"LIBRA OF THE' UNIVERSITY OF ILLINOIS 590.5 FI v.40 cop. 3 NATURAL H.'STORY SURVEY FT y '4- 00/3.3 A MONOGRAPH ON THE TERMITOPHILOUS STAPHYLINIDAE (COLEOPTERA) CHAKLKS H. SEEVERS FIELDIANA: ZOOLOCY VOLUMK / hy THE UMW OF WBICAGO NATURAL HISTORY MI- APRIL 9, 1957 MW S4 1957 NAT APR 13 i:37 FIELDIANA: ZOOLOGY A Continuation of the ZOOLOGICAL SERIES of FIELD MUSEUM OF NATURAL HISTORY VOLUME 40 CHICAGO NATURAL HISTORY MUSEUM CHICAGO, U.S.A. 1957 MAY 2 1957 ri A MONOGRAPH ON THE TERMITOPHILOUS STAPHYLINIDAE (COLEOPTERA) A MONOGRAPH ON THE TERMITOPHILOUS STAPHYLINIDAE (COLEOPTERA) CHARLES H. SEEVERS Research Associate, Division of Insects Professor of Zoology, Roosevelt University FIELDIANA: ZOOLOGY VOLUME 40 Published by CHICAGO NATURAL HISTORY MUSEUM APRIL 9, 1957 Editors AUSTIN L. RAND Chief Curator, Department of Zoology LILLIAN A. ROSS Associate Editor, Scientific Publications PRINTED IN THE UNITED STATES OF AMERICA BY CHICAGO NATURAL HISTORY MUSEUM PRESS Contents PAGE I. TERMITES AND TERMITOPHILES: GENERAL CONSIDERATIONS 9 Introduction 9 Acknowledgments 12 Organization of Termite Societies and Termite Evolution 13 General Remarks on the Biogeography of Termites 16 Termitophily in General 17 Wasmann's Classification of Termitophiles 18 Major Adaptations Associated with Termitophily 20 II. TERMITOPHILOUS STAPHYLINIDAE: PHYLOGENETIC AND BIOGEOGRAPHI- CAL CONSIDERATIONS 23 Termitophilous Staphylinidae Associated with Nasutitermitinae . . 24 Phylogeny and Evolution of the Nasutitermitinae 24 The Termitophilous Tribe Corotocini 25 Tribal Characteristics 26 Phylogenetic and Biogeographical Considerations 28 The Termitophilous Tribe Termitonannini 42 The Termitophilous Tribe Termitohospitini 48 Termitophilous Staphylinidae Associated with Macrotermitinae . . 51 The Macrotermitinae 51 The Termitophilous Tribe Termitopaediini 52 The Termitophilous Tribe Termitodiscini 54 Termitophilous Staphylinidae Associated with Rhinotermitidae . . 54 The Termitophilous Tribe Philotermitini 54 The Termitophilous Subfamily Trichopseniinae 54 III. TERMITOPHILOUS STAPHYLINIDAE: SYSTEMATIC REVISION 59 Material and Methods 59 The Staphylinidae; General Considerations 60 Subfamily Aleocharinae 61 Key to Tribes Comprised of Termitophilous Species 62 Tribe Corotocini, including Subtribes Timeparthenina, Termito- gastrina, Abrotelina, Termitocharina, and Corotocina . . 63 Tribe Termitonannini, including Subtribes Perinthina and Termitonannina . .164 8 CONTENTS PAGE Tribe Termitohospitini, including Subtribes Termitohospitina, Hetairotermitina, Termitospectrina, and Termitusina . . . 191 Tribe Termitopaediini 214 Tribe Myrmedoniini (s. lat.), including Subtribes Myrmedoniina, Feldina, Termitondina, and Termitozyrina 232 Tribe Pseudoperinthini 247 Tribe Athetini, including Subtribes Coptotermoeciina and Termi- totelina 248 Tribe Philotermitini 250 Tribe Termitodiscini 258 Subfamily Pygosteninae 265 Subfamily Trichopseniinae 266 Subfamily Tachyporinae 281 Subfamily Hypocyphtinae 282 IV. SYSTEMATIC CATALOGUES 285 A Classification of the Termitophilous Staphylinidae, Including Host Records 287 A List of Termites and Their Staphylinid Guests 302 Regional List of Termitophilous Staphylinidae 311 An Annotated List of Doubtfully Termitophilous Species 318 V. BIBLIOGRAPHY 321 VI. INDEX ... . 329 I. Termites and Termitophiles: General Considerations INTRODUCTION This monograph represents a departure from a majority of systematic revisional works in that ecological relationships rather than systematic position determined the species to be included. The systematic section is concerned with revisions of groups of staphylinid beetles having in common the fact that they are ob- ligatory inhabitants of termite societies. The central theme is the evolutionary history of one mode of life within a very large family of organisms, the Staphylinidae. My studies on termitophilous insects began about twenty years ago, when Dr. Alfred E. Emerson very generously provided me with a substantial collection accumulated during his termite investi- gations in South America. This collection was augmented from a number of sources until a systematic revision of the world's termitophilous Staphylinidae became feasible. Even though the termite nests of extensive tropical areas have not been thoroughly explored and many genera and species are still to be discovered, a synthesis of the substantial body of knowledge relating to this group of insects and a discussion of evolutionary implications seemed justifiable. The objectives of these studies may be summarized as follows: taxonomic revisions of the various categories of termitophilous Sta- phylinidae; presentation of a list of host termites; formulation of hypothetical phylogenies for the larger groups of termitophilous Staphylinidae; and, lastly, a synthesis of the taxonomic, phylo- genetic, biogeographical, and host data for each group in an attempt to elucidate its evolutionary history. These goals have frequently been somewhat elusive owing to the difficulty of obtaining critical specimens and the unsatisfactory status of staphylinid classification. As the studies progressed it became apparent that the termitophilous Staphylinidae are exceptionally satisfactory for certain evolutionary studies, particularly when interpreted as part of relatively stable interspecies relationships, i.e., the relationships of termites and their 10 FIELDIANA: ZOOLOGY, VOLUME 40 associated organisms. Opportunities are rarely provided to study the history of two diverse genetic systems that evolved under very similar ecological circumstances for long periods of geologic history. The evolution of staphylinid termitophiles, as evidenced by data to be presented later, was quite remarkable for the high degree of host specificity that characterized it. There are probably few cases to be found among host-parasite or host-commensal relationships in which the integrity of host relationships was maintained so faithfully over such long periods of evolutionary history. If the evolutionary histories of termitophiles have been very closely linked with those of their hosts, hypotheses relating to the phylogenetic and biogeographical histories of hosts and guests should be mutually supportive. One of the major conclusions of this work is that such hypotheses, worked out independently for termites and Staphylinidae, have a very high degree of concordance. Progress in preparing hypothetical phytogenies of the termitophilous groups has been slow, due in large part to the artificial nature of the classification of the subfamily Aleocharinae. Before aleocharine categories containing termitophiles could be reorganized so as to be defensible on phylogenetic grounds, it was necessary to devote considerable study to the entire subfamily. Fortunately, on the other hand, the phyletic history of termites has been extensively studied by Dr. Emerson and several of his students, Dr. Laura Hare, and Dr. Muzaffer Ahmad. Emerson has clarified many aspects of termite evolution and has proposed important principles relative to the biogeography of termites. At the outset it is proper to take cognizance of earlier works relating to the evolution of termitophily. The large majority of papers on termitophilous Staphylinidae have been taxonomic in character and have devoted little attention to evolution. The pioneering studies on myrmecophilous and termitophilous insects made by the Rev. E. Wasmann may have been important in some respects, but they contributed little of significance to our knowledge of the phylogeny, biogeography, and evolution of these insects. For more than thirty years Wasmann wrote voluminously and repetitiously about certain aspects of the evolution of myrmecophily and termitophily, but emphasis on mystical concepts such as "amical selection" and "symphilic instincts" did much to discredit his theoretical views. Wasmann's opinions have little bearing on this monograph and need not be reviewed. In 1920, Warren published a diagram purporting to show the independent evolution of physogastric Aleocharinae in the three SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 11 major continental tropic areas of the world. This fanciful tree was quite obviously based on a superficial examination of the literature; Warren was not a specialist on the Staphylinidae. His "phylo- genetic tree" is probably one of the most absurd on record; genera of different tribes and subtribes are arranged in three lines with complete disregard of the facts of morphology. Wheeler (1928, pi. 41) published Warren's tree and, tacitly at least, approved it. Warren derived the termitophilous Staphylinidae of each major tropical area from generalized Myrmedonia-Yike stock and showed in each case a series of genera presumably leading to a highly developed physogastric genus at the apex of the tree. At the apices of his independent evolutionary lines he had Spirachtha (South America), Termitomimus (Africa), and Termitoptochus (Indomalaya). It will be shown later that these genera were derived from termi- tophile stock that spread throughout the tropics of the world with its hosts. It will also be shown that the genera of Warren's "evolutionary lines" belong to a series of tribes and subtribes and could not conceivably bear the relationships indicated. Warren concluded that the termitophilous Staphylinidae gradu- ally evolved from free-living species of the different faunal regions after the major termite areas became geographically isolated and the various termite genera had evolved and spread. He stated that the termitophilous faunas of the major termite regions have a striking external similarity but that systematically they do not appear to be more closely allied than the ordinary free-living faunas. Warren reasoned that a greater common element would exist throughout the termitophilous faunas of the world if the termi- tophiles had evolved before the general distribution of termites. I can only conclude from my studies that Warren's generalizations are completely erroneous when applied to the major groups of termitophilous Staphylinidae. Warren accepted the highly artificial aleocharine classification without critical analysis and thus failed to recognize the "common elements" in the termitophilous faunas of the major tropical areas. One of the major thesis of the present work is that a number of termitophilous groups originated early in the evolutionary history of their host group and dispersed with their hosts to far distant areas. In recent years new methods of approach to problems of insect evolution have been utilized with considerable success. For many years attempts to analyze the evolutionary history of insects have relied almost exclusively on fossil evidence. Paleozoic fossils have given some insight into the nature of very ancient insects, and 12 FIELDIANA: ZOOLOGY, VOLUME 40 significant information about more recent forms has resulted from studies of Tertiary fossils, but, except in a few groups, there have been no Mesozoic fossils to illuminate that important period, when many modern subfamilies, tribes, and genera were differentiating. But even without fossil evidence it now seems possible to deduce past histories of some insect groups after a careful synthesis of various areas of evidence: their phyletic lines as judged by compar- ative morphology; their present-day geographical distributions and ecological limitations; and knowledge and hypotheses about ancient land masses, land bridges, and climate. Some very important contributions to our knowledge of insect evolution have resulted from this type of analysis (Emerson, 1949, 1952a, 1955, on termites; Darlington, 1950, on paussid beetles; Ross, 1951, on caddiceflies) . Evidence is accumulating to demonstrate that many insect genera, both primitive and derived, have existed without important changes since Cretaceous times or earlier. The evidence for this hypothesis is largely circumstantial but as the evidence grows to significant proportions the probability that the hypothesis is valid increases. ACKNOWLEDGMENTS I am very greatly indebted to Dr. Alfred E. Emerson of the University of Chicago, whose numerous contributions to this project made it possible. Through his own collecting efforts and through contacts with other termite specialists, Dr. Emerson has been the most important source of material. Even more significant as a contribution to our knowledge of termitophile-termite evolutionary interrelationships, Dr. Emerson has made it possible to present a host list of unprecedented accuracy. The termites that became available were identified by him, with the exception of certain Brazilian specimens determined by Mr. Renato L. Araujo. Dr. Emerson also checked the host records cited by earlier investigators and in many cases suggested changes of zoological or nomenclatural significance (the actual specimens on which these earlier records were based were not examined, however). Finally, discussions of various termitophile problems that Dr. Emerson and I have had over a period of years contributed significantly to the improvement of the monograph. It is a pleasure to express appreciation to the following persons through whose courtesy and generosity I was able to study im- portant materials: Dr. A. Reichensperger, Bad Godesberg, Germany; Mr. R. L. Araujo, Institute Biologico, Sao Paulo, Brazil; Drs. E. A. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 13 Chapin and R. E. Blackwelder, United States National Museum; Drs. E. Ross and H. B. Leach, California Academy of Sciences; Drs. T. Borgmeier and Helmut Sick, Brazil; Dr. W. M. Mann, Washington, D.C.; Dr. P. J. Darlington, Museum of Comparative Zoology; Mr. W. O. Steel, Maidenhead, England; Messrs. R. L. Wenzel and H. Dybas, Chicago Natural History Museum. After completing what I considered to be a final draft of this monograph, I had an opportunity to visit several European museums in connection with other studies on the systematics of Staphylinidae. This trip, made possible by a grant-in-aid from the National Science Foundation, enabled me to examine types and other specimens of termitophilous Staphylinidae not previously available. Some of the results of these studies have been incorporated in this manuscript. In addition to gratefully acknowledging the aid from the National Science Foundation, I wish to thank the following men for their kind co-operation in helping to make the study trip successful: Dr. E. M. Kruytzer, Curator of the Natuurhistorisch Museum, Maastricht, The Netherlands, and Dr. H. Schmitz, Curator of the Wasmann collection in the Natuurhistorisch Museum; Dr. A. Reichensperger, Bad Godesberg, Germany; Messrs. N. D. Riley, E. B. Britton, and J. Balfour-Browne of the British Museum (Natural History); Mr. Gaston Fagel of the Institut Royal des Sciences Naturelles de Belgique. ORGANIZATION OF TERMITE SOCIETIES AND TERMITE EVOLUTION It is appropriate at this point to review some of the general features of termite evolution as background for subsequent discus- sions of host-guest interrelationships. This section deals with the organization of termite families and their phylogenetic relationships. Later in the paper, those termite families and subfamilies that con- tain important groups of termitophiles are considered in greater detail. The material for this summary was drawn for the most part from the publications of Emerson; these include discussions of the evolution of termite nesting behavior (Emerson, 1938), termite ecological and evolutionary interrelationships (Emerson, 1949), and the biogeography of termites (Emerson, 1952, 1955). Discussions of termite phylogeny in this and later sections are based on the above works, on several of Emerson's taxonomic papers, and on contributions by Hare (1937) and Ahmad (1950). 14 FIELDIANA: ZOOLOGY, VOLUME 40 The 1800 named species of living termites are grouped into about 150 genera and 5 families. Primarily tropical insects, relatively few termites occur even in warm temperate regions, and only about nineteen species of four genera are adapted to a temperate climate. The general relationships of the five termite families are shown in a diagram (fig. 1) designed to illustrate independently derived stocks of staphylinid termitophiles and their primary host associations. A short resume" of the social systems of these families is presented below, although we are chiefly concerned with the Termitidae and their guests. Kalotermitidae: The termites of this family are primitive, as evidenced by their morphology, social organization, and nesting behavior. The kalotermitids, frequently called dry-wood termites, excavate irregular channels in decayed portions of tree trunks galleries which require no soil connections. The kalotermitids do not construct definite termitaria and most of their galleries result from feeding. The colonies of these termites consist of reproductive males and females, soldiers, and nymphs which perform the work of the colony; no adult worker caste is differentiated. The queen is not physogastric and no royal cell is constructed. The Kalotermitidae apparently do not have staphylinid guests and will not be discussed further. Mastotermitidae: This family is represented by a single living species in Australia. Fossil evidence shows that species of this family were widely distributed in past periods. These termites show many primitive morphological features and their social system contains no worker caste. The nesting behavior is more specialized than that of the Kalotermitidae and a colony may contain more than a million individuals. Hodotermitidae: A small family of primitive termites regarded as relicts of an earlier, more widely distributed group. Rhinotermitidae: Although not as a rule having more elaborate nesting behaviors than the Kalotermitidae, the termites of this family exhibit a more highly developed social system, as evidenced by differentiation of a worker caste, development of a physogastric queen with greater reproductive capacity, and increase in colony size. Nests of most species are found in excavated galleries in wood, but the galleries often extend into the ground and external gallery- tunnels are present. Coptotermes, a large tropical genus, may construct large mounds of triturated wood or clay mixed with excreta. Rhinotermes and close relatives are noteworthy for their SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 15 Aleocharinae Ami- termitinae Termitonannina Termltonannini Perinthina Corotocmi __ ' Pseudoperinthini ' Termitohospitini Termitozyrina Macrotermitinae \ Rhmotermitidae Hodoterm'tidae Kolotermitidoe Pygosteninae Trichopseniinae FIG. 1. Primary host relationships of major categories of termitophilous Staphylinidae. dimorphic soldiers, the minor soldiers having a prolonged labrum for ejecting a repellent fluid. Reticulitermes is of special interest in that most of its species are adjusted to temperate conditions. Termitidae: About three-fourths of the termite species belong to the Termitidae, a family widely distributed in the tropics. Their social life is the most advanced in the Isoptera, and its members are 16 FIELDIANA: ZOOLOGY, VOLUME 40 noted for morphological specializations. The many nest types and nesting behaviors make generalization difficult; some species reside in nests no more elaborate than those of the simplest Rhinotermi- tidae, while others construct termitaria of remarkable complexity. Reference to the diagram of termite relationships (fig. 1) will show that the Termitidae includes four important subfamilies, two of which (Nasutitermitinae and Macrotermitinae) contain a large majority of the staphylinid termitophiles. Discussions of the evo- lution and biogeography of these subfamilies are deferred for later sections. The evidence suggests that successful establishments of various termitophile groups occurred after the subfamilies had differentiated. However, it is possible that early generalized members of the Termitidae may have had guest species that were passed on to several subfamilies. The Termitinae and Amitermitinae have relatively few staphylinid termitophiles and some of these may be the result of secondary adaptations on the part of nasutitermitine guests. Nests of the Amitermitinae should be thoroughly investi- gated in the hope of finding evidence bearing on the origin of some termitophile groups. General Remarks on the Biogeography of Termites The following summary is abstracted from Emerson's paper on the biogeography of termites (1952a). Many of the generalizations are relevant to the discussions of this monograph and will be dis- cussed later in context. All five termite families were in existence and had dispersed to the major tropical areas of the earth by late Mesozoic times. The subfamilies of Termitidae differentiated in the Mesozoic and in the Tertiary. No major changes in the present continental land masses need be postulated to account for the distribution of termites, but several hypotheses pertaining to land bridge connections are necessary. It is necessary to postulate that tropical conditions existed in Mesozoic times and that a Bering or Aleutian land bridge was available to tropical insects, thus enabling termites originating in South America to introduce elements into the Australian, Indo- malayan, and Ethiopian faunas, and termites originating in the Old World to introduce elements into the Neotropical fauna. It seems very unlikely that this bridge was available to tropical termites during the Tertiary. South America and North America were connected presumably at the Isthmus of Panama before Eocene and after late Pliocene times. Opportunity for the spread of termites SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 17 from the Indomalayan region to Australia existed in the Cretaceous Period but not later. Access from tropical Asia to tropical Africa probably existed under moist conditions in the Pliocene or earlier, and Madagascar was probably contiguous to the African continent in the late Cretaceous or Eocene but not later. In discussing the dispersal of termites Emerson emphasized the importance of such factors as food, defense against predators, and ability to compete against ecological equivalents, and he notes that many genera failed to move far from their place of origin because of these biotic factors. Accidental or chance dispersals were kept at a minimum by the fact that termites were dispersed only by land or over narrow water gaps in a large majority of cases. Termites are relatively weak fliers and after the nuptial flight the males must find the females to mate, a situation that precludes long, successful flights over water. TERMITOPHILY IN GENERAL During the long evolutionary history of termites and ants, their societies have been invaded continually by predatory animals of great diversity. The probability that a particular species of staphy- linid beetle could adapt to the exacting set of conditions imposed by social life was extremely small, yet a few were able to do so. The nature of the integrative mechanisms of insect societies was such that some alien species were able to adapt to them and to become termitophiles or myrmecophiles. The integration of termite popu- lations is achieved through behavior mechanisms initiated by various sensory stimuli, particularly those of a chemical nature (see Emerson, 1949, pp. 419-435, for a discussion of the organization of insect societies). Colony odors are evidently quite specific in character; some may be characteristic of the species, others of the individual colony, and termites are probably accepted or rejected on the basis of their odors. The difficulties involved in adapting to the narrow limits of the society are emphasized by the fact that the present-day fauna of termitophilous Staphylinidae represents per- haps a dozen establishments of individual species, for each natural group of termitophiles (tribe, subfamily, etc.) stemmed, of course, from one established species. Inasmuch as the term "termitophile" has been broadly and loosely applied to almost any animal found in or near termite nests, it is essential to define the term as used in this monograph. Un- critical use of the term has resulted in a superficially impressive list 18 FIELDIANA: ZOOLOGY, VOLUME 40 of termitophiles ranging from annelid worms to reptiles (see Warren's 1920 list of the kinds and number of species of animals recorded from termite nests). Inclusion of all manner of organisms in the category of termitophiles has tended to obscure generalizations that might be reached. I propose to restrict the use of "termitophile" to those species of staphylinid beetles that are obligatory members of termite societies at some stage in their life history. This implies that termitophilous species are unable to survive without their host societies, and that, as a rule, they leave the colony only for dispersal purposes. Whether or not this definition is applicable to termitophiles in general is debatable, and no attempt will be made to justify it as a general definition. It is admittedly difficult in certain cases to determine whether a species has an obligatory relationship to termite societies or not. Experience has shown that certain groups of Staphylinidae are always found in termite nests and we may assume that newly discovered members of these groups are termitophilous. Physo- gastric species, even when collected for the first time, may be presumed to have an obligatory association with termites. On the other hand, one may be justifiably skeptical of species of large free-living genera, such as Atheta, which are found in termitaria. Various species of staphylinids are attracted to termitaria when they are broken open for investigation, and if the collector is not careful some of these species may soon be on the list of termitophiles. In doubtful cases, repeated captures are necessary to establish the status of a species. Wasmann's Classification of Termitophiles Wasmann, recognizing the fact that all organisms associated with social insects do not have the same relationships to the colony, proposed that termitophiles be placed in three categories: synech- thrans, synoeketes, and symphiles. Synechthrans were defined as poorly adjusted, often persecuted predators living in close association with termite nests; presumably these species survive in sparsely inhabited areas of the nest or in the vicinity of nests. The synoeketes were considered by Wasmann to be tolerated guests with no obvious adaptation for exudate production; they were believed to be scaven- gers in the nest. Most synoeketes were said to have defense-type body forms, usually limuloid. Symphiles were defined as "true guests," inasmuch as they have some morphological adaptation (physogastry or trichomes) presumed to be for the elaboration of exudates. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 19 Wasmann's categories have been generally accepted (Wheeler, 1928, Emerson, 1949, and others). Nevertheless, it seems that the classification should be examined critically, and the desirability of retaining it discussed. Emerson (1949, p. 719) stated that "Was- mann's classification seems to have stood the test of time," but I suspect that the categories have been used without being considered very carefully. I have no objection to the use of the term "synechthrans" for termitophagous predators, but I believe that they should not be regarded as termitophiles. They exhibit no integration into the social systems that they prey on, and it is doubtful, at least in many cases, that this is an obligatory relationship. The staphylinid synechthrans are morphologically generalized, and there is no evidence whatsoever that this category provides material for the evolution of true termitophiles. Inasmuch as a division of termitophiles into symphiles and synoeketes in accordance with Wasmann's definitions requires some knowledge of the behavior and physiology of each species, such a division is virtually impossible. In practice, definitions have been ignored and all physogastric species have been called symphiles and all non-physogastric species referred to as synoeketes. It has been assumed that all physogastric species (symphiles) have established with the termites a mutualistic relationship not attained by the synoeketes; this carries the implication that they are better adapted and more successful than the synoeketes. In all probability this implied difference in success exaggerates the actual gap between these two categories. If success is measured in terms of ability to survive in the social systems, there is probably little to choose between in the two categories; the non-physogastric termitophiles have, on the whole, probably survived for as long a period of evo- lution, and at present inhabit as many colonies and in as great numbers as the symphiles. It seems to me that all termitophilous Staphylinidae, as defined in this paper, have achieved physiological adaptation to the biotic conditions of termite societies, with all that this implies. By one device or another they have achieved a method of escaping detection in societies highly sensitive to foreign organisms and well prepared to cope with them. Both so-called synoeketes and symphiles are remarkably well adjusted physiologically to termite societies, and even though the latter exhibit adaptive features presumably not possessed by the former they cannot justifiably be differentiated as "true guests." In order to avoid 20 FIELDIANA: ZOOLOGY, VOLUME 40 unjustified connotations the terms "physogastric" and "non- physogastric" termitophiles will be used in this paper. Before we terminate a discussion of Wasmann's categories, there is one matter that calls for comment. Wheeler (1928, p. 265) stated that "in both cases (myrmecophiles and termitophiles) there is the same origin from synechthrans and synoeketes, which, as they become more intimate with their hosts, gradually take on the peculiar characters of symphiles." This statement seems to be completely without foundation and probably gives the incorrect impression of the evolution of staphylinid guests. The non-physo- gastric synoeketes are specialized for termitophily in their own fashion, and many genera have apparently existed with little change since Cretaceous times. The evidence, to be discussed in more detail later in the paper, suggests that the specialized physogastric termitophiles evolved rather rapidly from generalized aleocharine stocks and became physogastric early in their evolution, probably during Cretaceous times or earlier. There are a few cases in which physogastric genera have evolved within a group of non-physogastric species some time after the group had become established, e.g., Termitonilla of the Termitonannini, but in the large majority of cases a major category of termitophiles is comprised of physogastric or non-physogastric species but not a combination of the two types. The present-day termitophilous groups probably originated early in the evolutionary history of their host groups, attained their adaptive features in a relatively short time, and have since changed only in relatively minor ways. Major Adaptations Associated with Termitophily The advantages that termitophilous insects may have in living in well-established termitaria seem to be these: galleries and chambers provide secluded places even for some poorly adapted insects; physical conditions of temperature and humidity are controlled and uniform; available food is plentiful and varied; the colony offers protection against predators. The physical factors of termite nests could scarcely have offered difficult problems of adjustment to potential termitophiles but the biotic factors with which they had to cope were indeed formidable. Only by adjustment to factors that serve to integrate the colony could resistance be overcome. The most conspicuous adaptive features of termitophiles are those that relate to trophallaxis. The physiological mechanisms of social integration are not well known, but it is presumed that morphological specializations such as physo- SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 21 gastry and exudatory appendages are concerned with them in some way. Physogastry and exudatory appendages. The physogastric ab- domens of termites and termitophiles (figs. 14, 24, c) are swollen to some degree and the sclerites are separated by areas of non-sclero- tized, membranous integument. Physogastry reaches its maximum development in certain termite queens, the abdomen of some Macrotermes queens attaining lengths of more than five inches and volumes some 125 times greater than that of stenogastric individuals. Wasmann and Holmgren were early proponents of the idea that one function of the physogastric abdomen is to secrete exudates that serve an important role in colony integration. Holmgren (1909), in seeking a physiological mechanism of exudation, decided that the abdominal adipose tissue of termites is a source of a fatty secretion carried by the blood to the integument, through which it exudes. Wheeler proposed the theory of trophallaxis to account for social integration and attributed considerable significance to abdominal exudates. Escherich and others had noted that the physogastric abdomen of queens was continually licked, and even nibbled at, by workers tending the queen and progeny. Although there seems to be agreement that physogastric ab- domens of termites secrete substances avidly sought by workers, there is as yet no satisfactory explanation for the physiological mechanism involved. Hypertrophy of the queen's abdomen is primarily due to increase in size of the ovaries and is basically an adaptation for increased egg production. Blood tissue increases and long columns of royal adipose tissue are well developed. Grasse" and Lesperon (1938) and Grasse" (1949) pointed out that these fat bodies have no secretory function and do not participate in exuda- tion. Grass also stated that there are no glandular hairs or in- tegumental glands on the distended termite abdomen, yet in some cases droplets of clear fluid appear on its surface. Wheeler (1928) observed termites transporting staphylinids of the genus Spirachtha and licking their remarkable abdominal ap- pendages and other membranous parts of the body. Wheeler explained the physogastry of termitophiles as the result of adaptation to the peculiar trophallactic habits of termites and to similar physical and trophic conditions. He believed that confinement to narrow, crowded galleries, limited oxygen supply, absence of light, and an abundance of carbohydrate food are conducive to adiposity and that physogastry may result from these conditions if the organisms are predisposed to it. 22 FIELDIANA: ZOOLOGY, VOLUME 40 No adequate explanation of the physiological mechanism of exudate secretion in termitophiles is available. Mclndoo (1923) made a histological examination of the Spirachtha abdomen and appendages but came to no important conclusions; he suggested only that materials could pass from the blood into hypodermal cells and through the cuticle. Tragardh (1907) reported that the physo- gastry of Termitomimus was due primarily to hypertrophy of fat bodies and to a lesser extent to the gonads, and that the cuticle is penetrated by many extremely fine pores. Grasse" and Lesperon (1938) studied the secretory mechanism in exudatory appendages of a termitophilous beetle larva, Troctontus silvestrii Grasse" and Lesperon. They reported that termites licked the appendages and bodies of the larvae and that in return the larvae received stomodeal aliment. The secretion was elaborated by special open hairs (adenotrichs) having a saccule at their base, which opens into the hair cavity. This is evidently a special case and is perhaps not typical of mechanisms in other termitophilous larvae (Lepidoptera, Diptera, Coleoptera). Studies on physogastry and its physiological role in termitophily are greatly needed if this important feature is to be fully appreciated. Limuloid body form. Inasmuch as numerous termitophilous and myrmecophilous groups are characterized by the limuloid body-form of their members, Wasmann presumed that it has much adaptive significance. He believed that limuloid species are not particularly well adjusted to social conditions and rely on their form for pro- tection. There is little evidence bearing on this hypothesis but it will be discussed at appropriate places later in the paper. II. Termitophilous Staphylinidae: Phylogenetic and Biogeographical Considerations It is now evident from host-termitophile data that the evolution of each termitophile category (subfamily, tribe, subtribe) occurred almost entirely in conjunction with the phylogenetic history of a natural group of termites. Speciation and generic differentiation of termites within the termite groups doubtless provided oppor- tunities for the isolation of various termitophile populations and for subsequent speciation phenomena. A high degree of host specificity was necessary for evolution of termitophiles to be restricted to phy- letic lines of termites. Many species of present-day termitophiles are monoxenous, and the others generally occur with one or more allied species of the same host genus. Most of the records of termi- tophilous species occurring with several genera of termites are unsubstantiated ; many of these records are probably due to careless and uncritical collecting techniques. Even though host specificity is strong in most termitophile relationships, it is to be expected that some cases of termite speciation would have resulted in the occurrence of a staphylinid species with two host species for some period of time. Of course we may have difficulty in determining the true state of affairs, as our judgments are based on morphological criteria. Individuals occurring with two termite host species may be judged to belong to the same species, yet may have undergone physiological speciation. Such cases could hardly be detected without experimental analysis. It is probable that some species of termitophile have been able to adjust to the social conditions of a second termite host rather rapidly, even in rare cases to the societies of distantly related termites. We may conclude that staphylinid guests, in general, have been very conservative in their host relationships and that transfer of host affinities has been very difficult. The host-guest patterns that are now discernible could hardly have been attained under other circumstances. 23 24 FIELDIANA: ZOOLOGY, VOLUME 40 TERMITOPHILOUS STAPHYLINIDAE ASSOCIATED WITH THE NASUTITERMITINAE Phylogeny and Evolution of the Nasutitermitinae The discussion below reflects the phylogenetic concepts of Emerson (1945, 1949, 1952a, 1952b) and Ahmad (1950). The Nasutitermitinae, the largest subfamily of termites, are very widely distributed in tropical regions. Their most noteworthy evolutionary advance was the differentiation of nasute soldiers, an achievement presumably contributing greatly to their success. The primitive nasutitermitine genera, restricted to South America, have conventional soldiers with biting mandibles, whereas the much more numerous derived genera have soldiers with prolonged snouts and functionless mandibles. Forceful ejection of a sticky repellent from their snouts constitutes their mode of defense. Emerson's studies have led him to conclude that nasute soldiers evolved independently in two phylogenetic sequences of Nasutiter- mitinae (fig. 3), each line containing, among others, a large tropi- copolitan genus, Nasutitermes and Subulitermes, respectively. Inasmuch as all the present-day primitive mandibulate genera (Syntermes, Procornitermes, Cornitermes, etc.) are restricted to the Neotropical Region, it is reasonable to believe that the Nasutiter- mitinae differentiated in this area. The Nasutitermes branch. This phylogenetic sequence, com- prised of some nineteen genera, is widely distributed in the tropics of both hemispheres. Its largest genus, Nasutitermes, contains about 195 named species; they occur in the Neotropical Region, the Oriental Region, the Ethiopian Region, Madagascar, Australia and the Papuan Region, Micronesia, and Polynesia as far as the Fiji Islands. Emerson postulates that this line of nasute genera stemmed from Syntermes-\ike stock during Cretaceous times in South America and subsequently spread to the Oriental Region by way of an Alaskan land bridge. This dispersal was early enough to allow the group to reach Australia and the Pacific Islands. After the Alaskan portal became ecologically unsuitable for these genera tropical conditions were essential several endemic genera arose in each hemisphere during the Tertiary. Such endemic genera as Velocitermes, Tenuirostritermes, Diver sitermes, and Constricto- termes appeared in the American tropics. The Palaeotropical Region evolved a series of genera including Grallatotermes, Lacessiti- termes, and Trinervitermes. The last-named genus is a large and SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 25 important one occurring in grasslands of Africa and Indomalaya. Emerson believes that it originated in Africa during Miocene times, soon after the tropical savannas were established. The Subulitermes branch. This sequence of genera was probably derived from Syntermes-Procornitermes stock in South America. Several neotropical genera Paracor niter mes, Labiotermes, Armi- termes, and Curvitermes have well-developed mandibles as well as long frontal protuberances. The most advanced genera of this evolutionary line have nasute soldiers with reduced, functionless mandibles. Subulitermes is found rather widely in both Neotropical and Palaeotropical Regions and other genera are endemic to Africa (Mimeutermes, Eutermellus) or other areas. Inasmuch as the genera of this branch of Nasutitermitinae have relatively few termitophiles, they do not enter into subsequent discussions as frequently as do those of the Nasutitermes branch. The Termitophilous Tribe Gorotocini As constituted in this monograph the Corotocini is the largest and most highly specialized tribe of termitophilous Aleocharinae. Originally proposed by Fenyes (1918) to include a series of termi- tophilous and dorylophilous genera having 4, 4, 4-segmented tarsi, the tribe was reduced to the status of a subtribe of Hygronomini by Bernhauer and Scheerpeltz (1926). I cannot concur with the previous treatments of the group for these reasons: all aleocharines with 4, 4, 4-segmented tarsi should not be grouped in one tribe; African dorylophiles do not belong in the same group with termi- tophilous genera; the Corotocini should be expanded to include genera formerly in other tribes. More than 120 species, or almost one-half of the known termitophilous Aleocharinae, belong here. Even though our knowledge of this tribe is far from complete, the body of information now available provides a satisfactory basis for speculation concerning its evolutionary history. General intra- tribal relations are illustrated by figure 2, and hypothetical phyletic lines within subtribes are shown in a series of diagrams (figs. 4, 5, 7). The evidence, discussed in more detail later, suggests that the Corotocini originated in the Neotropical Region from generalized aleocharine stock. It is hardly to be expected that any present-day genus would fulfill all the criteria as an ancestor for the Corotocini. The large free-living genus Oxypoda is doubtless a generalized aleocharine and Oxypoda-\ike stock could well have been ancestral 26 FIELDIANA: ZOOLOGY, VOLUME 40 to this tribe. While it has been conventional to place Oxypoda near the end of aleocharine lists, this only reflects the artificiality of the classification. In subsequent discussions Oxypoda is used to represent the generalized Aleocharinae for comparative purposes but this does not imply that it was the ancestral genus. Corotocina Pantroplcal Abrotelina Neotropical Term/toga strina Old World Termitogastrina M Neotropical Termitocharina Madagascar ^-^ yyy \ i / Timeparthenino Neotropical Corotocini FIG. 2. Subtribes of Corotocini. Tarsal formulae are indicated. A. Tribal characteristics. In support of the viewpoint that the Corotocini are a monophyletic group is the fact that its genera are characterized by four traits of an unusual nature. Each of these characters appears rarely in other Aleocharinae but no group possesses all of them. The tribe Termitopaediini, associated with Macrotermitinae, is comprised of physogastric genera that have acquired several of the same adaptive traits, e.g., the free mesocoxae and triangular hind coxae. Before proceeding to a discussion of the SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 27 phylogenetic relationships of the Corotocini an analysis of the most important tribal characteristics is desirable. 1. Physogastry: This adaptation evolved independently among termitophilous insects on a number of occasions (see general dis- cussion of physogastry, p. 21). All corotocines have this feature to some degree, although there is a surprising diversity of expression within the tribe. Physogastry is an important character in differ- entiating the Corotocini from all non-termitophilous Aleocharinae, but not from all termitophilous groups. Several subtribes of Coro- tocini have distinctive patterns of physogastry. 2. Mentum fused to submentum: Although the Aleocharinae in general have the mentum free and movable, the corotocines have the mentum fused to the submentum to form a shield-like sclerite (fig. 18, q). It is reasonable to believe that the stationary condition of the mentum has some significance in feeding behavior; if these beetles receive liquid aliment from the hosts, this condition may facilitate the process. A few genera of termitophilous Aleocharinae have independently acquired a similar condition (Lauella of the Termitonannini, Termitobaena of the Termitopaediini), but this is exceptional among non-corotocine termitophiles. Furthermore, all myrmecophilous Aleocharinae have the conventional labial con- dition. 3. Mesocoxal acetabula unmargined; mesocoxae free. Hind coxae triangular; metepimera abbreviated: The evolution of physo- gastry evidently necessitated adaptive changes in the legs and thorax to provide better balance for the hypertrophied abdomen. A series of adaptations seem designed to lengthen the legs and to spread them so as to balance the voluminous abdomen (fig. 21, a, 6). Femora, tibiae, and tarsi of many corotocines are rather long, but this alone does not seem to serve the purpose. The mesocoxae are free; they are not set in deep, well-defined acetabula in typical aleocharine fashion. The unusual length of the coxae, their freedom, and their widely separated points of artic- ulation seem to suggest that the middle legs are capable of being widely spread. The hind legs are somewhat lengthened and more widely spread by several modifications. The typical aleocharine condition places the proximal ends of the hind femora rather close together, but the corotocine femora articulate with elongated, tri- angular coxae. The subtribe Corotocina carries this adaptation farther, and the legs are even more widely separated due to pro- longations of the metasternal articulating processes (fig. 21, 6). It 28 FIELDIANA: ZOOLOGY, VOLUME 40 is significant that the Corotocina have the most voluminous ab- domens in the tribe. Abbreviation of the metepimera is apparently correlated with change in form of the hind coxae, but the significance of this is not apparent. 4. Terminal antennal segments with two or more coeloconic sensilla: The species of this tribe, as well as those of the termi- tophilous tribe Termitonannini, bear distinctive coeloconic sensilla on the terminal antennomeres. These specialized receptors, usually visible only in cleared specimens, consist of an oval pit into which sensory pegs extend. Inasmuch as sensilla of this exact type and position are not commonly found in the Aleocharinae, they are a useful supplementary character in diagnosing Corotocini and Termitonannini. Similar sensilla occur in other Staphylinidae; Sym- polemon Wasmann, an African genus of dorylophilous Pygosteninae, has them. Coeloconic sensilla have been noted on antennae of insects of various groups; the antennae of the house fly have pits with 10-20 sensory pegs (Snodgrass, 1935, p. 521). There can be no doubt that these sensilla are chemoreceptors, but there are of course no clues to the nature of the chemicals to which they are sensitive. They are among the three types of sensilla (placoid, basiconic, and coeloconic; Snodgrass, 1935, pp. 514-523) usually associated with olfaction. B. Phylogenetic and biogeographical considerations. Phylogen- etic, biogeographical, and host data may reasonably be inter- preted to show that the history of the tribe Corotocini began with the establishment of an aleocharine species in societies of a general- ized species of Nasutitermitinae and that this event occurred in South America during the Cretaceous Period or earlier. The fol- lowing discussion in support of this statement is accompanied by a series of diagrams and maps. The diagrams are designed to illus- trate phyletic relationships within the Corotocini (figs. 2, 4, 5, 7) and to show how closely associated the phyletic history of the tribe was with that of the Nasutitermitinae (fig. 3). The maps (figs. 6, 8) present hypothetical dispersal patterns to account for the present distribution of corotocine termitophiles. 