TRUE BUGS OFTHE WORLD (Hemiptera: Heteroptera) CLASSIFICATION AND NATURAL HISTORY Randall T. Schuh Department of Entomology American Museum of Natural History, New York James A. Slater Department of Ecology and Evolutionary Biology (Professor Emeritus) University of Connecticut, Storrs Comstock Publishing Associates a division of Cornell University Press \ ithaca and London Ped ro W. Wygodzinsky, 1916-1987 Rene H. Cobben, 1925-1987 Occasionally, remarkable individuals emerge in a given field of study, leaving for posterity writings transcendent in their scope and organization. We dedi¬ cate this volume to two such individuals, our late colleagues Rene Cobben and Pedro Wygodzinsky. Although they died prematurely with much impor¬ tant work unfinished, both left a legacy of knowledge on the heteropterans of their special interest as well as a profound influence on our conception of the broader aspects of heteropteran morphology and classification. As authors of the present volume, we have been inspired by the activities and influenced by the opinions of these two great heteropterists, a fact reflected by our frequent citation of their works. We regret only that they could not be with us now to influence the next generation with their unbounded enthusiasm for the study of bugs. CONTENTS Preface xi Chapter 1. A History of the Study of the Heteroptera 1 Chapter 2. Major Workers on the Heteroptera 6 Chapter 3. Sources of Information 14 Chapter 4. Collecting, Preserving, and Preparing Heteroptera 17 Chapter 5. Habitats and Feeding Types 20 Chapter 6. Wing Polymorphism 23 Chapter 7. Mimicry and Protective Coloration and Shape 27 Chapter 8, Heteroptera of Economic Importance 32 Chapter 9. Historical Biogeography 38 Chapter 10. General Adult Morphology and Key to Infraorders of Heteroptera 41 HETEROPTERA Chapter 11. Enicocephalomorpha by Pavel Stys 67 Chapter 12. Aenictopecheidae by Pavel Stys 68 Chapter 13. Enicocephalidae by Pavel Stys 70 EUHETEROPTERA Chapter 14. Dipsocoromorpha by Pavel Stys 74 Chapter 15. Ceratocombidae by Pavel Stys 75 Chapter 16. Dipsocoridae by Pavel Stys 78 Chapter 17. Hypsipterygidae by Pavel Stys 80 Chapter 18. Schizopteridae by Pavel Stys 80 Chapter 19. Stemmocryptidae by Pavel Stys 82 NEOHETEROPTERA Chapter 20. Gerromorpha 84 Mesoveloidea Chapter 21. Mesoveliidae 88 Hebroidea Chapter 22. Hebridae 90 Contents vii Chaoter 23. Paraphrynoveliidae 92 Chapter 24. Macroveliidae 93 Hydrometroidea Chapter 25. Hydrometridae 95 Gerroidea Chapter 26. Hermatobatidae 97 Chapter 27. Veliidae 98 Chapter 28. Gerridae 102 PANHETEROPTERA Chapter 29. Nepomorpha 107 Nepoidea Chapter 30. Belostomatidae 111 Chapter 31. Nepidae 114 Ochteroidea Chapter 32. Gelastocoridae 116 Chapter 33. Ochteridae 118 Corixoidea Chapter 34. Corixidae 119 Naucoroidea Chapter 35. Potamocoridae 122 Chapter 36. Kaucoridae 124 Chapter 37. rsphelocheiridae 126 Notonectoidea Chapter 38. Notonectidae 127 Chapter 39. Pleidae 129 Chapter 40. Helotrephidae 130 Chapter 41. Leptopodomorpha 134 Saldoidea Chapter 42. Aepophilidae 136 Chapter 43. Saldidae 137 Leptopodoidea Chapter 44. Omaniidae 141 Chapter 45. Leptopodidae 142 Chapter 46. Cimicomorpha 146 Reduvioidea Chapter 47. Pachynomidae 148 Chapter 48. Reduviidae 150 Velocipedoidea Chapter 49. Velocipedidae 161 Microphysoidea Chapter 50. Microphysidae 161 viii Contents Joppeicoidea Chapter 51. Joppeicidae 164 Miroidea Chapter 52. Thaumastocoridae 165 Chapter 53. Miridae 169 Chapter 54. Tingidae 180 Naboidea Chapter 55. Medocostidae 184 Chapter 56. Nabidae 186 Cimicoidea Chapter 57. Lasiochilidae 190 Chapter 58. Plokiophilidae 190 Chapter 59. Lyctocoridae 194 Chapter 60. Anthocoridae 195 Chapter 61. Cimicidae 199 Chapter 62. Polyctenidae 202 Chapter 63. Pentatomomorpha 205 Aradoidea Chapter 64. Aradidae 208 Chapter 65. Termitaphididae 214 Pentatomoidea Chapter 66. Acanthosomatidae 215 Chapter 67. Aphylidae 218 Chapter 68. Canopidae 219 Chapter 69. Cydnidae 220 Chapter 70. Dinidoridae 225 Chapter 71. Lestoniidae 227 Chapter 72. Megarididae 228 Chapter 73. Pentatomidae 229 Chapter 74. Phloeidae 234 Chapter 75. Plataspidae 236 Chapter 76. Scutelleridae 238 Chapter 77. Tessaratomidae 241 Chapter 78. Thaumastellidae 243 Chapter 79. Urostylidae 245 Lygaeoidea Chapter 80. Berytidae 246 Chapter 81. Colobathristidae 249 Chapter 82. Idiostolidae 251 Chapter 83. Lygaeidae 251 Chapter 84. Malcidae * 264 Chapter 85. Piesmatidae 266 Pyrrhocoroidea Chapter 86. Largidae 268 Chapter 87. Pyrrhocoridae 270 Contents IX Coreoidea Chapter 88. Alydidae 271 Chapter 89. Coreidae 274 Chapter 90. Hyocephalidae 279 Chapter 91. Rhopalidae 281 Chapter 92. Stenocephalidae 283 Literature Cited 285 Giossary 317 Index 323 About the Authors 337 X Contents PREFACE The Heteroptera, or true bugs, are the largest and most diverse group of insects with incomplete metamorphosis. They are generally treated as a suborder of the Hemiptera, and a majority of the 75 families occur on all continents (except Antarctica) and on many islands. Their great age and apparent adaptability have resulted, over evolutionary time, in extreme structural and biological diversity. The earliest volume to deal solely with the Hemiptera (including Heteroptera) was Fabricius’s Systema Rhyngo- torum (1803), It was followed 40 years later by the much more comprehensive and influential Histoire mturelle des insectes hemipteres of Amyot and Serville (1843). In sub¬ sequent years many treatments have appeared, most of them dealing with a single family or a specific region. More general treatments have usually appeared in text¬ books, wherein individual families are given spare cov¬ erage and less familiar families are often excluded com¬ pletely because of their rarity or a lack of space or both. The number of taxa known by Fabricius was compara¬ tively small, and thus he was able to treat the world fauna at the species level. Today, with over 38,000 known species of Heteroptera, such an undertaking would take several lifetimes and many volumes. Until now, the most recent attempt to treat all fami¬ lies of Heteroptera was The Biology of the Heteroptera (Miller, 1956a, 1971). This work concentrated heavily on the Reduviidae, the family of Miller’s primary interest. His treatments of most of the other families were gener¬ ally too brief to be adequately informative and in some cases contained numerous factual errors. The present volume was conceived as a way of provid¬ ing a general summary of what is currently known about the Heteroptera. We first offer a nonsystematic introduc¬ tion to the group in chapters that cover the history of its study; a review of the major workers, techniques, and sources of specimens; attributes of general biologi¬ cal interest; selected taxa of economic importance; and basic morphology. Second, we present a current classifi¬ cation of the Heteroptera, synthesizing to the subfamily and sometimes tribal level the enormous and scattered lit¬ erature and supplying diagnoses, keys, figures, general natural history information, a summary of distributions, and a listing of important faunistic works. Third, we list references for over 1350 published works dealing with Heteroptera. Finally, we provide a glossary as an aid to organizing and interpreting the welter of terms that have appeared over the years and that often differ for the same structure from family to family. Restating the view of Cobben (1968:360): “Authors of textbooks and faunistic works are . . . faced with a bewildering array of ‘higher classification[s]’ and proper selection is extremely difficult. Confusion and error [are] bound to occur.” Although a great number of classifica¬ tions have been proposed for the Heteroptera, many are inadequately documented. We have therefore attempted to provide a classification for the group which is best sup¬ ported by existing information—down to the subfamily level. This classification, we hope, will serve all biologists studying the Heteroptera as a framework for the presenta¬ tion of other comparative information. Taxa are grouped by infraorder, superfamily, family, and subfamily. Although we provide diagnoses for infra¬ orders, families, and subfamilies, we do not provide them for superfamilies because superfamily concepts, particu¬ larly in the Pentatomomorpha, are in flux, with little agreement in the literature among the various authors. The sequence of family presentation is usually alphabetic in those groups for which no credible phylogenetic evi¬ dence has been published. The lack of such evidence is most glaringly obvious in the Pentatomomorpha. The structure of information presented in the family treatments is parallel for the most part. For a few of the larger families, however, we have included information on natural history and distribution and faunistics under each subfamily, because we found it impossible to pro¬ duce useful generalizations at the family level, in contrast to the approach taken for most of the smaller families. Pavel §tys, Charles University, Prague, deserves spe¬ cial thanks for his substantial contribution to this work. He wrote the sections on Enicocephalomorpha and Dip- socoromorpha and prepared the keys for the Nepomor- pha chapters. Dr. Stys provided critical commentary on several of the introductory chapters as well as all chap- Preface XI ters dealing with the Cimicomorpha. Last, but not least, he made many helpful suggestions regarding the general organization and contents of the volume. The following colleagues assisted in the preparation of this work. For reading and commenting on the en¬ tire manuscript we thank W. R. Dolling, 1. M. Kerzh- ner, G. M. Stonedahl, and M. H. Sweet. For reading and commenting on portions of the manuscript we thank N. M. Andersen, J. Grazia, T. J. Henry, D. A. Polhemus. J. T. Polhemus, C. W. Schaefer, and M. D. Schwartz. For assistance in the library and with securing references we thank S. O. Fischl, C. Chaboo Michalski, R. Pack- auskus, J. T. Polhemus, M. D. Schwartz, G. M. Stone¬ dahl, the staff of the American Museum of Natural His¬ tory library, and the many colleagues who have faithfully sent us papers over the years. For the loan of original figures we thank R. C. Froeschner, J. D. Lattin, A. S. Menke, C. W. Schaefer, Kathleen Schmidt, G. M. Stone¬ dahl, and the Division of Entomology, CSIRO, Canberra, Australia. For preparation of original figures we thank Kathleen Schmidt. For the loan of specimens used in preparation of the scanning micrographs of Nepomorpha we thank J. T. Polhemus. Donna Englund and Beatrice Brewster assisted with editing and printing various drafts of the manuscript. Pel- ing Fong Melville and William Barnett, Interdepartmen¬ tal Laboratory, American Museum of Natural History, assisted in the preparation of the scanning electron micro¬ graphs. M. D. Schwartz assisted in scanning and assem¬ bling many of the black-and-white line illustrations, with help from J. M. Carpenter, G. Sandlant, and S. Stock. Lee Herman facilitated preparation of the index. We especially thank our wives. Brenda Massie and Elizabeth Slater, for attentively listening to progress re¬ ports and other tedious details concerning the preparation of this volume. Robb Reavill, Cornell University Press, worked patiently with us and offered many encouraging words during the lengthy preparation of the manuscript. Helene Maddux and Margo Quinto helped bring the text into its final form. All previously published figures are used with permis¬ sion. We gratefully acknowledge the generous coopera¬ tion of publishers, authors, and artists for allowing us to reproduce their work. The College of Agriculture and Life Sciences Fund, Cornell University, helped support preparation of the art¬ work for this volume. Randall T. Schuh James A. Slater New York, New York Storrs, Connecticut xii Preface 1 A History of the Study of the Heteroptera Classification of the Heteroptera has reached its present state through a long evolutionary process beginning, in¬ sofar as modem systematics is concerned, with the work of Linnaeus. Taxonomic studies did not take place in an intellectual vacuum, but rather were the result of forces arising from both the general scientific community and society at large. Reviews of this historical development have been published by Stys and Kerzhner (1975) and Gollner-Scheiding (1991). Early Attempts at Higher Classification The first recognized higher group to include the true bugs was the Hemiptera of Linnaeus, a group that also included thrips, aphids, scale insects, and cicadas. Although, as the name indicates, Linnaeus based his group on the structure of the wings, he also recognized the distinc¬ tive namre of the hemipteran rostrum, subdividing the group into those insects with the “rostrum inflexum” (true bugs, cicadas, and other Auchenorrhyncha) and “rostrum pectorale” (scales and some other Stemorrhyncha). The true bugs were divided by Linnaeus in the tenth edition (1758) of the Systema naturae into three genera: No- tonecta, Nepa, and Cimex. These are all familiar modern- day generic names, but the concepts attached to them have become more restricted over time, particularly for Cimex. Fabricius, a student of Linnaeus, placed those insects with distinctive sucking mouthparts in his group Rhyn- gota (Rhynchota of later authors) and was the first to prepare a “monograph” of the group, the Systema Rhyn- gotorum (1803), in which he recognized 29 genera. Fabri¬ cius’s greatest misconception, from the view of modern classifications, was the inclusion of Pulex (Siphonaptera), the fleas, in the Rhyngota. The French naturalist Latreille used the Linnaean term Hemiptera to refer to the Rhyngota of Fabricius (but excluded the fleas). He formally named the subgroups Homoptera and Heteroptera (Latreille, 1810) and later divided the Heteroptera into the Geocorisae and Hydro- corisae, groupings based on the structure of the antennae. Dufour (1833) subsequently divided the Geocorisae of Latreille, recognizing the Amphibicorisae (modern-day Gerromorpha). Fieber (1861) introduced the redundant descriptive terms Gymnocerata and Cryptocerata for the Geocorisae and Hydrocorisae, respectively. A decade after the appearance of Dufour’s (1833) monograph, the first comprehensive family-group classi¬ fication of the Heteroptera was published by Amyot and Serville, their Histoire naturelle des insectes Hemipteres (1843). Many of the included names between the level of order and modern families have fallen into disuse be¬ cause they were descriptive, rather than following the modern convention of being based on generic names. Nonetheless, the Amyot and Serville classification was a fundamental advance, and it has had a lasting impact. Not all subsequent authors followed the nomencla¬ ture of Latreille, causing confusion as to which names should be applied to the subgroups of insects with their distinctive sucking mouthparts. The term Hemiptera has been applied by many North Americans only to the true bugs (the Heteroptera of Latreille) as an order coordi¬ nate with the Homoptera, rather than treating the two as subgroups of the more inclusive Hemiptera as done by most Europeans. Modern textbook authors such as Hor¬ ror, Triplehorn, and Johnson (1989) have argued that the Heteroptera should be called Hemiptera and treated at the ordinal level because they are sufficiently morphologi¬ cally distinct from the Homoptera. We recognize what appear to be monophyletic groups and apply to them names of longest standing, irrespective of categorical rank. Thus, whereas the terms Hemip¬ tera sensu law, Coleorrhyncha, and Heteroptera iden¬ tify monophyletic groups, it now seems clear that the Homoptera are not a natural group (e.g., CSIRO, 1991), and would better be referred to as Stemorrhyncha and Auchenorrhyncha (see Fig. 1.1). Descriptive Foundations of Heteropteran Classification The establishment of specific entities and the means for their recognition form the building blocks upon which all meaningful phylogenetic, biological, ecological, and physiological studies are based. Early work, including that of Linnaeus and Fabricius, in addition to establish- History of Study of Heteroptera 1 ing what has come to be recognized as a monophyletic assemblage, began the long tradition of describing, in a formal system, genera and species from all parts of the world. Their work was strongly influenced by the many exotic insects that were brought to Europe by explorers and early scientific expeditions. The impetus given to basic taxonomic work by these hitherto unknown species persists to the present day. For each family of any size there exist descriptive de¬ velopments, sometimes of great complexity, which mir¬ ror developments in higher classification. We have at¬ tempted to summarize these in discussions of individual families, and discuss only more general trends in the following paragraphs. Slater (1974), in a discussion of the South African Heteroptera fauna, recognized four periods, which can also be applied to the development of basic knowledge of Heteroptera throughout the world; Classical Period, Period of European Specialists, Intermediate Period, and Contemporary Period. The classical period, beginning with Linnaeus, ex¬ tends roughly to 1870. Most of the workers during this time studied the Heteroptera in general. Although many published works were important syntheses encompassing the first major higher classifications, nearly all contained large amounts of basic descriptive work. Most works up to this point, with some notable exceptions, were devoid of illustrations, even up to and through the magnificently detailed and influential works of Carl Stal. A series of specialized works began to appear primarily in Europe in the 1870s. Knowledge of some families had reached a point where what we today call revisional studies became imperative, while at the same time the opening of many previously inaccessible parts of the world brought rich collections into European centers and stimulated a great deal of descriptive work. Some of this work was faunistic, as for example the great Biologia Cen- trali Americana and Fauna of British India. It was a period of dominance for systematics, and some of the best scien¬ tific minds were involved. During this time Lethierry and Severin (1893-1896) produced the first and only world catalog of the Heteroptera. Their catalog was followed by comprehensive catalogs for the Palearctic (Oshanin, 1906-1909, 1912) and Nearctic (Van Duzee, 1916, 1917) faunas. This period of descriptive intensity, which con¬ tinued until about 1920, was marked by the publication of impressive faunal compendia, and the inclusion of illustrations became much more common. European dominance of systematics began to decline after World War I, and heteropteran studies greatly in¬ creased in the United States and Japan. Many impor¬ tant works were published during this period, including scores of papers by prolific authors such as C. J. Drake, H. H. Knight, H. B. Hungerford, and T. Esaki. Novel approaches were seen in works of authors such as H. H. Knight, who consistently applied the use of male genitalic characters in species recognition. The modern period, beginning in about 1950, is re¬ markable in the history of heteropteran classification. It embraces the completion of many fundamental revisional studies, some covering the entire world, such as those of Usinger and Matsuda (1958) on the Aradidae, Usinger (1966) on the Cimicidae, and Andersen (1982a) on the Gerromorpha. World catalogs of several families also ap¬ peared, for example, Miridae (Carvalho, 1957, 1958a, b, 1959, 1960), Lygaeidae (Slater, 1964b), Tingidae (Drake and Ruhoff, 1965), and Reduviidae (Putchkov and Putch- kov, 1985, 1986-1989; Maldonado, 1990), as well as the only up-to-date regional catalog, that of Henry and Froeschner (1988) on the North American fauna. The First World Specialists Many concepts of heteropteran classification up through the time of Fieber and his Die europdischen Hemiptera (1861) were based largely on the European fauna. With the appearance of the works of Carl Stal in the late 1850s. things began to change. In the course of two decades Stal monographed in a series of papers of increasing geographic scope—the best known and most sweep¬ ing being the Enumeratio Hemipterorum (1870-1876)— the Reduviidae, Lygaeidae, Coreidae, Pentatomidae, and several other families. In some groups such as the Corei¬ dae, no subsequent equivalent work has appeared, and StaTs keys still serve as important aids to identification. StaTs successor as dean of heteropteran taxonomy was O. M. Reuter. Whereas Stal had concentrated primarily on the Pentatomomorpha and Reduviidae, Reuter devoted most of his taxonomic efforts to the Miridae, Saldidae. and Cimicoidea. He authored the definitive works on the Palearctic fauna for the first two groups (Reuter, 1878- 1896, 1912b) and produced a world classification of the Miridae (Reuter, 1910). Reuter’s contemporaries such as C. Berg, E. Bergroth, G. Breddin, G. C. Champion, W. L. Distant, G. Hor¬ vath, V. E. Jakovlev, G. W. Kirkaldy, J.-B. A. Puton, P. R. Uhler, and many others were involved mostly in the production of descriptions of new taxa in faunistic works, tremendously broadening knowledge of the world fauna. Comparative Morphology in the Study of the Heteroptera Beginning with the work of Tullgren (1918) on abdomi¬ nal trichobothria, followed by the works of Singh-Pruthi (1925) on male genitalia, Poisson (1924) and Ekblom 2 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) (1926, 1930) on a variety of structural systems, and Spooner (1938) on the head capsule, comparative mor¬ phological studies in the Heteroptera began to develop in a way not encountered since the work of Dufour, and they began to be integrated into heteropteran clas¬ sification. Although the impact of our understanding of morphological variation on classifications is in most cases discussed below under the relevant section dealing with morphology, some areas merit special mention. Genitalia. Genitalia were first used in heteropteran classification by Verhoeff (1893) for the females and Sharp (1890) for the males. Although Poisson (1924) made an important contribution to detailing the structure of male genitalia in the Gerromorpha and Nepomorpha, Singh- Pruthi (1925) was the first to examine comparatively the detailed structure of the phallus for nearly all major het¬ eropteran groups. His study has some limitations from a modern perspective, because he did not examine all major groups—notably the Enicocephalidae—and exam¬ ined only gross structure. He nevertheless demonstrated that there are distinctive genitalic types within the Het¬ eroptera, and he proposed a scheme of interrelationships based on those characters. Notable was his recognition of the pentatomoid genitalic type, although his reduvioid genitalic type almost certainly forms a group based on symplesiomorphy. With the exception of the works of Ekblom (1926, 1930), the conclusions of Singh-Pruthi were not reexam¬ ined or extended until the comparative studies of Kullen- berg (1947) on the Miridae and Nabidae. Further broad- based studies on genitalia (Pendergrast, 1957; Scudder, 1959), as well as those for individual families (e g.. Slater, 1950: female Miridae; Ashlock, 1957: male Ly- gaeidae; Kelton, 1959: male Miridae; Carayon: many papers on male and female Naboidea and Cimicoidea), produced results that greatly advanced our understanding of heteropteran relationships at all levels. Internal anatomy. It was not until about 1940 that significant additional comparative documentation of ana¬ tomical details beyond that acquired by Dufour began to appear. Notable works include those of Baptist (1941) on the scent-gland system and Miyamoto (1961a) and Goodchild (1963) on the alimentary canal. Whereas many anatomical studies have not produced information on variation that proved to be of value in higher-level clas¬ sification, existing knowledge makes it clear that the gut type found in the Pentatomomorpha is clearly distinctive and, along with many other characters, argues for the monophyly of that group. Other internal structures, such as the dorsal vessel and nervous system, are relatively homogeneous across all Heteroptera. Ovariole numbers were first investigated on a com¬ parative basis by Woodward (1950) and Carayon (1950a) and later by Miyamoto (1957, 1959) and Balduf (1964). Testis follicle numbers were initially studied by Wood¬ ward (1950) and later for the Miridae by Leston (1961). Although both sets of structures suggest certain patterns of relationships, they are nonetheless relatively simple and possess substantial variability. It is only in the Cimi¬ coidea. a group first investigated through studies of the human bed bug and later broadened by Carayon (e.g.. 1977) to include all cimicoids, that internal reproduc¬ tive anatomy has offered a wealth of detail pertinent to classification of the group. Methodological Issues in Heteropteran Classification The earliest heteropteran classifications were based largely on single character systems, such as the structure of the wings or mouthparts. Additional discriminating characters—such as color, size, pronotal shape, antennal length—were largely those used to differentiate species. Some classifications, such as that of Schipdte (1869. 1870) founded on coxal types, were based not only on single character systems, but also on ones that showed little concordance with other available information; con¬ sequently they were adopted by only a few subsequent authors. Others, as pointed out by Stys and Kerzh- ner (1975), were totally undocumented, introducing new' taxonomic concepts in conjunction with the publication of checklists or catalogs. Reuter (1905, 1910, 1912a) stated in his treatises on heteropteran phylogeny that the early classifications of the Heteroptera were linear in character, and it was therefore not appropriate to make deductions about phylogenetic relationships. As Reuter (1910:31) noted, when Fieber (1861) was “compelled to select characters for his key, which permitted him to put related families together in a series, it is certain that the selection of the structure of the tarsi, as a consequence of which he put the fami¬ lies Phymatidae, Aradidae, Tingididae, and Microphysae [j/c] after each other, was a mistake” (our translation). Reuter further rejected the idea of deriving one group from another, as in the classifications of Kirkaldy. Reuter’s phylogenetic work incorporated several im¬ portant innovations: first, he reviewed all of the previ¬ ously proposed classifications of the group; second, he summarized and discussed the eharacters on which those classifications were based; and third, he made a list of characters he believed to be diagnostic for the groups he proposed. His effort was very nearly the preparation of a synapomorphy scheme. For example, with reference to the structure of the antennae in the Nepomorpha (Crypto- cerata) he said (Reuter, 1910:26): “The water bugs have short, concealed antennae. This. . . type is doubtless. . . History of Study of Heteroptera 3 an adaptation to life in the water that was acquired later [from long exposed antennae] and it, might not be at all excluded that it represents a heterophyletic homomor¬ phism unless other conditions were at hand which made it probable that at least most of the so-called Cryptocerata were homophyletic” (our translation). Reuter was also consistent in his reference to the composition of groups, using the terms homophyletic and heterophyletic, to refer to what in modern parlance would be monophyletic and polyphyletic. Character polarity. Unfortunately, as has been the case with the work of many other authors old and new, Reuter’s method for determining character polarity was probably the weakest aspect of his work. In some cases— such as number of tarsal segments—he used the “com¬ mon equals primitive” principle. In other cases he treated some groups as more ancient than others because they appeared to bear a greater number of primitive charac¬ ters. In this vein, Reuter’s thinking was pervaded by the idea that the Heteroptera are strictly diagnosable on the structure of the “hemelytra.” He stated on more than one occasion that it is clear that the corium, clavus, and membrane were present in the primitive forms. Thus, in his view the undifferentiated mesothoracic wings in the Enicocephalomotpha, Dipsocoromorpha, and Gerromor- pha were explicitly derived from the differentiated type (Reuter, 1910). This point of view contrasts markedly with recent studies involving the' search for congruence among more characters, which suggest that the relatively undifferentiated wings represent the primitive condition, the true “hemelytron” being derived. Cobben (1978:5) commented that some authors who had worked on feeding behavior and mouthpart structure in phytophagous Hemiptera “failed to consider informa¬ tion previously published in a wide variety of papers on comparative morphology and systematics ... or [be¬ lieved] . . . that the Homoptera are more generalized (or more symplesiomorphous) than the Heteroptera.” Although Cobben was at times inconsistent in his argu¬ ments-—as for example, in treating the Gerromorpha as the basal heteropteran group—his criticism can be ap¬ plied to the works of China and Myers (1929) and China (1933), who were of the opinion that the primitive Het¬ eroptera were phytophagous and were homopteroid in character, and to Miles (1972) and Sweet (1979), who asserted that, because many pentatomomorphs produce a salivary sheath or feeding tube similar to that found in the Sternorrhyncha and Auchenorrhyncha, they must possess the primitive heteropteran feeding type. Homology problems. Some authors arrived at erro¬ neous conclusions concerning homology of structures. For example, China (1933) noted the “absence of aro- lia” in Leotichius Distant, which meant that there were no fleshy pads between the claws. Such remarks, which are widespread in the literature, overlook the fact that nearly all Heteroptera possess similarly placed—although sometimes differently formed—structures between the claws. Fleshy structures were arolia by definition, bur setiform structures (parempodia) of similar position were not treated as homologous. Tullgren (1918) attempted to- resolve this problem, but as indicated by the labeling in his figure 11, his concept of an arolium was based solely on a definition without respect to structural homoiogy, and because he reasoned from a false premise he ar; ved at a false conclusion. Adaptationist arguments. Character analysis in much of the earlier literature was permeated by adaptationist arguments. For example, China and Myers (1929), China (1933), and Kullenberg (1947) argued that Reuter had placed an “exaggerated importance” on the arolia and claws as a guide to relationships in the Heteroptera—and particularly the Miridae—because “these organs are far too plastic to serve as a fundamental group character” (China, 1933:192). Such arguments overlooked the fact that if characters did not vary they would be of no use in forming groups and that only by examining this variability for congruence with other characters does one arrive at a hierarchic classification. Autapombrphy. Several authors have been impressed with the number of autapomorphous characters possessed by certain heteropteran taxa and have selectively used novelty of appearance as a measure for determining clas- sificatory rank. Possibly the earliest of these was Lin¬ naeus, in his recognition of Nepa and Notonecta as dis¬ tinctive from his omnibus Cimex, the last clearly a group devoid of defining characters other than those of the true bugs as a whole. Distant (1904) originally (and in our view correctly) placed Leotichius in his Leptopinae (Lep- topodidae). China (1933) elevated the group to family status, concluding that the taxon could not possibly be¬ long to the Leptopodidae, not because it did not have many of the attributes of other members of that group, but rather because it had several distinctive features of its own including, among others, only 3 veins in the mem¬ brane with no closed cells, and a 1-2-2 tarsal formula. Along similar lines Froeschner and Kormilev (1989) and Maldonado (1990) argued that the Phymatinae are so dis¬ tinctive as to merit family status, whereas all the features that diagnose the Reduviidae are also found in the Phy¬ matinae, although sometimes in a more highly modified form. The advent of cladistics and its more rigorous approach to character analysis is beginning to have an impact on heteropteran classification (Schuh, 1986c). As noted by Stys and Kerzhner (1975), the dismemberment of the classic Geocorisae comes from the realization that the 4 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) group was based on only a few symplesiomorphies and that groups such as the Enicocephalomorpha and Dip- socoromorpha are not closely related to the two major groups of land bugs, the Cimicomorpha and Pentatomo- morpha. Modern Higher Classification of the Heteroptera The classic subdivisions of Latreille and Dufour were used in heteropteran classifications well into the twentieth century. Although Reuter and others proposed subordinal names, most were never consistently adopted. Leston, Pendergrast, and Southwood (1954) introduced the terms Cimicomorpha and Pentatomomorpha in the first formal attempt to recognize natural groups within the poly- phyietic Geocorisae. Their conclusions were based on accumulated evidence from comparative studies of inter¬ nal anatomy and external morphology of the Heteroptera. The influence of their work was widely felt; other authors adopted these groups and applied the “morpha” nomen¬ clature to additional higher groups in the Heteroptera, although not without some variant spellings along the way (see, e.g,, Popov, 1971). More important, the work of Leston et al. (1954) spurred the attempt to document the monophyly of higher groups within the Heteroptera, with the eventual recognition of seven such groups—termed infraorders—all with typified names, as outlined by Stys and Kerzhner (1975). We present below diagnoses and references for each of these infraordinal groupings. The evidence for the relationships among them is discussed in the following paragraphs. The first documented higher-level scheme for the seven heteropteran infraorders was that of Schuh (1979). The character data were drawn mostly from information as¬ sembled by Cobben (1978). Cobben (1981a, b) criticized Schuh’s scheme because all the characters used to support it showed some within-group variability, even though he had argued that many of those same characters were pos¬ sibly diagnostic for certain groups. Other authors found certain portions of the scheme of interest, as for ex¬ ample Andersen (1982a) who agreed on the placement of the Gerromorpha among the more primitive heteropteran infraorders. Slater’s (1982) arrangement of the seven infraorders was in line with that of Schuh’s scheme. Stys (1985a) provided a set of names for the basal inclusive groups on the cladogram (Fig. 1.1). More recently the authors of the Insects of Australia (CSIRO, 1991) have portrayed the Coleorrhyncha (Peloridiidae) as the sister PANHETEROPTERA I •' NEOHETEROPTERA I !• EUHETEROPTERA J-' HETEROPTERA HETEROPTERODEA Fig. 1.1. Phylogenetic relationships of heteropteran infraorders (after Wheeler et at, 1993). group of the Heteroptera (Heteropterodea; see Schlee. 1969), and the Auchenorrhyncha as the sister group of those two combined (e.g., Emel’yanov, 1987). thereby treating the classic Homoptera as paraphyletic. These out¬ group relationships were not explicitly specified by Schuh (1979), who did not provide arguments for the polarity of characters used in his scheme. Little new evidence for higher-group relationships within the Heteroptera was adduced since the work of Schuh (1979), until the publication of 18s nuclear rDNA sequences by Wheeler et al. (1993) for 29 hemipteran taxa, representing all infraorders and six outgroup taxa, including the Psocoptera, Sternorrhyncha, Auchenor¬ rhyncha, and Coleorrhyncha. Their scheme is shown in Fig. 1.1, indicating substantial congruence between the molecular data and most of the morphological data used by Schuh (1979). Certain things about this scheme are at variance with traditional ideas concerning phylogenetic relationships within the Heteroptera. First, the Heteroptera are primi¬ tively predaceous, contrary to the beliefs of China and Myers (1929), Miles (1972), Sweet (1979), and others. Second, the Enicocephalomorpha are the basal heterop¬ teran lineage, rather than the Nepomorpha (Hydrocori- sae) as proposed by Reuter (1910) and China (1933), or the Gerromorpha as proposed by Cobben (1978), or the Pentatomomorpha as strongly implied by Sweet (1979). Third, the “hemelytron” is not a ground-plan character for the Heteroptera, but rather a synapomorphy for the Panheteroptera. We now await additional evidence to test this scheme, around which much of the remainder of the present work is based. History of Study of Heteroptera 5 2 Major Workers on the Heteroptera The following section provides brief descriptions of now deceased influential or controversial workers on the Het¬ eroptera from the time of Linnaeus. Where possible we provide dates and places of birth and death, a short biographical sketch, citations of particularly influential contributions to the field, and references to published bibliographies. Most of the information was located by consulting the works of Derksen and Gollner-Scheiding (1963-1975) and Gilbert (1977). We have not covered all workers because of limitations on space and avail¬ able information. Names that could be considered of equal significance to many of those cited might in¬ clude: G. C. Champion, A. Costa, E. F. Germar, F. E. Guerin-Meneville, R. F. Hussey, C. L. Kirschbaum, F. L. Laporte, I. LaRivers, O. Larsen, O. Lundblad, S. Matsumura, G. Mayr, W. L. McAtee, L. R. Meyer- Dur, P. Montrouzier, V. Motschulsky, E. Mulsant, E. C. Reed, C. P. Thunberg, F. Walker, J. O. Westwood, and A. Wroblewski. Living workers are dealt with by reference to their works at appropriate places in the text. Amyot, Charles Jean-Baptiste. b. Vandeuvre, Aisne, France, September 23, 1799; d. Paris, Octo¬ ber 13, 1866. Orphaned at early age, taken in by wealthy businessman Pavet with home in Paris opposite that of Audinet-Serville. Lawyer and influential person in French society. Coauthor with Serville of Histoire natu- relle des insectes Hemipteres (1843). Had no personal collection; worked with that of Serville and Museum National d’Histoire Naturelle, Paris. Bibliography pub¬ lished by Signoret (1866). Ashlock, Peter D. b. San Francisco, California, USA, August 22, 1929; d. Lawrence, Kansas, Janu¬ ary 26, 1989. Ph.D., University of California, Berkeley. Curator, Bishop Museum, Honolulu (1964-1967); pro¬ fessor, University of Kansas (1968-1989). Specialist on Orsillinae (Lygaeidae) and author of major classifica¬ tion of group (1967); also strong interest in systematic methodology. Collection deposited University of Kansas. Bibliography and list of names proposed published by Slater and Polhemus (1990). Barber, Harry G. b. Hiram, Ohio, USA, April 20. 1871; d. Washington, D.C., January 27, 1960. Second¬ ary school teacher in New York City. After retirement a specialist in Heteroptera, U.S. Department of Agricul¬ ture, Washington, D.C. Expert on Lygaeidae, particu¬ larly North American and Caribbean faunas. Collection deposited National Museum of Natural History, Wash¬ ington, D.C. Bibliography of over 100 papers and list of names proposed published by Ashlock (1960). Berg, Carlos, b. Tuckum (Tukums), Curlandia, Lat¬ via, April 2, 1843; d. Buenos Aires, Argentina, Janu¬ ary 19, 1902. First employed as biologist at museum in Riga, Latvia, later in Baltic Polytechnic, Riga. Moved to Argentina June, 1873, to fill position in museum in Buenos Aires, of which H. Burmeister was then director. Lectured in local colleges and universities, succeeding Burmeister as director of museum in 1892. Published widely in biology; best known to heteropterists ioxHemip- tera Argentina (1879, 1884). Majority of types deposited in Museo de la Plata, a small number in Museo Nacional de Historia Natural, Buenos Aires. Bibliography pub¬ lished by Gallardo (1902). Bergevin, Ernest de. b. 1859; d. 1933. Author of many alphataxonomic, faunistic, and bionomic papers on fauna of North Africa. Collection deposited in Museum National d’Histoire Naturelle, Paris. Bergroth, Ernst Evald, b. Jakobstad (Pietarsaari), Finland, April 1, 1857; d. Ekenas, Finland, Novem¬ ber 22, 1925. Swedish-speaking Finn, trained as a medi¬ cal doctor. Lived in United States 1905—1911 (Alaska, Minnesota, Massachusetts), remainder of life in Finland. Known for linguistic ability, mastery of many heterop- teran families (notably Aradidae), and biting critiques of work of others. Did little field work but mostly identi¬ fied collections of others; numerous types deposited in University Zoological Museum, Helsinki, and elsewhere. Bibliography of 317 papers published by Lindberg (1928). Blatchley, W. S. b. North Madison, Connecticut, USA, October 6, 1859; d. Indianapolis, Indiana, May 28, 1940. Master’s degree from Indiana University with thesis on butterflies of Indiana. State geologist of Indiana 1894- /-N, 6 TRUE BUGS OF THE \A/ORLD (HEMIPTERA; HETEROPTERA) 1910. Individualist, all-around naturalist, entomological generalist, prolific publisher, and sometime antagonist of several heteropterist contemporaries (e.g., Knight, 1927; Blatchley, 1928). His Heteroptem of Eastern Norih America (1926), a singular and still widely respected work. Collection deposited at Purdue University, Lafay¬ ette, Indiana. Bibliography and list of new taxa published by Blatchley (1930, 1939). Bliven, B. P. Resident of Eureka, California, USA; d. Eureka, ca. 1980. Between 1954 and 1973 described numerous species of Auchenorrhyncha and Heteroptera from western North America in his own journal. Occiden¬ tal Entomologist. R. L. Usinger and others argued that the names of the reclusive Bliven should be suppressed be¬ cause contemporaries were unable to examine specimens on which his taxa were based. Collection now available in California Academy of Sciences. Breddin, Gustav, b. Magdeburg, Germany, Febru¬ ary 25, 1864; d. Oschersleben, Germany, February 25, 1909. Director of a secondary school. Author of about 70 papers, mainly on Pentatomomorpha and Reduviidae from Orient, South America, and Africa. Collection de¬ posited in Deutsches Entomologisches Institut, Berlin- Dahlem; some material in Senkenberg Museum, Frank¬ furt. Burmeister, Hermann Carl Conrad, b. Stralsund, Germany, January 15, 1807; d. Buenos Aires, Argen¬ tina, May 2, 1892. Professor of zoology. University of Halle (1837-1861). Traveled and collected extensively in Brazil (1850-1852) and later Argentina, Peru, Panama, and Cuba (1857-1860). Left wife and family, moved to Buenos Aires, Argentina, in 1861 to begin new life as director of fledgling Museo Publico de Historia Natural, where he studied primarily fossil vertebrates. Prodigious worker, best known to entomologists and heteropterists for Handbuch der Entomologie (1832-1839; Heterop¬ tera in vol. 2, 1835). Collection in Zoological Museum, Martin Luther University, Halle-Wittenburg, Germany. Butler, Edward A. b. Alton, Hants, England, March 17, 1845; d. Clapham, England, November 20, 1925. Schoolmaster who studied natural history of Het¬ eroptera, summarizing his life’s observations in A Biology of the British Hemiptera-Heteroptera (1923). Collections deposited in The Natural History Museum, London. China, William Edward, b. London, England, De¬ cember 7, 1895; d. Mousehole, Cornwall, England, September 17, 1979. University education Trinity Col¬ lege, Cambridge University, D.Sc. 1948. Curator of Het¬ eroptera (and Keeper of Entomology, 1955-1961), The Natural History Museum, London. General heteropterist: described new species, genera, and suprageneric taxa; elucidated genitalic and other morphology; established relationships among poorly known taxa; and resolved nomenclatorial problems. Known especially for assist¬ ing other heteropterists and for keys to world families and subfamilies of Heteroptera (China and Miller. 1959). Bibliography of 260 papers published by W. J. Knight (1980). CoBBEN, Rene H. b. Netherlands, 1925; d. Rhenen. Netherlands, 1987. Gifted morphologist and illustrator; specialist in taxonomy and morphology of Leptopodo- morpha; professor of entomology, Netherlands Agricul¬ tural University, Wageningen, from 1954 until death. Author of monumental monographic treatises on egg structure and embryogenesis (1968a) and feeding struc¬ tures (1978), each with voluminous additional observa¬ tions and evolutionary syntheses; other important papers detailing classification and description of Leptopodomor- pha. Collections from Caribbean, Africa, and Europe de¬ posited in Agricultural University, Wageningen, Nether¬ lands. Bibliography of 64 papers published by de Vrijer (1988). Dallas, William Sweetland. b. London, England, January 31, 1824; d. London, May 28, 1890. Worked for a time in The Natural History Museum, London (around 1850), producing list of Hemiptera in its collections (1851-1852). Published additional books and translations and single-handedly prepared first five volumes of Insecta for Zoological Record. Collection in The Natural History Museum, London. De Geer, Carl. b. Finspang, Sweden, February 10, 1720; d. Stockholm, March 8, 1778. Wealthy Swedish nobleman of Dutch ancestry. Author of now rare work Memoires pour servir a I'histoire naturelle des insectes (1752-1771). Collection in Swedish Museum of Natural History, Stockholm. Distant, William Lucas, b. Rotherhithe, England, November 12, 1845; d. Wanstead, Essex, England, Feb¬ ruary 4, 1922. Businessman in hide-tanning industry. Traveled to Malaya and South Africa. From 1899 to 1920 part-time assistant. The Natural History Museum, Lon¬ don. Prolific species describer, ardent student of Cicadi- dae and Heteroptera. Author of Rhopalocera Malayana and A Naturalist in the Transvaal; best known to heterop¬ terists for sections of Biologia Centrali Americana (1880- 1893) and Fauna of British India (1902-1918). Criticized by Bergroth, Kirkaldy, Horvath, and Reuter (e.g., 1905) for failing to appreciate their works on heteropteran clas¬ sification. Many types, deposited in The Natural History Major Workers on the Heteroptera 7 Museum, London; additional types in Genoa, Italy. Bib- eroptera. Status of Fabrician types and other information liography of several hundred papers published by Dolling summarized in Zimsen (1964). Most material deposited (1991a). in Zoological Museum, Copenhagen. Douglas, John William, b. Putney, England, No¬ vember 15, 1814; d. Harlesden, England, August 28, 1905. Worked briefly at Kew Gardens, and then for over 50 years as a customshouse officer. Author of numer¬ ous papers on British Heteroptera, including The British Hemiptera (1865), coauthored with John Scott, first com- prv:nens:ve work on subject. Types in Hope Museum, Oxford. Drake, Carl J. b. Eaglesville, Ohio, USA, July 28, 1885; d. Washington, D.C., October 2, 1965. Ph.D. from Ohio State University under direction of Herbert Osborn. Longtime head of the Department of Zoology and Ento¬ mology, Iowa State College. World authority on Tingi- dae, and prolific publisher on Saldidae, Gerromorpha, and aspects of applied entomology. Primary interest in description of new species; included many excellent habi¬ tus illustrations of Saldidae and Tingidae in later papers. Collection and considerable estate willed to National Mu¬ seum of Natural History, Washington, D.C. Bibliography of 519 papers and list of names proposed published by Ruhoff(1968). Dufour, Leon. b. St. Sever, France, April 11, 1780; d. St. Sever, April 18, 1865. Published on a great diver¬ sity of insect groups. Author of first detailed anatomical work on Heteroptera, Rccherches anatomiques et physio- logiques sur k:: Hemipu"-cs (1833), a magnificently illus¬ trated study of salivar glands, alimentary tract, and re¬ productive and respiniory systems. Bibliography of more than 230 papers published by Laboulbene (1865). Collec¬ tion in Museum National d’Histoire Naturelle, Paris. Esaki, Teiso. b. Tokyo, Japan, July 15, 1899; d. De¬ cember 14, 1957. Professor of entomology, Kyushu University, Kyoto, Japan. Traveled widely in Europe, working with G. Horvath in Budapest and at The Natural History Museum, London. Best known for extensive work on aquatic Heteroptera, and particularly for discovery of Helotrephidae. Bibliography published by Hasegawa (1967). Fabricius, Johann Christian, b. Tonder, Denmark, January 7, 1745; d. Kiel, Germany, March 3, 1808. Trained as physician, close associate of Linnaeus, from early age interested in entomology. Described nearly 10,000 species. Appointed professor of natural history at University of Kiel at early age by patron. Author of Systema Rhyngotorum (1803), first monograph of Het- Fallen, Carl Friedrich, b. 1764; d. 1830. Profes¬ sor of Natural History, Lund University, Lund, Sweden. Provided first detailed treatments of Heteroptera of Scan¬ dinavia in Monographia Cimicum Sueciae (1807) and subsequent publications. Types in Swedish Museum of Natural History, Stockholm. Fieber, Franz Xavier, b. March 1, 1807; d. Chru- dim, Bohemia, Czech Republic, February 22, 1872, Director of Royal Imperial District Court at Chmdim. Author of earliest comprehensive publications on Euro¬ pean fauna, culminating in Die europdischen Hemiptera (1861). Established modern generic classification of Miri- dae based on European fauna. Most types in Museum National d’Histoire Naturelle, Paris and Naturhistorisches Museum, Vienna. Gmelin, Johann Friedrich, b. Tubingen, Germany, August 8, 1748; d. Gottingen, Germany, November 1, 1804. Professor of chemistry in Gottingen. Compiler of final (13th) edition of Systema naturae of Linnaeus. Hahn, Carl Wilhelm, b. December 16, 1786; d. 1836. German naturalist who worked in Nuremberg, on spiders as well as Heteroptera. Author of volumes 1- 3 of Die wanzenartigen Insecten (1831-1835), in which were described as new large numbers of European and extra-european genera and species. Work completed by G. A. W. Herrich-Schaeffer. Handlirsch, Anton, b. Vienna, Austria, January 20, 1865; d. Vienna, August 28, 1935. Longtime em¬ ployee of Naturhistorisches Museum, Vienna. Specialist in taxonomy of Sphecidae and later Heteroptera (particu¬ larly Phymatinae), insect paleontology, and higher-level classification. Bibliography published by Beier (1935). Heidemann, Otto. b. Magdeburg, Germany, Sep¬ tember 1, 1842; d. Washington, D.C., USA, Novem¬ ber 17, 1916. Emigrated to United States 1876, employed as an illustrator and engraver and later by U.S. Bureau of Entomology (USDA), Washington, D.C. Student of North American Heteroptera, mostly Aradidae, Miridae, and Tingidae, Collection deposited in Cornell University Insect Collection, Ithaca, New York. Bibliography of 34 papers published by Howard et al. (1916). Herrich-Schaeffer, Gottlieb A. W. b. Regens¬ burg, Germany, December 17, 1799; d. Regensburg, 8 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) April 14, 1874. As did father, served as physician in courts in Regensburg, from 1833 until retirement in 1856. Author, among other works on Heteroptera, Lepidoptera, and so on of volumes 4-9 of Die wanzenartigen Insecten (1839-1853), completing work of C. W. Hahn. Types destroyed during World War 11. Horvath, Geza. b. Csecs, northern Hungary, No¬ vember 23, 1847; d. Budapest, September 8, 1937. Trained as M.D., University of Vienna. First worked in control of pest insects; instrumental in elimination of Phylloxera from European vineyards; later appointed head. Department of Zoology, National Museum, Buda¬ pest. Traveled widely, active in international scientific affairs and influential in careers of other well-known heteropterists. Published on systematics of Heteroptera, Homoptera, and other subjects. Like A. N. Kiritshenko, reputedly made all observations with a hand lens. Bibli¬ ography of 467 papers published by Csiki (1944). Most types in Budapest. Hsiao, Tsai-yu. b. Shandong (Shantung) Province, China, July 25, 1903; d. Tianjin, China, June 27, 1978. Basic education in China, graduate work in United States at University of Illinois, Oregon State College (M.S.), and Iowa State College (Ph.D.) under H. H. Knight. Re¬ turned to China in 1946 after 10 years in United States and became dean and professor of biology at Nankai University, Tianjin, and head of Tianjin Natural His¬ tory Museum. Suffered during Cultural Revolution of late 1960s and early 1970s. Published on systematics of Het¬ eroptera, most comprehensive being two volumes of A Handbook for the Determination of Chinese Hemiptera- Heteroptera {W-si&o, 1977, 1981). Bibliography published by Schaefer and Sailer (1980). Hungerford, Herbert Barker, b. Mahaska, Kan¬ sas, USA, August 30, 1885; d. Lawrence, Kansas, May 13, 1963. Ph.D., Cornell University. Professor of ento¬ mology, University of Kansas, Lawrence; mentor of many students in Gerromorpha and Nepomorpha. Author of The Biology and Ecology of the Aquatic and Semiaquatic Hemiptera (1919) and comprehensive works on Notonecta of the world (1933) and Corixidae of the Western Hemi¬ sphere (1948), as well as numerous other papers. Out¬ standing collection of Gerromorpha and Nepomorpha de¬ posited University of Kansas. Bibliography published by Woodruff (1956; 1963). Jaczewski, Tadeusz L. b. St. Petersburg, Russia, February 1, 1899; d. February 25, 1974. Studied Uni¬ versity of St. Petersburg; moved to Warsaw University, Poland, 1920. Employed Warsaw Zoological Museum and Warsaw University. Organized several Polish faunaf series. Primary systematic interests in Nepomorpha, par¬ ticularly Corixidae, often in cooperation with O. Lund- blad. Prepared sections on aquatic and semiaquatic bugs for Keys to the Insects of the European USSR (Kerzhner and Jaczewski, 1964). Bibliography published by Wrob- lewski(1974). Jakovlev, Vasiliy E. b. Tsaritsyn (now Volgograd), Russia, February 9,1839; d. Eupatoriya, Ukraine, August 15, 1908. A government official living in different loca¬ tions, for 20 years in Astrakhan, for 12 years in Irkutsk; upon retirement in 1898 lived briefly in St. Petersburg and then in Eupatoriya (Crimea) until death. Conducted some vertebrate studies; best known to heteropterists for work on Russian fauna. Collections deposited in Zoological Institute, St. Petersburg. Bibliography and list of names proposed published by Semenov-Tian-Shanski (1910). Jeannel, Rene. b. 1879; d. Paris, February 20, 1965. One-time director of Museum National d’Histoire Naturelle, Paris; ardent collector, coleopterist, biogeog¬ rapher, and monographer of Enicocephalidae (1941). Bibliography published by Delamare Deboutteville and Paulian (1966). Collections in Museum National d’His¬ toire Naturelle, Paris. Kiritshenko, Alexandr Nikolayevich, b. Berd¬ yansk, Zaporozh’ye Province, Russia (now Ukraine), September 9, 1884; d. St. Petersburg, January 23, 1971. Longtime curator of Heteroptera, Zoological Institute, St. Petersburg; worked actively to time of death. Author of numerous papers on Palearctic fauna. Built extensive collection of Palearctic Heteroptera at Zoologieal Insti¬ tute, St. Petersburg. Bibliography published by Kerzhner and Stackelberg (1971). Kirkaldy, George Willis, b. Clapham, England, July 26, 1873; d. San Francisco, California, USA, Feb¬ ruary 2, 1910. Educated in England, moved to Hono¬ lulu, Hawaii, 1903. Worked initially for USDA Board of Agriculture and Forestry, later for Hawaiian Sugar Planters’ Experiment Station. Author of numerous works on Auchenorrhyncha and Heteroptera, mainly on tax¬ onomy, nomenelature, bibliography, and natural history, possibly best known among them a world catalog of Pen- tatomoidea (1909). Bibliography published by Dolling (1991a). Collections deposited The Natural History Mu¬ seum, London, Bishop Museum, Honolulu; Snow Ento¬ mological Museum, University of Kansas, Lawrence; and National Museum of Natural History, Washington, D.C. Major Workers on the Heteroptera 9 Knight, Harry Hazelton. b. Koshkonong, Mis¬ souri, USA, May 13, 1889; d. Ames, Iowa, Septem¬ ber 6, 1976. Ph.D.. Cornell University. Faculty mem¬ ber, University of Minnesota 1919-1924, then Iowa State University until retirement, mid-1950s. Nearly single- handedly detailed genera and species of North Ameri¬ can Miridae, describing in excess of 1300 species in 182 papers, notable among them Miridae of Connecticut (1923), Miridaeof Illinois (1941), and Miridae of Nevada Test Site (1968). Prolific collector in early years. First American worker to use male genitalia for species recog¬ nition. Collection, including nearly all types, deposited in National Museum of Natural History, Washington, D.C., with limited material also at Texas A&M University and Biosystematics Research Centre, Ottawa. Latreille, Pierre-Andre. b. Brive, Correze, France, November 29, 1762; d. Paris, February 6, 1833. Primarily an entomologist, having worked in Museum National d’Histoire Naturelle, Paris, and as a professor of zoology. Particularly worthy of mention in this volume for his proposal of term Heteroptera and classificatory work on the Heteroptera. Leston, Dennis, b. ca. 1919, England; d. 1979, Florida, USA. An iconoclastic commoner of flashing intellect. Author of numerous papers on Pentatomoidea, particularly of Africa, and Cimicoidea, testes follicle numbers, stridulatory mechanisms, and general classifi¬ cation of Heteroptera. Coauthor with T. R. E. Southwood of Land and Water Bugs of the British Isles (1959). Tropi¬ cal ecologist and collector stationed in Ghana and Brazil. Collections deposited in The Natural History Museum, London, with limited material in American Museum of Natural History, New York. Lethierry, Lucien. b. Lille, France, 1830; d. Lille, April 4, 1894. Entomologist of diverse interests, includ¬ ing Heteroptera, best known for his joint publication with M. Severin of Catalogue general des Hemipteres (3 vol¬ umes, 1893-1896), only nearly complete world catalog of group ever published, but not including Miridae. Types in Museum National d’Histoire Naturelle, Paris. Lindberg, Hakan. b. Joroinen, Finland, May 24, 1898; d. Helsinki, August 6, 1966. Swedish-speaking Finn educated at and longtime professor in Department of Zoology, University of Helsinki. Best known for work on Heteroptera fauna of Canary (1953) and Cape Verde (1958) islands. Collection deposited in University Zoo¬ logical Museum, Helsinki. Linnaeus, Carl. b. Rashult, Swaland, Sweden, May 24, 1707; d. Uppsala, January 10, 1778. Professor, Uni¬ versity of Uppsala. Father of systematic biology, author of earliest systematic concepts in Heteroptera. Matsuda. Ryuichi. b. Kajiki, Kagoshima Prefec¬ ture, Japan, July 8, 1920; d. Ottawa, Canada, June 19. 1986. Ph.D., Stanford University, under direction of G. F. Ferris. Held positions at University of Kansas. Univer¬ sity of Michigan, and Biosystematic Research Centre. Ottawa. Published extensively with H. B. Hungerford on Gerridae, R. L. Usinger on Aradidae (1959), as well as on general insect morphology. Bibliography published by Ando (1988). Miller, Norman Cecil Egerton. b. Ramsgate, Kent, England, July 13, 1893; d. Sturminster, Newton, England, May 26, 1980. Without formal training in ento¬ mology, worked as entomologist for Department of Agri¬ culture, Kuala Lumpur, Malaysia, 1928-1947; retired to Zimbabwe, 1947-1949; joined staff of Commonwealth Institute of Entomology in London, 1949-1958. Pri¬ mary interest in Reduviidae (and Acrididae); probably best known for his book The Biology of the Heteroptera (1956a, 1971), a work heavily weighted to his anecdotal observations on Reduviidae, and “Notes on the Biology of the Reduviidae of Southern Rhodesia” (1953). Collec¬ tions deposited in The Natural History Museum, London. Bibliography published by Dolling (1987b). Montandon, Arnold Lucien. b. Besan?on, France, 1852; d. Iasi (Jassy), Romania, March 1, 1922. Author of over 120 descriptive taxonomic papers on many groups of Heteroptera, especially Nepomorpha, Reduvi¬ idae, Lygaeidae (especially Geocorinae), Coreidae, and Plataspidae. Collection sold in parts, most in The Natural History Museum, London, and Natural History Museum. Bucharest. OsHANiN, Vasiliy F. b. Politovo, Lipetsk Province, Russia, December 21, 1844; d. St. Petersburg, Janu¬ ary 22, 1917. Educated Moscow University and later in western Europe. Worked briefly in Moscow and from 1872 to 1906 in Tashkent as director of a silk mill, and later as a teacher and director of Tashkent Girls’ School. Participated in cultural life in Tashkent, mostly with political deportees, among them G. A. Lopatin, a friend of Marx and Engels. Upon retirement in 1906 worked as an associate in Zoological Institute, St. Petersburg. Best known to heteropterists for his Verzeichnis der paldark- tischen Hemipteren (1906-1909), Katalog der paldark- tischen Hemipteren (1912), and 'Wade mecum" (1916), a guide to literature on identification of Heteroptera; also 10 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) translated rules of zoological and botanical nomenclature into Russian. Many Central Asia specimens destroyed by dermestids; some specimens in Zoological Institute, St. Petersburg, others in Zoological Museum, Moscow Uni¬ versity. Bibliography published by Kiritshenko (1940). Parshley, Howard Madison, b. Hallowell, Maine, USA, August 7, 1884; d. May 19, 1953. Ph.D., Har¬ vard University; professor and administrator. Smith Col¬ lege, Amherst, Massachusetts; accomplished musician. Early student and author on Aradidae and other groups in eastern North America, but possibly best known for A Bibliography of North American Hemiptera-Heteroptera (1925). Poisson, Raymond A. b. Briouze (Orne), France, February 18, 1895; d. Lucon, France, November 28, 1973. Professor of zoology. Faculty of Sciences, Rennes. Notable among more than 300 publications, those on Ger- romorpha and Nepomorpha of Palearctic and Ethiopian regions as well as papers on morphology, anatomy, and biology of Nepomorpha. Author of Heteroptera chapter in Traite de zoologie (Poisson, 1951). Collection pur¬ chased by Zoological Museum, Copenhagen (Veliidae) and National Museum of Natural History, Washington, D.C. (other groups). Bibliography published by Grasse (1974). Poppius, Robert Bertil. b. Kyrkslatt (Kirkko- nommi), Finland, July 28, 1876; d. Copenhagen, Den¬ mark, November 27, 1916. Swedish-speaking Finn, educated at University of Helsinki; worked University Zoological Museum, Helsinki, 1900-1914. Important revisionary work on Miridae, particularly Ethiopian re¬ gion (1912, 1914) and cimicomorphan fauna of Old World tropics; also an accomplished student of Cole- optera. Most types in University Zoological Museum, Helsinki, although many specimens originally from other museums. PUTON, Jean-Baptiste Auguste. b. Remiremont, France, 1834; d. Remiremont, April 8, 1913. M.D.; stu¬ dent of Palearctic and particularly French Heteroptera. Published over 150 papers, including taxonomy, faunis- tics, and biology of western European fauna, including several catalogs. Described numerous new species. Col¬ lection in Museum National d’Histoire Naturelle, Paris. Reuter, Odd Moranal. b. April 28, 1850, Turku, Finland; d. Turku, September 2, 1913. One of most pro¬ lific and influential heteropterists of all time. Ph.D. from and longtime professor at Imperial Alexander University (University of Helsinki); student of Miridae, Anthocori- dae, Saldidae, and many other families of Heteroptera and other orders. Notable works include lavishly illus¬ trated monograph of European Miridae entitled Hemip- tera Gymnocerata Europae (1878-1896). first character- based phytogeny of Heteroptera (1910), and first world classification of Miridae (1905, 1910). A gifted poet, who wrote of his own late-in-life blindness. Many types in Zoological Museum, Helsinki, and other institutions. Bibliography of more than 500 papers published by Pal- men (1914). Ruckes, Herbert, b. New York, New York, USA. February 1, 1895; d. New York City, December 23, 1965. Ph.D., Columbia University; professor of biology. City College of New York; Research Associate, Ameri¬ can Museum of Natural History. Student of Pentatomidae and osteology of Chelonia. Author of approximately 50 papers, most published in Journal of the New York Ento¬ mological Society and Bulletin and Novitates of American Museum. Say, Thomas, b. Philadelphia, Pennsylvania, USA, June 21 y 1787; d. New Harmony. Indiana, October 10, 1834. Father of entomology in North America. Spent short time as druggist; at 25 devoted himself to natu¬ ral history, working at Academy of Natural Sciences. Philadelphia. Made two major expeditions to western U.S. territories. Left Philadelphia permanently in 1825 to live in New Harmony, an ill-fated utopian commu¬ nity in southern Indiana. Most of Say’s writings rare in original; reprinted completely by LeConte (1859). De¬ scribed numerous heteropterans in a variety of families, mostly in his Descriptions of New Species ofHeteropterous Hemiptera of North America (1831). Most of collections destroyed by dermestids. ScHOUTEDEN, Henri. b. Brussels, Belgium, May 3, 1881; d. Brussels, November 15, 1972. D.Sc., Free Uni¬ versity, Brussels, 1905. Soon joined staff of Musee Royal du I’Afrique Centrale; appointed head of section 1919; director 1927-1946; worked actively to time of death. Student of Pentatomoidea, Coreidae, Tingidae, and Re- duviidae. Authored several lavishly illustrated volumes in Wytsman’s Genera insectorum (for example, Schouteden, 1905b, 1907, 1913). Possibly best known for work on birds of the former Belgian Congo (Zaire). Scott, John. b. Morpeth, England, September 21, 1823; d. Morpeth, August 30, 1888. Civil engineer; au¬ thor of papers on Tineidae, Psyllidae and other Auchen- orrhyncha, and Heteroptera (frequently with J. W. Doug¬ las), particularly of Great Britain. To heteropterists, best known for contribution on Miridae to The British Hemip- Major Workers on the Heteroptera 11 tera, Vol. 1: Hemiptera-Heteroptera (1865), coauthored with J. W. Douglas. Collection deposited in The Natural History Museum, London. Seidenstucker, Gustav, b. Nuremburg, Germany, June 1, 1912; d. November 18, 1989. Worked as health insurance official in Bavaria. Published 99 papers, many reporting results of his travels in Mediterranean and Tur¬ key, particularly on Miridae and Lygaeidae, including concise descriptions, keys, and useful illustrations. Col¬ lection deposited in Zoologische Staatssammulung, Mu¬ nich. Bibliography published by Heiss (1990). Serville, Jean-Guillaume Audinet. b. Paris, France, November 11, 1775; d. Gerte sous-Touarre, March 27, 1858. Born into wealthy French family; father secretary to a prince. Became acquainted with wealthy Mada'he Tigny, who entertained influential biologists of the time, including Latreille. Among many works, most important contribution on Heteroptera Histoire naturelle des insectes Hemipteres (1843) coauthored with C. J.-B. Amyot. Possessed one of finest insect collections of his time, later sold to a number of individuals. Signoret, Victor, b. Paris, France, April 6, 1816; d. Paris, April 3, 1889. Pharmacist and medical doc¬ tor. Among most important works “Revision du groupe des Cydnides” (1881-1884). Collection of Heteroptera deposited in Natural History Museum, Vienna. Bibliog¬ raphy published by Fairmaire (1889). Spinola, Maxmillian. b. Toulouse, France, July 1, 1780; d. Tassarolo, Italy, November 12, 1857. Author of 54 papers and books, mainly on Coleoptera, Hymenop- tera, and Heteroptera. including Essai sur les genres . . . des Heteropteres . . . (1837). Some of later works con¬ fusing for inclusion of genera without species and other features not in conformity with modern practice. Collec¬ tion deposited in Museo Regionale di Scienze Naturali, Turin, Italy (see Vidano and Arzone, 1976). Stal, Carl. b. Castle of Carlberg, Stockholm, Swe¬ den, March 21, 1833; d. Stockholm, June 13, 1878. One of most respected and influential heteropterists of all time. Primarily a student of Pentatomomorpha and Reduviidae, but also worked extensively in other groups. Studied under C. H. Boheman at Uppsala University and later received Ph.D. from Jena. Appointed assistant in entomology. National Zoological Museum, Stockholm, 1859; head of section and professor, 1867. Traveled widely in Sweden, continental Europe, and England, studying collections and conferring with colleagues. Ac¬ cording to O. M. Reuter, Stal had a well-nigh inspired eye for “essential characters significant of natural affinity.” Published numerous works on Heteroptera, notable among them compendious Enumeratio Hemipterorum (1870-1876), and significant works on Orthoptera and Chrysomelidae. Bibliography published by Spangberg (1879). Collections in Swedish Natural History Museum. Stockholm. Torre-Bueno, Jose R. de la. b. Lima, Peru, Octo¬ ber 6, 1871; d. Tucson, Arizona, USA. May 3. 1948. Graduated from School of Mines, Columbia University, New York City, 1894, later worked for General Chemi¬ cal Company, New York. Longtime editor of Bulletin of the Brooklyn Entomological Society, author of A Glos¬ sary of Entomology (1937), “Synopsis of North American Heteroptera” (1939, 1941), and numerous smaller papers and notes. Collection deposited Snow Entomological Mu¬ seum, University of Kansas, Lawrence. Uhler, Phillip Reese, b. Baltimore, Maryland, USA, June 3, 1835; d. Baltimore, October 21, 1913. Graduate of Harvard University; student of Louis Agas¬ siz. Librarian at Peabody Institute, Baltimore, and as¬ sociate professor at the newly formed Johns Hopkins University. Father of North American heteropterology. Monographer of North American Cydnidae and Saldidae; describer of numerous species from newly explored west¬ ern territories of United States. Collection deposited National Museum of Natural History, Washington, D.C. Bibliography published by Schwarz et al. (1914). UsiNGER, Robert L. b. Fort Bragg, California, USA, October 24, 1912; d. Berkeley, California, October 1, 1968. Student (B.S., Ph.D.) and longtime professor of entomology. University of California, Berkeley. Gifted and internationally recognized entomological organizer and administrator. Author or coauthor of many papers and several books, including. Methods and Principles of Systematic Zoology (Mayr et al., 1953), Aquatic Insects of California (1956), Monograph of Cimicidae (1966), and, with R. Matsuda, Classification of the Aradidae (1959). Life and personality chronicled in autobiography (1972). Bibliography and list of names proposed published by Ashlock (1969). Van Duzee, Edward Payson. b. New York, New York, USA, April 6, 1861; d. June 2, 1940. Librarian at Grosvenor Library, Buffalo, New York, from 1885 to 1912; later assistant librarian and curator of entomology, California Academy of Sciences, San Francisco. Ardent collector, bibliographer, and author of over 250 pub¬ lications. Probably best known for Catalogue of North American Hemiptera (1917); author of many new taxa. 12 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) particularly from California. Extensive collections from western United States form core of extensive Hemiptera collections of California Academy of Sciences. ViLLiERS, Andre, b. 1915; d. June 8, 1983. Worked in laboratories of Jeannel and T. Monod; traveled ex¬ tensively in Africa. Became subdirector of Laboratory of Entomology, Museum National d’Histoire Naturelle, Paris, 1956, with responsibility for Coleoptera. Pub¬ lished 661 papers, majority on Cerambycidae; also pro¬ duced immense body of work on Enicocephalidae and Reduviidae of Africa and Madagascar. Bibliography pub¬ lished by Quentin (1983). Collection in Museum National d’Histoire Naturelle, Paris. Wagner, Eduard, b. Hamburg, Germany, June 20, 1896; d. September 11, 1978. Educated and worked as a schoolteacher. Student of western European and Medi¬ terranean fauna, particularly of Miridae, important works including Die Tierwelt Deutschlands (1952, 1966, 1967) and “Die Miridae des Mittelmeerraumes” (1971-1978). First to consistently illustrate detailed structure of aedea- gus in Miridae. Collection deposited University of Ham¬ burg Zoological Museum. Bibliography of 553 papers and list of taxa proposed published by H. H. Weber (1976). Woodward, Thomas Emmanuel, b. Auckland, New Zealand, June 8, 1918; d. Brisbane, Australia, Novem¬ ber 22, 1985. Ph.D., Imperial College, London, under direction of O. W. Richards. Worked briefly in New Zea¬ land, then as professor. Department of Entomology, Uni-' versity of Queensland, Brisbane. Primarily a student of Australian and New Zealand Lygaeidae. Bibliography of 64 papers published by Monteith (1986). Wygodzinsky, Petr (Pedro) Wolfgang, b. Bonn, Germany, October 5, 1916; d. Middletown, New York, USA, January 27, 1987. Ph.D., University of Basel, Switzerland; student of Eduard Handschin. Lifelong stu¬ dent of Microcoryphia and Zygentoma. Emigrated to Brazil in 1940, began study of Heteroptera; moved to Argentina in 1948, began study of Simuliidae at Instituto Miguel Lillo, Tucuman; moved to American Museum of Natural History, New York City in 1962, publishing on all groups of interest. Specialist on Enicocephalomorpha, Dipsocoromorpha, and Reduviidae; published scores of papers (many heavily illustrated), including “Monograph of Emesinae” (1966) and “Revision of the Triatominae” with Herman Lent (1979). First to examine detailed struc¬ ture in Dipsocoromorpha, revealing wealth of morpho¬ logical detail (e.g., Wygodzinsky, 1947b, 1948a). Col¬ lection deposited American Museum of Natural History, New York. Bibliography published by Schuh and Herman (1988). Zetterstedt, J. W. b. Goteborg, Sweden, May 24, 1785; d. December 23, 1874. Professor, Lund Univer¬ sity, Sweden. Studied for doctorate under Retzius, a con¬ temporary of Linnaeus. Student of Scandinavian fauna; produced some of earliest faunistic treatments of area, including Fauna insectorum Lapponica (1828). Major Workers on the Heteroptera 13 3 Sources of Information Literature The present volume is designed to provide a window on the literature dealing with heteropteran biology and clas¬ sification. All text citations are organized into a list of references at the back of the book, and most of those are derived from the family treatments. The following introduces the most general literature on the Heteroptera, primarily that which is not mentioned elsewhere in this volume. General sources. Presented with only the name of a taxon, such as a family, the simplest approach to find¬ ing literature is through the Zoological Record (ZR). First published in 1864, the ZR annually indexes the litera¬ ture on systematics and other subject areas. Beginning in 1972 the Hemiptera are treated in a separate volume. Possessed by most university and specialty libraries, the ZR offers relatively easy access to the literature if one has the time to search it systematically. Beginning with the 1970 volume it is available for on-line computer searches. Earlier literature can be located using bibliographies pub¬ lished by Horn and Schenkling (1928-1929), Derksen and Gbllner-Scheiding (1963-1975), and Gaedike and Smetana (1978-1984). Biological Abstracts first appeared in 1926, reviewing the literature on a monthly basis. It is less strongly ori¬ ented toward systematics than is the ZR and covers all of biology. Therefore it is more cumbersome to use than ZR, albeit more current. Biological Abstracts is available in most university libraries and beginning in the early 1970s is available for on-line computer searches. Catalogs. Entomologists, and particularly system- atists, have long relied on systematic catalogs to aid them in understanding classifications and for gaining access to the massive literature on insects. The most basic catalogs are in the form of checklists, some are synoptic and a bit more complete, and the most thoroughgoing include sub¬ stantial numbers of references from the general biological as well as systematic literature. The only world catalog ever completed for the Heterop¬ tera was that of Lethierry and Severin (1893-1896). This once valuable work is now badly outdated and presents antiquated classifications in most groups. All other gen¬ eral heteropteran catalogs are regional in coverage. World catalogs at the family level are mentioned elsewhere in the text. One of the earliest regional catalogs was that of Os- hanin (1906-1909), also published as a checklist (1912). for the Palearctic fauna. The only other Palearctic work of similar coverage is the checklist of Stichel (1956-1962). a work that is sparely documented, although remarkably comprehensive as a listing of available names. Two North American catalogs exist—Van Duzee, 1916 as a checklist and Van Duzee, 1917 as a comprehensive catalog, works that are badly out of date. Henry and Froeschner, 1988 carries on the Van Duzee tradition, pro¬ viding synoptic coverage of the North American fauna north of Mexico. Australia, the Orient, Africa, and Central and South America are without regional catalogs, and workers must therefore refer to family catalogs for groups or go to the primary literature. Bibliographies. Two historical bibliographies exist for the Heteroptera. The Vade mecum (Oshanin, 1916) and the Bibliography of the North American Hemiptera-Heter- optera (Parshley, 1925). Both are still valuable for access¬ ing earlier literature. The recent paper by Stonedahl and Dolling (1991), “Heteroptera Identification: A Reference Guide, with Special Emphasis on Economic Groups,” lists many papers in addition to those dealing strictly with groups of economic importance, and as such serves as a valuable general reference. General biological and morphological treatments. Few general treatments have appeared on bugs. Weber’s (1930) Biologic der Hemipteren treats general biological phenomena and morphology in the Hemiptera in a way not seen in any other book. The Handbuch der Zoologie (Beier, 1938) and the Heteroptera chapter in the Trade de zoologie (Poisson, 1951) also offer excellent general treat¬ ments. The Biology of the Heteroptera (Miller, 1956a, 1971) offers some general observations and summary in¬ formation, with emphasis on the Reduviidae. China and Miller (1959) provided keys to the heterop¬ teran families and subfamilies on a world basis. Few texts deal with the group comprehensively, nearly all being regional, as for example Borror et al. (1989) with its emphasis on the North American fauna and The Insects of Australia (Carver et al., 1991). The Classification of Living Organisms (Slater, 1982) provided diagnoses for 14 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) all families, a limited number of references, and some helpful habitus illustrations. Monographs. Only the earliest works on Heteroptera pretended to be monographic on a world basis, as for example Amyot and Serville (1843). Thus, there is no up-to-date volume that treats all Heteroptera. Faunistic studies. Faunistic studies are concentrated on the Northern Hemisphere fauna. Notable in its com¬ pleteness is The Land and Water Bugs of the British Isles (Southwood and Leston, 1959), with its species- by-species treatment, keys, and habitus illustrations. More recent is Dolling’s (1991b) The Hemiptera, which deals with the Heteroptera and other hemipterans of the British Isles at the family level. The western Palearctic is covered in the Die Tienx’elt Deutschlands (Wagner, 1952, 1966, 1967), IllustrierteBestimmungstabellen der Wanzen (Stichel, 1956-1962), and Keys to the Insects of the European USSR (Kerzhner and Jaczewski, 1964, 1967), whereas the Heteroptera of Yakutia (Vinokurov, 1979, 1988) and Insects of the Far Eastern USSR (Vinokurov et al., 1988) treat the fauna of the eastern Palearctic. The serious student will wish to inquire into the several additional available treatments. The North American fauna was most recently covered by Slater and Baranowski (1978). The classic Heteroptera of Eastern North America (Blatchley, 1926) dealt with the eastern fauna in detail. Three historical studies are deserving of mention here. The Enumeratio Hemipterorum (Stal, 1870-1876) was a singular work in its time, and it is still useful in the iden¬ tification of some groups that have not been the subject of subsequent faunistic studies, such as the Coreidae. The Biologia Centrali Americana (Distant, 1880-1893; Cham¬ pion, 1897-1901) and the Fauna of British India (Distant, 1902-1918) dealt with the fauna of their respective re¬ gions in a way that has never again been seen for a tropical area. Although now out of date in terms of classification and nomenclature, both are extensively illustrated and are still useful for identification of members of these faunas. Collections The museums of the world are repositories for collec¬ tions made from the time of Linnaeus until the present day. We describe below the holdings of some of those in¬ stitutions that have particularly significant collections of Heteroptera. Information is available in Horn and Kahle (1935-1937), Sachtleben (1961), and Horn et al. (1990). It might be noted that in the United States most collec¬ tions segregate holotypes from the remaining material, whereas many of the major European collections store holotypes and other material together. Many universities and other organizations maintain im¬ portant regional collections and should be contacted by anyone interested in revisionary or faunistic studies. The following are notable in the United States and Canada: Cornell University, Ithaca. New York; Oregon State University, Corvallis; University of California, Berkeley; Texas A&M University, College Station; the J. A. Slater Collection, University of Connecticut, Storrs; University of Michigan Zoological Museum, Ann Arbor; Florida State Collection of Arthropods, Gainesville. Carnegie Museum, Pittsburgh; and University of British Columbia. Vancouver. A number of European museums not listed below also contain valuable material, including those in Berlin, Geneva. Leiden, Munich, and Oxford. Finally, any study of a regional nature will almost certainly bene¬ fit from consulting the collections maintained by local institutions. North America American Museum of Natural History, New York. Rich in relatively modem collections, but with limited historical material. Incorporates collections amassed by Pedro Wygodzinsky, Randall Schuh, much material col¬ lected or acquired by J. A. Slater, and the collection of Rauno Linnavuori (including most types), as well as the Heteroptera formerly deposited in the Museum of Com¬ parative Zoology, Harvard University, including Meyer- Diir material. Strongest in the New World, but also with substantial material from New Guinea and Africa as well as other areas. Biosystematics Research Centre, Agriculture Canada, Ottawa. A collection with strong holdings from North America, including Mexico, with particular emphasis on the Miridae of Canada. Also occasional series of exotic material. Bishop Museum, Honolulu. The single most impor¬ tant repository for material from the western Pacific, including Hawaii. Contains unparalleled collections from New Guinea, Borneo, and the Philippines. Much un¬ worked material. Strongest in groups that can be collected through the use of lights. California Academy of Sciences, San Francisco. Con¬ tains the collections of E. P. Van Duzee from the western United States (including nearly all of his types), 1. LaRi- vers, B. P. Bliven, and the types of R. L. Usinger. Particularly rich in the Miridae. National Museum of Natural History, Smithsonian Institution, Washington, D.C, Extensive holdings in nearly all major groups, with special strength in the North American fauna. Repository for the collections of P. R. Uhler, C. J. Drake, H. H. Knight, R. A. Poisson (part), J. C. M. Carvalho (part), and J. T. Polhemus (although still maintained by him). Rich in types. Collections origi¬ nally begun by the U.S. Department of Agriculture and still maintained in close association with that organiza¬ tion. Sources of Information 15 Snow Entomological Museum, University of Kansas, Lawrence. Possibly the single most important collection of aquatic and semiaquatic Heteroptera. including numer¬ ous types, amassed by H. B. Hungerford. Also contains significant material in other groups, including the collec¬ tions of J. R. de la Torre-Bueno and P, D. Ashlock. Europe Lund University Museum of Zoology and Ento¬ mology, Lund, Sweden. Extensive collections from Cen¬ tral America, West Africa, South Africa, and Sri Lanka. Musee Royal de I’Afrique Centrale, Tervuren, Bel¬ gium. Repository for collections from the former Bel¬ gian Congo, especially those involving the work of Henri Schouteden. Important material also present in the Insti- tut Royal de Sciences Naturelles de Belgique, Brusselles. Museum National d’Histoire Naturelle, Paris. Impor¬ tant collections for continental Europe and the former and present French colonies. Alone among most world class collections in maintaining the physical integrity of collec¬ tions of early workers, including, for example, those of Bergevin, Dufour, and Puton, with many types. National Museum, Prague. Collection assembled largely through efforts of Ludvik Hoberlandt, with exten¬ sive holdings from Czechoslovakia, the Balkans, Turkey, and Iran. Natural History Museum, Budapest. One of the most important European repositories, containing much ma¬ terial worked by G. Horvath. The Natural History Museum, London. Formerly British Museum (Natural History), containing collections on which Fauna of British India and Biologia Centrali Americana were based. Extremely rich in types, number¬ ing in the many thousands. Most species represented by short series or by types alone. Modem collections of Het¬ eroptera limited, but with significant material collected by Dennis Leston in Ghana. Naturhistorisches Museum, Vienna. A historical col¬ lection, containing comparatively large numbers of types, including those of Signoret and some of Stal. Swedish Museum of Natural History, Stockholm. Re¬ pository of the collections studied by Carl Stal. Extremely rich in type material, but with limited recent acquisitions. University Zoological Museum, Helsinki. One of the three or four most important historical collections of Het¬ eroptera, containing many types and other material of O. M. Reuter, B. Poppius, and J. R. Sahlberg. Particu¬ larly rich in Miridae. Zoological Institute, St. Petersburg. The national in¬ sect collection of the former USSR. One of the truly great collections of Heteroptera, developed by A. N. Krritshenko, containing material of most of the important Russian heteropterists and long series of many species. Zoological Museum, Copenhagen. Repository for col¬ lection of J. C. Fabricius. Contains much material in the Gerromorpha resulting from the work of N. M. Ander¬ sen. Zoological Museum, University of Hamburg, Ham¬ burg. Repository for the collection of Eduard Wagner. Central and South America Departamento de Entomologia, Museo de La Plata, La Plata, Argentina. Repository for most Carlos Berg types. A general Heteroptera collection of limited scope. Institute de Biologia, Universidad Autonomo de Mexico, Mexico, D.F. A recent collection amassed pri¬ marily through the efforts of Harry Brailovsky. Contains a broad representation of the extremely diverse Mexican fauna, including much unworked material. Museo Argentine de Ciencias Naturales “Bernardino Rivadavia,” Buenos Aires. Repository for the Nepomor- pha collection of Josd de Carlo, with a limited amount of other material, including a small number of Carlos Berg types. National Museum of Natural History, Rio de Janeiro. Repository for many of the types and other material of Jose Carvalho. Africa National Collection of Insects, Plant Protection Re¬ search Institute, Pretoria. Probably the most important collection of Heteroptera on the African continent. Con¬ tains much material, although often unworked. Other South African institutions of significance include the South African Museum (Cape Town), Natal Museum (Pietermaritzburg), and Transvaal Museum (Pretoria). The Orient Department of Biology, Nankai University, Tianjin, People’s Republic of China. The most extensive and well-organized collection of Heteroptera in China. Entomological Laboratory, Kyushu University, Kyoto, Japan. Repository of types and other material studied by T. Esaki, S. Miyamoto, and co-workers. Institute of Zoology, Academia Sinica, Beijing. A relatively large but mostly unworked collection of Het¬ eroptera from China. National Science Museum (Natural History), Tokyo. Repository of Collections of M. Tomokuni and others. Australia Australian National Insect Collection, Canberra. One of three collections in Australia containing significant holdings of Heteroptera (others are South Australian Mu¬ seum, Adelaide, and Queensland Museum, Brisbane). 16 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) 4 Collecting, Preserving, and Preparing Heteroptera Heteroptera occur in a wide variety of habitats and there¬ fore are most effectively collected through the application of a diversity of methods. The following section briefly describes collecting equipment, techniques, and meth¬ ods for mounting and preservation. Explanations of tech¬ niques and instructions on how to construct equipment can be found in most general entomology texts. Collecting Equipment and Techniques Aquatic nets. As with any group of aquatic insects, water bugs are best collected with a net designed spe¬ cifically for the task. A net with a relatively long sturdy handle, a heavy hoop, and a heavy, coarse-mesh bag will usually work best, especially in larger bodies of water. For collecting in shallow water, or in confined areas such as rock pools, and among aquatic vegetation, a metal or plastic strainer is often most effective. Many stream¬ dwelling species, particularly in the tropics, can be found only by disturbing the marginal ground and vegetation and then collecting the drifting debris and bugs in a net or strainer. Aspirator. The aspirator is probably the single most effective tool available for capturing small Heteroptera. It can be employed when collecting from a light trap sheet, beating net, sweeping net, when sorting litter by hand or collecting directly in the litter, and in many other situations. It should not be used when collecting around carrion, dung, bat guano, and similar substrates. Beating. For groups that occur on woody vegetation, sweeping is a relatively ineffective method of collection, although often used. A preferable method employs a beat¬ ing sheet or net. Beating sheets usually consist of a piece of canvas stretched on a lightweight frame; an umbrella with a short handle and a light-colored covering can also be used. They offer the advantage of considerable surface area, but for groups such as the Miridae, which often fly soon after they are dislodged from the foliage or other plant parts, many specimens will escape before they can be captured. An alternative, compact, and often more effective de¬ sign is a shallow oval net made of light or dark material and fitted with a short handle. When rapidly flying species are knocked into the net, they generally fly and alight on the sides of the net first and can be more readily captured than from a large flat sheet. Berlese funnels. Funnels provide one of the best meth¬ ods of extracting Heteroptera from forest litter. Although litter (or litter concentrated by a sifter) can be collected and examined by hand on a white sheet or in a yellow pan (or similar substrate), many heteropterans will be difficult to see (e.g., Dipsocoromorpha, small tingids) or capture (fast running or flying species). Funnels are used most effectively by first concentrating the litter through the use of a concentrating sifter and then placing the residue in the funnel to be processed for a day or two, depending on the amount of moisture in the soil-litter layer. Funnels may be metal and stationary or built of fabric and easily transportable. Canopy fogging. Use of pyrethrin mist blowers to dislodge insects from the forest canopy has become a standard technique among workers in the tropics. Large numbers of heteropterans have been discovered using this approach. Flight intercept traps. These are of two types. Malaise and pan traps. They are not the most effective meth¬ ods for collecting large numbers of Heteroptera, but are more commonly and effectively used for Diptera and Hy- menoptera. Nonetheless, certain groups will be difficult to find by other means, and some taxa or morphs, such as winged Vianaidinae (Tingidae), have been collected only by this method. Killing bottles. Many groups of Heteroptera can be collected directly into 70% ethanol, probably the sim¬ plest and safest approach. Others, particularly the Miridae and some of the Pentatomoidea with coloration derived from plant pigments, are better killed by other means. Two obvious choices are ethyl acetate and potassium or ^sodium cyanide. The former works well for nearly all het¬ eropterans, has the advantage of being relatively safe to use, and keeps the specimens moist and relaxed during the time they are in the killing jar. Ethyl acetate bottles always “sweat” to a certain extent, and therefore, espe¬ cially for specimens of Miridae and other tiny cimicomor- phans, such as Microphysidae and Cimicoidea, cyanide is the preferred killing agent. One must use great care with Collecting, Preserving, and Preparing Heteroptera 17 cyanide because it is a lethal poison; also, if pigmented specimens are left in cyanide for long periods they will change color. Specimens can be carefully layered in “cel- lucotton” packing material or pinned directly from the killing bottle. Lights. Many winged insects—and the Heteroptera are no exception—can be collected most easily through the use of lights or light traps, particularly sources con¬ taining a certain amount of ultraviolet. Self-starting mer¬ cury vapor lamps are e.specially effective. Groups such as the Dipsocoromorpha, which are difficult to see under the best of conditions, can often be easily collected at lights; the Cydninae (Cydnidae) frequently come to lights in large numbers in the tropics but may be virtually im¬ possible to locate by other means. Many Heteroptera are crepuscular, so the light must be set up and running just at sundown. However, some species, particularly of Reduviidae, often appear later in the evening. Collecting aquatic heteropterans at lights requires the use of a trap or a sheet arranged in such a way that when the insects fly to the light but are unable to cling to the vertical surface, they are not missed when they fall to the ground. Many interesting taxa can be easily overlooked because of their inability to grasp the surface of the sheet or light trap. Pitfall traps. Burying a plastic container so that the top is flush with the soil level and placing a centimeter or two of ethylene glycol in the bottom is an effective way to capture many ground-dwelling species. In areas of high rainfall, it is imperative that the traps be changed frequently or that a “roof” be provided to keep the trap from filling with water. Sweep nets. Many groups of Heteroptera occur on the foliage of herbaceous plants or grasses and can be effec¬ tively collected using a typical insect net with a sturdy bag. Mounting and Preservation Techniques Pin mounting. Many larger heteropterans can be pinned directly. In tropical environments only stainless steel pins should be used. Members of the Pentatomoidea (and others with a large scutellum) are usually pinned through the right side of the scutellum. In taxa with a small scutellum (such as many Reduviidae), the pin many be run through the right hemelytron anteriorly or placed near the posterior margin of the pronotum. Card mounting. Gluing specimens on small rectangu¬ lar cardboard mounts is the traditional method of prepa¬ ration among European workers. This approach has the advantage of protecting the specimens from breakage and allows for attractive arrangement of the appendages. It has the disadvantage of obscuring the ventral surface of the bug, including the labium and bucculae, and makes study of the pretarsus, and at times the genitalia, diffi¬ cult. The use of water-soluble adhesive makes removal of specimens relatively easy. Point mounting. Most American workers use “card points” (small elongate triangular mounts), for which punches are available in a variety of sizes and shapes for preparation of smaller specimens. This method does not protect the specimens directly (as card mounting does) although the locality and identification labels generally do. It also does not obscure the ventral body surface or the pretarsi of the specimen. As with card mounting, the use of water-soluble (or at least softenable) adhesive simplifies removal of specimens, should the need arise. Most workers follow the convention of placing the point on the right side of the specimen. If there is a question about whether a specimen should be point mounted or pinned directly, we recommend point mounting whenever possible. Ethanol preservation. Ethyl alcohol (70%) is an ex¬ cellent short- and long-term preservative for nearly all groups of Heteroptera, with the obvious exception of the Miridae (which will shed their legs) and species whose coloration is derived from plant pigments. For best pres¬ ervation, alcohol should be changed a few hours after col¬ lecting. Members of the Enicocephalomorpha and Dip¬ socoromorpha should be collected directly into alcohol, and are best stored that way, until special preparation such as slide mounting or for use on the scanning electron microscope (SEM) is required. Ultrasonic cleaning. Some heteropterans, including the Gerromorpha and Leptopodomorpha, have a dense vestiture and often become covered with soil particles or other debris during the collecting process. These and other dirty specimens often benefit greatly from use of the ultrasonic cleaner before mounting. Specimens can be placed in some dilute household ammonia or laboratory detergent for removal of debris and then returned to alco¬ hol after removal from the ultrasonic cleaner to remove water and facilitate drying. The ultrasonic cleaner can also be used with great success to clean most specimens for SEM observation. Critical point drying. This process is beneficial in the preparation of delicate specimens from alcohol (most Enicocephalomorpha, Dipsocoromorpha, Microphysi- dae, Cimicoidea, and many nymphs), in that body shape is retained, whereas when such specimens are simply air dried they collapse. The process requires specialized equipment, a carbon dioxide tank, and anhydrous alco¬ hol. Slide mounting. Certain groups of Heteroptera can be studied most efficiently when slide mounted whole or in part, for example, small Enicocephalomorpha, most Dip- 18 TRUE BUGS OF THE WORLD (HEMiPTERA: HETEROPTERA) socoromorpha, and some Cimicomorpha (notably Cimi- cidae and Polyctenidae). All of these groups are best collected directly into alcohol, if possible, in anticipation that slide mounting might be required. Specialized Preparation and Observation Techniques Scanning electron microscopy. Since the mid-1970s the use of the scanning electron microscope has become commonplace in the study of insects. The instrument is perfectly suited for the observation of details of the exo¬ skeleton. Air-dried specimens may be mounted whole or dismembered. Material stored in alcohol may benefit from critical point drying in that specimens should be as dry as possible. Dissection and preparation of genitalia. Techniques for the study of male and female genitalia, including in¬ flation of the phallus in some taxa, have been described in the literature by many authors (e.g., Slater, 1950; Kelton, 1959; Ashlock, 1967). The abdomen of dried specimens must first be softened with dilute alcohol or other suit¬ able relaxing fluid so that the pygophore can be teased from the abdomen or the abdomen removed entirely. The muscle tissue can then be removed by hydrolysis in potassium hydroxide (5-10% solution). The prepa¬ ration should then be washed in dilute acetic acid and distilled water, dehydrated in alcohol, and set up for ob¬ servation. Minute structures will require slide mounting; temporary mounts in lactophenol, glycerine, or glycerine jelly are often effective and allow for easy reorientation of complex structures, after which the specimens can be permanently stored in glycerine. Permanent slide mounts are optically better and allow for observation at very high magnification, but preclude reorientation. Carayon (1969) described a technique for staining with chlorazol black, an approach that allows visualization of delicate chitinous structures that would otherwise be vir¬ tually impossible to see. Great care should be taken when using this chemical because of its recognized toxicity. Collecting, Preserving, and Preparing Heteroptera 19 5 Habitats and Feeding Types No other major group of insects successfully utilizes such an enormous array of different habitats as do the Heterop- tera. They live as parasites of birds and mammals, feed on all parts of seed plants—and a very few ferns—from roots to pollen grains, feed on the mycelia of fungi, prey on other arthropods, live in spider and embiopteran webs and in the water and on its surface; a few species even occupy the open ocean. Only in their limited ability to burrow into woody tissues or to internally parasitize other organisms are heteropterans more limited than insects in some other orders. This diversity is especially remarkable when one realizes that in numbers of species the Heterop- tera constitute one of the smallest of the “major” orders or suborders of insects. Their numbers dwindle into in¬ significance as compared with the 300,000 or so species of beetles. Yet this relatively small monophyletic group of approximately 38,000 described species must be consid¬ ered one of the most successful of the Exopterygota, and their habitat diversity suggests a very long evolutionary history. Heteropterans are phytophagous, predatory, or hema- tophagous. Significant numbers of representatives of each type are known, and it is this diversity that has led many authors to comment on the success with which the Heteroptera have exploited the range of possible food sources—as well as habitats. The individual feeding habits of the various groups are discussed in the family and subfamily treatments. Feeding Types Phytophagy. Plant-feeding species make up the ma¬ jority of Heteroptera. As indicated by the results of phylo¬ genetic studies discussed elsewhere in this volume, the phytophagous habit has been acquired independently at least twice from predatory precursors over the long mil¬ lennia in which heteropterans have evolved. Phytophagous feeding types may also be categorized in a functional manner, these being referred to as stylet- sheath feeders and lacerate-flush feeders (see discussion in Cobben, 1978). Miles (1972) believed incorrectly that the first type was restricted to the Pentatomomorpha, based on the fact that the salivary sheath—widespread in the Sternorrhyncha and Auchenorrhyncha—was thought to be secreted in the Heteroptera only by members of that group, although it is known to occur in the Reduviidae (Friend and Smith, 1971) and probably many other fami¬ lies. In this type, the salivary glands produce a feeding cone that attaches the apex of the labium to the feeding substrate, and sometimes they also produce a salivary sheath that lines the puncture through which the stylets are inserted. It has been observed that a given species of lygaeine lygaeid may secrete a feeding cone only when feeding on seeds and using the lacerate-flush method, whereas when feeding on plant sap will produce a stylet- sheath. This observation probably applies to many other pentatomomorphans, including true predators such as the geocorine Lygaeidae and asopine Pentatomidae. Lacerate-flush feeders use the barbed apical portion of the mandibles to macerate tissues within the host (plant or animal), which are then mixed with saliva (usually containing digestive enzymes) and sucked up through the food canal. In the Pentatomomorpha plant feeding appears to be the ancestral condition, and the great majority of species possess it, albeit feeding occurs on many parts of the plant. Most Cydnidae live in the ground—sometimes at considerable depths—feeding upon roots. Several of the chinch bugs occur about the roots. Stem, leaf, and bark feeding also occur, and members of at least three fami¬ lies—Aradidae, Termitaphididae, and Canopidae—feed on fungi. Many species whose habits have been studied feed on portions of the plants where the nutritional return for a given amount of feeding is high. Thus the great majority of Lygaeoidea, Pyrrhocoroidea, and Rhopalidae feed on mature seeds and are what Cobben (1978) referred to as “lacerate flush” feeders. The pentatomoids, remaining coreoids, and Piesmatidae may feed on developing fruits or on the flowers, but most appear to gain their nourish¬ ment by extracting plant sap directly from the vascular system, particularly the phloem vessels (e.g. Maschwitz et al., 1987). These bugs produce from salivary secre¬ tions a feeding cone formed on the plant surface and in the feeding lesion. Phytophagy is by no means limited to the pentato- momorphan evolutionary line. Within the Cimicomorpha the largest family is the Miridae. While some mirids 20 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) are predaceous, probably the majority are phytophagous. These species tend to concentrate on the growing portions of the plant, chiefly the flowers, buds, and new foliage, but some have specialized on pollen. In this great family we also find species that feed on both plant and animal material. Likewise the Tingidae are entirely phytopha¬ gous, feeding more frequently on leaf tissue than do the Miridae, and usually on mature foliage. These two fami¬ lies (and the Thaumastocoridae) contain a vast number of species in an otherwise predaceous evolutionary line. The method of feeding in phytophagous cimicomorphans is quite different from that in many pentatomomorphans. They slash through the tissue with their stylets and in a sense whip a liquid from the cells in a fashion similar to that of their predatory relatives (Cobben, 1978). The Aradoidea have succeeded in utilizing the mycelia of various fungi. This ability has been accomplished by a remarkable elongation of the stylets, which lie in a vast coil within the head of the insect when it is not feeding and extend for a long distance into the slender threads of the fungus when the insect is feeding. Predation. Despite the numerous phytophagous het- eropterans, the majority of families are predaceous upon insects and other arthropods. Among these there are ex¬ quisite specializations for success. There are not only predators searching along the ground and in vegetation for prey, but also species that live in webs of spiders and Embiidina, feeding upon the small insects trapped there. Others conceal themselves in flowers, acting much as small praying mantids, seizing the bee, wasp, fly or butterfly that comes within reach. Some are very general predators, feeding upon almost anything small enough to be overwhelmed, others are highly specialized to the point of feeding only upon a single prey species. The carnivorous habit has arisen secondarily more than once in the Pentatomomorpha from phytophagous ances¬ tors. The pentatomid subfamily Asopinae is an obvious example, most species of which feed upon soft-bodied caterpillars. Similarly, in the Lygaeidae, the majority of whose species are seed feeders, carnivory has become at least the most common feeding habit in the subfamily Geocorinae. It has been shown that at least some predaceous taxa, such as members of the Reduviidae and Nabidae, will at times feed on plant substances, probably to secure mois¬ ture or to tide them over in times of shortage of suitable prey. They are not able to complete development on such a diet and die in a relatively short time (e.g., Stoner et al., 1975). Hematophagy. Perhaps the most remarkable develop¬ ment has been the ability of heteropterans to utilize the blood of vertebrates. This feeding type has arisen at least three and possibly four times independently. As in the bed bug family, Cimicidae, all of whose members feed on bats and birds, most blood-feeding Heteroptera live in the nests of their hosts. In the otherwise predominantly arthropod-feeding assassin bugs of the family Reduvi¬ idae, blood feeding is dramatically demonstrated in the Triatominae, many species of which carry the dread Cha¬ gas’ disease, which is of great importance to health in South America. In the small rhyparochromine lygaeid tribe Cleradini, blood feeding has evolved in an otherwise seed-feeding lineage. This habit reaches its greatest de¬ gree of specialization in the Polyctenidae (bat bugs), all species being permanent ectoparasites on bats. Habitat Types Inquilinous or commensal. Species in a number of families are associated with ants and ant nests. Most of these do not prey on the ants. Included are species of Enicocephalidae, Tingidae (Vianaidinae), Anthocoridae, Cydnidae, and Lygaeidae. All species of Termitaphididae live in obligate associa¬ tion with termites. Members of at least four families have perfected the ability to live in webs. Plokiophiline Plokiophilidae, phy- line mirids of the genus Ranzovius Distant, nabids in the genus Arachnocoris Scott, and some emesine Reduviidae live in the webs of spiders, usually feeding on insects ensnared there. They coexist with the spiders, and in the case of a few emesines, apparently feed on them (Snoddy et al., 1976). Embiophiline Plokiophilidae, on the other hand, inhabit the webs of the Embiidina, feeding on eggs and weak and dead individuals. Water surface-dwelling. Heteroptera of the infraorder Gerromorpha compete for the water surface only with the beetle family Gyrinidae. Members of the group have modifications, including specialized pretarsi, unwettable body surfaces, and novel communication mechanisms, that enable them to thrive in this habitat. Aquatic. Only the Heteroptera and Coleoptera have successfully adapted to a true aquatic existence in the adult stage. In the Nepidae and Belostomatidae air is breathed directly from the atmosphere through a “si¬ phon,” through the use of an air bubble captured on the venter in the Notonectidae, Pleidae, Helotrephidae, and some Naucoridae (as in the Hydradephaga), and through the use of a plastron in certain Naucoridae and all Aphelo- cheiridae, a condition found elsewhere only in dryopoid beetles. Intertidal. A number of species of Leptopodomor- pha live only in the intertidal zone, the most specialized among them being Aepophilus bonnairei Signoret with its plastron respiration and greatly reduced compound eyes and hemelytra. Habitats and Feeding Types 21 Competition Much has been written concerning competition, or means of avoiding it, between closely related species. In the Het- eroptera such subtle interactions have not been studied for most species, but there are some striking examples. Blakley (1980) studied the relationships of two species of the lygaeid genus Oncopeltus Stal in the West Indies. He found that Oncopeltus fasciatus (Dallas) is able to survive only upon plants of the milkweed Asclepias curassavica when seeds and pods are present, whereas O. cinguUfer Stal is able to pass its entire life cycle on the vegetative tissues of the plants. Thus while O. fasciatus has greater flight ability, it is more restricted in local distribution be¬ cause of the limitations placed upon its available breeding sites. A striking case of two species not at all related taxo- nomically but feeding upon some of the same host plants also involves a species of Oncopeltus. Blakley and Dingle (1978) reported that on the West Indian island of Barba¬ dos the monarch butterfly has completely eradicated the milkweed bug O. sandarachatus (Say). What appears to have happened in this case was that the monarch built up large populations on the introduced milkweed Calo- tropis procera, a plant not suitable for O. sandarachatus. Since the monarch also feeds on A. curassavica, the large populations eliminated the latter plant, upon which the milkweed bug is dependent. Hamid (1971b) studied three Cymus spp. that live on the identical sedges and rushes in the same habitats. Their life cycles are largely nonoverlapping so that when one species is adult the others are either in the egg stage or early nymphal stage and thus utilize the hosts in seasonal succession. It is known that in the extensive fauna of the ground¬ living Lygaeidae that feeds on fallen figs there is also a succession of species: from those that appear as soon as the fruits fall to the ground, through a changing fauna, to finally a group of minute species that appear after most of the seeds have been “eaten” and search out seeds that have fallen into cracks and crevices and thus, in a sense, “scavenge” for the leftovers. Taylor (1968) and Streams and Newfield (1972) studied the habits of backswimmers (Notonectidae) that occur in the same ponds and discovered that they divide the available niches both in time and space. Stonedahl (1988a) found that in the mirid genus Phy- tocoris Fallen a number of species occur simultaneously on the same host plant. In most such cases resource partitioning occurs, wherein different species occupy dif¬ ferent parts of the host plant such as the cones, foliage, branches, boles, and bark. These preferences may be in part due to the different degrees of phytophagy and pre¬ dation in different species. Cooper (1981) found that the bark-inhabiting species Phytocoris neglectus Knight and P. nobilis Stonedahl appeared to be occupying the same part of the host plant. Abies procera, at the same time in Oregon. However, they are actually temporally isolated and occur together for only a brief period in mid-August. Dispersal It is apparent that to succeed, populations of insects must be able to move from an unsuitable habitat to a suitable one. For the Heteroptera this ability is a remarkable phe¬ nomenon and involves evolutionary time adaptations as well as contemporary ecological ones. The evolutionary consequences of dispersal or the lack of need for it are discussed in Chapter 6. Here we will be concerned with the ecological adaptations. Dispersal and dispersion are terms that are used loosely in the literature. Perhaps it is proper to restrict the term dispersal to the extension of the range of a species across a barrier that is unsuitable for it to live in or on. If this is done, then the movement of individuals within a popula¬ tion can be considered dispersion and does not imply an increase in range. One must recognize of course that the actual range of a given species is not constant and that there is continual increase and decrease along the margins of a range depending upon better or worse conditions for the population at any given time. Nevertheless, species do have ranges that can be stated rather accurately in a general sense. One has only to go to a light at night in summer or in the warm tropics to see that heteropterans are dispersing almost constantly. Indeed all we know of many species that live in very specialized or inaccessible habitats is that they actually exist, because they have been collected at lights and thus far their actual habitats are unknown. Monteith (1982b) reported a remarkable aggregation of insects in northeastern Australia. Here the dry season is very harsh, and insects migrate to the pockets of mon¬ soon forest. He noted four species of Heteroptera (two alydids, a coreid, and a plataspid), some of which occur in dense clusters comprising large numbers of individu¬ als. Monteith believed that these bugs do not feed and that the aggregating behavior is an example of group re¬ inforcement of natural chemical defenses, in the case of the Heteroptera of course the scent-gland secretions. He reported that when sudden massed flights were elicited by disturbance of aggregations of Leptocorisa acuta (Thun- berg) and Coptosoma lyncea Stal flight was accompanied not only by discharge of the glands but also by a sud¬ den loud buzzing similar to that of a disturbed paper- wasp nest. This combination yielded considerable fright reaction in humans coming suddenly on an aggregation. 