The Biotic Associations of Cockroaches

Edwin R. Willis, Louis M. Roth

The Biotic Associations of Cockroaches

Sharp Ink Publishing

2024

Contact: info@sharpinkbooks.com

ISBN 978-80-282-3883-4

Table of Contents

FOREWORD

LIST OF PLATES

LIST OF FIGURES

I. INTRODUCTION

HISTORICAL

METHODS

FUTURE WORK

ILLUSTRATIONS

II. SPECIES OF COCKROACHES

III. ECOLOGICAL RELATIONSHIPS

CAVE HABITATS

CAVERNICOLOUS COCKROACHES

COCKROACHES FROM THE BURROWS OF VERTEBRATES

DESERT HABITATS

DESERT COCKROACHES

AQUATIC HABITATS

AMPHIBIOUS COCKROACHES

OUTDOOR HABITATS

COCKROACHES FROM OUTDOOR HABITATS

STRUCTURAL HABITATS

LAND-BASED STRUCTURES

COCKROACHES ASSOCIATED WITH LAND-BASED STRUCTURES

SHIPS

COCKROACHES ASSOCIATED WITH SHIPS

AIRCRAFT

COCKROACHES ASSOCIATED WITH AIRCRAFT

IV. CLASSIFICATION OF THE ASSOCIATIONS

V. MUTUALISM

BACTEROIDS

COCKROACHES IN WHICH BACTEROIDS HAVE BEEN FOUND

BACTERIA

PROTOZOA

VI. VIRUSES ASSOCIATED WITH COCKROACHES

POLIOMYELITIS VIRUSES

OTHER VIRUSES

VII. BACTERIA ASSOCIATED WITH COCKROACHES

ADDITIONAL BACTERIA WHOSE TAXONOMIC POSITION IS UNKNOWN

VIII. FUNGI AND YEASTS

FUNGI ASSOCIATED WITH COCKROACHES

INCERTAE SEDIS

IX. HIGHER PLANTS

DAMAGE TO PLANTS BY COCKROACHES

X. PROTOZOA ASSOCIATED WITH COCKROACHES

NEGATIVE FINDINGS

XI. HELMINTHS

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

RECORDS OF UNIDENTIFIED MITES

CONTROL OF MITES IN COCKROACH COLONIES

PREDATORS AND PARASITES OF COCKROACH EGGS

HOST SELECTION BY EGG PARASITES

COCKROACH-HUNTING WASPS

WASPS THAT PROVISION THEIR NESTS WITH COCKROACHES

ANTS PREDACEOUS ON COCKROACHES

XIII. VERTEBRATA

XIV. CHECKLIST OF COCKROACHES AND SYMBIOTIC ASSOCIATES

XV. COCKROACHES AS COMMENSALS

CHECKLIST OF COMMENSAL COCKROACHES WITH THEIR HOSTS

OBSCURE ASSOCIATIONS

XVI. COCKROACHES AS PREDATORS

INTERSPECIES PREDATION

INTRASPECIES PREDATION

XVII. ASSOCIATIONS AMONG COCKROACHES

FAMILIAL ASSOCIATIONS

GREGARIOUSNESS

INTRASPECIES FIGHTING

INTERSPECIES COMPATIBILITY

INTERSPECIES ANTAGONISM

XVIII. DEFENSE OF COCKROACHES AGAINST PREDATORS

XIX. THE BIOLOGICAL CONTROL OF COCKROACHES

INVERTEBRATES

VERTEBRATES

ACKNOWLEDGMENTS

PLATES

INDEX

FOREWORD

Table of Contents

People having only casual interest in insects usually express amazement 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 studied museum collections and the percent of new species found in recent acquisitions, particularly from tropical and subtropical countries, 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.

The present work is such a compilation, for it assembles what has been gleaned from approximately 1,700 sources, including correspondence 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 are especially well qualified to appraise previous work. Seldom has a compilation been done so thoroughly or a single large group of insects 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 regarded only as disagreeable pests; many species appear to be important, 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 conclusions 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 review 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 Smithsonian 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

LIST OF PLATES

Table of Contents

LIST OF FIGURES

Table of Contents

THE BIOTIC ASSOCIATIONS OF COCKROACHES

[1]

Table of Contents

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 of us collectors the life history of an insect begins in the net and ends in the bottle.