1. The origin of the Corotocini: Inasmuch as all corotocine genera have the specialized characteristics of the tribe, links with generalized Aleocharinae such as Oxypoda are very difficult to detect, if, indeed, they exist. Termitophya (subtribe Termito- gastrina) and Termituncula (subtribe Timeparthenina) have features SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 29 Tenuirostritermes >y Velocitermes Diversitermes i-AJ Nasutltermes ^ Trinervitermes Armitermes Termitogastrin a Paracornitermes f X""* Cornitermes --~~_ Procornitermes Nasutitermitinae FIG. 3. Host relationships of tribe Corotocini. indicating that they are the most generalized members of the tribe. At least each is near the base of a phyletic sequence within the Corotocini. On morphological evidence alone it would be exceed- ingly difficult to determine which of the present-day groups first differentiated from basic corotocine stock. However, the host relationships of these subtribes seem to provide the answer. All genera of Termitogastrina occur with derived genera (Nasuti- termes, Trinervitermes, etc.), whereas most species of Timepar- thenina are associated with primitive mandibulate Nasutitermi- tinae (Cornitermes, Procornitermes, Paracornitermes}. We know, 30 FIELDIANA: ZOOLOGY, VOLUME 40 then, that the Corotocini had become established early in the history of the Nasutitermitinae. The present-day Timeparthenina (figs. 4, 14) evidently represent a fragment of early corotocine stock that became isolated with the Cornitermes complex of genera and had no opportunity to become associated with the Nasutitermes branch of Nasutitermitinae. Both the Timeparthenina and the primitive mandibulate genera of Nasutitermitinae are restricted to South America, a fact that suggests a neotropical origin for the Corotocini. The subtribe Timeparthenina is distinctive in its pattern of physogastry; the basal abdominal segments undergo the greatest increase in size, and the abdomen cannot be recurved above the thorax. Termiluncula, Autuoria, and Timeparthenus illustrate a tendency for the second, third, and fourth abdominal segments to become progressively larger so large in extreme cases that the basal part of the abdomen overlies the thorax. As evidenced by its relatively small physogastric abdomen, 5, 5, 5-segmented tarsi, and 11-segmented antennae, Termituncula is the most generalized mem- ber of the subtribe. Among the derived genera cf Timeparthenina certain trends are noteworthy: increased hypertrophy of the abdo- men, decrease in tarsal segmentation from 5, 5, 5 to 4, 5, 5 and then to 4, 4, 5, and reduction in number of antennal segments to ten. ' Although it appears that the primary association of the Time- parthenina was with the generalized mandibulate genera of Nasuti- termitinae, the fact that the two most highly specialized genera, Timeparthenus and Ptocholellus, occur with Anoplotermes (Amiter- mitinae) merits attention. The occurrence of a natural group of termitophiles with two subfamilies of termites seems to require one of these alternative explanations: (1) a species of an established termitophilous group may have been able to adapt rather rapidly to a termite species of a different subfamily; (2) the termitophilous group may have been established with termite stock ancestral to both termite subfamilies. Available evidence in the case of the Timeparthenina seems to suggest the first alternative. The pri- mary evolution of the group probably took place in association with several primitive genera of Nasutitermitinae, followed by a secondary establishment of one of its species in societies of a species of Anoplo- termes. This secondary establishment led to subsequent differentia- tion of several specialized genera, Timeparthenus and Ptocholellus. One can only surmise the difficulty of the major host change required by this hypothesis. At the present time earthen mounds frequently harbor species of Anoplotermes in close proximity to various species SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 31 of Nasutitermitinae. Presumably, it would be no more difficult for a termitophilous species to adapt to a new host genus than for a free-living species to adapt to termite societies. It is perhaps of interest to note that the societies of Anoplotermes lack a soldier Ptocholellus Tarsi 4-4-5 Sublribe Timeporthenina FIG. 4. Subtribe Timeparthenina. caste, but it would probably be a mistake to infer that their societies are susceptible to the establishment of alien organisms because of this. As an alternative hypothesis, it is conceivable that the differentia- tion of Corotocini occurred earlier in the history of Termitidae than 32 FIELDIANA: ZOOLOGY, VOLUME 40 postulated early enough, in fact, to permit descendants of early corotocines to evolve with Amitermitinae as well as Nasutitermi- tinae. There are several objections to this theory: In the first place, only Anoplotermes among Amitermitinae is known to harbor Coro- tocini; secondly, only the most specialized members of the subtribe Timeparthenina are known to occur with Anoplotermes. 2. The differentiation and dispersion of sub tribes: After the differentiation of the Timeparthenina, generalized corotocine stock evidently became associated with the newly evolved Nasutitermes branch of Nasutitermitinae and gave rise to the other subtribes. The Termitogastrina evidently originated in the New World tropics and spread to the Indomalayan and Ethiopian Regions with the Nasutitermitinae. The Corotocina, occurring as they do in almost all tropical areas of the world, probably had the same general dispersal pattern as the Termitogastrina. The Abrotelina are probably restricted to the Neotropical Region, although Termi- tochara of Madagascar may have been derived from this group. The very poor representation of Corotocini in termite colonies of the Subulitermes branch raises interesting questions. The few species of Abrotelina known to occur with Subulitermes may be the result of a secondary establishment rather than the product of an evolutionary sequence within the Subulitermes branch. (a) Subtribe Termitogastrina: The main features of the origin and differentiation of this subtribe in the Neotropical Region seem relatively simple, but they become more complex in the Old World tropics. In the American tropics all genera and species occur with Nasutitermes, so there can be little doubt that this genus or its immediate precursor was the original subtribal host. Termitophya, the least specialized member of the Termitogastrina, retains many generalized features: The head and thorax, although having the special tribal features, are relatively generalized; the abdomen is at most feebly physogastric; and the fourth and fifth tarsal segments are free and movable. Other genera of South American Termitogastrina are more strongly physogastric than Termitophya but in no case does the abdomen have extensive areas of non-sclerotized integument as in certain Old World genera. Apart from being considerably larger than in free-living species, the abdominal sclerites are not unusually modified in the South American genera of the subtribe, nor do extreme post-imaginal changes occur as in the Corotocina. The Old World genera of Termitogastrina do not comprise as homogeneous a group as do their American allies. There is little SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 33 Generalized Corotocini FIG. 5. Subtribe Termitogastrina. doubt that they were derived from subtribal stock that entered the Palaeotropical regions with Nasutitermes-like hosts. Several of these genera Termitotima (India) and Idiogaster (Ethiopia) are rather like certain American genera, but others have undergone appreciable specialization. One generic group including Affino- ptochus (Java), Termitella (Belgian Congo), and Idioptochus (Tan- ganyika) has much in common with the above-mentioned genera 34 SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 35 except that the sixth and seventh tergites (fig. 19, d, e) are distinc- tively modified. Another group Termitoptochus (Indomalaya), Millotoca (Africa, Madagascar), and Paracorotoca (South Africa) includes the most highly specialized species of the subtribe. They are noteworthy for having large ovate abdomens with large areas of non-sclerotized integument (fig. 19, e) as well as the distinctive sixth and seventh tergites referred to above. The tarsal situation in the Old World genera merely reflects the trend, initiated in South America, toward fusing of the fourth and fifth tarsal segments; in many derived genera the segmentation is 4, 4, 4. The strongly physogastric genera, e.g., Termitoptochus, frequently have reduced prementums and labial palpi, the latter consisting of one or two very tiny segments. Our knowledge of the host relationships of palaeotropical Termitogastrina is fragmentary yet in some aspects instructive. The record is consistent in that all species occur with genera of the Nasutitermes branch of Nasutitermitinae. Approximately one-half of the species occur with Nasutitermes and the others with endemic Old World genera Trinervitermes, Bulbitermes, Grallatotermes. On the basis of available evidence, it may be postulated that ancestral stock from which the palaeotropical representatives of the subtribe were derived entered Asia at the time of the dispersal of certain Nasutitermitinae from the American tropics. Inasmuch as the American tropics are believed to have been continuous with those of the Old World by way of an Alaskan bridge during the Cretaceous Period, this may well have been the dispersal route. The hosts were probably very similar to the present-day Nasutitermes. Thereafter, the Termitogastrina were evidently successful in keeping pace with the geographic expansion of the Nasutitermitinae as well as in following their phyletic ramifications, if we may judge by present-day associations with several endemic Old World genera. (b) Subtribe Corotocina (figs. 7, 8, 21-26): The most highly specialized of all staphylinid termitophiles belong to the Corotocina. They are especially noteworthy for the remarkable post-imaginal development that results in such bizarre abdominal and thoracic modifications. Generalizations with respect to physogastry in this group are not easily made due to the diversity of abdominal types, but a correlated character involving the hind coxal articulation is important for diagnostic purposes. As pointed out in the section on tribal characteristics (p. 27), one means of improving balance in those species with hypertrophied abdomens was to spread the hind 36 FIELDIANA: ZOOLOGY, VOLUME 40 Eburmola Termitopullus Termitoptocinus Generalized Corotocini Sub tribe Cor of o tin a FIG. 7. Subtribe Corotocina. legs by various devices. Apparently the distinctive hind coxal articulation (fig. 21, 6) in the Corotocina is such an adaptation. The complexity of post-imaginal development in such genera as Thyreoxenus, Spirachtha, and Termitomimus is indeed astonishing. This process has yet to be analyzed in detail, and some of its aspects are extremely puzzling. The problem of the attainment of physo- gastry has received almost no consideration and those who have believed physogastry to be a post-imaginal development have doubtless thought only in terms of hypertrophy of the abdomen. rt C a ~ o o I .1 5 37 38 FIELDIANA: ZOOLOGY, VOLUME 40 Two series of specimens that I have examined show clearly that the problem is far more complex. Reference to photographs of four individuals of Thyreoxenus major Mann (fig. 26) and three of Termitomimus emersoni Seevers (fig. 24) will show that other characteristics, even proportions of the legs, are modified. Steno- gastric individuals are frail, slender, winged forms with abdomens of typical aleocharine appearance. Their hind coxae and metasterna do not have the distinctive characteristics of the physogastric imagos. The stenogastric individuals would not, in fact, be identi- fiable as members of the subtribe Corotocina although perhaps as Corotocini. In the case of Thyreoxenus major several intermediate stages are available, suggesting that the changes from stenogastric to physogastric condition are gradual. If intermediate stages ex- hibiting various degrees of change were not available, genetic dimorphism rather than post-imaginal development would seem to be a more attractive theory to account for these phenomena. The chief difficulty with the post-imaginal development theory is to account for the many body changes in the absence of a moult. Larvae of the Corotocina present several features of exceptional interest. As early as 1854, Schi0dte reported the presence of well- developed larvae in abdomens of female Corotoca, and Reichensperger (1936) published a photograph of a female with a larva protruding from her abdomen. A number of Corotoca females that I have examined contained larvae of various sizes. Larvae of this genus are released in an advanced stage and probably pupate almost immediately. By contrast, larvae of several species of Thyreoxenus have been collected from termite colonies and have never been noted in female abdomens. Thyreoxenus larvae, by way of further contrast with those of Corotoca, are characterized by the presence of long lateral abdominal appendages. Evidence relating to larval development in the Corotocina, inconclusive as it is, suggests the following hypothesis. If abdominal appendages of larval Thyreoxenus and other genera are exudatory structures of adaptive significance, it may be that species of these genera are capable of living freely among the termites because of them. On the other hand, viviparity in Corotoca, and perhaps in Spirachtha, may be an adaptation to offset the lack of exudatory appendages. Corotocina are known to occur in South America, Australia, the Solomon Islands, and Africa, and it is predictable that a moderate number will be discovered in the Indomalayan Region. A majority of the species occur with Nasutitermes, but others are associated with SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 39 Trinervitermes and Constrictotermes. It is difficult at present to relate the Corotocina to other subtribes very precisely. Stenogastric individuals may logically be expected to provide more clues to relationship than physogastric individuals, but so far I can only conclude that the Corotocina are more similar to some Timepar- thenina and Abrotelina than to the Termitogastrina. The Coro- tocina evidently differentiated from generalized tribal stock early in the history of the Nasutitermes branch. Judging by abdominal structure, three very closely allied genera Eburniola (South America), Termitopullus (Belgian Congo), and Termitoptocinus (Aus- tralia) are closest to the base of this phyletic sequence. Inasmuch as these genera occur with Nasutitermes it is probable that their precursor originated with that host genus in South America and accompanied it to various tropical areas of the Old World. A pre- Tertiary origin of the Eburniola-complex is suggested by the presence of Termitoptocinus in Australia as it is very doubtful if an opportunity existed in post-Cretaceous times for Nasutitermes and its guests to reach that continent. The African genus Termitomimus (fig. 24) surely originated in the Ethiopian Region from Termitopullus-like stock. The head and thorax of Termitomimus are very similar to those of the Eburni- ola group but the abdomen is remarkably distinctive. Two Belgian Congo species of Termitomimus occur with Nasutitermes and a South African species with Trinervitermes. The latter genus is restricted to the Old World tropics and differentiated according to Emerson (1952a) in Miocene times when the African savannas were developed. The evolutionary histories of the South American Constricto- termes guests Corotoca and Spirachtha are somewhat difficult to reconstruct. Apart from its astonishing abdomen with unique exudatory appendages (fig. 22), Spirachtha bears considerable resemblance to Eburniola and may have evolved from Eburniola-like stock. Corotoca, on the other hand, has departed to a greater degree from the cephalic and thoracic pattern that characterizes the majority of the subtribe. It can hardly be stated with confidence that Corotoca and Spirachtha descended from one species of the subtribe that became ecologically isolated with Constrictotermes; perhaps they originated from different segments of the Corotocina. The present- day distribution and host relationship of these genera provide few clues to their history. Constrictotermes is a small genus of three described species occurring in Brazil, the Guianas, and eastern Bolivia, Peru, and Ecuador. The small size of the genus has served to limit the number of guest species, but it is surprising that as many 40 FIELDIANA: ZOOLOGY, VOLUME 40 as three species of Corotoca are known to occur in a single colony of Constrictotermes cyphergaster. This suggests that geographic isolation of populations of C. cyphergaster has provided an opportunity for speciation of Corotoca and that subsequent reunion of the termite populations allowed the ranges of the Corotoca species to overlap. Unfortunately, the geographic ranges of the different species of Corotoca are not well enough known to provide evidence for this theory. Clues to the origin of Corotoca and Spirachtha may ultimately be provided through study of guests of the closest relatives of Con- strictotermes. Emerson (1949) and Ahmad (1950) conclude that Constrictotermes is the American representative of a group of Oriental Nasutitermitinae including Grallatotermes, Bulbitermes, Hospitali- termes, and Lacessititermes. The nests of these genera have not yet been examined for guests, except possibly in the most cursory fashion. A most interesting line of divergence within the Corotocina is represented by Thyreoxenus and Oideprosoma genera noteworthy for their physothoracic as well as physogastric condition (figs. 25, 26). Post-imaginal development of the images of these genera results in a large membranous prothorax upon which the pronotum rests, saddle-like. Ten neotropical and one Solomon Islands species of Thyreoxenus are now known. Oideprosoma is as yet represented by a single French Guinea species. The American species of Thy- reoxenus reveal little concerning the origin and history of this generic complex; the species are too uniform in structure and all occur with the host genus Nasutitermes. On the other hand, the Old World species, by virtue of their interesting localities and host associations, do provide a few clues to this evolutionary history. The Solomon Islands and French Guinea records can only indicate that the Thyreoxenus-Oideprosoma generic group has at one time or another occurred in many areas of the palaeotropical region. Indeed, it may be predicted with considerable assurance that species of this group will be found in a number of Indo-Australian and African localities, although perhaps somewhat less frequently on the latter continent. The fragmentary evidence now available points to a close association of the phyletic histories of the Thyreoxenus generic group and the Nasutitermes branch of Nasutitermitinae. All species of Thyreoxenus have been collected with Nasutitermes, and Oide- prosoma was found with Trinervitermes. As a working hypothesis, SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 41 it may be reasoned that Thyreoxenus differentiated in societies of Nasutitermes in the American tropics prior to the dispersal of these termites to the Oriental Region in pre-Tertiary times. The very close relationship of T. major of Panama and T. solomonensis of Melanesia attests to the high degree of specialization attained at this early date and the minor changes that have occurred since. (c) Subtribe Abrotelina (fig. 20) : At present this group contains about twenty-five neotropical species, associated, except in a few cases, with genera of the Nasutitermes branch: Abroteles with Nasu- titermes; the Termitoiceus complex with Diversitermes and Veloci- termes; Eburniogaster and Termitonidia with Tenuirostritermes. The Abrotelina are not characterized by a single outstanding trait, but their frail build and pattern of physogastry indicate that they comprise a distinct phyletic line within the Corotocini. The time of origin of the subtribe is somewhat doubtful, as there is still a question of whether or not the group reached the Old World. The Madagascar genus Termitochara, which I have provisionally placed in a separate subtribe, may belong to the Abrotelina. If so, this would indicate an origin early in the history of the Nasutitermes branch. Morphologically, there is much to indicate that Termito- chara is of the same phyletic line as this group; on the other hand, its occurrence (not verified) with two genera of termites of other subfamilies (Capritermes of the Termitinae and Microcerotermes of the Amitermitinae) is quite incompatible with the host relationships of the Abrotelina. The Termitoiceus generic group seems to have differentiated in stock of the Nasutitermes branch ancestral to two endemic South American genera, Diversitermes and Velocitermes. Several species have, however, been reported in association with societies of Subuli- termes (Subulitermes branch of Nasutitermitinae). As in similar cases (see discussion, p. 30) it may be necessary to postulate a host change at some time in the evolutionary history of the subtribe. An interesting group of species comprising the genera Eburnio- gaster and Termitonidia occur with colonies of the subterranean genus Tenuirostritermes of the southwestern United States, Mexico, and Central America. Inasmuch as these specialized termites excavate galleries that reach the surface of the ground beneath stones or other objects, large concentrations of termites and some guests may be exposed by turning stones. Emerson (1952a) postu- lated a Tertiary origin for Tenuirostritermes in Central America at a time when South America was isolated. Tenuirostritermes was 42 FIELDIANA: ZOOLOGY, VOLUME 40 able to enter South America to a very limited extent following the Pliocene re-establishment of a connection between South and Central America. The Central and South American Tenuirostri- termes colonies have not yet been searched for termitophiles. The Termitophilous Tribe Termitonannini In marked contrast to the Corotocini, the members of this tribe are limuloid in form and with the exception of the Brazilian genus Termitonilla are non-physogastric. It is probable that the Termitonannini evolved from free-living aleocharine stock inde- pendently of any other termitophilous group. The presence, on the terminal antennal segments, of coeloconic sensilla almost identical with those of the Corotocini may suggest a closer relationship between the two tribes than seems likely on other grounds. These sensilla, rare as they are in the Aleocharinae, may be the result of convergent development. As organized in this monograph, the tribe is comprised of the subtribes Perinthina and Termitonannina groups formerly placed in different sections of the Aleocharinae by virtue of their different tarsal and antennal segmentations. If we discount these easily- accounted-for differences, there is little to justify recognition of two tribes. A limuloid body-form does not by itself suffice to distinguish this tribe, for there are a number of other termitophilous and myrmeco- philous groups with similar body form. But termitophilous limuloid species with 4, 4, 4- or 4, 4, 5-segmented tarsi and with one or more pairs of coeloconic sensilla on their terminal antennomeres, prob- ably belong here. In numerous papers Wasmann characterized limuloid myrmeco- philes and termitophiles as "defensive type" guests and placed great emphasis on the protection that this form allegedly afforded them. The fact that at least five independently evolved groups of termi- tophilous Staphylinidae have limuloid forms would seem to be significant, but there is no reason to believe that the most obvious explanation that this form affords protection from attack by termites or ants is necessarily the correct one. It seems unlikely that present-day guests rely chiefly, or at all, on body-form for protection from termites, although it is conceivable that during an early phase of their history large pronota and elytra may have provided important protection to head, antennae, mouth parts, and legs. It is doubtful that the limuloid termitophiles could have SEEVERS: TERMITOPHILOUS STAPHYLINIDAE Perinffiodas x Porapennthus 43 Tribe Termitonannini FIG. 9. Tribe Termitonannini. survived over a long period of time and speciated so successfully without a more subtle and effective protective device. It is unlikely that the Termitonannini are synoeketes in Wasmann's sense; it seems more reasonable to believe that they have evolved a rather high degree of physiological integration with the colony. A hypothetical history of the Termitonannini may be partially constructed from available phylogenetic, biogeographical, and host data. The subtribe Perinthina is pantropical in distribution and restricted in host associations to the Nasutitermitinae. The subtribe Termitonannina is apparently limited to the American tropics and 44 FIELDIANA: ZOOLOGY, VOLUME 40 TUnervitarmos Tentarostriter/nas Velocitormes Diversitar/MS --- ( Nasutitermos Porinthina Term 'ton annino Nosutifermifinaa FIG. 10. Host relationships of Termitonannini. has a host pattern that includes Anoplotermes (Amitermitinae) and several genera of Nasutitermitinae. The following discussion pre- sents data that may be interpreted as evidence for an early tribal origin, probably in the American tropics during the Cretaceous Period. The events prior to the differentiation of the present-day subtribes are conjectural, but it is clear that only the first phases in the evolution of the limuloid form had taken place. The more specialized aspects of this modification apparently took place in- dependently in each subtribe. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 45 A. Subtribe Perinthina (figs. 9, 10, 11, 27). The evolutionary history of the Perinthina is so closely linked with that of the Nasu- titermitinae, and the Nasutitermes branch in particular, that much of the previous discussion in connection with the Corotocini is ap- plicable here. It is of particular interest to note how closely the evolutionary pattern parallels that of the Corotocini. The association of the Brazilian genus Termitopelta with the primitive nasutitermitine genus Syntermes is interpreted as evidence that the Perinthina differentiated soon after the origin of the Nasu- titermitinae if not sooner. There are certain morphological features of Termitopelta that indicate it as possibly the most generalized of present-day Perinthina. Termitopelta seems to represent the level of evolution within the subtribe prior to acquisition of the special- ized limulbid facies that characterizes almost all of its members. In contrast to the derived genera with large convex pronota that almost cover their hypognathous heads, Termitopelta has a large, relatively flat pronotum and an exposed, prognathous head. In- asmuch as the latter condition more closely resembles the typical aleocharine structure, it is presumed to be more generalized. With the exception of Termitonicus, the neotropical genera Termitocola, Perinthus, Macrognathellus, and Poduroides have a limuloid, Perinthus-\ike appearance and are evidently offshoots from the main phyletic sequence. Termitonicus, a specialized genus of unusual facies, retains the generalized prognathous condition of the head. Of the derived limuloid genera, Termitocola exhibits fewer specializations of antennae and mouth parts and has departed to a lesser degree from the main course. The neotropical limuloid genera occur with termites of the Nasutitermes branch and were doubtless closely associated with that phyletic line of termites from the time of its differentiation. Palaeotropical Perinthina are now known to occur in India, Java, Palua Islands, Fiji Islands, and the Belgian Congo. The four genera involved could have evolved from Termitocola-like stock from the Neotropical Region; all are certainly derived genera of the main phyletic sequence within the subtribe. The Indomalayan- Papuan genera Lauella and Eutermitophila differ from Termitocola of Panama in minor respects; the African genera Perinihodes and Paraperinthus are more highly specialized in some respects. The palaeotropical species have been found in societies of Nasutitermes and Trinervitermes. Available evidence thus suggests that Termi- tocola-\ike stock entered the Oriental Region during the dispersal of 46 FIELDIANA: ZOOLOGY, VOLUME 40 the Nasutitermitinae, probably during Cretaceous times. There is doubtless a moderately large undiscovered perinthine fauna in the Indomalayan Region, judging by the occurrence of Lauella in such outlying areas as the Fiji and Palau Islands. The two Belgian Congo genera, the first of the Perinthina to be recorded from the Ethiopian Region, are notable for modifica- tions of the hind coxae from the typical transverse type of the Aleocharinae to a "triangular" form characteristic of certain other termitophilous tribes (Corotocini and Termitopaediini). B. Subtribe Termitonannina (figs. 9, 10, 30). Twenty neotrop- ical species of this subtribe have now been recorded and there is as yet no evidence that the group occurs in the Old World tropics. The host relationships of its species are perhaps the most interesting, and certainly the most puzzling, feature of the Termitonannina. The host pattern includes Anoplotermes and closely allied Speculi- termes of the Amitermitinae as well as a series of genera of Nasu- titermitinae, the latter including Syntermes, Procornitermes, Corni- termes, Labiotermes, and Subulitermes. It is especially noteworthy that the nasutitermitine hosts belong either to primitive mandibulate genera or to genera of the Subulitermes branch. No species has been found with a host of the Nasutitermes branch. The more generalized genera of the subtribe Nannellus, Eunannodes, and Macrotrichurus and the remarkable physogastric genus Termitonilla occur with Anoplotermes. The large genus Termitonannus, the most highly specialized limuloid genus of the group, occurs with a variety of hosts; a majority of its species are associated with Anoplotermes while the others are recorded as guests of the nasutitermitine genera named above. The phyletic relationships of the subtribe may be summarized rather briefly. As in the case of the Perinthina, a prognathous head is presumed to be more generalized than the hypognathous one. Nannellus and Eunannodes are believed to be the most generalized of the subtribe. The physogastric genus Termitonilla has generalized head and thoracic structures and evidently differentiated from relatively generalized stock. In appearance Termitonilla is strikingly like Eburniola of the Corotocini, so that this is a most interesting case of the convergent development of physogastry. The limuloid genus Termitonannus has very specialized head structures; its head is greatly shortened and hypognathous and almost covered by the large pronotum. In view of the difficulties in correlating phyletic and host pat- terns, the evolutionary history of the Termitonannina is not easily . ~ i /. b 47 48 FIELDIANA: ZOOLOGY, VOLUME 40 reconstructed. This is especially true if we bear in mind the different host pattern of the subtribe Perinthina. Inasmuch as a large majority of its species, including the more generalized ones, occur with Anoplotermes, one can hardly escape the conclusion that this genus was probably the primary host genus for the subtribe. In the preceding section a common termitophile ancestor for the subtribes Perinthina and Termitonannina was postulated; yet the host patterns of the two tribes are hardly compatible with such a hypothesis and it is most difficult to decide what the host of such an ancestor might have been. Several possibilities are available. To account for the two host patterns without postulating one or more major changes in host associations from Nasutitermitinae to Amitermitinae or vice versa, it would be necessary to place the common ancestor in societies of Termitidae ancestral to both subfamilies. If this had been the case, one would expect the Termi- tonannina to be more widely distributed in the Amitermitinae than present evidence suggests. Future considerations of the Perinthina and Termitonannina should be carried on with two possibilities in mind. The first is that these groups may have evolved independently from different free-living aleocharine stocks. Secondly, the records of species of Termitonannina with hosts other than Anoplotermes may simply reflect faulty collecting procedures. Anoplotermes colonies, in a high percentage of cases, occur in mounds occupied by other termites, such as Syntermes and Subulitermes. Inasmuch as Anoplotermes lacks the soldier caste it would have been easy in at least some cases to have collected soldiers of the wrong host. The Termitophilous Tribe Termitohospitini Although originally proposed (Seevers, 1941) for a small group of neotropical species, the tribe Termitohospitini (figs. 12, 31) is expanded in this monograph to include a number of Old World genera formerly placed in the heterogeneous tribe Bolitocharini. It is not unlikely that stock similar to that from which this tribe of termitophiles originated may be found among the present-day free- living bolitocharines. With the exception of the subtribe Termitospectrina, whose members are perhaps slightly physogastric, this tribe is comprised of limuloid species that may be grouped in three subtribes. Tribal members may be recognized by their 4, 4, 5-segmented tarsi and their distinctive heads and mouth parts. I - g I a '~ 9 -= 1 49 50 FIELDIANA: ZOOLOGY, VOLUME 40 The South American subtribe Termitohospitina is comprised of five genera that occur with primitive mandibulate genera of Nasu- titermitinae as well as with derived genera of the Nasutilermes branch. It is difficult to arrange the Termitohospitina in a phyletic series at present. Termitohospes occurs with Cornitermes and Nasu- titermes; Termilosocius and Paratermitosocius are guests of Nasuti- termes; Blapticoxenus and Termitosodalis are associated with two derived endemic neotropical genera Velocitermes and Diver sitermes. This pattern is quite similar to the patterns of certain Corotocini and Termitonannini discussed in previous sections. The subtribe Termitohospitina almost certainly differentiated in association with primitive Nasutitermitinae; in fact, the entire tribe may have originated with the establishment of a free-living ancestor in such a colony. After the hypothetical origin of the tribe in South America, it seems probable that tribal stock dispersed to the Oriental Region with the Nasutitermitinae, as described for other groups. A small group of Indomalayan and Australian species belonging to Hetairo- termes and Termitobra comprise the subtribe Hetairotermitina. This group presumably evolved from American stock; the morphological characteristics of the species are quite consistent with this pre- sumption. In only a few instances is the host of these species known, but the occurrence of two Palau Islands species with Nasutitermes is convincing evidence that the group at least reached the Indo- malayan Region with Nasutitermitinae. Hetairotermes apparently speciated rather extensively in Australia, judged by the material now available. Three genera from widely separated parts of the world are herein grouped to form the subtribe Termitospectrina. The species of this group differ from those of other subtribes in having a typical aleocharine body-form (non-limuloid) . Their heads are prognathous, the pronota do not cover the head, and their abdomens are recurved over the foreparts. Despite these important differences, I tentatively place Termitospectrum and related genera in this tribe because of marked similarities in the structure of their mouth parts. The diver- gence of the Termitospectrina from the other subtribes, if indeed they had a common ancestry, must have been very early in tribal history certainly before the limuloid body-form was evolved in the tribe. The Termitospectrina is comprised of Termitospectrum of South America, Termitoecia of Australia, and Termitana of Madagascar. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 51 In the current classification of Aleocharinae these genera are placed in different tribes; in the case of Termitana an incorrect tribal position resulted in part from Fairmaire's miscounting of the tarsal segments. There is little about the structure, distribution, or host relationships of these genera to suggest a place of origin for the group or to account for the present distribution. Termitospectrum is known to occur with Nasutitermes in South America, but the hosts of the other genera have not been recorded. The subtribe Termitusina is provisionally established for a single African genus that occurs with Cubitermes and Noditermes of the Termitinae. Eight species of Termitusa have been proposed, but in view of the fact that about seventy species of Cubitermes have been named this may be far short of the number in existence. The species of Termitusa are limuloid in form and have mouth parts like those of Termitohospitina in many respects. Yet I am not com- pletely convinced that the genus belongs here, inasmuch as the host relationships do not conform to the tribal pattern. I have been unable to discover an adequate basis for its separation, but I have not subjected the genus to as careful a morphological study as is desirable to establish its true relationship to the Termitohospitini. TERMITOPHILOUS STAPHYLINIDAE ASSOCIATED WITH MACROTERMITINAE The Macrotermitinae The Macrotermitinae are fungus-growing termites of the Old World tropics and according to Emerson (1955) originated in the Ethiopian Region during Tertiary times. All ten genera of the subfamily occur in Africa and the more primitive genera such as Acanthotermes and Pseudacanthotermes are endemic to that continent. Only the largest and most advanced genera Macrotermes, Odonto- termes, and Microtermes reached the Indomalayan Region, pre- sumably during the Miocene. The fungus-growing termites occur as far east as Java, Sumatra, Celebes, Borneo, and the Philippines, but apparently had no opportunity to reach the Australian and Papuan regions. Some of the most spectacular termitaria in the world are con- structed by certain African Macrotermes. The mounds of M. bellicosus in the Cameroons and Belgian Congo attain heights greater than eighteen feet and diameters of more than seventy- 52 FIELDIANA: ZOOLOGY, VOLUME 40 five feet. Within the termitaria workers construct a complex system of chambers and cells in which eggs, fungus masses, and wood dust are stored. Fungus gardens are constructed by workers who masticate wood to a paste and roll it into pellets that are pressed together to form convoluted structures upon which fungus growth occurs. Fungus mycelia may bear small white spherules that are fed to reproductives and nymphs. Workers and soldiers feed primarily on grass, leaves, and wood. The termitophile fauna associated with societies of Macro- termitinae evidently originated independently of those groups that live with Nasutitermitinae. The staphylinid fauna, consisting of members of several independently derived tribes, is in no way shared by the termites of these major subfamilies. It is noteworthy, though, that the staphylinid guests of both termite subfamilies have acquired almost all the termitophile characteristics presumed to be adaptive. The termitophilous Aleocharinae occurring with Macrotermitinae are assigned in this monograph to the following tribes: Termito- paediini, Termitodiscini, and Myrmedoniini. In addition, some species of Indomalayan Pygosteninae, a predominantly myrme- cophilous subfamily, are guests of fungus-growing termites. The following discussion of macrotermitine guests is limited to the Termitopaediini and Termitodiscini. The Termitophilous Tribe Termitopaediini The tribe Termitopaediini is proposed as a new assemblage in this paper and its composition is very likely to require revision when new material becomes available. The tribe is based on a generic complex including the African genera Termitopaedia, Termitopulex, and Termitolinus, as well as the Indomalayan Dioxeuta (=Jacobson- ella), and Neodioxeuta. A second generic group is comprised of three Ethiopian genera, Prolermitobia, Termitotecna, and Termitobia. How closely allied these two groups are is still to be determined. Two other genera, Termitotropha (South Africa) and Termitobaena (Borneo), are tentatively placed in the Termitopaediini. These genera may be closely allied to three Indomalayan genera, Felda, Zunia, and Termitobiella, which I have left in the "catch-all" tribe Myrmedoniini because I have not had an opportunity to study them. The Termitopaedia group of genera occurs with Pseudacantho- termes, Macrotermes, and Microtermes, thus conforming to a con- sistent pattern. Pseudacanthotermes is a relatively primitive, SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 53 endemic African genus of fungus-growing termites; it provides some of the evidence linking the origin of this termite subfamily to the Ethiopian Region. The occurrence of Termitopaedia with Pseuda- canthotermes suggests that the Termitopaediini were established early in the evolutionary history of the Macrotermitinae. Termi- topulex, Termitolinus, Dioxeuta, and Neodioxeuta are associated with the derived genus Macrotermes; Termitopulex also occurs with Microtermes, the most specialized genus of fungus-growing ter- mites. The Termitobia group of genera presents a less consistent host situation. It is true that Macrotermes, the host of Termitobia, and Odontotermes, with which Termitotecna occurs, are fungus-growing termites, but Cubitermes, the host genus for the two known species of Protermitobia, belongs to the subfamily Termitinae. Unless one falls back on the explanation of a major shift in host affinities at some point in the history of the group, this situation is difficult to explain. Additional discoveries may help to clarify these host relationships. Our present knowledge of the hosts of Termitotropha and Termi- tobaena is even less satisfactory. Termitotropha was stated to be a guest of Amitermes (Amitermitinae), but the single, nineteenth century record requires confirmation. The host of Termitobaena is not known. The three Indomalayan genera Felda, Zunia, and Termitobiella that may be allied to these genera occur with Capri- termes (Termitinae). This fact, by itself, would not lead one to suspect close relationship. It is clear that our knowledge of the Termitopaediini is too fragmentary to allow for more than brief glimpses into the tribe's evolutionary history. There is no evidence to contradict Emerson's (1952a, 1955) view that the Macrotermitinae originated in Africa and that Macrotermes and Odontotermes moved eastward into Indo- malaya during the Tertiary, too late to reach the Australian and Papuan regions. The Indomalayan genus Dioxeuta (India, Malaya, Java, Sumatra, Borneo, the Philippines) is certainly the counterpart of the Ethiopian genus Termitopulex. These genera are relatively generalized, but Dioxeuta gave rise to a group of physogastric species (Neodioxeuta) with very remarkable abdomens (fig. 33, 6). The Neodioxeuta abdomen exhibits the same type of enlargement, in- volving chiefly the basal segments, as that of the bizarre genus Termitobia of Africa, but this is doubtless a convergence within the tribe. 54 FIELDIANA: ZOOLOGY, VOLUME 40 The Termitophilous Tribe Termitodiscini As I have indicated in the systematic section of the monograph, there is little justification for separating this group from the Aleo- charinae as Wasmann did when he proposed the subfamily Termi- todiscinae. The Termitodiscini are minute, limuloid species that occur with the fungus-growing species. Termitodiscus has been found with Odontotermes in both Ethiopian and Indomalayan regions, Discoxenus with Odontotermes in India and Ceylon, and Termitogerrus with Macrotermes in the Ethiopian Region. There is as yet hardly an adequate basis for profitable speculations about the evolutionary history of the group. TERMITOPHILOUS STAPHYLINIDAE ASSOCIATED WITH RHINOTERMITIDAE The Termitophilous Tribe Philotermitini The aleocharine tribe Philotermitini is a small group of species occurring with two genera of New World Rhinotermitidae six species of Philotermes with Reticulitermes in the eastern United States, and one species of Neophilotermes with Central American Coptotermes. The Philotermitini resemble free-living Staphylinidae in appearance, and exhibit no adaptive features usually associated with termitophily. There can be little doubt, however, that the species are well integrated with termite societies and none at all about the obligatory nature of the relationship. Speculation concerning the origin and evolution of this small group is probably fruitless at this time. The tribe appears to be an endemic American group but this may merely reflect the small attention given to colonies of Old World Rhinotermitidae. Inas- much as Reticulitermes is a Holarctic genus with about as many species in Eurasia as in the United States and Mexico, and Copto- termes is a tropicopolitan genus of nearly fifty recorded species, it is difficult to believe that the present distribution of the tribe represents its true range. The Termitophilous Subfamily Trichopseniinae All members of this unusually interesting subfamily are guests of termites. Although only nine genera have as yet been discovered, the subfamily is known to occur in every major zoogeographic area (fig. 13) : Neotropical, Nearctic, Palaearctic, Indomalayan, Papuan, OS 1 c H 55 56 FIELDIANA: ZOOLOGY, VOLUME 40 Australian, and Ethiopian Regions. Its species have been recorded with termites of several families but it seems likely that the primary association was with the Rhinotermitidae. It would be very difficult if not impossible to describe a sub- family facies, inasmuch as the genera present a diversity of body- forms. The less specialized genera are non-physogastric while others exhibit moderately strong physogastry. One form of speciali- zation was the development of a pronounced limuloid form. A series of photographs shows this diversity to good advantage: Trichopsenius (fig. 40, a, b), Hamitopsenius (fig. 41, c), Rhino- termopsenius (fig. 40, d), Xenistusa (fig. 40, c), and Mastopsenius (fig. 41, a). Trichopsenius of North America and Japan may possibly be the most generalized of the known genera; its species are semi-limuloid in form, with one species showing incipient physo- gastry. Its North American relative Xenistusa is relatively general- ized as to head structure and has a feebly physogastric abdomen. Mastopsenius of Australia has a remarkable fusiform body, including a physogastric abdomen. Schizelythron of Java is noteworthy for its bizarre elytra, which are split for almost their entire length. The extreme limuloid body-form is displayed by the Indomalayan, Papuan, and Australian Hamitopsenius as well as by its American ally, Termitopsenius. These two genera are so close that it is questionable whether they should be separated. An extremely interesting new Ethiopian genus, received too late to describe in this paper, is the first representative in that region. The Trichopseniinae, although diverse in appearance, are very easy to recognize by the very distinctive relationship of the hind legs to the metasternum (fig. 42). The large hind trochanters appear to articulate with the metasternum, but this can only mean that the coxae are indistinguishably fused to the metasternum. The reader who is interested in more details of this unique characteristic is referred to an earlier paper (Seevers, 1941). The head of the Trichopseniinae also has some unusual features. The adaptive significance of the hind leg modification is not evident from its morphology, but a clue may be provided by a be- havior trait observed in British Guiana by Dr. Emerson some years ago. He noticed that two species of Rhinotermes guests (Termito- psenius acanthoscelis Seevers and Rhinotermopsenius saltatorius Seevers) were able to leap for surprising distances, considering their very small size. Leaps up to three inches were observed. It is difficult to appraise the value of this behavior as an escape mechan- SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 57 ism, but it might have played an important role in the evolution of the subfamily. It is not known how widespread this behavior is within the family, but it might well have been associated, at least early in the evolution of the group, with the singular mode of leg articulation. It is difficult, with the information now available, to reconstruct the phyletic history of the subfamily, or to relate this group to free- living Staphylinidae. Inasmuch as fourteen of the seventeen known species of the subfamily occur with species of Rhinotermitidae, it is reasonable to consider this group their primary hosts. Two of the three records involving termites of other families (Mastotermes, Amitermes) are based on single records and require confirmation; the other record (Neocapritermes) is probably correct but is rather easily accounted for. There seems to be no evidence at present favoring any one region as a center of origin for the Trichopseniinae, although the alternatives almost certainly include only the Americas and the Indomalayan Region. Although most conclusions made at this time concerning the evolution of this subfamily are likely to be premature, it is in- structive to examine the host relationships of the Trichopseniinae to determine whether or not a pattern is discernible. Emerson (1955) places the center of origin of the Rhinotermitidae in the Indomalayan Region during pre-Cretaceous times. The family is comprised of two major phyletic lines (Ahmad, 1950); one branch contains Prorhinotermes, Schedorhinotermes, Rhinotermes, and allied genera; the other line gave rise to Heterotermes, Coptotermes, Reticu- litermes, and relatives. Trichopseniine guests are found with mem- bers of both phyletic lines, a fact suggesting an ancient origin for the subfamily. The first of the above-mentioned phyletic lines has more recorded termitophile genera at present: Schedorhinotermes is the host of Schizelythron in Java as well as an undescribed genus of the Belgian Congo; Prorhinotermes harbors a species of Hamitop- senius in Micronesia; and the neotropical Rhinotermes has three associated genera in British Guiana Megaxenistusa, Termitopsenius, and Rhinotermopsenius. Of the other phyletic line, Reticulitermes is the only genus known to harbor trichopseniine termitophiles; Xenistusa occurs in the southern United States and Trichopsenius in North America and Japan. There is no satisfactory explanation as to why such large tropical genera as Heterotermes and Coptotermes do not have guests of this subfamily. Inadequate collecting could easily be responsible for failure to record them. 58 FIELDIANA: ZOOLOGY, VOLUME 40 The three records of Trichopseniinae with termites other than Rhinotermitidae do not seriously distort the host pattern that is beginning to become apparent. Two of these records involve the Hamitopsenius-Termitopsenius branch of Trichopseniinae. Species of these extremely specialized limuloid genera are known for certain to occur with Prorhinotermes in the Palau Islands (Micronesia) and with Rhinotermes in British Guiana. A Sumatran species of Hami- topsenius was recorded with Amitermes by Wasmann, but this could easily reflect uncritical collecting of host material and requires substantiation. On the other hand, the occurrence of the Argentine species Termitopsenius limulus with Neocapritermes (Termitidae) has been confirmed, and inasmuch as the British Guiana species of Termitopsenius is a Rhinotermes guest, it seems likely that in this case a host change was involved. The most puzzling case of all is that of the Australian species Mastopsenius australis, which occurs with Mastotermes darwiniensis, the only living species of the primitive family Mastotermitidae. This situation seems to defy explanation at this time. III. The Termitophilous Staphylinidae: Systematic Revision MATERIAL AND METHODS This revision deals with approximately three hundred species of termitophilous Staphylinidae, exclusive of synecthrans such as Zyras. At least nine-tenths of the genera and four-fifths of the species were examined during the course of the studies. Perhaps one-half of the unavailable species were relatively well described and figured by Silvestri and present few problems of placement. Type material of almost all species described by Wasmann, Mann, Borgmeier, Bernhauer, Cameron, Casey, Fenyes, Reichensperger, Sharp, Fauvel, and Lea was examined at one time or another. My own collection, consisting of a high percentage of the termitophilous species treated in the revision, is now incorporated in the collection of Chicago Natural History Museum. Dr. Alfred Emerson provided a substantial part of the material for these studies; some he collected and some he acquired from other termite specialists and collectors. The following is a summary of the principal sources of the termitophilous specimens and the regions from which the material came: A. E. Emerson (British Guiana, Panama, Belgian Congo, United States) ; A. Reichensperger (Brazil, Tanganyika); R. L. Araujo (Brazil); A. M. Adamson (Trinidad); W. M. Mann collection, United States National Museum (Bolivia, Central America, Liberia); H. S. Dybas collection, Pacific Science Board Expedition (Micronesia); M. Bernhauer collection, Chicago Natural History Museum (Indomalaya, Africa, Brazil) ; M. Cameron collection, British Museum (Natural History); E. Wasmann col- lection, Natuurhistorisch Museum, Maastricht, The Netherlands (world-wide); A. Fauvel collection, Institut Royal des Sciences Naturelles de Belgique. All available host termites (except as noted in the systematic section) were determined by Dr. Emerson. Host records are sum- marized in lists at the end of the paper. Many generic names and some specific names of termite hosts are not the same as those cited 59 60 FIELDIANA: ZOOLOGY, VOLUME 40 by Wasmann and other early writers due to subdivision of the broad generic categories then in use. The termite nomenclature agrees closely with that of Snyder's (1949) catalogue of the termites. The specimens studied have been on points, in alcohol, and on slides after treatment with potassium hydroxide. Measurements of parts of the body are generally given in terms of an arbitrary unit (0.009 mm.), the measurements having been made with an ocular grid calibrated with a stage micrometer. The measurements are useful primarily for comparative purposes and enable one to judge the relative lengths of such structures as antennal segments. The actual lengths of such small structures are generally of little concern, but they may easily be obtained by converting the units to fractions of a millimeter. Although the measurements are useful for comparative purposes it should be borne in mind that slight differences in antennal and tarsal lengths, for example, are difficult to measure. There are not as many figures of structural details as I should have liked to provide. Inasmuch as a high percentage of previously described genera and species were illustrated in some detail, it is not deemed necessary to repeat the figures here. Reference should be made to papers by Silvestri, Wasmann, Kemner, Warren, and Seevers. The termitophilous Staphylinidae are far better illustrated than are most free-living species of the family. Some readers may wonder why the aedeagi of males were not utilized to a greater extent for diagnostic purposes. These structures have not proved to be especially useful either for taxonomic purposes or for studies of phylogeny. There are some differences in the aedeagi that may prove to be of greater significance in future studies. THE STAPHYLINIDAE: GENERAL CONSIDERATIONS The Staphylinidae comprise one of the largest groups of insects, numbering more than 25,000 species distributed throughout the world. A large majority of these species are predators and feed chiefly on other insects that occur in carrion, dung, and decaying organic materials. The ecological role of the Staphylinidae must be of considerable importance in many microhabitats, inasmuch as their habits of predation must influence the populations of those habitats to a significant degree. Some genera are fungivorous and others apparently saprophytic. The family Staphylinidae is divided into about twenty sub- families; some are well-defined categories, others of dubious merit. