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SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. 141

“EVERY MAN IS A VALUABLE MEMBER OF SOCIETY WHO, BY HIS OBSERVATIONS, RESEARCHES, AND EXPERIMENTS, PROCURES KNOWLEDGE FOR MEN’’—JAMES SMITHSON

(Pusiication 4470)

CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION 1961

PORT CITY PRESS, INC. BALTIMORE, MD., U. S. 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 An- nual Report and occasional publications of a special nature. As the name of the series implies, its scope is not limited, and the volumes thus far issued relate to nearly every branch of science. Papers in the fields of biology, geology, anthropology, and astrophysics have predominated.

LEONARD CARMICHAEL, Secretary, Smithsonian Institution.

(iii)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 141 (WHOLE VOLUME)

THe BIOTIC: ASSOCIATIONS: OF COCKROACHIES

(WitTH 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

SMITHSONIAN MISCELLANEOUS COLLECTIONS VO aay Peed

Blaberus craniifer, c. X 2. 1. (Photograph by Jack Salmon, Philadelphia Quartermaster Depot. )

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 141 (WHOLE VOLUME)

CRE PIGTIC ASSOCIATIONS OF COCKROACHES

(Wit 37 PLATES)

By LOUIS M. ROTH AND

EDWIN R. WILLIS

Pioneering Research Division, United States Army Quartermaster Research and Engineering Center Natick, Mass.

(Pusiication 4422)

CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION DECEMBER 2, 1960

THE LORD BALTIMORE PRESS, INC. BALTIMORE, MD., U. S. A.

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 prede- cessors, 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 in- formation 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 in- formation 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 studied museum collections and the percent of new species found in recent acquisitions, particularly from tropical and subtropical coun- tries, indicate that at least 4,000 species remain unnamed. Although the group is well known in general terms to nearly all entomologists, there is an almost complete void of information about all except the 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 known about cockroaches.

With the exception of mosquitoes and a few other comparatively small groups of insects on which work has been concentrated, it is doubtful if 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 serious obstacle to progress. Progress can continue only if the scattered literature resulting from the diversified labors of hundreds of contributors is brought together and summarized in thorough and well-organized compilations that can serve as a solid basis for future research.

iii

iv SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

The present work is such a compilation, for it assembles what has been gleaned from approximately 1,700 sources, including correspond- ence with a large number of other workers. Original observations during some eight years of concentrated effort in U. S. Army Quar- termaster research laboratories are a valuable supplement to what others have done, and with this background of experience the authors are especially well qualified to appraise previous work. Seldom has a compilation been done so thoroughly or a single large group of in- sects been the subject of such uninterrupted effort.

The 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 longer are cockroaches re- garded only as disagreeable pests; many species appear to be im- portant, actually or potentially, as carriers of disease. Recognition of this 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 information about cockroach bionomics must be kept in mind. The cited observations of many writers were fragmentary, or their con- clusions disagreed. But it is fundamental to scientific inquiry that we should know and attempt to evaluate the results of previous study, and that is what Drs. Roth and Willis have done. Fortunately, their re- view is readily available. Sometimes, a piece of work fails to be of maximum 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 Natick, 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 financial support of this publication.

ASHLEY B. GURNEY Entomology Research Division United States Department of Agriculture

CONTENTS

Page

[Day eenordil Ueto Cin MnAinn cnbiolo bia oigob ogo UOOe DES DODUISSEe Hon Got Dotna iii ep lntroductOomeusn soci serie ee cto et ers ieiscle te ele cieteiercs c c)steleterel sce I le hvoratcal be ee hen yar b tomo Om OOOO DOCS SUGS aoe bone Ne oetos 2 IMetOdS eyes Sa aioe ae re et cle Sebati stators diets aceite tous 4 ate ODIC SA Re ae ree shoe arco aialaia eee crater e leche e cielo sists 5 TIS Er AHONS) Wa rare eRe Ae wich een re aa ae chet feat aie 7

Te eSpeciesi-of (cockroachess)avsd Sac Secioe nes ho boats clon ele es rie oe 7 IE Keolocical), relationships) fascias eels hats Wolens oreets pire ll ataimrniatet ss es 14 (Cave: Habitats). 3 cave cassie ow c.ae Hee On nee eee eters an eta eres eer 16 CGaveriicolousecockroachestaeeeeeee eee omer: 17 @ockroaches) from buLrOwS ei. \o sale Croletele sate! slarevstereeel stove ols 6 0)2 23