Hanitsch (1928)

I. INTRODUCTION

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Recently we brought together much of the literature linking cockroaches 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 associations 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 inferred 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 satisfactory 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

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Chopard (1938) in his book La Biologie des Orthoptères reviewed much of the literature on cockroaches, but of the many biotic associations that exist he discussed only the commensal cockroaches, gregariousness, 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 surinamensis) 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. Linstow (1878, 1889) recorded 14 species of helminths from cockroaches. Van Zwaluwenburg (1928) listed 33 names of roundworms which 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 relationships 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 important 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 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 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 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 synonyms), 1 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

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We have listed the organisms known to be associated with cockroaches 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 cockroaches are listed by the currently accepted name. Unidentified cockroaches 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[3] 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 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 apparently 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 cockroaches have been changed. Although it would have been comparatively simple to list the names as they appeared in the original references, 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. 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 responsibility for the names.

FUTURE WORK

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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. 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 relationships has barely 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 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 parasites 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 informative 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 oöthecae 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 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 frequently 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.

ILLUSTRATIONS

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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.

II. SPECIES OF COCKROACHES

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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 illustrations 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) [Ceratinoptera 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 records = Ceratinoptera diaphana R. and H.; Hebard (1917)]

vegeta Rehn

ypsilon Princis

Allacta similis (Saussure) [Phyllodromia obtusata Brunner; Zimmerman (1948)]

Alluaudellina cavernicola (Shelford) [Alluaudella cavernicola Shelford; 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 [Acanthogyna deplanata Chopard; Princis (1957)]

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

Apteroblatta perplexa Shelford

Archiblatta hoevenii Vollenhoven

Archimandrita marmorata (Stoll)

tessellata Rehn

Arenivaga apacha (Saussure)

bolliana (Saussure)

erratica (Rehn)

floridensis Caudell

grata Hebard

roseni (Brancsik) [Heterogamodes roseni; Bei-Bienko (1950). Polygamia roseni is undoubtedly an erroneous citation of Polyphaga roseni, as there is no genus Polygamia (Gurney, personal communication, 1957)].

tonkawa Hebard

Aristiger histrio (Burmeister) [Plumiger histrio (Burm.); Bruijning (1948). Hemithyrsocera histrio Burm.; Hebard (1929)]

Aspiduchus boriquen J. W. H. Rehn [In Puerto Rico records = Aspiduchus deplanatus R. and H.; Rehn, J. W. H. (1951a)]

cavernicola J. W. H. Rehn

deplanatus (Saussure)

Attaphila aptera Bolívar

bergi Bolívar

flava Gurney

fungicola Wheeler

schuppi Bolívar

sexdentis Bolívar

Atticola mortoni Bolívar

Audreia bromeliadarum Caudell

jamaicana Rehn and Hebard

Balta godeffroyi (Shelford)

patula (Walker)

platysoma (Walker) [Temnopteryx platysoma (Walker); Hebard (1943)]

quadricaudata Hebard

scripta (Shelford)

torresiana Hebard

verticalis Hebard

Bantua stigmosa (Krauss) [Derocalymma 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; Hebard (1917)]

(Shelfordella) lateralis (Walker) [Shelfordella tartara (Saussure); Princis (1957). Periplaneta tartara Saussure; Bei-Bienko (1950)]

Blattella germanica (Linnaeus) (pls. 5, A, B; 31, F) [Blatella germanica; Gurney (1952). Phyllodromia germanica; Hebard (1917). Ectobius germanicus; Gurney (personal communication, 1957)]

humbertiana (Saussure) [Blatta humbertiana; Phyllodromia humbertiana; Hebard (1929)]

lituricollis (Walker) (fig. 7, A) [Blattella bisignata (Brunner); Bei-Bienko (1950)]

schubotzi Shelford

vaga Hebard (pl. 5, C, D)

Buboblatta armata (Caudell) [Latindia 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 (Beauvois); Rehn and Hebard (1927)]

hylaea Rehn

imitans Hebard

insularis (Walker)

landalei Rehn and Hebard

lutea lutea (Saussure and Zehntner)

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

parishi Rehn

specilliger Hebard

texensis Saussure and Zehntner [Chorisoneura plocea Rehn; Rehn and Hebard (1916)]

translucida (Saussure)

Choristima sp.