22 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) 6 Wing Polymorphism Biologists usually equate the success of insects with their ability to fly. Yet, when flight is no longer advantageous, insects frequently lose this ability and often the wings themselves. Nowhere in the insect world is this more evi¬ dent than in the Heteroptera. In family after family the ability to fly has been lost. The usual result is the reduc¬ tion, modification, or loss of the wings. In some cases flight loss is less evident; the wings appear normal, but flight muscles become reduced after a migration period, as in some species of Dysdercus Guerin-Meneville (F. J. Edwards, 1969; Dingle and Arora, 1973). There are many reasons why heteropterans have lost the ability to fly. In the ectoparasites (Cimicidae, bed bugs; Polyctenidae, bat bugs), termitophiles (Termitaphi- didae, some Lygaeidae, and others), and myrmecophiles, flightlessness is obviously associated with the adaptation to living on the bodies of warm-blooded vertebrates or in the nests of termites or ants. Darwin (1859) considered the problem of why species living on the ground, under bark, or on the water sur¬ face should develop flightless morphs. He interpreted the phenomenon in what today’s jargon calls the “energy budget”; that is, an individual, or a population, has a given amount of energy and if that energy is not needed for “making” wings, it can be better used for other pur¬ poses such as increased reproduction. Thus, if it is not an advantage to be able to fly, individuals in the population that do not utilize energy in developing wings capable of flight will be favored, and eventually the entire population will become flightless. One sex may become flightless in some species while the other will remain fully capable of flight. This phe¬ nomenon is not well understood experimentally, but in some myrmecomorphic Miridae, where only the female is strongly mimetic (and flightless), fewer eggs need to be produced because of the reduction in mortality (Mclver and Stonedahl, 1993). In many semiaquatic heteropter¬ ans, populations will be flightless during most of the year but have a winged generation during one season. Heteropterans may be entirely wingless or the wings may appear normally developed but flight muscles art reduced. In a number of ground-living species the fore¬ wings become shell-like and closely resemble the elytra of beetles, an adaptation that reduces water loss in arid habitats (Slater, 1975). The different degrees of wing modification may be categorized as follows. These descriptions apply most easily to the Panheteroptera, which have a distinct clavus, corium, and membrane in the forewing (see Slater, 1975). Aptery: Fore- and hind wings completely absent. Microptery: Forewings reduced to minute leathery pads covering, at most, only the base of the abdomen. Hind wings either minute flaps or absent. Staphylinoidy: Forewings covering no more than basal half of abdomen; clavus and corium indistinguishably fused; membrane, if present, only a narrow marginal rim; distal ends truncate. Hind wings reduced to flaps or absent. Brachyptery: Forewings reduced, not covering abdomi¬ nal terga 6 and 7. Clavus and corium fused or not, but elements recognizable. Hind wings reduced but usually not flaplike. Coleoptery: Forewings shell-like, with clavus and co¬ rium elongated, usually fused and meeting evenly along midline; membrane of forewing reduced. Hind wing absent or reduced to small flaps. Anterior abdominal terga often membranous. Submacroptery: Clavus and corium distinct and usually not reduced. Membrane of forewing reduced leaving pos¬ terior abdominal segments exposed. Hind wings either slightly reduced or elongate. Macroptery: Clavus and corium distinct, membrane well developed. Hind wing elongate. Caducous (= deciduous): Wings fully developed, but broken off or eliminated at some time during adult life. A condition commonly found in Enicocephalomorpha, Mesoveliidae, Veliidae, and occurring in some Aradidae. There is relatively little experimental evidence for the mechanisms producing flightless morphs. The one feature that seems to be common to most of the flightless species is relative permanency of habitat (Brown, 1951; South- wood, 1962; Sweet, 1964; Fujita, 1977; Slater, 1977; Vepsalainen, 1978). Many islands also contain a high proportion of flightless morphs (see Darlington, 1971, 1973 for general discussion). Wing reduction in the Nepomorpha was long thought not to exist except where there was an adaptation for underwater respiration. Several species of the naucorid Wing Polymorphism 23 genus Cryphocrkos .■ignoret display wing reduction (Sites, 1990). In ( nungerfordi Usinger in Texas, a species that lives in permanent streams. Sites found that in a sample of 790 adults only six were macropterous and one submacropterous. The others (98%) were strongly brachypterous, the wings being what Slater (1975) termed staphylinoid, reaching only over the basal third of the abdomen and truncate or subtruncate along the distal mar¬ gins with the membranes absent. J. T. Polhemus (1991a) described the Neotropical naucorid Procryphocricos per- plexus Polhemus as having the wings reduced to small pads that reach only the second abdominal tergum. Both of these naucorid genera must have plastron respiration, allowing them to stay rcrmanently submerged. The literature shov. , however, that several genera of Nepidae also have reduced hind wings; in one case these wings are leathery. These nepids breathe atmospheric air by means of the long terminal appendages. Brown (1951) observed that, in Britain, Corixidae that were brachyp¬ terous were confined to permanent habitats. He also pointed out that species of water boatmen normally found in temporary habitats migrated a great deal, whereas those in permanent habitats did not, and he implied that brachyptery was alvvuys associated with permanent habi¬ tats. Thus, it seems that adaptation to underwater respi¬ ration is not necessary for flightlessness to evolve in the Nepomorpha. Slater (1977) noted that permanency of habitat may be thought of in an evolutionary-time sense as well as in a shorter ecological-time sense. He observed that in old stable areas, such as the southern Cape region of South Africa and southwestern Australia, the majority of ground-living Lygaeidae have a flightless morph, and that in such areas the degree of wing modification is greater than in less stable areas. Similarly, Malipatil (1977) noted that the New Zealand ground-living lygaeid fauna is com¬ posed almost entirely of members of the tribe Targaremini and that 95% of the species are flightless, with coleoptery being the predominant condition. He observed high pro¬ portions of flightlessness in several other orders of New Zealand insects. The genetic or environmental factors involved in species with wing polymorphism are still understood for only a very few species. Solbreck (1986) showed that the ground-living lygaeid Horvathiolus gibbicollis (Costa) from the Mediterranean region possessed both macrop¬ terous and submacropterous forms. In the latter the fore¬ wings are about two-thirds the length of those in mac¬ ropterous forms and the hind wings less than one-third the length of those of macropterous forms. He found that each morph bred true to wing length when reared under variable density, food, and temperature conditions for several generations. All FI offspring between crosses of the two morphs were submacropterous. In the F2 generation approximately 25% were macropterous and 75% submacropterous, implying a monogenic control of the wing form. His study also demonstrated that short¬ winged morphs reach adulthood more rapidly than do macropterous forms and have a distinctly shorter adult preoviposition period. Solbreck’s study suggested that a simple gene modification may be involved and offered a possible explanation for why macropterous forms will persist in populations for a long period of time. The ability to develop flightless populations, but also to re¬ tain a proportion of fully winged individuals, gives such populations the ability to make maximal use of a limited food supply for which they may be competing and still retain the ability in the population to colonize new food sources at some distance from the original population. Klausneret al. (1981) studied a population of brachyp¬ terous Oncopeltusfasciatus (Dallas) on Guadeloupe. They found brachyptery to be caused by a single recessive Men- delian unit and that neither temperature nor photoperiod had any effect on the expression of the brachypterous trait in the homozygous short-winged genotype. Southwood (1961) believed that wing polymorphism in Heteroptera was under hormonal control. He felt that the short-winged condition resulted from an excess of juve¬ nile hormone and thus wing reduction was essentially a neotenic condition. Southwood noted that more species living at high altitudes were flightless than were species of the same groups at low altitudes. He suggested that at high altitudes low temperatures acted on the hormone bal¬ ance or that a longer time was spent in the nymphal stage and therefore the insect was under the influence of the juvenile hormone for a longer period of time. Although this theory may account for the occasional production of short-winged morphs, it cannot account for most species with wing reductions. First, while low temperature—as shown by Brinkhurst (1961)—certainly influences wing development in some taxa, it is not clear that this is due to excess juvenile hormone. Second, the short-winged adults are not neotenous, as the short wing is not the wing pad of the nymph but morphologically an adult wing. Third, the presence of reduced wings in many terres¬ trial insects living at high altitudes may be the result of living in areas of long-term habitat stability. Such insects are basically in stable environments, for when conditions change in montane habitats the insect populations need move only a relatively short distance up or down to find themselves in the same habitat they were in before. By contrast, were the same variation in conditions to take place on a level surface, it would require movement of an insect population hundreds of miles to find ecologically similar conditions. An appreciable number of flightless heteropterans live 24 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) in extremely hot situations such as deserts, tropical savan¬ nahs, and beaches. The last may not at first glance appear to be a stable habitat, but it actually is, in the same sense as mountains are, in that insects living in beach habitats must move only a short distance to remain in the same habitat as the beach expands or retreats over a period of time. The most important consideration, we believe, is to understand the genetic system at work and to realize that what is a stable habitat for one group may not be for another. Heteropterans living upon the surface of the water, especially in temperate habitats, face a somewhat dif¬ ferent ecological challenge. Brinkhurst (1961) and Vep- salainen (1978) showed that gerrids and veliids living in rivers and streams (relatively permanent habitats) con¬ sist of populations that are univoltine and predominantly apterous. Univoltine populations living in ponds, how¬ ever, are usually long-winged. In bivoltine species the situation is much more complicated. Most species of gerrids studied by Brinkhurst are polymorphic. The over¬ wintering generation is chiefly long-winged, and summer generations of short-winged individuals are the offspring of long-winged overwintering mothers. Brinkhurst found that wing development was under the control of a single gene and that short-winged homozygotes were lethal but that short-winged heterozygotes were under environ¬ mental influence so that expression of a “switch gene” came into play. For example, when he crossed short¬ winged with short-winged forms he obtained either all long-winged forms or 2.5:1 ratio of short-winged to long¬ winged forms depending upon the temperature. Vepsalai- nen (1978), however, indicated that the most important factor in determining wing length was photoperiod. Not all semiaquatic bugs behave under the model de¬ scribed above. For example, Muraji et al. (1989) showed that in the veliid Microvelia douglasi Scott, which lives in relatively permanent habitats as well as ephemeral ones, the percentage of macropterous forms declines dramati¬ cally in autumn, and the populations are then almost en¬ tirely flightless. Temporary habitats such as rice paddies are apparently recolonized each year, and the flightless fall generation hibernates only near permanent water. These authors found that the strongest determinant for an increase in the proportion of macropterous forms is density, but that this factor is modified by photoperiod, temperature, and abundance of food. Muraji and Naka- suji (1988) showed in a comparative study of three species of Microvelia Westwood, that the species that lived in the most permanent habitats developed a macropterous morph only when populations reached high densities, and it was the slowest of the three species to respond by production of macropterous forms as density increased. Zera and Tiebel (1991) studied the effects of photoperiod on wing length in the pterygopolymorphic gerrid Limno- porus canaliculatus (Say). They found that reduction in photoperiod had a strong positive effect on the production of the long-winged overwintering morph. Brinkhurst’s belief that wing polymorphism is under the control of a single gene was challenged by Andersen (1973), who discussed the condition in two species in Denmark, noting that in bivoltine species there is a loss of wing muscles in the overwintering generation after a flight period in the spring. Vepsalainen (e.g., 1971a, b) demonstrated that, at least in temperate-latitude species, the development of short- and long-winged forms was due to changes in photoperiod. Andersen (1973) reviewed the evidence in detail and presented experimental data on changes in fifth-instar nymphs in several water striders that are correlated with pigmentation, aspects of repro¬ ductive dimorphism, and diapause. Andersen (1993) used cladistic techniques to corre¬ late wing polymorphism and habitat type in temperate- latitude Gerridae. He found that species with the ancestral pattern of wing polymorphism occupy the most stable habitats. He therefore hypothesized that the ancestral type of habitat was relatively permanent (streams, permanent ponds, lakes) and the ability of water striders to colo¬ nize less stable or temporary water bodies had evolved more recently. Andersen (1993) concluded that wing di¬ morphic or short-winged morphs are the ancestral states and that where only long-winged morphs exist it is be¬ cause the short-winged morph has been lost. He further reasoned that two distinct types of short-winged gerrids exist, one emerging from fifth-instar nymphs with distinct wing pads, the other from nymphs with reduced wing pads. He suggested that the two kinds of short-winged conditions are associated with different mechanisms of morph determination involving a polygenic system and environmental switches adapted to features of the habitat. Waloff (1983) pointed out that in arboreal species, habitat architecture may militate against the development of flightless morphs. She noted that in the British Het- eroptera 75 of the species associated with grasses or forbs show wing polymorphism, whereas only 11 of the 111 tree-living species have short-winged forms. Moreover 10 of these 11 species occur outside of the niches occu¬ pied by typically phytophagous insects (9 are predaceous and 1 lives under bark), the 11th species lives on dwarf willow and heather. She believed this may be true only of temperate populations and cited the high incidence of flightless arboreal tropical Acridoidea. There is no evi¬ dence at present, however, that tropical arboreal Heterop- tera tend to become flightless to any greater degree than do temperate ones. Waloff’s paper would be even more compelling if she had distinguished between those forb- inhabiting species that live above the ground (arboreal in Wing Polymorphism 25 the sense of Slater [1977]) and those that are geophilous for the most part. Roiston (1982a) described a strongly staphylinoid pen- tatomid from an elevation of about 1800 meters in Haiti. He assumed it was arboreal because all other members of the tribe are. If Roiston is correct, this is an example of wing reduction in a group where the phenomenon is rare. 26 TRUE BUGS OF THE WORLD (HEMIPTERA; HETEROPTERA) 7 Mimicry and Protective Coloration and Shape The questions of mimicry, aposematic coloration, and protective resemblance have created much controversy and a large literature over the years. The Heteroptera con¬ tain striking examples of each. Mimicry has traditionally been grouped into distinct types, although there are cases that are somewhat intermediate or involve more than one of the classical categories. Mimicry Batesian mimicry. An insect that is not protected comes to look like a distasteful or dangerous insect such as a wasp, bee, or ant. This phenomenon is widespread in the Heteroptera; ant mimicry alone occurs in no fewer than seven families. Mullerian mimicry. Several distasteful or dangerous species come to look alike in color or structure or both to reduce predation as the potential predator learns to avoid a given color or form pattern. This phenomenon also is widespread in the Heteroptera. Documented cases are difficult to establish experimentally, and most evidence is based upon taxa not closely related, coming to look like other species in sympatry but unlike each other in areas of allopatry. Mertensian mimicry. The model is moderately dis¬ tasteful or dangerous, whereas at least some of its mimics are lethal. This type of mimicry has not been established in the Heteroptera; but it probably occurs. Wasmannian mimicry. Mimicry in which an insect attempts to “fool” the ant with which it is associated is not yet documented in the Heteroptera, although it is common in the Staphylinidae and related beetles. Aggressive mimicry. A predator looks like a prey species to “fool” the latter and enable predation to be" enhanced. Mclver and Stonedahl (1993) note this type may occur in some species that feed on ants. This phe¬ nomenon should be studied carefully as the reduviid- pyrrhocorid mimetic situation discussed below might ap¬ pear to be such a situation, but is probably a case of Batesian mimicry. Aposematism and Mimicry Among the Miridae, Lopidea Uhler, Hadronema Uhler, and most Resthenini exhibit aposematic coloration. Mc¬ lver and Lattin (1990) studied Lopidea nigridia Uhler in detail, noting that it possessed a clumped distribution, was relatively sedentary, and fed on a plant species {Lupi- nus caudatus) known to produce several alkaloid com¬ pounds. They experimentally tested the palatability of L. nigridia, using spiders as predators, and found it to have limited acceptance as a prey item when compared with a nonaposematically colored mirid, Coquillettia insignis Uhler. Many species of lygaeids in the genera Lygaeus Fabri- cius and Oncopeltus feed on milkweed plants that contain cardenolides. Such species have been shown to be ex¬ tremely distasteful to vertebrate predators. It has long been believed that brightly colored insects living in simi¬ lar habitats would be protected by resembling these showy red and black, and presumably aposematically colored, milkweed-feeding insects. The problem with this concept has always been to ac¬ count for the great similarity of many of the presumably protected species. For example, in Africa some species of Spilostethus Stal (a genus closely related to Lygaeus) are almost carbon copies of one another in color pattern. Similarly several species of Oncopeltus are also almost identical in color pattern, and indeed TxLwy (1990b) marshaled evidence from study of the west Palearctic fauna to indicate that similar color patterns have evolved many times. Such close resemblance suggests that greater protection from predators is obtained the closer the re¬ semblance of one group of species is to another (i.e., less learning needed by the potential predator) and that, although bright color may afford some protection, it is only partially effective. Yet, when species from a num¬ ber of families resemble one another, several will have well-developed scent glands (supposedly rendering them distasteful), and determination of who is the model and who is the mimic awaits experimental work. McLain (1984) showed that the red and black mirid Lopidea instabile (Reuter) and the similarly colored ly- gaeid Neacoryphus bicrucis (Say) are both unpalatable to Anolis lizards when fed on the alkaloid-containing com¬ posite Senecio smallii. When fed on sunflower seeds the bugs are palatable to anoles. However, the lizards rarely Mimicry and Protective Coloration and Shape 27 attack either bug species, once they have attempted to eat a bug that had fed on an alkaloid-containing plant, thus suggesting a Mullerian mimicry complex. Sillen-Tullberg et al. (1982) demonstrated that in the case of the Palearctic lygaeid Lygaeus equestris (Lin¬ naeus), two aposematic forms must be very similar to gain full mutual protection. This study provided evidence why not only Batesian mimics, but also Mullerian mim¬ ics, and their aposematic models develop such extremely close resemblances. A striking example of what appears to be a combi¬ nation of Batesian and Mullerian mimicry occurs in the milkweed and related bugs of the family Lygaeidae and similar-appearing species in the Rhopalidae, Largidae, Pyrrhocoridae, and Reduviidae. Van Doesburg (1968) noted that many of the Dys- dercus spp. (Pyrrhocoridae) in the Western Hemisphere appear to form mimicry “rings.” He pointed out that in a given geographic area several species have a similar color pattern. In the case of Dysdercus obscuratus Distant five subspecies are recognized, four of these having adult color patterns very different from each other and each resembling another species of the genus that occurs in the same region. This case appears to be one of classic Miil- lerian mimicry, but it is complicated by the resemblance of some of these insects to species of Lygaeidae {Oncopel- tus) and to species of Coreidae and Miridae. Complex combinations of Mullerian and Batesian mimicry may be involved. Involvement of such combinations is fur¬ ther suggested by conflicting evidence in the literature concerning the palatability of members of the genus and whether the scent glands produce distasteful liquids. The relationship of the insects to potential predators has not been investigated. The nymphs of most species are bright red, conspicuous, almost certainly aposematic, and do not vary in coloration geographically, suggesting that the adult coloration has evolved through terminal addition in ontogeny. Further developments of this type occur in a number of African pyrrhocorids and some of their assassin bug predators. These cotton stainers often occur on or near malvaceous host plants in aggregations of hundreds of individuals. Stride (1956) showed that for several differ¬ ent species of Pyrrhocoridae an assassin bug of the genus Phonoctonus Stal lives in the aggregations and feeds upon the cotton stainers. Each of these assassin bugs closely re¬ sembles the particular cotton Stainer species with which it lives. Stride studied three species of Phonoctonus in West Africa, each of which was found most frequently feeding on the pyrrhocorid that it most closely resembled. How¬ ever, the assassin bugs also occurred regularly both as nymphs and adults, laid eggs with, and fed upon species of cotton stainers with which they showed lesser degrees of resemblance. Stride further found that one Phonocto¬ nus sp. was extremely variable in color, one color morph resembling a brown species of pyrrhocorid and the other a red and white species. When he fed nymphs of this re- duviid species upon both red and white and brown pyrrho¬ corids the resulting adults were always red and white. What this mimetic situation almost certainly suggests is that the predator is protected from vertebrate predation by developing a close resemblance to its aposematically colored prey. Other examples include an African assassin bug of the genus Rhynocoris Hahn, which bears red and black stripes and closely resembles the lygaeid species of Spilostethus Stal that live in the same area. A Neotropical assassin bug of the genus Zelus Fabricius is strikingly like the aposematically colored species of Oncopeltus. Although these cases have not been supported by ex¬ perimental work, it is difficult to believe that assassin bugs whose relatives are dull brown or gray would develop such exacting patterns of black and red coloration except as a mimetic resemblance. There is no evidence, as had previously been hypothesized, that these assassin bugs are exhibiting aggressive mimicry of the phytophagous heteropterans with which they are associated. Poppius and Bergroth (1921) suggested that African species of Poeantius Stal, ground-living myrmecomor- phic lygaeids, were predatory on ants and that this was a case of aggressive mimicry. They believed that the ly¬ gaeids lived in ant nests. However, the lygaeid bugs are not predators but seed feeders, and it is probable that the mimicry is simple Batesian. Many studies suggest or demonstrate that vertebrate predators, especially birds and lizards, are the principal agents involved in the devel¬ opment of mimicry. However, Berenbaum and Miliczky (1984) reported that the Chinese mantid Tenodera ardi- folia sinensis learns to avoid the toxic Oncopeltusfasciatus after several trials. The mantid regurgitates and shows signs of poisoning when cardenolides are present and will not attack the milkweed bugs when they are reared with¬ out cardenolides. Mclver and Stonedahl (1993) stated that the reduviid Sinea diadema (Fabricius) will accept nonmimetic species of Lopidea much more often than they will the myrmecomorphic Coquillettia insignis (both Miridae). Tyshchenko (1961) believed that spiders and predatory insects were at least as important as vertebrates in mimicry development. Ant Mimicry, Myrmecomorphy, and Other Insect Resemblances One of the most striking mimetic conditions found in the Heteroptera is the development of myrmecomor¬ phy. This phenomenon has developed independently not only in more than seven different families (Mclver and Stonedahl, 1993), but also in different phylogenetic lines 28 TRUE BUGS OF THE \«ORLD (HEMIPTERA: HETEROPTERA) of the same family. Furthermore, the degree of mimetic resemblance is extremely diverse, ranging from cases where even the experienced investigator in the field can¬ not always differentiate the hemipteran from the ant, to situations where in museum specimens there would ap¬ pear to be no mimicry at all. Yet the movements of the insects in the field are distinctly antlike, apparently suf¬ ficiently so to provide the potential prey species with a degree of protection when disturbed by a vertebrate (or other) predator. The Neotropical lygaeid Neopamera bilobata (Say) does not appear to be ant mimetic in collections, but in the field when a large population under a food plant is disturbed, the insects rush out in all directions with jerky irregular motions that make them difficult to dis¬ tinguish from ant species that live in similar habitats. It is our belief that such “action mimicry” will often cause a predator to hesitate for a period long enough to allow most of the lygaeids to escape. Frequently myrmecomorphy involves the loss of flight (and possibly the reverse). The most common condition is a shortening of the wings, an enlargement of the ab¬ domen with a concomitant constriction at the base, and a modification of the shape of the head. This condition may be enhanced by various color patterns, especially but not exclusively, in the Miridae by streaks of silvery hairs that break up the body shape into a three-lobed ant form. In macropterous forms myrmecomorphy is often achieved by patches of white color on a dark insect, usually involv¬ ing the forewings—but sometimes also the pronotum— which have patches of color placed so as to give the illu¬ sion of a “wasp waist,” but which still allow the insect to fly. In some of the most striking mimics the head may be modified to possess ridges along the lateral margins that closely resemble the mandibles of ants. Ant mimicry is widespread in the Lygaeidae and has developed many times in different phyletic lines. Ble- dionotus systellonotoides Reuter, a ground-living species from the Near East that occurs under plants growing on rocks, is said to sometimes occur with, and closely re¬ semble, the mimetic mirid Mimocoris rugicollis (Costa). The Pamphantini, which are often brightly colored with orange and black, are speciose in Cuba and Hispaniola, where they form mimicry complexes with beetles and spiders, all of which mimic species of Solenopsis ants (Myers and Salt, 1926). The Pamphantini, in contrast to Bledionotus, frequently occur on trees; Brailovsky (1989) described several species from the Brazilian rainforest canopy. Many Rhyparochrominae are myrmecomorphic and, because of the large incrassate forefemora, were long thought to be predaceous, at least in part on ants, but all evidence indicates strict seed-feeding habits. Some Heterogastrinae are also strikingly myrmecomorphic. Ant mimicry appears to take two basic evolutionary pathways. The first is to live in a habitat frequented by many species of ants. The mimicry often is not to any particular species but to a general antlike habitus. The second is probably less common but has the most exqui¬ site adaptations, in which a specific ant appears to be the model. Sometimes more than one species of insect or spider will mimic the same ant species. Myers and Salt (1926) noted that each of several species of Solenopsis ants in Cuba was mimicked by a spider, a beetle, and at least one pamphantine lygaeid. They commented that the mimicry was so striking that even after a period of study, when they attempted to pick a lygaeid bug mimic from a branch, they were astonished to see it drop by a thread from the branch and realized that it was a spi¬ der. Some of these Solenopsis ants are strikingly colored orange and black. In addition to the pamphantines there are two lygaeid bugs belonging to different genera that develop this striking orange and black color on Cuba, but elsewhere in the Neotropics other species of these genera are dull brown and black. Parenthetically it may be added that such information can be useful in interpreting ques¬ tions of vicariance versus dispersal for given taxa (Slater, 1988). In the case above, the lygaeid Heraeus triguttatus (Guerin-Meneville) is the only orange and black lygaeid occurring in extreme southern Florida as well as on Cuba. Since the ant model is not present in Florida the obvious conclusion would seem to be that this species has reached Florida by overwater dispersal from Cuba subsequent to its evolution of mimetic coloration. Mclver and Stonedahl (1987a, b) conducted experi¬ mental studies on myrmecomorphic species of Coquillet- tia Uhler and Orectoderes Uhler and showed that their arthropod predators were capable of learning the antlike habitus and associating it with unpleasant experiences. In some cases it is clear that myrmecomorphic mirids are associated with ants, and with a particular ant species, whereas in other cases, it is not obvious that such an association exists (see Fig. 53.2C, D). Heteropteran nymphs often mimic different species of ants as they grow (Mclver and Stonedahl. 1993). Species of the mirid genera Coqidllettia and Paradacerla Car¬ valho and Usinger—and the lygaeid genus Slaterobius Harrington—have two color morphs that resemble two different ant species. In many mirids only the female is strongly mimetic, whereas in alydids and lygaeids both sexes are mimetic. Mclver and Stonedahl (1993) believed that within given phyletic lines of orthotyline and phyline mirids the most derived taxa are myrmecomorphic and the most plesiomorphic are nonmimetic. Many genera of Alydidae contain species that are strik¬ ing ant mimics as nymphs, but most species as adults have behavioral and color patterns of wasps. This phe- Mimicry and Protective Coloration and Shape 29 nomenon may best be illustrated by an experience that we had in South Africa some years ago. We were attracted to the behavior of a species of alydid that was alighting on a sandy area in bright sunlight. When the insects landed they moved their bodies in a jerky fashion and opened the forewings to display their bright orange dorsal abdominal coloration. The movements and color pattern closely re¬ sembled the actions of sand wasps working in the same area. Interestingly, adjacent to this site was a stand of Acacia trees under which was a large population of alydid nymphs that were, as usual, strongly ant mimetic. The irony of this whole association was that upon returning to the laboratory with a sample of nymphs we discovered that among them were several adult specimens of a tiny lygaeid that so closely resembled the ant-mimetic, early- instar alydid nymphs that we had not differentiated them in the field. Possible beetle mimicry is less well understood. Many ground-living and some arboreal hemipterans develop co¬ riaceous front wings that meet down the middle of the back in a coleopteroid fashion, but it has not been shown that these bugs are beetle mimics. Some of these cases are associated with flightlessness, and it appears to be an adaptation to prevent water loss in xeric habitats. Never¬ theless, there are peculiar longitudinal markings on the shell-like covering of the scutellum of some Scutelleri- dae that greatly enhance the beetle-like appearance of the insects and do not seem to have any other protective value. Protective Coloration and Shape Protective coloration is present to some extent in almost every family of Heteroptera. Frequently this is a simple matching of the color of the insect to that of its environ¬ ment. In many Miridae that feed on flowers, the nymphs in particular are the same color as the flowers on which they develop. The occurrence of green coloration in phy¬ tophages and brown or gray coloration in bark-dwelling predators is widespread in the family. More striking is the modification of body shape to “mimic” the substrate on which the insect lives. In the South American pentatomoids of the family Phloeidae, which live on tree bark, the sides of the head, thorax, and abdomen are expanded into great flattened lobes that make the insect almost indistinguishable from the back¬ ground on which it lives. A convergent condition occurs on Borneo, where the pentatomid genus Serbana Distant, whose habits appear to be unknown, has developed simi¬ lar flattened, expanded body lobes and presumably also will prove to live on bark. Insects of several families, such as the stenodem- ine mirids, monocot-living alydids, tingids, lygaeids. predatory reduviids, and even pentatomids living among grasses and sedges develop elongate slender bodies that resemble closely the elongate slender grass and sedge stems and leaves among which they live. The lygaeid bugs of the subfamilies Cyminae and Pachygronthinae live chiefly in the seed heads of sedges. Both nymphs and adults closely resemble the shape and color of the seeds on which they feed. Some heteropterans have the ability to change color with the seasons or the places where they live to increase their protective (or hiding) ability. Species of the mirid genus Stenodema Laporte are green during the early part of the host growing season, but pale brown later as the grasses on which they feed mature and become brown. This color change can also be seen in such predators as the ambush bug Phymata pennsylvanica Handlirsch. It is green and black early in the season, but in the au¬ tumn, when it lives most frequently in the heads of yellow goldenrod {Solidago spp.), it becomes a bright yellow and black. This latter change may be considered protective coloration but it also is concealing coloration for these “sit and wait” predators, making them much less con¬ spicuous to potential prey insects. Some tropical assassin bugs exhibit this concealing pattern in color and some¬ times in structural modifications that make them virtually indistinguishable from the flowers on or in which they sit. Even more striking color modifications occur in toad bugs of the genera Gelastocoris Kirkaldy and Nerthra Say. We have observed a species of the former living along the banks of the Platte River in Nebraska, where in the same area individuals were black when on a dark substrate, tan when on a brown substrate, and speckled when on sand of white and dark grains. In Western Australia a species of Nerthra that lives in dry habitats among rock chips becomes reddish, black, or dull white depending on the color of the rocks among which it lives. To our knowl¬ edge there has been as yet no study to determine whether such striking color variations are genetically controlled or whether individuals are able to change color to match the substrate on which they live. Protective coloration also may take the form of de¬ flective patterns which are perhaps best illustrated by the white banding on the terminal segments of many ground-living heteropterans and by the occurrence of white patches near the end of the abdomen. There has been littie experimental work done on this phenomenon, but to the field collector it becomes obvious that these are deflective markings that cause the potential vertebrate predator either to miss the rapidly moving heteropteran by striking at the white marking at the posterior end, or in the case of the antennal marking striking at the area anterior to the insect’s body and at best obtaining the terminal antennal segment. These color patterns occur in 30 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) family after family of geophilous heteropterans and merit serious experimental work. The leaflike broadened and flattened antennal segments and hind tibiae of several Coreidae may be either a type of camouflage to break up the insectlike profile or another example of deflective patterning that causes the predator to focus on an appendage rather than on the body of the prey species. Bright coloration, especially iridescence, might seem at first glance to be aposematic. However, it occurs most frequently in tropical forest species and is probably pro¬ tective in habitats where light and dark contrast strongly. Protective coloration may also involve the use of for¬ eign substances. The masked bedbug hunter, Reduvius personatus (Linnaeus), a black shining insect when adult, is pale and soft-bodied as a nymph. These nymphs, which live in houses, habitually cover themselves with a thick layer of dust particles so that they look exactly like a small piece of dust moving about in corners and under furni¬ ture. They are often called dust bugs for this reason. The origin of this habit presumably is not due to their asso¬ ciation with humans, because a similar phenomenon also occurs in some species in dry dusty habitats, where the nymphs cover themselves with small particles of sticks and other debris and become almost invisible among the detritus in which they live (see Chapter 48, Reduviidae). Mimicry and Protective Coloration and Shape 31 8 Heteroptera of Economic Importance Although it is probably true that no single species of Het¬ eroptera is as economically important as some species in other orders, it is also true that the variety of ways in which heteropterans affect humans and their environment is certainly as great as, if not greater than, that of any other major insect group. Economic importance traditionally has been viewed in terms of the direct damage that insects do to the crops upon which humans depend for existence and of the diseases that are transmitted to humans and their crops and domestic animals by insect vectors. As ecological sophistication has increased in recent decades so has our understanding of the exquisite interr-. iationships that exist between all of the organisms in an area, with ecologists now speaking of the impact that species and communi¬ ties have on the ecosystem. It is thus more difficult to speak of heteropterans in traditional economic terms, as the removal of what today may be thought of as a species of relatively little direct consequence to human welfare may in fact set in motion a chain of events leading to irreversible damage to the habitat in which we live. It is also true that our knowledge of the biology of most Heteroptera is still fragmentary or nonexistent. There is a great opportunity here for the field biologist (Gross, 1975-1976). In this discussion we shall limit ourselves chiefly to the more traditional concepts of economic importance—that is, the immediate damage or benefit caused by heteropter¬ ans to humans and to their associated crops and animals— and we will confine the discussion to a review of only a few of the heteropterans involved. Otten (1956) should be consulted for a detailed discussion of economically important plant-feeding Heteroptera. Because the majority of heteropteran species are phyto¬ phagous, it is obvious that among these plant-feeding species some will feed on the crops that humans utilize for food, medicines, and esthetic pleasure, and some will feed on plants that are essential for the continuation of the habitat. Plant damage by Heteroptera is, in most cases, di¬ rect damage. Although some plant diseases are carried b>’ Heteroptera, the various species that transmit diseases do so to a much lesser extent than do the Auchenorrhyncha and Stemorrhyncha. The economic importance of various Heteroptera also involves many species that are beneficial in that they feed on economically destructive insects. Some species are ectoparasitic on humans and domestic animals, and a few carry serious diseases of humans. The following discussion is organized alphabetically by family. Alydidae. Camptopus lateralis Germar is an alfalfa seed pest of considerable importance in southern Eurasia and North Africa (Mukhamedov, 1962). The elongate slender Leptocorisa acuta (Thunberg) is often a major pest of rice in the Orient. In India it is known as the Gandhi bug. Other species of Leptocorisa Latreille such as the rice ear bug, L. oratorius (Fabricius) (Rothschild, 1970), and L. chinensis Dallas sometimes cause serious injury in Asian paddies. Various species of Riptortus Stal, the “pod-sucking bugs,” are serious pests of beans {Phaseolus and other genera) from West Africa to Japan. Mirperus jaculus (Thunberg) is a pest of beans throughout tropical Africa. Alydids feed on seeds of rice and beans at the “milky grain” stage, when they are fully formed but unripe. Anthocoridae. Various species of Orius Wolff are im¬ portant predators on Thysanoptera, mites, and the eggs o; Lepidoptera, in both greenhouse and field crop situa¬ tions. Montandoniola maraquesi (Puton) has been widely introduced as a biocontrol agent of Thysanoptera on figs and olives. Berytidae. Wheeler and Henry (1981) summarized in¬ formation on Jalysus wickhami Van Duzee (the tomato stilt bug) and J. spinosus (Say) in North America. The former feeds on a great variety of dicotyledonous plants and sometimes is destructive to tomatoes. Jalysus spi¬ nosus feeds chiefly on monocotyledonous plants. Elsey and Stinner (1971) considered J. wickhami to be an im¬ portant predator of aphids and the eggs of Lepidoptera on tobacco. They stated that development could not be completed without utilization of some animal food. Cimicidae. All species feed on vertebrate blood, and among them are the common temperate and tropical bed bugs, Cimex lectularius Linnaeus and C. hemipterus (Fabricius). It has not been established that either of these species is a disease carrier, although both may cause nu¬ tritional deficiencies and allergies (Ryckman et ah, 1981). Human bed bugs as well as such species as the poultry 32 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) bug, Haematosiphon inodorus (Duges), are often serious pests in poultry houses. Bed bug bites have been one of the minor plagues of human societies for untold centuries. It is surprising how recently society has been able to free itself to a consider¬ able extent from these pests through the use of insecti¬ cides. (One of us remembers vividly as a young student going on a “field trip” to a house, only three blocks from his large campus, where bed bugs were so abundant that when wallpaper was stripped back there was an almost solid mass of swollen red bugs.) Oeciacus vicarius Horvath is a vector of Fort Morgan virus found in Cliff Swallows and House Sparrows and related to western equine encephalitis. Coiobathristidae. Several species of Phaenacantha Horvath are important pests of sugar cane in the Orient (Kormilev, 1949). Coreidae, A number of species are of considerable economic importance. For example, the North Ameri¬ can squash bug Anasa tristis (De Geer) often is a serious pest of cucurbits. Other Anasa spp. become destructive to cucurbits in localized areas. In East Africa Pseudotheraptus wayi Brown causes “gummosis,” or early fall, of coconuts (Brown, 1955a), and in the Solomon Islands the same type of damage is caused hy Amblypelta cocophaga China (Brown, 1955). Leptoglossus gonagra (Fabricius) is an important pest of garden and truck crops throughout the Pacific and in southern Africa (see Slater and Baranowski, 1990). Lep¬ toglossus phyllopus (Linnaeus) is a pest on a variety of cultivated plants in the southern United States, and L. clypealis (Heidemann) causes a high percentage of loss to the pistachio crop in California. Leptoglossus corculus (Say) and L. occidentalis Heidemann are often destructive to the seeds of various species of Pinus in North America, at times destroying over 50% of the seed crop (Krugman and Korber, 1969). Anoplocnemis curvipes (Fabricius) is a polyphagous pest of field crops in tropical Africa. Corecoris spp. attack sweet potato and other crops in the tropics and subtropics of the Western Hemisphere. Phthia picta (Drury) is a pest of tomatoes in the Caribbean. In the Orient Clavigralla gibbosa Spinola, the tur pod bug, has been reported in an abundance of 10 adults per plant on pigeon pea {Cajanus cajan), where it can cause almost complete destruction of the crop. Clavigralla elon- gata Signoret is a pest of certain legumes in East Africa (most of the economic literature is under the name Acan- thomia horrida), and C. tomentosicollis Stal is common throughout most of subsaharan Africa and is particularly injurious to beans. In Australia the crusader bug, Mictis profana (Fabri¬ cius), is often a serious pest in warm areas. Several species of the genus Chelinidea Uhler were introduced into Australia in an attempt to control the prickly pear cactus but with minor success. Cydnidae. Aethus indicus (Westwood) is sometimes a pest of cereal crops in tropical Africa and Asia. Scapto- coris castaneus Perty is a pest of sugar cane and has been reported feeding on the roots of cotton, bananas, toma¬ toes, and pimentos, A much more complex relationsh: is that of S. divergens Froeschner to the production of bananas. This insect injures banana plants by feeding on the roots. In many parts of Central America the banana crop has been destroyed by the presence of Fusarium wilt. Roth (1961) summarized what is known concerning the relationship of Fusarium wilt to populations of S. di¬ vergens. This cydnid is found primarily in sandy soils. Where the cydnids were abundant, feeding on the roots of bananas, the plants were healthy, whereas nearby plants with few cydnids were dying from Fusarium wilt. Roth noted records of populations as high as 76 individuals per square foot, with 532 insects feeding around the roots of a single healthy plant. The scent-gland secretions of nymphal and adult cydnids are toxic to Fusarium. This is an excellent example of how complex the relationships of insects to humans can be, for, while in many areas this cydnid is still considered harmful, in other areas it is possible to grow bananas only when the bugs are present. Dinidoridae. Coridius Janus (Fabricis) is a serious pest of cucurbits in the Orient, where it attacks both pumpkins and gourds of the genera Cucumis, Luffa. and Lagenaria (Rastogi and Kumari, 1962). Cyclopelta obscura (Lepele- tier and Serville) is a pest of legumes in Southeast Asia. Lygaeidae. Historically the most destructive plant¬ feeding North American heteropteran was Blissus leucop- terus (Say). As early as the mid-1700s, this small black and white insect was the scourge of corn fields and did considerable damage to other grain crops. Unlike many major insect pests it is a native species, and presumably before the advent of vast monoculture techniques it fed on native grasses. In the course of adapting to human’s crops it apparently shifted from being predominantly flightless to always being capable of flight and migrating to and from the fields each year. The ravages of this insect in the midwestem United States in the 1930s enlisted most of the farming community in burning, trenching, and spray¬ ing. The chinch bug is still an important pest, but its abundance seems to be tied to drought conditions, for in wet summers its numbers are held in check by a disease organism. At present an eastern subspecies (B. leucop- terus hirtus Montandon) and a southern relative {B. insu- laris Barber) are of great importance because they attack lawn grasses. In Florida whole fleets of trucks with their “chinch bug control” logos may be seen, from which professional workers busily attempt to rid householders of this lawn destroyer. Heteroptera of Economic Importance 33 In South Africa the chinch bug Macchiademus diplop- terus (Distant) is a serious pest of wheat. It appears to have adapted to this plant from a native grass in a man¬ ner similar to that of the North American chinch bug. The Oriental chinch bug Caveleriiis sweeti Slater and Miyamoto is a serious pest of sugar cane. In Africa, the Orient, and Australia several species of Oxycarenus Fieber, especially O. hyalmipennis (Costa), are important pests of cotton, where they damage the crop not only by feeding but also by staining the bolls. Oxycarenus hyalipennis has been introduced into South America. In Australia the rutherglen bug, Nysius vinitor Ber- groth, is a major pest as are Nysius ericae Schilling ( = niger Van Duzee) and N. raphanus Howard in the United States. These oligophagous plant feeders damage culti¬ vated crops most when native plants in arid regions dry up. At times the numbers become extremely high and the insects even become nuisances as they swarm into human habitations. In southern Africa a similar phenomenon occurs. Species of Spilostethus, especially S. pandurus (Sco- poli), damage many different crops in Europe, Africa, and Asia. Elasmolomus sordidus (Fabricius) is a pest of various seed crops, especially peanuts (groundnuts) both in the field and in storage situations in Africa and Asia. Some species of Geocorinae are considered of value in biological control of crop pests. A substantial litera¬ ture on North American species of Geocoris Fallen has developed in recent decades. Malcidae. Several species of Chauliops Scott have been reported as destructive to cultivated legumes in the Orient (see Slater, 1964b). Miridae. This large family contains many species that damage crops. Problems range from occasional outbreaks or injury to crops of minor importance, to serious injury to crops of great economic value. At least 40 species of the subfamily Bryocorinae are known to feed on cacao. Members of the subtribes Odo- niellina and Monaloniina are particularly destructive. In Africa the principal species are Sahlbergella singularis Haglund and Distantiella theobroma Distant. At least 11 species of Monalonion Herrich-Schaeffer are serious pests in South and Central America; in southeast Asia and Africa species of Helopeltis Signoret may be equally or more destructive; and in Madagascar the principal damage is done by Boxiopsis madagascariensis Lavabre (Lavabre, 1977). Several species of the genus Helopeltis in the Orient are serious pests of crops other than cacao. The primary destruction is to tea, cashew, and cinchona, but other economically important plants including allspice, black pepper, apples, grapes, and guava are also attacked. Be¬ fore the use of modern insectieides crop losses on tea plantations often reached 100% (Stonedahl, 1991). Calocoris norvegicus (Gmelin) is a serious potato pest in many temperate areas. Adelphocoris lineolatus (Goeze) was accidentally introduced into North America from Europe, spread rapidly, and is a very serious pest in areas where alfalfa (Medicago) is grown for seed (Wheeler, 1974). It is also a serious pest in the Palearctic. The tarnished plant bug, Lygus lineolaris (Palisot de Beauvois), and closely related species are very general feeders, attacking a wide variety of plants. When feed¬ ing on fruits such as peaches and pears they cause a blemish called “cat facing.” that seriously damages the fruit. There is an extensive literature dealing with this group as pests of many garden and crop plants (see Scott, 1977, 1981; Graham et al., 1984; O. P. Young, 1986). Lygus hesperus Knight and L. elisus Van Duzee are also of major importance, the latter causing serious damage to cotton crops in the western United States. Other cul¬ tivated plants damaged by these species include beans, strawberries, peaches, and many seed crops, the most important being alfalfa and rape seed grown for oil. Stern (1976) stated that in California alone in 1974 L. elysus and L. hesperus caused an estimated loss of S56 million, more than 10% of the total loss caused by all insects and mites to all California crops for that year. The history of control of these two pests offers a striking example of the importance of knowledge of the ecological relation¬ ships of insect pests. Stern (1976) found thatL. hesperus actually does not attack cotton if other suitable plants are available. When alfalfa was interplanted as a “trap crop,” cotton was raised without any insecticide application. Ly¬ gus rugulipennis Poppius and L. pratensis (Linnaeus) are destructive to many cultivated plants in the Palearctic. The predatory nature of several Lygus spp. as well as other mirids further complicates the picture. Wheeler (1976) concluded that many Miridae usually considered to be phytophagous feed to some extent on small arthro¬ pods. Taylorilygus pallidulus (Blanchard) is a polyphagous pantropical pest. Creontiades pallidus (Rambur) injures sorghum, Solanum spp., and other crops in southern Eu¬ rope, Africa, and Asia. Cyrtopeltis (Nesidiocoris) tenuis Reuter is a pantropical pest of tomato and other solana- ceous crops. Several mirids do significant damage to rangeland grasses and small grain crops. Among these are species of Irbisia Reuter, Labops Burmeister, Leptopterna Fieber, and Trigonotylus Fieber. Tytthus mundulus (Breddin) is well known among predatory mirids, because of its successful use in Hawaii as a biological control agent of the sugar cane leafhopper Perkinsiella saccharicida Kirkaldy (Zimmerman, 1948). 34 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Members of the deraeocorine genus Stethoconus Flor, are either primary or exclusive predators of Tingidae. Stethoconus frappai Carayon (1960) has been recorded as an important population control agent of Dulinius uni¬ color (Signoret) on coffee in Madagascar and S. japonicus Schumacher of the Rhododendron lace bug Stephani- tis rhododendri Horvath in Japan and the United States (Henry et al., 1986). Pentatomidae. Most stink bugs are plant feeders, and several of them are of great economic importance. Nezara viridula (Linnaeus) is a large green species found in the warmer parts of the Old and New Worlds. It attacks over 90 species of plants, including many vegetables, and is often a serious pest of beans and tomatoes. Murgantia histrionica (Hahn), the brightly colored har¬ lequin cabbage bug, is a serious pest of some crucifers in the United States and Mexico (Paddock, 1918). This species is native to Mexico but has spread northward within the past hundred years. Stink bugs severely damage cacao in the Old and New Worlds, not only by direct feeding but, more impor¬ tant, by introducing pathogens. Among the more de¬ structive species are Mecistorhinus tripterus (Fabricius) (Sepulveda, 1955) and Bathycoelia thalassina (Herrich- Schaeffer). The latter species has become increasingly important, apparently because of the destruction of its natural enemies by the use of insecticides, the destruc¬ tion of its wild host plants, and the introduction of new and more susceptible varieties of cacao (Owusu-Manu, 1977). Pentatomids have also been implicated in the transmis¬ sion of pathogens to palms (Dolling, 1984). Other destructive pentatomids include the following: Scotinophara spp. are pests of cereals, particularly S. lurida (Burmeister) on rice (Kiritani et al., 1963), and grasses in Africa and Asia; Eysarcoris ventralis (West- wood) is sometimes a pest of rice in Southeast Asia; Oebalus pugnax (Fabricius) is a rice pest in the Western Hemisphere (Sailer, 1944); Piezoderus hybneri (Fabri¬ cius) feeds on various legumes in Africa and Asia; Ba- grada cruciferarum Kirkaldy feeds on crucifers in the Old World; Biprorlulus bibax Breddin is a pest of citrus (McDonald and Grigg, 1981); Antestiopsis lineaticollis (Stal) is a widespread coffee pest (Greathead, 1966); vari¬ ous species of Euschistus Dallas are descructive to soy¬ beans in North America (Boas et al., 1980); Agonoscelis versicolor (Fabricius) attacks millet, cotton, and cacao; and Aelia rostrata (De Geer) is a serious wheat pest in the Near East. The last species, like several scutellerids and some other pentatomids has an unusual life cycle, in that the insects leave the wheat fields in late summer and migrate as much as 50 km to overwinter in the mountains at elevations of 1500-1800 meters in Quercus scrub. The migration is not a continuous one, but occurs in stages over several days (Brown, 1965). In Morocco the related Aelia germari Kuster is known to migrate several hun¬ dred kilometers. Similar migrations in the Near East have been observed in species of Dolycoris Mulsant and Re\'. Carpocoris Kolenati, Codophila Mulsant and Rey, and Eurydema Laporte. The Asopinae are predaceous pentatomids, some of which are significant biological control agents. Particu¬ larly important among these is Perillus bioculatus (Fabri¬ cius), which feeds almost exclusively on the Colorado potato beetle. The beetle, which apparently originally lived on wild solanaceous weeds in the western United States, adapted early to potatoes and spread rapidly east¬ ward across the country with the predatory stink bug following along. Unfortunately the beetle was introduced into Europe, where it has become even more destructive than it is in North America. Perillus Stal has been intro¬ duced into Europe as a control measure, but apparently it has been a failure. Several species of Podisus Herrich-Schaeffer are con¬ sidered valuable predators, but since most of them are rather general feeders they are for the most part not capable of major control. Eocanthocona furcellata (Wolff) preys on larvae of Limacodidae on palms in Southeast Asia. In Australia the pentatomid Agonoscelis rutila (Fabricius) is widely known for its control of horehound. Piesmatidae. Piesma cinereum (Say) transmits the virus known as “savoy” that causes serious damage to sugar beets. Piesma capitatum (Wolff) is a pest of sugar beets in Russia, but does not transmit any disease. Piesma quadratum (Fieber) transmits a leaf-curl disease of sugar and fodder beets in Germany. Pyrrhocoridae. Several species of the genus Dysder- cus Guerin-Meneville are serious pests of cotton in tropi¬ cal areas (Fuseini and Kumar, 1975). They frequently form large aggregations both on cotton and on wild hosts such as Hibiscus and Ceiba. The colonial nature of these aposematically colored bugs aids to warn predators away from the distasteful clusters. Van Doesburg (1968) re¬ ported that in the New World several species are some¬ times present in feeding .clusters that are confined almost entirely to ripening fruits and seeds. Also, it has been demonstrated that when several insects feed on a seed, an enzyme pool is secreted that makes feeding more ad¬ vantageous for the entire group. These aggregations are formed by visual, tactile, and olfactory cues. Reduviidae. Perhaps the most destructive heteropteran is not a plant feeder but an assassin bug that feeds upon the blood of humans and other mammals. Throughout the warmer parts of the Western Hemisphere, but especially in South America, various species of Triatoma Laporte Heteroptera of Economic Importance 35 and related genera transmit the trypanosome that causes the debilitating Chagas’ disease in humans. These insects are thought to be one of the most important reasons for the impoverishment of many areas of South America. The insects often live in thatched houses, and they transmit the trypanosome parasite to the inhabitants by subsequent defecation in the wound that they make when feeding on the sleeping victim. Whole communities may be infected and, as with African sleeping sickness, infected people become lethargic, are unable to function efficiently, and are susceptible to a variety of other diseases. It is inter¬ esting to note that Charles Darwin wrote in his journal of being bitten by the “great black bug of the pampas” when he lived in Argentina with the gauchos. For years there has been controversy concerning the lifelong illness of Darwin, with theories ranging from psychosomatic to genetic, and including the possibility that Darwin suffered from Chagas’ disease. Some Reduviidae are beneficial. Tinna wagneri Villiers preys on mosquitoes in dwellings in East Africa; Pho- noctonus spp. feed on Dysdercus spp.; Sycanus collaris (Fabricius) preys on Limacodidae larvae in Southeast Asia; and Amphibolus Venator (Klug) and Peregrinator binnnulipes (Montrouzier and Signoret) prey on stored- prv;ducts pests. Rhopalidae. The box elder bug, Boisea trivittata (Say), i.s D. ten abu\aant on box elder trees {Acer negundo) in N'; -\h Amer: but it is chiefly a nuisance insect for its habit of entering houses in large numbers for hibernation. Liorhyssus hyalinus (Fabricius) is a cosmopolitan pest of many low-growing crops, especially Asteraceae. Lep- tocoris hexophthalmus (Thunberg) often destroys coffee and cotton in Africa. Scutelleridae. The most destructive species are mem¬ ber; of the ge .IS Eurygaster Laporte. In the Near E' t they destroy large quantities of wheat and related g> ■ crops. Most infamous is the soun bug, Eurygaster inttgri- ceps Puton. In the southern part of the former USSR, Turkey, Iraq, and Iran this species is a major pest of wheat, and there is an enormous economic literature. The Russian literature is summarized in Fedotov 1947-1960. Studies by Brown (1962a, b, 1963, 1965) are especially important. In the spring these insects attack wheat and other cereal crops, sucking the green shoots and causing the terminal buds to die. Later they feed on the seeds. The crop may be damaged so that it is not worth harvest¬ ing, and even when the damage is of a lesser magnitude the feeding affects palatability and lowers baking quality. These insects have a remarkable life cycle. In late summer they fly en masse from the wheat fields to mountainous areas, where they overwinter. The flights may be as long a: 00 km, but usually they average 20-30 km. The mi- gi on from the lowlands is to areas as much as 1000 or even 2000 meters higher in elevation. The insects re¬ main there for nine months, massed under cushionlike or dome-shaped ground-hugging plants such as species of Astragalus and Artemisia. Brown (1962b) reported popu¬ lations of up to 900 bugs per plant and up to 1000 bugs per square meter. Overwintering consists of two stages. The first is actually an aestivation while the weather is relatively warm. It terminates, at least in Turkey and Iran, in a second movement, to an elevation as much as 100 meters lower. In the late spring the population migrates back to the wheat fields. Thus, a given individual lives for an entire year. Although E. integriceps is by far the most important species economically, it is only part of a more widespread phenomenon in the Near East that also includes pen- tatomids, coreids, alydids. and rhopalids. At least three species of Odontotarsus Laporte and Ventocoris fischeri (Herrich-Schaeffer) also migrate from the mountains to cultivated fields, but they feed to a large extent on weeds rather than on wheat and other cereals. Brown (1965) showed that migration is influenced by wind conditions and to some extent by the direction of the sun. In the case of E. integriceps there are periods of enormous abundance and periods of relatively low popu¬ lations. During outbreak periods there is increased homo¬ geneity in the populations and increased resistance to environmental conditions. These changes in the popu¬ lation structure are passed on for several generations, a situation rather similar to that found in migratory lo¬ custs. It seems likely that the original populations of E. integriceps and the other migratory scutellerids lived on mountain grasses at middle elevations and moved to abun¬ dant food sources in the lowlands as agriculture became more concentrated. Tessaratomidae. Musgraveia sulciventris (Stal), known as the bronze orange bug in Australia, is a pest of citrus in the northeastern part of the continent. Its native host plants are also members of the Rutaceae. It is a very large insect (over 20 mm) with flattened ovoid nymphs. In China Tessaratoma papillosa Drury is destructive to the fruit of the litchi (Yang, 1935). Tingidae. Lace bugs of the genus Stephanitis Stal are frequently serious pests of ornamental azaleas, rhododen¬ drons, and andromedas. Stephanitis pyri (Fabricius) and Monosteira unicostata (Mulsant and Rey) are both pests of rosaceous orchard trees, especially pear, in Europe. Corythuca ciliata (Say), the sycamore lace bug, dam¬ ages Platanus spp. in the eastern United States and is now causing concern in southern Europe, where it was recently introduced. Urentius hystricellus (Richter) dam¬ ages aubergine in the Old World tropics. Corythuca gossypii (Fabricius) is a pest of cotton, beans, citrus, and solanaceous crops in the Caribbean and Central and 36 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) South America. Gargaphia torresi Costa Lima attacks cotton in Brazil. Corythaica cyathicollis (Costa) is a pest of Solanaceae in the Caribbean and South America, and Corythaica monacha (Stal) is destructive to cotton and beans in South America. Several phytophagous heteropterans have been used as biological control agents against invasive weeds. None of these has been spectacularly successful. Among the best known is the tingid Teleonemia scrupulosa (Stal), which has been widely introduced in an attempt to control Lan- tana. For additional information see Drake and Ruhoff (1965). Veliidae. Some species of Microvelia Westwood have been employed as predators in biological control schemes; for example, Microvelia pulchella Westwood has been used to control mosquitoes (Miura and Takahashi, 1987. 1988). Microvelia spp. are effective predators of rice planthoppers (Reissig et al., 1985). Heteroptera of Economic Importance 37 9 Historical Biogeography Our understanding of the meaning of heteropteran dis¬ tributions has undergone revolutionary change since the 1960s. Two factors have been of particular importance. First, most working scientists have accepted the con¬ cept of continental drift, despite continuing controversy over details. Our knowledge of the heteropteran fossil record—no matter how fragmentary—makes it evident that most major lineages diverged early in the Mesozoic, from Pangaean, Laurasian, or Gondwanan patterns, and therefore under.sta;.ding the former relationships of con¬ tinents is crucial ;o understanding many present-day dis¬ tributions. Second, the rise of cladistic methodology has provided a more rigorous means of recognizing mono- phyletic group:- and the relationships among them. This acceptance of continental movement and cladis¬ tic methods shifted biogeographic emphasis from centers of origin, classification of subregions, and methods and abilities of dispersal, to attempts to establish areas of endemism and degrees of distributional concordance be¬ tween different taxa and the areas they occupy (e.g., see Nelson and Platnick, 1981). It must be kept in mind that historical biogeography can be no better than the facts upon which interpreta¬ tions are made. There is a great deal that we do not know about heteropteran distributions both factually and causally. Not only are there still large numbers of un¬ described taxa in every large family, but knowledge of the distributions of most species and even of some higher groups is fragmentary at best. Thus, there is a continuing need for careful faunistic studies of local areas. Although most groups of Heteroptera are reasonably well known in Europe and North America, most of the rest of the world is in desperate need of additional study, in both temper¬ ate and tropical areas. This is true because of the danger of irrecoverable loss of information due to extinction, particularly at the hand of humans. Because most families are almost worldwide in distri¬ bution, it is at the level of the subfamily, tribe, and below that we ordinarily begin to perceive and attempt to under¬ stand patterns of endemism. In the chapters that follow dealing with the various families of Heteroptera, the dis¬ tributions of families, subfamilies, and sometimes tribes are summarized. Because the meaning of distributions is subject to in¬ terpretation, it seemed to us important to first enumerate some working principles, especially today, when so much controversy exists as to the most productive approaches to the study of historical biogeography. We then provide some interpretive remarks regarding apparent general pat¬ terns of distribution within the Heteroptera. A first principle is that whether or not one wishes to recognize paraphyletic groups, only monophyletic groups can be the subject of biogeographic analysis. A second principle is that in interpreting disjunct dis¬ tributions, a vicariance model should be considered first, and only when this model proves to be unsatisfactory should dispersalist hypotheses be considered. A third principle is that, even though only positive occurrences provide information, distributions of lin¬ eages of many taxa, including recent representatives, changed dramatically during the Tertiary and before, as is clearly shown by the amber-fossil record, for example. Thus, extinction should never be discounted as a possible explanation for seemingly anomalous patterns. A fourth principle in interpreting any distributional pat¬ tern is to ask what geologic or climatic event might best explain the observed pattern. Perhaps a final principle of any meaningful distribu¬ tional study should be a clear statement of the method¬ ological basis of the study. The classical separation of the terrestrial faunas by Sclater (1858) into Nearctic, Palearctic, Neotropical, Ethiopian, Oriental, and Australian still has merit, but it tends to mask relationships across continental lines. We give here only a few examples from the Heteroptera which more or less conform to Sclater’s system but which also help to point out its defects. Neotropical Region. This area has long been treated as including all of Central and South America and the West Indies. This well-known pattern has been documented in the Miridae by Schuh and Schwartz (1985, 1988). The far southern part of South America, however, has a het¬ eropteran fauna that is either depauperate or more closely allied to that of Australia and New Zealand, with some elements extending far northward at higher elevations in the Andes. The latter situation is well demonstrated by 38 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) the Saldidae. Many elements of the Nearctic fauna extend southward along the cordilleras through Mexico deep into Central America and in some cases Colombia. The West Indies are for the most part composed of a mixture of South American elements in the Lesser Antilles and Cen¬ tral American elements in the Greater Antilles, with only a small part of the fauna being derived from temperate North America. Slater (1988) believed that much of the West Indian lygaeid fauna is the result of over-water dis¬ persal despite the complex geologic history of the Greater Antilles and the strong evidence for the influenee of plate movements in their formation. Ethiopian Region. Like that of South America, the African fauna is composite. Southern Africa has a degree of relationship with Australia and New Zealand, as shown by the Plinthisini and Stygnocorini (Lygaeidae). Tropical Africa has a high proportion of Paleotropical elements found in family after family. There is also close relation¬ ship with the Palearctic, particularly north of the Sahara, but also in the east African savannah fauna. There is evidence, however, of an older relationship between the Macchia plant community of South Africa and the Ma- greb of North Africa. Africa does not have the mountain chains so characteristic of most other continents. None¬ theless, the isolated volcanic peaks of East Africa have a fauna showing relationships to the Palearctic, to the Drakensberg range of South Africa, and to Madagascar. Madagascar shows a high degree of endemicity, which increases as the fauna becomes better known. This en¬ demicity, however, is overlaid with a recent invasive Afri¬ can savannah fauna. The vaunted Oriental relationship is not yet well documented in the Heteroptera, although it is known to occur in the Ptilomerinae (Gerridae). Australian Region. Australia contains many Paleo¬ tropical elements in the north, often shared with New Guinea. The temperate Bassian subregion shows a high degree of endemicity. In some groups such as the udeo- corine Lygaeidae a marked radiation has occurred in this area (Slater, 1975). Small families such as the Lestoni- idae and Aphylidae are endemic here. As in the case of South America and Africa, the Australian Heterop¬ tera fauna is to a great degree a composite, despite the relatively homogeneous mammal fauna on which most classical biogeographic scenarios have been based. Holarctic. The faunas of the northern Hemisphere are much better understood taxonomically than those of the Southern Hemisphere. There is a marked Hol¬ arctic element in the Miridae, Saldidae, Gerridae, and other families (see Wheeler and Henry, 1992). Genera in many families show considerable endemicity (Holarctic). Nonetheless, in many families the faunas of both the Nearctic and Palearctic comprise northern extensions of the rich tropical faunas to the south. Other types of intercontinental patterns are also readily found in heteropteran distributions. Gondwana patterns. The term Gondwanan has often been used in reference to taxa found in Australia, New Zealand, southern South America, and southern Africa. In fact, many far southern distributions do not include Africa, apparently because the northward movement of Africa has eliminated cold-adapted elements from the southern part of the continent. There is no known heterop¬ teran distribution similar to the classic Gondwana pattern proposed by Brundin (1966) for the Chironomidae. Slater and Sweet (1970) documented a similar pattern for styg- nocorine lygaeids, but the group is not known from South America (perhaps because of a lack of nymphs, which are critical for its recognition). But there are several groups with distributions that are difficult to interpret as other than Gondwanan. The Enicocephalomorpha, the apparent sister group of all other Heteroptera (Fig. 1.1), have their greatest diver¬ sity in New Zealand and the southwest Pacific, with the fauna of the remainder of the world being relatively mo¬ notonous and composed almost entirely of members of the Enicocephalidae. Similarly, the Aradidae are diverse in New Zealand, with all eight subfamilies occurring there, while the Northern Hemisphere fauna is composed of a limited number of phyletic lines. A similar pattern occurs in the Acanthosomatidae. Monteith (1980) documented an apparently ancient pattern in the Chinamyersiinae (Aradidae), a group that occurs in the mountains of eastern Australia, New Cale¬ donia, and New Zealand. A similar pattern was found in some cylapine Miridae, which also occur on Lord Howe Island, although not in New Zealand (Schuh. 1986a). For such taxa as the Idiostolidae, considered by several authors to be the most primitive of the lygaeoid families, and the aradid subfamily Isoderminae, the situation is less clear. Some would consider such groups to represent a classic East Gondwanaland pattern including southern Australia, Tasmania, New Zealand, and southern Chile. But, because there is no cladistic analysis of these groups, it is impossible to exclude the possibility of a more recent relationship through Antarctica long after the breakup of East Gondwanaland. There is, however, strong evidence in the Heterop¬ tera of West Gondwana relationships—that is. between tropical and subtropical Africa and South America. Ex¬ amples are the Pachynomidae, with the Aphelonotinae restricted to Africa and South America, and the Pachy- nominae, with one South American genus and two Old World genera (these extending from Africa east into India also). Similar patterns can be found in the Cetherinae and Holoptilinae (Reduviidae), Plokiophilinae (Plokio- philidae), and Madeoveliinae (Mesoveliidae). Historical Biogeography 39 The family Pyrrhocoridae demonstrates a possible ori¬ gin in the Eastern Hemisphere with subsequent migration to and diversification of one taxon in the New World. Van Doesburg (1968) noted that of the 30 recognized genera, all Western Hemisphere species belong to one subdivi¬ sion of Dysdercus that otherwise occurs only in Africa. If van Doesburg is correct in believing that the family reached the New World by over-water migration, then it must have been sufficiently long ago for the elaborate mimicry complexes to have evolved. To believe that this is a vicariance pattern involving West Gondwanaland means that the other African pyrrhocorids were not present or subsequently have become extinct in the New World. Paleotropical patterns. Distributions encompassing the Old World tropics and subtropics are extremely com¬ mon in the Heteroptera. Such patterns frequently in¬ clude tropical northern Australia and the western Pacific Islands—sometimes as far east as the Marquesas. Some taxa are absent from Africa, and others, while present there, are restricted to West Africa or also the Congo basin. Schuh (1984), Schuh and Stonedahl (1986), and Stonedahl (1988b) have termed such patterns in the Miri- dae “Indo-Pacific.” At the family level only a few groups show such a pattern (Plataspidae, Malcidae), but there are untold numbers of examples at the subfamily, tribal, and generic levels. Transpacific patterns. Several groups of Heteroptera occur exclusively or primarily in the Southern Hemi¬ sphere but do not occur in Africa. This striking distri¬ bution is well illustrated by the Colobathristidae (Stys, 1966a, b) and triatomine Reduviidae (Lent and Wygod- zinsky, 1979); both show substantial diversity in the New World tropics and range in the Old World from India to northern Australia. In the Triatominae this distribu¬ tion may even be present at the specific level, as a single species is tropicopolitan, but may have been spread by humans as may also be the case with the only known Eastern Hemisphere species of the coreid genus Lepto- glossus Guerin-Mendville. Several other reduviid distri¬ butions appear to be transpacific (Peiratinae, Physoderi- nae, and Vesciinae). The Thaumastocoridae may show such a pattern, although of a more relictual nature. There is insufficient information in all these cases to identify the next area of relationship. Nevertheless, in the eccritotar- sine Miridae, Africa appears to contain the sister group of a clade whose distribution is otherwise transpacific. Ly- gaeid clades such as the Antillocorini and Ozophorini also appear to have such a pattern, although the higher-level cladistic relationships are not yet sufficiently well under¬ stood. This transpacific pattern is widespread in flowering plants. Relict patterns. When we look at present-day distri¬ butions, it is evident that monotypic groups (such as the Joppeicidae and Medocostidae) and small families with limited distributions (Canopidae, Megarididae, Phloei- dae, and Lestoniidae) provide little information of general interest. On the other hand, small families such as the Paraphrynoveliidae, with only two known species, one in the Cape region of South Africa and the other in Lesotho, show a distribution similar to that of many other insect and plant taxa, suggesting a localized vicariance pattern, perhaps due to aridity as recently as the Pleistocene, but conceivably much older. Oceanic islands. Traditionally the faunas of oceanic islands have been thought to be solely the result of long¬ distance dispersal over water. This explanation fails to appreciate the great age of many lineages within the Het¬ eroptera and discounts movements of land masses as a causal factor in explaining distributions. Consequently, the origins of island faunas must be reexamined. As an example, the Hawaiian fauna had been thought to have arrived within the last 5 million years. Yet the recognition of the Emperor Sea Mount chain, which con¬ tains the remnants of formerly larger islands that passed over the Hawaiian “hot spot,” suggests that the fauna may well be much older. Nevertheless, these islands also possess an imbalanced fauna, suggesting that over-water dispersal may have been important in populating them. This is well illustrated by the mirid fauna which Zimmer¬ man (1948) placed in several subfamilies, whereas Schuh (1974) demonstrated that all groups that have diversified belong to a single tribe in the Orthotylinae. The diverse lygaeid fauna belongs almost entirely to the subfamily Orsillinae, a group notorious for its ability to colonize remote islands. The Nabidae also radiate remarkably in these islands, but within only one or a few lineages. The history of other islands such as New Caledonia, the New Hebrides, the Galapagos, and the Juan Fernan¬ dez remains enigmatic because the relationships of most groups occurring there are still ambiguous. New Zea¬ land may be an exception. It shows ancient Gondwana relationships, more recent ones through a warm Antarc¬ tica, and evidence of recent over-water dispersal. For example, the previously mentioned diversity of the ara- did and enicocephalid faunas is in striking contrast to the imbalanced lygaeid and mirid faunas and the com¬ plete absence of Coreidae and Reduviidae (other than Emesinae). ; The Heteroptera will be important to any understand¬ ing of historical biogeography on a world scale. Further progress toward such understanding will come only with an improved knowledge of distributions and cladistic re¬ lationships. 40 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) 10 General Adult Morphology and Key to Infraorders of Heteroptera Head The heteropteran head ranges from elongate both anterior and posterior to the large compound eyes, as in the Enico- cephalomorpha (Fig. 13. lA), to barely longer than the compound eyes, as in many Miridae (Fig. 10.1). The compound eyes are generally well developed, and in many groups such as the Nepomorpha (Fig. 33.1) and Lepto- podomorpha (Fig. 45.1 A), they are large in comparison with the total size of the head. In other groups such as the Gerromorpha (Fig. 20.2C) and many members of the Pentatomomorpha (Fig. 64.1 A) they are relatively much smaller. In some groups such as the Vianaidinae (Tin- gidae) (Fig. 54.1) they are reduced, whereas in some Enicocephalomorpha and all Polyctenidae and Termi- taphididae they are completely absent (Fig. 65.1 A, B). In first-instar nymphs the number of ommatidia ranges from 4 or 5 to many, and 1 or 2 setae arise from the center and/ or periphery of these facets (Fig. lO.lOE). Primitively 2 ocelli are situated between or sometimes slightly behind the compound eyes in adults (Fig. 13.1 A) and may be close to one another or widely separated; these tue lost in many Nepomorpha, consistently in some families of other infraorders, often in forms with reduced wings, and are always absent in nymphs. Posteriorly the head forms a neck that inserts into the pronotum. Anteriorly there is a more or less prominent clypeus (or tylus), which is subdivided into an antecly- peus (tylus) and postclypeus in some taxa, such as most Leptopodomorpha. Adjacent—and in most taxa poste¬ rior—to the clypeus, lie dorsally the mandibular plates (or juga) and ventrally the maxillary plates (or lora) (Fig. 10.1), which internally form the attachments for the man¬ dibular and maxillary levers and their associated stylets" (Spooner, 1938; Cobben, 1978). The labrum, triangular in form, is generally situated below the clypeus. The mandibles and maxillae are modi¬ fied—as in all Hemiptera—into tubular concentric stylets (Fig. 10.2B), the former surrounding the latter. The man¬ dibular stylets serve a cutting and lacerating function and are often adorned apically with minute barbs or teeth. The maxillary stylets—like the mandibular—are held together by interlocking keyways and contain the salivary canal and food canal (Fig. 10.2B). The labium, which is tubular, either straight or curving, and in many taxa capable of substantial flexion during feeding, possesses 3 or4 segments (Fig. 10.2A). The 3-segmented condition is always the result of reduction of the basal segment. The labium varies from relatively short and barely reaching the posterior margin of the head—as in many Nepomor¬ pha and Reduviidae—to relatively long and reaching to near the apex of the abdomen—as in some Pentatomo¬ morpha; distally it bears a field of specialized sensors (Fig. 10.7B) (Cobben, 1978). Maxillary and labial palpi are completely absent. The epipharyngeal sense organ lies between the base of the labrum and the labium. In the family chapters we refer to the ultimate labial segment as 4 and count backward, so that the basal segment in the case of a 3-segmented labium will be segment 2. Ventrally the head possesses a distinct gula. The buc- culae are usually well developed, lying along each side of the labium and ordinarily extending to near the posterior margin of the head. Posteriorly the gula may be open and conjoin the neck, or it may be closed by a distinctively sclerotized buccular bridge. The heteropteran antenna consists of 4 basic segments: the scape (1), pedicel (2), basiflagellum (3), and disti- flagellum (4). During postembryonic development addi¬ tional sclerites may be formed or subdivisions may take place. Most obvious among these are the presence of a prepedicellite between segments 1 and 2. which forms the fifth segment in the Prostemmatinae (Nabidae) (Fig. 56.1), and the subdivision of the pedicel to form 5- segmented antennae in the Pachynomidae (Fig. 47.1) and many Pentatomoidea (Fig. 66.1). A few taxa, including some Nepomorpha and Phloeidae show segmental loss or fusion (Figs. 40.1C, 74.2A). Antennal structures, in¬ cluding other less obvious intersegmental sclerites were described in detail by Zrzavy (1990c). Thorax The prothorax may possess anteriorly a distinct rounded or flattened collar (or collum) or may take the form of a finely reflexed margin. Dorsally there is usually a transverse impression delimiting the anterior lobe and General Adult Morphology and Key to Infraorders 41 labrum- labium coxa ostiolar peritrem abdomen pygophore genitalia frons antennal insertion J, mandibular plate gula clypeus — maxillary plate buccula antenna propleuron mesepistemum mesepimeron 'mesoscutum 'Scutellunfi •'Clavus- claval suture 'medial trochanter ioxa femur ^ fracture R+M - costal fracture cuneus tibia tarsus pretarsus pulvillus parempodium vertex collar callus Fig. 10.1. General heteropteran morphology. Lygus sp. (Miridae) (modified from Schwartz and Foottit, 1992). 42 TRUE BUGS OF THE WORLD (HEMiPTERA: HETEROPTERA) Fig. 10.2. Mouthpart structures. A. Longitudinal section of head of Aradidae (from Weber, 1930). B. Cross section of mandibles and maxillae, Oncopeltus tasciatus (modified from Forbes, 1976). Abbreviations: cd, centrai duct; cdm, cibarial dilator muscles; fc, food canal; mds. mandibular stylets; mms, mandibular and maxillary stylets; mxs, maxillary stylets; sc, salivary canal. posterior lobe, the former usually possessing a pair of somewhat protuberant calli, which are grossly developed in some Saldidae and Miridae into conical structures, the external expression of foreleg muscle attachments. The humeral (posterolateral) angles of the pronotum may be rounded or projecting and sometimes spinelike as in many Reduviidae, Coreidae, and Pentatomidae. Laterally the prothorax may be either rounded or carinate, explanate, or reflexed. The dorsum of the mesothorax is usually visible in the form of the triangular scutellum (mesoscutellum), which is bordered anteriorly by the sometimes visible mesoscutum; both may be occasionally completely ob¬ scured by the posterior lobe of the pronotum; in some Pentatomoidea the scutellum is greatly enlarged, entirely covering the wings and abdomen (Fig. 68.1 A). The thoracic pleuron is structurally comparatively mo¬ notonous and assumes the form similar to that found in most other pterygote insects. The metepisternum is dis¬ tinguished by the presence of the peritreme, the actual point of release onto the body surface of fluid from the metathoracic scent glands, and the associated evapora- tory area (evaporatorium) found in most Cimicomorpha and Pentatomomorpha (see also Exocrine Glands). The thoracic sternum may be relatively broad or almost completely obscured by closely spaced coxae. The pro¬ sternum sometimes is produced ventrally in the form of a xyphus, or it may be longitudinally grooved for reception of the labium. The mesosternum and metasternum may bear longitudinal spines in some Pentatomoidea. Spiracles are located dorsally on the pleuron on the meso- and metathorax, the anterior pair usually being obscured. Wings Wing venation in the Heteroptera has been investigated by Tanaka (1926), Hoke (1926), Davis (1961), Leston (1962), and Wootton and Betts (1986), among others. There is no absolute agreement on the homologies of all veins. The wings are folded flat over the abdomen. The forewings may be either of uniform texture (tegminal) (Fig. 10.3A)—as in the phylogenetically more primitive groups—or in the form of hemelytra. that is. divided into a distinctly coriaceous anterior portion and a membranous posterior portion (Fig. 10.3C) (with sometimes reduced venation), as in the Panheteroptera. The anterior margin of the wing, which lies laterally in repose, is bordered by the subcosta (Sc) or costa plus subcosta (C + Sc), and often reflexed and thickened ventrally into a hypocostal ridge or lamina. Although venation in all Heteroptera is somewhat reduced, it is most well developed in the Enicocephalomorpha, with a distinct radius (R), media (M), and cubitus (Cu), all attaining the posterior margin of the wing. R and M are often fused and accompanied by a fracture line known as the medial furrow, which may be continuous with the costal fracture, a break in C-I-Sc, and distal to which may be formed a distinct triangular cuneus. The clavus, containing the Jst and 2nr/ anal veins. lies adjacent to the scutellum, and in the Panheteroptera General Adult Morphology and Key to Infraorders 43 Fig. 10.3. Wings and wing coupling. A. Forewing, Australostolus monteithi Slys (Aenictopecheidae) (from Stys, 1980). B. Hind wing, A. monteithi (from Stys, 1980). C. Forewing, Triatoma rubrotasciata (De Geer) (from Lent and Wygodzinsky, 1979). D. Hind wing, Notonecta undulata (Say) (from Davis, 1961). E. Wing-to-body coupling in Panheteroptera (from Weber, 1930). Abbreviations: af, anal furrow; al, anal lobe; An. anal vein; C. costa; cs, ciaval suture; Cu, cubitus; cu-pcu, cross vein; h, hemelytron; M, media; m-cu, cross vein; Pcu, postcubitus; R, radius; Rs, radial sector; Sc, subcosta. (see Fig. 10.1) is interlocked with the latter and usually to the opposite wing along the ciaval commissure by the frenum. The clavus is separated from the corium by a line of weakness, the ciaval furrow (ciaval suture), which runs obliquely from the basal articulation of the wing toward the posterodistal margin between the cubitus and the anal veins. The outer portion of the corium (morphologically anterior to R+M) is often expanded and reflexed and referred to as the embolium (or exocorium). Membrane venation may be totally lacking, consist of a few veins, 1 to 5 closed cells, or in some cases—such as some Nepo- r- rpha and many Pentatomomorpha—consist of numer¬ ous anastomosing veins; homology of membrane veins in most Panheteroptera is the subject of substantial dis¬ agreement. The corium plus the membrane is sometimes referred to as the remigium. Hind wing (Fig. 10.3B, D) venation is similar to that of the forewing, but often shows even greater reduction, and the wing is largely membranous except for the veins. In those taxa with the scutellum greatly enlarged the wings are often rather elaborately folded. The hind wings are frequently subject to substantial reduction in size or complete loss. Wing-to-body coupling mechanisms are not elabo¬ rately developed in the Enicocephalomorpha, Dipsocoro- morpha, and Gerromorpha. The forewings in the Pan- 44 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) heteroptera are coupled to the body anterolaterally by the Druckknopfsystem (Cobben, 1957) (Fig. 10.3E) and posteriorly (mesially) by the above-mentioned frenum. Wing-to-wing coupling mechanisms consist of a holding structure on the posterior margin of the forewing (Fig. 12.1C), which provides a sliding attachment for the re¬ curved anterior margin of the hind wing (Teodoro, 1924; Schneider and Schill, 1978; Wygodzinsky and Schmidt, 1991). Variation in wing development is treated in Chapter 6. Legs Heteropteran legs show substantial structural variation within and among groups. Much of the diversity is cor¬ related with life habits (see review by Putchkova, 1979), although certain easily observed structures, such as tri- chobothria in the Miridae, serve functions that are as yet unknown. We treat leg structure in substantial detail, be¬ cause in addition to the above-mentioned points, it is of great importance in the classification of true bugs. Cursorial. The legs of many terrestrial bugs are of the obviously cursorial type, with femora and tibiae elon¬ gate and relatively slender. They serve the functions of walking and running. Saltatorial. Some bugs are capable of jumping. For example, species of Halticus Hahn and Coridromius Sig- noret (Miridae: Halticini) have the hind femora con¬ spicuously enlarged and capable of propelling the bug a substantial distance. In the Saldidae, Gelastocoridae, and Ochteridae the legs are not obviously enlarged, but nonetheless provide substantial propulsive force. Raptorial. The term raptorial has been applied to the forelegs of many heteropteran groups, most par¬ ticularly the obvious predators such as the Phymatinae (Reduviidae), Prostemmatinae (Nabidae), and the true aquatic bugs (Nepomorpha) (Fig. 10.4A). The forefemur is weakly to strongly enlarged and often armed with spines (Fig. 10.4B); the foretibia is also often enlarged or otherwise modified so as to be adpressed against the forefemur in repose. In Phymatinae the foretarsus is non¬ functional in the prey-grasping process, either lying in a scrobe (Fig. 10.4G, H) or being completely absent. The femur in species of Carcinocoris Handlirsch (Carcinoco- rini) projects beyond the tibiofemoral articulation to form a chela in conjunction with the opposable tibia. Modifications of the forelegs in the Enicocephalomor- pha are restricted to the distal end of the tibia and the tarsus; these structures form an opposable grasping appa¬ ratus (Figs. 12. ID, 13.1C, D). As in many of the truly aquatic bugs, the claws are either reduced to only one or are apposed in such a way as to function as a single claw. In many phytophagous Heteroptera, notably the Blissi- nae and Rhyparochrominae (Lygaeidae), the forelegs have the basic structure of the raptorial type. Rather than being used to capture and hold animal prey, they manipu¬ late seeds or other vegetable foodstuffs in the latter group. Their function in the former group is not easily explained by prevailing theories. Fossorial. Only a limited number of bugs spend any part of their lives underground. Most obvious are mem¬ bers of the Cydninae (Cydnidae) (Fig. 69.1C, D). In this group the forelegs are armed with heavy spines, and in some cases the tarsus is inserted proximal to the tibial apex. The tarsus may be absent from the propulsive hind legs, as it is in some species of Scaptocoris Petty. Rowing. All members of the Gerromorpha are capable of walking on the water surface film, but rowing legs are restricted to some members of the Veliidae (Rhagoveli- inae: Fig. 20.3A; some Veliinae) and all Gerridae (Fig. 20.3B-D). The middle legs are longer than the hind legs and serve the function of propulsion on the water surface; also, the claws are usually inserted before the apex of the tibia. Natatorial. Modifications for swimming are of several types. In the Belostomatidae, the middle and hind femora and tibiae are usually flattened (Fig. 30.1) and fringed with hairs (except in the African snail predator Limno- geton Mayr), and both pairs of legs participate in the swimming function. The same is true of most Naucori- dae. The legs of Nepidae show little modification for true swimming, but appear to be better suited to assist the animals in “crawling” through the water (Fig. 31.1). The same can be said of the Pleidae and Helotrephidae. In the Notonectidae (Fig. 38.1) and Corixidae (Fig. 34.1), the hind legs are flattened and fringed with setae and function like oars. Coxae. Schipdte (1869, 1870), followed by Kirkaldy (1906), classified bug coxae as rotatory and cardinate (trochalopodous and pagiopodous, respectively). Most modem authors (e.g., Cobben, 1978) have abandoned this system, observing that the elongate and globose types are not discrete, but are actually the extremes of a con¬ tinuum, and they certainly do not correspond to phyletic lines. The coxae of some Veliidae and all Gerridae are distinctive by virtue of their rotation into the horizontal plane. Trochanters. The heteropteran trochanter is usually simple. In the Miridae it appears to be consistently di¬ vided and forms the point at which the legs can be autotomized (see Dolling, 1991). Femora. The heteropteran forefemur is often swollen and fitted with spines or tubercles associated with grasp¬ ing. The hind femur may be ornamented with large spines (Coreidae) (Fig. 89.1) or enlarged to serve in jumping. Tibiae. The heteropteran tibia is usually straight and cylindrical, although particularly the hind tibia may be fo- General Adult Morphology and Key to Infraorders 45 liaceous in some Coreidae (e.g., species of Diactor Stal) or flattened and bladelike in some Miridae (e.g., species of Pilophorus Hahn, Diocoris Kirkaldy). The tibial shaft is usually adorned with scattered or regularly arranged spines, as long as or longer than the tibial diameter. At least in most Miridae, it is also adorned with several, longitudinal, parallel rows of tiny spicules. Distally on one or more tibiae there may be a grooming comb (Fig. 10.4C, D). The Heteroptera appear to be unique in their posses¬ sion of not only a well-sclerotized tibial flexor sclerite at the base of the tibia, but also a tibial extensor pendant sclerite (Furth and Suzuki, 1990). Tarsi. The heteropteran tarsus shows substantial vari¬ ability (Cobben, 1978). The number of segments is usu¬ ally the same for all legs, although some taxa show variation in this regard. First-instar nymphs of the Enico- cephalomorpha, Dipsocoromorpha, Gerromofpha, and many Nepomorpha have 1 tarsal segment, whereas in the Leptopodomorpha, Cimicomorpha, and Pentatomomor- pha the tarsi are generally 2-segmented. Some groups add segments during nymphal development. Adults most commonly have 3 tarsal segments, although there is sub¬ stantial variation. In a few cases {Scaptocoris spp., some Macrocephalinae, and other Reduviidae) the tarsi may be lost; in others (e.g., Polyctenidae), a fourth (supernu¬ merary) segment is present (Fig. 62.1 A). In many true bugs the ventral surface of the first tarsal segment bears a long slender seta (Figs. 10.4D, 24. ID); the dorsal surface may bear a campaniform sensillum (Fig. 24. ID). Pretarsi. The greatly variable heteropteran pretarsus has long been utilized in classifications, although cer¬ tain structural details are still poorly understood or mis¬ understood. Terminology was reviewed by Crampton (1923) and Dashman (1953), but was not consistently applied within the Heteroptera until about 1970. The most thorough comparative review of these structures was offered by Cobben (1978). The basic units of the pretarsus are two; the unguitrac- tor plate, which is positioned internally at the apex of the tibia (Figs, 29.2D; 53.5C, F) with a proximally attached retractor tendon, and the claws (ungues), whose bases have a flexible attachment to the unguitractor plate and the distal end of the tibia (Fig. 53.5E). These structures are common to all Heteroptera (where a tarsus is present), although in some groups such as the Enicocephalomor- pha and Nepomorpha there is often only a single claw. Several other structures may or may not be present. Pulvilli. Pulvilli are padlike structures arising from the claws (Figs. 10.5G-I, 53.5E); with light micros¬ copy they usually appear white in contrast to the claws themselves, whereas with scanning electron microscopy they are often difficult to distinguish from the claws ex¬ cept for subtle differences in surface texture. Pulvilli are present in nearly all pentatomomorphans (Fig. 10.5G- I), their structure being rather monotonous (Goel and Schaefer, 1970). They are also present in many Miridae (Fig. 53.5E) (where they were referred to as pseudarolia by Knight), Oriini (Anthocoridae), and in Xylastodori- nae (Thaumastocoridae) (Fig. 52.3E) (but see discussion under Accessory Parempodia). Pentatomomorphan pulvilli often possess a distinctly lamellar, grooved, or striate surface. It has been sug¬ gested that this grooved surface produces and serves to conduct lipids across the pulvillus (Ghazi-Bayat and Has- senfuss, 1979, 1980a, b), providing the pads with an adhesive function. The evidence for the origin and nature of these secretions is very limited. Parempodia. Parempodia are usually a symmetrical pair of setiform structures arising from the distal surface of the unguitractor plate between the claws (Figs. 10.5A, 53.5F); some Nepomorpha have either 3 parempodia (Fig. 29.2B) or 2 pairs (Fig. 29.2E). Parempodia are part of the ground plan of the Heteroptera and are observ¬ able in some form in nearly all families, although they are occasionally completely absent (e.g., some Plokio- philidae) (Fig. 10.5E), are modified into low elevations or protuberances (adult Saldidae), or exist in the form of fleshy pads in many Miridae (Fig. 53.5C). The term arolium has been used to refer simply to a “fleshy pad.” In the writings of most European authors, the statement “arolia absent” meant that no fleshy parem¬ podia or accessory parempodia were present (see, e.g., discussion of the construction of the claws in Reuter, 1910), even though homologous setiform parempodia are often present; in some cases, such as in the Tingidae and Thaumastocorinae (Thaumastocoridae), the failure to ob¬ serve the parempodia appears to have been the result of their small size. In the North American literature (e.g., Knight, 1941) dealing with the Miridae, the term arolium was applied to a parempodium of either setiform or fleshy structure, recognizing that the differently formed struc¬ tures were homologous, but using the term arolium in a way not applied to most other insect groups. The function of the parempodia is not well under¬ stood. The setiform type suggests simple mechanorecep- tors associated with the placement of the tarsi and claws. Parempodia in many Miridae are fleshy and have surface structural details similar to those of pentatomomorphan pulvilli and accessory parempodia of other Miridae and some other heteropterans. If the adhesive function theory is correct, then one might postulate a similar function for fleshy parenfpodia. Arolia. These unpaired medial structures arise be¬ tween the bases of the claws, dorsal to the unguitractor plate (contra Goel and Schaefer, 1970; Goel, 1972) and 46 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 10.4. Leg structures. A. Right foreleg, Cryphocricos latus Usinger (Naucoridae). B. Grasping setae on forefemur, Ranatra sp. (Nepidae). C. Tibial combs, apex of middle tibia, C. latus (Naucoridae). D. Apex of tibia and base of tarsus of foreleg, Prostemma sp.; fossula spongiosa, tibial comb, and elongate seta ventrally on tarsal segment 1. E. Detail of fossula spongiosa, Prostemma sp. (Nabidae). F. Fossula spongiosa and tibial comb, foretibia, Nabis sp. (Nabidae). G. Foreleg, Phymata sp. (Reduviidae). H. Same as G, detail. I, J. Foreleg antennal cleaner (I, tibiae; J, femur), Gelastocoris oculatus (Fabricius) (Gelastocoridae). Abbreviations: fs, fossula spongiosa; s, seta; tc, tibial comb; ts, tarsus. General Adult Morphology and Key to Infraorders 47 Fig. 10.5. Pretarsal structures. A. Adult middle leg pretarsus, Oncvlocotis curculio (Kirschbaum) (Enicocephalidae). B. Detail of dorsal arolium, adult middle leg, O. curculio (Enicocephalidae). C. Pretarsus, adu': •^aravelia rescens (Drake and Harrisj, showing dorsal and venr. al arolia (Veli- idae). D. Hino ^eg pretars :., adult Rhagovelia sp. (Veliidae). E. Nymphal pretarsus showing asymmetrical claws and greatly reduced parempodia, Upokophita sp, (Plokiophiiiuae). F. Adult pretarsus, Zetekella minuscula (Barber) (Tingidae) (trom Schuh, 1976), G. Adult pretarsus, Mezira sp. (Aradidae). H. Adult pretarsus, Termitaradus guianae (Morrison) (Termitaphididae). 