Desert: Hab itatsw icin cveicis si ciccuerot ie share SSR he are oP eo olle ata 25 DESertrCOCKMOAGCIES s aaa yt che tear te Ee P octane eater ets ele welts 27

Aveatic, Hapitats) sacs an wor’ 2 hat basa arora, Heer eae Me eiatein, e's ole 30 Amaplitbious), COCKEOaCHES y 49 a2 «ic Poneto Via elevabetetand ele) Hale aca we. 31

Outdoors habitats yc ey eo A heer cae atererait ue el Leaors eltay averaie 33 Cockroaches from) outdoor shabitats2e/je-eiesin-) cee eles ees oes 35 Stauctinal habitats escrito crate a tere crete retererc eet trevess ete 70 Wand=hasedistractunesu ste itee creel tec ce ereroterstetotey actors tots reks 73 Cockroaches associated with land-based structures....... 74

SAIS oh ccio ia wich aie Siere wis’ aaa > dialele MRNAS aeiaReTA, emia nals aa ese slaiaars &2 Cockroaches) associated: withhishipstetacciast «tie seit score ole 85

AICTE He oi Sa horas RTA ae ore a el Naha tee onahc matee tan eR P ana arcs 87 Cockroaches associated) withyaincralte. sce ost eee sle- 88

TiVen Classification ot uchewassOciatlOlS\tem aeicterasiniiote eters aeteretieieler =e QI Ne INMst ITU LE STIR cits) 2 aiarala a ere cn easier etaels Shaver staiaa tis a mia wlan aehereRe a 96 RACHEL OLUS mavater terete calexorc ton srateuerrs ld iro avorer otal Mar clever ha sone siahet Aiatonetetee 96 Cockroaches in which bacteroids have been found........... 99

RACER Teas ere eee eee eer ea Seen rat aeTn ielsioharsrels stayarelerese 100

IP KOLOZOA aise Tee ee ee eat atalls oh Paterotete sear aiehel sheto-tasanehenctsiaions oye 101

Nise Virisesr associated, wathucocis Oa CheSmre iin aieiciraiieiiatc i tetorelele = elare 103 WIL Bacteria: associated wath) cocktoachesw.-\-isc0 seas 22 4 «cites = 104 NAlMnE ned alicaveAaStSeatmerercmieier rami ravereietcnectolskellslcrororshoveretegsloceie oretaletstats) = 127 Bunei associated, with (cOckraaches.. i... seen cane aeeccesaces 129

IX. Higher plants associated with cockroaches.............+++++04- 139 Damage to plants by cockroaches............+.eeeeeeeeeeeee 162

xe Protozoa associated) with’ Cockroaches: 1.5... ses 0 cle ele cele = = 1606 XI. Helminths associated with cockroaches.............-eeeeeeeeees 190 Helminths for which cockroaches serve as primary hosts...... 192 Helminths for which cockroaches serve as intermediate hosts.. 203 Helminths whose eggs have been carried by cockroaches....... 208

XII. Arthropoda associated with cockroaches.............seeeeeeees 210 DAtNGHaTii nee Ree a etete erento retell sei ayoroiseustoierstaicretene(oleieysucrausiefatersl's 211 Chilopoda:. \ovenesccess deen e EP Wemete Ine Arai Sart nu ate ce diese weatere he 222

vi SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. IAI Page

SEC tay ree cise sarees ate rar ere cia ralene oie losaycuetete ler hrs clare) satel ereeeteencsT stare terete 224 EVymienopterat ce. dice seh o's aires naive treelovele, stelraretom tates ean tarss 234

Predators and parasites of cockroach eggs............... 234

Host selection *by, eee: parasites sie lack oes «mtv ce 254 Cockroach-hunting wasps: ayesieicleisioriclsterseietielsieierciatieteye 255

Ants predaceous, On: COCKrOACHES oarcia ere) aye ctaie) Veieisiaiss sey ile 266

XIII. Vertebrata associated with cockroaches..............0- eee Colt SOS: IPISCESH ciataitere a tlateiers, steterelsicveleraleranes asin: wet Shay elisa btoke etatensterarar e.evaters 268

