The Biotic Associations of Cockroaches

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

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

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

FOREWORD

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

CONTENTS Page

Foreword iii

I. Introduction 1 Historical 2 Methods 4 Future work 5 Illustrations 7

II. Species of cockroaches 7

III. Ecological relationships 14 Cave habitats 16 Cavernicolous cockroaches 17 Cockroaches from burrows 23 Desert habitats 25 Desert cockroaches 27 Aquatic habitats 30 Amphibious cockroaches 31 Outdoor habitats 33 Cockroaches from outdoor habitats 35 Structural habitats 70 Land-based structures 73 Cockroaches associated with land-based structures 74 Ships 82 Cockroaches associated with ships 85 Aircraft 87 Cockroaches associated with aircraft 88

IV. Classification of the associations 91

V. Mutualism 96 Bacteroids 96 Cockroaches in which bacteroids have been found 99 Bacteria 100 Protozoa 101

VI. Viruses associated with cockroaches 103

VII. Bacteria associated with cockroaches 104

VIII. Fungi and yeasts 127 Fungi associated with cockroaches 129

IX. Higher plants associated with cockroaches 139 Damage to plants by cockroaches 162

X. Protozoa associated with cockroaches 166

XI. Helminths associated with cockroaches 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 210 Arachnida 211 Chilopoda 222 Insecta 224 Hymenoptera 234 Predators and parasites of cockroach eggs 234 Host selection by egg parasites 254 Cockroach-hunting wasps 255 Ants predaceous on cockroaches 266

XIII. Vertebrata associated with cockroaches 268 Pisces 268 Amphibia 269 Reptilia 272 Aves 276 Mammalia 283

XIV. Checklist of cockroaches and symbiotic associates 290

XV. Cockroaches as commensals 310 Hosts of commensal cockroaches 310 Checklist of commensal cockroaches with their hosts 315 Obscure associations 316

XVI. Cockroaches as predators 319 Interspecies predation 319 Intraspecies predation 322

XVII. Associations among cockroaches 324 Familial associations 325 Gregariousness 330 Intraspecies fighting 336 Interspecies compatibility 337 Interspecies antagonism 341

XVIII. Defense of cockroaches against predators 343

XIX. The biological control of cockroaches 348 Invertebrates 349 Vertebrates 353

Acknowledgments 354

References 356

Index 441

THE BIOTIC ASSOCIATIONS OF COCKROACHES

BY LOUIS M. ROTH AND EDWIN R. WILLIS

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 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.

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