Choristimodes sp.

Chromatonotus infuscatus (Brunner)

notatus (Brunner)

Compsodes schwarzi (Caudell)

Comptolampra liturata (Serville) [Compsolampra liturata; Comptolampra 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 (Bolívar) (pl. 9) [Determined by Dr. A. B. Gurney from photographs.]

Cyrtotria capucina (Gerstaecker)

Dendroblatta sobrina Rehn

Derocalymma cruralis (Stål) [Homalodemas cruralis (Stål); Gurney (personal communication, 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 (Serville); Zimmerman (1948)]

Dryadoblatta scotti (Shelford) [Homalopteryx scotti Shelford; Rehn (1930)]

Ectobius africanus Saussure

albicinctus (Brunner)

duskei Adelung

lapponicus (Linnaeus) [Ectobius perspicillaris Herbst, as used by Lucas (1920); Blair (1934)]

lucidus Hgb.

nicaeensis (Brisout)

pallidus (Olivier) (pls. 7, C; 29, A) [Ectobius lividus (Fabricius); Ectobius livens (Turton); Kevan (1952); Princis (in Roth and Willis, 1957)]

panzeri Stephens [Ectobius ericetorum (Wesmaël); Ramme (1923)]

panzeri var. nigripes Stephens

semenovi Bei-Bienko

sylvester (Poda) [Ectobius sylvestris (Poda); Ramme (1951)]

tadzihicus Bei-Bienko

vittiventer (Costa) [Ectobius vittiventris (Costa); Ramme (1951)]

Ellipsidion Saussure [Apolyta Brunner; Hebard (1943)]

affine Hebard

australe Saussure [Ellipsidion pellucidum (Brunner); Hebard (1943)]

bicolor (Tepper)

simulans Hebard

variegatum (Fabricius) [Ellipsidion aurantium Saussure; Hebard (1943)]

Epilampra 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

wheeleri Rehn

sp. (fig. 7, B, C)

Eremoblatta subdiaphana (Scudder)

Ergaula capensis (Saussure) [Dyscologamia capensis Saussure; Dyscologamia wollastoni Kirby; Princis (1957)]

scarabaeoides Walker [Dyscologamia piolosa (Walker); Princis (1957). Parapolyphaga erectipilis Chopard; Princis (1950). Dyscologamia chopardi 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 decastigmata Hebard; Princis (1959)]

Eurycotis bananae Bei-Bienko

biolleyi Rehn [Eurycotis carbonaria Biolley; Rehn (1918)]

caraibea (Bolívar)

decipiens (Kirby)

dimidiata (Bolívar)

ferrum-equinum Rehn and Hebard

floridana (Walker) (pl. II) [Platyzosteria ingens Scudder; Platyzosteria sabalianus Scudder and hence, by inference, Eurycotis 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 (Stål) [Blatta notulata Stål; 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)]

tristis Hanitsch

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

Hemiblabera brunneri (Saussure)

Henicotyle antillarum (Brunner)

Heterogamodes krügeri (Salfi)

rugosa (Schulthess)

Holocompsa azteca (Saussure)

cyanea (Burmeister)

fulva (Burmeister)

metallica Rehn and Hebard

nitidula (Fabricius)

zapoteca Saussure

Hololampra bivittata (Brullé)

chavesi (Bolívar)

maculata (Schreber) [Aphlebia maculata Schreber; Harz (1957); Gurney (personal communication, 1959)]

marginata (Schreber)

punctata (Charpentier) [Aphlebia punctata Charpentier; Ramme (1951)]

Hololeptoblatta sp.