7 -Xvanh 0) 061 YC PR eI A AS CO aAEES Oe OG oC Dob ao so lSocmpods 269

esata Tags Seatapeconers seine vate’ gusce avails: she! crevasse coheree tote re Chem systetenonerNeneke ctereratts 272

DANES sce ia ots esoaverte a: o evs eis.e,ev nkayevereefel ors /skeie eveveveraley sxcvepe te toueied enue rabereiceone 276 Mammatnial lita sis)c s sseveverss auc alesse oo wsevovelevayesiensoterclate ciare choke peeyeealerainee che 283

XIV. Checklist of cockroaches and symbiotic associates.............6. 290 DEV. ACOCKCGACHES, AS. COMMENSAIS << 5:.)s.c, <cs.cie sino clniers olaalerea arte aiece alors 308 flosts) of commensal ‘cockroaches. 14+ a atetieeioereers sites 309

Checklist of commensal cockroaches with their hosts......... 313

Obscure vassocsiations, a 4. dieda.cc esereioknararee eee i acals onan

Vel Cockroaches| vas) predators sje sciecieee ee ee eee events 317 Interspecies -PKECATIOM {s,.</c)s)s/oco1s) oiepstctovauctarentarsreteteererane otto craverene 317 Intraspecies predation’ cara gac dee seiea Tee eine eee 320

DOV Associations) aniong cockroaches 1s. sanen pomereine aloes 6 eine 322 Bamilial associationsisncm en seeds eee ee eee 323 GKEGAPIOUSHESS:: 5 a5)5)5;5 4's) asa wins dhevora saber Cena a aeRO Oe Sele eR 328 Intraspecies fighting, aisciep« sn occ ela ee aera ee eee eine 334 Interspecies compatibility i. .sm see ee oon iene 335 Interspecies: antagyonism 3.01 cisacia da coctmadeeeieen ee ee 339

XVIII. Defense of cockroaches against predators.............2+eeeeees 341 EX. he, biological ‘control. of cockroaches iiss sesame ewan nese 346 Invertebrates: \\s.35:/tadid.eweenrcnee taste rate ee oeeie eee & 347 SVICHEEI TA ECS fi ics, :'s(aivicia che, siete RSA Wee PRE doy senate tere Se sccle ae!) BO BAUCLETIO WLC ETACTIES oc 5-95 io .5: 4:82, <; sos) b'5/- 6) os) caro) al Oren Toe ke meee he es 352 BR CREE CIIC OS) 5) aa) ohs: 6/6 oral nit e/eiosoro;e.acuinj sve, o\ ev aso) ofs\ syarard Cloke aR RET IeT: etnte 354 Pndex joes strere cle aloiata s]c?ehotalo evsje.s aisle ateractevelelaletatenn atte stenatateneReralevelicrer aie ie letar ais 439

THE BIOTIC ASSOCIATIONS OF COCKROACHES:

By Louis M. Rory anv Epwin R. WILL Is ?

Pioneering Research Division, United States Army Quartermaster Research and Engineering Center Natick, Mass.

(With 37 Plates) With most of us collectors the life history of an insect begins in

the net and ends in the bottle. HanitscH (1928)

I. INTRODUCTION

RECENTLY we brought together much of the literature linking cock- roaches with the transmission of certain organisms that cause disease in man and other vertebrates. In that paper (1957a) we concluded that cockroaches, being potential vectors of pathogenic agents, should not be regarded simply as minor annoyances. Obviously the associa- tions of cockroaches with agents of vertebrate diseases are of more immediate importance than their relations with pathogens of lower animals or with nonpathogens. On the other hand, cockroaches are of general economic as well as medical importance, and their control is sought by many who are unaware of their medical significance. That the control of domiciliary cockroaches is far from satisfactory may be ‘nferred from current entomological and pest-control journals in which new insecticides are continually advocated to replace others found to be inadequate. Possibly new approaches to the control of cockroaches are needed. Whether these lie in the direction of increased use of parasites and predators for the biological control of these insects remains to be seen. In any event, the more we know about any insect, especially its ecology, the greater the likelihood of achieving satis-

1 Part of the cost of publication of this monograph was borne by the Office of Naval Research, Department of the Navy (through the American Institute of Biological Sciences), and by the Quartermaster Research and Engineering Cen- ter, Department of the Army.

