. p. 185. p. ovalis, p. sp., p. Plistophora kudoi, Plistoplwra sp., p. Retortamonas Stenophora, 188. p. 177. p. 185. p. p. 186. 186. blattae, p. sp., 293 166. Plislophora periplanetae, Herpetomonas periplanetae, p. 167. Hexamita periplanetae, p. 171. Lophomonas blattariim, p. 172. Lophomonas striata, p. 173. 167. 181. Tetratrichomastix blattidarmn, sp., p. 167. p. Monocercomonoides & WILLIS Peltomyces periplanetae, Grcgarina blattanun, p. 181. Haplosporidium periplanetae, Hartmannella blattae, p. 177. Monas —ROTH 170. orlhopteroruin, 169. Helminths Ascaris sp., p. 209. Enterobius vermicularis, Leidynema appendiculata, Spirura gastrophila, 209. p. Gongylonema neoplasticum, p. 206. Gordius blattae orienfalis, p. 202. ammerschmidtiella diesingi, p. 195. Hajnmerschmidtiella neyrai, p. 196. Mite Undetermined, pp. 220, 222. H Thelastoma pachyjuli, Trichostrongylus p. 197. p. 207. p. 201. 210. sp., p. Trichtiris trichiura, p. 210. : Insects Dermestes ater, p. 234. Dermestes sp., p. 234. Evania appendigaster, p. Evania dimidiata, p. 239. Bird: Qiicken, Mammal : p. Homo Prosevania punctata, worm : p. 248. Tetrastichits hagenozmi, p. 250. 236. Tetrastichus sp., p. 254. 278. sapiens, p. 286. Blatta Hair p. 240. Systellogaster ovivora, Gordius aquations, sp. p. 2or. Insects Evania appendigaster, p. 236. Tetrastichus hagenoivii, p. 250. Blattella germanica and/or Blattella vaga Viruses: Unspecified strains of poliomyelitis virus, p. 103. Blattella germanica Bacteria Achromobacter sp., p. Aerobacter aerogenes, Aerobacler cloacae, p. Alcaligenes jae calls, Alcaligenes viscosus, Bacillus circulans, Escherichia p. 1 10. p. in. 112. p. iii. p. iii. 120. coli, p. 112. Escherichia freundii, p. 113. Micrococcus aurantiacus, p. 106. Micrococcus epidermidis, p. Micrococcus pyogenes var. p. 106. albus, 107. Micrococcus pyogenes var. aureus, p. 107. Micrococcus ureae, p. 107. Micrococcus sp., p. 108. Mycobacterium leprae, p. 123. Paracolobactrum aerogenoides, p. 113. :: : : :: SMITHSONIAN MISCELLANEOUS COLLECTIONS 294 Paracolobactriim colijorme, Paracolobactrum p. 113. Pscudomonas aeruginosa, p. Sarcina symbiotica (host may have been B. oricntalis), Sahuonclla typhimurium, Serratia marcescens, sp., p. 113. 104. VOL. I4I Streptococcus jaecalis, Streptococcus p. 116. 117. p. p. 109. no. sp., p. 108. p. Fungi Aspergillus flavus, Aspergillus Cordyccps Protozoa Bodo Herpomyces 130. p. Aspergillus tamarii, p. 130. sp., p. 132. p. 133. blattae, p. 134. may have (host sp. P. americana), sp. p. 185. p. 177. Eiidamoeba blattae (host may have been P. americana) p. 177. Endolimax sp. (host may have been , P. americana), Entamoeba may have P. americana) Entamoeba , been 178. p. (host histolytica have been P. americana) Helminths , may p. 179. sp. p. 179. may have (host been P. americana) 169. p. , Gregarina blattarum, p. 182. lodamoeba may have been sp. (host P. americana) , p. 180. Lophomonas blattarum, p. 172. Lophomonas striata, p. 173. Nyctotherus 180. p. coli (host Entamoeba thomsoni, Eiitrichomastix may have been (host P. americana), been p. 167. Coelosporidium periplanetae, Dobellina ectobiae, p. 135. Memnoniella echinata, Saccharomyces sp., p. 130. ovalis, p. 188. Peltomyces periplanetae, p. 177. Plistophora periplanetae, p. 186. Tetratrichomastix blattidarum, p. 170. Blatfelicola blattelicola, p. 193. Moniliformis kalahariensis, Blatticola blattae, p. 193. Prosthenorchis elegans, Enterobius vermicularis, Galebia aegyptiaca, p. 209. p. 195. Gongylonema neoplasticum, p. 206. p. 203. p. 203. Prosthenorchis spirula, p. 203. Tetrameres americana, p. 207. Trichuris trichiura, p. 210. Mites Blattisocius tincivorus, Caloglyphus sp., p. p. 216. Undetermined, p. 220. 218. Insects Brachygaster minuius, Eupelmus p. 235. atriflagellum, p. 247. Dolichurus corniculus, Ripidius pectinicornis, p. 261. p. 232. Mammals Callithrix chrysoleucos, Callithrix jacchus, Cebus Lemur Lemur p. 285. apella, p. 284. coronatus, Leontocebus ursulus, p. 283. fulvus, p. 283. Leontocebus oedipus, p. 285. Leontocebus rosalia, p. 285. Perodicticus potto, Saimiri sciurea, Blattella humbertiana Fungus: Herpomyces gracilis, p. 135. Protrellus phyllodromi, Nematode : p. 200. Mite: Uropoda sp., p. 217. p. 285. Macaca sylvanus, p. 286. Nasua narica, p. 288. Nasua nasua, p. 288. Pon sp., p. 286. p. 285. p. 284. p. 285. BIOTIC ASSOCIATIONS OF COCKROACHES Blattella Insect Dolichurus stantoni, : —ROTH & WILLIS 295 lituricoUis p. 262. Blattella sp. Reptile : Anolis cnstatcllus, p. 273. Cariblatta delicatula Hyptia Insect: Reptile : Anolis sp., p. 240. cristatellus, p. 273. Cariblatta lutea lutea Plants Sarraccnia flava, Sarraccnia purpurea, p. 154. Chorisoneura p. 154. sp. Insects Stylogaster sp., p. Tachysphex 228. Choristima blatticidus, p. 264. sp. Insect: Rhipidioides rubricatus, p. 231. Choristimodes sp. Insect: Rhipidioides rubricatus, p. 231. Cryptocercus punctulatus Bacterium: Bacillus subtilis, p. Protozoa Adelina cryptocerci, p. 184. 122, Oxymonas Barbidanympha estaboga, p. 173. Barbulanympha laurabuda, p. 174. Barbulanympha uj alula, p. 174. Barbulanympha wenyoni, p. 174. Eticomonympha imla, p. 176. Hexamita cryptocerci, p. 171. Idlonympha perissa, p. 174. Leptospironympha eupora, p. 172. Leptospironympha rudis, p. 172. Leptospironympha wachida, p. 172. Macrospironympha xylopletha, p. 172. Monocercomonoides globus, Notila proteus, Oxymonas Amphibian p. 269. : p. 169. p. 170. doroaxostylus, Plethodon p. 170. glutinosiis, nana, p. 170. Prolophomonas tocopola, Rhynchonympha p. 173. 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 :: : : : : SMITHSONIAN MISCELLANEOUS COLLECTIONS 296 VOL. I4I Cutilia soror Insects Dolichurus stantoni, S::epligetella sericca, p. 242. p. 262. Evania appendigaster, Phastamis sp., p. Bird p. 236. 277. : Cutilia sp. Nematode: Undetermined, p. 201. Insects Undetermined strepsipteron, Riekella anstralis, p. 231. p. 234. Diploptera punctata Bacterium Serratia marccsccns, : 117. p. Fungus Herpoinyccs : diploptcrac, 135- p. Mites Locuslacanis lolina nana, p. 219. Nauphocta cinerca, Amphibian Bufo marimis, Insect p. : Bird : Phasianiis sp., p. 219. 324. p. : 270. 277. sp., p. Ectobius lapponicus Protozoa Camocystis tenax, p. Monocercomonoidcs orlhopterorum, 184. p. Nematode : 169. Blatticola blaitac, p. 193. Insects Brachygaster minutus, p. Dolichurus corniculus, p. 261. Tachysphex 235. lativalvis, p. 264. Ectobius pallidus Protozoan: Helminths Camocystis tcnax, p. 184. Undetermined mcrmithid, Blatticola blattae, p. 193. p. 192. Insects Ampidcx Ripidius denisi (presumptive record), fasciata, p. 259. Aphaenogaster picea, p. 267. Dolichurus cornicidus, p. 261. p. 232. Tachysphex lativah'is, p. 264. Lasitts alicmis, p. 268. Ripidius boissyi (presumptive rec- ord), p. 231. Ectobius panzeri Centipede: Scolopendra sp., p. 224. Insects Dolichurus corniculus, p. 261. Tachysphex lativalvis, p. 264. BIOTIC ASSOCIATIONS OF COCKROACHES Ectobius —ROTH & WILLIS 297 sp. Insects Brachygastcr miniitus, Tachysphex p. 235. Dolichurus cornictilus, p. lativalvis, p. 264. 261. Ellipsidion affine Insect Rhipidioidcs fuscatus, : p. 230. Ellipsidion australe Insects: Agamer'wn Mcstocharomyia oophaga, mclallica, p. 243. Chciloncnrns viridiscutnin, p. p. 248. 244. Epilampra abdomen -nigrum Podium flavipcnne, Bufo marinus, Insect: Amphibian : 265. p. p. 270. Epilampra conferta Insect Podium : 266. sp., p. Epilampra Fungus p. Herpomyces : sp. tricuspidatus, 138. Podium hacmatogastrum, Insect: p. 266. Bird : Gymnasio nudipcs, p. 279. Epilampra wheeleri Reptiles Ameiva Mammal : Anolis cristatellus, exsul, p. 275. p. 273. Herpestes javanicus auro- pnnctatus, p. 289. Escala (?) Insect: Rhipidioidcs adynatus, p. sp. 230. Eurycotis floridana Fungi Aspergillus flavus, p. Aspergillus sydowi, 130. p. 130. Helminths Euryconema paradisa, p. 194. Leidyncma appendiculata{ ?), Protrelloides paradoxa, p. 198. p. 200. Undetermined gordian worm, p. 202. Insects Anastatiis floridamis, p. 245. Undetermined Eurycotis improcera Mammal : Herpestes javanicus auro- punctatus, p. 289. tachinid, p. 228. SMITHSONIAN MISCELLANEOUS COLLECTIONS 298 VOL. I4I Eurycotis manni Fungus Herpomyces : sanzibarinus, 138. p. Graptoblatta notulata Tachysphex Insect: jamiiensis, p. 264, Gromphadorhina portentosa Undetermined gregarine, Protozoan: 184. p. Mite: Coleolaelaps ? sp., p. 216. Gyna Fungus sp.(?) Herpomyces zanzibarinus, : 138. p. Holocompsa fulva : Phcidole megaccphala, Insect: Dolichurus corniculus, Insect p. 268. Hololampra punctata p. 261. Ischnoptera deropeltiformis Plant Sarracenia flava, : Amphibian Mammal : p. 1 54. Hyla cinerea, p. 270. Dasypus novemcinctus, : p. 287. Ischnoptera rufa rufa Fungus Spicaria prasina, : Mammal p. 130. Herpestes javanicxis auro- : punctatus, p. 289. Ischnoptera Fungus Insects : Herpomyces sp. arietiniis, p. 134. : Ampulex canaliciilata, p. 257. Syntomosphynim p. 249. Karnyia discoidalis Spider : Latrodectus indistinctus, p. 215. Kuchinga hemerobina Hair worm p. 202. : Parachordodes raphaells, ischnoptcrae, BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS Leucophaea maderae Bacterium: Serratia marcescens, p. 117- Fungus Herpomyces : tricnspidahis, 138. p. Protozoa Gregarina rhyparobiae, : Hexamita Nematodes ( ?) Hammerschmidtiella p. Retortamonas (?) sp., p. 167. Undetermined gregarine, p. 184. p. 183. 171. sp., p. Protospirura bonnei, diesingi, Leidynema Mite: p. 206. Protospirura miiricola, 195. p. 206. delatorrei, p. 198. Chactodactyliis sp., p. 218. Mammals Aotes zonalis, Cebus capucinus, p. 284. Ateles dariensis, Leucophaea Nematode tum, : p. p. 284. Rattus norvegicus, p. 284. p. 287. sp. Cephallohellus brevicauda194. Leurolestes pallidus Fungus : Herpomyces leurolestis, p. 136. Loboptera decipiens Insects Dolichurus corniculus, Zeuxevania splendidula, 261. p. Loboptera Fungus : Herpomyces p. sp. lobopterae, p. 136. Lobopterella dimidiatipes Insect: Ampulex Bird Coturnix : canaliciilata, p. 257. coturnix japonica, 277. p. Melanosilpha capensis Protozoa Gregarina impetuosa, p. Gregarina sandoni, 183. Monastria Insect : Triatoma arthurneivai, p. 183. sp. p. 227. Nauphoeta cinerea Bacteria Salmonella typhimnrium, p. 116. Serratia marcescens, p. 117. 243. 299 : : : SMITHSONIAN MISCELLANEOUS COLLECTIONS 300 Fungus Herpomyces : VOL. I4I tricuspidatus, 138. p. Mites Blattilaclaps naiiphoetae, p. 216. Insect Pheidole megacephala, : Locnstacanis sp., p. 219. p. 268. Neostylopyga rhombifolia Bacterium Serratia marcescens, : p. 117. Insects Evania appendigaster, Ssepligetella sericca, Mammal Homo : p. Tefrastichus hagenowii, 237. p. 250. p. 242. sapiens, p. 286. Nyctibora obscura Fungus Herpomyces amasonicus, : 134- p. Nyctibora Fungus Herpomyces : sp. nyctoborae, 136. p. Nyctibora tomentosa Fungus Herpomyces nyctoborae, : p. 136. Oniscosoma granicoUis Insect Parancphrites xemis, : p. 230. Panchlora exoleta Protozoan p. : Gregarina panchlorae, 183. Panchlora nivea Bacterium: p. Fungus p. Serratia marcescens, 117. : Herpomyces pancJilorae, 136. Mammal: Herpestes pimctatus, p. javanicus auro- 289. Panesthia angustipennis Protozoa Clevelandella constricta, p. 189. Clcvclandclla hastula, p. 189. Clevelandella contorta, p. 189. Clevelandella pancsthiae, Clevelandella elongata, p. 189. Clevelandella parapanesthiae, p. 189. p. iJ BIOTIC ASSOCIATIONS OF COCKROACHES Endamoeba javanica, p. 178. Endamoeba philippinensis, p. Hexamita cryptocerci, p. 171. —ROTH & WILLIS Nyctotherus uichancoi, 178. p. 188. Paraclevelandia brevis, Undetermined amoeba, 190. p. Paraclevelandia simplex, Monocercomonoides panesthiae, 3OI 190. p. 181. p. p. 170. Nematodes Aorurus philippinensis, Blattophila Leidynemella fusiformis, p. 193. sphaerolaima var. ja- vanica, p. 194. Leidynema nocalum, p. p. 198. Leidynemella paracranijera, p. 198. Thelastoma palmcttum, p. 201. 198. Panesthia australis Fungus p. : Metarrhisium anisopliae, 131. Mite: Hypoaspis sp., p. 217. Undetermined tachinid, Undetermined diplogyniid, Insect: p. 217. p. 228. Panesthia laevicoUis Nematode : Blattophila sphaerolaima, p. 194. Panesthia laevicoUis (?) Nematode p. : Leidynemella fusiformis, 198. Panesthia lobipennis Fungus p. : Herpomyces panesthiae, 136. Panesthia spadica Protozoa Clevelandella constricta, p. 189. Endamoeba Clevelandella contorta, Nyctotherus uichancoi, p. 188, Paraclevelandia brevis, p. 190. 189. p. Clevelandella nipponensis, Clevelandella panesthiae, 189. p. p. 189. Panesthia Nematode p. : javanica, p. 178. Paraclevelandia simplex, p. 190. sp. Leidynemella panesthiae, 198. Parahormetica bilobata Helminths Agamospirtira parahormeticae. Undetermined gordian worm, p. 205. Parcoblatta lata Sarracenia flava, Protozoan: Leptomonas Plant: Nematode : p. sp., 154. ,,. Protrellus aiirifiuus, 167. p. 199. p. 202. :: SMITHSONIAN MISCELLANEOUS COLLECTIONS 302 VOL. I4I Parcoblatta pensylvanica Protozoa Gregarina blattarum, p. 182. Gregarina illinensis, p. 183. Gregarina parcoblattae, p. 183. Nematode Rictularia : Gregarina thomasi, p. 184. Leptomonas sp., p. 167. Nyctotherus ovalis, p. 188. coloradensis, 205. p. Centruroides vittatus (?), Scorpion: p. 212. Insects : Camponotus pennsylvanicus, p. 267. Hyptia dorsalis, p. 239. Hyptia harpyoides, p. 239. Hyptia reticulata, p. 240. Amphibian: Rana (?) sp., p. 272. Hyptia thoracica, Podium p. 240. Carolina, p. 265. Systellogaster ovivora, 248. p. Parcoblatta uhleriana Fungus Herpomyces : arietimis, 134- p. Protozoan : Gregarina parcoblattae, p. 183 Nematode : P rot reikis aurifluus, p. 199- Mite: Undetermined, p. 220. Hyptia harpyoides, p. 239. Insect : Parcoblatta virginica Fungus : Herpomyces arietinus, p. 134. Protozoa Gregarina ohioensis, Nematode : Leptomonas p. 183. sp., p. 167. Rictularia coloradensis, p. 205. Insects : Hyptia harpyoides, p. Podium 239. luctiiosum, p. 266. Parcoblatta sp. Fungus : Herpomyces arietinus, p. 134. Plant: Sarracenia flava, p. 154. Pimeliaphilus podapolipopJiagus, Mite : p. 219. Insects : Coenosia basalts, Syntomosphyrum p. 229. Dolichurus greenei, Tetrastichus hagenowii, Hyptia sp., p. 240. Megaselia sp., p. 227. Periplaneta americana Viruses: Unspecified strain(s) poliomyelitis virus (host been P. brunnea), blattae, p. 248. Systellogaster ovivora, p. 261. p. 103. of may have p. p. 248. 250. BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 303 Bacteria Achromohacter hyalinum, p. no. Aerobacter aerogenes, p. in. Aerohacter cloacae, p. 112. Aerobacter sp., p. 112. Alcaligenes faecalis, Bacillus cereus, p. Pseudomonas 120. Bacillus niegaterium, p. Bacillus subtilis, p. 122. Bacterium alkaligenes, Clostridium sp., Salmonella 119. p. p. Salmonella 112. colt, p. 112. Escherichia jrcundii, sp. (Type Kentucky), sp. (Type Meleagris), p. 116. Salmonella 113. p. Escherichia intermedium, , Salmonella Pscudomonas aeruginosa, Newport), 113. (Type Oranienburg), sp. p. 116. Salmonella (Type sp. Panama), p. 116. Salmonella (Type sp. Rubislaw), p. 116. Salmonella (Type Tennessee), sp. p. 116. Sarcina 113. Paracolobactrum coliforme, p. Paracolobactrum sp., p. 113. Proteus mirabilis, p. 114. Proteus morganii, p. 114. Proteus rettgeri, p. 114. Proteus vulgaris, p. 114. Proteus sp., p. 114. (Type sp. p. 116. p. 113. Mycobacterium jricdmannii, 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. Bredeny), p. 115. Eberthclla oedematicns, Escherichia (Type sp. p. 115. 122. p. 115. Salmonella 121. p. 114. p. Salmonella morbificans, p. 115. Salmonella schottmuelleri, p. 115. Salmonella sp. (Type Bareilly), iii. p. fluorescens, p. 105. Salmonella anatis, sp. p. 108. Serratia marc esc ens, p. 118. Shigella alkalescens, p. 118. Spirillum periplaneticum, 105. p. Streptococcus faecalis, p. 109. Streptomyces leidynematis, pp. 196, 197. Tetragenous 127. sp., p. Veillonella parvida, p. 109. p. 104. Fungi Aspergillus flavus, p. Aspergillus niger, p. 130. Aspergillus sp., Cephalosporimn Herpomyces Herpomyces p. chactophilus, p. 135. periplanetae, p. (host 137. p. Endamoeba 187. Endolimax Endolimax 187, may have been 5. Dobellina sp., p. sp. p. 185. 181. 181. (host 5. germanica), p. p. may have been 177. 131. p. 133. 132. blattae, p. sp. , been 180. p. coli (host p. 180. may have (host 5. germanica), Entamoeba 177. blattae, p. 5. germanica) Entamoeba gertnanica), p. 167. Diplocystis schneideri, 132, sp., p. p. 131. Coelosporidium periplanetae Diplocystis sp., p. Rhisopus sp., p. 133. Syncephalastrum sp., Torula acidophila, p. 131. sp., p. Metarrhisiicm anisopliae, sp. guilliermondii, p. 132. Penicillium 131. Protozoa Balantidium blattarum, Balantidium ovatum, p. Balantidium sp., p. 187. j5ocfo Mucor Mucor 130. may have been 178. histolytica (host have been 5. germanica) Entamoeba sp. p. 179. Entamoeba thomsoni, p. , 179, p. may 179. :: :: : : : : SMITHSONIAN MISCELLANEOUS COLLECTIONS 304 may have Eutrichomastix sp. (host been B. germanica), p. 169. Gregarina blattarum, p. 182. Gregarina legeri, p. 183. Gregarina neo-brasiliensis, p. Hexamita periplanetae, p. 171. lodcmoeba sp. (host may have been B. germanica) Helminths Ancylostoma duodenale, simm, p. Ascaris sp., p. p. 206. Leidynenia appendiculata, 197. p. dubitis, p. 203. Moniliformis moniliformis, p. 204. Necator americanus, p. 210. Protrellus kUnckeli, 199. p. Schivenkiella icemi, p. 200. 202. Hammerschmidtiella p. p. 184. 170. Moniliformis 209. Bincma mirzaia, p. 193. Gongylonema neoplasticum, Gongylonema sp., p. 206. Gordiiis Protomagalhaesia serpentida, p. 209. or lumbricoides 188. p. Plistophora periplanetae, p. 186. p. Isotricha caulleryi, p. 187. Ascaris 169. Nyctotheriis ovalis, Tetratrichomastix blattidantm, 180. p. Lophomonas blattarum, p. 172. Lophomonas striata, p. 173. Monocercomonoides orthopterorum, p. 183. VOL. I4I diesingi, Splrura gastrophila, p. 207. Thelastoma pachyjiili p. 201. 195- Hymenolepis Trichuris trichiiira, 208. sp., p. 210. p. Mites Caloglyplnis 218. sp., p. Rhisoglyphus tarsalus, Pimeliaphilus podapolipophagus, p. 219. Tyrophagus noxius, p. 218. p. 218. Undetermined, p. Avicularia p. 214. 220. Spiders Avicularia avicularia, Centipedes p. 214. Undetermined, : sp., p. 222. Insects Ainpulex amoena, p. 256. Ampulex compressa, Ripidius pectinicornis, p. 259. Anastatus tenuipes, p. 246. Calodexia (?) venteris, -p. 22%. Evania appendigaster, p. 237. Melittobia chalybii, p. 248. Prosevania punctata, Amphibian Hyla cinerea, p. 241. Spiniger domesticus, p. 232. p. 227. Ssepligetella sericea, p. 242. Tetrastichus hagenowii, p. Tetrastichns periplanetae, Tetrastichus Trirhogma sp., p. 254. caeridea, p. 262. 270. p. Reptiles Ameiva Anolis exsiil, p. 275. cristatellus, p. 273. Birds Chicken, p. Sparrow, 278. p. 282. Mammals Fclis catus, p. 289. Homo Herpestea javanicus auropunctatus, Rattus norvegicus, Herpestes sapiens, p. 286. p. 287. Rattiis rattus, p. 287. p. 289. sp., p. 289. Periplaneta australasiae Bacteria Mycobacterium leprae, p. 123. 250. p. 253. Serratia marcescens, p. 118. BIOTIC ASSOCIATIONS OF Fungus Heipomyccs : COCKROACHES —ROTH & WILLIS 305 periplanetae, 137. p. Plant: Sarracenia vtinor, p. Protozoa Etidamoeba blattae, p. 177. Nematodes Gongylonema p. 206. diesingi, p. 195. Moniliformis dubius, blattae, p. 180. Protrcllus australasiae, p. 199. Protrellns kiinckeli, p. 200. 203. p. Undetermined, : Endolimax neoplasticiim, Hammerschmidtiella Centipedes 154. p. 222. Insects Ampulex amoena, p. Ampulex compressa, Tetrastichus australasiae, 256. Evania appcndigaster, Tetrastichus hagenozvii, 259. p. Tetrastichus, p. 237. Ssepligetella sericea, p. 242. sp., p. p. 249. p. 250. 254. Trirhogma caertdca, p. 262. Chrysemys picta, p. 272. Reptiles Anolis cristatellns, p. 273. Bird Callus sp., p. 277. : Mammals Herpestcs sp., p. Homo 289. sapiens, p. 287. Periplaneta brunnea Viruses: Unspecified strain (s) poliomyelitis virus (host of may have been P. americana) p. 103. Bacterium Serratia marccsccns, , : 118. p. Fungus Herpomyces : Nematode Insect periplanetae, 137- P- : Schwenkiella icemi, p. 200. Tetrastichus hagenowii, p. 251. : Periplaneta fuliginosa Insect: Ampulex amoena, p. 256. Periplaneta Bacterium : sp. Serratia marcescens, p. 118. Fungi Herpomyces chactophilus, p. 135. Lophomonas blattarum, Protozoan p. Herpomyces periplanetae, p. 137. : 172. Helminths : Thelastoma rivcroi, p. 201. Moniliformis moniliformis, p. 204. Insects Ampulex compressa, p. 259. Evania subspinosa (presumptive record?), p. 239. Amphibian: Bufo marimis, p. 270. Ssepligetella sericea, p. 242. : : : : : SMITHSONIAN MISCELLANEOUS COLLECTIONS 3o6 VOL. I4I Reptiles Anolis grahami, Bird: Anolis leachi, p. 274. Vireo latimeri, p. 274. p. 281. "Phyllodromia" sp. Herpomyces phyllodromiae, Fungus: ^27. P- Insects Dicarnosis alfierii, p. Dolichurtis stantoni, 245. p. 262. Platyzosteria castanea Insect Riekella : sp., p. 231. Platyzosteria novae seelandiae Protozoan : Monocercomonoides inelo- lonthae, p. 169. Platyzosteria scabra Fungus Herpomyces appendiculatus, : p. 134. Platyzosteria Riekella nitidioides, Insect: 231. p. Plectoptera Bird: Todus mexicamis, p. sp. sp. 279. Polyphaga aegyptiaca Nematodes Hammerschmidtiella diesmgi, Oxyuris (?) heferogamiae, p. Blatticola blattae, p. 193. Galebia aegyptiaca, p. 195. p. 195. 199. Polyphaga saussurei Helminths Hyvtenolepis Insects: p. sp., p. Undetermined tapeworm ova, 208. p. 208. Undetermined reduviids, 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 Undetermined ciliate, p. 190. flagellate, p. 176. Undetermined gregarine, p. 184. BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS Nematodes Oxyspirura mansoni, Severianoia severianoi, p. 204. p. 200. Mites Caloglyphns spinitarsus, Histiostoma feroniarum, Tyrophagus 217. p. lintneri, p. 218. Undetermined, p. 217. p. 220. Insects Pheidole megacephala, Amphibian p. Bufo marinus, : Anolis sagrei, Reptile: Undetermined 268. ants, p. 350. p. 270. p. 274. Birds Acridothercs Passer domesticus, 281. tristis, p. Ducks, p. 277. Melanerpes portoricensis, p. 280. Meleagris yallopavo, p. 278. Mammal Herpestes javanicns aiiro- Phasianiis sp., p. p. 282. 