Homalopteryx laminata Brunner

Hoplosphoropyga babaulti Chopard

Hormetica apolinari Hebard

laevigata Burmeister

ventralis Burmeister

Ignabolivaria bilobata Chopard

Ischnoptera deropeltiformis (Brunner) (pl. 12A) [Temnopteryx deropeltiformis Brunner; Hebard (1917)]

panamae Hebard

podoces Rehn and Hebard

rufa occidentalis Saussure

rufa rufa (De Geer)

schenklingi Karney

Karnyia discoidalis (Brunner)

Kuchinga hemerobina (Gerstaecker) [Phyllodromia hemerobina Gerstaecker; Rehn (1932)]

remota Hebard

Lamproblatta albipalpus Hebard

meridionalis (Brunner)

Latiblattella chichimeca (Saussure and Zehntner) [Blattella chichimeca S. and Z.; Hebard (1932)]

lucifrons Hebard

rehni Hebard

vitrea (Brunner)

zapoteca (Saussure)

Leucophaea maderae (Fabricius) (pl. 13) [Rhyparobia maderae; Hebard (1917). Panchlora maderae; Kirby (1904). Very probably Blaberus maderae is a careless reference to this species; Gurney (personal communication, 1957)]

Leurolestes pallidus (Brunner)

Litopeltis biolleyi (Saussure)

bispinosa (Saussure) [Audreia marginata Caudell; Hebard (1920)]

deianira Rehn

musarum Rehn

Lobolampra subaptera Rambur

Loboptera decipiens (Germar)

thaxteri Hebard

Lobopterella dimidiatipes (Bolívar) [Loboptera dimidiatipes (Bolívar); Princis (1957a). Loboptera sakalava (Saussure); Hebard (1933a). Loboptera extranea Perkins; Hebard (1922). Princis (1957a) in erecting Lobopterella pointed out that only the nontypical variety of sakalava is identical with dimidiatipes.]

Lophoblatta arawaka Hebard

Macropanesthia rhinocerus Saussure

Mareta acutiventris Chopard

Maretina uahuka Hebard

Megaloblatta blaberoides (Walker) [Megaloblatta rufipes Dohrn; Hebard (1920)]

Megamareta verticalis Hebard

Melanosilpha capensis Saussure and Zehntner

Methana canae Pope

curvigera (Walker)

marginalis (Saussure)

Moluchia (?) dahli Princis

Monastria biguttata (Thunberg)

Muzoa madida Rehn

Myrmeblattina longipes (Chopard)

Myrmecoblatta rehni Mann

wheeleri Hebard

Namablatta bitaeniata (Stål)

Nauclidas nigra (Brunner) [Poroblatta nigra Brunner; Rehn (1930)]

Nauphoeta cinerea (Olivier) (pl. 14) [Nauphoeta bivittata Burmeister; Zimmerman (1948)]

flexivitta (Walker) [Nauphoeta brazzae (Bolívar); Rehn (1937)]

punctipennis Chopard

Nelipophygus ramsdeni Rehn and Hebard

Neoblattella brunneriana (Saussure) [Blattella brunneriana; Gurney (personal communication, 1959)]

carcinus Rehn and Hebard

celeripes Rehn and Hebard

detersa (Walker)

dryas Rehn and Hebard

eurydice Rehn and Hebard

fratercula Hebard

fraterna (Saussure and Zehntner)

grossbecki Rehn and Hebard

laodamia Rehn and Hebard

nahua (Saussure) [Blattella nahua Saussure and Zehntner of Caudell (1914); Hebard (1920)]

proserpina Rehn and Hebard

semota Rehn and Hebard

tridens Rehn and Hebard

vatia Rehn and Hebard

Neostylopyga rhombifolia (Stoll) (pl. 15) [Dorylaea rhombifolia; Rehn (personal communication, 1956)]

Nesomylacris cubensis Rehn and Hebard

relica Rehn and Hebard

Nocticola bolivari Chopard

caeca Bolívar

decaryi Chopard

simoni Bolívar

sinensis Silvestri

termitophila Silvestri

Nothoblatta wasmanni (Bolívar)

Notolampra antillarum Shelford

Nyctibora azteca Saussure and Zehntner

brunnea (Thunberg)

laevigata (Beauvois)

lutzi Rehn and Hebard

mexicana Saussure

noctivaga Rehn

obscura Saussure

sericea Burmeister

stygia Walker

tomentosa Serville [Nyctibora latipennis Burmeister; Hebard (1917, p. 263)]

Oniscosoma granicollis (Saussure)

Opisthoplatia maculata Shiraki

orientalis (Burmeister)