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 61 Species of at least nine subfamilies have been listed as termitophiles, but only the Aleocharinae, Trichopseniinae, and Pygosteninae have termitophilous groups of significance. The recorded species of other subfamilies, with the exception of a few species of Hypocyphtinae and Tachyporinae, do not conform to the definition of a termitophile given earlier in the paper. A list of the doubtful species is given at the end of the paper; some of these are probably synecthrans but most are accidental associations. Single records of species belonging to large genera of free-living forms are particularly suspect. KEY TO SUBFAMILIES WITH TERMITOPHILOUS SPECIES 1. The large hind trochanters appearing to articulate directly with metasternum (fig. 4'2); metasternal plates extending laterally from mid-line over bases of femora; paratergites absent Trichopseniinae Relationship of hind leg to metasternum generalized; no metasternal plates present ; paratergites present 2 2. Antennae inserted medial to the eyes 3 Antennae inserted at front or side margin of head 4 3. Antennae thick, spindle-shaped, apex tapering to point. Pygosteninae (and some Termitodiscini) Antennae otherwise Aleocharinae 4. Antennae 10-segmented Hypocyphtinae Antennae 11-segmented Tachyporinae Subfamily ALEOCHARINAE The subfamily Aleocharinae, to which a large majority of the termitophilous Staphylinidae belong, is an extremely large assem- blage of relatively small species, averaging perhaps 5 mm. in length. So great is the number of aleocharine species and individuals that they frequently dominate microhabitats. The aleocharines are doubtless of major importance as predators in a variety of decaying animal and vegetable materials where they feed on dipterous larvae and other insects. They are abundant on cadavers, in dung, in decaying fruit and fungi, in the leaf mold of the forest floor, in de- caying logs and under bark, on stream and pond shores, in the intertidal zone of ocean shores, in nests of some birds and mammals, on fresh mushrooms, and in a variety of other habitats. The Aleocharinae, in the course of their evolution, have moved into ecological niches and have speciated to a degree rivaled by few insect groups at the subfamily level. In 1831 Mannerheim proposed the Aleocharides as a sixth tribe of Brachelytra. Soon after, Erichson (1837, 1840) established the classification of the Staphylinidae in essentially its present form, and recognized the Aleocharini as a major subdivision. Aleocharine 62 FIELDIANA: ZOOLOGY, VOLUME 40 specialists since Erichson's time have contributed little of funda- mental importance to the organization of generic groups within the subfamily and have retained the artificial features on which the tribal classification was based. The tribal organization currently in use was proposed by Ganglbauer (1895) and the twentieth century specialists have expanded but scarcely modified the tribal arrangement (Fenyes, 1918-21; Bernhauer and Scheerpeltz, 1926; Cameron, 1939). The first termitophilous Staphylinidae to be discovered were the highly specialized South American genera, Corotoca and Spir- achtha. Schi0dte (1853) was even able to report that Corotoca females produce living larvae. Kraatz added the American genus Philotermes in 1857 but it was nearly forty years later that Wasmann published the first of an extensive series of reports dealing with termite and ant guests. At the beginning of the present century Silvestri also began a series of noteworthy publications on termite guests. It is somewhat surprising that many of the men who described genera and species of staphylinid termitophiles Schi0dte, Wasmann, Silvestri, Tragardh, Warren, Kemner, Reichensperger, Mann, and Borgmeier were not primarily students of the Staphy- linidae, whereas the aleocharine specialists Casey, Fenyes, Bern- hauer, and Cameron did not make large contributions to our knowledge of the termite guests. The present-day organization of aleocharine genera into tribes and subtribes is so artificial that it almost completely obscures interesting evolutionary relationships. A complete revision of generic groups with phylogenetic considerations uppermost in mind is badly needed. The termitophilous genera have been assigned in bewildering fashion to tribes that included free-living as well as myrmecophilous species. I have found it necessary to shift the positions of so many genera that there is little resemblance between my proposed classification and that currently in use. KEY TO TERMITOPHILOUS TRIBES OF ALEOCHARINAE OCCURRING IN THE NEW WORLD 1. Mentum fused to submentum; hind coxae triangular; metepimera ab- breviated; mesocoxal acetabula not margined; physogastric Corotocini Species without the above combination of characters 2 2. Tarsi 4, 4, 4-segmented; stenogastric species Perinthina (Termitonannini ) Tarsi 4, 4, 5 or 4, 5, 5-segmented 3 3. Tarsi 4, 5, 5-segmented Termitozyrina ( Myrmedoniini ) Tarsi 4, 4, 5-segmented . . 4 SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 63 4. Antennae with 10 segments or less Termitonannina (Termitonannini) Antennae 1 1-segmented 5 5. Labial palpi very much elongated, filiform Termitohospitini Labial palpi neither elongated nor filiform ... ... Philotermitini KEY TO TERMITOPHILOUS TRIBES OF ALEOCHARINAE OCCURRING IN THE OLD WORLD 1. Mentum fused to submentum; hind coxae triangular; metepimera ab- breviated; mesocoxal acetabula not margined; physogastric Corotocini Species without the above combination of characters . . 2 2. Tarsi 4, 4, 4-segmented ; limuloid species Perinthina (Termitonannini) Tarsi 4, 4, 5 or 4, 5, 5-segmented 3 3. Tarsi 4, 4, 5-segmented Termitohospitini Tarsi 4, 5, 5-segmented 4 4. Head covered by large, shield-shaped pronotum Termitodiscini Head visible from above 5 5. Body-form limuloid 6 Body-form otherwise 7 6. Front of head with transverse cariniform line Pseudoperinthini Front of head without such a line Termitendina (Myrmedoniini) 7. Physogastric species Termitopaediini or Feldina (Myrmedoniini) Non-physogastric species 8 8. Galea and lacinia elongated; maxillary sinuses very large. . . . Myrmedoniina (Myrmedoniini) Galea and lacinia not elongated; maxillary sinuses not appreciably enlarged. Athetini Tribe COROTOCINI Tribe Corotocini: Fenyes, 1918-21, Gen. Insect., 173: 17, 61. The distinctive characteristics of this tribe were discussed at some length in an earlier section of the paper (p. 26). The species are grouped in five subtribes. KEY TO SUBTRIBES OF COROTOCINI 1. Abdomen (fig. 14, a-d) not recurved over foreparts; abdomen strongly inflated and very membranous Timeparthenina Abdomen recurved over foreparts . . 2. Hind coxae (fig. 21, b) loosely articulated to metasterna; median metasternal articulating processes prolonged so that hind coxae are not contiguous with metasternum . Corotocina Hind coxal-metasternal relationship generalized 3 64 FIELDIANA: ZOOLOGY, VOLUME 40 3. Sixth and seventh tergites as in fig. 19, d; seventh unusually large, shield- shaped; some genera with large, ovate, membranous abdomens; Old World tropics Termitogastrina (part) Sixth and seventh tergites otherwise; never with large, ovate, membranous abdomens 4 4. Tarsi 4, 4, 4-segmented 5 Tarsi 5, 5, 5-segmented (the two terminal tarsal segments frequently fused or partly so, but recognizable as segments) 6 5. Antennae cylindrical, 10-segmented; neotropics Genus Abroteles Casey Antennae 11-segmented; Madagascar Termitocharina 6. Frail species; integuments weakly sclerotized; tarsi 5, 5, 5-segmented, the fourth and fifth segments free Abrotelina (part) Relatively robust species with heavily sclerotized integuments; the terminal segment of the 5, 5, 5-segmented tarsi partly fused to fourth segment and not movable Termitogastrina (part) Subtribe TIMEPARTHENINA The genera that comprise the Timeparthenina are distinguished primarily by their pattern of physogastry. The abdomens of these genera are strongly inflated and present appreciable areas of mem- branous integument, but they are especially noteworthy because they are not recurved over the foreparts as are the abdomens of the other subtribes of Corotocini. The known genera illustrate various degrees of physogastric development, showing a definite trend toward enlargement of the basal abdominal segments (figs. 4, 14). The least specialized condition is that of Termituncula, while the most specialized condition is illustrated by Timeparthenus. The basal abdominal segments of the latter genus are extremely large. The variations in tarsal and antennal segmentation within the subtribe are rather striking; the tarsal segments number 5, 5, 5 or 4, 5, 5 or 4, 4, 5; the antennae consist of eleven or ten segments. A minor deviation from the pattern of characters that distinguish the Corotocini is shown by Termituncula and Termitozophilus, which have free mentums in contrast to the usual mentum-sub- mentum condition. Fenyes (1921, p. 34) first proposed the tribe Timeparthenini to receive Timeparthenus, but this served only to isolate the genus and to obscure its relationship to other termitophilous genera. TERMITUNCULA Borgmeier Termituncula Borgmeier, 1950, Rev. Ent., 21 : 662. Type species: Termitun- cula gracilipes Borgmeier. Head, pronotum, and elytra generalized in form; body rather frail. Antennae 11-segmented; elongated; segments 1-7 distinctly longer than broad; 8-10 sub- SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 65 equal in width and length. Mentum-submentum distinctive. Maxillary palpi slender; segment 3 moderately thick, segment 4 slender, elongated, as long as 3. Abdomen distinctive; elongato-ovoidal, two to three times as broad as fore- parts; width less than the dorso-ventral thickness; with moderately deep inter- segmental constrictions. Segments 3-7 physogastric; membranous integument extensive; all sclerites widely separated. Second abdominal segment not inflated, second sternite not present; segments 3-5 increasing in size, 6-7 decreasing in size. Tergites small, the seventh a little longer than the others; paratergites narrow, strap-like. Sternites generalized, larger than tergites. Legs long and slender; femora and tibiae nearly equal in length. Tarsi 5, 5, 5-segmented. Termituncula gracilipes Borgmeier. Figure 14, a. Termituncula gracilipes Borgmeier, 1950, Rev. Ent., 21 : 664, text figs. 39-51 (Campinas, Goids, Brazil; Corniiermes similis Hagen; Borgmeier coll.). Material examined. One paratype from same termite colony as holotype. The host is Procornitermes araujoi Emerson, determined by Emerson after Borgmeier's paper was published. Head, thorax, elytra, and tergites bright yellow-brown, sternites yellow, legs and antennae yellowish. Head shining, not punctured; pronotum and elytra very finely and sparsely punctured, without pubescence. Head with 2 vertexal setae; pronotum with 6 setae on apical margin, 2 along lateral border, and 2 tiny hairs in the middle of the basal margin. Elytra with one bristle near anterior angle and one on side border near hind angle. Tergites 2-6 with 6 bristles on apical margin; tergite 7 with one at each hind angle; tergite 8 with one sub-basal seta and an apical row of 8 bristles. Each paratergite with 2 or 3 bristles. First antennal segment thickened, longer than the following segments; segment 3 shorter than second and a little longer than fourth; segments 2-7 longer than broad, 8-10 subquadrate. Pronotum subequal in length and width. Length, 2.6-3 mm. TERMITOZOPHILUS Silvestri Termitozophilus Silvestri, 1901, Boll. Mus. Zool. Torino, 16: 7; 1903, Redia, 1: 192; Fenyes, 1920, Gen. Insect., 173B: 348. Type species: Termi- tozophilus laetus Silvestri. Corymbogaster Mann, 1923, Zoologica, 3: 346; Borgmeier, 1950, Rev. Ent., 21 : 640. Type species: Corymbogaster miranda Mann. Head and thorax slender; abdomen extremely broad, about four times as broad as foreparts. Head slightly longer than broad; dorsum unmodified. Gula broad, its sides scarcely converging in front; submentum and mentum distinct. Labial palpi slender. Maxillary palpi slender, segments 2 and 3 almost spindle- shaped. Antennal segments verticillate. Pronotal length subequal to maximum width; base and apex arcuate, sides sinuate, broadest at anterior angles. Legs long and slender; tarsi 5, 5, 5-segmented. Abdomen very distinctive (fig. 14, d); twice as broad as thick dorso-ventrally; segment 2 not inflated; second sternite present. Tergites, paratergites, and sternites augmented by secondary sclero- tizations of the adjacent integument; the sclerites proper are clearly defined by the fact that they bear setae. 66 FIELDIANA: ZOOLOGY, VOLUME 40 Termitozophilus laetus Silvestri. Figure 14, c, d. Termitozophilus laetus Silvestri, 1901, Boll. Mus. Zool. Torino, 16, no. 398, p. 8 (Tacurupucu, Paraguay, and Coxipo, Mato Grosso, Brazil; Corni- termes similis Hagen); 1903, Redia, 1 : 193, pi. 5, fig. 252, pi. 6, figs. 255- 256; Bruch, 1931, Rev. Ent., 1 : 388, fig. 2. Material examined. A large series of specimens from 25 colonies of Cornitermes cumulans Kollar, one colony of Cornitermes snyderi Emerson, and one colony of Cornitermes bequaerti Emerson, collected by R. L. Araujo at various localities near Sao Paulo, Brazil; a series from four colonies of Cornitermes cumulans Kollar, from Passa Quatro, Minas Gerais, Brazil. This species was recorded from Posadas, Misiones, Argentina, by Bruch (1931). The Cornitermes similis Hagen as determined by Silvestri was Cornitermes cumulans Kollar. Color reddish-brown, sternites sometimes paler; membranous integument white. Head with one seta medial to each eye and a pair of occipital setae. Pronotum with 8 setae on apical margin, 2 on each lateral margin, and 2 on disk. Each elytron with about 8 long setae. Tergites 3-6 with apical rows of 6 setae; tergite 7 without setae; tergite 8 with two rows of short setae, about 6 in each row. Inner paratergites of segments 3-7 with 5, 10, 9, 11, and 8 setae, respectively; outer paratergites with 5, 9, 9, 9, and 6 setae, respectively. Sternites with three rows of macrosetae; an apical row of about 10, a middle row of 6-8, and a basal row of about 8. First antennal segment a little more than twice as long as broad, thicker than the other segments; segments 2-4 long and slender, subequal, each a little shorter than the first; segments 5-10 increasing very slightly in width, each about twice as long as broad; segment 11 more than twice as long as segment 10, almost four times as long as broad. Length, 3 mm. (abdomen extended). Termitozophilus mirandus Mann Corymbogasler miranda Mann, 1923, Zoologica, 3: 347, text fig. 35 (Kartabo, British Guiana; Cornitermes pugnax Emerson; United States National Museum). Material examined. Type series. Head, pronotum, and elytra light reddish-brown, sternites rufo-testaceous, their anterior margins brown; tergites pale reddish-brown. Head with two setae behind and slightly medial to each eye; pilosity of pronotum, elytra, and tergites as in laetus. Inner paratergites 2-7 with 2, 3, 2, 4, 3, 4 setae, respectively; outer paratergites 3-7 with 3, 4, 4, 4, 5 setae, respectively. Antennae (12: 6: 6: 4: 4: 5: 5: 4: 4: 4: 16) strongly incrassate, segments 1-4 very slender, 5-10 increasing FIG. 14. (a) Termituncula gracilipes Borgmeier. (b) Autuoria elegantulum Silvestri. (c) Termitozophilus laetus Silvestri; stenogastric, winged individual, (d) Termitozophilus laetus Silvestri; physogastric individual. 67 68 FIELDIANA: ZOOLOGY, VOLUME 40 in width, segment 3 only two-fifths as broad as segment 10; segment 1 slender, its maximum width less than one-half its length, its posterior surface flattened; segments 2 and 3 slightly longer than broad; segment 4 small, subquadrate; 5 and 6 larger than 4, subquadrate; 7-10 slightly broader than long. Length, 2.1 mm. AUTUORIA Silvestri Autuoria Silvestri, 1946, Comm. Pont. Acad. Sci., 10: 309. Type species: Autuoria elegantulum Silvestri (not A. orthocephali Silvestri, MS., desig- nated by Silvestri). Differing from Termituncula and Termitozophilus in having 10-segmented antennae, 4, 5, 5-segmented tarsi, and distinctive abdominal structure. The abdomen is intermediate in degree of physogastry between Termituncula and Timeparthenus; the second segment is much inflated and pushes the elytra into a vertical position, the second tergite is small, but the second sternite is apparently absent. The abdomen is more compressed dorso- ventrally than in Timeparthenus, and the sternites and tergites are more generalized and not spaced so far apart. Autuoria elegantulum Silvestri. Figure 14, 6. Autuoria elegantulum Silvestri, 1946, Comm. Pont. Acad. Sci., 10: 310, text figs. V, 1-17 (Jabaquara, Sao Paulo, Brazil; Cornitermes "autuorii Sil- vestri"; Silvestri coll.). Material examined. Ten specimens, Sao Paulo (Ipiranga), Brazil, collected by R. L. Araujo, with Paracornitermes emersoni Araujo (determined by R. L. Araujo). It is possible that "Corni- termes autuorii" Silvestri (a manuscript name) is the same as Para- cornitermes emersoni Araujo, as this termitophilous species occurs in colonies of both. Head, thorax, elytra, and tergites bright yellow-brown, sternites yellow, legs and antennae yellow. Head shining, not punctured; pronotum and elytra very finely and sparsely punctured, without pubescence. Head with 2 vertexal setae; pronotum with 6 setae on apical margin, 2 on lateral border, and 2 tiny hairs in the middle of the basal margin. Elytra with one bristle near anterior angle and one on side border near hind angle. Tergites 2-6 with 6 bristles on apical margin; tergite 7 with one at each hind angle; tergite 8 with one sub-basal seta and an apical row of 8 bristles. Each paratergite with 2 or 3 bristles. Antennae not incrassate, the scape about equal to segments 2-4 combined, segments 2 and 3 subequal, perhaps slightly longer than broad, segment 4 a trifle shorter than 3; segments 4 and 5 subequal; segments 2-7 longer than broad, 8-10 subquadrate. Pronotum as broad as long. Length, 2.6-3 mm. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 69 TIMEPARTHENUS Silvestri Timeparthenus Silvestri, 1901, Boll. Mus. Zool. Torino, 16, no. 398, p. 10; 1903, Redia, 1 : 196; 1946, Comm. Pont. Acad. Sci., 10: 299; Fenyes, 1918, Gen. Insect., 173A: 75. Type species: Timeparthenus regius Silvestri. This genus has not been available for study. Silvestri's figures are so complete that identification of the genus should not be difficult. The distinctive abdomen and 4, 4, 5-segmented tarsi distinguish this genus from the preceding genera of the subtribe. The antennae are 10-segmented, and the terminal segment has two pairs of polytrichous sensilla. The abdomen is strongly hyper- trophied, ovate, and incapable of being reflected over foreparts. Segments 2-5 are greatly expanded, segments 6 and 7 are less so, and segments 8 and 9 are small. The second segment is so large and extends so far forward that it obscures a view of the thorax from above. Tergites and sternites 3-6 are small, the paratergites are very small, and all sclerites are widely separated. Timeparthenus regius Silvestri Timeparthenus regius Silvestri, 1901, Boll. Mus. Zool. Torino, 16, no. 398, p. 11 (Coxip6, Cuyaba, Mato Grosso, Brazil; Silvestri coll.; Anoplotermes tenebrosus Hagen); 1903, Redia, 1: 197, pi. 6, figs. 261-264; Hegh, 1922, Les Termites, fig. 412; Silvestri, 1946, Comm. Pont. Acad. Sci., 10: 301, text figs. I, 1-3, II, 1-14. Silvestri presented detailed figures of this and the following species but did not contrast them very well. There should be little difficulty in identifying specimens by comparing them with the figures. Anoplotermes hageni Snyder and Emerson (= tenebrosus Silvestri) is the new name for the host. Timeparthenus oglobini Silvestri Timeparthemus (sic) oglobini Silvestri, 1946, Comm. Pont. Acad. Sci., 10: 303, text fig. Ill, 1-11 (Loreto, Misiones, Argentina; Anoplotermes pacificus Fr. Miiller; Silvestri coll.). Anoplotermes bequaerti Snyder and Emerson (= pacificus Fr. Miiller) is the new name of the host. PTOCHOLELLUS Silvestri Ptocholellus Silvestri, 1946, Comm. Pont. Acad. Sci., 10: 305. Type species: Ptocholellus mimus Silvestri. Ptocholellus has not been available for study. Silvestri's diagnosis of this genus is not convincing but the long description and detailed 70 FIELDIANA: ZOOLOGY, VOLUME 40 figures suggest a number of small differences that may separate it from Timeparthenus. I shall make no attempt to evaluate these differences or to present a diagnosis. Silvestri states that the antennae are closer together than in Timeparthenus and the tergal glands of the seventh abdominal segment are widely separated. Ptocholellus minuis Silvestri Ptochokllus mimus Silvestri, 1946, Comm. Pont. Acad. Sci., 10: 308, text fig. IV, 1-28 (Lagoa Santa, Minas Gerais, Brazil; Anoplotermes ? turricola Silvestri; Silvestri coll.). Subtribe TERMITOGASTRINA The neotropical genera of this subtribe were formerly placed in the Caloderae group of Oxypodini (Fenyes, 1918-21), or in the subtribe Termitogastri of the Aleocharini (Bernhauer and Scheer- peltz, 1926). The palaeotropical genera were scattered through several tribes, including the Hygronomini and Termitoptochini. Although the group doubtless forms an evolutionary unit it is not easy to characterize; this is due to the fact that the Termitogastrina include such different genera as the generalized Termitophya and the highly specialized Paracorotoca. A discussion of the phylogenetic relationships within the subtribe is presented elsewhere in the paper. GENERIC GROUPS A. Genera having the sixth and seventh tergites in a relatively generalized condition: The Termitophya group, including all the neotropical genera and two Old World genera, Termitotima Wasmann and Idiogaster Wasmann. B. Genera having distinctively specialized sixth and seventh tergites (fig. 19, rf) but not having the abdomen greatly inflated or predominantly membranous: The Termitella group, including four Old World genera, Termitella Wasmann, Termitellodes Seevers, Affinoptochus Kemner, and Idioptochus Seevers. C. Genera having the sixth and seventh tergites as above and having large, ovate, predominantly membranous abdomens (fig. 19, e): The Termitoptochus group, including the Indomalayan Termitoptochus Silvestri and the African Paracorotoca Warren and Millotoca Paulian. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 71 KEY TO NEOTROPICAL GENERA OF TERMITOGASTRINA 1. Inner margins of elytra contiguous; apex of elytra moderately to strongly emarginate (sutural length in Termitopithus only about one-half as long as outer margin) 2 Inner margins of elytra divergent a short distance behind scutellum 7 2. Pronotum generalized, i.e., transverse, evenly convex, not impressed, sub- glabrous; seventh and eighth sternites with finely fluted sculpture. Termilophya Wasmann Pronotum and sternites otherwise 3 3. Pronotum so impressed as to be saddle-shaped; pronotum reddish-brown, subglabrous, elongated Termitopithus, new gen. Pronotum otherwise 4 4. Abdomen strongly inflated; outer paratergites separated from inner para- tergites and from sternites by membranous integument 5 Abdomen broad and flat, with little membranous integument exposed; paratergites, tergites, and sternites contiguous; each inner paratergite about two-fifths as broad as tergite 6 5. Mentum-submentum as in fig. 18, k; pronotum with a narrow transverse impression terminating in pits Neotermitogaster Seevers Mentum-submentum as in fig. 18, a-j; pronotum impressed differently. Termitogasler Casey 6. Seventh tergite one-third as long as broad Termitosyne, new gen. Seventh tergite two-thirds as long as broad Termitosynodes, new gen. 7. Pronotal disk not impressed; gula very narrow at some point. Termitomorpha Wasmann Pronotal disk impressed; gula moderately broad 8 8. Elytra with a longitudinal carina laterally Xenopelta Mann Elytra without carina 9 9. Clypeus with a prominent medial horn Termitoides Seevers Clypeus without medial horn 10 10. Entire pronotal disk impressed Xenogaster Wasmann Pronotal disk not broadly impressed although with several minor impressions. Trachopeplm Mann TERMITOPHYA Wasmann Termitophya Wasmann, 1902, Ent. Tijd., 45: 95; Fenyes, 1920, Gen. Insect., 173B: 346; Seevers, 1937, Ann. Ent. Soc. Amer., 30: 19. Type species: Termitophya heyeri Wasmann. Head capsule relatively generalized; clypeus strongly declivous or deflexed from a point behind the antennal fossae. Antennae geniculate; scape compara- tively short, about equal to segments 3 and 4 together; segments 2-10 variable. Mentum-submentum (fig. 17, a-i) uniform in shape except for that of T. arnica. Maxillary palpi relatively slender; segment 2 cornucopia-shaped, somewhat compressed; segment 3 ovoidal, usually more than twice as long as broad. Pronotum about one-fifth broader than long; uniformly convex, never im- pressed; sides moderately arcuate, basal angles obsolete; two submarginal and one marginal seta present; surface smooth, shining, very minutely punctulate, glabrous except for extremely fine, sparse stubble. Elytra contiguous; apex shallowly emarginate. Abdomen feebly to moderately physogastric; non-sclero- 72 FIELDIANA: ZOOLOGY, VOLUME 40 tized integument exposed to a limited extent in some species. Inner paratergites elevated moderately; ventral surface (sternites and outer paratergites) uniformly convex. Surface of sternites 7 and 8 distinctively fluted, i.e., with numerous longitudinal ridges and furrows. Remarks. Termitophya is comprised of the most generalized species of the subtribe Termitogastrina as evidenced by the following features: Head capsule without modifications; feebly geniculate antennae; gula and mentum-submentum not separated by a suture; maxillary palpi relatively generalized ; pronotum and elytra general- ized; abdomen not strongly physogastric. The species of Termitophya present a uniform appearance but may be distinguished by variations in the following: proportions of antennal segments, sculpture and pilosity of head, pilosity of elytra, pilosity and sculpture of sternites, and shape and pilosity of mentum-submentum. KEY TO SPECIES OF TERMITOPHYA 1. Each elytron with two pairs of erect setae; two supraorbital setae present. punctata Mann Each elytron with one pair of erect setae (three setae in heyeri); one supra- orbital seta or none present 2 2. Head coarsely reticulated; sternites 3-6 reticulated 3 Head not reticulated (finely and obsoletely so in flaviventris) ; sternites 3-6 not reticulated 4 3. Fine punctures of head obscured by reticulation; sternites 3-5 glabrous, or nearly so, at middle arnica Mann Coarse punctures of head clearly visible; sternites 3-5 with a vestiture of pale erect hairs throughout piliventris Seevers 4. Median area of sternites 3-5 not different in vestiture from more lateral areas: the hairs moderately dense, uniformly distributed, recumbent, and rather pale; surface of sternites 3-5 distinctly punctulate. holmgreni Wasmann A wide median area (about two-thirds) of sternites 3-5 differing in vestiture from more lateral areas: either glabrous or with sparse, semi-erect hairs. ... 5 5. Head coarsely punctate, the punctures rather deep and clear-cut; surface between punctures shining and without sculpture; median areas of sternites 3-5 with a vestiture of sparse, erect hairs (obvious only when abdomen is viewed in profile) 6 Punctures of head not coarse, deep, and clear-cut (in some species they may be moderately coarse, but are shallow and indistinct); median area of ster- nites 3-5 almost glabrous (third sternite with a few short semi-erect hairs) . . 7 6. Chaetotaxy of mentum-submentum as in fig. 17, a heyeri Wasmann Chaetotaxy of mentum-submentum as in fig. 17, i araujoi Borgmeier 7. Second antennal segment longer than third (4:3); clypeus and anterior part of vertex finely reticulated flaviventris Mann Second antennal segment shorter than third ; head not finely reticulated ... 8 SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 73 8. Elytra with two setae in a submarginal row; second antennal segment one- half as long as scape i nornata Seevers Elytra with one seta on disk; second antennal segment less than one-half as long as scape 9 9. Median two-thirds of fifth sternite glabrous except for a very sparse stubble; mentum-submentum with pilosity as in fig. 17, c; fourth antennal segment one-half longer than third and broader emersoni Seevers Median area of fifth sternite with a vestiture of medium-sized pale, erect hairs; mentum-submentum with setae as in fig. 17, f; fourth antennal segment possibly a little longer and broader than third but not distinctly so. ecuadoriensis Seevers Termitophya heyeri Wasmann. Figure 17, a. Termitophya heyeri Wasmann, 1902, Tijd. Ent., 45: 97, pi. IX, fig. 1 (Sao Leopoldo, Rio Grande do Sul, Brazil ; Eutermes fulviceps Silvestri; Was- mann coll.). Material examined. Brazil: Sao Leopoldo (type); Blumenau, Santa Catarina (ex Reichensperger coll.). Host records. The identity of Eutermes fulviceps Silvestri is not known to termite specialists. The Blumenau specimens were accompanied by specimens of Nasutitermes jaraguae Holmgren and N. itapocuensis Holmgren. One supraorbital seta and three elytral setae present. Head coarsely punctate, the rather deep and clear-cut punctures moderately dense although irregularly distributed. Head shining, with scarcely perceptible sculpture except along occipital border. Antennae (16, 9, 11, 11, 9, 8, 8, 8, 8, 8, 16) with segments 3 and 4 longer than any of segments 2-10; segment 2 a little more than one-half as long as scape. Sternites 3-5 with a uniformly distributed sparse vestiture of fine, erect hairs, the tips of which are bent (obvious in profile); sternites 3-6 smooth, shining. Termitophya araujoi Borgmeier. Figure 17, i. Termilophya araujoi Borgmeier, 1950, Rev. Ent., 21 : 670, figs. 62-67 (Parada Deodoro, Santo Amaro, Sao Paulo, Brazil; host not cited; Institute Biol6gico, Sao Paulo). Host record. Nasutitermes proximus Silvestri (host of the holotype; determined by Emerson after publication of Borgmeier's paper). This species has not been available for study, but judging from Borgmeier's description it is very close to T. heyeri. A slight difference in the chaetotaxy of the mentum-submentum is indicated in figures 17, a and 17, i. Borgmeier reported that T. araujoi may be distinguished from T. heyeri by its shorter basal segment of labial palpi and by the longer antennae (longer than head and pronotum), but it is very doubtful if Wasmann's description was accurate enough to allow a comparison. 74 FIELDIANA: ZOOLOGY, VOLUME 40 Termitophya emersoni Seevers. Figures 15, a, 17, c. Termitophya emersoni Seevers, 1937, Ann. Ent. Soc. Amer., 30: 14, pi. Ill, fig. 12 (Barro Colorado Island, Panama Canal Zone; Nasutitermes ephratae Holmgren; Chicago Natural History Museum). Material examined. Panama: Barro Colorado Island (type series). British Guiana: Kartabo. Host records. Nasutitermes ephratae Holmgren (Panama, British Guiana). No supraorbital setae present; one elytral seta near outer apical angle. Head punctures medium-coarse and not very densely arranged ; surface shining, intervals smooth except for a few almost imperceptibly fine punctules. Head with sparse, pale, short hairs. Antennae (16, 6, 6, 9, 8, 7, 6, 6, 6, 6, 14) with segment 2 less than one-half as long as scape; segments 2 and 3 subequal; segment 4 the longest of segments 2-10. Sternite 3 with sparse, short, semi-erect hairs; sternites 4-6 glabrous. Mentum-submentum as in figure 17, c. Termitophya ecuadoriensis Seevers. Figure 17, /. Termitophya ecuadoriensis Seevers, 1937, Ann. Ent. Soc. Amer., 30: 16, pi. Ill, fig. 14 (Gualaquiza, Ecuador; Nasutitermes peruanus Holmgren; American Museum of Natural History). No supraorbital setae present; one elytral seta on disk. Head moderately densely punctate, the punctures of medium coarseness, shallow, and not clear-cut (possibly elongated); intervals semi-opaque, due to very fine, irregularly-spaced punctules; hairs short, pale, fine. Antennae (16, 6, 8, 8.5, 7, 7, 7, 7, 7, 7, 14) with segment 2 less than one-half as long as scape and a little shorter than any of segments 3-10; segments 3 and 4 subequal and longer than any of segments 5-10. Sternite 3 with sparse, fine, semi-erect hairs; sternite 4 with a wide median area almost glabrous; sternite 5 with sparse, semi-erect hairs, its surface almost without sculpture. Mentum-submentum as in figure 17, /. Termitophya inornata Seevers. Figure 17, g. Termitophya inornata Seevers, 1937, Ann. Ent. Soc. Amer., 30: 18 (Guala- quiza, Ecuador; Nasutitermes dendrophilus Holmgren; American Museum of Natural History). No supraorbital setae present; two elytral setae in submarginal row. Head with punctures of medium coarseness, shallow, their outlines obscure; intervals semi-opaque, surface irregularly sculptured with confluent punctules or obso- lescent reticulation (visible only with high magnification). Antennae (16, 8, 9, 9, 9, 8, 8, 7, 7, 7, 16) with segment 2 about one-half as long as scape; segments 3, 4, and 5 subequal and a little longer than segment 2 or any of segments 6-10. Ster- nites 3-6 smooth, very shining; sternite 3 with very sparse, short hairs; sternites 4 and 5 glabrous. Mentum-submentum as in figure 17, g. FIG. 15. (a) Termilophya emersoni Seevers. (b) Termitopithus crassiusculus Seevers. (c) Termitosyne plalygastra Seevers. (d) Termitosyne plalygastra Seevers; dorsal view of abdomen, (e) Termitomorpha fissipennis Casey, (f) Termito- morpha meinerli Wasmann. 75 76 FIELDIANA: ZOOLOGY, VOLUME 40 Termitophya flaviventris Mann. Figure 17, e. Termitophya flaviventris Mann, 1923, Zoologica, 3: 352 (Kartabo, British Guiana; Nasutitermes costalis Holmgren; United States National Mu- seum). Material examined. British Guiana: Kartabo (type series). Trinidad, British West Indies: Arena Forest and Northern Range (ex Adamson coll.). Host records. Nasutitermes costalis Holmgren (British Guiana, Trinidad). No supraorbital setae present; two elytral setae in submarginal row. Head with punctures of medium coarseness, not densely arranged. Clypeus and anterior part of vertex finely reticulated, but more posterior surface almost smooth. Head with sparse vestiture of fine short, pale hairs. Antennae (15, 8, 6, 7, 7, 6, 6, 6, 6, 6, 14) with segment 2 a little more than one-half as long as scape and longer than any of segments 3-10; segment 3 a little shorter than either 4 or 5. Sternites 3-6 smooth, very shining; sternite 3 with very sparse, pale hairs; sternites 4 and 5 glabrous. Mentum-submentum as in figure 17, e. Termitophya holmgreni Wasmann. Figure 17, d. Termitophya holmgreni Wasmann, in Wasmann and Holmgren, 1911, Zool. Anz., 38: 429 (Mojos, Bolivia; Eutermes mojosensis Holmgren; Wasmann coll.). Termitophya wasmanni Holmgren, in Wasmann and Holmgren, 1911, Zool. Anz., 38: 429 (Chaquimayo, Peru; Eutermes chaquimayensis Holmgren and Eutermes minimus Holmgren). New synonymy. Material examined. Three syntypes in Wasmann collection labelled "types" of Termitophya holmgreni Wasmann; two "bei Eutermes minimus Holmgren, Chaquimayo, Peru, Holmgren, 1909" and one "bei Eutermes chaquimayensis Holmgren, Chaquimayo, Peru." I have designated the latter specimen as lectotype. One specimen in Wasmann collection is labelled "Termitophya wasmanni Holmgren, type; bei Eutermes ripperti; Mojos, Bolivia," with an additional label "N. B., bei Eutermes mojosensis Holmgren (nicht ripperti; vgl. Holmgren)." Inasmuch as there are probably additional specimens in the Holmgren collection, the above specimen may be regarded as a syntype. It may be noted that the locality and host records cited by Wasmann and Holmgren do not agree with the data accompanying the specimens. Evidently the data for the two species were inter- changed in the manuscript before publication. It seems to me that the data in the collection must be assumed to be correct. In addition to the specimens in the Wasmann collection, I have studied specimens from Rurrenabaque, Beni, Bolivia, collected with FIG. 16. (a) Termitogaster insolens Casey. (b) Termitogasler nigricollis Silvestri. (c) Xen- opelta cornuta Mann, (d) Neotermitogaster colonus Seevers. (e) Xenogaster inquilina Borgmeier. 77 78 FIELDIANA: ZOOLOGY, VOLUME 40 NasiUitermes chaquimayensis Holmgren (ex W. M. Mann coll., USNM). Remarks. Careful examination of specimens of T. holmgreni and T. wasmanni in the Wasmann collection did not reveal the diagnostic features given by the two authors. The species were distinguished from one another by different antennal lengths, proportions of antennal segments, and pronotal and elytral pro- portions. Wasmann was very inaccurate in judging measurements and proportions and in this instance he erred in all of his estimates. The differences do not exist. Inasmuch as this is true, the confusion of locality and host data does not assume much importance. No supraorbital setae present; elytra with two setae. Head with medium- coarse punctures densely arranged; the punctures not clearly distinct from each other, apparently because of an obsolescent reticulation of broken lines; surface dull. Antennae (15: 6: 8: 8: 8: 7: 7: 7: 7: 6: 14) with segment 2 less than one-half as long as scape, and not as broad as the following segments; segments 3, 4, and 5 subequal, only a little longer than any of segments 6-9. Sternites 3-5 with a rather conspicuous vestiture of moderately dense, pale, recumbent hairs; the median and lateral areas similarly clothed. Mentum-submentum as in figure 17, d. Pronotum about one-sixth broader than long. Elytra shorter than pro- notum. Termitophya piliventris Seevers. Figure 17, b. Termitophya pilicentris Seevers, 1937, Ann. Ent. Soc. Amer., 30: 17, pi. Ill, fig. 13 (Barro Colorado Island, Panama Canal Zone; Nasutitermes colum- bicus Holmgren; Chicago Natural History Museum). One supraorbital seta present; two elytral setae in marginal row. Head punctures coarse and moderately dense, concentrated chiefly in a large circular area of vertex and becoming finer and less dense laterally and posteriorly. Head with a close-meshed coarse reticulation which does not obscure the punctation. Antennae (18, 7, 9, 10, 9, 9, 9, 8, 8, 8, 15) with segment 2 less than one-half as long as scape and shorter than any of the following segments; segment 4 the longest of segments 2-10; segments 3, 5, 6, and 7 subequal. Sternites 3-5 with a sparse vestiture of erect, pale hairs (conspicuous in profile), the median and lateral areas similarly clothed; Sternites 3-6 finely reticulated. Mentum-submentum as in figure 17, 6. Termitophya arnica Mann. Figure 17, h. Termitophya arnica Mann, 1923, Zoologies, 3: 351 (Kartabo, British Guiana; Nasutitermes guayanae Holmgren; United States National Museum); Emerson, 1935, Ann. Ent. Soc. Amer., 28: 369, fig. 4; Seevers, 1937, Ann. Ent. Soc. Amer., 30: 19. Material examined. British Guiana: Kartabo (type series). Trinidad, British West Indies: Valencia Forest (ex Adamson coll.). /I I I I i \ FIG. 17. Submentum-mentum and gula. (a) Termitophya heyeri Wasmann. (b) Termitophya piliventris Seevers. (c) Termitophya emersoni Seevers. (d) Termitophya holmgreni Wasmann. (e) Termitophya flavivenlris Mann, (f) Termi- tophya ecuadoriensis Seevers. (g) Termitophya inornata Seevers. (h) Termitophya arnica Mann, (i) Termitophya araujoi Borgmeier (after Borgmeier, 1950). (j) Termitosynodes williamsi Seevers. (k) Xenogaster inflata Wasmann. (1) Xeno- gaster pilosula Seevers. (m) Xenogaster reichenspergeri Seevers. (n) Termitopithus crassiusculus Seevers. (o) Termitosyne platygastra Seevers. (p) Xenogaster glabriventris Seevers. (q) Xenogaster nana Seevers. (r) Xenogaster subnuda Seevers. (s) Xenogaster inquilina Borgmeier. (t) Termitoides marginatus Seevers. 79 80 FIELDIANA: ZOOLOGY, VOLUME 40 One supraorbital seta present; two elytral setae in submarginal row. Head punctures fine, sparsely and irregularly arranged; punctation almost obscured by a close-meshed coarse reticulation. Head dull ; its fine, pale hairs short and sparse. Antennae (19, 7, 11, 11, 10, 9, 8, 8, 8, 8, 17) with segment 2 less than one-half as long as scape, and shorter than any of the following segments; segments 3 and 4 subequal in length, longer than any of segments 2-10. Sternite 3 with very sparse, semi-erect, short hairs; 4 and 5 glabrous; 3-6 with reticulated surface. Mentum-submentum distinctive in form arid pilosity (fig. 17, h). Termitophya punctata Mann Termitophya punctata Mann, 1923, Zoologica, 3: 349, text fig. 36 (Kartabo, British Guiana; Nasutitermes guayanae Holmgren; United States National Museum); Emerson, 1935, Ann. Ent. Soc. Amer., 28: 369, text fig. 3. Material examined. British Guiana: Kartabo (one paratype). Host record. Nasutitermes similis Emerson (Emerson, 1935). Two supraorbital setae and two pairs of elytral setae present. TERMITOPITHUS, new genus Type species: Termitopithus crassiusculus, new species. Termitopithus seems to be closely allied to Termitophya, from which it differs in pronotal structure, elytra, and distinctive abdom- inal form. The head is in most respects like that of Termitophya, but the antennae are more geniculate. The elongated, saddle- shaped pronotum and the ovate, very membranous abdomen are very distinctive. Head similar to that of Termitophya; vertex flat or feebly concave; clypeus moderately declivous from approximately the level of the posterior margins of antennal fossae. Antennae strongly geniculate, scape about equal to the three following segments combined; scape somewhat thickened in its basal two-thirds when viewed from side, anterior face of scape produced basally to an acute ridge and deeply concave in its apical one-third. Antennal segment 2 subequal to segments 3 and 4 combined, its anterior face concave; segments 3-10 short, cylindrical. Mentum-submentum (fig. 17, n) not especially distinctive, not separated from gula by a suture. Maxillary palpi unusually small for the subtribe; segment 2 cornucopia-like, its basal one-third attenuated; segment 3 oval, about three- fourths as wide as long; segment 4 awl-shaped. Pronotum distinctively saddle-shaped, one-tenth longer than maximum width; entire disk shallowly concave, without pits or elevations; apical margin strongly arcuate, anterior angles almost obsolete. Pronotum widest midway between apex and base; sides arcuately convergent in front and posteriorly sinuate to the base; basal angles almost rectangular, base about five-sixths the maximum width. Inner margins of elytra contiguous; sutural length only about one-half as long as outer margins; apical margin of each elytron strongly oblique. Elytra and pronotum about equal in length. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 81 Abdomen strongly physogastric, ovate in form and permanently recurved over foreparts. Tergites 3-6 moderately large, separated by membranous integu- ment, tergites 6 and 7 contiguous, tergite 7 much larger, about four-sevenths as long as broad. Sternites widely separated by membranous cuticle; sternite 3 very large, shield-like, separated from metasternum by considerable non-sclerotized cuticle; sternites 4-6 very broad, strap-like. Termitopithus crassiusculus, new species. Figures 15, b, 17, n. Holotype from Blumenau, Santa Catarina, Brazil (ex Reichens- perger collection). In collection of Chicago Natural History Museum. Paratypes. Six specimens, same data as holotype. One speci- men, Ilha Grande, Rio de Janeiro, Brazil, collected October 9, 1944, by Helmut Sick. Paratypes in Chicago Natural History Museum, Reichensperger collection, United States National Museum, and Wasmann collection. Host. Nasutitermes aquilinus Holmgren (both localities). Head, pronotum, and elytra light reddish-brown, tergites light-brown, sternites pale testaceous. Head smooth, not reticulated, but with an almost imperceptibly fine punctulation. Pronotal disk smooth, shining, impunctate, and with only traces of reticulation medially; sides coarsely reticulated, sparsely punctate, the punctures obscured by reticulation. Elytral disk smooth, shining, not reticulated, but with a sparse fine punctation in basal half; sides coarsely reticulated and sparsely punctate on basal two-thirds. Tergites with a reticulation of moderately coarse raised lines, most conspicuous on tergites 7 and 8. Sternites with a broad medial smooth area without reticulation, and bearing only scattered setigerous punctures; the sternites are dull in appearance on the sides of the abdomen as the setigerous punctures become coarse and densely arranged and as the surface becomes coarsely reticulated (the general appearance is that of coarse granulation). Head with scattered, very inconspicuous, fine short hairs. Pronotal disk with extremely few, very fine hairs; sides with a moderate number of fine pale hairs, especially along lateral margins near humeri. Elytra with sparse, fine hairs near base and on sides. Tergites 3-5 with few hairs except in the apical row; 6-8 with moderately dense vestiture of fine pale hairs; sternites with vestiture of erect pale hairs, among which are a number of longer, darker setae. Sternites 4-7 sparsely clothed with erect pale hairs medially, and with a few longer darker hairs in basal and apical rows. Sternites with a much denser pubescence laterally, sternite 8 rather hairy. Postgenae (lateral to gula), prosternum, and metasternum with a number of moderately long pale hairs. Antennae (42, 18, 9, 9, 9, 8, 8, 8, 8, 8, 18) with the scape and segment 2 modi- fied as for the genus; segment 2 twice as long as any of segments 3-10; segments 3-5 subequal in length. Length of head and thorax, 2 mm.; abdomen, 2.6 mm. TERMITOSYNE, new genus Type species: Termitosyne platygastra, new species. 