HE 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 reflects the extensive efforts undertaken to find chinks in the armor of problematic cockroaches, and the fact that certain species are ideal for physiological and behavioral investigations 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 americana 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, surprisingly, light up.

The publication most responsible for alerting the scientific community to the diversity exhibited by the 99+% 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 extraordinary 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 relies heavily on the information contained in its progenitor. Our update, however, narrows 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 behavioral ecology and evolution. Even we were surprised at some recent findings, such as the estimate by Basset (2001) that cockroaches constitute approximately 24% of the arthropod biomass in tropical tree canopies worldwide, and hints from various studies suggesting that cockroaches may ecologically replace termites in some habitats (Chapter 10). We address previously unexplored aspects of their biology, such as the relationship with microbes that lies at the heart of their image as anathema to civilized households (Chapter 5). As our writing progressed, some chapters followed un-predicted paths, particularly evident in the one on mating strategies (Chapter 6). We became fascinated with drawings of male and female genitalia that are buried in the taxonomic literature and that suggest ongoing, internally waged battles to determine paternity of offspring. It is the accessibility of this kind of information that can have the most impact on students searching for a dissertation topic, and we cover it in detail at the expense of addressing more familiar aspects of cockroach mating biology. We planned the book so that each chapter can be mined for new ideas, new perspectives, and new directions for future work.

An interesting development since Roth and Willis (1960) was published is that the definition of a cockroach is somewhat less straightforward than it used to be. Cockroaches are popularly considered one of the oldest terrestrial arthropod groups, because insects with a body plan closely resembling that of extant Blattaria dominated the fossil record of the Carboniferous, "The Age of Cockroaches." The lineage that produced extant cockroaches, however, radiated sometime during the early to mid-Mesozoic (e.g., Labandeira, 1994; Vrsansky, 1997; Grim-aldi and Engel, 2005). Although the Carboniferous fossils probably include the group that gave rise to modern Blattaria, they also include basal forms of other taxa. Technically, then, they cannot be considered cockroaches, and the Paleozoic group has been dubbed "roachoids" (Grimaldi and Engel, 2005), among other things. Recent studies of extant species are also blurring our interpretation of what may be considered a cockroach. Best evidence currently supports the view that termites are nested within the cockroaches as a subgroup closely related to the cockroach genus Cryptocercus. We devote Chapter 9 to developing the argument that termites evolved as eu-social, juvenilized cockroaches.

Roth (2003c) recognized six families that place most cockroach species: Polyphagidae, Cryptocercidae, Nocti-



CTryonicus Lauraesilpha

— Lamproblatta j— Polyphaga

— Cryptocercus

— Mastotermitidae j— Termopsidae

— Kalotermitidae Anaplecta Nahublattella Supella

Euphyllodromia j— Parcoblatta T— Nyctibora Nauphoeta Phoetalia Blaberus Blaptica

Blattidae: Archiblattinae Blattidae: Polyzosteriinae Blattidae: Blattinae Blattidae: Blattinae Blattidae: Tryonicinae Blattidae: Tryonicinae Blattidae: Lamproblattinae Polyphagidae: Polyphaginae Polyphagidae: Polyphaginae Cryptocercidae: Cryptocercinae


Blattellidae: Blattellidae: Blattellidae: Blattellidae: Blattellidae: Blattellidae: Blaberidae: Blaberidae: Blaberidae: Blaberidae:











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 hypotheses of higher-level relationships. Phylogenetic trees of Vrsansky 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.

colidae, Blattidae, Blattellidae, and Blaberidae; the majority of cockroaches fall into the latter three families. His paper was used as the basis for assigning the cockroach genera discussed in this book to superfamily, family, and subfamily, summarized in the Appendix. Despite recent morphological and molecular analyses, the relationships among cockroach lineages are still very much debated at many levels; Roth (2003c) summarizes current arguments. For general orientation, we offer a recent, strongly supported hypothesis by Klass and Meier (2006) (see fig. P.1). In it, there is a basal dichotomy between the family Blattidae and the remaining cockroaches, with the rest falling into two clades. The first consists of Cryptocerci-dae and the termites as sister groups, with these closely related to the Polyphagidae and to Lamproblatta. The other clade consists of the Blattellidae and Blaberidae, with the Anaplectinae as most basal and Blattellidae strongly pa-raphyletic with respect to Blaberidae. One consequence of the phylogenetic uncertainties that exist at so many taxonomic levels of the Blattaria is that mapping character states onto phylogenetic trees is in most cases premature. An analysis of the evolution of some wing characters in Panesthiinae (Blaberidae) based on the work of Mae-kawa et al. (2003) is offered in Chapter 2, a comparative phylogeny of cockroaches and their fat body endosym-bionts (Lo et al., 2003a) is included in Chapter 5, and key symbiotic relationships are mapped onto a phylogenetic tree of major Dictyopteran groups in Chapter 9.