1. Adult pretarsus, Edessa sp. (Pentatomidae). Abbreviations: cl, claw: da, dorsal arolium; pe, parempodium; pv, pulvillus; va, ventral arolium. are usually present in conjunction with parempodia. The dorsal arolium is found in nymphs and adults of Enico- cephalomorpha (Fig. 10.5A, B), Gerromorpha (Fig. 10.5D), some Dipsocoromorpha and Nepomorpha (Fig. 29.2F), and nymphs and adults of most Leptopodomor- pha (Fig. 41.2A, C). The ventral arolium is present in some Dipsocoromorpha, all Gerromorpha (Fig. 10.5C), and probably in all nymphal Nepomorpha (Fig. 29.2D, F, G). The occurrence of arolia may vary between sexes in the Dipsocoromorpha. Furthermore, arolia may not be present on all pairs of legs in Dipsocoromorpha and Nepomorpha. Arolia are absent in all life stages of all members of th ■ Cimicomorpha and Pentatomomorpha (Cobben, 1978). Accessory parempodia (pseudopulvilli). Some heter- opterans possess paired fleshy structures at the base of the claws. These structures appear to be connected, at least in part, to the unguitractor plate. They were observed by Schuh (1976) and Cobben (1978) in the Bryocorini (Fig. 53.5E, F), Dicyphina (Fig. 53.5F), Monaloniina, and Odoniellina (Miridae). Schuh referred to them as pseudo- pulvilli, and Cobben as accessory parempodia. Cobben (1978) also believed that the “pulvilli” found in Xylasto- dorinae (Thaumastocoridae) were accessory parempodia, a term he also used to describe pretarsal structures found in the Rhagoveliinae (Veliidae). Fossula spongiosa (spongy fossa). This pad of special¬ ized setae is found in many Reduvioidea, Naboidea, and 48 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Cimicoidea. The structure is located apicoventrally on the tibia (usually foretibia, and less commonly on the middle and hind tibiae) (Fig. 10.4D-F). It presumably serves to assist members of these predatory groups in prey capture. Tibial appendix. This structure arises from the distal end of all tibiae in the Thaumastocorinae (Thaumastoco- ridae) (Fig. 52.3C, D) (incorrectly stated as foretibia only in Schuh and Stys, 1991). It was previously referred to as a fossula spongiosa, although clearly the two are not homologous. Drake and Slater (1957) used the term “lo- bate sensory appendage.” The tibial appendix apparently functions in grasping the substrate. Trichobothria. Femoral trichobothria are known in two groups of Heteroptera. In the Gerridae and some Veli- idae a seta arises ventrodistally from the trochanter and one or more setae arise from the femur. They may func¬ tion in communication between conspecifics on the water surface and in prey location. In the Miridae, 3 to 8 (rarely more) trichobothria are located on the lateral and ven¬ tral surfaces of the meso- and metafemora (Figs. 10.81, 53.3E, F) (Schuh, 1976). Viscid setae. Sticky setae are known to occur on at least the foretibiae of some harpactorine Reduviidae. Miller (1956a) described how the bugs smeared resins from trees on their forelegs, the resulting sticky setae aid¬ ing in prey capture. It appears that other taxa actually produce viscid secretions that serve to accomplish the same purpose (Cobben, 1978). Abdomen In its simplest form the pregenital abdomen in the Het¬ eroptera consists of terga 1-7 and sterna 2-7 in females and terga 1-8 and sterna 2-8 in males; these are com¬ monly divided into mediotergites and dorsal laterotergites and sometimes ventral laterotergites (paratergites) and/ or laterosternites, collectively forming the connexivum. More rarely they also are divided into inner laterotergites lying between the dorsal laterotergites and the medio- tergite (Sweet, 1981). Primitively there are 8 abdominal spiracles, spiracle 1 always located on tergum 1, spiracles 2-8 ordinarily located on the respective sternum or latero- stemite; spiracle 1 may be nonfunctional or absent (e.g., Gerromorpha and many Cimicomorpha) as is spiracle 8 in many groups, or the total number of functional spiracles may be further reduced to as few as 3 in some Schizop- teridae. One or more of spiracles 2-8 may be located dorsally, usually on the laterotergites, as in most Lepto- podoidea and many Lygaeidae. Segments 8 and 9 in females and segment 9 in males are incorporated into the genitalia. Segment 10 forms the proctiger, containing the anus, which also bears the remnants of segment 11. Male genitalia. Sternum 9 is developed in most taxa into a distinct genital capsule (pygophore, pygopher) (Fig. 10.1) containing the phallus. The phallus was first studied comparatively by Singh-Pruthi (1926) and subse¬ quently has been examined at a more restricted taxonomic level by Galliard (1935; Reduviidae), Larsen (1938; Nepomorpha), Kullenberg (1947; Miridae, Nabidae), Bonhag and Wick (1953; Lygaeidae), Ashlock (1957; Lygaeidae), Kelton (1959; Miridae), and Davis (1966; Reduviidae). The literature and terminology were re¬ viewed by Dupuis (1970). A welter of terms has been proposed for its various parts. Although showing sub¬ stantial variability, the phallus is usually composed of a basal articulatory apparatus (incorporating the basal plates) to which is attached the aedeagus (intromittent organ) and the parameres (claspers or harpagones). The ductus ejaculatorius passes through the basal foramen into the lumen of the aedeagus. The intromittent organ is formed proximally in most taxa of an at least partially sclerotized phallotheca (theca or phallosoma) into which much of the membranous distal portion of the aedeagus, the endosoma, is often drawn in repose. The endosoma is sometimes formed of a proximal conjunctiva and dis¬ tal (often tubular) vesica, the former of which may bear lobes, spines, and other ornamentation when inflated. Upon passage into the phallotheca, the ductus ejacula¬ torius forms the ductus seminis, which proximally may be modified into an ejaculatory reservoir. The position of the secondary gonopore determines the termination of the ductus seminis and may be in the conjunctiva (e.g., Pentatomoidea), tubular vesica (e.g., Lygaeoidea), or membranous vesica (e.g., most Miridae). Portions of the phallus may be asymmetrical, as also are the parameres in most Nepomorpha, some Cimico¬ morpha, and occasionally in some other groups; one or both parameres may be lost. Asymmetries of pregeni¬ tal abdominal segments occur in Dipsocoromorpha and Nepomorpha. There are normally 2-7 testis follicles. Female genitalia. The hetCropteran ovipositor is com¬ posed of the first (anterior) and second (posterior) val- vulae (gonapophyses, gonostyli), usually fused with the first and second valvifers (gonocoxae, gonocoxopodites), respectively (Scudder, 1959; Dupuis, 1970) (Fig. 10.6). The valifers are derived from abdominal segments 8 (first valifer) and 9 (second). Segment 9 may also bear a pair of third valvulae (gonoplacs, styoids), which form an ovipositor sheath, although these frequently are absent in the Nepomorpha and always are absent in the Penta- tomomorpha and a few other taxa. In many groups the valvulae are elongate and laterally compressed to form a laciniate ovipositor suited for depositing eggs in plant tis- General Adult Morphology and Key to Infraorders 49 Fig. 10.6. Male and female genitalia. A. Aedeagus, Lygaeidae (from Ashlock, 1957). B. Female abdomen and external genitalia, lateral view, Miridae (modified from Davis, 1955). C. Ibid, ventral view. Abbreviations: be, bursa copu atrix; ct, conjunctiva; en, endosoma; ov, ovipositor; ph, phallotheca (phallosoma); rm 1, first ramus; sgp, subgenital plate; sp, spiracle: vlfl, first valifer; vlf2, second valifer; vivl, first valvula; vlv2, second valvula; vlv3, third valvula; vs, vesica. sue, whereas in others the valvulae are greatly reduced or absent and the eggs are laid free or cemented to surfaces, as on plants. Although the ovipositor is morphologically associated only with abdominal segments 8 and 9, it may encroach on sternum 7 and partially or completely divide it. The portion of the internal genitalia associated with copulation, sperm reception, and sperm storage is some¬ times referred to as the gynatrial complex. The sper- matheca (Figs. 12.IG, 16.IB, 40. IJ, 78.2E), or sperm- storage organ, is primitively present in the Heteroptera and shows distinctive structural variation from group to group (Pendergrast, 1957). It is modified into a vermi¬ form gland (spermathecal gland), lost, or nonfunctional in all Cimicomorpha, being replaced by alternative sperm- storage organs, such as paired pseudospermathecae in 50 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) the Pachynomidae, Reduviidae (Fig. 48.3L), and Tingi- dae, and the bursa copulatrix in the Miridae (Fig. 53.3G, I). Heteropteran ovariole numbers vary from at least 2 to as many as 17. Insemination and fertilization are un¬ remarkable, except in the Cimicoidea, in which sperm are usually introduced into the abdominal cavity and mi¬ grate to the oviduct. Fertilization in cimicoids usually takes place before completion of chorionic development, as does some development (Larsen, 1938; Bonhag and Wick, 1953; Davis, 1955, 1966; Scudder, 1959; Dupuis, 1970; Dolling, 1991b). Sound Production and Reception Organs The first systematic survey of sound-producing devices in the Heteroptera was that of Handlirsch (1900a, b). Many individual notices and several summary papers have ap¬ peared since. The function of sounds produced by Heteroptera is less well understood than are the structures that produce them. Sound production in some cases clearly is associated with courtship by males and the readiness of females to mate (Corixidae), agonistic behavior (Corixidae) (Finke, 1968; Jansson, 1976), a defensive repertoire (Reduviidae), and aggregative behavior. Stridulatory Structures Observations of sound production exist for taxa other than those for which putative stridulatory devices are listed below, but no physical mechanism has been found and certainly many additional structures remain to be discovered or are not yet documented in the literature. Most of the following information is derived from the works of Torre-Bueno (1903, 1905), Kormilev (1949), Usinger (1954a), Leston (1957b), Miller (1958), Ash- lock and Lattin (1963), §tys (1961a, 1964a), Jansson (1972), Schuh (1974, 1984), Wilcox (1975), Akingbo- hungbe (1979), Schaefer (1980b), Andersen (1981a), Schaefer and Pupedis (1981), Gogala (1984), J. T. Polhe- mus (1985), Pericart and Polhemus (1990), and J. T. Polhemus (1994). Stridulatory structures in the Heteroptera are usually composed of a movable and a stationary portion. The former was early on termed the plectrum (Fig. 10.7H), and that term has been widely adopted among heteropter- ists. The terminology referring to the stationary portion is more complicated; Ashlock and Lattin (1963) proposed using stridulitrum (Fig. 10.7F, G, I) in place of the many terms that had gone before, such as “pars stridens,” and stridulitrum is now widely used. Forewing edge-hind femur mechanism. This is the most widely distributed and easily observed type of stridu¬ latory apparatus in the Heteroptera. It is known to occur' in some members of the Dipsocoromorpha. Leptopodo- morpha, Cimicomorpha (Miridae), and Pentatomomor- pha (Lygaeidae: Orsillinae, Rhyparochrominae; Largi- dae; Alydidae). Ordinarily the corial margin is either finely transversely striate or finely tuberculate. and the mesal surface of the metafemur is longitudinally ridged or finely tuberculate. Whereas in most known occurrences the stridulitrum is located at the extreme margin of the hemelytron, in the Saldidae it may be located either on the hypocostal or secondary hypocostal ridge, and the peg field of the plectrum may be either proximal or distal on the femur. Connexival margin-hind femur mechanism. Males and females of the veliid genera Angilovelia Andersen and Stridulivelia Hungerford have the stridulitrum located on the connexival margin and the plectrum on the inner surface of the hind femur. Abdominal sternum-hind tibia or femur mechanism. Within a substantial number of Lygaeidae (Cleradini, Myodochini, Ozophorini), Pentatomidae (Macideini), Scutelleridae (Tetyrinae), and some Aradidae (Fig. 64.2L, M) the stridulitrum is composed of striae or minute tubercles located lateroventrally on one or more abdominal sterna, and the plectrum is positioned on the mesal surface of the metafemur or in some Aradidae on the metatibia. A single example in the lygaeid subfamily Blissinae (Heteroblissus anomilis Barber) has a stridu¬ litrum consisting of a series of coarse elongate ridges on the abdomen, while the plectrum appears to consist of a series of small spines on the mesal surface of the metatibia. Propleuron-forefemur mechanism. In nymphs and adults of the North American rhyparochromine lygaeid Pseudocnemodus canadensis (Provancher) the striduli¬ trum consists of an elongate cross-striate area beginning over the foreacetabulum and extending forward to the an¬ terior prothoracic margin; the plectrum is located on the basal third of the forefemur. Metapleuron-middle femur mechanism. This type of stridulatory apparatus is found in some calisiine Aradi¬ dae. Head-forefemur mechanism. A lunate stridulitrum occurs laterally on the head in several genera of the Colo- bathristidae (e.g., Peruda Distant, Trichocentrus Hor¬ vath), the plectrum consisting of a small field of scattered tubercles on the mesal surface of the forefemur. Forecoxa-forecoxal cavity mechanism. In Ranatra spp. (Nepidae) a roughened elevated area on the outer surface of the forecoxa is rubbed against the striate inner surface of the foreacetabulum. Tibial comb-labial prong mechanism. Males of most species of Buenoa Kirkaldy and Anisops Spinola (Noto- General Adult Morphology and Key to Infraorders 51 stridulatory structures. A. Apex of nymphal foretibia, Laccocoris sp. (Nauooridae). B. Apex of labium, Prostemma sp. (Nabidae). C. Peg plate, Paravelia rescans (Veliidae). D. Pit receptors with associated specialized setae, on thorax, Rhagovelia sp. (Veliidae). E. Pregenital ventral abdominal organ, Hydrometra sp. (Hydrometridae). F. Prosternal stridulatory sulcus, Phymata sp. (Reduviidae). G. Same as F, detail. H. Metafemoral stridulatory plectrum, Hallodapus albofasciatus (Motschulsky) (Miridae) (from Schuh, 1984). I. Ventral abdominal stridulitrum, Ugyrocoris diffuses (Uhler) (Lygaeidae). Abbreviations; pp, pegplate; pr, pit receptor (pit organ); ss, specialized seta. 52 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) nectidae) have an elongate elevated area proximally on the inner surface of the foretibia, on which lies a comb of flattened setae. This comb is apparently rubbed against prongs located on either side of labial segment 3. Femoral ridge-coxal peg mechanism. Males of most Buenoa spp. (Notonectidae) have on the inner surface of the forefemur a series of approximately parallel ridges. These apparently are rubbed against a short heavy peg located on the lateral face of the forecoxa. Base of labium-femoral apex mechanism. Males of most Buenoa spp. (Notonectidae) possess several stout setae distally on the inner surface of the forefemur. These may be plucked against the base of the labium. Maxillary plate stridulitrum. Males and females of most Corixinae have the stridulitrum formed by the acute edge of the maxillary plate. The plectrum consists of a field of specialized pegs on the basomesal surface of the forefemur. The device is usually better developed in males, which are capable of making sounds louder than those made by the females. Metathoracic wing vein stridulitrum. A variety of pentatomomorphans, and some other bugs, possess stri- dulatory structures combining a striate vein or accessory vein of the hind wing with a striate area on the tho¬ racic or abdominal dorsum (Leston, 1954a). All species of Kleidocerys Stephens (Lygaeidae: Ischnorhynchinae) have an arcuate veinlike ridge located on the underside of the hind wing. This ridge contacts a transversely stri¬ ate longitudinal ridge located near the lateral margin of the metanotum. In Piesma quadrate Fieber (Piesmatidae) the underside of the cubitus is transversely striate basally, making contact with a striate area located anterolaterally of abdominal tergum 1. In the Tessaratomidae, Cydnidae, some Scutelleridae, Thaumastellidae, and the Leptopodi- dae, the postcubitus, vannus, or other area of the hind wing has a striate area that interacts with a striate area on abdominal tergum 1. Hypocostal lamina (or articulatory sclerite) stridu¬ litrum. Males and females of some Coreidae have a stri¬ ate ridge located posterolaterally on the undersurface of the pro thoracic foramen. It functions in conjunction with a stridulitrum located either on the hypocostal lamina or on the articulatory sclerites of the forewing. Underside of clavus stridulitrum. Rhytocoris spp. (Coreidae) have the stridulitrum located on the underside of the clavus; the plectrum is situated on the articulatory region of the hind wing. Posterior margin of pygophore stridulitrum. In the Scutellerinae (Sphaerocorini) the posterodorsal margin of the pygophore is fitted with six rows of short stout setae, which may produce sound in conjunction with the use of vesical conjunctival appendages as a plectrum. Prosternal groove stridulitrum. A transversely stri¬ ate prostemal stridulitrum exists as a groove in nearly all members of the Reduviidae, the apex of the rostrum functioning as the plectrum. Other Sound-Producing Organs Tymbals. The dorsal surface of the abdomen may function as a tymbal (or drum), whereby terga 1 and 2 are usually fused together and vibrate at low frequen¬ cies over a hollow chamber within the abdomen. This feature occurs widely in those Pentatomidae and Acan- thosomatidae that lack stridulatory structures associated with the abdominal dorsum, as well as in the Cydnidae, which have a stridulitrum anteriorly on the abdominal dorsum. Similar tymbals also appear to exist in at least some members of the Lygaeidae, Coreidae, and possibly Reduviidae. Sound Reception Scolopophorous organs. Scolopophorous organs are known to occur in many groups of insects. Those located on the meso- and metathoracic pleuron and on abdomi¬ nal terga 1 and 2 of most nepomorphan families (Larsen, 1957; Mahner, 1993), including the Gelastocoridae (Par¬ sons, 1962), appear to represent a set of structures dis¬ tinctive to the infraorder. They are composed of sensory sensilla (scolopidia), which may be single or united into a single strand, whose distal end attaches to a membrane on the body wall or to the body wall itself. The proximal end is innervated and sometimes associated with the tracheal system. These organs are thought by some to be auditory in function, although Larsen believed that they might be involved in orientation in the aquatic Nepomorpha. The scolopophorous (scolopidial) organ, located on the tarsus of all 3 pairs of legs in Notonectidae, is sensitive to wave frequencies of 20-50 Hz, a range typical of that produced by prey of some Notonecta spp. (Wiese, 1972). Tympanal organs and physical gills. Tympanal organs exist on either side of the body in the mesothorax under the hind wing articulation in the Corixidae (Hagemann, 1910) and are in contact with the physical gill air bubble (Prager and Streng, 1982). They are innervated by two scolopidia, which are stimulated by airborne sound, those on opposite sides of the body differing in their responses (Prager, 1973, 1976). One of the scolopidia in each tym¬ panal organ is sensitive to stridulation frequencies pro¬ duced by conspecific bugs. Thus, there appears to be a credible description for a hearing method in the Corixi¬ dae. Equally plausible hearing methods are not available for most other families that are known to stridulate. Mechanoreceptors The Heteroptera possess a wide variety of mechano¬ receptors, most of which are of a type found in nearly General Adult Morphology and Key to Infraorders 53 Fig. 10.8. Trichobothria. A. Cephalic trichobothrium, Rhagovelia sp. (Veliidae). B. Scutellar trichobothria, Prostemma sp. (Nabidae), C. Ab¬ dominal trichobothria, Edessa sp. (Pentatomidae). D. Medial abdominal trichobothria, Paragonatas divergens (Distant) (Lygaeidae). E. Detail of base of lateral abdominal trichobothrium, P. divergens (Lygaeidae) (from Schuh, 1975a). F. Trichobothrium, ventrally on posterolateral margin of laterotergite of abdominal segment 7, Nabis sp. (Nabidae). G. Antennal segment 2 showing trichobothria, Barce sp. (Reduviidae, Emesinae) (from Wygodzinsky and Lodhi, 1989). H. Base of antennal trichobothrium, Aphe/onofus sp. (Pachynomidae) (from Wygodzinsky and Lodhi, 1989). I. Base of femoral frichobothrium, Sthenaridea australis (Schuh) (Miridae) (from Schuh, 1975a). Abbreviations: sp, spiracle; tb, trichobothrium. 54 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) mcc Fig. 10.9. Metathoracic scent-gland configurations (after Carayon. 1971a). Abbreviations: ab. abdomen: ag. accessory gland: Ig. lateral gland: mcc, metacoxal cavity; o, orifice; r, reservoir; th, thorax. all other groups of insects. The function of many of these is poorly understood or unknown. Notable among the mechanoreceptors are trichobothria (described earlier), specialized setae that have been implicated in sound re¬ ception, but for which function there is no solid evidence. They arc distributed on many body regions in a great number of heteropteran families. Other mechanorecep¬ tors worth mention are the static sense organs found on the abdominal sternum of Nepidae and Aphelocheiridae. Most such structures are documented and discussed in the family treatments. Although no comprehensive review of mechanoreceptors exists specifically for the Heteroptera. Grasse (1975) provided a good general introduction to the subject. Exocrine Glands The monophyly of the Heteroptera is based in part on their pos.session of scent glands, located on the abdominal dorsum in nymphs and in the metathorax of adults (Fig. 10.9). Like most insects, the Heteroptera communicate through the use of chemicals, and additional glandular structures continue to be discovered in the group. Most of the following summary is derived from information con¬ tained in the excellent review articles of Carayon (1971a). Cobben (1978), Staddon (1979), and Aldrich (1988). Additional references are cited below in the text. Scent reception is much less well understood than are glandular structure and glandular products. Calla- General Adult Morphology and Key to Infraoraers 55 Fig. 10.10. Glandular and other structures. f/tetathoracic pleuron showing scent-gland auricle and evaporatory area, Pilophorus sp. (Miridae) (from Schuh, 1984). B. Metathoracic seen; “land evaporatory area, Pronotacantha sp. (Berytidae). C. Scent-gland channel on metathoracic pleuron. Termitaradus guianae (Termitaphiaiaae). D. External manifestation of coriai glands, Upokophila eberhardi Schuh (Plokiophilldae), E. Ocular seta of nymphal instar 1, L eberhardi (Plokiophilldae). Abbreviations: au, auricle; eg, corial gland; ea, evaporatory area; os. ocular seta; sgc, scent-gland channel. han (1975) reviewed available evidence and proposed the theory that specialized setal receptors on the antennae of insects serve as “antennae” to receive specific frequency emissions of various semiochemicals. Grasse (1975) pro¬ vided a general review of chemoreceptive organs in in¬ sects. although nothing exists specifically for the Heterop- tera. Nymphal dorsal abdominal scent glands. The nymphs of nearly all Heteroptera have scent glands located on the abdominal dorsum. These structures appear to have first been investigated by Guide (1902); Cobben (1978) summarized their number and distribution in heteropteran families. They vary in number from 1 to 4 and are located at the anterior margin of terga 4-7. The most widespread condition is to have 3 functional glands, with their ori¬ fices located on the anterior margins of abdominal terga 4, 5, and 6 (3/4, 4/5, 5/6), this arrangement being found commonly in the Pentatomomorpha, but also occurring in many cimicomorphan families (except Miridae, 3/4, and Tingidae, 3/4, 4/5; Reduviidae show substantial varia¬ tion). The Gerromorpha, Leptopodomorpha. and Nepo- morpha, have either a single functional gland (3/4) or no glands at all, with the exception of the Corixidae, which typically have 3 functional glands (3/4. 4/5, 5/6). Ordinarily, functioning glands are lost from posterior to anterior, with the exception of the Coreoidea. where only glands 4/5 and 5/6 are functional. No taxa have 4 func¬ tional glands, and only Joppeicidae and some Schizop- teridae have functional glands between terga 6/7. The scent glands are multicellular, representing invagi- 56 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) nations of the cuticle. They may be single or divided and may open through a single ostiole or a pair of ostioles. Muscles open the ostiole as well as compress the gland. Some taxa are capable of ejecting secretions from the glands as a spray. Chemical components produced by these glands vary, but typically they include compounds such as 2-hexenal. 4-oxo-2-hexenal (or the corresponding octenal), and 2- hexenyl-acetate; the list of other substances is large and constantly growing. The glands appear for the most part to serve defensive or alarm functions (e.g., Calam and Youdeowei, 1968; Ishiwatari. 1974). although they may also be involved in the aggregation behavior seen in the nymphs of some taxa or serve a fungistatic role, as in the subterranean cydnid Scaptocoris divergens (Roth, 1961). Dorsal abdominal glands in adults are usually absent or represented only by a scar, or they are nonfunctional and become smaller over time. In a few taxa they are well developed and functional. For example, in males of species of the predatory asopine pentatomid Podisus the 3/4 dorsal abdominal glands are enormous, producing a mixture of chemicals that serve as a powerful sex phero¬ mone. These substances also function as kairomones, attracting scelionid egg parasitoids (Aldrich, 1988). The dorsal abdominal glands also remain active in adult Ser- inethinae (Rhopalidae) of the genera Boisea Kirkaldy and Jadera Stal (Aldrich et al., 1990). The two glands pro¬ duce different compounds in adult Jadera spp., but the composition remains the same throughout nymphal and adult life and is the same for males and females, sug¬ gesting a pheromonal role, possibly in aggregation, in these aposematically colored bugs. Sexual attraction by functional abdominal scent glands in adults has been sug¬ gested for some Miridae, but Aldrich et al. (1988) were unable to experimentally substantiate that thesis with Ly- gus spp. and suggested that other as yet unrecognized glands were performing this function. Adult metathoracic scent glands. Most adult Heterop- tera have active scent glands located ventrally in the metathorax. The glands are integumentary invaginations. There may be a single gland with a single opening, a single gland opening into a pair of ostioles, or paired glands opening into closely to widely spaced ostioles. The glands themselves may consist of a single cuticle-lined corpus that produces and stores the scent fluid. More fre¬ quently, however, there is a reservoir with glands empty¬ ing into it. Sometimes there is also an accessory gland attached to the reservoir; this gland apparently serves to deliver enzymatic reaction products important in produc¬ tion of the final scent-gland chemicals. Dimorphism may exist between the sexes. For example, in several groups of Lygaeidae (Carayon, 1948b) and in the Lethocerinae (Belostomatidae) the glandular structures of the males are more highly developed than are those in the females'. In some Enicocephalomorpha the glands are apparenth present only in the males (Carayon. 1962). The gland ostioles are always located ventrallv on the metathorax and have a.ssociated with them a \a!ve and a valve-opener muscle. The orifice is connected to a groov e or canal that runs laterally onto the metathoracic pleuron. ordinarily terminating in an area termed the peritremc. that area usually surrounded by the cvaporarory area. which is particularly well developed and of distinctive mushroomlike microsculpture (termed "flake cuticle" by Johansson 11957a]) in the Cimicomorpha and Penta- tomomorpha (Fig. lO.lOA, B) (Filshie and Waterhouse. 1969; Johansson and Braten, 1970; Carayon. 1971a). The distinctive structure of the evaporatorium is sometimes found on areas of the body such as the region around the thoracic spiracles (Remold. 1962. l963;Schuh. 1984; Carver. 1990). Two theories have been proposed for the function of the evaporatory area. One suggests that the modified cuticle offers a greater surface area from which the scent fluid can evaporate, thus increasing the effectiveness of the glands in defense. The other contends that the specialized evaporatory surface serves in restricting spread of the scent fluids to a circumscribed area of the body because the scent fluids are known to be toxic to the bugs that secrete them as well as to potential predators (Remold. 1962, 1963; Johansson. 1957a; Johansson and Braten, 1970; Carayon. 1971a; Carver, 1990). The observations of Falkenstein (1931) on Tessaratoma papillosa Drury in¬ dicate clearly that although the scent-gland secretions of both nymphs and adults serve a defensive function, they are also extremely toxic to both life stages when coming into contact with the cuticle of the individual that secretes them. The main function of the adult scent glands appears to be defensive, as is the case in nymphs, although sexual, alarm, and aggregation functions may also exist. The composition of the glandular products is complex, but they often comprise substances similar to those listed above for the nymphs. Several authors have shown that the shape of the peri- treme and the shape and distribution of the evaporatorium have systematic value, as for example in the Cydnidae (Froeschner, 1960; Dethier, 1974) and .Miridae (Cassis. 1984). Brindley’s glands. This pair of simple glands is located in many adult reduviids and pachynomids dorsolaterally in the anterior region of the abdomen: a gland of similar structure and location is found in the Tingidae. Brindley’s gland opens onto the metepisternum. where in Rhodnius prolixus Stal a channel with a spongy surface runs some distance from the orifice of the gland. The main compo- General Adult Morphology and Key to Infraorders 57 nent of the glandular secretion in Rhodnius appears to be isobutyric acid, although paraffins may also be present. The secretions may serve to alert other individuals of danger in the case of aggregations of Rhodnius; they are nontoxic to adults, but toxic when administered to the cuticle of nymphs (Kalin and Barrett, 1975). Brindley’s glands are not present in all reduviids, although when present the metathoracic scent glands are often inactive. Ventral abdominal glands (paragenital glands; ura- denies) of the Pentatomomorpha. “Ventral abdominal glands” were observed in female Pyrrhocoris apterus by Dufour (1833), who concluded that they were part of the reproductive system. They are now known to exist in the females of some members of the Lygaeidae (Lygaeinae, Orsillinae, Rhyparochrominae). Largidae, Pyrrhocori- dae, Stenocephalidae, Coreidae, Alydidae, and Rhopali- dae and in males of at least the Lygaeidae, Coreoidea, and Stenocephalidae, although apparently not in the Pyrrho- coroidea. These large multicellular glands have a single ventral opening in the intersegmental membrane between sterna 7 and 8, 8 and 9, or more rarely 9 and 10, depend¬ ing on the species and the sex (Thouvenin, 1965). Gland secretions have been identified in the males of at least 10 species of Coreidae and a few Rhopalidae. Males produce volatile compounds, whereas females apparently do not. Aromatic compounds with the odors of cherries, roses, and other familiar substances are common in Leptoglossus spp., but aliphatic alcohols and esters are also produced by the coreids (Aldrich, 1988). The secretions of the ventral abdominal glands play a role in mate recognition, at least in the Rhopalidae (Aldrich et al., 1990). Ventral abdominal glands of male Anthocoridae (Fig. 60.2E). Some male Anthocoridae of the tribe Scolopini possess internally paired glands with a single or double orifice opening onto abdominal sternum 4 or 5. At present nothing is known of the chemicals they produce or their function, although the latter is presumed to be sexual. At the orifice or between the orifices there is a brush of setae. Carayon considered these structures to be serially ho¬ mologous with the ventral abdominal glands (uradenies) occurring in certain female Pentatomomorpha. He also suggested that they might be derived from the secretory integumentary cells found in some members of the Lasio- chilidae and Lyctocoridae (Carayon, 1954, 1972a; Peet, 1979). Eversible glands in abdomen of Saldidae. In both sexes of all members of the family Saldidae there exists a pair of glands that arise ventrolaterally in the membrane between abdominal segments 7 and 8. First observed by Cobben (1961), they can be everted by gently squeez¬ ing the abdomen of live or alcohol-preserved specimens. To date no function has been ascribed to them, and the composition of their secretions has not been determined. Ventral (sternal or Carayon’s) glands in Reduviidae. These paired, simple, saclike glands, known from the Elasmodeminae, Holoptilinae, and Phymatinae. are lo¬ cated in a position similar to the metathoracic scent glands, but each individual gland opens ventrally into the membrane between the thorax and abdomen. Despite the similarity of position to the metathoracic scent glands, the location of the glandular exit, structure of glands, and existence of both ventral glands and metathoracic glands in adults of Phymata Latreille (although both glands are not present in all taxa possessing ventral glands) attest to their distinctness. The secretions of these glands are described as oily, although their specific composition and function are unknown (Carayon et al., 1958; Staddon. 1979). Subrectal gland in the Harpactorinae. Many female Harpactorinae (Reduviidae) have large bilobed saclike glands opening into the membrane between the styloids and the anus (Barth, 1961). When the bugs are disturbed, the bright orange glands are everted like a caterpillar osmeterium, producing a nauseatingly sweet odor (Barth. 1961; Davis, 1969; Aldrich, 1988). Dermal glands. So-called type B dermal glands were first discovered and described in Rhodnius (Reduviidae: Triatominae) (Wigglesworth, 1933, 1948). They are com¬ posed of four cells (Lai-Fook, 1970), one of them assum¬ ing the secretory function, and they show their greatest activity at the time of molting. They have therefore been thought by most authors to play a role in the process of ecdysis or of cuticle formation (Wiggles¬ worth, 1933, 1948; Baldwin and Salthouse, 1959; Lai- Fook, 1970). Such glands appear to be widespread over the cuticle of Rhodnius, and similar glands occur in other orders of insects, lending credence to the ecdysial- function theory. “Floral glands” and “socket glands” have been de¬ scribed in Dysdercus fasciatus Signortl by Lawrence and Staddon (1975). Floral glands are multicellular, whereas the socket glands are unicellular. Both types are most common on the abdominal venter, and in Dysdercus ap¬ parently are confined to the adult male, strongly suggest¬ ing a pheromonal function. Males and some females of most groups of Pentatomoi- dea (e.g., Pentatomidae, Plataspidae, Acanthosomatidae. and Scutelleridae) possess pheromone-producing unicel¬ lular tegumentary glands laterally on 3 or more abdominal sterna (Carayon, 1981). A special type of such a gland is found in males of several genera of Scutelleridae (e.g.. Tectocoris Hahn, Psacasta Germar, Odontoscelis Laporte. and Irochrotus Amyot and Serville) and in at least some Oxycareninae (Lygaeidae). Referred to as an androco- nium, each gland consists of a single blind cell set in an alveolus. The contained aphrodisiacal substance, which 58 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 10.11. Salivary glands and alimentary canal. A. Salivary gland, Dictyonota strichnocera Fieber (Tingidae). B. Salivary gland, Stollia fabricii Kirkaldy (Pentatomidae) (A, B from Southwood, 1955). C. Alimentary canal, Hypselosoma hirashimai Esaki and Miyamoto (Schizopteridae). D. Alimentary canal, Megymenum gracilicorne Dallas (Dinidoridae) (C, D Miyamoto, 1961a). Abbreviations: ag. accessory gland; e, esophagus (foregut); gc, gastric caeca; hg, hind gut; mdg, midgut; mg, main gland: mt, Malpighian tubules; p, pylorus; sg, salivary gland. is produced by an endocuticular cell, can be released only if the androconium is ruptured in a fragile zone (Carayon, 1984). Corial glands (Figs. lO.lOD, 58.2B). These unicel¬ lular glands cover the corium, clavus, and cuneus in the Plokiophilidae. They were first observed by China (1953), who termed them “tubercular sense organs,” their glandular function having been determined by Cara¬ yon (1974). They may play a role in maintaining a com¬ mensal relationship with the bugs in the webs of spiders and embiids. Salivary Glands and Alimentary Canal The classical work on the salivary glands is that of Baptist (1941) with additional useful comparative and structural information in the works of Barth (1954), Kumar (1967), Nuorteva (1956), and Southwood (1955), among others. The broad outlines of gut structure in the Heteroptera were presented by Dufour (1833). Details of the anatomy and function of the alimentary canal can be found in the works of Goodchild (1963) and Miyamoto (1961a). Salivary Glands The salivary glands (Fig. 10.11 A, B) in the Heterop¬ tera are positioned anteriorly in the thorax and usually are adpressed to the alimentary canal. The general struc¬ ture consists of a main gland with 1-several lobes, most commonly 2—4, connected by an elongate slender duct to a tubular or vesicular accessory- gland. The ducts of the main glands unite to form a common salivary duct, which leads to the salivary pump and hence to the salivary canal formed by the maxillary stylets. The main glands of the Heteroptera, as opposed to those of the Sternorrhyncha and Auchenorrhyncha, possess a distinct lumen that holds the secretory products produced in the gland walls. The anterior portion of the main gland appears to pro¬ duce sheath material that fixes the labium to the substrate at the point of insertion, although the glands may also produce substances that serve to lubricate the stylets. Such sheaths, or at least attachment flanges, appear to be best developed in the Heteroptera among the Penta- tomomorpha, but they are also known in other groups, for example, predators such as the Reduviidae (Friend and Smith, 1971). The anterior lobes, which appear to pro¬ duce zootoxic secretions important in immobilizing prey General Adult Morphology and Key to Infraorders 59 in the Reduviidae, are often greatly reduced in hemato- phagous species (Haridass and Ananthakrishnan, 1981). The posterior lobe (or lobes) of the main gland apparently produce hydrolyzing enzymes, which are proteolytic in the case of obligate predators. In the case of hemato- phagous species such as the Triatominae (Reduviidae), the saliva does not cause paralysis, but serves as an anti¬ coagulant, the bite of the bug being undetected by the vertebrate host. The glands of taxa that feed primarily in the plant vascular system (most Sternorrhyncha and Auchenor- rhyncha, many Pentatomoidea and Coreidae) generally contain no hydrolyzing enzymes (at least when feeding in the phloem vessels), whereas those of taxa that feed on plant cell contents and seeds (e.g., Miridae, Tingidae, many Lygaeoidea, some Coreoidea) generally produce such enzymes. Substances that initiate gall formation are also produced in the salivary glands, such as in certain Tingidae (see Chapter 54). The accessory glands may function as excretory or¬ gans, usually dispensing a dilute liquid, which apparently is extracted from the hemolymph. The function of this substance is not well understood, although it may serve to suspend materials being imbibed by so-called lacerate- flush feeders, in which case the accessory glands are usually better developed than in the stylet-sheath feed¬ ers; it might alternatively serve to dilute the concentrated secretions of the main glands. Maxillary glands. These glands lie anteriorly in the head in a cavity formed by the proximal end of the maxilla and mesad of the maxillary protractor muscles. The glands consist of large peripheral cells and numer¬ ous smaller central cells. Chitinous ducts lead from the peripheral cells to a collecting reservoir on the floor of the preoral cavity which opens into the space occupied by the mandibular and maxillary stylets (Linder, 1956). Nymphal thoracic glands. All nymphal instars of all Heteroptera possess thoracic glands, elongate bodies of heavily nucleated cytoplasm lying in close association with the salivary glands and attached to the accessory gland or the main gland. They are derived from an em¬ bryonic invagination of the second maxillary segment. During each nymphal instar the glands go through a phase of swelling, glandular discharge, and shrinkage. These cyclic changes appear to be mediated by secretions from the brain, the glands functioning in the hormonal control of molting. They atrophy and disappear within 72 hours of the imaginal molt (Wells, 1954). Gut The heteropteran alimentary canal (Fig. 10.11C, D) is formed of 3 principal divisions: foregut, midgut, and hindgut. The structure in nymphs and adults is essentially similar. The foregut consists of an elongate, chitin-lined esophagus connected to the food canal (Fig. 10.2A) of the maxillary stylets via the cibarial pump located dorsally in the head. The esophagus proper varies greatly in length, but it is never differentiated into a true crop as found in many insects. The midgut, which is endodermal in origin, is vari¬ ously subdivided, depending on the group. It comprises 2 or 3 sections in most groups, but in the Pentatomo- morpha it often is divided into 4 or rarely 5 sections. The most anterior portion of the midgut forms a capa¬ cious chamber, or stomach, in all bugs. The remainder of the midgut may take on various forms, the penultimate section bearing 2-4 rows of gastric caeca in most Tricho- phora, except in the predatory Asopinae (Pentatomidae) and many seed feeders. The midgut of the Phyllocephali- nae (Pentatomidae) forms 2 i filter chamber similar to that found in many Auchenorrhyncha, apparently uniquely among the Heteroptera. Some pentatomomorphans have a discontinuous mid¬ gut. This situation was investigated in detail by Goodchild (1963), who prepared histological sections of the guts of a relatively large number of taxa. The midgut in many sap-feeding taxa is occluded at a point just anterior to the section that contains the gastric caeca. There is a liga¬ mentous connection between the anterior and posterior sections, but Goodchild found no lumen. This distinctive gut morphology exists in both nymphal and adult forms, but how it functions is not understood. The pylorus (Miyamoto, 1961a), or ileum (Goodchild, 1963), is not lined with chitin, and therefore appears to represent part of the true midgut, rather than a por¬ tion of the hind gut as asserted by some authors. It may be well developed, particularly in the Pentatomomorpha, to virtually absent. The Malpighian tubules, which vary greatly in length and general conformation, arise from the pylorus, as an anterior and posterior pair in the Penta¬ tomomorpha and as a dorsal and ventral pair in most other Heteroptera. Between the pylorus and the rectum there is generally a conspicuous valve, the pyloric valve (Myiamoto, 1961a:201), or ileorectal valve (Goodchild, 1963), composed of a cone of small columnar cells and surrounded by many strong circular muscle fibers. The chitin-lined true hind gut consists primarily of the rectum, which is often pear-shaped. At least anterodor- sally, if not to a much greater extent, it is modified into a rectal gland, or rectal pad, a relatively extensive area of specialized cells, which are thought to play an important role in extraction of water from the hemolymph. Symbiotic Microrganisms Gut symbionts. The gastric caeca in the Pentatomo¬ morpha are filled with bacteria, which serve an undeter¬ mined function. Bacteria from a sample of bugs were 60 TRUE BUGS OF THE WORLD (HEMIPTERA; HETEROPTERA) described as species of Pseudomonas by Steinhaus et al. (1956). These symbionts are generally transmitted by the females’ depositing symbiont-laden material in cap¬ sules among eggs or on freshly laid eggs (Carayon, 1951; Buchner, 1965). In the ovoviviparous rhyparochromine (Lygaeidae) Stilbocoris, however, the female deposits this material on the head of the freshly laid nymphs. The nymphs subsequently use the rostrum to imbibe the sym¬ bionts from the droplet (Carayon, 1963). Mycetomes. Although gastric caeca are present only in most Pentatomomorpha, symbiotic microorganisms nonetheless appear to be present in most Heteroptera. They are harbored in the mycetomes (Buchner, 1921), which occur midlaterally in the abdomen in Cimicidae. In the male they are attached to the vas deferens, and in the female they lie unattached (Usinger, 1966). Trans¬ mission is apparently transovarial, and the identity of the symbionts is not agreed upon by all authors. Intracellular symbionts. Intracellular symbionts are also known from a few Heteroptera, although, as for those described above, their function is not well understood. Nervous System The gross structure of the heteropteran nervous system was first studied comparatively by Brandt (1878) and later in greater detail by Pflugfelder (1937). The following ac¬ count has been prepared from the works of Johansson (1957b), Parsons (1960a), and Livingston (1968). Other basic studies include those of Hamilton (1931), Rawat (1939), and Guthrie (1961). Central nervous system. Like other phylogenetically more advanced insects, the Heteroptera show a great amount of ganglionic fusion. The so-called brain, or supraesophageal ganglion, is vaguely differentiated into a protocerebrum, deutocerebrum, and tritocerebrum. The first of these includes the optic lobes, which innervate the compound eyes and ocelli. From the second ema¬ nate both sensory and motor nerves for the antennae. The tritocerebrum and subesophageal ganglion show a strong degree of fusion in most Heteroptera. It is from this re¬ gion that the mouthparts, salivary glands, and associated musculature are innervated. The first thoracic ganglion is more or less distinct from the subesophageal ganglion, depending on the taxon. From it emanates foreleg innervation. The “posterior ganglionic mass,” or central ganglion, represents the fusion of the mesothoracic ganglion, metathoracic gan¬ glion, and all abdominal ganglia, supplying nerves to the wings, posterior 2 pairs of legs, and to the abdomen. There are fewer obvious pairs of nerves than abdominal segments, the largest pair being that which innervates the genitalia. Stomodaeal nervous system (stomogastric or sympa¬ thetic nervous system) (Fig. 10.12B). The. frontal gan¬ glion, which has a nervous connection with the trito¬ cerebrum, innervates the pharyngeal dilator muscles via the procurrent nerve. It is connected via the recurrent nerve with the hypocerebral ganglion, which may inner¬ vate the esophagus, and in some taxa is reported to have a nervous connection with the corpus cardiacum. Endocrine glands (neurosecretory system). This spe¬ cialized portion of the nervous system consists of the corpora cardiaca and the corpora allata; in some groups each exists as a single structure rather than in the paired condition. These structures are connected to the central or stomodaeal nervous system or both. During the first four nymphal stadia the corpora allata secrete juvenile hormone, which causes the insect to retain its imma¬ ture form. The corpora cardiaca produce prothoracotropic hormone, which acts on the prothoracic gland to produce ecdysone, which causes metamorphosis into the adult form (Wigglesworth, 1984). Circulatory System Information on the insect dorsal vessel is taken from the following references; Hamilton, 1931; Malouf, 1933; Wooley, 1951; Hinks, 1966; and Miyamoto, 1981. The insect circulatory system shows great similarity across a wide range of taxa. In the Heteroptera it consists of the dorsal vessel (Fig. 10.13A, B), which lies close against the abdominal and thoracic dorsum. This struc¬ ture may be divided into a posterior heart and an anterior aorta. Hemolymph enters the heart through 2-7 pairs of ostia, which may lie in abdominal segments 1-7. The pumping action is peristaltic, being mediated by the alary and intrinsic musculature. Most terrestrial Heteroptera have 3 pairs of alary muscles, whereas the Nepomorpha usually have more, with a maximum of 8. The intrinsic musculature may be arranged in a circular or helical fash¬ ion. The Hemiptera and some other Acercaria are unique among the Insecta in generally having the heart restricted to the posterior portion of the abdomen. The anterior aorta is devoid of ostia and extends forward into the head capsule, often lying close along the dorsal surface of the alimentary canal and emptying over the supraesophageal ganglion of the nervous system. The dorsal vessel is generally lined with specialized cells known as nephrocytes; these often are several layers thick and more prominent in the posterior region. They apparently function in the removal of waste substances from the hemolymph. Innervation of the heart wall and the alary muscles was reported by Hinks (1966), although the source of the nerves was not mentioned. Tracheation of the dorsal vessel is restricted to the heart region and may be largely dorsal or ventral, depending on the taxon. The Malpighian tubules are usually intertwined with the alary muscles in close association with the dorsal vessel. General Adult Morphology and Key to Infraorders 61 Fiq, 10.12. Nervous system. Gelastocoris oculatus (Fabricius) (Irom Parsons, 1960a). A. Central nervous system. B. Detailed ventral view o! cerebral area. Abbreviations: eg, central ganglion; d, deutocerebrum; fg, frontal ganglion; ftg, first thoracic ganglion; ol, optic lobe; p. protocerebrum; sg, subesophageal ganglion. Eggs Heteropteran eggs have been known since the time of Leuckhart (1855) to show substantial morphological vari¬ ability. and particularly those of the pentatomoids have attracted attention because of their bright coloration and deposition on plant surfaces. Major modern reviews, from which most of the following material is taken, have been prepared by Southwood (1956) for terrestrial Het- eroptera and Cobben (1968a) for all groups (see also works of Putchkova referenced in Cobben). The work of Ren (1992) contains many superb scanning micrographs of eggs, showing the details of their surface structure. The eggs may be elongate (Fig. 10.14C), cylindri¬ cal and sometimes weakly curving (Fig. 10.14A, B), or barrel-shaped (Fig. 10.14D). Eggs are inserted into plant tissue, cemented to a surface, or laid free. The shell, or chorion, is laid down by follicle cells in the telotrophic ovaries, after the yolk has completely de¬ veloped, and except in the case of the Cimicoidea, before fertilization. The chorionic surface may range from shal¬ lowly pitted to a well-formed honeycomb, or it may even be rugose. There may or may not be a distinct operculum, or egg cap, present at the cephalic pole, a feature strongly correlated with the type of eclosion mechanism. A true operculum, with a line of weakness referred to as a seal¬ ing bar, is present in the Cimicomorpha, all of which lack a true egg burster; eclosion takes place via pressure from within the egg and has been described by Cobben (1968a) as either embryonic or serosal. Most other groups have an egg burster present on the head of the ecloding embryos, and according to Cobben these may be gener¬ ally classified as clypeal or frontal. Although the eggs of some Pentatomomorpha have a true operculum, eclosion in many members of the group takes place by an irregular rupture of the egg, rather than by lifting off a cap. The cephalic pole of the egg of most bugs bears from 62 TRUE BUGS OF THE WORLD (HEMIPTERA; HETEROPTERA) Fig. 10.13. Circulatory system (from Hinks, 1966). A. Dorsal vessel and alary muscles, Phonoctonus fasciatus (de Beauvois) (Reduviidae). B. Dorsal vessel and alary muscles, Aquarius lacustris (Linnaeus) (Gerridae). 1 or 2 to many micropyles (Fig. 10.14F, G), and in the case of most Cimicomorpha also bears what have been referred to as aeropyles, which apparently serve the func¬ tion of gaseous exchange. In the case of the Pentatomo- morpha, including the Aradoidea, the cephalic pole of the egg is also ornamented with micropylar processes (Fig. 10.14D, E), at which point the micropylar canals exit the egg shell. In all bugs except the Cimicoidea, sperm enter the egg during fertilization through the micropyles. Sperm and Fertilization Heteropteran sperm, like those of most other insects, are long and slender, with the nucleus present in the head (Phillips, 1970). On the basis of at least one representative of all infraorders except Enicocephalomorpha and Dipso- coromorpha, they appear to have three characteristics that do not occur in most other insects—additional evidence for the monophyly of the group. First, there are 2 or 3 crystalline bodies in the mitochondrial derivatives, rather than 1, as found in other Pterygota; second, 2 bridges join the mitochondrial derivatives to the axoneme at the level of doublets 1 and 5; and third, the axoneme lacks the longitudinal accessory bodies found in many closely related groups of insects (Dallai and Afzelius, 1980; Af- zelius etal., 1985; Jamieson, 1987). Some Pentatominae (Pentatomidae) produce sperm with abnormal chromosomal complements. These always come from one lobe of the testes, referred to as the har¬ lequin lobe, and they do not participate in reproduction. The functional significance of their existence has not been satisfactorily explained. Most male Heteroptera apparently deposit sperm in the female as a spermatophore (Ambrose and Vennison. 1990). Sperm are stored by the female in the spermatheca, or its functional equivalent in the Cimicomorpha. Fer¬ tilization and oviposition are ordinarily simultaneous in the Heteroptera. In most Cimicoidea, however, sperm are deposited in the abdominal cavity by injection through a specialized area of the body wall known as the ecto- spermalege (Carayon, 1966, 1977). From that point the sperm migrate to the base of the ovaries, and in many taxa embryogeny precedes oviposition. Chromosomes and Karyotypes Heteroptera, in common with other Hemiptera, have holocentric (holokinetic) chromosomes, in which the centromere is distributed along the entire length of the chromosome. Diploid chromosome complements range from 4 to 48 or 49, with some of the lowest and highest numbers occurring in the Nepomorpha. The total amount of genetic material is about the same in members of the same family with very different chromosome numbers, suggesting fragmentation rather than polyploidy as the ordinary method of increase in chromosome numbers. The Pentatomomorpha generally have diploid comple¬ ments ranging from 12 to 18, whereas all other major groups (except Enicocephalomorpha, in which they are unknown) have diploid complements of 20 or above, but with substantial variation. Sex chromosome systems may be either XY or XO The XO system is found in Dipsocoromorpha, Gerromo’ pha, a few Nepomorpha, a few Lygaeidae, and all Lepto- General Adult Morphology and Key to Infraorders 63 Fig. 10.14. Eggs and egg structures. A. Egg, lateral view Saldula fucicola (Sahiberg) (Saldidae). B. Egg, S. fucicola (A, B from Cobben, 1968a). C. Egg, Nabis limbatus Dahibom (Nabidae). D. Egg, showing micropylar processes at cephalic pole. Nezara viridula (Linnaeus) (Pentatomidae). E. Micropylar process, Oncopeltus fasciatus Dallas (Lygaeidae). F. Operculum and micropylar region, Rhinocoris sp. (Reduviidae). G. Base of micropylar region, Rhinocoris sp. (C-G from Southwood, 1956). podomorpha studied. Nearly all known Cimicomorpha and Pentatomomorpha have XY sex-determination sys¬ tems, suggesting that XY is the derived condition. Many groups have multiple X chromosomes, with a maximum of 5 in some Reduviidae, and indeed, the greatest varia¬ tion in chromosome complement is often the result of variation in the number of X chromosomes. One or more supernumerary “m” chromosomes occur in many members of the Nepomorpha and in lygaeoid, pyrrhocoroid, and coreoid Pentatomomorpha (Ueshima, 1979). Nymphs Heteropteran nymphs typically resemble adults and live in similar environments. There are ordinarily five instars, although some variation does occur—as for example the consistent occurrence of four instars in Mesovelia fur- cata Mulsant and Rey (Zimmermann, 1984)—but with¬ out any apparent systematic pattern (Stys and Davidova, 1989). Especially in the Pentatomomorpha, the first- instar nymphs may be red and black, whereas in most other groups later instars are usually colored in a manner similar to the adults. Aside from the lack of ocelli, wings, and genitalia, nymphs are distinguished primarily by the presence of one less tarsal segment than found in adults, and in the case of the Pentatomoidea the subdivision of the pedicel to form the 5-segmented antennal condition usually appears at the imaginal molt. Yonke (1991) gave an excellent general treatment of heteropteran nymphs, including keys to all families, diagnoses, representative illustrations, and citations for additional sources. Cobben (1978) provided the best survey of nymphal character¬ istics of phylogenetic value, with illustrations, descrip¬ tions, and tabular summaries. Keys to families occurring in North America were published by DeCoursey (1971) and Herring and Ashlock (1971). 64 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Key to Infraorders of Heteroptera Diagnosis. Mouthparts typically hemipteran with mandibular stylets concentric and surrounding maxillary stylets; labium inserted anteriorly on head, a distinct gular area always present, often closed behind to form a buc- cular bridge; scent-gland structures, often paired, located and exiting ventrally in metathorax of adults, channels moving fluid to variously elaborated metathoracic pleu- ron; nymphs with paired scent glands located at junction of one or more of the following abdominal terga: 3/4, 4/ 5,5/6, 6/7. 1. Compound eyes usually present, sometimes reduced, very rarely absent (a few Enicocephalomor- pha); wings occasionally reduced or absent; mostly free-living . 2 - Compound eyes always absent; apterous; obligate parasites of bats or termitophilous inquilines .... . 14 2. Head constricted transversely, divided into 2 distinct lobes, ocelli (when present) placed on posterior lobe posteriorto compound eyes (Figs. 12.1 A, 13.1 A); foretibiae flattened, with distinct distal spines (Fig. 13.1 A, C, D); foretarsus usually 1-segmented, sometimes 2-segmented. always opposable to apex of foretibia (Figs. 12. ID, 13.1C, D); forewings, when present, of uniform texture, not differen¬ tiated into distinct corium and membrane, lacking claval commissure (Fig. 13.1 A); compound eyes sometimes greatly reduced or absent . Enicocephalomorpha - Head generally not constricted and divided into lobes; if divided, prostemum with a distinct sulcus for reception of apex of labium (Fig. 10.7F) and foretibia and foretarsus never as above; fore¬ wings, if fully developed, of variable structure, divided into a distinct corium and membrane or not; compound eyes only very rarely reduced or absent (Figs. 42.1,54.1) . 3 3. Head with 3 or 4 pairs of conspicuous trichobothria placed near inner margin of compound eyes (Figs. 10.8A, 20.2C), inserted in distinct pits, bothrium obscured in pit. hair pile obviously devel¬ oped around pit margin; forewings, when present, lacking claval commissure and not divided into a well demarcated corium-clavus and membrane (Figs. 20.2A, B, 20.3B); part or most of body always covered with a distinct pile of microsetae and spicules . Gerromorpha - Head without trichobothria, or if with one or more pairs of trichobothria, these never placed in deep pits; forewings, when developed, with or without claval commissure; venter infrequently with pile of microsetae and spicules (Nepomorpha: Aphelocheiridae; some Naucoridae; some Leptopodomor- pha) . 4 4. Antennae shorter than head and folded beneath it, usually concealed (Fig. 29.1 A) in grooves (except Ochteridae lacking groove; Fig. 29. ID), at most apex of antenna slightly exposed beyond margin of large compound eyes (Fig. 29. ID); compound eyes present, usually large (Figs. 30.1, 33.1); prostemal sulcus never present; usually macropterous, rarely staphylinoid (some Naucoridae, Aphe¬ locheiridae; Fig. 37. IB), claval commissure always present in macropterous forms (Fig. 30.1) .... . Nepomorpha - Antennae usually longer than head and never concealed in grooves below compound eyes, even when shorter than length of head; if antennae shorter than head, prostemal sulcus present; forewings variously developed, claval commissure present or absent . 5 5. Abdominal sterna 3-7 usually with 2 or 3 (or rarely 1 or more than 3) trichobothria placed sub- laterally (Pentatomoidea) (Fig. 10.8C) or submedially on sterna 3 and 4 and sublaterally on sterna 5-7 (other groups; Fig. 10.8D), occasionally present sublaterally only on sterna 5-7, or very rarely completely absent (some Piesmatidae); elongate pulvillus always attached near base of each claw (Fig. 10.51) . Pentatomomorpha (less Aradoidea) - Abdomen at most with a single trichobothrium-like seta on either side of midline of one or more sterna and never arranged as described above, or with a single trichobothrium on ventral lateroter- gite 7 (some Nabidae; Fig. 10.8F); claws sometimes with pulvilli (most Aradoidea: Fig. 10.5G, H; some Miridae: Fig. 53.5E; and some Anthocoridae) . 6 6. Antennal segments 1 and 2 short, subequal in length, segments 3 and 4 very long and slender and clothed with erect setae of length much greater than segmental diameter (Fig. 17.1); length usually under 2.5 mm; hemelytra, when present, tegminal orcoleopteroid (Figs. 15.lA, 17.1, 18.lA) .... . Dipsocoromorpha (less Stemmocryptidae) - Antennal segment 2 usually longer than segment 1, if not (most Tingidae; Fig. 54.2B), then seg¬ ments 3 and 4 lacking erect setae longer than segmental diameter and pronotum and hemelytra General Adult Morphology and Key to Infraorders 65 covered with areoles; hemelytra, when developed, never tegminal and undifferentiated . . 7 7. Claws usually with distinct elongate pulvilli free from claw except at base (Fig. 10.5G); tarsi 2- segmented (Fig. 64.1 A); body conspicuously flattened, sometimes encrustate (Fig. 64. IB); wings, when present, covering only discal area of abdomen, connexivum exposed (Fig. 64.1 A); eyes small relative to size of head, pebblelike, but not with reduced numbers of ommatidia (Fig. 64.1 A, B) . Aradidae - Claws sometimes with pulvilli (some Miridae: Fig. 53.5E; some Anthocoridae), but if present pulvilli rarely free from claw over most of length (some phyline Miridae) and then tarsi always 3- segmented; tarsi usually 3-segmented, less frequently 2-segmented; wings, when present, usually covering connexivum (except some Reduviidae); eyes often comparatively large relative to size of head (Fig. 53.2B), rarely pebblelike or with reduced numbers of ommatidia . 8 8. Membrane of forewing present, in at least a reduced form, 3, 4. or 5 closed cells usually visible (Figs. 43.1 A, B, 45.1 A, B), never with veins emanating from posterior margin of cells . . Leptopodomorpha (part) - Forewing membrane usually present, often with closed cells, usually 2 (Fig. 47.1), if more than 2 then always with veins emanating from their posterior margins . 9 9. Forewings present in the form of hemelytra with a differentiated corium-clavus and membrane; compound eyes well developed . Cimicomorpha (part) - Forewings either tegminal and undifferentiated (Stemmocryptidae: Fig. 19.1 A, B), coleopteroid (Omaniidae; Fig. 44.1 A; Tingidae: Vianaidinae, Fig. 54.1), or greatly reduced (Aepophilidae; Fig. 42.1; some Cimicomorpha) . 10 10. Forewings coleopteroid . 11 - Forewings tegminal or greatly reduced . 12 11. Eyes very large, occupying nearly entire sides of head; body length approximately 1 mm (Omaniidae; Fig. 44.1) . Leptopodomorpha (part) - Eyes consisting of only a few ommatidia; body length greater than 2 mm (Vianaidinae; Fig. 54.1) . Tingidae (part) 12. Forewings developed, largely membranous with a few relatively strong veins (Stemmocryptidae; Fig. 19.1 A) . Dipsocoromorpha (part) - Forewings completely absent or in the form of small pads . 13 13. Most of body covered with a short hair pile; padlike forewings as in Fig. 42.1 (Aepophilidae) .... . Leptopodomorpha (part) - Body not covered with a hair pile; forewings greatly reduced or absent, but not exactly as in Fig. 42.1 . Cimicomorpha (part) 14. Body very strongly flattened, with marginal laminae (Fig. 65.1 A, B) (Termitaphididae) . . Pentatomomorpha (part) - Body not so flattened, and without marginal laminae (Fig. 62.1 A) (Polyctenidae) . .. Cimicomorpha (part) TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) HETEROPTERA 11 Enicocephalomorpha by Pavel Stys General. The unique-headed bugs are also called four- winged flies for their habit of forming mating swarms and their similarity in appearance to the Nematocera when flying. These delicate, elongate bugs range in length from 2 to 15 mm and, although seldom collected, are widely distributed, even in temperate areas. Diagnosis. Head elongate, porrect, subdivided into anterior and postocular lobes by a usually conspicuous postocular constriction (Fig. 13.1 A); ocelli, if present, on posterior lobe, well removed from eyes (Fig. 13.1 A); gula long; labium 4-segmented, short, straight to arcu¬ ate, never exceeding length of head; antennae flagelli- form to terete, moderately long (Fig. 13.1 A); forewings always completely membranous (tegminal) (Figs. 12.1 A, 13.1 A); ambient vein in remigium marginal or slightly submarginal, but if venation reduced, then vein repre¬ sented at least by a continuous row of macrotrichia; forewings sometimes reduced or absent, occasionally ca¬ ducous; medial fracture situated in front of R on fore¬ wing, in common only with some Dipsocoromorpha; base of forewing with bracelike cross vein connecting marginal veins with R and M-f-Cu; foreleg usually rap- Key to Families of Enicocephalomorpha torial, tibia distoventrally produced, usually dilated, with' 1 or 2 clusters of spiniform setae, opposed to ventral face of 1- or 2-segmented foretarsus, usually also with spines (Figs. 12.ID, 13.1C, D); male genitalia always symmetrical, with paired genital plates as in Auchen- orrhyncha, greatly reduced to racquet-shaped guide in Enicocephalidae (Fig. 13. IE); ovipositor present (Fig. 12. IF) to absent; subgenital plate formed by sternum 8 rather than 7 as in other Heteroptera; spermatheca present (Fig. 12. IG). Discussion. The Enicocephalomorpha, whose unique structure and phylogenetic position in the Heteroptera have been recognized by most modem authors (Stys and Kerzhner, 1975), is currently divided into two families— Aenictopecheidae and Enicocephalidae (Stys, 1989)— rather than the single family Enicocephalidae (e.g., Stys, 1970b). Enicocephalomorphans were at one time placed in the Reduvioidea e.g., Usinger, 1943), but they were combined with dipsocoromorphans by Miyamoto (1961 a; and Popov (1971). Grimaldi et al. (1993) illustrated and described aspects of morphology of a Lower Cretaceous amber fossil enico- cephalomorph from Lebanon, helping to establish the great antiquity of this group. They also described in two Recent genera species of Oligocene-Miocene amber fos¬ sil Enicocephalidae from the Dominican Republic, noting that these taxa are virtually identical to living forms. Major classical students of the family were Bergrotli, Eckerlein, Breddin, and Usinger. Jeannel (1941) mono¬ graphed the group, and Usinger (1945) provided a higher classification. Villiers (1958) monographed the fauna of Madagascar and later that of the Afrotropical and Mala¬ gasy regions (1969a) Usinger and Wygodzinsky (1960) revised the fauna of Micronesia. Stys (1969) revised the extinct Enicocephalomorpha and in many subsequent papers described new higher taxa and provided morpho¬ logical information, listed the world supraspecific taxa (Stys, 1978), and established the current higher classi¬ fication of the group. Wygodzinsky and Schmidt (1991) monographed the New World fauna (excluding Systello- deres Blanchard), adding much new structural data based on SEM observations. 1. Males with a distinct inflatable or noninflatable phallus protruding out of the pygophore (Fig. 12. IE); parameres distinct, movable, articulated with phallus; ovipositor usually present, rarely strongly re¬ duced; forewing in macropterous forms usually with a short costal fracture (Fig. 12.1 A) (except Murphyanellinae) . Aenictopecheidae - Males with noninflatable intromittent organ never resembling typical heteropteran phallus (Fig. 13. IE); parameres distinct, but always immobile (Fig. 13. IE), hooklike, tuberculiform, plate-shaped to reduced to paired lateral parameral sclerites associated with supradistal plate or its vestige; oviposi¬ tor vestigial or wanting; forewing in macropterous forms without costal fracture (Fig. 13.1 A) . .. Enicocephalidae 67 Enicocephalomorpha 12 Aenictopecheidae by Pavel Stys General. The aenictopecheids are inconspicuous, little known, and rarely encountered bugs. Some species have retained plesiomorphic characters not encountered in other enicocephalomorphans, or among Heteroptera in general. They range in length from approximately 3 to 10 mm. Diagnosis. Posterior lobe of pronotum often abbrevi¬ ated, poorly defined, not demarcated by sinuate lateral margin (Fig. 12.1 A); Rs in forewing branching in Maori- stolinae, costal fracture short (Figs. 12.1 A, B); wing coupling mechanism as in Fig. 12.1C; pygophore never subdivided into tergum, laterotergites, and sternum; phal¬ lus typically heteropteran (Fig. 12. IE), inflatable or not, with movable parameres; ovipositor usually fully devel¬ oped (Fig. 12. IF); nymphs with normally developed wing pads, not contiguous along midline. Classification. The present conception of this family is on the basis of plesiomorphic characters. Nonetheless, none of its subfamilies possesses any apomorphies in common with the Enicocephalidae. This group is clearly relict, and many taxa badly need more detailed morpho¬ logical study. Four subfamilies are currently recognized comprising 10 genera and 20 species. Key to Subfamilies of Aenictopecheidae 1. Eyes reduced to a single ommatidium; all legs fossorial; spiniform setae (markedly thicker than other macrotrichia) situated on ventral face of middle and hind femora, along the length of all tibiae, and on ventral apex of all tarsi, but not markedly clustered at the ventral apex of foretibia; micropterous. transverse scale-shaped forewings not extending onto abdomen: tarsal formula 1-1-1; posterior lobe of pronotum extremely short and not demarcated in lateral view; New Zealand and Tasmania . . Nymphocorinae - Eyes normally developed; forelegs raptorial, middle and hind legs cursorial; spiniform setae situated only on apices of tibiae in most taxa and ventrally on foretarsi (foretarsal spines absent in Mur- phyanellinae), those of ventral apex of foretibia concentrated into 1 or 2 clusters; macropterous, or if micropterous, then forewings not transverse, with remnants of venation, and extending onto abdomen . 2 2. Tarsal formula 1-2-2; pronotum seemingly of 2 lobes only, posterior lobe usually reduced in size and/ or not distinctly delimited in lateral view; collar rarely vaguely delimited and then pronotum seemingly not subdivided into lobes; macropterous forms with a short costal fracture, just interrupting forewing marg in; foretarsus with ventral spiniform setae . 3 - Tarsal formula 1-1-1; pronotum with 3 distinct lobes; macropterous forms (only macropterous males known) without costal fracture; foretarsi without ventral spiniform setae; minute, less than 2 mm long; Singapore . Murphyanellinae 3. Forewing with R branching into Ri and Rs; foretibial ventral apical armature formed by clustered simple spiniform setae situated on dilated, but not projecting, tibial apex; male with a pair of inflatable pygophoral “vesicles”; ovipositor almost completely absent, no external traces of first valvulae; New Zealand . Maoristohnae - Forewing with simple R; at least part of the tibial ventral apical armature formed by modified (broad, flattened, abbreviated, apically often rounded or bilobate) spiniform setae, situated on projecting tibial apex and subdivided into 2 clusters by a notch; no inflatable pygophoral “vesicles”; ovipositor retained, at least as internal remnants of valvulae . Aenictopecheinae AENiCTOPECHEiNAE (FiQ. 12.1A). Small to medium-sized, sometimes robust {Gamostolus Berg); macropterous to strongly brachypterous, or sometimes dimorphic {Bo- reostalus Wygodzinsky and Stys); posterior lobe of pro¬ notum reduced, usually not distinguishable in lateral out¬ line (Fig. 12. lA); apical armature of foretibia subdivided into 2 clusters of abbreviated, thickened, often rounded or bilobate, peglike spiniform setae (Fig. 12. ID); tar¬ sal formula 1-2-2; phallus noninflatable, permanently erected, and always strikingly protruding from lower part 68 TRUE BUGS OF THE WORLD (HEMIPTERA; HETEROPTERA) Fig. 12.1. Aenictopecheidae. A. Tomocrust/s pena; Wygodzinsky and Schmidt (from Wygodzinsky and Schmidt, 1991), B. Forewing, Gamostolus subantarcticus (Berg) (from Wygodzinsky and Schmidt, 1991). C. Wing-to-wing coupiing mechanism, G. subantarcticus (from Wygodzinsky and Schmidt, 1991). D. Foreieg, distai portion, generalized Aenictopecheidae (A-D from Wygodzinsky and Schmidt, 1991). E. Maie genitalia, Monteithostolus genitalis Stys (from Stys, 1981 a). F. Female genitalia, ventral view apex abdomen, Australostolus monteithi. G. Spermatheca, A. monteithi (F, G from Stys, 1980). Abbreviation: cf, costal fracture. of posterior foramen of pygophore (Fig. 12. IE); oviposi¬ tor either well developed with first valvifer fused with broad sternum 8 forming triangular submedial processes, or strongly reduced. Six genera are grouped in two tribes. The Aenic- topecheini includes Aenictopechys necopinatus Breddin from Indonesia and Lomagostus jeanneli Villiers from Madagascar. These are small species with complex pat¬ terns of vein fusion in the cubitoclaval area. Members of the Gamostolini are usually larger and with more typical venation (Fig. 12. IB). Gamostolus Berg includes a single species, G. subantarcticus (Berg), from southern Argentina and Chile, but unidentified nymphs of the genus are known (Wygodzinsky and Schmidt, 1991) from the Venezuelan and Colombian cordilleras. Tornocrusus Kritsky includes several species from Central and South America. Boreostolus Wygodzin¬ sky and Stys (1970) includes two boreal species. Austral¬ ostolus monteithi Stys (1980) is Australian. MAORiSTOUNAE. Slender, medium-sized, macropterous to moderately brachypterous; foretibia with simple spini- form apical armature; tarsal formula 1-2-2; posterior lobe of pronotum strongly reduced, pronotum apparently formed of 2 lobes; forewings with short costal fracture Aenictopecheidae 69 and branching Rs; phallus inflatable; pygophore with a pair of large eversible vesicle-like structures; ovipositor strongly reduced. This group comprises a single genus, Maoristolus Woodward (1956), with two species from New Zealand. MURPHYANELLINAE. Pronotum formed of 3 distinct lobes; only macropterous males known; forewings lack¬ ing costal fracture; phallus distinct, inflatable in Mur- phyanella aliquantula Wygodzinsky and Stys, beaklike in Timahocoris paululus Wygodzinsky and Stys; para- meres uniquely associated with base of phallus by pair of Y-shaped connectives resembling those in Auchen- orrhyncha, tubercle-shaped and probably immobile in Timahocoris Wygodzinsky and Stys; Murphyanella Wy¬ godzinsky and Stys with spiracle of abdominal segment 3 enlarged to about 11 times diameter of other abdominal spiracles. This enigmatic group includes two tiny species de¬ scribed by Wygodzinsky and Stys (1982) from Singapore. NYMPHOCORINAE. Small, yellowish, micropterous; eyes with a single ommatidium; feme; .: and tibiae with many spiniform setae; foretibia with apical cluster of spiniform setae spreading along tibia; forelegs appearing fossorial; middle and hind legs cursorial; tarsal formula 1 -1-1; phal¬ lus complex, inflatable, with lateral appendages; proc- tigeral region complicated; ovipositor complete. This subfamily includes only Nymphocoris Woodward, with two species, N. maoricus Woodward (1956) from New Zealand and N. hilli Stys (1988) from Tasmania. Specialized morphology. The Aenictopecheidae are most obvious for their lacking characters distinctive to the Enicocephalidae as well as having genitalia that are typi¬ cally heteropteran in form (Fig. 12. IE, F) as opposed to the greatly reduced and modified structures found in most Enicocephalidae. Natural history. Few details are known of the life habits of the Aenictopecheidae. Gamostolus has been found swarming, under stones, and in forest litter. Aus- tralostolus Stys is periodically attracted to lights in semi- desert areas. Boreostolus spp. live under big stones on gravel and sand substrates along mountain streams in habitats characteristic of Cryptostemma spp. (Dipsocori- dae). Maoristolines are found in litter, under bark, and in mosses. The Nymphocorinae are known to live in soil, litter, among tussocks of grass, and are found through the use of Berlese funnels and pitfall traps. Distribution and faunistics. Boreostolus has a typical boreal amphipacific distribution; B. americanus Wygod¬ zinsky and Stys in North America ranging from Wash¬ ington and Oregon to Colorado, and B. sikhotalinensis Wygodzinsky and Stys occurring in eastern Russia from Vladivostok to Magadan, the latter locality being the northernmost record for any enicocephalomorphan. The New World representatives are included in Wygod¬ zinsky and Schmidt (1991). Old World taxa are treated in the papers referred to above. 13 Enicocephalicdae by Pavel Stys General. The Enicocephalidae include about 95% of the species and 99.5% of known specimens of Enico- cephalomorpha. The general facies of most species re¬ sembles small to medium-sized reduviids. As in the Aenictopecheidae, most species are dull-colored, usually uniformly yellow, brown, or blackish, less frequently with contrasting color patterns, sometimes of bright red. Length ranges from about 2 to 15 mm. Diagnosis. Pronotum usually subdivided into 3 dis¬ tinct lobes (except Megenicocephalus and Alienates Bar¬ ber); wing polymorphism common, males sometimes macropterous and females brachypterous to apterous; costal fracture absent (weakly indicated in Megenico- cephalinae); nymphal wing pads of later instars large, contiguous, and sometimes slightly overlapping along midline; foretibia and tarsus as in Fig. 13.1C. D; phallus in primitive taxa unlike that of other Heteroptera, formed from structures including paired elements homologous to the genital plates of Auchenorrhyncha, often transformed from condition found in Monteithostolus genitalis Stys to arcuate or racquet-shaped, distally perforated “guide” associated with the ventral margin of posterior foramen of pygophore (Fig. 13. IE); parameres always immobile, fused at bases or reduced to flat sclerites; external female genitalia absent or retained as remnants in Systelloderini and Megenicocephalinae; female genital opening covered by extensive subgenital plate formed by sternum 8. Classification. Five subfamilies and several additional tribes are recognized (Stys, 1970c, 1989; Wygodzinsky and Schmidt, 1991). At present 33 genera and approxi¬ mately 400 species are known. Because of the difficulty in diagnosing some taxa, distribution and wing development are emphasized in the key. 70 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Key to Subfamilies of Enicocephalidae 1. Pronotum either with collar and middle lobe, or middle lobe and posterior lobe, if of 3 lobes then middle and posterior lobes distinguished by sinuation of lateral margin: males always with extremely simplified genitalia formed by “guide” and parameral sclerotizations only; forelegs variable; tarsal formula 1-1-1 or 1-2-2 . 2 - Pronotum of all wing morphs usually subdivided in 3 well-delimited lobes (Fig. 13.1 A); if lobes not distinct, then apterous/micropterous taxa from SW Pacific (including New Zealand) and subantcU'ctic islands; male genitalia complex to simplified; forelegs always raptorial, with dilated ventral apicotibial projection and its clustered spiniform armature; tarsal formula 1-2-2 . 3 2. Large, length 9-17 mm, robust; coloration reddish; pronotum formed of 2 lobes, collar undiffer¬ entiated; eyes with many ommatidia; forelegs incrassate (particularly femora), cursorial; apicotibial projection and its armature undifferentiated; tarsal formula 1-2-2; female forelegs with spinous pro¬ cess on trochanter, and spiniform tubercles on tibia and sometimes femur; forewing with complete venation and indication of costal fracture; antennae flagelliform; female with a minute knoblike struc¬ ture with paired clawlike appendages in ventral abdominal intersegmental membrane 8-9; Oriental . Megenicocephalinae - Minute, length under 2 mm, slender; coloration dull; pronotum in macropterous males with collar well delimited, middle and hind lobes differentiated by sinuation of lateral margin only; pronotum in apterous females formed of collar and a single lobe (Fig. 13. IB); eyes of females with 0-5 omma¬ tidia (Fig. 13. IB); forelegs raptorial, with apicotibial projection and its armature differentiated (Fig. 13.ID); tarsal formula 1-1-1; male foretarsus without spines; forewing venation of males reduced to 2-3 longitudinal veins and 0-1 cross veins; antennae incrassate (Fig. 13. IB); female abdomen strongly sclerotized with sharply delimited sets of dorsal external and inner laterotergites (Fig. 13.1B); Neotropical . Alienatinae 3. Upper and lower surfaces of fore and hind wings with numerous macrotrichia on veins and mem¬ branous areas (only macropterous males known); male genitalia formed by large bulbous closed projecting “phallandrium” with long tapering or subtruncate apex; parameres large, partly projecting out of the pygophore; Oriental . Phallopiratinae - Most macrotrichia on upper surface of forewings occurring on veins and wing margin; lower surface of forewings and both surfaces of hind wings without macrotrichia; macropterous to apterous; male genitalia complex to simple, but not as above . 4 4. Micropterous to apterous, no sexual pterygodimoiphism; middle lobe of pronotum without paired, in¬ versely Y-shaped impressions; male genitalia complex, never represented by a simple racquet-shaped guide, parameral sclerotizations, and a medial sclerite only; subantarctic islands (Crozet; Auckland), New Zealand, New Caledonia, New Guinea . Phthirocorinae - Macropterous to apterous, sexual pterygodimoiphism common; if micropterous to apterous, then either not occurring in SW Pacific area and on subantarctic islands, or with paired, inversely Y- shaped impressions or vestiges on middle lobe of pronotum; male genitalia reduced, usually formed by a usually racquet-shaped guide (Fig. 13. IE), paired parameral sclerotizations (rarely somewhat protruding), and a simple, plate-shaped supradistal lobe, the last sometimes prominent, arcuate or spiniform; worldwide . Enicocephalinae ALIENATINAE (FIG. 13.1 B). Males with posterior pronotal lobe virtually absent; forewing venation strongly reduced; females apterous (Fig. 13.IB), largely desclerotized; middle and hind tarsi l-segmented. This small subfamily comprises only the speciose genus Alienates Barber, discovered originally in the Bahama Islands, but widely distributed in the Caribbean, Meso- america, and northern South America. These tiny bugs superficially resemble small midges. All species appear to live in soil. ENICOCEPHALINAE (FIG. 13.1A). Male genitalia greatly simplified (Fig. 13. IE); females sometimes micropterous to apterous; micropterous and apterous Old World enico- cephalines restricted to Oncylocotis Stal; some taxa with caducous wings, for example, Nesenicocephalus Usinger. This is the most speciose group in the family, with two tribes recognized. At present 26 genera and about 250 species have been described. Most species occur in the tropics and subtropics, but the range of the subfamily ex¬ tends south to southern Chile and Tasmania and north to Enicocephalidae 71 Fig. 13.1. Enicocephalidae (from Wygodzinsky and Schmidt, 1991). A. Hymenocoris brunneocephalis Wygodzinsky and Schmidt. B. Alienates elongatus Wygodzinsky and Schmidt. C. Foreleg, distal portion, generalized Enicocephalinae. D. Foreleg, generalized Alienatinae. E. Male genitalia, Urnacephala californica Wygodzinsky and Schmidt. the northern United States and the southern fringes of the Palearctic, Japan being the northernmost part of the range in the Old World. The most speciose genera within the Enicocepha- lini are the New World Neoncylocotis Wygodzinsky and Schmidt and Enicocephalus Westwood and the Old World Henschiella Horvath, Hoplitocoris Jeannel, Steno- pirates Walker, and particularly Oncylocotis Stal. The last genus includes some micropterous to apterous species or morphs, some of them truly neotenous; an undescribed species from New Caledonia has minute club-shaped forewings similar to dipteran halteres. The genus Codes Bergroth (Madagascar; Seychelles Islands) is holoptic, the eyes occupying most of the head and obscuring the usual distinction between anterior and posterior cephalic lobes. 72 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) The Systelloderini comprise only the cosmopolitan genus Systelloderes Blanchard, but the group is well char¬ acterized only for the New World taxa, for which many of species await description. Some of its species retain plesiomorphic genitalia rather resembling those of the Phthirocorini. It is possible that some of the Old World genera of Enicocephalini actually belong in the Systello¬ derini. MEGENicocEPHALiNAE. Large, broad bodied, mostly reddish; costal margin of forewing with a small break re¬ sembling an incipient costal fracture; pronotum formed of 2 lobes, resembling superficially the condition in Aenic- topecheinae, but collar undifferentiated, and middle and hind lobes normally developed; forelegs strongly devel¬ oped, essentially cursorial, lacking usual apicotibial ar¬ mature, but often with variously tuberculate to spinose tibia, femur, and trochanter (sexually dimorphic). This monotypic subfamily contains only the Oriental genus Megenicocephalus (Malay Peninsula; Indonesia). PHALLOPIRATINAE. Forcwings and hindwings generally covered by macrotrichia on both the upper and lower surfaces; foretibia in males compressed in two different planes; male genitalia represented by a bulbous “phallan- drium” (§tys, 1985b) produced into a spine, resembling a scorpion telson and including paired elements homolo¬ gous to genital plates of other Heteroptera. Represented only by Phallopirates §tys with four spe¬ cies, this Oriental group ranges from Malaya and Borneo to the Philippine Islands. Males have been collected at light, but females are unknown and probably nonflying. §tys (1985b) found no gonopore in the males and sug¬ gested that “androtraumatic” insemination takes place, the male having to break off the apex of the phallandrium in order to pass the spermatophore to the female. PHTHiROCORiNAE. Apterous or micropterous; male genitalia structurally diverse, sometimes greatly reduced into an enicocephaline-like condition. Two tribes are recognized from the southwest Pacific and subantarctic islands. The two genera and three species of Monteithostolini, all from New Caledonia, are apterous. Monteithostolus genitalis Stys has the pygophore subdivided into a distinct tergum, laterotergites, and sternum (Stys, 1982b). The Phthirocorini comprises two genera and four spe¬ cies from New Guinea, New Zealand. Auckland Islands, and Crozet Island. Phthirocoris subantarcticus Enderlein occurs on Crozet Island in penguin and albatross nests. Specialized morphology. The division of the pro¬ notum into 3 apparent lobes, absence of the costal frac¬ ture, and the greatly modified and often reduced male and female genitalia are distinctive for the Enicocephalidae. Natural history. Most species of Enicocephalidae are found in leaf litter or loose soil, in moss, rotting wood, under bark, and in similar microhabitats, usually in the humid tropics or subtropics; species from arid zones and temperate regions probably live mostly in soil crevices. All enicocephalids are undoubtedly generalized preda¬ tors, but few observations are available. The most thor¬ ough review of the biology of the group is that of Wy- godzinsky and Schmidt (1991). Hickman and Hickman (1981) studied the life history of Oncylocotis tasmanicus (Westwood) and described the nymphal stages. Adults can be collected by use of Berlese funnels, pit- fall traps, or lights. All taxa in which both sexes are capable of flight form nuptial swarms, which can be diurnal or crepuscular. These swarms are mixed or uni¬ sexual (Stys, 1981b), but the details of mating habits are unknown. The method of copulation has not been described for any species. The swarms may be formed by an extremely high number of individuals and with a strongly skewed sexual ratio {Systelloderes, Oncylocotis, Stenopirates). Distribution and faunistics. New World taxa, with the exception of Systelloderes spp., can be identified using Wygodzinsky and Schmidt (1991). The Palearctic was treated by Stys (1970b). Villiers (1958, 1969a) mono¬ graphed the Malagasy and African and Malagasy faunas, respectively. For other areas in the Old World the works of Stys cited above are essential. Enicocephalidae 73 EUHETEROPTERA 14 Dipsocoromorpha by Pavel Stys General. The Dipsocoromorpha, which have no com¬ mon name, include the smallest tme bugs. They range in length from about 0.5 to 4.0 mm and range in appear¬ ance from tiny beetles to small, dull-colored, free-living members of the Cimicoidea. Diagnosis. Minute, 4 mm or less; head often strongly declivous (Fig. 18.IB); antennae flagelliform, with very short segments 1 and 2, segment 2 at most twice as long as 1, segments 3 and 4 usually very long, much thinner than segments 1 and 2, and with many long, thin, semierect to erect setae (except Stemmocryptidae; Fig. 19.1 A); pronotum variable, with 1 or 2 lobes; pro- epistemum often inflated and produced below ventral margin of eyes, sometimes reduced; forewing usually tegminal (Figs. 15.lA, 17.1, 18.lA, 19. IB); hind wing usually with several deeply incised lobes; pretarsus with 2 equally developed claws, pair of setiform parempodia, and usually on at least one pair of legs with a dorsal and ventral arolium, the arolia sometimes very large; number of abdominal spiracles often reduced to 3-6; male subgenital plate, when present, represented by ster¬ num 7; male genitalia, including pregenitalic abdominal segments, usually (except some Ceratocombidae) asym¬ metrical and extremely complex (Figs. 15.ID, 18.ID), some laterotergites appendage-like (Fig. 18. IE); bases of parameres directly associated with articulatory apparatus Key to Families of Dipsocoromorpha of phallus; nymphs with dorsal abdominal scent-gland openings on as many as 4 segments. Classification. The most recent classification (Stys, 1970a, 1983a) includes five families: Ceratocombidae, Dipsocoridae, Hypsipterygidae, Schizopteridae. and Stemmocryptidae. Hypsipterygids and stemmocryptids were discovered only relatively recently (Drake, 1961; Stys, 1983a; respectively). Formerly the names Dipso¬ coridae or Cryptostemmatidae were used either for all the dipsocoromorphans or for Dipsocoridae and Ceratocom¬ bidae (e.g., China and Miller, 1959) or for Dipsocoridae, Ceratocombidae, and Hypsipterygidae (Emsley, 1969). Dipsocoromorphans have been regarded as a group of uncertain affinity or considered jointly with enico- cephalomorphans in a group called either Dipsocoromor¬ pha (Miyamoto, 1961a) or Enicocephalomorpha (Popov, 1971). Nonetheless, early on, Reuter (1910, 1912a) and later Stys (1970a) used the series name Trichotelocera for the group, setting these unusual bugs aside from other terrestrial Heteroptera. The usage of Dipsocoromorpha in the present sense was stabilized by Stys and Kerzhner (1975). The authors who have contributed most to knowledge of Dipsocoromorpha are Reuter, McAtee and Malloch, Wygodzinsky, Emsley, Stys, Linnavuori, Kerzhner, Josi- fov, Miyamoto, Hill, and Ren. Reuter (1891) provided the first monograph of the group. Available taxonomic information was summa¬ rized, with particular reference to the American fauna, by McAtee and Malloch (1925), who also provided a key to the described genera. In a series of papers on the South American and African faunas Wygodzinsky set a high standard for describing and illustrating structures in the group, particularly the genitalia, and his papers on the fauna of Angola are still essential references (Wy¬ godzinsky, 1950, 1953). Emsley (1969) monographed the schizopterid fauna of Trinidad and provided at the same time an excellent morphological overview of the Schizop¬ teridae and, to a lesser extent, of the whole infraorder. Stys (1970a), independent of Emsley, established a higher classification and surveyed the morphology of the infra¬ order, in particular the complex morphology of the male abdomen and terminalia. Later, in association with his description of the family Stemmocryptidae, Stys (1983a) provided an account of the comparative morphology of the group. ’ 1. Antennae not conspicuously flagelliform, segment 1 rather short, segments 2-4 moderately long, subequal in length, 3 and 4 only slightly thinner than 2. without strikingly long erect setae (Fig. 19.1 A); ocelli situated behind eyes; macropterous; forewing tegminal, distal part of claval suture dis- 74 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) tant from wing margin, costal fracture long (Fig. 19. IB); male genitalia asymmetrical (Fig. 19. ID); New Guinea . Stemmocryptidae - Antennae flagelliform, segments 1 and 2 short and thick, segment 2 at most twice as long as seg¬ ment 1, segments 3 and 4 strikingly longer and much thinner than 1 and 2, provided with many long, thin, erect to semierect setae; ocelli, if present, interocular; forewing tegminal. hemelytraceous. ely- traceous, or reduced; distal part of claval suture usually meeting or closely approaching wing margin, costal fracture usually short or absent (Fig. 15.1 A, B) . 2 2. Pronotum with 3 longitudinal carinae meeting basally in a well-defined marginal ambient ridge; general facies tingidlike (Fig. 17.1); forewing without costal fracture, with remigial veins forming a system of about 10 large rectangular to polygonal cells (Figs. 16. IE, 17.1); labium straight, thin, segments 1. 2, and 4 minute, 3 strikingly long; head porrect, with distinctly delimited cephalic re¬ gions, including frons; macropterous to submacropterous; male genitalia asymmetrical; African and Oriental . Hypsipterygidae - Pronotum without longitudinal ridges and without distinct marginal ridge, general facies seldom tingidlike (Figs. 15.1 A, 18.1 A); forewing not as above; labium not as above, number of segments sometimes reduced; head declivous to porrect, frons never delimited; male genitalia symmetrical or asymmetrical . 3 3. Proepisternal lobe narrow in lateral view, not inflated and not extending cephalad; articulation of forecoxa laterally exposed, supracoxal cleft extremely short to hardly indicated, proepimeron large; costal fracture present, except in extreme brachypterous forms . 4 - Proepisternal lobe broad in lateral view, mostly inflated and extending below eye; articulation of forecoxa and basal part of latter covered, supracoxal cleft long . 5 4. Costal fracture short, just interrupting forewing margin; metapleuron without evaporator!um; male genitalia and abdomen symmetrical or asymmetrical; forewing tegminal to hemelytraceous; macrop¬ terous to brachypterous, or rarely coleopteriform . Ceratocombidae - Costal fracture reaching to about middle of width of forewing (Fig. 16.ID); metapleuron with evaporatorium; male genitalia and abdomen asymmetrical; forewing tegminal; macropterous to brachypterous . Dipsocoridae 5. Coleopteriform, forewings elytraceous; proepisternal lobe large, but not inflated; head porrect; male genitalia symmetrical (Fig. 15.ID); foretarsus with ventral spines (male Feshina §tys). or head decli¬ vous and pronotum strikingly transverse (female Kvamula Stys); hind coxae without adhesive pads . Ceratocombidae (part) - Macropterous to brachypterous, rarely micropterous, often coleopteriform, forewing tegminal to mostly hemelytraceous, often elytraceous, rarely micropterous; proepisternal lobe usually strongly inflated, and extending below eyes, and head strongly declivous; male genitalia, often including ab¬ domen, always asymmetrical (Fig, 18.1C-E); hind coxae with adhesive pads mesoventrally; costal fracture usually absent, or interrupting costal margin only, long and transversing the remigium in Guapinannus Wygodzinsky . Schizopteridae 15 Ceratocombidae by Pavel Stys General. These little-known bugs, which range in length from 1.5 to 3.0 mm, are mostly dull-colored, ranging from yellowish to dark brown. Ceratocombini and Trichotonanninae often resemble small long-legged drymine lygaeids with more or less tegminal forewings and flagelliform antennae; Issidomimini are more similar in appearance to small anthocorids. Diagnosis. Antennae flagelliform, second segment 1.5-2.5 times as long as segment 1; labium ranging from thin and elongate to short and thick in Feshina Stys; pro¬ notum ecarinate; propleuron with reduced proepistemum and exposed procoxal articulation except in rare coleop- teroid morphs; forewing in macropterous forms always with distinct, but very short, costal fracture just inter¬ rupting marginal vein (Fig. 15.IB), and 2-4 large cells distaliy; tarsal segmentation often sexually dimorphic, with nearly all possible combinations of 2 or 3 segments Ceratocombidae 75 A B C Fig. 15.1. Ceratocombidae. A. Ceratocombus mareki Stys. B. Forewing, C. mareki (A, B from Stys, 1977). C. Male abdomen, Trichotonannus dundo Wygodzinsky. D. Male genitalia, Muatianvuaia barrosmachadoi Wygodzinsky. E. Aedeagus, M. barrosmachadoi. F. Male genitalia, T. dundo (C-F from Wygodzinsky, 1953). for all pairs of legs; spiracles present on abdominal seg¬ ments 2-8 or 3—8, usually dorsal; male genitalia some¬ times symmetrical (Fig. 15.ID), but laterotergites 9 sec¬ ondarily associated with tergum 8 and appendage-like, unlike all other Heteroptera; ovipositor well developed; spermatheca always present (Fig. 16. IB, C). Classification. §tys (1970a) first recognized the u: ueness of the Ceratocombidae, raising the group to family level and establishing a suprageneric classification (modified by Stys [1982a, 1983a]). Stys (1983a) pro¬ vided updated keys to genera and higher taxa. Previously, members of the group were usually regarded as belong¬ ing to the Dipsocoridae (= Cryptostemmatidae) or Dip- socorinae (= Cryptostemmatinae) in a broadly defined Dipsocoridae. Two subfamilies are currently recognized, containing approximately 8 genera and 50 species. 