2 Present address of both authors, Central Research Laboratories, United Fruit Co., Upland Road, Norwood, Mass.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 141 (WHOLE VOL.)

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. TAT

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. We hope that such inadvertent omissions will not seriously impair the usefulness of this compilation. Whatever its defects, this review should be a unified source of information for all who are interested in the biotic associates of cockroaches.

In addition to previously published information, this monograph also contains original records and observations on the associations of cockroaches that are reported here for the 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 communications.

HISTORICAL

Chopard (1938) in his book La Biologie des Orthoptéeres reviewed much of the literature on cockroaches, but of the many biotic associa- tions that exist 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 that attack cockroaches. Thompson (1951) in his Parasite Host Catalogue, which was based mainly on papers abstracted or noted in the Review of Applied Entomology, listed only 19 insect parasites of cockroaches. Eighteen of these were Hymenoptera which attack only cockroach eggs; the single dipteron listed (Sarcophaga lambens Wiedemann, supposedly parasitic on Pycnoscelus surinamensts) is not a parasite in this case, but deposits its eggs on the dead insects (see p. 229). Cameron (1955) listed as parasites and predators of the cockroach 24 species of hymenopterous egg parasites, 7 species of Ampulex which hunt nymphs and adults, 17 Protozoa, 13 nematodes, 5 bacteria, 2 mites, and a few other miscellaneous predators. In his classified list of the protozoan parasites of the Orthoptera of the world, Semans (1943) listed about 26 species from cockroaches. Lin- stow (1878, 1889) recorded 14 species of helminths from cockroaches. Van Zwaluwenburg (1928) listed 33 names of roundworms which

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 list with 13 additional species. Chitwood (1932) recognized 24 species of nematodes which are primary parasites (probably commensals) of blattids. Steinhaus (1946) gave many instances of biological rela- tionships between cockroaches and bacteria, fungi, and yeasts, but the cockroaches were not discussed as an entity and the information is scattered throughout the book.

In surveying the literature on this subject we have collected a far more extensive list 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 im- portant organisms associated with the Blattaria, we pointed out that in addition to many experimental associations cockroaches have been found to harbor, naturally, 4 strains of poliomyelitis virus, about 4O 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 carry, on occasion, 3 species of Protozoa that are pathogenic to man and 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 cock- roaches. Of the arthropods there are about 2 species of scorpions, 4 spiders, 15 mites, 4 centipedes, and go insects. Of vertebrates there are 4 species of fish, 16 amphibians, 12 reptiles, 20 birds, and 27 mam- mals. Besides these there are many records of experimental associa- tions 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

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

synonyms), I mite, 1 wasp, and 1 beetle. In addition, they mentioned as other foes of the cockroach: monkeys, hedgehogs, polecats, cats, rats, birds, chameleons, and frogs.

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 species. Under each organism 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.” The name of each cockroach is 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 (e.g., species of Ampulex, Evania, Podium) probably prey upon or parasitize cockroaches exclusively.

The validity of a host-parasite or predator-prey record is dependent upon the accuracy and knowledge 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 sub- stantially complete.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 5

some work on their hymenopterous parasites, we have questioned certain records in the literature. Other dubious records which have been perpetuated from one publication to the next, but which ap- parently were not based on fact, have also been questioned or have been clarified with the aid of specialists in particular groups.

Because the records cover a period of many years, the names of many of the organisms as well as the names of some of the cock- roaches have been changed. Although it would have been compara- tively simple to list the names as they appeared in the original ref- erences, this would 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. How- ever, 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 IJ. 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 responsibility for the names.

FUTURE WORK

After having examined thousands of references on cockroaches, we are impressed by how little is known about the biology of most 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. Un- fortunately, 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 bately progressed beyond the taxonomic stage. The total effect 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

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. IAI

among all other insects. Secretions produced by certain cockroaches (e.g., 2-hexenal by Eurycotis floridana) will ward off certain predators. The identities and biological activities of most cockroach secretions are unknown, but the use of protective chemicals against predators may be widespread among cockroaches. If so, how effective are these repellents in protecting the individual or the species? It is not known whether cockroaches are protected by apparent mimetic resemblances to other arthropods. There is no experimental proof that insect para- sites can successfully attack the eggs of cockroaches that incubate their eggs while they are being carried by the female.