82 FIELD IANA: ZOOLOGY, VOLUME 40 Head subequal in length and width or slightly longer than broad; clypeus feebly declivous, its border slightly emarginate; head feebly elevated between antennal fossae; vertex with a shallow, V-shaped impression. Antennae geniculate; scape relatively long, about equal to segments 2-5 combined. Mentum-sub- mentum distinctive in form (fig. 17, o). Pronotum transverse; disk broadly impressed but more superficially than in Xenogasler. Elytra contiguous. Abdomen distinctive; very broad, dorsum almost flat; sclerites of each segment contiguous; relatively little non-sclerotized cuticle exposed. Inner paratergites very large, two-fifths or more as broad as tergites, in the same plane as tergites; outer paratergites small. Convex ventral surface of abdomen occupied by the large sternites. Termitosyne platygastra, new species. Figures 15, c, d, 17, o. Holotype from Blumenau, Santa Catarina, Brazil (ex Reichens- perger collection). In collection of Chicago Natural History Museum. Paratypes. Nineteen specimens, same data as holotype. One specimen, Ilha Grande, Rio de Janeiro, Brazil, collected October 9, 1944, by Helmut Sick. Four specimens, Passa Quatro, Minas Gerais, Brazil (ex Reichensperger collection). Paratypes in collec- tions of Chicago Natural History Museum, Reichensperger, Borg- meier, and United States National Museum. Head coarsely reticulated and with a moderately dense covering of short pale hairs. Pronotum reticulated, moderately densely clothed with short pale hairs; the impressed area with feeble asperities. Elytra with a fairly dense vestiture of pale semi-recumbent hairs. Tergites glabrous except for a very few pale hairs and with almost no traces of reticulation; inner paratergites reticulated; sternites smooth and shining, with a sparse vestiture of pale hairs, more dense on terminal segments. Antennae (46, 6, 10, 10, 10, 10, 10, 9, 8, 8, 20) with segment 2 unusually small, only one-seventh as long as scape; segments 3-8 not appreciably different. Segment 2 of maxillary palpi cornucopia-like, its inner surface concave; distinctive segment 3 slightly wider than long, its surface spongy except for a conspicuous flattened area dorsally. Mentum-submentum distinctive (fig. 17, o). Pronotum a little less than one-fifth broader than long; disk broadly and shallowly impressed, deepest near middle. Elytra contiguous for a distance and diverging feebly behind middle. Proportions of tergites and inner paratergites of abdomen as follows: Fourth paratergite (42 X 38) almost one-half as broad as tergite (90 X 36) ; fifth para- tergite (42 X 40) almost one-half as broad as tergite (90 X 36) ; sixth paratergite (42 X 36) about one-half as broad as tergite (90 X 40) ; seventh paratergite (36 X 30) about four-fifths as broad as tergite (86 X 30). Seventh tergite trapezoidal, almost three times as wide as long, its surface slightly convex medially and slightly reflexed laterally; seventh paratergite triangular, its inner margin oblique, its outer margin arcuate. Length, 4 mm. (with abdomen extended). SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 83 TERMITOSYNODES, new genus Type species: Termitosynodes williamsi, new species. Similar to Termitosyne, especially in the form of the abdomen, but differing in a number of respects: clypeus strongly deflexed; maxillary palpi distinctive; men tum-submen turn distinctive; anten- nal proportions different; pronotum broader and distinctively impressed; seventh tergite distinctively elongated (two- thirds as long as broad; only one-third as long as broad in Termitosyne); seventh inner paratergite distinctive. Head slightly broader than long; clypeus almost vertically deflexed to the apical margin, and elevated to form a transverse ridge between antennal fossae; vertex broadly concave. Maxillary palpi not modified as in Termitosyne. Men- tum-submentum distinctive (fig. 17, j). Pronotum transverse; disk broadly impressed except for a central eminence. Elytra contiguous, except diverging slightly behind middle. Abdomen similar in form to that of Termitosyne; very broad, with little non- sclerotized cuticle and with very large inner paratergites. Seventh tergite ex- tremely long, about two-thirds as broad as long; seventh paratergite distinctive in form. Termitosynodes williamsi, new species. Figure 17, ;'. Holotype from Guayaquil, Ecuador, collected by F. X. Williams. In the collection of United States National Museum. Paratype. One specimen, same data as type; in collection of Chicago Natural History Museum. Host. Nasutitermes nigriceps Haldeman. Head strongly shining, weakly reticulated, obsoletely so near base; sparsely setose, with a few clypeal setae and one occipital pair. Pronotum shining, feebly and irregularly reticulated; setae on anterior and lateral margins and a few on disk. Tergites and paratergites smooth, shining, with a few traces of reticulation and only feebly punctulate; with a row of long, pale, semi-erect hairs. Sternites sparsely and irregularly punctulate and reticulate; with a sparse vestiture of pale, erect hairs, conspicuous only when viewed in profile. Antennae (42, 10, 11.5, 11.5, 10, 9, 9, 8, 8, 8, 18) with segment 2 about one- fourth as long as scape; segments 3 and 4 equal in length; segments 5-10 not appreciably different. Mentum-submentum distinctive (fig. 17, j). Pronotum about three-tenths broader than long; entire disk impressed except for a central eminence. Relative widths of tergites and inner paratergites of abdomen as follows: fourth paratergite (30) almost one-half as wide as tergite (65); fifth paratergite (32) less than one-half as wide as tergite (70); sixth paratergite (32) more than two-fifths as wide as tergite (73); seventh paratergite (26) two- fifths as wide as tergite (68). Seventh tergite and inner paratergites very long, the latter with a distinctive form, its outer margin broadly arcuate in front, becoming deeply emarginate and sinuate behind the middle. Outer paratergites 84 FIELDIANA: ZOOLOGY, VOLUME 40 extremely thin, often located with difficulty. Sternites convex; seventh broadly and deeply emarginate medially. Length, 2.5 mm. TERMITOMORPHA Wasmann Termitomorpha Wasmann, 1894, Krit. Verz., p. 210; Fenyes, 1920, Gen. Insect., 173B: 346. Type species: Termitomorpha meinerti Wasmann. Thaxteria Fenyes, 1921, Bull. Mus. Comp. Zool., 65: 17. Type species: Thaxteria insularis Fenyes. New synonymy. Termitosomus Seevers, 1939, Amer. Mus. Nov., no. 1018, p. 3; 1946, Rev. Ent., 17: 253; Borgmeier, 1950, Rev. Ent., 21: 638. Type species: Termi- tospmus fissipennis Casey. Clypeus moderately declivous or abruptly and strongly deflexed some distance in front of antennal fossae. Antennae strongly geniculate, rather slender. Gula (fig. 18, TO) very slender, widest at base. Mentum-submentum variable (fig. 18, I, m); in general longer than broad and in most cases with parallel sides; surface often broadly impressed near base. Surface of maxillae broadly impressed basally, maxillary palpi large, segment 2 approximately triangular, its upper margin arcuate; segment 3 oval in outline, segment 4 subulate. Pronotum transverse, evenly convex, not impressed, its sides nearly parallel. Elytra not contiguous, their inner margins diverging strongly behind the scutellum; inner one-half to one-third of elytra coriaceous to hyaline; elytral apices narrow. Wings vestigial. Prosternum very short, usually with a median elevation. Abdomen only moderately physogastric, and with limited areas of mem- branous integument; rather termite-like in appearance in some species. Tergites strongly convex, paratergites contiguous with each other and with sternites and tergites; abdomen usually medio-laterally compressed. Remarks. -Termitosomus Seevers was, at my suggestion, placed in synonymy by Borgmeier (1950). Thaxteria Fenyes is also a syno- nym of this genus. Fenyes incorrectly determined the tarsal formula of that genus to be 4, 4, 4 and placed Thaxteria near Coro- toca. Examination of the type of Thaxteria insularis Fenyes con- firmed my suspicion that it is a synonym of Termitomorpha meinerti Wasmann. KEY TO SPECIES OF TERMITOMORPHA 1. Outer margin of hind femora obtusely dentate due to an arcuate emargination of the proximal two-thirds; clypeus with two cariniform tubercles medially. fissipennis Casey Hind femora not dentate; clypeus without tubercles 2 2. Mesothoracic trochanter with a dentiform tubercle 3 Mesothoracic trochanter without a dentiform tubercle 4 3. Second antennal segment compressed and as long as segments 3 and 4 com- bined huachii Seevers Second antennal segment not compressed and about three-fourths as long as segments 3 and 4 combined meinerti Wasmann SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 85 4. Clypeus flat for a short distance in front of antennal fossae to an arcuate "margin" and then sharply deflexed to its anterior border; lateral and apical margins of mentum-submentum almost straight costaricensis Seevers Clypeus declivous to its anterior border, but not abruptly deflexed; apical margin of mentum-submentum strongly arcuate, the apical angles obsolete. manni Seevers Termitomorpha meinerti Wasmann. Figures 15, /, 18, /. Termitomorpha meinerti Wasmann, 1894, Krit. Verz., p. 211 (Las Trincheras, Venezuela; Eutermes meinerti Wasmann; University of Copenhagen Museum). Thaxteria insularis Fenyes, 1921, Bull. Mus. Comp. Zool., 65: 18 (Grand Etang, Grenada; Museum of Comparative Zoology). New synonymy. Termitogaster simulans Mann, 1923, Zoologica, 3: 338, text fig. 32 (Kartabo, British Guiana; Nasutitermes costalis Holmgren). New synonymy. Termitosomus simulans Mann, Seevers, 1939, Amer. Mus. Nov., no. 1018, p. 3, text figs. 7, 21, 24; 1946, Rev. Ent., 17: 253. Material examined. British Guiana: Kartabo (type series of T. simulans Mann). Grenada: Grand Etang (type of T. insularis Fenyes). Trinidad, British West Indies: Northern Range (ex Adamson coll.). Host records. Nasutitermes meinerti Wasmann (Venezuela); Nasutitermes costalis Holmgren (British Guiana, Trinidad, Grenada). Color reddish-brown. Clypeus elevated to form a transverse ridge and then rather steeply declivous to its feebly emarginate border. Antennal scape longer than segments 2, 3, 4 combined; segment 2 about three-fourths longer than 3; segment 3 longer than 4; segments 4-10 gradually decreasing in length. Mentum- submentum distinctive (fig. 18, I). Pronotum with sides converging slightly toward base; its apex broadly arcuate; base arcuate but with a medial shallow emargination and lateral sinuations. Prosternum broadly elevated at middle but not tuberculate or carinate. Hind femora not modified. Mesothoracic trochanter with a dentiform tubercle. Remarks. Although I have not examined the type of T. meinerti there is little doubt that T. simulans is a synonym of it. This, however, should be checked as soon as possible. Fenyes' insularis is a synonym of Mann's simulans and presumably of meinerti also. Termitomorpha huachii Seevers Termitomorpha huachii Seevers, 1946, Rev. Ent., 17: 254 (Huachi, Beni, Bolivia; Nasutitermes sp. indet., worker caste only; United States National Museum). Closely related to T. meinerti from which it differs in antennal structure: Antennae with a very long scape, equal to segments 2-5 combined; segment 2 compressed and elongated, equal to segments 3 and 4 combined. 86 FIELDIANA: ZOOLOGY, VOLUME 40 Termitomorpha fissipennis Casey. Figures 15, e, 18, m. Termitogaster fissipennis Casey, 1890, Ann. N. Y. Acad. Sci., 5: 187 (Panama; no host cited; United States National Museum). Termitogaster simopelta Mann, 1923, Zoologica, 3: 341, text fig. 33 (Kartabo, British Guiana; Nasutitermes costalis Holmgren; United States National Museum). Termitosomus fissipennis Casey, Seevers, 1939, Amer. Mus. Nov., no. 1018, p. 3, text figs. 8, 20, 23; Seevers, 1946, Rev. Ent., 17: 253. Material examined. Panama: (Casey's type). In addition, Barro Colorado Island, Canal Zone. British Guiana: Kartabo (types of T. simopelta Mann). Trinidad, British West Indies: Northern Range (ex Adamson coll.). Costa Rica: Hamburg Farm (ex Nevermann coll., USNM). Host records. Nasutitermes corniger Motschulsky (Panama, Costa Rica). N. costalis Holmgren (British Guiana, Trinidad). Color flavo- testaceous; each elytron with a broad, reddish, longitudinal stripe. Clypeus almost flat for a short distance and then strongly deflexed to its anterior border, which has a moderately deep median notch flanked by fine, acute processes. Clypeus at point of deflexion with two broad cariniform tubercles. Antennal scape equal to segments 2, 3, and 4 combined; segments 2 and 3 subequal in length; segment 4 a little more than one-half as long as 3; 4-10 very similar, decreasing very slightly in length. Mentum-submentum distinctive (fig. 18, TO). Pronotal apex arcuate, base nearly straight, and sides almost parallel. Prosternum with an acute tubercle at middle. Outer margin of hind femora obtusely dentate due to an arcuate emargination of the proximal two-thirds. Mesothoracic tro- chanter with small dentiform tubercle. Termitomorpha costaricensis Seevers Termitosomus costaricensis Seevers, 1946, Rev. Ent., 17: 255 (Hamburg Farm, Santa Clara Prov., Costa Rica; Nasutitermes sp. ; United States National Museum). Material examined. Costa Rica: Hamburg Farm (type) and Ibera, Santa Cruz, with Nasutitermes corniger Motschulsky. Color light reddish-brown. Clypeus almost flat for a distance in front of antennal fossae and then strongly deflexed to its anterior border, the deflexion producing an arcuate "margin" which is evenly rounded at middle but obliquely carinate laterally; with a convex longitudinal elevation between antennal fossae. Mentum-submentum distinctive, more strongly setose than in meinerti; its basal one-half transversely concave, and the apical half strongly reflexed dorsad. Pronotal base feebly arcuate but with a relatively deep medial notch due to an impression of adjacent surface and moderately strong lateral sinuations. Hind femora and middle trochanters not modified. FIG. 18. Submentum-mentum and gula. (a) Termitogaster insolens Casey, (b) Termitogaster puncticeps Seevers. (c) Termitogaster impressicollis Seevers. (d) Termitogaster beniensis Seevers. (e) Termitogaster nigricollis Silvestri. (f) Termitogaster wenzeli Seevers. (g) Termitogasier emersoni Mann, (h) Termito- gaster chavantinae Seevers. (i) Termitogasier bicolor Seevers. (j) Termitogaster dicersicollis Seevers. (k) Neotermitogaster colonus Seevers. (1) Termitomorpha meinerii Wasmann. (m) Termitomorpha fissipennis Casey, (n) Trachopeplus disjunctipennis Seevers. (o) Xenopelta cornuta Mann, (p) Termitella foveolata Reichensperger. (q) Termitella lujae Wasmann, under side of head illustrating the deep postgenal sulci. (r) Idioptochus lehmensicki Seevers. 87 88 FIELDIANA: ZOOLOGY, VOLUME 40 Termitomorpha rnanni Seevers Termitosomus manni Seevers, 1946, Rev. Ent., 17: 253 (Rurrenabaque, Beni, Bolivia; Nasuiitermes peruanus Holmgren and N. pilosus Snyder; United States National Museum). Color light reddish-brown. Clypeus with a low, transversely arcuate, smooth elevation, and then strongly declivous to the feebly arcuate anterior border. Antennae with a very long scape, about as long as segments 2-5 combined; scape somewhat compressed, especially near base; segment 2 about one-fourth longer than 3; segment 3 about one-third longer than 4; segment 4 a little longer than 5; 6-10 subequal. Pronotum with strongly arcuate apical margin, straight sides, and broadly arcuate base, which is slightly notched at middle and sinuate laterally. Prosternum broadly and convexly elevated at middle. Hind femora not modified; middle trochanters without dentiform tubercle. TRACHOPEPLUS Mann Trachopeplus Mann, 1923, Zoologica, 3: 353; Seevers, 1939, Amer. Mus. Nov., no. 1018, p. 6. Type species: Trachopeplus setosus Mann. Head wider than long; feebly elevated between antennal fossae; clypeus feebly declivous, its apical margin strongly arcuate; vertex with shallow, narrowly confluent depressions behind antennal fossae. Gula moderately broad, not separated from mentum-submentum by a suture. Mentum-submentum distinc- tive (fig. 18, n). Maxillary palpi with an unusually long cornucopia-shaped segment 2, very narrow at base and broad at apex; segment 3 large, broad at base, its sides convergent to the slender apex; segment 4 subulate. Antennae geniculate, scape somewhat longer than the three following segments combined. Pronotum transverse; its basal and apical margins bisinuate, lateral margins almost straight, and disk with a deep transverse impression in front of the middle and usually with other smaller depressions and elevations. Elytra large; strongly divergent except for a short distance at base; the medial margins deflexed. Wings vestigial. Abdomen similar to that of Termitogasler; dorsum slightly convex, venter strongly so; sclerites of a given segment almost contiguous, but with some non- sclerotized cuticle exposed between segments; inner paratergites moderate in size, not elevated; outer paratergites smaller. Trachopeplus setosus Mann Trachopeplus setosus Mann, 1923, Zoologica, 3: 354, text fig. 37 (Kartabo, British Guiana; Nasutitermes acajutlae Holmgren; United States National Museum). Host record. Nasutitermes nigriceps Haldeman (= acajutlae Holmgren). This species has not been available for study and the original description did not characterize the species very thoroughly. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 89 Trachopeplus disjunct ipennis Seevers. Figure 18, n. Trachopeplus disjunctipennis Seevers, 1939, Amer. Mus. Nov., no. 1018, p. 6, text figs. 13, 17, 25, 28, 32, 34 (El Coco, near Bolivar, Ecuador; Nasu- titermes pilifrons Holmgren; American Museum of Natural History). Host record. Nasutitermes nigriceps Haldeman (= pilifrons Holmgren). This species is closely allied to T. setosus Mann but seems to differ in details of chaetotaxy and in the form and contours of the pronotum. Color dark brown; abdomen reddish-brown, darker apically. Head, pronotum and elytra densely, evenly, very minutely punctulate and microscopically pu- bescent. Clypeus with about 13 setae, vertex with two pairs of medial setae; occiput and postgenae with a row of about 9 setae extending forward below the eye. Pronotum with 4 setae on apical margin, 8 on each lateral margin, about 8 in a semi-circular subapical row, about 4 in a submarginal row on each side, and about 12 in the area behind the U-shaped impression. Elytra with 3 marginal setae and 6 discal setae. Third tergite with an apical row of 8 black setae and a few pale ones; tergites 4-7 with an apical row of 9 and a subapical row of 8 stout setae; eighth tergite with 6 apical setae and 8 in a row about the middle; ninth tergite with an apical row of 6 and a middle row of 4; paratergites with 3 or 4 stout setae. Sternites with three or four rows of stout black setae. Pronotum about three-fourths as long as wide; anterior and basal borders bisinuate; the latter more strongly arcuate medially; anterior angles truncate; sides nearly parallel, slightly produced at the insertion of each bristle so as to appear feebly scalloped. Pronotal disk in front of middle with a broad, deep, transverse impression which bends and extends to basal border; this impression encloses a broad, low eminence; surface adjacent to apical margin shallowly impressed on each side. Length, 3-3.5 mm. TERM ITOG ASTER Casey Termitogaster Casey, 1889, Ann. N. Y. Acad. Sci., 4: 63; Fenyes, 1920, Gen. Insect., 173B: 345; Mann, 1923, Zoologica, 3: 338; Seevers, 1937, Ann. Ent. Soc. Amer., 30: 11; 1939, Amer. Mus. Nov., no. 1018, p. 1; 1946, Rev. Ent., 17: 248. Type species: Termitogaster insolens Casey. Dorsum of head having only minor elevations and depressions; clypeus slightly declivous to shallowly concave but never abruptly deflexed, its margin straight to rather deeply emarginate. Antennal scape about as long as segments 2, 3, and 4 combined; antennal segments variable, especially 2-4. Maxilla with cardo prolonged laterally or not; palpi variable. Mentum-submentum of distinc- tive form as a rule (fig. 18, a-j); separated from gula by a suture. Pronotum, as a rule, robust, strongly convex, black in color, and with only minor impressions and elevations; but in a few cases lighter in color, not as convex, and with the disk broadly impressed. Pronotum averaging about one- tenth broader than long, but subequal in length and width in at least one case. Elytra with inner margins contiguous throughout. 90 FIELDIANA: ZOOLOGY, VOLUME 40 Abdomen strongly physogastric, probably held in a vertical position. Sides of abdomen with moderately extensive areas of white non-sclerotized integu- ment. Tergites subequal in length, strongly transverse; inner paratergites triangular in shape, the fifth strongly attenuated. Outer paratergites very slender, oblique. Remarks. The following structures seem to exhibit little varia- tion within the genus: abdominal structure and chaetotaxy, pronotal chaetotaxy, elytra, sternum, legs. The antennae are variable in the proportions of their segments, and the following table may be useful for comparative purposes. These are relative lengths and are subject to minor intraspecific variation. Scape 2 3 4 56789 10 11 insolens 34 10 12 8 8 7 7 7 7 7 18 testaceus 34 12 11 8 8 7 7 7 7 7 18 puncticeps 42 12 13 10 9 9 8 8 8 8 18 brevis 32 9 9 7 6 7 7 6 6 6 17 emarginatus 40 12 12 9 9 8 7 7 6 6 18 impressicollis 40 14 14 12 9 8 8 8 8 8 18 bicolor 34 11 12 8 8 7 7 6 6 6 17 emersoni 38 10 11 9 8 7 7 7 7 7 18 chavantinae 36 10 12 9 8 8 8 8 7 7 20 wenzeli 36 13 9 9877666 16 nigricollis 40 14 11 10 8 8 7 7 7 6 16 beniensis 36 12 12 10 9 8 8 8 7 7 18 diversicollis 34 10 12 9 8 8 8 8 8 7 18 magdalenae 32 10 9 8 8 8 7 7 6 6 18 The dimensions of the pronotum are variable: the pronotum is one-tenth to one-fifth wider than long in most cases impressicollis, emarginatus, emersoni, chavantinae, nigricollis, beniensis, bicolor, insolens, testaceus, magdalenae; less than one-tenth wider than long in a few species puncticeps, wenzeli, and brevis; the width and length nearly equal diversicollis. KEY TO SPECIES OF TERMITOGASTER 1. Cardo of maxilla acuminately produced laterally 2 Cardo of maxilla not produced 4 2. Dorsum of head densely and coarsely punctate puncticeps Seevers Dorsum of head finely punctulate 3 3. Pronotum piceous, strongly convex, its disk with only feeble impressions; antennal segment 3 longer than 2 insolens Casey Pronotum testaceous, its disk strongly impressed; antennal segment 2 slightly longer than 3 testaceus Seevers 4. Legs relatively short; neither metathoracic femora nor tibiae longer than pronotum; surface of head between antennae elevated; clypeus concave, its border moderately emarginate brevis Mann SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 91 Legs longer; both metathoracic femora and tibiae at least one-third longer than pronotum; surface of head otherwise 5 5. Clypeus very deeply emarginate emarginalus Seevers Clypeus at most shallowly emarginate 6 6. Vertex coarsely and uniformly punctate 7 Vertex finely and uniformly punctulate, or if the surface behind the eyes is moderately coarsely punctate, the vertex is almost impunctate medially ... 8 7. Pronotal disk more or less flattened, and with irregular impressions; disk brown, outer one-fourth of pronotum black; mentum-submentum distinctive (fig. 18, i); head with one occipital seta on each side bicolor Seevers Disk of pronotum not flattened appreciably but with several moderately deep impressions; pronotum not conspicuously bicolored, although areas of disk may be rufo-piceus; mentum-submentum distinctive (fig. 18, c); head with two or three occipital setae on each half impressicollis Seevers 8. Pronotum distinctly transverse, usually about one-tenth broader than long; pronotum convex and with only minor impressions 9 Pronotum nearly equal in length and width; pronotum brown, its entire disk strongly impressed, and with a sub-basal eminence within the impression. diversicollis, new sp. 9. Antennal segment 2 longer than 3 10 Antennal segment 2 subequal to segment 3 or shorter 11 10. Chaetotaxy of mentum-submentum as in fig. 18, e; dorsum of head moder- ately coarsely and densely punctate except for medial areas of vertex and occiput nigricollis Silvestri Chaetotaxy of mentum-submentum as in fig. 18, /; dorsum of head very finely punctulate wenzeli, new sp. 11. Vertex with a pair of setae between antennal fossae in a line with their posterior margins emersoni Mann Vertex without the above-mentioned vertexal setae 12 12. Antennal segment 3 longer than 2; mentum-submentum distinctive (fig. 18, /t) chavantinae, new sp. Antennal segment 3 equal to 2; mentum-submentum not as above 13 13. Mentum-submentum distinctive; one occipital seta on each side. magdalenae Seevers Mentum-submentum distinctive (fig. 18, d); two or three occipital setae on each side beniensis Seevers (Termitogaster wasmanni Holmgren is not included in the above key) Termitogaster insolens Casey. Figures 16, a, 18, a. Termitogaster insolens Casey, 1889, Ann. N. Y. Acad. Sci., 4: 65 (Panama; with termites; United States National Museum); Seevers, 1937, Ann. Soc. Amer., 30: 14, pi. 2, fig. 5; 1939, Amer. Mus. Nov., no. 1018, text figs. 1, 10, 14, 22, 26, 27, 33; 1946, Rev. Ent., 17: 248. Material examined. Barro Colorado Island, Panama Canal Zone (CNHM); Estrella, Costa Rica (USNM); Hamburg Farm, Costa Rica (USNM); Zent, Costa Rica (USNM); Progress and Malon, Honduras (USNM) ; all with Nasutitermes corniger Motschulsky. Color reddish-brown, pronotum black. Vertex not elevated between antennae; clypeus slightly declivous, its margin feebly emarginate. Distance 92 FIELDIANA: ZOOLOGY, VOLUME 40 between antennal fossae only two-fifths the width of head at eye level. Three to five occipital setae on each side; vertexal setae absent. Head finely punctulate. Antennae with segment 3 longer than 2 and 4-10; segments 4 and 5 subequal. Cardo of maxilla acuminately produced laterally, visible from above. Segment 2 of maxillary palpi with surface of its broader basal half flattened; segment 3 oval, about two-thirds as broad as long. Mentum-submentum distinctive (fig. 18, a). Pro no turn about one-seventh broader than long; robust, convex; its surface contours variable, with several shallow impressions and deeper pits. Length, 3 mm. Termitogaster testaceus Seevers Termitogaster testaceus Seevers, 1946, Rev. Ent., 17: 250 (Zent, Costa Rica; Nasutitermes corniger Motschulsky; United States National Museum). Material examined. The holotype. Closely allied to insolens but differing in these respects: Coloration testaceous, sides of pronotum reddish-brown; antennae with segment 2 slightly longer than 3; pronotal disk strongly and rather uniformly impressed, basal one-half with a low, triangular elevation. Termitogaster puncticeps Seevers. Figure 18, b. Termitogaster puncticeps Seevers, 1946, Rev. Ent., 17: 249 (Rurrenabaque, Beni, Bolivia; Nasulitermes peruanus Holmgren; United States National Museum). Material examined. The type series. Color light-brown, pronotal sides piceous, disk mottled-brown and reddish- brown. Vertex elevated somewhat between antennae; clypeus moderately declivous, its margin feebly emarginate. Three to five occipital setae on each side; vertexal setae absent. Head coarsely punctate except for a median V-shaped impunctate area. Antennae with segment 2 slightly shorter than 3; segment 3 the longest of segments 2-10; segment 4 slightly longer than 5. Cardo acuminately produced. Segment 2 of maxillary palpi bent at middle to form a right angle, proximal half with its broad flat surface in a horizontal plane, distal half vertical. Mentum-submentum distinctive (fig. 18, b). Pronotum only about one-twentieth broader than long; its disk delimited from the smooth sloping sides by rather deep, although irregular, impressions that converge midway between basal and apical borders, and again near base; disk with a prominent median eminence before base; disk may also have a shallow median impression near apical border that terminates in a pit in front of above-mentioned eminence. Termitogaster wasmanni Holmgren, new comb. Xenogaster wasmanni Holmgren, 1911, in Wasmann and Holmgren, Zool. Anz., 38: 429 (Mojos, Bolivia; Eutermes mojosensis Holmgren; Holmgren coll.). Wasmann and Holmgren placed this species in Xenogaster, distinguishing it from X. nigricollis Silvestri by the fact that the SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 93 second antennal segment is longer than the third. This one line "diagnosis" was the only information provided by the authors. It has been evident to me for some time that nigricollis, and presumably wasmanni, belong to Termitogaster. Recently when I had the opportunity to examine the Wasmann types I thought it would be possible to place this species more accurately. Un- fortunately, the specimens in the Wasmann collection labeled "types" of this species are from nests of Nasutitermes chaquimayensis Holmgren, Chaquimayo, Peru (not the recorded host or the type locality), and they do not conform to the one diagnostic character given by Wasmann and Holmgren. The type, if such exists, is probably in the Holmgren collection. Termitogaster impressicollis Seevers. Figure 18, c. Termitogaster impressicollis Seevers, 1937, Ann. Ent. Soc. Amer., 30: 11, pi. 2, figs. 3, 6 (Hacienda de Tenguel, Ecuador; Nasutitermes peruanus Holm- gren; American Museum of Natural History); 1939, Amer. Mus. Nov., no. 1018, text figs. 2, 15; 1946, Rev. Ent., 17: 248. Material examined. Ecuador specimens: Hacienda de Tenguel; Gualaquiza; Faijoo; Hacienda de Santo Domingo; San Miguel; Bucay Chico (USNM); all with Nasutitermes peruanus Holmgren. Rurrenabaque, Beni, Bolivia (USNM). Rio Frio and Santa Anna, Colombia (USNM), with Nasutitermes meinerti Wasmann. Head and elytra brown, pronotum black, abdomen reddish-brown. Head transversely elevated between antennae; surface behind this elevation moderately deeply concave, and clypeus moderately declivous. Distance between antennal fossae less than two-fifths the width of head at eye level. Two or three occipital setae on each side; vertexal setae absent. Dorsum coarsely and densely punctate behind eyes, the punctures finer and less dense at middle; surface between an- tennae impunctate, occipital border very sparsely punctate. Antennae with segments 2 and 3 equal, longer than segment 4, which is longer than 5. Cardo not produced laterally. Segment 2 of maxillary palpi with its ventral surface flattened. Mentum-submentum as in figure 18, c. Pronotum about one-fifth broader than long; disk not deeply or uniformly impressed, but with irregular contours as follows; two rather deep pits at either side of center and a transverse groove, interrupted medially, in front of basal border; an impression before apical border, giving rise to a rather sharp ridge (visible in profile). Termitogaster bicolor Seevers. Figure 18, t. Termitogaster bicolor Seevers, 1937, Ann. Ent. Soc. Amer., 30: 12, pi. 2, figs. 4, 7 (Gualaquiza, Ecuador; Nasulitermes dendrophilus Holmgren; Ameri- can Museum of Natural History); 1946, Rev. Ent., 17: 248. Material examined. The type series. 94 FIELDIANA: ZOOLOGY, VOLUME 40 Color reddish-brown; pronotum black, disk light-brown. Vertex slightly elevated between antennae; clypeus feebly declivous. Distance between antennal fossae only one-third the width of head at eye level. One occipital seta on each side; no vertexal setae. Head punctate as in T. impressicollis. Antennae with segment 3 slightly longer than 2; segments 4 and 5 subequal, each only about two-thirds as long as segment 3. Cardo not produced laterally. Segment 2 of maxillary palpi triangular in outline, its ventral surface flattened. Mentum- submentum as in figure 18, i. Pronotum one-fifth broader than long; disk broadly impressed and much lighter in color than the sides; discal surface irregular due to a number of grooves, pits, and low elevations. Termitogaster emersoni Mann. Figure 18, g. Termitogaster emersoni Mann, 1923, Zoologica, 3: 342, text fig. 34 (Kartabo, British Guiana; Nasutitermes ephratae Holmgren; United States National Museum; Seevers, 1937, Ann. Ent. Soc. Amer., 30, pi. 2, fig. 8; 1939, Amer. Mus. Nov., no. 1018, text fig. 3; 1946, Rev. Ent., 17: 248. Material examined. A series of specimens, including types, from Kartabo, British Guiana; also Rio Claro, Glyan Road, and Northern Range, Trinidad, British West Indies (CNHM, ex Adamson coll.) ; all with Nasutitermes ephratae Holmgren. Color brown, pronotum black. Vertex not elevated between antennae; clypeus feebly declivous, its border slightly emarginate. Distance between antennal fossae about one-half the width of head at eye level. Two or three occipital setae on each side; vertex with a pair of setae medial to antennal fossae at level of their caudal margins. Head very finely punctulate, the punctules often indistinctly defined; occiput and vertex very sparsely and indistinctly punctulate at middle. Pronotum dull, surface with a fine reticulation, often obsolescent, and extremely fine and dense punctulation, but without distinct, easily observed punctures. Antennae with segment 3 a little longer and stouter than 2; segments 4, 5, and 6 decreasing slightly in length. Cardo not produced laterally. Segment of maxillary palpi triangular; segment 3 oval, little more than one-half as broad as long. Men tum-submen turn distinctive (fig. 18, g). Pronotum about one- tenth wider than long; strongly convex; contours irregular, with variable impressions and pits, but without distinctive pattern. Termitogaster chavantinae, new species. Figure 18, h. Closely allied to emersoni, but lacking vertexal setae, and having moderately coarse, distinct head punctures (very fine and indistinct in emersoni}, a distinctive mentum-submentum, and distinct punctures on the pronotum in addition to the very minute reticu- lation. Color brown, pronotum black. Vertex of head not elevated between antennae; clypeus slightly declivous, its margin feebly emarginate. Distance between antennal fossae about one-half the width of head at eye level. Three occipital setae; vertexal setae absent. Head with fine to moderately coarse, distinct SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 95 punctures; these are moderately dense except between antennae. Antennae with segment 3 longer than segment 2 or any of segments 4-10. Cardo not produced laterally. Pronotum moderately densely punctate; punctures distinct and easily discernible; in addition, with a much finer sculpture of fine lines and extremely minute punctules. Mentum-submentum as in figure 18, h; its apical margin more deeply emarginate than in emersoni and wenzeli. Pronotum almost one-tenth wider than long; structure as in emersoni. Length, 2.5 mm. (with abdomen extended). Holotype from Chavantina, Mato Grosso, Brazil, collected January 7, 1947, with Nasutitermes bivalens Holmgren, by Helmut Sick. In collection of Chicago Natural History Museum. No paratypes. Termitogaster magdalenae Seevers Termitogaster magdalenae Seevers, 1946, Rev. Ent., 17: 252 (Puerto Berrio, Antioquia, Colombia; Nasutitermes ephratae Holmgren; Chicago Natural History Museum). Color brown, pronotum black, abdomen reddish-brown. Vertex not elevated between antennae; clypeus slightly declivous. Distance between antennal fossae about one-half the width of head at eye level. One occipital seta on each side; vertexal setae absent. Head finely punctate. Antennae with segment 2 a little longer than any of segments 3-10; segments 2, 3, and 4 decreasing in length. Cardo not produced. Mentum-submentum with only two apical setae. Pronotum more than one-tenth broader than long; a shallow median impression in the apical half, a low, rather broad elevation behind it, and a U-shaped impression caudad and laterad of the medial elevation. Termitogaster nigricollis Silvestri, new comb. Figures 16, b, 18, e. Xenogaster nigricollis Silvestri, 1901, Boll. Mus. Zool. Torino, 16, no. 398, p. 3 (Santa Ana, Misiones, Argentina, and Coxip6, near Cuyaba, Mato Grosso, Brazil; Eutermes arenarius proximus Silv. and E. a. pluriarticulatus Silv.; Silvestri coll.); 1903, Redia, 1: 189, pi. 5, figs. 246-248; Wasmann and Holmgren, 1911, Zool. Anz., 38: 429. Material examined. Birigui, Sao Paulo, Brazil (R. L. Araujo); Jabaquara, Sao Paulo, Brazil (R. L. Araujo); with Nasutitermes bivalens Holmgren. Distribution. Misiones, Argentina, with Nasutitermes sanctae- anae Holmgren; Sao Paulo, Brazil, with Nasutitermes bivalens Holmgren; and Mato Grosso, Brazil, with Nasutitermes pluri- articulatus Holmgren. Wasmann and Holmgren recorded this species from Mojos, Bolivia, but their identification is questionable. Color reddish-brown, pronotum black. Surface of head scarcely elevated between antennal fossae; clypeus feebly declivous. Distance between antennal fossae about one-half the width of head at eye level. Head with moderately 96 FIELDIANA: ZOOLOGY, VOLUME 40 dense and coarse punctation behind eyes, otherwise sparsely and finely punctulate. Two occipital setae on each half, vertexal setae absent. Antennae with segment 2 longer than 3; segments 3, 4, and 5 decreasing in length, 5 and 6 equal. Cardo not produced. Segment 3 of maxillary palpi about two-thirds as broad as long. Mentum-submentum distinctive (fig. 18, e). Pronotum more than one-tenth broader than long; with a transverse sub-basal impression, the ends of which extend forward as weak longitudinal impressions to terminate as moderately deep pits; and with a small eminence in front of transverse impression. Length, 3 mm. Termitogaster wenzeli, new species. Figure 18, /. Closely related to nigricollis Silvestri but distinguished by the structure and chaetotaxy of the mentum-submentum and by the punctation of the head. Head reddish-brown, pronotum black, elytra light-brown, abdomen reddish- brown. Head very sparsely and finely punctulate, the pale V-shaped area between antennae im punctate; vertex and occiput almost impunctate medially. Head between antennal fossae very feebly elevated; clypeus very slightly declivous, its border feebly emarginate. Distance between antennal fossae about one-half the width of head at eye level. Two occipital setae on each side; vertex without a pair of setae at level of posterior margins of antennal fossae. Antennae with segment 2 longer than 3; segments 3 and 4 equal; 5-10 decreasing very slightly. Mentum-submentum distinctive (fig. 18, /). Cardo not produced. Segment 3 of maxillary palpi three-fifths as broad as long. Pronotum strongly convex, only one-twentieth broader than long; with a pair of moderately deep discal pits, and other minor irregularities. Length, 3.5 mm. Holotype from Recife, Pernambuco, Brazil; collected July 3, 1944, with Nasutitermes bivalens Holmgren, by Rupert L. Wenzel. In collection of Chicago Natural History Museum. No paratypes. Termitogaster beniensis Seevers. Figure 18, d. Termitogaster beniensis Seevers, 1946, Rev. Ent., 17: 250 (Huachi, Beni, Bolivia; Nasutitermes chaquimayensis Holmgren; United States National Museum). Material examined. Bolivia: Huachi; Rurrenabaque; Rio Colo- rado; and Cachuela Esperanza; all with N. chaquimayensis Holm- gren. Closely related to nigricollis; distinguished by having its second and third antennal segments equal in length, by having more occipital setae, and by the more slender third segment of the maxil- lary palpi. Color light reddish-brown; pronotum piceus, abdomen dark reddish-brown. Head not elevated between antennae, clypeus not declivous, its border feebly SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 97 emarginate. Distance between antennal fossae about one-half the width of the head at eye level. Two or three occipital setae; vertexal setae absent. Antennae with segments 2 and 3 subequal in length; 4, 5, and 6 decreasing slightly in length. Cardo not produced. Segment 3 of maxillary palpi twice as long as broad. Men- tum-submentum distinctive (fig. 18, d). Pronotum more than one-tenth broader than long; convex, with relatively minor surface contours; a shallow arcuate impression near basal border with ends extending forward as lateral impressions to terminate as rather deep pits. Length, 3 mm. Termitogaster emarginatus Seevers Termitogaster emarginatus Seevers, 1946, Rev. Ent., 17: 251 (Mogi-mirim, Sao Paulo, Brazil; Nasutitermes globiceps Holmgren; Chicago Natural History Museum). Material examined. The type and additional specimens from lepe and Santa Ernestina, Sao Paulo, Brazil, collected by R. L. Araujo with Nasutitermes globiceps Holmgren (determined by R. L. Araujo). Head reddish-brown, pronotum black, elytra brown, abdomen light reddish- brown. Vertex not elevated between antennae, clypeus very feebly declivous, its margin with a deep, arcuate emargination. Five occipital setae on each side; not vertexal setae; only 6-8 clypeal setae on each half. Antennae with segments 2 and 3 about equal in length, but segment 2 distinctly broader; segments 4 and 5 subequal, 6-10 decreasing in length. Cardo not produced. Pronotum almost one-fifth broader than long; very strongly convex; surface with a feebly longi- tudinal impression on each side, a shallow, broader median impression, and a few other minor irregularities. Elytra with only one humeral seta. Length, 3 mm. Termitogaster brevis Mann Termitogaster brevis Mann, 1923, Zoologica, 3: 344 (Kartabo, British Guiana; Nasutitermes costalis Holmgren; United States National Museum); Seevers, 1937, Ann. Ent. Soc. Amer., 30, pi. 2, fig. 9; 1939, Amer. Mus. Nov., no. 1018, text fig. 4; 1946, Rev. Ent., 17: 248. Material examined. Kartabo, British Guiana (type series). Manzanilla (CNHM), Arena Forest (CNHM, ex Adamson coll.), and Northern Range (CNHM, ex Adamson coll.), Trinidad, British West Indies; all with N. costalis Holmgren. Color brown, pronotum black. Surface of head between antennae convexly elevated; clypeus concave, its border moderately emarginate. Head finely and inconspicuously punctulate. Antennae with segments 2 and 3 equal, segments 4-10 subequal; segments 2 and 3 relatively short, segment 10 two-thirds as long as segment 2. Cardo with a suggestion of lateral production. Pronotum only a trifle broader than long. Legs relatively short; metathoracic legs, for example, with neither femora nor tibiae longer than pronotum. 98 FIELDIANA: ZOOLOGY, VOLUME 40 Termitogaster diversicollis, new species. Figure 18, j. Distinguished from all other members of the genus by its dis- tinctive pronotum, and by its uniform light-brown coloration. The general appearance is that of a Xenogaster, but except for the unusual pronotum this species agrees with the pattern of characters of Termitogaster. Color light-brown throughout. Head not elevated between antennal fossae; clypeus flat, its border moderately emarginate. Head with low, oblique ridges extending caudad and mesad from antennal fossae to occipital border where they join, enclosing a triangular impression. Head subopaque, very fine and moderately densely punctulate. One occipital seta on each side. Antennae with segment 3 longer than 2 or 4. Mentum-submentum as in figure 18, j; postgenae lateral to gula with a suggestion of sulci. Pronotum as broad as long; basal and anterior margins arcuate; broadest near anterior margin; sides arcuate in front and con- verging basally, slightly sinuate. Pronotal disk broadly and strongly impressed, the impression broadest near base and narrower anteriorly; basal half of the impression with a very conspicuous, triangular eminence. Abdomen of holotype stenogastric (atypical in appearance, but with the structure and chaetotaxy of the genus). Length, 2 mm. (stenogastric individual). Holotype from Villavicencio, Meta, Colombia; collected July 11, 1938, with Nasutitermes ephratae Holmgren, by Charles H. Seevers. In collection of Chicago Natural History Museum. NEOTERMITOGASTER Seevers Neotermitogaster Seevers, 1939, Amer. Mus. Nov., no. 1018, p. 8. Type species: Neotermitogaster colonus Seevers. Most closely related to Termitogaster, from which it differs in the form and pilosity of the mentum-submentum, in not having the gula and submentum separated by a suture, and in the dentition of the mandibles, the distinctive chaetotaxy of head and pronotum, distinctive punctation, and pronotal contours. Head feebly impressed between antennae; clypeus slightly convex, its margin feebly emarginate. Mandibles broader than in Termitogaster, each with a single broad, blunt median tooth. Antennae geniculate, rather stout; scape almost as long as segments 2-5 combined; thickened distally; segments 2-10 cylindrical. Mentum-submentum distinctive (fig. 18, k); not separated from gula. Maxilla with large, subtriangular cardo that is not produced laterally. Segment 2 of maxillary palpi large, transverse, moderately arcuate; segment 3 oval. Pronotum slightly transverse; moderately convex; sides rather strongly deflexed; disk with a transverse impression, the ends of which deepen to form crater-like pits; the impression with a shallow, elongated pit at middle. Elytra contiguous; as broad as pronotum but somewhat shorter, the sutural length a little more than one-half the pronotal length; apical margin of elytra deeply emarginate. Abdomen similar to that of Termitogaster; the last paratergites fused at middle. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 99 Neotermitogaster colonus Seevers. Figures 16, d, 18, k. Neotermitogaster colonus Seevers, 1939, Amer. Mus. Nov., no. 1018, p. 9, text figs. 9, 11, 19, 29, 36 (Hacienda de Santo Domingo, Ecuador; Nasutitermes peruanus Holmgren; American Museum of Natural History). Head and thorax fuscous, abdomen brown; strongly shining. Head, prono- tum, elytra, and abdomen finely punctate, the punctures arranged in dense clusters, especially around setiferous punctures. Head with 7 clypeal setae in two rows; vertex with one median seta; dorsum and genae with numerous short, fine hairs. Antennae with segment 2 about three-fourths as long as 3; segment 2 subequal to each of segments 4-7; segments 8-10 a trifle shorter than the preceding. Pronotum with 8 rather stout .setae on apical margin (the median 6 longer) ; disk with 6 setae in two longitudinal rows, and with numerous fine, suberect pale hairs. Elytra with one seta on lateral border and one on disk close to anterior border; otherwise glabrous except for very sparse hairs near pronotum. Tergites 3-7 with a subapical row of 4 erect setae; tergites 4-6 with an additional basal row of 4; eighth tergite with an apical row of 4 and a basal row of 2; tergites 3-6 with 12-16 semi-recumbent bristles on apical margin. TERMITOIDES Seevers Termitoides Seevers, 1939, Amer. Mus. Nov., no. 1018, p. 7. Type species: Termitoides marginatus Seevers. This genus is distinguished by its clypeal horn and the form of its mentum-submentum. The pronotum and elytra resemble those of Xenogaster, except that they do not have the granulose sculpture. Clypeus with a large median "horn," a broad acute process extending above and beyond the apical border; clypeus lateral to the horn, somewhat depressed. Antennae geniculate; scape longer than the three following segments combined; segments 2-11 slender, cylindrical. The slender mandibles with a moderately strong subapical tooth, and a larger medial tooth having a slightly hooked tip. Mentum-submentum (fig. 17, <) distinctive, its narrow apex almost straight, its anterior angles strongly truncated and its sides sinuate for a distance and then converging to the gula. Segment 2 of maxillary palpi about as broad as long, segment 3 oval, elongated. Pronotum approximately one-fifth broader than long, its apical border moderately bisinuate, sides evenly rounded from base to apex, and posterior border feebly arcuate. Pronotal disk strongly impressed medially, with two very deep pits. Elytra not contiguous, their inner margins diverging posteriorly. Termitoides marginatus Seevers. Figure 17, t. Termitoides marginatus Seevers, 1939, Amer. Mus. Nov., no. 1018, p. 8, text figs. 6, 12, 18, 31, 35 (Hacienda de Santo Domingo, Ecuador; Nasutitermes peruanus Holmgren; American Museum of Natural History). Head and pronotum smoky-brown, pronotal disk lighter; abdomen reddish- brown. Head, pronotum, and elytra finely, sparsely punctulate. Pronotum with an oblique row of 4 setae bordering disk; elytra with one seta on lateral margin. Antennal segment 2 nearly twice as long as broad; segments 2-6 de- 100 FIELDIANA: ZOOLOGY, VOLUME 40 creasing in length and increasing in width; segment 6 broader than long; 6-7 subequal, 8-10 slightly shorter, subequal. Chaetotaxy of abdomen as follows: Tergites 3-6 with about 12 semi-erect setae on posterior margin and 4 in a sub- apical row; tergite 7 with 2 apical and 2 subapical setae; tergites rather densely clothed with fine, suberect hairs. Sternite 3 with three rows of 4, 4, 12 setae; sternites 4-6 with 8 in a basal row and 14 in an apical row; sternite 7 with 8 in a basal row and 10 in an apical row; sternite 8 with 8 in an apical row. Length, 3-4 mm. XENOGASTER Wasmann Xenogaster Wasmann, 1891, Verh. Zool. Bot, Ges. Wien, 41: 651; Fenyes, 1920, Gen. Insect., 173B : 349. Type species: Xenogaster inflata Wasmann. Distinguished from Termitogaster, with which it is often confused, in these respects: Inner margins of elytra separated for almost their entire length; pronotal disk broadly impressed; submentum and gula not separated by a suture; coloration brown; head, pronotum, and elytra asperately punctate and reticulated so as to appear granulose. Surface of head not elevated between antennal fossae. Clypeus strongly deflexed immediately in front of antennal fossae to produce a transverse "carina" connecting the anterior margins of the fossae (inflata, nana, subnuda, pilosula) ; or clypeus not deflexed and at most feebly declivous (fossulata, reichenspergeri, glabriventris, inquilina). Antennae strongly geniculate; scape often longer than the next four segments combined; the segments variable: Scape 2 3 4 5 6 789 10 11 inflata 50 10 11.5 11.5 9 9 8.5 8 8 8 20 pilosula 50 14 14 14 12 10 10 9 8 8 22 subnuda 40 9 12 9 8 8 8 7 7 7 18 nana 36 8 6 6 6 6 6 6 6 6 16 reichenspergeri 58 17 12 10 10 8 8776 18 glabriventris 54 18 12 12 10 9 8888 19 fossulata 35 10 8 7 7 7 7 7 7 7 20 inquilina 50 14 12 10 8 8 8 7 7 7 18 Mentum-submentum variable in form and pilosity (fig. 17, fc-m, p-s); not separated from gula by a suture. Maxillary palpi with segment 2 shaped like a compressed cornucopia; segment 3 variable in size: as broad as long (nana); four-fifths as broad as long (subnuda) ; two-thirds as broad as long (inflata, pilosula, inquilina, reichenspergeri, fossulata); one-half as broad as long (glabriventris). Pronotal disk broadly and deeply impressed ; usually brown in color and with a granulose texture. Pronotal proportions variable: one-fifth broader than long (subnuda); one-eighth broader than long (pilosula and nana); one-tenth broader than long (inflata); subequal in length and width (glabriventris, inquilina, and fossulata); one-tenth longer than broad (reichenspergeri). Elytra separated for almost their entire length, their inner margins usually pale and somewhat hyaline. Abdomen very similar to that of Termitogaster, the sclerites usually separated by areas of non-sclerotized cuticle. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 101 Remarks. It may be justifiable to separate the four species with a feebly declivous clypeus (reichenspergeri, glabriventris, inquilina, and fossulata) from those with a strongly deflexed clypeus, but I believe that we know too little about this group to do so at present. I am not sure that the four species are actually as closely allied as this one character may suggest. The condition of the clypeus is quite variable in the subtribe Termitogastrina and there is about as much variation among the species of Termitomorpha as among those of Xenogaster. Xenogaster seems to occur primarily in the southern Neotropical Region, seven of its species having been found in southern Brazil and one in British Guiana. Termitogaster, by contrast, has nine of its fourteen species in northern South America and Central America, three in Bolivia, and only two in southern Brazil. KEY TO SPECIES OF XENOGASTER 1. Clypeus strongly deflexed anterior to antennal fossae; pronotum broader than long 2 Clypeus not deflexed anterior to antennal fossae, at most moderately de- clivous; pronotum subequal in length and width or longer than broad 5 2. Carina between anterior margins of antennal fossae almost straight 3 Carina of front of head distinctly arcuate pilosula, new sp. 3. Third antennal segment distinctly longer than any of segments 2-10. subnuda, new sp. Third antennal segment not longer than some of segments 2-10 4 4. Antennal segments 3 and 4 subequal, each longer than segment 2 or any of segments 5-10 inflata Wasmann Antennal segments 3 and 4 subequal, each shorter than segment 2 and not longer than any of segments 5-10 nana, new sp. 5. Antennal scape extremely long, more than one-half as long as segments 2-11 combined; distal half of its anterior surface broadly concave, the proximal half of scape conspicuously broader reichenspergeri, new sp. Antennal scape less than one-half the combined length of segments 2-11; scape nearly uniform in width 6 6. Second antennal segment one-half longer than segment 3. glabriventris, new sp. Second antennal segment at most one-fifth longer than segment 3 7 7. Sides of pronotum dark reddish-brown, tinged with black, the disk lighter brown ; coloration of body reddish-brown inquilina Borgmeier Pronotum uniform in color; coloration of body pale testaceous. fossulata Mann Xenogaster inflata Wasmann. Figure 17, k. Xenogaster inflata Wasmann, 1891, Verh. Zool. Bot. Ges. Wien, 41: 652, pi. 6, figs. 16-24 (Santa Catarina, Brazil; Eutermes arenarius fulviceps Silvestri; Wasmann coll.); Silvestri, 1903, Redia, 1: 190, pi. 5, figs. 250- 251, pi. 6, fig. 260. 