Since the inception of this book nearly 15 years ago, the world of entomology has lost two of its giants, William J. Bell and Louis M. Roth. It was an enormous responsibility to finish the work they initiated, and I missed their wise counsel in bringing it to completion. If just a fraction of their extraordinary knowledge of and affection for cockroaches shines through in the pages that follow, I will consider the book a success. This volume contains unpublished data, observations, and personal communications of both men, information that otherwise would have been lost to the scientific community at large. Bill Bell's observations of aquatic cockroaches are in Chapter 2, and his unpublished research on the diets of tropical species is summarized in Chapter 4. Lou Roth was the acknowledged world expert on all things cockroach, and was the "go to" man for anyone who needed a specimen identified or with a good cockroach story to share. The content of his conversations and personal observations color the text throughout the book. Bill's and Lou's notes and papers were kindly loaned to me by their colleagues at the University of Kansas and Harvard University, respectively. I found it revealing that on Lou's copy of a paper by Asahina (1960) entitled "Japanese cockroaches as household pest," the s in the last word was rather emphatically scratched out.

A large number of colleagues were exceedingly generous in offering their time and resources to this project, and without their help this volume never would have seen the light of day. For advice, information, encouragement, references, photographs, illustrations, permission to use material, or for supplying reprints or other written matter I am glad to thank Gary Alpert, Dave Alexander, David Alsop, L.N.Anisyutkin, Jimena Aracena, Kathie Atkinson, Calder Bell, David Bignell, Christian Bordereau, Michel Boulard, Michael Breed, John Breznak, Remy Brossut, Valerie Brown, Kevin Carpenter, Randy Cohen, Stefan Cover, J.A. Danoff-Burg, Mark Deyrup, R.M. Dobson, C. Durden, Betty Faber, Robert Full, César Gemeno, Fabian Haas, Johannes Hackstein, Bernard Hartman, Scott Hawkes,W.F. Humphreys, T. Itioka, Ursula Jander, Devon Jindrich, Susan Jones, Patrick Keeling, Larry Kipp, Phil Koehler, D. Kovach, Conrad Labandeira, Daniel Lebrun, S. Le Maitre, Tadao Matsumoto, Betty McMahan, John Moser, I. Nagamitsu, M.J. O'Donnell, George Poinar, Colette Rivault, Edna Roth, Douglas Rugg, Luciano Sacchi, Coby Schal, Doug Tallamy, Mike Turtellot, L. Vidlicka, Robin Wootton, T. Yumoto, and Oliver Zompro.

I am particularly indebted to Horst Bohn, Donald Cochran, Jo Darlington, Thomas Eisner, Klaus Klass, Donald and June Mullins, Piotr Naskrecki, David Rentz, Harley Rose, and Ed Ross for their generosity in supplying multiple illustrations, and to George Byers, Jo Darlington, Lew Deitz, Jim Hunt, Klaus Klass, Nathan Lo, Kiyoto Maekawa, Donald Mullins, Patrick Rand, David Rentz, and Barbara Stay for reviewing sections or chapters of the book and for spirited and productive discussions. Anne Roth and the Interlibrary Loan and Document Delivery Services at NCSU were instrumental in obtaining obscure references. I thank Vince Burke and the Johns Hopkins University Press for their patience during the overlong gestation period of this book. I am sure that there are a great number of people whose kindness and contributions eased the workload on Bill Bell and Lou Roth during the early stages of this endeavor, and I thank you, whoever you are.

Christine A. Nalepa

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