76 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Key to Subfamilies of Ceratocombidae, Based on Males 1. Abdominal segments 7-9 and parameres asymmetrical; laterotergites 7-9 asymmetrical, 7 and 8 appendage-like, 9 transformed into processes of pygophore; head, pronotum, and legs with long, erect, black bristles; eyes with a central bristle; distal part of forewing in macropterous forms with 4 large cells; macropterous to brachypterous; Paleotropical . Trichotonanninae - Abdominal segments 7-9 and their laterotergites symmetrical; parameres symmetrical or asymmetri¬ cal; laterotergite 7 not appendage-like, 8 variably developed (from appendage-like to tubercle-shaped to undifferentiated), 9 appendage-like and secondarily articulating with tergum 8; if bristles different from other setae, then concolorous and not strikingly strong and long; eyes without a strong central bristle; distal part of macropterous forewing with 2-3 large cells (rarely with additional small ones); macropterous. brachypterous, or coleopteriform . . . Ceratocombinae CERATOCOMBINAE, Macroptcrous, forewing with only 2—3 large cells distally, in contrast to Trichotonanninae; body devoid of strong bristles; no central eye bristle; male abdomen and laterotergites 7-9 always symmetri¬ cal; laterotergite 7 not appendage-like, 8 appendage-like or not, 9 always appendage-like; parameres symmetrical or asymmetrical. Two tribes are recognized. The Ceratocombini includes the cosmopolitan Ceratocombus Signoret, with 3 sub¬ genera and 25 valid species and literally hundreds of undescribed species, particularly from the Indo-Pacific. Many species are pterygopolymorphic (macropterous to strongly brachypterous) and are usually dull-colored (yel¬ lowish to dark brown), though some Madagascan, Ori¬ ental, and Australian species have distinct color patterns. The genus Leptomnnus Reuter is known from three macropterous Nearctic, Neotropical, and Afrotropical species, Feshina schmitzi §tys from Zaire is coleopteroid, the male resembling a hydraenid beetle; its foretarsus bears an armature of spines similar to that found in females of Trichotonannus Reuter, The Issidomimini includes three genera, all from the Eastern Hemisphere: Issidomimus Poppius, one Oriental and one Papuan species; Kvamula Stys, four species from Vietnam; and Muatianvmia Wygodzinsky, five Afro- tropical species. Numerous Indo-Pacific species remain undescribed. Most Issidomimini are macropterous, but the female of Kvamula coccinelloides Stys is coleopteroid and resembles a scymnine coccinellid beetle, TRICHOTONANNINAE. Superficially similar to robust and strongly setose Ceratocombus; conspicuous central eye bristle; macropterous to moderately brachypterous (ptery¬ gopolymorphic); forewing with 4 large cells distally; setae on legs; foretarsi sexually dimorphic, 3-segmented in male, 2-segmented in female, with armature of spines on lower surface; metathoracic scent glands absent; male abdomen asymmetrical, laterotergites 7 and 8 appendage¬ like; pygophore, proctiger, and parameres strongly asym¬ metrical and remnants of laterotergites 9 developed as pygophoral processes, but never appendage-like. This subfamily comprises only the genus Trichotonan¬ nus Reuter, with six species from the Afrotropical, Mada¬ gascan, and insular Oriental regions (Philippines, Nico¬ bar Island). Although the general facies resembles that of Ceratocombini, the structure of the male abdomen is different. Trichotonannus oidipos §tys from Madagascar has the eyes composed of only a few isolated ommatidia. Specialized morphology. Ceratocombids are unique among the Dipsocoromorpha in that all Ceratocom¬ bini and some Issidomimini retain a symmetrical male abdomen and genitalia. Although some ceratocombids are macropterous, most species are pterygopolymorphic, some possibly always brachypterous. Males and females of Issidomimini are usually macropterous. Natural history. Most species live in moderately hu¬ mid leaf litter, decaying wood, mosses, sphagnum, and similar habitats with interstitial spaces. Ceratocombus species in Europe are typically found in bracken leaf litter mixed with needles in coniferous woods, drying sphag¬ num mixed with leaf litter and needles in wet woods, and reed litter. Ceratocombids can best be collected in pitfall traps, at light, and by berlesating litter. All ceratocombids are probably generalized predators on small arthropods. The labium in Ceratocombus sug¬ gests a searching predator, while that of some other genera suggest active prey hunting or totally different habits, such as feeding on molds. Ceratocombus, Lepto- nannus, and Trichotonannus are fast runners but do not jump. Distribution and faunistics. The family is cosmo¬ politan, but its diversity is poor in cold and temperate zones, though some species (e.g., Ceratocombus cortica- lis Reuter in the Palearctic) are restricted in cold regions. Maximum diversity is reached in tropical areas, and the number of undescribed species is enormous. Palearctic Ceratocombus species were revised by Ceratocombidae 77 Kerzhner (1974). Useful information about the Afrotropi- cal and Madagascan faunas was provided by Wygodzin- sky (1953), Linnavuori (1974), and Stys (1977, 1983b). American species were treated and keyed by McAtee and Malloch (1925). Outdated global treatments of species are available by Reuter (1891) and McAtee and Malloch (1925). Keys to world genera were provided by Stys (1982a, 1983a). Data on morphology and biology are contained in papers by Emsley (1969), Hill (1980), Stys (1958, 1959, 1970a, 1977, 1982a, 1983b), and Wygodzinsky (1953). 16 Dipsocoriidae by Pavel Stys General. These are usually somewhat flattened, elon¬ gate bugs that never exceed 3 mm in length. They have the general habitus of many free-living Cimicoidea. Diagnosis. Head porrect; labium thick, not exceeding procoxae, 4-segmented, first segment well-developed, other segments subequal in length; pronotum without well defined collar; proepistemum reduced, procoxal articulation expo,sed; scutellum large; macropterous to moderately brachypterous; forewings tegminal, mem¬ brane defined only by absence of veins, claval angle and claval commissure not formed; deep costal fracture almost reaching M (Fig. 16. ID); legs rather thick, tar¬ sal formula usually 3-3-3 or in females of some species 2-2-3, possibly also 2-2-2 (Emsley, 1969); metapleu- ral scent-gland evaporatorium present, in common with Stemmocryptidae; abdominal venter with dense pilosity; mediotergites largely desclerotized, a single (or double) set of laterotergites bearing spiracles ventrally, spiracles developed on segments 3-8, 4—8, or 4-7, the anterior ones probably absent. Nymphs with scent-gland orifices between terga 3/4, 4/5, 5/6, and 6/7; male abdomen always strongly asymmetrical, sinistral in contrast to other Dipsocoromorpha, asymmetry affecting in extreme cases segments 2-9, sometimes some of the pregeni¬ tal segments symmetrical; one or more of laterotergites 3 left, 6 left, 7 right, and 8 right sometimes appendage¬ like; parameres asymmetrical; phallus long, sometimes with coiled processus gonopori; female subgenital plate 7 terminating in medial complex laminate structure; ovi¬ positor developed, valvulae with microtrichial armature resembling filtering device. Classification. The Dipsocoridae as here restricted comprise the genera Cryptostemma Herrich-Schaeffer and Pachycoleus Fieber (Stys, 1970a). The family name Dip¬ socoridae Dohrn, 1859, is based on a junior synonym, Dipsocoris Haliday, and was later replaced by Crypto- stemmatidae McAtee and Malloch, 1925. Nonetheless, the former name has won wide acceptance and is now in general use (Stys, 1970a). The family was covered in now outdated monographs by Reuter (1891) and McAtee and Malloch (1925). More recent summaries are those of Emsley (1969) and Stys (1970a). Two genera are recognized. Cryptostemma Herrich- Schaeffer, with 3 subgenera and 26 described and many undescribed species, is known from all zoogeographical regions except New Zealand. Pachycoleus Fieber (often regarded as a subgenus of Cryptostemma) comprises four Palearctic and many undescribed Neotropical species. Specialized morphology. The presence of a meta- thoracic scent-gland evaporatory area and the sinistral asymmetry of the male genitalia are both distinctive for the family, although the former also occurs in the Stemmocryptidae. Natural history. Cryptostemma species inhabit a moist zone within gravel-sand banks of clear streams and rarely lakes, and may be found under stones on such a sub¬ strate. When disturbed, the bugs immediately exhibit an escape reaction, either creeping into the ground or simul¬ taneously jumping and flying. Habitats of Cryptostemma are subject to periodic flooding. The bugs survive either in trapped air bubbles or by using plastron respiration. Dipsocorids are undoubtedly generalized predators of small arthropods and have been observed to feed on dead mayflies. Pachycoleus spp. in the Palearctic inhabit wet moss and Sphagnum on the banks of ponds and lakes and in fens and bogs. The typical microhabitat is a small mound of Sphagnum and other mosses formed around tussocks of hygrophilous grasses, sedges, and other plants. All macropterous dipsocorids may be attracted to lights. Distribution and faunistics. Regional revisions have been provided by Josifov (1967) for Palearctic and Hill (1987) for Australia, but particularly the latter is not ex¬ haustive since numerous species remain to be described. Systematic collecting in suitable habitats will increase the number of known species many times. 78 TRUE BUGS OF THE WORLD (HEMIPTERA; HETEROPTERA) Fig. 16.1. Ceratocombidae. A. Female abdomen, Trichotonannus dundo. B. Spermatheca, T. dundo. C. Spermatheca, T. oidipos (A-C from Wygodzinsky, 1953). Dipsocoridae. D. Forewing, Cryptostemma incurvatum Stys (from Stys, 1977). Hypsipterygidae. E. Forewing, Hypsipteryx machadoi Drake. F. Maie genitalia, H. machadoi. G. Female genitalia, H. machadoi. H. Spermatheca, H. ungadensis Stys (E-H from Stys, 1970a). Dipsocoridae 7S 17 Hypsipterygidae by Pavel Stys General. Members of this group bear a striking re¬ semblance to many Tingidae. They are rather flattened dorsally and are 2-3 mm long. Diagnosis. Head porrect; all regions of head capsule sharply delimited; antennal segment 4 relatively short and thick, segment 3 very long (Fig. 17.1); labium straight, reaching or almost reaching base of metaster¬ num, segments 1, 2, and 4 distinct, very short, seg¬ ment 3 very long; pronotum with expanded lateral mar¬ gins and 3 longitudinal carinae (Fig. 17.1); forewing broadly explanate, tegminal, clavus sharply delimited, no costal fracture, no medial fracture, all veins raised, forming 9 large cells, entire wing finely areolate; legs strikingly slender; tarsi 2-segmented in both sexes; most abdominal sterna divided into mediotergites and latero- tergites; spiracles situated on laterotergites on segments 2-8 in males, on 3-7 in female; male pregenital abdomen symmetrical, pygophore, parameres, and laterotergites 9 asymmetrical, the last appendage-like; phallus com¬ plex, with long, coiled processus gonopori (Fig. 16. IF); Fig. 17.1. Hypsipterygidae. Hypsipteryx ectpaglus Drake (from Drake, 1961). female with large subgenital plate 7; all components of ovipositor present, modified; spermatheca present (Fig. 16.1H). Classification. The family was described by Drake (1961) as a subfamily Hypsipteryxinae [s/c] of Dipsocori- dae, and later it was raised to family rank by Stys (1970a). A single genus Hypsipteryx Drake includes H. ecpaglus Drake from Thailand, H. machadoi Drake from Angola, and H. ugandensis Stys from Uganda. Specialized morphology. The tingidlike appearance, thinness of the legs, and straightness of the labium are unique to the group. Natural history. Hypsipteryx ecpaglus, with fully de¬ veloped fore- and hind wings, was collected at light. All other Hypsipteryx specimens from Africa have the fore¬ wings slightly abbreviated and the hind wings minute, laceolate, and nonfunctional, and were collected from leaf litter or decaying wood. The slender legs suggest that Hypsipteryx species do not jump. Unlike other dip- socoromorphs from the Paleotropics, hypsipterygids are virtually absent from Berlese samples, suggesting they may have somewhat different life habits. Distribution and faunistics. This little-known Paleo- tropical group is treated by Drake (1961) and Stys (1970a). 18 Schizopteridae by Pavel Stys General. These cryptic bugs are sometimes remark¬ ably similar to certain beetles and members of the Omani- idae, owing to their often near-black coloration and uni¬ formly sclerotized, coleopteroid forewings. They range in length from 0.8 to 2.00 mm. Diagnosis. Small, compact, rotund; integument with pits, tubercles, a dense layer of microtrichia, or other ornamentation; body usually lacking conspicuous elon¬ gate vestiture common in most other Dipsocoromor- pha; head usually strongly declivent (Fig. 18.IB); com¬ pound eyes varying from relatively small to very large, sometimes extending posteriorly along nearly entire lat¬ eral margin of pronotum; ocelli present or absent in adults, either proximal to or distant from compound eyes; antennal segments 1 and 2 very short, subequal in length; labium with 3 or 4 segments, varying from short 80 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 18.1. Schizopterldae. A. Hypselosoma matsumurae Esaki and Miyamoto. B. Lateral view head, H. hirashimai Esaki and Miyamoto (A, B from Esaki and Miyamoto, 1959c). C. Male abdomen, dorsal view, Machadonannus ocellatus Wygodzinsky. D. Male abdomen, lateral view, M. ocellatus (C, D from Wygodzinsky, 1950). E. Male genitalia, Semangananus mirus Stys (from Stys, 1974). F. Spermatheca, Seabranannus ;m/77/fafor Wygodzinsky (from Wygodzinsky, 1950). and not extending beyond forecoxae to very long and 6/7; male pregenital abdominal segments sometimes all reaching well onto abdomen; proepistemal lobe inflated, symmetrical, more commonly segments 6-8 showing extending anteriorly to cover posteroventral surface of some modifications on the right-hand side (Fig. 18.1C, head; metastemum often with a median spine or pair D); male genitalia, including parameres, usually strongly of V-shaped processes, sometimes extending onto ab- asymmetrical (Fig. 18.1C, E); ovipositor developed or domen; hind coxa with adhesive pad on mesal margin, not; spermatheca present (Fig. 18. IF), apparently facilitating jumping; macropterous to strongly Classification. Reuter (1891) first accorded the taxon coleopteroid, forewings tegminal and nearly uniformly higher-group status, Schizopterina, as a subdivision of sclerotized, medial and costal fractures usually lacking; the Ceratocombidae. Reuter (1910) treated the group as a hind wings developed or not in nonmacropterous forms; family, a designation that has been followed by all subse- nymphal dorsal abdominal scent gland between terga quent authors. The Schizopterldae, as treated herein, are Schizopterldae 81 divided into two subfamilies. Although Emsley( 1969), in terinae, many genera in his scheme were left incertae his monographic work on the family, provided clear diag- sedis and have not yet been placed. At least 35 genera noses for his new subfamily Ogeriinae and the Schizop- and about 120 species have been described. Key to Subfamilies of Schizopteridae 1. Eyes exceedingly large, broadly overlapping anterolateral margins of pronotum (Fig. 18.1 A); oviposi¬ tor well developed, with first and second valvulae retained . Hypselosomatinae - Eyes of moderate size, overlapping at most anterolateral angles of pronotum; ovipositor reduced to a varying extent, sometimes completely absent . Schizopterinae HYPSELOSOMATINAE. Abdominal spiracles present on segments 2-8; ovipositor well developed. Three genera were originally included, Glyptocombus Heidemann and Ommatides Uhler from the New World and Hypselosoma Reuter, primarily from the Old World tropics. Hill (1980, 1984) described seven additional genera from the Australian mainland and Tasmania. SCHIZOPTERINAE. Abdominal spiracles usually present on sterna 6 and 7 and tergum 8, but other patterns known; ovipositor absent or poorly developed. The remaining genera of Schizopteridae are placed in this subfamily. Schizoptera Fieber, with several recog¬ nized subgenera, is by far the largest genus. The group is distributed worldwide, excluding the Palearctic. Specialized morphology. The typically globular bod¬ ies, relatively short first and second antennal segments, metasternal spine, and adhesive pads of hind coxae are all characteristics distinctive of the Schizopteridae. As with most other dipsocoromorphs, the male genitalia show strong dextral asymmetry. The male pregenital abdominal segments are often strongly modified, apparently forming secondary genitalia in Semangananus mirus §tys. Natural history. Emsley (1969), in the most complete account of schizopterid biology, showed that many mem¬ bers of the group are ground- and litter-dwelling. Yet, many taxa have never been found by collecting in these habitats but only by collecting at lights. Their attrac¬ tion to lights suggests broad-scale occupation of arboreal habitats. Schizopterids habitually jump and fly (winged morphs) when disturbed rather than run, as do other dip¬ socoromorphs. It appears that they develop only one egg at a time, which occupies nearly the entire abdomen. Hill (1980, 1984) aptly summarized information for the Tasmanian and mainland Australian hypselosomatines, indicating that the microhabitats include leaf litter, moss, grass tussocks, sedge and rush sods, and grass and fern foliage. Known major habitat types include rain forests, other wet forests, tropical palm swamps, elevated bogs, and sod tussock grasslands. Distribution and faunistics. As do all other dipso- coromorphan groups, the Schizopteridae show by far their greatest diversity in the tropics, although the group is by no means restricted to the strictly tropical latitudes, having been taken in Tasmania and as far north as south¬ ern Michigan in the United States. Many taxa remain to be described from the Indo-Pacific. The major comprehensive works on the group are those of McAtee and Malloch (1925), with keys to genera and emphasis on the North American species, and Emsley (1969), which includes a catalog to the world fauna and details on the fauna of Trinidad, with an excellent treat¬ ment of morphology for the group as a whole. Much of the remaining descriptive work has been done by Stys and Wygodzinsky, some of whose works are cited elsewhere in the text. Hill (e.g., 1980, 1984, 1990) has made sub¬ stantial contributions to the literature on the Australian fauna. 19 Stemmocryptidae by Pavel Stys General. Elongate, soft-bodied, light brown, 2.0-2.4 mm long, these obscure insects vaguely resemble some flattened, lightly sclerotized, free-living Cimicoidea. Diagnosis. Macropterous; no conspicuous sexual di¬ morphism; head porrect; ocelli postocular, adjoining hind margins of eyes; gula long; antennal segment 2 subequal in length to segments 3 and 4, the latter 2 only slightly thinner and devoid of long, thin, erect setae; labium about as long as head, directed forward (Fig. 19.1 A), segment 1 very short, segment 2 short, segments 3 and 4 longer; pronotum simple, subtrapezoidal, without dorsal collar; forewings membranous, venation reduced, medial frac¬ ture running in front of R (as in Enicocephalomorpha and Dipsocoridae), costal fracture long (Fig. 19.ID), clavus not delimited; forewings freely overlapping, not forming 82 TRUE BUGS OF THE WORLD (HEMIPTERA. HETEROPTERA) Fig. 19.1. Stemmocryptidae. Stemmocrypta antennata Stys (from Stys, 1983a). A. Habitus. B. Forewing. C. Hind leg, with detail ot adhesive pad on coxa. D. Male genitalia. E. Spermatheca. Abbreviation; ap, adhesive pad. claval commissure; legs short; tarsi swollen, tarsal for- Stemmocrypta antennata Stys, 1983a, from Laing Island, mula male 3-3-3, female 2-3-3, pretarsus with 2 long Papua New Guinea. setiform parempodia and a dorsal and ventral arolium; Specialized morphology. This peculiar group of in- metathoracic scent gland unpaired, with single opening, sects combines characteristics of the Dipsocoromorpha, evaporatorium present; abdominal spiracle 1 present in including the asymmetrical male genitalia, spermatheca, membrane posterior to metapostnotum, spiracles 2-7 on dorsal rmd ventral arolium, and tegminal forewings, with dorsal laterotergites; no remnants in adults of nymphal those of some Cimieoidea, including habitus and aiitennal dorsal abdominal scent glands; male abdominal seg- structure. ment 8 asymmetrical, with appendage-like laterotergites Natural history. Stemmocrypta antennata was col- (Fig. 19. ID); pygophore, laterotergites 9, and parameres lected in UV light traps and by berlesating leaf litter, asymmetrical; phallus complex; female sternum 7 form- Stemmocryptids are probably searching predators, and ing subgenital plate; ovipositor plate-shaped, modified; their short thick legs suggest they do not jump, spermatheca present (Fig. 19. IE). Distribution and faunistics. Known only from New Classification. This is the most recently discovered Guinea (Stys, 1983a). family of Heteroptera, being based on and including only Stemmocryptidae 83 NEOHETEROPTERA 20 Gerromorpha General. The Gerromorpha, or semiaquatic bugs, have been recognized as a monophyletic group since the time of Dufour (1833), but until recently they generally were called the Amphibicorisae (or Amphibiocorisae). Most members have the ability to walk on the water sur¬ face film, and some, such as most Gerridae and many Veliidae, spend nearly the entire active period of their lives on the water surface, a few even living on the open ocean. All taxa are predaceous, feeding upon other in¬ sects or arthropods. Most of the following material, in¬ cluding the key, is derived from the monographic work of Andersen (1982a; see also Spence and Andersen, 1994). The Gerromorpha, and particularly the Gerridae, have been the subject of study for their methods of loco¬ motion (Miyamoto, 1955; Andersen, 1976), methods of ripple communication and prey location on the water surface film (Wilcox, 1972, 1979, 1980; Wilcox and Spence, 1986; J. T. Polhemus, 1990b), wing polymor¬ phism and its determination (Brinkhurst, 1959, 1963; Andersen, 1973; Vepsalmnen, 1974a, b), and spatial competition and coexistence (e.g., Spence and Scudder, 1980; Spence, 1983). Diagnosis. Head in most families elongate and often more or less cylindrical and protruding distinctly anterior to the eyes, head much shorter in Veliidae and Gerridae, appendages elongate and slender (Fig. 20.2B), and col¬ oration somber. Characters that Andersen (1982a) listed as synapomorphic are, among others; 3 pairs (rarely 4) of cephalic trichobothria inserted in deep cuticular pits in the adult (Fig. 10.8A); epipharynx with a long, nar¬ row, external projection; mandibular levers quadrangular; maxillary levers absent; rostral groove distinct and ac¬ cepting labium at rest (Fig. 22. IB); labium elongate. segments 1 and 2 very short, segment 3 very long, much longer than segment 4 (Figs. 21.IB, 22.IB), except in Veliidae, Gerridae, and Hermatobatidae, in which labium generally much shorter, segment 3 usually not conspicu¬ ously longer than remaining segments, and rostral groove not developed; forewings usually not differentiated into anterior coriaceous and posterior membranous portions (Figs. 20.2B, 21.1C); pretarsus with dorsal and ven¬ tral arolium (Fig. 10.5C, D), ventral arolium sometimes highly modified; peg plates (sieve pores) generally dis¬ tributed on body surface (Fig. 10.7C), except in Gerridae and Hermatobatidae; 2-layered hair pile usually covering head, thorax, and basal segments of the abdomen, con¬ sisting of short, densely placed microtrichia, and longer, less densely set macrotrichia; female gynatrial complex with long, tubular, entirely glandular spermatheca and secondary fecundation canal (Figs. 21.2K, 26.2F). Pterygopolymorphism is common, it sometimes shows continuous variation, or there may be only macropter- ous and/or micropterous or apterous morphs. Numerous species and many suprageneric taxa are known only from the apterous morph. The details of the structure and function of the specialized body hair layers found in all Gerromorpha have been studied by Andersen (1977b). Discussion. General treatments include The Semi¬ aquatic Bugs (Andersen, 1982a), which is the most comprehensive reference available, including sections on morphology, classification, phylogeny, biogeography, functional adaptations, list of genera and suprageneric taxa, and keys for their identification. Andersen’s dado- gram of family relationships is shown in Figure 20.1. The Semiaquatic and Aquatic Hemiptera of California (Menke, 1979a) is a useful reference for North American Fig. 20.1. Phylogenetic relationships otfamiiies of Gerromorpha (after Andersen, 1982a). 84 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 20.2. Gerromorpha. A. Hebrus sobrinus Uhler (Hebridae). B. Macrovelia hornii (Uhler) (Macroveliidae) (A, B from Usinger, 1956). C. Oravelia page Drake and Chapman (Macroveliidae) (from Drake and Chapman, 1963). D. Paravelia sp. (Veliidae: Veliinae) (from Slater, 1982). C Fig. 20.3. Gerromorpha. A. Rhagovelia distincta Champion (Veli- idae: Rhagoveliinae) (from Usinger, 1956). B. Gerris incurvatus Drake and Hottes (Gerridae: Gerrinae). C. Metrobates trux (Torre-Bueno) (Gerridae: Trepobatinae) (from Usinger, 1956). D. Halobates sericeus Eschscholtz (Gerridae: Halobalinae) (from Zimmerman, 1948). 86 TRUE BUGS OF THE WORLD (HEMIPTERA; HETEROPTERA) genera and for the species known to occur in California. Other regional treatments for North America are those of Brooks and Kelton (1967), Bobb (1974), and Bennett and Cook (1981). The West Indian fauna is covered in part by Cobben (1960b, c). The fauna of the western Palearctic is treated in the works of Stichel (1955), Poisson (1957b), Kerzhner and Jaczewski (1964), and Savage (1989) and that of the eastern Palearctic by Kanyukova (1988). Pois¬ son (1957a) prepared a preliminary account of the fauna of South Africa. The work of Linnavuori (1971, 1981, 1986) covers much of northern Africa and the Arabian Peninsula. The last comprehensive work for India was that of Distant (1904, 1910), for Indonesia Lundblad (1933), and for Australia that of Hale (1926). The habi¬ tats, habits, and distributions of the marine Gerromorpha were treated by Andersen and Polhemus (1976). Key to Families of Gerromorpha 1. Macropterous . 2 - Apterous or short-winged . 7 2. Scutellum exposed, forming subtriangular, rounded, or transverse plate behind pronotal lobe (Fig. 20.2A) . 3 - Scutellum not exposed, covered by posteriorly extended pronotal lobe (Fig. 20.2B) . 4 3. Ventral surface of head with a pair of well-developed bucculae covering base of labium (Fig. 22.1B); tarsi 2-segmented, first segment very short . Hebridae (part) - Bucculae not so well developed and covering base of labium as above; tarsi 3-segmented . . Mesoveliidae (part) 4. Bucculae covering first and sometimes also second labial segments laterally; claws inserted apically on last tarsal segment (Fig. 25.1C) (slightly preapically in Limnobatodes Hussey, Hydrometridae) . 5 - Bucculae not so well developed and not fully covering basal labial segments; claws inserted distinctly before apex of last tarsal segment (Figs. 27. IG, H) (with a few exceptions) . 6 5. Head not distinctly prolonged behind eyes; length of postocular area less than the diameter of an eye; forewing with 6 closed cells (Figs. 20.2B, 24. IB) . Macroveliidae - Head distinctly prolonged behind eyes; length of postocular area longer than diameter of an eye; forewing with 3 closed cells (Fig. 25,1 A) . Hydrometridae (part) 6. Head with a distinct longitudinal, median impressed line on dorsal surface (Figs. 20.3A, 27.1 A); foretibia of male usually with a grasping comb of short spines along inner margin; hind femur usually stouter than middle femur . Veliidae - Head without a median impressed line on dorsal surface; foretibia of male without grasping comb; hind femur usually more slender than middle femur . Gerridae 7. Abdominal scent-gland orifice absent from metastemum . 8 - Abdominal scent-gland orifice present on metastemum . 9 8. Head much more than 3 times as long as wide (Fig. 25.IB), eyes far removed from base of head . Hydrometridae (part) - Head at most 3 times as long as wide, eyes situated very close to or at base of head . 6 9. Pronotum very short, exposing both meso-and metanotum . 10 - Pronotum longer, at least covering mesonotum . 11 10. Head longer than wide, porrect (Fig. 21.1 A); all claws apical (Fig. 21. IF) . . Mesoveliidae (part) - Head shorter than wide, subvertical (Figs. 26.1 A, B); claws of forelegs subapical, claws of middle and hind legs apical (Fig. 26.1C) . Hermatobatidae 11. Bucculae well developed, covering base of labium; tarsi 2-segmented . Hebridae (part) - Bucculae not so well developed and not obscuring basal labial segments; tarsi 3-segmented . ..12 12. Antennal segment 4 not subdivided in middle by membranous zone . 13 - Antennal segment 4 subdivided in middle by a membranous zone . Paraphrynoveliidae 13. Head not distinctly prolonged behind eyes; length of postocular region longer than diameter of an eye . Macroveliidae (part) - Head distinctly prolonged behind eyes; length of postocular region longer than diameter of an eye . Hydrometridae (part) Gerromorpha 87 Mesoveloidea 21 Mesoveliidae General. This small group of insects is thought to be the relatively most primitive of all semiaquatic bugs. They vary greatly in habitus (Fig. 21.1 A) and degree of wing development and range from 1.2 to 4.2 mm in length. Members of the widely distributed genus Meso- velia Mulsant and Rey are sometimes referred to as water treaders. Diagnosis. Micro- and macrohair layer restricted to head and prosternal region of thorax; ocelli present or absent; head without dorsal indentations and correspond¬ ing internal apodemes; base of labium not obscured by bucculae (Fig. 21. IB); pronotum truncate posteriorly; scutellum developed and exposed (Fig. 21.IH); fore¬ wing venation reduced (Fig. 21.1C); tarsi 3-segmented (Fig. 21. IE, F); pretarsus inserted apically or subapically; first abdominal mediotergite with a pair of longitudi¬ nal ridges in macropterous forms (Fig. 21. IH); aedeagus (Fig. 21.11) with specialized ejaculatory pump and bulb; parameres symmetrical; ovipositor usually laciniate (Fig. 21.IJ); gynatrial complex as in Fig. 21 IK; anterior end of egg obliquely truncate with an egg cap developed in many species: eclosion by means of an embryonic bladder rather than an egg burster. Classification. The most important single paper on the group is that of Andersen and Polhemus (1980), in which four new genera were described and a phylogeny and a classification for the family were presented, with two subfamilies being recognized. Previously China and Miller (1959) recognized a more broadly conceived group with the subfamilies Mesoveliinae, Mesoveloideinae, and Macroveliinae. Stys (1976) recognized the Macroveliidae as a distinct family, as had earlier authors. The name Madeoveliidae was proposed by Poisson (1959) and is a senior synonym of Mesoveloideinae of China and Miller (1959). Horvath (1929) provided a catalog of the species. Eleven genera and about 39 species are currently recog¬ nized. Key to Subfamilies of Mesoveliidae 1. Macropterous . 2 - Wingless . Mesoveliinae 2. Ocelli lacking; claws inserted subapically on tarsi (Fig. 21.IE); forewing with 2 basal cells and one apical cell (Fig. 21.1C) . Madeoveliinae - Ocelli present; claws inserted apically on tarsi (Fig. 21. IF); forewings with 2 or 3 basal cells but no apical cell (Fig. 21. ID) . Mesoveliinae MADEOVELIINAE. Head deflected in front of eyes; ocelli absent; scutellum subtriangular; forewing with 2 basal cells and one apical cell (Fig. 21.1C); pretarsus inserted anteapically (Fig. 21. IE). Included genera are Mesoveloidea Hungerford, two species from tropical America, and Madeovelia Poisson, one species from Guinea. MESOVELIINAE. Ocelli present in macropterous forms; scutellum reduced, apically rounded (Fig. 21. IH); pre¬ tarsus inserted apically (Fig. 21. IF); forewing with 3 basal cells, no apical cells (Fig. 21.ID); most genera apterous. Included genera are Austrovelia Malipatil and Mon- teith, two species from Australia and New Caledonia; Cavaticovelia Andersen and Polhemus, one species from Hawaii; Cryptovelia Andersen and Polhemus, one species from the lower Amazon basin; Darwinivelia Andersen and Polhemus, two species from northern South America and the Galapagos Islands; Mesovelia Mulsant and Rey, about 25 species, cosmopolitan; Mniovelia Andersen and Polhemus, one species from New Zealand; Nereivelia J. and D. Polhemus, one species from Singapore and Thailand; Phrynovelia Horvath, three species from New Guinea and New Caledonia; and Speovelia Esaki, two species from Mexico and Japan. Specialized morphoiogy. Cryptovelia terrestris An¬ dersen and Polhemus, among the smallest of all known Heleroptera, is apterous and has compound eyes consist¬ ing of only 3 or 4 facets (Fig. 21. IB). On the other hand Mniovelia kuscheli Andersen and Polhemus from New 88 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 21.1. Mesoveliidae (from Andersen, 1982a). A. Adult female, Cryptovelia terrestris Andersen and Polhemus. B. Lateral view. C. terrestris. C. Forewing, Mesoveloidea williamsi Hungerford. D. Forewing, Mesovelia mulsanti White, E. Left middle pretarsus, Mesoveloidea williamsi. F. Left middle pretarsus, C. terrestris. G. Unguitractor plate and parempodia, Mesoveloidea williamsi. H. Dorsal view, thorax and base of abdomen, Mesovelia mulsanti. I. Aedeagus, Mesovelia mulsanti. J. Female terminal abdominal segments, ventral view, Mesovelia mulsanti. K. Gynatrial complex, Mniovelia kuscheli Andersen and Polhemus. Abbreviations: cs, campaniform sensillum; gs, gynatrial sac; st, spermathecal tube. Mesoveliidae 89 Zealand—also very small—has very large eyes com¬ pared with other Gerromorpha. Three thoracic morphs associated with degree of fore¬ wing development were recognized by Galbreath (1975) in Mesovelia mulsanli White, but she was not able to demonstrate definitively the mechanism for their determi¬ nation, although environmental factors appeared to play an important role. Natural history. The life history of several Mesovelia spp. in the Holarctic has been studied in detail (Hunger- ford, 1917; Ekblom, 1930; Zimmermann, 1984). The females insert the eggs into plant tissue with their elon¬ gate ovipositor, laying 100 or more eggs. In very cold regions the bugs apparently overwinter as eggs, whereas in warmer climates adults may be present and active year round. Most Mesovelia spp. inhabit the margins of ponds of streams, some leading a rather secretive exis¬ tence, others spending most of their time in the open. They are extremely agile on the open water surface, although they are usually associated with some form of aquatic vegetation. Mesoveloidea has been found living on wet moss near streams; Cryptovelia. Mniovelia, and Phrynovelia are known to inhabit forest leaf litter, often at a considerable distance from the nearest body of water. Cavaticovelia and some Speovelia spp. are cave- inhabiting (Esaki, 1929; Gagne and Howarth, 1975). All Speovelia spp., some Mesovelia spp., Nereivelia, and Darwinivelia are associated with intertidal marine habi¬ tats (Andersen and Polhemus, 1980; Carvalho, 1984a). Female Mesovelia furcata Mulsant and Rey possess symbiotic inclusions in the midgut wall. These are appar¬ ently transmitted by the female to the egg shortly after oviposition, as the female taps the opercular end of the egg with the anus (Cobben, 1965). The macropterous morph in some species of Mesovelia (e.g., M. furcata; Jordan, 1951) mutilates the wings by tearing the distal portions with the legs, thus losing the ability to fly. Mesovelia amoena Uhler is parthenogenetic in Hawaii and tropical areas, males having been collected only in North America, Mexico, and Hispaniola (J. T. Polhemus and Chapman, 1979c). Distribution and faunistics. Mesovelia is widely dis¬ tributed, with a few of the included species also having very wide distributions. All other taxa are of limited dis¬ tribution and widely scattered. This fact and the basal phylogenetic position within the family suggest an ancient group. Andersen and Polhemus (1980) included a checklist of species and distributional record and Andersen (1982a) provided much additional information on the family. J. T. Polhemus and Chapman (1979c) supplied a useful review of the group for North America, with emphasis on the fauna of California. Hebroidea 22 Hebri(jae General. Sometimes referred to as velvet water bugs, hebrids are some of the smallest members of the Ger¬ romorpha, ranging in length from 1.3 to 3.7 mm, and having the general appearance of very small veliids (Figs. 20.2A, 22.1 A) They are infrequently encountered by the general collector because of their minuteness and often secretive habits. Diagnosis. Micro- and macrohair layer covering body, except abdomen, and appendages; antennae often rather short, segment 2 subequal to or shorter in length than segment 1 (Fig. 22. ID, E), segment 4 undivided in He- brus (Subhebrus), Lipogomphus, Merragata, and Hyrca- nus and with a median constricted membranous zone in all other taxa, giving 5-segmented appearance (Fig. 22. IE); ocelli present, each with a deep pit anterior to it, corresponding to an internal apodeme; bucculae promi¬ nent, obscuring 2 basal segments of labium (Fig. 22. IB, C); labium short in Hyrcanus; pronotum truncate along posterior margin, usually exposing short transverse scu- tellum and posteriorly adjoining triangular metanofum (Fig. 20.2A); thorax ventrally with a pair of longitudi¬ nal carinae between coxae, forming rostral groove (Fig. 22.1C); adult tarsi 2-segmented, segment 2 representing fusion of segments 2 and 3 of other Gerromorpha (Fig. 22. IG); pretarsus inserted apically (Fig. 22. IG); fore¬ wing with 1 or 2 basal cells, no venation on distal portion (Fig. 22.1 A, F); genitalia apparently inserted anteapically on abdomen in both male and female; pygophore small; parameres usually symmetrical (Fig. 22. IH); aedeagus as in Fig. 22.11; ovipositor valves much reduced, weakly sclerotized, more platelike than in Mesoveliidae; gyna- trial complex as in Fig. 22. IJ. Classification. Two subfamilies are currently recog¬ nized, following the classification proposed by Andersen (1981b, 1982a). Approximately 160 species are placed in seven genera. 90 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 22.1. Hebridae (from Andersen, 1982a). A. Merragata brunnea Drake. B. Lateral view, head, Timasius ventralis Andersen. C. Ventral view, Hebruspusillus (Falien). D. Antenna, T. ventralis. E. Antenna, Hyrcanus capitatus Distant. F. Forewing, H. capitatus. G. Foretarsus. H. capitatus. H. Dorsal view, pygophore and proctiger, T. ventralis. i. Aedeagus, H. capitatus. J. Gynatrlal complex, T. ventralis. Abbreviations; da, dorsal arolium; fc, fecundation canal; pa, paramere; rg, rostral groove; st, spermathecai tube; va, ventral arolium. Hebridae 91 Key to Subfamilies of Hebridae 1. Antennal segments slender, segment 4 much longer than segment 1 . Hebrinae - Antennal segments stout, segment 4 subequal in length to segment 1 (Fig. 22. ID) . 2 2. Large species more than 2.5 mm. in length; head narrow and pointed apically; labium not reaching mesosternum . Hyrcaninae - Small species not more than 2.5 mm. in length; head broad not pointed apically; labium reaching metasternum . Hebrinae HEBRINAE. Antennal segment 4 with preapical cluster of modified blunt setae; eyes situated near base of head (Fig. 20.2A); antennae distinctly longer than head; labial segment 3 not surpassing hind margin of head, dorsal and ventral arolia subequal in length; parameres symmetrical, except in Timasius. Included genera are Hebrometra Cobben, four species from Ethiopia (Cobben, 1982); Hebrus Curtis, largest genus in the family, with approximately 110 described and many undescribed species distributed on all major land masses; Lipogomphus Berg, four species from South and Central America and the southern United States (Andersen, 1981b); Merragata Buchanan-White, several species, cosmopolitan; Neotimasius Andersen, one spe¬ cies from Southern India; and Timasius Distant, 15 spe¬ cies from Sri Lanka, parts of India, Taiwan, southeast Asia and Java (Andersen, 1981b). HYRCANINAE. Eycs distinctly removed from anterior margin of the pronotum; length of antenna subequal to or shorter than length of head; labial segment 3 not surpass¬ ing hind margin of head; dorsal arolium distinctly shorter than ventral arolium (Fig. 22. IG); parameres symmetri¬ cal. Hyrcanus Distant, with four species from the Oriental region, is the only included genus (Andersen, 1981b). Specialized morphology. At least in Hebrus ruficeps, there exist, in addition to the peg plates, some small (6- 10/xm) cup-shaped or mushroomlike structures, scattered among the hair layer. In addition, in most hebrids, an¬ tennal segment 4 possesses subapically a small group of short, blunt setae of unknown sensory function (Cobben, 1978). The structure and function of the male and female reproductive systems have been studied in detail for He¬ brus pusillus and H. ruficeps (Heming-Van Battum and Heming, 1986, 1989). Natural history. Hebrids live on vegetated margins of ponds or similar permanently damp habitats, some¬ times deep in mats of moss or frequently in interstices, as well as among sparse vegetation on sloping stream banks. Some species are more specialized in their habitat requirements, such as members of the ge.ne.v 2 i Hebrometra and Timasius, which live on rocks in streams or torrents or on seeping rock faces or those splashed by waterfalls. A few species tolerate brackish, saline, or marine condi¬ tions. Hebrus ruficeps can overwinter frozen in ice among Sphagnum. As far as is known, hebrids lay their eggs superficially on a substrate, such as on mosses, attaching them in a lengthwise position with a gelatinous substance. Distribution and faunistics. The Hebridae are nearly worldwide in distribution, with the greatest generic di¬ versity in the Asian tropics. Drake and Chapman (1958) provided a checklist of New World species. Stichel (1955) treated the European fauna, Miyamoto (1965a) the spe¬ cies from Taiwan, Poisson (1944) some species from Africa, and Cobben (1982) those known from Ethiopia. 23 Paraphrynoveliidae General. This family, with no common name, in¬ cludes only one genus and two species, which range in size from 1.7 to 2.4 mm. Its members, which have the general appearance of wingless hebrids or Microvelia spp. (Fig. 23.1 A), have rarely been collected. Diagnosis. Microvelia-like in general appearance; all known specimens apterous; ocelli absent (Fig. 23.IB); antennae flagelliform, segments 1 and 2 elongate, sub¬ equal in length, segments 3 and 4 longer, segment 4 with a medial membranous area with ringlike cuticular thickenings; pronotum short (Fig. 23. lA), produced pos¬ teriorly to cover mesonotum but not metanotum; tarsi 3- segmented (Fig. 23. IE); pretarsus inserted apically, with well-developed dorsal and ventral arolia (Fig. 23.IE); abdominal dorsum as in Fig. 23.ID; ovipositor rather weakly developed (Fig. 23.1C); pygophore protruding from end of abdomen; parameres (Fig. 23. IF) symmetri¬ cal; gynatrial complex as in Fig. 23. IG. 92 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Rft. PMptV>no^icto« dftwtf RbIMW (from 166 ( 2 at A laierat vi»» t v>ftw. heiad C V^tral Ihom and iMtanian Di knamaJ view. praganM MQvnanto of afadowniit ddnum inoMo^ scent appereius. E. MiddAa tarsus F. Left parameie Ql Qyfuitntf oompieai. dorwd viee AOCmvtelkm da. ddrsil arotlum, va. venM aroftum Ctftftftificftllon. PKimphytwvrHa Pbsvvm wus ortg»> naJiv placed in die MeMweiiidac Mon: dcUiled viudy ftuggetiited. htiwever, that the kmi^n Kpecic^ Kbciuld he placed tn a dts(tiK( family in a clade including the Macniveliidae and Ilydnwneiridac fAndet%en« 197H) Nntumi Mttory. Pam/xArytimWrci live in the Iran- 24 Macroveliidae Oanarai. Memliers of this small family hase no com- nam name They ate similar to M>mr meuiveliMli^ in gen • end appearance (Fig. 20.2B. C) and rapge in length fitnn 2 ^ to 5.6 mm. Dtagnonla. Micro* and macrotuir layer generally dts* tnbuled. but reaching only onto segment 1 of abdomen; ocelli present in macroptcrotis Mocrovtlia (F^. 20.2Bh siUon rone hcliseen land and water iti wet debris and water*Miaked moss on nicks (Andersen. 1*^8y DistribuBona a«>d teuryaUcs. This group is rcsirictcd to sotnhem Atnca. Informatum on the details of distrihu* tinsi were given hy Ponacm { 1^7a)^ and Anderven (1978« 1982a) (sec Chapter 9t. reduced to Ontvriia, and absent in Che/Mrue/io; antennae fUgclliform. segments 1 and 2 ciongatc (Fig. 20.2B. C); Labial segments I and 2 very short, segmcni 3 approxi* mately 2 5 times length of segment 4, labiuin reaching to about mesocoxac (Fig 24,1 A), prtmotum in Mocro\'efia extending postenorly to cover nidimcnlary tcutellum. meaonocum. and metanotum (Fig. 20.2H), hut truncate in apterous Chepuvrita and Ora\'ttia (Fig. 20.2C); (orewmg venatioQ in Macrav^fki as in Figs. 20.2B, 24.IB; tarsi 3‘Scgmenced, pretarsus inserted apically (Fig 24. ID); pygophore inserted apically on abdomen; poramcrcstHg. 24.IE) symmetncal. aedeagus as in Fig. 24. IF; Macro- vMa ovipositor platc-shapcd. similar to Hcbridae. but Macrovakidae B9 MftcfOMtMM homif \JNm ltm» A, uumi hMd 0.ntg|WlMMrtng D.Lail rTK3i^ tarsos. A* Py^ophoraandp>octiqBr. IMnl nm. P, AadMQUlL wMraJ vww O. G ynairia)oompleK.itoraaJ viaw. AUbfvnaiKin: pa, paramare kKoicd 01 apex itf oMmiCQ (Fig^ 24, IC); gyoalnaJ com* plex aA in Fig 24, IQ. ClnaatflCAtlon. The composition of the Macroveliidae has varied Loiisider'ahly at the lunds i>( dilTcrctU authors. The current classitkaiion was established by ficy^ (1976) and Andcn^n 1197K. t982a). It comprises C/iepuiWiu China (1962). one species from southern Chile; Mainp- le/tiii Uhkr. one species from western North Amerka; and Orux'rtia Drake and Chapman, one species from Frrsno County. Calilomia Spaciaiiamd morphology. iformvWio hfirmi Uhkr may be either rnacnifMetous or hrachyptcroos Onnriia and ChrfHn^lMi are knmn only (nun apterous individu* ak, f4atural hlatory, Chfpti\Hia usinneri China is found in ykKt forest litter, all known spccimem having been colkcted by silting and using Bcrksc tunnels. Macro- \rlia hymii and <>ru^c^i^i pei^c l>rake and Chapman in* habit sprii^ts or seeps with abundant xegetatHin; they ane negatively pboioiiophic and often secrete thcmsclrvcs in shaded areas. Matrosdiu hornti oserwinters as an adult and may be active during warm periods even in midw inter tMcKinsiry'. 1942: Anderson. 1963; Drake and Chapman. 1963; J. T. R)lhcmus and Chapman, 197%) The eggs of Macroretm are glued to the substnile in a loQgihiidigrf pcnilkm. Distribution and taunlatica. J. T Poliemus and Chapnun i IM7V)h) sunmtan/ed availabk inliirtiuiliun on the Nordi American taxa. rnuE eucs Of vvorlo hctgropt&U) Hydrometroidea 25 Hydrometridae General. This is one of the most distinctive heterop- teran groups, many members having an extremely elon¬ gate body and appendages; all taxa have the eyes far removed from the anterior margin of the pronotum (Fig. 25.1 A, B). Commonly called marsh treaders or water measurers, they range in length from 2.7 to 22 mm. Diagnosis. Extent of micro- and macrohair layer vari¬ able; head elongated in front of and behind compound eyes (Fig. 25.1 A, B); posterior pair of cephalic tricho- bothria inserted on tubercles (Fig. 25.1 A), except in Hydrometra; ocelli present or absent; antennal segment 4 with an apical invagination, generally bordered by modified setae; tarsi 3-segmented (Fig. 25.1C); pretarsus usually inserted apically, ventrally in Limnobatodes, dor¬ sal and ventral arolium usually both well developed (Fig. 25.1C); forewing development variable, venation usually somewhat reduced (Fig. 25.1 A); pygophore protruding apically from abdomen; phallus as in Fig. 25.IE; para- meres symmetrical (Fig. 25.ID); ovipositor reduced in most species; gynatrial complex as in Fig. 25. IF. Classification. Although it has long been clear that Hydrometra was a member of the Gerromorpha (Ekblom, 1926; Andersen, 1982b), the position of all members of the group was not always so obvious. For example, Heterocleples was originally placed in the Reduviidae by Villiers (1948). Three subfamilies—comprising seven genera and more than 110 species—are currently recog¬ nized, following the classification of Andersen (1977a, 1982b). Key to Subfamilies of Hydrometridae 1. Antennal segment 1 subequal to or shorter than segment 2. usually at most barely exceeding apex of head; body length 6 mm or more; metasternum lacking scent-gland orifices . Hydrometrinae - Antennal segment I much longer than segment 2, surpassing apex of head by more than half its length; body length 3-5 mm; metasternum with scent gland orifices present . 2 2. Ocelli present; posterior pair of cephalic trichobothria inserted on prominent rounded elevations (Fig. 25.1 A); head and pronotum clothed only with simple setae; articulation between antennal segments 1 and 2 subapical . Heterocleptinae - Ocelli lacking; posterior pair of cephalic trichobothria inserted on small tubercles; head and pronotum clothed with stout black spinules; articulation between antennal segments 1 and 2 apical . . Limnobatodinae HETEROCLEPTINAE (FIG. 25.1 A). Micro- and macrohair layer covering head, thorax, and base of abdomen; poste¬ rior cephalic trichobothria very long, inserted on promi¬ nent round elevations; ocelli present (even in apterous morphs of Veliometra)\ antennal segment 1 longer than segment 2, antennal segment 2 inserted laterally and somewhat anteapically on segment 1, antennal segment 4 in Heterocleptes with a membranous ring distad of midpoint, producing 5-segmented appearance; pronotum relatively short and broad, produced posteriorly over the metanotum, even in apterous morphs; arolia rudimentary, parempodia padlike in Heterocleptes; peg plates present. Included genera are Heterocleptes Villiers, four species from Africa and Borneo (Villiers, 1948; China and Usin- ger, 1949b; China, Usinger, and Villiers, 1950; Ander¬ sen, 1982b), and Veliometra Andersen, one species from the central Amazon basin of Brazil (Andersen, 1977a). HYDROMETRINAE. Micro- and macrohair layer covering entire body; anteocular portion of head much longer than length of pronotum; antennal segment 1 subequal to or shorter in length than segment 2; metasternal scent glands absent. Included genera are Bacillometra Esaki, four species from tropical South America (Esaki, 1927); Chaetome- tra Hungerford, one species from the Marquesas Islands; Dolichocephalometra Hungerford, one species from the Marquesas Islands; and the cosmopolitan Hydrometra Latreille, with at least 80 species (Andersen, 1977a). LIMNOBATODINAE. Sharing many features with the Hy¬ drometrinae; distinguished by micro- and macrohair layer covering head, thorax, and base of abdomen; anteocu¬ lar portion of head shorter than pronotal length; antennal segment 1 distinctly longer than segment 2; claws in¬ serted slightly before tarsal apex; metasternal scent glands present. Limnobatodes paradoxus Hussey, known from Belize, Hydrometridae 95 - cs Fig. 25.1. Hydrometridae (from Andersen, 1982a). A. Heterocleptes hobeiiandti China and Usinger. B. Lateral view, head and thorax, Hydro- metra sp., macropterous female. C. Left middle tarsus, Veliometra schuhi Andersen. D. Pygophore and parameres, dorsal view, V. schuhi. E. Phallus, Hydrometra stagnorum (Linnaeus), lateral view. F. Gynatrial complex, Heterocleptes hoberlandti. Abbreviations: da, dorsal arolium; cs, campaniform sensillum; pa, paramere; va, ventral arolium. Brazil, and Peru (Hussey, 1925; Andersen, 1977a) is the only included taxon. Specialized morphology. The hydrometrids, particu¬ larly the Hydrometrinae, possess many modifications as¬ sociated with elongation of the body and appendages, these particularly affecting the head and thorax (Fig. 25. IB). Other novel structures are mentioned in the diag¬ nosis. Males of some Hydrometra spp. have a clump of modified setae ventrally on abdominal segment 7 (Fig. 10.7E). A thorough review of the morphology of Hydro¬ metra martini Kirkaldy was provided by Sprague (1956). Natural history. Veliometra has been collected in the marginal vegetation of an impondment of an Amazonian forest stream. The scant data available for Heteroclep¬ tes and Limnobatodes suggest that at least the former is semiterrestrial, most collections of the latter being from lights. Hydrometra spp. are usually found on or around quiet bodies of water and generally are associated with 96 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) marginal vegetation but may also be found on damp rock walls. Hydrometra spp. can walk on the water surface with great agility and apparent effortlessness. The eggs of Hydrometra are placed some distance above the water level and are cemented to the substrate in an upright position by their base (Sprague, 1956). Similarities in structure suggest a like habit for Limno- bates. The eggs of Veliometra are more similar in struc¬ ture to those of other nonhydrometrid Gerromorpha and are probably cemented in a horizontal position in closer association with water (Andersen, 1982a). Distribution and faunistics. The family is most di¬ verse in the tropics, with only species of Hydrometra occurring elsewhere. The group is notable for the occur¬ rence of the endemic genera Chaetometra and Dolicho- cephalometra in the Marquesas Islands, an area from which nearly all other gerromorphans are absent. Hunger- ford and Evans (1934) and Drake and Lauck (1959) offer the most comprehensive distributional surveys, for the world and the Western Hemisphere, respectively. Gerroidea 26 Hermatobatidae General. Members of this small, seldom-collected group of ovoid-bodied marine bugs closely resemble short-bodied apterous gerrids. Sometimes referred to as coral treaders, they range in length from 2.7 to 4.0 mm. Diagnosis. Micro- and macrohair layers covering en¬ tire body; peg plates absent; head extremely short and broad, anteclypeus and antennal fossae hidden in dorsal view (Fig. 26.1 A, B); ocelli absent; 3 pairs of cephalic trichobothria; antennae relatively long, segment 2 long¬ est, inserted relatively close together and far from anterior margin of eye; labium short and stout, segments 1 and 4 subequal in length to segment 3, segment 2 shorter (Fig. 26.1 A, B); always apterous; prothorax short; meso- and metathorax indistinguishably fused, except laterally; pro- thoracic sternum narrow, meso- and metathoracic sterna broad; all coxae inserted in a horizontal position and directed caudad (Fig. 26. ID); forefemur incrassate, fore- tibia slightly curved, with an elongate apical groom¬ ing comb; femur and tibia of middle and hind legs more elongate and slender than foretibia; all tarsi 3- segmented, pretarsus of foreleg inserted anteapically. claws slender, strongly developed, pretarsus of middle and hind legs inserted apically (Fig. 26.1C); dorsal aro- lium flattened horizontally and tapering, ventral arolium 3-branched (Fig. 26.1C); parempodia long, setiform. and symmetrically developed (Fig. 26.1C); metathoracic scent-gland apparatus present; abdomen highly modified, greatly shortened (Fig. 26. ID); scent-gland apparatus present on abdominal segment 4 in nymphs; parameres reduced, left fused to margin of pygophore, right in form of a small plate loosely connected with pygophoral mar¬ gin; aedeagus as in Fig. 26.IE; female genital segments simple, ovipositor absent; gynatrial complex as in Fig. 26. IF; 4 ovarioles. Classification. Carpenter (1892) described Hermato- bates in the Gerridae. Coutiere and Martin (1901) recog¬ nized it as the subfamily Hermatobatinae. Poisson (1965) accorded the group family status, an action supported by more recent authors (§tys and Kerzhner, 1975; Ander¬ sen and Polhemus, 1976; Andersen, 1982a). Andersen (1982a) argued that Hermatobates Carpenter is the sister group of Veliidae + Gerridae. Most of the eight known species were described from only one or a few specimens. Specialized morphology. The head of Hermatobates is extremely short and broad, only apterous forms are known, and the abdomen is strongly shortened and shows substantial fusion of terga and sterna, especially in females. Natural history. Hermatobates spp. are associated with coral reefs and rubble. They secrete themselves in an air bubble at high tide, coming out to feed during low tide. Those that do not retreat before the rising tide are able to continue skating or resting on the water surface until the next tidal cycle. They move very rapidly on the water surface and can jump a considerable distance. There are apparently only four nymphal instars (Andersen, 1982a; Foster, 1989; D. A. Polhemus, 1990a). Distribution and faunistics. Most of the known spe¬ cies occur in the Indian or Pacific ocean. A single species. Hermatobates breddini Herring, is known from the West Indies. In addition to the work of Andersen (1982a), useful papers on the group include those of Coutiere and Martin (1901), China (1956), Herring (1965), Cheng (1977), and J. T. Polhemus and Herring (1979). Hermatobatidae 97 Fig. 26.1. Hermatobatidae. Hermatobates weddi China (from Andersen, 1982a). A. Lateral view, body. B. Ventral view, head and protho¬ rax. C. Pretarsus, middle leg. D. Ventral view, male pterothorax and abdomen. E. Aedeagus, dorsal view. F. Gynatrial complex, dorsal view. Abbreviations: da, dorsal arolium; pa, parempodium; va, ventral arolium. r 27 Veliidae General. Sometimes referred to as broad-shouldered water striders, small water striders, or riffle bugs, the Veliidae range from about 1 to 10 mm in length. Most species are rather stout-bodied. They occupy a wide range of habitats, including the surface of the ocean, and, along with the Gerridae, represent the epitome of adaptation for life on the water surface film. Diagnosis. Entire body surface (exeept abdomen in Ocellovelia China and Usinger), covered with macro- and microhair layer; all members with “grooved setae”; head rather short and broad, with a median impressed dorsal line and with a pair of deep pits near postero- mesal angle of eyes, representing external evidence of internal apodeme for attachment of antennal muscula¬ ture; compound eyes usually large (Figs. 20.2D, 20.3A), sometimes reduced; ocelli absent (except in Ocellovelia)-, cephalic trichobothria as in Fig. 10.8A; antennal sock¬ ets usually located beneath eyes and obscured in dorsal view (Fig. 27.1A-C) (in contrast to foregoing families of Gerromorpha); bucculae relatively small, not obscuring ^ basal segments of labium (Fig. 27.1C); labium reach¬ ing to mesosternum, segments 1 and 2 short, segment 3 longest, segment 4 short (Fig. 27.IB, C); pronotum enlarged, obscuring meso- and metanotum and part of ab¬ dominal mediotergite 1 (Fig. 20.2D) (at least in macrop- terous morphs, variously reduced in apterous morphs); r- coxae of meso- and metathorax widely separated; meso¬ sternum with a mesal longitudinal impression, but with¬ out a distinct rostral groove (Fig. 27.11); metasternum '' with characteristic pair of strongly arched grooves lead¬ ing from metasternal scent orifice to a subovate swelling with a tuft of long hairs on metasternopleuron; forewing with a variable number of closed cells (Fig. 27.1D-F); alary polymorphism common; all legs usually of similar structure, sometimes metathoracic pair longest; femora variously elongated, dilated, or spinose; male foretibia 98 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) usually with grasping comb of short, stout spines on pos¬ terior, innermost margin; mesotibia usually with row of erect trichobothrium-like setae on posterior surface; tarsi 1-, 2-, or 3-segmented; last tarsal segment always cleft over one-fourth to nearly two-thirds of length, pretarsus situated in cleft (Fig. 27. IG, H); parempodia asymmet¬ rically developed; pretarsus of middle legs often highly modified (e.g., claws flattened and asymmetrical and ventral arolium modified into “swimming fan”) (Fig. 27.IH); abdomen generally elongate, sometimes short¬ ened; abdominal mediotergites 1-3 with a pair of medial longitudinal ridges; dorsal abdominal scent-gland appara¬ tus absent in both nymphs and adults; pygophore usually large, protruding from abdomen; parameres usually sym¬ metrically developed (Fig. 27. IJ); aedeagus as in Fig. 27.IK; testes as in Fig. 27. IL; female usually with a" reduced, plate-shaped ovipositor (Fig. 27.11), with a me¬ dian connection between second valvulae as in Gerridae; gynatrial complex as in Fig. 27. IM; 4 ovarioles. Classification. The Veliidae comprise six subfami¬ lies, following the classifications of Stys (1976) and Andersen (1982a), who treated the controversial Macro- veliidae and Ocelloveliinae as a distinct family and sub¬ family, respectively. Hebrovelia Lundblad is treated as belonging to a distinct tribe within the Microveliinae (see Stys, 1976), and the Haloveliinae, which have been vari¬ ously placed by prior authors, are treated as a subfamily within the Veliidae. The family currently includes 38 genera and nearly 600 species. Key to Subfamilies of Veliidae 1 . 2 . 3. 4. 5. Ocelli present; forewings with 6 closed cells (Fig. 27. ID) . Ocelloveliinae Ocelli absent; forewings with fewer than 6 closed cells (Figs. 20.2D, 27. IE, F) . 2 Middle tarsus deeply cleft with a fan of plumose or hairy swimming fans arising from base of cleft . Rhagoveliinae Middle tarsus not so deeply cleft, lacking plumose or hairy swimming fans . 3 Middle tarsus 3-segmented (first segment short) . 4 Middle tarsus 2-segmented . 5 Foretarsus 2-segmented; forewing divided into a proximal coriaceous portion with 2 closed cells and membranous distal portion . Perittopinae All tarsi 3-segmented (basal segments short); fotewings not divided into coriaceous and membranous portions, but with 4 closed cells (Fig. 20.2D) . Veliinae Foretarsus 1-segmented; middle tarsus rarely more than twice as long as hind tarsus . . Microveliinae Foretarsus 2-segmented; middle tarsus 3 or more times length of hind tarsus . Haloveliinae HALOVELIINAE. Middle legs greatly elongated; poste¬ rior pronotal lobe strongly reduced in apterous forms; macropterous Entomovelia Esaki and Stongylovelia Esaki with forewing venation reduced to some short basal thickenings and pair of obsolete longitudinal veins. Included genera are Entomovelia Esaki, Halovelia Ber- groth, Halovelioides Andersen, Strongylovelia Esaki, and Xenobates Esaki. All Halovelia are intertidal, living on relatively protected oceanic areas such as coral reefs; the remaining taxa live in mangroves and estuaries in the Orient, including New Guinea. Certain authors have placed this highly modified group within the Gerridae (e.g., Esaki, 1924, 1930) or in its own family. Halovelia has been studied in detail by Andersen (1989a, c), with 30 species being recognized; Andersen (1991c) offered a revised key to the genera. MICROVELIINAE. Tarsal formula unique, 1-2-2. This is the largest subfamily of Veliidae, with 21 genera placed in 3 tribes—Hebroveliini, Veliohebriini, Micro- veliini (Stys, 1976; Andersen, 1982a). The first two tribes are monotypic, have the pretarsus inserted near the apex of the tarsus (subapical), and are probably largely terres¬ trial. The largest genus of Microveliini, Microvelia West- wood, contains approximately 170 species. These are the classic small veliids that occur around the shores of ponds and springs and in quiet stream-margin habi¬ tats. Drake and Hussey (1955) provided a checklist to the New World species, Poisson (1940) treated some of the species known from Africa, Esaki and Miya¬ moto (1955) dealt with the Japanese species, and J. T. Polhemus (1974) dealt with the M. austrina group from Central America and Mexico. Xiphovelia Lundblad front the Orient (Japanese species treated by Esaki and Miya¬ moto, 1959b), Xiphoveloidea Hoberlandt from Africa, and Euvelia Drake from Bolivia and Brazil inhabit flow¬ ing mountain streams or the margins of rivers (J. T. Polhemus and D. A. Polhemus, 1984) and, along with the Veliidae 99 Fig. 27.1 . Veliidae. A. Head and thorax, Microvelia reticulata (Burmeister). B. Lateral view, body, Entomovelia doveri Esaki. C. Lateral view, head, Ocellovelia german (Distant). D. Right forewing, O. germari. E. Right forewing, Perittopus sp. F. Right forewing, Microvelia pulchella Westwood. G. Left middle tarsus, Perittopus sp. H. Left middle pretarsus, Xiphoveloidea major (Poisson). I. Ventral view, body, O. germari. J. Pygophore and parameres, dorsal view, Neoalardus typicus (Distant). K. Aedeagus, lateral view, Velia caprai Tamanini (A-K, M from Andersen, 1982a). L. Testes, V. caprai (from Pendergrast, 1957). M. Gynatrial complex, Tenagovelia sjoestedti Kirkaldy. Abbreviations: fc, fecundation canal; fp, fecundation pump; gs, gynatrial sac; oa, occipital apodeme; pa, paramere; st, spermathecal tube; va, ventral arolium. 100 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) marine Neotropical Husseyella Herring, have 4-bladed swimming fans. Andersen (1981a) reviewed the genus Pseudovelia Hoberlandt in the Orient, documenting the presence of 11 species, all of which live near the margins of streams or rivers, some on the riffle zone. Four species of Baptista Distant and two of Lathrovelia Andersen live in secluded aquatic environments and water-filled cavities (Andersen, 1989b). Veliohebria Stys from New Guinea is apparently largely terrestrial in habits (Stys, 1976). Tonkuivelia Linnavuori (1977) is unusual in the Micro- veliini, having a subapical insertion of the pretarsus, and is also probably terrestrial. OCELLOVELIINAE. Ocelli present; forewing with 6 cells (Fig. 27. ID); tarsi 3-segmented; pretarsus inserted pre- apically. Ocellovelia China and Usinger (1949a), with two spe¬ cies from South Africa, is a group that at various times has been placed in the Mesoveliidae, Macroveliidae, and an omnibus Veliidae. PERiTTOPiNAE. Foretarsus 2-segmented; venation of forewing reduced, with 2 closed cells (Fig. 27. IE). Contains only Perittopus Fieber with about five species from the Orient. RHAGOVELiiNAE. Pretarsus of middle leg with swim¬ ming fan formed from dorsal branch of ventral arolium inserted at base of deep cleft in distal segment of tarsus. Included genera are the widely distributed Rhagovelia Mayr, with many species, commonly referred to as riffle bugs; Tetraripis Lundblad; and the marine Trochopus Carpenter from the Caribbean (Lundblad, 1936; Bacon, 1956; Matsuda, 1956; Hungerford and Matsuda, 1961; J. T. Polhemus and D. A. Polhemus, 1989a; J. T. Polhe- mus, 1990a). VELiiNAE. Tarsi 3-segmented on all legs; pretarsus simple in most genera; thorax unmodified in apterous morphs; parameres large and symmetrical; female with foretibial grasping comb (Andersen, 1982a). Included genera are Angilia Stal, Angilovelia Ander¬ sen, Oiovelia Drake and Maldonado, Paravelia Breddin, Strididivelia Hungerford, Velia Latreille (see Tamanini, 1947, 1955, for European taxa), and Veloidea Gould. Velia spp. and Paravelia spp. are often found on slow- moving streams and are some of the largest members of this family of generally rather small bugs. Specialized morphology. The “pit organs” described in Halovelia by Cobben (1978) resemble the punctures common in most Gerromorpha (Fig. 10.7D) (except Ger- ridae and Hematobatidae). Also the Veliidae possess what Andersen (1977c) referred to as “thornlike outgrowths” (the “grooved setae” of Cobben [1978]), straight or slightly curved structures thickened at the base and taper¬ ing apically and projecting above the microhair layer. The Veliidae, in common with the Gerridae, and as opposed to all other Gerromorpha, do not have an iso- radial salivary pump, but rather the pump is inflected from behind on both sides; rather than being entirely membranous, the dorsal surface is sclerotized (Andersen. 1982a). The posterior margin of the head capsule is provided with a pair of slender occipital apodemes (Fig. 27.1C), which serve as attachments for the maxillary retractor muscles inserting on the prolonged maxillary stylets. The veliine genera Angilovelia Andersen and Stridu- livelia possess stridulatory structures incorporating the metafemur and the connexival margin (Andersen. 1981a) (see also Chapter 10). Many veliids possess striking modifications of the pre- tarsus, and indeed the family contains more variation in these structures than all the other Gerromorpha com¬ bined. The claws and dorsal and ventral arolia are often modified, as for example in Xiphoveloidea (Fig. 27. IH). where they are in the form of four leaflike structures, whereas in the Rhagoveliinae a branching process arising dorsally from the ventral arolium forms a swimming fan. In contrast to all other members of the family, males of some Microvelia spp. have asymmetrically developed parameres, the left paramere being greatly reduced. The gynatrial complex has a glandular gynatrial sac. and (except in Ocellovelia) the basal thickening of the fe¬ cundation canal is modified into a “fecundation groove.” or as in the Gerridae, a “fecundation pump” may be present on the fecundation canal (Fig. 27. IM). Natural history. Most Veliidae live on or near stand¬ ing water, some are marine, and Hebrovelia, Tonkuivelia, and Veliohebria are apparently semiterrestrial. The group does not show as strong a relationship with the open water surface as do the Gerridae. At least four species each of Paravelia and Microvelia live in the tanks of Bromeliaceae or occasionally in tree holes in the New World tropics (J. T. Polhemus and D. A. Polhemus, 1991a). A few species of veliids have been recorded from foam masses in tropical streams. Among these \% Oio¬ velia spumicola Spangler (1986) (Veliinae) from southern Venezuela, which lives secreted inside the foam masses; it is a member of a genus whose other two described species are recorded as free-living. Afrovelia phoretica J. T. Polhemus and D. A. Polhemus (1988) (Microveli- inae) was recorded as occurring on the surface of foam masses caught among debris in rivers in western Mada¬ gascar. The locomotory mechanisms of most genera are like those in the foregoing families of Gerromorpha, whereby the legs are moved alternately and no single pair predomi¬ nates. Some groups, such as Velia live on slow-moving water, have the middle legs slightly elongate, and move them in a rowing motion—in contrast to the alternating Veliidae 101 triangle gait practiced by most insects—but have retained the ability to walk on land. Many species can move ex¬ tremely rapidly on the water surface. Stridulivelia spp. form fast-swirling swarms. Rhagovelia spp., which often form huge schools, represent the extreme of modifica¬ tion within the family, possessing adaptations for life on extremely fast-flowing and disturbed water. In the last group, the middle legs are the primary locomotory ap¬ pendages, the pretarsus is modified into a swimming fan, and the contralateral legs are moved simultaneously in a rowing motion; the ability to walk on land has been lost (Andersen, 1976). Many veliids are known to practice ‘‘expansion skat¬ ing.” This phenomenon is produced by the ejection of a droplet of fluid from the rostrum; the surface tension of the water is momentarily lowered, causing the bug to move away much more rapidly than is possible simply by the use of the legs (Linsenmair and Jander, 1963; Andersen, 1976). Veliids and other tiny aquatic bugs are capable of grasp¬ ing the water surface for ascension of the meniscus, an obstacle often of greater size than the bug itself. This phenomenon was apparently observed independently by Baudoin (1955) and Miyamoto (1955). Sexual dimorphism is present in some veliids, usually with males being smaller than females. In the most ex¬ treme cases the female may be nearly twice the size of the male. Males are sometimes phoretic and may spend long periods riding on the backs of the females. This behav¬ ior was described in detail by Miyamoto (1953; see also Andersen, 1982a:310) for Microvelia diluta Distant, and has also been noted in Afrovelia phoretica (J. T. Polhe- mus and D. A. Polhemus, 1988), and in Halovelia spp. (Kellen, 1959). J. T. Polhemus (1974) described modifi¬ cations of the thorax in Microvelia to accommodate the male, whose ability to grasp his mate is apparently facili¬ tated by combs located on the tibiae of the fore- (and sometimes also middle) legs. Many species of Veliidae self-mutilate their wings. The eggs of the Veliidae are glued lengthwise to the substrate, attached by their less convex side, often to marginal vegetation. Distribution and faunistics. Identification aids for the western Palearctic include those by Stichel (1955; Velia prepared by Tamanini; see also Tamanini, 1947) and Kerzhner and Jaczewski (1964). The eastern Palearc¬ tic has been treated by Kanyukova (1988). Smith and Polhemus (1978) keyed the North American species, and J. T. Polhemus and Chapman (1979d) keyed the North American genera and California species. The species of the Great Lakes region were keyed by Bennett and Cook (1981). Cobben (1960b) treated the fauna of the Netherlands Antilles. New World Rhagovelia were mono¬ graphed by Bacon (1956). Useful treatments for Asia include Esaki and Miya¬ moto (1955) on Microvelia and Pseudovelia from Japan and (1959b) on Xiphovelia from Japan; Lundblad (1936) on Old World Rhagovelia and Tetraripis: J. T. Polhemus and D. A. Polhemus (1989a) and J. T. Polhemus (1990a) on Rhagovelia from the Malay Archipelago and South¬ east Asia; and Andersen (1989a, c, 1991c) on Halovelia and Halovelioides. The African Microvelia fauna has been treated by Poisson (1940) and Linnavuori (1977), and Poisson (1957a) dealt with some taxa in the South African fauna. 28 Gerridae General. The Gerridae, known as water striders, pond skaters, or wherrymen, spend—with few exceptions— nearly their entire lives on the open water surface. They range in length from some diminutive Rheumatobates spp. (Fig. 28.1 A) of 1.6 mm to the 36-nnm long Giganto- metra. They are always long-legged, and their bodies vary from nearly globular (Fig. 20.3C) to elongate and cylindrical (Fig. 20.3B). Diagnosis. Entire body and appendages covered with micro- and macrohair layer; peg plates absent; head usually somewhat extended beyond anterior margin of compound eyes, antennal fossae often visible from above (Fig 28.1B); ocelli absent; 4 pairs of cephalic trichoboth- ria (Fig. 28.1C) (rather than 3 as in other Gerromorpha), all inserted in pits in the Rhagadotarsinae and Eurygerris (Gerrinae), other groups with fourth pair projeeting from body surface as in nymphs; antennal segment 1 usually longer and somewhat stouter than remaining 3 segments (Fig. 20.3C); labium usually surpassing prostemum, seg¬ ment 1 longer than segment 2, segment 3 longest, seg¬ ment 4 short (Fig. 28. IB); pronotum without collar, never punctured, extending posteromesally to obscure meso- notum in macropterous forms (Fig. 20.3B); mesothorax elongated relative to the condition found in other Ger¬ romorpha, particularly so in larger taxa, ratio of meso- to metathoracic lengths ranging from 1.2:1 to 10:1 (Fig. 28. IF); lateral channels of metathoracic scent-gland sys¬ tem, when present, similar in structure to those of Veli- 102 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) idae; forelegs relatively short and stout, middle and hind legs slender and greatly elongated, femora and tibiae gen¬ erally subequal in length (Fig. 20.3B, C); forecoxa small and inserted near midline of thorax; middle and hind coxae rotated and extended caudad (Fig. 28. IF), middle coxa large and usually greatly extended along latero- ventral margin of thorax, hind coxa somewhat smaller than midcoxa and not similarly prolonged; all tarsi 2- segmented, segments of varying lengths, with large cam- paniform sensillum on segment 1, suggesting fusion of primitive segments 1 and 2; pretarsus inserted anteapi- cally, usually laterally; dorsal arolium usually bristlelike and short, ventral arolium usually well developed; parem- podia setiform; claws of middle and hind legs usually smaller than those of forelegs; forewing venation vari¬ able, never with more than 4 cells, 2 apical cells each terminating in a relatively long free veini (Figs. 20.3C, 28. ID, E); hind wing lacking a distinct first anal vein; alary polymorphism common, macropterous and apter¬ ous morphs predominating; abdominal leiagth variable; pygophore usually protruding from apex of abdomen (Fig. 20.3C), sometimes obviously asymmetrical, includ¬ ing segment 8; parameres usually present and symmet¬ rically developed; aedeagus as in Fig. 28. IG; ovipositor generally relatively short with plate-shaf©d valvulae; 4 ovarioles; gynatrial complex as in Fig. 28. iH. Classification. Following the classification of Ander¬ sen (1982a), we recognize eight subfamilies containing 60 genera and approximately 500 species. Key to Subfamilies of Gerridae 1. First abdominal sternum present and distinct; ovipositor serrate, elongate; bucculae produced (Fig. 28.1C) . Rhagadotarsinae - First abdominal sternum absent, or fused with metasternum; ovipositor not serrate, short; buecialae not produced (Fig. 28. IB) . 2 2. Forewing differentiated into a coriaceous basal portion and membranous distal portion (Fig. 28. fE); middle femur stout, shorter than mesotibia and usually shorter than hind femur; metathoracic scent- gland orifices usually absent . Trepobalinae - Forewing not differentiated into a thickened basal and membranous distal portion (Figs. 20.3B, 28. ID); middle femur slender, usually distinctly longer than middle tibia; metathoracic scent-gland orifices present . 3 3. Metastemum greatly reduced, represented by a very short subtriangular plate enclosing the scent- gland orifice; if metasternum reaching metacetabular region, then middle tibia with distinct fringe of setae; claws of hind tarsus straight or S-shaped . Halobaiinae - Metastemum well developed, reaching metacetabular region laterally; claws of hind tarsus falcate orabsent . 4 4. Metacetabular groove distinct, dorsally reaching anterior end of abdominal tergum 1; foretarsus very long, at least one-half length of foretibia . Ptilomerinae - Metacetabular groove often indistinct dorsally, if distinct, not reaching anterior end of abdominal tergum 1; foretarsus less than one-half length offoretibia . 5 5. Metacetabular groove distinct, dorsally connected with posterior margin of mesopostnotum; anten¬ nal segment 4 curved; labium not extending posteriorly beyond prosternal margin; first segment of foretarsus less than one-half length of second segment . Cylindrostethinae - Metacetabular groove indistinct dorsally; antennal segment 4 straight; labium distinctly surpassing prosternal margin; first segment of foretarsus usually more than one-half length of second segment . 6 6. Metastemum with lateral scent channels leading from scent-gland orifices to metacetabular region; secondary straight transverse line present across metanotum; no connexival spines present . . Charmatometrinae - Metastemum usually lacking lateral scent channels! if present then insects 30 mm in length; connexival spines present; metanotum without a transverse line . 7 7. Pronotal lobe reduced in apterous individuals; arolia present; lateral suture between meso- and metathorax distinct . Eotrechinae - Pronotal lobe usually not reduced in apterous individuals, but if reduced then arolia absent or lateral suture between meso- and metathorax indistinct . Gerrinae Gerridae 103 Fig. 28.1. Gerridae. A. Rheumatobates meinerti Schroeder, male (from Hungerford, 1954). B. Lateral view, head and pronotum, Onychotrechus rhexenor Kirkaldy. C. Lateral view, head, Rhagadotarsus kraepelini Breddin. D. Right forewing, Aquarius paludum (Fabricius). E. Right forewing, Trepobates taylori (Kirkaldy). F. Ventral view, body, apterous male, Eurymetra natalensis (Distant). G. Aedeagus, inflated, lateral view, Neogerris parvulus (Stal). H. Gynatrial complex, dorsal view, Cylindrostethus palmaris (Drake and Harris) (B-H from Andersen, 1982a). Abbreviations; fc, fecundation canal; fp, fecundation pump. CHARMATOMETRINAE. Lateral scent-gland channels little studied. Available observations suggest its members present (found elsewhere only in Cylindrostethinae and live on quiet areas of streams. Gigantometra); distinct sutures between the meso- and cylindrostethinae. As in Ptilomerinae and Halobati- metathorax; secondary transverse line in front of first nae, antennal segment 2 with trichobothrium-like setae; abdominal tergum. labium short, not surpassing posterior margin of proster- Included genera are Brachymetra Mayr (Shaw, 1933), num; uniquely with distinctly flattened middle tibia and Charmatometra Kirkaldy, and Eobates Drake and Harris. tarsus; heavily sclerotized second valvulae. The group is restricted to the Neotropics and has been Included genera are Cylindrostethus Mayr, Platygerris 104 TRUE BUGS OF THE WORLD (HEMiPTERA: HETEROPTERA) Buchanan-White, and Potamobates Champion. They are found in the Neotropics, with Cylindrostethus also occur¬ ring in the Old World (Hungerford and Matsuda, 1962). All appear to be stream dwellers, the last two genera occupying broad flat areas of flowing streams. EOTRECHINAE. Shares with Gerrinae sclerotized and setose inner lobes of first valvulae and 2 distinctly glan¬ dular areas in wall of gynatrial sac; unique in possessing spinose setae on tibiae and tarsi of middle and hind legs; pretarsus of Eotrechus inserted apically. Included genera are Amemboa Esaki, Chimarrhotnetra Bianchi, Eotrechus Kirkaldy, Onychotrechus Kirkaldy (Andersen, 1980), and Tarsotrechus Andersen (Ander¬ sen, 1980); all occur in India and the Orient and are generally hygropetric or semiterrestrial. A generic phy- logeny was published by J. T. Polhemus and Andersen (1984). GERRINAE (FIG. 20.3B). In addition to characters shared with the Eotrechinae mentioned above, also recognized by the following: wing venation relatively complete; dor¬ sal and ventral arolia usually strongly reduced; parem- podia both rather short; metathoracic scent-gland channel usually absent; abdomen usually elongate; vesica with sclerotized dorsal plate. Two tribes are currently recognized. The Tachyger- rini comprise the New World genera Tachygerris Drake and Eurygerris Hungerford and Matsuda. The remain¬ ing 10 genera are placed in the cosmopolitan Gerrini and include large commonly encountered members of the genera Aquarius Schellenberg (Andersen, 1990), Ger- ris Fabricius, Limnogonus Stal (Andersen, 1975), and Limnoporus Stal (Andersen and Spence, 1992); all are frequently associated with quiet waters, such as ponds. Tenagogonus Stal from the Old World tropics (Hunger¬ ford and Matsuda, 1958b), by contrast, is usually found in quiet places on streams. The Western Hemisphere fauna was treated by Drake and Harris (1934). HALOBATiNAE. Head and thorax short and wide; meta- stemum reduced; abdominal segments shortened; tho¬ racic and abdominal segments strongly fused. Members of the 10 halobatine genera, placed in two tribes, represent some of the most highly modified Ger- ridae. The tropicopolitan Halobatini comprise Asclepios Distant and Halobates Eschscholtz (Herring, 1961; An¬ dersen, 1991b; J. T. Polhemus and D. A. Polhemus, 1991b; Andersen and Foster, 1992), most species of which are marine. The remaining genera—placed in the Paleotropical Metrocorini and comprising, for example, the Oriental Esakia Lundblad (Hungerford and Matsuda, 1958a; Cheng, 1966), Metrocoris Mayr (Boer, 1965; D. A. Polhemus, 1990b; Chen and Nieser, 1993), and Ventidius —are found on slow-flowing waters, usually in heavily shaded areas (Hungerford and Matsuda, 1960a; Cheng, 1965). An important historical paper on the group is that of Esaki (1926). Metrocoris has been revised for the Malay Archipelago and the Philippine Islands (D. A. Polhemus, 1990b), with an additional large and mostly undescribed fauna known from tropical continental Asia. PTILOMERINAE. Antennal segment 4 with an elongate trough ventrally; first segment of foretarsus longer than second. This subfamily comprises eight genera distributed from Madagascar to New Guinea. Ptilomera Amyot and Ser- ville (Hungerford and Matsuda, 1965). is a group of often large insects, all members of which inhabit swiftly flow¬ ing waters, often near waterfalls. Potamometra Bianchi was revised by Drake and Hoberlandt (1965). RHAGAD0TARSINAE(FIG.28.1A). Bucculae well developed (Fig. 28.1C), in contrast to other Gerridae; metathoracic scent apparatus absent (also in some other Gerridae); Rhagadotarsus with pretarsus inserted ventrally and with thornlike cuticular outgrowths found in the Veliidae; laterotergites and true abdominal sternum 1 distinct; ovi¬ positor long, laciniate, and serrate. Rhagadotarsus Breddin, with five species from the Old World tropics (J. T. Polhemus and Karunaratne, 1993), is ordinarily found in large schools on the surface of Standing water, as for example in fish ponds and on quiet reaches in streams. Rheumatobates '&ttgxoth, comprising more than 30 species, is widely distributed in the New World (Hungerford, 1954; J. T. Polhemus and Spangler, 1989), occupying a variety of habitats, including muddy pools, quiet portions of large rivers, and even estuaries; they often assemble in large swarms. Substantial infor¬ mation on the group was provided by Esaki (1926). TREPOBATiNAE (Fig. 20.3C). Middle femur relatively short and stout, always shorter than middle tibia; forewing with 2, closed, basal cells, and transverse line of weakness (Fig. 28. IE). This subfamily comprises nineteen genera. A world re¬ view was provided by J. T. and D. A. Polhemus (1993). The New World genus Metrobates Uhler usually lives on streams or rivers. It is distinctive in the subfamily for its flattened body, modified second and third antennal segments, relatively long middle tibia, and conspicuous claws on the middle and hind legs (Anderson, 1932; Drake and Harris, 1932a). Trepobates Uhler (Drake and Harris, 1932b), also from the New World, ordinarily inhabits quiet or slow-moving water. Telmatometra Ber- groth from the New World tropics was revised by Kenaga (1941). Specialized morphology. The Gerridae have several novel morphological features in common with the Veli¬ idae, including posterior margin of head capsule with pair of long occipital apodemes for attachment of maxillary retractor muscles (Fig. 28.1C); salivary pump laterally inflected, membranous in forms studied, but sclerotized Gerridae 105 dorsally in Eurymetra (Halobatinae); and gynatrial com¬ plex usually with a fecundation pump and glandular gyna¬ trial sac (Fig. 28. IH). The single most distinctive feature of the Gerridae. one unique in the Heteroptera, is the modification of the thorax in association with life on the water surface film, wherein the mesothorax is always elongated, sometimes greatly so, and the middle and hind coxae are oriented in a horizontal plane (Fig. 28. IF). Details of complex tho¬ racic fusions in apterous morphs and allometric growth of appendages were reviewed in detail by Matsuda (1960). The antennae of some male Rheumatobates spp. are irregularly thickened and bent and covered with tufts of setae (Fig. 28. lA). They are used to grasp the female during copulation, a function performed by the foretibiae in all other members of the family (Hungerford, 1954). The hind legs of Rheumatobates are also highly modi¬ fied (Fig. 28.1 A) compared with all other Gerridae (Fig. 20.3B-D). Males of the genus Ptilomera have a brush of long setae on the posterior surface of the middle tibia. Asymmetry of male terminal abdominal segments oc¬ curs in most Halobates spp. and in some Cylindrostethi- nae. Although the parameres are always symmetrically developed, they are very small in the Gerrinae, Eotrechi- nae, and Cylindrostethinae and are absent in the Rha- gadotarsinae and Halobates. Natural history. Gerrids commonly occupy bodies of standing water, ranging from relatively small pools to lakes, and indeed they are most easily observed in such quiet situations. Nonetheless, many taxa live on running waters, some of them capable of gliding easily on the rapids or torrents of mountain streams, in which situa¬ tions they are much less easily observed. The remaining species inhabit marine environments, the majority found in relatively protected areas such as estuaries, mangroves, and lagoons, with a few species of Halobates completing their entire life cycle on the open ocean. “Ripple communication” is well known in the Gerridae and has been studied in some detail; it is not well known to what degree it plays a role in the lives of other ger- romorphans, although it has been recorded in the veliid Microvelia longipes (J. T. Polhemus, 1990b). Gerrids use waves on the water surface for prey location and for com¬ municating with each other. Prey on the water surface are located by orienting to the waves they make (Murphey, 1971). Gerrids themselves also produce surface waves of a species-specific frequency by vertical movements of one or more pairs of legs. “Calling signals” are produced by males, and both sexes may produce additional “courtship signals” (Wilcox, 1972, 1980). Such signals allow for sex discrimination within a species (Wilcox, 1979) and play a role in territory delimitation and defense as well as species recognition as a defense against cannibalism (Wilcox, 1980; Wilcox and Spence, 1986). The reception of such surface waves is thought to occur either in the stretch receptors located in the tibiotarsal joint (Murphey, 1971) or in specialized setae located on the tarsi (see Lawry, 1973). Andersen (1982a) suggested that vision is also important in prey location. Most members of the Eotrechinae, including species of Chimarrhometra, Eotrechus, and Onychotrechus, have moved from life on the water surface film to being virtu¬ ally terrestrial or hygropetric, living in situations typical of those often occupied by the Hebridae. They are capable of extremely rapid and agile movements in such habitats, even on almost vertical surfaces. Andersen (1982c) de¬ scribed the novel orientation of the middle and hind legs in Eotrechus, the tibiae being positioned nearly perpen¬ dicular to the substrate, with the weight of the animal resting on the relatively short tarsi. The evidence sug¬ gests that the apically inserted claws of the Eotrechinae represent a secondarily derived condition. Eggs are fixed to floating objects in a lengthwise posi¬ tion, with the convex side uppermost. Rhagadotarsus is known to embed its eggs in plant tissue, and the structure of the ovipositor suggests that Rheumatobates does the same (Andersen, 1982a). Description of wing polymorphism and methods of its determination have been intensively studied in the Ger¬ ridae. They are discussed in Chapter 6. Whereas most gerrids are either macropterous or apterous, intermediate wing morphs are known in Gerris. Distribution and faunistics. Keys to the higher groups and genera and illustrations of representatives of most genera can be found in Hungerford and Matsuda (1960b) and in Andersen (1982a); Matsuda (1960) pro¬ vided diagnoses of all genera. Faunistic treatments are available for Europe (Stichel, 1955; Wagner and Zimmer- mann, 1955), the former Soviet Union (Kanyukova, 1982), California (J. T. Polhemus and Chapman, 1979e), Argentina (Bachmann, 1966), the Malay Peninsula and Singapore (Cheng and Fernando, 1969; see also Miya¬ moto, 1967), Japan (Miyamoto, 1961b), and Africa and Madagascar (Poisson, 1965), with many more-restricted surveys also present in the literature. 106 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) PANHETEROPTERA 29 Nepomorpha General. The water bugs were first recognized as a group by Latreille (1810) under the name Hydrocorisae. All except some Corixidae are predators. The short an¬ tennae, which are concealed—often completely—below the eyes, are common to all families and have traditionally been treated as an adaptation to the aquatic mode of life. The Ochteridae and Gelastocoridae are usually riparian, however, and they do not possess the streamlined bodies and swimming legs found in the remaining nine fami¬ lies. The Nepomorpha range from very small, as in some species of Helotrephidae and Pleidae, to the largest of all bugs, the Neotropical belostomatid Lethocerus maximus DeCarlo, with a length of 110 mm. The painful nature of the bite of the aquatic bugs is the subject of folklore— and reality. Diagnosis. Compound eyes usually very large and occupying nearly entire sides of head in dorsal and lateral views (Fig. 33.1); ocelli absent (except Ochteroidea, Co¬ rixidae: Diaprepocorinae); cephalic trichobothria always absent; labium usually short and very stout (Fig. 32.2A) (except Ochteridae: Fig. 33.2A; Aphelocheiridae; Fig. 37.1C), often with 3 apparent segments (except Corixi¬ dae), segment 1 greatly reduced; antennae at most as long as head, often thickened (Fig. 29.1 A, D), provided with processes (Fig. 31.2B), or with segments fused, situated posteroventrally on head, often hidden in groove or concavity, musculature reduced; middle and hind legs often flattened (Fig. 30.1) and fringed with setae; fore¬ legs often raptorial with tibia apposed to femur; pre¬ tarsus, particularly of foreleg (Fig. 10.4A), sometimes with one claw reduced or absent; parempodia setiform. Keys to Nepomorpha prepared by Pavel Stys. number variable from 2 to 4 (Fig. 29.2B, E, G); dor-' sal and ventral arolia usually present in both nymphs and adults (Fig. 29.2A-1); forewings with well-developed wing-to-body coupling mechanism and frena along scu- tellum and claval commissure, in the form of hemely- tra with coriaceous anterior portion formed of distinct corium and clavus and posterior membranous portion (Fig. 33.1), with none to many veins; scolopophorous organs usually present in meso- and metathorax as well as on abdominal tergum 1 (Larsen, 1957; Parsons, 1962; Mahner, 1993); spiracle present dorsally on abdominal tergum 1, spiracles on segments 2-8 usually ventral; ab¬ domen of fully aquatic forms with various modifications for respiration under water; male genitalia symmetrical in Nepoidea, asymmetrical in most other groups except some Naucoridae (Cheirochelini), Potamocoridae, and many Notonectidae, asymmetry often including some pregenital abdominal segments; ovipositor often weakly developed; spermatheca tubular, without a terminal bulb; number of micropyles variable. Discussion. We recognize 11 families of Nepomor¬ pha, following the classification of Stys and Jansson (1988). China (1955c) proposed a scheme of relation¬ ships among them, treating the Ochteridae as the rela¬ tively most primitive, on the basis of its possession of ocelli and the respiratory system typical of terres¬ trial bugs. More recent authors, including Popov (1971), Rieger (1976), and Mahner (1993), have treated the Nepoidea as the sister group of remaining Nepomorpha. Popov (1971) documented that the fossil record for many families dates back to the mid-Mesozoic, and he pro¬ vided a detailed discussion of the morphology of fossil and recent taxa. The character analyses of Rieger (1976) and Mahner (1993) show a substantial increase in breadth of coverage and methodological sophistication over pre¬ vious efforts. Their conclusions contradict one another mainly in the placement of the Corixidae and relation¬ ships among those families we place in the Naucoroidea, differences that can be appreciated by comparing Figures 29.3 and 29.4 (Figure 29.3 incorporates some modifica¬ tions of family-group status from the work of Stys and Jansson [1988]). The taxonomy of parts or all of many nepomorphan families has been the subject of detailed publications, par¬ ticularly by H. B. Hungerford and his many students. The attention that has been paid to the group probably results in large part because the habitat of most aquatic bugs is obvious, they often have long adult life spans, and in temperate areas they can be collected even in winter. Excellent morphological studies have been published on several groups, and many families are also the subject of detailed life history investigations. The western Palearctic fauna was treated by Poisson Nepomorpha 107 Fig. 29.1. Morphology of Nepomorpha. A. Ventral view, head and antenna, Gelastocoris oculatus (Gelastocoridae). B. Ventral view, male genital segments, G. oculatus (Gelastocoridae). C. Base of adult labium, Hydrotrephes sp. (Helotrephidae). D. Ventral view, head and antenna, Ochterus caffer (Stal) (Ochteridae). E. Ventral view, male genital segments, O. caffer (Ochteridae). F. Ventral abdominal hair pile, including spiracle, adult Hydrotrephes sp. (Helotrephidae). G. Abdominal venter, adult male Cryphocricos latus (Naucoridae), H. Detail, spiracle and modified setae, C. latus (Naucoridae). I. Abdominal static sense organ, Ranatra sp. (Nepidae). Abbreviations: lb, labium; so, static sense organ; sp 6, spiracle 6; sp 7, spiracle 7; sp 8, spiracle 8; spc, spiracle cover. 108 TRUE BUGS OF THE WORLD (HEMIPTERA- HETEROPTERA) Fig. 29.2. Pretarsus of Nepomorpha. A. Foreleg pretarsus, nymph, Gelastocoris oculatus (Gelastocoridae). B. Foreleg pretarsus, adult, G. oculatus (Gelastocoridae). C. Foreleg pretarsus, nymph, Ochterus sp. (Ochteridae). D. Nymphal pretarsus, Laccocoris hoogstraali La Rivers (Naucoridae). E. Hind leg pretarsus, adult, Cryphocricos latus (Naucoridae). F. Middle leg pretarsus, adult, Aphelocheirus lahu Polhemus and Polhemus (Aphelocheiridae). G. Prefarsus, nymph, Hydrotrephes sp. (Helotrephidae). H. Hind leg pretarsus, nymph, Diplonychus sp. (Belostom- atidae). I. Hind leg pretarsus, nymph, Laccotrephes sp. (Nepidae). Abbreviations: cl, claw; da, dorsal arolium; pe, parempodium; ut, unguitractor; va, ventral arolium. Nepomorpha 109 Fig. 29.3. Phylogenetic relationships of families of Nepomorpha Fig. 29.4. Phylogenetic relationships of families of Nepomorpha (modified from Rieger, 1976). (modified from Mahner, 1993). (1957b), Kerzhner and Jaczewski (1964), and Savage (1989). Lundblad (1933) dealt with the faunas of Java, Sumatra, and Bali. Poisson (many papers) has dealt with much of the fauna of Africa. North America was first treated in detail at the generic level by Usinger (1956) and again by Menke (1979a). Nieser (1975) treated the fauna of Surinam. Key to Families of Nepomorpha 1. Labium broadly triangular, short, nonsegmented, with transverse sulcation (except Cymatiainae); foretarsus unsegmented (sometimes fused with tibia), spoon-, scoop-, or sickle-shaped (Fig. 34.2A), rarely cylindrical, with ventral fringe of long setae; head overlapping pronotum; pronotum and fore¬ wings often with linear or hieroglyphical transverse light and dark patterns (Fig. 34.1) . . Corixidae - Labium cylindrical to conical, obviously segmented (e.g.. Figs. 31.2A, 32.2A). without transverse sulcation; foretarsus segmented or not. without a fringe of setae; head never overlapping pronotum anterodorsally; coloration never as above . 2 2. Apex of abdomen with paired respiratory processes (Figs. 30.1, 30.2. 31.1); body cylindrical or ovoid and flat; medium-sized to gigantic; forelegs raptorial {excepl Liinnogeion: Fig. 30.2) .... 3 - Apex of abdomen without respiratory processes; body never eylindricul. dorsum flat to extremely convex; very small to medium-sized; forelegs raptorial or cursorial, often modified in males .... 4 3. Respiratory siphon nonretractile, usually long and filiform (Fig. 31.1), sometimes short and straplike; all tarsi 1-segmented; hind tibiae simple (Fig. 31.1); metacoxae short, free . Nepidae - Respiratory air straps short, retractile, often only apices visible (Fig. 30.11; tarsi 2- or 3-segmented, rarely foretarsus 1-segmented; hind tibiae usually flat (Fig. 30.1), with swimming setae; metacoxae conical, firmly united with metapleuron . . Belostomatidae 4. Ocelli present (Fig. 33.1), if obsolete or absent, then head transverse and eyes pedunculate or subpedunculate (submacropterous Gelastocoridae); legs cursorial . 5 - Ocelli absent; eyes sessile; hind and/or middle legs usually flattened, fringed with setae (except minute Pleidae) . 6 5. - Antennae relatively long, filiform, partially visible in dorsal view (Fig. 33.1); head moderately transverse; eyes sessile; scutellum flat; legs homomorphous, cursorial (Fig. 33.1) .... Ochteridae - Antennae short and incrassate, concealed in cavity between eyes and prothorax (Figs. 29.1 A, 32.2A); head strongly transverse; eyes subpedunculate to pedunculate (Fig. 32.1); scutellum tumid; fore femur incrassate, anterior face sulcate to carinate . Gelastocoridae 6. Dorsum flat to moderately convex (Figs. 35.1 A, 36.1, 37.1 A); head and p;' lorax never fused; forelegs strikingly raptorial or antennae relatively long, protruding beyond outf of head . 7 110 TRUE BUGS OF THE WORLD (HEMIPTERA; HETEROPTERA) - Dorsum usually strongly convex, inversely boat-shaped to tectiform (Figs. 38.1, 39.1, 40.1 A), if flattened, then head and pronotum fused and cephalonotal sulcus incomplete; forelegs never raptorial, although sometimes modified in males . 9 7. Antennae long, extending beyond lateral margins of head (Fig. 35.IB); labium slender, reaching from prostemum to mesosternum (Fig. 37.1C); head narrow, elongate, strongly produced in front of eyes; foretarsus 3-or 1-segmented, mobile; 2 well-developed claws . 8 - Antennae short, not extending beyond head, usually not visible from above (except many Sagocorini and Tanycricini); labium short and thick, not surpassing prosternum (Fig. 36.2A); head usually transverse, anteocular portion only slightly produced in front of eyes (Fig. 36.1); foretarsus 2- or 1-segmented. immobile; 2, 1, or 0 small claws . Naucoridae 8. Labium just reaching on prosternum, segment 3 shorter than or subequal to segment 4 (Fig. 35. IB); tarsal formula 1-2-2; macropterous to coleopteroid; corium-membrane boundary in macropter- ous forms sinuate; lateral margins of head, pronotum, and corium proximally with erect setae; Neotropical . Potamocoridae - Labium extending at least one-half length of mesosternum. segment 3 at least twice as long as segment 4 (Fig. 37.1C); tarsal formula 3-3-3, tarsomeres sometimes fused in small species; macrop¬ terous to micropterous; corium-membrane boundary straight in macropterous forms; outer posterior angle of corium rounded; lateral margins of head, pronotum. and forewings without erect setae; Eastern Hemisphere . Aphelocheiridae 9. Elongate, wedge-shaped, usually over 4.0 mm long; eyes large (Figs. 38.1. 39.1 A, B); vertex nar¬ row; legs often markedly heteronomous; hind legs long, oar-shaped, with reduced and inconspicuous claws (Fig. 38.1); head not fused with prothorax; midline of pterothoracic venter ecarinate; midline of abdominal venter with a flattened to sharp keel, never with an irregular carina .... Notonectidae - Broadly oval, robustly built species (Figs. 39.1, 40.1 A) under 4 mm long; eyes small to medium¬ sized; vertex broad; legs homonomous; hind legs not oar-shaped, usually with 2 claws; head firmly and immovably associated with prothorax; basal abdominal segments with an irregularly shaped carina . 10 10. Dorsum of head not fused with pronotum; head-pronotum boundary straight (Fig. 39.1); antennae 3-segmented; dorsum strongly convex . Pleidae - Dorsum of head fused with pronotum; cephalonotal sulcus not straight (Fig. 40.1 A); antennae 2- segmented or nonsegmented (Fig. 40.1C); dorsum rather flat to extremely convex . . Helotrephidae Diagnosis. Antennae short, concealed in grooves on underside of head, usually 4-segmented, segments 2 and 3 with lateral projections (Fig. 30.3A); membrane of forewing with reticulate venation; forefemur usually en¬ larged with tibia apposed, foretarsus 2- or 3-segmented. 1 claw slightly to greatly reduced; middle and hind legs usually flattened (Fig. 30.1) and fringed with setae; ab¬ dominal tergum 8 modified into a pair of “airstraps”; nymphs with metepisterna extending posteriorly to cover 2 or 3 proximal abdominal sterna; dorsal abdominal scent glands nonfunctional in nymphs; static sense organs as¬ sociated with spiracles 2-7; male genitalia symmetrical, aedeagus as in Fig. 30.3B-D; testes as in Fig. 30.3E; spermatheca as in Fig. 30.3F, G. Classification. This group was first recognized as a higher taxon by Leach (1815) as Belostomida. The clas¬ sification of Lauck and Menke (1961) recognized three subfamilies comprising nine genera and 146 species. Nepoidea 30 Belostomatidae General. Often referred to as giant water bugs, this group, whose species range in length from 9 to 110 mm, includes the largest true bugs. All species are flattened dorsoventrally, ovoid to elongate ovoid (Figs. 30.1,30.2), and brown in color. Only Limnogeton Mayr, the African snail predator (Fig. 30.2), lacks the greatly enlarged rap¬ torial forefemora and middle and hind legs adapted for swimming. Belostomatidae 111 Fig. 30.1. Belostomatidae. Abedus indentatus (Haldeman) (from Fig. 30.2. Belostomatidae. fimnogefon sp. Usinger, 1956). Key to Subfamilies of Belostomatidae 1. Abdominal sterna 3-7 with midplate and paired lateral plates, the latter not differentiated by a mesal sulcus; sulci differentiating lateral plates terminating near proximal angles of mesal plate 7; abdominal spiracles located near center of lateral plates, far removed from lateral sulci . Belostomatinae - At least abdominal sternum 6 and usually also some other segments (from 3 to 7) with mesal, sub¬ lateral, paired spiracle-bearing lateral plates; lateral sulci terminating near proximal angles of lateral plate 7; abdominal spiracles on mesal margins of lateral plates close to lateral sulci . 2 2. Proximal portions of lateral lobes 7 (laterad of subgenital plate) subdivided into sublateral and lateral plates; lateral sulci on lateral lobes 7 either parallel to margins of subgenital plates and terminating discally, or converging toward tip of subgenital plate and disappearing underneath latter; antennal segments 2 and 3 each with one fingerlike projection, 4 with 2; hind tibia and tarsus thinly com¬ pressed, much more dilated than middle tibia and tarsus; foretarsus 3-segmented, although often appearing 2-segmented externally; anterior claw of foreleg large, posterior claw vestigial to absent; length 37-115 mm . Lethocerinae - Lateral lobes 7 formed entirely or largely by lateral plates, sublateral plates being absent or not visible externally, or developed as minute triangular sclerites; lateral sulci terminating at basilateral angles of subgenital plate (not continuing onto lateral lobes 7) or only slightly distad of them; antennal segment 2 with one fingerlike projection, 3 with a large expanded dorsal lobe, 4 short and transverse; tibiae and tarsi of middle and hind legs similar, flat, narrow; foretarsus 2-segmentcd (externally appearing 1-segmented); claws of foreleg paired, vestigial; length 25-30 mm . Horvathiniinae Key to subfamilies of Belostomatidae adapted from Lauck and Menke, 1961. 