102 FIELDIANA: ZOOLOGY, VOLUME 40 Material examined. The specimens in the Wasmann collection, mounted and in alcohol, which were determined as this species by Wasmann, belong to two species; one of these species is described elsewhere in this paper as X. subnuda. From the four mounted specimens bearing "type" labels, I have selected as lectotype the one with the label "Santa Catharina (Hetscko)" as it is evidently the specimen upon which Wasmann based X. inflata. A series of 34 mounted and unmounted specimens from Joinville, Santa Catarina, collected by Schmalz (1891, 1903, 1904), consists of 22 X. inflata and 12 X. subnuda. Specimens from Joinville were distributed to other collections by Wasmann as "cotypes" of X. inflata; some of these are probably X. subnuda. Other specimens studied: Two specimens from Joinville, Santa Catarina, labelled cotypes, accompanied by Nasutitermes aquilinus Holmgren (USNM). A series from Blumenau, Santa Catarina (ex Reichensperger coll.), with Nasutitermes itapocuensis Holmgren; and specimens from Ilha Grande, Rio de Janeiro, Brazil (ex Helmut Sick), with Nasutitermes itapocuensis Holmgren. Coloration of head, pronotum, and elytra dark reddish-brown, abdomen reddish-brown. Clypeus deflexed in front of antennal fossae, the deflexed part in a vertical plane; clypeus thus with a transverse "carina" between the antennal fossae. Vertex between antennal fossae broadly and shallowly concave; vertex and occiput transversely convex. Antennae with scape five times as long as seg- ment 2; segments 3 and 4 subequal, each longer than segment 2 or any of segments 5-10. Mentum-submentum distinctive in form and pilosity (fig. 17, k). Seg- ment of maxillary palpi two-thirds as broad as long. Pronotum about one-tenth broader than long, its disk broadly and rather deeply impressed. Head, pronotum, and elytra coarsely reticulated and asperately punctate. Head without erect setae; pronotum with 3 setae along the lateral rim of the discal impression; elytra with a few feeble setae. Tergites 3-6 with a marginal row of semi-erect dark bristles and tergites 4-6 with an additional submarginal row of 6 long, erect bristles; tergite 7 with 4 long submarginal bristles. Tergites with numerous fine, erect pale hairs. Sternites with marginal and sub-basal rows of long, erect black bristles. Sternites moderately densely clothed with pale hairs with curved tips. Length, 3-3.5 mm. Xenogaster pilosula, new species. Figure 17, I. Coloration of head, pronotum, and elytra dark reddish-brown, abdomen light-brown. Clypeus strongly deflexed, the resulting transverse "carina" arcuate. Antennae with scape as long as in inflata but only a little more than three times as long as segment 2; segments 2, 3, and 4 subequal in length; each longer than any of segments 5-10. Mentum-submentum distinctive (fig. 17, I). Maxillary palpi as in inflata. Pronotum about one-eighth broader than long, impressed as in inflata. Head, pronotum, elytra, and tergites coarsely reticulate and asperately punctate. Tergites 3-6 with apical row of about 15 long, coarse, recumbent setae SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 103 and usually with a few black, semi-erect subapical bristles; tergites 7 and 8 with 4 long erect subapical bristles. Tergites with a moderately dense vestiture of pale, semi-erect hairs. Vestiture of sternites as in inflata. Length, 3 mm. Holotype from Blumenau, Santa Catarina, Brazil (ex Reichens- perger collection), with Nasutitermes sp. In collection of Chicago Natural History Museum. Paratypes. Ten specimens, same data as holotype; four speci- mens, Passa Quatro, Minas Gerais, Brazil (ex Reichensperger collection), with Nasutitermes ehrhardti Holmgren. Paratypes in collections of Chicago Natural History Museum, United States National Museum, and A. Reichensperger. Xenogaster subnuda, new species. Figure 17, r. Head and pronotum brown, elytra dark brown with paler inner margins; abdomen reddish-brown. Clypeus deflexed in front of antennal fossae, resulting in an almost straight "carina" as in inflata. Antennae with scape relatively short, less than five times as long as segment 2; segment 3 distinctly longer than any of segments 2-10. Mentum-submentum distinctive (fig. 17, r). Segment 3 of maxillary palpi about four-fifths as broad as long. Pronotum about one-fifth broader than long, impressed as in inflata. Head, pronotum, and elytra coarsely reticulate, and asperately punctate. Tergites with marginal rows of long re- cumbent bristles, but otherwise almost devoid of setae, except that tergites 3-5 have one subapical seta near each lateral margin, and tergites 6-8 have a few additional subapical bristles; tergites shining, with only traces of reticulation. Tergites with a sparse vestiture of short hairs; sternites with a basal and a sub- apical row of erect bristles; sternites almost glabrous, having only a sparse, short stubble. Length, 3-3.5 mm. Holotype from Blumenau, Santa Catarina, Brazil (ex Reichens- perger collection), with Nasutitermes itapocuensis Holmgren. In collection of Chicago Natural History Museum. Paratypes. Forty-four specimens, same data as holotype; nine specimens, Passa Quatro, Minas Gerais, Brazil, with Nasutitermes sp.; one specimen, Ilha Grande, Rio de Janeiro, Brazil, with Nasu- titermes sp. In collections of Chicago Natural History Museum, United States National Museum, Reichensperger, and Borgmeier. Xenogaster nana, new species. Figure 17, q. Color reddish-brown, elytra a little darker. Clypeus sharply deflexed in front of antennal fossae, resulting in a carina with a medial incision. Antennae with scape four and one-half times as long as segment 2, segment 2 longer than any of segments 3-10, which are subequal in length. Mentum-submentum 104 FIELDIANA: ZOOLOGY, VOLUME 40 distinctive (fig. 17, q). Segment 3 of maxillary palpi subequal in length and width. Pronotum about one-eighth broader than long, its discal impression broad and deep; shallow subapically, deep at middle, broad and shallow sub- basally. Head, pronotum, and elytra coarsely reticulated and asperately punctate; elytral granulations quite coarse. Tergites shining, with an inconspicuous reticu- lation or very fine asperate punctures. Tergites 3-8 with the usual marginal row of setae, and tergites 5-8 with an additional submarginal row; sternites 3-8 with erect bristles; glabrous except for an extremely sparse short stubble. Length, 2.5 mm. Holotype from the state of Minas Gerais, Brazil (ex Reichens- perger collection); with Nasutitermes jaraguae Holmgren. In col- lection of Chicago Natural History Museum. Pamtypes. Six specimens, same data as holotype. In collections of Chicago Natural History Museum, United States National Mu- seum, and Reichensperger. Xenogaster reichenspergeri, new species. Figure 17, m. Distinguished from the four preceding species by the non- deflexed clypeus; distinguished from the following species by its distinctive antennal scape, the proportions of its antennal segments, the proportions of its pronotum, the distinctive men tum-submen turn, the glabrous sternites, and the more reduced elytra. Color reddish-brown; lateral and anterior margins of pronotum black. Clypeus slightly declivous, its margin straight; vertex shallowly concave, with a low, smooth ridge between antennal fossae. Antennae with an unusually long scape which is more than one half as long as segments 2-11 combined; segment 2 almost one-half longer than segment 3; segments 3-10 decreasing in length and in- creasing in width. Antennal scape distinctive; with one margin of its anterior face elevated as a conspicuous "tooth" near middle, and the anterior surface of its distal half broadly and shallowly emarginate. Mentum-submentum dis- tinctive (fig. 17, m). Maxillary palpi very similar to those of inflata. Pronotum about one-tenth longer than broad; discal impression narrower than in the other species, most deeply impressed in the middle one-third; basal one-third weakly impressed, with a smooth medial elevation. Inner margins of elytra strongly divergent behind scutellum (elytra exhibit more degenerative changes than in the other species), each elytron so decreasing in width that the apex is approximately one-half as broad as base. Head reticulated and asperately punctate; pronotum reticulated only at sides, with the impressed area asperately punctate; elytra reticulated and asperately punctate. Sternites glabrous except for an almost imperceptible stubble of fine hairs. Clypeus with 6 or more long black setae and many finer hairs; surface between antennae with a pair of black bristles. Pronotum with sparse, pale, erect setae along anterior and lateral margins of the impression. Tergites with an inconspicuous pilosity; sternites with basal and marginal rows of bristles. Length, 3.25 mm. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 105 Holotype from Blumenau, Santa Catarina, Brazil (ex Reichens- perger collection); with Nasutitermes jaraguae Holmgren. In collection of Chicago Natural History Museum. Paratypes. Fourteen specimens, same data as holotype; two specimens, Passa Quatro, Minas Gerais, Brazil (ex Reichensperger collection), with Nasutitermes, new sp.; two specimens, Rio Negro, Parana, Brazil (ex Reichensperger collection), with Nasutitermes jaraguae Holmgren. Xenogaster glabriventris, new species. Figure 17, p. Color reddish-brown; lateral margins of pronotum rufo-piceous. Clypeus very feebly declivous, its anterior margin arcuate; surface between antennal fossae not at all elevated. Antennae with scape very long, approximately one- half as long as segments 2-11 combined; segment 2 one-half longer than 3; seg- ments 3 and 4 subequal, segments 5-10 decreasing in length. Mentum-sub- mentum distinctive (fig. 17, p). Maxillary palpi as in inflata except that segment 3 is only one-half as broad as long. Pronotum subequal in width and length; disk broadly impressed. Head reticulated, only feebly and sparsely asperate; surface more shining than usual; pronotum with a dense asperate stubble (appearing granulose) ; elytra with a very coarse reticulation of raised lines; tergites reticulated and in places with short, fine, recumbent hairs. Tergites with marginal rows of pale recumbent bristles; tergites 3-5 without erect bristles, 6-8 with one con- spicuous black bristle near each lateral margin, 7 and 8 with an additional medial pair. Sternites smooth, shining, with no sculpture except a very faint reticulation ; 6 long marginal bristles and a pair of sub-basal ones but no finer hairs of any kind. Length, 3-3.5 mm. Holotype from Blumenau, Santa Catarina, Brazil (ex Reichens- perger collection), with Nasutitermes jaraguae Holmgren. In col- lection of Chicago Natural History Museum. Xenogaster fossulata Mann Xenogaster fossulata Mann, 1923, Zoologica, 3: 359 (Kartabo, British Guiana; Nasutitermes intermeditis Banks; United States National Museum). Material examined. The holotype. Color pale testaceous. Clypeus horizontal, not deflexed; surface between antennae not elevated. Antennae with scape about two-fifths as long as segments 2-11 combined; segment 2 longer than 3; segment 3 a little longer than 4; segments 4-10 subequal. Mentum-submentum finely setulose (not studied in detail). Maxillary palpi with segment 3 about two-thirds as broad as long. Pronotum subequal in length and width; disk deeply impressed but the impression more restricted than in inflata, its deepest part beginning about one-third of the way back and terminating short of the base; impression with a deeper pit on each side near its anterior limit. Head sparsely and finely setulose, vertex almost glabrous; pronotum sparsely and finely setose, with no apparent pattern; elytral surface closely granulose, and moderately densely setose. Tergites smooth, shining, 106 FIELDIANA: ZOOLOGY, VOLUME 40 feebly reticulated, each with one or two rows of long, pale, semi-recumbent hairs. Sternites with an apical row of sparse, long, pale to brown setae, and with two or three irregular rows of short bristles. Xenogaster inquilina Borgmeier. Figures 16, e, 17, s. Xenogaster inquilina Borgmeier, 1950, Rev. Ent., 21: 667, text figs. 52-61 (Parada Deodoro, Sto. Amaro, Sao Paulo, Brazil; Nasutitermes sp.; Institute Biologico, Sao Paulo). Material examined. Four paratypes from same colony as holo- type, with Nasutitermes proximus Silvestri (determined after pub- lication of Borgmeier's paper); 134 specimens, Santo Amaro, Sao Paulo, Brazil, April 10, 1951 (R. L. Araujo), with Nasutitermes itapocuensis Holmgren (determined by R. L. Araujo). Related to glabriventris and fossulata, from which it differs in characteristics of antennae and submentum-mentum. Color brown. Clypeus moderately strongly declivous from the anterior margin of antennal fossae, resulting in an obtuse "ridge" between the antennae. Antennae with scape about two-fifths as long as segments 2-11 combined; seg- ments 2-5 decreasing in length, 5-10 not appreciably different. Mentum-sub- mentum distinctive (fig. 17, s). Maxillary palpi with segment 3 about two-thirds as broad as long. Pronotum approximately equal in length and width; its sides darker brown than the disk. Pronotum deeply impressed at middle, with two large, circular pits at deepest point; impression about one-half as broad as pro- notum. Head with a coarse reticulation of raised lines, and with fine asperate punctures laterally; pronotum coarsely reticulated and with very fine, asperate punctulation; elytra coarsely reticulated and asperately punctate. Sternites glabrous, strongly shining, with obsolescent reticulation. Sternites 3-6 with two or three rows of long erect setae, as follows: third: 2 sub-basal, 2 at middle, 8 subapical; fourth: 4 sub-basal, 10 subapical; fifth: 4 sub-basal, 10 subapical; sixth: 4 sub-basal, 8 subapical. XENOPELTA Mann Xenopelta Mann, 1923, Zoologica, 3: 356. Type species: Xenopelta cornuta Mann. Ceratoxenus Mann, 1923, Zoologica, 3: 360; Emerson, 1935, Ann. Ent. Soc. Amer., 28: 369. Type species: Ceratoxenus tricornis Mann. Head triangular in profile (margin of pronotum forming base of triangle, the sloping dorsum of head one of the sides, and ventral margin of postgena the other). Clypeus with a carina medial to each antennal fossa and a smaller medial pro- tuberance; abruptly deflexed in front of these processes. Pronotal disk strongly impressed; one larger, deeper basal impression, and a smaller shallow, apical impression. Elytra strongly divergent a short distance behind scutellum; elytra broadly impressed, the impression bounded laterally by longitudinal carinae. Gula moderately broad, short, not separated from submentum by a suture. Mentum-submentum distinctive (fig. 18, o); large, robust, trapezoidal in form, SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 107 and densely setose. Maxillary palpi with segment 2 cornucopia-like, and segment 3 broad at base and narrow at apex. Abdomen with moderate areas of non-sclerotized integument. Inner para- tergites 3-6 about one-fifth as broad as tergites, separated only slightly from them. Seventh paratergite long, slender, arcuate. Outer paratergites about three-fourths as broad as inner ones, separated from them to a slight degree but contiguous with sternites. Remarks. Emerson (1935) synonymized Ceratoxenus, pointing out that cornuta and tricornis are closely allied species. Xenopelta cornuta Mann. Figures 16, c, 18, o. Xenopelta cornuta Mann, 1923, Zoologica, 3: 357, text fig. 38 (Kartabo, British Guiana; Nasutitermes guayanae Holmgren; United States National Museum); Emerson, 1935, Ann. Ent. Soc. Amer., 28: 369, text fig. 5. Material examined. Type material from British Guiana. In addition, specimens from Barro Colorado Island, Panama Canal Zone; with Nasutitermes Columbians Holmgren. Head, pronotum, and the elytral sides closely reticulated, the surface some- what dull. Clypeus setose, dorsum of head without setae; pronotal sides with scattered short setae; elytra with a pair of long setae in front of impressed disk; sides of elytra with short, stubby bristles. Tergites 3-5 with marginal rows of long, pale, recumbent setae and numerous short, erect hairs; tergites 6-8 without the above vestiture; tergite 7 with a pair of long, dark setae and tergite 8 with two pairs. Sternite 3 with numerous moderately long, erect setae, sternites 4-7 with two or three irregular rows of moderately long, pale setae. Vertex of head with a large shallow V-shaped impression, bounded by smooth, low elevations; head with an oblique, cariniform elevation medial to each antennal fossa, each carina sloping toward the median line to a small, rounded clypeal eminence. Clypeus strongly deflexed, clothed with a vestiture of dense white pubescence. Antennae (44: 10: 9: 12: 8: 8: 8: 8: 7: 7: 20) with segment 4 longer than any of segments 2-10. Maxillary palpi with segment 2 roughly triangular, its upper margin slightly arcuate; segment 3 twice as long as broad, attenuated, its apex one-half as broad as base, segment 4 short, only one-fifth as long as third. Men- tum-submentum as in figure 18, o. Length, 2.5 mm. Xenopelta tricornis Mann Ceratoxenus tricornis Mann, 1923, Zoologica, 3: 360, text fig. 39 (Kartabo, British Guiana; Nasutitermes guayanae Holmgren; United States National Museum). Xenopelta tricornis Mann, Emerson, 1935, Ann. Ent. Soc. Amer., 28: 369, text fig. 6; with Nasutitermes similis Emerson. Material examined. Paratypes. Dorsum of head as in cornuta except that it is tricornuate: the carinae medial to antennal fossae strongly produced as long acute 108 FIELDIANA: ZOOLOGY, VOLUME 40 processes, and the clypeus produced medially as a stout obtuse process. Antennae (36: 10: 8: 9: 8: 8: 7: 6: 6: 6: 16) differing from cornuta in the proportions of some segments, chiefly in the shorter first and fourth segments. Maxillary palpi with segment 3 less than twice as long as basal width; its round apex narrower than the base. TERMITOTIMA Wasmann Termitotima Wasmann, 1916, Zool. Jahrb. Syst., 39: 188; Cameron, 1939, Fauna Brit. India, Coleopt., 4: 40. Type species: Termitotima assmuthi Wasmann. Termitotima seems to have been derived from Xenogaster-\ike stock and shares numerous characters with the present-day species of Xenogaster. With respect to tergite characters, it occupies an intermediate position between the neotropical genera of Termito- gastrina and most of the paleotropical species. Head capsule unmodified; clypeus moderately declivous. Antennae strongly geniculate, scape longer than three following segments combined. Gula moder- ately broad. Postgenae without sulci. Second and third segments of maxillary palpi subequal in length. Pronotum a little more than one-third broader than long. Pronotal apex almost straight; sides sinuate, the basal one-half somewhat narrower than apical half (11: 12); base arcuate but straight in medial area. Pronotal disk broadly but shallowly depressed, the depressed area with a low median elevation. Inner elytral margins divergent from a short distance behind scutellum. Elytra hyaline. Wings vestigial. Tarsi 4, 4, 4-segmented; the point of fusion of original fourth and fifth segments visible. Abdomen broad; in appearance much like that of Xenogaster. Third sternite very large; its length at median line approximately one-half the width; its pos- terior margin straight. Sixth tergite bent near each lateral margin; perhaps representative of the first stage in the extreme condition of the sixth tergite found in most palaearctic genera of Termitogastrina. Seventh tergite obtrapezoidal. Termitotima assmuthi Wasmann Termitotima assmuthi Wasmann, 1916, Zool. Jahrb. Syst., 39: 188, pi. 4, fig. 15 (Khandala, India; Eutermes biformis Wasmann; Wasmann coll.). Material examined. India: Khandala (type). Host record. The host of the type is Trinervitermes biformis Wasmann. Head, pronotum, and elytra pale testaceous; abdomen light rufo-testaceous. Head and pronotum finely granulose. Elytra with a covering of extremely fine microsetae. Body for the most part devoid of macrosetae. Seventh and eighth tergites with moderately dense vestiture of black hairs; sternites 3-6 with an apical row of very short, erect hairs and extremely few elsewhere. Inner para- SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 109 tergites of segment 7 with an apical cluster of about 10 moderately long, brown hairs. Seventh sternite produced near each lateral margin to form rounded lobes over eighth sternite and these processes bear clusters of perhaps 15-20 brown setae. Relative lengths of antennal segments as follows: 44: 10: 12: 13: 10: 9: 9: 8: 8: 8: 17. Scape one-half broader than the other segments, which are subequal in width. Relative lengths and widths of tergites: third (8, 90), fourth (16, 100), fifth (16, 105), sixth (14, 100), seventh (70, maximum width 90, minimum width 45), eighth (50, 65). Width of inner paratergites 20. Outer paratergites slender; third distinct, 4-5 almost completely fused to sternites, sixth partially fused, seventh distinct. IDIOGASTER Wasmann Idiogaster Wasmann, 1912, Zeitschr. wiss. Zool., 101: 89; Fenyes, 1918, Gen. Insect., 173A: 62. Type species: Idiogaster escherichi Wasmann. This genus appears to be more closely related to Termitotima than to any of the known African genera. The abdomen of the type specimen has the terminal segments telescoped to a degree that makes it difficult to examine the sixth and seventh tergites accur- ately. The sixth tergite is probably relatively short (it is partially covered by the fifth) and does not seem to be bent at all. The seventh tergite is obtrapezoidal, and it is probable that the anterior end is almost transverse. I am unable to state in what respects Idiogaster is different from Termitotima, except that the pronotum is relatively broader (three- fifths broader than long and not twice as broad as long, as Wasmann indicated). Idiogaster escherichi Wasmann Idiogaster escherichi Wasmann, 1912, Zeitschr. wiss. Zool., 101: 90, pi. 5, figs. 3, 3, a (Urwald Dongola, Eritrea; Eutermes rapulum SjOstedt; Was- mann coll.). Material examined. Eritrea: Urwald Dongola (type). Host record. The host is Trinervitermes rapulum Sjostedt. Head finely granulated; pronotum smooth, almost impunctate. Seventh ter- gite with several irregular rows of moderately long setae; terminal tergites setose; sternites 3-6 with very inconspicuous stubble. Relative lengths of antennal segments as follows: 38: 12: 12: 13: 12: 11: 11: 10: 9: 9: 15. Pronotum three-fifths broader than long. Abdomen more than twice as broad as elytra at scutellum; relative widths of fifth tergite and fifth paratergite 90: 26. 110 FIELDIANA: ZOOLOGY, VOLUME 40 TERMITELLA Wasmann Termitella Wasmann, 1911, Rev. Zool. Afr., 1: 170; 1912, Zeitschr. wiss. Zool., 101: 90; Fenyes, 1918, Gen. Insect., 173A: 65. Type species: Termitella lujae Wasmann. Termitissa Reichensperger, 1912, Ent. Mitteil., 11: 78. Type species: Termi- tissa foveolata Reichensperger. New synonymy. Termitoctesis Bernhauer, 1938, Ann. Mus. Stor. nat. Geneva, 60: 123. Type species: Termitoctesis gridelli Bernhauer. New synonymy. The Termitella generic group, comprised of this and the three following genera, have the abdomen as in the preceding genera except that the sixth and seventh tergites are distinctively modified (fig. 19, d). Termitella is easily distinguished from all other genera of the subtribe by its remarkable postgenal sulci (fig. 18, FIG. 25. (a) Thyreoxenus cucullalus Seevers; dorsal view, (b) Thyreoxenus cucullatus Seevers; lateral view, (c) Thyreoxenus parviceps Mann. 159 160 FIELDIANA: ZOOLOGY, VOLUME 40 segments 2-10 as in parviceps. Legs relatively long; anterior leg, 0.67 mm., middle leg, 0.85 mm., hind leg, 1 mm. Femora constricted slightly near middle; outer margin of hind femora somewhat arcuate proximally so that they appear somewhat bowed. Tibiae two-thirds as long as femora. Median non-setose areas of the upper surface of abdominal segments 4 and 5 narrowly margined on three sides by a dark band. Abdomen lateral to these areas rather heavily sclero- tized. Abdominal collar extremely developed, produced as a hood that conceals the terminal segments of the abdomen from above; the collar notched medially. Length, 2.5-3 mm. Thyreoxenus brevitibialis Seevers Thyreoxenus brevitibialis Seevers, 1946, Rev. Ent., 17: 259 (Villavicencio, Meta, Colombia; Nasutitermes costalis Holmgren; Chicago Natural His- tory Museum). Head mottled brown and yellow, postgenae very dark; pronotum and proster- num reddish-brown; elytra brown; femora and tibiae very dark reddish-brown; tarsi light brown; membranous areas of prothorax white, streaked with rust; abdominal sclerites brown; membranous areas of abdomen straw-colored. Eyes shallowly sulcate, weakly and irregularly pigmented. Two postgenal setae present. Antennal scape as long as segments 2-6 combined; segments 2-10 transverse, subequal. Legs relatively long; anterior leg 0.9 mm., middle leg 1.1 mm., hind leg 1.35 mm. Tibiae relatively short, only a little more than one-half as long as femora, at most three-fifths as long. Tibiae broadly spindle- shaped; hind tibiae one-third as broad as long. Abdomen rather heavily sclero- tized and strongly pigmented. The median non-setose areas of segments 4 and 5 margined along their anterior margins but for only one-half of the lateral margin; median area of segment 4 three times as long as broad; that of segment 5 almost twice as long as broad. Abdominal collar moderate in size, uniformly rather broad; its base not constricted. Length, 3 mm. Thyreoxenus albidus Seevers Thyreoxenus albidus Seevers, 1946, Rev. Ent., 17: 258 (Villavicencio, Meta, Colombia; Nasutitermes ephratae Holmgren; Chicago Natural History Museum). Head light brown, darker beneath; femora and tibiae dark brown; tarsi light brown; abdomen feebly sclerotized; membranous integuments of prothorax and abdomen milky-white, paler than usual. Eyes shallowly sulcate. Two postgenal setae present. Antennal scape shorter than segments 2-5 combined; segments 2-10 subequal. Legs moderately long; femora relatively short, the middle and hind tibiae three-fourths as long as femora. Hind femora not arcuate; not con- stricted near middle. Tibiae slender, scarcely spindle-shaped; hind tibiae only one-fifth as broad as long. Median non-setose areas of abdominal segments 4 and 5 narrowly margined by well-defined dark bands along three borders; median area of segment 4 two-fifths as broad as long. Abdominal collar low and very broad, obsolescent; broadly impressed medially. Length, 3 mm. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 161 Thyreoxenus convexinotus Seevers Thyreoxenus convexinotus Seevers, 1946, Rev. Ent., 17: 260 (Rurrenabaque, Beni, Bolivia; Nasutitermes chaquimayensis Holmgren; United States National Museum). Head light brown, postgenae darker, pronotum brown, elytra light brown, legs dark brown, tarsi pale, abdominal sclerites light smoky-brown, membranous integuments white to ivory. Eyes not sulcate. Two postgenal setae present. Antennal scape longer than segments 2-4 combined; segments 2-10 subquadrate, segment 3 a little longer than the others. Median ventral lobe of prothorax smaller than in the other species. Pronotum longitudinally convex, the crest of its convexity, a transverse, arcuate carina, attained about one-fourth the distance from base to apex; surface from crest to base strongly declivous; apical margin of pronotum almost straight; median incision relatively broad at level of transverse carina. Legs stout, moderately long; anterior leg 0.85 mm., middle leg 1 mm., hind leg 1.22 mm. Tibiae about two-thirds as long as femora. Tibiae broad, hind tibiae one-third as broad as long. Abdomen rather heavily sclerotized; median non-setose areas of segments 4 and 5 margined with dark bands along only their anterior borders (in a few cases for a very short distance along sides). Abdominal collar distinctive; very tall and slender; strongly constricted along its base. Length, 3 mm. Thyreoxenus major Mann. Figure 26. Thyreoxenus major Mann, Zoologica, 3: 332 (Kartabo, British Guiana; Nasutitermes guayanae Holmgren; United States National Museum); Emerson, 1935, Ann. Ent. Soc. Amer., 28: 369, text fig. 1; Seevers, 1937, Ann. Ent. Soc. Amer., 30: 3; Silvestri, 1946, Mem. Atti Accad. Naz. Lincei, (8), 1: 21, text fig. 12. Material examined. A series from Kartabo, British Guiana, with Nasutitermes guayanae Holmgren; and from Barro Colorado Island, Panama Canal Zone, with N. columbicus Holmgren. T. major differs from the other neotropical species in a number of charac- teristics: larger size; more robust build; postgenae with 3 macrosetae and finer setae; antennal segments 4-10 subquadrate; relative lengths of antennal segments: 22:4:7:5:5:5:5:5:5:5 :15; pronotum with arcuate apex, straight base, obtuse posterior angles, sides converging posteriorly; second tarsal segment not produced; tibiae and femora subequal in length; prosternum distinctive, produced posteriorly to form a large median triangular area not present in the other species; median non-setose areas of upper surface of abdominal segments 4 and 5 more heavily sclerotized and pigmented; abdominal collar obsolete. Length, 3-3.5 mm. (abdomen recurved). Remarks. The closest known relative of this series is solomon- ensis of the far distant Solomon Islands. 162 FIELDIANA: ZOOLOGY, VOLUME 40 Thyreoxenus solomonensis Seevers Thyreoxenus solomonensis Seevers, 1937, Ann. Ent. Soc. Amer., 30: 2, pi. 1, figs. 1, 2 (Auki, Malaita, Solomon Islands; Nasutitermes novarumhebri- darum N. and K. Holmgren; Chicago Natural History Museum). Head, thoracic sclerites, legs, except tarsi, dark brown to black; abdominal sclerites brown; elytra pale brown. Head with many short setae; postgenae with numerous long setae. Antennae similar to those of major; scape slightly longer than segments 2-4 combined; segments 2-10 subquadrate, segment 3 a little longer than the others. Prothorax three times as broad as head; sides straight, not impressed; pronotum with its anterior border strongly emarginate, the border within arcuate; anterior and posterior angles rounded; pronotum with two trans- verse rows of 4 erect setae on disk and scattered fine hairs. Legs short and stout; anterior femora with a distinct tooth on outer ventral margin near middle. Tarsi with basal segment twice as long as segment 2 or 3; segment 2 not produced at apex. Length, 2.8 mm. OIDEPROSOMA Silvestri Oideprosoma Silvestri, 1920, Boll. Lab. Zool. Portici, 14: 313; Kemner, 1925, Ark. Zool., ISA, no. 10, p. 17. Type species: Oideprosoma mirandum Silvestri. Oideprosoma is without doubt most closely related to Thyreoxenus, inasmuch as both genera have a remarkably hypertrophied pro- thorax. The genus is apparently represented in collections only by the somewhat damaged type in the Silvestri collection. Judging from Silvestri's figures, Oideprosoma differs from Thyreoxenus chiefly in the structure of its abdomen, but how much of this is due to the condition of the specimen cannot be determined now. Silvestri's illustrations show that the abdominal segments 3-5 are only moderately inflated and in a horizontal position, that seg- ment 6 is turned up, and that the terminal segments are directed cephalad. Abdominal segments 3-7 gradually increase in size, with the seventh segment the largest of the abdomen. The Oideprosoma abdomen seems to represent an intermediate stage between that of generalized subtribal stock and Thyreoxenus, but this interpretation is very tentative. The single specimen may not represent the definitive condition of the physogastric abdomen of Oideprosoma; it may be an intermediate form in post-imaginal development. Silvestri figured a larva which he believed to be that of Oidepro- soma, but this is open to serious question. The larva was not col- lected from the same termite nest as the Oideprosoma imago, but from a nest of the same termite species about five hundred miles from the type locality. The larva seems to resemble that of Para- \ FIG. 26. Thyreoxenus major Mann; four individuals from the same termite colony, illustrating post-imaginal development, (a) Stenogastric individual; winged, (b, c) Intermediate stages, (d) Physogastric individual. 163 164 FIELDIANA: ZOOLOGY, VOLUME 40 corotoca, figured by Warren, more than the larvae of Thyreoxenus that are in my collection. Oideprosoma mirandum Silvestri Oideprosoma mirandum Silvestri, 1920, Boll. Lab. Zool. Portici, 14: 315, text figs. 29-32 (Kakoulima, French Guinea; Eutermes trinervius Rambur; Silvestri coll.). The alleged larva of this species was collected at Thies, Senegal, French West Africa. The host 0. mirandum is Trinervitermes trinervius Rambur. Silvestri's description of this species was very limited. Tribe TERMITONANNINI Tribe Termitonannini: Fenyes, 1918, Gen. Insect., 173A: 17, 75. This tribe was proposed by Fenyes for the genera of Aleocharinae with 4, 4, 5-segmented tarsi and 10-segmented antennae. It happens that only termitophilous genera related to Termitonannus have since been added to the tribe, so that it is a useful category. At this time I propose to broaden the concept of the tribe by including the Perinthina, a group of termitophilous genera with 4, 4, 4-segmented tarsi and 11-segmented antennae. This group almost certainly originated from the same stock of termitophilous Aleocharinae as the Termitonannina. The phylogenetic relationships of the Termi- tonannini have been discussed in an earlier section of the paper (see p. 42). With the exception of one physogastric genus (Termilonilla), the species of the tribe are non-physogastric and in the majority of cases are limuloid. Their heads are either prognathous or hypog- nathous, the antennae 9-11-segmented, the terminal antennal segments bear one or more pairs of polytrichous coeloconic sensilla, the mentum is free except in certain derived genera, the mesocoxal cavities are margined, the hind coxae and metepimera are generalized except in certain derived genera, and the tarsi are either 4, 4, 4- segmented or 4, 4, 5-segmented. Subtribe PERINTHINA Tribe Hygronomini, subtribe Perinthi: Bernhauer and Scheerpeltz, 1926, Coleopt. Cat., 82: 521. The tribe Hygronomini has served merely as a convenient category into which all aleocharine genera with 4, 4, 4-segmented SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 165 tarsi might be placed, and should now be broken up to provide a more natural classification. Some of the variable characters of the Perinthina may be sum- marized as follows: Head prognathous in Termitopelta, Macro- gnathellus, and Termitonicus, and hypognathous in all other genera; a frontal cariniform line present in Perinthus; mentum and sub- mentum distinct except in Lauella, Termitonicus, Poduroides, and probably Macrognathellus; ligula bifid except in Perinthodes and Paraperinthus; gula broad in front (narrow in Perinthodes and Paraperinthus) ; apex of galea slender and acute (broad and ciliated in Termitopelta); antennal segments cylindrical (compressed in Perinthus and Lauella, and some segments compressed in Eutermi- tophila); galea and lacinia subequal in all genera except Macro- gnathellus and Poduroides, which have the galea elongated; pro- notum strongly convex and with deflexed sides except in Termito- pelta, Termitonicus, and probably Macrognathellus; pronotal base bisinuate except in Lauella and Termitonicus; hind coxae transverse except in Paraperinthus, which has triangular coxae; anterior coxae broad and convex in Perinthus, Termitocola, Termitopelta, and Macrognathellus, and elongated and narrow in the other genera. KEY TO GENERA OF PERINTHINA 1. Head prognathous; clypeus and labrum visible from above 2 Head hypognathous; only vertex and eyes visible from above 4 2. Pronotum cordiform; elytra very broad, their sides explanate. Termitonicus Mann Pronotum not cordiform; strongly transverse, about twice as broad as long; sides of elytra not explanate 3 3. Galea much longer than lacinia; body above densely, finely pubescent; pos- terior elytral angles not produced Macrognathellus Silvestri Galea and lacinia subequal in length; body above not pubescent, strongly shining; posterior elytral angles produced, acute Termitopelta Borgmeier 4. Elytral epipleurae visible; sides of elytra not deflexed against side of body. .5 Elytral epipleurae not visible; sides of elytra deflexed against side of body; Africa 9 5. Labial palpi 2-segmented (not verified); India Enter mitophila Cameron Labial palpi 3-segmented 6 6. Galea much longer than lacinia, its slender apex extending beyond the other mouth parts; antennae short and compact, segments 2-11 slightly telescoped, fusiform Poduroides Mann Galea not as above; antennae otherwise 7 7. Front of head with a complete transverse cariniform line. . . .Perinthus Casey Front of head without a cariniform line 8 8. Base of pronotum bisinuate; mentum and submentum distinct. Termitocola Seevers Base of pronptum almost straight; mentum and submentum fused, no suture evident; Pacific Islands Lauella Mann 166 FIELDIANA: ZOOLOGY, VOLUME 40 9. Pronotum and elytra robust, rufo-piceous, differing in color and texture from abdomen; pronotum and elytra without fine pubescence. Paraperinthus, new gen. Pronotum and elytra moderately large, testaceous, with the same texture and color as abdomen; pronotum and elytra with dense, fine pubescence. Perinthodes, new gen. TERMITOPELTA Borgmeier Termitopelta Borgmeier, 1950, Rev. Ent., 21: 656. Type species: Termitopelta fulgens Borgmeier. Termitopelta is noteworthy for its prognathous head; broad, almost horizontal pronotum (sides not deflexed); glabrous integu- ments that are neither densely reticulate nor pubescent; and large size. Head prognathous; occiput covered by pronotum; vertex, clypeus, and labrum visible from above; frontal cariniform line absent, clypeus not declivous. Antennae elongated; all segments, except tenth, longer than broad, segments 1-10 gradually decreasing in length; segments subcylindrical, verticillate. Man- dibles robust. Gula moderately wide throughout, its sutures subparallel. Sub- mentum and mentum distinct. Labial palpi 3-segmented, the cylindrical third segment longer than the others. Galea and lacinia subequal in length; lacinia with its spinose apex incurved, its medial border with numerous setae; galea densely and finely setose at apex, its apex not produced to a very acute point. Maxillary palpi 4-segmented; second segment compressed, strongly incrassate, third strongly incrassate. Pronotum strongly transverse, more than twice as wide as long; only feebly convex, its hyaline lateral margins explanate; apex evenly and moderately deeply bisinuate. Elytra very large, even wider than pronotum; its lateral margins arcuate, explanate; posterior angles produced, acute, its apical margins bisinuate. Epipleurae large, in a horizontal plane. Tarsi 4, 4, 4-segmented; terminal seg- ment subequal to the other segments combined. Anterior coxae convex, but less robust than in other genera. Termitopelta fulgens Borgmeier Termitopelta fulgens Borgmeier, 1950, Rev. Ent., 21: 657, text figs. 18-31 (Campinas, Goias; Syntermes molestus Burmeister; Borgmeier coll.). Material examined. One paratype. Rufescent throughout, strongly shining. Head, pronotum, and elytra in- distinctly and sparsely punctulate and finely pubescent. Head without macro- setae; pronotum (each half) with 11 marginal setae along anterior and lateral borders and with 2 or 3 setae on disk. Elytra with 8 marginal setae and one pair of long setae and several lesser ones on disk. Second tergite with one pair of setae; tergites 3-6 with 4 strong apical bristles, 3-5 with one pair and the sixth with 4 additional setae; tergite 7 without apical setae but with 4 sub-basal ones; tergite 8 with one seta on each lateral margin. Sternites with 6 long, black, pale-tipped setae on apical margins and 6 to 8 sub-basal setae; sternites with a moderate number of pale, recumbent hairs. Length, 3.5 mm. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 167 PERINTHUS Casey Perinthus Casey, 1890, Ann. N. Y. Acad. Sci., 5: 192; Wasmann, 1902, Boll. Mus. Zool. Torino, 17, no. 427, p. 1; Fenyes, 1918, Gen. Insect., 173A: 64; Seevers, 1937, Ann. Ent. Soc. Amer., 30: 10. Type species: Perinthus dudleyanus Casey. Head strongly deflexed, only a small part of its vertex and eyes visible. Head with a supra-antennal cariniform line extending across the vertex. Clypeus saddle-shaped. Entire body with a very fine-meshed reticulation and a dense vestiture of very fine pubescence. Antennae slightly geniculate, their scape short; segments 3-11 compressed, thin in profile; pedicels visible. Pronotum broad and convex, one-half broader than long; anterior border emarginate; apical angles broadly rounded; sides arcuate; pronotum widest behind middle; basal angles broadly rounded, base bisinuate. Elytra as wide as pronotum; upper surface only moderately convex; evenly so to the sharp marginal line delimiting the upper surface from the epipleuron. Epipleura strongly reflexed but almost in a hori- zontal plane, and easily seen in ventral view; epipleura form an acute angle with elytra. Remarks. With the exception of tarsatus, the species of Perinthus are very closely allied. Differences in pronotal and elytral chaeto- taxy (fig. 29) seem to be satisfactory for diagnostic purposes. Some of the differences are slight, but the setal patterns are apparently constant within a population. Species of Perinthus more often than not occur with several host species of Nasutitermes. The generic positions of P. silvestrii Wasmann and P. crassicornis Wasmann have long been uncertain, but having recently had an opportunity to examine the types of these species I am transferring the former to Termitocola and the latter to Macrognathellus. Perinthus dudleyanus Casey. Figures 27, a, 28, 29, a. Perinthus dudleyanus Casey, 1890, Ann. N. Y. Acad. Sci., 5: 194 (Panama; with termites; U. S. Nat. Mus.); Wasmann, 1902, Boll. Mus. Zool. Torino, 17, no. 427, p. 2; Seevers, 1937, Ann. Ent. Soc. Amer., 30: 10, pi. 3, fig. 19. I include in this species at present all individuals of the genus having the pronotal and elytral chaetotaxy shown in figure 29, a. Specimens from British Guiana, Trinidad, Colombia, Panama, and Ecuador are very similar except in size. In general, the smallest individuals occur in British Guiana and Trinidad at the eastern end of the range, intermediate ones in Colombia and Panama, and the largest in Ecuador. The three size groups are tentatively recognized as subspecies. Inasmuch as telescoping of abdominal segments makes an accurate measurement of body length almost impossible, the pro- notal dimensions were measured, as they seem to reflect the body 168 FIELDIANA: ZOOLOGY, VOLUME 40 size quite well. The results of my analysis are graphically illustrated in figure 28, and a summary of means follows: Pronotal width Pronotal length P. dudleyanus wasmanni Mann mm - mm. Trinidad (36) 0.707 0.443 British Guiana (27) 0.711 0.441 P. dudleyanus dudleyanus Casey Puerto Berrio, Colombia (9) 0.745 0.480 Panama (32) 0.790 0.503 Villavicencio, Colombia (4) 0.855 0.524 Santa Anna, Colombia (8) . 876 . 540 P. dudleyanus major, new subsp. Ecuador (16) 0.923 0.581 Perinthus wasmanni Mann was proposed as a full species, but this hardly seems justifiable, considering the overlap in size with dudleyanus. Considerably more material from various sections of the range of dudleyanus is required before the validity of the above hypothesis can be determined. Eight species of host termites are involved. In only one case is there a record of two subspecies with the same host: dudleyanus s. str. and wasmanni occur with Nasutitermes ephratae. This is due in part to the fact that some of the host species do not occur through- out the range of P. dudleyanus and in part to inadequate collecting. Perinthus dudleyanus dudleyanus Casey Perinthus dudleyanus Casey, 1890, Ann. N. Y. Acad. Sci., 5: 194 (Panama; with termites; United States National Museum). Material examined. Casey's types. Barro Colorado Island, Panama Canal Zone; with Nasutitermes ephratae Holmgren, N. corniger Motschulsky, and N. columbicus Holmgren. Puerto Berrio, Antioquia, Colombia, and Villavicencio, Meta, Colombia; with N. ephratae Holmgren. Santa Anna, Colombia; with N. meinerti Wasmann. Perinthus dudleyanus wasmanni Mann Perinthus wasmanni Mann, 1923, Zoologica, 3: 337 (Kartabo, British Guiana; Nasutitermes ephratae Holmgren; United States National Museum); Seevers, 1937, Ann. Ent. Soc. Amer., 30: 10. Material examined. Types and other specimens from Kartabo, British Guiana; with N. ephratae Holmgren and N. costalis Holm- gren. St. Joseph, Glyan Road, Arena Forest, St. Augustine, Rio Claro, and Northern Range, Trinidad, British West Indies; with N. ephratae Holmgren and N. costalis Holmgren. FIG. 27. (a) Perinthus dudleyanus Casey, (b) Perinthus tarsatus Mann. (c) Lauella palauensis Seevers. (d) Poduroides bovingi Mann, (e) Termitonicus mahout Mann. 169 170 FIELDIANA: ZOOLOGY, VOLUME 40 Perinthus dudleyanus major, new subspecies Distinguished from the other subspecies by the larger size of its individuals (see fig. 28 for pronotal dimensions). Holotype from Gualaquiza, Ecuador, collected October 29, 1934, by Wolfgang Hagen; with Nasutitermes dendrophilus Holmgren. In collection of Chicago Natural History Museum. Paratypes. Fifteen specimens, same data as holotype, except that some occurred with Nasutitermes peruanus Holmgren. Perinthus xenocostalis Seevers. Figure 29, e. Perinthus xenocostalis Seevers, 1937, Ann. Ent. Soc. Amer., 30: 9, pi. 3, fig. 18 (St. Joseph, Trinidad, British West Indies; Nasutitermes costalis Holm- gren; Chicago Natural History Museum). Material examined. A series from St. Joseph, Toco, Northern Range, and Manzanilla, Trinidad; with N. costalis Holmgren. Bacolet, Tobago; with N. costalis Holmgren. Kartabo, British Guiana; with N. costalis Holmgren. Huachi and Cachuela Esper- anza, Bolivia; with N. pilosus Snyder. Distinguished by its chaetotaxy (fig. 29, e): pronotum with marginal row of 8 (6 long, 2 short), its disk with 3 : 2 : 1; elytra with marginal row of 2, the disk with 1 : 1. Perinthus bolivari Seevers. Figure 29, f. Perinthus bolivari Seevers, 1946, Rev. Ent., 17: 264 (Huachi, Beni, Bolivia; Nasutitermes chaquimayensis Holmgren; United States National Museum). Material examined. Huachi, Beni, and Rurrenabaque, Beni, Bolivia; with N. chaquimayensis Holmgren. Rio Claro, Glyan Road, Mayaro, and Arena Forest, Trinidad, British West Indies; Bacolet, Tobago, British West Indies; all with N. ephratae Holmgren. Chavantina, Mato Grosso, Brazil (H. Sick); with N. bivalens Holmgren. Distinguished by its chaetotaxy (fig. 29, /) : pronotum with marginal row of 8 (6 long, 2 short), its disk with 3 : 2 : 2; elytra with marginal row of 2, the disk with 1 : 1. Perinthus mayae Seevers. Figure 29, c. Perinthus mayae Seevers, Rev. Ent., 17: 263 (Malon, Honduras; Nasutitermes corniger Motschulsky; United States National Museum). Material examined. Malon and Progresso, Honduras; Bobas, Guatemala; all with N. corniger Motschulsky. Distinguished by its chaetotaxy (fig. 29, c) : pronotum with marginal row of 9 (5 long, 4 short), its disk with 3:2:0; elytra with marginal row of 2, the disk with 1. Ecuador PERINTHUS DUDLEYANUS CASEY Santa Anna Colombia Villavicencio Colombia Barro Colorado Island Panama .91*5 .927 .909 .891 .873 .855 .837 .819 .801 .783 .765 .71*7 .729 .711 .693 .675 .657 .639 .396 .101* .1*32.1*50 .1*68 .1*86 .501* .522 .51*0 .558.576 .591* Pronotal width and length (fractions of ram.) FIG. 28. Perinthus dudleyanus Casey; variation in size of individuals (based on pronotal proportions) in different localities. Puerto Berrio Colombia British Guiana and Trinidad 171 172 FIELDIANA: ZOOLOGY, VOLUME 40 Perinthus guatemalae Seevers. Figure 29, b. Perinthus guatemalae Seevers, 1946, Rev. Ent., 17: 262 (Izabel, Guatemala; Nasutitermes ephratae Holmgren; Chicago Natural History Museum). Material examined. Type series. Distinguished by its chaetotaxy (fig. 29, 6): pronotum with marginal row of 7 (5 long, 2 short), its disk with 3 : 2 : 1; elytra with marginal row of 2, the disk with 1. Perinthus hageni Seevers. Figure 29, d. Perinthus hageni Seevers, 1937, Ann. Ent. Soc. Amer., 30: 9, pi. 3, figs. 15, 17 (Barro Colorado Island, Panama Canal Zone; Nasutitermes ephratae Holmgren; Chicago Natural History Museum). Material examined. A series from Barro Colorado Island, Panama Canal Zone; with N. ephratae Holmgren and N. corniger Motschulsky. Hamburg Farm, Costa Rica; with N. corniger Mot- schulsky. Hacienda de Tenguel and Bucay Chico, Ecuador; with N. peruanus Holmgren. Distinguished by its chaetotaxy (fig. 29, d) : pronotum with marginal row of 4, its disk with 3:1; elytra with marginal row of 2, the disk with 1. Perinthus tarsatus Mann. Figures 27, 6, 29, g. Perinthus tarsatus Mann, 1923, Zoologica, 3: 336 (Kartabo, British Guiana; Nasutitermes surinamensis Holmgren; United States National Museum). Material examined. A series of paratypes. Distinguished by the following characteristics: pronotum with a marginal row of 5 setae but without discal setae; elytra with 2 marginal setae and one on disk; antennal segments 3-7 slightly longer than broad, 8-10 subequal in length, 1-5 gradually broader, 5-11 subequal in width; pronotal sides more arcuate, pronotal base more strongly bisinuate; hind femora and tibiae subequal in length to pronotum (hind femora only four-fifths as long as pronotum in other species, and tibiae only three-fifths as long); hind tarsi three-fourths as long as tibiae (two-thirds as long as tibiae in other species). TERMITOCOLA Seevers Termitocola Seevers, 1937, Ann. Ent. Soc. Amer., 30: 7. Type species: Termitocola cylindricornis Seevers. Distinguished from Perinthus in these respects: Front of head without a transverse cariniform line; antennae elongated, segments not compressed; elytral epipleura less strongly reflexed, forming an obtuse angle with the upper surface. