Cockroaches : Ecology, Behavior, and Natural History / William J
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Cockroaches This page intentionally left blank Cockroaches ECOLOGY, BEHAVIOR, AND NATURAL HISTORY William J. Bell Louis M. Roth Christine A. Nalepa Foreword by Edward O. Wilson The Johns Hopkins University Press Baltimore © 2007 The Johns Hopkins University Press All rights reserved. Published 2007 Printed in the United States of America on acid-free paper 9 8 7 6 5 4 3 2 1 The Johns Hopkins University Press 2715 North Charles Street Baltimore, Maryland 21218-4363 www.press.jhu.edu Library of Congress Cataloging-in-Publication Data Bell, William J. Cockroaches : ecology, behavior, and natural history / William J. Bell, Louis M. Roth, Christine A. Nalepa ; foreword by Edward O. Wilson. p. cm. Includes bibliographical references and index. ISBN-13: 978-0-8018-8616-4 (hardcover : alk. paper) ISBN-10: 0-8018-8616-3 (hardcover : alk. paper) 1. Cockroaches. I. Roth, Louis M. (Louis Marcus), 1918– II. Nalepa, Christine A. III. Title. QL505.5.B43 2007 595.7Ј28—dc22 2006033232 A catalog record for this book is available from the British Library. To the families, friends, and colleagues of William J. Bell and Louis M. Roth This page intentionally left blank Contents Foreword, by Edward O. Wilson ix Preface xi ONE Shape, Color, and Size 1 TWO Locomotion: Ground, Water, and Air 17 THREE Habitats 37 FOUR Diets and Foraging 61 FIVE Microbes: The Unseen Influence 76 SIX Mating Strategies 89 SEVEN Reproduction 116 EIGHT Social Behavior 131 NINE Termites as Social Cockroaches 150 TEN Ecological Impact 165 Appendix 177 Glossary 179 References 183 Index 225 This page intentionally left blank Foreword Let the lowly cockroach crawl up, or, better, fly up, to its rightful place in human esteem! Most of us, even the entomologists in whose ranks I belong, have a stereotype of revolt- ing little creatures that scatter from leftover food when you turn on the kitchen light and instantly disappear into inaccessible crevices. These particular cockroaches are a problem, and the only solution is blatticide, with spray, poison, or trap. I developed a better understanding when I came to realize that the house pests and feces-consuming sewer dwellers are only the least pleasant tip of a great blattarian bio- diversity. My aesthetic appreciation of these insects began during one of my first excur- sions to the Suriname rainforest, where I encountered a delicate cockroach perched on the leaf of a shrub in the sunshine, gazing at me with large uncockroach-like eyes. When I came too close, it fluttered away on gaily colored wings like a butterfly. My general blattarian education was advanced when I traveled with Lou Roth to Costa Rica in 1959, and further over the decades we shared at Harvard’s Museum of Compar- ative Zoology, as he worked as a taxonomist through the great evolutionary radiation of the blattarian world fauna. This volume lays out, in detail suitable for specialists but also in language easily un- derstood by naturalists, the amazing panorama of adaptations achieved by one impor- tant group of insects during hundreds of millions of years of evolution. Abundant in most terrestrial habitats of the world, cockroaches are among the principal detritivores (their role, for example, in our kitchens), but some species are plant eaters as well. The species vary enormously in size, anatomy, and behavior. They range in habitat preference from old-growth forests to deserts to caves. They form intricate symbioses with micro- organisms. The full processes of their ecology, physiology, and other aspects of their bi- ology have only begun to be explored. This book will provide a valuable framework for the research to come. Edward O. Wilson This page intentionally left blank Preface The study of roaches may lack the aesthetic values of bird-watching and the glamour of space flight, but nonetheless it would seem to be one of the more worthwhile of human activities. —H.E. Evans, Life on a Little Known Planet Most available literature on cockroaches deals with domestic pests and the half dozen or so other species that are easily and commonly kept in laboratories and museums. It re- flects the extensive efforts undertaken to find chinks in the armor of problematic cock- roaches, and the fact that certain species are ideal for physiological and behavioral in- vestigations under controlled conditions. These studies have been summarized in some excellent books, including those by Guthrie and Tindall (1968), Cornwell (1968), Huber et al. (1990), Bell and Adiyodi (1982a), and Rust et al. (1995). The last two were devoted to single species, the American and the German cockroaches, respectively. As a result of this emphasis on Blattaria amenable to culture, cockroaches are often discussed as though they are a homogeneous grouping, typified by species such as Periplaneta amer- icana and Blattella germanica. In reality the taxon is amazingly diverse. Cockroaches can resemble, among other things, beetles, wasps, flies, pillbugs, and