112 true bugs of the world (HEMIPTERA: HETEROPTERA) Fig. 30.3. Belostomatidae. A. Antenna, Horvathinia sp. (from Lauck and Menke, 1961), B. Aedeagus, Diplonychus rusticum (Fabricius) (from Kumar, 1961). C. Aedeagus, lateral view, Lethocerus sp. D. Aedeagus, sagittal view, Lethocerus sp. (C, D from Lauck and Menke, 1961). E. Testes, Diplonychus rusticum. F. Spermatheca, Lethocerus niloticus Stal. G. Spermatheca, L griseus (Say) (E-G from Pendergrast, 1957). tennal segments 2-4 with dorsal appendages; hind tibia and tarsus broadly flattened; airstraps long, mesal mar¬ gins nearly contiguous. This subfamily includes only the cosmopolitan genus Lethocerus Mayr, comprising approximately 25 species. They are the largest of all heteropterans. Cummings (1933) and DeCarlo (1938) dealt with the New World fauna and Menke (1960a) with that of the Old World. Specialized morphology. The “airstraps” of adults, derived from abdominal tergum 8, are the most distinc¬ tive feature of the group. They do not form a tube, as in the Nepidae; instead, air is transmitted to the sub- hemelytral airstore by a channel, formed by the setae (which converge mesioventrally). Static sense organs are present in the Belostomatidae. They are associated with the spiracles of abdominal sterna 2-7 (Moller, 1921), and although not as large and com¬ plex, apparently serve the same function as those occur¬ ring in the Nepidae (Figs. 29. II, 31.2C). Natural history. Many species of Belostomatidae live in standing water. Most are excellent swimmers, although they generally lie in wait for their prey. A few taxa, such as Abedus Stal in the New World and some Diplonychus Laporte in the Old World tropics, live in slow-flowing portions of streams. Most species are probably general predators, eating whatever they are capable of catching. BELOSTOMATINAE (FIGS. 30.1, 30.2). Relatively small to large, ovoid, elliptical, or elongate; antennal segments 2 and 3 each bearing a long, dorsal, fingerlike process, segment 4 long; middle and hind tibia and tarsus usually flattened, not broadly dilated; pretarsus of hind leg of Diplonychus sp. as in Fig. 29.2H; airstraps variable; metathoracic scent glands absent. Included genera are Abedus Stal (10 species, south¬ western United States to Central America; Menke, 1960b); Belostoma Latreille (approximately 60 species. New World; Lauck, 1962, 1963, 1964); Diplonychus Laporte (approximately 20 species, Africa east to Aus¬ tralia; Poisson, 1949); Hydrocyrius Spinola (five species, Africa and Madagascar; Brown, 1948; Poisson, 1949); Limnogeton Mayr (four species, Africa; Poisson, 1949). HORVATHiNiiNAE. Medium-sized, elliptical; antennal segment 2 long, flattened, segment 3 flattened with a large dorsal lobe, segment 4 dorsoventrally elongated (Fig. 30.3A); hind tibia and tarsus flattened, not broadly dilated; airstraps short, mesal margins remote. Only Horvathinia Montandon, with nine nominal spe¬ cies from central and southern South America, is in¬ cluded. Members of the group are rarely encountered in nature, and their habits and habitats are poorly known. The group was revised by DeCarlo (1958). LETHOCERINAE. Large to very large, elongate ovoid; an- Belostomatidae 113 including vertebrates such as fishes and frogs. Limno- geton, which is unique in not having any of the legs modi¬ fied for swimming, is apparently an obligate freshwater snail predator (Voelker, 1966), Adult belostomatids protrude their airstraps through the surface, while lying motionless in the water. Respi¬ ration in adults takes place primarily through the dorsal first abdominal spiracles, the remaining ventral spiracles playing a lesser role (Miller, 1961; Parsons, 1972, 1973). Respiration in nymphs takes place from the airstore on the abdominal venter. Uniquely among the Insecta, female Belostomatinae deposit their eggs on the dorsal surface of the males (Fig. 30.2). Males that are carrying eggs engage in brooding behavior, including exposing the eggs to atmospheric air (Smith, 1976). Most belostomatids are capable of ejecting a foul¬ smelling liquid from the anus, apparently as a defensive reaction (Harvey, 1907). With the exception of the Be¬ lostomatinae, male giant water bugs have well-developed metathoracic scent glands and are said to produce a strong smell, although certainly not of the quantity or intensity of many large Pentatomoidea. When Lethocerus is eaten by humans, a common practice in Thailand and other parts of Asia, the scent-gland odor and taste are very obvious. Although many nepomorphans, and in fact most het- eropterans, are attracted to lights, the belostomatids have in some areas been dubbed “electric light bugs” be- -cause of their conspicuous presence, especially at mer¬ cury vapor lamps. Distribution and faunistics. The Belostomatidae are worldwide in distribution, although their greatest diver¬ sity is in the tropics. Aids to identification are cited under the subfamily treatments and Nepomorpha. 31 Nepidae General. The water scorpions are brown, elongate- ovoid to tubular-bodied aquatic bugs that range in body length from 15 to 45 mm. Their caudal breathing siphon may be as long or longer than the body (Fig. 31.1). Diagnosis. Eyes relatively small within Nepomorpha (Fig. 31.2A); antennae usually 3-segmented, segment 2 and sometimes 3 with fingerlike projections (Fig. 31.2B); membrane of forewing with numerous cells (Fig. 31.1); I h j ?• I \ - \ \ Fig. 31.1. Nepidae./.accofrep/ies sp. (from Slater, 1982). # all legs elongate, slender, forelegs raptorial (Figs. 10.4B, 31.1), all tarsi 1-segmented; adult metathoracic and nymphal dorsal abdominal scent glands absent; static sense organs near spiracles on ventral laterotergites 4-6 (Figs. 29.11, 31.2C); male genitalia symmetrical, para- meres and genital capsule as in Fig. 31.2E; aedeagus as /—\,-- /'■N. ! 114 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 31.2. Nepidae. A. Head and pronotum, lateral view, Ranatra compressicollis Mo.itandon, B. Antenna, Ranatra compressicollis (A, B from Lansbury, 1974b). C. Ventral view, abdomen, Ranatra sp. D. Ventral view, abdomen, nymphal Ranatra sp,, showing air channels, (C, D from Menke, 1979a). E. Genital capsule, lateral view, R. drakei Hungerford. F. Aedeagus., lateral view, R. drakei (E, F from Lansbury, 1974b). G. Spermatheca, Nepa cinerea (Linnaeus) (from Larsen, 1938). Abbreviations: hso, static sense organ; pa, paramere. in Fig. 31.2F; subgenital plate forming an ovipositorlike structure in some taxa; spermatheca as in Fig. 31.2G; eggs uniquely with from 2 to 26 respiratory horns on anterior pole. Classification. The Linnaean genus Nepa was first recognized as a higher taxon by Latreille (1802) as Nepa- riae. Modem treatments on the higher classification of the group are those of Menke and Stange (1964) and Lans¬ bury (1974a), the latter author having done much recent revisionary work on the group. Currently two subfamilies comprising 14 genera and 231 species are recognized. Nepidae 115 Key to Subfamilies of Nepidae 1. Laterosternites flat, visible, not concealed by ventral laterotergites; ventral laterotergites not longitu¬ dinally subdivided; female subgenital plate (= operculum) broad and short, not exceeding apex of abdomen, if triangular, not keeled; body flattened; mesocoxae separated by more than coxal width . Nepinae - Parasternites concave, infolded, concealed by ventral laterotergites (Fig. 31.2D); ventral laterotergites longitudinally subdivided; female subgenital plate either laterally compressed and keeled or extending well beyond apex of abdomen (Fig. 31.2C); body nearly cylindrical in cross section; mesocoxae separated by less than coxal width . Ranatrinae NEPINAE (FIG. 31.1). Laterosternites not concealed by folding of abdomen; ventral laterotergites divided by a crease tangential to spiracles; female subgenital plate broad and flattened, never extending beyond end of abdo¬ men; eggs with 5 or more respiratory horns. Two tribes are recognized (Lansbury, 1974a). TheCu- rictini includes only the primarily tropical Curicta Stal from the New World. The Nepini includes seven genera, the best known being the paleotropical Laccotrephes Stal and the Holarctic Nepa Linnaeus. RANATRINAE. Laterostemites concealed by folding of abdomen (Fig. 31.2D); ventral laterotergites not divided by a crease tangential to spiracles; female subgenital plate laterally compressed, keeled (Fig. 32.1C), and often ex¬ tending beyond apex of abdomen; eggs with 2 respiratory horns. Three tribes are recognized (Lansbury, 1974a). The Austronepini and Goondnomdanepini, each including only a single genus from Australia, respectively, Austro- nepa Menke and Stange and Goondnomdanepa Lansbury. The Ranatrini comprise the tropical Asian Cercotmetus Amyot and Serville and the widespread Ranatra Fabri- cius, the Oriental species of which were reviewed by Lansbury (1972). Specialized morphology. The water scorpions are noted for their anteriorly projecting head, the elongate prothorax (Fig. 31.2A) with the proximal coxae and the anteriorly directed legs, the respiratory siphon, and the presence of 2—26 respiratory horns on the anterior pole of the eggs. The modified abdominal tergum 8 is formed into a paired structure, forming a tube, which directs air from the atmosphere to abdominal spiracle 8, which is placed dorsally. The static sense organs in the Nepidae, located near the spiracles on ventral laterotergites of abdominal segments 4-6, are large and conspicuous (Figs. 29.11, 31.2C) (Baunacke, 1912). These structures have been shown experimentally to function to keep the insect correctly oriented in the water (Thorpe and Crisp, 1947). The mor¬ phology of Nepa cinerea Linnaeus was treated in detail by Hamilton (1931). Natural history. Water scorpions are poor swimmers and tend to “crawl” through the water. Most inhabit quiet water in ponds or streams. They often “hang” in vegeta¬ tion, with the respiratory siphon protruding through the water surface and the forelegs outstretched, waiting for prey. The foretibia has an apical sense organ, which ap¬ parently senses prey vibrations, although visual responses seem also to be involved in prey capture (Cloarec, 1976). Life histories oiRanatra spp. have been studied by Torre- Bueno (1906) and Radinovsky (1964). The abdominal dorsum of some nepids, including Laccotrephes spp., is bright red, in strong contrast with the dull greenish brown of the body surface. When the bugs are in flight the red coloration is obvious, but when they alight they become nearly invisible because the bright color disappears under the hemelytra. Distribution and faunistics. This worldwide group shows its greatest diversity in the tropics. Sources are noted above. Ochteroidea 32 Gelastocoridae General. The toad bugs, which range in length from 7 to 15 mm, like the Ochteridae, are usually found in riparian situations. They very much resemble small Bu- fonidae, and all are capable of jumping. The coloration and texture often match remarkably the background upon which they are found. Diagnosis. Body surface often roughened and “warty” (Fig. 32.1); eyes large, with a distinct mesal 116 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) emargination on dorsal surface, appearing reniform (Fig. 32.1); adults with ocelli; labrum broad and flaplike; antennae 4-segmented, without fingerlike projections (Figs. 29.1 A, 32.2A); membrane of forewing some¬ times reduced, if developed, usually with numerous veins (Fig. 32.1); forefemora greatly enlarged, interior surface grooved for reception of tibia and tarsus; foretarsus 1- segmented or sometimes fused with tibia, middle tarsus 2-segmented, hind tarsus 3-segmented, elaws unequally developed on forelegs, equally developed on middle and hind legs; foreleg pretarsus of nymphs as shown in Fig. 29.2A, adult as in Fig. 29.2B; antennal cleaner on fore¬ leg as in Fig. 10.41, J; nymphs lacking dorsal abdomi¬ nal scent glands; spiracles on abdominal sterna 3 and 4 shifted toward midline; male genitalia asymmetrical, ab¬ dominal venter as in Figs. 29.IB, 32.2E; aedeagus as in Fig. 32.2B; right paramere as in Fig. 32.2C, D; left paramere reduced or absent; female abdominal venter as in Fig. 32.2F; spermatheca as in Fig. 32.2G. Classification. This group was first recognized as a higher taxon by Billberg (1820) under the name Galgu- lides, after Galgulus Latreille, a preoccupied name re¬ placed much later by Gelastocoris Kirkaldy (1897). Two genera are currently recognized. Gelastocoris from the New World, contains approximately 15 species. Nerthra Say, which contains approximately 85 species, is cosmo¬ politan and is a senior synonym of Mononyx Laporte. The taxon has frequently been treated as a family group under the name Mononychidae. These genera, which have sometimes been accorded subfamily status, can be separated according to the following key. Fig. 32.1. Gelastocoridae. Nerthra sp. Fig. 32.2. Gelastocoridae. A. Lateral view, head, Gelastocoris oculatus (from Popov, 1971). B. Aedeagus, G. oculatus (from Kumar 1961) C. Right paramere, Nerthra hungerfordi Todd. D. Right paramere, G, peruensis Melin. E. Male abdomen, N. amplicollis (Stal). F. Female’abdomen' N. amplicollis (C-E from Todd, 1955), G. Spermatheca, N. annuUpes (Horvath) (from Pendergrast, 1957). Gelastocoridae 117 Key to Genera of Gelastocoridae 1. Foretarsus 1-segmented, freely articulating with tibia, with 2 claws; forefemur moderately incrassate, about twice as long as wide, its apposable face flat and bordered by two rows of short spines; labium arising near apex of head, stout, directed caudad (Fig. 32.2A) . Gelastocoris - Foretarsus fused with tibia, tibiotarsus terminating in a single claw (Fig. 32.1); forefemur subtri- angular, very broad at base, about as wide as long, its apposable face with a flangelike extension projecting over tibiotarsus when apposed; labium appearing to arise from ventral surface of head, slender, inversely L-shaped, apex directed ventrad to anteroventrad . Nerthra Specialized morphology. The toadlike appearance and the modified forelegs, with unusually large femora (Fig. 32.1), are distinctive for the group. The forewings in some Nerthra spp. are immovably fused (Todd, 1955). Parsons reviewed the morphology of the head (1959), tho¬ rax (1960b), nervous system (1960a), and abdomen and thoracic scolopophorous organs (1962) of Gelastocoris oculatus (Fabricius). Natural history. Gelastocorids are generally found in riparian situations, although Nerthra spp. are not infre¬ quently encountered a long distance from water. Some species can be extremely numerous at a given locality. 33 Ochteridae General. Sometimes referred to as velvety shore bugs (Menke, 1979b), members of this group—which range in length from 4.5 to 9 mm—appear remarkably like Sal- didae with short antennae, generally dark coloration, and pruinose markings on the dorsum. Diagnosis. Eyes large, mesal margin emarginate dor- sally (Fig. 33.1); ocelli present; antennae 4-segmented, visible dorsally, projecting, filiform (Figs. 29. ID, 33.1); clypeus transeversely rugose; labium long, slender, taper¬ ing, reaching hind coxae, segment 3 much longer than others (Fig. 33.2A); medial and costal fracture continu¬ ous and well developed, membrane of forewing with sev¬ eral closed cells (Fig. 33.1), but without anastomosing veins found in many other Nepomorpha; all legs slen¬ der, forefemora not enlarged or otherwise modified, tar¬ sal formula 2-2-3; foreleg pretarsus of nymph as in Fig. 29.2C; nymphs lacking dorsal abdominal scent glands; male abdomen ventrally as in Fig. 29.IE; aedeagus as in Fig. 33.2C; genital capsule and right paramere as in whereas others seem to lead a solitary existence. Al¬ though most Gelastocoris spp. are usually found in the open, Nerthra spp. are capable of at least some burrowing and are frequently found beneath stones or other objects. The life history of Gelastocoris oculatus was reviewed by Hungerford (1922b). Distribution and faunistics. Although toad bugs oc¬ cur worldwide, they are most diverse in the tropics. The genus Nerthra is especially diverse in Australia and Mela¬ nesia. The major source on the group is by Todd (1955). Fig. 33.2B; left paramere reduced; ovipositor reduced; spermatheca as in Fig. 33.2D. Classification. This group was first recognized as a higher taxon under its current name by Kirkaldy (1906). Latreille (1809) proposed the unnecessary replacement name Pelogonus for Ochterus Latreille, and the family was often referred to as Pelogonidae in the older litera¬ ture. Three genera and 55 species are currently recognized. Specialized morphology. Ochterids are notable for their saldidlike appearance (Fig. 33.1), but they nonethe¬ less have a preponderance of obvious nepomorphan char¬ acters, including the short, ventrally inserted antennae, asymmetrical male genitalia, and tubular spermatheca. Rieger (1976) reviewed the morphology of the head and thorax of Ochterus marginatus Latreille in detail. Natural history. These bugs are usually found along the shores of ponds or streams, and in the tropics they largely replace the Saldidae; they may also inhabit more open sandy habitats. Life histories of ochterids have been published by Takahashi (1923) and Bobb (1951). Distribution and faunistics. Ochterids are found worldwide, but are most diverse in the tropics. The West¬ ern Hemisphere fauna was revised by Schell (1943), the Eastern Hemisphere fauna by Kormilev (1971b), and Ochterus in Australia by Baehr (1989). 118 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 33.1. Ochteridae. Ochterus barberi Schell (from Usinger, 1956). Corixoidea 34 Corixidae General. The water boatmen represent the most speci- ose of the families of aquatic bugs. Because of their oarlike hind legs (Fig. 34.1), they have the general ap¬ pearance of flattened backswimmers, but swim with the dorsal side up. They range in length from 2.5 to 15 mm. Diagnosis. Small to medium-sized, dorsum flattened; head broad, strongly hypognathous; ocelli absent ex¬ cept in Diaprepocorinae; antennae 3- or 4-segmented, hidden between eyes and prothorax; labium modified from typical heteropteran condition, broad at base, taper¬ ing distally, unsegmented, immovably fused to head, usually with transverse grooves and a longitudinal chan¬ nel; labrum reduced, covered by labium; scutellum ex- A Fig. 33.2. Ochteridae. A. Lateral view head, Ochterus marginatus (La- treille) (from Popov, 1971). B. Genital capsule, O, barberi (from Schell, 1943). C. Aedeagus, 0. seychellensis D, A. Polhemus (from D. A. Polhemus, 1992). D. Spermatheca, O. marginatus (from Pendergrast, 1957). posed or hidden by pronotum; forewings of uniform tex¬ ture, membrane area without veins (Fig. 34.1); forelegs relatively short; foretarsus with a single segment, usually modified into a pala in the form of a spoon or a scoop, fringed with long setae (Fig. 34.2A), sometimes fused with tibia; middle legs very long and slender, tarsi 1- or 2-segmented, pair of claws very long (Fig. 34.1); hind legs flattened, oarlike, fringed with setae, tarsi 2- segmented (Fig. 34.1); stridulatory structure formed by field of pegs on basomesal surface of forefemur apposed to edges of maxillary plates (in males only of Micronecti- nae); metathoracic scent glands present in adults; larval dorsal abdominal scent glands present between terga 3/4, 4/5, and 5/6; male abdomen with a “strigil” posterolater- ally on tergum 6 (Fig. 34.2B, C); male distal abdominal Corixidae 119 terga and sterna and genital segments usually strongly asymmetrical (Fig. 34.2B); aedeagus as in Fig. 34.2D, E; parameres as in Fig. 34.2F; spermatheca as in Fig. 34.2G, H. Classification. The Corixidae were first treated at the family rank by Leach (1815) under the name Corixida. Their unusual morphology has attracted the attention of many authors, and Borner (1934) went so far as to accord them subordinal rank under the name Sandaliorrhyncha. There seems little doubt, however, that they belong to the Nepomorpha, in spite of their obvious specializa¬ tions; more recent opinions on their phylogenetic position can be seen in Figs. 29.3 and 29.4. The classification of Hungerford (1948) recognized six subfamilies, which comprise 34 genera and 556 species. Key to Subfamilies of Corixidae 1. Scutellum broadly exposed, only anterior margin covered by short, transverse pronotum; female foreleg (and sometimes male foreleg) with fused tibiotarsus; color patterns never as below . 2 - Mesoscutellum covered by arcuate to subtriangular posterior extension of pronotum (Fig .34.1), rarely its extreme apex exposed, although often fully exposed in preserved specimens owing to deflection of pronotum; foreleg with tibia and pala separate in both sexes; pronotum and hemelytra mostly with complex mosaic of alternating transverse, linear, undulating, hieroglyphical, or zig-zag light and dark patterns, these rarely obsolete or absent .. 3 2. Ocelli present; middle tarsus 2-segmented; male foreleg with fused tibiotarsus; antennae 4-segmented; Australia and New Zealand . Diaprepocorinae - Ocelli absent; middle tarsus 1-segmented; male foreleg with pala and tibia separate; antennae 3- segmented; tropicopolitan . Micronectinae 3. Exocorial pruina present on small basal portion of forewing, extremely short, about as wide as and similar in length to endocorial pruina; embolar groove practically undeveloped; proximal sector of Cu on forewing indistinguishable, Cu developed as a distinct ridge only distad to meeting with costal fracture; Afrotropical . Stenocorixinae - Exocorial pruina present along costal margin in long embolar groove, long and narrow, much longer than endocorial pruina, usually terminating distad of costal fracture, rarely at costal fracture or just proximad of it (the latter structure absent in some taxa); Cu present from near base of forewing as distinct ridge delimiting emboliar groove and exocorial pruina; junction of Cu and costal fracture (or projected intersection) situated at border of exocorial pruina or nearby . 4 4. Labium without transverse striation on ventral face; male pala cylindrical or terete, without dif¬ ferentiated palmar face and without palar pegs; male palar claw long, thick, appendage-like, not resembling rake setae; media of forewing usually meeting apex of costal fracture, forming closed Fig. 34.1. Corixidae. Trichocorixa reticulata (Guerin-Meneville) (from Lauck, 1979). 120 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEHOPTERA) field within pruinose embolar groove, inner angle of field remote from Cu ridge; cosmopolitan . Cymatiainae - Labium with transverse striation on ventral face; male pala variously shaped, palmar face differenti¬ ated and palar pegs developed, if pegs, poorly defined or reduced in number, then pala very broad; male palar claw usually resembling a thickened rake hair, if appendage-like then short; exocorial pruina without a closed field within it, as above . 5 5. Infraocular (postocular) portion of gena very broad in lateral view, ventrally delimited by a hypocular sulcus arising near subacutely produced inferior angle of eye and running toward posterolateral mtu'gin of head; posteroventral margin of eye markedly concave in lateral view; media running very close to and parallel with Cu, often indistinct; Neotropical . Heterocorixinae - Infraocular portion of gena very narrow to broad in lateral view, hypocular sulcus present or absent; if infraocular portion of gena broad and hypocular sulcus present, then the latter arising almost midway along posteroventral margin of eye, this margin shallowly concave, inferior angle not subacutely produced, usually broadly rounded; media variously developed, but not as above, usually running midway between costal margin and Cu, only apically turning toward Cu or the inner portion of costal fracture; cosmopolitan . Corixinae coRixiNAE (FIG, 34,1), Labium with transverse sulcations; antennae 4-segmented; scutellum completely covered by pronotum or nearly so; embolar groove and nodal furrow (costal fracture) present; female pala spoon-shaped, male pala variable; claw of hind leg inserted subapically. This is by far the largest of the subfamilies and contains the majority of known species placed in four tribes and 26 genera, with the large and widespread genus Sigara Fabricius being divided into many subgenera. The group is worldwide in distribution, CYMATIAINAE, Labium without transverse sulcations; antennae 4-segmented; scutellum covered by pronotum; embolar groove present, nodal furrow absent; pala elon¬ gate, nearly cylindrical in both sexes, not fused with tibia; hind leg claw inserted apically. This group, whose name has been emended from Cymatiinae because of homonymy, contains only two genera, the Holarctic Cymatia Flor and Cnethocymatia Jansson from northern Australia and New Guinea. DiAPREPOCORiNAE. Ocelli present; antennae 4-seg¬ mented; scutellum exposed; male and female foreleg with fused tibiotarsus; middle tarsus 2-segmented. These water boatmen have been thought of as the most primitive members of the group because they have ocelli, an unusual trait within the Nepomorpha. The group con¬ tains only Diaprepocoris Kirkaldy from Australia and New Zealand. HETEROCORIXINAE. Labium with transverse sulcations; scutellum covered by pronotum; embolar groove present, nodal furrow present; tibia of foreleg nearly as long as palm-shaped pala. This New World tropical group contains only Hetero- corixa Buchanan-White. MiCRONECTiNAE. Labium with transverse sulcations; an¬ tennae 3-segmented, last segment broad and lying in a pocket in head; scutellum exposed; embolar groove short, shallow, its pruinose area not crossed by nodal furrow'; pala fused to foretibia in female; claw of hind leg inserted apically. This group contains the Old Wodd Micronecta Kirkaldy —with many subgenera—which is diverse in the Palearc- tic and the Orient. Tenagobia Bergroth is restricted to the Neotropics. STENOCORixiNAE. Body more elongate and slender than in other Corixidae; labium with transverse sulcations; pala slender; scutellum covered by pronotum; embolar groove absent, nodal furrow present; male abdomen only slightly asymmetrical. This subfamily is represented by the single African species Stenocorixa protrusa Horvath. Specialized morphology. The morphology of the labium in the Corixidae is unique within the Heteroptera. It was long thought to be associated strictly with feeding habits, but the correlation may not be so strong as sup¬ posed. The forelegs also are highly modified, the femur functioning in stridulation, and the tarsus existing in the form of a spoon-shaped pala (Fig. 34.2A), which func¬ tions in food gathering. The brown and pale hieroglyphic patterning of the dorsum is characteristic of most mem¬ bers of the family and is important in the recognition of some taxa. The so-called strigil on abdominal tergum 6 (Fig. 34.2B, C) in the males is misnamed and appears to have nothing to do with stridulation. It has been implicated in holding the female during mating (Larsen, 1938) and in helping to maintain the subhemelytral air stores (Popham et al., 1984). Natural history. In addition to being the most speci- ose of all the families of Nepomorpha, the Corixidae are also the most widely distributed, occupy a relatively Corixidae 121 Fig. 34.2. Corixidae. A. Male foreleg, Cor/'xa dentipes Thunberg, B. Male abdominal dorsum, C. jakowleffi (Horvath). C. Strigll, C. dentipes (A- C from Jansson, 1986). D. Aedeagus, Agraptocorixa hyalinipennis (Fieber). E. Aedeagus, Hesperocorixa interrupta (D, E from Kumar, 1961). F. Parameres (right above, left below), C. dentipes (from Jansson, 1986). G. Spermatheca, A. hyalinipennis (from Kumar, 1961). H. Gynatrial complex, including spermatheca, SIgara sahibergi (Fieber) (from Larsen, 1938). papers by Hutchinson (1929) for southern Africa and Lundblad (1933) for the Orient should be consulted. Naucoroidea | -^^^- 3 35 j ■"'I Potamocoridae J General. These small bugs, which are between 2.5 - 1 and 3.0 mm long, have the appearance of small naucorids (Fig. 35.1A). J wide range of habitats, and are sometimes extremely nu¬ merous. Some are known to inhabit saline waters on an exclusive basis (see Lauck, 1979). Water boatmen, unlike other nepomorphans, eat pri¬ marily plant material, particularly algae, using their palae to forage on bottom ooze. They are also known to feed on animal matter as well, such as nematoceran larvae, including mosquitoes (Lauck, 1979). Several species are eaten by humans in Mexico, and those same species are important in the pet-food trade. Stridulation in the Corixidae and its role in courtship and mating have been studied in detail by Jansson (1972, 1976). Distribution and faunistics. The New World fauna was treated in its entirety by Hungerford (1948), and that work is still the single most important reference on the fauna. The European fauna has been reviewed exhaus¬ tively by Jansson (1986). The remainder of the world is covered in a somewhat more piecemeal manner, but 122 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 35.1. Potamocoridae. A. Potamocoris parvus Hungerford (from Hungerford, 1941). B. Ventral view, body, P. nieseri van Doesburg (from van Doesburg, 1984). C. Ventral view, head, P. parvus (from Hungerford, 1941). D. Foreleg, P. nieseri (from van Doesburg, 1984). E. Middle leg, P. parvus. F. Hind leg, P. parvus. G. Male abdomen, ventral view, Potamocoris parvus. H. Genital capsule, P. parvus. I. Female abdomen, ventral view, Pofamocoris parvus (E-l from Hungerford, 1941), Diagnosis. Small; eyes not overlapping anterolateral angles of pronotum; antennae relatively long, visible dor- sally, filiform (Fig. 35.1 A, B); labium short, just reaching onto prosternum (Fig. 35.IB, C); membrane of fore¬ wing without veins; forefemora only slightly enlarged, with tibia and tarsus opposable, middle and hind femora without obvious swimming modifications; foretarsus 1- segmented, middle and hind tarsi 2-segmented; 2 claws on all legs; nymphs with dorsal abdominal scent glands between terga 3/4 (Hungerford, 1942); male genital seg¬ ments as in Fig. 35. IG, H; female as in Fig. 35.11. Classification. In his original description of Potamo¬ coris, Hungerford (1941) noted the unusual combination of characters in this group of bugs. In the same jour¬ nal issue, Usinger (1941) accorded the group subfamily status in the Naucoridae. Cobben (1978), in a footnote, indicated that the group deserved family status, a position followed by Stys and Jansson (1988). Two genera, Coleopterocoris Hungerford (1942) and Potamocoris, each with four species, are recognized. Specialized morphology. The attributes of the Po¬ tamocoridae are seemingly a mixture of some of those possessed by the Aphelocheiridae and Naucoridae, in¬ cluding the elongate antennae of the former and the short labium of the latter. Natural history. Nothing appears to be known of the biology. Most specimens have apparently been collected at lights, although Hungerford (1942) had access to a nymph. Distribution and faunistics. This group is restricted to the Neotropics. The main sources are those of Hunger¬ ford and Usinger mentioned above, DeCarlo (1968), La Rivers (1950, 1969), J. T. Polhemus and D. A. Polhemus (1982), and van Doesburg (1984). Potamocoridae 123 36 Naucoridae General. Naucorids, or creeping water bugs, are usu¬ ally ovoid (Fig. 36.1) to slightly elongate, flattened bugs ranging in length from 5 to 20 mm. Most have the general appearance of tiny belostomatids, in both coloration and shape. Diagnosis. Eyes often overlapping anterolateral angles of pronotum (Fig. 36.1); antennae 4-segmented, short, hidden, not projecting laterally; labium short and stout (Fig. 36.2A); forewing membrane without vena¬ tion (Fig. 36.1); forefemora conspicuously enlarged (Fig. 10.4A); apex of foretibia as in Fig. 10.7A; hind legs often modified for swimming; foretarsus usually fused with tibiae, 1- or 2-segmented, with 1 or 2 claws or with¬ out claws, middle and hind tarsi with 2 distinct segments (basal segment very small or absent), with 2 equally developed claws; pretarsal structure variable (see Fig. 29.2E); nymphal pretarsus as in Fig. 29.2D; metathoracic scent glands present; posterolateral angles of abdominal connexiva sometimes produced; nymphal dorsal abdomi¬ nal scent gland between terga 3/4; some taxa with paired sublateral sense organs on abdominal sternum 2; male and female venter in copulatory position as in Fig. 36.2B; aedeagus as in Fig. 36.2C; spermatheca vermiform as in Fig. 36.2D. Classification. This group was first treated at family rank by Leach (1815) as the Naucorida and has been Fig. 36.1. Naucoridae. Pelocoris shoshone La Rivers (from Usinger, 1956). so recognized by most subsequent authors, although the limits of the family have been subject to dispute, as explained in the discussions under Aphelocheiridae and Potamocoridae. The most recent classification (Stys and Jansson, 1988) included five subfamilies—separated as in the follow¬ ing key—comprising 40 genera and approximately 395 species. Key to Subfamilies of Naucoridae 1. Labium arising from a deep excavation on underside of head, its insertion markedly distant from apex of head owing to more or less horizontal, laminate extension of latter; Oriental and Papuan . . Cheirochelinae - Labium arising freely, not from an excavation, usually near ventral apex of head; sometimes true apex of head markedly folded over and insertion of labium distinctly ventra . 12 2. Lateral margins of pronotum crenulate, dorsal surface roughly granulate; head rather narrow, anterior portion longer than wide, distinctly projecting in front of subglobular eyes; head deeply inserted into anteriorly strongly concave pronotum; maxillary plates well delimited and anteriorly reaching or exceeding level of apex of head; New World . Cryphocricinae (part) - Lateral margins of pronotum smooth, dorsal surface smooth to punctate, not granulate; head broad, outline streamlined jointly with pronotum; interocular region usually much broader than long (narrow in Ambrysini), only slightly exceeding eyes, the latter never subglobular and usually not projecting beyond outline of head, eyes rarely somewhat rounded and interocular portion slightly projecting; maxillary plates variously developed, often incompletely or not at all delimited, not reaching level of apex of head . 3 3. Proepimeral lobes strongly expanded, meeting to cover posterior portion of prosternum: head deeply inserted into concave and markedly trisinuate anterior pronotal margin (Ambrysini) . . Cryphocricinae (part) 124 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) - Proepimeral lobes not in mutual contact, posterior portion of prosternum exposed; head usually not deeply inserted into pronotum, anterior pronotal margin usually shallowly concave to almost straight . 4 4. Foreleg pretarsus in females always with 2 claws, often minute and closely appressed, sometimes resembling a single structure; males of most genera with a tomentose adhesive patch on ventral sur¬ face of middle tibiae, weakly developed in females; morphological apex of head folded over, oriented posteriorly, insertion of labium markedly posterior to topographical apex of head; middle and hind femora with 2 longitudinal rows of conspicuous bristles or spinelike setae on ventral face, in addition to 2 usual rows on posterior face .. Laccocorinae - Foreleg pretarsus with or without 1 minute claw; middle tibia without distoventral adhesive tomentose patch; labium usually inserted apically or subapically on ventral apex of head, rarely on anteroventral surface of head; middle and hind femora without additional rows of bristles or spiniform setae on ventral surfaces, or with just an indication of upper row, or with some scattered additipnal setae . 5 5. Meso- and metastema with prominent, broad, laterally expanded median longitudinal carinae bearing foveae or otherwise excavate; inner margins of eyes (dorsal view) distinctly diverging anteriorly; gula short; apex of head (anterior view) simply arcuate, not notched; lateral margins of abdomen with fine serration; body broadly oval to subcircular, flattened; New World . Limnocorinae - Meso- and metasternal longitudinal carinae inconspicuous, thin, platelike, or absent; inner margins of eyes in dorsal view usually distinctly converging anteriorly, infrequently parallel-sided to slightly- diverging; gula usually long; apex of head straight, delimited laterally by distinct notches or indenta¬ tions laterad of basal angles of labrum; lateral margins of abdomen without fine serration; body more elongate, robust, dorsum moderately convex; worldwide . Naucorinae CHEiROCHELiNAE. Labium inserted in a deep excavation well removed from anterior margin of head; labrum often greatly reduced; anterior margin of pronotum excavated in region adjoining interocular space of head. Three tribes are recognized in this Oriental (includ¬ ing New Guinea) group: Cheirochelini (three genera), Sagocorini (eight genera), and Tanycricini (three genera). CRYPHOCRiciNAE (FIG. 36.1). Labium inserted near an¬ terior margin of head, not in a deep excavation; labrum well developed; anterior margin of pronotum excavated in region adjoining interocular space of head. Three tribes are recognized in this New World group; Ambrysini (two genera), Cataractocorini (one genus), Cryphocricini (two genera). LACCOCORINAE. Labium inserted near anterior margin of head, not in a deep excavation; labrum well devel¬ oped; anterior pronotal margin nearly straight; foretarsus always with 2 claws. Seven genera are currently recognized in this primarily Paleotropical group, which is marginally represented in the Palearctic (Stys and Jansson, 1988). LIMNOCORINAE. Labium inserted near anterior margin of head, not in a deep excavation; labrum well developed; anterior pronotal margin nearly straight; foreleg with at most 1 claw; meso- and metasterna with prominent, broad, laterally expanded median longitudinal carinae bearing foveae or otherwise excavate; lateral abdominal margins with fine serrations; abdominal terga 2-5 fused; stridulatory structure in all species involving hind femur and connexival margin. This New World group currently contains two genera. NAUCORINAE. Labium inserted near anterior margin of head, not in a deep excavation; labrum well developed; anterior pronotal margin nearly straight; foreleg with at most 1 claw; meso- and metasterna not prominent, with¬ out expanded longitudinal carina; no stridulatory struc¬ tures. This worldwide group appears for the most part to be diagnosed by the absence of characters found in the Lim¬ nocorinae, a point of view supported by the character analysis of Mahner (1993). The seven included genera are restricted primarily to the tropics of either the Old or New World. Specialized morphology. Researchers studying nau- corids have tended to treat the Potamocoridae and Aphe- locheiridae as having more primitive attributes than the Naucoridae as here construed. Thus, they have consid¬ ered the short, concealed antennae and short labium to be apomorphic for the Naucoridae, although the antennae project beyond the eyes in the Tanycricini. The Cryphocricini have plastron respiration (Fig. 29. IG, H) and pressure receptors (Parsons and Hewson, 1975), as possibly do some other Naucoridae. Natural history. Naucorids live in a variety of aquatic environments. Some, such as Pelocoris femoratus (Pali- sot de Beauvois) in North America live in quiet pond waters. Limnocoris spp., and indeed many naucorids and aphelocheirids, are found in marginal riffles in streams on gravel substrates. A few taxa, such as Cryphocricos Signoret in the New World tropics and Idiocarus Montan- Naucoridae 125 Fig. 36.2. Naucoridae. A. Lateral view head, Ambrysus magniceps La Rivers (from Parsons, 1969). B. Copulation, Naucoris cimicoides (Linnaeus). C. Phallus, N. cimicoides. 0. Spermatheca, N. cimicoides (B-D from Larsen, 1938). don and Cheirochela Hope in the Old World tropics live in torrential streams, situations where it would be difficult to come to the surface at regular intervals, suggesting a lifestyle befitting their plastron respiration. Distribution and faunistics. Although the group oc¬ curs worldwide, the greatest diversity is conspicuously in the tropics. La Rivers (1971) treated some members of the New Guinea fauna and compiled a world catalog, with corrections published by La Rivers (1974, 1976). 37 Aphelocheiri(jae General. Aphelocheirids have the general appearance of many naucorids (Fig. 37.1 A, B), with their ovoid flattened bodies, but they differ in some characters, par¬ ticularly the relatively long labium and antennae. They range from 3.5 to 11.5 mm in length. Diagnosis. Head produced anteriorly, inserted into and surrounded posterolaterally by pronotum (Fig. 37.1 A, B); antennae elongate, slender, 4-segmented, fili¬ form; labium relatively long, reaching well onto meta¬ sternum, segment 2 very short, segment 3 very long. segment 4 less than one-half length of segment 3 (Fig. 37.1C); alary polymorphism common (Fig. 37.1 A, B); all tarsi 3-segmented; all legs with a pair of equally devel¬ oped claws; middle leg pretarsus as in Fig. 29.2F; meta- thoracic scent glands lacking, dorsal abdominal scent glands persistent in adults; posterolateral angles of con- nexiva often produced and spinelike (Fig. 37.lA, B); abdomen with simple dorsal and ventral plates; venter of thorax and abdomen with a plastron; abdominal spiracles 2-7 uniquely in Nepomorpha surrounded by rosettes (see Larsen, 1938:126; Thorpe and Crisp, 1947); abdomi¬ nal sternum 2 with a pair of sublateral sense organs of unknown function; aedeagus as in Fig. 37. ID; female subgenital plate as in Fig. 37. IE; spermatheca as in Fig. 37.1F, G. Classification. This group was first treated at family rank by Fieber (1851) as Aphelochirae, within the Nauco- roidea. It has been treated as a subfamily of Naucoridae (D. A. Polhemus and J. T. Polhemus, 1988), or a distinct family (Stys and Jansson, 1988), by most subsequent au¬ thors. Comparison of the arguments of Hoberlandt and Stys (1979) and Mahner (1993) with the conclusions of Stys and Jansson (1988) and D. A. Polhemus and J. T. Polhemus (1988) indicates that there is still little agreement about the ranking and relationships of the Aphelocheiridae, Naucoridae, and Potamocoridae. The cladogram of Rieger (Fig. 29.3) indicates that the Nauco- roidea of Stys and Jansson (1988) is not a monophyletic group, whereas that of Mahner (1993; Fig. 29.4) suggests otherwise but leaves the Potamocoridae unplaced. 126 TRUE BUGS OF THE WORLD (HEMIPTERA; HETEROPTEHA) Fig. 37.1. Aphelocheiridae. A. Male habitus, Aphelocheirus malayensis Polhemus and Polhemus. B. Female habitus, A. malayensis (A, B from D. A. Polhemus and J. T. Polhemus, 1988). C. Lateral view, head, A. aestivalis (Fabricius) (from Parsons, 1969). D. Phallus, A. malayensis. E. Female subgenital plate, A. malayensis (D, E from D. A. Polhemus and J. T. Polhemus, 1988). F. Spermatheca, A. aestivalis (from Larsen, 1938). Only the genus Aphelocheirus Westwood, with two subgenera and approximately 55 species, is currently rec¬ ognized. Specialized morphology. The Aphelocheiridae have relatively long slender antennae and a long labium, at¬ tributes not encountered in most other Nepomorpha. All species have a plastron, and they uniquely possess “ros- settes” surrounding the ventral abdominal spiracles (Lar¬ sen, 1938) as well as pressure receptors (Thorpe and Crisp, 1947; Parsons and Hewson, 1975). Natural history. Aphelocheirids inhabit the benthos of streams and lakes. Some species tolerate rather cold tem¬ peratures, and some are capable of living several meters below the water surface. Respiration via a plastron en¬ ables them to extract oxygen directly from the water and remain submerged for their entire lives (D. A. Polhemus and J. T. Polhemus, 1988). Distribution and faunistics. This is essentially a Paleotropical group, extending from Africa in the west to Northern Queensland in the east, with several species in Madagascar. As with most such distributions, which are remarkably common in the Heteroptera, a few species occur in the Palearctic. Particularly useful references are by Kanyukova (1974), Hoberlandt and §tys (1979), and D. A. Polhemus and J. T. Polhemus (1988). Notonectoidea 38 Notonectidae General. The backswimmers are all elongate and fusi¬ form (Fig. 38.1). Superficially they resemble members of the Corixidae because of the oarlike hind legs. Back- swimmers are relatively large among the Notonectoidea, ranging in length from 5 to 15 mm. Notonectidae 127 Diagnosis. Dorsum strongly convex, venter concave, abdomen with a median keel; compound eyes large; an¬ tennae 3- or 4-segmented, partially concealed between head and prothorax (Fig. 38.1); labium 4-segmented, short; membrane of hemelytra without veins, subdivided in 2 parts disposed in a tentlike fashion; fore- and middle legs adapted for grasping, hind legs oarlike, hind tibia and tarsus fringed with long setae (Fig. 38.1); tarsi of fore- and middle legs usually apparently 2-segmented, first segment always greatly reduced, sometimes absent; hind tarsi always 2-segmented; all pretarsi with 2 claws, those of hind legs reduced; abdomen with a median longitudi¬ nal keel and heavy fringes of setae medially and laterally forming air chambers; abdominal spiracles 2-8 located lateroventrally on abdomen, within the ventral air cham¬ ber; thoracic and first abdominal spiracles elongate and slitlike; abdominal sternum 5 strongly produced anteri¬ orly at midline, sternum 4 very narrow at midline; male genital segments symmetrical or nearly so; aedeagus as in Fig. 38.2D (see also Truxal, 1952); spermatheca ver¬ miform, as in Fig. 38.2F, G; ovipositor present, varying in degree of development (Fig. 38.2E). Classification. The Notonectidae were first recog¬ nized as a family group under the name Notonectariae by Latreille (1802), and until 1928 often were treated as including also the Pleidae (and Helotrephidae). Two subfamilies are recognized (Hungerford, 1933), comprising 11 genera and 343 species. Rg. 38.1. Notonectidae. Martarega mexicana Truxal (from Menke, 1979a). Key to Subfamilies of Notonectidae 1. Claval commissure simple, without a sensory pit; antennae 4- or 3-segmented; male parameres symmetrical or asymmetrical . Notonectinae - Claval commissure with a large, proximal, sharply delimited, seta-lined, sensory pit; antennae 2- or 3-segmented; male parameres usually asymmetrical .. Anisopinae ANISOPINAE. Claval commissure basally with a pit; male foretibia usually {Anisops, Buenoa) with a stridulatory comb (plectrum) located proximally on interior surface (Fig. 38.2C), labium with an apposing “prong” (Fig. 38.2B); metathoracic scent glands absent. This group, which was first recognized as a tribe by Hutchinson (1929), contains four genera, most members of which are rather small: Anisops Spinola from the Old World, Buenoa Kirkaldy from the New World, Parani- sops Hale from Australia, and Walambianisops Lansbury from Australia. NOTONECTINAE (FIG. 38.1). Claval commissure without a pit basally; males lacking foreleg stridulatory structures; metathoracic scent glands present. Obviously none of the above characters are unique to the Notonectinae, and as noted by Mahner (1993), the monophyly of the group cannot be supported. Members of the seven included genera are usually somewhat larger than those of the Anisopinae. Notonecta Linnaeus, although worldwide in distribution, is most di¬ verse in the New World. Enithares Spinola occurs in the Old World. The remaining genera contain many fewer species. Specialized morphology. The Anisopinae have a con¬ spicuous stridulatory plectrum proximally on the tibia (Fig. 38.2C). In Anisops and Buenoa the structure is ap¬ posed to the rostral prong (Fig. 38.2C). On the forefemur in Buenoa the plectrum is apposed to coxal pegs. All Ani- 128 TRUE BUGS OF THE WORLD (HEMIPTERA; HETEROPTERA) Fig. 38.2. Notoneotidae, A. Lateral view, head, Enithares maai Lansbury (trom Lansbury, 1968). B. Lateral view head, Anisops megalops Lans- bury. C. Foreleg with detail of stridulatory structure, Anisops megalops (B, C from Lansbury, 1962). D. Male genitalia, A. megalops. E. Female genitalia, Enithares sp. (D, E from Lansbury, 1968). F. Spermatheca, Notonecta obllqua (from Pendergrast, 1957). G. Spermatheca, N. glauca Linnaeus (from Larsen, 1938). sopinae have a seta-lined pit located at the anterior end of the claval commissure. Hemoglobin was discovered in Buenoa by Hunger- ford (1922a) and also occurs in Anisops. Hemoglobin- containing cells are concentrated in abdominal segments 3-7. The hemoglobin functions in providing neutral buoy¬ ancy, allowing the insects to remain at a constant level in the water column (Bare, 1928; Miller, 1966). Natural history. One of the most obvious features of the Notonectidae (and other Notonectoidea) is their habit of swimming on their backs. The resonant quality of stridulation in Buenoa was studied by Wilcox (1975). References to habitat partitioning in temperate-latitude species are given in Chapter 5. Distribution and faunistics. The back-swimmers are worldwide in distribution and well represented in temper¬ ate as well as tropical regions. Keys to the world genera are given by Lansbury (1968). Notonecta was revised by Hungerford (1933), Oriental Enithares by Lansbury (1968), Anisops by Brooks (1951), Buenoa by Truxal (1953), and Paranisops by Lansbury (1964). 39 Pleidae General. Sometimes referred to as pygmy backswim- mers, members of this group range in length from 1.5 Pleidae 129 Fig. 39.1. Pleidae. Plea sp. (drawn by T, Nolan; from CSIRO, 1991). to 3.0 mm. They have the general appearance of tiny Notonectidae, but lack the oarlike hind legs (Fig. 39.1). Diagnosis. Small; body globular (Fig. 39.1), heavily punctured; head very broad and short, immobile rela¬ tive to thorax; frontoclypeus medially with a distinctive sensory organ (Cobben, 1978; Mahner, 1993); labium 4-segmented, short, with a small apical labellum as in Helotrephidae; antennae 3-segmented; scutellum rela¬ tively large; forewings elytraceous, meeting along mid¬ line; hind wings developed or not; all legs apparently cursorial; fore- and middle tarsi 2- or 3-segmented, hind tarsi 3-segmented; all legs with 2 claws; nymphal dorsal abdominal scent gland located between terga 3/4; tho¬ racic venter and abdominal venter with a laminate keel on segments 2-5 or 2-6; male genitalia not strongly ro¬ tated as in Helotrephidae, and only weakly asymmetrical; aedeagus as in Fig. 39.2A; spermatheca as in Fig. 39.2B; ovipositor well developed. Classification. Fieber (1851) first recognized the group as a family Pleae, but the group was ranked as a subfamily of Notonectidae by most authors until Esaki and China (1928) argued for recognition of the group as a family. The world fauna includes three genera (Plea Fig. 39.2. Pleidae. A. Aedeagus, Neoplea striola (Fieber) (from Kumar, 1961). B. Spermatheca, Plea atomaria (Pallus) (from Pender- grast, 1957). Leach, Old 'World; Neoplea Esaki arid China, New World; and Paraplea Esaki and China, widely distributed) and 37 species. Specialized morphology. Although pleids are in gen¬ eral structure similar to the Notonectidae, they do not have the oarlike hind legs, but rather have all legs some¬ what homomorphous in form (Fig. 39.1), the hind tibia and tarsus usually not being fringed with long setae. Wefelscheid (1912) studied the anatomy and biology of Plea minutissima. Natural history. Like the Notonectidae, the Pleidae swim in the inverted position, using the legs in a row¬ ing fashion. They frequent vegetated areas in quiet-water habitats and are reported to feed on mosquito larvae, ostracods, and other small arthropods. The air store func¬ tions in a manner similar to that in the Notonectidae (Gittelman, 1974, 1975). Distribution and faunistics. Pleidae are broadly dis¬ tributed but show their greatest diversity in the tropics. The main works on the group are those of Esaki and China (1928) for the world and Drake and Chapman (1953) on the New World fauna. 40 Helotrephi(dae General. These small, globular to somewhat flattened bugs range in length from 1.0 to 4.0 mm. Their habitus (Fig. 40. lA) is similar to that of members of the family 130 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Pleidae, but they have many unique characteristics, most notable being the more complete fusion of the head and prothorax. They have no common name. Diagnosis. Body generally globular (Fig. 40.1 A); head and prothorax fused into cephalonotum, demar¬ cation indicated by a finely impressed line; compound eyes relatively small (Fig. 40.IB); antennae 1- or 2- segmented, the terminal segment furnished with numer¬ ous long setae (Fig. 40.1C); labrum reduced, nearly membranous; labium short, 4-segmented (Fig. 40.ID) with a small apical labellum; scutellum very large (Fig. 40. lA); forewings elytraceous, without distinct veins, slightly overlapping; hind wings often absent; tarsal for¬ mula variable from 3-3-3 to 1-1-2; all pretarsi with 2 equally developed claws, a pair of equally developed, short parempodia, and at least a ventral arolium in both nymphs and adults (Fig. 40. IE); venter of thorax and abdominal segments 2-5 or 2-6 with a laminate keel; nymphs with a single dorsal abdominal scent gland; male genitalia strongly asymmetrical (Fig. 40. IF), genital cap¬ sule rotated 90° to the left, left paramere lying ventrally, right paramere dorsally; aedeagus as in Fig. 40.IG, H; parameres as in Fig. 40.IK; spermatheca vermiform, as in Fig. 40. IJ; ovipositor present (Fig. 40.11). Classification. The genus Helotrephes Stal was placed in the Notonectidae by its author and subsequent workers, until Esaki and China (1927) recognized members of the group as belonging to a distinct family, primarily on the basis of the nearly complete fusion of the head and pro¬ thorax. Esaki and China (1927) recognized two subfami¬ lies to contain the three genera known at that time. Cur¬ rently this small family is divided into four subfamilies, which comprise 16 genera and 44 species (J. T. Polhe- mus, 1990c). Papacek, Stys, and Tonner (1988) provided characters for a cladistic analysis of relationships among the subfamilies. Keys to Subfamilies of Helotrephidae 1. Tarsal formula 3-3-3 or 2-2-3; scutellum wider than long . 2 - Tarsal formula 1-1-2; scutellum at least as long as wide . 3 2. Tarsal formula 3-3-3; cephalonotum small, posterior margin straight to moderately convex, lateral margins posterior to eyes insinuate; propleuron easily visible in lateral view; Neotropical . . Neotrephinae - Tarsal formula 2-2-3; cephalonotum extremely large, posterior margin strongly convex and broadly extending over proximal portion of scutellum and hemelytra, lateral margins posterior to eyes convex; propleuron concave, not visible in lateral view; Oriental . Trephotomasinae 3. Dorsum strongly convex, “pleoid”; lateral cephalic carina distinct; antennae 2-segmented; cepha- lonotal suleus complete, W-shaped, moderately postocular in position; scutellum usually moderately longer than wide, rarely about as long or strikingly longer, its sides convex; lateral postocular margins of cephalonotum usually convex . Helotrephinae — Dorsum rather flat, “naucoroid”; lateral cephalic carina indistinct; antennae 1-segmented; lateral sectors of cephalonotal sulcus absent, only medial, inversely U-shaped portion situated far behind eyes retained; scutellum strikingly long and narrow, sides straight; lateral postocular margins of cephalonotum insinuate, indented, or sharply constricted . Idiocorinae HELOTREPHINAE. Body globose; antennae 2-segmented; tarsal formula 1-1-2. This group was revised by Esaki and China (1928) and divided into two tribes by J. T. Polhemus (1990c). Cur¬ rently 11 genera, broadly distributed in the Paleotropics, are included. IDIOCORINAE. Body at least somewhat flattened; anten¬ nae 1-segmented; tarsal formula 1-1-2. This group was first recognized by Esaki and China (1927) to include the genera Idiocoris Esaki and China and Paskia Esaki and China from tropical Africa. For both genera, the authors provided exquisite illustrations that have seldom been matched for quality or detail in studies of the Nepomorpha. NEOTREPHINAE. Cephalonotum small; antennae 2-seg¬ mented in macropterous forms, 1-segmented in brachyp- terous specimens; tarsal formula 3-3-3. This group, which was first recognized by China (1940), contains the Neotropical genera Neotrephes China and Paratrephes China. TREPHOTOMASINAE. Cephalonotum very large; anten¬ nae 2-segmented; tarsal formula 2-2-3, This is the most recently described subfamily of Helo¬ trephidae and contains only the Oriental genus Trephoto- mas Papacek, Stys, and Tonner. Specialized morphology. The helotrephids are dis¬ tinguished primarily by the complete or nearly complete fusion of the head and the prothorax into a cephalonotum Helotrephidae 131 Fig. 40.1. Helotrephidae. A. Heterotrephes admorsus Esaki and Miyamoto. B. Lateral view, Helotrephes admorsus {A, B from Esaki and Miya¬ moto, 1959a). C. Antenna, Helotrephes bouvieri Kirkaldy. D. Labium, Helotrephes bouvieri (C, D from Esaki and China, 1928). E. Pretarsi, Idiocoris lithophilus Esaki and China. F. Male genitalia, /. lithophilus (E, F from Esaki and China, 1927). G. Aedeagus, Mixotrephes hoberlandti Papacek, Stys, and Tonner. H. Aedeagus, M. hoberlandti (G, H from Papacek et al., 1989). I. Female genitalia, I. lithophilus (from Esaki and China, 1927). J. Spermatheca, Neotrephes usingeri China (from Pendergrast, 1957). K. Paraineres, Helotrephes admorsus Esaki and Miyamoto (on left dorsal paramere, ventral view; on right, ventral paramere, ventral view) (from Esaki an.J Miyamoto, 1959a). (Fig. 40.1 A, B). The 1- or 2-segniented antennae, with the covering of elongate setae on the terminal segment are also distinctive (Fig. 40.1C). J. T. Polhemus (1990c) described the occurrence of a stridulatory mechanism in the Helotrephinae, consisting of a serrated lateral margin of the hemelytron plus a ridge dorsally on the hind femur. He also noted that other stridulatory structures might exist in the group (see Miyamoto, 1952). Natural history. Helotrephids live in a variety of habi¬ tats, ranging from the quiet, nearly stagnant waters of ponds to much better aerated backwaters in streams; some have been taken from hot springs. They apparently swim with the venter up, as do other Notonectoidea, or they may swim with the venter down and use only the hind legs for propulsion, in a rowing motion (Miyamoto, 1952). They feed on small invertebrates. 132 TRUE BUGS OF THE WORLD (HEMIPTERA:HETEROPTERA) Distribution and faunistics. This group is restricted Esaki and China (1927), China (1935, 1940), Esaki and- to the tropics and shows its greatest diversity in the Old Miyamoto (1959a), Papacek, Stys, and Tonner (1988), World. Although there is no comprehensive reference for and J. T. Polhemus (1990c). identification of species, important works include those of Helotrephidae 133 41 Leptopodomorpha General. Bugs in this group are of very small to mod¬ est size. They vary in shape from nearly globose (Fig. 44.lA), to weakly flattened and ovoid (Fig. 43.IB), to elongate and parallel-sided (Fig. 45. IB). All species are predatory. The majority inhabit damp areas adjacent to water, and a relatively large number are intertidal. Many members of the Leptopodidae have no apparent asso¬ ciation with water and as a consequence have habitats that are less easily characterized. Those species are more rarely collected. Diagnosis. Head usually relatively short and broad; compound eyes usually very large, occupying nearly en¬ tire side of head (Figs. 43.1 A, 45.1 A, B); antennae in most groups with all segments of nearly equal diameter, segment 1 short, segments 2-4 longer and of more or less equal diameter, segment length and diameter variable in the Leptopodinae; labium inserted ventrally, short (Fig. 45.2B) or long (Fig. 43.2A), with 3 or 4 obvious seg¬ ments; ocelli usually present; head with at least 3 pairs of trichobothria dorsally; forewings in the form of hemely- tra with a conspicuously coriaceous anterior portion and membranous posterior region, often with a distinct me¬ dial fracture combining with costal fracture (Fig. 43.2E) and attaining corial margin in macropterous forms (except Leptopodinae), membrane usually with 3, 4, or 5 closed cells in macropterous forms (Figs. 43.1 A, B, 45. lA, B); legs usually relatively short, slender, and mutic, or in Leptopodinae longer, more slender, and femora some¬ times armed with spines (Fig. 45.2A); adult tarsal for¬ mula usually 3-3-3, \-2-2 in Leotichius, nymphal formula usually 2-2-2; parempodia usually reduced (Fig. 41.2D), sometimes setiform (Fig. 41.2B), accessory parempodia present; small ventral arolium present in some nymphs, absent in most adults; metathoracic scent-gland system with 1-4 reservoirs and 1 or 2 ostioles; abdominal sterna composed of a single plate, dorsal laterotergites present or absent, 8 abdominal spiracles, usually located dorsally on laterotergite (Fig. 45.2D); copulation always in the side-by-side position (Fig. 43.2A), male holding fore¬ wing margin of female with grasping apparatus present between laterotergites of abdominal segments 2 and 3, pegs present on this structure in Jialdidae (Fig. 41.2,!), absent in all other groups; nympiu.; dorsal abdominal scent-gland opening present between abdominal terga 3/ 4; male genitalia symmetrical, parameres either hook¬ shaped with a distinct processus sensualis or club-shaped (Figs. 44.IE, 45.2E); ovipositor valvulae laciniate, or reduced and platelike; spermatheca with bulb and flange (Figs. 43.21, 45.2G,H). Discussion. The overall composition of the Lepto¬ podomorpha has been generally agreed upon by most modem authors, although categorical rank and phylo¬ genetic relationships of several taxa have been debated. Following the classification of Schuh and Polhemus (1980b), we recognize four families—Aepophilidae, Sal- didae, Omaniidae, and Leptopodidae (including Lepto- saldinae and Leotichius Distant); their cladogram is shown in Figure 41.1. In contrast Cobben (1971) grouped the Leptopodomorpha as follows: Saldidae (including Aepo¬ philidae, Leptosaldinae), Omaniidae, Leptopodidae, and Leotichiidae. Popov (1985) transferred the Leptosaldinae to the Omaniidae as a subfamily. The catalog of Schuh et al. (1987) is a comprehensive reference to the classifi¬ cation and literature on the group. Fig. 41.1. Phylogenetic relationships of family-level taxa of Leptopodo¬ morpha (from Schuh and Polhemus, 1980b). 134 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 41.2. Leptopodomorphan morphology. A. Foreleg pretarsus, nymph, with dorsal arolium, Saldula pallipes (Saldidae). B. Hind leg pretarsus, nymph, S. pallipes (Saldidae), C. Foreleg pretarsus, adult, with dorsal arolium and reduced parempodia, S. pallipes (Saldidae). D. Adult pre¬ tarsus, Omania coleoptrata (Omaniidae). E. Foreleg pretarsus, nymph, Valleriola sp. (Leptopodidae), F. Hind leg pretarsus, adult, Leotichius Shiva (Leptopodidae). G. Metacoxal adhesive pads, O. coleoptrata (Omaniidae). H. Same as G, detail, I. Larval organ on abdominal sternum 3, S. pallipes (Saldidae). J. Male abdominal grasping apparatus, pegs on laterotergite of abdominal segment 3, S. pallipes (Saldidae). Abbreviations: aga, abdominal grasping apparatus; da, dorsal arolium; lo, larval organ; It 2, laterotergite 2; It 3, laterotergite 3; pe, parempodium; sp, spiracle; ut, unguitractor plate. Leptopodomorpha 135 Key to Families of Leptopodomorpha 1. Labium tapering, long, reaching to base of hind coxae orbeyond (Fig. 43.2A) . 2 - Labium much shorter, reaching at most to apex of forecoxae, often reaching only base of forecoxae (Fig. 45.2B) . ,. Leptopodidae 2. Compound eyes large, covering most of head in lateral view; forewings covering abdomen, macrop- terous, submacropterous, orcoleopteroid . 3 - Compound eyes small (Fig. 42.2), covering a small portion of head in lateral view; wings greatly reduced, in form of pads without obvious venation, covering only anterior portion of abdomen (Fig. 42.1) . Aepophilidae 3. Compound eyes reaching posteriorly only to level of pronotal collar or very slightly beyond (Fig. 43.1 A, B); macropterous or submacropterous, seldom coleopteroid; body length always greater than 2.2 mm . Saldidae - Compound eyes reaching posteriorly about one-third length of pronotum, distinctly surpassing pro¬ notal collar (Fig. 44.1 A); always coleopteroid; body length always less than 2 mm .... Omaniidae j Saldoidea 42 Aepophilidae General. The Aepophilidae comprises Aepophilus bonmirei Signoret, sometimes referred to as the marine bug. This intertidal creature, slightly more than 2 mm long, has the general appearance of a tiny bed bug (Fig. 42.1). Diagnosis. Respiratory plastron covering pronotum, scutellum, and hemelytra; compound eyes very small, with approximately 50 ommatidia (Figs. 42.1, 42.2A); ocelli possibly represented by minute remnants; head pro¬ jecting anteriorly, clypeus visible from above; labium long, segment 1 greatly abbreviated, segment 2 short, segments 3 and 4 subequal in length, each much longer than segment 2; forewings padlike (Fig. 42.1), hind wings absent; as in Saldidae, adult and nymphal pretarsus with, “accessory parempodia” (Cobben, 1978) and rudimen¬ tary parempodia, ventral arolium present in adult and nymphal stages (Schuh and Polhemus, 1980b); meta- thoracic scent gland with 2 reservoirs and 2 ostioles (Cobben, 1970); abdomen with both dorsal and ventral Rg. 42.1. Aepophilidae. Aepophilus bonnairei Signoret. 136 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) A Fig. 42.2. Aepophilidae. Aepophilus bonnairei. A. Head (from Schuh and Polhemus, 1980b). B. Testes (from Pendergrast, 1957). laterotergites; spiracles located on simple sterna; nymphal abdominal scent glands with 2 reservoirs and 2 ostioles; abdominal grasping apparatus present, but lacking pegs found in Saldidae; eversible glands and nymphal organ found in Saldidae absent; phallus similar to Saldidae but lacking reel system; parameres hook-shaped with a dis¬ tinct processus sensualis; testes as in Fig. 42.2B; oviposi¬ tor valvulae laciniate, serrate. Classification. Aepophilus has often been treated as a member of the Saldidae (e.g., Leston, 1957a; Cobben, 1959), but most work suggests that it is the sister group of all remaining Saldoidea. We therefore follow Schuh and Polhemus (1980b) in giving it family status. Specialized morphology. Early investigators were not certain by what mechanism Aepophilus was able to acquire oxygen when submerged. Most assumed that an air bubble was trapped in the rock crevices in which the bugs secreted themselves. SEM examination re\'eals, however, that the surface of the pronotum, scutellum. and hemelytra is densely covered with fine, branching micro- trichia, which have been observed to maintain a constant air covering and which apparently serve as a physical gill in the form of a plastron (King and Ratcliffe, 1970; King and Fordy, 1984), allowing the bugs to remain submerged for prolonged periods. Natural history. Aepophilus bonnairei lives in rock crevices in the lowest reaches of the intertidal zone—the so-called Fucus zone—where it feeds on small inverte¬ brates (e.g., Baudoin, 1946). Distribution and faunistics. Aepophilus is known from the coast of Ireland, the Low Countries, north¬ ern France, southwestern England, the Channel Islands, Spain, Portugal, and possibly Morocco. This area is not known for harboring phylogenetic relicts, placing Aepo¬ philus in something of a unique position. 43 Sal(jidae General. Shore bugs, which range in length from 2.3 to 7.4 mm, are usually ovoid in outline (Fig. 43.1 A, B). As the family name suggests, they can jump, and in fact most are extremely agile through a combination of jump¬ ing and flight. Many species vary greatly in the extent of hemelytral pigmentation, and therefore identification is often difficult. Diagnosis. Compound eyes large, reniform, posterior margin usually reaching, but never surpassing, anterior margin of pronotum (Fig. 43.1 A, B); 3 pairs of cephalic trichobothria; labium long, tapering, nearly reaching onto abdomen, segment 1 greatly reduced, segment 3 much longer than either segment 2 or 4 (Fig. 43.2A); me¬ dial fracture in forewing well developed and in combi¬ nation with costal fracture attaining costal margin (Fig. 43-2B); 4 or 5 closed cells in the membrane (Fig. 43.1 A, B); subcostal region of the forewing in female modi¬ fied to accommodate abdominal grasping apparatus of male; parempodia developed and setiform in nymphs. Saldidae 137 Fig. 43.1. Saldidae. A. Pentacora signoreti (Guerin-MeneviHe) (Chilo- xanthinae) (from Usinger, 1956). B. Pseudosaldula chilenis (Blan¬ chard) (Saldinae) (from Drake, 1962). greatly reduced and rudimentary in adults, ventral aro- lium present in nymphs (Schuh and Polhemus, 1980b, contra Cobben, 1978), present on some legs in adults of at least some taxa (Fig. 41.2A-C); metathoracic scent gland with 1 reservoir and 1 ostiole; abdomen with dor¬ sal and ventral laterotergites, spiracles located on sterna; nymphs of most Saldidae (except Enalosalda Polhemus and Evans, Orthophrys Horvath, and all Saldini) with “larval organ” on either side of midline of sternum 3 near spiracle (Figs. 41.21, 43.2J-M) (Cobben, 1957); adults with eversible glands located laterally on abdomen between segments 7 and 8; parandria in males well de¬ veloped and sclerotized (Fig- 43.2F); parameres hooked and with a conspicuous processus sensualis (Fig. 43.2E); phallus with a reel system (Fig. 43.2C, D); spermatheca as in Fig. 43.21; ovipositor valvulae well developed and serrate. Classification. The major classical worker on the Sal¬ didae was O. M. Reuter, who described many species and published a catalog of the Palearctic fauna (Reuter, 1895). C. J. Drake described many species from the New World (e.g., 1949a, b, 1955) and published some of the first illustrations of the male parameres. More recent work has been dominated by the contributions of the late R. H. Cobben on the morphology (Cobben, 1957, 1968a) and higher classification of the group (Cobben, 1959). Nomenclature in the Saldidae was confused well into the twentieth century because of differences of opinion regarding the type genus of the family. Reuter consis¬ tently considered Acanrfti'a Fabricius to be the type, while most other authors (e.g., Lethierry and Severin, 1896) believed Saida Fabricius was the type. The current classification is primarily the work of Cob¬ ben (1959) and is based heavily on the structure of the male genitalia. Two subfamilies comprising 26 genera and 265 species are recognized. 138 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Keys to Subfamilies of Saldidae 1. Hemelytra with long medial fracture reaching at least to level of posterior end of claval suture (fracture lost in Enalosalda): female sternum 7 truncate, usually square, or if produced caudad along midline (Fig. 43.2G); membrane with five cells (Fig. 43.1 A) . Chiloxanthinae - Hemelytra with short medial fracture, not reaching anteriorly more than half the distance from costal fracture to posterior end of claval suture (fracture lost in Orthophiys)', female sternum 7 produced caudad along midline (Fig. 43.2H); membrane usually with 4 cells (5 cells in Pseudosaldula) . . Saldinae CHILOXANTHINAE (FIG. 43.1 A). Membrane with 5 closed cells (Fig. 43.2B); base of penis filum forming at most 1 coil; ductus ejaculatorius with 2 ventral accessory glands; median sclerotized structure of penis paired; apicolateral sclerotized stmetures of penis absent; sternum 7 in female square. This group of 22 Recent and four fossil species placed in four Recent and two fossil genera is largely restricted to the Northern Hemisphere and is best known for its as¬ sociation with very high latitudes {Chiloxanthus Reuter), saline habitats (Pentacora Reuter), or intertidal habitats {Enalosalda Polhemus and Paralosalda Polhemus and Evans). Pentacora signoreti Guerin-Meneville is probably the largest species of saldid, measuring up to 7.4 mm in length. SALDINAE (FIG. 43.1 B). Membrane with 4 closed cells (ex¬ cept Pseudosaldula Cobben with 5; Fig. 43. IB); base of penis filum coiled like a watch spring (Fig. 43.2D); me¬ dian sclerotized structure of penis unpaired; apicolateral sclerotized structure of penis present; mesal portion of sternum 7 in female produced caudad (Fig. 43.2H). Three tribes are currently recognized. Most species of Saldini are large, often exceeding 5 mm in length; they are black, with few if any markings, and often are brachypterous. Species of Teloleuca Reuter have varie¬ gated markings and are usually macropterous. Saldini occur at moderately high latitudes in the Northern Hemi¬ sphere in wet freshwater environments, although some species—for example, Saida littoralis (Linnaeus)—occa¬ sionally may be found in salt marsh or other saline situa¬ tions. The Saldoidini are a group of worldwide distribution, with the greatest numbers of species in temperate regions and a lesser number in lowland tropical habitats. Most Saldoidini are associated with fresh water, although Halo- salda Reuter contains only halophilous species; a few, such as Saldula laticolUs (Reuter) and Orthophiys pyg- maeum (Reuter) are intertidal. The genus Saldula Van Duzee contains a relatively high proportion of all de- Keys to subfamilies of Saldidae adapted from J. T. Polhemus, 1985. scribed species of shore bugs, as well as some of the most complex taxonomic problems in the family; Lindskog and Polhemus (1992) noted that the group is apparently para- phyletic. Saldoida Osborn is remarkable among the Saldi¬ dae for the pronotal calli produced into conical tubercles and for antlike behavior. The Saldunculini contain only the Indo-Pacific inter¬ tidal genus Salduncula Brown, whose seven described species were keyed by J. T. Polhemus (1991b). Specialized morphology. The well-developed peg plate of the abdominal grasping apparatus in the male is one of the unique features of the Saldidae (Fig. 41.2J). This structure was first described by Drake and Hottes (1951) as stridulatory in function. Cobben (1957) and Leston (1957b) correctly observed that is was actually used during copulation, the male abdomen grasping the anterolateral margin of the female hemelytron. Actual stridulatory structures occur in the genera Chartoscirta Stal, loscytus Reuter, Lximpracanthia Reuter, Macrosal- dula Leston and Southwood, Rupisalda Polhemus, and Saldoida Osborn, all members of the Saldinae. In all cases the stridulitrum is located on the anterior margin of the forewing, and the plectrum is located on the hind femur (J. T. Polhemus, 1985). The larval organ, of unknown function, is located on abdominal sternum 3 (Figs. 41.21, 43.2J-M). The ever- sible glands, also of unknown function, are found be¬ tween the laterotergites of abdominal segments 7 and 8. Both structures are unique to the saldids, although the former is absent in some taxa. Sperm ultrastructure has been studied in two species by Afzelius et al. (1985). Naturtil history. Many saldids are extremely agile, moving through a combination of jumping and flight. All are predators of small invertebrates, seeking out live food or scavenging on dead individuals. They can be maintained in captivity on immobilized flies or other small arthropods and have been shown to locate subsur¬ face prey through antenna-based chemoreception. Sur¬ face prey may be located visually as well as by chemo¬ reception (J. T. Polhemus, 1985). Most members of the Saldini appear to overwinter as Saldidae 139 Fig. 43.2. Saldidae. A. Mating position, Saldidae (from Schuh and Polhemus, 1980b). B. Forewing, Pentacora grossi Cobben (from Cobben, 1980b). C. Phallus, dorsal view, Saldula laticollis (Reuter). D. Phallus, lateral view, S. laticollis (C, D from J. T. Polhemus, 1985). E. Paramere, S. sibiricola Cobben (from Cobben, 1985). F. Parandria, S. dentulata (Hodgden). G. Subgenital plate, P. signoreti. H. Subgenital plate, S. dentulata (F-H from Cobben, 1960d). I. Gynatrial complex, S. sibiricola (from Cobben, 1985). J. Larval organ, Chartoscirta cocksii (Curtis). K. Larval organ, C. cocksii. L. Larval organ, C. cocksii. M. Larval organ, S. tucicola (Sahiberg) (J-M from Cobben, 1957). Abbreviation: of, costal fracture. 140 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) eggs and produce only a single generation each year, whereas, in the Northern Hemisphere at least, most mem¬ bers of the Saldoidini probably overwinter as adults, some producing multiple generations in a given season. The life cycles of tropical species are poorly known. Most temperate-region shore bugs are associated with damp substrates along the margins of ponds and streams and occasionally along the margins of larger bodies of water. Some species practice a truly terrestrial existence, as for example members of the Saldula orthochila species group and some members of the Saldini. The majority of tropical saldids are saxicolous, usually living in asso¬ ciation with large stones, a habitat peculiar to only a few temperate-latitude species. There is a significant high- altitude fauna in the Andes of South America; most of the known species are associated with seeps or pond margins and a few occur on rocks in streams. The eggs of most species appear to be laid in vegetation with the laciniate ovipositor. Intertidality, one of the most conspicuous adaptations in the shore bugs, occurs in nearly every phyletic line (Schuh and Polhemus, 1980b; J. T. Polhemus, 1985) as well as in the related families Aepophilidae and Omani- idae. Life history studies of the intertidal Saldula laticollis in western North America indicated that it can remain submerged for substantial periods (Stock and Lattin, 1976) without the use of a plastron. Distribution and faunistics. Saldids are known from all major land areas except Antarctica, including remote islands such as St. Helena (one species; Cobben, 1976) and the Hawaiian Islands (eight species; Gobben, 1980a). A preponderance of the described species occurs in the Northern Hemisphere, but only with additional knowl¬ edge of phylogenetic relationships within the group at the species level will a more mature understanding of the distributional history of the group come into focus. Regional treatments exist for the faunas of the Palearc- tic (Cobben, 1960a, 1985; Kerzhner and Jaczewski, 1964; Pericart, 1990); the Nearctic (e.g., Usinger, 1956; Chap¬ man, 1962; Brooks and Kelton, 1967; Schuh, 1967; J. T. Polhemus and Chapman, 1979a), and Middle America (J. T. Polhemus, 1985). Cobben (1987b) provided an annotated checklist of the African species (see also Cob¬ ben, 1987a; J. T. Polhemus, 1981). Only scattered papers are available for South America, Australia, and New Zea¬ land. Leptopodoidea 44 Omaniiijae General. The Omaniidae, or intertidal dwarf bugs, are the smallest members of the Leptopodomorpha, ranging in length from 1.15 to 1.59 mm. They are similar in size and appearance (Fig. 44.1 A) to many Schizopteridae. Diagnosis. Eyes very large, projecting posteriorly along anterolateral margins of pronotum (Fig. 44.1 A); vertex and frons with 4 pairs of trichobothria; labium long, with 4 obvious segments, 1 and 2 short, 4 longer, 3 distinctly longer than 4 (Fig. 44.IB); coleopteroid, hemelytra meeting along midline in the form of elytra, no medial fracture, membrane absent; as in most Saldi- dae, parempodia well developed and setiform in nymphs, greatly reduced and budlike in adults (Fig. 41.2D); exis¬ tence of arolium in nymphs unconfirmed; hind coxae with adhesive pads (Figs. 41.2G, 44.1C); metathoracic scent glands with 4 reservoirs in 2 pairs and a single ostiole; dorsal (inner) laterotergites absent; abdominal spiracles dorsal, situated in membrane adjacent to outer latero- tergite; abdominal grasping apparatus present between laterotergites of abdominal segments 2 and 3, but without pegs; aedeagus as in Fig. 44. ID; parameres club-shaped (Fig. 44. IE); female gynatrial complex and spermatheca as in Fig. 44. IF; ovipositor reduced and platelike. Classification. Omania Horvath, with a single species from the Red Sea and Gulf of Oman was originally placed in the Saldidae. Cobben (1970) elevated the group to family status, placing four species in a new genus, Coral- locoris Cobben, which is distributed from Aldabra Atoll off the coast of Africa east to Samoa and Kwajelein Atoll. Specialized morphology. Although schizopterid in general appearance and size, the leptopodomorphan af¬ finities of this group have been recognized by all workers. The Omaniidae are notable among the Leptopodomor¬ pha for the coleopteroid hemelytra and the presence of adhesive pads on the hind coxae. Natural history. Although the Omaniidae are all inter¬ tidal, they—like many Saldidae—have no obvious mor¬ phological specializations for this mode of life (see Cob¬ ben, 1970). They apparently feed on small organisms in the intertidal zone and can be found crawling around on rocks at low tide. They secrete themselves in crevices during high tide, apparently using a trapped air bubble Omaniidae 141 Fig. 44.1. Omaniidae. A. Habitus, Corallocoris nauruensis Herring and Chapman. B. Frontal view, head, C. nauruensis (A, B from Schuh and Polhemus, 1980b). C. Hind ieg, Omania coleoptrata Horvath. D. Phalius, O. coleoptrata (C, D from Cobben, 1970). E. Paramere, C. nauruen¬ sis (from Schuh and Poihemus, 1980b). F. Gynatrial complex, including spermatheca, O. coleoptrata (from Cobben, 1970). Abbreviation; ap, adhesive pad. as a source of oxygen. The eggs, which are very large relative to the size of the animal, apparently develop one at a time, and a single egg is laid every few days (Kellen, 1960). Distribution and faunistics. The basic reference for this Indo-Pacific group is Cobben (1970). 45 Leptopodi(jae Generai. This primarily tropical group has no com¬ mon name. Its members are very active and agile. Lepto- podids range from 1.8 to 7.0 mm in length and are of variable habitus (Figs. 45.lA, B, 45.2A), but they are distinctive for the relatively short, 4-segmented labium and deeply punctured dorsum. 142 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Diagnosis. Compound eyes of varied shape and ocelli of varied position; labium 4-segmented, short, at most reaching to apex of forecoxae (Fig. 45.2B); at least cla- vus, and sometimes most of dorsum, deeply and densely punctured (Figs. 45.1 A, 45.2A); parempodia present and setiform in nymphs (Fig. 41.2E), greatly reduced or absent in adults (Fig. 41.2F); arolia absent in those taxa examined; abdominal grasping apparatus present but without pegs; parameres club shaped (Fig. 45.2E); ovi¬ positor platelike (Fig. 45.2D, F); spermatheca with bulb and flange (Fig. 45.2G, H). Classification. The earliest comprehensive work on the Leptopodidae was that of Horvath (1911). The clas¬ sification of Schuh and Polhemus (1980b) presented here recognized two subfamilies, some members of which were placed in other families by prior authors. Ten genera and 37 species are included. Key to Subfamilies of Leptopodidae 1. Ocelli proximate and situated on a low prominence (Figs. 45.IB, 45.2A); compound eyes hemi¬ spherical, with ommatidia on dorsal surface of eyes or not . Leptopodinae - Ocelli widely separated, not situated on a prominence (Fig. 45.1 A); compound eyes distinctly reniform, ommatidia always present on dorsal surface (Fig. 45.1 A) . Leptosaldinae LEPTOSALDiNAE (FIG. 45.1 A). Habitus saldidlike; eyes large, reniform, extending well back onto lateral pro- notal margins (similar to Omaniidae); 3 cells in mem¬ brane; body flattened dorsoventrally; appendages reia- •tively short; spiracles located dorsally on laterotergites. Leptosalda chiapensis Coboen is known only as an amber fossil from southern Mexico (Cobben, 1971). The other member of the subfamily, Saldolepta kistnerorum Schuh and Polhemus, is known from two Recent speci¬ mens, one taken from a termite nest in the tropical low- Leptopodidae 143 Fig. 45.2. Leptopodidae. A. Patapius spinosus (Rossi) (from Schuh and Polhemus, 1980b) B. Ventral view, Saldolepta kistnerorum (from Schuh and Polhemus, 1980a). C. Forewing, Leptosalda chiapensis Cobben (from Schuh and F'olhemus, 1980b). D. Dorsal view, abdomen, female S. kistnerorum (from Schuh and Polhemus, 1980a). E. Paramere, Erianotus lanosus (Duiour) (from Schuh and Polhemus, 1980b). F. Female genitalia, P. thaiensis Cobben (from Cobben, 1968b). G. Spermatheca, Leptopus marmoratus (Goeze) (from Pendergrast, 1957). H. Gynatrial complex, including spermatheca, P. thaiensis (from Cobben, 1968b). Abbreviation: sp, spiracle. lands of western Ecuador (Schuh and Polhemus, 1980a), the other recovered by port inspectors on plant material from Colombia. LEPTOPODINAE (FIGS. 45.1 B, 4S.2A). Eyes protruding, nearly hemispherical; ocelli proximate to one another, situated on low prominence; costal fracture absent; fore¬ femora and often dorsum with very long setae or heavy cuticular spines; tarsi elongate, slender; spiracles located dorsally on laterotergites in Leptopodini, ventrally on abdominal sternum in Leotichiini. Two tribes are recognized, with all members occurring in the Old World, primarily in the tropics. The Leotichi- 144 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) ini, with three species of Leotichius Distant, range from Burma to Bali. The Leptopodini comprises seven genera. Valleriola Distant includes 11 species, most living on vertical rock surfaces around the shores of ponds and streams, some¬ times being very common. Genera such as Erianotus Fie- ber, Leptopus Latreille, Martiniola Horvath, and Patapius Horvath, each with only a few species, are often found under stones in screes or in other dry situations some dis¬ tance from water. The group is distributed in the southern Palearctic, throughout Africa and Madagascar, and east to New Caledonia. Specialized morphology. The leptopodids, and par¬ ticularly the Leptopodinae, are distinctive for their modi¬ fication of certain body ptu'ts. All have at least the clavus, and often much of the dorsum, heavily punctured, in con¬ trast to other Leptopodomorpha. The Leptopodinae have the ocelli situated close together on a low tubercle or prominence (Figs. 45.IB, 45.2A). In Patapius Horvath much of the body surface—including the compound eyes—is covered with spines (Fig. 45.2A); in Valleriola Distant the legs are very long and covered with a fringe of long setae (Fig. 45. IB); and in Leotichius the omma- tidia on the upper surface of the compound eyes are not developed. Pericart and Polhemus (1990) described a stridulatory structure involving abdominal tergum 1 and the vannus of the hind wing for several species of Leptopodinae. They' asserted that this structure represented a synapomorphy for the Leptopodinae and that it was absent in Leotichius either because that genus represents a distinct subfamily or family or because the structure had been lost. A simpler explanation interprets this structure as a synapomorphy of the Leptopodini sensu Schuh et al. (1987). Natural history. No simple description serves to cate¬ gorize the habitats of the Leptopodidae. All members are presumably predatory, although their feeding approach may differ from that of other leptopodomorphans in that they have a very short rostrum. Leotichius spp. were long thought to inhabit caves, but more recent collections suggest that they are simply denizens of extremely dry tropical habitats. The most ex¬ tensive collections have been made from an area of ant lion pits under the overhanging roofs of Hindu temples on the island of Bali (J. T. Polhemus and Schuh, 1995). Distribution and faunistics. The Leptopodidae were long known only from the Old World, until the two New World species, Leptosalda and Saldolepta —now included in the Leptosaldinae—were described. Patapius spinosus Rossi has been introduced near San Francisco, California, in North America. Most species live within the tropical latitudes. Horvath (1911) provided the only comprehen¬ sive treatment for the Leptopodini; J. T. Polhemus and D. A. Polhemus (1991c) provided a key to most genera. Leptopodidae 145 46 Cimicomorpha General. Among the 16 families currently placed in the Cimicomorpha are the two largest families of true bugs, the Miridae and Reduviidae, as well as two of the most obscure, the monotypic Joppeicidae and Medocosti- dae. Although it can be argued that the primitive cimico- morphan stock was predaceous, the group also contains large numbers of strictly phytophagous species in the families Miridae, Tingidae, and Thaumastocoridae. The most important classical authors working in the group were Fieber on the fauna of the western Palearctic, Stal on the Nabidae and Reduviidae, and Reuter on the Anthocoridae, Microphysidae, and particularly Miridae. Modem authors have generally become family special¬ ists, because of the sheer numbers of taxa involved. Diagnosis. Head often prognathous, with labium in¬ serted anteriorly, and clypeus lying more or less dor- sally and in a nearly horizontal position, or head not so strongly oriented anteriorly, labium inserted antero- ventrally, clypeus positioned anteriorly and more or less vertical in orientation; head often with cephalic tricho- bothria; antennae often flagelliform; labium with 3 or 4 segments; forewings in macropterous forms always in the form of a hemelytron, with a coriaceous anterior region and a membranous posterior region, often with a long medial fracture and a costal fracture; membrane often with 1, 2, or 3 closed cells, or with a few free veins, or no veins, or veins emanating from closed cells; hind wings usually with a simple, nonbranching distal sec¬ tor of R-f M; tarsi usually with 3 segments, sometimes 2; pretarsus symmetrically developed in most families, lacking arolia in all life stages; claws simple and with¬ out pul villi or other ornamentation (except many Miridae and some Thaumastocoridae and Anthocoridae); meta- thoracic scent-gland evaporatory area usually with dis¬ tinctive mushroom bodies (Fig. 10. lOA); spiracle 1 some¬ times present, remaining spiracles always ventral, located either on distinct laterosternite or on mediostemal plate with laterotergite present or not; spermatheca nonfunc¬ tional as a sperm storage organ, being rudimentary, modi¬ fied into a vermiform gland, or absent; eggs usually with micropyles distinct from aeropyles and both arranged in a ring outside the operculum. Discussion. First recognized by Leston et al. (1954), the Cimicomorpha have been conceived differently by various authors. Cobben (1968a, 1978), who had serious doubts about the placement of the Reduviidae and Thau¬ mastocoridae within the group, consistently associated the Pachynomidae with the Nabidae rather than with the Reduviidae, as argued by Carayon (1950b), Davis (1969), and Carayon and Villiers (1968). We follow the classifica¬ tion presented by Schuh and §tys (1991). Their cladogram is shown in Figure 46.1. Fig. 46.1. Phylogenetic relationships o1 families of Cimicomorpha (after Schuh and Stys, 1991). Key to Families of Cimicomorpha 1. Mandibular plates greatly enlarged and conspicuous, usually exceeding and surrounding apex of clypeus (Figs. 52.1 A, 52.2A, 52.3A, B) . Thaumastocoridae — Mandibular plates not conspicuously enlarged, never obvious in dorsal view, never exceeding apex of clypeus . 2 146 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) 2. Labium conspicuously 4-segmented, inserted ventrally on head, segment 1 reaching posterior margin of head or nearly so (Figs. 53.3D, 54.3A); fossula spongiosa never present . 3 - Labium usually with 3 obvious segments, inserted anteriorly on head (Figs. 47.2A, 49. IB, C), if 4- segmented, segment 1 never approaching posterior margin of head: fossula spongiosa often present on 1 or more pairs of legs . 5 3. Pronotum and hemelytra areolate (Fig. 54.2B); hemelytra of nearly uniform texture throughout, without obvious corium-clavus and membrane; antennal segment 2 short (Figs. 54.2B. 54.3A); ocelli always absent; tarsi 2-segmented (Cantacaderinae, Tinginae) . Tingidae (part) - Pronotum and hemelytra never areolate, although sometimes heavily punctured; hemelytra usually with obvious corium, clavus, and membrane, although rarely coleopteroid; antennal segment 2 more elongate, usually much longer than segment 1; ocelli present or absent; tarsi 2- or 3-segmented 4. Macropterous or brachypterous, rarely coleopteroid; R+M of forewing never raised and keel-like; compound eyes always normally developed; trichobothria present on meso- and metafemora (Fig. 53.3E, F); male genitalia always asymmetrical; tarsi 2- or 3-segmented; ocelli present or absent . Miridae - Usually coleopteroid, heavily punctured, compound eyes greatly reduced and ocelli absent (Fig. 54.1), or, if macropterous, R-HM in forewing elevated and keel-like, compound eyes developed and ocelli present; trichobothria never present on meso- and metafemora; male genitalia symmetrical; tarsi 2-segmented (Vianaidinae) . Tingidae (part) 5. Prosternal sulcus present, usually in the form of a stridulitrum (Fig. 10.7F, G). receiving apex of labium; labium usually short, stout, and strongly curving (Fig. 48.3A, C), sometimes more slender and nearly straight (Fig. 48.3B); head necklike behind eyes (Fig. 48.2A-D), frequently with a trans¬ verse impression anterior to ocelli; membrane usually with 2 large cells (Fig. 48.2C) (sometimes more) or rarely with a few longitudinal veins (Fig. 48.2A) . Reduviidae - No prosternal sulcus receiving apex of labium; labium straight or curving; head not conspicuously necklike behind eyes, never with a transverse impression anterior to ocelli; membrane venation variable . 6 6. Antennae with 5 apparent segments (Figs. 47.1,56.1) . 7 - Antennae with 4 segments . 8 7. Scutellum with 1-7 pairs of trichobothria laterally; membrane with a “stub,” usually most readily visible on ventral surface (Prostemmatinae) . Nabidae (part) - Scutellum without lateral trichobothria; membrane without a “stub” . Pachynomidae 8. Hemelytra usually well developed'; never ectoparasitic . 9 - Hemelytra always staphylinoid (Fig. 50. IB), in the form of small pads, or absent (Figs. 61.1 A. B, 62.1 A)'; frequently ectoparasitic . 18 50.1 A. ... 10 ... 16 9. Costal fracture present in macropterous forms, usually demarcating a distinct cuneus (Figs 53.2A, 57.1, 58.1) . - Costal fracture absent in macropterous forms, no cuneus (Figs. 51. 2C. 55.lA, 56.1) .... 10. Large species, 10-15 mm long; exocorium expanded into broad embolium (Fig. 49.1 A); veins ema¬ nating from posterior margin of cells in membrane; penultimate labial segment much longer than other 2 combined . Velocipedidae - Much smaller species, usually under 4 mm long; exocorium not greatly expanded; membrane without many emanating veins; proportions of labial segments variable, penultimate segment rarely longer than combined lengths of other segments . 11 11. Membrane with a single cell with very heavy veins and a wed-developed “stub” on posterior angle (Fig. 50. ID); all tarsi 2-segmented . Microphysidae (part) - Membrane with weakly developed venation, stub always close to posterior margin of corium; tarsal formula variable . . 12 12. Tarsi 2-segmented; male genitalia symmetrical or nearly so, parameres elongate and nearly equally developed . Plokiophilidae (part) - Tarsi 3-segmented; male genitalia symmetrical or asymmetrical . 13 13. Male genitalia symmetrical or nearly so, parameres elongate, nearly equally developed, genital cap¬ sule tubular (Fig. 58.2C); corial glands present (Fig. 10. lOD); spider web commensals {Lipokophila Stys) . Plokiophilidae (part) Cimicomorpha 147 14. 15. 16. 17. 18. 19. Male genitalia strongly asymmetrical, left paramere variable in shape (Figs. 59.1C, 61.1C), right paramere often greatly reduced, genital capsule frequently short and broad; corial glands absent; free-living . 14 Abdominal terga 1 and 2 with laterotergites, terga 3-8 forming simple plates; no traumatic insemi¬ nation . Lasiochilidae All abdominal terga with laterotergites; insemination traumatic . 15 Female with internal apophysis on anterior margin of abdominal sternum 7 - . Lyctocoridae Female lacking apophysis on abdominal sternum 7- . Anthocoridae R+M in forewing elevated and keel-like (Fig. 51.2C); small species, about 2.0 mm long . . Joppeicidae R-l-M in forewing not elevated and keel-like; larger species, always over 4 mm long . 17 Ultimate labial segment longest, labium rather straight (Fig. 55.1B) . Medocostidae Ultimate labial segment not the longest, labium at least weakly curving (Fig. 56.2A) (Nabinae) . Nabidae (part) All tarsi 2-segmented; hemelytra staphylinoid (Fig. 50. IB) . Microphysidae (part) All tarsi 3-segmented or middle and hind tarsi 4-segmented; hemelytra in form of small pads or absent . 19 All tarsi 3-segmented; hemelytra in the form of small pads; no ctenidia (Fig. 61.1 A, B); compound eyes small; temporary ectoparasites . Cimicidae Middle and hind tarsi 4-segmented; hemelytra absent; ctenidia present (Fig. 62.1 A, F, G); compound eyes absent; permanent ectoparasites of bats . Polyctenidae ' Members of many families may be brachypterous or apterous, such as Nabidae and Anthocoridae, although the more common condition is to have well-developed forewings. Thus, specimens with short wings should be run through both halves of couplet 8. 2 Members of the Lasiochilidae, Lyctocoridae, and Anthocoridae are very similar in general appearance and share many attributes. Correct family placement may require reference to faunistic treatments containing detailed keys for identification of genera and species. Reduvioidea 47 Pachynomitdae General. This small group of seldom-collected tropi¬ cal ground-dwelling predators has no common name. They range in length from 3.5 to 15 mm and often have a general facies of prostemmatine Nabidae (Fig. 47.1). Diagnosis. Dorsum varying from highly polished (Pachynomus Klug) to dull, glabrous to highly setose; compound eyes large, head without constricted postocu¬ lar region (Fig. 47.1); ocelli present or absent; antennae apparently 5-segmented, pedicel subdivided, distal por¬ tion usually with a single trichobothrium (Fig. 10.8H), intrapediceloid uniquely present (Fig. 47.1); labium thick, strongly curving, segment 1 obsolete, segment 2 shortest (Fig. 47.2A); bucculae directed anteriorly, ob¬ scuring base of labium; gula long; forefemur greatly en¬ larged; foretibia bearing a fossula spongiosa (Fig. 47.1); forewing with short costal fracture in Pachynomus and Punctius Stal, membrane with 2 elongate cells with a few to many radiating veins, stub (processus corial) absent (Figs. 47.1, 47.2B, C); metathoracic scent glands with strongly reduced grooves; Brindley’s gland present; ab¬ domen with dorsal and ventral laterotergites; abdominal spiracle 1 present, spiracles 2-8 located on mediostemite; pair of trichobothrium-like setae on either side of midline of abdominal segments 3-8 or 4-8 (Fig. 47.2D); para- stigmal pits located on ventral laterotergites of abdomi¬ nal segment 2 (except in Punctius)', apophysis present on anterior margin of abdominal sternum 7 in female; dorsal abdominal scent-gland scars present on anterior margin of mediotergites 4-6 of adults; abdominal seg¬ ment 8 in males reduced and telescoped within segment 7; male genitalia symmetrical, pygophore well devel¬ oped and inserted apically on abdomen (Fig. 47.2E, F); ovipositor platelike; spermatheca absent, paired pseudo- spermathecae situated on medial ectodermal portion of female gonoducts (Fig. 47.2H). Classification. Pachynomus was treated by its au¬ thor Klug (1830) as a subgenus of Reduvius; Stal later treated Pachynomus and Punctius Stal as members of the Nabidae, a position followed by many authors (but see 148 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Reuter, 1908), until Carayon (1950b) raised the group to viidae. Two subfamilies comprising 4 genera and about family rank, pointing out that on the basis of reproductive 15 species are recognized, following the classification of anatomy, the taxon has its closest affinities with the Redu- Carayon and Villiers (1968). Key to Subfamilies of Pachynomidae 1. Ocelli absent (Fig. 47.1); posterior margin of corium obliquely angled posterolaterally (Figs. 47.1, 47.2B);corium largely glabrous . Pachynominae - Ocelli present; posterior margin of corium tmncate (Fig. 47.2C); corium distinctly pubescent . . Aphelonotinae APHELONOTINAE. Ocelli present; endocorium truncate posteriorly and of same length as clavus; corium pubes¬ cent and without a trichobothriiform seta; abdominal sterna 4-8 each bearing a trichobothrium on either side of midline. Aphelonotus Uhler, the only included genus, is found in tropical America and central Africa. Most species are less than 5-6 mm long and are light brown. PACHYNOMINAE (FIG. 47.1). Ocelli absent; posterior mar¬ gin of corium obliquely angled posterolaterally; corium glabrous except for a trichobothriiform seta at anterolat¬ eral angle of clavus; abdominal sterna 3-8 each bearing a trichobothrium located on either side of midline (Fig. 47.2D). The group comprises the genera Camarochilus Harris, Pachynomus Klug, and Punctius Stal, the first being re¬ stricted to the New World tropics, the last two to the Old World, ranging from Africa to India, Members of all three genera are relatively large, in the case of Pachynomus picipes Klug attaining a length of 10.0-14.0 mm. Specialized morphology. The antennal pedicel (seg¬ ment 2) is subdivided, resulting in apparently 5-seg- mented antennae (Fig. 47.1). Along with the Redu viidae, the Pachynomidae are novel among the Heteroptera for the occurrence of antennal trichobothria (Fig. 10.8H) in most species. They also possess trichobothria on either side of the ventral abdominal midline (Fig. 47.2D), which occur elsewhere only in the Velocipedidae, but the phylogenetic analysis of Schuh and Stys (1991) suggests that they are not homologous in the two groups. All Pachynomidae except Punctius possess parastigmal pits (fossettes parastigmatiques) on the ventral lateroter- gites of abdominal segment 2; these apparently are not homologous with similar-appearing structures located on the ventral laterotergites of one or more of abdominal seg¬ ments 3-7 in the Nabidae. Paired pseudospermathecae of the type found in the Reduviidae—and Tingidae—are also present in the family (Fig. 47.2H). Natural history. Nothing is known of the life habits of the group. Nymphs are unknown, and most specimens have been collected at lights. Fig. 47.1. Pachynomidae. Punctius alutaceus (Stal). Distribution and faunistics. Carayon and Villiers (1968) provided a generic revision, synonymic, and dis¬ tributional information for the known species. Harris (1931b) revised the New World Aphelonotus spp., but substantial numbers of species remain to be described. Pachynomidae 149 Fig. 47.2. Pachynomidae. A. Lateral view, head, Camarochilus sp. B. Forewing, Camarochilus sp, C. Forewing, Aphelonotus sp. (A-C from Schuh and Stys, 1991). D. Abdominal venter, Camarochilus americanus (Harris). E. Male genitalia, Aphelonotus africanus Carayon and Viiliers. F. Aedeagus, A. fuscus Carayon and Viiliers. G. Paramere, A. fuscus. H. Gynatrial complex, A. fuscus (D-H from Carayon and Viiliers, 1968). Abbreviations: ps, pseudospermatheca: tb, trichobothrium. 48 Reduviidae General. The predatory assassin bugs are one of the largest and morphologically most diverse families of true bugs. Reduviids range in size from relatively small tuid extremely delicate insects of only a few millimeters in length, such as members of the genus Empicoris Wolff (Emesinae), to very large and formidable animals such as Arilus Hahn (Harpactorinae), which may attain a length ^ of nearly 40 mm. Diagnosis. Compound eyes large, protuberant, head behind them usually elongate and constricted behind, ^ usually bearing 2 ocelli (Fig. 48.2A-D); antennae some¬ times with flagellar segments fusiform (e. g., Phymatinae; Fig. 48.1), prepedicellite present, not noticeably elon- gate, pedicel in nymphs and adults with at least 1 tri¬ chobothrium, adults sometimes with 20 or more (Fig. 48.3D, E); labium usually with 3 (rarely 4) apparent ^ segments, short, stout, curving, often inflexible (Fig. 150 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 48.1. Reduviidae; Phymatinae. Oxythyreus cylindricomis (West- wood) (from Slater, 1982). 48.3A, C), sometimes longer, straight, and flexible (Fig. 48.3B); forewings lacking costal fracture, membrane usually with 2 elongate cells and a few veins emanating posteriorly (Fig. 48.2C, D), sometimes cells subdivided or open (Fig. 48.2A); stub (processus corial) absent; fos- sula spongiosa usually present on one or more pairs of legs; tarsal formula variable, foretarsi sometimes absent; prostemum usually with a transversely striate stridulatory groove (Fig. 10.7F, G); metathoracic scent-gland chan¬ nels and evaporatory areas strongly reduced or absent; paired Brindley’s gland present in first abdominal seg¬ ment, opening dorsolaterally at thoracicoabdominal junc¬ tion; paired ventral glands present in some taxa, opening ventrally at thoracicoabdominal junction; dorsal and ven¬ tral laterotergites present; abdominal spiracle 1 present (except phymatine complex), spiracles 2-8 located on mediosternite; nymphal dorsal abdominal scent glands— if present—on anterior margin of abdominal terga 4, 5. and 6; abdominal segment 8 in male largely telescoped within segment 7; male genitalia (Fig. 48.3F-J) usually symmetrical; ovipositor usually platelike; female internal genitalia with paired pseudospermathecae and accessory gland (Fig. 48.3L—M); eggs with 3 or more micropyles. Classification. The classification of the Reduviidae derives originally from the work of Amyot and Serville (1843) and Stal (1859a, 1866a, b). More recent contribu¬ tions by Usinger (1943), Davis (1957, 1961, 1966, 1969), and Carayon et al. (1958) inquired into the basis of those early systems. N. C. E. Miller and A. Villiers established several subfamilies on the basis of unique characters. Two up-to-date catalogs by Putchkov and Putchkov (1985, 1986-1989) and Maldonado (1990; see also Kerzhner, 1992) are available, although based on substan¬ tially different classifications. Maldonado recognized 25 subfamilies and treated the Elasmodeminae and Phymati¬ nae as separate families; Putchkov and Putchkov recog¬ nized 21 subfamilies, including the Elasmodeminae and Phymatinae. We treat the Reduviidae in a manner con¬ sistent with the classification of Putchkov and Putchkov, except we place Carayonia Villiers in the Saicinae, fol¬ lowing Maldonado. The approximately 930 genera and 6500 species are placed in 22 subfamilies, which—except for the phymatine complex—are arranged alphabetically in the following text. Key to Subfamilies of Reduviidae 1. Antennal segments 3 and 4 incrassate, of greater diameter than segments 1 and 2; forefemora tremendously dilated, the tibiae chelate (Fig. 48.1) . Phymatinae (part) - Antennal segments 3 and 4 more slender than remaining segments; forefemora not tremendously dilated (Fig. 48.2) ...:. ’. . .1. . 2 2. Membrane with 3 simple veins attaining posterior margin (Fig. 48.2A); body extremely flattened . Elasmodeminae - Membrane with 1 or more closed cells (Fig. 48.2C, D); body generally not extremely flattened . 3 Key to subfamilies of Reduviidae modified from Usinger, 1943. Reduviidae 151 3. Abdominal sternum 3 usually produced anteriorly to form a trichome; antennal segments 3 and 4 inserted proximal to apex of segments 2 and 3, respectively . Holoptilinae - Abdominal sternum 3 without trichome; antennal segments 3 and 4 inserted on apex of segments 2 and 3, respectively . 4 4. Ventral surface of head produced to form a more or less distinct rostral (buccal) groove . 5 - Ventral surface of head not produced to form a rostral groove . 6 5. Ocelli absent; antennal segments 2, 3, and 4 usually slender; body usually clothed with a dense vestiture of apically curved setae . Tribelocephalinae - Ocelli present; antennal segments 2, 3, and 4 not long and slender; body without a dense vestiture . Phimophorinae 6. Forecoxae elongate, usually at least four times as long as wide, usually extending well beyond cly- peus anteriorly (Fig. 48.2B); body elongate, slender; hemelytra almost entirely membranous except for veins; forelegs raptorial . 7 - Anterior coxae usually less than twice as long as broad and not extending beyond clypeus; body, if elongate, not extremely slender; clavus and corium coriaceous, membrane well differentiated; forelegs not conspicuously raptorial ... 8 7. Ocelli and nymphal dorsal abdominal scent-gland openings absent . Emesinae - Ocelli and 3 nymphal dorsal abdominal scent-gland openings present . Bactrodinae 8. Pronotum with a distinct constriction behind middle . 9 - Pronotum without a distinct constriction at or behind middle . 10 9. Ocelli present; fore- and middle tibiae not curved distally, provided with fossula spongiosa distally . Peiratinae - Ocelli absent, foretibia curved apically and produced beyond insertion of tarsus as a stout spine; fossula spongiosa absent . Vesciinae 10. Ocelli absent . 11 - Ocelli present (except in rare brachypterous forms) . 12 11. Second labial segment swollen proximally; membrane with at least 2 closed cells . Saicinae - Second labial segment not swollen proximally; membrane with a single large cell .... Chryxinae 12. Cubitus branching to form an additional four- to six-angled cubital cell between corium and mem¬ brane (Fig. 48.2C) . 13 - Cubitus simple, not forming such a cubital cell . 14 13. Cubital cell usually hexagonal (Fig. 48.2C); antennal segment 1 stout, porrect; nymphs with 2 dorsal abdominal scent glands . Stenopodainae - Cubital cell usually quadrangular, antennal segment 1 usually relatively slender; nymphs with 3 abdominal scent glands . Harpactorinae 14. Antennae with 4 segments . 16 - Antennae usually with more than 4 apparent segments . 