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 173 Termitocola cylindricornis Seevers. Figure 29, h. Termitocola cylindricornis Seevers, 1937, Ann. Ent. Soc. Amer., 30: 8, pi. 3, figs. 16, 20 (Barro Colorado Island, Panama Canal Zone; Nasutitcrmes pilifrons Holmgren; Chicago Natural History Museum). Color reddish-brown, abdomen slightly paler. Pronotum (fig. 29, h) with a marginal row of 8 setae, its disk with longitudinal rows of 3 : 3 : 2; elytra with FIG. 29. Chaetotaxy of pronotum and elytra, (a) Perinthus dudleyanus Casey, (b) Perinthus guatemalae Seevers. (c) Perinthus mayae Seevers. (d) Perinthus hageni Seevers. (e) Perinthus xenocostalis Seevers. (f) Perinthus bolivari Seevers. (g) Perinthus tarsatus Mann, (h) Termitocola cylindricornis Seevers. a marginal row of 2, the disk with rows of 1, 2 setae. Antennae very long, approxi- mately one-half as long as body, their segments slender, cylindrical (14 : 6 : 8 : 7 : 7 : 7 : 7 : 7 : 6 : 6 : 12); first segment the broadest, segments 2 and 3 slightly broader than segments 4-11, which decrease in width. Pronotal base distinctly bisinuate, moderately deeply incurved medial to each basal angle. Length, 2-2.2 mm. Termitocola silvestrii Wasmann, new comb. Perinthus silvestrii Wasmann, 1902, Boll. Mus. Zool. Torino, 17, no. 427, p. 1 (Urucum, Corumba, Mato Grosso, Brazil; Wasmann coll.; with termites); Silvestri, 1903, Redia, I: 198, pi. 6, figs. 267-272. Material examined. Two specimens on pin bearing type label in Wasmann collection. A series from Rio Pardo de Minas and Montes Claros, Minas Gerais, Brazil, collected January 8-10, 1952, 174 FIELDIANA: ZOOLOGY, VOLUME 40 by R. L. Araujo; with Constrictotermes cyphergaster Silvestri (deter- mined by R. L. Araujo). Coloration pale reddish-brown. Dorsum without setae, but with the usual very fine pubescence. Sternites with 6-8 setae in marginal row. Antennae cylindrical in form (10 : 6 : 6.5 : 6 : 7 : 5.5 : 5.5 : 5 : 5 : 5 : 11); segments 2-4 about two-thirds as broad as long, each tapering a little from apex to base; segment 5 longer than any of segments 2-10, subcylindrical; segments 5-10 increasing in width and decreasing in length (segment 5 about two-thirds as broad as 10). Pronotal base only feebly bisinuate, slightly incurved near basal angles. Length, 1.75-2 mm. Remarks. Wasmann's description of this species was short and vague, and Silvestri's comments and figures lacked important details. Two important clues to its identification were mentioned: the absence of pronotal and elytral macrosetae, and its occurrence with Constrictotermes. After examining Wasmann's specimens and an additional series from Brazil, I have decided to transfer this species to Termitocola. It may be desirable to erect a new genus for it eventually, for its presence in Termitocola does broaden that genus appreciably. MACROGNATHELLUS Silvestri Macrognathellus Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 20. Type species: Macrognathellus crassicornis Wasmann (= paraguayensis Silvestri). Silvestri separated this genus from Perinthus because of its elongated galea, which is longer than the lacinia. With the exception of P. tarsatus, which has a moderately elongated lacinia, the species of Perinthus have the galea and lacinia subequal. The pronotum of this genus is nearly twice as broad as long, about one-half broader than long in Perinthus. The elongated, subcylindrical antennae are unlike the compressed antennae of Perinthus. Silvestri's figure may give one the impression that the head of this genus is prognathous, but it is actually very similar to the condition in Perinthus. Macrognathellus crassicornis Wasmann, new comb. Perinthus crassicornis Wasmann, 1902, Boll. Mus. Zool. Torino, 17, no. 427, p. 1 (Paraguay, Heterotermes longiceps Snyder; Wasmann coll.). Macrognathellus paraguayensis Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 21, text fig. 9 (Sapucai, Paraguay; Nasutitermes sp.; Silvestri coll.). New synonymy. Silvestri's figures should be consulted for details of this species. The host recorded by Wasmann for the species is so inconsistent SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 175 with the host pattern for the Perinthina that it is almost certainly an error. Silvestri's record of Nasutitermes sp. as host of his para- guayensis should be regarded as correct until additional collections are made. PODUROIDES Mann Poduroides Mann, 1926, Proc. Ent. Soc. Wash., 28: 151. Type species: P. bovingi Mann. Poduroides has the appearance of a small Perinthus, but several features suggest closer relationship to Macrognathellus. The under side of the head and the mouth parts are apparently similar to the conditions in the latter genus; the elongated galea, the gula, the mentum, the submentum, the prementum and the labial palpi, and the unusual setose lines on the postgenae are especially suggestive. On the other hand, the head is probably more deflexed than in most genera of Perinthina and only the vertex is visible from above. Head hypognathous, mouth parts elongated, especially the galea, the pre- mentum, and the labial palpi. Postgenae (as seen in slide preparations) with an arcuate cariniform line almost paralleling the gular sutures; each of these lines bears about 35 long hairs. Gula not much narrower in front; submentum ex- panding rapidly in front of gula and becoming rather broad; mentum with sides converging rapidly for a short distance and then subparallel to apical margin. Mentum and submentum doubtfully distinct. Prementum very long; ligula bifid; labial palpi very long, basal segment twice as long as broad, second segment small, subquadrate, third more slender. Galea extending far beyond lacinia, its apex very slender and acute. Antennae geniculate, scape short; segments 3-11 cylin- drical, as a group compact and fusiform; segments slightly telescoped, their pedicels not visible. Pronotum about two-thirds as long as broad, the apical margin almost straight, the sides arcuate, the base strongly bisinuate. Poduroides bovingi Mann. Figure 27, d. Poduroides bovingi Mann, 1926, Proc. Ent. Soc. Wash., 28: 151, text figs. 1, 2 (Kartabo, British Guiana; Nasutitermes gaigei Emerson; United States National Museum). Material examined. Paratypes. Color testaceous throughout; finely pubescent and punctulate; pronotum with a marginal row of 5 setae, its disk with rows of 3, 2, 2, 1 setae; elytra with one or two marginal setae, disk with 1, 2, 2, 2 setae; tarsi all very short. TERMITONICUS Mann Termitonicus Mann, 1926, Proc. Ent. Soc. Wash., 28: 153; Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 22. Type species: Termitonicus mahout Mann. 176 FIELDIANA: ZOOLOGY, VOLUME 40 Termitonicus is a very distinctive genus and bears little super- ficial resemblance to the other Perinthina; nevertheless its character- istics definitely place it in this subtribe. The prognathous head, non-limuloid form, and distinctive pronotum and elytra indicate that Termitonicus differentiated from basic subtribe stock. Head prognathous, slightly broader than long. Antennae rather long, sub- cylindrical; intermediate segments elongated, more distal segments transverse; very feebly incrassate. Pronotum cordiform and distinctive; moderately convex but feebly explanate laterally; apex broadly and deeply emarginate, anterior angles broadly rounded, sides strongly arcuate and continuous with the oblique basal margin, base strongly produced medially. Elytra very broad; at base only as broad as pronotum, but the arcuate sides diverging strongly; elytra convex but becoming explanate laterally; epipleura very broad, in a horizontal plane. Anterior coxae long and slender; hind coxae transverse. Ninth tergite with a dense vestiture of unusual setae, the enlarged distal ends of which are heart- shaped. Termitonicus mahout Mann. Figure 27, e. Termitonicus mahout Mann, 1926, Proc. Ent. Soc. Wash., 28: 153 (Kartabo, British Guiana; Velocitermes beebei Emerson; United States National Museum); Emerson, 1949, Prin. Anim. Ecol., p. 720, text fig. 257. Material examined. A series from the type locality. Color flavo-testaceous, abdomen slightly darker. Pronotum with 4 setae on anterior margin, 5 on each lateral margin, and each half of disk with 2, 1, 1 setae. Elytra with 2 small marginal setae, the disk with 2 setae in a transverse row. Mentum with about 16 setae in irregular rows. Tergites with a subapical row of 4 setae. Relative lengths of antennal segments: 15 : 6 : 10 : 8 : 8 : 7 : 7 : 6 : 6 : 5 : 13; scape two and one-half times as long as segment 2; the latter about three-fifths as long as segment 3. Length, 2 mm. Termitonicus uroclaviger Silvestri Termitonicus uroclaviger Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 23, text fig. 12 (Jabaquara, Sao Paulo; Nasutitermes diversimilis Silvestri; Silvestri coll.). Record. Loreto, Misiones, Argentina (Silvestri, 1946b). This species is apparently very close to T. mahout Mann. Sil- vestri differentiated uroclaviger on the basis of its color and its having the second antennal segment a little shorter than the third. T. uroclaviger may be darker than mahout but the other character is a dubious one inasmuch as the second antennal segment of mahout is also shorter than the third. However, there are apparently two transverse rows of setae on the elytral disk of uroclaviger while there SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 177 is only one row in mahoui, and the former has only 4 setae on the mentum while mahout has about 16. LAUELLA Mann Lauella Mann, 1921, Psyche, 28: 54. Type species: Lauella vitiensis Mann. Closely related to Termilocola of the Neotropical Region, es- pecially in the structure of its head and mouth parts; distinguished from that genus by having the mentum fused to the submentum and some antennal segments compressed. The pronotal form differs from that of Termitocola, the anterior coxae are more slender and elongated, and the elytra are narrower and more convex. Pronotum very convex, sides more strongly deflexed than in Termitocola, apex only slightly emarginate, sides almost straight, base feebly arcuate but not bisinuate, posterior angles obsolete. Elytra strongly convex, only three-tenths as broad as long (three-fifths as broad as long in Termitocola); epipleura visible. KEY TO SPECIES OF LAUELLA 1. Head, pronotum, and elytra almost devoid of punctation, pubescence, and ground sculpture; only antennal segments 9-11 somewhat compressed. vitiensis Mann Head, pronotum, and elytra densely pubescent and punctulate; antennal segments 2-11 compressed 2 2. Pronotal disk with longitudinal rows of 2, 2, 3, 3 setae, beginning with median row (excluding setae of anterior and lateral margins); pronotum less than two-thirds broader than head palauensis, new sp. Pronotal disk with longitudinal rows of 2, 2, 2, 3 setae; pronotum nearly three-fourths broader than head javana, new sp. Lauella vitiensis Mann Laitella vitiensis Mann, 1921, Psyche, 28: 54 (Vunisea, Kadavu, Fiji Islands; Nasutitermes olidus Hill; United States National Museum). Hetairotermes leai Cameron, 1927, Rec. S. Australian Mus., 3: 269 (Taveuni, Fiji Islands; "with a termite;" South Australian Museum). New synonymy. Material examined. The holotype of L. vitiensis and a paratype of H. leai in the Cameron collection, British Museum (Natural History). In addition, a specimen from Suva, Viti Levu, Fiji Islands (Chicago Natural History Museum, K. P. Schmidt leg.), collected with N. olidus Hill. Head, pronotum, and elytra rufous, abdomen testaceous. Head, pronotum, and elytra with few traces of sculpture, punctation, or pubescence; abdomen minutely and densely punctulate and densely pubescent. Head with 18 or more short, erect hairs. Pronotum with 8 marginal setae and about 20 setae in 4 or 5 irregular rows on each half of disk. Elytra with a marginal row of 7 setae and disk with 18-20 in irregular longitudinal rows. Antennal scape stout, its upper 178 FIELDIANA: ZOOLOGY, VOLUME 40 surface shallowly concave; segments 2-11 slightly telescoped, pedicels invisible, segments 2-7 stout, cylindrical, 8-11 compressed, eleventh thin in profile; segments elongated except segment 2, which is three-fifths as long as third; segments 3-8 subequal in size, 9 and 10 a little shorter. Length, 2.75 mm. Lauella palauensis, new species. Figure 27, c. Distinguished from vitiensis by its smaller size, coloration, chaetotaxy, and antennal structure, and the presence of the punctu- lation and pubescence so characteristic of a majority of species of Perinthina. Pale brown; minutely, densely punctulate, and densely, finely pubescent. Head with a few short hairs; pronotum with 4 setae on each lateral margin, 6 on anterior margin, and with longitudinal rows of 2, 2, 3, 3 setae on each half of disk, beginning with median row; elytra with 2 marginal setae and 3 in a transverse row on disk. Antennal scape slightly concave above, broader than segments 2-11 ; segments 2-11 compressed, all except segment 2 elongated; segments slightly telescoped, pedicels not visible. Length, 1.8-2 mm. Holotype from Garakayo Island, Palau Islands, Micronesia; collected August 9, 1945, by Henry S. Dybas; with Nasulitermes brevirostris Oshima. In collection of Chicago Natural History Museum. Paratypes. Ten specimens, same data as holotype. Lauella javana, new species Closely allied to palauensis; differentiated by the characters of the key and by its broader, more robust build and paler coloration. Testaceous; densely punctulate and pubescent. Pronotum with 4 setae on lateral margins, 6 on anterior margin, and rows of 2, 2, 2, 3 setae on each half of disk, beginning with median row. Elytra with 2 setae on lateral margin and 3 in a transverse row on disk. Pronotum almost three-fourths broader than head (81 : 47); base with feeble sinuation. Antennae long and slender, scape broader and longer than other segments; segments 2-11 compressed, all except segment 2 longer than broad; segments 4-10 about two-thirds longer than broad. Length, 1.9 mm. Holotype from Preange, Tjigembong, Java; collected by J. B. Corporaal on September 28, 1915; with Nasutitermes corporaali Wasmann. In Wasmann collection, Natuurhistorisch Museum, Maastricht, The Netherlands. Paratype. One specimen, same data as type, in Chicago Natural History Museum. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 179 EUTERMITOPHILA Cameron Eutermitophila Cameron, 1939, Fauna Brit. India, Staphyl., 4: 41. Type species: Eutermitophila fletcheri Cameron. Eutermitophila has not been available, but there is no doubt that it is very closely related to Lauella. Cameron stated that Eutermi- tophila differs from Lauella in having 2-segmented labial palpi. Cameron's description mentions several characteristics of interest: Antennae with segments 8-11 slightly compressed, similar to those of Lauella vitiensis; anterior border of pronotum broadly emarginate, base bisinuate (base straight in Lauella) ; elytra strongly emarginate (not so in Lauella), anterior coxae almost as long as femora (longer in Lauella). Eutermitophila fletcheri Cameron Eutermitophila fletcheri Cameron, 1939, Fauna Brit. India, Staphyl., 4: 42, text fig. 6, pi. 1, fig. 3 (Coimbatore, Madras, India; Eutermes biformis Wasmann; Cameron coll.). Color ferrugineous; head, pronotum, and elytra extremely finely, closely punctulate and pubescent. Antennae with first segment a little longer than second; segment 3 slightly longer than 2; segments 3 and 4 equal, 5-7 decreasing, 8-10 subequal, slightly compressed. The host is Trinervitermes biformis Wasmann. PERINTHODES, new genus Type species: Perinthodes africanus, new species. Perinthodes resembles Perinthus, Termitocola, and Eutermitophila in appearance and seems to be most closely allied to the last-named genus. It differs from Eutermitophila in having the elytra strongly convex and closely appressed to the body, and their epipleura small and not visible, even in ventral view. The hind coxae are intermediate in form between those of the foregoing genera and Paraperinthus, new genus; these coxae are transverse, but the medial portion is somewhat produced so that the coxae are three- fourths as long as broad (one-half as long as broad in other genera). Body densely, finely punctulate and pubescent. Head hypognathous. An- tennal segments cylindrical, feebly telescoped, pedicels not visible; none of the segments compressed. Mentum separated from submentum by a very weak suture; gula narrower in front than in the foregoing genera; labial palpi 3-seg- mented, small, basal segment longer than broad, second segment short, third segment longer, cylindrical. Galea slender, the apex not acutely pointed. Pro- notum strongly convex, anterior margin almost straight, sides arcuate, base strongly bisinuate. Elytra strongly convex, sides closely appressed to body, epipleura small and not visible; posterior margin of elytra almost straight, not 180 FIELDIANA: ZOOLOGY, VOLUME 40 emarginate. Anterior coxae longer than femora, relatively slender. Hind coxae with thicker medial part somewhat produced, their maximum length about three-fourths the width; laterally, the coxae are nearly contiguous with the elongated metepimera. Perinthodes africanus, new species Head, pronotum, and elytra light brown, abdomen testaceous. Pronotum with 9 or 10 setae on each lateral margin, and 3 small setae on each half of the disk; elytra with 5 marginal setae and one pair of discal setae. Tergites 3-6 with an apical row of 6 fine setae, tergite 7 without marginal setae but with one sub- marginal seta on each side; tergite 8 with two rows of 4 setae each; tergite 9 with a cluster of long bristles. Antennal scape moderately broad, its upper surface concave; segments 2-11 cylindrical, segment 2 shorter than 3 but no less robust, segments 3-5 slightly elongated, 6-10 slightly transverse, decreasing a little in length. Length, 2.1-2.6 mm.; slender in build. Holotype from Camp Putnam, Epulu, Belgian Congo, collected May 15, 1948, by A. E. Emerson; with Nasutitermes latifrons Sjostedt. In collection of Chicago Natural History Museum. Paratypes. Seven, same data as holotype. Twelve, Camp Put- nam, Belgian Congo, collected May 16, 1948, by A. E. Emerson: with N. latifrons Sjostedt. Seven, Yangambi, Belgian Congo, collected May 29, 1948, by A. E. Emerson; with N. incurvus Sjostedt. PARAPERINTHUS, new genus Type species: Paraperinlhus pauciseta, new species. Most closely allied to Perinthodes, from which it is distinguished by the following characteristics: Pronotum and elytra distinctive in form, size, texture, color, punctulation, pubescence, and chaetotaxy; hind coxae triangular and metepimera short. The hind coxae and metepimera are unique within the Perinthina, and bear very close resemblance to those which distinguish the Corotocini. Head finely punctulate and minutely pubescent, but the pronotum, elytra, and abdomen with only faint traces of punctulation and pubescence. Upper surface of the body without setae and sternites with a few inconspicuous hairs. Head hypognathous. Pronotum two-thirds as long as broad, robust, somewhat coriaceous in texture, very shiny; disk strongly convex, moderately reflexed laterally so as to be feebly explanate; anterior margin almost straight, anterior angles evenly and broadly rounded, sides almost straight, basal angles rounded, base strongly bisinuate, its medial two-thirds more arcuately produced than usual. Elytra similar in robustness and texture to pronotum ; sides almost straight, apical angles almost rectangular, apical margins feebly oblique; epipleura not visible. Anterior coxae long and relatively slender, their upper surface convex; front femora five-sixths as long as coxae; front tibiae four-fifths as long as femora. Hind coxae with medial part somewhat produced, the apical margin strongly SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 181 oblique so that the general form is triangular; coxal length equal to width; laterally the hind coxae scarcely extend beyond the point of articulation. Metepimera short. Paraperinthus pauciseta, new species Head brown, pronotum and elytra rufo-piceous, abdomen testaceous. An- tennae with segments 2-11 robust, cylindrical; scape moderately broad, slightly concave above; segment 2 three-fourths as long as third, but not as broad; seg- ments 3 and 4 slightly longer than broad, 5-10 subquadrate to slightly transverse. Hind femora and tibiae equal in length; hind tarsi about two-thirds as long as tibiae. Length, 2.5-3.25 mm. Holotype from Yangambi, Belgian Congo, collected May 29, 1948, by A. E. Emerson; with Nasutitermes incurvus Sjostedt. In collection of Chicago Natural History Museum. Paratypes. Seven, same data as type, from two colonies of N. incurvus Sjostedt. One pupa was collected with this series. Subtribe TERMITONANNINA Tribe Termitonannini: Fenyes, 1918, Gen. Insect., 173A: 17, 75. Distinguished from the subtribe Perinthina by their 9- or 10- segmented antennae and 4, 4, 5-segmented tarsi. All genera except Termitonilla stenogastric. Derived stenogastric species usually limuloid in form. Head prognathous to hypognathous; clypeus and labrum visible from above in Nannellus, not in some species of Termilonannus. Head capsule generalized in Nannellus; short, transverse, and specialized in other genera. Antennae 10-segmented as a rule, 9-segmented in Eunannodes, moderately elongated in Nannellus to short and compact in Termitonannus; in many species, segments 4 and 5 extremely short. Antennae usually inserted under vertexal arcade; antennal fossae facing cephalad, shallow in Nannellus, deep in most genera. Gula and postgenae relatively short in species with hypognathous heads; mentum distinct in some species, fused to submentum in others. Labial palpi 3-segmented. Ligula incised except acutely produced in Macrolrichuris. NANNELLUS Silvestri Nannellus Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 17. Type species: Nannellus anoplotermitis Silvestri. This genus has not been available for study; the brief charac- terization is based on Silvestri's figures. Nannellus is apparently the most generalized known member of the subtribe. Body form generalized, slender, not limuloid. Head prognathous, clypeus and labrum visible from above; head transverse but not unusually short, only partly covered by pronotum; antennal fossae shallow, not beneath a vertexal 182 FIELDIANA: ZOOLOGY, VOLUME 40 arcade. Gula broad; mentum and submentum not separated by a suture in Silvestri's figure. Labial palpi small, slender. Antennae 10-segmented, relatively long; segments 1-3 elongated, 4 and 5 subquadrate, segment 5 a little larger than 4, segments 6-9 elongated, increasing in length. Pronotum relatively small, transverse, one-half broader than long; apex broadly emarginate. Nannellus anoplotermitis Silvestri Nannellus anoplotermitis Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 19, text fig. 10 (Lagoa Santa, Minas Gerais, Brazil; Anoplotermes sp.; Silvestri coll.). The structural details of this species were well illustrated by Silvestri. EUNANNODES Silvestri Eunannodes Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 15. Type species: Eunannodes reconditi Silvestri. Eunannodes is evidently most closely related to Nannellus, from which it was distinguished by Silvestri on the basis of its 9-segmented antennae. The head is short and transverse, more deeply set in the pronotum, and has the vertex more strongly deflexed than in Nannellus. Eunannodes reconditi Silvestri Eunannodes reconditi Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 16, text fig. 9 (Jabaquara, Sao Paulo, Brazil; Anoplo- termes sp.; Silvestri coll.). This species was well illustrated by Silvestri. TERMITONANNUS Wasmann Termitonannus Wasmann, 1902, Boll. Mus. Zool. Torino, (2), 17, no. 427, p. 2; Silvestri, 1903, Redia, 1: 199; Fenyes, 1918, Gen. Insect., 173A: 75; Seevers, 1941, Ann. Ent. Soc. Amer., 34: 341; Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 1. Type species: Termitonannus schmalzi Wasmann. Tetraphilus Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 2 (subgenus). Type species: Termitonannus (Tetraphilus) brachy- cerus Silvestri. Body form limuloid; head more or less hypognathous, to a degree that the anterior margin of the body is broadly arcuate. Head usually broad and short; vertex strongly deflexed in front, clypeus and labrum usually not visible from above. Antennae inserted under a vertexal arcade, usually about one-half of scape visible. Antennae 10-segmented, cylindrical in form, segments 4-9 trans- verse, 4 and 5 usually extremely short. Gula usually very short, mentum ap- parently fused to submentum in most species (distinct in vagans Silvestri). SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 183 Remarks. Only a few of the species of Termitonannus have been available for study, and thus a revision of the genus is not feasible at this time. Wasmann's species were inadequately described, and I am not confident that specimens at hand are all correctly determined. Silvestri's species were well described and figured and should be recognized as they become available. Species of Termitonannus are very small and slide preparations of material are almost essential for detailed studies. Silvestri's figures were evidently prepared after study of slide mounts, and satisfactory comparison of dry specimens is often not possible. The best diagnostic characters for species differentiation seem to be the following: chaetotaxy of pronotum, elytra, tergites, ster- nites, mentum-submentum, and legs; proportions of antennal seg- ments; head proportions; variations in galea and lacinia; form of eighth sternite and ninth tergite; size. Termitonannus schmalzi Wasmann Termitonannus schmalzi Wasmann, 1902, Boll. Mus. Zool. Torino, (2), 17, no. 427, p. 3 (Joinville, Santa Catarina, Brazil; Anoplotermes ater Hagen; Silvestri coll.); Silvestri, 1903, Redia, 1: 200, pi. 6, figs. 274-279. Material examined. One cotype, Joinville, Brazil (Bernhauer coll., CNHM). Fourteen specimens, Blumenau, Santa Catarina, Brazil (ex Reichensperger coll.), with Anoplotermes sp.; Rio Negro, Parana, Brazil (ex Reichensperger coll.), with Anoplotermes sp.; and Ilha Grande, Rio de Janeiro, Brazil (H. Sick), with Anoplotermes pacificus Fr. Miiller. Flavo-testaceous. Head with one seta behind each eye; pronotum with one seta near each apical angle and 3 very close to each lateral margin; without discal setae; elytra with 3 curved marginal setae and one on disk near humerus. Tergites sparsely setose, 3-6 with one curved bristle on each lateral margin, 5-6 with an additional pair on apical margin, tergite 7 with 4 short setae on apical margin. Sternites with one long seta on each lateral margin, sternites 5-7 with about 4 black setae on apical margin. Head twice as broad as long. Antennae with segments 6-9 transverse, more than twice as broad as long; segment 10 equal to segments 8 and 9 combined. Length, 0.8 mm. Termitonannus silvestrii Wasmann Termitonannus silvestrii Wasmann, 1902, Boll. Mus. Zool. Torino, (2), 17, no. 427, p. 3 (St. Catilinas, Buenos Aires, Argentina; Anoplotermes morio Hagen; Wasmann coll.); Silvestri, 1903, Redia, 1: 200, pi. 6, fig. 280. There is little information available about this species; it is a small species, about the same size as schmalzi, and is said to differ 184 FIELDIANA: ZOOLOGY, VOLUME 40 from that species in some details of antennal structure. According to Silvestri, antennal segments 8 and 9 are unusually large, and the terminal segment is twice as long as the ninth, but much narrower. The host is Anoplotermes meridianus Emerson. Termitonannus major Wasmann Termitonannus major Wasmann, 1902, Boll. Mus. Zool. Torino, (2), 17, no. 427, p. 3 (Tacurupucu, Paraguay; Anoplotermes pacificus F. Mtiller; Wasmann coll.); Silvestri, 1903, Redia, 1: 200, pi. 6, figs. 281-282; Hegh, 1922, Les Termites, p. 613, text fig. 418, a. This species was very poorly characterized by Wasmann. The dorsum was stated to be devoid of setae but it is doubtful if this can be taken literally. The antennal figure given by Silvestri is not very satisfactory and distinctive qualities are not evident. Length, 1.8 mm. Termitonannus vagans Silvestri. Figure 30, b. Termitonannus vagans Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 3, text fig. 1 (Jabaquara and Itapetininga, Sao Paulo, Brazil; Anoplotermes reconditus Silvestri; Silvestri coll.). Material examined. Specimens from Santo Amaro, Sao Paulo, Brazil; collected by J. F. Prado; probably with a species of Speculi- termes. Head not as short as that of most species of Termitonannus. Gula broad but not very short; mentum distinct; galea shorter than lacinia. Chaetotaxy dis- tinctive; pronotum with 5 setae on each lateral margin, 4 on anterior margin, and 2 on disk; elytra with 3 on lateral margin and 3 on disk. Eighth sternite produced at middle to an acute apex. Length, 2 mm. Silvestri's excellent figures should be consulted. The host is Speculitermes reconditus Silvestri. Termitonannus domunculi Silvestri Termitonannus domunculi Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 4, text fig. 2 (Saladillo, Santa Fe, Argentina; Corni- termes striatus Hagen; Silvestri coll.). Pronotum with 4 short setae on each lateral margin; elytra with 3 lateral setae and 2 short setae near apical margins. Antennae very similar to those of vagans. Galea and lacinia subequal. Mentum apparently fused to submentum. Eighth sternite not as in vagans. Length, 1.95 mm. FIG. 30. (a) Macrotrichurus brasiliensis Silvestri. (b) Termitonannus vagam Silvestri. (c) Termilonilla luteola Borgmeier. 185 186 FIELDIANA: ZOOLOGY, VOLUME 40 Silvestri's figures must be examined for diagnostic characters. The host is Procornitermes striatus Hagen. Termitonannus proximatus Silvestri Termitonannus proximatus Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 6, text fig. 3 (Lagoa Santa, Minas Gerais, Brazil; Anoplotermes sp.; Silvestri coll.). Pronotum with 5 short setae on each lateral margin; elytra with 3 lateral setae and 2 setae near apical margins. Antennae very similar to those of vagans. Head short, gula short and broad; mentum fused to submentum. Chaetotaxy of abdominal segments distinctive. Length, 1.6 mm. Silvestri's figures must be examined for diagnostic characters. Termitonannus microsomatis Silvestri Termitonannus microsomatis Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 7, text fig. 4 (Cagapava, Sao Paulo, Brazil; Nasutitermes microsoma Silvestri, var.; Silvestri coll.). Pronotum with 5 setae on each lateral margin; elytra with 3 moderately long lateral setae and many shorter ones, and 2 short setae on apical margins. Head short, gula short and broad, mentum fused to submentum. Abdominal chaetotaxy distinctive. Antennae similar to those of proximatus but with segments 7-9 shorter. Length, 1.4 mm. Silvestri's figures must be consulted for diagnostic features. The host is Subulitermes microsoma Silvestri. Termitonannus parvulus Silvestri Termitonannus parvulus Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 9, text fig. 5 (Alto da Serra, Sao Paulo, Brazil; Anoplotermes ? minimus; Silvestri coll.). Pronotum with 5 setae on each lateral margin and 8 discal setae; elytra with 3 long lateral setae and numerous short setae and 2 sub-basal setae on disk. Antennae rather similar to those of microsomatis. Abdominal chaetotaxy dis- tinctive. Length, 1.5 mm. Silvestri's figures must be consulted for diagnostic characters. The host is apparently a new species of Anoplotermes. Termitonannus brachycerus Silvestri Termitonannus (Tetraphilus) brachycerus Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 10, text fig. 6 (Jabaquara, Sao Paulo, Brazil; Anoplotermes sp.; Silvestri coll.). SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 187 Silvestri placed this species in the subgenus Tetraphilus because of the presence of two pairs of polytrichous sensory foveae on the terminal antennal segment (in contrast to the usual pair). Pronotum with 5 short setae on each lateral margin; elytra apparently without setae. Antennae short, segments 3-9 very short. Abdominal chaetotaxy dis- tinctive. Length, 1.4 mm. Silvestri's figures should be consulted for diagnostic characters. Termitonannus echinoides Seevers Termitonannus echinoides Seevers, 1946, Rev. Ent., 17: 264 (Ivon, Beni, Bolivia; Anoplotermes sp.; United States National Museum). Color testaceous. Pronotum with 28-30 long, stout, yellow-tipped bristles, about one-half as long as pronotum. Elytra with about 22 irregularly dispersed setae. Tergites 3-6 with a row of 8 very long bristles and numerous short pale hairs; tergite 8 with 6 bristles. Sternites 3-7 with a subapical row of about 16 long, slender setae and numerous pale hairs in addition. Lateral plates of ninth abdominal segment with numerous long, curving bristles. Antennae with seg- ments 1 and 2 short, elongated; segment 3 small, segments 4 and 5 very short, transverse, 6-9 transverse. Length, 1.2 mm. Termitonannus setosus Seevers Termitonannus setosus Seevers, 1941, Ann. Ent. Soc. Amer., 34: 344, pi. 3, figs. 42, 43 (Barro Colorado Island, Panama Canal Zone; Anoplotermes, new sp. [Emerson cat. no. 152b]; Chicago Natural History Museum). Color testaceous, polished. Pronotum with one long seta near each apical angle and 3 on each lateral margin; elytra with 3 conspicuous setae on lateral margins, one small seta on posterior margin, and one on disk. Dorsum of abdomen with 4 longitudinal rows of long, stout bristles. Head very short and broad; gula and mentum-submentum short. Galea slender, longer than lacinia and with a few fine hairs. Antennae with segments 1 and 2 short and stout, segment 3 small, obtrapezoidal, segments 4 and 5 extremely short, transverse, 6-9 transverse, moderately long, segment 10 about as long as segments 7-9. Length, 1.2 mm. Termitonannus gatuni Seevers Termitonannus gatuni Seevers, 1941, Ann. Ent. Soc. Amer., 34: 345 (Barro Colorado Island, Panama Canal Zone; Anoplotermes, new sp. [Emerson cat. no. 152b]; Chicago Natural History Museum). Color flavo-testaceous. Pronotum with 3 very feeble setae on each lateral margin; elytra with one seta at outer apical angles. Tergite 6 with one small seta on lateral margins; tergite 7 with an apical row of 6 small setae; tergite 8 with two transverse rows of 4 setae. Sternites 3-8 with one stout seta at outer apical angles and 4 fine setae on apical margin. Antennae as in selosus. Length, 1 mm. 188 FIELDIANA: ZOOLOGY, VOLUME 40 TERMITOCOMES Seevers Termitocomes Seevers, 1941, Ann. Ent. Soc. Amer., 34: 342. Type species: Termitocomes wasmanni Seevers. Distinguished from Termitonannus by these characters: Antennal segments 6-10 strongly compressed; segment 4 of maxillary palpi distinctive, large, conoidal in form; ligula acute, not bifid; galea unusually long. Head short, transverse, a small area of occiput covered by pronotum. Vertex strongly deflexed, forming a rounded frontal arcade. Antennae inserted in broad, deep fossae; basal half of scape concealed. Antennae with first and second seg- ments moderately long; segment 3 small, conical; segments 4 and 5 very short, transverse; 6-9 longer than broad; segment 10 about as long as segments 8 and 9; segments 6-10 strongly compressed. Gula large, transverse, submentum short, strongly transverse; mentum trapezoidal. Ligula long, slender, not incised. Galea somewhat longer than lacinia, apex slender and finely ciliated. Maxillary palpi with segment 4 large, conoidal. Pronotum large, convex, its anterior margin feebly emarginate, its sides diverging, nearly twice as broad at base as at apex; base bisinuate. Abdomen conical, tapering strongly at apex. Ninth tergite of male divided to form two long, slender, styliform processes. Termitocomes wasmanni Seevers Termitocomes wasmanni Seevers, 1941, Ann. Ent. Soc. Amer., 34: 344, pi. 3, figs. 40, 41 (Kartabo, British Guiana; Speculitermes silvestrii Emerson; Chicago Natural History Museum). Color testaceous; shining. Pronotum with 6 setae on apical margin, 4 on each lateral margin and 10 on disk (8 in a transverse row at middle, and one sub-basal pair). Elytra with one basal and one apical seta on disk and 2 long setae and a number of fine hairs on lateral margins. Tergite 3 with 4 dark setae; tergites 4-6 with 6 setae in subapical row; tergite 7 with a row of 6 setae near middle; tergite 8 with a basal row of 4 bristles and a subapical row of 4. Sternites 4 and 5 with subapical rows of 8 setae; sternite 6 with 6 setae; sternite 7 with 4 setae; sternite 8 with a medial transverse row of 4 setae. Tergites and sternites densely clothed with pale yellow hairs, those of sternites rather long. Surface of abdomen reticulated, appearing finely rugulose. Length, 2.6 mm. Remarks. The original description of the pronotal chaetotaxy was incorrect; some of the setae were missing from the holotype. Termitocomes trinidadensis, new species Distinguished from wasmanni by pronotal, elytral, and abdominal chaetotaxy. Color rufo-testaceous. Head, pronotum, and elytra smooth and shining, not punctulate or pubescent. Abdomen not reticulated or rugulose. Pronotum with 4 setae on lateral margins, 4 on anterior margin, and 8 on disk (6 in transverse SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 189 row about middle, 2 sub-basal). Elytra with 3 setae on lateral margins, 2 basal setae, and 2 apical setae on disk. Tergites with strong, black, pale-tipped setae as follows: 3-7 with five rows of 4, 6, 6, 6, 6 setae, respectively; tergite 8 with two rows of 4 setae each. Tergites rather densely clothed with pale, recumbent hairs. Sternites densely clothed with fine, yellow hairs, many of which are moderately long; sternites with transverse row of about 4 dark setae. Antennae very similar to those of wasmanni; relative lengths of segments: 8 : 7 : 6 : 2 : 3 : 7 : 8 : 8 : 9 : 18. Antennae strongly compressed, segment 3 obconical, segments 4 and 5 more than twice as broad as long, segment 6 a little broader than long, segments 7-9 sub- quadrate, segment 10 more slender than 9. Length, 2.5 mm. Holotype from Arena Forest, Trinidad, British West Indies; collected February 22, 1945, by S. A. Omenai and F. 0. Iwenjora (A. M. Adamson coll.); with Labiotermes labralis Holmgren, new subspecies. In collection of Chicago Natural History Museum. MACROTRICHURUS Silvestri Macrotrichurus Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 11. Type species: Macrotrichurus brasiliensis Silvestri. Macrotrichurus apparently differs from Termitonannus in having an unusually long prementum; an acute ligula; and stout labial palpi. Other differences include the distinctive galea and lacinia; an elongated, more generalized head; an emarginate mentum; the cylindrical ninth tergite, which is not deeply divided as in Termi- tonannus; and the vestiture of the ninth tergite. Antennae 10-segmented; segments 4-8 transverse; 4 and 5 very short; segment 9 elongated. Head prognathous, slightly transverse but not very short, vertex declivous in front, antennae moderately deeply inserted under vertexal arcade. Gula short, broad, mentum fused to submentum; mentum deeply emarginate in front. Ligula acute. Ninth tergite distinctive, its lateral margins deflexed; thus the sclerite is somewhat cylindrical, narrow, and stout in dorsal view. Macrotrichurus brasiliensis Silvestri. Figure 30, a. Macrotrichurus brasiliensis Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 13, text fig. 7 (Jabaquara, Sao Paulo; Anoplotermes pacificus Miiller, var.; Silvestri coll.). Material examined. A series from Sao Paulo, Brazil; with Corni- termes cumulans Kollar (R. L. Araujo); and Morumbi, Sao Paulo, Brazil; with undetermined host. Head with 2 long vertexal setae and 2 behind each eye; pronotum with 2 long setae and several short ones on each lateral margin, disk with three longitudinal rows of 3 setae on each half; elytra with 5 or 6 long setae and numerous shorter ones on lateral margins or submarginal, disk with three longitudinal rows of 3, 2, 2 setae. 190 FIELDIANA: ZOOLOGY, VOLUME 40 Length, 2.2 mm. Silvestri's figures should be consulted. Macrotrichuris notabilis Silvestri Macrotrichuris notabilis Silvestri, 1946, Rend. Accad. Sci. Fis. Math. Soc. Reale Napoli, (4), 14: 14, text fig. 8 (Tabalao, Brazil; Anoplotermes ? pacificus Miiller; Silvestri coll.). Distinguished from brasiliensis by chaetotaxy of pronotum, elytra, and abdomen, and by proportions of antennal segments. Silvestri's figures must be consulted for diagnostic details. TERMITONILLA Borgmeier Termitonilla Borgmeier, 1950, Rev. Ent., 21 : 651. Type species: Termitonilla luteola Borgmeier. Due to its physogastry this is perhaps the most noteworthy genus of the subtribe. Superficially, Termitonilla bears a strong resemblance to Eburniola of the Corotocini, but this is without doubt due to convergence. Head, pronotum, and elytra generalized in appearance. Head prognathous, vertex and clypeus moderately declivous from eye level; opening of antennal fossae cephalad, but antennae not inserted under a vertexal arcade. Head trans- verse, occiput covered by pronotum. Antennae 10-segmented, segments 1 and 2 moderately long, segment 3 short, obtrapezoidal, segments 4 and 5 very short, transverse, 6-9 transverse. Gula moderately broad, its sides subparallel; mentum distinct; labial palpi small, ligula incised, galea and lacinia subequal. Pronotum large, convex, anterior margin straight, sides and base arcuate, posterior angles almost obsolete; pronotal hypomera relatively small, visible from side. Abdomen very large, strongly physogastric, almost all sclerites except those of terminal segments isolated by non-sclerotized integument. Tergites, sternites, and para- tergites generalized. Termitonilla luteola Borgmeier. Figure 30, c. Termitonilla luteola Borgmeier, 1950, Rev. Ent., 21: 652, text figs. 1-8 (Cam- pinas, Goias, Brazil; Anoplotermes sp.; Borgmeier coll.). Material examined. One paratype, same data as holotype. The hosts have been determined as Anoplotermes ? ater Hagen. Head, pronotum, and elytra reddish-yellow; under side of thorax and ab- dominal sclerites pale yellow, somewhat hyaline; terminal abdominal segments darker. Head with an irregular transverse row of 5 or 6 bristles between eyes, and a pair of occipital bristles. Pronotum with 8 setae on anterior margin, 3 on lateral margins, and about 10 on each half of disk. Elytra with 4 lateral bristles and about 4 irregular rows on disk. Tergite 3 with 4 apical bristles, tergites 4-6 with 6 apical bristles; sternites 4-6 with 12 basal setae and 14-16 on apical margins. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 191 Tribe TERMITOHOSPITINI Tribe Termitohospitini: Seevers, 1941, Ann. Ent. Soc. Amer., 34: 331. This tribe was proposed to separate a small group of neotropical termitophiles from the large heterogeneous tribe Bolitocharini. These species were characterized by limuloid body-form, large pronota that cover or almost cover the deflexed head, very large, deep antennal fossae, distinctive mouth parts, and 4, 4, 5-segmented tarsi. Study of termitophilous Aleocharinae with the same tarsal for- mula from other zoogeographic regions reveals that most of them belong to this tribe. Four subtribes are proposed: the Termito- hospitina of the Neotropical Region, the Hetairotermitina of the Oriental and Australian Regions, the pantropical Termitospectrina, and the Ethiopian Termitusina. The first three of these subtribes occur with the subfamily Nasutitermitinae, but the Termitusina are guests of Cubitermes of the Termitinae. The phylogenetic and evolutionary interrelationships with the host termites are discussed elsewhere (p. 48). Tribal characteristics: Tarsi 4, 4, 5-segmented. Antennae 11 -segmented; without polytrichous foveae on terminal segment. Mentum distinct; prementum elongated, ligula slender, nearly filiform; labial palpi 2- or 3-segmented, very long and slender, segments filiform, basal segment often greatly elongated. Galea and lacinia very long and slender. Antennal fossae large and usually deep; antennae inserted under a vertexal arcade. Anterior tentorial pits located at medial, inferior angles of antennal fossae. Middle coxae contiguous, mesosternal process very short, metasternum not produced between coxae; mesocoxal aceta- bula very large; metasternum behind them very short. Abdomen not physo- gastric. Subtribe Termitohospitina: Body-form limuloid. Head strongly deflexed; concealed by large, shield-like pronotum except in Paratermitosocius (in which a small part of the vertex is visible). Pronotum with anterior and lateral margins continuously arcuate; surface not impressed. Anterior margin of submentum straight or slightly arcuate but not emarginate; anterior angles of mentum pro- duced, apex of mentum emarginate. Abdomen slender, acuminate. Ninth tergite bifid, but otherwise unmodified. Subtribe Hetairotermitina: Body-form more or less limuloid. Head slightly deflexed; covered to the eyes by pronotum. Pronotum large, transverse; anterior border almost straight, not continuously arcuate, with lateral margins. Anterior margin of submentum arcuately emarginate. Anterior angles of mentum pro- duced, apex emarginate. Abdomen slender, acuminate. Subtribe Termitospectrina: Body-form not limuloid. Head scarcely deflexed, almost prognathous. Pronotum distinctive (fig. 31); its surface with strong sulci. Anterior angles of mentum not produced, apex straight. Abdomen not acuminate, recurved. 192 FIELDIANA: ZOOLOGY, VOLUME 40 Subtribe Termitusina: Body-form limuloid. Head strongly deflexed, usually with the long axis vertical; a small part of vertex visible from above. Pronotum large, shield-like; its sides strongly arcuate, converging anteriorly but anterior margin transverse, anterior angles evident. Anterior margin of submentum emarginate. Anterior angles of mentum not produced, apex straight. Abdomen acuminate. Ninth tergite deeply bifid, the two parts elongated as stout processes. KEY TO NEOTROPICAL GENERA OF TERMITOHOSPITINI 1. Pronotum with two deep longitudinal sulci on disk; pronotal apex very deeply emarginate Termitospectrum Mann Pronotum not as above 2 2. Pronotum completely covering head 3 Pronotum truncate in front, a small area of deflexed head visible from above. Paratermitosocius Seevers 3. Antennae elongated; segments longer than broad or subquadrate but never transverse; segments 3-10 not incrassate 4 Antennae shorter; some or all of the segments transverse, segments 3-10 to some degree incrassate 5 4. Antennae very slender; all segments elongated, subcylindrical; abdominal bristles extremely long and conspicuous Blapticoxenus Mann Antennae more robust; segments 4-10 subquadrate to feebly elongated; abdominal bristles moderately long Termitosodalis Seevers 5. Mouth parts extremely elongated, galea very long and slender, lacinia strongly sclerotized, spinose; labial palpi extremely long and thin, second palpomere seven times as long as broad; third antennal segment longer than broad Termitosocius Seevers Mouth parts moderately long, galea and lacinia not unusually long; labial palpi moderately long, second palpomere only slightly longer than broad; third antennal segment usually transverse, at most subquadrate. Termitohospes Seevers Subtribe TERMITOHOSPITINA TERMITOHOSPES Seevers Termitohospes Seevers, 1941, Ann. Ent. Soc. Amer., 34: 333. Type species: Termitohospes miricorniger Seevers. Head deflexed, completely covered by pronotum. Head short, transverse; labrum, maxillae, and labium not greatly elongated. Antennal segments 3-10 usually transverse, in some species a few segments subquadrate; segments 3-10 incrassate. Pronotum very large, its lateral and anterior margins continuously arcuate; pronotum from one-tenth to seven-tenths broader than long. KEY TO THE SPECIES OF TERMITOHOSPES 1. Pronotum approximately two-thirds broader than long limulus, new sp. Pronotum not more than one-half broader than long and usually somewhat less 2 2. Smaller species, 1.25-1.5 mm. in length 3 Larger species, 1.8-2.25 mm. in length 5 SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 193 3. Third antennal segment very short, transverse, produced medially and bearing a long bristle miricorniger Seevers Third antennal segment subquadrate, not produced medially 4 4. Antennal scape about as long as segments 2-5 combined. panamensis Seevers Antennal scape about as long as segments 2 and 3 combined. guianae Seevers 5. Pronotal base almost straight, basal angles rounded; pronotum one-tenth to one-fourth broader than long 6 Pronotal base emarginate or sinuate; pronotum two-fifths to one-half broader than long 7 6. Pronotum one-tenth to one-seventh broader than long. tacky poroides, new sp. Pronotum one-fifth to one-fourth broader than long silvestrii, new sp. 7. Antennal segments 3-5 very short, about one-fourth as long as second; tergites 3-6 with 4 erect marginal setae brasiliana, new sp. Antennal segments 3-5 about one-half as long as second; tergites 3-6 with one seta near each apical angle unicolor Silvestri Termitohospes miricorniger Seevers Termitohospes miricorniger Seevers, 1941, Ann. Ent. Soc. Amer., 34: 335, pi. 3, figs. 33, 37, 44, 45 (Kartabo, British Guiana; Nasutitermes ephratae Holmgren; Chicago Natural History Museum). Material examined. Kartabo, British Guiana, with Nasuti- termes ephratae Holmgren and N. costalis Holmgren. Manzanilla, Mayaro, Mt. Tabor, and Northern Range, Trinidad, British West Indies (ex Adamson coll.); with N. ephratae Holmgren. Manzanilla and Northern Range, Trinidad, with N. costalis Holmgren. Testaceous; strongly shining. Front of head between antennal fossae at narrowest point one-sixth as broad as head. Antennal scape ovoidal; segment 2 obconical; segment 3 very short and broad, its medial apical surface produced and bearing a long seta; segment 4 very short; segments 4-10 transverse, increasing in length and slightly incrassate; segment 11 as long as segments 8-10 combined. Head with a sparse vestiture of fine hairs which are longer on front of head. Pronotum and elytra minutely punctulate and pubescent. Pronotum with 2 short setae on lateral margins. Tergites 3-8 sparsely pilose and with a marginal row of pale setae. Sternites 3-6 with a pair of setae on apical margin near each side and 1 medial seta; eighth sternite with 8 setae (2 on apical margin, 4 in subapical row, and 1 on each lateral margin); all sternites densely clothed with recumbent hairs. Length, 1.25-1.5 mm. Termitohospes brasiliana, new species Related to miricorniger, from which it differs in larger size, antennal structure, chaetotaxy, and slightly broader pronotum. 