limpets. Some are hairy, several snorkel, some chirp, many are devoted parents, and males of several species, sur- prisingly, light up. The publication most responsible for alerting the scientific community to the diver- sity exhibited by the 99ϩ% of cockroaches that have never set foot in a kitchen is The Bi- otic Associations of Cockroaches, by Louis M. Roth and Edwin R.Willis, published in 1960. Its encyclopedic treatment of cockroach ecology and natural history was an extraordi- nary achievement and is still, hands down, the best primary reference on the group in print. Now, nearly 50 years later, we feel that the subject matter is ripe for revisitation. The present volume was conceived as a grandchild of the Roth and Willis book, and re- lies heavily on the information contained in its progenitor. Our update, however, nar- rows the focus, includes recent studies, and when possible and appropriate, frames the information within an ecological and evolutionary context. This book is intended primarily as a guided tour of non-domestic cockroach species, and we hope that it is an eye-opening experience for students and researchers in behav- ioral ecology and evolution. Even we were surprised at some recent findings, such as the estimate by Basset (2001) that cockroaches constitute ap- is somewhat less straightforward than it used to be. Cock- proximately 24% of the arthropod biomass in tropical roaches are popularly considered one of the oldest terres- tree canopies worldwide, and hints from various studies trial arthropod groups, because insects with a body plan suggesting that cockroaches may ecologically replace ter- closely resembling that of extant Blattaria dominated the mites in some habitats (Chapter 10). We address previ- fossil record of the Carboniferous, “The Age of Cock- ously unexplored aspects of their biology, such as the re- roaches.” The lineage that produced extant cockroaches, lationship with microbes that lies at the heart of their however, radiated sometime during the early to mid- image as anathema to civilized households (Chapter 5). Mesozoic (e.g., Labandeira, 1994; Vrsˇansky´, 1997; Grim- As our writing progressed, some chapters followed un- aldi and Engel, 2005). Although the Carboniferous fossils predicted paths, particularly evident in the one on mat- probably include the group that gave rise to modern ing strategies (Chapter 6). We became fascinated with Blattaria, they also include basal forms of other taxa. drawings of male and female genitalia that are buried in Technically, then, they cannot be considered cockroaches, the taxonomic literature and that suggest ongoing, inter- and the Paleozoic group has been dubbed “roachoids” nally waged battles to determine paternity of offspring. It (Grimaldi and Engel, 2005), among other things. Recent is the accessibility of this kind of information that can studies of extant species are also blurring our interpreta- have the most impact on students searching for a disser- tion of what may be considered a cockroach. Best evi- tation topic, and we cover it in detail at the expense of ad- dence currently supports the view that termites are nested dressing more familiar aspects of cockroach mating biol- within the cockroaches as a subgroup closely related to ogy. We planned the book so that each chapter can be the cockroach genus Cryptocercus. We devote Chapter 9 mined for new ideas, new perspectives, and new direc- to developing the argument that termites evolved as eu- tions for future work. social, juvenilized cockroaches. An interesting development since Roth and Willis Roth (2003c) recognized six families that place most (1960) was published is that the definition of a cockroach cockroach species: Polyphagidae, Cryptocercidae, Nocti- Fig. P.1 A phylogeny of cockroaches based on cladistic analysis of 175 morphological and life history characters; after Klass and Meier (2006), courtesy of Klaus Klass. Assignation of genera to subfamilies is after Roth (2003c) and differs somewhat from that of K & M, who place Archi- blatta in the Blattinae and Phoetalia in the Epilamprinae. Pseudophyllodromiinae used here is Plecopterinae in K & M. Based on their results, K & M suggest that Lamproblattinae and Try- onicinae be elevated to family-level status. Mukha et al. (2002, Fig. 2) summarize additional hy- potheses of higher-level relationships. Phylogenetic trees of Vrs˘ansky´ et al. (2002, Fig. 364) and Grimaldi and Engel (2005, Fig. 7.60) include fossil groups. Lo et al. (2000), Klass (2001, 2003), and Roth (2003c) discuss major issues. xii PREFACE colidae, Blattidae, Blattellidae, and Blaberidae; the major- at the University of Kansas and Harvard University, re- ity of cockroaches fall into the latter three families. His spectively. I found it revealing that on Lou’s copy of a pa- paper was used as the basis for assigning the cockroach per by Asahina (1960) entitled “Japanese cockroaches as genera discussed in this book to superfamily, family, and household pest,” the s in the last word was rather em- subfamily, summarized in the Appendix.