15 15. Antennal segment 2 subdivided into 4-36 pseudosegments; eyes located posteriorly, ocelli placed between them; scutellum without 2 posteriorly projecting prongs . Hammacerinae - Antennal segment 3 or segments 3 and 4 usually subdivided, forming a total of 7 or 8 apparent antennal segments; eyes not located posteriorly; ocelli placed posterior to eyes; scutellum with 2 posteriorly projecting prongs . Ectrichodiinae 16. Head rarely transversely constricted behind eyes, ocelli usually located on oblique elevations at posterolateral angles of long, cylindrical head (Fig. 48.2D); dorsal abdominal scent-gland openings absent in nymphs . Triatominae - Head transversely constricted behind eyes, ocelli variously positioned; dorsal abdominal scent gland(s) present in nymphs . 17 17. Eyes strongly pedunculate .. Cetherinae - Eyes not stalked or pedunculate . 18 18. Body surface tuberculate or spiny . 19 - Body surface not tuberculate or spiny . 21 19. Penultimate labial segment very long and straight, much longer than other 2 segments . . Physoderinae - Penultimate labial segment not elongate, about as long as ultimate segment, antepenultimate segment longest . 20 Reduviidae 153 20 . 21 . 22 . 23. Tarsi 3-segmented; fossula spongiosa present on fore- and middle tibiae; head and pronotum bearing large spines (Fig. 48.3A); pronotum without longitudinal carinae . Centrocneminae Tarsi 2-segmented; fossula spongiosa absent from all tibiae; body without large spines, only setiger- ous tubercles; pronotum with a longitudinal carina on either side of midline . . Phymatinae (Themonocorini) Scutellum with a long posteriorly projecting spine . Manangocorinae Scutellum without a long spine .. 22 Antennae inserted anteriorly, or more commonly laterally, but not on long anteriorly projecting tubercles . Reduviinae Antennae inserted on prominent, anteriorly projecting tubercles at front of head . 23 Foretarsi 2-segmented; middle and hind tarsi 3-segmented . Salyavatinae All tarsi 3-segmented . Sphaeridopinae Phymatine Complex of Subfamilies Although the phylogenetic relationships of the redu- viid subfamilies as a whole are not well understood, four groups—the Phymatinae, Holoptilinae, Elasmodeminae, and Centrocneminae—are drawn together by the follow¬ ing unique characters: presence of ventral glands near posterior border of metathoracic sternum; basal plate stmts of male phallus prolonged apically into pair of slen¬ der filaments partially or completely enclosed by endo- soma; duck-head-shaped parameres; and reduced external female genitalia (Carayon et al., 1958). CENTROCNEMINAE. Adults with a Single trichobothrium on antennal pedicel; labium obviously 4-segmented (Fig. 48.3A); membrane with 2 elongate cells; body highly tuberculate (Fig. 48.3A), as in many other members of phymatine complex, and somewhat flattened. Currently known from four genera and 33 species dis¬ tributed from India to the Philippines, the Centrocnemi¬ nae were revised by Miller (1956b). Most specimens have been found on the trunks of trees, suggesting that the group may be subcorticolous. ELASMODEMINAE (FIG. 48.2A). Membrane with a few lon¬ gitudinal veins reaching posterior margin, no closed cells; body strongly flattened, appendages spinose. Elasmodema Stal, with two Neotropical species, has been treated as a distinct family by some authors (Wy- godzinsky, 1944a; China and Miller, 1959; Maldonado, 1990), but its closest affinities are with the Holoptilinae (Carayon et al., 1958). Elasmodemines live under loose bark, apparently preying on other insects that occur there (Wygodzinsky,. 1944a). HOLOPTILINAE. Body and appendages with long to very long setae; membrane venation often with a single cell or 2 longitudinal veins; antennal segments 3 and 4 some¬ times fused; abdominal sternum 3 often with a trichome and associated gland. This group of 15 genera and 78 species is divided among three tribes (Holoptilini, Dasycnemini, and Ara- dellini) according to Wygodzinsky and Usinger (1963). who keyed the known genera. Most taxa occur in the southern Palearctic, Old World tropics, and Australia (Malipatil, 1985), with Neolocoptiris Wygodzinsky and Usinger (1963) occurring in Guyana. PHYMATINAE (FIG. 48.1). Forefemur often greatly en¬ larged; foretibia and tarsus fused (except in Themonoco¬ rini) and often lying in scrobe on femur (Fig. 10.4G, H). or occasionally absent; forewing membrane often without distinct cells, or if with cells, with several supernumerary veins radiating from them posteriorly. Commonly known as ambush bugs, this group of 26 genera and 281 species of morphologically distinctive bugs has been monographed by Handlirsch (1897), Kor- milev (1962), and Maa and Lin (1956). Four tribal group¬ ings are recognized; Carcinocorini, Macrocephalini, Phy- matini, and Themonocorini (Carayon et al., 1958). The most recent source on the group, including keys to tribes, genera, and species of all genera except Lophoscutus Kormilev and Phymata Latreille, is the conspectus of Froeschner and Kormilev (1989), who, like many other authors, have accorded the group family rank. Other Reduviid Subfamilies BACTRODINAE. Elongate, slender; ocelli present; mem¬ brane with a single closed cell; forelegs with elongate coxae and spinose trochanters, femora, and tibiae; tarsal segment 3 slightly swollen on middle leg with a short stout spine near middle of ventral surface; claws on all legs unequally developed, one claw large and with large basal tooth, other claw much smaller, about as large as basal tooth of larger claw. The Neotropical genus Bactrodes Stal, with four in- 154 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 48.3. Reduviidae. A. Lateral view, head, Neocentrocnemis signoreti (Stai) (from Schuh and Stys, 1991). B. Lateral view, head, Triatoma rubrofasciata (from Lent and Wygodzinsky, 1979). C. Lateral view, head, Kodormus sp. D. Antennal trichobothrium, Phymata pennsylvanica Handlirsch. E. Antennal trichobothria, Rasahus sulcicollis (Serville) (D, E from Wygodzinsky and Lodhi, 1989). F. Aedeagus, sagittal view. Tribe- mula pilosa Horvath (from Wygodzinsky, 1966). G. Aedeagus, lateral view, Basmodema erichsoni (from Carayon et al., 1958). H. Aedeagus, lateral view, Mesosepis papua Wygodzinsky (from Wygodzinsky, 1966). I. Paramere, £ erichsoni (from Carayon et al., 1958). J. Paramere, M. papua (from Wygodzinsky, 1966). K. Right paramere. Pirates hybridus Scopoli (from Ghauri, 1964). L. Gynatrial complex, Phymata erosa (Lin¬ naeus) (from Pendergrast, 1957). M. Female reproductive system, Themnocoris kinkaianus Carayon, Usinger, and Wygodzinsky (from Carayon et al., 1958). Abbreviations: ps, pseudospermatheca; tb, trichobothrium; vg, vermiform gland. Reduviidae 155 eluded species, was placed in a monotypic subfamily by Stal (1862). Davis’s (1969) analysis indicated that, even though it differed in several characteristics from other re- duviids he placed in the Harpactorinae. Bactrodes was probably a member of the clade containing the Harpacio- rini, Rhaphidosomini, and Tegeini. We treat Bactrodes as a subfamily, awaiting improved knowledge of character polarity in the Reduviidae. The group was diagnosed and keyed by McAtee and Malloch (1923). Nothing is known of the life habits. CETHERINAE. Head short, transverse; transverse sulcus running between eyes on vertex; eyes strongly peduncu¬ late. The group, comprising five genera and 23 species, was first treated as a tribe within the Reduviinae by Jeannel (1919). Putchkov and Putchkov (1985) accorded subfamily status with three tribes—Cetherini, Euphenini, Pseudocetherini. Four genera, including Cethera Amyot and Serville, are known from tropical Africa and Mada¬ gascar, with only Eupheno Gistel occurring in the New World tropics. Villiers (1948) provided a comprehensive treatment of the tropical African fauna. These are active insects, some of which are known to feed on termites (Miller, 1956a). CHRYXiNAE. Head short, transverse, barely projecting in front of compound eyes; corium distinctly delimited from membrane, no cubital cell; membrane with a single cell. The monotypic Neotropical Chryxus Champion was originally given subfamily status (Champion, 1897- 1901); subsequently, two additional monotypic genera have been placed in the group, Lentia Wygodzinsky (1946) and Wygodzinskyella Usinger (1952). Chryxine morphology was well illustrated by Lent and Wygod¬ zinsky (1944) and Wygodzinsky (1946). Wygodzinsky (1946) noted that the movements and coloration of these small insects are reminiscent of some Anthocoridae and particultirly Fulvius quadristillatus Stal (Miridae), in whose company Lentia corcovadensis Wygodzinsky was taken in the field. ECTRiCHODiiNAE. Often heavy-bodied with red and black coloration; scutellum with two short prongs pro¬ jecting posteriorly; sexual dimorphism very common, females often with greatly reduced wings; antennae usu¬ ally with 7 or 8 apparent segments, resulting from frag¬ mentation of flagellum; 4-20 antennal trichobothria in nymphs and adults (Wygodzinsky and Lohdi, 1989); fore¬ wing membrane with 2 or 3 cells; fossula spongiosa present on fore- and middle tibiae. This circumtropical group comprises approximately 75 genera and 300 species. Cook (1977) dealt with the Asian fauna, and Dougherty (1980) the Neotropical fauna. EMESINAE (FIG. 48.2B). Foreacetabulum opening anteri¬ orly; pterostigma carried beyond apex of discal cell (Wy¬ godzinsky, 1966); body elongate, slender; appendages often threadlike; wing polymorphism common, often sexual, occasionally both sexes having wings greatly re¬ duced and nonfunctional; ocelli absent except in Aus¬ tralian Armstrongocoris Wygodzinsky; fossula spongiosa absent from all legs; claws of foretarsus often unequally developed and parempodia frequently reduced; claws and parempodia on middle and hind legs fully developed; claws sometimes toothed, undersurface sometimes with membranous lamella. The group was monographed by Wygodzinsky (1966), who recognized six tribes—Collartidini, Leistarchini, Emesini, Ploiariini, Deliastini, and Metapterini—com¬ prising approximately 86 genera, some 20 of which are monotypic and known from a single locality. The Emesinae, the apparent sister group of the Saicinae (Wygodzinsky, 1966), are most diverse in the tropics and show substantial diversity in the islands of the Paci¬ fic. HAMMACERINAE. Head Strongly produced anteriorly, not constricted or elongated behind eyes; ocelli present; labium stout, curving, inflexible; membrane 2-celled; fossula spongiosa on fore- and middle tibiae; antennal segment 2 annulated, containing 4-36 pseudosegments. First proposed by Stal (1859b) as a subfamily level taxon, this group has often been referred to as Micro- tominae. Hammacerines comprise the New World genera Homalocoris Perty and Microtomus Illiger with a total of 19 described species. According to Readio (1927) most records indicate they live under bark. Microtomus was revised by Stichel (1926). HARPACTORINAE. Head usually with cylindrical post¬ ocular region; antennal segment 1 strikingly elongate; foretibia usually with a well-developed apical spur; meta- coxal comb absent; metathoracic scent glands reduced or absent; hemelytron with quadrate or pentagonal corial cell at base of cubital cell, formed uniquely by anterior and posterior cross vein between Cu and Peu; hind wing with submarginal Sc, postcubital sector relatively broad, Peu slightly curved and well separated from first anal vein; ventral connexival suture usually poorly formed or absent; vermiform gland absent in females; subrec- tal gland ofte.n present, opening into membrane between styloids and anus; pseudospermathecae usually greatly reduced or absent (Davis, 1969). First recognized in the classification of Amyot and Ser¬ ville (1843), this is the largest reduviid subfamily. We follow Davis (1969) and Putchkov and Putchkov (1985) in treating as tribes several groups that have been accorded subfamily status by Maldonado, Miller, and Villiers. 156 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Key to Tribes of Harpactorinae 1. Ocelli lateral, more widely separated from one another than from eyes . 2 - Ocelli less widely separated from one another than from eyes . 4 2. Pronotum completely covering scutellum . Diaspidiini - Scutellum at least partly exposed . 3 3. Foretarsi 1-segmented; membrane with 3 cells . Ectinoderini - Foretarsi 2-segmented; membrane with 2 cells . Apiomerini 4. Labial segments 3 and 4 (penultimate and ultimate) slender, straight or slightly procurved, and of uniform thickness or very slightly tapered; labial segment 2 always shorter than 3; antennal segment 1 not strikingly elongate .. 5 - Labium usually distinctly recurved and tapering from base to apex; labial segment 2 often subequal in length to segment 3; antennal segment 1 strikingly elongate . Harpactorini 5. Body and legs long, very slender; postocular region of head long, cylindrical; forecoxal cavities usually closed .. Rhaphidosomini - Body more robust, with dense areas of glandular setae; postocular region of head usually short, subhemispherical; forecoxal cavities open . Tegeini APIOMERINI. The genera were keyed by Costa Lima et al. (1948). The largest and probably best known genus is Apiomerus Hahn (see Costa Lima et al., 1951), with 110 described species, often of conspicuous black and red coloration. Twenty-eight Heniartes spp. were treated in the well-illustrated work of Wygodzinsky (1947a). These diurnal predators live on plants, and at least some use sticky material on the anterior tibiae to hold prey. DIASPIDIINI. A grouping of three Ethiopian genera— Cleontes Stal, Rhodainiella Schouteden, and Diaspidius Westwood—all species of which apparently live in leaf debris or under loose bark. ECTINODERINI. Containing only the genera Amulius Stal and Ectinoderus Westwood (Maldonado, 1990), this group of 20 species is distributed from Sri Lanka to New Guinea. Nothing appears to be known of their habits. HARPACTORINI, INCLUDING RHYNOCORINI VILLIERS (1982). This is a very large group—containing hundreds of gen¬ era—that, according to Davis (1969), possesses few dis¬ tinctive characters. Included are such conspicuous taxa as the wheel bugs of the New World genus Arilus Hahn and some of the largest genera in the Reduviidae, such as Rhynocoris Hahn and Sphedanolestes Stal from the Old World and Zelus Fabricius from the New World (see Hart 1986, 1987). The feeding and reproductive behav¬ ior of Rhynocoris and Pisilus spp. have been studied by Edwards (1962) and Parker (1965, 1969), and Louis (1974) described the habits of additional West African species. RHAPHIDOSOMINI. Comprises the genera Rhaphidosoma Amyot and Serville, Lopodytes Stal, Leptodema Carlini, Hoffmanocoris China, and Harrisocoris Miller—inhabi- Key to tribes of Harpactorinae adapted from Davis, 1969. tants of the deserts and savannas of the Old World— where in the hottest periods of the day they may be found basking at the tips of grasses, fully exposed to the sun. TEGEINI. Comprising seven genera from the Old World tropics, these diurnal predators inhabit the trunks and foli¬ age of trees and are known to feed on termites (Miller. 1952; China and Miller, 1959). MANANGOCORiNAE. Head short, globose, barely pro¬ jecting anteriorly beyond relatively small eyes; scutellum with long spine; membrane with 2 cells, inner trapezoi¬ dal; tarsi 2-segmented; tibiae without fossula spongiosa. Containing only the Malaysian species Manangocoris horridus Miller (1954a), this taxon may not merit sub¬ family status. Nothing is known of its habits. PEiRATiNAE. Antennal segment 1 shorter than segment 2; apparent first labial segment (2) shortest; anterior pro- notal lobe longer than posterior lobe; forecoxae more strongly enlarged than middle and hind coxae and with lateral surface flattened and weakly concave: foretibia di¬ lated distally, with a fossula spongiosa; middle tibia with or without fossula spongiosa; males of some Peirates spp. dimorphic, one morph possessing a large asymmetrical sinistral process on sternum 7 and small symmetrical parameres, the other morph lacking process and possess¬ ing large (Fig. 48.3K) asymmetrical parameres (Ghauri, 1964). This worldwide group (often referred to as Piratinae) comprises approximately 31 genera, including Ecfomo- coris Mayr and Peirates Serville-^which are diverse and widely distributed in the Old World— Rasahus Amyot and Seri’ille from the New World, and the circumtropical Sirthenea Spinola. Peiratines appear to be primarily ground-dwelling, feeding on a variety of arthropods. In general facies, habi- Reduviidae 157 tat, and behavior they resemble Prostemmatinae (Nabi- dae). They are often attracted to lights. These are fast¬ running bugs whose bite can be extremely painful. PHIMOPHORINAE. Habitus aradid-like; body and append¬ ages covered with patches of waxy secretions; all an¬ tennal segments conspicuously thickened with antennal insertions protected by lateral shieldlike structures; buc- culae present; labial segment 2 much longer than 3 and 4 combined; forewings largely membranous; legs short and stout, tarsi minute, 2-segmented; prosternum with 2 large shieldlike structures adjacent to stridulatory groove. Comprises Mendanocoris Miller, with two species from the Solomon Islands and Malaya, and Phimophorus Ber- groth, with a single species from the New World tropics (Usinger and Wygodzinsky, 1964). No information exists on their biology. PHYSODERiNAE. Body surface tuberculate, with spatu- late setae; head elongate; eyes relatively small, far re¬ moved from the anterior pronotal margin; ocelli present; labial segment 3 (penultimate) very long and straight; apex of pronotum spatulate; membrane with 2 cells; fos- sula spongiosa absent from all legs; nymphs with 3 dorsal abdominal scent glands. Comprises 11 Indo-west Pacific genera (including Phy- soderes Westwood with 37 species) and the monotypic Cryptophysoderes fairchildi Wygodzinsky and Maldon¬ ado (1972) from Panama. The subfamily status of the group was established by Miller (1954b). Known habi¬ tats include vegetable debris, the bases of banana and Pandanus leaves, hollow trees, and caves. REOUViiNAE. Recognized primarily by the absence of characters occurring in other reduviids; ocelli usually present; discal cell usually absent; tarsi 3-segmented; fos- sula spongiosa on fore- and middle tibiae; nymphs with 3 dorsal abdominal scent glands. This worldwide group comprises at least 140 genera, including Acanthaspis Amyot and Serville, Reduvius Fabricius, and Zelurus Burmeister. Many, if not most, members are thought to be general insect or arthropod predators. Most are nocturnal. Some, such as many Re¬ duvius spp., live in animal burrows, whereas Reduvius personatus (Linnaeus) is synanthropic. SAiciNAE. Ocelli absent; second apparent labial seg¬ ment (3) expanded and bulbous basally; opposing sur¬ faces of head and labium armed with stiff setae or spines; fossula spongiosa absent. This tropicopolitan group was first recognized by Stal (1859b) and currently contains two tribes—Saicini and Visayanocorini (Putchkov, 1987)—and 21 genera, by far the largest of which are Polytoxus Spinola from the Old World and Saica Amyot and Serville from the New World. It is composed primarily of elongate slen¬ der species and was treated as the sister group of the Emesinae by Wygodzinsky (1966). Maldonado (1981) keyed the American genera, and Villiers (1969b) treated the African fauna, including a key to genera. Little ap¬ pears to be known of their habits, and they have been most commonly collected at lights. SALYAVATINAE. Head relatively small, somewhat globu¬ lar, constricted just behind eyes; neck short; eyes rela¬ tively small; antennae sometimes apparently 3-segmented (e.g., in Salyavata), inserted on prominent, anteriorly projecting tubercles; labium moderately long, weakly- curving; pronotum, scutellum, and abdomen often with slender, strongly acuminate spines; fore- and middle tibiae with fossula spongiosa; foretarsi 2-segmented, middle and hind tarsi 3-segmented; membrane with 2 cells; nymphs with 3 dorsal abdominal glands. This circumtropical group comprises 16 genera, the largest being Lizarda Stal from Africa and the Orient; only Salyavata Amyot and Serville is known from the New World tropics (Wygodzinsky, 1943). Most species are nocturnal, although at least some Patalochirus Palisot de Beauvois are diurnal. Some salyavatine nymphs have a vestiture of sticky setae and cover themselves with debris. SPHAERiDOPiNAE. Head projecting only slightly beyond anterior margin of eyes; eyes large, nearly contiguous ventrally; antennae inserted on anteriorly projecting tu¬ bercles; rostrum straight; all tarsi 3-segmented. This Neotropical group comprises the monotypic gen¬ era, Eurylochus Torre-Bueno, Sphaeridops Amyot and Serville, Veseris Stal, and Volesus Champion. Pinto (1927) speculated, probably on the basis of the labial structure, that the group fed on vertebrate blood, although there seems to be no direct evidence for this theory. STENOPODAiNAE (FIG. 48.2C). Elongate, dull-colored; large cell, often pentagonal or hexagonal, formed on co- rium by cubital and postcubital veins, with 2 cells of membrane posteriorly adjacent (Fig. 48.2C); antennifer- ous tubercles sometimes strongly produced anteriorly; an¬ tennal segment 1 elongate, rather strongly developed, re¬ maining segments more slender and in repose folded back against first (Figs. 48.2C, 48.3C); ocelli present; legs long, slender; hemelytral development variable, males macropterous, females often with more reduced fore¬ wings. Approximately 113 genera have been described, most of them from the tropics. Before 1969, this group was referred to as the Stenopodinae. Many species appear to be closely associated with the soil—often being covered with soil or sand—and at least some insert their eggs in the soil. Most species are known from collections made at lights, with males being collected much more com¬ monly than females. It seems certain that the group is largely nocturnal, even though there are almost no avail¬ able direct observations of their behavior. Barber (1930) 158 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) revised the New World fauna, and Giacchi (e.g., 1984) has subsequently added considerably to our knowledge of diversity of the group in South America. TRIATOMINAE (FIG. 48.2D). Labium elongate, nearly straight, with a-flexible membranous connection between segments 3 and 4 (Fig. 48.3B); antennal segment 2 in adults bearing from 3 to 7 trichobothria; head not constricted behind compound eyes; dorsal abdominal scent glands absent in nymphs; connexivum often broadly membranous, allowing for abdominal expansion during consumption of blood meal. Lent and Wygodzinsky (1979) recognized five tribes— Rhodniini, Cavernicolini, Bolboderini, Alberproseniini, and Triatomini—containing 14 genera and 111 species. The group is most diverse in the New World, ranging from the southwestern United States to Argentina. Triatoma rubrofasciata (De Geer) is tropicopolitan, five species of Linshcosteus Distant are restricted to India, and seven species of Triatoma Laporte are restricted to an area ranging from southern India and Burma to New Guinea. The obligate blood-feeding triatomines are well known for their ability to transmit the debilitating Chagas’ disease, whose causative agent. Trypanosoma cruzi, is widely distributed in the New World (see Chapter 8). The literature on the Triatominae is immense, includ¬ ing numerous references to Rhodniusprolixus Stal, which is an important vector of Chagas’ disease and a widely used laboratory animal. Summary sources include com¬ pilations by R. E. Ryckman and coauthors (see Literature Cited). TRIBELOCEPHALINAE. Head, body, and veins of corium densely tomentose, body surface obscured; eyes flattened and not strongly projecting laterally, sometimes nearly contiguous dorsally; ocelli absent; antennal segment 1 thickened and much longer than head; corium narrow and elongate, membrane very broad. First recognized by Stal (1866a), this broadly dis¬ tributed paleotropical group comprises 13 genera placed in two tribes—Opistoplatyini and Tribelocephalini—the largest genus being Tribelocephala Stal. A few specimens have been taken in leaf litter, but most have been collected at lights, suggesting nocturnal habits. VESCiiNAE. Head sometimes with an elongate median spine between antennae; forefemora strongly swollen; pronotum constricted posterior to midpoint. First recognized by Fracker ^d Bruner (1924), this small group comprises five genera from Africa and the Neotropics, the New World Vescia Stal being the largest. Nothing is known of the life habits. Specialized morphology. Assassin bugs are a mor¬ phologically diverse group of dramatically varied facies. Not only do they possess a number of phenotypes unique to the group, but they “mimic” such diverse groups as the Aradidae, Berytidae, Coreidae, Enicocephalidae, Hydro-' metridae, Nabinae, Prostemmatinae, and Pyrrhocoridae. Most are distinctive for the necklike structure of the head behind the eyes (e.g.. Fig. 48.2B-D) and the strongly curving, short, inflexible labium (Fig. 48.3A, C), features that occur consistently elsewhere in the Het- eroptera only in the Enicocephalidae and that once were used to unite these two distantly related groups. Reduviids are further distinguished by their almost universal posses¬ sion of a prosternal stridulatory sulcus (Fig. 10.7F, G). As predators, reduviids often use their forelegs dur¬ ing prey capture (see Putchkov, 1987). Most Phymatinae have the femora greatly enlarged (Fig. 48.1), as they are to a lesser degree in the Peiratinae. Chelae formed by the juxtaposition of the femur and tibia have developed more than once in the group, and the tarsus is sometimes lost. In groups such as the Emesinae, the forefemur, although serving a raptorial function, is elongate and slender (Fig. 48.2B). The use of glandular setae on the forelegs for prey capture is unique to the assassin bugs. The fossula spongiosa, arising from the apex of the tibiae, shows a great diversity of occurrence and development in the Reduviidae. The Reduviidae share the possession of antennal tri¬ chobothria only with the Pachynomidae and Gerridae (Zrzavy, 1990a). The primitive reduvioid condition ap¬ pears to be a single trichobothrium (Fig. 48.3D), the situation found in the Pachynomidae and a relatively large number of reduviid subgroups and in the first-instar nymphs of some reduviids whose adults have multiple trichobothria; the maximum number may exceed 20 (Fig. 48.3E) (Wygodzinsky and Lodhi, 1989; Zrzavy, 1990a). The true spermatheca is transformed into a vermi¬ form gland (Fig. 48.3L, M) and is functionally replaced by paired pseudospermathecae (Fig. 48.3L, M). Many, but not all, reduviids possess Brindley’s glands (Brind¬ ley, 1930) (an analog of which apparently occurs in the Tingidae) and ventral glands (Carayon et al., 1958). Egg structure was examined by Haridass (1985, 1986a, b, 1988), including use of scanning electron microscopy. Miller ()956a) gave a survey of the many egg shapes found in the family. Natural history. All reduviids are predators. Most ap¬ pear to orient to their prey visually. The Triatominae are distinctive for the blood-sucking habits. Valuable sources on reduviid biology include the works of Miller (1953; based on the fauna of Zimbabwe), Miller (1956a, 1971), Readio (1927) on North America, Putchkov (1987) on the fauna of the Ukraine, and Louis (1974) on reduviids from cacao farms in Ghana. The works of Wygodzinsky (1966) and Lent and Wygodzinsky (1979) contain much valuable information of the Emesinae and Triatominae, respectively. Reduviidae 159 Most Phymatinae live on vegetation where, well cam¬ ouflaged, they lie in wait of their prey, toward which they appear to orient visually. Feeding and oviposition habits and nymphal morphology in Macrocephalus no- tatus Westwood have been described by Wygodzinsky (1944b). The life history of Phyrmta pennsylvanica americana Melin has been studied in detail by Balduf (e.g., 1941). Ectrichodiines are apparently all obligate predators of millipedes, rejecting insects and other arthropods, even when starved (Louis, 1974). Most live in leaf litter and are nocturnal (Miller, 1953; Louis, 1974). Ectrichodia gigas Herrich-Schaeffer in the Ivory Coast preys exclu¬ sively on the iulid millipede Peridontopyge spinosissima. This diurnal predator apparently does not produce a ven¬ omous salivary secretion, as the millipedes were not sub¬ dued for some time after attack commenced. Interaction between individual reduviids during rest and feeding peri¬ ods appears to be mediated by chemical stimuli, the bugs producing a distinctive odor, unlike that found in other heteropterans (Cachan, 1952a). Free-living Emesinae may be found in litter, under stones, on living vegetation, or in dead vegetation such as hanging ferns or palm fronds. Some species are domestic or peridomestic, a few being widely distributed, suggest¬ ing transport by humans. Some species are cavemico- lous but do not possess characteristics common to many cave dwellers, such as eyelessness or depigmentation. A substantial number of emesines are obligate inhabi¬ tants of spider or psocopteran webs, living there without becoming entangled or incurring the wrath of the web makers. Wygodzinsky (1966) thought that most spider¬ web-dwelling species are not obligate spider predators, but rather opportunists, whereas, Snoddy et al. (1976) reported Stenolemus lanipes Wygodzinsky feeding exclu¬ sively on the Achaearanea tepidariorum, and Cob- ben (1978) concluded that Stenolemus spp. are obligate spider predators. Triatomines commonly inhabit the nests or dwellings of their hosts, such as wood rats or other rodents in the American Southwest and Mexico. They may also be found under fallen logs, in caves, hollow trees, among palm fronds, and in bromeliads. Some species have be¬ come domestic or semidomestic, and may be found in chicken coops or the habitations of other domestic ani¬ mals. In association with humans they often inhabit the thatch of roofs of rustic dwellings or, in more modern structures, live in nearly any dark place where they can secrete themselves. They are strictly nocturnal, taking a blood meal for 20-30 minutes and inflicting no pain on the host. Trypanosomiasis is caused by the spreading of fecal material on the feeding lesions rather than by direct transmission. The nymphs of some Reduviinae, and some other re¬ duviids (Cetherinae, Salyavatinae, Triatominae), have the habit of covering the body with debris—including soil particles, pieces of termite nest carton, and corpses of ants and other insects—which is affixed by sticking ob¬ jects to the viscid secretions of specialized setae located on the dorsum (e.g., Odhiambo, 1958a). Camouflaged nymphs of Acanthaspis petax Stal stalk their prey; they commonly feed on ants, as well as other insects (Odhi¬ ambo, 1958b). Salyavata variegata Amyot and Serville was recorded eating the termite Nasutitermes exitiosus. Nymphs covered themselves with crumbs of nest carton, sat by perforations in the nest, and with their forelegs captured individual workers as they arrived to repair the nest. For bait, the bugs used the carcasses of termites on which they had fed, dangling them in the opening to at¬ tract workers, which were then dragged from the nest and eaten (McMahan, 1982). Nymphal and adult reduviids stridulate by rubbing the apex of the labium against the transversely striate proster- nal sulcus. All other Heteroptera capable of stridulation produce sound only in the adult stage. Reduviid stridula¬ tion has usually been regarded as defensive in function; however, reduviids emit other, nondefensive, noises (Go- gala and Cokl, 1983; Gogala, 1984), some of which are produced by the labial-prosternal mechanism and some by unknown means. They also probably emit low- frequency vibrations from an abdominal tymbal formed by fused abdominal terga 1 and 2. Phymata crassipes (Fabricius) is known for its alternating calls (Gogala and Cokl, 1983; Gogala et al. 1984) imitating in their dura¬ tion and frequency various acoustic stimuli and ranging in character from other insect calls to the human voice. Reduviids usually have platelike ovipositors, and many species—particularly in vegetation-dwelling groups such as the Harpactorini—glue their eggs to the plant, often in a group, sometimes covering the eggs with a gelati¬ nous material. A few species are known to guard their eggs. Ground-dwelling species may insert their eggs into the soil, or the eggs may be laid loose. Some reduviines have more well-developed ovipositors, suitable for the insertion of eggs into cracks or crevices. The glandular secretions from the abdominal trichome of the Holoptilinae are attractive—and apparently toxic— to ants, the prey of these bugs, as reported by Jacobson (1911) for Ptilocerus odiraceMj Montandon. Distribution and faunistics. The work of Readio (1927) on tlie biology of the North American fauna serves not only as a summary of biological information, but also as a review of the literature on the group for the region. The fauna of the western Palearctic is probably best docu¬ mented in the works of Dispons and Stichel (1959) for Europe and Putchkov (1987) for the Ukraine. Several fau- 160 TRUE BUGS OF THE WORLD (HEMIPTERA; HETEROPTERA) nistic works provide valuable aids in understanding the African fauna, notable among them Miller (1953), which contains many field observations as well as nine colored plates of habitus views. Other works include those of Vil- liers (1948) on West Africa, Schouteden (1931, 1932) and Villiers (1964) on the fauna of the Congo basin, and Villiers (1968, 1979) on the fauna of Madagascar. Many new tropical Asian taxa were described and illus¬ trated by Miller (1940, 1941). The fauna of China was detailed by Hsiao et al. (1981), a volume that includes many photographic illustrations. The catalogs of Putch- kov and Putchkov (1985, 1986-1989) and Maldonado (1990) should also be consulted. Velocipedoidea 49 Velocipediidae General. The velocipedids appear to have no common name, although they might be referred to as the fast¬ footed bugs. These tropical Asian bugs range in length from 10 to 15 mm and are distinctive for their elongate head and broadly expanded exocorium (Fig. 49.1 A). Diagnosis. Pronotum, scutellum, corium, and clavus rather heavily punctured; vestiture inconspicuous; head elongate with a neck behind eyes about length of diame¬ ter of an eye; eyes hemispherical, protuberant; ocelli present; antennal prepedicellite present; bucculae con¬ spicuously developed, more or less obscuring labial seg¬ ments 1 and 2; labium straight except for rather sharp angle between segments 2 and 3, reaching to about apex of mesocoxae, segment 1 very short, segment 2 short, segment 3 extremely long, segment 4 short (Fig. 49. IB, C); pronotum collarlike anteriorly, lateral margins of anterior lobe usually produced; exocorium explanate, medial fracture very long, costal fracture present (Fig. 49.1 A); membrane with 3 short, closed, basal cells with many simple or bifurcating veins emanating from them, and a stub (processus corial) located near apex of anterior cell (Fig. 49. ID); legs cursorial, femora and tibiae rela¬ tively long and nearly cylindrical; no fossula spongiosa; abdomen with distinct dorsal laterotergites in both sexes, ventral laterotergites developed only in males; abdominal spiracle 1 absent, spiracles 2-8 located on discrete ventral laterotergite in males and on sternum in females; abdo¬ men with a pair of trichobothrium-like setae on either side of ventral midline of segments 3-7; pygophore in male well developed and located at apex of abdomen (Fig. 49. IE); parameres symmetrical (Fig. 49. IG); vesica symmetrical, endosoma extremely simple (Fig. 49.IF); ovipositor laciniate, first valvula associated by ramus with first valvifer; spermatheca in form of a vermiform gland (omitted from Fig. 49. IH). Classification. Stal (1873) described the genus Sco- tomedes and placed it in the Nabidae. Bergroth (1891) subsequently described Velocipeda, placing it as a distinct subfamily, Velocipedinae, within the Saldidae. Later it was recognized that the two genera (as well as Godefri- dus Distant of the Reduviidae, Apiomerinae and Bloeteo- medes van Doesburg) were synonyms. At least six species have been described. Specialized morphology. The structure of the labium in the Velocipedidae (Fig. 49.IB, C) is similar to that found in the Mesoveliidae, Ochteridae. Aphelocheiri- dae, Saldidae, and Lasiochilidae (Rieger, 1976; Kerzh- ner, 1981). The broad expansion of the exocorium (Fig. 49.1 A) appears to be virtually unique within the Heterop- tera. Natural history. No direct observations have been published regarding the life habits of this group, although nymphs have been collected. Most known specimens have probably been collected at lights. Distribution and faunistics. Velocipedids are distrib¬ uted from Assam in Northeast India to New Guinea. No comprehensive papers exist on the group, although in addition to those cited above, one might see those by Blote (1945), van Doesburg (1970, 1980), and Kerzhner (1981). Microphysoidea 50 Micrcphysi(dae General. This group of tiny insects ranges in size from 1.5 to 3.0 mm, with many species resembling small an- thocorids (Fig. 50.1 A, B). They have no common name and are known in the field to only a few heteropterists. The small size, restricted distribution, and obscure habits of the Microphysidae have left the group little studied. Microphysidae 161 Fig. 49.1. Velocipedidae. A. Habitus, Scotomedes borneensis van Doesburg. B. Latetai view, S. borneensis (A, B from van Doesburg, 1970). C. Lateral view, head, S. alienus (Distant). D. Forewing, S. alienus (C. D from Schuh and Stys, 1991). E. Pygophore, S. alienus. F. Aedeagus, S. alienus. G. Parameres, S. alienus. H. Gynatrial complex, S. alienus (E-H from Kerzhrer, 1981). Diagnosis. Head weakly prognathous (Fig. 50.1C); macropterous forms with large ocelli; relative lengths of labial segments usually (shortest first) 1-4-3-2, with seg¬ ment 1 well developed in Palearctic species, reduced in Nearctic species; metapleuron without ostiolar groove or evaporatory area; single, heavy-veined cell in membrane of forewing, with a distinct stub (processus corial) in posterolateral angle (Fig. 50. ID); costal fracture well de¬ veloped, forming a distinct cuneus; distal sector of R+M in the hind wing branching to a fork (shared with the Nabinae); females of Palearctic taxa flightless, hemely- tra often staphylinoid; tarsi 2-segmented; male genitalia (Fig. 50. IE, F) symmetrical; ovipositor laciniate. Classification. The Microphysidae were first treated as a separate family by Fieber (1861) but were placed in an omnibus AnthoCoridae by Reuter (1884). Micro- physa Westwood, upon which the family name is based, is a junior synonym of Loricula Curtis. Although micro- 162 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 50.1. Microphysidae. A. Habitus, male, Loricula pselaphiformis Curtis. B. Habitus, female, L. pselaphiformis (A, B from Kelton, 1980a), C. Lateral view, head, Chinaola quercicola Blatchley, D. Forewing, Myrmedobia exilis (Fallen) (C, D from Schuh and Stys, 1991). E. Aedeagus, L pselaphiformis (from Pericart, 1972). F. Paramere, L. pilosella (from Miyamoto, 1965b). Abbreviation: pc, processus corial. physids superficially resemble members of the Cimico- idea, a relatively large number of characters suggest they are actually members of the Miriformes (Schuh and Stys, 1991). Five genera are currently recognized: Ciorulla Peri¬ cart (1974) (one species; Uzbekistan), Loricula Curtis (12 species; Palearctic), Myrmedobia Baerensprung (eight species; Palearctic), Chinaola Blatchley (one species; eastern North America), and Mallochiola Bergroth (one species; eastern North America and eastern Mexico). Specialized morphology. The Palearctic genera of Microphysidae are notable for their distinctive sexual di¬ morphism, manifested particularly in the extreme reduc¬ tion of the forewings in the females (Fig. 50. IB) of many species with complete loss of the membrane and exposure of 3-7 abdominal terga, giving a staphylinoid or myrme- comorphic habitus. The female abdomen is often greatly broadened and rounded laterally, caudad of the posterior margin of the hemelytra. Nearctic taxa are macropter- ous in both sexes. The structure of the labium of the taxa from the two areas is also different, with segment 1 greatly reduced in the New World taxa (Fig. 50.1C) and conspicuously present in Palearctic taxa. Ciorulla is unique among the microphysids in having the labial formula 2-3-1-4. Natural history. Most female microphysids are en¬ countered on the bark of trees, often those covered with lichens or mosses (Wheeler, 1992). The males and some¬ times females can often be collected on herbaceous vege¬ tation under trees harboring the females. Females of the Microphysidae 163 Palearctic species may also be found in decaying wood, and a few are found in wet litter, particularly Sphagnum containing fallen leaves and needles. Males and female are often found in separate habitats except during the period of reproductive activity. All species for which life histories are known are predatory and overwinter as eggs, with nymphal development taking place in the spring (Carayon, 1949; Kelton, 1980a, 1981). Distribution and faunistics. Published information shows the Microphysidae as being restricted to the Hol- arctic. Schuh and Stys (1991), however, noted examining specimens of an undescribed species from Namaqua- land. South Africa, considerably extending the previously known range. The work of Pericart (1972) is the most comprehensive treatment available for the family; other useful papers in¬ clude another study by Pericart (1969). Two Palearctic species, Loricula pselaphiformis Curtis and Myrmedo- bia exilis (Fallen), have been introduced into the eastern Nearctic (Kelton, 1980a, 1981). Joppeicoidea 51 Joppeicicjae General. This family includes only Joppeicus para¬ doxus Puton, a tiny species about 3.0 mm long and of anthocorid-like habitus (Fig. 51.1). The group has no common name and has long perplexed heteropterists with its peculiar combination of morphological attributes. Diagnosis. Dorsal surface weakly tuberculate; com¬ pound eyes relatively small, placed near posterior margin of head (Figs. 51.1, 51.2A, B); ocelli present; antennal segments 1 and 2 relatively short (Fig. 51.2A, B), seg¬ ment 4 with a stout apical seta; antenniferous tubercles set slightly anterior to eye at about level of ventral margin; labium in repose bent at right angle between segments 2 and 3, segment 1 very small, segment 3 about 3 times length of segment 2 and 1.5 times length of segment 4 (Fig. 51.2A, B); bucculae obscuring base of labium; pronotum carinate laterally, with a median keel on an¬ terior two-thirds (Figs. 51.1, 51.2A); forewing trough¬ like at Sc, R-l-M raised and keellike; costal and medial fractures absent (Figs. 51.1, 51.2C); membrane with- out stub (processus corial) and with very weakly devel¬ oped veins (Fig. 51.2C); M-Cu cross vein absent in hind wing (Fig. 51.2D); tarsi 2-segmented, fossula spongiosa absent; metathoracic scent gland with paired reservoirs and widely separated ostioles, grooves present on meta- pleuron; abdomen broadly membranous at junction with thorax, capable of 90° rotation, terga 1-3 membranous, sternum 2 a narrow strip (Fig. 51.2E), abdominal spiracle 1 absent, spiracles 2-8 located on abdominal sterna; nymphal dorsal abdominal scent glands located on an¬ terior margins of terga 4, 5, and 6 (Cobben, 1978); func¬ tional spermatheca absent (Fig. 51.2H); genital capsule distinct, parameres symmetrical; phallus in the form of sclerotized tube (Fig. 51.2F); ovipositor greatly reduced (Fig.51.2G). Classification. Joppeicus was originally placed in the Aradidae by its author Puton (1881) and later was trans¬ ferred to the Lygaeidae by Bergroth (1898). Reuter (1910) established a separate family within the Aradoideae and later (1912a) placed it in the Reduvioideae. Schuh and §tys (1991) treated it as a member of the Miriformes. 164 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 51.2. Joppeioidae. Joppeicus paradoxus. A. Dorsal view, head, pronotum, sculellum. B. Lateral view, head (A, B from China, 1955b). C. Forewing (from Davis and Usinger, 1970). D. Hind wing (from China, 1955b). E. Lateral view, abdomen. F. Phallus. G. Female genital segments, lateral view. H. Ovaries and nonfunctional spermatheca (E-H from Davis and Usinger, 1970). Specialized morphology. Two intensive studies of the morphology of Joppeicus (China, 1955b; Davis and Usinger, 1970) revealed that many of the attributes of Joppeicus, such as the structure of the wings, phallus, and the large intersegmental membrane between abdominal segments 3 and 4, are unique to the taxon. Natural history. Joppeicus is a general predator, in culture thriving on a wide range of prey items (Davis and Usinger, 1970). The rostrum is held out straight in front of the body when prey is being attacked. In nature, Joppeicus prefers marginal, often dusty situations, little frequented by other insects. It has been collected on the surface of the ground, under stones, in shallow caves, under bark, and also in colonies of bed bugs associated with bats (Stys, 1971). When mating, the male first mounts the dorsum of the female, rotates to the right or left—as seen in profile— then rotates the abdomen 90° at the large intersegmen¬ tal membrane between segments 3 and 4 and moves the dorsal surface under the abdominal venter of the female, at which time copulation actually takes place (Davis and Usinger, 1970). Distribution and faunistics. Joppeicus is restricted to southern Israel and the Nile and Blue Nile drainages in Egypt and the Sudan. It has been collected most com¬ monly in the Nile delta region (Stys, 1971). Miroidea 52 Thau mastoco ri d ae General. This group of morphologically unusual and seldom collected bugs has no common name, although the New World members might be collectively referred to as palm bugs. All species are small, ranging from 2.0 Thaumastocoridae 165 to 4.6 min, and are distinguished by the strongly anteri¬ orly produced mandibular plates (Fig. 52.1 A, B) and the grossly asymmetrical male genitalia. Diagnosis. Body moderately to strongly flattened; dorsum often heavily punctured, pubescence short and inconspicuous; head with mandibular plates and clypeus projecting in front of eyes, mandibular plates often ex¬ ceeding clypeus (Figs. 52.1 A, 52.3A); ocelli present; antennae relatively short, segments 3 and 4 at most weakly fusiform; labium inserted on ventral surface of head, reaching from anterior margin of prostemum to near base of abdomen (Fig. 52.3A, B); prostemal stridu- latory sulcus incorrectly indicated by Drake and Slater (1957; see Fig. 52.3B); legs homomorphous, relatively short, femora mutic; tarsi 2-segmented; macropterous or brachypterous; costal fracture absent; stub (proces¬ sus corial) absent; metathoracic scent-gland chaimels present; dorsal and ventral laterotergites absent, abdomi¬ nal spiracle 1 present; abdominal spiracles 2-8 situated on sternum; dorsal abdominal scent glands present in nymphs on anterior margin of terga 4 and 5; genital capsule grossly asymmetrical (Fig. 52.3F) (usually dex- tral or sinistral in a given species, rarely direction vari¬ able within a species); one or both parameres lost (Fig. 52.2G); phallus as in Fig. 52.2F; ovipositor completely absent; spermatheca absent. Classification. The first described thaumastocorids were treated as a subfamily of the Lygaeidae (Kirkaldy, 1908). The group was accorded family status by Reuter (1912a). The classification presented here follows Drake and Slater (1957) in placing the Thaumastocoridae in the Cimicomorpha, whereas Cobben (1978) treated the group as of problematic position. Two subfamilies are recog¬ nized, the Xylastodorinae and Thaumastocorinae. Viana and Carpintero (1981) treated these as distinct families, an approach rejected by Slater and Brailovsky (1983), be¬ cause this action failed to establish new criteria for group recognition and obscured the novel morphological fea¬ tures shared by the two groups. Drake and Slater (1957) related the Thaumastocoridae to the Cimicoidea, whereas Kerzhner (1981) treated them as the sister group of the Tingidae, and Schuh and Stys (1991) as the sister group of the Miiidae -I- Tingidae. Fig. 52.1. Thaumastocoridae. A. Onymocoris izzardi Drake and Slater (from Drake and Slater, 1957). B. Oiscocoris drake/ Slater and Ashlock. Fig. 52.2. Thaumastocoridae. A. Habitus, Xylastodoris luteolus Barber (from Slater and Baranowski, 1978). B. Dorsal view nymph, X. luteolus. C. Ventral view head and thorax, X. luteolus. D. Forewing, X. luteolus. E. Hind wing, X. luteolus (B-E from Schaefer, 1969). F. Aedeagus, Baclozygum depressum Bergroth. G. Paramere, Thaumastocoris hacker! Drake and Slater (F, G from Drake and Slater, 1957). Key to Subfamilies of Thaumastocoridae 1. Claws bearing large, basally attached pulvilli (Fig. 52.3E); tibiae apically without a lobate appendage (tibial appendix); males without parameres; costa extending to near apex of forewing (Figs. 52. IB, 52.2D) . Xylastodorinae - Claws without pulvilli; tibiae bearing a lobate appendage apically (tibial appendix) (Fig. 52.3C, D); males with one peiramere (Fig. 52.2G); membrane extending greatly beyond apex of corium . . Thaumastocorinae dominal scent glands with paired openings (Fig. 52.2B); both parameres lost. Two genera are currently recognized; Discocoris Kor- milev (five species, South America) and Xylastodoris Bar¬ ber (one species, Cuba and Florida). They were originally described in separate subfamilies but treated as belonging to a single subfamily by Drake and Slater (1957). Specialized morphology. The thaumastocorids are tiny, strongly flattened bugs that hold tenaciously to the substrate (Slater, 1973; Schuh, 1975b), apparently with the aid of the accessory pretarsal structures such as the tibial appendix (Fig. 52.3C, D) in the Thaumastocorinae and the large pulvilli (Fig. 52.3E) in the Xylastodori¬ nae. Most species have greatly enlarged and extended mandibular plates (Fig. 52.1 A, B), which surpass the THAUMASTOCORINAE. Eyes strongly protuberant (Fig. 52.1 A); tibial appendix apically on all tibiae (Fig. 52.3C, D); nymphal dorsal abdominal scent glands with a single opening (Slater, 1973); one paramere lost. Currently four genera are included: Baclozygum Ber¬ groth (two species; Tasmania and mainland Australia), Onymocoris Drake and Slater (three species; Australia), Thaumastocoris Kirkaldy (three species; Australia), and Wechina Drake and Slater (one species; southern India). XYLASTODORINAE. Antcnniferous tubercle conspicu¬ ously projecting (Figs. 52.2C, 52.3B); large pulvilli aris¬ ing from base of claw (Fig. 52.3E); nymphal dorsal ab- The key to the subfamilies of Thaumastocoridae is modified from Drake and Slater, 1957. Thaumastocoridae 167 168 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) apex of the clypeus, a situation generally found elsewhere in Aradoidea and Pentatomoidea. The asymmetrical and otherwise highly modified male genitalia (Figs. 52.2F, 52.3F), with the parameres being completely lost in the Xylastodorinae, are unique. Natural history. Thaumastocorids are phytophagous. Thaumastocorinae are known to feed on a variety of dico¬ tyledonous plants. Baclozygum depressum Bergroth and B. brevipilosum Rose have been collected on Eucalyp¬ tus trachyphloia, B. depressum on E. globosus (Myrta- ceae) (Hill, 1988), Onymocoris hackeri Drake and Slater on Banksia sp. (Proteaceae) (Drake and Slater, 1957; Rose, 1965) and Elaeocarpus obovatus (Elaeocarpaceae) (Rose, 1965), and Thaumastocoris australicus Kirkaldy on Acacia cunninghami (Fabaceae) (Kumar, 1964) and A. maidenii (Slater, 1973). Xylastodorinae feed only on palms, Discocoris spp. on a variety of taxa, including Eutorpe edulis (Kormilev, 1955a), Phytelephas sp. (Schuh, 1975b), and Socratea montana (Slater and Schuh, 1990), apparently always on the inflorescences (Kormilev, 1955a) or infructescences (Schuh, 1975b; Slater and Schuh, 1990). Xylastodoris feeds on the developing fronds of the royal palm, Roy- stonea regia (Barber, 1920; Baranowski, 1958), where it sometimes causes serious damage. Distribution and faunistics. The work of Drake and Slater (1957) is still the single most important publication on the Thaumastocoridae, but considerable knowledge concerning the diversity and distribution, particularly of the New World fauna, has been gained since the publi¬ cation of that work, the most important references being those cited above. 53 Miri(dae General. The Miridae is the largest family of true bugs. In the United States and Canada members of the group are usually called plant bugs, in Britain capsids, and in Germany Blindwanzen. Ranging in size from less than 2 mm to about 15 mm, they are among the most delicate of all bugs, and often their coloration blends well with the foliage, flowers, and bark on which they rest or feed. The habitus varies from simple ovoid to remarkably myrmecomorphic. Diagnosis. Size and appearance variable; ocelli absent (except Isometopinae); antennal segments 3 and 4 usually Fig. 53.1. Miridae, A. Magnocellus transvaalensis Slater and Schuh (isometopinae) (from Slater and Schuh, 1969). B. Carvalhoma mal- colmae Slater and Gross (Cylapinae) (from Schuh and Schwartz, 1984). Miridae 169 Fig. 53.2. Miridae. A. Froeschnerana mexicanus Schaffner and Fer¬ reira (Deraeocorinae: Clivinemini) (from Schaffner and Ferreira, 1989). B. Phytocoris populi (Linnaeus) (Mirinae) (from Stonedahl, 1983). C. Orectoderes obliquus Uhler (Phyiinae), male. D. O. obliquus, female (C, D from Mclver and Stonedahl, 1987b). Fig. 53.3. Miridae. A. Psallops oculatus Carvalho. B. Frontal view, head, P. oculatus (A, B from Carvalho, 1956). C. Frontal view, head, Pilophorus /(ocir^ocorinae 5. Metastemum produced anteriorly onto mesosternum or rarely prosternum; labium not surpassing mesocoxae . Edessinae - Mefesternum rarely produced anteriorly onto mesosternum, if so then labium extending onto abdo¬ men; labium usually reaching at least to metacoxae . 6 6. Trichobothrium nearest spiracle on sternum 7 laterad of spiracular line by distance at least equal to greatest diameter of spiracular opening . 7 - At least one trichobothrium on sternum 7 on or mesad of spiracular line . 8 7. Base of abdominal venter with mesal tubercle; metasternum produced, flattened . . Pentatominae (part) - Abdominal venter rarely tuberculate at base, if so then metasternum thinly carinate mesally . . Discocephalinae (part) 8. Labium arising on or behind imaginary line traversing head at anterior limit of eyes and/or superior surface of tarsal segment 3 of hind legs shallowly excavated in females .... Discocephalinae (part) - Labium arising before such a line; superior surface of tarsal segments convex or flattened . 9 9. Tibiae sulcate on outer surface; labial segment 1 longer than bucculae; trichobothria paired; frena one-third or more length of scutellum; scutellum not reaching apex of abdomen . . Pentatominae (part) - Tibiae not sulcate on outer surface; labial segment 1 not longer than bucculae; trichobothria single; frena short, less than one-third length of scutellum; scutellum usually U-shaped, reaching apex of abdomen . Podopinae ASOPINAE. Labial segment 1 thickened, usually free from bucculae, rarely lying within bucculae for its entire length, in which case foretibiae foliate {Cecyrina Walker, Heteroscelis Latreille); males often with pair of large 230 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 73.2. Pentatomidae. A. Genital capsule, Heteroscelis lepida Stal. B. Genital capsule, Sciocoris microphthalmus Flor, C. Aedeagus, lateral view, H. lepida. D. Aedeagus, sagittal view, S. microphthalmus (A-D from McDonald, 1966). E. Aedeagus, sagittal view, Stagonomus amoenus (Brulle). F. Ejaculatory reservoir, S. amoenus (E, F from Seidenstiicker, 1965b). G. Paramere, H. lepida. H. Paramere, Sciocoris microphthalmus (G, H from McDonald, 1966). I. Spermatheca, Theseus modestus (Stal) (from Pendergrast, 1957). brown because of dense secretion covering body; tarsi 2-segmented. First recognized as a higher group by Distant (1880- 1893), the systematic position of this group is ambiguous. We treat it as a highly modified subfamily of Pentatomi¬ dae, although Brailovsky et al. (1988), who provided a modern description, treated it as a distinct family. The group is Neotropical and consists of four genera and 11 species. DiscocEPHALiNAE. Body flattened; coloration mottled brown, black, fuscous, or brown and black mottled; an¬ tennae either 4- or 5-segmented; labium usually arising on or posterior to an imaginary line traversing head at anterior margin of eyes; metasternum not produced an¬ teriorly onto mesosternum; tarsi 3-segmented; dorsal sur¬ face of tarsal segment 3 of hind legs usually excavated pilose sensory patches, these often extending across at least part of last 3 pregenital abdominal sterna; pygopho- ral plate located interior to (mesad of) each paramere. This group was first recognized by Amyot and Ser- ville (1843) as the “Spissirostres,” although most authors credit it to Spinola (1850) as the Asopoideae. Asopines are found in all faunal regions. There are ap¬ proximately 63 genera and 357 species known (Gapud 1981). The work of Thomas (1992) is the most current for the New World fauna. All whose biology is known are predatory. CYRTOCORiNAE. Cryptically colored, resembling tree bark; antennae 4-segmented; pronotum and abdomen strongly expanded laterally, mostly covered by enlarged scutellum, the latter produced mesally into a strong, pro¬ jecting spine; adults bright black, but appearing dull Pentatomidae 231 in females; trichobothrium nearest spiracle on sternum 7 usually laterad of spiracular line; phallotheca, ejaculatory duct, median penial lobes, and conjunctival appendages (when present) heavily sclerotized. last fused to margin of phallotheca and permanently exserted. First recognized as a higher group by Fieber (1861), the Discocephalinae, which are speciose in the Neotropics, comprise 71 genera and at least 263 species. Two tribes are recognized: the Ochlerini. erected by Rolston (1981) (23 genera), and the Discocephalini (48 genera). EDESSiNAE. Large to very large; smooth, polished dorsal surface; appearance rather streamlined; antennae either 4- or 5-segmented; metasternum strongly pro¬ duced, extending anteriorly onto mesosternum and sometimes prostemum and laterad between meso- and metacoxae; anterior projection of metastemal plate bi¬ fid (except in Pantochlora Stal), labium terminating in this notch; posterior margin of metasternum notched to receive mesal tubercle of abdomen; tarsi 3-segmented. First recognized as a higher group by Amyot and Ser- ville (1843), edessines until recently were considered a tribe of the Pentatominae, but were elevated to subfamily status by Rolston and McDonald (1979), although Gapud (1991) treated them at a tribal level. These large, robust stink bugs are abundant and diverse in the Neotropics, where four genera and 269 species are known. PENTATOMINAE (FIG. 73.1). Varied in form and color, usu¬ ally obovate, often with prominent caudolateral pro- notal angles; scutellum never attaining apex of abdomen; frena extending two-fifths or more length of scutellum; metastemum rarely produced anteriorly onto mesoster¬ num, when so labium reaching metacoxae; trichobothria, usually at least one or a pair on each side of sterna 3-7, on or near spiracular line; dilation of spermathecal duct fusiform. (For more details regarding genitalic structures see Rolston and McDonald, 1979.) This is the largest subfamily of Pentatomidae. Eight tribes are recognized: Aeptini (11 genera, 30 species); Diemeniini (13 genera, 47 species); Halyini Amyot and Serville (82 genera, 361 species); Lestonocorini (5 gen¬ era); Mecideini Distant (1 genus, 17 species); Myrocheini (14 genera, 45 species); Pentatomini Leach (404 genera, 2207 species); Sciocorini Amyot and Serville (10 genera, 107 species). Tribal relationships are under active investi¬ gation. PHYLLOCEPHALiNAE. Large, flattened; labium not ex¬ tending beyond posterior margin of forecdxae. Thirty-one genera and 175 species are known. Most species appear to live on the bark of trees. Miller (1956a) stated that Tetroda histeroides (Fabricius), an important rice pest, belongs to this subfamily. PODOPiNAE. Coloration generally dark yellow brown to dark brown or nearly black; antenniferous tubercles visible from above; antennae either 4- or 5-segmented; lateral margins of pronotum usually toothed or tubercle¬ bearing; tarsi 3-segmented; trichobothria usually paired, single in some Podopini. behind each spiracle on or near spiracular line; scutellum enlarged, always attaining membrane of forewing and often reaching apex of abdo¬ men. covering most of forewings; frena well developed but extending less than one-third length of scutellum; hamus absent; R-fM and Cu of hind wing parallel (Les- ton. 1953a); parameres biramous; pygophore often with appendages on caudolateral margin. This widely distributed subfamily has had a checkered history varying from subtribe to family status. Many lit¬ erature references are under the name Graphosomatinae, a polyphyletic group with some of its members belonging to the Podopinae and most of the others to the Penta¬ tominae. Schaefer (1981c) discussed details of the group, recognizing two tribes, the Podopini and the Graphoso- matini. Sixty-four genera and 255 species are known. Podopines are found chiefly in damp marshy or muddy habitats. Some species frequently come to lights. Certain species of Scotinophara Stal are serious pests of rice and other cereal crops in Asia and Africa. SERBANiNAE. Body flattened, with broad lateral folia¬ tion on head, pronotum, and abdomen; coloration cryptic (resembling Phloeidae in habitus); antennae 4-segmented; tarsi 3-segmented; compound eyes divided into dorsal and ventral portions as in Phloeidae; male conjuncti¬ val appendages largely membranous, paired, phallotheca basally membranous, distally sclerotized. The subfamily was originally described by Distant in the Phloeidae but later was removed and established as a higher taxon within the Pentatomidae by Leston (1953c). A single species is known, Serbana borneensis Distant, from Borneo. Nothing appears to be known of the habits of these rare insects, but they probably will be found to live on the bark of trees. Specialized morphology. Pentatomids (and aphylids) are distinguished from all other Pentatomomorpha by the form of the spermatheca, which is invaginated proximal to the pump with a sclerotized median wall (Figs. 67.2, 73.21). The scutellum of pentatomids is triangular (Fig. 73.1). although it is sometimes enlarged to cover almost the entire abdomen, as in Podopinae. The 5-segmented antennae found in most taxa are also a specialized feature, as is the flattened platelike female ovipositor. Some stink bugs have stridulatory structures involving the abdominal dorsum and hind wing. Wing reduction is very rare in the Pentatomidae. In the Western Hemisphere it occurs in only two species— Brachelytron angelicas Ruckes from Brazil and Alathe- tus haitiensis Rolston from Haiti. The latter species has short, truncate padlike wings. Rolston (1982a) stated that 232 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) members of the tribe (Ochlerini) live in the canopy of Neotropical forests and surmised that with the deforesta¬ tion of Haiti A. haitiensis may be extinct. But because it was collected by P. J. Darlington at 5000 feet, the pre¬ sumed arboreal habitat is in need of verification. Miller (1956a) illustrated Tahitocoris cheesmanae Yang (Asopi- nae), which is cimicoid in outline and apparently apter¬ ous. Natural history. Most pentatomids are plant feeders and have a distinct preference for immature fruits and seeds. Many also feed in the plant vascular system. Only members of the Asopinae are predaceous, apparently sec¬ ondarily. They feed on a great variety of insects and other small arthropods; the North American Podisus maculiven- tris (Say) has been reported feeding on over 90 species of insects. The majority prefer the larvae of Lepidoptera and Coleoptera. Several species of Asopinae are of consider¬ able importance as biological control agents of destruc¬ tive insects. Perillus bioculatus (Fabricius) is extremely variable in color, the red color being a carotin pigment obtained from the Colorado potato beetle. If the pigment is not metabolized rapidly enough, the body color will change from white or yellow to red. The deposition of the pigment is primarily controlled by temperature, which af¬ fects the physiological activity of the bug (Knight, 1922, 1924; Palmer and Knight, 1924). The phytophagous pentatomine Thyanta calceata (Say) has a green summer generation with short pubescence and a brown autumnal-vernal generation with long pubes¬ cence. This variation is due to a developmental photo¬ period response. The adults are capable of color reversal in both directions (McPherson, 1977). One of the striking biological features of some penta¬ tomids (and other pentatomoids) is the protection of the eggs by the females. In several species this behavior is elaborately developed: the adult female not only wards off potential predators but moves her body from side to side to inhibit egg laying by small parasitic wasps. After the eggs hatch, the females of some species guard the first-instar nymphs, which apparently do not feed. Egg guarding has arisen several times independently within the family. Eberhard (1975) stated that it is known in 12 genera and 14 species, but this figure is certainly too small because he mentions parental behavior in five species of Antiteuchus Dallas alone. The most detailed work on maternal care is that of Eberhard (1975) on Antiteuchus tripterus limbativentris Ruckes (Discocephalinae) in Colombia. He found that these insects protected both the egg masses and first- instar and early second-instar nymphs. The protection is effective against general predators. But interestingly it appears to be detrimental in the case of two scelio- nid wasp egg parasites that are attracted by the females over the eggs; despite defensive behavior by the females a high proportion of the eggs are parasitized. Eberhard presented detailed information concerning differences in wasp attack behavior. This pentatomid. as well as sev¬ eral other species in the genus, is a serious pest of cacao and mango and is a vector of a fungus that attacks cacao. Interestingly, the bug also is a folk cure of intestinal para¬ sites of humans. Oddly, Eberhard found the bugs to be more common in the city of Cali than in the countryside, and they were most common where heavy pedestrian and automobile traffic was present. There are many striking cases of protective resem¬ blance in the family, especially in those species that live on bark, some of which assume bizarre shapes as well as highly cryptic patterns. This adaptation reaches its apo¬ gee in the Cyrtocorinae, which cover themselves with a dense brownish white secretion that, together with the great expansions of the pronotum and abdomen, renders them nearly invisible on the bark of the principle host plants. Species of many other genera sueh as Brochy- mena Amyot and Serville are cryptically colored and also may resemble the bark of trees. Brightly colored species are thought to be aposematic, although the basis for the warning coloration has not been well established for most of them. In Cyrtocoris trigonus (Germar), which feeds primarily on Acalifa diversifolia (Euphorbiaceae), nymphs form large aggregations of up to 700 individuals on a single plant. Some maternal care appears to occur (Brailovsky etal., 1988). Distribution and faunistics. The family is worldwide in distribution and well represented in all of the major faunal regions. As with most of the large families of Het- eroptera, the tropical and subtropical faunas are the most extensive. Useful identification aids include Gross (1975-1976) for Australia, Cachan (1952b) for Madagascar; Rolston and McDonald (1979, 1981, 1984) and Rolston et al. (1980) for keys to Western Hemisphere subfamilies and for tribes of Pentatominae; McPherson (1982) for the east¬ ern North American fauna; Schouteden (1903, 1905bj for world genera and African species of Podopinae; Ga- pud (1991) for world genera of Asopinae; and Thomas (1992) for the Asopinae of the Western Hemisphere. Schouteden’s (1907) revision of the world genera of Aso¬ pinae contains excellent color plates. Other useful studies of smaller groups include Freeman’s (1940) revision of Nezara Amyot and Serville, Rolston’s (1974, 1982b, 1984) revisions of Euschistus, and Ruckes’s (1947) re¬ vision of B rochymena. Pentatomidae 233 74 Phloeidae General. These are bizarre, large, flattened, brownish insects with the lateral body margins strongly expanded (Fig. 74.1 A, B) and with obvious protective shape and coloration associated with life on the bark of trees. They range in size from 20 to 30 mm and have no common name. Diagnosis. Body extremely depressed, with outer margins of mandibular plates, pronotum, base of corium, and abdomen broadly foliate (Fig. 74.1 A, B); eyes di¬ vided into a dorsal and ventral portion; bucculae long, low posteriorly, with labial channel very elongate; an¬ tennae 3-segmented, with segment 1 very long, segment 3 somewhat curved (Fig. 74.2A), antennal segments almost hidden below expanded mandibular plates; post- frenal portion of scutellum very elongate; scutellum not covering forewing; membrane extensively reticulate; hind wing with hamus; tarsi 3-segmented; abdominal sterna 3-7 with trichobothria arranged longitudinally mesad of spiracular line; metathoracic scent-gland opening near lateral margin of pleuron; dorsal abdominal scent-gland openings of nymphs present between terga 3/4, 4/5, 5/ 6 (Fig. 74. IB), those between 3/4 and 4/5 paired, those between 5/6 coalesced into a single opening (anterior gland sometimes lacking); abdominal connexivum with terga and sterna fused, no inner laterotergites; spiracle 2 present and partially exposed; ninth paratergites greatly elongated; genital capsule as in Fig. 74.2B; aedeagus with 3 pairs of conjunctival appendages (Fig. 74.2G; see Lent and Jurberg, 1965 for details); ovipositor plate¬ like, second valvifers fused medially (contra Lent and Jurberg [1965] and Gapud [1991]); spermatheca with well-defined pump region with flanges (Fig. 74.2F, H); parameres as in Fig. 74.2E; egg lacking a pseudopercu- lum. Classification. This group was first recognized as a taxon above the generic level as the “Phleides” by Amyot and Serville (1843) and was treated as a sub- 234 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 74.2. Phoeidae. A. Antenna, Phloeophana longirostris (Spinola). B. Male genital capsule, Phloeophana longirostns. C. Aedeagus, sagittal view, Phloeophana longirostris. D. Aedeagus, lateral view, Phloeophana longirostris (A-D from Leston, 1953c). E. Parameres, Phloea corticata (from Lent and Jurberg, 1965). F. Spermatheca, Phloeophana longirostris (from Pendergrast, 1957). G. Conjunctiva, sagittal view, Phloeophana longirostris. H. Spermatheca, Phloea subquadrata Spinola (G, H from Lent and Jurberg, 1965). family of Pentatomidae (the Phloeina) by Stal (1872) and others. Leston (1953c), Lent and Jurberg (1965), and Rolston and McDonald (1979) treated it as worthy of family status. Phloeophana Kirkaldy and Phloea Lepele- tier and Serville—together containing three species—are recognized. Specialized morphology. The 3-segmented antennae (Fig. 74.2A), strongly foliate body margins (Fig. 74.1 A, B), and unique aedeagal morphology (Fig. 74.2C, D, G) are novel to the group (Lent and Jurberg, 1965). Natural history. These curious insects are apparently phytophagous. They are wonderfully camouflaged to ap- Phloeidae 235 pear to be pieces of lichen on tree trunks or branches (Lent and Jurberg, 1966). They defend themselves when disturbed by “shooting” a liquid from the scent-gland ostioles a considerable distance from the body. Hussey (1934) summarized the early accounts for Phloeophana longirostris (Spinola). He quoted Magalhaes (who first reported maternal care in this species in a local news¬ paper) as stating that the females protect not only the eggs but also the first-instar nymphs, which attach themselves to the venter of the parent and are carried by her for many days. Magalhaes thought that the young nymphs were fed by the parent, but it is probable that this is merely another case of nonfeeding by the first-instar nymphs. Other ac¬ counts indicate maternal care through three instars (see Leston, 1953c, for a detailed discussion of conflicting biological observations). Distribution and faunistics. The group is restricted to South America. Basic sources are those by Hussey (1934), Leston (1953c), and Lent and Jurberg (1965). 75 Plataspidae General. The members of this family are beetlelike in appearance, being ovoid or suborbicular and strongly convex, with the scutellum greatly enlarged to cover almost the entire abdomen (Fig. 75.1). Many species are as broad or broader than long—ranging in length from 2 to 20 mm—and some have the mandibular plates devel¬ oped into horns (Fig. 75.1). Diagnosis. Head flattened and keeled; antenniferous tubercles located below lateral margins of head, not visible from above; antennae appearing 4-segmented (division between segments 2 and 3 weakly developed); labial segment 2 sometimes much enlarged, flattened, and saclike, with stylets partially coiled (Fig. 75.2A); labium often swollen and sometimes also clypeus; scu¬ tellum completely covering abdomen (Fig. 75.1); wings complexly modified, forewings much longer than body Fig. 75.1. Plataspidae. Ceratocoris sp. (from Slater, 1982). (Fig. 75.2B); hind wings specialized to allow folding below scutellum by transverse constrictions (Fig. 75.2C); tarsi 2-segmented; abdominal sterna with a straight trans¬ verse sulcus on each side; nymphs with dorsal scent-gland openings between terga 3/4, 4/5 and 5/6, those between 3/4 sometimes much reduced; aedeagus as in Fig. 75.2D, E; parameres as in Fig. 75.2F; spermatheca with well- developed pump and 2 flanges as in Fig. 75.2G, H. Classification. This taxon was first mentioned as a higher group by Dallas (1851-1852). Fieber (1861) used the name Arthropteridae, Kirkaldy (1909) the name Cop- tosominae, and Leston (1952) the name Brachyplatidae. Coloration, enlarged scutellum, and wing folding suggest relationships to the Aphylidae, Canopidae, Lestoniidae, Megarididae, and Scutelleridae. The origins and occur¬ rences of these attributes, as well as other characteristics shared by at least some of these groups, have not been investigated using cladistic methods, and thus their status as synapomorphies remains unclear. Key to Generic Groups of Plataspidae 1. Ocelli placed near eyes; ratio of distance between eyes and ocelli to interocellar distance less than 1:2; abdominal sterna usually convex; head usually narrow, approximately 0.3-0.5 times width of pronotum; base of scutellum (pseudoscutellum) usually raised, demarcated from rest of scutellum by an impressed line .. Coptosoma Group 236 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) ■( Fig. 75.2. Plataspidae. A. Mouthparts, Bozius respersus Distant (from China, 1931). B. Forewing, Neocratoplatys salvazai Miiler. C. Hind wing, N. salvazai (B, C from Miller, 1955). D. Aedeagus, laterai view, Coptosoma scutellatum (Geoffroy). E. Aedeagus, sagittai view, C. scutellatum. F. Paramere, C. scutellatum (D-F from Davidova and Stys, 1980). G. Spermatheca, C. inclusa Stal (from Pendergrast, 1957). H. Spermatheca, C. scutellatum (from Davidova and §tys, 1980). - Ocelli placed near one another, ratio of distance between eyes and ocelli to interocellar distance greater than 1;2; abdominal sterna not, or very slightly, convex; head transverse, usually 0.5-0.7 times width of pronotum; pseudoscutellum absent or weakly developed . 2 2. Body flattened; color black, sometimes spotted with yellow, often with a yellow submarginal line on head, pronotum and scutellum . Brachyplatys Group - Body usually convex; color pattern red, yellow, or brown, maculated with dark brown to black punctures; dark areas sometimes extensive and spotted or flecked with yellow, without a yellow submarginal line on head, pronotum, and scutellum . Libyaspis Group Specialized morphoiogy. In some species, the labial segments, and sometimes clypeus, are swollen, in which case the stylets extend backward through the thorax into the base of the abdomen (China, 1931). The complex folding of the wings (Fig. 75.2B, C) and the greatly enlarged and subtruncate scutellum are obvious specializations of the family. Several tropical African species reach a size of 15- 20 mm, and many of these species are strongly sexu- Key to generic groups of Plataspidae adapted from Jessop. 1983. ally dimorphic. The males have enormously produced mandibular plates in the form of “horns” (Fig. 75.1). which may be bifid as in the genus Ceratocoris White or enormously prolonged as in Severiniella Montandon. Although the function of these horns is unknown, it seems likely that they are involved in territorial defense. Natural history. China (1931) believed that some plataspids must be mycophagous because they possess very long stylets similar to those found in the Aradi- dae and Termitaphididae. But Maschwitz et al. (1987) showed a symbiotic relationship between two Tropidoty- Plataspidae 237 lus spp. and the ant Meranoplus mucronatus, wherein the ants tended and protected the bugs for their substantial honey dew secretions, indicating that the bugs are phloem feeders. These authors postulated that the long stylets function in reaching the phloem through the thick bark of the dicotyledonous host tree. Other species of the family are known to attack cowpeas and other legumes in Asia, the Pacific, and Australia. Monteith (1982b) reported aestivation in large clus¬ ters by Coptosoma lyncea Stal in the monsoon forests of northern Australia. He described one aggregation on a tree 5 meters in height on which every leaf was covered with the insects. They assumed a regular spacing on the underside of the leaves and were arranged in rows along the petioles, giving the impression of galls or blemishes on the plants. When disturbed large numbers took flight instantly, buzzing loudly and producing a discharge of the stink glands. After several minutes all returned to their original roosting tree and became quiescent again. (See also Natural History, Chapters 88 and 89). Distribution and faunistics. Plataspids are largely re¬ stricted to the tropics and subtropics of the Eastern Hemi¬ sphere, but a few species of Coptosoma Laporte occur in the temperate Palearctic. A basic source on the group is that of Jessop (1983), which provides keys to generic groups and a detailed dis¬ cussion of the Libyaspis Group. Davidova-Vilimova and Stys (1980) keyed the Western Palearctic Coptosoma spp. 76 Scutelleri(dae General. These insects, which range in length from 5 to 20 mm, are often known as shield bugs because of the greatly enlarged convex scutellum that usually en¬ tirely covers the abdomen (Fig. 76.1 A, B). Some tropical species are vividly colored, even becoming iridescent, and others have a striking variety of strongly contrast¬ ing reds, blues, and yellows. Thus, some scutellerids are among the most spectacularly colored of all Heteroptera. Diagnosis. Antennae 3- or 5-segmented (Fig. 76.1 A); scutellum greatly enlarged and convex, covering or nearly covering entire abdomen (Fig. 76.1 A, B); frena obsolete or lacking; strongly laminate propleural carinae present; forewing membrane with numerous veins; tarsi 3-seg- mented; prosternal sulcus present; sutures of abdominal venter extending to lateral margins; trichobothria paired; Fig. 76.1. Scutelleridae. A. Pachycoris torridus (Scopoli). B. Acantho- lomidea sp. aedeagus as in Fig, 76.2A, B; parameres as in Fig. 76.2C; second valvifers completely fused; first valvulae with re¬ duced rami and largely membranous (Fig. 76.2D); base of spermatheca with a sclerotized groove and an enlarged bulb proximal to flanges (Fig. 76.2E, F). Classification. The taxon was first established by 238 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Leach (1815) as Scutellerida. Lattin (1964), Kumar (1965), and McDonald and Cassis (1984) reviewed the history of the group. They noted that Amyot and Ser- ville (1843) were the first to subdivide the family (as a subfamily) and that they were followed by Dallas (1851- 1852), Stal (1872), and Schouteden (1904-1906). Leston (1952) discussed the tribal arrangement, but treated the taxon as a subfamily, as did Lattin (1964). McDonald and Cassis (1984) stated that Van Duzee (1917) was the first to accord the group family status. But Fieber (1861) recog¬ nized a family level taxon as Tetyrae, and Stal (1867) used Tetyridae for the family name. Uhler (1863) recognized the group under the family name Pachycoridae. Dupuis (1947), Pendergrast (1957), and Kumar (1962) all treated the group as of family status. The question of whether scutellerids should be con¬ sidered a family or a subfamily of Pentatomidae has re¬ mained controversial. Kumar (1965) discovered that the internal male genitalia of scutellerids contain 6 ‘‘eca- deme” tubules, four paired and two unpaired, opening into the “bulbus ejaculatorius.” This feature is not found in any other pentatomoid, and at least suggests scutel- lerid monophyly. Gaffour-Bensebbane (1990) discovered that the spermatozoa of scutellerids are also distinctive and apparently derived within the Pentatomoidea. They possess a “plumed” structure, aiding the motility and thus providing additional evidence of the integrity of this family. A position outside the Pentatomidae is justified by the fact that scutellerids lack the distinctive spermathecal structure found in the Pentatomidae. The family contains approximately 80 genera and at least 450 species (Lattin, 1964). Key to Subfamilies of Scutelleridae 1. Antennae usually 5-segmented but sometimes 3-segmented; lateral portion of posterior margin of abdominal sterna emarginate. recurved anteriorly; metathoracic wing with trace of antevannal vein; 2 intervannal veins . Scutellerinae - Antennae 5-segmented (Fig. 76.1 A); lateral portion of posterior margin of abdominal sterna not emarginate or recurved; metathoracic wing with antevannal vein absent; a single intervannal vein present . 2 2. Abdominal venter with striated areas present on sterna 4, 5, and 6 . Pachycorinae - Abdominal venter lacking striated areas . 3 3. Scutellum only moderately enlarged laterally, hemelytra exposed for entire length; connexivum broadly exposed; metathoracic wing with intervannal vein well developed . Eurygastrinae - Scutellum broadly developed, hemelytra exposed laterally only near base; connexivum at most narrowly exposed; metathoracic wing with intervannal vein greatly reduced . Odontotarsinae EURYGASTRINAE. Moderate size; scutellum not strongly convex; dorsum punctate and glabrous; scutellum attain¬ ing end of abdomen, usually parallel-sided and leaving hemelytra and connexivum exposed laterally for almost entire length. The subfamily is chiefly Old World, with a single Hol- arctic genus Eurygaster Laporte occurring in the Western Hemisphere. ODONTOTARSINAE. Small to moderate size, length up to 11 mm; punctate, usually with considerable pubescence; head usually transverse and broadly rounded; hemelytra exposed only basally, laterad of large scutellum. Unlike most other scutellerids the members of this sub¬ family are chiefly Holarctic with the largest fauna occur¬ ring in the Palearctic. However, representatives have been assigned to this subfamily from all major faunal regions. It is sparsely represented in Australia and the Indo-Pacific. PACHYCORINAE (FIG. 76.1 A). Size range small to large; strongly convex dorsally; abdominal sterna striated. Key to subfamilies of Scutelleridae modified from Lattin, 1964. The subfamily is primarily New World in distribution and constitutes the majority of New World Scutelleri¬ dae, with 27 genera and 125 species (Lattin, 1964). Two genera {Hotea Amyot and Serville and Deroplax Mayr) are African and are the only Old World representatives. SCUTELLERINAE. Characterized by features in preced¬ ing key; antennae 5- (rarely 3-) segmented; many species large and colorful. This subfamily is diverse in the Eastern Hemisphere, with only Augocoris Burmeister in the Western Hemi¬ sphere. Three tribes are recognized; the Elvisurini, Scu- tellerini, and Sphaerocorini, comprising five genera, all confined to the eastern Hemisphere, with three restricted to Australia. Their distribution extends far into Oceania and westward into Africa and the Middle East. The Elvi¬ surini were recognized as a distinct subfamily by McDon¬ ald and Cassis (1984) and Gross (1975-1976). McDonald and Cassis (1984) recognized the Australian genus Tec- tocoris Hahn as a distinct subfamily. The genus contains only a single species, Tectocoris diophthalmus (Thun- berg). Scutelleridae 239 Rg. 76.2. Scutelleridae. A. Aedeagus, lateral view, Eupychodera corrugata (Van Duzee). B. Distal portion of aedeagus, sagittal view, E. corru- gata. C. Paramere, £ corrugata. D. Terminal female abdominal segments, Diolcus irroratus (Fabricius). E. Spermatheca, Pachycoris torridus (A-E from McDonald, 1966). F. Spermatheca, Sphaerocoris annulus (Fabricius) (from Pendergrast, 1957). Specialized morphology. The enlarged scutellum covering the entire dorsal surface of the abdomen, and often also the forewings, may have evolved independently more than once in the Pentatomoidea. Other specialized features of the Scutelleridae include the bizarre color pat¬ terns and unique characters of the male genitalia and sperm noted above. The 3-segmented antennae of the Scutellerinae are a derived feature. Natural history. Despite the frequent large size and brilliant coloration of many scutellerids, biological infor¬ mation on the group is surprisingly sparse. All species are plant feeders and some are of economic importance, especially species of Eurygaster Laporte in the Middle East (see Chapter 8). Several species exhibit maternal care. Hussey (1934) discussed this phenomenon in the Neotropical bug Pachycoris torridus (Scopoli) in Para¬ guay. He noted lack of hatching of eggs at the periphery of the mass guarded by the female and suggested para¬ sitism, which has subsequently been well established by Eberhard (1975) for a South American pentatomid (see Natural History, Chapter 73). Hussey also noted reports of egg guarding by Tectocoris diophthalmus (Thunberg) in Australia and Cantao ocellatus (Thunberg) in the Ori¬ ent. Miller (1956a) referred to an observation of the latter species as being the only pollinator of the Moon Tree (Macaranga roxburghi) in India. Tectocoris diophthal¬ mus often does serious damage to cotton bolls and other Malvaceae in Australia, where it is known as the cotton harlequin bug. Shield bugs, because of the large, usually convex scu¬ tellum, often are mistaken for beetles. Several species have a longitudinal stripe down the middle of the scu¬ tellum, which increases the similarity of appearance to beetle elytra. This phenomenon merits careful field ob¬ servations as it is not evident if, or why, this would be a selective advantage for the shield bugs. The majority of scutellerids are not brightly colored, but the family does include some of the most strik¬ ingly colored of all heteropterans. Many are iridescent blue and green, others a rainbow of red, yellow, blue, and green markings. Some of the most bizarrely colored genera are Callidea Laporte, Chrysocoris Hahn, Crypta- crus Mayr, Cosmocoris Stal, Poecilocoris Dallas, and Scutellera Lamarck. Distribution and faunistics. The family is represented in all major faunal regions, but it is most varied and nu¬ merous in the tropics and subtropics, where all of the spectacularly colored species occur. McDonald and Cassis (1984) revised the Australian fauna and provided a key to the four recognized subfami- 240 TRUE BUGS OF THE WORLD (HEMIPTERA'. HETEROPTERA) lies. Lattin (1964) provided a key to subfamilies, detailed discussion of morphology, biological notes, and a treat¬ ment of the entire North American fauna. The last world key to genera was presented in the beautifully illustrated work of Schouteden (1904-1906). 77 Tessaratomiidae General. These are large to extremely large (often over 15 mm), robustly ovate or elongate-ovate bugs. They resemble large pentatomids (Fig. 77.1) in general habitus and as a group have no common name. Diagnosis. Head small, triangular, much narrowed to apex, mandibular plates meeting mesally in front of cly- peus; antennae usually 4-segmented, but if 5-segmented then segment 3 very short; head laterally keeled, anten- niferous tubercles not visible from above; labium short, not exceeding forecoxae; pronotum extending over base of scutellum; scutellum triangular, not covering corium; hamus present on hind wings; strigil present on Pcu vein of hind wing and plectrum on abdominal tergum 1; meta- stemum produced laterad between coxae and anteriorly onto mesosternum, most strongly produced as an anterior wedge reaching nearly to front coxae with posterior mar¬ gin truncate at its junction with abdomen; tarsi either 2- or 3-segmented; 6 pairs of abdominal spiracles usually visible, spiracle of segment 2 strongly exposed on an un¬ differentiated portion of sternum; abdominal trichoboth- ria posterior to spiracles on sterna 3-7, arranged trans¬ versely mesad of spiracle; nymphs with dorsal abdominal scent-gland openings present between terga 3/4,4/5, and 5/6 (that between 3/4 sometimes small); a small “scar” present between terga 6/7; ninth paratergites greatly en¬ larged; pygophore as in Fig. 77.2A; aedeagus as in Fig. 77.2B-D; parameres as in Fig. 77.2E, F; spermatheca as in Fig. 77.2G. Classification. The group was first recognized as a higher taxon by Stal (1864—1865). Leston (1954b), while Fig. 77.1. Tessaratomidae. Musgraveia sulciventris (Stal) (drawn by S. Monteith; from CSIRO, 1991). treating it as a subfamily of a very inclusive Pentatomidae, noted resemblances to the male genitalia of the Scutelleri- dae, such as third dorsal conjunctival appendages short and sclerotized, dorsal seminal duct, canal at junction of phallotheca and reservoir always somewhat undulat¬ ing, reservoir and vesica merging imperceptibly into one another, and second conjunctival appendages ventral to vesica and membranous. He later (Leston, 1956) elevated the group to family status without comment, a status that has been maintained by Kumar (1969) and subsequent authors. Three subfamilies, comprising 49 genera and about 235 species, are recognized (Rolston et al., 1993). Key to Subfamilies of Tessaratomidae 1. Scutellum distinctly longer than wide; tarsi 3-segmented . 2 - Scutellum subequilateral; tarsi 2-segmented . Natalicolinae 2. Membrane scarcely areolate basally; longitudinal veins arising from base of wing membrane . . Oncomerinae - Membrane areolate basally with longitudinal veins arising from this basal cell-like area . . Tessaratominae Tessaratomidae 241 Fig. 77.2. Tessaratomidae. A. Genital capsule, Piezostemum subulatum (Thunberg). B. Aedeagus, lateral view, P. subulatum (A, B from McDon¬ ald, 1966). C. Aedeagus, sagittal view, P. calidium (Fabricius) (from Leston, 1954c). D. Aedeagus, sagittal view, Tessaratoma papillosa (Drury) (from Leston, 1954b). E. Paramere, P. calidium (from Leston, 1954c). F. Paramere, P. subulatum (from McDonald, 1966). G. Spermatheca, T. javanica (Thunberg) (from Pendergrast, 1957). NATALICOLINAE. Tarsi 2-segmented; male genitalia of type found in Pentatominae (Leston, 1954b). This is an almost exclusively Ethiopian subfamily. ONCOMERiNAE. Membrane lacking basal cells or with single elongate and narrow but feebly defined cell; hind wing with R+M and Cu parallel and contiguous on proxi¬ mal two-thirds. Leston (1955b) included 11 genera separated into two tribes, the Piezosternini and Oncomerini. TESSARATOMINAE. Membrane with basal cells and lon¬ gitudinal veins emanating posteriorly. This taxon is widely distributed in the Old World tropics, with some of the species being among the largest of the Heteroptera outside the Belostomatidae and Corei- dae. Leston (1955b) treated this group as a tribe within his subfamily Tessaratominae. He reduced several species previously considered as tribes to subtribal level. .The elevation of the tessaratomids to family rank returns the following to tribal status in the subfamily Tessaratominae; Prionogastrini Stal, Sepinini Horvath, Eusthenini Stal, Tessaratomini Stal, and Platytatini Horvath. Specialized morphology. The extremely large size, the produced and enlarged wedgelike metasternum, and exposed second abdominal spiracle appear to be special¬ ized features, as may be the brightly colored nymphs. Natural history. All species so far as known are phyto¬ phagous (see also Chapter 8). Distribution and faunistics. The family is primarily Old World tropical, with the widely distributed Piezoster¬ num Amyot and Serville also having three species in the Neotropics. Rolston et al. (1993) summarized the current classifica¬ tion. Kumar and Ghauri (1970) provided a key to the sub¬ families of the world. Yang (1935) discussed the Chinese fauna. Kumar (1974b) keyed the genera of Natalicoli- 242 TRUE BUGS OF THE WORLD (HEMIPTERA; HETEROPTERA) nae. Leston (1955b) treated the genera of Oncomerini. Schouteden (1905a) provided keys to the African tribes, genera, and species with color plates. Horvath (1900) keyed family groups and many genera and species. 78 Thaumastellidae General. These small, somewhat flattened, elongate, brown pentatomoids closely resemble some ground-living Lygaeidae (Fig. 78.1). They are never more than 3.5 mm long and occur as brachypterous and macropterous forms. They have no common name. Diagnosis. Head porrect with rounded lateral mar¬ gins; labium 4-segmented, segment 1 obscured in buccu- lar groove; antennae 5-segmented, articulation between segments 2 and 3 not capable of flexure (Fig. 78.1); scu- tellum triangular, not enlarged (Fig. 78.1); scent-gland channel elongate, sometimes nearly reaching dorsal mar¬ gin of metapleuron, adjacent evaporative area large, ex¬ tending onto posterior area of mesopleuron; corium of macropterous morph divided into exo- and endocorium with furrow running along M vein, both parts separately rounded posteriorly, membrane penetrating a short dis¬ tance between them; claval commissure present; mem¬ brane of forewing with reduced venation, no branching veins; flightless morphs with forewings reduced to short truncate (staphylinoid) pads, posterior margin straight, membrane absent (Fig. 78.1) (resembling lygaeids of tribe Plinthisini); coxal combs present, composed of flat¬ tened setae; foretibia spinose; tarsi 3-segmented; abdo¬ men lacking inner laterotergites on segments 3-7; all ab¬ dominal spiracles ventral; nymphs with 3 pairs of dorsal abdominal scent glands, between terga 3/4, 4/5 and 5/6; abdominal sterna 3-6 with 2+2 trichobothria laterally in oblique rows, sternum 7 with 1 + 1 trichobothria in Thau- mastella aradoides and T. elizabethae Jacobs (Fig. 78.2F) but 2+2 in T. mmaquensis Schaefer and Wilcox (Jacobs, 1989); aedeagus as in Fig. 78.2A, B; parameres as in Fig. 78.2C; abdominal sternum 7 of female not divided; ovipositor valvulae platelike (Fig. 78.2D); second valvi- fers fused at midline; spermatheca with long coiled duct, pump flanges variable, both distal and proximal flanges present or absent (Fig. 78.2E). Classification. The genus Thaumastella Horvath was originally described as a lygaeid and remained in that family until Seidenstiicker (1960) noted that it lacked Fig. 78.1. Thaumastellidae. Thaumastella namaquensis Schaefer and Wilcox (from Schaefer and Wilcox, 1971). the laciniate ovipositor characteristic of most Lygaeidae. Stys (1964a) emphasized its pentatomoid relationships and erected the family Thaumastellidae for the then single known species, Thaumastella aradoides Horvath, from North Africa and the Middle East. Dolling (1981) con¬ sidered the group to be a subfamily of Cydnidae, whereas Jacobs (1989) treated it as a distinct family. Although we treat this group at the family level, it is clear that the coxal combs, spinose foretibia, and stridulatory structures ally it with the Cydnidae. Jacobs (1989) found the chromosome number to be 2n = 20 (16XY + m + XY) but also found conditions of 18 and 17 in different populations of 7. namaquensis. The presence of an m-chromosome is particularly important, because it is unique in the Pentatomoidea, although it does occur frequently in the Lygaeidae. The presence of a claval commissure is also uncommon in the Pentatomoi¬ dea, occurring only in the Amnestinae (Cydnidae) and Urostylidae. Clearly, the Thaumastellidae is a family of great phylogenetic importance, but precise resolution of its relationships will require additional cladistic analysis. Three species—all placed in the genus Thaumastella — Thaumastellidae 243 Fig. 78.2. Thaumastellidae. Thaumastella aradoides Horvath. A. Aedeagus, sagittal view. B. Aedeagus, lateral view. C. Paramere. D. Female terminal abdominal segments. E. Spermatheca (A-E from Stys, 1964a). F. Ventral view of abdomen with trichobothria (from Seidenstucker, 1964). Abbreviations: sp, spiracle; tb, trichobothria. are known, the fully winged North African T. aradoi¬ des and the completely flightless T. mmaquensis and T. elizabethae from Namaqualand, the Richterveld, and Na¬ mibia. Specialized morphology. All species have a well- developed stridulatory mechanism (Jacobs, 1989). Schae¬ fer (1980b) believed that the hind wings were absent in the flightless species from southern Africa and that the stridu- litrum was situated on the hemelytra or possibly on the third abdominal tergum. Jacobs (1989), however, showed conclusively that very short, subtriangular hind wings have a stridulitrum formed of a well-sclerotized longitudi¬ nal ridge on the underside, situated above the abdominal tergum 1, which bears an extremely finely, transversely ridged, suboval stridulitrum situated near its anterolateral margin. Jacobs argued the “limae” were not involved in stridulation as was suggested by Seidenstucker (1960), Stys (1964a), and Schaefer (1980b). Natural history. Jacobs (1989) found that the two Na¬ maqualand species live chiefly in cavities under large stones. Despite having di.^covered a few individuals adja¬ cent to these stones at dusk, he rarely found them leaving these sheltered cavities and believed that they may feed chiefly on seeds that accumulate in such areas by the force of the strong winds. Jacobs also gave an account, how¬ ever, of T. elizabethae leaving its stone shelters to feed on the seeds of Pharnaceum aurantium (Aizoaceae), which it dragged along the ground, with the seeds attached to its stylets. The eggs are large, ovoid, and half the size of the abdomen. Distribution and faunistics. The family has a disjunct distribution including North Africa, the Near East, and the semidesert areas of southwestern Africa. The major sources on the group are by Stys (1964a) and Jacobs (1989). 244 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Urostylidae General. These pentatomoids are usually relatively elongate, ranging in length from 3.5 to 14 mm, with elongate legs and a small head (Fig. 79.1). Many have a coreidlike habitus. They have no common name. Diagnosis. Head small, peltoid, with lateral margins not keeled; antennae 5-segmented, segment 1 much longer than head; antenniferous tubercles broad, strongly exerted, often appearing annulate, placed on or slightly above line running through middle of eye; ocelli placed very close to one another (Fig. 79.2A) (closer than to the eyes, a unique condition in Pentatomoidea); bucculae small; claval commissure reduced (Fig. 79.1) or obsolete (Fig. 79.2A); radial and medial veins of forewings di¬ vergent from base; membrane usually with only 4 or 5 longitudinal veins (Figs. 79.1, 79.2A); frenum extend¬ ing to apex of scutellum; metasternal scent-gland orifice frequently with an elevated spinose auricle; middle and hind coxae widely separated; tarsi 3-segmented; nymphal scent-gland openings present between abdominal terga 4/ 5 and 5/6; pygophore as in Fig. 79.2C; vesica short with conjunctival appendages (Fig. 79.2D); second valvifers fused to form an M-shaped or W-shaped sclerite (but said to be separate by Gapud, 1981); chromosome number 2n = 16. Classification. The systematic position of this family has long been ambiguous. Kumar (1971) reviewed the early literature, noting that Singh-Pruthi (1925) related it to the Acanthosomatidae, Yang (1938a, b, 1939) and Pendergrast (1957) to the Pyrrhocoridae, and Miyamoto (1961a) to the Pentatomidae. Kumar (1971) believed that the group represents an early divergence from the other pentatomomorphans, possibly together with the Pyrrho¬ coridae with which they share uniquely the fused second valvifers that form an M- or W-shaped sclerite. China and Fig. 79.1. Urostylidae. Urolabida sp. (drawn by S. Monteith; from CSIRO, 1991). Slater (1956:411) indicated that the urostylids “must represent the Proto-Trichophora at the base of the Penta¬ tomidae, Coreidae and Lygaeidae.” Schaefer and Ash- lock (1970:629) noted that, although primitive, they are “some distance from the origin of the Pentatomoidea.” Since most of the above was based on study of only one or a very few species and the language mostly “precladis- tic,” clearly a comprehensive study of the family and its relationships is badly needed. Whatever its more precise position, the trichobothrial number and pattern suggest placement in the Pentatomoidea. Key to Subfamilies of Urostylidae 1. Body relatively large; well over 5 mm in length; spiracles ventral; scent-gland auricle with a spine; hamus present in hind wing . UrostyUnae » - Body very small, at most scarcely exceeding 4 mm in length; spiracles lateral (except segment 2 ventral when present); scent-gland auricle not spined; hamus absent . Saileriolinae SAiLERiouNAE. Very small, unlike Urostylinae in habi¬ tus; head strongly declivent, eyes close to base; antennal segment 3 very short; legs simple, mutic; meso- and metacoxae separated by distance nearly twice their length; tarsal segments 1 and 3 each longer than segment 2; scu¬ tellum swollen anteromesally; corium semihyaline, apex Urostylidae 245 Fig. 79.2. Urostylidae. Saileriola sandakanensis China and Slater (from China and Slater, 1956). A. Habitus, B. Venter. C. Male genital capsule D. Aedeagus, sagittal view. extending beyond apex of abdomen; trichobothria appar¬ ently always absent on abdominal sterna 3 and 4; meta- thoracic scent-gland orifice slitlike, lacking a peritreme and evaporative area. Three genera are known: Saileriola China and Slater (two species; Borneo and Vietnam) and the monotypic Bannacoris Hsiao (1964) from Yannan Province, China, and Ruckesona Schaefer and Ashlock from Thailand. UROSTYLiNAE (FIGS. 79.1,79.2A). Relatively large; habitus somewhat coreoid; antennae usually long, sweeping; legs long; body more elongate than in most pentatomoids; abdominal trichobothria paired, transverse, present on sterna 3-7. Approximately four genera and more than 80 species are known. Specialized morphology. The reduced third antennal segment in the Saileriolinae and perhaps the fused second valvifers are certainly novel features, as presumably is the elongation of the corium in the Saileriolinae. Natural history. Little is known of the biology of these pentatomoids other than a collection record of Ruckesona vitrella Schaefer and Ashlock on “palm at water edge.” Nymphs of several instars were taken, suggesting that this unidentified palm is a true host plant. Schaefer and Ash¬ lock (1970) noted that both adults and nymphs of this species had what appeared to be fragments of chloro- plasts in the gut, suggesting that the insects do not feed exclusively upon sap. Distribution and faunistics. Urostylids occur in southern and eastern Asia, reaching northward into the eastern Palearctic and southwest into New Guinea. A basic reference on the group is by Yang (1939). Lygaeoidea 80 Berytidae General. Commonly known as stilt bugs, these in¬ sects, which range in length from 2.5 to 11 mm, are usually elongate and slender, with threadlike legs and antennae (Fig. 80.1 A, B). Many superficially resemble species of Hydrometridae or the reduviid subfamily Emesinae, but lack the raptorial forelegs of the latter. The majority are rather dull yellowish or reddish brown and mutic, but some species are bizarrely ornamented with spines and other protuberances (Fig. 80. IB). Diagnosis. Head subspherical, often with clypeus 246 TRUE BUGS OF THE WORLD (HEMiPTERA: HETEROPTERA) Fig. 80.1. Berytidae. A. Gampsocoris panormimus Seidenstticker (from Seidenstucker, 1965a). B. Acanthoberytus wygodztnskyi Stusak (from Stusak, 1968). C. Lateral view head, thorax, scutellum, G. panormimus (from Seidenstucker, 1965a), D. Aedeagus, sagittal view. Berytinus hir- ticornis (Brulle). E. Aedeagus, sagittal view, G. culicinus Seidenstucker (D, E from Pericart, 1984). F, Spermatheca, B. minor (Herrich-Schaeffer) (from Pendergrast, 1957). Abbreviation: per, peritreme. produced anteriorly (Fig. 80.1C); antennae located above a line through middle of eye; antenniferous tubercles re¬ duced; antennal segment 4 usually short and somewhat swollen; distal ends of femora often swollen (Fig. 80.1 A, B); scutellum pointed posteriorly; peritreme of metatho- racic scent gland usually uniquely produced, often as an elongate spine (Fig. 80.1 A); corium usually in part de- sclerotized; all abdominal spiracles dorsal; adults usually with 3 trichobothria (sometimes 2) on abdominal sternum 3; abdominal mediotergites fused; nymphs with dorsal abdominal scent-gland openings between terga 3/4 and 4/5 or only between terga 3/4; aedeagus as in Fig. 80. ID, E; ovipositor reduced; sternum 7 of females entire; apical spermathecal bulb large, ovoid, and globular with distal pump flange well developed, proximal flange reduced or absent (Fig. 80. IF); nymphs usually with glandular setae. Classification. The taxon was first established by Fie- ber (1851) as Berytidea. Costa (1853) (as Berytini) and Uhler (1876) (as Berytina) treated them as a subfamily of the Coreidae. Stal (1874) (as Berytina) considered them Berytidae 247 to be a subfamily of Lygaeidae. Kirkaldy (1902) recog¬ nized a subfamily Neidinae and treated the entire taxon at family rank. Because Neides Latreille, 1802 was consid¬ ered to be a senior synonym of Berytus Fabricius. 