It is conceivable that biological control of cockroaches might be achieved in limited areas such as man-made structures or sewers, but this possibility has not been thoroughly explored. It would be informa- tive to know what effects, if any, organisms such as bacteria, Protozoa (e.g., gregarines), intestinal nematodes, or other helminths have on cockroaches. Possibly pathogenic microorganisms can be used for biological control of cockroaches ; this approach seems to have been little explored.

Associations of colonial cockroaches (e.g., Cryptocercus spp.) may be truly familial or they may merely result from gregariousness. Newly hatched nymphs of species that carry their odthecae 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 (thig- motaxis? 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 this claim has been brought forward since that time. The apparent supersedure of one species of domiciliary cockroach by another may result from antagonism between different species, or it may result from more rapid breeding and more effective utilization of available food and space; but which? Several species of cockroaches are fre- quently found associated with certain plants (e.g., bromeliads and bananas) ; the ecological relations in these associations remain to be determined. Many of the obscure associations between cockroaches and other insects, spiders, birds, and burrowing animals have never been adequately defined. The 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. We hope that these suggestions as well as other questions that may occur to readers will stimulate further research in areas where it is obviously needed.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 7

ILLUSTRATIONS

Unless otherwise credited, the illustrations were prepared from photographs taken by the authors. Except where otherwise stated, all photographs were taken of unposed living specimens.

ih) SPEGibS, OF COCKROACHES

The cockroaches referred to in this paper are listed below. The currently accepted name for each species is given alphabetically by genus and species irrespective of its taxonomic affinities. Synonyms used by certain authors whose work we have quoted are given in brackets under the respective species; the synonymy is 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 describers.

Agis orientalis Chopard Aglaopteryx absimilis Gurney diaphana (Fabricius) [Ceratinop- tera diaphana Fabricius; Rehn and Hebard (1927) ] facies (Walker) [Aglaopteryx devia Rehn; Princis (1929). A. diaphana (Fabricius) in records from Puerto Rico only; Rehn (1932b) ; Gurney (1937) ] gemma Hebard [In Florida rec- ords = Ceratinoptera diaphana R. and H.; Hebard (1917) ] vegeta Rehn ypsilon Princis

Allacta similis (Saussure) [Phyllo- dromia obtusata Brunner; Zim- merman (1948) ]

Alluaudellina cavernicola (Shelford) [Alluaudella cavernicola Shel- ford; Chopard (1932) ]

Amazonina emarginata Princis

Anaplecta asema Hebard

azteca Saussure decipiens Saussure and Zehntner fallax Saussure hemiscotia Hebard lateralis Burmeister mexicana Saussure Aneurina tahuata Hebard viridis Hebard

Apotrogia angolensis Kirby [Acantho- gyna deplanata Chopard; Princis (1957) ]

Aptera fusca (Thunberg) [Aptera cingulata (Burmeister) ; Gurney (personal communication, 1957) ]

Apteroblatia perplexa Shelford

Archiblatia hoeventi Vollenhoven

Archimandrita marmorata (Stoll)

tessellata Rehn

Arenivaga apacha (Saussure)

bolliana (Saussure)

erratica (Rehn)

floridensis Caudell

grata Hebard

rosem (Brancsik) [Heterogamodes rosem; Bei-Bienko (1950). “Polygamia”’ roseni is undoubt- edly an erroneous citation of Polyphaga roseni, as there is no genus Polygamia (Gurney, per- sonal communication, 1957) ].

tonkawa Hebard

Aristiger histrio (Burmeister) [Plu- miger histrio (Burm.) ; Bruijn- ing (1048). Hemithyrsocera histrio Burm.; Hebard (10929) ]

Aspiduchus boriquen J. W. H. Rehn [In Puerto Rico reccrds= Aspiduchus deplanatus R. and Hee Rehn, Jo We i. Crosta)]

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS

cavernicola J. W. H. Rehn deplanatus (Saussure) Attaphila aptera Bolivar bergi Bolivar flava Gurney fungicola Wheeler schuppi Bolivar sexdentis Bolivar Atticola mortomi Bolivar Audreia bromeliadarum Caudell jamaicana Rehn and Hebard Balta godeffroyt (Shelford) patula (Walker) platysoma (Walker) teryx platysoma Hebard (1943) ] quadricaudata Hebard scripta (Shelford) torresiana Hebard verticalis Hebard