277. Streptopelia chinensis, p. 278. : piinctatiis, p. 289. Robshelfordia circumducta or Robshelfordia longiuscula Insects Neorhipidius neoxenus, Rhipidioides helenae, Rhipidioidcs mollis, p. 230. p. 231. p. 230. Steleopyga (?) sinensis Nematode Suifunema : candelli, p. 200. Supella supellectilium Viruses: Unspecified poliomyelitis virus, strain(s) p. of 103. Bacterium: Serratia marcesccns, p. 117. Fungus Herpomyces supellae, p. 138. Nematode Blattophila supellaima, : : p. 194. Insects Comperia merceti, Anastatiis tenuipes, p. 246. Symploce Reptile : Anolis flagellata or Symploce cristatelliis, p. 273. Symploce parenthesis Hair worm : Parachordodes raphaelis, p. 202. Temnopteryx phalerata Protozoan : Cregarina gibbsi, p. 182. ruficollis p. 244. 3O7 : : : : :: SMITHSONIAN MISCELLANEOUS COLLECTIONS 3o8 VOL. I4I Undetermined Cockroaches Bacteria Arthromitiis intestinalis, p. 124. p. 122. Bacillus suhtilis, p. 122. Clostridium perjringens, Escherichia Proteus vulgaris, Spirillum 112. coli, p. Paracolon bacilli, p. 113. Proteus morganii, p. 114. Fusiformis lophomonadis, p. 119. Micrococcus citreus, p. 106. Micrococcus pyogenes var. aureus, p. 107. leprae, p. Streptococcus Vibrio I Heiberg, 106. 123. diploptcrae, p. 135. forficularis, p. 135. Metarrhisium anisopliae, blattina, p. 139. sp., sp., p. 139. p. 133. Cordyceps amazonica, Herpomyces Herpomyces Herpomyces Herpomyces Types p. 125. no. and II sp., p. 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. Amphoromorpha Amphoromorpha Blastocystis 114. p. 105. Spirochaeta periplanatae , p. Mycobacterium Fungi sp., p. 134. p. anaplectae, p. 134. chilensis, p. 135. p. 131. Plants Nepenthes ampularia, Nepenthes gracilis, p. Protozoa p. 154. Diplocystis sp., p. Endamoeba Entamoeba blattae, p. Giardia sp., p. 182. Hexamita periplanetae, p. 171. Lophomonas blattarum, p. 172. Lophomonas striata, p. 173. Monas orthopterorum, p. 169. 177. 172. Gregarina blattarum, sp., p. 154. Monocercomonoides 181. histolytica, p. 179. p. Nepenthes 154. sp., p. 167. Nyctotherus buissoni, Nyctotherus ovalis, p. Nyctotherus viannai, 188. p. 188. 189. p. Oikomonas blattarum, p. Oikomonas sp., p. 166. Paramecium sp., p. 186. Trichomonas sp., p. 171- 166. Helminths Cephalobellus magalhdesi, p. 194. Chordodes morgani, p. 201. Gordius sp., p. 202. Hammerschmidtiella diesingi, p. 195. Leidynema appendiculata, p. 197. Moniliformis moniliformis, p. 204. Schwenkiella icemi, p. 200. Severianoia magna, p. Severianoia severianoi, 200. p. 200. Spirura gastrophila, p. 207. Undetermined gordian worms, p. 202. Protrellus galebi, p. 199. Scorpion: Heterometrus longimanus, p. 213. Spiders : Latrodectus mactans Ctenid, p. 214. p. 216. Heteropoda venatoria, p. 215. Pimeliaphilus podapolipophaMite : gus, p. 219. Centipedes Allothereua maculata (circumstantial evidence), p. 223. Scolopendra morsitans, p. 223. Scolopendra subspinipes, p. 224. Scutigera coleoptrata (circumstantial evidence), p. 222. BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 3O9 Insects Acanthinevania princeps, Agamerion p. 235. Amptilex compressa, Ampule X ruficornis, p. 259. Ampulex sibirica, p. 260. Ampulex somwrati, p. 260. 247. p. 246. Pompiliis bracatus, Blatticidella ashmeadi, p. 243. Pompilus Calodexia Salius verticalis, spp., p. 228. Dicarnosis p. 256. p. 256. p. 233. Solindenia piciicornis, 245. alfierii, p. p. 247. Diestrammena apicalis, p. 226. Diestrammena japanica, p. 226. Stylogaster spp., Dolichiirus bicolor, 260. Syntomosphyrum p. 261. Systellogaster ovivora, p. Dolichurus corniculns, Dolichurus p. 261. 262. sp., p. Dorylus nigricans sjostedi, Doryhts sp., p. 267. Dorylus wilverthii, p. 267. Eciton burchelli, p. 228. Stylogaster stylata, p. 228. blattae, p. 248. p. 248. Tachysphex blatticidus, p. 264. Tachysphex coriaceus, p. 264. Tachysphex fatmiensis, p. 264. gilberti, p. 261. Dolichurus ignitus, Dolichurus 230. p. 256. sp., p. Ripidius scutellaris, 226. p. 268. p. Podium abdominale, p. 265. Podium dubium, p. 265. Podium rufipes, p. 266. Blatticida pulchra, p. 243. Clerada apicicornis, p. 245. 268. p. Neonephrites partinigcr Anastatiis blattidifiirax p. 245. Anastatus tenuipes, sp., p. "Formica omnivora," Iridomyrmex humilis, 259. p. Eupelmus Eutrichosomella blattophaga, mctallica, p. 243. p. 267. p. 268. Tetrastichus hagenozvii, 251. p. Tetrastichus periplanetae, p. 253. Trirhogma sp., p. 264. Undetermined tachinid, 229. p. Fish: Chalceus macrolepidotus, p. 269. Cyrtocharax magdalenae essequiben- Potamotrygon humboldti, Rhamdia sebae, p. 268. p. 268. sis, p. 269. Amphibians Arthroleptis variabilis, p. 270. Hyla cinerea, p. 270. Hyper alius picturatus, Leptodactylus pentadactylus, Leptopelis calcaratus, Bufo ictericus, p. 270. Bufo funereus, p. 270. Bujo marinus, p. 270. Leptopelis rujus, p. 271. Megalixaliis jornasinii, 7?ona catesbeiana, p. 271, Leptodactylus albilabris, p. 271. p. 271. p. 271. 7?aMa mascareniensis, Tree frogs, p. p. 271. p. 271. p. 351. Reptiles Ameiva Ameiva Anolis cristatellus, p. 273. Anolis leachi, p. 274. sp., p. p. 273. Cnemidophorus Leiolopisma laterale, p. p. 274. sp., p. p. 272. p. 274. 275. Skinks, p. 274. Sphaerodactylus 274. Anolis stratulus, p. 272. Hemidactylus frenatus, Lizards, p. 274. Anolis pulchellus, p. 273. Gekko gecko, 275. sp., p. Anolis grahami, Anolis Geckos, exsul, p. 275. Thecadactyliis sp., p. sp., p. 273. 273. 275. Tropidophorus grayi, 281. Bambusicola thoracica, p. 274. Birds Acridotheres tristis, p. Agelaius xanthotntis, p. 282. Chickens, p. 278. p. 277. 271. ; : SMITHSONIAN MISCELLANEOUS COLLECTIONS 310 Colurnix coturnix japonica, Dendrocopos mahrattensis, Ducks, p. 278. Tiaris bicolor omissa, p. 282. Tockus 280. Holoquiscalus brachypterus, Icterus portoricensis, p. Phasiamis sp., p. 277. Sparrow, p. 282. p. 279. p. 277. Gymnasia nitdipes, p. 279. Gymnopiihys leucaspis, p. Owl, Partridge, 277. p. VOL. I4I 282. p. p. 280. birostris, p. 279. Troglodytes aedon, p. 281. Troglodytes aiidax, p. 281. 351. Mammals Bassariscits astutiis, p. 288. Homo Callithrix jacchus, p. 285. Loris tardigradus, Erinaceus Molossus sp., p. 283. Monodelphis sp., p. 283. sp., p. 283. Felis catus, p. 289. sapiens, p. 287. p. 284. Felis paradalis mearnsi, p. 290. Nasiia nasita, p. 288. Herpestes javanicus auropimctatus Herpcstes sp., p. 289. Nasua sp., p. 289. Ratins 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 com- mensal 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 a specific habit, individual cases are generally is, however, unwise to disregard such isolated observations or dismiss them lightly. fixed as dismissed as coincidences. It Rothschild and Clay (1957) The following social insects have been found harboring cockroaches commensalism by acquiring food from which the cockroaches presumably bene- in a state of in fit their hosts. are not apparent. Benefits accruing to the hosts Unfortunately, biological details are not always sufficient to substantiate the suspected association. commensals of the However, it seems 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 significant that the cockroach insects listed pointed out that the myrmecophilous cockroaches are only a very few millimeters long their eyes are reduced ; ; all small, being they are apterous or subapterous and they are all of American origin. HOSTS OF COMMENSAL COCKROACHES ISOPTERA RHINOTERMITIDAE Order Family Coptotennes ceylonicus Holmgren Commensal. —Sphecophila ravana, Ceylon females, 50 males, and (Fernando, 1957) : Six nymphs of both sexes were found among de- BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 3II caying timber in the ground in association with a colony of this termite. The antennae of most specimens were mutilated unsymmetri- cally. Family TERMITIDAE Macrotermes barneyi Light Commensal. — Nocticola Among Kowloon sinensis, (Silvestri, 1947) : specimens of termites collected from a nest. Macrotermes bellicosus (Smeathman) or Macrotermes natalensis (Haviland) — — Synonymy. Termes bellicosus [Snyder, 1949]. Commensal. Sphccophila 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 Commensals. —Nocticola sp. Kowloon sinensis, : (Silvestri, 1946) : In termite nest. Nocticola termitophila, Penang (Silvestri, 1946) Termes Commensal. —Nocticola vestri, 1946) : sinensis, : In termite nest. sp. Repulse Bay, Australia (?) (Sil- In a termite gallery. Termites Commensal. — Ergaula capensis [=Dyscologamia zvollastoni] French Equatorial Africa, Brazzaville (Rehn, 1926; Chopard, 1949). Order HYMENOPTERA FORMICIDAE Subfamily FORMICINAE Family Camponotus femora tus (Fabricius) Commensal. —Phorticolea holiviae, Bolivia, Cachuela Esperanza Three males collected in the joint nests of C. 1923) femoratus and Crematogaster limata. (Caudell, : SMITHSONIAN MISCELLANEOUS COLLECTIONS 312 Camponotus maculatus (Fabricius) Note. — Dr. that this ant VOL. I4I ? W. is L. Brown (personal communication, 1957) states 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 Commensals. rufipes (Fabricius) —Atticola mortoni, Nothoblatta wasmanni, and Phor- ticolea testacea, Brazil, San Leopoldo (Bolivar, 1905) : Found in the formicaries of C. rufipes. Formica rufibarbis Fabricius and Formica subcyanea Wheeler — Commensal. Myrmecoblatta rehni, Mexico (Mann, 1914) were very abundant, several occurring in almost every nest." Subfamily "They MYRMICINAE Acromyrmex lobicornis Emery — Commensal. Attaphila bergi, or possibly a variety of Huasan, Argentina? (Bruch, 1916). Acromyrmex Synonyjny. — Atta lundi Commensal. —Attaphila : lundi this species, (Guerin) [Brown, personal communication, 1957]. Argentina and Uruguay (Bolivar, The cockroach was found in the nests of the ants sitting on 1901) 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 oi A. lundi. bergi, : Acromyrmex — — niger (F. Smith) Synonymy. Atta nigra Schupp [Brown, p.c, 1957]. Commensal. Attaphila schuppi, Brazil, Porto Alegre (Bolivar, Found outside the nest of the ant and mixed in the columns 1905) : of ants on the march. BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 3I3 Acromyrmex octospinosus (Reich) —Atta octospinosa [Brown, p.c, 1957]. Commensals. —Attaphila fungicola, Panama (Wheeler, Synonymy. 1928) fungus gardens of the ant. Attaphila aptera, Esperanza, Dibulla, Colombia (Bolivar, 1905). Taken : in the Acromyrmex Emery silvestrii — Commensal. Attaphila hergi, or possibly a variety of this species, San Luis Province, Argentina (Bruch, 1916) According to Bruch, : the behavior of this species of Attaphila countered in Huasan in the nests of it differed from A. hergi in size and is identical with the Acromyrmex lobicornis one en- Emery color. Atta cephalotes (Linnaeus) —Attaphila Commensal. Taken 1928) : Attaphila sp., in the Guiana British fungicola, (Wheeler, fungus gardens of the ant. Guiana (Beebe, 1921) British : 7 of 12 queens in one nest had cockroaches hanging on them. Atta sexdens (Linnaeus) Commensal. —Attaphila sexdentis, 1905) Found : Brazil, San Leopoldo (Bolivar, in nests of the ant. Atta texana (Buckley) — — Synonymy. Atta fervens Say [Wheeler, 1910]. Commensal. Attaphila fungicola, U.S.A., Texas (Wheeler, 1900, The cockroach does not feed on the fungus in the ants' nest, 1910) 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 Wheeler (1910) tional, it is of the cockroach are clipped short. Although stated that this is probably accidental or uninten- same invariable Wheeler (1900) had peculiar that Bolivar (1905) noticed the mutilation of the antennae of Attaphila hergi. 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 terms with the ants. tunnels. It is found and maintains very intimate fungus cavities and living in the SMITHSONIAN MISCELLANEOUS COLLECTIONS 314 VOL. I4I Crematogaster limata parabiotica Forel Commensal. —Phorticolea (Caudell, 1923) Bolivia, holiviae, Cachuela Esperanza Collected in joint nests of C. limata and : Compo- notus femoratiis. Solenopsis geminata Commensal. 1917a) : —Myrmecohlatta Collected (Fabricius) wheeleri, from a colony of this ant shores of Lake Atitlan, altitude 11,719 Unknown (Hebard, Guatemala under a stone on the feet. host — Cockroach. Attaphila flava, British Honduras (Gurney, 1937) Because the known hosts of the other five species of Attaphila are ants, : we presume mecine that this species also lives in the nest of some myr- ant. PONERINAE Subfamily Odontomachus Commensal. —Myrmehlattina affinis (Gueiin) 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. — Sphccophila 1906a) living : polyhiarum, French Guiana (Shelford, Shelford stated that on the and spiders floor of the that it was probable that the cockroaches paper nest fed on small fragments of insects were dropped by the wasp larvae feeding in the cells above. Family MEGACHILIDAE Melipona nigra Lepeletier — Commensal. Oulopteryx m,eliponarum, 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 page 318.] : bees. [See comment by Sonan (1924) on BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 315 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 Myrmecoblatta wheeleri Acromyrmex : octospinosus, Ant: Solcnopsis gcminata, p. 314. 313- P- !Nocticola sinensis Attaphila bergi Termites Ants : Acromyrmex p. : lobicornis, Macrotermes barncyi, p. 311. 312. Odontoiermcs Acromyrmex Acromyrmex sp., p. 311. lundi, p. 312. silvestrii, p. 313. Nocticola termitophila Termites Attaphila flava Host unknown, presumably an : Macrotermes malaccen- sis, p. 311. ant, Odontoteruies p. 314. Termcs sp., p. sp., p. 311. 311. Attaphila fungicola Ants : Acromyrmex P- oclospinosus, Ant: Camponotus 313- Atta cephalotes, p. 313. Atta texana, p. 313. Attaphila Nothoblatta wasmanni Oulopteryx meliponarum Bee schuppi Ant: Acromyrmex niger, p. 312. Attaphila sexdentis p. Melipona nigra, : : 314. Camponotus femoratus, p. 311. 313. Crematogaster Attaphila p. Phorticolea boliviae Ants Ant: Atta sexdens, rufipes, p. 312. sp. limata para- biotica, p. 312. Ant: Atta cephalotes, p. 313. Phorticolea testacea Atticola mortoni Ant: Camponotus Ant: Camponotus rufipes, p. 312. rufipes, p. 312. Sphecophila polybiarum Ergaula capensis Wasp: Polybia pygmaca, Termites, p. 314. p. 311. Sphecophila ravana Myrmeblattina longipes Ant: Odontomachus affinis, p. 314. Termite : Coptotermcs ceyJonicus, p. 310. Myrmecoblatta rehni Ants: Camponotits maculatus(?) p. Termite 312. Formica rufibarbis, Formica subcyanea, Sphecophila termitium p. 312. p. 312. : Macrotermes bcllicosus or Macrotermes natalensis, p. 311. SMITHSONIAN MISCELLANEOUS COLLECTIONS 3l6 VOL. I4I OBSCURE ASSOCIATIONS Cockroaches that are sometimes found in the nests of, or in asso- ciation 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. hundred or more lively cockroaches occupied a part A Near the cockroaches McCook of the nest that contained few ants. also found a colony of Terines flavipcs. Ischnoptera deropeltiformis has been found in the company of ants, but cophilous (Donisthorpe, 1900). tera" sp. it is probably not myrme- Mann (1911) found an "Ischnop(probably a species of Parcohlatta) abundant in the nests of, and tolerated by, Camponotus maccooki Forel in California. Dead and common in the nests of common in the nests of Vero- mutilated specimens of this cockroach were "Formicas." "Ischnoptera" sp. was also messor andrei (Mayr) [ = Stenamma andrei]. Hebard (1917) reported that W. M. Wheeler collected Eremohlatta subdiaphana in Arizona as an ant guest. Rehn (1906a; Rehn and Hebard, 1927) reported that Pholadohlatta inusifafa had also been taken by Wheeler from the galleries of a jumping ant, Odontomachiis darns Roger pallens Wheeler; Brown [ = 0. haematodes insularis Guerin var. (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 Dendrohlatta Amazon ant nest in a tree in the Basin. sohrina as taken in an Tivia australica was taken in an ant nest in Australia (Princis, 1954). The male of Compsodes was taken in an ant nest in the Santa Rita Mountains of Arizona (Ball et al., 1942). A male and female of Stilpnohlatta schivarzi minuta were taken lensis in a Princis cautioned that this, migrating column of the ant Myrmicaria nata- Sm. subsp. eiimenoidcs it is Gerst. in Nyasaland (Princis, 1949). premature to derive any inference from Four females of Parcohlatta Ischnomyrmex sp. (Henymph of Parcohlatta virgimca was found in a nest possibly accidental, association. desertae were taken about a nest of an ant, bard, 1943a). A of Formica sp. (Hauke, 1949). Chorisoneura texensis has been found in nests of Florida (Rehn and Hebard, 1916). an ootheca of Aristiger histrio (sp. Karny (1924) ?) in webworm in Malaya found between leaves (Cost us sp.) BIOTIC ASSOCIATIONS OF —ROTH & WILLIS 317 had been stuck together by a thysanopteron, Anaphothrips that He COCKROACHES sp. pointed out that the ootheca would not adhere to leaves that were not stuck together but would Sein {in the ground. fall to Rehn and Hebard, 1927) in Puerto Rico found Aglaop- tcryx fades in abandoned cocoons of Megalopyge kriigii (Dewitz) webbed together by caterpillars and in abandoned spiders Wolcott (1950; and in Rehn and Hebard, 1927) also found A. fades in the empty cocoons of M. krugii and in the larval tents of Tetralopha scabridella Ragonot on Inga vera (coffee shade tree) and and in leaves nests. ; "Where there are no butterfly-nests, abandoned spider-nests on the leaves of other forest trees." Cotton (in Wolcott, 1950) found the type of Aglaoptcryx absiniilis also living in the abandoned cocoon of M. krugii on bucare trees in Puerto Rico. Wolcott (1950) reported that Plectoptera dorsalis, Plectoptera infulata, and Plectoptcra rhahdota 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 (Moschler) in the leaves of Petitia domingensis in the mountains of Puerto Rico. Sein {in Rehn and Hebard, 1927) had collected P. rhahdota in the nest of larvae of T. scabridella. Wolcott (1950) reported that Nyctibora lutzi had been found in a large rotten stump associating with " 'come j en' termites [Nasutitermes costalis (Holmgren) ], yellow wood-ants and rhinoceros beetle grubs." Rehn and Hebard (1927) found S'miblcrastes 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 it lives in other structures remains to be determined." In Virginia Cryptocercus punctulatus has been found living in the same galleries with Reticuliteruies sp., and on the Pacific Coast been found occupying the same log with Termopsis sp. it has (Cleveland et 1934). Shelford (1909) found one male and one female of Balta platysov.m al., in a nest of a spider of the genus Phryganoporiis and assumed a sym- Chopard (1924) recoxde^d Marcta acutivcntris hora 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 obliquetntncafa 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 rebiotic association. ; ; treat in the nests. SMITHSONIAN MISCELLANEOUS COLLECTIONS 3l8 VOL. I4I McClure (1936) obtained a large nest of Vespula maculata (LinIn it were living 65 [ = Vespa maculata] in March in Illinois. nymphs of Parcohlatta pensylvanica, 3 spiders {Philodromus pernix naeus) Blackwall), 2 immature spiders (Drassus sp.), and 6 mites. Balduf (1936) observed four individuals of Parcohlatta pensylvanica in a nest of Vespula inacidafa; 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 Parcohlatta and the wasp, it is well to recall a statement by Roth- and Clay (1957): "A commensal relationship is potentially even more dangerous than a merely social tie, for by nature it is more schild intimate. One set. The closer the association, the more easily is the balance up- mean advantage partner can then suddenly take a of the other." Cockroach nymphs may enter bees' nests where, according to Ima(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 mura nest ; the bees are not disturbed by the cockroaches. Cockroaches that have been found listed on pages 23-25. in the burrows of vertebrates are Paulian (1950) found immature cockroaches in the nests of birds (Ploceinae) in Madagascar and Ivory Coast. All nests of Fondia sp. Madagascar contained many cockroaches, and Paulian bewas 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 fades examined in lieved that the blattid ; "in the nests of the grey kingbird, in the region of the Cartagena 'living among the twigs.' " In Trinidad, Kevan Lagoon [Puerto Rico] , found a male of Blaherus discoidalis Kevan, 1955). in a bird's nest (Princis and BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 3I9 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 Peri- planeta 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. associations exist in which many on the feces of man and domestic and other animals (Roth and Similar of the domiciliary cockroaches feed 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 close proximity; one rarely meets several in 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 and that Australian species of Cutilia 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 (191 5) attributed to Tepper the statement that cockroaches, like Epilampra notahilis, which are found out-of-doors in Australia, are carnivorous and feed on caterpillars and other soft-bodied insects the foliage of shrubs, [ = Drymaplaneta, ; but Froggatt (1907) believed that this alleged behavior needed confirmation. A number of observations have been recorded which indicate that sometimes cockroaches may be (191 5), nymphs of the cockroach gentine ant Iridomyrmex humilis. predatory. Pseudomops According to Ealand cincta fed on the Ar- In the laboratory, Eurycotis flori- dana 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 ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 320 VOL. I4I long. Aiinandale (1910) described the destruction in Calcutta of ter- many mites by Periplaneta americana. During a heavy rain storm mites flew into the dining which seized them with mens. If disturbed, room and were their ter- upon by the cockroaches mandibles and began to gnaw their abdoset 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. Blattclla 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 (Uichanco, in Williams et al., upon leafhoppers 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, Blatgermanica, and/or Periplaneta americana) dislodge and eat the tclla Sonan (1924) reported that cockroaches {P. americana and P. australasiae) devoured the egg mite Pirneliaphilus podapolipophagus. clusters and first instar larvae of Prodenia litula and the first instar larvae of Attaciis atlas which were being reared in the laboratory. Lederer (1952) stated that Periplaneta americana ate reptile eggs in am Main. Pettit (1940) stated that cock- the aquarium at Frankfurt roaches "are said to have destroyed a large colony of dermestids used to skeletonize carcasses at the University of DeFraula (1780) believed that his Kansas." silent "gryllon" [obviously 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. Blatta orientalis Gilbert from his White (1905 ed.), writing in England in the late i8th century, "Poda says that these [Blatta orientalis] and house crickets stated that will not associate together however, in ; but he is August 1792 White noted mistaken in that assertion" that "Since the hlattac 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 kitchen in France. relative He orientalis and Acheta domesticiis in an old suggested that the cyclical fluctuations in the abundance of the cockroaches and crickets might be caused by reciprocal predatism with one species more susceptible than the other BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 32I MalHs (1954) has stated that crickets prey on other on one another. Lheritier (1951) had also observed crickets becoming rare in bakeries in France, having been superseded everywhere by B. orientalis; however, he doubted that Joli vet's hyat certain stages. insects as well as 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. pothesis Platysosteria novae seelandiae New was found under the bark of trees in Zealand devouring bugs (Walker in Shelford, 1909b). For years it has been believed that cockroaches feed on bedbugs {Cimex lectularhis 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 The all the little basis for this belief parasites they can capture" may have Webster (1834) who wrote that (Gaul, 1953). originated with a statement by bedbugs disappeared aboard "H.M. Sloop Chanticleer" when cockroaches made their appearance. New- man (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 cock- roach 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 numbers the bugs apparently were not attacked and their numbers increased greatly over a period of a few weeks (Johnson and in large ; 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 1916). In laboratory experiments Gulati (Cornwall, (1930) found that Peri- SMITHSONIAN MISCELLANEOUS COLLECTIONS 322 VOL. I4I planeta americana ate young bedbugs which had soft, blood-filled ab- domens adult bedbugs with harder exoskeletons sometimes were ; The maximum number of bedbugs rejected. was eaten by a cockroach 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 oricntalis or that bedbugs are eaten to any extent by them. little The existing evidence indicates that there is basis for the often repeated statement that cockroaches destroy bedbugs As Lorando (1929) in nature. 