Oulopteryx meliponarum Hebard

Oxyhaloa buprestoides (Saussure)

deusta (Thunberg)

Panchlora antillarum Saussure

exoleta Burmeister

fraterna Saussure and Zehntner

nivea (Linnaeus) (pl. 16) [Panchlora cubensis Saussure; Gurney (1955). Pycnosceloides aporus Hebard; Hebard (1921c)]

peruana Saussure

sagax Rehn and Hebard

virescens (Thunberg)

Panesthia angustipennis (Illiger) [Panesthia javanica Serville; Hebard (1929)]

australis Brunner (pl. 8, B)

laevicollis Saussure

lobipennis Brunner

spadica (Shiraki)

Parahormetica bilobata (Saussure)

Parcoblatta americana (Scudder)

bolliana (Saussure and Zehntner) [Kakerlac schaefferi Rehn; Hebard (1917)]

caudelli Hebard [♀♀ of Ischnoptera insolita R. and H.; Ischnoptera uhleriana fulvescens S. and Z. (in part); Hebard (1917)]

desertae (Rehn and Hebard) [♂♂ of Ischnoptera insolita R. and H.; Hebard (1917)]

divisa (Saussure and Zehntner) [Ischnoptera divisa S. and Z.; Hebard (1917)]

fulvescens (Saussure and Zehntner) [Ischnoptera uhleriana fulvescens S. and Z. (in part); Hebard (1917)]

lata (Brunner) [Ischnoptera couloniana R. and H. (not Saussure); Ischnoptera major R. and H. (not S. and Z.); Hebard (1917)]

notha Rehn and Hebard

pensylvanica (De Geer) (pl. 17, A) [Ischnoptera pennsylvanica Saussure; Hebard (1917)]

uhleriana (Saussure) (pl. 18) [Ischnoptera uhleriana Saussure; Hebard (1917)]

virginica (Brunner) (pls. 17, B; 27, A; 33, C; fig. 6) [Ischnoptera borealis Brunner; Hebard (1917)]

zebra Hebard

Pelmatosilpha coriacea Rehn

kevani Princis

marginalis Brunner

purpurascens (Kirby)

rotundata Scudder

vagabunda Princis

Periplaneta americana (Linnaeus) (pls. 19, 35) [Stylopyga americana; Blatta americana L.; Hebard (1917)]

australasiae (Fabricius) (pls. 20, 32)

brunnea Burmeister (pl. 21)

cavernicola Chopard

fuliginosa (Serville) (pl. 22)

ignota Shaw

lata (Herbst)

Perisphaerus armadillo Serville

glomeriformis (Lucas)

Phaetalia pallida (Brunner)

Phidon (?) dubius Princis

Phlebonotus pallens (Serville)

Pholadoblatta inusitata (Rehn)

Phorticolea boliviae Caudell

testacea Bolívar

Phyllodromia treitliana Werner

Phyllodromica brevipennis (Fischer)

graeca (Brunner)

irinae (Bei-Bienko)

maculata (Schreber)

megerlei (Fieber)

polita (Krauss)

pygmaea (Bei-Bienko)

tartara (Saussure)

tartara nigrescens Bei-Bienko

Platyzosteria analis (Saussure) [Polyzosteria analis Saussure; Kirby (1904)]

armata Tepper

bifida (Saussure)

castanea (Brunner)

novae seelandiae (Brunner) (pl. 23) [Periplaneta fortipes Walker; Shelford (1912); Platyzosteria novae-zealandiae]

scabra (Brunner)

Plectoptera dorsalis (Burmeister)

infulata (Rehn and Hebard)

lacerna Rehn and Hebard

perscita Rehn and Hebard

poeyi (Saussure) [Plectoptera floridana Hebard; Rehn and Hebard (1927)]

porcellana (Saussure)

pygmaea (Saussure)

rhabdota (Rehn and Hebard)

vermiculata Rehn and Hebard

Polyphaga aegyptiaca (Linnaeus) [Blatta aegyptiaca L.; Bei-Bienko (1950). Heterogamia aegyptiaca (L.); Gurney (personal communication, 1957). Polygamia aegyptiaca; according to Gurney (p. c.), there is no genus Polygamia and almost surely the reference is to Polyphaga aegyptiaca.]