194 FIELDIANA: ZOOLOGY, VOLUME 40 Pilosity of head as in miricorniger. Pronotal margin with a continuous row of about 16 short setae; elytral margin with 3 setae. Tergites 3-6 with 4 marginal setae; tergite 7 not setose; tergite 8 with two pairs of setae. Sternites 3-7 with 6 erect setae on apical margins; sternite 8 with a prominent subapical pair of setae and 6 on arcuate apical margin. Sternites with a moderately dense covering of fine, pale, recumbent hairs, becoming shorter and finer on more posterior segments. Antennal segments with the following relative lengths: 12 : 4 : 1 : 1 : 1.5 : 2 : 2 : 2 : 2 : 2.5 : 7. Antennae with segments 3-10 short, transverse; segment 3 very short, but not produced medially as in miricorniger; segments 4-10 gradually incrassate; segment 10 twice as long and broad as segments 3 or 4. Pronotum two-fifths broader than long. Length, 1.9 mm.; width, 0.85 mm. Holotype and one paratype from Blumenau, Santa Catarina, Brazil (ex Reichensperger coll.) ; with Nasutitermes aquilinus Holm- gren. In collection of Chicago Natural History Museum. Termitohospes guianae Seevers Termitohospes guianae Seevers, 1941, Ann. Ent. Soc. Amer., 34: 335, pi. 3, fig. 34 (Kartabo, British Guiana; Nasutitermes surinamensis Holmgren; Chicago Natural History Museum). Distinguished from miricorniger by having antennal segments 3-7 subquadrate, not as short or as broad; segments 8-10 slightly transverse, but not as short, and segment 3 not produced medially. Termitohospes panamensis Seevers Termitohospes panamensis Seevers, 1941, Ann. Ent. Soc. Amer., 34: 336 (Barro Colorado Island, Panama Canal Zone; Nasutitermes columbicus Holmgren; Chicago Natural History Museum). Closely related to guianae from which it is distinguished by slight antennal differences and distinctive chaetotaxy: Antennal scape longer and more slender, almost equal to segments 2-5 combined (in guianae equal to segments 2 and 3); pronotum with 2 setae on anterior margin and 5 on lateral margins (middle 3 setae much longer) ; elytra with 3 strong setae on lateral margins; tergites 3-6 with 4 setae on apical margin; tergite 7 without setae; tergite 8 with 2 small apical and 2 subapical setae; sternites 3-7 with 6 apical setae (middle pair weak except on sternite 7). Termitohospes unicolor Silvestri, new comb. Figure 31, a. Termitomus unicolor Silvestri, 1947, Arch. Zool. Ital., 31 : 142, fig. 8 (Jabaquara, Sao Paulo, Brazil; Cornitermes similis Silvestri; Silvestri coll.). FIG. 31. (a) Termitohospes unicolor Silvestri. (b) Termitoecia fabulosa Bernhauer; the holotype without elytra, (c) Termitoecia wilsoni Cameron. (d) Termitana perrieri Fairmaire. (e, f) Termitospectrum thoracicum Mann; dor- sal and lateral views. d 195 196 FIELDIANA: ZOOLOGY, VOLUME 40 Material examined. Thirty-four specimens from 10 colonies of Cornitermes cumulans Kollar (determined by R. Araujo) from localities in the vicinity of Sao Paulo, Brazil (Ibirapue'ra, Morumbi, Eldorado, Interlagos, Sacoman, Congonhas, Vila Formosa); col- lected by R. L. Araujo, J. F. Prado, and W. A. Maluf. Silvestri's Cornitermes similis is the same as C. cumulans Kollar. Rufo-castaneous to rufo-testaceous. Head with about 5 medially directed pale setae along inner margins of antennal fossae, vertex finely setose and pubescent except between antennal fossae. Pronotum microscopically punctulate and pubescent; anterior and lateral margins bearing 25-30 short, fine hairs. Elytral margins with 2 moderately long, black setae. Tergites 3-6 with a long seta near each apical angle. Chaetotaxy of terminal abdominal sclerites figured by Silvestri (1947). Front of head between antennal fossae at narrowest point almost one- third as broad as head. Antennal segments with the following relative lengths: 18 : 6 : 3 : 2 : 2.5 : 3 : 3.5 : 3.5 : 4 : 4.5 : 10. Antennal segments 4-10 trans- verse, incrassate, segment 10 twice as broad as segment 4. Pronotum about one-half broader than long. Length, 2-2.2 mm. Termitohospes limulus, new species Distinguished from other species of the genus by the different body proportions: limulus is short and very broad; the pronotum is one-third broader than long; the abdomen is broad at base but tapers rapidly to a narrow apex; the abdomen comprises only one- third the total length of the body (one-half the length of the body in the other species). Rufo-testaceous, shiny. Head with 5 or more fine, pale hairs along inner margins of antennal fossae. Apical margin of pronotum with 25-30 very short, fine setae. Elytral margins with 2 conspicuous setae and several finer ones. Abdominal setae unusually long; tergites 3-6 with one seta near each posterior angle; tergite 7 without setae; tergite 8 with 2 setae on each half; tergites 3-7 with a few recumbent hairs, but without the moderately dense vestiture of the other species; sternites with a dense covering of long, pale hairs and with a few longer, dark bristles near lateral margins. Pronotum and elytra microscopically punctu- late and with an almost imperceptible pubescence; tergites very minutely and sparsely punctulate; tergite 7 longitudinally strigulose. Head between antennal fossae relatively broad, at narrowest point one-fourth to one-third as broad as head. Antennal fossae relatively long and narrow, reaching apical margin of head. Eyes moderately large but smaller than antennal fossae. Antennal segments with the following relative lengths: 14 : 5 : 3 : 3 : 3 : 3 : 3 : 3 : 3.5 : 3.5 : 10; segments 3-10 strongly incrassate. Pronotum about one-third broader than long, base bisinuate (broadly emarginate, middle of emargination feebly arcuate), posterior angles rounded, apical margin of elytra bisinuate, but anterior angles not trun- cated. Length, 1.4 mm.; width, 1 mm. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 197 Holotype from Passa Quatro, Minas Gerais, Brazil (ex Reichens- perger collection), with Cornitermes sp. In Chicago Natural History Museum. Paratypes. Thirteen specimens, same data as holotype, in Chicago Natural History Museum and Reichensperger collections. Termitohospes silvestrii, new species Rufo-testaceous, very shiny. Pronotum and elytra minutely punctulate, and with extremely fine pubescence; intervals smooth. Tergites uniformly, minutely punctulate and pubescent, and with extremely fine, transverse strigulation. Front of head with a number of fine, pale hairs; pronotum with 2 lateral and 2 short anterior setae; elytral margin with a row of about 18 closely placed, moder- ately long, obliquely directed setae. Tergites 3-6 with 4 moderately long apical setae; tergite 7 without setae; tergite 8 with one long and one short seta on lateral margins and one additional pair of short setae. Sternites 3-6 with 8 long, erect setae on apical margin (some of these setae may be small and indistinct on sternites 3 and 4); sternite 7 with 6 setae; tergite 8 with 2 setae. Sternites with numerous pale setae, recumbent to semi-erect; near base of each stemite these hairs are short, but they are quite long near apex. Head short and broad; front between antennal fossae one-fifth as broad at narrowest point as head width. Eyes larger than antennal fossae, strongly convex. Antennal fossae broad and deep, but not elongated. Antennal segments with relative lengths as follows: 14 : 7 : 3.5 : 3 : 4 : 4 : 4 : 4 : 4 : 4 : 15; segment 2 obconical ; segment 3 narrow at base, at apex one-third broader than long; segments 5-8 increasing in width; 8-10 subequal in width. Pronotum one-fifth to one-fourth broader than long; base straight, posterior angles broadly rounded. Outer apical angles of elytra truncated; maximum length of each elytron about one-fourth as great as distance from lateral margin to suture. Length, 2-2.2 mm. Holotype from Blumenau, Santa Catarina, Brazil (ex Reichens- perger collection), with Nasutitermes aquilinus Holmgren. In Chicago Natural History Museum. Paratypes. Seven specimens, same data as type; in Chicago Natural History Museum and Reichensperger collection. Termitohospes tachyporoides, new species Closely related to silvestrii, from which it differs in these respects: narrower, more strongly deflexed pronotum, the sides of which are more closely appressed against the body (pronotum only one-tenth to one-sixth as broad as long) ; sides of pronotum more nearly parallel near base, the apical margin thus more elliptical; width of head between antennal fossae only one-seventh as broad as head. Chaeto- taxy similar to that of silvestrii except that the vestiture of the sternites is more dense and conspicuous; the fine, pale bristles are numerous and many are quite long; the erect setae of the sternites are not, as a rule, as well developed as in silvestrii. 198 FIELDIANA: ZOOLOGY, VOLUME 40 Length, 2-2.2 mm. Holotype from Passa Quatro, Minas Gerais, Brazil (ex Reichens- perger collection, with Cornitermes sp. (according to accompanying label; no specimens of the host were available). In collection of Chicago Natural History Museum. Paratypes. Forty- three specimens, same data as type; in col- lections of Chicago Natural History Museum, Reichensperger, Borgmeier, and Institute Biologico, Sao Paulo, Brazil. Remarks. Several larger species of Termitohospes resemble some species of the tachyporine genus Tachyporus, and this species has the facies of that genus to a marked degree. TERMITOSOCIUS Seevers Termitosocius Seevers, 1941, Ann. Ent. Soc. Amer., 34: 338. Type species: Termitosocius microps Seevers. Closely related to Termitohospes, from which it is distinguished by the structure of its head and mouth parts. Head long and slender, subtriangular in outline; labium, maxillae, and labrum much elongated, somewhat beak-like in appearance. Eyes small. Antennal fossae relatively small; vertex between fossae about one-fourth as broad as head. Labium similar in structure to that of Termitohospes but its parts very much elongated; labial palpi extremely long and filiform, basal segment seven times as long as broad, segments 2 and 3 longer than broad. Mandibles about two-thirds as long as head capsule, straight except for slightly incurved apices. Galea and lacinia very long and slender, the former with an apical tuft of hairs, the latter strongly sclerotized, dentate (laciniae extending beyond mandibles, for which they may be mistaken). Termitosocius microps Seevers Termitosocius microps Seevers, 1941, Ann. Ent. Soc. Amer., 34: 339, pi. 3, figs. 31, 35, 38, 46 (Kartabo, British Guiana; Nasutitermes octopilis Banks; Chicago Natural History Museum). Coloration uniformly flavate. Head clothed with semi-erect yellow hairs. Pronotum and elytra very minutely and sparsely punctulate and pubescent. Pronotum with three longitudinal rows of setae on each half; an inner row of 3, a middle row of 5, and a marginal row of 5 setae. Elytra with similar rows of 2, 2, 1 setae, respectively, and a comb of about 10 moderately long, yellow hairs on the margin. Tergites 3-6 with 4 stout black setae on posterior margins; tergite 8 with 2 apical and 2 subapical setae. Sternites 3-8 with 6 black setae on apical margin. Antennal segments with relative lengths as follows: 10 : 8 : 5 : 3:3:3:3:3:3:3:8; scape ovoidal; segment 3 slender, longer than broad; segments 4-10 subequal in length but somewhat incrassate; segment 10 more than twice as broad as segment 3. Length, 1.75 mm. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 199 PARATERMITOSOCIUS Seevers Paratermitosocius Seevers, 1941, Ann. Ent. Soc. Amer., 34: 340. Type species: Paratermitosocius vestitus Mann. Closely related to Termitosocius; distinguished by having the smaller pronotum truncate at the apex so that the head is visible from above, and by having antennal segments 5-10 longer than broad. Paratermitosocius vestitus Mann Perinlhus vestitus Mann, 1923, Zoologica, 3: 337 (Kartabo, British Guiana; Nasutitermes octopilis Banks; United States National Museum). Paratermitosocius vestitus Mann, Seevers, 1941, Ann. Ent. Soc. Amer., 34: 340. Material examined. Type series. Color brown, integuments rather dull. Pronotum and elytra with a dense vestiture of pale hairs. Pronotum with 4 moderate setae on each lateral margin ; elytral margins with 3 longer, stouter setae. Tergites 3-6 with one prominent seta near each apical angle; sternites 3-7 with 2 apical setae near each side; sternite 8 with 2 apical and 4 subapical setae; tergites and sternites with a dense covering of fine, pale, semi-erect hairs. Length, 2 mm. BLAPTICOXENUS Mann Blapticoxenus Mann, 1923, Zoologica, 3: 362. Type species: Blapticoxenus brunneus Mann. Termitocolonus Seevers, 1941, Ann. Ent. Soc. Amer., 34: 336. Type species: Termitocolonus ericiogaster Seevers. New synonymy. Distinguished from Termitohospes by its distinctive antennal structure and distinctive abdominal setae. Head deflexed, completely covered by the large pronotum; head smaller in proportion to body size than in Termitohospes. Antennal fossae broad and deep; vertex between the fossae narrow. Eyes prominent. Antennae geniculate, very much elongated (three times as long as the antennae of Termitohospes); scape fusiform, segments 2-10 subcylindrical, twice to three times as long as broad, terminal segment shorter than preceding segments. Gula large, a little more than one-half as long as broad; submentum shorter than gula; mentum with apical border emarginate, its anterior angles acutely produced. Abdomen beset with extremely long, aciculate bristles, most of which are thickened at base. Blapticoxenus brunneus Mann Blapticoxenus brunneus Mann, 1923, Zoologica, 3: 363 (Kartabo, British Guiana; Velocitermes beebei Emerson; United States National Museum). Termitocolonus ericiogaster Seevers, 1941, Ann. Ent. Soc. Amer., 34: 337, pi. 3, figs. 32, 39 (Kartabo, British Guiana; Velocitermes beebei Emerson; Chicago Natural History Museum). New synonymy. 200 FIELDIANA: ZOOLOGY, VOLUME 40 Color flavo-testaceous. Head, pronotum, and elytra without setae; head with sparse, recumbent pubescence, pronotum and elytra sparsely and very minutely punctulate and pubescent. Abdominal macrosetae about one-half longer than the corresponding abdominal segment; stout and dark at base, pale and aciculate distally. Tergites 3-6 with 4 setae on apical margins; tergite 7 without setae; tergite 8 with 2 setae on apical margin and 2 on each lateral margin; tergite 9 with 3 setae on each half, lateral plates of ninth segment with 3 setae. Sternites 3-6 with 10 to 12 macrosetae; sternite 7 with 8 setae; sternite 8 with 6 subapical and 2 apical setae. Length, 1.5 mm. TERMITOSODALIS Seevers Termitosodalis Seevers, 1941, Ann. Ent. Soc. Amer., 34: 337. Type species: Termitosodalis barticae Seevers. Termitomus Silvestri, 1947, Arch. Zool. Ital., 31: 138; Borgmeier, 1950, Rev. Ent. 21: 639. Type species: Termitomus fasciatus Silvestri. New synonymy. Distinguished from Termitohospes by its antennal structure: Antennae three times as long as those of Termitohospes; scape moderately long; segments 2-10 cylindrical, elongated, or subquadrate but not transverse, segment 3 usually the longest. Remarks. Borgmeier placed Termitomus Silvestri in synonymy with Termitohospes Seevers but that is incorrect as Termitomus fasciatus is congeneric with Termitosodalis barticae. Termitosodalis barticae Seevers Termitosodalis barticae Seevers, 1941, Ann. Ent. Soc. Amer., 34: 338, pi. 3, fig. 36 (Kartabo, British Guiana; Velocitermes beebei Emerson; Chicago Natural History Museum). Coloration dull brown. Antennae with segment 2 smaller than scape, cylin- drical; segment 3 elongated, cylindrical, equal to segments 1 and 2 combined; segments 4-10 slightly elongated, subequal except segment 5, which is a little longer. Pronotum, elytra, and abdomen with a dense covering of fine, pale hairs. Pronotum with three irregular longitudinal rows of setae on each half comprised of 2 (inner), 2 (middle), and 3 (submarginal) setae; elytra with similar rows of 1, 2, 2 setae. Tergites 3-6 with 4 rather long, stout setae in subapical rows; tergite 7 without setae; tergite 8 with a basal pair of setae and one on each side. Sternite 3 with 6 erect setae and numerous long, recumbent hairs on apical margin; sternites 4-6 with 8 submarginal setae; sternite 7 without setae; sternite 8 with 2 setae near apex and 6 in a transverse row near middle. Length, 1.5 mm. Termitosodalis fasciatus Silvestri, new comb. Termitomus fasciatus Silvestri, 1947, Arch. Zool. Ital., 31: 140, fig. 7 (Pernam- buco, Brazil; Nasutitermes diversimilis Silvestri; Silvestri coll.). Termitohospes fasciatus Silvestri, Borgmeier, 1950, Rev. Ent., 21 : 647. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 201 Distinguished from barticae by the chaetotaxy of pronotum, elytra, and abdominal sclerites, and by the proportions of the antennal segments. Silvestri's figures should be consulted. Color reddish-brown. Antennae with segments 2 and 3 elongated, apparently subequal in length; segments 3-10 appear to decrease gradually in length. Pro- notum covered with very short hairs and about 24 longer, dark setae; elytra each with about 11 setae. Tergites 3-6 with 6 long, black setae in apical rows; tergite 7 without setae; tergite 8 with 2 sub-basal and 2 subapical setae. Sternites with 10 setae in apical rows. Length, 2 mm. Subtribe HETAIROTERMITINA HETAIROTERMES Cameron Termophila Lea., 1910, Proc. Roy. Soc. Victoria, 23: 136; Fenyes, 1918, Gen. Insect., 173 A: 91. Type species: Termophila latebricola Lea. Hetairotermes Cameron, 1920, Trans. Ent. Soc. London, 1920: 223 (for Termophila Lea, preoccupied, Termophila Grassi, 1887); 1921, Trans. Ent. Soc. London, 1921: 357, 387. Head, pronotum, and elytra smooth, highly polished; abdomen less so. Head prognathous, scarcely deflexed; covered to eyes by pronotum. Dorsum rather strongly declivous between antennal fossae and margined in front by a fine, U-shaped line. Clypeus almost vertical, convex at middle and becoming concave laterally, where it forms one surface of the moderately deep antennal fossae. Antennae inserted under a short arcade. Eyes large. Antennae robust, moder- ately long. Gula very broad, scarcely narrower in front. Submentum broad, its apex emarginate. Mentum very broad and short, its apex concave, its apical angles narrowly produced. Labial palpi 3-segmented, very long and slender, filiform. Galea and lacinia subequal, long and slender. Pronotum one-half to two-thirds broader than long; anterior margin slightly arcuate to moderately emarginate, base arcuate, sides strongly rounded, basal angles rounded. Meso- sternal process slender, acute; middle coxae nearly contiguous. Legs densely pubescent, usually bearing long setae. The species of Hetairotermes have been very superficially de- scribed. Perhaps the following notes may aid in determinations, but the genus needs a thorough revision. Hetairotermes leai Cameron is a synonym of Lauella vitiensis Mann. Hetairotermes latebricola Lea Termophila latebricola Lea, 1910, Proc. Roy. Soc. Victoria, 23: 137 (Galston, New South Wales; "from a termite nest;" Lea coll.). Hetairotermes latebricola Lea, Cameron, 1920, Trans. Ent. Soc. London, 1920: 223. Material examined. Two cotypes, New South Wales; Cameron collection, British Museum (Natural History); and other specimens. 202 FIELDIANA: ZOOLOGY, VOLUME 40 Testaceous-brown, pronotal and elytral margins and abdomen paler. Ver- texal setae short and inconspicuous; one pair of clypeal setae. Pronotum with irregular longitudinal rows of relatively short setae arranged 4, 3, 4 (on each half) ; no fine hairs present. Dorsum impunctate. Antennae relatively short for the genus; less than twice as long as pronotum (95 : 50). Pronotum one-half broader than long (75 : 50); anterior margin slightly arcuate; pronotum broadest in front of middle, the sides converging moderately in front; base moderately arcuate; side margins moderately reflexed, base very feebly so. Elytral side margins only one- fourth longer than sutural length (behind scutellum). Tergites and sternites not pubescent. Length, 1.1 mm. Hetairotermes agilis Cameron Hetairotermes agilis Cameron, 1920, Trans. Ent. Soc. London, 1920: 223 (Woodlands, Singapore Island, Malaya; "with a wood-dwelling termite;" Cameron coll.). Material examined. The type and one other specimen, Cameron collection. Stated by Cameron to differ from latebricola in having the sides of the elytra punctured, but there are much more important dif- ferences: The antennal segments have different proportions; the chaetotaxy of pronotum and head is different; the pronotum and elytra have different proportions; the sternites are densely pu- bescent. Castaneous, abdomen pitchy-testaceous. Head with one pair of setae be- tween antennae and a row of about 11 setae between eyes. Pronotum with four irregular longitudinal rows of macrosetae on each half, comprised of 4 or 5, 4, 4, and 3, respectively. Pronotum with a moderate number of fine, short hairs. Elytra with about 15 macrosetae in no perceptible pattern; sides with dense, recumbent pubescence ending at upper surface. Antennae approximately twice as long as pronotum (100 : 48); relatively short and compact; scape moderately stout, compressed; segments 4-10 incrassate; segment 3 about one-half longer than broad; segments 4-10 transverse. Antennal segments of these relative lengths: 14:10:9:5:6:7:7:7:9:7:16. Pronotum three-fifths broader than long (76 : 48); anterior margin straight; sides evenly arcuate; base slightly arcuate; side margins and base narrowly reflexed. Elytral side margins a little less than one-half longer than sutural length. Tergites 3-7 with dense, recumbent pubescence; each with 4 long, black (pale-tipped) macrosetae, middle pair with 2 shorter, pale setae between them. Sternites densely pubescent. Hetairotermes piceus Cameron Hetairotermes piceus Cameron, 1920, Trans. Ent. Soc. London, 1920: 224 (Bukit Timah, Singapore, Malaya; "with a wood-dwelling termite;" Cameron coll.); 1921, Trans. Ent. Soc. London, 1921: 387. Material examined. The type specimen. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 203 Cameron's differential diagnosis of this species is very poor. It is closely related to agilis and may be distinguished by the appreciably longer antennae, the proportions of the antennal seg- ments, and minor differences in chaetotaxy. Pitchy black. No pubescence on head, pronotum, or elytra. Head setae as in agilis; pronotum with about 26 macrosetae in roughly five longitudinal rows, although a pattern is difficult to discern; elytra with several dozen macrosetae and many shorter finer hairs, but these grade into one another and are difficult to count. Antennae long and slender, two and three-fifths longer than pronotum; relative lengths of segments: 16 : 10 : 12 : 10 : 9 : 9 : 9 : 8 : 8 : 8 : 15; basal three segments of antennae slender; 4-10 gradually but not markedly incrassate; 2 and 3 about twice as long as broad; 4-8 longer than broad; 9-10 subequal in length and width; segment 11 twice as long as 10. Pronotum similar to that of agilis; three-fifths broader than long (80 : 50); anterior margin slightly emarginate; sides evenly arcuate. Elytra as in agilis. Chaetotaxy of abdomen as in agilis. Hetairotermes bryanti Cameron Hetairotermes bryanti Cameron, 1950, Ann. Mag. Nat. Hist., (12), 3: 98 (Penang, Malaya, no host cited; Cameron coll.). Material examined. The holotype. Cameron compared this species with agilis, but many of the statements that he made are incorrect. In antennal characters this species is intermediate between agilis and piceus. The pronotal apex is more emarginate than in the foregoing species, and the pronotum is widest in front of the middle. Head with 16-18 medium-length hairs; these tend to be in rows. Pronotum with 12 or more dark macrosetae and many finer but not much shorter hairs scattered among them. Antennae two and two-fifths longer than pronotum (110 : 45); relative lengths of segments: 14 : 8 : 9 : 9 : 8 : 8 : 7 : 7 : 6 : 7 : 14; three basal antennal segments slender; 2-6 not appreciably different in length but slightly incrassate; 6-10 incrassate; 7-10 broader than long. Pronotum almost two-thirds broader than long (74 : 45); its apical margin moderately deeply emarginate; broadest in front of middle; base quite arcuate, with an indication of sinuation; side and basal margins reflexed. Lateral elytral margin about two-thirds longer than sutural margin (56 : 34). Abdominal chaetotaxy much as in agilis and piceus. Hetairotermes punctiventris Lea Termophila punctiventris Lea, 1910, Proc. Roy. Soc. Victoria, 23: 137, fig. 4 (Bridgetown, West Australia; "in a nest of white ants;" Lea coll.). Hetairotermes punctiventris Lea, Cameron, 1920, Trans. Ent. Soc. London, 1920:223. According to Lea, this species is distinguished from H. latebricola by its peculiarly punctate seventh tergite and sternite, and by having 204 FIELDIANA: ZOOLOGY, VOLUME 40 its pronotum slightly upturned or flattened at the hind angles but not reflexed along the lateral margin. Testaceous-brown, pronotum slightly paler. Head glabrous, pronotum and elytra with short, sparsely distributed setae; abdomen with rather sparse but more noticeable setae, more or less confined to apex and sides of segments. An- tennae extending almost to apex of elytra; segments 2-10 subequal in length, gradually incrassate. Pronotum twice as broad as long; anterior angles strongly rounded; hind angles moderately so. Abdomen with sixth tergite longitudinally strigose; seventh tergite and sternite with distinct punctures. Hetairotermes formicicola Oke Hetairotermes formicicola Oke, 1933, Proc. Roy. Soc. Victoria, 45: 135 (Gyp- sum, Victoria, Australia; Iridomyrmex nitida; Oke coll.). Oke's description of this species was very general. He stated that H. formicicola differs from the two Australian species "in not having any 'longer setae' anywhere. Under a hand lens it appears to be almost glabrous, but under the microscope it is seen to have a close, minute pubescence. The fine puncturation of most of the surface will also distinguish this species." Two matters relating to this species require confirmation: First, that it belongs to this genus, and, secondly, that it occurs with ants. Hetairotermes insulanus, new species This species apparently differs from the other members of the genus in its chaetotaxy, especially in the presence of extremely long marginal setae on pronotum and elytra. Rufo-testaceous; abdomen paler. Head, pronotum, and elytra smooth, shiny, glabrous, impunctate. Head without setae. Lateral pronotal margins with 7 and elytral margins with 4 extremely long, curve-tipped setae. Pronotum with 2 setae on anterior margin, and its disk with 12 setae as follows: each de- flexed side with a pair about the middle, a transverse row of 4 at the center of the disk just behind the middle, a pair behind this row, and a pair in front of the basal border. Elytral disk with 11-12 erect setae irregularly arranged. Tergites with numerous fine setae of varying lengths. Sternites smooth, shiny and subglabrous, but with numerous long, curve-tipped setae. Antennal segments with the following relative lengths: 9:8:6:6:7:6: 6:6:6:6:8; scape and segment 2 more slender than the following segments; segment 5 broadest, segments 6-10 a little less broad. Pronotum slightly less than twice as broad as long; anterior border shallowly emarginate; base arcuate medi- ally, nearly straight near sides; sides arcuate, basal angle rounded. Elytra generalized. Eighth sternite very strongly produced to form a large arcuate lobe. Legs densely hairy and bearing long setae; middle tibiae with about 6 and hind tibiae with about 8 very long, fine, curve-tipped setae on their outer margins; femora with similar setae on outer margins near their apices. Length, 1.1 mm. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 205 Holotype from Koror Island, Palau Islands, Micronesia; collected January 16, 1948, by Henry S. Dybas (Pacific Science Board Expedition), with Nasutitermes rufirostris Hill. In collection of United States National Museum. No paratypes. TERMITOBRA, new genus Type species: Termitobra perinthoides, new species. Termilobra differs from Hetairotermes in having the galea con- siderably shorter than the lacinia, the pronotum only one-half broader than long, and the integuments densely pubescent. Head moderately deflexed, covered by pronotum to the eyes; vertex declivous between antennal fossae, which are moderately deep. Gula very broad, its sides converging moderately in front; submentum broad and short, its apex deeply emarginate; mentum distinctive in structure (its apical angles as in Termitohospes); prementum very long; ligula short, filiform; labial palpi 3-segmented, extremely long; relative lengths of segments 14 : 3 : 2, the basal segment about seven times as long as the maximum width; segments 2 and 3 more slender, filiform. Galea somewhat shorter than lacinia, its oblique apex setose; lacinia dentate and setose. Segment 2 of maxillary palpi club-shaped; segment 3 fusiform. Mandibles falcate, the incurved apices long and slender. Antennae feebly incrassate; seg- ments cylindrical, slightly elongated. Pronotum transverse, one-half broader than long, anterior border almost straight, sides converging slightly in front, base slightly arcuate. Elytra generalized. Mesocoxae large, contiguous. Meta- sternum very short. Hind coxae large. Tarsi 4, 4, 5-segmented. Abdomen tapering, ninth tergite deeply bifid. Remarks. Termitobra perinthoides bears such a close resemblance to Perinthus and Lauella of the Termitonannini that it may easily be mistaken for a species of that tribe. However, the similarities are only superficial and the head capsule and mouth parts establish the relationship with the Termitohospitini. Termitobra perinthoides, new species Head, pronotum, elytra, tergites, and sternites densely punctulate and with a dense covering of very fine pubescence. Pronotum with four longitudinal rows of long setae on each half comprised of 3, 2, 1, 3 setae, respectively, beginning with a medial row. Elytra with similar rows of 1, 1, 2, 3 setae. Tergites 3-6 with a row of 4 subapical setae; tergites 7 and 8 each with a pair of widely sepa- rated sub-basal setae. Sternites with an apical row of about 8 long, fine setae and a similar row about the middle of each sternite. Length, 1.2 mm. Holotype from Peleliu Island, Palau Islands, Micronesia; collected August 4, 1945, by Henry S. Dybas, with Nasutitermes rufirostris Hill. In collection of Chicago Natural History Museum. Paratypes. Three paratypes, same data as holotype. 206 FIELDIANA: ZOOLOGY, VOLUME 40 Subtribe TERMITOSPECTRINA TERMITOSPECTRUM Mann Termitospectrum Mann, 1926, Proc. Ent. Soc. Wash., 28: 153. Type species: Termitospedrum thoracicum Mann. Head not deflexed, only occipital border covered by pronotum. Vertex forming an obtuse angle between antennae, then declivous to front margin; clypeus short; antennal fossae moderately large, somewhat elongated, antennae inserted under a feeble vertexal arcade. Head broader than long; labrum large, arcuate, not emarginate, mouth parts elongated and somewhat beak-like in appearance. Antennal scape short and broad, segments 2 and 3 slightly elongated, 4-10 subquadrate. Gula very broad, its sides scarcely converging in front; submentum very broad, its sides subparallel; mentum much narrower than submentum, semi-tubular due to inflexion of its sides; prementum very much elongated; ligula nearly filiform. Labial palpi apparently 2-segmented; filiform, first segment much longer than second. Maxillae elongated; galea and lacinia rather long, the former slender, curved, its apex with a cluster of hairs; the latter more robust and rather heavily sclerotized. Maxillary sinus very large. Segment 2 of maxil- lary palpi medio-laterally compressed, segment 3 longer and somewhat fusi- form. Pronotum distinctive: strongly transverse, its apical border very deeply emarginate and anterior angles produced along sides of head as far as eyes; dorsum with two broad, deep sulci extending forward from near base to vicinity of apical angles; surface lateral to sulci very convex and membranaceous in texture. Pro- notal hypomera broad, reflected against pronotum, but partially visible from side. Elytra membranaceous in part, especially near apices, apical border emargi- nate. Abdomen about as broad as foreparts; recurved over thorax; not obviously physogastric but with a tendency for small areas of membranous integument to show along the sides near the paratergites. Termitospectrum thoracicum Mann. Figure 31, e, f. Termitospectrum thoracicum Mann, 1926, Proc. Ent. Soc. Wash., 28: 154 (Kartabo, British Guiana; Nasutitermes gaigei Emerson; United States National Museum). Flavo-testaceous; head, sides of pronotum, tips of elytra, and parts of abdomen translucent, pale. Clypeus and anterior part of vertex with a few moderately long, dark setae; vertex with sparse, short hairs. Mentum with 4 black setae at apex and 2 in middle; submentum with a few inconspicuous hairs. Pronotum with a pair of setae on anterior margin, two pairs on disk and 4 setae on each convex area lateral to sulci. Elytra with a number of fine, pale setae. Sternite 3 with a moderate number of setae; sternite 4 with 10 or more unusually long bristles with curved tips; other sternites with setae only on apical margins. An- tennal segments with the following relative lengths: 10 : 5 : 5.5 : 4 : 4 : 4 : 4 : 4 : 4 : 4 : 8; scape not much longer than broad; segments 2 and 3 subequal in width; segments 4-10 slightly transverse, not incrassate; segments verticillately setose. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 207 TERMITOECIA Bernhauer Termitoecia Bernhauer, 1921, Ark. Zool., 13: 20. Type species: Termitoecia fabulosa Bernhauer. Differs from Termitospectrum in its larger size, more robust build, distinctive elytra, pronotal form, distinctive mentum, and scaphoidal abdomen. Moderately robust in build. Head transverse, not deflexed; vertex forming an obtuse angle between antennae, then so abruptly declivous as to be almost vertical; clypeus short. Labrum large, deeply emarginate. Antenna! fossae relatively small. Antennae elongated, most segments longer than broad. Mouth parts elongated, somewhat beak-like in appearance. Gula broad, its sides scarcely converging in front; submentum broader, its apex broadly emarginate; ligula entire, moderately long, rounded; mentum large, trapezoidal, its apex straight. Labial palpi probably 2-segmented; segment 1 much elongated, segment 2 shorter and more slender, less than one-half as long as basal segment. Galea and lacinia equal in length; the former very long and slender, heavily sclerotized, its apex with a few hairs; the latter with an acute apex, spinose, its medial surface with a row of setae. Maxillary palpi with segment 2 broader distally, medio-laterally compressed; segment 3 elongated, sub-fusiform. Pronotum three-fifths broader than long (fabulosa) to twice as broad as long (wilsoni); about three-fifths broader than head. Pronotum broadly cordiform; apex broadly, shallowly emarginate, apical angles produced a little, sides strongly arcuate, basal angles obsolete, base arcuate. Pronotal disk flat to convex; outer one-fourth explanate, membranaceous, and very feebly pigmented; explanate sides delimited from disk by shallow, incomplete sulci terminating in basal foveae. Pronotal hypomera large, visible from side. Elytra distinctive: broader than pronotum, sides strongly arcuate, apex strongly emarginate (maximum length two-fifths greater than sutural length), disk with a deep triangular impression in the distal two-thirds, the sides of elytra much swollen and convex. Abdomen not physogastric or limuloid; capable of being reflected over fore- parts, somewhat scaphoidal in dorsal view; fourth segment about as broad as elytra, segments 5-8 decreasing in width. Paratergites large, especially the inner ones. Termitoecia fabulosa Bernhauer. Figure 31, b. TermUoecia fabulosa Bernhauer, 1921, Ark. Zool., 13, no. 8, p. 21 (Herberton, Queensland, Australia; no host indicated; Chicago Natural History Museum). Material examined. The holotype. This specimen is accom- panied by a termite, Schedorhinolermes intermedius Brauer, which may be presumed to be the host of this species. Head and pronotum rufo-testaceous, abdomen reddish-yellow, elytra brown; pronotal sides membranaceous, very pale flavate. Head with a small, shallow pit in center of vertex, flanked by a seta on each side; head sparsely punctate and finely reticulated, and with sparse, short hairs. Pronotum with 4 apical 208 FIELDIANA: ZOOLOGY, VOLUME 40 setae, 2 or more on lateral margins, 2 on membranaceous sides, 2 along basal border; pronotum with some extremely fine punctules and pubescence, but subglabrous in appearance. Elytra very minutely punctulate and pubescent. Tergites with a marginal row of setae, usually 4 dark setae and some finer ones; tergites very finely reticulated but not punctulate or pubescent. Sternites 4 and 5 with a marginal row of about 20 long, fine setae, sternite 6 with about 10, and tergite 7 with about 6. Sternites extremely minutely punctulate and pubescent and finely reticulated. Pronotum and elytra subequal in length. Antennal segments with the following relative lengths: 30 : 18 : 18 : 13 : 12 : 12 : 12 : 12 : 12 : 12 : 25; segments 2 and 3 subequal in length and width, about three times as long as broad; segments 4-10 incrassate; segment 4 only three-fifths as broad as 10; all segments longer than broad. Length, 3.2 mm. Termitoecia wilsoni Cameron. Figure 31, c. Termiloecia wilsoni Cameron, 1943, Ann. Mag. Nat. Hist., (11), 10: 349 (Bogan River, New South Wales, Australia; with termites; Cameron coll.). This species has not been available; apparently it is distinguished from fabulosa by its darker coloration, different proportions of antennal segments, broader pronotum, and impunctate, glabrous, non-reticulated head. Blackish-brown to dark reddish-brown, pronotal sides brownish-yellow. Head impunctate, glabrous, and without ground sculpture; pronotum with a few extremely fine punctures and very short hairs; elytra very finely and sparingly punctured and with short stiff hairs. Elytra a little longer than pronotum. Pronotum twice as broad as long. Basal segment of antennae stout, segment 3 a little longer and stouter than 2, segments 4 and 5 a little longer than broad, 6-10 subequal in length and width. TERMITANA Fairmaire Termitana Fairmaire, 1899, Bull. Soc. Ent. France, 1899: 316; Wasmann, 1912, Zeitschr. wiss. Zool., 101: 88; Fenyes, 1920, Gen. Insect., 173B: 288. Type species: Termitana perrieri Fairmaire. Head covered to eyes by pronotum; vertex probably unmodified (strongly impressed according to Fairmaire, but this is probably due to collapse of the thin integument, as is evidently the case in the specimen at hand). Antennal fossae and clypeus typical of the subtribe, the latter strongly convex transversely. Gula very broad, about two-fifths as broad as head, its sides scarcely converging in front; submentum broader than gula, almost one-half as broad as head and moderately long, its apex broadly concave; mentum large, trapezoidal, its apex straight. Labial palpi long and filiform (not determined whether two- or three- segmented). Pronotum almost one-half broader than long, strongly convex at middle, broadly but not deeply impressed laterally (the area adjacent to each apical angle not involved). Pronotum entirely corneous, its sides not a different texture. Pronotal shape distinctive: apex almost straight, broadest point about one-half the distance from base to apex; sides in front of broadest point converging SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 209 * moderately; sides behind broadest point converging at a forty-five degree angle to form a broad arc with the base. Sutural length of elytra only about one-half that of lateral margins. Elytral disk with a moderate longitudinal, submarginal impression (not as deep or as conspicuous as in Termitoecia). Coxae very large, anterior coxae long and spindle-shaped, middle coxae almost attaining posterior margin of metasternum, hind coxae very broad, somewhat produced beyond lateral articulation. Tarsi 4, 4, 5-segmented (not 4, 5, 5-segmented as stated by Fairmaire). Abdomen scaphoidal, broadest at level of fifth segment (about one-half broader than pronotum), sclerites not separated and with almost no exposed membranous integument. Paratergites large, elevated to a very marked degree to form a conspicuous margin. Remarks. The position of Termitana has been incorrectly de- duced due to the fact that Fairmaire miscounted the tarsal segments. Study of a "cotype" in the Bernhauer collection reveals the relation- ship indicated here. Termitana perrieri Fairmaire. Figure 31, d. Termitana perrieri Fairmaire, 1899, Bull. Soc. Ent. France, 1899: 316 (Betsi- boka, Madagascar; "with termites;" Fairmaire coll.). Material examined. One specimen, labeled cotype, from Mada- gascar, in Bernhauer collection, Chicago Natural History Museum. Rufo-flavate. Head without macrosetae except for those on clypeus. Pro- notum with 6 fine setae on apical margin, 5 moderately long and 5 very short hairs on lateral margins, a pair of basal setae, and three longitudinal rows of 3, 2, 1 setae, on each half of disk. Elytra with a moderately dense vestiture of fine, pale hairs, more numerous on sides, disk with a few darker, erect setae. Tergites, as a rule, with only an apical row of rather long bristles; tergites im- punctate and glabrous. Sternites with an apical row of setae; surface of third sternite finely pubescent, others with only sparse, very minute stubble. Head impunctate, pronotum subimpunctate, with only sparse punctures bearing short stubble. Antennae with relative lengths of segments as follows: 17 : 13 : 9 : 8 : 8:8:8:8:8:8: 18; segments 1-3 longer than broad; segment 4 broader than 3; segments 4-10 similar in size and form, increasing slightly in width. Length, 2.1 mm. Subtribe TERMITUSINA TERMITUSA Wasmann Termilusa Wasmann, 1905, Rev. d'Ent., 24: 199; 1911, Rev. Zool. Afr., 1: 162; Fenyes, 1918, Gen. Insect., 173A: 106. Type species: Termilusa sjostedti Wasmann. Glaphyrinus Bernhauer, 1942, in Bernhauer and Paulian, Rev. Zool. Bot. Afr., 35: 375. Type species: Glaphyrinus mundus Bernhauer. New synonymy. Head strongly deflexed, its long axis almost vertical; a small area of vertex visible from above. Head subtriangular in outline; mouth parts prolonged, 210 FIELDIANA: ZOOLOGY, VOLUME 40 somewhat beak-like; vertex large, convex, rather broad between the moderately large antennal fossae. Clypeus moderately long; labrum large. Antennae elongated, slender, its segments subcylindrical and usually elongated, never appreciably incrassate. Gula broad, its margins not converging much in front; submentum moderately long broad, its apex arcuately emarginate; mentum long, subtrapezoidal; ligula elongated, nearly filiform. Labial palpi 2-segmented; basal segment extremely long, robust; segment 2 more slender, slightly bowed, about one-half as long as first. Mandibles very long, falcate. Maxillae much elongated; galea and lacinia subequal in length; lacinia terminating in a long, slender, in- curved spine, its medial surface with about 10 long, stout, spinose setae; apex of galea membranous, bearing many fine, ciliate hairs. Pronotum about one-half broader than long, large, convex, shield-like, its sides deflexed; lateral margins arcuate, apex almost straight, base broadly emar- ginate or bisinuate. Hind coxae large, very broad, strongly produced laterally; metepimera very large. Abdomen long and slender as a rule, but rather broad in one species. Ninth tergite bifid, each half forming a long slender setose process. Remarks. This genus needs a complete revision of its species, but adequate material for such a task has not been available. At one time or another I have examined most of the species, but this has not been particularly satisfactory, as material for dissection purposes is required before the species can be properly compared and diagnostic features determined. It should be noted that the dorsal bristles of Termitusa break off very easily and many specimens lack nearly all of them. Slide preparations reveal setigerous punctures on specimens almost com- pletely lacking the bristles. Wasmann indicated in a key that escalerae lacks dorsal setae but this is not so. Termitusa sjoestedti Wasmann Termitusa sjostedti Wasmann, 1905, Rev. d'Ent., 24: 200 (Cap Debundscha, Cameroons; Cubitermes fungifaber Sjostedt; Naturh. Reichsmus. Stock- holm); 1911, Rev. Zool. Afr., 1 : 163, pi. 5, fig. 9; Hegh, 1922, Les Termites, text fig. 421. Material examined. Two specimens in Wasmann collection on pin bearing type label (the head of one specimen and antennae of the other are missing). In addition, a series from Longuyi, Cameroons, French Equatorial Africa; with Cubitermes fungifaber Sjostedt. Brownish, very shiny; extremely finely punctulate and pubescent. Pronotum with irregular longitudinal rows of 4, 3, 2, 4 setae on each half, beginning with medial row. Elytra with about 6 discal setae and one on lateral margins. Tergites 3-6 with 4 very strong bristles set in prominent, elongated tubercles (bristles stout, black at base, pale distally); tergite 7 with 4 slender setae; tergite 8 with a dense brush of long, fine, pale hairs; each half of tergite 9 bearing 5 or more very long, dark bristles. Lateral plates of ninth segment also attenuated and bearing bristles. Sternites with about 6 setigerous tubercles with long, black bristles; SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 211 paratergites usually with one similar bristle. Sternites finely reticulated, very finely pubescent, and finely setose. Antennae long enough to reach apex of elytra; segments with the following relative lengths: 8:8:5:8 .-8:8:8:8:8:8:12; all segments except third twice as long as broad. Length, 2-2.4 mm. Termitusa escalerae Fauvel Termitusa escalerae Fauvel, 1906, Mem. Soc. Esp. Hist. Nat., 1: 289 (Cabo San Juan, Biafra, Rio Muni; Cubitermes sp.; Fauvel coll.); Wasmann, 1911, Rev. Zool. Afr., 1: 163 (Gabon, French Equatorial Africa). Material examined. One specimen in Wasmann collection from type locality labeled type; presumably this is a syntype. This species is very closely related to sjoestedli and difficult to distinguish from it. The pronotal and elytral setae are considerably less conspicuous than in sjoestedti; its size is smaller; the scape is one-half longer than segment 2. Rufo-testaceous. Pilosity inconspicuous; setae of pronotum and elytra long but very fine and pale. Antennae with segments of these relative lengths: 9:6: 5 : 8 : 8 : 8 : 8 : 8 : 8 : 8 : 14. Antennae long and slender, feebly incrassate; all segments elongated but only about one-half longer than broad. Length, 1.9 mm. Termitusa hystrix Wasmann Termitusa hystrix Wasmann, 1911, Rev. Zool. Afr., 1: 159, 163, pi. 5, fig. 10 (Sankuru, Belgian Congo; Cubitermes fungifaber Sjostedt; Wasmann coll.); Hegh, 1922, Les Termites, p. 613, fig. 421. Material examined. Three specimens in Wasmann collection on pin bearing type label; all with head, pronotum, or parts thereof missing. Two other complete specimens from type locality. Flavo-testaceous, elytra darker. Integuments glabrous, except eighth tergite with a dense brush of long, pale hairs. Stout, rather long, dark, pale-tipped bristles as follows: pronotum with longitudinal rows of 3, 3, 3, 4 bristles; elytra with 2, 1, 3, 2 bristles; tergites 3-6 with 4 setae; ninth tergite with about 6 long bristles. Antennae with segments of following relative lengths: 10:7:6:8:9:9:9: 8 : 8 : 8 : 15. All segments longer than broad; segment 3 the most slender; seg- ments 3-11 incrassate. Termitusa lujae Wasmann Termitusa lujae Wasmann, 1911, Rev. Zool. Afr., I: 156, 159, 164 (Sankuru, Belgian Congo; Cubitermes sankurensis Wasmann; Wasmann coll.). Material examined. Three specimens on pin labeled type in Wasmann collection. Three additional topotypes, one with Cubi- 212 FIELDIANA: ZOOLOGY, VOLUME 40 termes fungifaber Sjostedt. One specimen, Kondue, Belgian Congo, in United States National Museum. This species and the following, T. cameroni Reichensperger, are distinguished by a moderately dense vestiture of nearly recumbent hairs on the head; these are almost in transverse rows and are directed caudad. The pronotum has a similar vestiture. Coloration uniform brown. Head and pronotum with hairs as described above, those of pronotum in four transverse rows of about 7 setae each. Tergites 3-7 with a moderately dense vestiture of pale semi-recumbent hairs; tergites 3-6 with an apical row of 4 very long bristles (pale-tipped) ; tergite 8 with a very dense brush of long, pale setae. Antennae shorter than in most other species, reaching only to middle of elytra; their segments with these relative lengths: 8:7:4:5: 5:6:6:5:5:5:1 0. Antennal segments 1 and 11 somewhat longer than broad; segments 3-10 subequal in length and width, slightly incrassate. Pronotum two- fifths broader than long; pronotal base strongly bisinuate. Termitusa cameroni Reichensperger Termitusa cameroni Reichensperger, 1929, Ent. Blatter, 25: 136 (Ngerengere, Tanganyika Territory; Cubitermes glebae Sjostedt; Reichensperger coll.). This species is evidently closely allied to lujae according to Reichensperger but may be distinguished from it by its paler, more yellow coloration. It is distinguished from the other species by its pilosity. The head and pronotum are yellow-brown and the elytra dark brown. Termitusa mundus Bernhauer, new comb. Glaphyrinus mundus Bernhauer, 1942, in Bernhauer and Paulian, Rev. Zool. Bot. Afr., 35: 375 (Edea, Cameroons, French Equatorial Africa; in the upper galleries of a termitary; Chicago Natural History Museum). Material examined. Holotype in Bernhauer collection, labeled "Glaphyrius mirandus Bernhauer, typus." Inasmuch as the type bears the correct locality data, it seems evident that Bernhauer was negligent in checking the type label against his manuscript name. A second specimen, labeled "cotype," from Tiko, Cameroons (ex Hamburg Museum) does not belong to Termitusa mundus, but I have not been able to identify it with any named species of the genus. It is impossible at present to determine whether T. mundus is a valid species or not as the identity of escalerae and hystrix has not been established. It is less than 2 mm. in length, light reddish-brown in color, has very long antennae resulting from the fact that segments 4-10 are almost twice as long as broad, has extremely finely and sparsely punctulate pronotum and elytra which lack pubescence, SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 213 and has longitudinal rows of setae on pronotum comprised of 3, 3, 3, 3 setae (beginning with median row); similar rows on elytra with 2, 1, 3, 2 setae. The host is probably a species of Cubitermes. Termitusa quadricollis Cameron Termilusa quadricollis Cameron, 1952, Rev. Zool. Bot. Afr., 46: 331 (Massif des Kundelungu; host not stated; Museum of Belgian Congo). Material examined. Two cotypes from type locality in Cameron collection, British Museum (Natural History). Distinguished from the other species of the genus by the pro- portions of the pronotum, which is only about one-fifth broader than long. Head with scattered punctures, but nearly glabrous. Pronotum minutely punctured but with little evidence of pubescence. Pronotal macrosetae fine; arranged in rows of 4, 3, 3, 3 setae, respectively, beginning with medial row. Elytral macrosetae in rows of 2, 2, 2 setae; elytra with almost no fine hairs. Ter- gites with few recumbent hairs; tergites 3-7 with 6 long, apical or subapical macrosetae. Pronotum one-fifth broader than long; base broadly and rather strongly arcuate; broadest near base, sides converging moderately in front of middle, apex nearly straight. Termitusa lativentris, new species. Figure 32. Distinguished from the other species of Termitusa by broad, distinctive abdomen; shorter antennae with segments of distinctive proportions; maxillary palpi; and pubescent elytra. This species may represent a new genus but it is desirable to know more about Termitusa before forming more generic categories. Coloration of head, pronotum, and elytra light reddish-brown; abdomen testaceous. Antennae shorter than in most species of the genus; relative lengths of segments: 8 : 8 : 5 : 5 : 6 : 7 : 6 : 5 : 5 : 5 : 12. Segments 4-10 relatively short, only segment 6 a little longer than broad, the others subquadrate to slightly transverse; segments 2-10 moderately incrassate. Maxillary palpi unusually large, especially the fusiform third segment, which is one-third or more longer than second and twice as broad. Pronotum very finely and sparsely punctulate and very sparsely pubescent. Basal half of elytra with rather numerous fine, semi-recumbent hairs. Tergites glabrous, non-punctate and without pubescence. Pronotum with longitudinal rows of setae comprised of 4, 2, 2, 3 setae (beginning with medial row), and elytral rows of 2, 2, 2 setae. Tergites with an apical row of 4 pale, slender setae. Pronotum one-half broader than long, its base strongly bisinuate. Elytral suture about three-fifths as long as pronotum, elytral apex scarcely emarginate. Abdomen much broader than in the other species and not acuminate; possibly to some extent physogastric, although exhibiting no mem- branous areas. Abdomen broadest at fifth segment; upper surface somewhat 214 FIELDIANA: ZOOLOGY, VOLUME 40 scaphoidal in form, its margins somewhat raised and the paratergites broader than usual. Holotype from Brazzaville, French Equatorial Africa; collected April 1, 1948, by A. E. Emerson, with Noditermes cristifrons Was- mann. In collection of Chicago Natural History Museum. Paratypes. Seven specimens, same data as type. FIG. 