1803 and because Kirkaldy believed that genera in synonymy could not serve as the basis for higher group names, he used the name Neididae and was followed by many subsequent authors. Berytidae have long been considered to be closely re¬ lated to some subgroups of Lygaeidae, especially the sub¬ family Cyminae. Southwood and Leston (1959) placed the Cyminae in the Berytidae, presumably on the basis of the very short claval commissure, similarities of the egg, female genitalia, and chromosome numbers. Hamid (1975) noted that Southwood and Leston gave no char¬ acter evaluation, that the chromosome number was true for only one subfamily, and that some characters were obvious plesiomorphies. He also listed 15 character states in which the Cyminae and Berytidae differed. Hamid also disagreed with Stys's (1967c) discussion of the spiracle position in the two groups and concluded that the place¬ ment of the Cyminae in the Berytidae was unwarranted (see also Cyminae. Chapter 83), Pericart (1984) agreed, stating that some of the apparent similarities are the result of parallel evolution. Two subfamilies are recognized, the Berytinae and Metacanthinae, the latter established by Douglas and Scott (1865) as a separate family. They comprise about 39 genera and 150 species (Pericart, 1984). Key to Subfamilies of Berytidae 1. Frons usually produced above clypeus into a laterally compressed crest or cone; scutellum triangular without denticle or spine; abdomen ventrally punctate . Berytinae - Frons rounded anteriorly, without a crest or cone, but often spinose or tuberculate; scutellum more or less semicircular, usually with a long denticle or spine in middle, or with apex produced into a horizontal curved spine; abdomen ventrally impunctate . Metacanthinae BERYTINAE. Head truncate anteriorly or prolonged above postclypeus by an elongate process; pronotum ele¬ vated before the posterior margin in macropterous forms; scutellum lacking an upstanding spine; antennae and legs lacking extensive black annulations; frequently brachyp- terous or micropterous; hemelytral membrane with 5 dis¬ tinct veins; metathoracic scent-gland auricle variable, but never prolonged into fingerlike process; abdomen densely punctate below; 2 pairs of trichobothria on sternum 3; female terga 8 and 9 divided completely by a longitudinal groove; first valifer fused with paratergite 8 (except in ApoplymusVieher). This group of 11 genera is broadly distributed in the Old World, with only Neides muticus (Say) occurring natively in North America. METACANTHINAE. Head anteriorly annulate or subtrun¬ cate, but lacking a forward-projecting spinelike process; pronotum more or less trituberculate or trispinose pos¬ teriorly, sometimes spinose laterally; antennae and legs usually annulated with dark rings; usually macropterous; scutellum with an upstanding spine; metathoracic scent- gland auricle variable, frequently produced as an elongate spinelike process; abdomen impunctate ventrally or with at most scattered punctures; trichobothria on sterna 3-7; Key to subfamilies of Berytidae adapted from Kerzhner and Jaczewski, 1964, female terga 8 and 9 entire; first valifer not fused with paratergite 8. The Metacanthinae is a group of worldwide distribu¬ tion, with 28 genera currently recognized. Specialized morphology. The extended metapleural scent-gland auricle, elongate slender appendages, and especially the glandular setae of the nymphs are special¬ izations that are infrequently, or never, found in other Heteroptera. Natural history. The majority of species are thought to be phytophagous and live above the ground on plants; others are geophilous. At least some species of Jaly- sus Stal, Neides Latreille, and Berytinus Kirkaldy are in part predatory (Poisson and Poisson, 1931; Wheeler and Henry, 1981). Many species are known to live on plants covered with a sticky glandular pilosity (Pdricart, 1984). Pericart (1984) should be consulted for a summary of individual biological studies. Wheeler and Henry (1981) presented details of the biology of Jalysus wickhami Har¬ ris and J. spinosus (Say) (see Chapter 8). Wheeler and Schaefer (1982) presented a host list and discussed feed¬ ing trends. Distribution and faunistics. The family is represented in all major zoogeographic regions. Pericart’s (1984) study of the Palearctic fauna is the most exhaustive recent work. Stusak has published many descriptive papers on the world fauna, for example. 248 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) his recent paper on the Oriental fauna (Stusak, 1989). Froeschner (1981) provided a key to the South American fauna. 81 Colobathristidae General. The members of this family, which range in length from 6 to 20 mm, are generally very elongate in¬ sects with slender legs and antennae and a punctate dorsal body surface (Fig. 81.1). They have no common name. Diagnosis. Eyes substylate (Fig. 81.1; except in Dava- kiella Stys); antennae elongate, 4-segmented (Fig. 81.1), located above a line running through middle of eyes; an- tenniferous tubercles reduced; ocelli present; infraocular ridges sometimes strongly developed; distal end of scutel- lum narrowed, elongate, sometimes armed with an erect spine; forewing narrowed, lateral corial margin concave; corium at least in part transparent or translucent, usually with a triangular distal cell; membrane of forewing with veins reduced (Fig. 81.2A) or absent; clavi usually over¬ lapping, no claval commissure; abdomen constricted at base; abdominal spiracles 2, 3, and 4 dorsal, those on segments 5, 6, and 7 ventral; abdominal trichobothria as in Fig. 81.2B; nymphal scent-gland openings between terga 3/4, 4/5, 5/6, those between 3/4 and 4/5 reduced; aedeagus with elliptical phallobase and elongate descle- rotized tubular vesica (Fig. 81.2C, E); parameres as in Fig. 81.2D, F; ovipositor platelike, sternum 7 not split mesally; spermathecal duct elongate, coiled, bulb globu¬ lar, with reduced flanges (Fig. 81.2H); second valifers fused; eggs spindle-shaped with an obliquely set pseudo- perculum. Classification. The taxon was first recognized by Stal (1864-1865) as a subfamily of Lygaeidae and was ele- Fig. 81.1. Colobathristidae. Colobathristes chalcocephalus Burmeis- ter(from Slater, 1982), vated to family status by Bergroth (1910). Stys (1966a) considered it as belonging to the “malcid evolutionary line,” which included the cymine lygaeids, the Malcidae. and Berytidae. Kumar (1968) discussed the close relation¬ ship to the Lygaeidae and believed that the family might well be included in the Lygaeidae as a subfamily, and Stys (1966a) commented on intrafamilial relationships in de¬ tail. The relationships of the family are in need of further clarification. The group contains 23 genera and 83 species. Key to Subfamilies of Colobathristidae 1. Antennae relatively short, subequal to half length of body; antennal segment 1 shorter than width of head across eyes . Dayakiellinae - Antennae elongate, longer than or subequal in length to body length; antennal segment 1 longer than width of head across eyes .. Colobathristinae Colobathristidae 249 A C E Fig. 81.2. Colobathristidae. A. Forewing, Dayakiella brevicornis Horvath. B. Abdomen, with trichobothria, D. brevicornis (A, B from Stys, 1966b). C. Aedeagus, lateral view, Phaenacantha australiae Kirkaldy. D. Paramere, P. australiae. E. Aedeagus, Symphylax musiphthora Ghauri. F. Paramere, S. musiphthora. G. Spermatheca, S. musiphthora (E-G from Ghauri, 1968). H. Spermatheca, P. austraiiae (C, D, H from Stys, 1966a). Abbreviations: Itb, lateral trichobothria; mtb, mesial trichobothria. COLOBATHRISTINAE (FIG. 81.1). Antennal length greater than or subequal to body length; legs very long; tarsal segment 1 at least 1.66 times as long as (usually longer than) combined length of segments 2 and 3; hind wings lacking Cu vein; laterotergites of segment 7 “normally” developed, posteriorly free; male genitalia terminal (often telescoped). The group contains 11 genera and 44 species from the Oriental and Australian regions and 11 genera and 37 species from the Neotropics (Kormilev 1949, 1951). DAYAKIELLINAE. Antennae about one-half length of body; facies lygaeid-like; eyes sessile; head lacking infra¬ ocular ridges; scutellum with long horizontal spine; tarsal segment 1 at most 1.17 times as long as combined length of segments 2 and 3; hind wings with free distal part of Cu and with recurrent glochis; both dorsal and ventral laterotergites of segment 7 meeting posteriorly and fusing to form a genitoanal chamber. This subfamily contains only Dayakiella Horvath, with two species (D. brevicornis Horvath and D. sumatrensis Stys) known solely from Indonesia. Stys (1966b) believed that it possesses several plesiomorphic characters. SPECIALIZED MORPHOLOGY. The aedeagus is similar in form to the apparently derived condition found in the ly- gaeid subfamilies Pachygronthinae and Heterogastrinae. where the sperm reservoir is reduced and located near the distal end of the elongate membranous tubular phallus and in which holding sclerites are lacking. Several Neotropical genera possess densely striate infraocular ridges representing the stridulitrum of the stridulatory apparatus. Fine spines present on the inner ^ side of the anterior femora presumably function as the plectrum (Stys, 1966a). Natural history. All colobathristids whose habits are known feed exclusively on grasses. Phaenacantha saccha- ricida (Karsch) is destructive to sugar cane in Indonesia ^ (Miller, 1956a). Many species are myrmecomorphic; species of Tricentrus Horvath are strikingly so (Kormilev. 1949). C Distribution and faunistics. This family has exten- ^ sive Neotropical and Oriental faunas, but not a single ^ species occurs in Africa. Stys (1966a) believed that the Neotropical and Oriental faunas are quite distinct and may well be found to represent distinct subfamilies. Phae- nacantha, however, is represented by a single Neotropical ^ species {P. saileri Kormilev), whereas 27 other species of the genus are Oriental-Australian. Basic works on the ^ group include those of Horvath (1904) as well as those of Kormilev and Stys cited above. The key of Carvalho and Costa (1989) allows for identification of Neotropical ^ genera. 250 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) 82 Idiostolidae General. Members of this family range from 5 to 7 mm in length and resemble broad-bodied species of the ly- gaeid genus Ozophora Uhler (Fig. 82.1). They are of moderate size and have no common name. Diagnosis. Head porrect; antennae inserted at level below ventral margin of eye; ocelli present; membrane of forewing with 5 longitudinal veins or numerous cells (Figs. 82.1, 82.2A); legs slender and mutic; tarsi 3- segmented; abdomen with numerous trichobothria as fol¬ lows; adults with 7 on sterna 3 and 4 (positioned both medially and laterally) and 4 laterally on segments 5-7 (Fig. 82.2B); nymphs with 3 on segments 3 and 4, 1 on segment 5, 2 on segment 6, and 3 on segment 7 (Schaefer, 1966a); all spiracles ventral; dorsal inner laterotergites present (Fig. 82.2B); nymphal body shape ovoid; nym- phal abdominal scent glands between abdominal terga 4/ 5 and 5/6 and a scar representing a small gland between terga 3/4; abdominal terga 1 and 2 and terga and sterna 3-6 fused; aedeagus as in Fig. 82.2D; parameres as in Fig. 82.2E; females with sternum 7 completely divided medially; ovipositor laciniate; spermatheca absent. Classification. This taxon was originally described as a subfamily of the Lygaeidae by Scudder (1962a). Sub¬ sequently it was elevated to family and then superfamily status by Stys (1965) (see also Schaefer, 1966a; Stys and Kerzhner, 1975). Although we include the Idiostolidae within the Lygaeoidea, this may not correctly represent its phylogenetic position within the Pentatomomorpha, judging from the unusual combination of characters pos¬ sessed by the group. Three genera comprising four species are known; Idios- tolus insularis Berg from southern South America, Trise- cus pictus Bergroth from Tasmania, T. armatus Wood¬ ward from New South Wales, and Monteithocoris hirsutus Woodward from Tasmania. Specialized morphoiogy. The novel trichobothrial patterns (Fig. 82.2B), lack of spermatheca and vesica, and the broadly ovoid shape of the nymphs are all special¬ ized conditions (Schaefer, 1966b; Schaefer and Wilcox, 1969). Natural history. Idiostolids live in moss and litter in Nothofagus forests and are almost certainly phytopha¬ gous. They are most easily collected by processing in a Berlese funnel samples of litter and moss from the far southern forests. Distribution and faunistics. The family possesses a strikingly disjunct transantarctic distribution. Fig. 82.1. Idiostolidae. Trisecus pictus Bergroth (from Schaefer and Wilcox. 1969). Basic references in the group are by Scudder (1962a), Woodward (1968b), and Schaefer and Wilcox (1969). 83 Lygaeidae General. Members of this large and diverse family are extremely varied in size (1.2-12 mm) and form (Figs. 83.3-83.6). Most species are rather small and ob¬ scurely brown or black, but many are brightly colored red or yellow and black. Some have conspicuously enlarged forefemora. They are often referred to as seed bugs. Lygaeidae 251 Fig. 82.2. Idiostolidae. A. Hemelytron, Trisecus pictus (from Schaefer and Wilcox, 1969). B. Abdomen, ventral view, Idiostolus insularis Berg, trichobothrial positions marked by "x." C. Abdomen, dorsal view, I. insularis. D. Aedeagus, sagittal view, /. insularis (B-D from Schaefer, 1966a). E. Paramere, /. insularis (from Schaefer and Wilcox, 1969). Diagnosis. Ocelli present except in brachyterous forms; bucculae well developed; antennae located below a line drawn through middle of eye; forewing with 4- 5 veins in membrane; abdominal spiracle position ex¬ tremely variable; usually with 3 trichobothria submesally or laterally on abdominal sterna 3 and 4, and laterally on sterna 5 and 6, 2 trichobothria laterally on sternum 7 (Fig. 83. lA, B); aedeagus usually with conjunctival lobes and processes and a distinct vesica (Fig. 83.2A-F); sperm reservoir present (Fig. 83.1C); parameres variable in form, often elongate and slender or broadened (Fig. 83. ID); testes as in Fig. 83.2G; ovipositor usually lacini- ate (Fig. 83. IE); spermatheca usually with distinct bulb and flange (Figs. 83.IF, 83.2H, I); egg with 3 to 15 micropyles. Classification. The Lygaeidae are probably paraphy- letic, with some of the subfamilies presumably being the sister taxa of members of other groups such as Berytidae, Colobathristidae, and Malcidae (Southwood and Leston, 1959; Stys, 1967c). Consequently, the family is difficult to characterize, and the complex relationships have not yet been worked out. Many subfamilies will probably be elevated to family status in the future. Nonetheless, there has been a great deal of systematic work on the higher classification of the Lygaeidae in recent years, and the majority of subfamily and tribal taxa appear to be reasonably well established as monophyletic. The family possesses great diversity of some features found to be relatively constant in other lygaeoid (and pen- tatomomorphan) families. For example, the spermatheca is extremely variable and useful taxonomically chiefly at subfamUial levels, as are the position of the abdomi¬ nal spiracles, the dorsal abdominal scent-gland openings, and the number and placement of the abdominal tricho¬ bothria. The taxon was first recognized by Schilling (1829), and the most complete early synthesis was by Stal (1872). Slater (1964b) provided a modem world catalog. Kumar (1968) discussed the relationships relative to related superfamilies. At least 500 genera and 4000 species are known. 252 TRUE BUGS OF THE WORLD (HEMIPTERA; HETEROPTERA) Fig. 83.1. Lygaeidae. A. Female abdomen and trichobothria, lateral view, Targarema stali (White). B. Female abdomen and trichobothria. lat¬ eral view, Phasmosomus araxis Kiritshenko (A, B from Sweet, 1967), C. Sperm reservoir, sagittal view, Xenoblissus lutzi Barber (from Slater, 1979). D. Paramere, Ozophora singularis Slater (from Slater, 1983). E. Female abdomen, ventral view, Ischnobemus sp. (from Siater. 1979). F, Spermatheca, Plinthisus Hindersi Slater and Sweet (from Slater and Sweet, 1977). Abbreviations: Itb, lateral trichobothria; mtb. mesial trichobothria. Key to Subfamilies of Lygaeidae 1. Suture between abdominal sterna 4 and 5 usually curving forward laterally and rarely (e.g., Gastro- des Westwood, Phasmosomus Kiritshenko, Caenusia Strand) attaining lateral margins of abdomen (Fig. 83.1 A); if 4/5 suture complete, trichobothria usually present on head (Fig. 83.6A, B) . . Rhyparochrominae - Suture between abdominal sterna 4 and 5 not curving forward, attaining lateral margins of abdomen; head without trichobothria . 2 2. Spiracles on abdominal segments 2-7 all located dorsally' . 3 - At least one pair (and often more) of spiracles on abdominal segments 2-7 located ventrally (Fig. 83.1A, B) . 6 3. Clavus at least in part punctate; posterior margin of pronotum not depressed laterad of base of scutellum, or clavus and corium fused into a hard convex, coarsely punctate shell . 4 - Clavus impunctate; posterior margin of pronotum depressed between scutellum and humeral angles . 5 4. Forewings forming a convex, beetle-like shell, wings meeting evenly down midline (Fig. 83.5); tarsi 2-segmented; abdominal segment 5 with a single trichobothrium; no trichobothria present on abdominal segment 4 . Psamminae 'A few Cyminae key out at this point but are readily recognizable by the coarsely punctate body surface. Lygaeidae 253 Fig. 83.2. Lygaeidae. A. Aedeagus. lateral view, Cymus angustatus Stal. B. Aedeagus, lateral view, Oncopeltus fasciatus (Dallas). C. Aedeagus, lateral view, Orsil/us depressus Dallas. D. Aedeagus, lateral view, Oxycarenus hyalinipennis (Costa). E. Aedeagus. lateral view, Oedancala dorsalis (Say), F. Aedeagus, lateral view, Myodocha serripes Olivier (A-F from Ashlock, 1957). G. Testes, Nysius thymi (Wolff) (from Pendergrast, 1957). H. Spermatheca, Macropes femoralis Distant (from Slater, 1979). I. Spermatheca, Oxycarenus hyalinipennis (from Carayon, 1964b). - Forewings with a distinct clavus, corium, and membrane, membranes overlapping one another; tarsi 3-segmented; 2 or 3 trichobothria present on both abdominal segments 4 and 5 . . Ischnorhynchinae 5. Apical corial margin straight: hind wing with a subcosta but lacking intervannals; often brightly colored with red, yellow, orange, and black . Lygaeinae - Apical corial margin sinuate on mesal half; hind wing lacking a subcosta but with intervannals present; usually dull brownish yellow with hemelytra partially hyaline . Orsillinae 6. Spiracles of abdominal segment 7 ventral, all others dorsal . 7 - At least spiracles of abdominal segments 6 and 7 ventral ... 9 7. Hemelytra impunctate or at most with only weak, scattered punctures (Fig. 83.3A) .... Blissinae - Hemelytra coarsely punctate . 8 8. Bucculae short not extending caudad of base of antenniferous tubercles; trichobothria present laterally on abdominal sterna 3-7 . Cyminae - Bucculae elongate, extending to base of head; trichobothria present laterally only on abdominal sterna 5 and 6 . Cryptorhamphinae 254 TRUE BUGS OF THE WORLD (HEMIPTERA- HETEROPTERA) 9. Spiracles on abdominal segments 3 and 4 dorsal ... 10 - Spiracles on abdominal segments 3-7 ventral . 12 10. Abdominal sterna 2-4 with sutures fused and obliterated; no lateral trichobothria on segments 3 and 4; body appearing myrmecomorphic . Bledionotinae - Abdominal sterna 2-4 with distinct sutures present, trichobothria on sterna 3 and 4 lateral; body generally short and stout, not myrmecomorphic . 11 11. Spiracles on abdominal segment 5 dorsal; spiracles on segment 2 ventral . Henestarinae - Spiracles on abdominal segment 5 ventral; spiracles on segment 2 dorsal . Geocorinae 12. Abdominal sterna 3-5 lacking lateral trichobothria; females without a spermatheea . .. Henicocorinae - Abdominal sterna 3 and 4 each with 2 or 3 lateral trichobothria; females with a spermatheea . 13 13. Spiracles of abdominal segment 2 dorsal . 14 - Spiracles of abdominal segment 2 ventral . 15 14. Lateral pronotal margins explanate or laminate . Artheneinae - Lateral pronotal margins rounded or at most slightly carinate, never conspicuously explanate .... . Oxycareninae 15. Cross vein present in membrane of forewing creating a closed basal cell; forefemora at most weakly incrassate and with few spines; hamus of hind wing arising distad of point on discal cell where cubitus diverges as a free vein . Heterogastrinae - No cross vein or closed cell basally in membrane of forewing; forefemora strongly incrassate and heavily spinose (Fig. 83.4); hamus of hind wing arising in discal cell basad of divergenee of cubitus as a free vein . Pachygronthinae ARTHENEINAE. Pronotal margins explanate; hamus lack¬ ing; abdominal spiracles 3 through 7 located ventrally on sternal “shelf”; complete, unfused suture between ab¬ dominal sterna 4 and 5 in female; inner laterotergites lacking; trichobothrial pattern conventional, with those on sternum 5 located one above the other on a single elevation. Four tribes are recognized; Polychismini Slater and Brailovsky (1986), containing only Polychisme ferrugino- sus (Stal) from northern South America; Artheneini, con¬ taining five genera and 16 species; Dilompini, containing only Dilompus Scudder, with two known species from southeastern Australia and New Zealand; and Nothochro- mini Slater, Woodward, and Sweet, containing a single species, Nothochromus maoricus Slater, Woodward, and Sweet from New Zealand. BLEDIONOTINAE. Strikingly myrmecomorphic; prono- tum sometimes with bizarre ornamentation, lateral mar¬ gins rounded; anterior abdominal segments fused, sutures obliterated; spiracles dorsal on abdominal segments 2, 3 and 4, ventral on segments 5-7; dorsal abdominal scent- gland openings present between terga 4/5 and 5/6, these segments curving strongly posteriorly from lateral mar¬ gins to meson. Two tribes are recognized; Bledionotini, which in¬ cludes only the strongly myrmecomorphic Bledionotus systellonotoides Reuter from the Near East, and Pamphan- tini, which contains seven genera and 20 species. The Pamphantini are chiefly Neotropical; most species are known from Cuba and Hispaniola, but there are also a few South American species. Austropamphantus woodwardi Slater occurs in northern Australia. There is considerable doubt about the monophyly of the subfamily. Most of the characters used to associate the two tribes are the result of myrmecomorphic modifi¬ cations. BLissiNAE (FIG. 83.3A). Hemelytra not, or only weakly, punctate; body frequently covered with a pruinose layer formed of minute spicules; spiracles dorsal on abdominal segments 2-6, ventral on segment 7; nymphal scent- gland openings present between terga 4/5 and 5/6; body shape ranging from elongate and extremely slender to short and stout. This is the so-called chinch bug subfamily. Approxi¬ mately 50 genera and 385 species are currently recog¬ nized. The group is worldwide in distribution but most abundant in the tropics. Ischnodemus Fichei has a large element that shows close South American-African rela¬ tionships, but the group is absent from the rest of the Old World tropics. Slater (1979) monographed the subfamily, including keys to genera and species and cladograms of the genera and some species groups. All members of the subfamily breed only on monocotyledonous plants. Poaceae is the most frequent host group, although Cyper- Lygaeidae 255 aceae are common hosts and groups such as the Restiona- ceae are less common (Slater, 1976). CRYPTORHAMPHiNAE. Elongate, brownish yellow; body coarsely punctate; bucculae long, extending to base of head; pronotum lacking a median carina; corium with a distinct subcosta, radius, media, and cubitus; hind wing with well-developed hamus and anterior and posterior vannals; dorsal abdominal scent-gland scars present be¬ tween terga 4/5 and 5/6; ninth paratergite secondarily cleft; trichobothria present only on sterna 5 and 6 (Hamid, 1971a). Two genera and four species are currently known. Cryptorhamphus Stal (two species) from Australia and Tasmania and Gonystus Stal (two species) from Australia, with G. nasutus Stal occurring on Fiji. Nothing appears to be known of the biology. CYMINAE. Small, usually brownish yellow, body coarsely punctate; body shape elliptical; spiracles dorsal on abdominal segments 2-6 but usually ventral on seg¬ ment 7 (sometimes all spiracles dorsal); nymphal scent glands usually present between abdominal terga 3/4, and 4/5, sometimes also between terga 5/6 and occasionally only between terga 3/4 (Hamid, 1975). Fourteen genera and 76 species are currently recog¬ nized, segregated into three tribes—Ontiscini, Ninini, and Cymini. The group occurs worldwide, with the On¬ tiscini being restricted to the Australian and Oriental regions. The high chromosome number (20-28 autosomes -i- XY), the short uncoiled gonoporal process, and the side- 256 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) by-side method of copulation suggest that the sister taxon of the Cyminae may be the Berytidae, which currently are treated as a separate family (see South wood and Leston, 1959; Stys, 1967c). GEOCORiNAE. Eyes large, reniform, prominent, often projecting backward and usually overlapping or nearly overlapping anterolateral pronotal angles; body usually stout and ovoid; pronotum broad with a transverse fur¬ row; spiracles dorsal on abdominal segments 2-4, ventral on segments 5-7; nymphal scent glands present dorsally between abdominal terga 4/5 and 5/6, sutures between segments usually curving strongly backward from lateral margins to mesal scent-gland openings. The subfamily is distributed worldwide and contains 14 genera and approximately 219 species. Readio and Sweet (1982) revised the eastern Nearctic fauna. These insects are unusual in the Lygaeidae in being chiefly predaceous on other small arthropods. The “big-eyed bugs” have been studied intensively in recent years as possible bio¬ logical control agents against several destructive insects. HENESTARINAE. Body Strongly punctate; eyes stalked; pronotal margin sinuately convex posteriorly; hind wing with hamus and intervannals, latter basally fused; non- spinous forefemora; spiracles ventral on abdominal seg¬ ments 2, 6, and 7, dorsal on segments 3-5. This group contains three genera and 19 species. The distribution is chiefly southern Palearctic and African. Henestaris Spinola and Engistus Fieber are widespread in the Palearctic. HENICOCORINAE. Forewing with reduced membrane and tendency to coleoptery; hind wings absent; setae with strongly raised granular bases; metathoracic scent-gland auricle with a blocklike spout; lateral trichobothria lack¬ ing on sterna 3-5, sterna 6-7 each with 2 lateral tricho¬ bothria placed transversely behind level of spiracle; all abdominal spiracles ventral; inner laterotergites present on segments 2-4; no spermatheca; nymphal dorsal ab¬ dominal scent-gland openings present between terga 4/5 and 5/6; roof of genital chamber of female with a large membranous sac. This monotypic subfamily was established by Wood¬ ward (1968a) for Henicocoris monteithi Woodward from Victoria, Australia. HETEROGASTRINAE. Membrane of forewing with 1 or 2 closed cells at base; hamus and intervannals present in hind wing; all abdominal spiracles ventral; spermatheca elongate, coiled, nonflanged, in common with Pachy- gronthinae; nymphal abdominal scent-gland openings be¬ tween terga 3/4, 4/5, and 5/6. Twenty-two genera and 92 species are recognized. The group is widespread in the Old World tropics and has sev¬ eral Palearctic genera. The New World fauna consists of two Nearctic species of Heterogaster Schilling. ISCHNORHYNCHINAE. Small, usually dull brownish or reddish brown, frequently ovoid, sometimes shining or subshining; membrane often hyaline; corium frequently translucent; clavus punctate; posterior margin of prono¬ tum nondepressed; nymphs with dorsal abdominal scent- gland openings between terga 4/5 and 5/6 (except in Kleidocerys Stephens, with scent glands between terga 3/4). Acanthocrompus Scudder is unique within the sub¬ family in having the forefemora strongly incrassate and heavily spined below, although the condition is wide¬ spread in the Rhyparochrominae. Fifteen genera and 75 species are currently recog¬ nized. Kleidocerys Stephens is widespread in the North¬ ern Hemisphere, but the majority of Ischnorhynchinae are tropical or south temperate in distribution. Pvlorgus Stal is widespread in mountainous areas of Africa and the Orient. LYGAEINAE. Hemelytra impunctate; membrane of fore¬ wing usually possessing a distinct cell, subcostal vein, and hamus; all abdominal spiracles dorsal; nymphs with dorsal abdominal scent-gland openings between terga 4/ 5 and 5/6. This large subfamily is found worldwide. Fifty-eight genera and about 500 species are currently recognized. As with most lygaeid taxa, the greatest diversity is in the tropics and subtropics. A few genera occur in both the Old World and New World. Many species are large, with showy red and black or orange and black aposematic coloration, although other color combinations are also present. Most lygaeines feed above the ground, although many will live among ground litter when seeds from the host plant are abundant there. A small component appears secondarily geophi- lous; some of these species are flightless and cryptically colored. Examples include Apterola Mulsant and Rey (five species; Africa, southern Palearctic), Melanerythrus Stal (three species; Australia), Stenaptula Seidenstucker (two species; Africa and India), and Lygaeospilus Barber (four species; Nearctic). ORSILLINAE (FIG. 83.3B). Relatively small, dull, gray brown; hemelytra in large part impunctate; hind wing with hamus, lacking subcostal vein; abdominal spiracles all dorsal; inner laterotergites present on abdominal terga 2—6; nympnal dorsal abdominal scent glands present be¬ tween terga 4/5 and 5/6. Four tribes are currently recognized, comprising a total of 28 genera and 250 species: Lepionysiini, Metrargini, Orsillini, and Nysiini. Ashlock (1967) monographed the world genera. Although the orsillines occur worldwide, in temperate as well as tropical areas and on many oceanic islands, a striking feature of this group is its tremendous radiation on the Hawaiian Islands, with at least half of the world species occurring there (Usinger, 1942b). Species of the Lygaeidae 257 Fig. 83.4. Lygaeidae: Pachygronthinae. Pachygrontha sp. endemic genera Oceanides Kirkaldy and Neseis Kirkaldy tend to be restricted to one island and are often host- specific. By contrast, the 19 species of Nysius Dallas endemic to Hawaii occur on all of the major islands and are generalist feeders. OXYCARENINAE. Usually small; often flattened; some¬ times strongly myrmecomorphic; head commonly por- rect; hemelytra with explanate margins; pronotal mar¬ gins rounded, nonexplanate; hamus absent; abdominal spiracles 3-7 ventral; inner laterotergites absent; abdomi¬ nal sternum 5 with at most a single posterior trichoboth- rium. Twenty-three genera and 144 species are known. The group is most diverse in the Palearctic and is poorly repre¬ sented in the Western Hemisphere. Sarny (1969) revised the extensive African Oxycarenus Fieber fauna. PACHYGRONTHINAE (FIG. 83.4). Body coarscly punctate; forefemora armed; abdominal spiracles ventral; sperma- theca peculiar, elongate, nonflanged; nymphal scent glands present between terga 4/5 and 5/6. Slater (1955) treated the world fauna of this primarily tropical and subtropical group, providing keys to genera and species. The Pachygronthini, comprising elongate, slender, bright brown bugs with long sweeping anten- Fig. 83.5. Lygaeidae; Psamminae. Saxicoris verrucosus Slater (from Slater, 1970). nae, contain four genera and 50 species. The Teracriini are usually relatively short and stout, although elongate species occur in the genera Cymophyes Fieber and Steno- phyella Horvath, but like most other members of the tribe they have short, stout antennae. Nine genera and 28 species are known. PSAMMINAE (FIG. 83.5). Small, beetle-like; length not over 3-4 mm; body surface bearing waxy scalelike setae (Slater and Sweet, 1965); eyes large, reniform; ocelli lacking; antennae short and subclavate; hemelytra highly convex, completely covering abdomen and meeting along midline; membrane of forewing and entire hind wing lacking, clavus and corium indistinguishably fused; tarsi 2-segmenteil; spiracles dorsal on all abdominal segments; nymphal dorsal abdominal scent-gland openings between terga 4/5 and 5/6; trichobothria reduced, none on seg¬ ments 4 and 7, a single one located mesally on segment 5; inner laterotergites present. Three genera—all monotypic—are known. Psam- mium Breddin and Saxicoris Slater are found in the xeric regions of South Africa and Namibia, and Sympeplus Bergroth is known from India. RHYPAROCHROMINAE. Usually dull brown or mottled brown, black and white; frequently myrmecomorphic; 258 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) cephalic trichobothria usually present; forefemora usually incrassate, strongly armed below with stout spines; suture between sterna 4 and 5 fused, usually curving forward anterolaterally from midline of sternum, not reaching dorsal margin of abdomen. The subfamily was established by Amyot and Serville (1843) as Rhyparochromides. It was called Myodochina by Stal (1872), Pachymeriidae by Uhler (1860), Aphanini by Puton (1887), and Megalonotinae by Slater (1957). The basic separation into tribes was by Stal (1872), who recognized the Lethaeini, Drymini, Myodochini, Rhy- parochromini, and Gonianotini. Stal (1874) later added the Cleradini, and Guide (1936) the Stygnocorini. Scud- der (1957b) discussed tribal relationships, placing some of Stal’s tribes as subtribes, and expanded the concept of the Stygnocorini. Slater (1957) separated the Megalono- tini from the Rhyparochromini. Slater and Sweet (1961) established the Plinthisini. Ashlock (1964) recognized the polyphyletic nature of the Lethaeini and removed the Antillocorini and Targaremini as distinct tribes. Sweet (1964) recognized the Ozophorini and later (Sweet. 1967) reviewed the higher classification in detail and removed the Udeocorini from the Myodochini. Slater and Wood¬ ward (1982) established the Lilliputocorini, Because this subfamily is so large and diverse, and because the tribal classification is well established, we present a key to the tribes and briefly discuss each of them. Key to Tribes of Rhyparochrominae 1. All abdominal spiracles located on sternum . 2 - At least spiracle of abdominal segment 4 located dorsally on outer laterotergite . 12 2. Posterior pair of trichobothria on abdominal sternum 5 located one above the other. 3 - Posterior pair of trichobothria on abdominal sternum 5 located one in front of the other so that the 3 abdominal trichobothria of segments 4 and 5 occur as a linear series . 10 3. Females with a conjunctiva present between abdominal sterna 4 and 5; males with a stridulatory mechanism involving abdominal segment 1 (plectrum) and hind wing (stridulitrum); pronotum wider across anterior one-third than across humeral angles . Plinthisini - Both sexes with abdominal sterna 4 and 5 fused, lacking a conjunctiva between them; males lack¬ ing an abdominal and hind wing stridulatory mechanism; pronotum variable, usually wider across humeral angles than across anterior lobe . 4 4. Ocelli lateral, behind eyes; suture between abdominal sterna 4 and 5 attaining lateral connexival mar¬ gin; abdominal tergum 3 usually desclerotized; labial segment 2 usually not attaining base of head; nymphs lacking a Y-shaped suture but with a lateral suture along length of abdomen .... Cleradini - Ocelli located between and slightly posterior to eyes; suture between abdominal sterna 4 and 5 usually not attaining lateral connexival margin and usually markedly curving anteriorly from venter dorsally; labium variable, but usually with segment 2 reaching or exceeding base of head . 5 5. Nymphs lacking a Y-suture . 6 - Nymphs with a Y-suture . 7 6. Nymphal abdominal scent-gland openings only between terga 3/4 and 4/5; tarsi 2-segmented; inner laterotergites absent; metathoracic scent-gland auricle strongly curving anteriorly . . Lilliputocorini - Nymphal abdominal scent-gland openings between terga 3/4, 4/5 and 5/6; tarsi 3-segmented; inner laterotergites present; metathoracic scent-gland auricle straight or curving posteriorly . . Antillocorini 7. Abdominal trichobothria on sternum 5 closer to spiracle thin to posterior margin of segment 5; pronotum lacking a distinct anterior collar; hind wing usually with both hamus and secondary veins . 8 - Abdominal trichobothria on sternum 5 located closer to posterior margin of sternum 5 than to ' spiracle; distinct pronotal collar present; hind wing veins reduced, either hamus or secondary veins absent (usually both) . 9 8. All trichobothria on sterna 4 and 5 located anterior to spiracle of sternum 5; spiracle 5 located in central third of segment; pores present near spiracles of segments 3 and 4 . Drymini - Posterior trichobothria of sternum 5 located posterior to spiracle 5; spiracle 5 located in posterior third of segment; no pores present near spiracles 3 and 4 . Stygnocorini 9. Inner laterotergites present; suture between abdominal sterna 4 and 5 incised, straight and attaining Lygaeidae 259 10 . 11 . 12 . 13. connexival margin (Fig. 83. IB); spermathecal duct short, only 3 times as long as bulb; abdominal scent-gland scars small . Phasmosomini Inner laterotergites absent; suture between abdominal sterna 4 and 5 sometimes attaining con¬ nexival margin, usually remote from margin and strongly curving dorsoanteriorly (Fig. 83.1 A); spermathecal duct very elongate and coiled; abdominal scent-gland scars broad . Ozophorini All trichobothria on segment 5 located anterior to spiracle and usually equidistant from each other; nymphs with a Y-suture . Targaremini Usually with one trichobothrium on segment 5 posterior to spiracle; middle trichobothrium not equally distant from the other 2; nymphs lacking a Y-suture . 11 Apical corial margin deeply concave; inner laterotergites present; head lacking iridescent areas; abdominal scent-gland scars present between terga 3/4. 4/5, and 5/6 . Antillocorini Apical corial margin straight; no inner laterotergites present; head frequently with iridescent areas present basally; abdominal scent-gland scar between terga 5/6 minute or absent . Lethaeini Abdominal spiracles of segments 2-4 located dorsally . 13 Abdominal spiracles located dorsally only on either segments 3 and 4 or 4 only, always ventral on segment 2 . 14 Inner laterotergites absent; lateral pronotal margins almost always rounded; nymphs lacking large black sclerotized areas around dorsal abdominal scent-gland openings . Myodochini Inner laterotergites present; lateral pronotal margins variable from rounded to carinate; nymphs frequently with black sclerotized areas around dorsal abdominal scent-gland openings . . Udeocorini 14. Spiracles located dorsally only on abdominal segment 4 . Gonianotini - Spiracles located dorsally on abdominal segments 3 and 4 . 15 15. Nymphs with a distinct Y-suture present between abdominal terga 3/4 . Rhyparochromini - Nymphs lacking a Y-suture between abdominal terga 3/4 . Megalonotini ANTILLOCORINI (FIG. 83.6A). Very Small to minute; buccu- lae joined by a carina well behind labium; anterolateral pronotal trichobothria absent; apical corial margin deeply concave; inner laterotergites present; abdomen laterally with evaporative areas; well-developed nymphal scent glands between terga 3/4, 4/5, and 5/6; trichobothria linearly arranged on all segments. Twenty-nine genera and at least 93 species are known. This group is primarily tropical and subtropical, with a few species extending into the temperate Northern Hemi¬ sphere, and is well represented on oceanic islands in the Pacific. The group may not be monophyletic, because some Neotropical species lack the linear arrangement of abdominal trichobothria and do not have the deeply concave apical corial margin. Unfortunately, nymphs of these species are unknown, precluding determination of whether they possess the abdominal evaporative areas that are apparently a synapomorphy for the Antillocorini and Lethaeini. CLERADiNi. Ocelli located behind rather than between eyes; antennal segment 3 short; labium relatively short, segment 2 usually not exceeding base of head; forefemora usually slender, unarmed below; no inner laterotergites; connexival membrane greatly expanded; parameres bifur¬ cate. The body form ranges from short and stout to large with an elongate neck. Species of several genera pos¬ sess stridulatory mechanisms incorporating an abdominal stridulitrum and a femoral plectrum. Nymphs uniquely possess an impressed lateral suture running the length of the abdomen. Most remarkable is the presence of a dis¬ tinct pseudoperculum otherwise not present in the eggs of Lygaeidae. The Cleradini are confined to the tropics of the Eastern Hemisphere (except for the introduction of Clerada api- cicornis Signoret into the Western Hemisphere). Twenty genera and 50 species are currently recognized. Mali- patil (1981) revised the extensive Australian fauna, and Malipatil (1983) revised the world fauna and analyzed the cladistic relationships. DRYMiNi. Usually small to medium-sized; lateral prono¬ tal margins usually carinate or narrowly explanate; apical corial margin usually straight; posterior pair of tricho¬ bothria on sternum 5 located dorsoventrad of one another, placed anterior to spiracle 5; pore present near spiracles 3 and 4; usually with 9 chromosomes. Fifty-two genera and 269 species are currently known. The group is speciose in the Old World. There is a limited Nearctic fauna closely related to that of the Palearctic, but the group is absent from the Neotropics. GONIANOTINI. Body usually broad, elliptical, with strongly explanate lateral pronotal margins; spiracles of abdominal segment 4 dorsal; nymphal abdomen usually dark, heavily sclerotized; no Y-suture; nymphs with dor¬ sal abdominal scent-gland openings between terga 4/5 and 5/6. 260 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) t Fig. 83.6. Lygaeidae: Rhyparochrominae. A. Botocudo cavemicola Slater (Antillocorini) (from Slater, 1984). B. Primiews quadrispinosus Slater and Zheng (Ozophorini) (from Slater and Zheng, 1985). Twenty-two genera and approximately 123 species are currently recognized. Gonianotines are chiefly Paleetrc- tic and Ethiopian and, although present in other faunal regions, are absent from the Neotropical and Australian regions. LETHAEiNi. Usually shining to subshining; head with single or double iridescent area(s) at base; bucculae - joined posteriorly by carina immediately behind labial base; pronotal margins usually carinate or explanate; anterolateral angle of pronotum frequently with an elon- gate trichobothrium; nymphs lacking Y-suture but with y evaporative areas laterally; Y chromosome lacking (syna- pomorphy); sperm reservoir complex, highly modified (O’Donnell, 1991). Lethaeines comprise 33 genera and 150 species. They are chiefly tropical, although a few species extend north¬ ward into either the Palearctic or the Nearctic. -- LILLIPUTOCORINI. Minute, 2 mm long; body yellowish brown, somewhat flattened; antennae rather clavate; fore¬ femora mutic; tarsi 2-segmented; abdominal spiracles ventral; abdominal trichobothria variably placed, poste¬ rior pair on sternum 5 sometimes lacking, when present placed in dorsoventral position; inner laterotergites lack¬ ing; nymphal scent glands between terga 3/4 and 4/5, troughed groove present between terga 3/4 and 4/5 lead¬ ing to evaporative areas laterally on abdomen; ovipositor reduced. One genus and 10 species are known. The Lilliputo- corini are known from northern Australia, New Guinea, Borneo, Ceylon, Nepal, Seychelles, Ghana, South Af¬ rica, and Bra.zil (Slater and Woodward, 1982). Nothing is known of the biology of these minute insects other than that they live in ground litter. Wing reduction is common with both staphylinoid and micropterous forms known. Wing reduction is sexually dimorphic, a condi¬ tion common in some families of Heteroptera, but almost unknown in the Lygaeidae. MEGALONOTINI. Usually resembling Rhyparochromini, with no character yet discovered to differentiate these two tribes in the adult stage. Both have carinate or ex- Lygaeidae 261 planate pronotal margins, spiracles on abdominal seg¬ ments 3 and 4 dorsal, inner laterotergites present, and 3 well-developed nymphal abdominal scent-gland open¬ ings. Megalonotine nymphs have dark, heavily sclero- tized abdomens and lack a Y-suture, a character that is well developed in nymphs of the Rhyparochromini. Eighteen genera and approximately 87 species are known. The Megalonotini, most diverse in the Old World tropics and the Palearctic. are represented by a few taxa in the Nearctic and are absent from the Neotropics. MYODOCHiNi. Body form variable, ranging from short and stout to elongate and slender, from rather small to very large; often myrmecomorphic; myodochines share the following features with the Udeocorini; anterior pro¬ notal lobe almost always rounded laterally, abdominal spiracles 2-4 located dorsally. nymphs with Y-suture, 3 pairs of abdominal scent glands, inner laterotergites absent. Sixty-seven genera and approximately 307 species are known. Myodochini are abundant in all major zoogeo¬ graphic regions, sometimes reaching high latitudes in temperate areas. Some species also appear to be ex¬ tremely vagile, and there has been colonization of many oceanic islands. The Neotropical fauna is especially di¬ verse. Harrington (1980) provided a cladistic analysis of the world genera. 020PH0RINI (FIG. 83.6B). Medium-sizcd, rather slender; head generally porrect, grooved; pronotal collar usually well developed; legs and antennae elongate; hind wing lacking hamus and secondary veins; spiracles ventral; nymphal dorsal abdominal scent glands between terga 3/ 4, 4/5, and 5/6, strongly developed Y-suture in nymphs; inner laterotergites absent. Twenty-four genera and 173 species are currently rec¬ ognized. Some species are myrmecomorphic, and some show body shape development parallel to that of myodo¬ chines. Ozophorines are worldwide in distribution. They are most abundant in the Neotropics and in the New Guinea island arc. PHASMOSOMINI. Ocelli absent; cubital furrow absent; Y- suture present; inner laterotergites present; deeply incised suture between sterna 4 and 5; 4/5 abdominal suture complete although fused. This tribe contains two south-central Palearctic spe¬ cies. They are similar in habitus to many species of Ozophorini. PLiNTHisiNi. Small; shining or subshining; usually with pronotum expanded across anterior lobe (especially in flightless forms); forefemora heavily incrassate, spined; wings often greatly reduced, usually in a staphylinoid manner, posterior half of abdominal dorsum exposed; stridulatory mechanism novel (as noted in key); sperm reservoir continuous with body when not redueed. Uniquely within Rhyparochrominae, the Plinthisini have a conjunctival membrane present between abdomi¬ nal sterna 4 and 5 in the female, the lack of which feature has been used as a synapomorphy for the Rhy¬ parochrominae (Sweet, 1967). Thus, the Plinthisini may merit subfamily status. The presence of head trichoboth- ria, however, suggests that this may be the sister group of all other rhyparochromines. There are nearly 100 described species, with large numbers of undescribed species present in collections. The group occurs worldwide. RHYPAROCHROMINI. Often large and robust; lateral pro¬ notal margins carinate or explanate; nymphs frequently with variegated, lightly sclerotized abdomen and well de¬ veloped Y-suture; inner laterotergites present; 3 nymphal dorsal abdominal scent-gland openings between terga 3/ 4, 4/5, 5/6; spiracles of abdominal segments 3 and 4 located dorsally. Forty-one genera and approximately 350 species are currently recognized. The Rhyparochromini are most di¬ verse in the Old World tropics and subtropics, with an extensive Palearctic fauna. The group is poorly developed in the Nearctic and natively absent from the Neotropics. Eyles’s (1973) revision of the genus Dieuches Dohrn is the single most comprehensive work on this large and complex group. STYGNOCORiNi. Short, rather stout-bodied; vertex with¬ out longitudinal grooves; hamus and secondary veins strongly developed; abdominal spiracles ventral; inner laterotergites present; no pronotal collar; nymphs with Y- suture; nymphal dorsal abdominal scent-gland openings between terga 3/4, 4/5, and 5/6. Sixteen genera and 168 species are currently recog¬ nized. This group includes several Palearctic genera, a montane element in Africa and Madagascar, and a south- temperate group in Africa, Tasmania, and New Zealand (see Slater and Sweet, 1970; O’Rourke, 1974, 1975). TARGAREMiNi. Small to medium-sized; all abdominal spiracles ventral; trichobothria on abdominal sternum 5 located anterior to spiracle, usually equidistant from one another, in linear sequence as in Lethaeini and many Antillocoriini; nymphs with a Y-suture. Twenty-three genera and 59 species are known. The Targaremini appear to be an ancient group, being found in Australia, New Guinea, New Caledonia, the New Hebri¬ des, and New Zealand. On the last islands and associated islets there has been extensive radiation (Malipatil, 1977). UDEOCORINI. Medium-sized to large; body form vary¬ ing from elongate, slender, and long-legged, with rounded pronotal margins, to short, stout, and subflattened, with carinate, or even explanate, pronotal margins; spiracles dorsal on abdominal segments 2-4; Y-suture well devel¬ oped in nymphs; nymphal dorsal abdominal scent-gland 262 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) openings between terga 3/4, 4/5, 5/6; inner laterotergites present. Seventeen genera and 33 species are currently recog¬ nized. Udeocorines are found chiefly in Australia, where they have radiated. They have the appearance of other tribes in other regions of the world and (presumably) occupy the same niches. There are also three Neotropical taxa. Bathycles amarali Correa apparently is haematopha- gous and has converged remarkably in general habitus to species of Clerada Signoret. Specialized morphology. Nowhere else in the Het- eroptera do we observe such marked differences in the position of the spiracles or in the Trichophora such varia¬ tion in the number and position of the abdominal tricho- bothria (Sweet, 1967). Many lygaeids possess stridula- tory structures. These often occur as stridulitra along the lateral margins of the hemelytra, as crescent-shaped areas on the abdominal sternum, along the sides of the head and the prothorax, or on the hind wings. The forefemora are sometimes fossorial, but much more frequently are enlarged and spinose, appearing rap¬ torial, but actually serving either to hold and manipulate seeds or to aid in dragging the body in enclosed spaces. Many species are myremcomorphic, with constricted anterior abdominal segments and expanded posterior seg¬ ments as well as conical antlike protrusions on both the pronotum and scutellum. Brachyptery, coleoptery, and microptery are wide¬ spread (see Chapter 6). Natural history. Although the great majority of Ly- gaeidae feed on mature seeds (Sweet, 1964), there is sub¬ stantial diversity in the feeding habits for the family. The Blissinae are sap suckers, most Geocorinae are largely predaceous on other small arthropods, and the Cleradini feed on the blood of vertebrate hosts. There is a very large ground-living component. In some species, espe¬ cially those from stable habitats, the flying morph appears to have been completely eliminated. Although many members of the genus Nysius are gen¬ eralists, some orsillines are specialists. These often live on the seeds of trees, such as Belonochilus numenius (Say) on Platanus spp. (sycamores) in North America and Hyalonysius pallidomaculatus Slater on Buddleia spp. in South Africa. Species of the Holarctic genus Kleidocerys often are found in large numbers on catkins and seed heads of Be- tula. Rhododendron, Spiraea, Typha, and other plants. They produce sound by means of a stridulitrum on a vein of the hind wing that contacts a plectrum on the front wing (Southwood and Leston, 1959). Such strjdulation can be heard by the human ear by placing an insect in a small vial and shaking it vigorously. The various species of the genus Stilbocoris Bergroth (Rhyparochrominae: Drymini) are remarkable in being ovoviviparous. Males have a complex mating ritual in which they secrete salivary fluid into fig seeds, which they then offer to females to facilitate copulation. The males frequently fly about with the small fig seeds impaled upon their beaks (Carayon, 1964a). Aposematic coloration is widespread in the Lygaei- nae, many species of which feed on toxic or unpalat¬ able plants. They are involved in complex Mullerian and Batesian mimicry rings with beetles and moths, as well as other heteropterans (see Chapter 7). There is an interesting case of interspecific displace¬ ment of other insects from a host-plant by the apo- sematically colored red and black Neacoryphus bicrucis (Say) in North America. This species feeds on Senecio spp. (Asteraceae), from which it sequesters pyrrolizidine alkaloids and apparently is protected against attack by potential predators. It co-occurs with at least five other insect species, including members of the Miridae, Rho- palidae, and Coreidae. Males are extremely aggressive in attempting to copulate with other species as well as with females of their own species. This behavior drives the other species from the host plants. When Neacory’- phus males are removed, populations of the other species increase and vice versa (McLain and Shure, 1987). This interspecific displacement by pseudocompetition may be an important component in the success of several of the aposematically colored Lygaeinae. Oncopeltusfasciatus (Dallas), known as the large milk¬ weed bug, has been used for many years as a laboratory animal, and there is an enormous literature dealing with various aspects of its physiology and biochemistry. This species has a flight cycle and a reproductive cycle during which energy may be directed to flight or to egg produc¬ tion with concomitant loss of flight musculature in the latter case and marked dispersal in the former (Dingle, 1978, 1979, 1981, 1985). Work in the Western Hemi¬ sphere tropics has shown interesting aspects of habitat segregation even on the same host plants by closely re¬ lated species of Oncopeltus (Blakley, 1980; Dingle and Baldwin, 1983). For example, some species are depen¬ dent for reproductive success upon mature seeds and, in order to find new plants with mature seeds, have in¬ creased dispersal ability. Other species are able to breed on the host plant whether or not seeds are present, and such species are relatively more sedentary. Some lygaeids feed above ground in the seed heads of plants. Many of these, especially the nymphs of species of Cyminae and Pachygronthini, resemble the seed heads of the sedges and rushes upon which they feed. Others, however, live in such habitats but have the body flat¬ tened rather than resembling seeds. One of the best Lygaeidae 263 known of these is the Palearctic Chilacis typhae Perrin, all stages of which live on and overwinter in the catkins of Typha species. This species recently was introduced into the eastern United States, where it is now established (Wheeler and Fetter, 1987). In South Africa, several Oat- carenus spp. have adapted themselves for feeding on the deeply set seeds of Protea (Proteaceae) and have an enor¬ mously elongated labium that in nymphs may be nearly one-third longer than the body length of the insect. Many species of Heterogastrinae, especially those that feed on Ficus, have very elongate ovipositors. Two rather distinct groups occur within the subfamily. One occurs on the bark and trunks of Ficus spp. and feeds by inserting the very elongate rostral stylets through the synconium of the fig to reach the seeds within. These species, many of which belong to the genus Dinomachus Distant, seem to be gregarious, with large numbers of adults and nymphs occurring together on the host plants. The second is short and stout and occurs primarily on Labiatae (Slater, 1971). In Trinidad and Peru, Cligenes subcavicola Scudder lives in caves inhabited by fruit-eating bats, where it sometimes occurs in tremendous numbers, feeding on the seeds present in the guano. Scudder et al. (1967) reported numbers from 1000 to 100,000 per square meter in the Tamana Caves in Trinidad. We saw thousands of the bugs on the floor of an abandoned building at the Simla Tropi¬ cal Research Station on Trinidad, where there roosted a large bat population that was feeding chiefly on fruits of a Piper sp. At the Cueva de las Lechusas near Tingo Maria, Peru, the same lygaeid apparently fed primarily on fig seeds. Sweet (1964), in a major study of the biology of the Rhyparochrominae of northeastern North America, dis¬ covered that the fauna could be divided into two eco¬ logical groups. The first group was found in temporary habitats, had bivoltine life cycles, produced large num¬ bers of eggs, usually had weak diapause as adults, and was entirely macropterous. The second group was found in more permanent habitats, had univoltine life cycles, produced more limited numbers of eggs, had strong dia¬ pause, often in the egg stage, and most species were largely brachypterous. Distribution and faunistics. The family is worldwide in distribution. A number of taxa show distribution pat¬ terns that imply past continental connections, as between Africa and South America, transantarctic distributions between Australia and southern South America, trans- siberian distributions between boreal Asia and North America, and a few that appear to represent more wide¬ spread Gondwanaland relationships. Putchkov (1969) presented keys to the fauna of the Ukraine. This extensive treatment also includes detailed host plant and ecological data and figures of many nym- phal stages. Kerzhner and Jaczewski (1964) presented keys to the entire European fauna of the former USSR, including almost all of the Palearctic genera. Slater and Baranowski (1990) treated the fauna of Florida and con¬ sequently provided access to much of the eastern North American fauna. Slater (1964a) treated the South African fauna in detail. Many other papers should be consulted for identifica¬ tion of various groups within the family. Among them are those by Harrington (1980) and Malipatil (1978) for the Myodochini, Ashlock (1967) for the Orsillinae, A. Slater (1985, 1992) for the Lygaeinae, Gross (1965) for Aus¬ tralian Drymini, Gross and Scudder (1963) for Australian Rhyparochromini, Scudder (1962b) for the Heterogastri¬ nae. and Scudder (1962c) for the Ischnorhynchinae. 84 Malcidae General. Members of this family are small (3-4 mm), thick-bodied (Fig. 84.1 A), and coarsely punctate (Fig. 84. IB), with flattened scalelike or eurved glandular setae on the body surface. They have no common name. Diagnosis. Head strongly declivent; ocelli present; bucculae large; antennae placed above a line drawn through middle of e\'e; membrane of forewing with five nonbranching veins (Fig. 84.1 A, B); tarsi 3-segmented; abdominal sterna 2-5 fused; inner laterotergites absent; trichobothria located on “loaflike” tubercles, submedial on sternum 3 as well as laterally where 2 arranged diago¬ nally, missing on sternum 4 (Fig. 84. ID); lateral margins of abdominal segments 5-7 usually expanded into dis¬ tinct flanges; spiracles dorsal on abdominal segments 2- 6; nymphal dorsal abdominal scent glands small, situated between terga 3/4, 4/5, and 5/6 or between terga 4/5 and 5/6; aedeagus as in Fig. 84. IE; parameres as in Fig. 84. IF; ovipositor laciniate; sternum 7 entire; spermatheca often with elongate ductus (Fig. 84. IG); eggs quadrate in cross section. Classification. This taxon was established by Stal (1864-1865) as Malcida in a key to subfamilies of Ly- gaeidae. Despite the lack of included taxa, the name was obviously based on Malcus Stal (1859c). Stys (1967c) reviewed the history of the group in detail, noting that many authors credited authorship to Horvath (1904), who first used the term Malcinae. Subsequent authors usually treated the taxon as a lygaeid subfamily, although Lethierry and Severin (1893-1896) and Distant (1904) placed it in the Colobathristidae. 264 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 84.1. Malcidae (from Stys, 1967c). A. Malcus japonicus Ishihara and Hasegawa, adult, B. Forewing, M. furcatus Stys, C. Fifth-instar nymph, M. flavidipes Stal. D. Ventral view, male abdomen, M. furcatus. E. Aedeagus, sagittal view, M. furcatus. F. Paramere, M. furcatus. G. Spermatheca, M. furcatus. Abbreviations Itb, lateral trichobothria: mtb, mesial trichobothria. Stys (1967c) discussed the relationships of the Malci- eluded that the Chauliopinae and Malcinae were closely dae—and other lygaeoid groups—exhaustively. He con- related and together merited family status. Key to Subfamilies of Malcidae 1. Eyes stylate; ocelli widely separated from one another; body bearing waxlike scales; claval commis¬ sure much shorter than scutellum . Chauliopinae - Eyes sessile (Fig. 84.1 A); ocelli close together, situated on a common tubercle; body lacking waxlike scales; claval commissure subequal in length to scutellum (Fig. 84.1 A, B) .. Malcinae Malcidae 265 CHAULIOPINAE. Body short and stout; antenniferous tu¬ bercles large, spinous, produced; metathoracic scent- gland auricle not produced; no tubercle at apex of corium; abdominal terga 3-6 separate; nymphs lacking elongate spines; nymphal dorsal abdominal scent glands paired between terga 4/5 and 5/6. The Chauliopinae was erected as a subfamily of Ly- gaeidae by Breddin (1907) but was placed in the Hetero- gastrinae by Distant (1910), an action followed much later by Miller (1956a). Other authors, including Stys (1963), treated the group as a subfamily of Lygaeidae. Two genera are recognized, Chauliops Scott (seven species) and Neo- chauliops Stys (two species from Africa) (Stys, 1963). The Chauliopinae occur in both the Oriental and Ethio¬ pian regions. MALCINAE (FIG. 84.1A). Body relatively elongate; anten¬ niferous tubercles reduced; metathoracic scent-gland au¬ ricle protruding, dorsally perpendicular to metapleuron; corium with a conspicuous tubercle at apical angle; ab¬ dominal terga 3-6 fused; nymphs with numerous long spines over body surface (Fig. 84.1C); dorsal abdominal scent glands paired between terga 3/4 and 4/5, unpaired between terga 5/6; eggs with 3 club-shaped micropylar processes, lacking pseudoperculum. One genus, Malcus Stal containing 19 species from the Oriental region, is known. The subfamily was mono¬ graphed by Stys (1967c). Speciali2ed morphology. The elongate nymphal spines of the Malcinae (Fig. 84.1C), fused abdomi¬ nal sterna 2-5, tuberculate trichobothria (Fig. 84.ID), flanged abdominal segments, stylate eyes, and frequent occurrence of glandular setae are all specialized features. Natural history. The biology of Malcus spp. is essen¬ tially unknown. Ishihara and Hasegawa (1941) reported on Malcus japonic us Ishihara and Hasegawa on Morus bombycis. Stys (1967c) recorded Malcus flavidipes Stal as abundant on leaves of banana in Cambodia. Species of Chauliops apparently feed chiefly on Solanaceae but are sometimes destructive to beans (Sweet and Schaefer. 1985; see Chapter 8). Distribution and faunistics. Stys (1963:213) sug¬ gested “that Burma and the surrounding mountains on the west must be considered a center of speciation and probably also the original country of both subfamilies.” The basic reference on the group is Stys (1967c). 85 Piesmatidae General. These small insects, not over 5 mm in length, have reticulate cell-like forewings and prothorax (Fig. 85.1 A). Superficially they resemble small grayish or yel¬ lowish lace bugs. They are sometimes referred to as ash-gray leaf bugs. Diagnosis. Body surface reticulate (Fig. 85.1 A): ocelli present; mandibular plates elongate, projecting strongly forward (Fig. 85.1 A); scutellum exposed; meta¬ thoracic scent-gland openings obsolete; hind wing with stridulitrum on Cu vein and plectrum on first tergum; tarsi 2-segmented; nymphs with scent glands on abdominal terga 3/4 and 4/5 (small, sometimes visible only be¬ tween terga 3/4), these apparently functional in adults; one prespiracular trichobothrium present on abdominal sternum 5 and one on 6 in Piesma (sometimes one pair or none); some, or all, abdominal spiracles dorsal; aedeagus as in Fig. 85. IB, vesica greatly elongate; parameres as in Fig. 85.1C; spermatheca as in Fig. 85. ID, E; 2 pairs of Malphigian tubules opening into anterior end of rectum (apparent apomorphy for family). Classification. This taxon was first recognized as a higher group by Amyot and Serville (1843) as the “Pies- mides.” Early classifications placed the Piesmatidae with the Tingidae because of the reticulate wings. Reuter (1910) considered them a distinct family. Tullgren (1918) and Leston et al. (1954) removed the family from a posi¬ tion related to the Tingidae to the Pentatomomorpha. Drake and Davis (1958) further showed that the presence of trichobothria and details of genitalic morphology in¬ dicated that the resemblance to some Tingidae was only superficial. Both Stys (1961b) and Kumar (1968) con¬ sidered the family to be isolated and believed it merited superfamily status. Two subfamilies have been recog¬ nized. PIESMATINAE (FIG. 85.1 A). Piesma Lcpeletier and Serville, with 31 species, is widespread and has been segre¬ gated into three subgenera by Pericart (1974). Of these, Parapiesma Pericart is Holarctic, Piesma sensu stricto is Holarctic and African, and the monotypic Afropiesma Pericart is African. Two other genera have restricted distributions: Miespa Drake (one species; Chile) and Mcateella Drake (four species; Australia). THAIOCORINAE. Kormilev (1969) erected this subfamily for Thaicoris Kormilev, with one species from south¬ east Asia. If Schaefer (1972a, 1981b) is correct, how¬ ever, Thaicoris is actually the sister taxon of Miespa 266 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 8S.1. Piesmatidae. A. Piesma capitatum (Wolff). B. Phallus, sagittal view, P. cinereum (Say). C. Paramere, P, quadratum (Fieber), D. Spermatheca, P. maculatum (A-D from Heiss and Pericart, 1983). E. Spermatheca, P. quadratum (from Pendergrast. 1957). and Mcateella, and therefore not deserving of subfamily status. Specialized morphology. The unique position of the Malpighian tubules, functional abdominal scent glands in adults, and greatly reduced trichobothrial numbers are all specialized features. Natural history. All species are phytophagous. Schae¬ fer (1981b) summarized known host-plant associations of this strictly phytophagous group, listing 10 genera of Chenopodiaceae as being utilized by various species of Piesma. Species of the Australian genus Mcateella have been reported feeding on Acacia and Beyeria. There are also records of feeding on species of Caryophyllaceae, Amaranthaceae, and Cistaceae. Distribution and faunistics. This small family occurs in all major zoogeographic regions. Schaefer (1981b) be¬ lieved that the distribution of the family “is not inconsis¬ tent” with a Gondwana origin. The works of Drake and Davis (1958) and Heiss and Pericart (1983) serve as basic references on the group. Piesmatidae 267 Fig. 86.1. Largidae. A. Acinocoris stehliki van Doesburg {from van Doesburg, 1966). B. Aedeagus, lateral view, Physopelta famelica Stal. C. Aedeagus, detail, P. famelica (B, C from Kumar, 1967). D, Paramere, A. calidus (Fabricius). E. Paramere, rotated 90°, A. calidus (D. E from van Doesburg, 1966). F. Spermatheca, Largus rufipennis Laporte (from Pluot, 1970). Pyrrhocoroidea 86 Largidae General. These are moderately small to large insects (up to 55 mm), which are frequently brightly colored. The body is ovoid, oblong, or elongate (Fig. 86.1), ex¬ cept in myrmecomorphic species (Fig. 86,2). often with relatively short antennae and legs. The group appears to have no common name. Diagnosis. Antennae inserted below a line through middle of t;ye; ocelli absent; membrane of forewing with basal cells and at least 7 distally radiating veins (Fig. 86.1 A); metathoracic scent-gland openings reduced; sometimes an incomplete suture between abdominal sterna 4 and 5; 3 trichobothria on sterna 3-6, 2 on ster¬ num 7, those on segments 5-7 lateral and dispersed; nymphal dorsal abdominal scent-gland openings between terga 3/4, 4/5, and 5/6, 2 anterior gland openings re¬ duced; aedeagus as in Fig. 86. IB, C; parameres as in Fig. 86. ID, E; ovipositor laciniate; abdominal sternum 7 cleft mesally; spermatheca as in Fig. 86. IF. 268 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Classification. The taxon was first recognized as a higher group, the Largides, by Amyot and Serville (1843). Van Duzee (1916) treated it as a subfamily of the Pyrrhocoridae, calling it Euryophthalminae. but separate family status has been accepted by most recent workers on the basis of great differences in the female genitalia (Stys and Kerzhner, 1975; Henry, 1988). Hussey (1929), in a world catalog, treated the largids as a subfamily, but stated that the differences were of such a magnitude that family status was warranted. He recognized two tribes: the Largini (his Euryophthalmini), confined to the Western Hemisphere, and the Physopeltini (Old World tropics). Bliven (1973), apparently unable to distinguish myr- mecomorphy from fundamental morphological features, considered the largids to belong to the Alydidae. In so doing he erected a second subfamily, the Arhaphinae, which may have some validity as a higher taxon although it represents only one (or more) myrmecomorphic genera and certainly does not reflect a fundamental reorganiza¬ tion. The corial vein connecting M and Cu is lacking and is considered by some to be a synapomorphy with the Hyocephalidae. Reduction of the anterior nymphal scent- gland orifices suggests a possible relationship with the Colobathristidae. Schaefer (1964) suggested that the Pyr¬ rhocoridae evolved from the Largidae, basing the idea in large part upon the laciniate ovipositor of the latter. Kumar (1968) disagreed, noting a number of special¬ ized features in the internal anatomy of the Largidae, and believed that the laciniate ovipositor was a secondary development. Approximately 15 genera and over 100 species are known. Specialized morphology. The lack of ocelli (shared with Pyrrhocoridae), complex venation of the membrane of the forewing, and frequent fusion and obliteration of abdominal sutures are all specialized conditions. Natural history. All known species feed on seeds and plant juices. Two rather distinct elements are present: one group of genera lives on the ground and resembles ground-living Lygaeidae; a second group lives on forbs, shrubs, and trees and resembles species of Pyrrhocoridae. The species of Euryophthalmus Laporte show bewilder¬ ing variability in color, with yellow and black or red and black morphs and -tvith strikingly different metallic blue nymphs, which are presumably aposematic. Several genera of largids are striking, not only with Fig. 86.2. Arhaphe Carolina Herrich-Schaetfer (from Slater and Bara- nowski, 1978). regard to body shape, but because of their movements in the field. Hussey (1927) illustrated the remarkably myrmecomorphic (Neotropical) Thaumastaneis montan- doni Kirkaldy and Edwards, which has a swollen, antlike head, abdomen strongly constricted at the base, wings reduced to short pads, and a strong spine projecting from the posterolateral areas of the pronotum. Species of the genera Arhaphe Herrich-Schaeffer (Fig. 86.2), Japetus Stal, and Theraneis Spinola also are strongly myrmeco¬ morphic, presumably either Mullerian or Batesian mimics of species of velvet ants (Mutillidae). Distribution and faunistics. The family is represented in all major zoogeographic regions, but is most abundant and diverse in the tropics and subtropics. Hussey’s (1929) catalog remains the most important reference. Froeschner (1981) provided a key to South American genera. The confusing literature of Bliven is summarized by Henry (1988). Largidae 269 87 Pyrrhocoridae General. This family includes species often referred to as cotton stainers. Most are medium-sized to large (8- 30 mm), frequently colored with red or yellow and black. Many closely resemble in size and shape species of the lygaeid subfamily Lygaeinae (Fig. 87.1). Diagnosis. Ocelli lacking; metathoracic scent-gland openings reduced; membrane of forewing with 2 basal cells and a series of 7-8 anastomosing veins distally (Fig. 87.1); 3 trichobothria on abdominal segments 3-6 and 2 on segment 7; sometimes a curved suture between abdominal sterna 4 and 5 not attaining dorsal margin (similar to condition in most rhyparochromine lygaeids); inner laterotergites absent; abdominal spiracles ventral; nymphs with dorsal abdominal scent-gland openings be¬ tween terga 3/4, 4/5 and 5/6, posterior openings reduced; aedeagus as in Fig. 87.2B; parameres as in Fig. 87.2C- E; abdominal sternum 7 of female complete; ovipositor platelike rather than laciniate; spermatheca lacking a dis¬ tal pump flange (Fig. 87.2F-H); alimentary canal as in Fig. 87.2A. Classification. Although the recognition of this family as a higher taxon is usually attributed to Fieber (1861), the group was actually first recognized by Amyot and Serville (1843) as the “Pyrrhocorides.” For many years, the Lar- gidae were included in the Pyrrhocoridae as a subfamily, but recent authors have believed these two groups to be distinct (see Chapter 86). Approximately 30 genera and 300 species are known. Specialized morphology. The platelike ovipositor valvulae, reduced scent glands in the adult, and loss of ocelli (shared with Largidae) are specialized conditions. Natural history. Most species whose biology is known feed chiefly on seeds and fruits, particularly of the Mal- vales. While the most conspicuous species are arboreal as adults, there is a significant Old World ground-litter fauna that presumably feeds on mature seeds. In West Africa, studies have shown a yearly succession of species, with large colonies frequently being com¬ posed of more than one species (Fuseini and Kumar. 1975). These colonies have associated with them a group of reduviids of the genus Phonoctonus that feed on Dys- dercus. Each reduviid species mimics the coloration of its pyrrhocorid prey species (Stride, 1956) (see Chapter 7). Derr et al. (1981) demonstrated that larger species of Dysdercus had a greater migratory capacity and a greater degree of survival of diapause than smaller species. The larger species in an area were entirely or largely confined Fig. 87.1. Pyrrhocoridae. Dysdercus fasciatus Signoret. to woody Malvales, whereas smaller species utilized her¬ baceous plants. Thus larger size appeared to confine such insects to large oil-rich fruits of large trees that were widely separated, and the greater migratory ability of such species was selected for as a consequence. In most of the colonial Dysdercus species, the gravid females absorb the flight muscles after a migration flight and at the onset of oogenesis (Davis, 1975). They then are no longer capable of migrating from one host to another. Carroll and Loye (1990) studied dimorphism in wing muscle histolysis \n Dysdercus bimaculatus Stal. Females lose flight ability prior to egg laying, but not until they have fed and mated. Thus the ability to use muscle nu¬ trients and energy for increased egg productivity must be balanced against the danger of low reproduction if a poor habitat is utilized. Several Dysdercus spp. feed on malva- ceous seed crops that are ephemeral because of feeding pressure by the bugs, and eggs are laid in large numbers and in a very short period of time. Nymphs thus pre¬ sumably have a chance to mature before the seed crop is exhausted. Females live for a much shorter period than do males, presumably because of exhaustion i 'm such massive egg production, which may account fo: ; c male sex bias so often found in these insects. Pyrrhocoris apterus (Linnaeus) is a widespread Pale- arctic species. It is a ground-living insect, and most popu¬ lations are predominately brachypterous. This species has been studied with respect to the function of juvenile hor- 270 TRUE BUGS OF THE WORLD (HEMIPTERA-. HETEROPTERA) Fig. 87.2. Pyrrhocoridae. A. Alimentary canal, Pyrrhocoris apterus (Linnaeus) (from Dufour, 1833). B. Aedeagus, sagittal view, Paradindy- mus madagascariensis (Blanchard) (from Stehlik, 1966), C. Paramere, Dysdercus blotei van Doesburg. D. Paramere, D. blotei (C. D from van Doesburg, 1968). E. Paramere, Paradindymus madagascariensis (from Sfehlik, 1966). F. Spermatheca, D. discolor Walker (from Pluot, 1970). G. Spermatheca, D. faSciatus (from Pendergrast, 1957). H. Gynatrial complex, showing spermatheca and ring sclerites, Paradindymus madagascariensis (from Stehlik, 1966). Abbreviation: gc, gastric caecum. Coreoidea 88 Alydidae mone as well as inheritance of wing polymorphism. Sev¬ eral species of Pyrrhocoridae have been used as labora¬ tory animals. An extensive physiological and biochemical literature exists for such species as D. koenigii (Fabri- cius). Distribution and faunistics. Members of the family are chiefly tropical and subtropical, with only a very few species reaching into the temperate Holarctic. They are found, however, in all major zoogeographic regions. Freeman (1947) should be consulted for identification of the Old World Dysdercus fauna, and van Doesburg (1968) for the New World; Stehlik (1965) erected sub¬ genera for this large genus. Cachan (1952c) revised the Malagasy fauna. Hussey’s (1929) catalog remains the basic literature source. There is no general identification work for the other taxa. General. Members of the group are usually elongate and slender with disproportionately large heads (Fig. 88.1). They range in length from 8 to 20 mm. Many species are myrmecomorphic, particularly as nymphs. These insects are sometimes referred to as broad-headed bugs. Diagnosis. Bucculae very short, not extending poste¬ riorly beyond antennal insertion; antennae inserted dor- sally, segment 1 not constricted at base; ocelli not placed on sclerotized elevations; corium elongated on costal mar¬ gin; membrane of forewing with numerous veins (Fig. Alydidae 271 A Rg. 88.1 . Alydidae (from Slater and Baranowski, 1978). A. Stenocoris tipuloides (De Geer). B. Megalotomus quinquespinosus (Say). 88.1 A, B); metathoracic scent-gland auricle well devel¬ oped; tibiae nonsulcate; abdominal trichobothria placed laterally or sublaterally on segments 5-7, submedially on sterna 3-4, either clustered or dispersed; spiracle present on abdominal segment 8, all spiracles ventral; nymphal dorsal abdominal scent-gland openings between terga 4/ 5 and 5/6; aedeagus as in Fig. 88.2C; pygophore and parameres as in Fig. 88.2B; spermatheca lacking proxi¬ mal flange (Fig. 88.2E>-F); alimentary canal as in Fig. 88.2A. Classification. Amyot and Serville (1843) first char¬ acterized the group as “Alydides.” Stal (1867), who treated the group as a subfamily "Alydida” of the Corei- dae, was followed by many authors because of the multi- veined membrane of the forewing and the flattened plate¬ like ovipositor. Schaefer (1965b) and subsequent authors have treated the group as a family within the Coreoidea and elevated the tribes to subfamily status, but have com¬ bined the Micrelytrinae and Leptocorisinae into a single subfamily with two tribes. We follow Ahmad (1965) in recognizing three taxa at the subfamily level, a position accepted by Schaefer (1972b). The family contains approximately 42 genera and 250 species. Key to Subfamilies of Alydidae 1. Maximum width of pronotum at least 1.5 times maximum width of head and distinctly longer than head (Fig. 88.1 A) .;. Leptocorisinae - Pronotum at most only slightly wider and longer than head . 2 2. Hind femora swollen, with a series of ventral spines (Fig. 88. IB) (except in Euthetus Dallas); labial segment 2 distinctly shorter than combined length of segments 3 and 4; posterior margin of abdominal sternum 7 in females without a median split . . Alydinae - Hind femora never with ventral spines; labial segment 2 usually distinctly longer than combined length of segments 3 and 4; posterior margin of abdominal sternum 7 in females usually with a median split . Micrelytrinae 272 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Fig. 88.2. Alydidae. A. Alimentary canal, Camptopus lateralis (Germar) (from Dufour, 1833). B. Male genitalia, including parameres, Leptocorisa acuta (Thunberg). C. Aedeagus, sagittal view, Acestra malayana (Dallas). D. Spermatheca, Daclera punctata (Signoret). E. Spermatheca, L. chinensis Dallas, F. Spermatheca, Micrelytra fossularum (Rossi) (B-F from Ahmad and Southwood, 1964). Abbreviations: gc, gastric caecum; pa, paramere. ALYDINAE (FIG. 88.1 B). Head transverse, broader than tho¬ rax (Fig. 88. IB); hind femora with ventral spines (except in Euthetus)', metathoracic processes of scent-gland peri- treme fused and evaporative area not lateral to opening, smooth; abdominal sternum 7 with well-developed an¬ terior spur; second inner laterotergite absent; subcosta present and distinct on forewing. Some of the better known genera are Alydus Fabricius, Hyalymenus Amyot and Serville, Camptopus Amyot and Serville, and Riptortus Stal. The subfamily is found in all major zoogeographic regions. It contains many myr- mecomorphic species and others that appear to be wasp mimics. LEPTOCORisiNAE (FIG. 88.1 A). Head elongate and rela¬ tively slender; labial segment 1 extending well beyond posterior margin of compound eyes, segment 2 usually distinctly shorter than segments 3 and 4 combined, seg¬ ment 4 usually subequal in length to segment 3; posterior angles of metasternum acutely produced; legs lacking spines, usually very long and slender; hamus of hind wing at most only slightly separated from base of wing; poste¬ rior margin of abdominal tergum 2 truncate; spermatheca usually balloon-shaped with a coiled tube. Ahmad (1965) recognized two tribes. The Leptoco- risini, with four genera and a worldwide distribution, contains Leptocorisa Berthold from the Orient and Aus¬ tralia and Stenocoris Burmeister from the southern Nearc- tic. Neotropical, and Ethiopian regions. The Noliphini Alydidae 273 include the genus Lyrnessus Stal from South America, whereas the remaining two genera occur in Australia and in the Orient as far west as Sumatra. MiCRELYTRiNAE. Antennal segments 2 and 3 sometimes triquetrous; ratio length of head to antennal segment 1 not less than 1:1; labial segment 3 very short, less than one-half length of segment 4. both together shorter than segment 2; posterior margins of metathorax more or less produced, often acutely so; pronotal collar lacking; meta- thoracic scent-gland evaporative area well developed, occupying nearly one-half of metapleuron; moderately slender or strongly myrmecomorphic. Micrelytra Laporte is widely distributed in the southern Palearctic, and Protenor Stal in the Nearctic. Specialized morphology. Several genera possess a forewing stridulitrum and a hind femoral plectrum and also long spines on the dorsal surface of the male genital capsule {Alydus Fabricius, Megalotomus Fieber, Burtinus Stal, Tollius Stal) (Schaefer et al., 1989). The short buc- culae, which do not extend beyond the antennal insertion, appear to be unique to the group. Natural history. Most species of Alydinae live on legumes, whereas the Leptocorisinae and Micrelytrinae feed primarily on grasses (Schaefer, 1980a; Schaefer and Mitchell, 1983). There are reports of feeding on carrion, but this is not a general food source. Nymphs of Alydinae are strikingly antlike. The early instars are almost indistinguishable from ants in the field, not only in body form but in their jerky movements as well. Adults with reduced wings are also often strikingly antlike, whereas macropterous forms are remarkably like pompilid wasps (Southwood and Leston, 1959). This likeness may not be evident in museum specimens, but in the field the display of the brightly colored abdominal dorsum and the wasplike twisting and turning are remark¬ ably convincing. Some micrelytrines such as Trachelium Herrich-Schaeffer and Cydamus Stal are also strikingly antlike. Other members of the family for which observations are available live either on plant stems or on the ground among stem litter and are cryptic both in body shape and color. Some of these are associated with monocots, as for example three species of the Neotropical genus Bactro- phyamixia Brailovsky living on bamboos (Guadua spp.) in Mexico (Brailovsky, 1991). Monteith (1982) reported several species of Heterop- tera, including Leptocorisa acuta, aggregated in ‘large clusters during the dry season in monsoon forests in northern Australia. Apparently this is a phenomenon for protection during nonfeeding periods. They were found about a meter above the ground beneath leaves or in rows along twigs of low shrubs and ferns. When disturbed, the clusters burst into buzzing brief flight and discharged their repellent scents before settling .again in the original site. This aestivation probably covers a period of up to six months. A similar phenomenon has been observed in New Guinea, where with the onset of the dry season mi¬ gration from open grassland to shaded sites is triggered. Gregarious aestivation takes place for up to two months (Sands, 1978). Brown (1965) noted that in the Near East Camptopus lateralis GermSiT migrates from wheat fields, where it feeds chiefly on weeds, to montane overwinter¬ ing areas many kilometers away. Distribution and faunistics. Thefamii'. is represented in all major zoogeographic areas. Stys and Riha (1977) indicated Oligocene-Miocene records of all three sub¬ families from sites in the Holarctic. The extinct Monstro- coreinae Popov are recognized from the Upper Jurassic of Kazakhstan. Schaffner (1964) provided the most comprehensive taxonomic study of the group. Ahmad (1965) mono¬ graphed the Leptocorisinae: Froeschner (1981) keyed the South American subfamilies and genera; Gross (1963) keyed the fauna of Micronesia; Linnavuori (1987) treated the West and.Central African fauna; and Putchkov (1962) treated the fauna of the Ukraine. 89 Coreidae General. Most coreids are relatively heavy-bodied in¬ sects usually robustly elongate or broadly elliptical (Fig. 89.1). The family includes some of the largest of living heteropterans, as well as other species that are delicate (Fig. 89.2) or slender. Many have bizarre dilations and expansions of the hind femora or tibiae and antennal seg¬ ment 3. Body length ranges from 7 to 45 mm. Members of the group are sometimes referred to as leaf-footed bugs. Diagnosis. Very diverse in size and shape; head usu¬ ally small relative to body size; antennae inserted above a line running through center of eye; membrane of fore¬ wing multiveined; femora and tibiae of hind legs fre¬ quently incrassate or dilated (Fig. 89.1); inner lateroter- gites usually present; abdominal spiracles all ventral; 3 trichobothria on abdominal segments 3-6, 2 on segment 7, those on segments 5-7 sublateral, clustered; abdomi¬ nal spiracles ventral (but see Agriopocorinae); nymphal dorsal abdominal scent-gland openings between terga 4/ 274 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) 5 and 5/6; aedeagus as in Fig. 89.3A, B; parameres as in Fig. 89.3C; ovipositor valvulae flattened, platelike (Fig. 89.3D); sternum 7 usually cleft for about half its length (Fig. 89.3D), very rarely cleft throughout, sometimes not cleft; spermatheca usually with proximal pump flange but no distal flange, duct usually short (Fig. 89.3E, F); egg usually with pseudoperculum, usually as a well-defined circular cap, although nonoperculate in Pseudophloeinae and Hydara. Classification. The family was established by Leach (1815) and included Rhopalidae and Alydidae as sub¬ families. These groups were retained as subfamilies for many years. The basic suprageneric classification was established by Stal (1867, 1870). Although there has been considerable recent work on the higher classification of this great family, the infra- familial relationships remain surprisingly obscure, and relationships within the taxon are obviously in need of a modern synthesis. Schaefer (1964, 1965) provided the most comprehensive treatment. The phylogenetic scheme of Ahmad (1970) recognized the Coreinae, Colpurinae, Phyllomorphinae, Hydarinae, Pseudophloeinae, and Pro- camptinae. Schaefer (1982) discussed the status of several of these subfamilies. Other authors have placed some of them at the tribal level. As with the Reduviidae, we have adopted a conservative course with regard to the sub¬ family classification, treating all other proposed higher -taxa at the tribal level. The family contains at least 250 genera and 1800 species. Key to Subfamilies of Coreidae Fig. 89.1. Coreidae. Thasus acutangulus (Stal) (Coreinae) (from Slater, 1982). 1. Spiracles near lateral margin of abdomen; those on segments 2 and 3 visible from above; wings rarely present . Agriopocorinae - All spiracles ventral and located away from lateral abdominal margins, not visible from above; usually macropterous, sometimes wings reduced but always at least micropterous ... 2 2. Hind tibia with distal end produced into a tooth or spine . Meropachydinae - Hind tibia lacking a distal spine, or if spine present then head large and only slightly shorter and narrower than thorax .. 3 3. Median sulcus present on head before eyes; tibiae sulcate on outer surface; hind wing usually lacking an antevannal spur of cubitus . '..... .•.. Coreinae - Head lacking a median sulcus in front of eyes; tibiae not sulcate on outer surface; antevannal spur of cubitus usually present on hind wing ... Pseudophloeinae AGRIOPOCORINAE. Body flattened and aradid-like or elongate and sticklike; antenniferous tubercles occupy¬ ing entire anterodorsal head surface; usually apterous, rarely macropterous; macropterous forms with membrane of forewing reticulately multicellular; metathoracic scent- gland auricles bilobate; tibiae indistinctly sulcate; ab¬ dominal segment 8 of males with spiracles; some abdomi¬ nal spiracles marginal, those on segments 2 and 3 visible from above; valvulae of ovipositor intermediate between laciniate and platelike. Two genera are known from Australia. Agriopocoris Miller (five species) are flattened and broad-bodied and Coreidae 275 Fig. 89.2. Coreidae. Pephricus paradoxus (Sparrman) (Coreinae) (used with permission of Kathleen Schmidt). 276 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) D f t f Fig. 89.3. Coreidae. A. Aedeagus, lateral view, Agathyrna ceramica Dolling. B. Aedeagus, sagittal view, Agathyrna ceramica (A. B from Dolling, 1987a). C. Paramere, Anasa alfaroi Brailovsky (from Brailovsky, 1985). D. Female terminal abdominal segments, Catorhintha siblica Brailovsky and Garcia (from Brailovsky, 1987). E. Spermatheca, Anasa guayaquila Brailovsky (from Brailovsky. 1985). F. Spermatheca, Anoplocnemis sp, (from Pendergrast, 1957). have been taken on Acacia. The slender parallel-sided Tylocryptus egenus Horvath is the only other genus. It feeds on Casuarina branches, on which it is very cryptic. COREINAE (FIGS. 89.1,89.2). Usually medium-sized to very large; interocellar distance greater than that from eye to ocellus; anterolateral opening of the metathoracic scent gland well developed, peritreme with well-developed pro¬ jections, size of evaporative area usually twice that of scent-gland auricle; corial margins straight or slightly sinuate; membranal veins of forewing arising from a transverse vein near, or touching, corial margin; abdomi¬ nal terga 1-2 and 3-7 fused in both sexes; genital capsule of male without lateral prolongations; articulation of first valvifer and valvulae usually membranous; gonangulum usually flat and folded. This subfamily contains the vast majority of coreid bugs. It is worldwide in distribution, but most species occur in the tropics. Many of the currently recognized tribes, some of which may be polyphyletic, are either Old or New World in distribution. Only the Coreini (38 genera) and Hydarini (six genera) occur worldwide. Tribes occurring only in the Eastern Hemisphere are Acanthocorini (= Physomeraria) (eight genera); Amor- bini (seven genera); Anhomoeini (one genus); Clores- mini (four genera); Colpurini (19 genera); Cyllarini (one genus); Daladerini (= Brachytini) (nine genera); Dasy- nini (16 genera); Gonocerini (seven genera); Homoeo- cerini (12 genera); Latimbini (two genera); Manocoreoini (one genus); Mecocnemini (one genus); Mictini (47 genera); Petascelidini (12 genera); Phyllomorphini (four genera); Prionotylini (three genera); Procamptini (one genus); Sinotagini (one genus). Tribes occurring only in the Western Hemisphere are Acanthocephalini (19 genera); Acanthocerini (16 genera); Anisoscelidini (nine genera); Barreratalpini (one genus); Chariesterini (four genera); Chelinideini (one genus); Discogastrini (seven genera); Leptoscelidini (nine gen¬ era); Nematopodini (14 genera); Spartocerini (five gen¬ era). There are six described genera of uncertain tribal affiliation, Of the above-mentioned tribes, the following six may prove to merit subfamily status. COLPURINI. This taxon was raised to subfamily status from a tribe of the Coreinae by Ahmad (1970). Nine¬ teen genera are recognized. The genus Brachylybas Stal, which is widespread in the Pacific, is of special interest because 40% of its species have reduced wings (including coleopteroid types), a situation that, while widespread in the Heteroptera, is uncommon in the Coreidae. HYDARINI. This taxon has until recently always been considered a tribe within the Coreinae. Ahmad (1970) raised it to subfamily status. Schaefer (1982) suggested a close relationship with the Alydidae and indicated that in his view it is a relatively primitive taxon. Eight genera are currently recognized. PHYLLOMORPHINI (FIG. 89.2). This group was raised to Coreidae 277 subfamily status from a tribe of the Coreinae by Ahmad (19701. The insects are remarkably cryptic, with wide, flattened, often spinose marginal plates giving them somewhat the appearance of large Tingidae. Four genera are recognized. PROCAMPTiNi. This tribe was established by Ahmad (1964) for Procamptus segrex Bergroth from the Philip¬ pines and considered by him to probably merit subfamily status. MEROPACHYDINAE. Head Small', thorax narrow; poste¬ rior tibiae toothed or spined; hind femora large, clavate; metathoracic scent-gland opening located deep between coxae, opening anteriorly, projections of peritreme fused, evaporative area lacking ridges; abdomen with inner lat- erotergites fused to connexivum; sterna 2-5 fused in both sexes. Kormilev (1954) recognized three tribes: Merocorini (two genera), Meropachydini (six genera), and Spatho- phorini (three genera). The Meropachydinae are chiefly Neotropical, with Merocoris Hahn extending northward into the central and northern United States. PSEUDOPHLOEiNAE. Small to moderate-sizcd coreids; antennae inserted at sides of head, antenniferous tubercles provided with porrect or deflexed processes at outer api¬ cal angles; metathoracic scent-gland peritreme with dor¬ sal ridge entire or shortly bilobed, not produced into a Y-shaped auricle; membrane of hemelytron with a com¬ pound vein near base almost parallel with apical margin of corium; posterior coxae separated from one another by about width of a coxa; femora moderately to strongly clavate, hind femora typically with 2 or more large sub- distal spines ventrally on outer side; tibiae terete, never sulcate; female paratergite 8 without functional spiracle; spermatheca with lunate bulb and without a prominent flange; egg not operculate or pseudoperculate, opening by a transverse eclosion rent. The Pseudophloeinae, which comprise 28 genera and 166 species, are predominately Old World in distribution but are absent from temperate Australia, Only Vilga Stal occurs in the Neotropics, and two genera are found in the Nearctic, both of which also occur in the Palearctic. Dolling (1986) noted the apparently long isolation of the Neotropical fauna and believed it possible that the sepa¬ ration occurred prior to the opening of the Atlantic Ocean before the end of the Cretaceous. Specialized morphoiogy. Many coreids have strongly expanded, often leaflike hind tibiae (Fig. 89.1). Antennal segments 2 and 3 are also frequently dilated and flattened . Many species are ornamented with spines and tubercles, this being particularly true of the humeral angles of the pronotum, which are frequently produced into acute pro¬ cesses (Fig. 89.2). The eggs frequently have a large number of micropyles, up to a maximum of about 60. Natural history. Coreids are all phytophagous, and the majority live on plants above the ground. Most appear to feed in the plant vascular system (Mitchell. 1980b). Some are of considerable economic importance (see Chapter 8). Brown (1965, 1966) noted that in the Near East Aiio- plocerus elevaius (Fieber) and Ceraleptus obtusus (Brulle) show a migration similar to that found in several destruc¬ tive scutellerids and pentatomids—that is, from wheat fields to montane areas many kilometers away. These coreids, however, appear to feed mainly on weeds in the fields rather than on the wheat itself. All species of Pseudophloeinae whose food habits are known feed on herbaceous legumes. Schaefer and Mitchell (1983) gave a useful summary of the food plants of the family. They concluded that many groups of coreid bugs show definite associations with par¬ ticular plant groups; others, by contrast, contain members that feed on unrelated plant taxa. Within a given genus, some species may be host-specific, whereas others feed on a variety of plants. Mitchell (1980b) studied species of Leptoglossus in detail. In this genus, species such as L. phyliopus (Linnaeus) and L. gomgra (Fabricius) are very general feeders, whereas L. fulvicornis (Westwood), L. ashmeadi Heidemann, and L. corculus (Say) are ex¬ tremely restricted in their host associations. Polyphagous species tend to feed on annuals and have a labium of inter¬ mediate length, whereas specialists have an elongated labium, Mitchell (1980b) discussed host selection, sur¬ vival, and parasitism and provided an extensive analysis of the literature on feeding by phytophagous Hemiptera. She demonstrated that in Leptoglossus the specialist feed¬ ers did not feed on related plants, indicating that the plant-insect coevolutionary theory so strongly advocated by Ehrlich and Raven (1965) is not applicable to many piercing-sucking insects. Despite those observations, Schaefer and O’Shea (1979) noted that three coreine tribes (Mictini, Acantho- cerini, and Nematopodini) feed chiefly on plants of the family Leguminosae. They raised the possibility that such feeding habits could be used in phylogenetic reconstruc¬ tion—that is, that this habit constitutes a synapomor- phy. Schaefer and Mitchell (1983), however, noted that legume feeding may be primitive in the Coreoidea, being found in the Alydinae, Pseudophloeinae, Hydarini, and Hyocephalidae. It has recently been shown that, some of the large species of coreids defend territories on flower heads and fight vigorously with other males who attempt to enter their territories. This phenomenon may be widespread and could account for the sexually dimorphic hind legs of many coreids, in which the male hind femora are often much larger than those of the females and are provided with wicked series of sharp spines (Mitchell, 1980a). 278 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) Although the majority of Coreidae are dull-colored, many display strikingly bright coloration, some of which is presumably aposematic. Other coloration is probably deflective because it includes metallic hues and occurs chiefly in species found in tropical forest habitats. Members of the genus Thasus Stal are among the largest of the terrestrial Heteroptera. Thasus acutangulus Stal (Fig. 89.1) has aposematically colored bright orange, yellow, and black nymphs that form feeding aggregations. Apparently the aggregation pheromone is perceived by receptors on the distal segment of each antenna; thus nymphs displaced from the aggregation are able to re¬ aggregate. If these feeding aggregations are disturbed, the members pulsate, spray jets of anal fluid into the air, and exude scent-gland secretions over the abdominal terga (Aldrich and Blum, 1978). Distribution and faunistics. Coreid bugs are world¬ wide in distribution but are most abundant in the tropics and subtropics, where they also attain their largest size and most bizarre appearance. There is no modern key to the major groups. The works of Stal (1867, 1870) are still important. For special groups see those of O’Shea (1980a; Acanthocerini), O’Shea (1980b; Nematopodini), O’Shea and Schaefer (1978; Mictini), Yonke (1972; Chariesterini), Kormilev (1954; Meropachydinae), Herring (1980; Chelinldeini), Dolling (1978, 1979; Clavigrallini), Breddin (1900; Colpurini), Osuna (1984; Anisoscelidini), Gross (1963; Micronesian fauna), Putchkov (1962; fauna of Ukraine), Allen (1969; Leptoglossus), Brailovsky (1985; Anasa Amyot and Ser- ville), and Brailovsky and Garcia (1987; Catorhintha Stal). 90 Hyocephalidae General. These insects are rather large (length up to 15 mm), reddish brown or black, moderately elongate and parallel-sided (Fig. 90.1). They are dorsally flattened and resemble some pseudophloeine coreids and certain lygaeids. Diagnosis. Head very elongate, strongly tapered (Figs. 90.1, 90.2B, C), surface tuberculate; ocelli very small, placed near posterolateral margins of eyes; clypeus elevated, compressed in middle; antenniferous tubercles placed below a line drawn through middle of eye; buc- Fig. 90.1. Hyocephalidae. Maevius indecorus Stal (drawn by S. Mon- teith;from CSIRO, 1991). culae large, elongate, contiguous anteriorly, extending posteriorly to anterior margin of eyes (Fig. 90.2B. C); mandibular plates short; gula with a labial groove; mem¬ brane of forewing with basal cells formed by cross veins connecting 4 primary longitudinal veins, and with sev¬ eral distal veins (Fig. 90.2A); vein connecting M and Cu on corium lacking; metathoracic scent-gland ostiole with bristlelike processes; tibiae sulcate; base of abdo¬ men ventrally with an ovoid pore-bearing organ (sieve plate) on each side (Fig. 90.2D), comparable to porous area on sternum of some rhyparochromine lygaeids; tri- chobothria of abdominal sterna 3 and 4 mesal, 5 and 6 lateral, clustered (Fig. 90.2D); all spiracles ventral (Fig. 90.2D); inner laterotergites present; nymphs with 2 pairs of dorsal abdominal scent-gland openings between terga 4/5 and 5/6; ovipositor laciniate; spermathecal bulb lack- Hyocephalidae 279 Fig. 90.2. Hyocephalidae. Hyocephalus aprugnus Bergroth. A. Forewing, B. Lateral view head. C. Ventral view head. D. Lateral view abdomen, showing trichobothria and pore-bearing organ. E. Spermatheca (A-E from Stys, 1964b). Maevius indecorus Stal. F. Genital capsule. G. Aedea- gus, sagittal view. H. Aedeagus, lateral view. i. Paramere. J. Paramere, obverse view. (F-J modified from Schaefer, 1981a.) Abbreviation: pbo, pore-bearing organ. ing distinct flanges (Fig. 90.2E); eggs elongate, with 3 micropylar processes; nymphs with a U-shaped ecdysial line removed from eyes. Classification. The taxon was first established by Ber¬ groth (1906) as a subfamily of Coreidae but later was reduced to tribal status (Bergroth, 1912). Reuter (1912a) raised the group to family rank. Stys (1964b) reviewed the. position of the family, noting that it had been vari¬ ously placed in the Coreoidea and Lygaeoidea, a factor that influenced him (Stys, 1961b) to combine the two superfamilies. Two genera and two species —Hyocephalus aprugnus Bergroth and Maevius indecorus Stal—both from Aus¬ tralia, are known. Schaefer (1981a) gave evidence for 280 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) placing the Hyocephalidae and Stenocephalidae in the Coreoidea. Specialized naorphology. The bristlelike processes projecting from the scent-gland orifice appear to be an apomorphy for the family. The ovoid pore-bearing plate on the abdomen has been referred to as a “strainer” (Stys, 1964b). Natural history. Hyocephalids live on the underside of stones in sandy gravelly areas, where they are very cryptic. They feed on the ripe seeds of Acacia and Euca¬ lyptus. Distribution and faunistics. The family occurs only in Australia. The main references on the group are by Stys (1964b) and Schaefer (1981a). 91 Rhopalidae General. The members of this family range in length from 4 to 15 mm. They vary greatly in shape and color. The majority are dull brownish and resemble species of Orsillinae (Lygaeidae), with which one often finds them confused in collections. The remainder are much larger and similar in shape, body form, and bright color¬ ation (Fig. 91.1) to species of Lygaeinae (Lygaeidae) and many species of Pyrrhocoridae and Largidae. They are frequently called the scentless plant bugs. Diagnosis. Clypeus surpassing mandibular plates; ocelli situated on low tubercles; antennae never dilated, first segment constricted basally; metathoracic scent- gland openings usually obsolete or obsolescent; corium frequently with large hyaline areas; membrane of fore¬ wing always with numerous veins; trichobothria on ab¬ dominal sterna 3 and 4 mediolateral, those of 5, 6, and 7 lateral; abdominal spiracles ventral; inner laterotergites present; nymphs with dorsal abdominal scent-gland open¬ ings between terga 4/5 and 5/6, the latter displaced for¬ ward, a unique and universally occurring character in the family; pygophore with lateral, median, and paralateral lobes (Fig. 91.2A); aedeagus as in Fig. 91.28; parameres as in Fig. 91.2C; ovipositor platelike, abdominal sternum 7 of females entire; spermatheca consisting of a round bulb, small pump, and long, generally coiled duct. Classification. The taxon was first recognized as a higher group by Amyot and Serville (1843) as the “Rho- palides.” This name was used subsequently by many Fig. 91.1. Rhopalidae. Boisea trMttata (Say) (from Slater and Bara- nowski, 1978). Fig. 91.2. Rhopalidae (from Chopra, 1967). A. Genital capsule, Myr- mus miriformis (Fallen). B. Aedeagus, sagittal view, Uorhyssus hyali- nus (Fabricius). C. Paramere, Ithamar hawaiiensis Kirkaldy. Rhopalidae 281 authors including Dallas (1851-1852) and Stal (1862). Costa (1853) used Corizini as a tribe, and Douglas and Scott (1865) used Corizidae as a family. Mayr (1866) used “Corizida,” and much of the literature is under this name. The family has often been considered to be a subfamily of an inclusive Coreidae, but modern workers such as Chopra (1967) and Gollner-Scheiding (1983) treated it as Key to Subfamilies of Rhopalidae a distinct family. The last-mentioned work, a world cata¬ log, serves as a summary of the classification of the group and an exhaustive introduction to the literature. Schaefer and Chopra (1982) provided a cladistic analysis to the tribal level. Two subfamilies, comprising 18 genera and 209 species are recognized. 