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 we have found in which cockmentioned together is a paper by Whitroaches and Phlebotomus are tingham and Rook (1923) they fed ground-up cockroaches to larvae these statements ; the only reference ; Wharton (1951) reported that cockroaches and other predators attacked mosquitoes knocked down by insecticides and affected the number recovered. oi Phlebotomus papatasii. 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. Cockroaches will INTRASPECIES PREDATION Those who have reared cockroaches in the laboratory doubtedly seen cannibalism occur in the cultures. been observed among the common as well as laboratory colonies of have un- Cannibalism has domiciliary species of cockroaches Leucophaea maderae (Scharrer, 1953), and Blaherus craniifer^^ (Saupe, 1928). Edmunds (1957) 11 The West Indian "Blabera jusca Brunner" of Saupe (1928) is obviously Blaberus craniifcr as can readily be seen from a comparison of Saupe's figures of the pronotal shields of his species with the descriptions by Hebard (1917) of B. craniifer and Blaberus atropos (Stoll). The Chilean B. jusca Brunner is a junior synonym of B. atropos (Stoll), a South American insect (Hebard, 1917). BIOTIC ASSOCIATIONS OF COCKROACPIES— ROTII & WILLIS reported that cannibalism was common 323 colony of Peri- in a laboratory female were planeta hrimnea and that egg capsules deposited by a often eaten by the other cockroaches. their Periplaneta americana occasionally ate other cockroaches and (Lederer, species own their of members attacked also and oothecae the killing of male 1952). Griffiths and Tauber (1942) recorded American cockroaches by females of the species "One female was : and attacked each new male as he was introduced Jvlost of such males had molted less than 2 days container. the into themselves previously. Older males were more capable of defending adequate though Even females." cannibalistic these of attacks against especially vicious be present, females of Periplaneta americana may eat their own eggs (Klein, 1933). Some females may regularly eat their ooTo thecae as soon as they are dropped (Griffiths and Tauber, 1942). it before attacked be must be completely eaten an ootheca generally cockthe capsule, the of has hardened. If a hole is eaten in one side Freroach may devour the eggs and leave a portion of the ootheca. occurs this quently only the keel or a part of the keel is eaten and when because the eggs fail to hatch and usually do not complete development adults of of the rapid loss of water (Roth and WilHs, 1955). When both males food, of deprived were aiistralasiae and P. americana P. and females ate newly deposited eggs and, finally, the females ate the food may males (Sonan, 1924). Parcohlatta virginica in laboratory cultures also may eat part of its deposited in this species only the soft end of the recently oothecae ; ootheca was eaten (Roth, unpublished data, 1957)Cros (1942) observed oothecae-bearing females of Blatta orientalis attempting to attack and kill males of the same species which were these males were then eaten by the females. Cros also observed eaten by injured and recently molted nymphs of B. orientalis to be mate ; others of the same species. Blattclla (1940) noted that cannibalism in his culture of germanica occurred only when the insects were molting. Adult insects nymphs. attacked the molting cockroaches more often than did the Pettit However, nymphs after the fourth instar occasionally set upon other their molting nymphs. First- to third-instar nymphs rarely victimized stadia later the instar third than older all were mates. The victims were progressively more subject to attack, and molting adults suffered poputhe greatest mortality. No direct correlation was noted between ; lation density and cannibalism. SMITHSONIAN MISCELLANEOUS COLLECTIONS 324 VOL. I4I German cockroaches may attack newly molted nymphs of their own kind and cause them to deflate (Gould and Deay, 1938). Lheritier (1951) has observed the hatching nymphs of B. germanica being devoured by their congeners even before they have left the ootheca. Nauphoefa 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, A'^. cinerea may kill and eat the cypress cockroach, Diploptera punctata (Illingworth, 1942; Fullaway and Krauss, 1945). Bunting (1956) stated that species of Neohlattella are omnivorous with carnivorous and cannibalistic tendencies. An adult female and eaten by Neohlattella sp. in captivity. A A'^. celeripes, was killed and partly other males species. by two of the same eaten Panchlora sp. was killed male, provisionally identified as The factors influencing the extent among of cannibalism roaches are not completely known. According to Wille cock- 1920) hunger was not the cause of cannibalism in Blattella germanica. Wille ( claimed that the tendency toward cannibalism increased temperatures and decreased at low temperatures. Pettit high at (1940) also noted this efifect. Gould and Deay (1938) stated that under crowded laboratory conditions, when there was a scarcity of food, cannibalism among Periplaneta americana was and those unable lar observations. common. The injured cockroaches molt were often eaten. Adair (1923) made simiUndoubtedly, conditions of crowding, availability to of food, temperature and other factors practically all influence cannibalism, but no experimental work has been done on this subject. comparison with the above positive examples of cannibalism, that both Saupe (1928) and Roeser (1940) observed no cannibalism during extensive studies with Pycnoscelns 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 It is interesting, in insects. XVII. AMONG COCKROACHES ASSOCIATIONS Besides preying on their own species or on other roaches exhibit additional symbiotic relationships These relationships are offspring, ( i ) blattids, among the familial associations of parent (2) gregariousness, (3) intraspecies fighting, and (4) interThere are and (5) interspecies antagonism. some inconsistencies between observations made on the same species compatibility, cock- themselves. species BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 325 by different workers, which only further observation and experimentation will explain. is Some of the reported observations are unique; this especially true for the feral species. formation, is it Because of the paucity of impossible at this time to make in- 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 ootheca 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 oothecae, to the duration of embryogenesis in the so-called viviparous species, a period of over a month or more. Haber (1920a) observed groove in a piece of The ootheca her ootheca. a female of Periplaneta americana chew a pasteboard into which she attempted to deposit failed to adhere to the shallow hole and fell to the floor. After several futile attempts to replace the ootheca in the hole, the female finally left the coated it During tgg case on the floor of the cage and with an oral secretion to which she attached bits of trash. operation she chased other females this ventured near the site. away when they Qadri (1938) described the behavior of the female of Blatta orientalis in concealing her ootheca in a hole that she dug in sand ; she deposited the egg case in the hole, coated Rau (1943) it with saliva and sand, and then detail how females of P. americana and B. orientalis covered their wood dust or sand in holes they had prepared in the refilled the pit. described in oothecae with substrate. Both species placed a sticky oral secretion in the holes and then deposited their oothecae therein. After coating the oothecae with more sticky secretion, the females adjusted the oothecae so that the were uppermost and then carefully concealed the oothecae with the excavated debris. Both females spent over an hour in the act. Rau (1924) previously reported that of 90 oothecae 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 keels 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 cap- was deposited in the secretion and covered with additional froth, which hardened into a very strong cement. Some females spent as sule SMITHSONIAN MISCELLANEOUS COLLECTIONS 326 long as two hours coating the capsule after was it VOL. I4I deposited. not stated whether the ootheca was otherwise concealed. It was The female remained with her body over the ootheca for several hours and drove away other cockroaches which approached. Sonan (1924) observed that Periplaneta americana and Periplaneta australasiae spent and that if from 40 minutes returned again to complete strate in to an hour covering their oothecae, the females were frightened which to deposit it. its As away from this activity, they well as excavating holes in the sub- oothecae, P. americana also avails itself of readymade crevices of appropriate size (Ehrlich, 1943). Species of Epilampra Malaya were in said by Annandale (1900) to deposit above the water line. their oothecae in crevices in floating logs just However, Shelford (1906) stated that four genera (including £'/'f/ampra) of the subfamily Epilamprinae are "viviparous," in which event the females would carry their oothecae within their bodies during embryogenesis and would not place the oothecae in crevices in The female of Cryptocercus punctulatus was observed logs. make a from where to groove in a piece of w^ood, then carry her ootheca 6 inches she had dropped it and place it in the groove she covered the ootheca ; was so that only a portion visible (Cleveland in Cleveland et al., 1934). Dr. W. L. Nutting (personal communication, 1954) collected a number of oothecae of C. punctulatus in the field and found each one almost completely sealed with off The bits of wood in a deep groove in the was visible but the was well camouflaged. He stated that "The adult pair usually frequents the chamber at this time, while their broods of previous roof of a chamber in a log. keel of the ootheca rest years occupy neighboring galleries." Berland (1924) observed a female of Lohoptera decipiens filling a hole (the abandoned nest of a hymenopteron) with earth that she carried in her mouth ; he later found her ootheca behind the earthen barricade which she had erected. In summary, the following species of oviparous cockroaches have been observed concealing their oothecae (only those references not previously cited are given) Blatta orientalis; : Cryptocercus punctu- Epilampra sp. EuryLohoptera decipiens; Balta cotis floridana (Roth and Willis, 1954a) scripta, Methana curvigera, Methana marginalis, and Methana caneae latus; Ectobius Sylvester (Harz, 1956, 1957) ; ; ; (Pope, 1953a) ; Pelmatosilpha marginalis, Pelmatosilpha purpura- and Nauclidas nigra (Bunting, 1956) Periplaneta americana (Haber, 1919; Adair, 1923; Sein, 1923; Nigam, 1933; Gould and Deay, 1938; Rau, 1940a) Periplaneta australasiae (Girault, 1915b; Spencer, 1943; Pope, 1953) Periplaneta brunnea (Roth and Willis, scens, ; ; ; BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 327 unpublished data, 1958) Periplaneta fuliginosa (Gould and Deay, Supella supellectilimn Periplaneta ignota (Pope, 1953) 1940) ; ; ; (Flock, 1941). Undoubtedly other oviparous species that drop oothecae long before the eggs hatch also ceal the oothecae by placing them make some attempt their to con- them with in crevices or covering debris. Sometimes the oothecae are deposited but not concealed. Hafez and Afifi (1956) reported that in Egypt Supella supellectilium attaches its ootheca to a suitable substrate with a gummy but leaves the tgg capsule otherwise exposed. We oral secretion (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 oothecae in places which are dry and raised above the ground, although rarely one also find some oothecae scattered on the ground. For lack of suitable material females of Periplaneta americana sometimes did not conceal their oothecae (Nigam, 1933). Frequently in laboratory colonies P. americana merely drops the oothecae 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 ootheca anywhere (Bunting, may 1956). Rau (1940) stated that the female of Parcohlatta pensylvanica does not conceal her ootheca. However, Gould and Deay (1940) stated that this species deposits its oothecae loosely behind bark. Ellipsidion affine and Ellipsidion australe attach their oothecae to bark or the underside of leaves but apparently them (Pope, 1953a). The females of most of make no attempt to conceal the above species have no further familial 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 association with their offspring. such, after hatching. oothecae This may have been the eggs of the encountered first among is not unexpected, as several additional deposited by these oviparous females before oothecae hatch. However, a different behavior is form a second ootheca until have hatched (see below) and in the so- species that do not after the eggs of the first called colonial species. Shaw (1925) and Panesthia reported that in Australia both Panesthia australis laevicollis appear to live in families, and usually finds a pair of adults associated with from 12 to 20 that one nymphs in SMITHSONIAN MISCELLANEOUS COLLECTIONS 328 different stages of development ; he continued, "it is VOL. I4I 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 and 10 to 20 nymphs. A family communities of a pair of adults related colonial species, Cryptocercus punctu- both sound and rotten logs in colonies consisting of a latus, lives in two or Chopard pair of adults and 15 or 20 nymphs, probably representing (Cleveland three broods et al., 1934; Cleveland, 1948). (1938) has cited this association as an example of gregariousness, however, the presence of only one pair of which it may well be ; more intimate relationship. and certain other genera with adults in each colony suggests a Among species of Blattella similar reproductive habits the female carries her ootheca clasped in her genital cavity with the posterior portion projecting behind her. normal ootheca genesis and is is Each carried for approximately the duration of embryo- not dropped until, or shortly before, hatching. We have 65) newly hatched nymphs of Blattella vaga crawl over the body of the mother who stood quietly near the dropped seen (1954, ootheca ; fig. this female raised her wings and some of the nymphs crawled under them onto the dorsal svirface of her abdomen. The on the grease covering the mother's body. The 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 seemed association was nymphs to feed of 5. germanica remained together without shelter in a single, 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 oothecae together. In groups of 8 to 12 nymphs there was a perfect intermingling of the offspring from the two different females. sparse group. It is number among the so-called viviparous cockroaches that the greatest of observations have been made of postparturient associations between female cockroaches and their offspring. The females of these species carry their oothecae 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 oviShelford (1906, 1916) re- parity and viviparity in the Blattaria.) ported that he had captured a female of Psendophoraspis nebitlosa in Borneo with numerous young nymphs clinging to the undersurface of BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 329 He also recalled that there was in the Hope Museum (Oxford) a female of Phlebonotus pallens to which the following 'Ceylon carries its young beneath its wing label was attached: covers. 1878.'" Pruthi (1933) found in South India another female of P. pallens which was carrying over a dozen young nymphs on her her abdomen. *' . . . back beneath her wings. In his paper Pruthi reproduced a photograph nymphs in place on the back of of this specimen with the light-colored the female. Hanitsch (1933) reported having seen a museum speciPhilippine Islands, of the apterous female of Peris- men from Luzon, phaerus glomeriformis with nymphs face ; he also reported having seen a still clinging to her undersur- museum specimen of a female of from Australia with four young clinging to the upper side of the apex of her tegmina and six to the ootheca 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 en elongate face and specialized Ellipsidion variegatum Karny (1925) also observed that at the slightest alarm the young of some species of Phoraspidinae creep under the dome-shaped galeae. front wings of the mother. The newly hatched young of Lcucophaea maderae have also been seen congregated under the mother on several occasions. Sein (1923) nymphs of this species gather under accompany her at night in her excursions in search of food. Pessoa and Correa (1928) reported that "During the first days the free larvae hide under the adult cockroach which becomes restless stated that after being born, the the mother and usual slow gait." Wolcott (1950) stated and active in contrast to that "They are not only gregarious, but its 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 fiisca has been observed late summer and early winter in familial groups beneath loose during "Each party 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 fe- bark, under stones, and in dead leaves (Skaife, 1954) consists of a number of black young males of Perisphaerus young ; ariiiadillo : ones, together with one, surrounded by pale, yellowish-white similarly he had observed that Archiblatta hoevenii was found ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 330 VOL. I4I mostly in colonies made up of mothers and their young. The duration of these associations is not known. newly hatched nymphs of Blaberus body of their mother and stated that this is as pronounced a case of brood care as Zacher had observed with Pycnosceliis surinamensis. Nutting (1953) stated that "A degree of maternal solicitude is exhibited by this roach [5. craniifer], for many times I have observed the female to remain motionless for an hour or more with her unpigmented Saupe (1928) noticed that the craniifer (see footnote 11, p. 322) collected together under the We, brood clustered around and beneath her body." served similar behavior in laboratory colonies of J5. too, have ob- craniifer and Leitcophaea 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. lished data, 1958) have seen tcntosa also stay near their We (unpub- young nymphs of Gromphadorhina pormother for some time after birth; the produced a characteristic hissing sound when she 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 (WiUis et al., 1958). Bunting (1956) mother at this time was only slightly disturbed 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 Whole birth. families of cockroaches may be (Ohaus, 1900). Hebard (1920) observed a colony of adults and young of Dendrohlatta sobrina on a tree trunk in the Panama Canal Zone. Whether any of the above associations exemplify maternal care for found in bromeliads in Brazil the newly hatched young is beyond placing her eggs passive. The questionable. in first-instar The behavior of the mother, a suitable location, seems to be entirely nymphs are the active partners in these 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. associations, , GREGARIOUSNESS Casual statements that cockroaches are gregarious are often enThere has been some argument to the countered in the literature. BIOTIC ASSOCIATIONS OF COCKROACHES— ROTH & WILLIS numbers of these effect that large insects seeking the 33I same environ- ment in a Hmited space would appear to be gregarious, whereas there Reactions of cockis probably no true social tendency (Rau, 1924). do result undoubtedly environment the in stimuli certain roaches to has Chopard as However, (1938) individuals. of in aggregations pointed out, it is difficult to assign the respective parts played in asand by gregarious instincts. sembling by the attraction of the milieu Chopard (1938) also stated that Orthoptera with a gregarious tendthem, on the contrary found rarely isolated; one finds ency are 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, and offers if one places a large number of cockroaches in a container ; them similar shelters tubes, one finds that the individuals will assemble in one of the tubes, ignoring claimed that in Blattella germanica greothers. Pettit nearly the composed of cardboard all (1940) gariousness seemed to depend on the mutual attractiveness of body warmth. secretions as well as a thigmopositive behavior and love of Ledoux (1945) has studied experimentally gregariousness and so- Blatta orientalis and Blattella germanica. He containing also found that the cockroaches tended to collect in shelters reother cockroaches. He concluded that group formation is not the cial interattraction in a social phenomenon, and that interattraction is mainly olfactory, conditioned by (i) positive chemotaxis to odors emitted by the cockroaches themselves, (2) positive hygrotaxis, and large groups are not stable and (3) thigmotaxis. He found also that groups. tend to break into smaller Gregariousness in the Orthoptera varies in intensity according to sult of chance, but is the species and within a species according to the age or physiological by state of the insects (Chopard, 1938). This is well exemplified 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 cock"In the corroaches in workers' living quarters in the Ural region in a dense walls the ners near the stove, the cockroaches covered cockdead carpet." After fumigating he collected about 135,000 : We roaches from one barracks and about 475,000 from another. have reviewed a few other examples of heavy infestations in our 1957 (a) paper. SMITHSONIAN MISCELLANEOUS COLLECTIONS 332 VOL. I4I 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 harmony. Young nymphs have been underneath adults and crawling over the adults as sizes live together in perfect noted in clusters they wander about in rearing jars." In the monkey house of the Ham- burg 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 Frankfurt at individuals ; am Main, P. americana rested in groups of 20 to 30 it was remarkable that the "herd" divided he stated that 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 itself into ; these manholes were heavily infested with 100 or in each. more cockroaches Other heavy sewer infestations have been reviewed in our 1957(a) paper. Ehrlich (1943) has stated that Periplaneta americana exhibits social For instance, cockroaches of various ages inhabit a fairly behavior. 