[Temnop- (Walker) ;

Bantua stigmosa (Krauss) [Dero- calymma__stigmosa Krauss; Princis (1957) ]

Blaberus atropos (Stoll) [Blabera

fusca Brunner ; Hebard (1917) ] boliviensis Princis craniifer Burmeister (pls. 1, 2)

discoidalis Serville [Blaberus cubensis Saussure; Hebard (1916) ]

giganteus (Linnaeus) (pl. 3)

Blaptica dubia (Serville) [Blaberus clarazianus Saussure; Rehn, J. W. H. (1951) ]

Blatta orientalis Linnaeus (pl. 4) [Periplaneta orientalis; He- bard (1917) ]

(Shelfordella) lateralis (Walker) [Shelfordella tartara (Saus- sure); Princis (1957). Peri- planeta tartara Saussure; Bei- Bienko (1950) }

Blattella germanica (Linnaeus) (pls. 5, A, B; 31, F) [Blatella ger- manica; Gurney (1952). Phyillo- dromia germanica; Hebard (1917). Ectobius germanicus; Gurney (personal communica- tion, 1957) ]

VOL. I4I

hwmbertiana (Saussure) [Blatta humbertiana; Phyllodromia humbertiana; Hebard (1929) ] lituricollis (Walker) (fig. 7, A) [Blattella bisignata (Brun- ner); Bei-Bienko (1950) ] schubotzi Shelford vaga Hebard (pl. 5, C, D) Buboblatta armata (Caudell) [Latin- dia armata Caudell; Hebard (1920) ] Byrsotria cabrerae Rehn and Hebard fumigata (Guérin) (pl. 6) Cahita borero Rehn nahua (Saussure) Capucinella delicatula Hebard Cariblatta antiguensis (Saussure and

Zehntner )

cuprea Hebard

delicatula (Guérin) [Blattella delicatula’ Guérin; Cariblatta

punctulata (Beativois); Rehn and Hebard (1927) J hylaca Rehn imitans Hebard insularis (Walker) landalet Rehn and Hebard lutea lutea (Saussure and Zehnt- ner) lutea minima Hebard (pl. 7, A, B) nebulicola Rehn and Hebard orestera Rehn and Hebard punctipennis Hebard reticulosa (Walker) stenophrys Rehn and Hebard Cariblattoides instigator Rehn and Hebard suave Rehn and Hebard Ceratinoptera picta Brunner Chorisoneura barbadensis Rehn and Hebard flavipennis Saussure and Zehntner formosella Rehn and Hebard parisht Rehn specilliger Hebard texensis Saussure and Zehntner [Chorisoneura plocea Rehn; Rehn and Hebard (1916) } translucida (Saussure)

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 9

Choristima sp. Choristimodes sp. Chromatonotus infuscatus (Brunner) notatus (Brunner) Compsodes schwarzt (Caudell) Comptolampra liturata (Serville) [Compsolampra liturata; Comp- tolampra is the original spelling, which is followed by Dr. K. Princis, according to Gurney (personal communication, 1959) ] Cosmozosteria lateralis (Walker) Cryptocercus punctulatus Scudder (pl. 8, A) relictus Bei-Bienko Cutilia nitida (Brunner) soror (Brunner) sp. near sedilloti (Bolivar) (pl. 9) [Determined by Dr. A. B. Gurney from photographs. ] Cyrtotria capucina (Gerstaecker) Dendroblatta sobrina Rehn Derocalymma cruralis (Stal) [Ho- malodemas cruralis (Stal); Gurney (personal communica- tion, 1957) ] lampyrina Gerstaecker porcellio Gerstaecker Deropeltis autraniana Saussure erythropeza Adelung melanophila (Walker) nigrita Saussure Diploptera punctata (Eschscholtz) (pls. 10, 36) [Diploptera dytiscoides (Serville); Princis (1950). Eleutheroda dytiscoides (Ser- ville) ; Zimmerman (1948) ] Dryadoblatia scotti (Shelford) [Ho- malopteryx scottt Shelford; Rehn (1930) ] Ectobius africanus Saussure albicinctus (Brunner) duskei Adelung lapponicus (Linnaeus) [Ectobius perspicillaris Herbst, as used by Lucas (1920) ; Blair (1934) ] lucidus Heb. nicaeensis (Brisout)

pallidus (Olivier) (pls. 7, C; 20, A) [Ectobius lividus (Fabri- cius); Ectobius livens (Tur- ton); Kevan (1952); Princis (in Roth and Willis, 1957) ]

panzeri Stephens [Ectobius eri- cetorum (Wesmael); Ramme (1923) ]

panzeri var. nigripes Stephens

semenovi Bei-Bienko

sylvester (Poda) [Ectobius syl- vestris (Poda) ; Ramme (1951) ]

tadzihicus Bei-Bienko

vittiventer (Costa) [Ectobius vit- tiventris (Costa); Ramme (1951) ]