1. Lateral margins of pronotum straight or slightly sinuate, lacking a distinct notch immediately behind collar . Rhopalinae - Lateral margins of pronotum with a distinct notch behind collar . Serinethinae RHOPALINAE. Generally of small to moderate size; lat¬ eral pronottil margin lacking a distinct notch immediately behind collar; usually dull-colored; abdominal sterna 3 and 4 fused in both sexes; male and female genitalia also distinctive (see Chopra, 1967). Six tribes are generally recognized: Chorosomini, Corizomorphini, Harmostini, Maccevethini, Niesthreini, and Rhopalini. The European Corizus hyoscyami (Lin¬ naeus), a presumed mimic of Lygaeus equestris (Lin¬ naeus), is unusual in the subfamily, along with a few other species, for its bright coloration. SERINETHINAE (FIG. 91.1). Relatively large, elongate, elliptical; usually brightly colored; head broader than long; clypeus slightly raised; antenniferous tubercles with¬ out lateral projections; pronotum trapezoidal, lateral mar¬ gins notched just behind collar; metapleuron not, or only indistinctly, divided; a well-developed metathoracic third axillary sclerite spur; metathoracic scent-gland openings in coxal cavities; dorsal abdominal scent glands retained and functional in adults (Ribeiro, 1989); hind femora not incrassate or spined; abdominal sterna 3 and 4 not fused; phallus with sclerotized ventral conjunctival appendages. Three genera are currently recognized. Leptocoris Hahn, with 30 species from the Old World tropics, is par¬ ticularly speciose in Africa. Jadera Stal, with 17 species, is chiefly tropical and subtropical in the New World, with several species reaching into the southern United States. Boisea Kirkaldy, with two Nearctic, one central and west African, and one South Indian species, was until recently considered to be a junior synonym of Leptocoris. It was elevated to generic status by Gollner-Scheiding (1980) for the Western Hemisphere species previously placed in Leptocoris, a decision questioned by Schaefer and Cho¬ pra (1982). It includes the common box elder bug, Boisea trivittata (Say). Specialized morphology. The functional abdominal scent glands in the adults are a specialized feature in the Serinethinae. They may represent a neotenic condition (see Chapter 85 for a similar situation). Natural history. All species are phytophagous and feed on a variety of herbs and woody plants; host use was reviewed by Schaefer and Chopra (1982). None are of major economic importance, although Boisea trivittata often becomes a nuisance when it forms aggregations in the fall and enters houses in large numbers to hibernate. It is an arboreal species almost monophagous on box elder trees (Acer negundo). The soapberry bug, Jadera haematoloma Herrich- Schaeffer, feeds on three native species of Sapindaceae: Sapindus saponaria var. drummondii (the soapberry tree) in the south-central United States, Serjania brachycarpa (serjania vine) in southernmost Texas, and the peren¬ nial Cardiospermum corindum (balloon vine) in south¬ ern Florida (Carroll and Boyd, 1992). The insect has colonized three introduced species of Sapindaceae; Koel- reuteria paniculata (golden rain tree), K. elegans (flat- podded golden rain tree), and Cardiospermum halicaca- bum (heartseed vine) (Carroll and Loye, 1987). Carroll and Boyd (1992) discussed the establishment of recogniz¬ able populations upon these various hosts, distinguishable primarily by large differences in the length of the rostrum. Large fruits with deeply embedded seeds have popula¬ tions with very long rostra, whereas small fruits have populations with significantly shorter rostra. These au¬ thors noted that much of this differentiation must have occurred within the past 30 to 50 years because the intro¬ duced host plants were planted commonly only within that time period. This rapid establishment of host-plant races directly attributable to a selective trait has great evolutionary significance. Carroll (1988, 1991) reported that Jadera haematoloma has developed two different mating systems in different populations. Where females are abundant, promiscuous mating occurs, but where males are much more abun- 282 TRUE BUGS OF THE WORLD (HEMIPTERA; HETEROPTERA) tlant than females a guarding system has developed due to the increased cost of male searching in such popula¬ tions. Nymphs are reddish and aposematic and tend to congregate on the trunks and seeds, whereas the adults are cryptically colored and disperse widely when on tree hosts. The nymphs that form aggregations, especially during the time of molting, have lower mortality in early instars than do isolated nymphs, and they tend to molt earlier and with greater synchrony. Older nymphs do not show this advantage, possibly because of increased can¬ nibalism. Kxperiments on toads and blue jays show that all nytnphs are distasteful and indeed appear also to be distasteful to praying mantids. Interestingly, however, the adults also are distasteful. So the aposematic nymphal coloration appears to be associated with gregarious be¬ havior. serving especially as protection during periods of molting and being most significant in early-instar nymphs (Ribeiro. 1989). A similar phenomenon appears to obtain for the related box elder bug (see Chapter 8). Distribution and faunistics. Rhopalids are found in all major faunal regions. Some of their distributions are unusual. The Niesthreini contains three genera; two are ctinfincd to the Western Hemisphere, and the third is found in South Africa and India. The Chorosomini con¬ tains six genera; three are Palearctic (one also occurs in the Old World tropics), Xenogenus Berg is Neotropical, and Itiuinwr Kirkaldy is endemic to the Hawaiian Islands. Boisca contains two Nearctic species, one species from west and central Africa and one from southern India. Additional cladistic analyses should improve our under¬ standing of origins and relationships in rhopalid higher taxa. Chopra (1967) discussed the higher classification and provided valuable generic revisions (1968, 1973). Putch- kov (1986) monographed the fauna of the former Soviet Union. Gross (1960) revised the Australian and Pacific Leptocoris. and Gross (1963) provided keys to the Micro- nesian genera and species. Gollner-Scheiding (1979) re¬ vised the genus Jadera. and Gollner-Scheiding (1980) treated the African species of Lepwcoris and Boisea. The Gollner-Scheiding (1983) world catalog is a basic resource. 92 Stenocephalidae General. These bugs are relatively slender, elongate, parallel-sided, brown and yellow, and have a distinct coreidlike appearance (Fig. 92.1 A). They range in length from 8 to 15 mm. They have no common name. Diagnosis. Mandibular plates surpassing and usually contiguous anterior to apex of clypeus (Fig. 92.1 A); membrane of forewing opaque with a large and a small basal cell from which numerous radiating and anastamos- ing veins arise (Fig. 92.1 A); abdominal spiracles ventral; inner laterotergites present; trichobothria of abdominal segments 5 and 6 plaeed laterally and clustered posterior to spiracle (Fig. 92. IH); nymphs with dorsal abdominal scent-gland openings between terga 4/5 and 5/6; genital eapsule and abdominal organs as in Fig. 92. IB (Lans- bury, 1965); aedeagus as in Fig. 92.1C; parameres as in Fig. 92.ID, E; ovipositor laciniate, completely dividing sternum 7 (Fig. 92.IF); spermatheca as in Fig. 92.IG; m-chromosome present; egg oblong with 4-9 micropylar processes. Classification. This family is of special interest in that it shows characteristics that are transitional between the Coreidae and the Lygaeidae. Its relationships have been discussed in detail by Scudder (1957a) and Schaefer (1965a, b). The above characteristics as well as such specialized features as the four-lobed principal salivary gland, the structure of the phallus, and the coreidlike appearance of the nymphs all suggest close relationships to the Corei¬ dae. The ovipositor, however, is distinctly laciniate (Fig. 92. IF); the spermatheca has a spherical apical bulb and a differentiated pump and duct (Fig. 92. IG). and the egg is oblong with 4-9 micropylar processes grouped at the anterior pole. These features are definitely of the lygaeid type and represent conditions not generally found in the Coreoidea. Two genera, Dicranocephalus Hahn and Psotilnus Stal, and over 30 species are known (Lansbury, 1965-1966). The majority are from the tropics and subtropics of the Eastern Hemisphere, including Australia, but some species occur in the temperate Palearctic. Specialized morphoiogy. The unusual combination of features possessed by the group is discussed above under Classification. Natural history. Stenocephalids are frequently swept from various species of Euphorbiaceae, upon which they breed. The eggs are laid on the surface of the stems rather Stenocephalidae 283 Fig. 92.1. Stenocephalidae. A. Dicranocephalus setulosus (Ferrari) (from Lansbury, 1965-1966). B, Genilal capsule, showing abdominai organs, D. pallidus (Signoret) (from Lansbury, 1965, with permission from Nature, copyright 1965 Macmillan Magazines Limited). C. Aedeagus, sagittal view, D. albipes (Fabricius). D. Paramere, D. agilis (Scopoli). E. Paramere, D, albipes. F. Female terminal abdominal segments, D. agilis. G. Spermatheca, D. agilis. H. Ventral view, abdomen with trichobothria, D. albipes (C-H from Lansbury, 1965-1966). than inserted in the plant tissues, despite the fact that the ovipositor is laciniate. Distribution and faunistics. An endemic species has been reported in the Galapagos Islands and was treated as distinct in Lansbury’s (1965-1966) revisional study, the major work on the group. Usinger and Ashlock, the first heteropterists to do careful collecting in the islands, were not able to obtain this species, suggesting that this species is thus very scarce or has become extinct. Although origi¬ nally described from the Galapagos, it is now thought to be conspecific with an African species. It was certainly introduced, probably by sailing ships, not later than the early nineteenth century. 284 TRUE BUGS OF THE WORLD (HEMIPTERA: HETEROPTERA) LITERATURE CITED Abreu, J. M, de. 1977. Mirideos neotropicais associados ao cacaueiro. In: E. M. Lavabre (ed.), Les mirides du cacaoyer, pp. 85-106. Institut Fran^ais du Cafe et du Cacao, Paris. Afzelius, B. A., R. Dallai, and P. Lindskog. 1985. Spermatozoa of saldid bugs (Insecta, Hemiptera, Leptopodomorpha). J. Ultra- struct. Res. 90:304—312. Ahmad, I. 1964. 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Literature Cited 315 GLOSSARY accessory fecundation canal a slender, usually weakly sclero- tized, tube running along dorsal wall of common oviduct from gynatrial sac to point of entrance into common oviduct; divided into basal thickening and fecundation pump, accessory gland vermiform gland (q.v.) or accessory salivary gland (q.v.). accessory male genitalia in some Schizopteridae, complex dorsoabdominal pregenital structures assumed to perform actual insemination after receiving sperm from primary genitalia, accessory parempodium in Leptopodomorpha, e.g., small, sec¬ ondary parempodium; see also pseudopulvillus. accessory salivary gland tubular or vesicular gland associated by a duct with principal salivary gland, accessory scent glands small glands associated with primary reservoir of metathoracic scent gland, aedeagus that portion of phallus distal to phallobase, including proximal phallotheca (phallosoma) and distal endosoma. aeropyle fine pores connected to air spaces in outer and inner meshworks of chorion. airstraps in Belostomatidae, a pair of straplike appendages de¬ rived from abdominal segment 8 used like respiratory siphon in Nepidae to obtain atmospheric air. anal segment proctiger, q.v. anal tube proctiger, q.v. androconia in males of Scutelleridae and some other Pentato- momorpha, unicellular glands grouped in patches on abdominal venter, with hollow bristlelike androconium set in an alveolus, androtraumatic insemination in Phallopirates (Heteroptera: Enicocephalidae), presumed mode of insemination, in which male can pass sperm to female only after breaking off tip of his own copulatory organ. annulus (pi., annuli) a ring encircling an article or segment, as in antennae of some Reduviidae; proctiger, q.v. anteapicai claw see preapical claw. anteclypeus inferior (anterior) portion of clypeus, wheneverthere is a visible transverse line of demarcation. Adapted, with permission of the New York Entomological Society, from S. W. Nichols, The Torre-Buem Glossary of Entomology (New York: New York Entomological Society, 1989), xvii -l- 840 pp. antennal fossa a groove in which antenna is located or concealed, as in some Phymatinae (Reduviidae). antenniferous tubercle a protuberance of head which bears an¬ tenna. anterior gonapophyses first vaivulae, q.v. anterior pronotal lobe anterior portion of pronotum. bearing calli. anterior ramus first ramus, q.v. anterior vaivulae first vaivulae, q.v. apical bulb spermathecal bulb, q.v. apterous completely lacking wings. arolium (pi.. arolia) one or two unpaired bristle- or bladderlike, medial pretarsal structures, originating dorsad of unguitractor and between but isolated from bases of claws (sometimes incor¬ rectly called empodium, parempodium, or rarely pulvillus); see empodium, pulvillus. articulatory apparatus system of plates and apodemes for sus¬ pension of phallus and attachment of its motor muscles, drawn out along phallosoma into ligamentary processes, comprising (1) basal plates attached to suspensory apodemes and ponticu- lus transversalis and (2) dorsal connectives, ending in capitate processes. auricle(s) variously shaped structure on metapleuron of adult Cimicomorpha and Pentatomomorpha, assisting in spreading metathoracic scent-gland products from ostiole onto evaporato- rium. auxilia basipulvillus, q.v. basal apparatus articulatory apparatus, q.v. basal foramen entrance to phallic cavity surrounded by basal plates and ponticulus transversalis, closed or not by a septum, basal plates two major plates of articulatory apparatus; some¬ times applied to entire articulatory apparatus, basiconjunctiva distal membranous part of phallosoma reaching to. but not including, ejaculatory reservoir, basipulvillus basal portion of pulvillus in Pentatomomorpha; see distipulvillus. Berlese’s organ mesospermalege (q.v.) or spermalege (q.v.), as a whole. bothrium (pi., bothria) pit or tubercle from which a trichoboth- rium arises, brachypterous with shortened or abbreviated wings, usually in¬ capable of flight, Brindley’s glands in some adult Reduviidae and Pachynomi- dae. paired glands located in anteriortnost portion of abdomen, with openings situated dbrsolateraliy just posterior to thoracico- abdominal junction. buccula (pi., bucculae) a flange of gena, on each side of basal portion of labium. bug a term often loosely used for any insects, but strictly applied to members of suborder Heteroptera; true bug, q.v. bursa copulatrix (pi., bursae copulatrices) variously formed structure serving as a vagina, as in Miridae. callar area middle part of pronotum behind collar and containing calli, and corresponding in size to prothoracic body cavity, callus (pi., calli) paired or fused impression or elevation in an¬ terior part of pronotum behind collar, capitate processes mushroomlike or bladelike ends of dorsal connectives (apodemes) of basal plates, on which are inserted protractor muscles of phallus, capsid member of family Miridae. capsula seminalis (pi., capsulae seminales) spermathecal bulb, q.v, cardinate coxae hinged and elongate hind coxae in pagiopodous taxa. Glossary 317 cephalic glands maxillary glands, q.v. cephalic neck constricted posterior part of head, for most part inserted into prothorax. Chagas’ disease disease of humans and other mammals in South and Central America. Mexico, and Texas, caused by flagellate protozoan Trypanosoma cruzi (Trypanosomatidael and transmit¬ ted by assassin bugs (Reduviidae). especially Triaioma and Rhotl- nius claspers parameres, q.v. claval commissure junction of hemelytra along clavus on midline of body posterior to apex of scutellum and anterior to membrane, developed in most Panheteroptera. clavai iurrow claval suture, q.v. claval suture suture of forewing separating clavus from corium. clavopruina in Corixidae, a narrow, white frosted area along anterolateral margin of clavus. clavus (pi., clavi) usually parallel-sided and sharply pointed anal area of hemelytron. claw hairs setiform microtrichia on outer surface of a claw, e.g.. in some Miridae. claw plate unguitractor plate, q.v. clypeus that part of head below frons. to which labrum is attached anteriorly; usually weakly to strongly protruding, coiled duct in Gerromorpha, accessory fecundation canal, q.v. coleopteroid beetlelike in form, often referring to structure of forewings. collar rounded or flattened anterior margin of prothorax, collum (pi., collal collar, q.v. common oviduct proximal portion of female genital ducts, be¬ tween lateral oviducts (whether ectodermal or endodermal) and vagina. conceptaculum seminis (pi., conceptacula seminis) in many Cimicoidea. mesodermal organs of sperm storage, being a dif¬ ferentiation of mesodermal oviducts, copjunctiva (pi., conjunctivae) intersegmental membrane of ab¬ domen; proximal portion of endosoma of phallus in many Het- eroptera. conjunctival appendages lobes or processes arising from con¬ junctiva in expanded aedeagus of many Heteroptera. connexivum lateral margin of abdomen, formed by dorsal and ventral laterotergites or laterostemites. copulatory organ phallus, q.v. corial glands in Plokiophilidae. numerous, large, unicellular glands with low conical openings on dorsal surface of corium. coriopruina in Corixidae. a white frosted area between anterior apex of corium and clavopruina, q.v. corium (pL. coria) in Panheteroptera. proximal coriaceous or otherwise differentiated part of forewing exclusive of clavus and distinct from membrane, often being subdivided into anterior (lateral) exocorium and posterior (mesal) endocorium; see cuneus and embolium. costal fracture in many Heteroptera. a short, usually transverse line of weakness or break in costal margin of forewing separating sometimes well-differentiated cuneus from rest of corium. cotton Stainer species of Heteroptera that cause discoloration of cotton fibers by piercing unripe bolls for their sap, e.g., Dysder- cus spp. (Pyrrhocoridae). cricoid sclerite in many Pentatomomorpha, a parietal differentia¬ tion of endosoma delimiting conjunctiva from vesica. Cryptocerata Nepomorpha ctenidium (pi., ctenidia) in Polyctenidae, comblike rows of flat¬ tened spines. cubital furrow simple or forked furrow or plica on hind wing posterior to Cu. cultrate shaped like a pruning knife, cuneal incisure costal fracture, q.v. cuneus (pi., cunei) in some Heteroptera. usually triangular post¬ erolateral area of corium demarcated by costal fracture, diadenian type omphalian or diastomian type of metathoracic scent gland, with gland ceils concentrated within paired glandular components of system and with scent reservoir(s) differentiated; see diastomian type, omphalian type, diastomian type in adult Heteroptera, metathoracic scent-gland apparatus opening by paired, widely spaced orifices associated with metacoxal cavities on metapleuron; see also omphalian type, distipulvillus distal membranous or setiform portion of pulvillus in Pentatomomorpha; see basipulvillus. dorsal abdominal scent gland in nymphal Heteroptera, 1-4 paired or unpaired ectodermal abdominal glands with paired or unpaired orifices situated intersegmentally or intrasegmentally, structure and function sometimes persisting into adult stage, dorsal arolium arolium, q.v. dorsal laterotergite(s) lateral plate of an abdominal tergum, often subdivided into outer and inner laterotergites. dorsal paratergite dorsal laterotergite, q.v. ductus ejaculatorius (pi., duct! ejaculatorii) median ecto¬ dermal efferent duct proximal to phallus, merging into ductus seminis. ductus receptaculi spermatheca, q.v. ductus seminis median ectodermal duct in phallus, from fora¬ men ductus to secondary gonopore, frequently differentiated into ductus seminis proximalis and ductus seminis distalis, ductus spermathecae canal through which sperm enter sperma¬ theca from vagina or bursa copulatrix, q.v. ectospermalege (pi. ectospermalegia) in some Cimicoidea, ex¬ ternal pouchlike ectodermal part of spermalege, q.v. egg cap a lid, joined to body of egg along a line of weakness, that is forced off by hatching embryo. ejaculatory reservoir in Pentatomomorpha, Dipsocoromorpha, and Nepidae, complex differentiation of proximal end of ductus seminis in endosoma. embolar groove trough-shaped groove of forewing running par¬ allel with costal margin anterior to medial fracture and often delimiting embolium; see medial fracture, embolium in forewing of some Heteroptera. broadened submar¬ ginal part of corium proximal to costal fracture; exocorium, q.v. empodium (pi.. empodia) distal extension of unguitractor plate, or. often applied to any unpaired structure arising between claws; see arolium, parempodia, pulvillus. endocorium posterior (mesal in repose) part of corium between exocorium and clavus. endosoma (pi., endosomata) distal segment of phallus, free of ligamentary processes and surrounding ductus seminis distalis from ejaculatory reservoir (when present) to secondary gono¬ pore. epipharyngeal sense organ anterior (x organ) and posterior (y organ) groups of sensilla located in epipharynx, apparently with sensory function related to feeding. Euheteroptera that taxon including the Dipsocoromorpha, Gerromorpha, Nepomorpha, Leptopodomorpha, Cimicomor- pha, and Pentatomomorpha. evaporatorium evaporatory area, q.v. evaporatory area in most Cimicomorpha and Pentatomomorpha, area of specialized cuticle on metathoracic pleuron associated with, and usually surrounding, orifice and auricle of metathora¬ cic scent glands, possibly functioning in controlled dissemination and evaporation of scent-gland products. 318 Glossary eversible gland in adult Saldidae, a gland located in interseg- mental membrane of dorsal abdominal laterotergites 7 and 8. exocorium that part of corium lying between R or R+M and costal margin; see cuneus. emboiium. expansion skating in Gerroidea. dispersing of fluid (probably saliva) onto water surface, which lowers surface tension causing insect to move much more rapidly than is otherwise possible, extragenital insemination traumatic insemination, q.v. false spiracles in Nepidae, hydrostatic organs, q.v. fecundation canal accessory fecundation canal, q.v. fecundation pump in some Gerromorpha, a widened area of accessory fecundation canal provided with a pair of platelike flanges. first gonocoxae first valvifer; see valvifers. first gonocoxopodites first valvifer; see valvifers. first ramus connecting leaf of first valvulae. first valvulae the outer blades of the ovipositor, flagellum (pi., flagella) gonoporal process, q.v.; the 2 terminal segments of the antennae, forceps ( pi ., forcipes) parameres, q. v. fossula spongiosa in many Cimicomorpha, apically on one or more pairs of tibiae, a vesicular hemblymph-filled structure beset with adhesive setae. frenum (pi., frena) lateral groove in upper margin of scutellum into which fits or catches channeled locking device on lower edge of clavus. genital atrium vagina, q.v. i. , genital capsule pygophore, q.v. genital chamber vagina, q.v. genital segment(s) in males abdominal segment 9, in females abdominal segments 8 and 9; pygophore, q.v. glochis spur or short vein in hind wing arising distally from Cu. gonangulum in female, sclerite uniting first valvifer and lateroter- gite nine; attached internally to first valvula or at base of first ramus; obscured in external view by first valvifer. gonapophyses (sing., gonapophysis) valvulae, q.v. gonapophysis eight first valvula; see first valvulae. gonapophysis nine second valvula; see second valvulae. gonocoxite eight first valvifer; see valvifers. gonocoxite nine second valvifer; see valvifers. gonocoxites valvifers, q.v. gonocoxopodites valifers, q.v. gonoforcipes (sing., gonoforceps) parameres, q.v. gonopods parameres, q.v. gonoporal process elongate, sometimes coiled, distal portion of ductus seminis. gonopore secondary gonopore, q.v. gonostyli (sing., gonostylus) parameres, q.v, guide in some male Enicocephalidae. reduced remnants of exter¬ nal genitalia arising from posteroventral margin of pygophore. gustatory organ epipharyngeal sense organ, q.v. Gymnocerata a grade group of Heteroptera with freely movable, conspicuous antennae. gynatrial complex in Gerromorpha, a term referring to that por¬ tion of internal ectodermalia composed of gynatrial sac, sperma- thecal tube, and accessory fecundation canal; also applied to complex of homologous structures in other Heteroptera. gynatrial glands ringed glands, q.v. gynatrial sac vaginal pouch (q.v.) or bursa copulatrix (q.v.) gynatrium vagina, q.v. hamus (pi., hami) spur or short vein, sometimes pointed, pro¬ jecting into middle cell of hind wing and representing M. harpagones (sing,, harpago) parameres, q.v. helicoid process in some Pentatomomorpha, cricoid sclerite, q.v. hemelytron (pi., hemelytra) forewing of Heteroptera. especially in Panheteroptera. with distinctly thickened proximal portion and membranous distal portion. hemocoelic insemination traumatic insemination, q.v. hemoglobin respiratory pigment found in hemolymph of Aniso- pinae (Notonectidae). hood in Tinginae (Tingidae). elevated anterior part of prothorax, often covering head. humeral angle posterolateral angle of pronotum. humerus (pi. humeri) humeral angle, q.v. hydranapheuxis in Gerromorpha and Leptopodomorpha. pro¬ cess of deforming meniscus of water surface to allow ascension to adjacent substrate. hydrostatic organs in Nepidae. 3 pairs of ovoid structures on connexiva of abdominal sterna 3-5 near spiracles but not con¬ nected to tracheal system, which function in spatial orientation, hygropetric pertaining to life on a thin film of water on a rock surface, as in some Gerromorpha. hypandrium (pi., hypandria) process on ventroposterior margin of pygophoral rim (abdominal segment 9). hypocostal lamina ventrally deflected proximal part of costal margin of forewing. hypocostal ridge hypocostal lamina, q.v. hypocular suture in Corixidae. short sulcus on either side of head capsule posteroventral to eyes, hypopygium (pi., hypopygia) pygophore, q.v. intercalary sclerites in Gerromorpha, Nepomorpha, and a few other groups of Heteroptera. 2 minute sclerotized plates dorsally between segments 3 and 4 of labium, juga (sing., jugum) mandibular plates, q.v. lacerate-flush feeding in phytophagous Heteroptera, process of lacerating and macerating cells with stylets and then flushing out material with saliva and imbibing it; see sawing-clipping feeding, laciniate ovipositor ovipositor with elongate, often laterally com¬ pressed blades (valvulae). lamina an expanded or platelike region, as body margins of Ter- mitaphididae. larval organ in many nymphal Saldidae, an apparently sensory structure, in form of depression, located on abdominal sternum 3 just mesad of spiracle. lateral oviducts paired canals leading from ovaries to common oviduct, most frequently mesodermal, but in certain Heteroptera proximally mesodermal and distally ectodermal, laterosternites lateral subdivisions of sterna of pregenital ab¬ dominal segments (e.g,, in some aquatic bugs), laterotergites dorsal and ventral laterotergites, q.v. lima (pi., limae) stridulitrum, q.v. lorum (pi., lora) maxillary plate, q.v. m-chromosomes supernumerary autosomes, occurring most commonly in Lygaeoidea and in some Nepomorpha. macropterous with both fore- and hind wings fully developed and functional. macrotrichia a relatively large or elongate seta, as on abdomen of some Anthocoridae. male hooks parameres, q. v. mandibular plate that portion of head laterad of (posterior to)' clypeus and dorsad of maxillary plate, mating swarm a conspicuous cloud of insects, usually males, dancing or hovering over a marker or in lee of an obstruction, serving to attract solitary members of other sex, e.g., Enico- cephalomorpha. maxillary glands small paired glands opening near bases of max¬ illae. maxillary plate that portion of head ventral to mandibular plate. Glossary 319 medial fracture longitudinal furrow delimiting exocorium (or embolium) from endocorium; see embolar groove, mediotergite unpaired plate of an abdominal tergum with delim¬ ited, paired laterotergites. membrane membranous apical portion of hemelytron in Pan- heteroptera and some other Heteroptera, mesoscutellum scutellum, q.v. mesospermalege (pi., mesospermalegia) in some Cimicoidea, subintegumental mesodermal portion of spermalege into which spermatozoa are injected; see ectospermalege. metathoracic scent gland in Heteroptera, universally occurring adult system of paired or unpaired scent glands with single or paired opening on metastemum with external outflow chan¬ nels (ostiolar canals) that transmit glandular products to ostiolc located on metepistema. micropyle opening in chorion of egg through which sperm pass during process of fertilization; in most Pentatomomorpha mani¬ fested externally by elevated tubular or capitate processes, microtrichia in Gerromorpha, that portion of body hair layer composed of fine spicules, mutic without spines. natatorial, natatory fitted for swimming, being generally ap¬ plied to swimming legs in aquatic bugs (Nepomorpha). neck cephalic neck, q.v. Neoheteroptera that taxon including the Gerromorpha, Nepo¬ morpha, Leptopodomorpha, Cimicomorpha, and Pentatomomor¬ pha. nodal furrow in Corixidae, costal fracture, q.v. node costal fracture, q.v. nymph immature form; larva. ocular setae in many groups of Heteroptera, usually a pair of bristles located in disc of compound eye of early instars, often lost later in development. odoriferous giand metathoracic scent gland, (q.v.) or dorsal ab¬ dominal scent gland (q.v.). omphalian type in adult Heteroptera. metathoracic scent-gland apparatus usually with a single (rarely double) opening on meta¬ stemum (rarely on abdominal sternum 1), with paired or unpaired internal structures; see diastomian type, omphalium prominent metasternal opening of omphalian type (q.v.) of metathoracic scent gland, operculum (pi., opercula) egg cap (q. v.). as for example inmost Cimicomorpha. organ of Berlese spermalege, q.v. ostiolar canal external outflow pathway of metathoracic scent gland, usually leading from metathoracic venter to metepister- num. ostiolar groove ostiolar canal, q.v. ostiolar peritreme in many Cimicomorpha and Pentatomomor¬ pha, a calloused area of variable shape, surrounding the ostiole, and itself often surrounded by the evaporatorium, ostiole external opening of metathoracic scent gland, often refer¬ ring to opening on metepistemum. ovipositor organ by which eggs are deposited, formed in Het¬ eroptera by paired first and second valvulae. Pagiopoda that unnatural assemblage of Heteroptera in which posterior coxae are usually elongate and articulation is a hinge joint; see cardinate coxae, Trochalopoda, pagiopodous pertaining to Pagiopoda. pala (pi., palae) in Corixidae, tarsus of foreleg modified into a seta-fringed scoop for particle feeding, and in males for attach¬ ment to females during mating or sexual display, palm, palma in Corixidae, that portion of pala, usually pilose, lying between upper and lower row of palmar setae, sometimes furnished with stridulatory pegs. palmar hairs in Corixidae, usually a row of long setae on lower margin of pala and a row of short setae along upper margin of palm. Panheteroptera that taxon including Nepomorpha. Leptopodo¬ morpha, Cimicomorpha, and Pentatomomorpha. paraclypeal lobe mandibular plate, q.v. paragenital sinus in Cimicoidea. external pocket or channel leading to external aperture of ectospermalege, paragenital system in many Cimicoidea. various structural dif¬ ferentiations in females correlated with traumatic insemination; see spermalege. parameres paired male genital structures independent of phallus, arising postembryologically from lateral parts of 2 buds (primary phallic lobes), median parts of which give rise to phallus, parandrium (pi., parandria) one of a pair of expansions of ex¬ ternal wall of pygophore in lateroventral position, provided with setae but not muscles. paranota (sing., paranotum) in certain Tingidae. flattened or lamellate sides of pronotum. parasternites laterosternites, q.v. paratergites laterotergites, q.v. parempodia (sing., parempodium) paired setiform or lamellate processes arising distally from unguitractor plate, between claw- bases; see arolium. empodium, pulvillus. pars intermedialis spermathecal pump. q.v. pedicel the second antennal segment, sometimes subdivided, as in many Pentatomoidea, peg plates in Gerromorpha and Ochteridae, minute circular de¬ pressions bordered by a shallow rim and filled with subconical pegs, generally found on head and body, and in some species also on certain leg segments. Pendergrast’s organ specialized organ found on abdominal ven¬ ter of some female Acanthosomatidae. penis (pi., penes) phallus, q. v. penisfllum (pi., penisfila) in Saldidae, reel system, q.v. periadenian type omphalian or diastomian type of metathoracic scent glands, with gland cells uniformly distributed in paired or unpaired components of system without differentiation of scent reservoir; see diastomian type, omphalian type, peritreme ostiolar peritreme, q.v. phallandrium in Phallopnates (Enicocephalidae), conspicuous bulbous copulatory organ composed largely of novel components including genital plates, phallobase articulatory apparatus, q.v. phallosoma proximal portion of phallus supported by or incor¬ porating ligamentary process and surrounding ductus seminis proximalis to ejaculatory reservoir (if present), often referred to as phallotheca when sclerotized. phallotheca sclerotized proximal part of phallosoma (q. v.). espe¬ cially in Cimicomorpha and Pentatomomorpha. phallus (pi., phalli) intromittent organ, including phallobase. aedeagus, and its various processes; see endosoma, phallosoma, phallotheca, vesica. plastron in Aphelocheiridae and Cryphocricini (Naucoridae), a physical gill formed by a dense mat of microtrichia on ventral body surface. plate-shaped ovipositor ovipositor with shortened valvulae which may be fused, reduced, and dorsoventrally compressed; see laciniate ovipositor. plectrum ordinarily movable portion of stridulatory mechanism; see stridulitrum. 320 Glossary pleustonic of or pertaining to air-water interface, e.g., Gerridae are pleustonic. ponticulus transversalis large dorsal rodlike transverse superior connection between basal plates in male phallus; see articulatory apparatus, basal plates. pore-bearing plate in Hyocephalidae, an ovoid pore-bearing structure on each side of abdominal sternum 3. posterior gonapophyses second valvulae, q.v. posterior pronotal lobe posterior expansion of pronotum over¬ laying part or more rarely all of mesonotum. posterior ramus second ramus, posterior valvulae second valvulae, q.v. posterior wall in female Miridae. a sclerotized, platelike struc¬ ture lying between the rami of the second valvulae. postnodal pruina in Corixidae, a white, frosted area along lateral border of corium posterior to nodal furrow (costal fracture), preapical claw in Gerroidea, a condition in which pietarsus is inserted proximal to apex of last tarsal segment, primary gonopore distal end of ductus ejaculatorius before enter¬ ing phallus at level of basal foramen to merge into ductus seminis; see secondary gonopore. principal salivary gland major salivary gland of paired sali¬ vary system, with 2 or more lobes, always associated with an accessory salivary gland, processus gonopori flagellum, q.v. proctiger reduced abdominal segment 10, bearing anus, possibly surrounding invaginated abdominal segment 11. pronotal carina primarily in Tingidae. main or median carina or keel on pronotum. prosternal furrow in most Reduviidae. a cross-striated longitu¬ dinal groove in prostemum, by means of which stridulation is caused by rubbing apex of rostrum in it by up-and-down move¬ ments of the head. pseudaroUum (pi., pseudarolia) in Miridae, pulvlllus, q.v. pseudomicropyle in eggs of Cimicomorpha, hollow chorionic micropyle-like processes used for gas exchange, pseudoperculum an egg cap without a distinct sealing bar and in which eclosion is not result of fluid pressure, pseudoplacental viviparity viviparity in which eggs contain tittle or no yolk and embryo presumably receives nourishment from a pseudoplacenta, e.g., in Polyctenidae. pseudopulvilli in Miridae, paired pretarsal structures arising lat¬ erally from unguitractor plate, distinct from parempodia and often superficially resembling pulvilli; see also accessory parem- podium. pseudospermathecae in Pachynomidae. Reduviidae, and Tingi¬ dae, which lack functional spermatheca, 1 or 2 saclike or tubular diverticula arising from vagina or common oviduct, functioning as sperm-storage organs. pseudospiracle in Nepidae, hydrostatic organ, q.v. pulvillus (pi., pulvilli) in Miridae, some Anthocoridae, nearly all Pentatomomorpha, bladderlike pretarsal structures arising from ventral or mesal surfaces of claws; see arolium, basipulvillus, distipulvillus, empodium, parempodia, pseudopulvillus. pygofer pygophore, q.v. pygophore abdominal segment 9 of male, enclosing the phallus, ramus (pi., rami) connecting leaf (or arm) of ovipositor valvulae. raptorial adapted for seizing prey, e.g., forelegs of many preda¬ ceous Heteroptera. receptaculum seminis spermatheca, q.v. rectal organ proctiger, q.v. reel system in Saldidae, differentiation of ductus seminis at junc¬ tion of ductus seminis proximalis and distalis into a coiled tube. remigium anterior part of wing chiefly involved in flight; the wing anterior to the claval suture. respiratory siphon in Nepidae, paired caudal structures derived from abdominal tergum 8, forming a channel in nymphs and a long tube in adults, which connect with eighth abdominal spiracles, and which serve to replenish subhemelytral air store in these aquatic insects. Ribaga’s organ ectospermalege or spermalege. q.v. ringed glands in some Heteroptera, paired or unpaired glands, dorsally or ventrally on vagina or on vaginal pouch, or bursa copulatrix, sometimes ringed by annular sclerotizations known as ring sclerites. ring sclerite in some Lygaeidae, cricoid sclerite (q.v.); paired or unpaired annular sclerotization encircling ringed glands of vagina, vaginal pouch, or bursa copulatrix. rostrum combined labium and maxillary and mandibular stylets, rotatory coxae nearly globose hind coxae with a ball-and-socket articulation; see Trochalopoda. salivary sheath lipoprotein sheath left in plant tissue, formed from hardened salivary secretions, encasing stylets as they pene¬ trate plant tissue. sawing-clipping feeding method of feeding in which stylets are moved back and forth in a straight line; see lacerate-flush feeding, scape the basal segment of the antennae. scent glands dorsal abdominal scent gland in nymphs (some¬ times persisting to adulthood) and several types of scent glands in adults (see Brindley’s glands, metathoracic scent gland), pro¬ ducing pheromones, allomones, venoms, and other substances, with often notorious and unpleasant smell for humans, scent pore ostiole, q.v. scent reservoir paired or unpaired reservoir of metathoracic scent glands; see diadenian type, diastomian type, omphalian type, sclerotized ring ring sclerite, q.v. scrobe a groove, as in foretibia of Phymatinae (Reduviidae) for reception of tarsus. scutellum (pi., scutella) triangular part of mesothorax, generally placed between bases of hemelytra, but in some Pentatomoidea partly or completely overlapping them, sealing bar in eggs of Cimicomorpha. a bar joining cap to rest of chorion, consisting of a very thin layer of resistant endochorion and a thick amber layer. secondary gonopore opening of ductus seminis at or near apex of phallus. secondary hypocostal ridge m adult Heteroptera. a secondary modification of hypocostal lamina, second gonocoxae second valvifers, q.v. second gonocoxopodites second valvifers, q.v. second ramus connecting leaf of second valifer. second valvifers valifers arising from abdominal segment 9. second valvulae median blades of the ovipositor, seminal duct ductus seminis, q.v. seminal reservoir ejaculatory reservoir, q.v. semiring sclerite in Colobathristidae. cricoid sclerite, q.v. sieve pores peg plates, q.v. spermalege (pi., spermalegia) in some Cimicoidea, an organ on pregenital abdominal segments receiving sperm during traumatic insemination and lacking a direct communication with genital apparatus itself; usually consisting of an external integumental pouch (ectospermalege) and an internal mesodermal part (meso- spermalege). spermatheca (pi., spermathecae) median, dorsal, unpaired, sclerotized diverticulum of vagina serving as sperm-storage receptacle; receptaculum seminis (q.v.); vermiform gland (q.v.); Glossary 321 spermathecal tube (q.v.); see also concepiaculum seminis. pseudospermathecae. spermathecal bulb generally bulb-shaped terminal portion of spermatheca serving actual sperm-storage function, spermathecal duct ductus spermathecae, q.v. spermathecal gland vermiform gland, q.v. spermathecal pump part of spermatheca between ductus spermathecae and spermathecal bulb, frequently differentiated— in true spermatheca—into a muscular pump with flanges, spermathecal tube in infraorder Gerromorpha and some other Heteroptera, an elongate, looped spermatheca with glandular cells in its walls; see spermatheca. spermatic duct in many Cimicoidea, duct arising from fusion of vasa deferentia. spermatic furrow in many Cimicoidea, groove of left paramere in which runs interlocked phallus, spermodes in Cimicidae, intraepithelial network of canals in walls of pedicels and paired oviducts through which spermatozoa pass from conceptacula seminis to ovarioles. sperm reservoir ejaculatory reservoir, q.v. spiracular line a line drawn through the spiracles on the abdomi¬ nal venter of trichophoran Pentatomomorpha. used to refer to position of abdominal trichobothria relative to spiracles, spongy fossa fossula spongiosa, q.v. stapes basal plates, q.v. staphylinoid condition in which hemelytra are reduced and trun¬ cate. static sense organs see hydrostatic organs, strainer in Hyocephalidae. pore-bearing plate, q v. stridulitrum ordinarily stationary portion of stridulatory mecha¬ nism; see plectrum. Strigil, strigile, strigilis stridulitrum, q.v. styli (sing., stylus) parameres. q.v. subgenital plate in most female Heteroptera abdominal ster¬ num 7, in Enicocephalidae sternum 8. submacropterous condition of wings in which corium and clavus of forewings are fully developed, with membrane being slightly to greatly reduced, hind wings generally being functional; see brachypterous. supradistal plate in some male Enicocephalidae, dorsal cover of genitalia. suspensorial apodemes internal muscle attachments to which are affixed basal plates of articulatory apparatus, suspensory arms suspensorial apodemes, q.v. suspensory processes suspensorial apodemes, q.v. swarming in Enicocephalidae, aggregating in a mating swarm, q.v. swimming fan in some Veliidae, e.g., Rhagovelia (Veliidae), fan¬ like structure usually formed from a modified ventral arolium and which aids in swimming on flowing water, synthlipsis minimum interocular distance, tegmen (pi., tegmina), tegminai a forewing not differentiated into proximal coriaceous and distal membranous part, as for example in Enicocephalomorpha; see hemelytron. third valvulae a sheathlike structure of the ovipositor, fused with second valifers, absent in all Pentatomomorpha. traumatic insemination in many Cimicoidea and some Nabidae, puncturing of body wall or wall of inner genitalia by phallus dur¬ ing mating and deposition of sperm outside usual reproductive tract. trichobothrium (pi., trichobothria) specialized, slender, hair¬ like, sensory setae arising from and including tubercles or pits (bothria) on many body regions and appendages in Heteroptera; see bothrium. trichomes modified setae present on certain myrmecophilous in¬ sects which give off secretions that ants (Hymenoptera: Formici- dae) imbibe, e.g.. base of abdomen in Holoptilinae (Reduviidae). TVichophora those members of Heteroptera with trichobothria on pregenital abdominal sterna, i.e., Pentatomomorpha less Ara- doidea. Trochalopoda that grouping of Heteroptera in which posterior coxae are nearly globose and articulation is a ball-and-socket joint; see Pagiopoda. true bug a heteropteran. trypanosomiasis a disease caused by infection with Trypanosoma (Trypanosomatidae), transmitted by Triatominae (Reduviidae); see Chagas' disease. tyius (pi., tyli) distal part of clypeus; anteclypeal region, tymbal in some Heteroptera, a sound-producing membrane on abdominal segment 1 or segments 1 and 2. tympanal organ organ sensitive to vibrations, on mesothorax of some Nepomorpha. unguitractor plate sclerite lying between bases of claws, with which bases of claws articulate distally, to which retractor tendon is attached proximally, and from which parempodia arise distally. vagina (pi., vaginae) ectodermal genital duct distal to common oviduct. vaginal pouch variably formed pouch that may bear ringed glands, forming part of gynatrial complex, valvifers (1st and 2nd) in female Heteroptera, 4 plates or blades. 2 from abdominal segment 8, 2 from abdominal segment 9. articulating on corresponding paratergites and bearing first and second valvulae, respectively. valvulae the blades (in laciniate type) of the ovipositor, which in 2 pairs, form the egg-laying apparatus, and which proximally attach to the body wall via one (or 2) pairs of corresponding rami. ventral arolium arolium, q.v. ventral glands glands located in metathorax of some Reduviidae, distinct from metathoracic scent glands, ventral laterotergites ventrally situated laterotergites, distinct from dorsal laterotergites (q.v.) but generally fused with sternum and usually bearing spiracles, ventral lobe in Gerromorpha, buccula, q.v. ventral paratergites ventral laterotergites, q.v. ventral plates in some male Enicocephalidae, fused genital plates. ventral spine in Pentatomoidea, a spinelike projection anteriorly or third true abdominal sternum, directed toward head and lying at times between coxae. vermiform gland in Cimicomorpha, an organ homologous with spermatheca but without sperm-storing function, vesica (pi., vesicae) portion of endosoma of phallus, often scle- rotized, surrounding ductus seminus distalis. vesical process process arising from the vesica, water bug member of infraorder Nepomorpha. wing-coupling mechanism microtrichia-bearing stmeture at posteroventral margin of clavus of forewing, grasping leading edge of hind wing during expansion and flexion of wings and during flight. wing-locking mechanism in Euheteroptera, modification of cos¬ tal margin of forewing and mesothorax to retain wing firmly in postion in repose (see frenum); see wing-coupling mechanism. X organ epipharyngeal sense organ, q.v. xyphus, xiphus a spinous triangular process of prosternum or mesosternum, or both, y organ epipharyngeal sense organ, q.v. 322 Glossary Ml'' ' INDEX abdomen, 49; expansion of, 159, 201; membranous, 164 abdominal gland, 198 abdominal grasping apparatus, 134, 135. 137, 140, 141, 143 abdominal hair pile, 108 abdominal organ, pregenital ventral, 52 abdominal scent glands; functional, in adults, 126, 267, 282; nonfunctional, in nymphs, 111 abdominal sense organs: in Aphelocheiri- dae, 126; in Naucoridae, 124 abdominal spine, 231 abdominal spiracles, 74, 78 abdominal sutures, fusion of, 97, 182, 269 abdominal tymbal, 160 Abedus, 113; indentatus, 112 Abies procera, 22 Abulites, 216 Acacia, 30, 184, 267, 277, 281; c«n- ninghami, 169; maidenii, 169 Acalifa diversifolia, 233 Acalypta, 184 Acanthaspis, 158; petax, 160 Acanthia, 138 Acanthoberytus wygodzinskyi, 247 Acanthocephahni, 277 Acanthocerini, 277 Acanthocorini, 277 Acanthocrompus, 257 Acantholomidea, 238 Acanthomia horrida, 33 Acanthosoma haemorroidale, 217 Acanthosomatidae, 39, 58, 215, 216 Acanthosomatinae, 216 accessory gland, 55, 151 accessory parempodia, 48, 136, 168, 175 accessory salivary gland, 59, 60 Acer negundo, 36, 282 Acestra malayana, 273 Achaearanea tepidariorum, 160 Acinocoris calidus, 268; stehliki, 268 Acompocoris pygmaeus, 196 Acromyrmex lundi, 184 acus. 191, 194 adaptationist arguments, 4 Adelphocoris, 179; lineolatus, 34 Adenocoris, 212 adhesive pads, of hind coxae, 81-83, 135, 141, 142 Adrisa, 206 aedeagus, 49, 50 Aelia germari, 35; rostrata, 35 Aenictopecheidae, 68 Aenictopecheinae, 68 Aenictopecheini, 69 Aenictopechys necopinatus, 69 Aepophilidae, 136 Aepophilus, 137; bonnairei, 21, 136, 137 Aeptini, 232 aeropyles, 63, 146 Aesepus, 216 aestivation, 274 Aethus indicus, 33 Afrocimex, 201 Afrocimicinae, 201 Afropiesma, 266 Afrovelia phoretica, 101, 102 Agamedes, 216 Agathyma ceramica, 277 Agave, 179 aggregation, 22, 238 aggregation pheromone, 279 aggregations, dry season, 274 aggressive behavior, 51 agonistic behavior, 51 Agonoscelis rutila, 35; versicolor, 35 Agrammatinae, 182 Agraptocorixa hyalinipennis, 122 Agriopocorinae, 275 Agriopocoris, 275 airstore; abdominal, 114, 128, 130; sub- hemelytral, 121 airstraps, 113, 114 alary muscles, of heart. 61,63 Alathetus haitiensis. 232. 233 Alberproseniini. 159 alcohol preservation. 18 alfalfa. 34 alfalfa seed, 32 algae, as food, 122 Alienates. 70, 71; ehngalus. 72 Alienatinae, 71 alimentary canal, 59. 60. 271,273 alkaloids, 27, 263 Alloeorhynchus, 188. IS9; mabokei. 187 allspice. 34 Almeida, 197 Almeidini. 197 Alydidae, 29, 32, 58. 271,272 Alydinae, 273 Alydus. 273, 274 Amaranthaceae, 267 Amblypelta cocophaga, 33 Ambrysini, 125 Ambrysus magniceps. 126 Amemboa, 105 American Museum of Natural History. 15 Ammianus alberti, 181 Amnestinae, 222 Amnesius, 222; spinifrons, 221; subferrugi- neus. 224 Amorbini. 277 Amphaces, 216 Amphibicorisae. 1 Amphibolus Venator, 36 Amulius, 157 Amyot, Charles Jean-Baptiste, 1,6 anal furrow. 44 anal lobe. 44 anal veins, 43, 44 Anasa, 33, 279; alfaroi, 277; guayaquila, 277; tristis. 33 androconium, 58 Androctenes, 202 Andromeda, 36 Aneuraplera cimiciformis, 210 Aneurinae. 210 Aneurus. 208, 210. 213; laevis. 211 Angilia. 101 Angilovelia. 51. 101 Anhomoeini, 277 Anisopinae, 128 Anisops. 51, 128, 129; megalops, 129 Anisoscelidini, 277 Anolis. 27 Anommatocoris, 183; coleoptratus, 180, 184 Anoptocerus elevatus, 278 Anoplocnemis, 277; curvipes, 33 ant mimetic, 29 ant mimicry, models for, 29 anteclypeus, 41 antennae, 41, 149; antennae, concealed below head, 107, 108; flattened, 278; geniculate, 214; modified, 105, 106, 178; pseudosegmented, 156 antennal cleaner, 47, 117 antennal segmentation, 107 Antestiopsis lineaticollis, 35 Anthocoridae, 32, 195, 197 Index 323 Anthocorini, 197 Anthocoris, \97. nemorum, 196 Antillocorini, 40. 260 Anliieuchus, 233; iripterus limbativentris. 233 ants, attraction of. 160 anus, 49 aorta. 61 Aphelocheiridae, 55 Aphelocheirus, ITl', aestivalis, lll'.lalni. 109; trmlayensis, 127 Aphelonotinae, 39. 149 Aphetonotus, 54, 149, 150: a/ricanus. 150; fuscus. 150 aphids, predators of, 32 Aphylidae, 39, 218 Aphylum bergrolhi. 218; syntheticum. 218 Apiomerini, 157 Apiomerus. 157 Apodidae, 201 Apoplymus, 248 aposematic coloration, 27, 177, 257. 263. 269. 279, 283 apples, 34 Apierola, 257 aptery, 23 Aquarius, 105; lacustris, 63; paludum, 104 aquatic bugs. See Nepomorpha aquatic habitats, 21, 107 Araceae, 179 Arachnocorini, 188 Arachnocoris, 21, 186, 188, 192 Arachnophila cubana, 192 Aradacanthia, 210 Aradellini, 154 Aradidae, 39, 208, 209 Aradinae, 210 Aradiolus, 210 Aradus, 208, 210, 211; cinnamomeus. 211, 213 Arbela carayoni, 187 areolate sculpturing, 183 Arhaphe, 269; Carolina, 269 Arilus, 150, 157 Armstrongocoris, 156 arolia, 46, 146. See also dorsal arolium; parempodia; ventral arolium Artabanus lativenlris, 211 Artemisia, 36 Artheneinae, 255 Artheneini, 255 arthropod predators, 263 Asclepias curassavica, 22 Asclepios, 105 ash-gray leaf bugs. See Piesmatidae Ashlocic, Peter D., 6 Asopinae, 35, 230 aspirator, 17 Aspisocoris termitophilus, 213 Aspongopus, 206 assassin bugs. See Reduviidae Asterocoris australis, 211 Astragalus, 36 Atractotomus, 178 aubergine, 36 Auchenorrhyncha, 5 Augocoris, 239 Auricillocorini, 178 Australian National Insect Collection. 16 Australian Region, 38, 39 Australmeida, 197 Australostolus, 70; monteiihi. 44, 69 Austronepa, 116 Austronepini, 116 Austropamphanlus woodwardi. 255 Austrovelia, 88 autapomorphy, 4 Azalea, 36 Baccharis, 184 Bacillometra, 95 backswimmers. See Notonectidae Baclozygum, 167; brevipilosum, 169; de- pressum, 167-169 Bactrodes, 154, 156 Bactrodinae, 154 Bactrophyamixia, 274 Bagrada cruciferarum, 35 balloon vine, 282 bananas, 33, 184, 266 Banksia, 169 Bapiista, 10! Barber, Harry G., 6 Barce, 54 bark, as habitat, 163 bark bugs. See Aradidae Barreratalpini, 277 basal articulatory apparatus, 49 basal foramen, 49 basal plates, 49 basiflagellum, 41 basipulvillus, 205 Bassian Subregion, 39 bat parasites. See Polyctenidae Bathycles amarali, 263 Bathycoelia thalassina, 35 bats, as hosts, 201, 203 Beamerella balius, 171, personatus, 174 beans, 33-36, 266 beating sheet, 17 bed bugs. See Cimicidae beetles, myrmecomorphic, 29 Belonochilus numenius, 263 Belostoma, 113 Belostomatidae, 45, 111, 112, 114 Belostomatinae, 113 Berg, Carlos, 6 Bergevin, Ernest de, 6 Bergroth, Ernst Evald, 6 Berlese funnel, 17 Bertilia, 201 Berytidae, 32, 246, 248 Berytinac, 248 Berytinus, 248; hirticornis, 247; minor, 247 Be/ytus, 248 Betula, 263 Beyeria, 267 bibliographies, 14 big-eyed bugs. See Geocorinae Bilia, 197, 198 Bilianella, 197, 198 Biliola, 197, 198 biological control, 257 Biosystematics Research Centre, Agricul¬ ture Canada, 15 Biprorlulus bibax, 35 bird nests, as habitats, 195 birds, as predators, 28 Bishop Museum, 15 bivoltine lifecycle, 25, 179. 264 black pepper, 34 Blaptostethini, 197 Blaptostethoides, 197 Blaptostethus. 197 Blatchley, W. S., 6 Blaudusinae, 216 Blaudusini, 216 Bledionotinae, 255 Bledionotini, 255 Bledionotus, 29; systellonotoides, 29, 255 Blindwanzen. See Miridae Blissinae, 45, 255, 263 Blissus insularis, 33: leucopteruST-li^: ' leucopierus hirtus, 33 Bliven, B. P..7 Bloeteomedes, 161 blood feeding, 21, 159, 263 blue jays. 283 Boisea, 57, 282, 283; trivitiata, 36. 281, 282 Bolboderini, 159 Boraginaceae. 224 Boreostolus, 68-70; americanus, 70; sikh- otalinensis, 70 Bothriomirini, 176 Botocudo cavernicola, 261 box elder, as host. 282 box elder bug, 36, 282 Boxiopsis madagascariensis, 34 Bozius respersus, 237 Brachelytron angelicus, 232 Brachylybas, 277 Brachymetra, 104 Brachyplatys, 237 brachyptery. 23, 264 Brachysteles. 197 brain. See supraesophageal ganglion Breddin, Gustav, 7 bright coloration, 27, 240 Brindley's gland. 57, 148, 151, 159, 181 broad-headed bugs. See Alydidae broad-shouldered water striders. See Veli- idae Brochymena, 233 Bromeliaceae. as habitat, 101 bronze orange bug, 36 brooding behavior, 114 Bryocorinae, 34, 173 Bryocorini, 48, 173 Bryocoris, 173 bucculae, 41 buccular bridge, 41 Buchananiella, 197 Bucimex, 201 Buddleia, 263 Buenoa, 51. 53, 128, 129 Burmeister, Hermann Carl Conrad, 7 burrower bugs. See Cydnidae 324 Index bursa copulatrix, 50, 51 Burtinus, 274 Butler, Edward A., 7 cacao, 34, 35 Cacodminae, 201 Cacodmus bambusicola, 200 caducous wings. 23 Caenusia, 253 Cajanus cajan. 33 California Academy of Sciences. 15 Caliotis colarata. 198 Calisiinae, 210 Calisiopsis, 210 Calisius, 210 calli, 43 Callichilella grandis, 177 Callidea, 240 calling signals, 106 Callitris preissii, 228 Calocoris norvegicus, 34 Calotropis procera, 22 Camarochilus, 149, 150; americanus. 150 camouflage, 160, 235 campaniform sensillum, 46, 89, 96 Camptopus, 273; lateralis, 32, 273, 274 Campyloneura virgula, 179 Canopidae, 40, 219 Canopus, 219, 220; burmeisteri, 220; caesus, 219, 220; impressus, 220; orbicu¬ laris, 220 canopy fogging, 17 Cantacaderinae, 182 Cantacaderini, 182 Cantacader quadricornis, 183 Cantao ocellatus, 240 camharadin, attraction to, 180 capsids. See Miridae Caravalhoma malcolmae, 174 Carayonia, 151 Carayon’s glands. See ventral glands Carcinocorini, 154 Carcinocoris. 45 cardenolides, 27. 28 Cardiastethus, 197 cardinate coxae, 45 Cardiospermum corindum, 282; halicaca- bum, 282 Carpocoris, 35 Carrabas, 224 Carthasini, 188 Carthasis, 187 Carvalhoma, 179; malcolmae, 169 Carventinae, 210 Caryophyllaceae, 267 cashew, 34 Casuarina, 277 Catadispon, 216 catalogs, 14 Cataractocorini, 125 Catorhintha, 279; siblica, 277 Caulotops, 179 Cavaticovelia, 88, 90 Cavelerius sweeti, 34 Cavernicolini, 159 caves, as habitats, 90, 160, 224, 264 Cecyrina, 230 Ceiba, 35 central duct, 43 central ganglion, 61,62 central nervous system, 61 Centrocneminae, 154 cephalonotum, 131 Ceraleptus obtusus, 278 Ceratocapsini, 178 Ceratocapsus, 178 Ceratocombidae, 75, 77 Ceratocombinae, 77 Ceratocombini, 77 Ceratocombus, 77; corticalis, 77; mareki, 76 Ceratocoris, 236, 237 Cercotmetus, 116 cereal crops, 232 Cethera, 156 Cetherinae, 39, 156 Cetherini, 156 Chaetometra, 95, 97 Chagas’ disease, 36, 159 character polarity, 4 Chariesterini, 211 Charmatometra, 104 Charmatometrinae, 104 Chartoscirta, 140; cocksii, 139 Chauliopinae, 266 ChauUops, 34, 266 Cheirochela, 126 Cheirochelinae, 125 Cheirochelini, 125 chelae, on forelegs, 159 Chelinidea, 33 Chelinideini, 277 Chelonocorisferrugineus, 211; javensis, 211 Chenopodiaceae, 267 Chepuvelia, 93, 94; usingeri, 94 Chilacis typhae, 264 Chiloxanthinae, 140 Chiloxanthus, 140 Chimarrhometra, 105, 106 China, William Edward, 7 Chinamyersia, 212 Chinamyersiinae, 39, 212 Chinamyersiini, 212 Chinaola, 163; quercicola, 163 chinch bug, 33, 255 Chironomidae, 39 Chiton, 215 Chlamydatus, 178 chlorazol black, 19 chorion, 62 Chorosomini, 282 chromosome numbers, 63, 243, 245, 256 chromosomes, 63 Chrysocoris, 240 Chryxinae, 156 Chryxus, 156 cibarial dilator muscles, 43 cibarial pump, 60 Cimex, 201; hemipterus, 32, 201; lectu- larius, 32, 200, 201 Cimicidae, 32, 199. 200 Cimicinae, 201 Cimicoidea, 49, 51 Cimicomorpha, 5, 43, 46, 48, 57, 146 cinchona, 34 Ciorulla, 163 circulatory system, 61,63 Cistaceae, 267 citrus. 35. 36 cladistics, 4, 38 claspers. See parameres claval commissure, 44. 243 claval suture, 44 Clavigralla elongate, 33; gibbosa, 33; tomentosicollis, 33 clavus, 43 claws, 46, 48; cleft, 176. 177; spicules on, 176; with subapical tooth, 176-178; toothed, 156; unequal, 192 cleaning, ultrasonic, 18 Cleontes, 157 Clerada, 263; apicicornis, 260 Cleradini, 260 cliff swallows, 33 Cligenes subcavicola, 264 Clivinemini, 177 Cloresmini, 277 Closlerocoris, 173, 177 clypeus, 41; swollen, 237 Cnethocymatia, 121 Cobben, Rene H.. 7 Codes, 72 coconuts, 33 Codophila, 35 coffee, 35, 36 Coleoptera larvae, as prey. 233 Coleopterocoris, 123 Coleopterodes, 184 coleopteroid adults, 184 coleoptery, 23 Coleorrhyncha, 5 collar, 41 Collartidini, 156 collecting equipment, 17 collections, 15 collum. See collar Colobalhristes chalcocephalus, 249 Colobathristidae, 33. 40. 249 Colobathristinae, 250 color, changing, 30 Colorado potato beetle, 35 coloration; abdominal. 116; aposematic, 27. 177. 257, 263,-269, 279. 283; bright. 27. 240; cryptic, 283; deflective, 30, 279; metallic, in nymphs, 269; myrmecomor- phic, 29; protective, 30 Colpurini, 277 commensals, 21 competition, 22 compound eyes, 41; absent, 202, 214; reduced, 201 concealment, 31 conjunctiva, 49, 50 connexivum, 49 continental drift, 38 Copium, 184 Coptosoma, 231, 238; inclusa, 237; lyncea, 22, 238; scutellatum, 237 Index 325 copulation method, 257 copulatory position, 126, 134 copulatory tube, 191-193, 195, 198. 199 Coquillettia, 29; insignis, 27, 28 coral, as habitat, 97 Corallocoris, \Al , nauruensis, 142 coral treaders. See Hermatobatidae Corcovadocola, 179 Corecoris, 33 Coreidae, 33, 45, 46, 58, 274, 275 Coreinae, 277 Coreini, 277 corial glands, 56, 59. 190, 192, 193 Coridius, 206; janus, 33, 226; viduatus. 226 Coridromius, 45 Corimelaena lateralis, 223\pulicaha, 223 Corimelaeninae, 222 corium, 44 Corixadentipes, \22\jakowleffi, 122 Corixidae, 45, 120 Corixinae, 121 Corizomorphini, 282 Corizus hyoscyami, 282 com, 33 corpora allata, 61 corpora cardiaca, 61 Corythaica cyathicollis, 37; monacha, 37 Corythuca, 182; ciliala, 36; gossypii, 36 Cosmocoris, 240 costa, 43, 44 costal fracture, 43, 69, 139, 193 cotton, 33-36, 240 cotton Stainers. See Pyrrhocoridae coupling mechanisms, wing-to-body, 44 courtship, 51 courtship signals, 106 coxae, 45 coxal combs, 220, 224, 243 creeping water bugs. See Naucoridae Creontiades pallidus, 34 critical point drying, 18 crop damage, 32 crop plants, 34 Cruciferae, 35 crusader bug, 33 Cryphocricinae, 125 Cryphocricini, 125 Cryphocricos, 24, 125; hungerfordi, 24; latus,Al, 108, 109 Cryptacrus, 240 Cryptocerata, 1 Cryptophysoderesfairchildi, 158 Cryptorhamphinae, 256 Cryptorhamphus, 256 Crypiostemma, 70, 78; incurvatum, 79 Cryptovelia, 88, 90; terrestris, 88, 89 ctenidia, 202, 203 Ctypomiris, 179 cubitus, 43, 44‘ Cucumis, 33 Cucurbitaceae, 33, 226 cuneus, 43, 162, 171 Curicta, 116 Curictini, 116 cursorial legs, 45 Cydopelta obscura, 33 Cvdamus, 22A Cydnidae, 33, 220, 222 Cydninae, 45, 222 Cydnus alerrimus, 221 Cylapinae. 39, 176 Cylapini. 176 Cylapocoris, 179 Cylapus, 175, 179 Cylindrostethinae, 104 Cylindrostethus, 104, \0S; palmaris, 104 Cyllarini, 277 Cylopelta, 225 Cymalia, 121 Cymatiainae, 121 Cyminae, 30, 256, 263 Cymini, 256 Cymophyes, 258 Cymus, 22, angustalus, 254 Cyperaceae, 255 Cyphopelta, 173, 177 Cyrtocorinae, 231 Cyrtocoris trigonus, 233 Cyriomenus crassus, 223 Cyrlopeltis ebaeus. 175; tenuis. 34 Cyrtopeltocoris, 173 Daciera punctata, 273 Daladerini, 277 Dallas, William Sweetland, 7 damsel bugs. See Nabidae Darwinivelia, 88, 90 Dasycnemini, 154 Dasynini, 277 Dayakiella, 249, 250; brevicornis, 250; sumatrensis, 250 Dayakiellinae, 250 deciduous wings, 23 defensive behavior, 51 defensive reaction, 114 deflective coloration, 30, 279 De Geer, Carl, 7 Deliastini, 156 deotocerebrum, 61 Departamento de Entomologi'a. Museo de La Plata, 16 Department of Biology, Nankai University, Tianjin. China, 16 Deraeocorinae, 177. 179 Deraeocorini, 177 Deraeocoris, 177 dermal glands, 58 Deroplax, 239 deutocerebrum, 62 Diactor, 46 diapause, 264 Diaprepocorinae, 121 Diaprepocoris, 121 Diaspidiini, 157 Diaspidius, 157 Diconocoris capusi, 182 Dicranocephalus, 283; agilis, 284; albipes. 284; pallidus, 284; setulosus, 284 Dictyonota strichnocera, 59 Dicyphina, 48, 173 Dicyphini, 173 Dicyphus, 173, 178 Diemeniini, 232 Dieuclies. 262 Dilompini. 255 Dilompus. 255 Dinidor. 225. 226 Dinidoridae. 33. 225 Dinidorinae. 225 Dinidorini, 225 Dinomachus. 264 Diocoris. 46 Diolcus irroratus. 240 Diphlebini, 177 Diplonychus, 109, ] 13; rusiicum. 113 Diplura macrura, 192 Dipsocoridae, 78 Dipsocoris, 78 Dipsocoromorpha, 46, 48, 74 Discocephalinae, 231 Discocoris. 167, \69; drakei, 166. 168 Discogastrini, 277 disc-shaped organs, on female abdomen. 227 dispersal. 22, 263 dispersalist hypotheses, 38 Distant. William Lucas. 7 Distantiella iheobroma, 34 distasteful species, 27 distiflagellum, 41 distipulvillus, 205 distribution, 38 Ditomotarsinae, 216 . Ditomotarsini, 217 Doesbergiana, 225 Dolichiella, 190 Dolichocephalometra, 95, 97 Dolycoris, 35 dorsal arolium, 48, 91,93, 96, 98, 109, 135 dorsal vessel, 61,63 Douglas, John William, 8 Drake. Carl J., 8 Dmckknopfsystem, 45 Drymini, 260, 263 Dryophilocoris, 178 ductus ejaculatorius, 49 ductus seminis, 49 Dufour, Leon, 1, 8 Dufouriellini. 197 Dulmius unicolor, 35 dust bugs, 31 dwellings of humans, as habitats, 160 Dysdercus. 23, 28, 35, 36, 40, 58, 270, 271; bimaculatus, 270; blotei, 271; dis¬ color, 21\;fasciatus, 58, 270. 271; koenigii, 271; obscuratus, 28 Eccritotarsina, 40, 176, 179 Eccritotarsini, 176 Eccritotarsus, 179 economic importance, 32 Ectinoderini, 157 Ectinoderus, 157 Ectomocoris, 157 ectoparasites, 201,203 ectospermalege, 63, 198 Ectrichodia gigas, 160 Ectrichodiinae, 156, 160 326 Index Edessa, 48. 54 Edessinae, 232 egg burster, 62 egg eap. See operculum egg guarding. 160. 213. 233, 240 eggs. 62, 64; deposition of, 160 ejaculatory reservoir. 49 Ekblom’s Organ, 186. 188, 189 Elaeocarpus obovatus. 169 Elasmodema, 154; erichsoni, 152. 155 Elasmodeminae. 58. 154 Elasmolomus sordidus. 34 Elasmoslethus crucialus. 217 Elasmucha dorsalis. 217; putoni. 217 electric light bugs. See Belostomatidae EUenia, 173 Elvisurini, 239 Emballonuridae. 204 Embiidina, 193 Embiophila. 192; {Acladina), 192, africana, 193; myersi. 192 Embiophilinae, 192 embolium, 44 embryonic development; in Lyctocoridae, 195; in Plokiophilidae, 193 Emesinae, 156 Emesini, 156 Empicoris, 150 Enatosalda. 138, 140 endocrine glands, 61 endosoma, 49, 50 Engistus, 257 Enicocephalidae, 70, 71 Enicocephalinae, 71 Enicocephalini, 72 Enicocephalomorpha, 39, 43, 45, 46, 48, 67 Enicocephalus, 72 Enithares, 128, \29; maai, 129 Entomological Laboratory, Kyushu Univer¬ sity, 16 Emomovelia, 99; doveri, 100 Eobates, 104 Eocanthoconafurcellata, 35 Eocienes, 202; spasmae. 203 Eotrechinae, 105 Eotrechus, 105, 106 epipharyngeal sense organ, 41 Erianotus. \A5, hnosus. 144 Esaki, Teiso, 8 Esakia, 105 esophagus, 59, 60 Ethiopian Region, 38, 39 Eucalyptus, 218, 281; camaldulensis, 218; globosus, 169; trachyphloia, 169 Eugenia, 229 Eumenotes, 225, 226; obscura, 226 Eumenotini, 225 Euphenini, 156 Eupheno, 156 Euphorbiaceae, 233, 283 Eupychodera corrugata, 240 Eurydema, 35 Eurygaster, 36, 239, 240; integriceps, 36 Eurygastrinae, 239 Eurygerris, 102, 105 Eurylochus, 158 Eurymetra, 106; natalensis, 104 Euryophthalmus. 269 Euschistus, 35, 233 Eusolenophora, 190 Eusthenini, 242 Eulhetus, 272, 273 Eutorpe edulis, 169 Euvelia, 99 evaporatory area (evaporatorium), 43. 56. 57,78,83, 146,205 eversible glands, 58. 137, 138, 140 exocorium, 161 exocrine glands, 55 expansion skating, 102 Eysarcoris ventralis, 35 Fabaceae, 226 Fabricius, Johann Christian, 1, 8 Falconia, 173 Fallen, Carl Friedrich, 8 faunistic studies, 15 fecundation canal, 89, 91, 100. 104 fecundation pump, 100, 104 feeding, 20 feeding cone, 20 feeding habits, in Joppeicidae, 165 female genitalia, 50 femora, 45 fertilization, 51, 188, 195; in vitellarium. 190 Feshina, 75; schmitzi, 77 Ficus, 217, 264 Fieber, Franz Xavier, 8 figs, 32, 263 filter chamber, 60 first ramus, 50 first thoracic ganglion, 62 first valvifer, 50 first valvula, 50 flake cuticle, 57 flat bugs. See Aradidae flat-podded golden rain tree, 282 flight, loss of, 29 flight cycle, 263 flight intercept traps, 17 flightlessness, 23 floral glands, 58 flower bugs. See Anthocoridae flowers, feeding on, 179 flowing water, as habitat, 106 foam masses, as habitat, 101 food canal, 41,43 forefemora, 45; enlarged, 45, 117, 148, 159, 263 foregut, 60 foretarsi, 77 foretibiae, spinose, 243 foretibial sense organ, 116 forewing, 43, 44; folding, 219, 220, 228 Fort Morgan virus, 33 fossils, 67, 140, 143 fossorial legs, 45, 224 fossula spongiosa, 47, 48, 148, 151, 159, 186, 190, 195, 199 frenum, 44 Froeschnerana mexicanus, 170 frontal ganglion, 62 frontoclypeal sense organ, 130 fruit feeding, 270 Fucus. 137 Fulviini. 176 Fulvius. 176. 179: quadristillatus. 156 fungal mycelia, 213 fungi, as hosts, 21. 179. 213. 220 Fusariurn, 33 Galeatus. 182 Galgulus. 117; ovalis. 223 gall formation, 184 Gamostolini, 69 Gamostolus, 68-70; subaniarcticus, 69 Gampsocoris culicinus, 247; panormimus. 247 Gargaphia torresi, 37 Garsauria, 222 Garsauriinae, 222 gastric caeca, 59, 60. 205, 271,273 Gastrodes, 253 Gelastocoridae, 45. 118 Gelastocoris, 30, 117. 118; oculatus, 47, 62. 108, 109, 117, 118;pente/i5/s, 117 genital capsule, 49; rotated, in male Helo- trophidae, 130 genitalia; asymmetrical, 49; as characters, 3; dissection and preparation of, 19; female, 49; male, 49 Geocorinae, 34, 257, 263 Geocoris, 34 Geocorisae, 1,4 Gerridae, 25, 45, 49, 102, 103 Gerrinae, 105 Gerrini, 105 Gerris, 105, 106; incurvatus, 86 Gerromorpha, 45, 46, 48, 87 Ghandi bug, 32 giant water bugs. See Belostomatidae Gigamemetopus rossi, 177 Gigantometra, 102, 104 glandular cuticular structures, 183 glandular pilosity, of host plants, 248 glandular setae, 264, 266; in nymphs, 247, 248. See also viscid setae Gleditsia triacanthos. 184 Glyptocombus, i2: fluminensis. 211 Gmelin, Johann Friedrich. 8 Gnostocoris, 212 Godefridus, 161 golden rain tree, 282 gonangulum, 225, 277 gonapophyses. See valvulae Gondwanaland, 39, 40 Gondwana patterns, 39 Gonianotini, 260 Gonocerini, 277 gonocoxae. See valvifers gonocoxopodites. See valivifers gonoplacs. See third valvulae gonopore, secondary, 49 gonostyli. See valvulae Gonystus, 256; nasutus, 256 Goondnomdanepa, 116 Index 327 Goondnomdanepini, 116 Gorpini, 188 Gorpis, 188 gourds, 33 Gramineae. See Poaceae grapes, 34 grasses, 34, 177. 179. 184, 250, 274 gregarious nymphs, 184 grooming comb, 46 grooved setae, 98, 101 ground-living species. 263 Guadua, 214 Guapinatmm, 75 guava, 34 gula. 41 gummosis, 33 gut. See alimentary canal Gymnocerata, 1 gynatrial complex, 50 gynatrial sac, 89, 100 habitat partitioning, 129 habitats; permanent. 24, 264; temporary. 24 Hadronetm, 27, \&0,uhteri, 180 Haematosiphoninae, 201 Haematosiphon inodorus, 33 Hahn, Carl Wilhelm, 8 hair pile, 84 Hakea, 111 Hallodapini, 178 Hallodapus, 179; albofascialus, 52 Halobates, 105, \Q6, sericeus, 86 Halobatinae, 105 Halobatini, 105 Halosalda, 140 Halovelia, 99, 101, 102 Haloveliinae, 99 Halovelioides, 99, 102 Halticini, 45, 177 Halticotoma, 179 Halticus, 45, 177 Halyini, 232 Hammacerinae, 156 Handlirsch, Anton, 8 harlequin cabbage bug, 35 harlequin lobe, 63 Harmostini, 282 Harpactorinae, 58, 156, 157 Harpactorini, 157 harpagones. See parameres Harpocera thoracica, 178 Harrisocoris, 157 head, 41 hearing, 53 heart, 61 heartseed vine, 282 Hebridae, 90, 92 Hebrinae, 92 Hebrometra, 92 Hebrovelia, 99, 101 Hebroveliini, 99 Hebrus, 92; pusillus, 91,92; ruficeps, 92; sobrinus, 85; {Subhebrus), 90 Heidemann, Otto, 8 Heissia, 210 Hekista, 173 Helopeltis, 34, 176; westwoodi, 171 Helolrephes, 131', admorsus. 132; bouvieri. 132 Helotrephidae. 45. 131 Helotrephinae. 131 hematophagy. See blood feeding hemelytra, 5,43,44; padlike. 199 Hemiptera, classification of, 1 hemoglobin, 129 Henestarinae, 257 Henesiaris, 257 Heniaries, 157 Henicocorinae. 257 Henicocoris momeithi, 257 Henschielh, 72 Heraeus trigunatus, 29 Hermatobates, 97; breddini, 97; weddi, 98 Hermatobatidae, 97 Herrich-Schaeffer. Gottlieb A. W., 8 Hesperocorixa interrupta, 122 Hesperocienes, 202-204; fumarius, 203 Hesperocteninae, 202 Hesperolabops, 179 Heierobtissus anomiUs, 51 Heierocleptes, 95. 96; hoberlandli, 96 Heterocleptinae, 95 Heterocorixa, 121 Heterocorixinae, 121 Heterogaster, 257 Heterogastrinae, 257. 264 Heteroptera, 1,45; classification of. 3, 5; as prey. 188 Heteropterodea, 5 Heteroscelis. 230; lepida, 231 Heierosceloides lepida, 231 Heterotermes convexinotatus, 215 Heterotrephes admorsus, 132 heterozygotes, 25 Hibiscus, 35 hind femora, 45 hind gut, 59, 60 hind legs, modified, in Rheumatobaies, 106 hind tibiae, expanded, 278 hind wings, 44 Hipposideridae, 204 Hirundinidae, 201 historical biogeography, 38 Hoffmanocoris, 157 Holarctic, 39 Holoptilinae, 39, 58, 154 Holoptilini, 154 Homalocoris, 156 Homoeocerini, 277 homology, 4 Homoptera, 1,5 homozygotes, lethal, 25 Hoplilocoris, 72 Horcias scutellalus, 174 Horvath, Geza, 9 Horvathinia, 113 Horvathiniinae, 113 Horvathiolus gibbicollis, 24 host choice, in Coreidae, 278 host plants, with glandular hairs, 179 Hoiea, 239 hot springs, as habitats, 132 house sparrows, 33 Hsiao, Tsai-Yu, 9 human blood, as food, 195 human food: Belostomatidae as, 114: Corixidae as, 122 humans, as hosts, 201 humeral angles. See pronotum Hungerford, Herbert Barker, 9 Hussey ella, 101 Hyaliodini, 177 Hyalochloria, 179 Hyalonysius pallidomaculatus, 263 Hyalopeplini, 177 Hyalymenus, 273 Hydara, 275 Hydarini. 277 Hydrocorisae, 1 Hydrocyrius, 113 Hydrometra, 52, 95-97; martini, 96; stag- norum, 96 Hydrometridae, 95 Hydrometrinae, 95 hydrostatic organs. See static sense organs Hydrotrephes, 108, 109 hygropetric species, 106 Hymenocoris brunneocephalis, 72 Hyocephalidae, 279 Hyocephalus aprugnus, 280 hypocerebral ganglion, 61 hypocostal lamina (hypocostal ridge), 43 Hypoctenes, 202 Hypselosoma, 82; hirashimai, 59, 81; matsumurae, 81 Hypselosomatinae, 82 Hypsipterygidae, 80 Hypsipteryx, 80; ecpaglus, 80; machadoi, 79, 80; ugandensis, 79, 80 Hyrcaninae, 92 Hyrcanus, 90, 92; capitatus, 91 Idiocarus, 125 Idiocorinae, 131 Idiocoris, 131; lithopbilus, 132 Idiostolidae, 39, 251 Idiostolus insularis, 251,252 lella, 190 ileorectal valve. See pyloric valve ileum. See pylorus immature fruits, feeding on, 233 Indo-Pacific distribution patterns, 40 infraorders, phylogenetic relationships of, 5 inquilines, 21, 184; in Embiidina webs, 192; in spider webs, 192 insemination, 51; androtraumatic, 73; trau¬ matic, 188, 192, 194, 195, 201,202; traumatic intravaginal, 188 Institute of Zoology, Academia Sinica, Beijing, 16 Instituto de Biologi'a, Universidad Aut6- nomo de Mexico, 16 internal anatomy, 3 intertidal dwarf bugs. See Omaniidae intertidal zone, as habitat, 21,90. 137, 140, 141 intracellular symbionts, 61 intrapediceloid, 148 328 Index intrinsic musculature, of heart, 61 intromittent organ. See aedeagus loscytus, 140 Irbisia, 34, 177, 179, 180 Irochrotus, 58 Ischnodemus, 253, 255 Ischnorhynchinae, 257 Isoderminae, 39, 212 Isodermus, 212, 2\3; planus, 209 Isometoparia, 177 Isometopinae, 177, 179 Isometopini, 177 Issidomimini, 77 Issidomimus, 77 Ithamar, 283; hawaiiensis, 281 Jaczewski, Tadeusz L., 9 Jadera, 57, 282, 283; haematoloma, 282 Jakovlev, Vasiliy E., 9 Jalysus, 248; spinosus, 32, 248; wickhami, 32, 248 Japetus, 269 Jeannel, Rene, 9 Joppeicidae, 40, 164 Joppeicus, 164, 165', paradoxus, 164, 165 juga. See mandibular plates juvenile hormone, 61 kairomones, 57 Keltonia, 178 killing bottles, 17 Kiritshenko, Alexandr Nikolayevich, 9 Kirkaldy, George Willis, 9 kissing bugs. See Triatominae Kleidocerys, 53, 257, 263 Knight, Harry Hazelton, 10 Kodormus, 152, 155 Koelreuteria elegans, 282; paniculam, 282 Kumaressa, 212; scutellata, 209 Kvamula, 75, 77; coccinelloides. 77 Labiatae, 264 labium, 41, 121; swollen, 237 Labops, 34, 179 laboratory animals, 263, 271; bugs as, 159 labrum, 41 Laccocorinae. 125 Laccocoris. 52; hoogstraali, 109 Laccophorella, 216 Laccophorellini, 216 Laccotrephes, 109. 114, 116 lace bugs. See Tingidae lacerate-flush feeding, 20 Lagenaria, 33 Lamiaceae, 224 Lampracanthia, 140 Lanopini, 216 Lantana, 37, 184, large milkweed bug, 263 Largidae, 58, 268 Largus rufipennis, 268 Larix, 213 larval organ, 135, 138, 140 Lasiella, 190 Lasiochilidae, 190 Lasiochilus, 190, \9\-,fusculus, \9\',paHi- dulus, 191 Lasiocolpus, 190 lateral gland, 55 laterosternites, 49 laterotergites, 77; dorsal, 49. 190; inner. 49; ventral, 49 Lathrovetia, 101 Latimbini, 277 Latreille, Pierre-Andre, 1. 10 Lalrocimex, 201 Latrocimicinae, 201 lawn grasses, 33 leaf-curl disease, 35 leaf-footed bugs. See Coreidae legs, 45, 47 legumes, 274, 278 Leguminosae, 278 Leistarchini, 156 Lentia, 156; corcovadensis, 156 Leotichiini, 144 Leotichius, 134, l45-,shiva, 135 Lepidoptera: as predators of eggs, 32; larvae as prey, 233 Lepionysiini, 257 Leptocimex duplicatus, 200 Leptocoris, 282, 283; hexophthalmus, 36 Leptocorisa, 32, 273; acuta. 22, 32, 273, 274; chinensis, 32, 273; oratorios, 32 Leptocorisinae, 273 Leptocorisini, 273 Leptodema, 157 Leptoglossus, 40, 58, 278, 279; ashmeadi, 278; clypealis, 33; corculus, 33, 278; fulvicornis, 278; gonagra, 33, 278; occidentalis, 33; phyllopus, 33, 278 Leptonannus, 77 Leptophya capitaia, 183 Leptopodidae, 143 Leptopodinae, 144 Leptopodini, 145 Leptopodomorpha, 46,48, 134, 136 Leptopterna, 34 Leptopus, 145; marmoratus, 144 Leptosalda, 145: chiapensis. 143, 144 Leptosaldinae, 143 Leptoscelidini, 277 Leston, Dennis, 10 Lestonia grossi, 228; haustorifera, 227, 228 Lestoniidae, 39, 40, 227 Lestonocorini, 232 Lethaeini, 261 Lethierry, Lucien, 10 Lethocerinae, 113 Lethocerus, 113, \ \4:griseus, \ \3:maxi- mus, 107; niloticus, 113 Leucophoroptera, 178, 180 Leucophoropterini, 178 Libyaspls Group, 237, 238 life histories, in Miridae, 179 light traps, 18 Ligyrocoris diffusus, 52 Lilliputocorini, 262 Limacodidae, 35 Limnobates, 97 Limnobatodes, 87, 95, 96; paradoxus, 95 Limnobatodinae, 95 Limnocorinae, 125 Limnocoris, 125 Limnogeton . 45, 110-114 Limnogonus. 105 Limnonabis, 190 Limnoporus, 105; canaliculatus. 25 Lindberg, Hakan, 10 Linnaeus, Carl, 10 Linshcosteus, 159 Liorhyssus hyalinus, 36, 281 Lipogomphus, 90, 92 Lipokophila. 48, 147. 192: chinai, 192, 193: eberhardi, 56, 193 Lippomanus, 197 Litadeini, 182 litchi, 36 literature sources, 14 littoral zone, as habitat, 141 Lizarda, 158 lizards, as predators, 28 Llaimocorini, 212 Llaimocoris penal, 212 Lomagostus jeanneli, 69 Lophosculus, 154 Lopidea, 27, 28, 178; instabile, 27; nigridia. 27 Lopodytes, 157 lora. See maxillary plates Loricula. 162, 163: pilosella. 163: pselaphi- formis, 163, 164 low-frequency vibrations, 160 Lucerocoris brunneus, 256 Luffa, 33 Lund University Museum of Zoology and Entomology, 16 Lupinus caudatus, 27 Lyctocoridae, 194 Lyctocoris, 195; beneficus, 195; campestris, 194, 195; menieri, 194; nidicola, 195; luberosus, 194 Lygaeidae. 28, 33, 39, 40, 58, 251,253 Lygaeinae, 257, 263 Lygaeospilus, 257 Lygaeus. 27; equestris. 28, 282 Lygocoris, 177 Lygus. 34, 42, 57, 177, 179; elisus, 34; hes- perus. 34; lineolaris, 34; pratettsis, 34; rugulipennis, 34; spp., as predators, 34 Lyrnessus, 274 m chromosome. See supernumerary chro¬ mosomes Macaranga roxburghi, 240 Maccevethini, 282 Macchiademus diplopterus, 34 Machadonannus ocellatus, 81 Macrocephalinae, 46 Macrocephalini, 154 Macrocephalus notatus, 160 Macrocytus brunneus, 223 Macropes femoralis, 254 macroptery, 23, 264 Macrosaldula, 140 macrotrichia, 67, 84 Macrovelia, 93, 94; hornii, 85, 94 Index 329 Macroveliidae, 93 Madagascar. 39 Madeovelia. 88 Madeoveliinae, 39. 88 Maevius indecorus. 279. 280 Magnocellus transvaalensis, 169 Mahea, 216 main salivary gland. 59 Malcidae. 34. 40. 264. 266 Malcinae. 266 Malcus. 264. 266 Malcusflavidipes. 265. 266:furcaius. 265; japonicus. 265. 266 male genitalia. 50; asymmetrical. 78. 82. 169.172 Mallochiola. 163 Malpighian tubules. 59. 60. 267 Malvaceae. 240; as hosts. 28 Malvales. 270 Manangocorinae. 157 Mamngocoris horridus. 157 mandibles. See mandibular stylets mandibular levers. 41 mandibular plates. 41; produced. 166. 167. 237 mandibular stylets. 41.43; coiled. 208. 212.214 Manocoreoini, 277 Mantodea. as predators. 28 Maoristolinae. 69 Maoristolus, 70 marginal riffles, as habitats, 125 marine bugs. See Aepophilidae marine habitats, 105 marsh treaders. See Hydrometridae Martarega mexicana, 128 Martiniola, 145 maternal care. 184, 217. 224. 233, 236. 240 mating iiosition. in Joppeicidae, 165; in Saldidae. 139 mating ritual, 263 Matsuda. Ryuichi, 10 maxillae. See maxillary stylets maxillary glands, 60 maxillary levers. 41 maxillary plates, 41 maxillary stvlets, 41.43; coiled, 208, 212, 214 Mcaleelta. 266. 267 mechanoreceptors, 53 Mecideini, 232 Mecistorhinus tripterus. 35 Mecistoscelini, 177 Mecocnemini, 277 media, 43, 44 medial furrow, 43 Medicago, 34 mediotergites, 49 Medocostes. 184-186;/esM/it, 184-186 Medocostidae, 40, 184 Megaderma spasmae, 203 Megadermatidae, 204 Megalonotini, 262 Megalotomus, 274; quinquespinosus. 272 Megarididae, 40, 228 Megaris, 229; hemisphaerica, 228; taevicol- lis. 228; rmjusculus. 229; pueriohcensL';. 229; rotunda. 228: semiamicta. 229 Megenicocephalinae. 73 Megenicocephalus, 70. 73 Megymeninae, 225 Meaymenini, 225 Megymenum. 225, 226; gracilicorne. 59; iiuulare. 226 Melaneryihrus. 257 membrane, 44; deciduous, 212. 213 Mendanocoris, 158 meniscus, ascension of. 102 Meranoplus mucronaius. 238 Merocorini, 278 Merocoris. 278 Meropachydinae, 278 Meropachydini. 278 Merragata, 90, 92; brunnea, 91 Menila maiayensis, 174 mesoscutellum. See scutellum mesoscutum, 43 Mesosepis papua, 155 mesospermalege, 188 mesosternal carina. 215, 217 mesosternum, 43 Mesovelio, 88, 90; amoerm, 90: furcata, 64. 90; mulsanti. 89, 90 Mesoveliidae, 88 Mesoveliinae, 88 Mesoveloidea, 88, 90; williamsi. 89 Metacanthinae, 248 metamorphosis, 61 Metapterini, 156 metastemum, 43; produced, 232 Metatropiphorus, 186 metepisternura, 43 Metrargini, 257 Metrobates, 105; trux, 86 Metrocorini, 105 Metrocoris, 105 Mezira, 48; reducia. 213; tremulae, 211 Mezirinae, 212 Micrelytra, 224, fossularum, 273 Micrelytrinae, 274 Microchiroptera, 204 Micronecta, 121 Micronectinae, 121 Microphysa, 162 Microphysidae, 161 microptery, 23 micropylar canals, 63 micropylar processes, 63, 64, 205 micropyles, 63, 278 Microtomus, 156 microtrichia, 84 Microvelia, 25, 37, 92, 99, 101, 102; di- luta, \02: douglasi, 25: longipes, 106; pulchelia, 37, 100, reticulata, 100 Microveliinae, 99 Microveliini, 99 Mictini, 277 Mictis profana, 33 midgut, 59, 60; discontinuous, 60 Miespa, 266 migration, 270 Miller, Norman Cecil Egerton, 10 millet, 35 millipedes, as prey. 160 mimicry. 27. 270; aggressive, 27. 28; of ants, 29; Batesian, 27. 28. 263; of beetles, 30; Mertensian, 27; Mullerian, 27. 28. 263; in Rhopalidae. 282; of sub¬ strate. 30; Wasmannian, 27; of wasps, 29. 177 Mimocoris rugicoUis, 29 minute pirate bugs. See Anthocoridae Miraradus, 210 Miridae. 27. 34, 45, 46, 49, 169. 172 Mirinae, 177 Mirini, 177 Mirperus jaculus, 32 mites: as prey, 195; predators of. 32 Mixotrephes hoberlandli, 132 Mniovelia. 88, 90; kuscheli. 88. 89 Molossidae, 204 Monalocoris. 122: americanus. 175 Monaloniina, 34, 48. 176 Monalonion. 34. 176, 179 Monocotoledoneae, 255 monographs, 15 Mononyx, 117 monophagy, 179 monophyletic groups. 38 monophyly. 5 Monosteira unicostata, 36 Montandon, Arnold Lucien. 10 Momandoniola maraquesi, 32 Monteithocoris hirsutus, 251 Monteithostolini, 73 Monteithosiolus genitalis, 69, 70. 73 morphology, 42 Morus bombycis, 266 mosquito larvae, as prey, 130 mosquitoes, 36, 37 mosses, 184 mounting: card, 18; pin, 18; point, 18; slide, 18 mouthparts, 43; in Aradidae, 211; in Termi- taphididae, 211 Muatianvuaia, 77; barrosmachadoi, 76 multivoltine life cycle, 179 Murgantia histrionica, 35 Murphyanella, 70; aliquantula, 70 Murphyanellinae, 70 Musaceae, 179 Musee Royal de I’Afrique Centrale. 16 Museo Argentine de Ciencias Naturales “Bernardino Rivadavia,” 16 Museum National d’Histoire Naturelle, Paris, 16 museums, 15 Musgraveia sulciventris, 36, 241 Mutillidae, as models, 269 mycetomes, 61 mycophagy, 213, 215, 237 Myiomma, 177 Myiommaria, 177 Myodocha serripes, 254 Myodochini, 262 myrmecomorphic body shape, 29 myrmecomorphic coloration, 29 myrmecomorphic nymphs, 274 330 Index myrmecomorphy. 28, 179, 250, 255, 263, 269. 212,. 21‘i: sexual, 29 Myrmecophyes oregonensis, 171 Myrmedohia. 163; exilis. 163, 164 Myrmica, 217; ruginodis, 217 Mynnus miriformis. 281 Myrocheini, 232 Myrtaceae, 229 Nabicula propirujua. 188 Nabidae,40, 186 Nabinae, 187 Nabini, 188 Nabis. dl, 54. 188, lS9,americofeiiis. 187; ferus. 187; limbalus. 64 Naboidea, 48 Naniella, 173 Nasutitermes exitiosus. 160 Natalicolinae, 242 natatorial legs, 45 National Collection of Insects, Plant Pro¬ tection Research Institute, Pretoria, 16 National Museum of Natural History, Rio de Janeiro, 16 National Museum of Natural History, Smithsonian Institution, 15 National Museum, Prague, 16 National Science Museum (Natural His¬ tory!, Tokyo, 16 Natural History Museum, Budapest, 16 Natural History Museum, London, 16 Naturhistorisches Museum, Vienna, 16 Naucoridae, 23, 45, 124 Naucorinae, 125 Naucoris cimicoides, 126 Neacoryphus, 263; bicrucis, 27, 263 Neadenocoris, 212 Nearctic Region, 38 neck, 41 Neella, 179 negro bugs. See Cydnidae Neides. 248; muticus, 248 nematoceran larvae, as prey, 122 Nematopodini, 111 Neoalardus typicus, 100 Neocentrocnemis signoreli, 155 Neochauliops. 266 Neocratoplatys salvazai, 237 Neogerris parvulus, 104 Neogorpis, 188 Neolocoptiris. 154 Neoncylocotis, 12 Neopamera bilobaia, 29 Neoplea, \30, slriola, 130 neoteny, 24 Neotimasius, 92 Neotrephes, \3l; usingeri, 132 Neotrephinae, 131 Neotropical Region, 38 Nepa, 115, \ l6,cinerea, 115, 116 nephrocytes, 61 Nepidae, 24, 45, 55, 116 Nepinae, 116 Nepini, 116 Nepomorpha, 41,46, 48, 107, 110 Nereivelia, 88. 90 Nerthra, 30. 117, \\i:amplicotlis. 117; amulipes. Wl'.hungerfordi. 117 nervous system. 61,62 Neseis, 258 Nesenicocephahn:. 71 nests, as habitats. 160 nets, 17 Neuroclenus pseudonymus. 213 neurosecretory system. See endocrine glands Nezara. 233; viridula, 35. 64 Nichomachini, 178 Nichomachus. 178 Nidicola, 199 Niesthreini, 282 Ninini, 256 Noliphini, 273 Nothochromini, 255 Nothochromus maoricus, 255 Nothofagus, 251 Notonecta, 53. 128, \29, glauca, 129; undulata, 44 Notonectidae. 45, 127 Notonectinae, 128 Noualhieridia, 216 Nycteridae, 204 nymphal organ, 137 nymphal thoracic glands, 60 Nymphocorinae, 70 Nymphocoris, 70; hilli, 70; maoricus. 70 nymphs: first-instar, 46; myrmecomotphic. 29; of Reduviidae, 160 Nysiini, 257 Nysius, 258, 263; ericae, 34; raphanus. 34; thymi, 254; vinitor, 34 Oaxacacoris guadalajara, 174 occipital apodemes, 100, 101, 105 ocean, as habitat, 106 oceanic distributions, 40 Oceanides, 258 ocelli, 41 OcelloveUa,9%. lOl; germari, 100 Ocelloveliinae, 101 Ochteridae, 45 Ochterus, 109, IW, barbert, 119, coffer. 10%; marginalus, 118, 119; sevchellensis. 119 ocular seta, 56 Odoniellina, 34, 48, 176 Odontoscelis, 58 Odontotarsinae, 239 Odontotarsus, 36 odors, 160 Oebalus pugnax. 35 Oeciacus, 201; vicarius, 33 Oedancala dorsalis, 254 Oiovelia, 101; spumicola, 101 oligophagy, 179 olives, 32 Omania, 141; coleoptrata, 135, 142 Omaniidae, 141 Ommatides, 82 ommatidia, 41 omphalus, 199 Oncacontias, 216 Oncomerinae. 242 Oncomerini, 242 Oncopeltus. 22, 27, 28, 263; cingulifer. 22: fasciatus. 22. 24. 28. 43. 64. 254. 263; sandarachatus. 22 Oncylocotis, 71-73; curculio. 48; tasmani- cus. 73 Ontiscini, 256 Onychotrechus. 105. 106; r/te.venor, 104 Onymocoris. 161; hackeri. 169; izzardi. 166 operculum, 62. 64 optic lobes, 61,62 Opuntia. 179 Oravelia, 93, 94;pege, 85. 94 Orchidaceae, 179 Orectoderus, 29; obliquus, 170 Oriental Region. 38 Oriini, 46, 197 0ms. 32, 197-199 Orsillinae. 40, 257, 263 Orsillini, 257 Orsillus depressus. 254 Orrhophrys, 138. 140; pygmaeum. 140 Orthotylinae, 40, 111 Orthotylini, 178 Oshanin, Vasiliy F., 10 osmoregulation, 184 ostia, of heart. 61 ostracods, as food. 130 ovaries, 165 ovariole numbers, 3, 51 overwintering, 184 ovipositor, 49, 50; laciniate. 49; platelike, 50 ovoviviparous species, 263 Oxycareninae, 58, 258 Oxvcarenus, 34, 258, 264; hialinipennis, 24. 254 Oxythyreus cyiindricornis, 151 Ozophora, 251; singularis, 253 Ozophorinl, 40, 262 Pachycoleus, 78 Pachycorinae, 239 Pachycoris lorridus, 238, 240 Pachygrontha. 258 Pachygronthinae. 30. 258 Pachygronthini, 258, 263 Pachynomidae, 39, 41, 148, 149 Pachynomlnae, 149 Pachynomus, 148, 149;picipes, 149 Pagasa, 1%%; luticeps, 186 pagiopodous coxae, 45 pala, 119, 121 Palaucorina, 176 Palaucoris unguideniatus, 175 Palearctic Region, 38 Paleotropical, 40 palm bugs. See Xylastodorinae palms, 35, 169 Pameridea, 176 Pamphantini, 29, 255 Pandanaceae, 179 Pandanus, 158 Pantochlora, 232 Index 331 Parabryocoropsis, 171 Paracatisiopsis, 210 Paracalisius, 210 Paracimex, 201 Paracopium, 184 Paradacerla, 29 Paradindymus madagascariensis. 271 paragenital glands. See ventral abdominal glands Paragonatas divergens, 54 Paratosalda, 140 parameres, 49; articulated. 218; as copula- tory organs, 195, 199. 202 parandria, 138, 139, 208 Paranisops, 128, 129 paranoia, 183 Paraphrynovelia, 93; brincki, 93 Paraphrynoveliidae, 40. 92 paraphyletic group, Lygaeidae as. 252 Parapiesma, 266 Paraplea, 130 Pararachnocoris, 186 parasitoids, 57 parastigmal pits, 148, 149, 186-189 Parastrachia, 220, 222. 224,japoneiisis. 224 Parastrachiinae, 222 Paratrephes, 131 Paravetia, 85, 101; rescens, 48, 52 parempodia, 46, 48, 98, 109. 136. 137. 141, 168, 175, 178; asymmetrical. 176; fleshy, 171, 177; increased numbers of, 107; reduced, 135, 192 parental care. See maternal care Paropsis, 218 pars stridens. See stridulitrum Parshley, Howard Madison, 11 parthenogenetic species, 90, 179 Paskia, 131 Patalochirus, 158 Patapius, 145; spinoius, 144. 145; thoienr/r, 144 peaches, 34 peanuts, 34 pears, 34, 36 pedicel, 41, 148, 149 peg plates, 84 Peirates, 157 Peiratinae, 40, 157 Pelocorisfemoratus. \25'. shoshone. 124 Pelogonus, 118 Pendergrast’s organs, 215, 217 Pentacora, 140; grossi, 139; signoreli. 138-140 Pentatomidae, 35, 58, 229, 230 Pentatominae, 232 Pentatomini, 232 Pentatomoidea, 41,43 Pentatomomorpha, 5, 43, 46, 48, 57. 60, 205 Pephricus paradoxus, 276 Peregrinator biannulipes, 36 Peridontopyge spinosissima, 160 Perillus, 35; bioculatus, 35, 233 peritreme, 43, 57; produced. 247, 248 Peritropis, 172 Perittopinae, 101 Perittopus, 100, 101 Perkinsiella saccharicida, 34 Peruda, 51 pet food, Corixidae as, 122 Petascelidini, 277 Phaenacantha, 33, 250; australiae, 250; saccharicida. 250; saileri. 250 phaliandrium, 73 Phallopirates, 73 Phallopiratinae, 73 phallotheca (phallosoma), 49. 50 Pharnaceum aurantium, 244 Phaseolus, 32 Phasmosomini, 262 Phasmosomus, 253; araxis, 253 Phatnomini, 182 pheromones, 57 Phimophorinae, 158 Phimophorus, 158 Phloea, 235 Phloeidae, 40, 41,234 phloem vessels, 20 Phloeophana. 234, 235; longirostrits, 235. 236 Phonoctonousfasciatus, 63 Phonoclonus, 28, 36, 270 phoretic male, 102 Phorticini, 188 Phorticus, 188 photoperiod, 25 Phrynovella, 88, 90 Phthia picta, 33 Phthirocorinae, 73 Phthirocorini, 73 Phthirocoris subantarcticus, 73 Phylinae, 178 Phylini, 178 Phyllocephalinae, 60, 232 Phyllomorphini, 277 phylogenetic relationships: of Cimico- morpha, 146; of Gerromorpha, 84; of infraorders, 5; of Leptopodomorpha, 134; of Nepomorpha, 110 Phymaia, 47, 52, 58, 154; crassipes, 160; erosa, \55;pennsylvanica. 30, 155; pennsylvanica americana, 160 Phymatinae, 45, 58, 154, 160 phymatine complex, 154 Phymatini, 154 physical gill, 53, 137 physiology, 263 Physoderes, 158 Physoderinae, 40, 158 Physopelta famelica, 268 Phytelephas, 169 Phytocoris, 22, 177, 179; calli. 174; neglec- tus, 22; nobilis, 22; populi, 170 phytophagy, 20 Piesma, 266, 267; capiiatum, 35, 267; cinereum, 35. 267; maculatum, 267; quadratum, 35, 53, 267 Piesmatidae, 35, 266 Piesmatinae, 266 Piezoderus hybneri, 35 Piezosternini, 242 Piezosternum, 242; calidium, 242; subula- tum, 242 Pilophorini, 178 Pilophorus, 46, 56. 18Q: kockensis. 171 pimentos. 33 Pinus, 33. 179, 213 Piper, 264 Pirates hybridus, 155 Pisilus, 157 pistachio, 33 pit, at base of claval commissure. 128 pitfall traps, 18 pit organs, 52, 101 plant bugs. See Miridae plant pathogens, 35 plant vascular system, feeding in. 233, 278 plastron, 137 plastron respiration, 125-127, 136 Platanus, 36, 263 Plataspidae, 40, 58, 236 plates, 52 Platygerris, 104 Platylygus, 177, 179 Platytatini, 242 Plea, \'i0,aiomaria. 130; minurissi/tia. 130 plectrum, 51 Pleidae, 45 Plinthisini. 39, 262 Ptinihisusflindersi. 253 Plochiocoris, 190 Ploiariini, 156 Plokiophila, 192; cubana, 192 Plokiophilidae, 46. 59, 190, 192 Plokiophilinae, 39, 192 Plokiophiloides, 192; asolen, 193 Poaceae, 255 pod-sucking bugs, 32 Podisus, 35. 57; maculiventris, 233 Podopinae, 232 Poeanlius, 28 Poecilocoris, 240 Poecilometis eximus, 229 Poisson, Raymond A., 11 pollen, 179, 184, 199 Polychisme ferruginosus, 255 Polychismini. 255 Polyctenes. 202;.molossus. 203 Polyctenidae, 46, 202 Polycteninae, 202 polyphagy, 179 Polytoxus, 158 Pompilidae, as models, 274 pond skaters. See Gerridae Poppius, Robert Berth, 11 population density, 25 pore-bearing organ (pore-bearins plate). 279-281 postclypeus, 41 postcubitus, 44 postocular lobe, 67 Potamobates, 105 Poiamocoris, 123; nieseri, 123; pan us. 123 Potamomelra, 105 potatoes, 34 poultry bug, 33 predation, 21 332 Index predators, 217; learning by, 29; visual, 159 predatory habits, in Asopinae. 231 prenatal care ,213 prepedicellite. 41, 150, 18S pressure receptors, 125 pretarsus, 46, 48. 109, 135, 175; in Miri- dae, 178 prey location, 106 prickly pear cactus, 33 Primicimex. 201 Primicimicinae, 201 Primierus quadrispinosKS. 261 Prionogastrini. 242 Prionotylini, 277 Procamptini, 277, 278 Procamptus segrex, 278 processus corial. See stub processus gonopori. See acus Procryphocricos perplexus. 24 proctiger, 49 procurrent nerve, 61 Pronotacantha. 56 pronotum; anterior lobe, 41; humeral angles, 43; posterior lobe, 43 Propicimex, 201 Prostemma, 47, 52, 54, 188, 189; guttuki. 187 Prostemmatinae, 41,45, 188 Prostemmatini, 188 prosternum, 43 Prosympiestinae, 212 Prosympiestini, 212 Prosympiestus, 212 Protea, 264 Proteaceae, 264 protective resemblance, 263 Protenor, 274 prothoracotropic hormone, 61 prothorax, 41 protocerebrum, 61,62 Psacasta, 58 Psallopinae, 178 Psallops, Yl%'. oculatus, 171 Psamminae, 258 Psammium, 258 pseudarolia. See pulvilli Pseudatomoscelis, 178 Pseudocetherini, 156 Pseudocnemodus canadensis. 51 pseudocompetition, 263 Pseudomonas, 61 Pseudophloeinae, 278 Pseudopsallus, 178—180; lattini, 176 pseudopulvilli, 171, 175, 176, 178. See also accessory parempodia Pseudosalduia, 140; chilenis. 138 pseudospermatheca. 50, 148-151, 155, 159, 181, 183 Pseudotheraptus wayi, 33 Pseudovelia, 101, 102 Psocoptera, 5 psocopteran webs, as habitats. 160 Psotilnus, 283 pterygopolymorphism, 188 Ptiiocerus ochraceus, 160 Ptilomera, 105, 106 Ptilomerinae, 39, 105 pulvilli, 46, 48. 167, 168. 171, 175, 176, 197, 205; absent, in Aradidae, 210 pumpkins, 33 Punctius. 148, \d9,alutaceus. 149 Puton, Jean-Baptiste Auguste, 11 Pycnoderes, 175. 179 pygmy backswimmers. See Pieidae pygopher. See genital capsule pygophore. See genital capsule Pylorgus, 257 pyloric valve, 60 pylorus, 59, 60 Pyrrhocoridae, 28, 35, 40, 58, 270 Pyrrhocoris aplerus, 58, 270, 271 Quercus, 35 quiet water, as habitat, 106 Quilnus, 210 radial sector, 44 radius, 43, 44 Ranatra, dl, 51, 108, 115, \16>, compressi collis, liS.drakei, 115 Ranatrinae, 116 Ranatrini, 116 Raniovius, 21, 180 rape seed, 34 raptorial legs, 45. 67, 107 Rasahus, \S1 ■, sulcicollis, 155 rectal gland (rectal pad), 60 rectum, 60; eversible, 179 recurrent nerve, 61 Reduviidae, 28, 31,35, 49, 150, 151 Reduviinae, 158 Reduvioidea, 48 Reduvius, 148, }58;personatus, 3\, 158 reel system, of aedeagus, in Saldidae, 137 relict distributions, 40 remigium, 44 respiration, 114 respiratory siphon, 116 Resthenini, 177 Restionaceae, 256 reticulate body surface, 266 retractor tendon, 46 Reuter. Odo Moranal, 2,3, 11 Reuteroscopus, 178 Rhagadotarsinae. 105 Rhagadotarsus, 105, kraepelini, 104 Rhagovelia, 48, 52, 54, 101, 102; disiincia. 86 Rhagoveliinae, 101 Rhamphocoris, 188 Rhaphidosoma, 157 Rhaphidosomini, 157 Rheumatobates, 102, 105, 106; meinerti. 104 Rhinacloa, nS.forticornis, 179 Rhinocoris, 64 Rhinocylapus, 179 Rhinolophidae. 204 Rhodainiella, 157 Rhodniini, 159 Rhodnius, 58; proiixus, 57. 159 Rhododendron, 36, 263 Rhopalidae, 36. 58. 281,282 Rhopalimorpha. 216 Rhopalinae, 282 Rhopalini, 282 Rhopahmnrpha. 217 Rhynchota, 1 Rhyngota. See Rhynchota Rhynocoris. 28. 1.57 Rhyparochrominae, 29, 45, 258. 259 Rhyparochromini. 262 Rhytocoris, 53 rice, 32, 35. 232 rice planthoppers. 37 riffle bugs. See Veliidae ring gland. 184 riparian habitats, 116, 118 ripple communication, 106 Riptortus, 32, 273 rodent nests, as habitats. 195 rostral groove, 91 rotatory coxae, 45 rowing. 45 Roystonea regia, 169 Ruckes, Herbert, 11 Ruckesona. 246; vitrella. 246 Rupisalda, 140 rushes, 263 Rutaceae, 36 rutherglen bug, 34 Sagocorini, 125 Sagriva, 226 Sahlbergelia singular is, 34 Saica, 158 Saicinae, 158 Saicini, 158 Saileriola, 246; sandakanensis, 246 Saileriolinae, 245 Saida, 138; linoralis, 140 Saldidae, 39,45,46,58. 140 Saldinae, 140 Saldini, 140 Saldoida. 140 Saldoidini. 140 Saldolepta. 145; kistnerorum, 143, 144 Saldula. 140; denlulara. 139;fucicola. 64. 139; laticoUis. 139-141; orthochila. 141; pallipes. 135; sibiricola. 139 Salduncula, 140 Saldunculini, 140 salivary canal, 41,43 salivary glands, 59; function, 59; structure, 59 salivary pump, 59 saltatorial legs. 45 Saiyavata. 158; variegara. 160 Salyavatinae, 158 Sandaliorrhyncha, 120 ■ Sangarius. 217; paradoxus. 217 ’ sap feeding, 60, 263 Sapindus saponaria var. drummondii. 282 Saturniomirini, 177 savoy virus, 35 saxicolous, 141 Saxicoris. 258; verrucosus, 258 Say, Thomas, 11 Index 333 scale insects. 179 scanning electron microscopy. 19 scape. 41 Scaptocorinac. 774 Sea;)local’is. 4?. 4o; cdsuincii.s. 33. 224. dirt’it’cn.s. 33. 771 scioengcrs. in spider ueb,^. ISO Scclionidac. 57. 233 scent-gland channel. 5b scent-gland fluids: composition. 57; fungistatic role. 57: shooting. 236 scent-gland opening. 281; in coxal cavities. 287: obsolete. 781 scent glands: adult. 57: function. 57; meta- thoracic. 55; nvniphal. 56; structure. 56. 57 .scentless plant bugs. Sec Rhopalidae Schnffneria. 180 Schiodte. 3 Schizopwra. 82 Schizopteridae. 80. 82 Schizopterinae. 87 Schoepfia jasminodora. 224 Schouteden. Henri. 11 Sciocorini, 232 Sciocoris micropln/udmiis. 231 scolopidia. 53 Scolopini. 58. 198 scolopophorous organs (scolopidial organs). 53. 107 Scoloposcelis flaviconus. 198 Scotiiiophara. 35. 232; lurida. 35 Scoiomedes. 161. 1^4: alieinis. 162: bor- neensis. 162 Scott, John, 11 Scutellera, 240 bcutelleridae, 36, 58. 238, 239 .scutellerinae, 239 Scutellerini, 239 scutellum, 43, 232; enlarged. 218, 219. 228, 231,232. 236, 238. 240 Seabramnnnus immitator. 81 sealing bar, 62 second valifer, 50 second valvula, 50 secondary fecundation canal. 84 secretory cells, subhypodermal. 183 sedges. 263 seed bugs. See Lygaeidae seeds, feeding on, 28. 29, 233. 244. 263, 270,281 Sehirinae, 224 Sehirus. 224; bicolor. 224; cinctus, 223. 224; cinctus albonotatus. 221 Seidenstiicker, Gustav. 12 Setnangananus mints. 81 semiaquatic bugs. See Gerromorpha seminal conceptacles. 195, 199 Semium, 173 Senecio. 263; smallii. 27 sensory patches, on male abdomen. 231 Sepinini, 242 sequences, rDNA, 5 Serbana, 30; borneensis. 232 Serbaninae, 232 Serenthiinae, 182 Serinethinae, 282 Serjania brachycarpa. 282 Serville. Jean-Guillaume Audinet. 1. 12 Severiniella, 237 sex chromosomes. 63 sexual dimorphism. 163; of wings. 213 shell. See chorion shield bugs. See Scutelleridae shore bugs. See Saldidae sieve plate. See pore-bearing organ Sigara. \2\ : sahibergi. 122 Signorct. Victor. 12 Sinea diadema, 28 Sinotagini, 277 Sirthenea. 157 Slaterobius, 29 Slaierocoris. 178 small grains, 34 small water striders. See Veliidae snail predator, 114 Snow Entomological Museum. University of Kansas. 16 soapberry bug, 282 soapberry tree, 282 socket glands, 58 Socraiea montana, 169 Solanaceae, 37, 266 Solanum, 34 Solenopsis. 29 Solidago, 30 soun bug, 36 sound production, 51; in Phymata. 160 soybeans, 35 Spalacocoris philippinensis. 256 Spartocerini, 277 Spathophorini, 278 species recognition, 106 Speovelia, 88.90 sperm, 63, 140; storage of, 146 spermatheca, 50, 205, 229, 231; absent, in Idiostolidae. 251; invaginated. 232 spermathecal pump, 100 spermathecal tube, 89, 91 spermatic pocket. 195. 199 spermatophore, 63 spermatozoa, 239 Sphaeridopinae, 158 Sphaeridops, 158 Sphaerocorini, 239 Sphaerocorts annulus, 240 Sphagnum, 78, 92, 164 Sphedanolestes, 157 spider predators, 160 spiders, myrmecomorphic, 29 spider webs, as habitats. 160 Spilostethus, 27. 28, 34; pandurus. 34 spines, nymphal, 181.265 Spinola, Maxmillian, 12 spinose body, 278 spiracle cover, 108 spiracles, 43. 50, 108, 189; abdominal, 49 spiracuiar rosettes, 126, 127 Spiraea, 263 spongy fossa. See fossula spongiosa Squamocoris, 179 squash bug, 33 Stagonomus amoenus, 231 staining. 19 Stal. Carl. 2. 12 staphylinoidy. 23. 24. 163. 243 static sense organs. 55. 108. 111. 113-116 Siemmocrypta aniennata, 83 Stemmocryptidae. 82 Stenaptida. 257 Stenocephalidae. 58. 283 Stenocoris. 273; tipuloide.t. 272 Stenocorixa protrusa .121 Stenocorixinae. 121 Stenodema, 30 Stenodemini. 177 Stenolemus, 152. 160; lanipes, 160 Stenonabis, 188 Stenophyella. 258 Stenopirates, 12, IS Stenopodainae. 158 Stephanitis, 36. 179. 182; pvW, 36, 183; rhododendri, 35 sternal glands. See ventral glands Sternorrhyncha. 5 sternum 7 cleft, in female. 268 Slethoconus. 35. \19:frappai, 35; japoni- cus, 35 Sthenaridea australis■ 54. 175 sticky setae. See viscid setae Stilbocoris, 61. 188. 263 stilt bugs. See Berytidae stink bugs. See Pentatomidae Slolliafabricii, 59 stomach, 60 stomodeal nervous system (stomogastric nervous system). 61 stones, as habitat. 141 Stongylovelia, 99 strainer. See pore-bearing organ strawberries. 34 stretch receptors. 106 stridulation. 129. 224. 263; defensive, 160; by nymphs. 160 stridulatory structures: abdominal sternum- hind leg. 51. 52. 211; in Aradidae. 213; base of labium-femoral apex. 53; con- nexival margin-hind femur, 51, 101, 125; femoral ridge-coxal peg, 53: forecoxa- forecoxal cavity. 51; forewing edae-hind femur. 51.52.’l32. 140, 179; head- forefemur, 51; hypocostal lamina, 53; margin of pygophore, 53; maxillary plate, 119; maxillary plate-forefemur, 53, 121; metapleuron-middle femur, 51; metathoracic wing-abdomen, 53. 145, 224, 232, 244; propleuron-forefemur, 51; prosternal groove. 53, 151, 159, 166; tibial comb-labial prong, 51, 128, 129; underside of clavus. 53 stridulatory sulcus, 52 stridulitrum. 51 Siridulivelia. 51, 101. 102 strigil, 121. 122 Strombosoma, 224 Strongvlovelia, 99 stub, 162, 163, 171. 186 Stygnocorini, 39, 262 334 Index stylets. 41 stylet-sheath feeding, 20 styloids. See third valvulae Srytopomiris malayemis. 176 subcosta. 44 subesophageal ganglion. 62 subgenital plate. 50. 67. 74 submacroptery. 23 subrectal gland. 58. 156 sugar beets. 35 sugarcane, 33. 34. 250 sugar cane leafhopper. predator of. 34 supernumerary chromosomes. 64 supraesophageal ganglion. 61 Surinamellini. 177 swallows, as hosts, 201 swarming, 70, 73 sweating, 183 Swedish Museum of Natural History. 16 sweeping nets, 18 sweet potato, 33 swifts, as hosts, 201 swimming, inverted, 129. 130 swimming fan. 99, 101 sycamore lacebug, 36 Sycanus coUaris. 36 symbiont transmission, 61 symbionts, gut, 60. See also gastric caeca; intracellular symbionts; mycetomes sympathetic nervous system. See siomodcal nervous system Sympeplus, 258 Symphylax musiphthora. 250 syncytial bodies. 194 Systelloderes. 67. 73 Systelloderini, 73 Tachygerrini. 105 Tachygerris, 105 Tahitocoris cheesmanae, 233 Tanycricini, 125 Targarema stall. 253 Targaremini. 262 tarsi. 46, 47; cleft, 99; reduced. 224; swollen, 173. 176 Tarsotrechus. 105 Taylorilygus pallidulus. 34 tea, 34 Tectocoris, 58. 239; diophthalmus. 239, 240 Tegeini, 157 tegumentary glands, 58 Teleonemia scrupulosa, 37 Telmawmetra, 105 Teloleuca. 140 temporary habitats, 264 Tenagobia, 121 Tenagogonus, 105 Tengella radiata, 192 Tenodera ardifolia sinensis. 28 Teracriini. 258 Termatophylini. 177 Termitaphididae, 214 Termitaphis. 214 Termitaradus, 214, 215; guianae. 48, 56, 2\A\ jamaicensis. 215; panamensis. 211 termites: inquilines of, 213, 215; as prey. 1.56 territorial defense, 106; male. 278 Tessaratoma javanicu. 242: papillosa. 36. 57,242 Tessaratomidae, 36, 241 Tessaratominae. 242 Tessaratomini, 242 testes, 137, 196 testis follicle numbers, 3 Tetraphleps. 197; lalipennis. 196 Tetraripis. 101, 102 Tetroda histeroides. 232 Thaicoris. 266 Thaiocorinae, 266 Thalmini, 225 Thasus. 279; acuiangulus. 275, 279 Thaumamannia. 183; vanderdrifti. 183 Thaumastaneis montandoni. 269 Thaumastella. 243; aradoides, 243, 244; elizabethae, 243, 244; namaquensis. 243, 244 Thaumastellidae, 243 Thaumastocoridae, 165, 167 Thaumastocorinae, 49, 167 Thaumastocoris. \61. australicus. 169; hackeri. 167 theca. See phallotheca Themnocoris kinkalanus. 155 Themonocorini. 154 Theraneis. 269 Theseus modestus. 231 third valvulae, 49 thoracic ganglion, first, 61 thorax, modification of, !06 thornlike outgrowths. See grooved setae Thyanta calceata. 233 Thyreocoridae, 222 Thyreocorinae, 224 Thyreocoris. 224; scarabaeoides. 223. 224 Thysanoptera, predators of, 32 tibiae, 45; annulate. 202; spinose, 220 tibial appendix, 49. 167, 168 tibial comb, 47, 168 tibial extensor pendant sclerite. 46 tibial flexor sclerite, 46 Timahocoris. 70; paululus. 70 Timasius. 92; ventralis. 91 Tingidae, 36, 180. 182; as prey, 179 Tinginae. 182 Tingini, 182 Tingis ampliata. 183; cordu/', 183 Tinna wagneri. 36 toad bugs. See Gelastocoridae toads, as predators, 283 Totlius. 274 tomato stilt bug, 32 tomatoes, 32-35 Tonkuivelia. 101 Tornocrusus. 69; penai, 69 Torre-Bueno, Jose R., 12 torrential streams, as habitats, 126 torrents, as habitats, 106 Trachelium. 274 Transpacific, 40 tree holes, as habitats, 101 Trephotomas. 131 Trephotomasinae, 131 Trepobalcs. 105: laylori. 104 Trepobatinae. 105 Trctocorini. 212 Tretocoris. 212 Triatomo. 35, 159; nigromaculuta. 152; nibrofasciaia. 44. 155. 159 Triatominae, 40, 159. 160 Triatomini. 159 Trihelocephala. 159 Tribelocephalinae, 159 Tricentrus. 250 trichobothria: abdominal, 54. 148. 150. 161. 186, 188. 205. 227. 243. 244. 250. 252. 253. 265; antennal. 54, 148- 150. 155. 159; in Aphelonotinae, 149: cephalic. 54. 84. 95. 102, 134; femoral. 49. 54. 171, 178; in Miridae. 171. 178: in Nabidae. 54. 186; in Pachynomidae. 148, 150; in Pachynominae. 149; in Pen- tatomomorpha. 205; in Prostemmatinae. 188; in Reduviidae. 150. 159; scutellar, 54. 188; in Thaumastellidae. 243; in Velocipedidae, 161 Trichocentrus. 51 Trichocorixa reticulata. 120 trichome. abdominal. 160 Trichophora, 205. 263 Trichoielocera. 74 Trichotonanninae. 77 Trichotonannus. 77: dundo. 76. 79; oidipos. 77, 79 Tridemula pilosa. 155 Trigononius. 34 Trisecus armatus. 251; pictiis. 251.252 tritocerebrum, 61 trochalopodous coxae. 45 trochanters. 45 Trochopus. 101 Tropidotylus. 237 Trypanosoma cruzi. 159 tubercular sense organs. See corial glands Tullgren, A., 2, 4 tur pod bug. 33 Tylocrypnis egenus. 277 tylus. See clypeus tymbals, 53 ' tympanal organs. 53 Typha. 263, 264 Tyttlms. 178; mundulus. 34 Udeocorini, 39, 262 Uhler, Phillip Reese. 12 unguilractor plate, 109. 135 unique-headed bugs. See Enicocephalomor- pha University Zoological .Vluseum, Helsinki, 16 univoltine life cycle, 25, 264 unques. claws unquitractor plate. 46 uradenia, 198, 199 uradenies. See ventral abdominal glands Urentius hystricellus. 36 Urmcephala californica. 72 Index 335 Urolabida. 245 Urostylidae. 245 Urostylinae, 246 Usinger, Robert L.. 12 Valieriola. 135, 143. 145 valvifers: first, 49: second, 49 valvulae: first, 49; second. 49; third, 49 Van Duzee. Edward Payson, 12 Vanniiis. 172 vascular system, feeding in. 20 vectors. 159 Velia. 101, \(i2:caprai. 100 Veliidae. 25. 37, 45. 49. 98, 99 Veliinae. 101 Veliohebria, 101 Veliohebriini, 99 Veliometra. 95-97; schuhi, 96 Velocipeda. 161 Velocipedidae, 161 Veloidea. 101 velvet water bugs. See Hebridae velvety shore bugs. See Ochteridae Ventidiiis. 105 Vemocoris fischeri. 36 ventral abdominal glands. 58 ventral arolium, 48. 91.93, 96, 98, 100. 101. 109. 134, 138 ventral glands, 58. 151. 159 vermiform sland, 50, 146. 155, 159. 184- 187. 190' 191 Vernonia. 184 vescia. 49, 159 Vesciinae. 40, 159 Veseris. 158 vesica, 50 Vianaida. 183 Vianaidinae, 182 vicariance biogeographic model, 38 W/ga, 278 Villicrs, Andre, 13 Viola. 224 Visayanocorini, 158 viscid setae, 49. 158-160 viviparity, 204 Voles us. 158 Wagner. Eduard, 13 Walambianisops, 128 wasp mimics, 273 water boatmen. See Corixidae water bugs. See Nepomorpha water measurers. See Hydrometridae water scorpions. See Nepidae water slriders. See Gerridae water surface, as habitat, 21, 84 water treaders. See Mesoveliidae Waterhouseana, 178 waxy secretions, 184 web lovers. See Plokiophilidae webs: living in, 21; of spiders, as habitats. 188 Wechina. 167 western equine encephalitis, 33 wheat. 34-36 wherrymen. See Gerridae Whiieiella, 190; rosiralis. 191 wing coupling, 44, 69 wing dimorphism, sexual, 23 wing folding, 227, 228, 237 wing loss, 213 wing muscle histolysis, 270 wing polymorphism, 70; environmental determination, 24; genetic determination, 24, 25; hormomal control. 24; polygenic determination, 25 wing reduction. 23 wings. 43. 44: folding. 236. 237; reduced 277 Wollasioniella ferruginea. 196; ohesula. \96: punciam. 198 Wollasloniola. 197 Woodward. Thomas Emmanuel. 13 Wygodzinsky, Petr (Pedro) Wolfgang. 13 Wygodzinsh,ella, 156 Xenobates. 99 Xenoblissus lulzi, 253 Xenocylapus. 179 Xenogenus. 283 Xiphovelia. 99. 102 Xiphoveloidea. 99: major. 100 Xosa, 216 Xylastodonnae. 46. 48. 167 Xylastodoris. 167. 169; luieolus. 167 Xylocorini. 199 Xylocoris. \99:afer. 196: cacti. \96: galac tinus. 198 xyphus. 43 Ypsotingini. 182 Yucca, 179 Zelurus. 158 Zelus. 28. 157 Zetekella minuscula, 48 Zeiterstedt. J. W.. 13 Zoological Institute. St. Petersburg. 16 Zoological Museum, Copenhagen, 16 Zoological Museum, University of Ham¬ burg. 16 Zootermopsis nevadensis. 213 336 Index About the Authors O' O' 'W o O' O' ■ o O' O' 'O o o o o o o o' o O' o O' O' o 'O o Randall T. Schuh was born in Corvallis, Oregon, on May 11, 1943. He received his B.S. degree in business administration from Oregon State Univer¬ sity, Corvallis, his M.S. degree in entomology from Michigan State University, East Lansing, and his Ph.D. degree in entomology from the University of Connecticut. Storrs, under the direction of James A. Slater. His dissertation dealt with the systematics and phylogeny of Miridae from southern Africa. He is currently George Willett Curator of Entomology at the American Museum of Natural History, New York’, where he is responsible for one of the world’s largest collections of true bugs; Adjunct Professor, Department of Entomology, Cornell University, Ithaca, New York; and Adjunct Professor, Department of Biology, City College, City University of New York. His research is focused on the Miridae and Leptopodomorpha, as well as on the phylogenetic rela¬ tionships and historical biogeography of the Heteroptera. He has collected extensively in North America, South Africa, South America, and Malaya. James A, Slater was born in Belvidere, Illinois, on January 10, 1912. He received his B.A. and M.S. degrees in entomology from the University of Illinois, Urbana, maintaining as well a strong interest in snakes and other squamates. Under the direction of H. H. Knight, he earned his Ph.D. degree in entomology from Iowa State College, Ames, where he worked on the struc¬ ture and taxonomic value of female genitalia in the Miridae. As Professor of Biology, University of Connecticut, Storrs (1953-1988), and Research Asso¬ ciate, Department of Entomology, American Museum of Natural History, New York, he conducts research on the systematics of the Heteroptera and has advi¬ sed numerous M.S. and Ph.D. candidates. His primary interest has long been the Lygaeidae. He has collected in South Africa, Australia, and the Caribbean, and has published extensively on those faunas. v," ■ .w