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 the imagos drive older nymphs from their and the older nymphs drive out the younger ones, until cracks, depending on their size, are occupied by various age groups available living space ; resting places, all 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 of different ; the larger ones that could shelter more insects. roaches ceased to bite and fight each other Finally, the cock- when they crowded to- gether in the face of danger. Of Blatfa orientalis Marlatt (191 5) 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 re- BIOTIC ASSOCIATIONS OF COCKROACHES mained almost constantly in —ROTH groups during the but less SO during the third instar. He first & WILLIS and second 333 instars, believed that the aggregations of young occurred because they could occupy narrow crevices where the larger insects could not penetrate. older nymphs and At usual room temperatures the adults lived completely isolated, but at certain temperatures they gathered together in large, tightly pressed groups. Supella siipellectilkmi 1940). The smaller is said to be gregarious nymphs aggregate containers, but the older ones remain (Hafez and found alone Afifi, ; in small separate 1956). Leucophaea maderae is (Gould and Deay, groups in rearing from one another and rarely sociable in their favorite hiding places, hills of these cockroaches Wolcott (1950) also gregarious. Annandale (1900) observed that can be seen hanging together (Sein, 1923). maderae is "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 F. australasiae and Syniploce 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 fioridana fairly swarmed under the coquina boulders in the woods, in stated that L. in the 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. fioridana with the mature individuals. Hebard (1917) in his discussion of Lattiblattella rehni again mentioned finding frequent colonies of E. floridatm in Florida. He found many specimens of Blaberus craniifer under boards on Key West. He found Parcoblatta lata numerous under bark of dead pine trees in Alabama. However, Dowdy (1955), in an also the ground at ecological study of oak-hickory forest in Missouri, stated that ''Par- coblatta [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 rub- bish (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 servers also reported that all stages of development." Aspiduchns borinquen was found These obin Puerto SMITHSONIAN MISCELLANEOUS COLLECTIONS 334 VOL. I4I Rico in a limestone cavern by thousands in the grass and on the walls. 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 of these and, we infer, confirm the species. Rehn and Hebard (1927) J. in their numbers account of Simblerastes jammcanus reported finding in a termite nest. that Diploptcra punctata is it in Pemberton and Williams (1938) stated of gregarious habits in Hawaii. Saupe (1928) observed a strong "herd instinct" in all age groups of Blaherus craniifer. Bunting (personal communication, 1956) stated that large nymphs and adults of Blaherus discoidalis "congregate in cracks or on the underside of some low object. keep in close narrow The younger nymphs 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 of it, physiological or psychological effects of gregariousness, or lack Landowski are interesting aspects of the basic phenomenon. (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 i, 2, 4, 8, and 16 in jars of identical size and shape. Landowski found that (i) 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 produce an adult to in complete isolation extended the time required insect ; and (3) the mean weight of the adult sect was, generally, in inverse proportion to the raised together; number of in- nymphs isolated insects usually attained the greatest adult weight. Similarly, Griffiths and Tauber (1942a) found that isolation ex- tended 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 gerrnanica take longer to mature than those reared in groups. gerrnanica that there is Wallick (1954) found indications in B. an inverse relationship between population density and individual weight ; as the population decreased the weight BIOTIC ASSOCIATIONS OF He increased. also noted COCKROACHES —ROTH & WILLIS 335 an inverse relationship between population density and life expectancy in this species. We (Willis et 1958) have confirmed the above observations that and Periplaneta americana com- al., Blattella germanica, Blatta orientalis, plete nymphal development than individually. We in less time (loc. cit.) also following additional species matured groups : lectilium, when reared in groups rather found that nymphs of the more quickly when reared in Eurycotis floridana, Periplaneta fuliginosa, Supella supel- 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 madcrae. 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 "When two cockroaches, even of different and P. australasiae) were kept together, this in successive 24-hour cycles species {P. americana : depression did not appear to set in so readily." The associates ap- parently kept each other active. Isolated females of Periplaneta americana can be conditioned to less time and fewer errors per trial than when 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 run a simple maze with paired or 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 individuals. SMITHSONIAN MISCELLANEOUS COLLECTIONS 336 species are indications of gregariousness. However, VOL. I4I until gregarious- we ness has been proved experimentally for each species, concede that reactions to environmental stimuli might be sufficient to bring about some of the observed groupings without any interaction between in- dividuals. 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 Formosa hashi (1940) observed that in not have a tendency to throng together. in close proximity. Blattella Taka- humbertiana does Rau (1947) stated that the 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 adults of Ischnoptera deropeltiformis the restricted quarters of the cage. (1955) did not find Parcoblatta sp. to As mentioned above. be gregarious in the Dowdy field. INTRASPECIES FIGHTING among cockroaches of the same species over food 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. Fighting occurs or shelter or between males. Ehrlich (1943) stated that individuals of Periplaneta americana ward off intruders by spreading their wings and that are feeding will 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 to fight and bite each other. was sufficient to cause the cockroaches Biting and fighting also occurred when individuals of this species defended their daytime hiding places. position of attack is A 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 protrude ; when the insects jump at each other, they mouth may wound parts each other severely in the soft parts of the body. Fighting between sexually excited males resulted in injury to their legs, wings, cerci, parts of the body. and other Frequently an insect that could no longer defend BIOTIC ASSOCIATIONS OF itself was COCKROACHES —ROTH & WILLIS Lederer (1952) also made similar but killed. less 2)Z7 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 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 oi B. germanica that were crowded together quickly set substantiate such voracious attacks, although he upon, but did not always cage. When kill, other cockroaches introduced into their 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 Nauphoeta frequently observed aggressive behavior between males of cincrea, 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 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 species of Orthoptera, ; 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 of these associations are definitely ecological, particularly that certain among the SMITHSONIAN MISCELLANEOUS COLLECTIONS 338 feral species. As might VOL. I4I 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 detrees, and beneath corded wood. Adair (1923) stated that in house in Egypt Periplaneta americana, Blatta orientalis, and Blat- cayed his 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. tella Gould and Deay (1940) observed that in the Purdue University greenhouse Periplaneta fuUginosa 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 fuUginosa, 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 oi B. orientalis found in Tyler manholes were associated with larger colonies of F. americana. True breeding colonies oi B. orientalis appeared to be present since all the developmental stages were taken. The same situation existed with the P. fuUginosa and the two species of Parcoblatta. Larger colonies of P. americana were associated with the other species in each case. From our limited ob- servations the two species always appeared to be perfectly compatible." Eads et al. (1954) had found Periplaneta fuUginosa in three manholes, Parcoblatta holliana in one manhole and Parcoblatta pensyl- vanica in one manhole. were We assume spacially discrete so that they that the groups of each species 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 BIOTIC ASSOCIATIONS OF COCKROACHES Periplaneta americana on a quarter-boat. forficuHd Marava \_ = Prolahia] —ROTH They arachidis & WILLIS 339 also noted that the appeared (Yersin) 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 "The females were all taken in cupboards where Blattella germanica was found in swarms." The association in human habitations that of .S. supellectilium and B. germanica has been reported also by Sein Shaw (1924), Australia; Mallis (1954): "German and brownbanded roaches were often found in the same crevice."; Anonymous (1958), Texas; and Anonymous (1958a), Georgia. Gould and Deay (1940) stated that other species of cock(1923), Puerto Rico; may be found with S. supellectilium Yet Shaw (1925) stated that "when Supella supellectilium Serv. invades places already occupied by Blattella germanica L., it tends to oust the latter." roaches, especially B. germanica, in the same part of a building. 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 (191 5) in Hawaii found Symploce hospes associated with Nauphoeta cinerea, Graptoblatta notulata, and Diploptera punctata. Hebard (1917) found Aglaopteryx diaphana in a bromeliad on a and numerous numerous Aglaopteryx gemma under signs on long-leaf pines in Alabama with occasional specimens of Parcoblatta lata. In Virginia he found Parcoblatta uhleriana in a forest tree in Jamaica together with Nyctihora laevigata Cariblatta insularis. He also found decaying chestnut log with Cryptocercus punctulalus. In Florida he found Latiblattella rchni with Eurycotis floridana and, more with Periplaneta australasiae under bark of pine trees. In rarely, 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 : SMITHSONIAN MISCELLANEOUS COLLECTIONS 340 Jamaica They in VOL. I4I company with Neohlattella eurydice and N eohlattella dryas. most of the associations cited by Hebard (1917). (1923) reported that he found in Germany four species of also Hst Ramme Ectobius (lapponicus, hicidus, 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 gennanica, Periplaneta americana, and Periplaneta hrunnea 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 only. stated in my published note, the roaches were at The 83 Periplaneta subsequently first classified to genus identified to species represented 28 differ- from houses and 17 from privies, all in Pharr, Texas. Tabushows first that P. americana was taken only once in a house and that P. hrunnea 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. brimnea, however, was trapped with P. americana 2 of the 4 times it ent collections, 11 lation of the data 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 as- May 14 to June 22 In the houses, Periplaneta and Blattella were caught in the same jar sociated privies in Pharr, Texas, in weekly intervals (from [1948]). 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 (14.8). number calculated from the row and column frequencies 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 data. There is satisfactory if based on more extensive also a possible objection in that the traps were operated for at 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 least overnight, 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 BIOTIC ASSOCIATIONS OF COCKROACHES litter in —ROTH & WILLIS 34I Jamaica (Rehn and Hebard, 1927). In Florida Periplaneta was often taken in company with Pycnoscelus surina- australasiae mensis 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 Marlatt (1915) stated "Rarely do two of the domestic species occur together in the same house. Often, also, of two the same habitat niche. may neighboring districts one the other a distinct species is be infested with one species, while in the commoner one. The different species 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 are thus seemingly cockroach [Blattella germanica?] has everywhere driven before great congener [Blatta orientalis?]." Yet in it its 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 Wille (1920) in Germany found usually only one species of cockroach in a house. Yet when he placed B. orientalis and B. ger- size. manica together, there were no reciprocal attacks even by hungry in- dividuals. Wille concluded that because of their greater speed, smaller size, number of greater eggs, and faster development, the cockroaches eat the available food and so favorable for the oriental. in which the opposite is make German the environment un- However, he noted that cases may be seen Laing (1946; British Museum also possible. [Natural History], 1951) observed that in the British Isles B. orientalis seems recent years ; to it have was lost its dominant position to B. germanica stated that these species are not as a rule in found together and that the greater rapidity of breeding and ability to climb of B. germanica, as well as the layout of modem some Ledoux (1945) buildings, are of the factors that favor the spread of B. germanica. 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 SMITHSONIAN MISCELLANEOUS COLLECTIONS 342 nymphs oi B. geTmanica with which are fourth- and fifth-instar VOL. I4I nymphs of B. 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 orientalis, all practically of equal size, ; 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 Pope (1953) thought it doubtful in QueensWolcott (1950) stated that "The larger and more powerful Blattella germanica, but land. domestic cockroaches, Periplaneta americana (L.), P. aiistralasiae (F.) and P. hrimnea Burmeister have very hind in Puerto Rico in competition with the definitely fallen German little be- roach." Pessoa and Corea (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 Chopard (1932, 1938) company with P. apparently also driven out by P. americana. stated that the oriental cockroach does not exist in 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 sumably Periplaneta australasiae]. statement is Shaw (1925) in the same to displace our indigenous species" [pre- Tillyard (1926) noted that this incorrect as neither species stated that in Australia is native to Australia. "When both species places, australasiae Fabr. will probably be americana L." Local fluctuations live Yet together found gradually 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 BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 343 seemed to have overcome the American which had been more numerous in former years. In conclusion, we emphasize that many of the above observations who have are merely tentative impressions gathered by workers watched many species of cockroaches in nature. Obviously, additional observations coupled with appropriate experimentation will be needed association and to reto disclose the true structure of each presumed greatly indebted to are solve apparent discrepancies. Although we information, the cited authors for their contributions to the known exlaboratory designed we anticipate that future results of cleverly periments will do much to dispel the uncertainty that still surrounds our knowledge of the relations of the Blattaria to each other. DEFENSE OF COCKROACHES AGAINST PREDATORS XVIII. Irritating or repellent secretions provide to widely unrelated defence . many animals belonging groups with a more or less potent means of . . be seen that this method of defence does not rest merely attribute, but upon an active emission unpalatable substance which, since it occurs when the It will upon a passive unpalatable of the animal is seized or threatened by an enemy, enforces its effective- highest development we find different forms whose project specialized habits and modified structure enables them to secretion at the enemy, and thus to discourage attack. ness. In its 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 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 sughowever, there gested that the insects' odor made them repugnant am Main, ; are a number of positive records of monkeys feeding on cockroaches (see pp. 284-286). may escape capture by predators through evasive beprotective coloration, mimicry, or secretion of concealment, havior, Nocturnal cockroaches may avoid predators materials. malodorous day (Crawford, I934), but nocturnal predthe during that are active Cockroaches ators are apparently quite successful in capturing cockroaches. Some ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 344 VOL. I4I 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. cockroaches 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 rohustus, 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 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) obit served that Styphon hakeri Therca trees, etc. it, nuptialis, The is found in humus and rubble in the Dutch is sluggish in the open, but wedges into the found in India, conceals itself at the roots of fig West Indies where "It humus quite quickly." small hairs on or at any rate to render its it elytra retain sufficient dust to conceal inconspicuous, when not on the wing (Annandale, in Chopard, 1924c). Rehn and Hebard (1914) observed that the nymphs of Blaherus craniifer ^^ at Key West, Fla., "were usually found half buried in loose they remained motionless, looking damp much earth under boards, where like lumps of earth (with which they were usually much dusted) until disturbed." Hebard (1917) reported of Monastria higuttata 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." apparent from the numbers of predators reported herein that It is many animals are not deterred by the odorous secretions of cock- roaches ; these secretions, because they may seem repugnant are often claimed to be repellent to predators. points out that "There are many 12 The p. species was recorded by 204, footnote 327.) man, However, Cott (1940) instances in which protective devices and associated warning colours are known 1917 to to be ineffectual against these authors as Blabcnis atropos. (Hebard, BIOTIC