Ellipsidion Saussure [Apolyta Brun- ner; Hebard (1043) ]

affine Hebard

australe Saussure [Ellipsidion pel-

lucidum (Brunner); Hebard (1943) ]

bicolor (Tepper)

simulans Hebard

variegatum (Fabricius) [Ellip-

sidion aurantium Saussure; He- bard (1943) ] Eptlampra abdomen-nigrum (De Geer) annandalei Shelford azteca Saussure conferta Walker conspersa Burmeister grisea (De Geer) maya Rehn mexicana Saussure mona Rehn and Hebard notabilis Walker sodalis Walker tainana Rehn and Hebard zwheeleri Rehn Spx (ait. 7 iba C) Eremoblatta subdiaphana (Scudder) Ergaula capensis (Saussure) [Dys- cologamia capensis Saussure; Dyscologamia wollastoni Kirby ; Princis (1957) ] scarabaeoides Walker [Dyscolo- gamia_ _ piolosa (Walker) ; Princis (1957). Parapolyphaga

10 SMITHSONIAN MISCELLANEOUS COLLECTIONS

erectipilis Chopard; Princis (1950). Dyscologamia chopardt Hanitsch; Bruijning (1948).

Miroblatta silphoides Chopard; Hebard (1929) ]. Escala sp. Euandroblatta palpalis Chopard Eublaberus posticus (Erichson) Eudromiella bicolorata Hebard calcarata Bei-Bienko Euphyllodromia angustata (Latreille) liturifera [Euphyllodromia de- castigmata WHebard; Princis (1959) ] Eurycotis bananae Bei-Bienko biolleyi Rehn [Eurycotis carbo- naria Biolley; Rehn (1918) ] caraibea (Bolivar) decipiens (Kirby) dimidiata (Bolivar) ferrum-equinum Rehn and Hebard floridana (Walker) (pl. 11) [Platyzosteria ingens Scudder ; Platyzosteria sabalianus Scudder (and hence, by inference, Eury- cotis sabalianus (Scudder) ; Hebard (1917) ] galeoides Rehn and Hebard improcera Rehn kevani Princis lixa Rehn manni Rehn opaca (Brunner) Euthlastoblatta abortiva (Caudell) Euthyrrhapha nigra Chopard pacifica Coquebert Geoscapheus robustus Tepper Graptoblatta notulata (Stal) [Blatta notulata Stal; Hebard (1929). Phyllodromia hieroglyphica Brunner ; Kirby (1904) ] Gromphadorhina laevigata S. and Z. portentosa (Schaum) (pl. 12, A, B) Gyna kazungulana Giglio-Tos

maculipennis (Schaum) [Gyna vetula Brunner ; Shelford (1909b) J

tristis Hanitsch

VOL. I41

Hebardina concinna (Haan) [Blatta concinna Haan; Blattina con- cinna (Haan); _ Bei-Bienko (1950) ]

Hemiblabera brunneri (Saussure)

Henicotyle antillarum (Brunner)

Heterogamodes kriigeri (Salfi)

rugosa (Schulthess) Holocompsa asteca (Saussure) cyanea (Burmeister ) fulva (Burmeister ) metallica Rehn and Hebard nitidula (Fabricius) gzapoteca Saussure Hololampra bivittata (Brullé) chavesi (Bolivar) maculata (Schreber) [Aphlebia maculata Schreber; Harz (1957) ; Gurney (personal com- munication, 1959) ] marginata (Schreber) punctata (Charpentier) [Aphlebia punctata Charpentier; Ramme (1951) ]

Hololeptoblatta sp.

Homalopteryx laminata Brunner

Hoplosphoropyga babaulti Chopard

Hormetica apolinari Hebard

laevigata Burmeister