indica Walker [Polyphaga pellucida (Redtenbacher); Princis (1957)]

saussurei (Dohrn)

Polyzosteria limbata Burmeister

melanaria (Erichson)

Pseudoderopeltis aethiopica (Saussure) [Blatta aethiopica Saussure; Gurney (personal communication, 1957)]

Pseudomops cincta (Burmeister) [Thyrsocera cincta Scudder; Hebard (1917)]

laticornis Perty

septentrionalis Hebard

Pseudophoraspis nebulosa (Burmeister)

Pycnoscelus niger (Brunner)

striatus (Kirby) [Leucophaea striata Kirby; Gurney (personal communication, 1957)]

surinamensis (Linnaeus) (pl. 24) [Leucophaea surinamensis (L.); Hebard (1917). Blatta melanocephala Stoll; Kirby (1904)]

Rhicnoda natatrix Shelford

Rhytidometopum dissimile Princis

Riatia fulgida (Saussure) [Lissoblatta fulgida (Saussure); Gurney (personal communication, 1959)]

orientis Hebard

Robshelfordia circumducta (Walker) [Escala circumducta (Walker); Gurney (personal communication, 1957)]

longiuscula (Walker) [Escala longiuscula (Walker); Gurney (personal communication, 1957)]

Salganea morio (Burmeister)

Sibylloblatta panesthoides (Walker)

Simblerastes jamaicanus Rehn and Hebard

Spelaeoblatta gestroi Bolívar

Sphecophila polybiarum Shelford

ravana Fernando

termitium Shelford

Steleopyga (?) sinensis Walker [Dr. Gurney (personal communication, 1957) could not find a reference to this species. Walker described species named sinensis in three different genera of cockroaches, and it is uncertain which one this combination represents.]

Stictolampra buqueti concinula (Walker)

Styphon bakeri Rehn

Supella hottentotta (Saussure)

supellectilium (Serville) (pls. 25; 30, B-E; 31, A-E) [Phyllodromia supellectilium (Serv.); Bei-Bienko (1950)]

Symploce breviramis (Hanitsch)

cavernicola (Shelford) [Ischnoptera cavernicola (Shelford); Phyllodromia nigrocincta Chopard; Hebard (1929)]

curta Hanitsch

flagellata Hebard

hospes (Perkins) [Symploce lita Hebard; Hebard (1922)]

jamaicana (Rehn)

kevani Chopard

parenthesis (Gerstaecker) [Phyllodromia parenthesis Gerstaecker; Rehn (1932)]

remyi (Hanitsch) [Ischnoptera remyi Hanitsch; Chopard (1938)]

ruficollis (Fabricius) [Symploce bilabiata Rehn and Hebard; Princis (1949a)]

Tartaroblatta karatavica Bei-Bienko

Temnopteryx obliquetruncata Chopard

phalerata (Saussure)

Theganopteryx straminea Chopard

Therea nuptialis (Gerstaecker) [Corydia nuptialis Gerstaecker; Princis (1950)]

Tivia australica Princis

brunnea (Chopard)

fulva (Burmeister)

macracantha Chopard

obscura (Chopard)

Typhloblatta caeca (Chopard) [Spelaeoblatta caeca Chopard; Chopard (1924b)]

Typhloblattodes madecassus Chopard

Xestoblatta festae (Griffini)

immaculata Hebard

III. ECOLOGICAL RELATIONSHIPS

Table of Contents

The ecology of extinct cockroaches is necessarily a highly speculative subject. From the coexistence of fossil cockroaches and fossil plants in the 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 pusillus, which were attached to the leaves of Cordaites. Yet the proximity of fossil insects and plants in the same geological formation is hardly proof of a similar association during life. In fact, Sellards (1903), Bolton (1921), and Laurentiaux (1951) have all pointed out that the cockroach remains, particularly the more resistant wings, may have been washed into streams by heavy rains and transported with drifting plant material to places where permanent deposits were accumulating.

Some species of fossil cockroaches have long, well-developed ovipositors, very unlike present-day cockroaches whose ovipositors are small and nonprotruding. Brongniart (1889) and Zalesskii (1939, 1953) have suggested that certain Permian and Carboniferous