ASSOCIATIONS OF COCKROACHES certain enemies. on But —ROTH may 345 does not necessarily imply that they are not this Certain cockroach the whole beneficial to the species attacked." secretions & WILLIS well be repellent to many predators, but as this purely negative aspect of the predator-prey relationship little is a thus far has been observed or published. Potential prey that successfully deitself against attack is never found in a predator's stomach. Cockroaches have a variety of glands which secrete odorous ma- fends Certain secretions, produced by tergal or dorsal glands in terials. the females feed on the secrefrom these glands prior to copulating (Roth and Willis, 1954). However, other secretions which are produced by both sexes are males, are involved in sexual behavior ; tion ejected or given off when the insect is disturbed undoubtedly these ; are defensive weapons that are used against predators. experiments or observations are on record to show secretions may be in protecting the cockroach. how Very few effective these Although the mor- phology of some of the glands has been described, relatively known about Many little is the chemistry of their secretions. species of Australian cockroaches have been reported to emit "disgusting" odors, though the glands producing these secretions compounds known. Cosuwzosteria 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, Platysostcria castanca, 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 have not been described, nor is the chemistry of the ; 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 Cufilia nitida emits, when disturbed, a corrosive liquid with an extremely dis- agreeable odor. The seized adults of Eurycotis floridana emit (Rehn and Hebard, 1905). sensitive skin areas, may The an odorous fluid when which may irritate fluid, be ejected as a spray for a distance of several 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 Bnfo marinus and the frog Rana pipicus in- inches. variably spit out adults of £. floridana which they have seized. The odor of 2-hexenal was strongly apparent after these attacks, and the However, the lizard Anolis equestris insect was never damaged. SMITHSONIAN MISCELLANEOUS COLLECTIONS 346 seized and crushed E. floridana before releasing The the insect 5 to lO minutes later. readily attacked adults of E. floridana its VOL, I4I hold and dropping blue jay Cyanocitta cristata and killed them but did not odor had dissipated; however, the bird carried nymphs oi E. floridana to its perch and ate them. Nymphs of this species do not secrete 2-hexenal (Roth et al., 1956). Recently, eat the insects until after the 2-hexenal has been tested for found its antibacterial activity and has been to be active against seven species of pathogenic bacteria (Val- curone 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 Platysosteria novae seelandiae. Walker (1904) and Longstaff {in Shelford, 191 2) 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. spectrographic analyses show Both infrared and mass is a mixture containing that the secretion is found in E. floridana. Eisner (personal communication, 1958) observed that the lizard Anolis carolinensis immediately released Neostylopyga rhombifolia without injury, but that 2-hexenal, the aldehyde that and Cyanocitta Bufo marinus, Anolis equistris, despite the secretion several unidentified spiders nomyrmex ; cristata ate the insect and the ant Pogo- badius were not repelled by the secretion of N. rhonibi- folia. 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. have also found similar ventrally located glands in both Peri- We planeta australasiae, and P. brunnea. The reservoirs which store the secretion of the ventral glands are smaller in Blatta spp. than those found in Eurycotis, and Periplaneta 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 1898; Oettinger, 1906; Koncek, 1924), the functions of secretions of these nonepigamic dorsal glands and the ventral glands are still open to question. It is possible that (Haase, 1889, 1889a; Kul'vets, BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 34/ some of the odors produced by cockroaches have functions other than defense or sex attraction. For example, Ledoux (1945) showed that the species odor largely responsible for the gregarious behavior is shown by Blatta orientalis and Blattella germanica. The olfactory stimulus acts over a short distance only, and the source of this odor germanica in warm which had the odor of chloroform Dusham (1918) extracted a wax the German cockroach. However, there is no evidence to show that cockroaches respond to the same cockroach odors that are detected by in the insect is unknown. By washing Blattella 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 and adults are modified ture second abdominal spiracles of nymphs into odoriferous glands of 2-ethyl-i,4-benzoquinone ; which produce a mixand 2-methyl-i,4-benzoquinone ; means of defense. The offensive odor emitted by adults and nymphs of Leucophaea maderae also issues from the second abdominal spiracles (Roth and para benzoquinone ; this material is ejected as a Stay, 1958). Diploptera is capable of ejecting its quinones from either or left tracheal gland according to which side of the insect (pi. 36, A-B). Eisner (1958) found that the ant Pogonoinyrmex hadius (Latreille) Galerita janiis Fabricius when (pi. is its right attacked secretion repelled the 36, C) and the they attacked the cockroach. beetle The was repelled by large nymphs and of D. punctata but young nymphs were usually eaten promptly spider Lycosa helluo Walckenaer adults (Eisner, 1958). Bordas (1901, 1908) believed that the "conglobate" gland (Miall and Denny, 1886) found in males of Periplaneta americana and Blatta , was an odoriferous gland used for defense, but Gupta (1947) has shown that in all probability this gland (the phallic gland) orientalis, 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 SMITHSONIAN MISCELLANEOUS COLLECTIONS 348 VOL. I4I 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 vi'hich these insects assume that the crustacean when they immobilize themselves response to in 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 WiUiams (1928) mentioned diurnal cockroaches which by a combination of markings, shape, posture, and active flight taken for models. about vegetation suggest certain wasps. Unfortunately, practically nothing known about is the behavior of and models or their relationships with predators in the field. For the most part, the examples are based on a comparifor son of pinned insects from museum collections (Burr, 1899) 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 benethese so-called mimics ; fit in the survival of mimetic species. Certainly Cott's (1940) volu- minous 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 The purpose is to promote exit "shore leave for cockroaches." termination of cockroaches in a warship because, on the one hand, any warship suffers from numerous cockroaches, and, on the other The formalities for a hand, any seaman likes shore leave shore leave for cockroaches are as follows. A seaman keeps . . . which he captured (mainly B. gcrmanica, because p. americana and P. anstralasiae are seldom found in Japan) in a bottle or in a bag until the number reaches 300. Then he brings cockroaches 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 them BIOTIC ASSOCIATIONS OF deck officer COCKROACHES —ROTH & WILLIS confirmed the number of cockroaches. The petty name signs the seaman's name, on shore leave of division, rank, officer and date to be book for cockroach shore in the log 349 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 is a log book same as for the former, and there for the rat shore leave in the petty officer's quarters. The author took advantage of these systems frequently. SoNAN Little is known (1924) of the effects of predatism and parasitism on natural populations of cockroaches. Many are very general; few data on egg parasites yet there are a statements in the literature (e.g., Tetrastichus hagenotvii) which suggest that, in the absence of para- much larger than have summarized the literature on natural control and also that on the use by man of predators and parasites, populations of domestic cockroaches might be they are in certain areas. We sites in the biological control of paucity of information, cockroaches. However, because of the we have been unable to evaluate the effective- ness 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. The staple diet of scorpions in Arizona is the small cockroach commonly known as the water bug (Stahnke, 1949) in (Sein, 1923). ; the part of Arizona where he resides, Stahnke (personal tion, 1953) says that the "water-bug" is supellectilium although Blattella germanica communica- most generally Supella is also found, but less abundantly. Spiders. —Jefferys tected in the Antilles down and (1760) mentioned a large spider which was proand especially on Guadeloupe because it hunted fed on cockroaches in every house. Sir ; was reputed to be common (1725, in Cowan, 1865) reported the spider Hans Sloane that residents of Jamaica kept spiders in their houses to destroy cock- 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, 191 5) 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). roaches. SMITHSONIAN MISCELLANEOUS COLLECTIONS 350 Ants. —A Madam Merian noticed VOL. I4I that ants cleared houses of cock- (Kirby and Spence, 1822). A small reddish-yellow ant, called Pucchugigi by Peruvian Indians, pursued and destroyed a cockroach called Chilicabra which was a pest in native huts (Tschudi, roaches 1847). Schwabe (1950b) found swarms of ants attacking living Pycnoscelus surinameiisis and stated that ants are probably the chief enemy of this cockroach in Hawaii. Africa a band of driver ants may Wallace (1891) stated that enter a house and clear it in of cock- roaches 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 (191 7, in Asano, 1937) proposed that parasitic wasps such as Evania and Brachygastcr be protected in Japan as the natural enemies of cockroaches. In one area in France, 20 percent of the oothecae of Lohoptera decipiens were parasitized by Zeuxevania splendidula (Genieys, 1924). Edmunds (1952a) found that 12 percent of 459 oothecae 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 oothecae 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 oothecae of Periplanefa americana collected in Saudi Arabia were 29 percent parasitized in March and 25 percent parasitized in October by ^z^ama a/'/^^wrff^'ai'^^r. Sonan (1924) found I of 65 oothecae of P. americana and P. anstralasiae parasitized by E. appendigaster in Formosa. Cottam (1922) stated that the increase of Supella supellectilium ; in Khartoum was checked by a wasp identified as Anastatus tenuipes (see p. egg-parasite that was later 246) (Ferriere, 1930, 1935). In this country, this wasp seemed to be effective in decreasing the num- bers of Supella in certain areas in Arizona (Flock, 1941). In Formosa, Tetrastichns hagenozvii was an important parasite of cockroach eggs (Maki, 1937). Sonan (1924) reported 30 percent and P. anstra- parasitization of 65 oothecae of Periplanefa americana Formosa. In Bangalore, India, the natural pararandomly collected oothecae of P. americana varied from 21 percent (of 495 oothecae), July 1947-June 1948, and 43 percent (of 288 oothecae), July-December 1948, to 57 percent (of 178 oothecae), July-October 1949 (Usman, 1949). Cameron (1955) obtained T. hagenozvii from oothecae collected in Trinidad, B.W.I., and lasiae collected in sitization of COCKROACHES BIOTIC ASSOCIATIONS OF —ROTH & WILLIS 35I Saudi Arabia; some 15 percent of the oothecae of P. americana and P. australasiae collected in October in Trinidad were parasitized; a later collection (March) was 34 percent parasitized; a small sample was 65 percent parasitized. The oothecae collected in Saudi Arabia in March were 20 percent parasitized. Plank (1947) found that the eggs of the American cockroach in of P. americana oothecae 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 oothecae 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 was impaired by Tetrastichus acting as a hyperparasite (see stated that in Puerto 6, p. 236). However, Wolcott (1951) 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 control footnote are not supported by experimental evidence. smaller wasp egg parasites are more effective cockroaches. Only one evaniid decontrolling in evaniids than the but many individuals of the other ootheca, velops in a parasitized It is likely that the wasps develop in one ootheca and the number of females that emerge However, Cameron (1957) concluded that, with a is usually large. 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 effectiveness Zimmerman (1948) Coniperia merceti, practically did not report the parasite's reducing the cockroach population. in wiped out when According to accidentally imported, Siipella 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 oothecae of this cockroach were parasitized (Zimmer- man, 1944). oothecae We (1954b) ran some simulated field tests in which we Tetrastichus hagenowii in rooms artificially seeded with liberated ; from 28 to 83 percent of American cockroach oothecae and SMITHSONIAN MISCELLANEOUS COLLECTIONS 352 VOL. I4I 56 percent of oriental cockroach oothecae 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). introduced into This parasite was also successfully Samoa (Dumbleton, Cockroach-hunting zvasps. —An 1957). earnest attempt has been made to 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 191 7 and establish in spread to several of the Islands (Swezey, 1920, 192 1 1944). ; Williams, Bridwell (1920) stated that as a result of this introduction was a great decrease in cockroaches of the genus "Phyllodroniia." 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 there Williams ( Blattidae, 1928) who observed that Podium "destroyed innumerable 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; Wil1946; Van Zwaluwenburg, 1950). The compressa now found in the Hawaiian Islands are liams, three wasps captured in Noumea, New thousand of all A. descendants of Caledonia (Williams, 1944). number of cockroaches was noticeAccording to Williams ( 1941 ), ably 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 Isthe lands in 1955 against Periplaneta spp. parasite became established ; it is in either place not yet known whether (Dumbleton, 1957). the BIOTIC ASSOCIATIONS OF COCKROACHES —ROTH & WILLIS 353 VERTEBRATES ... on conserve avec soin les crapauds dans les maisons, et que les dames les tolerent, meme sous leurs robes, en raison de leurs continuels services, car ils se promenent sans cesse a 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 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 efifectively clear it of cockroaches (Marlatt, 191 5) on sugar plantations in Australia, these amphibians were encouraged in houses and kept as pets because they hunted and devoured large brown cockvector of the chicken eye — ; roaches (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 tilles else. In Hawaii (Zimmerman, 1948) and in the Lesser An- (Ballou, 19 1 2) 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 cock-roach" and suggested that lizards be used for — SMITHSONIAN MISCELLANEOUS COLLECTIONS 354 VOL. I4I cockroach control because the reptiles are docile and harmless. Arno On 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 PycnoAtoll geckos and night- feeding skinks eat large sceliis. Mammalia. — Cowan (1865) stated that in England hedgehogs were often kept domesticated in kitchens to destroy cockroaches. writer also stated that a lemur was This kept on board ship to destroy cockroaches. Large numbers of the American and Australian cockroaches were mongoose in Hawaii (Perkins, 1913). eaten by the ACKNOWLEDGMENTS We would have been unable help of many people who have complete this review without the to generously given us their time and the We benefit of their special experience. are exceedingly grateful to these individuals for they have contributed this work possesses ; we much to whatever merit 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. them for Many their many We are especially pleased to thank favors. persons have determined at our request the identity of specific These individuals are cited in the text and to them we extend our thanks. We thank our colleagues, cited in the text, who organisms. 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 T. Campbell, Commonwealth specimens of the species indicated: and Industrial Research Organization, Canberra, New Dr. L. R. Cleveland, Harvard South Wales (Panesthia australis) University (Cryptocercus punctulatus) Dr. N. T. Davis, University Dr. F. Englemann, Albert of Connecticut {Byrsotria fumigata) Dr. F, A. Einstein Medical School {Gromphadorhina portentosa) Scientific ; ; ; ; BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS Lawson, Kansas State College {Comperia merceti) ; 355 Dr. K. D. Dr. E. O. Roeder, Tufts University {Hierodula tenuidentata ( ?) ) deropeltiformis). {Ischnoptera Wilsoni Harvard University of thank the following individuals for checking the taxonomy ; We sections of the the following organisms or for reading the indicated of California; University Benjamin, Dt. R. K. manuscript: Fungi.— Laboratories. Protozoa. Dr. E. G. Simmons, Quartermaster Research of Illinois. HelUniversity Emeritus, —Dr. R. R. Kudo, Professor Lucker, U. S. T. Dr. and Chitwood J. minths.— Mrs. May Belle Dr. R. whipscorpions.— and Centipedes of Agriculture. Department U. S. National Museum. Scorpions.— Dr. J. L. Cloudsley-Thompson, University of London; Dr. R. E. Crabill, Jr. Dr. R. E. Spiders.— Dr. B. J. Kaston, University of Connecticut; Sciences of Academy Chicago Camin, Crabill, Jr. Mites.— Dr. J. H. A. B. Dr. Cockroaches.— Museum. Dr. E.'w. Baker, U. S. National of Museum Brown, L. Jr., Dr. W. Ants.— Gurney and J. A. G. Rehn. parasites of Hymenopterous College. Comparative Zoology, Harvard E. Crabill, Jr., Museum; Dr. cockroach eggs.— Dr. B. D. Burks, U. S. National zvasps. Cockroach-hunting Michigan. H. K. Townes, University of Lepidoptera.—Dr. Museum. National —Dr. K. V. Krombein, U. S. U. S. Clarke, U. S. National Museum; Dr. E. L. Todd, J. F. G. Beal, R. S. Dr. insects.— Miscellaneous of Agriculture. Department Dr. A. B. Gurney, C. Sabrosky, Dr. R. L Sailer, and J. T. SpilU. S. man, U. S. National Museum. Fishes.— Dr. L. P. Schultz, Cochran, M. Doris Dr. reptiles.— and National Museum. Amphibians Friedmann, U. S. U. S. National Museum. Birds.— Dr. Herbert U. S. National Johnson, H. D. Dr. National Museum. Mammals.— W. Museum. manuthank the following individuals for reading the entire Field, U. S. Gordon Maj. Gurney; Rehn; Dr. A. B. script J. A. G. The Massachusetts. of University Army and Dr. H. L. Sweetman, entomolthese of criticism friendly monograph has profited by the We : ; ogists. checking thank Dr. R. A. Howard, Harvard University, for of the formerly Torok, E. W. Maria Mrs. lists of plant names; obscure obtaining in assistance for Quartermaster Technical Library, Department of Agriculture literature; Miss Louise Bercaw, U. S. the paper by Vlasov containing identifying the journal We Library, for language and Miram the individuals who translated foreign Miss G. Lillian and bibliography for which they are credited in the the manutyping for Laboratories, Fede, Quartermaster Research articles, ; ; script. SMITHSONIAN MISCELLANEOUS COLLECTIONS 356 VOL. I4I REFERENCES Abdulali, H. The 1942. terns Journ. and edible-nest swifts Nat. Hist. 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Les insectes synanthropes comma botes intermediaires et botes vecteurs des helminthes au Tadjikistan. (In Russian, Frencb summary.) Trudy Tadzbiksk. Bazy Akad. Nauk SSSR, vol. 6, pp. 241-248. [Russian text translated by M. Yeas.] 1940. Certain points in tbe epidemiology of dysentery and its endemic foci in Central Asia connected with the cockroach Shelfordella tartara Sauss. (In Russian.) Second conference on Parasitological problems. November 1940, Leningrad, p. 35. Izd. Akad. Nauk SSSR, Moscow. [From Rev. App. Ent., 1946, ser. B., vol. 34, p. lie] 1936. ZULUETA, A. 1916. DE. Sobre la estructura y biparticion de Nyctothenis ovalis Leidy. Trab. Mus. Nac. Cienc. Nat., Madrid, ser. Zool., No. 26, pp. 1-16. PLATES VOL. 141, PL. 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS Blabcrus cnviiifcr. nympli. C Pliotograpli by Jack Salmon.) SMITHSONIAN MISCELLANEOUS COLLECTIONS Blabcnis (I'ujantcus, c. X 2.2. VOL. 141, PL. 3 (Photograph by Jack Sahnon.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 4 a; < 00 cq SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 5 B A-B, BlattcUa ycnnaiiica, c. X 5.2. A, Male. B, Female. C-D, BlattcUa vaga, c. X 5.2. C, Male. D, Female with ootheca. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 6 m y. cq SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 7 OIT u 03 c ^£ goo ed"5 < ><' < S SMITHSONIAM MISCELLANEOUS COLLECTIONS VOL. 141, PL. 8 y. ffl Ph y SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 9 X u SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 11 Ph OQ < X SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 12 !^ ^'Si SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 12A m < X SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 13 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 14 < X SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 15 m -j- ^ SMITHSONIAN MISCELLANEOUS COLLECTIONS X VOL. 141, PL. 16 Panchlora nivca. A, Dead individual showing normal, pale green coloration. 4.5. B, Dead individual sliowing the bright red coloration (very dark areas) characteristic of infection with Scrratia uiarccscciis. C, Living male. D, Living female. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 17 m -t CD :0 VOL. 141, PL. 1£ SMITHSONIAN MISCELLANEOUS COLLECTIONS \ CQ < X (x. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 19 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 20 < X CI. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 21 < y. ex, SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 22 03 X (Ih SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 23 < X Dh SMITHSONIAN MISCELLANEOUS COLLECTIONS X VOL. 141, PL. 24 A, Male from Hawaii. B, Macropterous parPycnoscdus siirijiainoisis, c. ;i.7. thenogenetic female from Florida. C, Brachypterous nonparthenogenetic female from Hawaii. D, Late instar nymph. (Photograph of nymph D, by Jack Salmon.) SMITHSOMIAIM MISCELLANEOUS COLLECTIONS VOL. 141, PL. 25 < X I SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 26 Bacteroids from Blattclla (jcniiaiiica. A, Part of abdomen showing mycetocytes in fat body, X 225. B, Lobe of fat body showing 3 mycetocytes, 750. C, Single mycetocyte; bacteroids appear hollow as result of fixation in Carnoy's fluid, X 1725. D, Smear of fat body showing bacteroids in various stages, X 1800. (All preparations and photographs through the courtesy of Dr. Marion A. Brooks.) X SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 27 Fungi parasitic on cockroaches. A, Hcrpoiiiyccs arictiniis growing on antennae, legs, body, and cerci of a nymph of Parcoblatta virginica. B, Hcrpo)ii\<ccs stylopygac 7. on antenna of Blatta orioitalis. X 35. (Reproduced from Richards and Smith [1955, 1956].) C, Hcrpoinyccs sp. [probably H. stylopygac] on antenna of B. oriciifalis, 132. (Photographs B and C through the courtesy of Dr. A. G. Richards.) X X SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 29 w 6 A, I'ndetermined mermithid that parasitizes Ectobius pallidus. X g. The worm has partly emerged of the cockroach. (Reproduced from Roth and Willis [igsr].) B. Undeterthat parasitized Eurycotis fJondaiia shown beside its host, X i.S (Specimen courtesy of Dr. T. Eisner. from the neck region mined gordian worm SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 30 A, Hctcropoda vcnatoria, a cockroach-hunting spider, slightly less than natural size, on bananas. (Reproduced from a Kodachrome transparency through the courtesy of Dr. B. J. Kaston.) B to E, Lycosa sp. {avida?) capturing and feeding on a nymph of Supella sitpclIcctilitim in the laboratory, X 1.4. VOL. 141, PL. 31 SMITHSONIAN MISCELLANEOUS COLLECTIONS ,>>o a! :=! U( "CI 03 O 7 C -^ 8U ' '''.'^\ / QQ < LiJ c5 ~ H SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 32 X Oh H SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 33 X:,. B, A Proscvania punctata c? beside an ootheca of Pcnplancta aincncaiia it had emerged, Hv^tia harhvoidrs with o5theca of Parcoblatta uhlcriana from which X 5 C Larva of a lampvrid beetle feeding on Parcoblatta virgimca in the laboratory, X4. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 34 B Uialcid parasites of cockroach eggs. A, Anastatus lioriihiiiiis ovipositing into an o^theca which is being carried by Eurycotts iloridana, c. X 4. B, Compcria moccti ovipositing into an o.Stheca of ^upclla supeflcctthum, c. X 13. C, Tetrastichus hagcnowii ovipositing into an ootheca of Pcriplaneta amcricaua, c. X 10. (C from Roth and Willis [1954b].) still VOL. 141, PL. 35 SMITHSONIAN MISCELLANEOUS COLLECTIONS impulcv cowprcssa attacking Pcriplancta sp.. (presumably '.i^^o"* ""'^^'-l™''"' "^t"^'-^ ^J^^j^^j^ ^,,^ SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 36 B Chemical defense of Dif'loptcra punctata against predators; the spray pattern is displayed on Kl-starch indicator paper. A, Spray pattern after right niesothoracic leg was pinched. Cumulative spray pattern after left niesothoracic leg of the same insect was pinched. 1>, C, The defensive glands of the cockroach on the left had been excised, and it is under persistent attack by ants from a laboratory colony of Pogonoinyrmcx badius (Latreille). The intact cockroach on the right was also attacked by the ants, but it discharged a spray of quinones and repelled the attackers. (From Eisner [1958], through the courtesy of Ur. T. Eisner.) INDEX Plate and page numbers in boldface type indicate illustrations. entries are placed in the index as unmodified substantives except In general, where a modifier contributes significantly to the identification of the item (e.g., blue heads, prairie dog). This index should be used in conjunction with the indexed check list of natural associations (pages 290 to 310) because these are not repeated here. All experimental associations are indexed below except those involving Blatta orientalis, Blattclla germanica, and Periplaneta americana that are cited on pages that also contain references to the natural associations of these three species. Abies sp., Agaricus, 164 257 Acacia, 56, 69 Acacia jarnesiana, 155, 164 harpophylla, 55 sp., Agave, 43, 57 Agelaius xanthomus, 282 Agis orientalis, 7, 27 Aglaopteryx absimilis, ISS Acanthinevania princeps, 235 Acanthocephala, 203 Acanthogyna dcplanata, 7 diaphana, Acaridae, 217 fades, gemma, sp., Achromobacteriaceae, Acrocomia tristis, no no 281 acnlcata, 142 Acromyrmex 7, 145, 146, 290, 35, 36, lobicornis, 312, 313 lundi, 312 vegeta, 7, 140, 36, 7, 143, 146 Alcaligenes jaecalis, recti, in III viscosus, III Algaroba, 45, 156, 164 Allacta similis, Allothereua macidata, 223 312 Alluadellina cavernicola, aciitinventris , 317 7, Actinocephalidae, 184 Actinomycetaceae, 124 Alsophila Actinomycetales, 123 Alsophilas, 164 Adeleidae, 184 Amasonina emarginata, Ameiva exstd, 275 Adelina cryptoccrci, 184-185 porteoides, 144 Aerobacter aerogenes, in cloacae, 112 sp., 112 metallica, 243 Agamidae, 275 Agamospirura parahormeticae, 205 17 139 7, 2'^, 146, 158 275 Amitermes, 102 Amoeba coli, Aeschnidae, 224 Agamerion sp., sp., 290 36, 75, 276, 7, Alluaudella cavernicola, 7 Alpinia, 163 Aechmaea 153, ypsilon, 7, 75 Aircraft, 87-90 octospinosns, 312 silvestrii, 144, 339 niger, 312 Acropyga 318 35, 156, 290, 317, 7, 158, 159, no Acridotheres 317 339 Acarina, 210, 216 Acheta domestica, 320 Achromobacter hyalinum, 35, 156, 7, devia, 7 blattae, lyy 178 undetermined sp., 181 Amoebidae, 177 Amoebina, 177 Amphibia, 269-272 441 — SMITHSONIAN MISCELLANEOUS COLLECTIONS 442 /4 w o/i.j Amphibians, 3, 353 Amphiboluriis barhatus, 275 Amphoromorpha sp., blattina, Continued. grahami, 274 leachi, 139 274 piilchellus, 139 256 amoena, 256-257 sagrei, 274 assimilis, 257 stratulus, Ampule X, 2, sp., 352 canalictilata, 94, 257-258, compressa, 258-259, 352; compressiventris, 260 pi. 35 carpenter, 267 predaceous, 266 Ananas comosus, 144 Anaphothrips sp., 317 Anaplecta ascma, 7, 36 36 hemiscotia, 7, lateralis, 36 36 diesingi, 195 145 7, 290 145, blattidifnrax, sphaerolaima, 194 Aotes sonalis, 284 Anastatus blattidarum, 246 245 floridanus, 94, 245, 255, 319; pi. 34 tenuipes, 94, 246, 255, 350 Anatidae, 277 Anax 267 philippinensis, 193 36 7, safari, wood, 317 Aorurus (Thelastoma) appendiculatus, 196 145 decipiens, 7, 88, Antagonism, interspecies, 341-343 Antbird, 280 hosts of cockroaches, 31 1-3 14 Ampulicidae, 256 Anacardiaceae, 157 spp., 274 Anseriformes, 277 army, 228, 267, 268, 280, 350 big-headed, 268 259 sonnerati, 260 sibirica, mexicana, 274 Ants, 57, 266-268, 316, 317, 350 Argentine, 268, 319 259 sibcrica, 260 ruficornis, jallax, 7, 273 Antibiotics, 97-98, 346 259 novarac, 256 fasciata, asteca, 7, strenuus, 224 Ancylosfoma caninunt, 209 ceylanicmn, 209 duodcnale, 209 Aotus, 284 Aphaenogaster Aphelocoma picea, 267 coerulescns, 280 Aphids, 62 Aphlebia maadata, 10 punctata, 10 Apocynaceae, 161 Apolyta, 9 Apotrogia angolensis, Apple, 166 Aptera cingulata, 7 Ancylostomidae, 209 fusca, 7, 290, 329 Aptcrohlatta perplexa, Androctonus australis, 211 Aneurina tahuata, 7, 160 Aquifoliaceae, 158 viridis, 7, 155, 160, 161 Anguillula macrnra, 195 Anisogamia, 26 Annate, 159 Annetta, Steamship, 149 Anoetidae, 217 Anolis carolinensis, 273, 346 cristatelliis, VOL. I4I 273 equestris, 273, 345, 346 7, 17 7, 18 Arachnida, 211 Arachnids, 34 Araceae, 143 Aranea, 214 Araneida, 214 Arbor vitae, 44 Archiacanthocephala, 203 Archiblatta hocvenii, 7, 329 Archlmandrita marmorata, 7, 147 tessellata, 7, 147 INDEX Arenivaga apacha, 7, 23, 26, 27 443 Continued. Associations bolliana, 7, 23, 26, 27, 37, 290 mutualism, 96-102 erratica, 7, 23, 26, 27 number floridcnsis, 7, 24, 37 obligate, 95 firrato, 7, 17, 18, obscure, 96, 316-319 37 with man, 70-90 roseni, 7, 24, 26, 27, 291 tonkawa, Astata, 255 24 7, of, 3, 5 Aristida pennata, 65, 141 Aristiger histrio, 7, 37, 159, 316 Armadillo, 287 Army transports, 84 Arthroleptis variabilis, 270 Arthromitaceae, 124 A teles dariensls, 284 Athel, 39 Atta cephalotes, 313 jervens, 313 lundi, 312 nigra, 312 Arthromitus intestinalis, 124 Arthropoda, 210-268 octospinosa, 313 scxdcns, 313 texana, 2,^2> Arum sp., 143 Ascalaphidae, 227 Attacus Ascaridae, 209 Attaphila aptera, Ascaris lumbricoides, 209 sp., atlas, 320 313 8, bergi, 8, 312, 313 209 flava, 8, suum, 209 314 jungicola, Ascaroidea, 209 schuppi, Aschelminthes, 192, 204, 209 Ascomycetes, 133 sexdenfis, 313 312 8, 8, 8, 313 Atticola mortoni, 313 sp., Asmorrhisa longistylis, 240 Asparagus officinalis, 235 8, 312 Attraction, intraspecies, 328-334 to decaying material, 53, 59-63 Aspen, 46, 152, 164 Aspergillaceae, 129 to honeydew, 62 Aspergillus flavus, 130 to lights, 52, 61, 64, 66, 79, 81 Audreia bromeliadarum, jumigahts, 130 jamaicana, niger, 130 sp., 130, 8, 8, 31, 37, 145 37 marginafa, 11 131 Aves, 276-282 Arncularia avicularia, 214 sydoKfi, 130 tamarii, 130 Aspiduchus borinquen, 7, 18, 37, 333- sp., 214 334 cavernicola, deplanatus, 8, 18, 334 37 7, 8, Aspleniiim nidus, 44 Assembling of cockroaches, 331-334 Associations, among 6, 15, 91-96 cockroaches, 324-343 biotic, 2, 91, 94-96 classification of, 91, 95-96 "B. aerobia del pseudoedema maligno," 125 B. alcaligenes beckeri, 125 faecalis, iii recti. III "B. del pseudoedema maligno," 125 B. violaceus, 106 Baboon, 286 ecological, 14-90, 324-343 Bacillaceae, 120 facultative, 95-96 Bacilli, colon, 126 familial, 325-330 gregarious, 330-336 paracolon, 113 "Bacillo del barbone dei bufali," 126 interspecies, 337-343 "proteisimile," 126 intraspecies, 336-337 "similcarbonchio," 126 — 1 SMITHSONIAN MISCELLANEOUS COLLECTIONS 444 Bamboo, Continued. "Bacillo" "tifosimile," 126 "Bacillus," 126 lactici, 112 120 albolactis, anthracis, 120 99 8, "sylvestre," 194, 199, 188, Barbulanympha, 102 coahoma, 173 circidans, 120 estaboga, 173 cloacae, 112 faecalis alkaligenes, 11 laurabuda, 174 175 sp., flacheriae, 121 aero genes, iii ufalula, 174 zvenyoni, 174 vtegateriiim, 121 Bark, 36-38, 42, 43, 45-65, 141, 152, 153, monachae, 121 160, 164, 165 periplanetae, 121 prodigiosus, 117 Bass, large-mouth black, 269 radiciformis, 121 Bassarisciis astutus, 288 stellatus, 121 Bat, 77, 283 caves, 17-18, 21-23 subtilis, 121 tritus, 122 Batrachians, unidentified, 272 Bacteria, 2-3, loo-ioi, 104-127, 346 "spirochaetoid," 127 Bayberry, 43, 44, 152 Beauveria bassiana, 131 Bacteriaceae, 119 Bedbug, 321-322 "Bacterium," 126 Beech, 47, 61, 62 Bacterium alkaligenes, 119 Bees, 314, 318 delendae-nmscae, 120 haemophosphoreum, 120 monache, 121 prodigiosum, 117 Badger, 289 black (see Blatta orientalis) rhinocerus, 317 Bertramia blatellae, 185 vulgaris cicla, 153 blattariim, 187 Binema 187 Birds, ovatum, 187 praenucleatum, 187 mirsaia, 193 3, 4, 276-282, 353 nests, 35, 318 Bites, cockroach, 83, 322, 336-33? 187 mite, 220 Ballota nigra, 38 5a/to godeffroyi, patula, 347, 348 Beta maritima, 47, 153 Balantidium, loi sp., Beetles, 26, 34, 57, 63, 91, 229-234, 318, black larder, 234 Bacteroides uncatus, 119 Bacteroids, 96-100; pi. 26 co/i, 200 189 "Baratas de pau podre," 200 120 ccreus, 120 lactis blossoms, 36, 41, 56, 57, 146-148 Bantua stigmosa, "Barata selvagem," alcaligeites faecalis, iii 82, 146- 215 151, 163, 165, "subtilis group," 122 biitschlii, 44, 141 Bambusicola thoracica, 277 Banana, 36, 40, S3, 54, 58, 66, "similtifo," 126 Bacillus acidi VOL. I4I 8, spider, 215 8, 37 Bixa 291 quadricaudata, 8, sp., 159 Bixaceae, 159 plat y soma, 8, 317 37, 141 scrip ta, 8, 37, 141, 326 torresiana, 8, 38, 141 verticalis, 8, 38, 141 Blabera fusca, 8, 322 Blaberus atropos, 8, 18, 203, 204, 291, 322, 344 boliviensis, 8, 147 38, 125, 147, 445 INDEX Blaberus — Continued, B la ttella—Con timed. germanica clarasianus, 8 craniifer 8, i8, 75. 97. 99. 116-118, 132, 138, 291, 322, 330, 333. 336, 339, 344; pis. I, 2, 28 discoidalis, 8, 38, 75. 85, I47. I5i. 163, 291, 318, 330. 334, 8, "maderae," 18, 38, 99, 291 pL ; 3 nahua, spp., 38, 290, 291 vaga, 292 133 americana," 108, no, 119, Blatticola, 199 blatticola, 193 "Blattidae sylvestres," 200 Blattilaelaps r.auphoetae, 216 Blattina concinna, 10 Blattisociiis tineivorus, 216 (Shelf ordella) lateralis, 8, 18, 38, triodons, 216 Blattophila sphaerolaima, 194 sphaerolaima var. javanica, 194 292 melanocephala, 13 supellaima, 194 notulata, 10 orientalis, 8, 18, 38. 70-72, 75-76. 84, 85, 88, 96, 99, 100, 118, 123, 128, 129, 131, 151, 153, 162, 163. 224190, 204, 205, 212, 219, 222, 226, 232, 235, 237, 245, 255. 269, 275, 276, 292-293, 320-323, 325327. 331. 332. 334-336, 338, 341, 342, 346, 347, 352 ; pis. A, 27 Blattotetrastichus hagenomi, 249 Bluegills, 269 Blue heads, 269 Blue jay, 280, 346 Bobolink, 282 Bodo blattae, 167 sp., 167 Bodonidae, 167 Bog, peat, 46 Boraginaceae, 161 292 224 Bounty, H. M. S., 83 Brachygasfer, 350 minutus, 235-236, 351 Blattartun alaianim, 83 Blattelicola blattclicola, 193 Blattclla bisignata, 8 Bracken, 45, 47 Brambles, 51. I55 Breadfruit, 162 brunneriana, il chichimeca, 10 delicattda, 8 83-85, I35. 141. 166, blattae, 193-194 humbertiana, 8 germanica, 39, 77, aegyptiaca, 195 caraibea, 79 concinna, 10 3, 27, pidchra, 243 americana, 12 sp., 8, Blatticidella ashmeadi, 243 aethiopica, 13 Blattaria, y() Blatticida ashmeadi, 243 121, 122 Blatta aegyptiaca, 13 71, 75, 263, 295 255. 293, 320, 328; pl. 5 Blastocystis hominis, 133 sp., I44. 294. spp. 88, 145. 295 Blastocystidaceae, 133 "Blatella 142, n schubotzi, 8, yellow-shouldered, 282 8, 39. 8, 336 littiricollis, 8, 248 11, 31 5, 26, humbertiana, 339 Blackbird, 282 Puerto Rican, 282 Blaptica dubia, 320, 321. 323. 324. 328, 331. 332. pis. 334. 335. 337-342. 347-349; cubensis, 8 giganteus, — Continued. 222, 225-227, 234-236, 240, 241, 293-294, 255, 272, 276, 277, 283, 8, 17, 18, 39, 87, 88, 96-101, 70-72, 7^, 103-105, 114, 118, 119, 124, 126, 131, 143, 171, 192, 204, 205, 211, 212, 219, Bream, 269 Bromeliaceae, 144 Bromeliads, 31, 36. 37. 4i, 42, 45, 49, 50, 54, 56, 57, 68, 144-146, 320 SMITHSONIAN MISCELLANEOUS COLLECTIONS 446 Brucella abortus, 119 Buboblatta armata, 8, 145 Camponotus VOL. I4I — Continued. pennsylvanicus, 267 Bucerotidae, 279 312 rnfipes, Bujo funereus, 270 ictericus, 270 Candida seylanoides, 129 marinus, 270, 345, 346, 353 valliceps, 270 Bufonidae, 270 Cants jamiliaris, 288 latrans, 288 Bulhoesia bulhoesi, 200 Cannaceae, 151 Cannibalism, 322-324 Cantao ocellatus, 320 Canidae, 288 Carina, 151, 163 icemi, 200 magalhdesi, 194 severianoi, 200 Bullfrog, 271 Canthium barbatum, 161 Capa bianco, 66 Burrows, 27 Capillar ia hcpatica, 210 cockroach, 55, 58, 67, 68, 328, 344 relative humidity in, 26 Capucinella delicatiila, 147 Carabidae, 229 vertebrate, 23-24, 26, 27, 29 Carboniferous, 14 Care, maternal, 325-330 Bursera simaruba, 157 Burseraceae, 157 Busaria blattariim, 188 Busariidae, Cariblatta antigucnsis, cuprea, 187 8, delicatula, 8, 40, 143, 147, 295 Buthidae, 211 hylaea, Buthus iniitans, 8, 41 australis, 211-212 8, 40, 147, 153 Butterfly nests, 66, 156, 317 insularis, 8, 41, 145, 147, Byrsotria cabrerae, landalei, 8, 41, 147 jumigata, 8, 8, 40 19, 40, 225, Z23, pi. 6 354 5 /wf£?a /Mf^ra, 8, 41, 153, 158, Cacao, 43, 52, 53, 55 nebulicola, Cacomistle, 288 or ester a, Cahita borero, 8, 8, punctulata, 40 spp., 42, 285 8, 147 42 8, 42, 142, Carica papaya, 159 Caricaceae, 159 Carnations, 163 Carnivora, 288 Carpinus orientalis, 38 spinitarsits, 217 218 Caryophanales, 124 Casuarina, 152, 164 Calyptracordia alba, 161 Casuarinaceae, 152 Camouflage, 344 Camponotus femoratus, 311, 314 maccooki, 316 Cat, 4, 289 maculatus, 312 340 143 43 Calluna vulgaris, 160 Calodexia venteris, 228 sp., 8, ring-tailed, 288 Caterpillars, 319 Catopsis fulgens, 144 8, 295 pi. 7 88 Callitris sp., 55 228 145, Cariblattoides instigator, suave, chrysolevca, 285 Caloglyphus 8, stenophrys, Callithrix chrysoleucos, 285 42, ; 151 reticulosa, 8, 40 Caladium, 66, 143 Callithricidae, 285 jacchiis, 8, 8, punctipcnnis, Cafards, 207 spp., 339 /Mf^a, 23 /m^^o minima, 8, 42, 143 nahua, 158, 40, 8, 40 156 43 161 INDEX Cattleya, 151, 162-163 Caudata, 269 Caves, 16-17 Cavia sp., Chordodcs morgani, 201 202 pilosiis, puerilis, 201 288 Chordodidae, 201 "Chordotes" puerilis, 201 Chorisoneura barbadensis, Caviidae, 288 Cebidae, 284 Cebiis apclla, 284 sp., 40, jorviosclla, 8, 43, 153 plocea, 8 Centruroides gracilis, 212 ^e^m.yw, sp., 295 212 translucida, 8, 44 Choristima magalhaesi, 194 Cephalosporitim sp., 131 Choristimodes Ceratinoptera diaphana, 7 picta, 8, 43, 159 Ceratonia siliqiia, 156 Cercopithecidae, 286 sp., 286, 343 Cestoda, 208 295 sp., 9, 295 Chromatonotus infuscatus, notatus, 9, 44, 9, 44 77 Chromobacterium violaceum, 106 Chrysanthemums, 165 Chrysemys picta, 272 Cicadas, 257 Ciliate, unidentified, 190 verticillata, 142 Chaetodactylus 218 Chalceus macrolepidotos, 269 Challenger, H. M. S., 83 sp., Chamaeleon chamaeleon, 275 oustaleti, 275 Chamaeleontidae, 275 Chameleon, 275 cat, sp., 9, Ciliata, 186 Chactidae, 212 Channel 152, 153, 157, 161, 8, 43, 316 Cephalobelliis brcvicaiidatum, 194 Chaetochloa 160 specilligcr, 8, 43 hentzi, 212 Cercopithecus 147 parishi, 8, 43 Cedar, Japanese, 164 Centipedes, 3, 222-224, 3^9 house, 222 vital tus, 8, flavipcnnis, 8, 43 capucinus, 284 Cecropia 447 269 Cimex lectularius, 321 Cinchona pitbescens, 162, 163 214 Citharexylmn villosmn, Ciniflo, sinensis, 157 sp., 157, 164 Cleonymidae, 243 Characidae, 269 Cheiloneurns virtdiscutiim, 244 Chelonia, 272 Chemotaxis, 331 Chenopodiaceae, 153 Chestnut, 44, 60-61 Chicken, 68, 82, 276, 278 Chilicabra, 281 Chilomastigidae, 169 Clepsidrina blattantm, 181 Clerada apicicornis, 226 Clerodendron irichotomum, 256-257 Clevelandella constricta, 189 contorta, 189 elongata, 189 hasfula, 189 nipponensis, 189 Chilomastix mesnili, 169 panesthiae, 189 Chilopoda, 222 parapancsthiae, 189 Clevelandellidae, 102, 189 Chimpanzee, 286 Clevelandia, 189 Chiroptera, 283 Chlamydosaurus 43, 161 Citrus aurantijolia, 157 maxima, i^y kingii, 275 Chlorion coinpressum, 258 Clitoris gayana, 141 Qevelandiidae, 189 Clostridiiiin fcscri, 122 Icntoputrcscens, 122 — SMITHSONIAN MISCELLANEOUS COLLECTIONS 448 Clostridium — Continued, Cockroach Continued. Madeira (see Leucophaea made- novyi, 122 rae) perjringens, 122 myrmecophilous, 310, 311-314 sporogenes, 122 spp., VOL. I4I odor, 122 84, 280, 344-347 6, oriental (see Blatta orientalis) tetani, 122 oviparous, 325-327 Clubiona, 214 Cnemidophorus sp., phytophagous, 68, 162-166 275 Coati, 288 predaceous, 319-324 Coccidia, 184 predators, escape from, 343 Coccidioides periplanciac, 133 ship, 82-87 Coccinellidae, 348 smoky "Coccobacillus," 126 brown (see Periplaneta fuliginosa) Coccobacillus cajae, 120 species, list of, 7-14 Cockroach, American (see Periplaneta number americana) amphibious, 30-33, 48 aposymbiotic, 97-99 Australian (see Periplaneta aus- known, of iii, 5 spotted Mediterranean (see Ectobius pallidiis) structural habitats of, 70-90 Surinam Pycnoscelus suri- (see namensis) tralasiac) beetle (see Diploptera punctata) undetermined, 308-310 brown viviparous, 31, 326, 328-330 (see Periplaneta brunnea) brown-banded (see Supclla snpcl- "wood" (except burrowing, 27, 29, 68, 328, 55, 58, 67, 30, 6, 14, 319-324, 336 cavernicolous, 16-25, 37 checklists of, 290-310, 315 cinereous Coconut, 35, 38, 49, 143 blattellae, 185 Cocnosia basalis, 229 Nauphocta (see Coelosporidium periplanctae, 168, 176, 177, 185 cine- rea) Coffea sp., 162 colonies, 75, 327-334 67 Coleolaelaps sp., 216; commensal, 310-315 Coleoptera, 229-234, 348 Cofifee, 66, control, Cuban I, 6, 84, 162, 331, "Colon 348 bacilli," 126 Colubridae, 276 Columbidae, 278 defenses, 343-348 Columbiformes, 278 cypress (see Diploptera punctata) Combretaceae, 159 desert, 25-30 disease relations, i, 3, domiciliary, 70-90, 51, 74 322, 331, enemies, 2-4, 91, 95-96 (see Blattella vaga) Florida (see Eurycotis floridana) German (see Blattella germanica) (see Neostylopyga rhombifolia) Key West Commensalism, 92 Commensals, 2-3, 95, cockroach, check 338, 349 harlequin pi. 12 Colonies, cockroach, 75, 327-334 (see Panchlora nivea) dendricoious, 51 field Par- Cocas nucijera, 143 344 carnivorous, here, usually coblatta spp.), 214, 266, 274 lectilium ) (see Blaberus craniifer) 173, 310-314 list of, 315 Compatibility, interspecies, 337-341 Comperia merccti, 244, 255, 351, 355; 34 merceti var. falsicornis, 244 Pl. Compositae, 162 Compsodes schivarzi, Compsolampra, 9 9, 27, 316 449 INDEX Comptolampra liturata, 9, Conocarpus erectus, 159 44. MO, 141 Cyperaceae, 142 Cypenis sp., 142 Cypress, 45, 63, 164 Conopidae, 228 Control, biological, i, 6, 348-354 Cypress pine, 55 Convolvulaceae, 161 Cyrilla, 42 Coptotermes ceyloniais, 310 Cyrtandra sp., 161 Cyrtocharax magdalenae essequibensis, Coraciformes, 279 269 Cordaites, 14 Cyrtotria capncina, Cordia dentata, 161 27 9, Cordyceps amazonica, 134 Dasypodidae, 287 Dasypus novcmcincius, 287 blattae, 134 Corn, 42, 48, 57, 58 Cornus tnas, 38 Corvidae, 280 Dates, 39 DDT, 278 Defenses, cockroach, 343-348 Corydia nuptialis, 14 Corynebacteriaceae, no Dematiaceae, 132 Corynehacterium diphtheriae, no Cosmozosteria lateralis, 9, 345 Costus 316 sp., Dendroblatta sobrina, Cotton, 66, 67, 158, 163 Coturnix coturnix japonica, 277 Derinestes ater, 234 Coxsackie viruses, 103-104 Coyote, 288 Dermestidae, 234 Dermestids, 320 Crappies, 269 Derocalymma Crataegus 234 Dcropeltis autraniana, Cristatithorax, 244 9, 44, 62, 67, Cryptocerctts punctulatus, 100-103, 175-229, 295, 317, 326, 328, relictus, 9, 9, 19 melanophila, 9, 28 45, 140, 164 Didelphidae, 283 Digenea, 208 Cupressiis, 164 "Diphtheroid macrocarpa, 9, Dinenymphidae, 170 I and 141, 164 226 II," 127 "Diplococci," 127 Diplococcus pneumoniae, 109 345 45, 77, 296 Diplocystidae, 181 sedilloti, 9; pi. 9 Diplocystis schneideri, 168, 181 pi. 28 sp., 181 spp., 45. 296, Cyanocitta apicalis, japanica, 226 Cunoniaceae, 15S Cupressaceae, 141 9, 245 alfierii, Diestrammena Cucumber, 164 near 28 Dictanimis fraxinella, 38 Cucarachero, 281 Cutilia nitida, 9, nigrita, 9, 28 Dicarnosis 44 34 Cryptomeria, erythropeza, Deserts, 25-30 339, 354; Pl- 8 sp. 27 9, stigtnosa, 8 stone, 226 Crinuni, 163 soror, cruralis, 9, 100 porcellio, 9, 28 Crickets, 320-321 spp., 6, sp., lampyrina, IS5 sp., 9, 45. 3i6, Dendrocopiis mahrattensis, 279 Dermaptera, 226 319 cristafa, 280, ; 346 Cyatheaceae, I39 Diplogasteridae, 192 Diplogyniidae, 217 Cycads, 166 Cynopotamus Diplogaster, 192 essequibensis, 269 Diplogyniid, undetermined, 217 330 1 SMITHSONIAN MISCELLANEOUS COLLECTIONS 450 Ectobius africanus, Diploptera dytiscoides, 9 punctata, 9, 45, 140-142, 152, 155- 157. 159, 164, 216, 224-226, 259, duskei, ericetorum, 9 germanictis, 8 347; pis. 10, 36 spectabilis spectabilis, 23 9, 46, 77 lapponicus, 164, 296, Diptera, 227 Disease organisms transmitted by cock- livens, 9, 46, 83, 100, 152, 9 Dobellina, 177 Incidus, 9, 340 Dog, 288 Dolichonyx oryzivorus, 282 Dolichurus bicolor, 260 nicaeensis, 9, 46 nigripes, 256 pallidus, 9, 19, 47, 77, 100, 140, 153, 161, 296, 340; pis. 7, 29 corniciilus, 260-261 gilbcrti, 261 panseri, greenei, 261 panzeri var. nigripes, 9 47, 153, 160, 9, haemorrhous, 260 perspicillaris, ignitus, 261-262 semenovi, stantoni, 262, 263, 352 spp., 19, 214, sp., Sylvester, 262 Dorylaea rhombifoUa, 1 Dorylus (Anomma) nigricans subsp. 152 297 47, 326, 340 sylvestris, 9 48 tadsihictis, 9, vittiventris , 9 267 Edentata, 287 wilvcrthi, 267 Dove, Chinese, 278 Dragonfly, giant Hawaiian, 224 9, 48, 297, 327 aurantiimi, 9 australe, 9, 48, 155, 160, 297, 327 214 Dnissus sp., 318 Dryadoblatta scotti, 9, 31, Drymaplaneta, 319 Ducks, domestic, 277, 343 Elcagnus, 48 Elentheroda dytiscoides, 9 Ellipsidion affine, Drake (Francis), 83 Dr ass odes, 9 9, 47, 9, 296 vittiventer, 9, 19, 48 sjostedi, 267 sp., 160, 340 9 lividus, roaches, 3 45 albicinctus, 9, 46 273, 280, 287, 296, 324, 334, 339, Dipodomys 9, VOL. I4I 45, 144, 145 Diiguay-Troiiin, Cruiser, 232 Dyscologamia capensis, 9 chopardi, 10 bicolor, 9, 48, 142 pellucidum, 9 simulans, 9, 48, 141 spp., 35, 48 varicgatum, Embadomonas pilosa, 9 wollasfoni, 9 9, 329 Elm, 38 blattae, 167 Embryophyta siphonogama, 140 Emydidae, 272 Dytiscidae, 230 Encephalomyelitis virus, mouse, 104 Earthworms, 319 Encyrtidae, 94, 243, 245 Earwigs, undetermined, 226 Endamoeba Eberthella oedematiens, 112 Echinococcus granulosis, 208 Eciion biirchelli, 268, 280 sp., 228 Ecology, 14-96, 324-343 "Ectobia" spp., 135 Ectobia livida, 232 blattae, loi, 102, 168, 177- 178 co/z, 178 javanica, 178 philippinensis, 178 thomsoni, 179 versus Entamoeba, 177 Endamoebidae, 177 INDEX Endolimax Escala circumducta, 13 blattae, loi, 180 nana, 180 sp., longiuscula, 13 180 297 sp., 10, Endomycetales, 133 Entamoeba 451 Escherichia 112 coli, coli var. acidilactici, 112 blattae, 177 blattarum, 177 coli var. coli, 178, freundii, 113 179 histolytica, lyg intermedium, 113 nana, 180 "Espino rubial," 66, 155 Estrildine sp., 318 Euandroblatta palpalis, 10, 28 pitheci, 179 sp., 179-180 thomsoni, 179 Eubacteriales, 104 Entedon hagenoivii, 236, Enterobacteriaceae, Eublaberus posticus, 249 Enterococcus, 109 Entomophthorales, 133 Ephestia kiihniella, 99 Eucharis, 163 Eucomonympha, , 9, 31, 48, 77, 141, 147, 160, 164, 297 conferta, conspersa, Euglenidae, 166 grisea, 9, 100 Euglenoidina, 166 maya, 9, 85, 145, 147 mexicana, 9, 147 mona, 9, 48, 144 Eulophidae, 94, 248 Eulophus, 250, 351 notabilis, 9, Eupelmus Eupelmidae, 319 sodalis, 9, 14s spp., 9, sp., 85, 32, 89, 14S-147, 263, 297, 326, 344 zvheeleri, 9, 49, 247 Euphorbiaceae, 157 10, 147 49 decastigmata, 10 297 Euryconema unidentified, 33 paradisa, 194 Eurycotis bananae, Epimys norvegicus, 287 44 245 liturifera, 10, Epilamprinae, 30, 326, 348 34, 94, 210, atriflagellum, 247 Euphyllodromia angustata, fainana, 9, 49 Epiphytes, 49 Eugenia aromatica, 160 Euglena sp., 166 145 9, 10, calcarata, 10, 147 297 9, Eucomonymphidae, 176 Eudromiella bicolorata, 31-32 48 a::teca, 9, 102 fm/a, 175, 176 Epilampra abdomen-nigruni 9, 89 10, 77, Eucalyptus, 48, 231 Eucalyptus sp., 160 1 1 Enterobiiis vermicidaris, 209 annandalei, cotmmmior, 113 (see 10, 47 biolleyi, 10, 49, 145, 158 also Brome- liads) caraibea, 10, 147 carbonaria, 10 Epomphaloides ischnoptcrae, 249 Eremoblatta subdiaphana, 9, 28, 316 Ergaida capensis, 9, 49, 77, 311 scarabaeoides, 9, 19 decipieus, 10, 49, 346 dimidiata, 10, 49, 147 jerrum-cquimim, floridana, 6, 10, 10, 49 49-50, 77, 100, 140, Ericaceae, 160 143, 144, 152, 162, 166, 171, 224, Erinaceidae, 283 255, 280, 297, 319, 326, 333, 334, Erinaceus europaeus, 283 338, 339, 341, 345, 346; pis. II, sp., 283 Erythrina glaiica, 35, 156 Erythrocebus patas patas, 180 29, 34 galeoides, 10, 50 improcera, 10, 297 — SMITHSONIAN MISCELLANEOUS COLLECTIONS 452 — Continued. Enrycotis Continued. Flies kevani, 10, 50, 158 phorid, 227 lixa, 10, 147 sand, 322 manni, sarcophagid, 229 298 10, opaca, 10, 50 tachinid, 228 sabalianiis, 10 Foeniculurn vulgare, 237 Eiiscorpius carpathicus, 212 Fondia germanus, 212 sp., 318 Food, cockroaches as human, 286-287 italicus italicus, 213 Euthlastoblatta abortiva, Euthyrrhapha nigra, 10, contamination 10, 24, 50, 143 of, 19 68, 72, 77, 83, 87, appendigaster, 235, 226-237, Forficulid, 339 Forficulidae, 226 238, Formica, 267 omnivora, 268 rufa, 316 239, 350-352 312 desjardinsii, 236 rtifibarbis, dimidiata, 239 impressa, 242 316 subcyanea, 312 laevigata, 236 Formicariidae, 280 minuta, 235 prince ps, 235 punctata, 240 Formicas, 316 Formicidae, 267, 311 Formicinae, 311 sericea, 242 Fossils, 14 sp., 237, 251 Fowl, guinea, 343 239 jungle, 277 Evaniidae, 93, 234-243 Fox, 288 Exothea paniculata, 36, 158 Ej^eworm, chicken, 68, 164, Freycinetia 204-205, 277, 278, 343 sp., Frogs, 4, 270-272 Fagaceae, 152 Fungi, Felidae, 289 catus, 289 Fig, 344 Fighting, intraspecies, 323, 336-337 Filarien, 202 Fir, Scotch, 140 Fish, 3, 268-269 Flacourtiaceae, 159 Flagellate, unidentified, 176 Flies, conopid, 228 ichneumon, 236 muscid, 229 3, 353 127-139 hyperparasitic in protozoa, 188 pardalis meanisi, 290 Fennel, 237 Fern, 44, 140, 164 141 Fringillidae, 282 tree, 270, F^/u 162-166, 314, 318, 319-324 Eutrichomastix sp., 169 Eutrichosomella blattophaga, 245 Evania, 237, 350 abyssinica, 239 sitbspinosa, 83 of cockroaches, 16, 21, 31, 32, 45, pacifica, 10, 77 sp., VOL. I4I Imperfecti, 129 Fungus gardens, 313 Fusiformis lophomonadis, 119, 173 Galebia aegyptiaca, 195 Galcrita janus, 347 Galliformes, 277 Callus sp., 277 Gamocystis tenax, Geckos, 272,, 354 Cekko gecko, 272 168, 184 178, INDEX 453 Gekkonidae, 272 Greenhouses, 68, 80, 81, 155, 157, 162- Geomys 165, 342 Gregarina blaberae, 184 24 sp., Geoscapheus robustus, 10, 319, 344 Geotrichmn candidiim, 131 blattae orientalis, 181 Geraniaceae, 156 blattarum, 168, 181-182 Geraniums, fastidiosa, 182 156, 164 Gesneriaceae, 161 gibbsij 182 Giardia intestinalis, 171 sp., illinensis, 183 172 impetuosa, 183 Ginger, wild, 151 legeri, 183 Glands, secretory, 345-347 Glomeropitcairnia erectiflora, 31, 144 Glyciphagidae, 218 nco-brasilicnsis, 183 panchlorae, 183 Goat, 319 parcoblattae, 183 ohioensis, 183 Goldenrod, 162 rhyparobiae, 168, 183 Gongyloncma sandoni, 183 inglnvicola, 205 neoplasticum, 191, 206, 287 serpenfida, 184 thomasi, 184 pulchrum, 192, 206 sp., Gregarine, 181 206 ; pi. 28 Gregarinida, 181 Gopher, pocket, 24 Gordiaceo, 202 unidentified, Gordian worms, undetermined, 202 ; pi, 184 Gregarlnidae, 181 Gordiidae, 201 Gregariousness, 328, 330-336 physiological effects, 334-335 Gordioidea, 201 Grevisia, 144 Gordius, 192 Gromphadorhina 29 orient alls, 202 Guano, raphaelis, 202 "Gram pi. 12 Gryllacrididae, 226 pilosus, 202 16, 18-23 Guanobies, 16-17, 19 202 Gossyptum ; Gru-gru, 142 Grubs, 317, 319 hlattae orientalis, 202 sp., laevigata, 10, 330 portentosa, 10, 298, 330, 354 aquaticus, 201-202 Guava, spp., 158 positive rods," 127 Guinea Gramineae, 141 Gum Grapefruit, 66, 67 Graptoblatta notidata, 54, dz, 67 Guepe ichneumon, 258 Gumbo 10, 50-51, 147, 151, 158-161, 163, 298, 141, 339 Grass, 37, 40, 42-53, 55, 59-62, 64-66, 69, 141-142 pigs, 288 tree, 36, 53, 59, 61, 62, I55 limbo, 43, 44, 157 Gymnasia niidipes, 279 Gymnopithys leucaspis, 280 Gyna kaaitngulana, maculipennis, bunch, 64 sp., desert, 27-30 tristis, feather, 46 vetula, 10 10, 19 10, 19 298 10, 20 marram, 47 Rhodes, 39, 141 Habitats, 15 saw, 42, 50 aquatic, 30-33, 48 wire, 41, 52 beach, 36, 42, 47, 52, 53, 58, 63, 69 cave, 16-17 Grassquit, Carib, 282 — — SMITHSONIAN MISCELLANEOUS COLLECTIONS 454 Helminths Continued, Habitats cultivated areas, 38, 39, 53, 58 desert, 25-30, 39, 59, 65, 67, 69 dumps and rubbish, 38-40, 50, 53, 69 55, 67, flatwoods, 37, 41, 42, 53, 60 forest, 36-38, 40-47, 49-57, 59-66 grassland, 41, 42, 52, 53, 64 hammock, 41-43, 50, 52, 53, 59, 60, 68 marsh, 42, 50 outdoor, 33-70 rotten wood, 34, 202 ova carried by cockroaches, 208210 Hemiblahera briinneri, 10, 51, 156 Hemidactylus frenatus, 272 Hemiptera, 226, 348 Hemithyrsocera histrio, 7 Hemlock, 44 Henicotyle antillarum, 35, 38, 44, 45, 47, sand dune, 47, 61 sandhills, 36, 41, 42, 47, 50, 52, 60 scrub, 36, 37, 40-42, 48, 50, 52, 55, 57-60, 62, 66, 67 68, Continued. cockroaches as primary hosts, 192- 39, 49, 53, 55, 58, 60, 67, sp., 289 Herpetomonas periplanetae, 167 Herpomyces, 93, 127-129 amasonicus, 134 anaplectae, 134 appendicidatiis, 134 arietinus, 128, 134-135 steppe, 46, 47 chilensis, 135 stones and rock, 29, 31-33, 36-40, diplopterae, 135 43, 45-57, 59, 60, 62, 64, 67-70 structures, 70-90 gracilis, 135 grenadinus, 136 ships, 82-87, 232-233 41, 42, 46, 51, 53, 58, 61, leurolestis, 136 loboptcrae, 136 macropus, 136 68 winter, 38-39, 61, 71 nyctoborae, 136 Hackberry, 50 panchlorae, 136 Hadrurus panesthiae, 136 arisonensis, 213 Hamamelidaceae, 155 HammcrschmidticUa paranensis, 136 dicsingi, 124, 190, periplanetae, 137, 138 phyllodromiae, 137 191, 195-196 platyzosteriae, 137 neyrae, 196 stylopygae, 128, 137-138; pi. 27 neyrai, 196 Haplosporidia, 185 supellae, 138 Haplosporidiwn periplanetae, 185 Harpalus pennsylvanicus, 229 sansibarinus, 138 blattae, 177 Heather, 46, 47, 160 Hebardina concinna, 10, 278 Hedgehog, 4, 283, 354 Hedychium, 163 Helianthiis sp., 162 2-3, 190-210 cockroaches as intermediate hosts, 203-207 tricuspidatus, 138 Heterodon contortrix, 276 platyrhinos, 276 Hcterogamia aegyptiaca, Heterogamodes kriigcri, 13 10, 20 roseni, 7 rugosa, 10, 28 Heterofnetrus longemanns, 211, 213 Heliconia, 40 Helminths, 27 ectobiae, 135 aircraft, 87-90, 232 Hartmanndla pi. forficularis, 135 land-based, 73-82 63, 66, ; chaetophihis, 135 344 swamp, 10, 51 Herpestes javanicus auropunc talus, 289 49-52, 56-63 soil, 37, VOL. I4I Heteropoda regia, 215 venatoria, 215 Heteroptera, 318 ; pi. 30 INDEX 455 Hylidae, 270 Hexamita, 102 cryptocerci, 171 Hymenolepididae, 208 periplanetae, 171 Hymenolepis dimimita, 203 171 sp., 2-hexenal, 234, 311 2, cockroach-hunting wasps, 255-266 280, 345-346 6, host Hibiscus, 36, 40, 68, 158 by selection Hickory, 53, 60, 61, 63 Hierodula tenuidentata, 224-225, 355 hosts of commensal cockroaches, 311-314 predators and parasites of cock- 32 roach eggs, 234-254 Hypermastigina, 172 Histiostoma feroniarum, 217 Holocompsa astcca, 10, yj cyanca, Hyper alius 78 10, Hypoaspis picturattis, 271 nitidula, 10, 78, 148, 339 217 Hypocreaceae, 134 Hypocreales, 134 zapoteca, 10, 20 Hyporliicnoda, 146 fulva, 10, 298 me tallica, Hololampra 10, 51 Hyptia bivittata, 10, 51 chavcsi, 10, 51, 155 marginata, punctata, sp., 33 240 thoracica, 239, 240 298 sp., Icteridae, 282 141 51, Icterus portoricensis, 280 Holomastigotidae, 172 Holoquiscaliis brachyptcnis, 282 Homalodcmas cruralis, perissa, 174 Ignabolivaria bilobata, 10, 52 9 Homalopteryx laminata, scotti, Idionympha 186 Holotricha, 10, 52, 151 sapiens, 286 Honeydew, Hookworm, 237, 242 ova, 210 Hoplonymphidae, 173 Hoplosphoropyga babaulti, 10, 20 Hormetica apolinari, 10, 51 laevigata, 10, 52, 143, 145, 148 spp., 148 ventralis, 10, 148 Hormurns Iguanidae, 273 Ilex cassine, 66, 158 coriacea, 41, 158 9 Hominidae, 286 caudicula, 213 Hornbill, gray, 279 Huckleberry, 43, 57 Humidity reactions, 35, 72, 129 Hygrocrocis intestinalis, 124 Hygroreceptors, 72 Hygrotaxis, 72, 331 Hyla pi. 51 Hololeptoblatta Homo dorsalis, 239 reticulata, 239, 240 10, 51 10, sp., harpyoides, 239; maciilata, 10, 51 sp., egg parasites, 254-255 Hibiscus rosa-sinensis, 68, 158 pi. 208 sp., Hymenoptera, Hexamitidae, 171 cinerea, 270 lucida, 41, 158 rotunda, 257 Incertae sedis, fungi, 139 protozoa, 176 Inermicapsifcr niadagascariensis, 203 Infestations of cockroaches, 331 Inga laurina, 35, 156 vera, 35, 66, 67, 156, 3^7 Insecta, 224-268 Insectivora, 283 Inuus sylvanus, 286 Invertebrates in biological control, 349- 352 lodamoeba sp., 180 lolina nana, 219 lolinidae, 219 Ipomoea tiliasea, 66, 161 Iridomyrmex hmnilis, 268, 319 SMITHSONIAN MISCELLANEOUS COLLECTIONS 456 Ironvvood, 164 Ischnomyrmex Lactobacillus jcmieiiti, 109 sp., 316 Lactobacteriaceae, Ischnoptera, 134-135, 319 borealis, 12 109 Laelaptidae, 216 Lamproblatta albipalpus, cavernicola, 14 coiiloniana, 12 52, 10, 142, 164, pL 12A 298, 316, 336, 355; divisa, 12 Lampyrid, undetermined, 230; Lampyridae, 230 33 Larvaevoridae, 228 Lasius aliemis, 268 panamae, Lastrea aristata, 163 Latiblattclla chichimeca, 10, 53 pennsylvanica, 12 10, 53, 340 10, 54, 145 lucifrons, 11, 54, 146, 255 remyi, 14 rehni, 11, 54, 140, 144, 339 occidentalis, 10, 78, 145 nt/a rw/a, pi. Laniidae, 281 major, 12 rx</fl 148 Lanius ludovicianns, 281 12 podoces, 10, 53, vieridionalis, 10, 53 deropeltiformis, insolita, VOL. I4I 10, 53, 78, 89, 148, 161, 298 spp., 85, 148 vitrea, 11, 143 sapoteca, 11, 54 rujesccns, 82 Latindia armata, 8 schenklingi, 10, 142 spp., 85, 89, 267, 298, Latrodectus indistinctus, 215-216 316 uhleriana, 12 uhleriana fulvcsccns, 12 Ischnuridae, 213 mac tans, 216 Lauraceae, 153 Leafhoppers, 320 Legumes, 165 Leguminosae, 155 Lcidynema appendiculata, Isoptera, 310 I so trie ha caulleryi, 187 Isotrichidae, 187 124, 190, 191, 196-198 appendiculata var. amcricana, 197 Jasmine, 45 Jasminum puhescens, 45 Jay, blue, 280 appendiculata var. hispana, 197 appendiculata var. indiana, 197 cranifera, 198 Florida, 280 delatorrei, 198 Jucaro, 66 nocalum, 198 Juniper, 59 Juniperus sp., 65, Leidynemclla fusiformis, 198 141 Jiissiaea natans, 160, panesthiae, 198 164 paracranifera, 198 Kakerlac schacfferi, 12 Kakkerlac, 260 Karnyia discoidalis, 10, 298 Kiawe tree, 164 Lemur laterale, 274 coronatus, 283 fulvus, 283 tardigradus, 284 Kingbird, grey, 35, 318 Klebsiella pneumoniae, 113 Kojukei, 277 Kuchinga hemerobina, Leiolopisma 10, 298 rcmofa, 10, 141, 148 Kiichenschaben, 100, 169, 171- 173, 188 Lemuridae, 283 Leontocebus oedipus, 285 rosalia, 285 ursulus, 285 Lepidoptera, 316-318 Lepomis Laboulbeniaceae, 127, 134 Laboulbeniales, 134 pallidas, 269 Leptodactyhts albilabris, 271 pentadactylus, 271 INDEX Lcptonwnas blaherae, 167 Lophomonas, loi 167 sp., blattariiin, lOi, 168, 172-173 Leptopelis calcaratus, 271 riifus, 457 striata, loi, 119, 168, 173 271 sulcata, 173 Leptospironympha, 102 Lorn tardigradus, 284 eupora, 172 Lorisidae, 284 rudis, 172 Lycolaimiis, 192 wachiila, 172 Lycosa avida, 216 Lesser apple worm, 65 Lejicophaea madcrac, 11, 86, 89, 119, 148, hclluo, 216, 347 54, 70, 71, 78, 168, 162, 171, 203, 216; sp., pi. 30 Lycosidae, 216 204, 213, 224, 236, 247, 299, 322, 329, Lygaeidae, 226 330, 333, 335, 339, 342, 347; pis. 13, Lymantria monacha, 121 28 sp., 299 Macaca striata, 13 surinamensis, 13 rhesus, 286 Leurolestcs circumvagans, 11, pallidus, II, 71, 78, 299, 78 sylvanus, 286 339 Macaque, 286 Macropanesthia rhinoccrus, Macrospironympha, 102 Lice, plant, 320 Lichens, 47, 5i, 5^, 64, I53 Liliaceae, 146 bellicosus, 311 Lime, 45, 66, 157, 164 Liopicus mahrattensis, 279 malacccnsis, 311 natalcnsis, 311 styraciflua, 155 Maize, Lissoblatta fiilgida, 13, 157 LitopeUis biollcyi, 145 11, 54, Malpighian tubules, parasites 185, 186 148 Malvaceae, 158 Mavicstra olcracea, 120 334 Lizards, 272-276, 353-354 Mammalia, 283, 354 Mammals, 3, 283 Lobogynioides, 217 Lobolanipra sttbaptcra, Loboptcra dccipicns, 2>^6, mallei, 119 Malojillo, 31, 48, 141 bispinosa, 11, 54, 58, 145, 148 11, 54, 55, 64, 142 Malleomyccs deianira, 11, 54 spp-, 11, 11, 55 55, 100, 299, 350 dimidiatipcs, 11 Mangifera indica, 157 Mangoes, 45, 157, 164 Mangrove, 36, 42, 53, 66 Mantidae, 224 cxtranca, 11 Mantis, Carolina, 225 sakalava, 11 Chinese, 226 European, 225 sp., 299 thaxteri, li, 55 Lohopterella dimidiatipcs, 338 Locustacarus 344 Macrotermes barneyi, 311 Lily, Easter, 146, 164 musarum, 11, 55, xylopletha, 172 Lily, Brig, 84 Liquidambar cyclopis, 179 mtdatta, 286 sp., 11, 55, 299, 219 Lophoblatta arazvaka, 11, 55, Lophocents birostris, 279 Lophomonadidae, 172 142 Mantis religiosa, 225 Maple, 61-63 Marava arachidis, 339 Marcta acntivcntris, 11, 317 Marctina uahnka, 11, Margattca sp., 317 Marsupialia, 283 142, 155 in, 177, SMITHSONIAN MISCELLANEOUS COLLECTIONS 458 Mines, 16, 19-21, 23 Miroblatta silphoides, 10 Mastigophora, 166 Mastigoproctns gigantcus, 211 sp., VOL. I4I Mirotermes, 102 211 Maze, 335 Mediterranean Mites, 211, 216-222, 318, 320 2, 3, cheese, 222 252 fruitfly, cockroach, 219, 221 Megachilidae, 314 Megalixahis Megaloblatta blahcroides, control, 222 fornasinii, 271 11, mushroom, 218 55 unidentified, 220 riifipes, II Mcgalopyge krugii, 35, 317 Megamareta verficalis, 11, 56, MegaseUa sp., 227 Melanerpcs portoricensis, 280 Melanosilpha capensis, 11, Molossidae, 283 141 Molossus sp., 283 Moluchia (?) dahli, Monadidae, 167 Manas 299 sp., 11, 167 Monastria biguttata, Meleagrididae, 278 sp., Meleagris gallopavo, 278 56 11, 344 299 Meles sp., 289 Mclipona nigra, 314 Mongoose, 289, 354 Monilaceae, 130 Mclittobia chalybii, 248 Moniliales, 129 Memnoniella echinata, 132 Mermis, 192, 202 Moniliformidae, 203 Moniliformis dubius, 203, 270, 271, 276 Mermithidae, 192 Mermithids, undetermined, 192; Mermithoidea, 192 Mesquite, 37, 58, 156 est char ornyia oophaga, 248 kalahariensis, 203 pi. 29 M Metallyticus semiacneus, 225 moniliformis, 191, 204 Monkey, 4, 284-286, 343 Canal Zone night, 284 Darien black spider, 284 white-faced, 284 Metarrhizium anisopliae, 131 Monocercomonas orthopterorum, 169 Metrosideros collina, 160 Monocercomonoides, 102 Methana canae, 11, 56, 326 curvigera, 11, 56, 155, 326 marg'malis, 11, 56, 79, 326 Methionine, loi Micrococcaceae, 106 globus, 169 melolonthae, 168, 169 orthopterorum, 169-170 panesthiae, 170 citreus, 106 Monodelphis sp., 283 Mononychoblatta, 26 Moraceae, 153 epidermidis, 106 Mosquitoes, 322 nigrofacicns, 106 Moss, Micrococcus anrantiacus, 106 46, 47, 56, pyogenes var. albus, 106-107 pyogenes var. aureus, 107 "Mouche spp., 107, 108 Mouse, 103-104 ureae, 107 70-74 Microtetrameres helix, 206 rosalia, Mimicry, 6, Mimosa, 57 white-footed, 24, 205 Miicor guilUermondii, 132 Microsporidia, 185 Midas bleue. La," 260 sand, 24 Microhabitats, 16-17, 25-26, 30-31, 3335, 64 Spanish, 54, 144 285 347-348 sp., 132 Mucoraceae, 132 Mucorales, 132 Muridae, 287 Murorinae, 129 INDEX Mus decumantis, 287 musculus, 287 459 Neisseriaceae, 109 Nelipophygus ramsdeni, 11, 56 Nematoda, 192, 204, 209 Nematodes, 2-3, 190-192, 204, 209 Musa (see banana) Musaceae, 146 Muscidae, 229 Mustelidae, 289 Mutant, white-eyed, 21 Mutualism, 92, 96-102 Mutuals, 95 Musoa madida, 11, 56 undetermined, 201 Nematomorpha, 201 Neoaplectana chresima, 192-193 sp., 192 Neoblattella brunneriana, 11, 145 carcinus, 11, 148 Mycetozoa, 176 celeripes, 11, 148, Mycobacteriaceae, 123 Mycobacterium avium, 123 detersa, 11, 56, 148 324 dryas, 11, 56, 145, 160, 340 eurydice, 11, 57, 145, 340 jricdmannii, 123 lacticola, 123 jratercula, 11, 86, 145, 148 leprae, 123 fraterna, 11, 86 lepraemurium, 123 grossbecki, phlei, 123 laodamia, piscimn, 124 nahna, 11, 86, 145 Proserpina, 11, 57, 145, 160, 339 semota, 11, 57, 148 spp., 124 tuberculosis, 124 Myna (mynah), Myrica 281 149, 324 vatia, II, 148 Myricaceae, 152 wheelcri, 11, 314 Myrmecophiles, 311-314, 316 Myrmicaria natalensis, 316 Myrmicinae, 312 bitaeniata, 11, Neonephrites partiniger, 230 Neorhipidius neoxcnus, 230 Neostylopyga rhombijolia, 11, 79, 70, yy, 86, 138, 209, 210, 224, 25s, 259, 272, 273, 280, 300, 346; pi. 15 Neotoma sp., 23-25, Z7, SO Nepenthaceae, 154 Nepenthes ampularia, 154 Myrtaceae, 160 28 gracilis, 154 sp., narica, 288 154 Nephrites australis, 231 nitidus Riek not Shuckard, 231 nasua, 288 sp., 148, tridens, 11, 148 Myrmeblattina longipes, 11, 314 Myrmccoblatta reJini, 11, 312 Nasua 145 148 spp., 79, 86, 89, cerifera, 43, 152 Namahlatta 11, 57, 11, 289 sp., 231 Nasutitermcs costalis, 317 Nauclidas nigra, 11, 56, 148, 326, 327 Ncpidae, 227 Nauphoeta Nesomylacris cubensis, bivittata, 11 11, 57 relica, 11, 57, 145 brassae, 11 cincrea, 11, 70, 79, 86, 89, 100, 115, Nests, ant, 27, 31 1-3 14, 316 318 116, 123, 162, 163, 192, 224-226, bird, 35, 259, 299, 324, 330, 335, 327, 339; insect, 35, 43, 66, 67, 69, 156, 317, pi. 318 14 flexivitta, 11, 148, 163 punctipennis, 11, 28 rodent, 23-25, 37, 50 spider, 215-216, 317 Necator americanus, 210 Nectandra coriacea, 43, 153 Neuroptera, 227 Nicotiana sp., 161 Neisseria meningitidis, 109 Nocardia sp., 124 SMITHSONIAN MISCELLANEOUS COLLECTIONS 460 Oikomonas "Nocticola" asteca, 149 Noetic ola, blattaniin, 166 167 sp., 166, 149 17, VOL. I4I bolivari, 11, 17, 57 Oligacanthorhynchidae, 203 caeca, 11, 20 Onagraceae, 160 deearyi, 11, 20 Oniscosoma granicoUis, simoni, 12, 20 344 Oothecae, concealment sinensis, 12, 311 Ophion 185, 186 Nosenia periplanctae, Nosematidae, 185 of, 47, 61, 12, liiieiis, 261 orientalis, 12, 33, 58 312 Notila proteus, 170 Notolampra antillarum, Nucleophaga sp., 178 Nyctibora asteca, 12, 144, 159 Orange, 45, 157, 164 Orangutan, 70 Orchidaceae, 151 Orchids, 151, 162-163, 164, 237 Origins of domiciliary cockroaches, 70- 149 12, 145 71 holoserieea, 12, 149 laevigata, 12, 57, 145, 149, 339 Oriole, Puerto Rican, 280 latipennis, 12 Oriolidae, 280 /m;si, 12, 57, mexicana, Orthoptera, 317 145, 12, 2, 15, 16, 25, 34, 85, 88, Otdopteryx meliponarum, 149 noctivaga, 12, 86, 100, 145, 149 Ovipositor, cockroach, 15 obscura, 12, 57, 149, 300 Owls, 276, 353 foment osa, Oxyhaloa bupr est aides, 300 jf^j^rfa, 12, 58, loi, 102, 168, 188 nana, 170 Oxyspiriira mansoni, 68, 191, 204-205, uichanci, 188 278 viannai, 189 Oxyuridae, 209 Nymphytria, 26 Oxyuris aegyptiaca, 195 anstralasiae, 199 Oak, 36-38, 41, 43, 44, 46, 47, 52, S3, 59-66, 152-153 blattae, 193, 196 blattae orientalis, 195, 196 Oats, wild, 52, 142 blatticola, 193 Ocelot, 290 btdhoesi, 200 Ocneria dispar, 126 diesingi, 195 Ocotea catesbyana, 43, 153 Octomitus periplanctae, 171 Odonata, 224 Odontomachus (?) heterogamiae, 199 kUnckeli, 199 macrura, 195 affinis, 314 darns, 316 haematodes insularis var. pallens, 316 Odontotermcs panesthiae, 198 Oxyuroidea, 193, 204, Palaninaeus, 211 sp., 86 doroaxostyltis, 170 188 biiissoni, 12, 70, 79, Oxymonadidae, 170 Oxymonas, 102 300 Nyctotheriis, 92, loi, 102 oralis, 224 314 deusta, 12, 149 156 12, 12, bare-legged, 279 sericea, 12, 149 spp., 86, 149, 63, Opisthoplatia maculata, 12, 30, 32-33 N othoblatta zvasmanni, 12, 300 325-327 tcrviitophila, 12, 311 brunnea, 12, spp., 58, 311 Oikomonadidae, 166 Palissota, 34 Paliurus acideatus, 38 209 INDEX Pallasiomys meridionalis pennicilliger, 461 Paramecium 186-187 sp., Paranephrites xenus, 230 24 Palmae, 141 Parapolyphaga Palmetto, 37, 49, 50, 57, 62, 64, 67, 166 Palms, 45, 50, 52, 58, 64, 143, 150, 164 Parasites, 91, 95-96 coconut, 36, 40, 42, 50, 53, 56, 58, cockroach egg, pis. 33, 9-10 pis. 16, 27, 29, 35 234-254, 350-352 2, 34 date, 143 Parasitism, 91-94 royal, 54, 143 Parcoblatta, 350 americana, sago, 166 sp., ; host selection by, 254-255 64, 68, 143 Pan erectipilis, 12, 267 bolliana, 12, 59, 338 286 Panchlora antillanim, 143, 149 12, 58, caudelH, 12, 59 desertae, 12, 25, 29, 59, 316 cubensis, 12 exoleta, 12, 150, 300 c?/wa, jraterna, 12, 150 julvescens, 12, 24, 59-60, 79, 140 maderae, 11 mwa, 12, 59, 140, 152, 153, /ato, 12, 60, 79, 12, 58, 79, 86, 89, 142, 150, 155 100, 140, 152, 301, 333, 339 notha, 12, 80 300; pi. 16 peruana, 12, 150 pensylvanica, 12, 15, 61, 80, 100, 150 152, 153, 257, 302, 318, 319, 327, virescens, 12, 150 spp., 17, 20, 63, 144, 206, 302, 316, sag ax, 12, 58, spp-, 150, 22Z, 339; Pl- 17 324 333, 336 Panchlorinae, 231 Pandanaceae, 141 Pandanus, 51, 141, 150 uhleriana, 12, 15, 61-62, 100, 302, Pancsthia, 102 virginica, 339; pis. 12, 58, 301, 327, 354; "brevicollis," 194 javanica, 12 laevicollis, 12, 58, 301, 327 lobipennis, 12, 301 Panicmn barbinode, 100, 230, pis. 17, 27, 33 155 Parinarium, 34 Parsley, 237 Parsnip, 240 Partridge, 278 Parvobacteriaceae, 119 328 Passer domesticns, 282 spadica, 12, 301 Passeriformes, 280 141 Passiflora purpurasceyis, 141 Papayas, 62-63, ::ebra, 12, 63, 140, pi. 8 spp., 301, 33 15, 255, 257, 258, 302, 316, 323, 333; angustipcnnis, 12, 100, 300 australis, 18, 12, sp., 37, 159 Passifloraceae, 159 45, 159, 164 Pasteurella mtiltocida, 119 Papio papio, 286 pestis, 119 Parabttthus capensis, 212 Parachordodes raphaclis, 202 Pastinaca sotiva, 240 Paraclevclandia brcvis, 190 Paxylomma Paracolobactrum aerogenoides, 113 p-chlorophenyl, p-chlorobenzene sulfonate, 222 coli forme, 113 Pedipalpida, 211 spp., 113 Pelmatosilpha coriacea, Paradisea papiiana, 280 Parahormetica bilobata, Parameciidae 186 buccata, 255 /'-benzoquinone, 216, 347 simplex, 190 12, 301 151, 160 kcvani, 12, G^ 12, 63, 142, 150, SMITHSONIAN MISCELLANEOUS COLLECTIONS 462 Pelmatosilpha — Continued, Phaetalia pallida, marginalis, 12, 150, 326 purpurascens, 12, 63, 150, Peltomyces 12, 326 sp., 150 blattcllae, 176 sp., Phidon (?) 131 Periplaneta aniericana, 277 Pheasants, 277 Pheidole megacephala, 268 periplanctae, 176-177 PenicilUum 81 12, Phasianidae, 277 Phasianus calchicus karpowi, 277 rotundata, 12, 145 vagabunda, VOL, I4I 12, 63- 17, 20, 64, 70, 71, 80, 83, 84, 87, 89, 98-101, diibius, 13, 64 Phileciton longipes, 314 Philip, Ship, 83 Philodromus pernix, 318 103, 104, 106, 126, 129, 143, 150, 151, Phlebonotiis pollens, 13, 329 156, 162, 163, 166, 190, 191, 20s, 211- 213, 223, 225, 226, 229, 24s, 255, 271, Phlebotomus papatasii, 322 Phoenix dactylifera, 143 272, 276, 280, 302-304, 318, 320, 321, Pholadoblatta inusitata, 323-327, 332, 334-336, 338-340, 342, Phoraspidinae, 329 346-348, Phoraspis 350-352, pis. 354; 19, 33. 316 142 sp., 64, Phoridae, 227 34» 35 australasiae, 12, 21, 55, 64, 70, 71, Phormictopus cancerides, 214 80, 87, 89, 100, 105, 115, 117, 123, Phorticolea boUviae, 143, 150, 151, 153, 158, 159, 163, testacea, 13, 312 209, 210, 214, 225, 304-305, 318320, 323, 326, 327, 333, 335, 338, 339, 341, 342, 348, 350, 351, 354; pis. 20, 32 bnmnca, 13, 12, 64, 70, 80, 150, 153, hieroglyphic a, 10 humbertiana, 8 pi. 21 nigrocincta, 14 cavernicola, 12, 16, 21 obtiisata, 7 jortipes, 13 parenthesis, 14 277,305, 327,335,338; pl. 22 ignota, 12, 81, 115, 327 352 245 Phyllodromica brevipennis, or lent alls, 8 graeca, picea, 12 spp., 21, 90, 305-306, 317, spp., 306, siipellectilium, 13-14 treitliana, 13, lata, 12, 21 352 tartara, 8 13, 13, 64 65 irinae, 13, 65 maculata, 13, 65 megerlei, 13, 65, 153 Periplaneticola mirsaia, 193 /jo/ifa, Perisphaerus armadillo, 12, 329 glomerijormis, 12, 329, 347 sp., 21, 329 pygmaea, Periwinkle, 47 314 Phryganoporus, 317 Phycomycetes, 132 Phyllodromia germanica, 8 hemerobina, 10 171, 305, 323, 325-326, 340, 346; fuliginosa, 12, 64, 81, 158, 164, 251, 13, 311, 13, 65 13, 65, 141 tartara, 13, 65 tartara nigrescens, 13, 65, 141, 162 Physaloptera hispida, 191, 207 Permian, 14 maxillaris, 191, 207, 288, 289 Perodicticus potto, 284 praeputialis, 207 Peromyscus leucopus noveboracensis, rara, 191, 207, 288, 289 205 polionoius rhoadsi, 24 turgida, 191, 207 Physalopteridae, 207 Petitia domingensis, 66, 317 Physignathiis lesueurii, 275 Petroselium crispum, 237 Phytoseiidae, 216 INDEX 463 Plectoptcra Picidae, 279 — Continued. rhabdota, Piciformes, 279 Pigeon, 279, 343 13, 66, 67, Pileocephaliis blabcrae, 184 sp., Pilocrocis secernalis, 66, 317 venniculata, Pimeliaphilus podapolipophagus, 219- 306 Plethodontidae, 269 Plistophora kiidoi, 185-186 Pinaceae, 140 Pine, 36, 41-44, 46, 47, 49-54, 57, 59-67, periplanetae, 177, 185, 186 sp., 140 186 Pineapple, 39, 45, 68, 144, 165 Ploceinae, 318 Finns Ploceus australis, 140 sp., Plumiger caribaea, 50, 52, 53, 59, 64, 140 318 histrio, 7 clausa, 53, 140 Plumilla, 35 echinata, 140 Pneumococcus, 109 Podapolipodidae, 219 40 Podium abdominale, 265 sylveslris, 140 Carolina, 265 Pisces, 268 Plagiotoma blattarmn, 188 Plants, carnivorous, 154 damage by cockroaches, 162- to, 166 dubium, 265 flavipenne, 265-266 haematogastrum, 263, 266, 352 luctuosum, 266 rufipes, 266 higher, 139-162 sp., pitcher, 154 Platyhelminthes, 208 13, Pogonomynncx 306 13, 36 Poinsettia 100 bifida, 13, 151 castanca, 13, 65, 306, novae scelandlae, 345 13, 65, 306, 321, 157 164 3, 103 Polistes, 318, 319 "Polygamia" aegyptiaca, 13 roseni, 7 scabra, 13, 306 Polymastigidae, 169 306 Plectoptcra dominicae, 66, 67, 81, Polymastigina, 169 Polymastix, loi 157 13, 66, 142, 143, 151, 155-158, 160, 161, 317 Polyphaga aegyptiaca, 29, 67, 81, 100, 190, 13, indica, 13, 24, 29, 317 injulata, 13, 66, 142, 155, 156-158, pellucid a, 13 saussurei, 317 lacerna, 13, 66, 140, 153 perscita, 13, 66, 157 13, sp., 21 porcellana, 13, 66, 157, 162 pygmaea, Polypodiaceae, 140 66 24, 25, 29, 67, 71, 114, IIS, 117-119, 306 Polyphagids, 344 Polyphaginae, 25 pocyi, 13, 66, 158, 159 24, 21, 306 floridana, 13 13, 347; pi, Polybia pygmaea, 314 sabalianus, 10 13, badius, 346, Polecat, 4 Poliomyelitis viruses, 346; pi. 23 novac-zealandiac , 13 dorsalis, sp., Poinsettias, ingens, 10 sp., 266 Podocarpus, 42 Platyzosteria analis, armata, 140 13, 67, Pleistophora periplanetae, 186 Plethodon glutinosus, 269 221, 320 Pimelodidae, 268 spp., 142, 155-158, 317 160, 81, SMITHSONIAN MISCELLANEOUS COLLECTIONS 464 VOL. I4I Polystichmn aristaimn, 163 Protomagalhaesia serpentnla, 168, 184 Polyzosteria analis, 13 Protoinonadina, 166 Protospiriira bonnci, 206 limbata, 13, 67 melanaria, 13, Columbiana, 206 306 Pomarrosa, 160 niuricola, 191, 206, Protozoa, Pompilidae, 256 Pompilus bracatiis, Protrclleta floridana, 199 Protrcllina, 199 aurifluus, 199 Ponies, pit (mine), 16, anstralasiae, 199 23 galebi, 199 diversifolia, 152 kUnckeli, 199 euphratica, 47, 152 sp., manni, 200 152 phyllodromi, 200 Poroblatta nigra, 11 spp., Protrelloidcs paradoxa, 200 67 Potamotrygon Jiuniboldti, 268 Protrellus aureus, 199 hystrix, 268 aurifluus, 199 Potamotrygontidae, 268 australasiae, 199 Potato, 165 galebi, 199 Potto, 284 kUnckeli, 199 Poultry, 276-278, 353 Prairie dog, 24 manni, 200 phyllodromi, 200 Predation, interspecies, 92, 319-322 13, 100 Pseudomonas aeruginosa, 104 Predatism, 91-95 Predators, 91, 96, 319 defense against, 343-348 eisenbergii, 105 fluorescens, 105 Pseudomopinae, of cockroach eggs, 234 Pseudomops Prey, experimental, 4 60, 230, cincta, 13, 231 319 laticornis, 13, 145 natural, 4 septentrionalis, 13, 67, 162 Primates, 283 Pscudoplioraspis nebtdosa, Pritchardia, 64, 163 Privies (latrines), cockroaches in, 74, 80, 81, 178, 340 Procyonidae, 288 litula, Pseudoderopeltis aethiopica, Pseudomonadaceae, 104 intraspecies, 92, 322-324 Prodenia 284 101-103, 166-190 cultured in vitro, 170, 178, 179, 188 256 256 Ponerinae, 314 Pongidae, 286 sp., Populus 2, 92, 328 Psocoptera, 318 Pteridophyta, 139 Pteromalidae, 247 320 Prolabia, 339 Prolophomonas tocopola, 173 Prosevania punctata, 240-242, 254-255, Pteromahis sp., 247, 248 Pterygosomidae, 219 Pycnosceloidcs aporiis, 351 pi. 33 Prosoplecta, 348 ; Prosthenorchis elegans, 203, 283, 285, 286, 288-290 spirilla, 203, 283, 286, 13, 81, Pseudosaccharomycetaceae, 129 Psiditmi guajava, 67, 160 12, 58 Pycnoscelus niger, 13, 22 striatus, 13, 22 surinamensis, 288-290 2, 13, 22, 25, 67-68, 70, 71, 81, 87, 90, 100, 138, 139, 142-146, 151, 152, 155-157, 161- Proteus mirabilis, 114 morganii, 114 165, 191, 212, 229, 259, 279, 280, rettgeri, 114 282, 306-307, 324, 330, 333, 335, spp., 338, 341-343, 350, 353; Pl- 24 114 vulgaris, 114 Pyrus sp., 62 465 INDEX Rhisoglyphus tarsabts, 218 Rliicophora mangle, 36, i59 Quail, Japanese, 277 Quercus alba, 152 Rhizophoracea, 159 piibescens, 38 Rhisopus nigricans, 133 rubra, 152 133 sp., spp., 153 opimus, 24 Rhombomys znrginiatia, 152 Rhynchonympha, 102 Quinones, 347 175 torc^a, 174, Rana catcsbeiana, 271 mascareniensis, 271 pipiens, 271, 345 Ranatra 4, 287 kangaroo, 23, 26 Rats, Riboflavin, loi Rictularia coloradcnsis, 192, 205 nitidioides, 231 wood, 23, 26 Rattus norvegicus, 287 ratlus, 287 sp., Rednvius christophi, 227 fcdtschenkianus, 227 boissyi, 231-232 denisi, sp., scutellaris, Ripiphoridae, 230 272-276, 353 of, 320 Roach (see cockroach) Robshelfordia circumdiicta, Reticulitennes sp., 317 Retortamonas blattae, 167, 168 orthopterortim, 169 307 167 Rodentia, 287 jRo.yo sp., 155 Rosaceae, ISS Rose, 155, 163-165 Roundworms, 2 (see Helminths) Rhabditoidea, 192 Rhamdia sebae, 268 Roystonea rcgia, 143 Rhantus pacificus, 230 Rhicnoda natatrix, 13, 33 Riibiis spp., 155 sp., 13, longiuscula, 13, 307 Reptilia, 272-276 sp., 233 232 151 Repellents, 345-347 eggs 232 pectinicornis, 84, 232-233 sp., 3, 231 Ripidiini, biology of Australian, 231 Ripidius, 230 287 Reduviidae, 227 Reduviids, tindetermined, 227 Reptiles, 158 Riekella anstralis, 231 Norway, 287 Renealmia 13, 68, orientis, 13, 68, 158 Ranidae, 270 spp., 11 Rhythms, diurnal, 335 Rhytidomctopnm dissimile, i^jaffa fulgida, 13, i57 227 sp., Rhyparobia maderae, 146 Rhinopsis caniculatus, 257 Rhinotermitidae, 310 Rhipidioidcs ableptus, 230 adynatus, 230 fuscatus, 230 helenae, 230 mollis, 231 rubricatus, 231 Rhipidiiis, 230 Rhipiphoridae, 230 Rhizobiaceae, 106 Rubiaceae, 161 Rushes, 46 Rutaceae, 157 5'aba/ palmetto, 50 Saccharomyces sp., cerevisiae, 133 133 Saccharomycetaceae, 133 Saccharomycetes, 129 Saccharum officinaruni, 141-142 Saccinobacnlus, 102 ambloaxostylus, 170 doroaxosfylus, 170 SMITHSONIAN MISCELLANEOUS COLLECTIONS 466 Saccinohaciilus lata, — Continued, 170 minor, 170 undescribed 170 spp., Saimiri sciurea, 285 Salamander, 269 Salganea morio, 13, 334 VOL. I4I Sarcophagidae, 229 Sarracenia flava, 154 minor, 154 purpurea, 154 variolaris, 154 Sarraceniaceae, 154 Sauria, 272 Salius verticalis, 256 Scale, diaspine, 237 Sceliphron caementarium, 248 Schistosoma haematobium, 208 S'alix sp., 152 Schistosomatidae, 208 Salicaceae, 152 Salientia, 270 Schizomycetes, 104 Salmonella anatis, 114 choleraesuis, 114 Schizophyta, 104 enteritidiis, Schwenkiella icemi, 200 114 Scincidae, 274 Scolopendra, 222 morbificans, 115 paratyphi, 115 schottmuelleri, 115 cingulata, 223 typhimurium, 116 morsitans, 223 typhosa, 117 subspinipes, 224 Salmonella (Type (Type (Type (Type (Type (Type (Type (Type (Type (Type (Type (Type (Type sp., sp. Adelaide), 115 Barellly), 115 Scolopendromorpha, 223 Bredeny), 115 Derby), 115 Kentucky), 115 Scorpionida, 211 Kottbus), 115 Meleagris), 116 Montevideo), 116 Newport), 116 3, 211, 349 Scorzonera acanthoclada, Scutigeromorpha, 222 Sea grape, 43, 69 Seaweed, 47, 55 Secretions, repellent, 35, 156 216, 225 Serpentes, 276 Serratia marcescens, 117-118; pi. 16 Setaria verticillata, 142 Sarcina alba, 108 Seurocyrnea colini, 207 Severianoia magna, 200 aurantiaca, 108 lutea, 108 severianoi, 200 108-109 Sewers, cockroaches symbiotica, 108 in, 74, 332, 338 Shelf ordella tartara, 8 Shigella alkalescens, 118 ventriculi, 108 Sarcodina, 176 229 6, Sedge, 66 Sapindaceae, 158 Sapotaceae, 160 omani, 229 pi. sexual, 235 Samui, Steamship, 232 sternodontis, 229 ; maculata, 223 Panama), 116 Sarcophaga lamb ens, 162 forceps, 222 Rubislaw), 116 Tennessee), 116 65, Scutigera coleoptrata, 222-223 Scutigeridae, 222 Samanea saman, spp., Scorpionidae, 213 Scorpions, Oranienburg), 116 Salsola kali, 65 spp., 224 Scolopendridae, 223 dysenteriae, 118 2, 229 parody senteriae, 119 Ships, 82-84 cockroaches on, slave, 79, 86, 87 79, 82-87 31 INDEX Shrike, 281 467 Spirillum Sibylloblatta panesthoides, 13, 151 Sida, 64, 159, 163 and /3, 7, 105 sp., 105 Spirocerca sangninolcnta, 203 Spirochaeta blattae, 125 Sideroxylon joetidissimum, 63, 160 Silkworm, bacterium from, 126 blattarum, 125 "disease bacillus," 127 Simbleraslcs jamaicamis, a, periplaneticum, 105, 127 13, 69, 317, periplanetae, 125 stylopygae, 125 334 Spirochaetaceae, 125 Spirochaetales, 125 Simla jacchus, 285 Sinanthropes, 71, 81 Skinks, 274, 354 Spirochaetes, unidentified, 125 Snakes, 276 "Spirochaetoid bacteria," 127 Spirorbis pusillus, 14 Sodium 278 fluoride, Spirostomidae, 188 Solanaceae, 161 Solanum tuberosum, Spirotricha, 187 161 Solenopsis qeminala, 314 Solindenia picticornis, 246, 247 Sound, cockroach, 330 Sow bugs, 347 Spirura gastrophila, 207 Spiruridae, 205 Spiruroidea, 204 Spondias, 67 mombin, 158 Sparassidae, 215 purpurea, 158 Sparrow, 282 Sporozoa, 181 English, 282 Spruce, 44 Squirrel, ground, 23, 24, 26 Spartan, Ship, 84 Spartina marsh, 42 Stagmomantis Spelaeoblatta caeca, 14 Staphylococci, 107 gcstroi, 13, 22 Carolina, 225-226 Staphylococcus 107 albtis, Spermophilopsis Icptodactylus, 24 Staurojoeninidae, 174 Sphaerita, 189 Stegobium paniccum, 99 Sphaerodactyliis sp., Steinernematidae, 192 273 Sphagnum, 52 Stele opyga sinensis, 13, 307 Sphecidae, 264 Sphecophila polybiarum, ravana, 314 310-31 13, termitium, Stenainma andrei, 316 13, 13, 311 Sphodromantis biocnlata, 225 viridis, 225 Spicaria prasina, 130 Spiders, 3, 211, 214-216, 317, 318, 347, 349 pi. 30 banana, 215 ; big, brown house, 215 bird-eating, 214 black widow, 216 button, 215 huntsman, 215 nests, 215-216 Spiniger domesticus, 227 Spirillochaeta blattae, 127 Stenobothrus vagans, 46 Stenophora sp., 181 Stenophoridae, 181 Sterculiaceae, 159 Stictolampra buqueti conciniila, Stigmatomyces sp., 129, 14, 33 138 Stilpnoblatta minuta, 316 Stipa lessingiana, 46 Strepsiptera, 234 Streptococcus jaecalis, 109 liquefaciens, microapoika, no no non-hemolyticus, pyogenes, spp., no no no Streptomyces leidynematis, 197 Streptomycetaceae, 124 124, 196, ; SMITHSONIAN MISCELLANEOUS COLLECTIONS 468 Streptopelia chinensis, 278 Streptostomum VOL. I4I Sysiellogaster ovivora, 248 Sysygium aromaticum, 160 gracile, 195 Strigidae, 279 Ssepligetella scricea, 242 Strigiformes, 279 Strongyloidea, 209 Tachinid, undetermined, 228 Sturnidae, 281 Tachysphex Stylogasler spp., 228 sty lata, 228 264 fanuiensis, 264 Stylopyga americana, 12 Styphon bakeri, blatticidiis, coriaceus, 264 lativalvis, 344 14, 69, 264-265 Taenia echinococcus, 208 saglnata, 208 Subiilura jacchi, 204 Subuluridae, 204 Taeniidae, 208 Sugarcane, 37-39, 45, 48, 64, 66-69, 141-142 51, 53-56, 58, Siiifuncma candelli, 200 Taenioidea, 208 Tamarind, 51 Tamarindus indica, Sulfur, flowers of, 222 Tapeworm, undetermined, 208 1 56 Sunfish, 269 Tarachodes tnaurus, 226 Sunflower, 67 Supella hottentotta, Tarantula, 214 Tarantula, 214 siipellectUimn, 29 14, 14, 69, 70, 71, yG, Tarsiidae, 284 82, 87, 89, 100, 115, 117, 142, 161, Tarsius, 284 165, 205, 216, 223, 232, 255, 307, Tartaroblatta karatavica, .327, 333, 335, 339, 342, 349-351 pis. 25, 30, 31, sp., 34 14, 70 Ta^M novemcinctiim, 287 Taxodiaceae, 140 Tcgenaria, 214 90 Swift, edible-nest, 318 Teiidae, 275 Swiss chard, Temnopteryx, 134-135 153 47, Symbiosis, 25-26, 92 dcropeltiformis, 10 Symbius blattarum, 232 Symploce bicolor, 14, 82 obliqiietruncata, 14, 317 phalerata, 14, 307 platysoma, 8 bilabiata, 14 curta, 14, 22 Temperature preference, 25, 71 Tenadores, Steamship Tenodera aridifolia sinensis, 226 flagcllata, 14, 69, 307 Termes breviramis, 14, 22 cavernicola, 14, 21, 22, 318, 333 hospes, 14, 82, 161, 339 jatimicana, 14, 69 sp., parenthesis, 14, 307 14, 22 riificollis, 14, 69, sp., 142, 161, 307 Termopsis Testudo 90 Syncephalastnim sp., blattae, glosshiae, 250 ischnopterae, 249 sp., 248 317 horsfieldi, Tetragenous 133 Synonyms, 5 Syntomosphymm 311 Termites, 57, 69, 102, 310-311, 317, 320 comejen, 317 Termitidae, 311 14 remyi, 311 316 malaccensis, 311 kevani, 14, 29 lita, bellicosiis, flavipes, sp., 24 127 Tetralopha scabridella, 35, Tetrameres americana, 207 66, 67, pattersoni, 207 Tetrastichodes asthenogmus, 249 317 INDEX Tetrastichiis aitstralasiac, 249 hagenozmi, 94, 249-253, 255, 349- Tody, 279 Torula acidophila, 132 gropengicsscri, 132 351; Pl- 34 pcriplanctae, 253 sp., 469 rosea, 132 Tondopsis, 130 249, 254, 351 Toxoplasma Tetratrichomastix blattidarum, 170 Tcttix kraussi, 46 Trapping, Thallophyta, 129 Trecferns, 36, 42, 50, 56, 57, 139 Thecadactylns 273 Theganopteryx straminca, gondii, 190 53, 59-63 Trees (see under sp., 14, Thelastoma aligarhica, 200 brevicaudatwn, 194 bidhocsi, 201 hcterogamiae, 199 icemi, 200 30 common or scientific name) in general, 35-68 Trematoda, 208 Treponema parviun, 125 stylopygae, 125 Treponemataceae, 125 Triatoma arthurneivai, 227 Indiana, 194 magalhaesi, 194 Tribuhis pachyjidi, 200-201 palmettum, 201 Trichomaslic orthopterum, 169 Trichomastix, 169 panesthiae, 198 Trichomonadidae, 171 river oi, 201 Trichomonas hominis, 171 sp., 63, 156, 163, Thelastomatidae, 190, 193 orthopteronmi, i6g Thelaziidae, 204 sp., Thelyphonidae, 211 171 Trichonympha, 102 Thclyphonus giganteus, 211 acuta, 174 Theobroma c/5'oa, cacao, 159 Theraphosidae, 214 Thcrea nuptialis, 14, 174, 175 chula, 176 344 Theridiidae, 215 grandis, 176 /a to, 176 Thistle, 60 okolona, 175, 176 giant, 51 Thyrsoccra parva, 176 cincta, 13 Tiaris bicolor ornissa, 282 Tillandsia fasciadata, 144 Trichonymphidae, 174 Trichostrongylidae, 210 Trichostrongylus sp., 210 nsneoidcs, 144 Trichotarsus, 218 uttriculata, 50, 144 Trichuridae, 210 sp., 31, 36, Trichuris trichiura, 210 144 Timulla eriphyla, 264 Tjwa auslralica, 14, 316 brimnca, 30 14, /i/Zro, 14, 28, macracantha , 30 14, Trichuroidea, 210 Trigonopsis abdominaUs, 265 Trirhogma cacrulca, 262 sp., 23 obscura, 14, 30 sp., 165 23 264 Troglobies, 17, 20 Troglodytes aedon, 281 audax, 281 Toads, 270, 353 Tobacco, 165 Troglodytidae, 281 Tockiis birostris, 279 Trogloxenes, 16 Todidae, 279 Todus mcxicamis, 279 Tropidophorns grayi, 274 Trypanosomatidae, 167 Troglophiles, 16, 17, 19, 20, 23 — — SMITHSONIAN MISCELLANEOUS COLLECTIONS 470 Tulip tree, Continued. poliomyelitis Lansing Turtle, desert, 24, 26 strain, 103 unspecified strains, 103 painted, 272 Typhloblatta caeca, 23 Typhloblattodcs madecassus, Tyrophagus Continued. Viruses S3 Turkey, 278 VOL, I4I yellow-fever, 104 14, lintncri, 14, 23 218 Viverridae, 289 Vtdpes sp., 288 noxius, 218 Wasps, 91, 94. 314, 318, 348 cockroach-egg parasites, 234-254, Urinympha, 102 talea, 174, 175 Urodacus novaehollandiae, 213 Uropoda sp., Water, balance, 72 drinking, 72 need for, 71 217 Uropodidae, 217 Vaccinium meridionale , Vanda, 151, 163 V eillonella 43, 160 parvida, 109 Vejovidae, 213 Veromessor andrei, 316 Vertebrata, 268-290 3, Water bug, 349 Wattle, 49 Web worm, 316 Weinmannia sp., 155 Whipscorpion, 211 Verbenaceae, 161 Vertebrates, 350, 352 cockroach-hunting, 255-266, 352 268 in biological control, 353-354 Vespa maculata, 318 Vespidae, 314 Vespiila maculata, 61, 318 Vibrio comma, 105 William Kieth, Steamship, 84 Willow, 47, 63, 152 Woodpeckers, 279-280 Wren, 281 Xerophiles, 25 Xestoblatta festae, Xylosma Deneke's, 105 14, 87 imniaciilata, 14, 23 suavcolcns, 159 Xysticus, 214 metschnikovii, 105 sp., 106 types I Yeasts, and II, 106 tyrogemis, 105 3, 127, 129, 132 Yellow-fever virus, 104 Yucca data, 54, 146 Vinca minor, 161 Vireonidae, 281 Zanthoxylnm caribaeiint, Zea m^iys, 142, 165 Viruses, 103-104 Zelotes, 314 Vireo latimeri, 281 Coxsackie, 103-104 encephalomyelitis, mouse, 104 poliomyelitis, 3, 103 Brunhilde type, 103 Columbia SK, 103 Zeuxevania splendidula, 350 Zingiberaceae, 151 Zootermopsis, 103 Zygophyllaceae, 156 66, 157 93, 243, 249, rr CO ± c/) NOIinillSNI NSTITUTION S3iyvyan LIBRARIES NVINOSHilkNS \ (/) SMITHSONIAI 2 CO 2 Z ;3iHvy9n — libraries CO Smithsonian ~ institution CO NoiiniiiSNi — NViNOSHiikN * (/) Z 2 -I NviNOSHims NOIinillSNI NSTITUTION -I S3idvaaii * 2 libraries smithsonia -^ m CO ;3iavdan libraries •^ CO __ Smithsonian ^ Z \x CO ;3idVHan -:, \ CO >> '" 2 CO NOIinillSNI NSTITUTION CO :z > Z S Z CO SMITHSONIA (/J nvinoshiiw NoiiniiiSNi r- CO _ _ NVINOSHIIWS S3ldVyan LIBRARIES SMITHSONIA CO z Z ^ ^ i'^i Biyvyan libraries r- NviNOSHim NoiiniusNi S3ldVyan — LIBRARIES libraries Smithsonian institution r- NSTITUTION S CO NVlNOSHimS NOIinillSNI NSTITUTION ^—-^ CO institution CO wiW z Smithsonian NOIinillSNI~'NVINOSHlll^S i CO NoiiniiiSNi NViNOSHim S3ldVaan LIBRARIES SMITHSONIA institution ri^. CO C/5 JTION c/) v^ ^ in ± £ NOIiniliSNI NVINOSHilWS S3ldVHan LIBRARIES z z ^ C/) c/J . 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