32. Termiiusa lativenlris Seevers. Tribe TERMITOPAEDIINI, new tribe This tribe is proposed for a group of Old World genera formerly placed in the Myrmedoniini. The latter group as now constituted is polyphyletic, and its genera must be reclassified. Some of the assignments to the Termitopaediini must be regarded as tentative; our knowledge of the palaeotropical termitophile fauna is probably too fragmentary to permit more than preliminary judgments. An earlier section of the paper (p. 52) is devoted to a discussion of the biogeography and host relationships of the Termitopaediini. The tribe as herein constituted contains three generic groups: Termitopaedia, Dioxeuta, Neodioxeuta, Termitopulex, and Termito- linus; Termitotropha and Termitobaena; Termitotecna, Prolermitobia, SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 215 and Termitobia. The tribe as a whole is not easily characterized. This may indicate that several independently derived phyletic lines of termitophiles are represented, and that a subdivision will be needed at a later date. The tribe is not characterized by a distinctive facies; the con- trast between Termitobia and Termitolinus, for example, is very striking. While the group as a whole may be said to be physogastric, some genera exhibit this character to a very minor degree. There is no common pattern of physogastry and the abdomens of various genera are appreciably different. The characters seemingly shared by the genera are as follows: tarsi 4, 5, 5-segmented; maxillary acetabula not extremely developed as in the Myrmedoniini; galea and lacinia moderate in length (in contrast to the Myrmedoniini), the galea usually foliate and membranous; terminal antennal seg- ments without coeloconic sensilla; mesocoxal acetabula large, shal- low, faintly margined externally or not at all ; the broad mesosternal process rounded or angulate; middle coxae narrowly or moderately widely separated; hind coxae subtriangular, not produced beyond lateral articulation; metepimera obliquely truncated posteriorly and not prolonged beyond coxal articulation. The men turn is free in Termilopaedia and related genera, fused with the submentum in the other genera. It may be noted that several characters listed above also typify the Corotocini: shallow mesocoxal acetabula, triangular hind coxae, abbreviated metepimera, and fused mentum. Without much doubt these are convergences associated with the habit of ter- mitophily and do not signify close relationship of these tribes. KEY TO GENERA OF TERMITOPAEDIINI 1. Abdomen extremely inflated, its basal segments protruding over elytra (fig. 33, b, e) Abdomen moderately inflated at most 3 2. Abdomen as in fig. 33, b; Indomalaya Neodioxeuta, new gen. Abdomen as in fig. 33, e; Africa Termitobia Wasmann 3. Antennae extremely long, capable of reaching beyond middle of abdomen. Termitopulex Fauvel Antennae not as above 4 4. Body very slender (about 0.4 mm. broad); head and pronotum longer than broad; abdomen only slightly broader than elytra; Africa. Termitolinus Wasmann Body appreciably broader; abdomen moderately inflated and distinctly broader than foreparts 5 5. Abdomen with inner paratergites forming oblique margins one-fourth to one-half as broad as the tergites 6 Abdominal inner paratergites somewhat less than one-fourth as broad as tergites 7 216 FIELDIANA: ZOOLOGY, VOLUME 40 6. Inner paratergites one-fourth as broad as tergites; Indomalaya. Termitobaena Bernhauer Inner paratergites one-half as broad as tergites; Africa. Termitotropha Wasmann 7. Pronotal margins strongly deflexed; pronotal hypomera not visible in lateral view; Africa Termitotecna Wasmann Pronotal hypomera visible in lateral view 8 8. Elytral apices modified, i.e., the outer apical angles separated from elytral disk by a short longitudinal incision; Africa Protermitobia, new gen. Elytral apices not modified 9 9. Pronotum unmargined; abdomen broadly inflated and strongly membranous; Africa Termitopaedia Wasmann Pronotum margined; abdomen moderately broad, but with little membranous integument exposed; Indomalaya Dioxeuta Sharp TERMITOPAEDIA Wasmann Termitopaedia Wasmann, 1911, Rev. Zool. Afr., 1: 114; Fenyes, 1920, Gen. Insect., 173B: 143. Type species: Termitopaedia kohli Wasmann. Head generalized, subequal in length and width; clypeus moderately declivous. Eyes small, one-third as long as head. Gula broad and long; mentum free. Second segment of maxillary palpi small, relatively slender, arcuate: third segment fusiform, two-thirds longer than second. Antennae geniculate; scape short, very thin in profile, its inner surface concave; segments 2-10 decreasing in length, incrassate. Pronotum small, moderately convex, unmargined; slightly broader than long, broadest in front of middle; side margins sinuate. Elytra unmodified. Wings present. Abdomen strongly inflated, broadest at level of fourth segment, two and one-half times as broad as pronotum. Abdomen with considerable mem- branous integument exposed; the sclerites of each segment contiguous, but the sclerites of segments 3, 4, 5, and 6 well separated from those of adjacent segments. Inner paratergites moderately broad, those of fifth segment one-seventh as broad as tergite; outer paratergites relatively broad, those of fifth segment one-third as broad as tergite. Termitopaedia kohli Wasmann Termitopaedia kohli Wasmann, 1911, Rev. Zool. Afr., 1: 115, pi. 5, fig. 8 (Stanleyville, Belgian Congo; Pseudacanthotermes spiniger Sjostedt; Was- mann coll.); Hegh, 1922, Les Termites, p. 613, fig. 420. Material examined. Belgian Congo: Stanleyville. Two speci- mens in Wasmann collection on pin bearing type label, and several additional specimens. Color testaceous. Head very sparsely and minutely punctulate, glabrous; with one pair of widely separated vertexal setae. Pronotum feebly and micro- scopically punctulate, glabrous; 6 setae on apical margin, 2 on lateral margins, and 2 on disk. Elytra with a marginal row of 3 setae, a submarginal row of 2, and 1 seta near inner margin. Tergites and sternites with an apical row of setae, but otherwise with very few hairs. Relative lengths of antennal segments: SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 217 11 : 8 : 9 : 8 : 8 : 7 : 5 : 6 : 6 : 6 : 14. Relative lengths of parts of hind leg: femur 60; tibia 54; tarsus 31; tarsomeres 8, 4, 4, 5, 10. Middle leg: femur 48; tibia 46; tarsus 23. Length, 2.5 mm. DIOXEUTA Sharp Dioxeuta Sharp, 1899, Ent. Monthly Mag., (2), 10: 205; Fenyes, 1920, Gen. Insect., 173B: 267. Type species: Dioxeula microps Sharp. Jacobsonella Silvestri, 1910, Boll. Lab. Zool. Portici, 5: 59; Fenyes, 1920, Gen. Insect., 173B: 276; Franssen, 1932, Tijd. Ent., 75, suppl., p. 170; Cam- eron, 1939, Fauna Brit. India, Staphyl., 4: 495. Type species: Jacobson- ella termitobia Silvestri. New synonymy. Distinguished from Termitopaedia by its margined pronotum and somewhat longer antennae, but chiefly by its scaphoidal, much more heavily sclerotized abdomen, with scarcely any membranous integument exposed. Head subequal in length and width; clypeus declivous. Eyes small, a little more than one-fourth as long as head. Gula broad and long; mentum free. Maxillary palpi long and very slender; second segment about two-fifths as broad as long; third segment spindle-shaped, three times as long as broad. Pronotum with a distinct raised margin; disk uniformly convex; broadest in front of middle; lateral margins sinuate; apex feebly emarginate; base arcuate. Pronotal hypomera visible in lateral view. Elytra two-fifths broader than pronotum ; lateral margins diverging behind humeri to the broadest point, about two-thirds the length of elytra. Middle coxae long, contiguous; mesocoxal acetabula large, shallow, faintly margined laterally; metasternum very short behind mesocoxae. Abdomen moderately enlarged, scaphoidal in form; heavily sclerotized; sclerites contiguous so that scarcely any membranous integument is exposed; paratergites form an elevated margin. Remarks. For more than fifty years there has apparently been no attempt to establish the identity of Sharp's Dioxeuta, and the species of this genus have been assigned to Silvestri's Jacobsonella. Sharp's description gave few clues to the identity of Dioxeuta, so that I was surprised upon examining the type of D. microps Sharp to find that Jacobsonella termitobia Silvestri is the same species. Of the specialists who dealt with Jacobsonella, Cameron was perhaps the only one who had ready access to Sharp's types. It is surprising that he did not detect the synonymy. Dioxeuta microps Sharp. Figure 33, a. Dioxeuta microps Sharp, 1899, Ent. Monthly Mag., (2), 10: 206 (Baram, Sarawak; Termes malayanus; British Museum, Natural History). Jacobsonella termitobia Silvestri, 1910, Boll. Lab. Zool. Portici, 5: 61, figs. I (1-10), II (1-9), III (3), (Samarang, Java; Macrotermes gilvus Hagen; 218 FIELDIANA: ZOOLOGY, VOLUME 40 Silvestri coll.); Wasmann, 1916, Zool. Jahrb. Syst., 39: 183, pi. IV, fig. 11; Kemner, 1925, Ent. Tidskr., 46: 122; Franssen, 1932, Tijd. Ent., 75, suppl., p. 170, text figs. 2, 4, 6; Cameron, 1939, Fauna Brit. India, Staphyl., 4: 497. New synonymy. Material examined. Sarawak: Baram (type). Java: Buitenzorg (cotypes of Jacobsonella termitobia in Wasmann and U. S. Nat. Mus. coll.); Gelangar; Poetjoeng; and Pamanoctlam (Cameron coll., ex Franssen). Sumatra: Songei Bamban (Wasmann coll.). Malaya: Parit Brintar, Malacca (Wasmann coll.). Philippines: Manila, Luzon (Chicago Nat. Hist. Mus., ex Light coll.). Host records. Macrotermes gilvus Hagen (Sarawak, Java, Su- matra, Malaya, Philippines). Franssen (1932, p. 170) reported one record with Odontotermes javanicus Holmgren at Buitenzorg, Java. Brown to reddish-brown; head and pronotum usually darker than elytra; abdomen usually reddish. Head, pronotum, and elytra smooth, with only traces of reticulation. Head with a pair of vertexal setae, a row of occipital setae, and a few hairs behind eyes. Pronotum with 2 or 3 setae on lateral margins, 4 on anterior margin, and 5 on each half of pronotal disk. Elytra with about 20 setigerous asperities, irregularly placed. Abdomen with a moderately dense vestiture of recumbent and semi-recumbent hairs, but without strong macro- setae. Pronotum slightly less than one-half broader than long. Relative lengths of antennal segments: 20 : 8 : 12 : 14 : 12 : 11 : 11 : 11 : 10 : 9 : 18. Segment 4 longer than either 3 or 5. Remarks. Silvestri based Jacobsonella termitobia on a series that included stenogastric (ovigera form) and physogastric (gestans form) individuals and identified the latter as gravid females. Frans- sen (1932) concluded that the "femina gestans" individuals belong to a different species which he named J. oudemansi. He observed that the physogastric and stenogastric individuals differ appreciably in morphological characteristics, that there are males and females of each form, and that no intermediate forms are known. I concur with Franssen in the belief that two species are involved and go one step farther by proposing a new genus, Neodioxeuta, for Jacob- sonella oudemansi Franssen. Dioxeuta flavescens Cameron, new comb. Jacobsonella flavescens Cameron, 1939, Fauna Brit. India, Staphyl., 4: 496, pi. II, figs. 11, 12 (Insein, near Rangoon, Burma; in nests of termites; British Museum, Natural History). FIG. 33. (a) Dioxeuta microps Sharp, (b) Neodioxeuta oudemansi Franssen. (c) Protermitobia kirbyi Seevers. (d) Termitobaena bryanti Bernhauer. (e, f) Termitobia burgeoni Cameron; lateral and dorsal views. 219 220 FIELDIANA: ZOOLOGY, VOLUME 40 Material examined. Burma: Insein (type). Very similar to microps in most respects, but differing in its paler coloration and in the proportions of its antennal segments. Color pale rufo-flavate; head and abdomen with more of a reddish tinge, pronotum less so; elytra very pale brown, with scarcely a reddish tinge. Relative lengths of antennal segments: 21 : 12 : 14 : 15 : 12 : 11 : 10 : 9 : 9 : 8 : 20. Anten- nae four times as long as pronotum: antennal length 170, pronotal length 42. In microps, antennae three and three-fourths times as long as pronotum: antennal length 150, pronotal length 40. Remarks. Cameron's "female" allotype of this species is herein made the holotype of a new species, Neodioxeuta burmae, and assigned to a different genus. Following Silvestri's example with respect to Jacobsonella termitobia, Cameron presumed that a strongly physogastric individual found in the same nest with Jacobsonella flavescens was the gravid female of the species. This interesting situation is discussed more thoroughly under the genus Neodioxeuta. Dioxeuta indosinensis Silvestri, new comb. Jacobsonella indosinensis Silvestri, 1946, Boll. Lab. Ent. Agraria Portici, 6: 317, fig. IV (1-13), (Yen Bay, Tonkino, Indo-China; Macrotermes malac- censis Haviland; Silvestri coll.). This species has not been available for study; according to Silvestri it may be distinguished from the other species by its distinctive chaetotaxy of head, pronotum, and elytra and its sub- equal second and third antennal segments. Silvestri's detailed figures should be adequate for identifying this species. NEODIOXEUTA, new genus Type species: Neodioxeuta oudemansi Franssen (= Jacobsonella oudemansi Franssen). Related to Dioxeuta as indicated by the structure of head and thorax but distinguished from that genus by its very distinctive abdomen (fig. 33, 6). Although the genus differs in specific details, the head and thorax are as in Dioxeuta. Abdomen very strongly inflated, more than four times as broad as thorax. Basal segments of abdomen most strongly inflated, the second and third segments protruding over thorax and pushing elytra into a vertical position; second and third tergites invisible from above. Tergites much broader than in Dioxeuta due to "secondary sclerotization" lateral to the "tergite proper," i.e., the setose central area. Area of membranous integument exposed variable; rather extensive in oudemansi, which has small paratergites, much less in burmae, which has large paratergites. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 221 Neodioxeuta oudemansi Franssen, new comb. Figure 33, 6. Jacobsonella termitobia Silvestri (female gestans form), 1910, Boll. Lab. Zool. Portici, 5: 61, pi. Ill (1-2). Jacobsonella oudemansi Franssen, 1932, Tijd. Ent., 75, suppl., p. 171, text figs. 1, 3, 5 (Gedangan, Java; Macrotermes gilvus Hagen). Material examined. Java: Gedangan (Cameron coll., British Museum, Natural History). Light-brown; tergites light reddish-brown. Head with one pair of setae between antennae, a semicircle of 5 setae on vertex behind eye level and a semi- circle of 4 setae on occiput. Relative lengths of antennal segments: 20 : 13 : 11 : 14 : 10 : 10 : 10 : 10 : 9 : 9 : 20; segment 4 distinctly longest of segments 3-10. Head subequal in length and width (42); pronotum one-fourth broader than long (52 : 42). Proportions of tergites as follows (length: total width: width of setose area): third (40 : 120 : 60), fourth (40 : 130 : 65), fifth (40 : 120 : 65), sixth (35 : 100 : 62). Proportions of inner paratergites (length : width): third (26 : 8), fourth (30 : 8), fifth (30 : 9), sixth (28 : 8). Remarks. Silvestri (1910a) believed specimens of this species to be gravid females of his Jacobsonella termitobia. Franssen pointed out that there was no evidence for this hypothesis and much to be said against it. There are many morphological differences that cannot be accounted for on the basis of post-imaginal development. There is no doubt that Dioxeuta and Neodioxeuta evolved from the same stock, but Neodioxeuta has undergone a remarkable series of abdominal changes. It is my opinion that these changes are so significant as to require generic separation. Neodioxeuta burmae, new species Distinguished from oudemansi by the following characteristics: pale coloration, head almost one-third broader than long, pro- portions of antennal segments, and marked differences in size of paratergites and tergites. Head, antennae, and pronotum pale flavate, elytra very pale, abdominal sclerites light rufo-flavate (for coloration and general appearance of this species, see Cameron's colored figure, 1939, pi. II, fig. 12). Chaetotaxy of head and pro- notum very similar to that of oudemansi. Relative lengths of antennal segments: 18 : 13 : 13 : 8 : 8 : 8 : 7 : 7 : 7 : 7 : 17. First two antennal segments a little broader than segments 3-11, which do not vary appreciably in width; segment 4 somewhat shorter than 3 and subequal to 5. Head almost one-third broader than long (46 : 36); pronotum one-fourth broader than long (48 : 38). Proportions of tergites (length : total width : width of setose area): third (40 : 82 : 75), fourth (40 : 130 : 78), fifth (40 : 140 : 80), sixth (30 : 140 : 80), seventh (40 : 80 : 55). Proportions of inner paratergites (length : 222 FIELDIANA: ZOOLOGY, VOLUME 40 width): third (42 : 23), fourth (40 : 28), fifth (32 : 26), sixth (24 : 16), seventh (22 : 6); paratergites much broader than in oudemansi. Seventh tergite lobated medially. Length, 2.8 mm.; width of abdomen, 1.75 mm. Holotype from Insein, Burma, collected February, 1914, by T. B. Fletcher, with termites. In collection of British Museum (Natural History). Remarks. The holotype of this species was the "female" type of Cameron's Jacobsonella flavescens. Cameron's presumption that this individual is a physogastric female of Dioxeuta flavescens was quite unwarranted; he was only following Silvestri's precedent with respect to Jacobsonella termitobia. It is difficult to understand how Cameron could have compared the "male" and "female" types of Dioxeuta flavescens very carefully and yet concluded that they are conspecific. They differ in numerous respects: the head of Dioxeuta flavescens is subequal in length and width, that of Neodioxeuta burmae is one-third broader than long; the relative lengths of antennal segments are very different (compare ratios given under the descriptions of the two species) ; the abdomens are, of course, very different in numerous respects (degree of physo- gastry, very different proportions of tergites, paratergites, and sternites, and chaetotaxy). TERMITOPULEX Fauvel Termitopulex Fauvel, 1899, Rev. d'Ent., 18: 37; Fenyes, 1920, Gen. Insect., 173B: 290. Type species: Termitopulex grandicornis Fauvel. Silvestrinus Bernhauer, 1932, Boll. Lab. Zool. Portici, 26: 14. Type species: Silvestrinus erythraeanus Bernhauer. New synonymy. Termitopulex is the African counterpart of the Indomalayan Dioxeuta. The two genera are very closely allied, Termitopulex being distinguished by its extremely long antennae, which reach to the seventh tergite, and by its very long and conspicuous abdominal bristles. Additional material from nests of the fungus-growing termites may bridge the gap between the two groups of species. Termitopulex grandicornis Fauvel Termitopulex grandicornis Fauvel, 1899, Rev. d'Ent., 18: 38 (Abyssinia; with termites; Brussels Museum); Wasmann, 1912, Zeitschr. wiss. Zool., 101: 87; Reichensperger, 1915, Ent. Mitteil., 4: 124. Silvestrinus erythraeanus Bernhauer, 1932, Boll. Lab. Zool. Portici, 26: 15 (Nefasit, Eritrea; no host given; Chicago Natural History Museum). New synonymy. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 223 Material examined. Ethiopia: Holotype (Fauvel coll.). Eritrea: Nefasit (holotype and paratype of S. erythraeanus Bernhauer). Host record. Macrotermes bellicosus Smeathman (Harar, Ethio- pia; Reichensperger, 1915). Dark testaceous. Head, pronotum, and elytra sparsely and finely punctulate, shining; not pubescent and not reticulated; with a moderate number of fine, pale hairs. Pronotal disk with 10 moderately long, stout setae; elytra with one pair of sublateral setae, and one sub-basal pair. Tergites 3-6 with an apical row of 6 very long, stout macrosetae (dark basally, pale-tipped); tergite 3 with 2 and tergites 4-6 with 4 additional setae near middle of tergite. Tergites with a number of shorter, finer setae. Sternites with a conspicuous vestiture of long, dark setae. Head very slightly longer than broad; pronotum one-fourth broader than long. Pronotal apex shallowly emarginate; apex four-fifths as broad as base; lateral margins arcuate, becoming sinuate near base; base arcuate. Antennae capable of reaching to apex of seventh tergite; all segments at least twice as long as broad and usually more; relative lengths and widths: 24 (9) : 10 (5) : 19 (5) : 18 (5) : 18 (6) : 18 (7) : 18 (7.5) : 17 (8) : 18 (8) : 17 (8.5) : 28 (10). Basal antennal segment subfusiform; segments 3-6 very slender; 7-11 incrassate; 3-10 narrow at base and thickened apically; segment 11 ovoidal. Remarks. Bernhauer's Silvestrinus erythraeanus is a synonym of this species. Termitopulex natalensis Wasmann Termitopulex natalensis Wasmann, 1912, Zeitschr. wiss. Zool., Itl: 87 (Natal, alt. 4,000 ft.; Macrotermes natalensis Haviland; Wasmann coll.). Material examined. Natal: Holotype in Wasmann collection. Head, pronotum, and elytra very minutely punctulate; somewhat dull; not pubescent or reticulated. Head, pronotum, and elytra with a sparse vestiture of very fine, pale hairs, longer and more conspicuous on sides of elytra. Abdomen with numerous long and moderately long bristles (with a more conspicuous array of pale, fine hairs than in grandicornis). Pronotum slightly less than one- half broader than long (58 : 40). Antennae of type specimen missing. Termitopulex sjoestedti Eichelbaum Termitopulex sjostedti Eichelbaum, 1908, in Sjostedt, Zool. Kilimandjaro-Meru Exped., 7: 91 (Mt. Kilimandjaro-Kibonoto-Tanganyika Territory, "in der Steppenzone und unteren Kulturzone, 1000-1300 meters"; Termes vad- schaggae Sjostedt). Material examined. Tanganyika: Ngerengere (one specimen, ex Reichensperger coll.). Host record. Microtermes vadschaggae Sjostedt (host of type). Light reddish-brown; margins of pronotum, apices of elytra, and basal areas of tergites lighter. Head with several short, fine setae on dorsum, the sides and 224 FIELDIANA: ZOOLOGY, VOLUME 40 ventral surface with moderately numerous, stronger macrosetae. Pronotum with 4 macrosetae on anterior margin and a pair of short setae between the medial pair; 1 macroseta on each lateral margin and 3 forming an equilateral triangle on each half of disk. Elytra with 15 or more macrosetae on disk and a moderate number on sides. Elytra densely, asperately punctate. Tergites with a conspicuous vestiture of macrosetae and finer hairs; apical row of tergites 3-6 usually consisting of 6 very long macrosetae (about two-thirds as long as tergite) interspersed with shorter setae. Head subequal in length and width (44 : 44) ; sides converging a little behind eyes. Pronotum (54 : 42) about one-fourth broader than long; apex emarginate, broadest in front of middle, side margins converging more in front than behind; apex only four-fifths (40) as broad as base (50); base arcuate. Antennae with following relative segmental lengths: 22 : 10 : 14 : 18 : 16 : 15 : 14 : 14 : 16 : 14 : 24 (total, 210). Segments 2 and 3 very slender, 4-11 in- crassate, segment 11 twice as broad as 3, segments 1-5 subcylindrical, 5-11 increasingly compressed; all segments elongated. Legs very long; relative lengths of parts of hind leg as follows: femur, 75; tibia, 62; tarsomeres, 19, 10, 8, 8, 16. Femora and tibiae compressed; tibiae very slender at base, twice as broad at apex; femora twice as broad as tibiae. TERMITOLINUS Wasmann Termitolinus Wasmann, 1911, Rev. Zool. Afr., 1: 97. Type species: Termito- linus natalensis Wasmann. Termitolinus is placed in the Termitopaediini with some hesi- tation. Specimens of the genus have not been available until very recently and the necessary comparative morphological studies pre- liminary to a proper placing of the genus have not been made. Although markedly in contrast with most genera by virtue of its very slender build and non-physogastric abdomen, Termitolinus has several qualities suggesting relationship to Termitopulex and Dioxeuta. The abdomens of these genera are basically similar although inflated to some degree in Termitopulex and Dioxeuta. Head one-third longer than broad (38 : 28) ; broadest at eye level, sides con- verging, the head narrowest at base. Eyes small, less than one-sixth as long as head. Head with infraorbital ridge. Pronotum one-fourth longer than broad (40 : 32); lateral margins bisinuate, the anterior one-half a little convex, the posterior one-half slightly concave; basal border feebly arcuate, the anterior border a little emarginate; basal angles slightly obtuse; pronotal hypomera completely visible; pronotum distinctly margined. Elytra subequal in length. Antennae relatively short, not capable of extending to base of pronotum; geni- culate, scape short; segments 2 and 11 a little longer than broad, 3-10 strongly incrassate, 4-10 broader than long. Gula moderately broad at base, its sides converging in front. Abdomen slender, not physogastric, only a little broader at fourth and fifth segments than elytra; tergites feebly convex; paratergites slender. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 225 Termitolinus natalensis Wasmann Termitolinus natalensis Wasmann, 1911, Rev. Zool. Afr., 1: 98, pi. 5, fig. 11 (Shivyre, Natal; Macrotermes natalensis Haviland; Wasmann coll.). Material examined. Type and five specimens in Wasmann col- lection. Head light-brown; pronotum darker, slightly reddish-brown ; elytra testaceous; abdomen light reddish-brown. Head smooth at center of vertex; a few moderately large punctures and fine setae laterally. Pronotum densely covered with coarse, elongated punctures, except for a narrow median area and a narrow basal area; pronotum clothed with fine, short hairs and a few longer setae. Abdomen with a moderate vestiture of fine, pale hairs and a few longer setae. Length, '2 mm.; width, 0.35-0.4 mm. TERMITOBAENA Bernhauer Termilobaena Bernhauer, 1915, Verh. Zool. Bot. Ges. Wien, 65: 155. Type species: Termitobaena bryanti Bernhauer. Termitobaena and the following African genus, Termilotropha, may not belong to the Termitopaediini but I have found no adequate basis for separating them. Their most distinctive feature is probably the broad, scaphoidal abdomen, with its wide lateral "margins" formed by the inner paratergites. Otherwise, these genera have no outstanding features to distinguish them. Head generalized, clypeus moderately declivous and moderately long. Gula narrow, only one-eighth as broad as head, scarcely converging in front. Eyes small. Pronotum with anterior margin slightly arcuate, broadest in front, sides converging slightly near base, basal angles broadly rounded, base straight, surface evenly convex. Elytra generalized. Abdomen somewhat recurved over foreparts; its dorsum almost circular, somewhat scaphoidal in appearance inasmuch as the very large inner paratergites are strongly elevated to form a broad "margin"; inner paratergites about one-fourth as broad as tergites. Eighth tergite unusually long. Relative lengths of tergites: third, 3; fourth, 8; fifth, 10; sixth, 11; seventh, 14; eighth, 22. Termitobaena bryanti Bernhauer. Figure 33, d. Termitobaena bryanti Bernhauer, 1915, Verh. Zool. Bot. Ges. Wien, 65: 156 (Sarawak; with termites; Chicago Natural History Museum). Material examined. The type series. In addition, two speci- mens, Mt. Matang, W. Sarawak (Cal. Acad. Sci. coll.). Head shining, nearly glabrous, with a few fine punctures; vertex with a few short, black setae. Pronotum with an apical row of 8-10 bristles and a few on disk; surface smooth, almost impunctate. Elytra smooth, shining, with few traces of sculpture; disk with scattered bristles; lateral margin with 3 or more setae. Tergites shining, weakly sculptured except that the eighth is closely, longitudinally strigulose. Relative widths of representative levels of body as 226 FIELDIANA: ZOOLOGY, VOLUME 40 follows: head, 48; pronotum, 52; elytra, 60; abdomen (maximum width), 95. Head one-fifth broader than long. Pronotum almost one-third broader than long. Abdomen a little broader than long. Relative lengths of antennal segments: 15 : 8 : 8 : 7 : 7 : 7 : 7 : 7 : 8 : 8 : 17. TERMITOTROPHA Wasmann Termitotropha Wasmann, 1899, Deuts. Ent. Zeitschr., 43: 178; Fenyes, 1920, Gen. Insect., 173B: 291. Type species: Termitotropha o'neili Wasmann. Foreparts slender; abdomen broad, scaphoidal in form. Head generalized, very slightly broader than long, its sides almost straight, and its base rounded; clypeus short, feebly declivous; eyes small. Pronotum one-fourth broader than long, apex almost straight, base arcuate, broadest in front of middle, sides con- verging in front and behind, basal angles rounded; disk ovally impressed at center, the impression with two elongated foveae. Elytra generalized, their apices almost straight. Abdominal sclerites feebly separated or contiguous; abdomen with very little non-sclerotized integument exposed. Tergites moderately broad, approximately the same width as pronotum; paratergites forming a very broad, elevated margin which at level of segments 4, 5, and 6 is at least one-half as broad as tergites; inner paratergites especially large, comprising about three- fourths of margin. Relative widths of representative levels of body as follows: head, 48; pronotum, 52; elytra, 60; third abdominal segment, 90; fifth segment, 103; seventh segment, 85. Remarks. The single available topotypic specimen is mounted on a card and the under side is not visible. Inasmuch as the specimen is in a fragile condition, it was not removed for a detailed study of the ventral structures. Termitotropha o'neili Wasmann Termitotropha o'neili Wasmann, 1899, Deuts. Ent. Zeitschr., 43: 179, pi. II, fig. 9, a-c (Dunbrody, Union of South Africa; Amitermes unidentatus Sjostedt; Wasmann coll.). Material examined. One specimen, Dunbrody, Union of South Africa, collected January 5, 1899; probably from the type series (Bernhauer coll., Chicago Nat. Hist. Mus.). Head, pronotum, elytra, and abdomen sub-impunctate; seventh tergite reticulated. Very sparsely and inconspicuously setose; vertex of head with 6 or more fine, short hairs; pronotum with several short marginal setae near apical angles, one or more on lateral margins and one at basal angle; elytra with a few fine, marginal hairs, and several scattered setae on disk; tergites with marginal rows of inconspicuous hairs and a few scattered elsewhere. TERMITOBIA Wasmann Termitobia Wasmann, 1891, Verh. Zool. Bot. Ges. Wien, 41: 647; 1912, Zeitschr. wiss. Zool., 101: 82; Fenyes, 1920, Gen. Insect., 173B: 289. Type species: Termitobia physogastra Wasmann. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 227 Distinguished from the other genera of the tribe by its remark- able abdomen. Robust species of distinctive facies (fig. 33, e). Abdomen greatly distended, in a horizontal position, and not recurved over foreparts. Abdominal sclerites large, heavily sclerotized, not widely separated from one another; very little membranous integument exposed. Abdominal segments 3, 4, and 5 greatly enlarged; anteriorly expanded to cover much of the thorax, which is scarcely visible from above. Tergite 3 extremely large, yet visible in dorsal view only where its apical margin curves over the anterior margin of the ovate abdomen. Tergites 4-8 large, scarcely overlapping, covering the very convex dorsal surface of the abdomen. Ventral surface of abdomen almost flat. Paratergites large; arcuate inner paratergites cover the lateral margin of the abdomen and are visible from above or below. Termitobia physogastra Wasmann. Figure 34, 6, c. Termiiobia physogastra Wasmann, 1891, Verh. Zool. Bot. Ges. Wien, 41: 649, pi. VI, figs. 1-15 (Akuse, Volta River region, Gold Coast; Macrotcrmes bellicosus Smeathman; Wasmann coll.); 1912, Zeitschr. wiss. Zool., 101: 82, pi. 5, fig. 1; Gridelli, 1928, Ann. Mus. Civ. Nat. Genova, 52: 410; Hegh, 1922, Les Termites, p. 611, fig. 411. Termitobia gastrophysa (misspelling) Eichelbaum, 1908, in Sjdstedt, "Kili- mandjaro-Meru Exp.," 7: 92. Material examined. The type (Gold Coast) in Wasmann col- lection. Color reddish-brown, pronotum black. Head with a semicircular row of about 8 vertexal setae, the inner ones farther apart than in T. burgeoni. Pronotum with about 18 moderately long hairs on anterior margin, 4 on lateral margin; discal hairs sparse but moderately long. Pronotum strongly shining; with circular areas of reticulated integument around each seta. Tergites with numerous elevated areas, each bearing a seta and reticulated integument. Form of head as in figure 34, b; eyes relatively small, the sides of head con- verging very strongly behind eyes. Head two-fifths broader than long (115 : 80). Antennal segments of the following relative lengths: 32 : 34 : 34 : 22 : 24 : 22 : 21 : 20 : 20 : 18 : 30; little variation in width. First three antennal segments three times as long as broad; subequal in length. Pronotum very strongly convex; with a broad, moderately deep impression on each side of middle. Third tergite distinctly separable into two areas: a much enlarged non-setose area, invisible from above, and a narrow arc, visible from above, bearing setigerous tubercles. The visible arc-like area has a non-sclerotized medial gap that gives it the appearance of two sclerites. Inner paratergites broad (almost twice as broad as in burgeoni), their relative widths as follows: 3 (70), 4 (80), 5 (80), 6 (70), 7 (40); outer paratergites: 3 (absent), 4 (30), 5 (28), 6 (16), 7 (fused to sternite). Remarks. This species has been recorded from a number of African localities but the identifications were probably incorrect in most instances. The Rhodesian record is certainly erroneous; 228 FIELDIANA: ZOOLOGY, VOLUME 40 the specimens on which this record was based belong to a new species herein described as T. rhodesiae. It is doubtful if the Transvaal specimens (Gridelli, 1928) collected with Macrotermes swaziae Fuller, the Tanganyika material (Eichelbaum, 1908) from nests of Macro- termes goliath Sjostedt, or the Groot-Fontain, Damaraland, South West Africa specimens (Wasmann, 1912) collected with Macro- termes natalensis Haviland are conspecific with T. physogastra. FIG. 34. (a) Termitobia rhodesiae Seevers, head, (b, c) Termitobia physogastra Wasmann, head and elytron. Termitobia rhodesiae, new species. Figure 34, a. Distinguished from T. physogastra by form of head, larger eyes, pronotal contours, differences in relative lengths of antennal seg- ments, and minor differences in chaetotaxy. Head, pronotum, elytra, and under side blackish, with tinges of red; foreparts of abdomen above brown or dark reddish-brown, becoming reddish black pos- teriorly. Not much membranous integument visible. Coloration in general darker than in physogastra. Pronotum with about 16 hairs on anterior margin and about 4 on lateral margins (shorter than in physogastra); discal hairs very sparse and short. Pronotal surface very dull. Tergites with elevated areas as in physogastra, but with the setae uniformly shorter and less conspicuous. Form of head distinctive (fig. 34, a); one-half broader than long (105 : 70). Eyes larger than in physogastra. Antennae with segments of the following relative lengths: 30 : 30 : 26 : 22 : 20 : 20 : 19 : 17 : 17 : 17 : 26; with little variation in width after the broad scape. Pronotum very strongly convex; with only faint traces of the surface im- pressions present in physogastra. Holotype. From Southern Rhodesia; collected by Dr. Brauns, with Macrotermes natalensis Haviland; in Wasmann collection, Natuurhistorisch Museum, Maastricht, The Netherlands. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 229 Paratype. One specimen, same data as type, in California Academy of Sciences. Termitobia burgeon! Cameron. Figure 33, e, f. Termitobia burgeoni Cameron, 1930, Rev. Zool. Afr., 19: 419 (Moto, on Uele River, Belgian Congo; Macrolermes natalensis Haviland; Cameron coll.). Material examined. One specimen, Niangara, Belgian Congo (Lang-Chapin Exp., 1910), with Macrotennes natalensis Haviland. A series of 25 specimens, Bendija, Liberia, with Macrotermes natal- ensis Haviland (W. M. Mann, Smithsonian Firestone Exp., U. S. Nat. Mus.). Head not as dorso-ventrally compressed as in physogastra; eyes smaller. Sides of head almost straight for a distance behind the eyes, then converging gradually to the base which is broader than in physogastra; head not triangular in outline. Head four-fifths as long as broad (65 : 80); in physogastra only seven- tenths as long as broad (55 : 78). Segment 2 of maxillary palpi not long and slender, medio-laterally compressed but not arcuate, broader at apex, its length less than twice its apical width; segment 3 almost twice as long as broad, some- what compressed, its inner face flattened, only a little longer than second, its apex a little broader than that of the second. Antennal segments with the following relative lengths: 36 : 32 : 24 : 19 : 18 : 16 : 16 : 15 : 14 : 14 : 24. Vertex of head with a semicircular row of 8 long, erect setae. Tergites setose but the setae not set in tubercles as in physogastra. Inner paratergites much smaller, as a rule, than in physogastra; outer paratergites larger and more distinct. Relative widths of inner paratergites: 3, 38; 4, 44; 5, 40; 6, 40; 7, 26; of outer paratergites: 3, 40; 4, 40; 5, 32; 6, 30; 7, 16. Remarks. The Liberia specimens differ in minor respects from the Belgian Congo examples studied, but little significance can be attributed to these at present. The pronotum has a median sulcus and lateral impressions that are not evident in the Belgian Congo specimens, the paratergites have more and longer bristles, and the abdominal bristles in general are longer and more conspicuous. Termitobia paolii Gridelli Termitobia paolii Gridelli, 1928, Ann. Mus. Civ. Nat. Geneva, 52: 403, text figs. (Villaggio Duca degli Abruzzi, Italian Somaliland; Macrotermes bellicosus Smeathman). TERMITOTECNA Wasmann Termitotecna Wasmann, 1912, Zeitschr. wiss. Zool., 101: 88; Fenyes, 1920, Gen. Insect., 173B: 348. Type species: Termitotecna braunsi Wasmann. Head short, transverse; eyes very large, prominent. Galea a little longer than lacinia but not large and foliate. Ligula bifid. Pronotum three-fifths broader 230 FIELDIANA: ZOOLOGY, VOLUME 40 than long; sides deflexed, hypomera not visible from side; pronotal apex broadly, shallowly emarginate; humeri depressed; lateral margins strongly arcuate, con- tinuously so with the strongly arcuate base; posterior angles obsolete. Elytra slightly longer than pronotum; apices with a slight indication of an incision just medial to the outer apical angles (this modification more pronounced in Pro- termitobia and Termitobia). Mesocoxal acetabula broad and shallow, distinctly margined laterally and caudally; mesosternal process short, with a median carina; metasternum behind middle coxae relatively short. Hind coxae moderately long; not produced lateral to articulation. Tarsi 4, 5, 5-segmented. Abdomen strongly physogastric (as in Protermitobia); abdomen about one-half broader than elytra at scutellum level. Tergites strongly convex, generalized; seventh a little longer than the others. Inner paratergites large, translucent; those of segments 4-6 about two-ninths as broad as tergites. Remarks. Termitotecna has always been assigned to the tribe Aleocharini on the basis of Wasmann's statement that the tarsi are 5, 5, 5-segmented. Recently I was able to verify a suspicion that the tarsi are 4, 5, 5-segmented and that the genus belongs here. Termitotecna braunsi Wasmann Termitotecna braunsi Wasmann, 1912, Zeitschr. wiss. Zool., 101: 89, pi. V, fig. 2, a-c (Bothaville, Orange Free State, Union of South Africa; Odonto- termes transvaalensis Sjostedt; Wasmann coll.). Material examined. Type and other specimens, Wasmann collection. Head and pronotum rufo- testaceous; elytra brown; tergites testaceous; paratergites pale translucent. Head and pronotum shining; punctulation and reticulation obsolescent. Pronotum with a sparse vestiture of fine, pale, erect hairs. Tergites glabrous, shining, very indistinctly reticulated; with apical rows of fine setae and a few on general surface. Relative lengths of segments of anten- nae: 20 : 8 : 10 : 8 : 8 : 10 : 10 : 10 : 10 : 9 : 28. Segment 3 longer than broad, segments 4-5 transverse, 6-9 subquadrate, segment 10 transverse, 11 more than twice as long as broad; antennae feebly incrassate from segment 2. PROTERMITOBIA, new genus Type species: Protermitobia kirbyi, new species. Protermitobia is most closely allied to Termitotecna but has much in common with stock from which Termitobia evolved. From Termitotecna it is distinguished by having the pronotal hypomera visible from the side; the pronotum not more than one-half broader than long; a broad, foliaceous galea; and an entire ligula. The abdomens of Protermitobia and Termitotecna are very similar. The head and thorax of Protermitobia and Termitobia are rather similar, but the abdomen of the latter is, of course, remarkably specialized. The very unusual modification of the elytral apices of Protermitobia SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 231 and Termitobia (fig. 34, c) is strongly indicative of the fact that they are of the same phyletic line. Robust in build, abdomen physogastric, but only moderately inflated. Head transverse, its dorsum generalized in appearance; sides converging strongly behind eyes, basal angles obsolete. Mandibles very strong, long, curved, piercing. Gula short and narrow, its sides converging so that in front it is only about one- ninth as broad as head. Submentum and mentum large, mentum apparently fused to submentum as no suture is visible. Labial palpi 3-segmented; segments elongated. Maxillae very similar to those of Termitobia; galea much longer and broader than lacinia, sclerotized at base, membranous distally, with outer apical margin sclerotized. Maxillary palpi with segment 2 compressed medio-laterally; segment 3 subcylindrical. Eyes small. Antennae moderately long, segments 3-10 subequal, slightly transverse, feebly obtrapezoidal ; eleventh segment as long as segments 8-10 combined. Pronotum two-fifths to one-half broader than long, surface moderately convex, its side margins perhaps feebly explanate; apex slightly bisinuate, apical angles broadly rounded, sides straight, base strongly arcuate, posterior angles rounded; hypomera entirely visible from sides. Elytra longer than pronotum; apical margins of elytra oblique, their apical angles produced and partially separated from disk by longitudinal incisions (somewhat as in Termitobia, fig. 34, c). Mesosternum and metasternum similar to those of Termilobia. Mesosternal process broad, margined; mesocoxae moder- ately separated; mesocoxal concavities very shallow, not margined. Metasternum behind coxae considerably longer than in Termitobia. Hind coxae shorter and broader than in Termitobia but not produced beyond lateral articulation. Metepi- sternum broader than in Termitobia. Tergites rather large, somewhat trans- lucent along margins in most cases; seventh tergite longer than others; posterior angles of tergites 3-5 rounded, their apical borders slightly emarginate; tergite 6 with medial two-thirds of apical margin strongly, arcuately produced. Para- tergites broad, only one on each side (indicating either fusion of inner and outer paratergites or the absence of one). Tarsi 4, 5, 5-segmented. Protermitobia kirbyi, new species. Figure 33, c. Ferrugineous, sternites and paratergites rufo-testaceous; outer apical angles of elytra flavous. Head, pronotum, and elytra very finely, very densely punctulate (fine asperities present), and in some areas very finely reticulated. Abdomen finely reticulated, the meshes not as close as on the foreparts. Vertex of head with about 18 fine, erect setae and other shorter, fine hairs; pronotum with about 15 fine hairs along anterior and lateral margins, about 10 on disk, and 6 or so along basal border; elytra with about 20 fine hairs; tergites with fairly numerous, fine hairs; sternites with a moderately dense vestiture of fine, erect hairs (mixed long and short) ; tibiae rather densely hairy, the hind tibiae with a posterior fringe of long, fine, curved-tipped hairs. Length, 4.2 mm. Holotype from Iringa, Tanganyika Territory; collected January 10, 1935, by Harold Kirby; with Cubitermes, n. sp. In collection of Chicago Natural History Museum. No paratypes. 232 FIELDIANA: ZOOLOGY, VOLUME 40 Protermitobia comes, new species Distinguished from kirbyi by the punctation of the head, the non-reticulated pronotum, and the pilosity of head and pronotum. Head and pronotum reddish-brown, elytra brownish, their outer apical angles flavate; abdominal sclerites light reddish-brown. Head with 40-50 moderately coarse, umbilicate punctures, each bearing a fine, pale, curved hair; surface reticulated. Pronotum with 14-16 moderately long, fine, curved hairs on anterior margin and one or two along apical half of lateral margins; disk with about 15 similar setae on each half, the pattern not regular. Pronotal surface not reticu- lated, but with exceedingly fine punctules that tend to dull the surface luster. Elytra reticulated, with about 8 fine, pale hairs on disk near base and a few others along deflexed elytral sides. Sternites with a moderately dense clothing of fine, pale, semi-recumbent hairs, which become sparse on the more apical segments. Length, 4 mm. Holotype and one paratype from Sierra Leone, collected July 8, 1924 (ex Imperial Bureau of Entomology); with Cubitermes sub- crenulatus Silvestri. In collection of Chicago Natural History Museum. Tribe MYRMEDONIINI (sens, lat.) The tribe Myrmedoniini of the Coleopterorum Catalogus (Bern- hauer and Scheerpeltz, 1926, pars 82, and Scheerpeltz, 1933-34, pars 129, 130) is a large group of approximately 175 genera. Students of the Aleocharinae have relegated almost all genera with 4, 5, 5- segmented tarsi to this category, so that it is certainly polyphyletic and should be subdivided as soon as possible. The separation of the Termitopaediini in this paper helps to reduce the heterogeneity. Several other termitophilous groups within the Myrmedoniini are recognized as subtribes pending more detailed studies; these are Termitondina, Termitozyrina, and Feldina. Bernhauer and Scheerpeltz divided the Myrmedoniini into five subtribes: Falagriae, Athetae, Schistogeniae, Hoplandriae, and Myrmedoniae. It is likely that each of these groups, with nu- merous modifications, will be elevated to tribal rank. The Myr- medoniae (Myrmedoniina of this paper) will be the basis for the Myrmedoniini (s. str.). This group should be restricted to Zyras, Drusilla, and their allies. Some termitophagous genera and a number of myrmecophilous genera are among these allies. Some doryline ant guests, but by no means all, are among the Myrmedoni- ina. Lomechusa, Atemeles, and Xenodusa belong in this category too. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 233 Subtribe MYRMEDONIINA Important subtribal characters are as follows: Maxillary sinuses extensive, larger than necessary to receive the maxillae, extending caudally beyond the cardo; galea and lacinia elongated, the former longer and usually very slender; middle coxae widely separated, mesosternal process broad and short, metasternal process broad and long; metepimera large, their posterior ends usually produced dorsad behind apices of elytra. In addition, the tarsi are 4, 5, 5-segmented, mentum free, infraorbital carina present or absent, terminal antennal segments without polytrichous coeloconic sensilla; hind coxae generalized. Although records are not available for a good many species, it is probable that a majority, if not all, of the species of the large genera, Zyras and Drusilla, are associated in some way with ant or termite colonies. Species of these genera are not modified in any discern- ible way for life with social insects and are probably poorly adjusted synecthrans. Many are large, robust species by aleocharine standards and resemble free-living predatory staphylinids. Cameron (1930a, p. 13) in a short discussion of ant and termite guests of India states that there are "species not living in the nests but in the vicinity and attacking the ant or termite in the open; amongst these is the large genus Zyras, and I have seen one seize an ant by the thorax and shake it much as a terrier does a rat." He states that "very shortly after exposing a comb from a termite's nest a number of species arrive which in my experience are never found otherwise; such are Quediosoma, Rhopalinda, Demerinda, Pelioptera, etc. If, however, the comb is removed to a zone above the limit of the termite, practically nothing is attracted to it until it is again brought into the region inhabited by the termites. These genera are then, I believe, in some way dependent on termites." ZYRAS Stephens Zyras Stephens, 1835, Illustr. Brit. Ent., Mandibulata, 5: 430; Bernhauer, 1928, Arch. Naturg., 92: 19; Cameron, 1939, Fauna Brit. India, Staphyl., 4: 497. Type species: Zyras haworlhi Stephens. As now constituted, Zyras is a very large genus of more than 300 species occurring in both hemispheres, especially in tropical areas. The only attempts to analyze this large complex have re- sulted in its subdivision into some 58 subgenera, most of which were proposed by Bernhauer. It is apparent that the genus has rather broad limits and numerous species groups, but the recognition of a large number of subgenera is not the best way to cope with such a complex. Zyras has been poorly defined and no satisfactory basis for distinguishing it from Drusilla Leach has been found. Cameron 234 FIELDIANA: ZOOLOGY, VOLUME 40 (1939, p. 498) states that Zyras is "very closely related to Astilbus (=DriLsilla), from which indeed it is doubtfully separable now that a wider knowledge of tropical forms is available, . . . authors indeed appear often to have referred their species to either Zyras or Astilbus, according to which genus the general facies most corresponded." Zyras is supposed to include those species that have the third segment of the labial palpi as long as or longer than the second segment, Drusilla those species that have the third segment shorter than the second segment. I have almost no acquaintance with the species of Zyras that have been recorded as inhabitants of termite nests, and can do no more than list those so recorded. It may be advisable to point out that Tottenham (The generic names of British insects, pt. 9, 1949) and Blackwelder (1955) agree that the species long known as Zyras collaris Paykull is the type species of Bolitochara Mannerheim and that as a result Zyras is a synonym of Bolitochara. There are, as Tottenham points out, very serious objections to substituting Bolitochara for Zyras. The large number of species involved, the substantial literature dealing with Zyras and its ecological relationships, and the fact that two large tribes are named after these genera would make this a confusing change. Tottenham petitioned the International Commission on Zoological Nomenclature to set aside all type designations for Bolitochara and to designate Aleochara pulchra Gravenhorst (1806) as type species, and to add Bolitochara Mannerheim and Zyras Stephens (type species, Aleochara haworthi Stephens, 1832) to the Official List of Generic Names in Zoology. Inasmuch as I concur completely with Tottenham's viewpoint, Bolitochara will not be substituted for Zyras, pending a decision by the Committee. LIST OF SPECIES OF ZYRAS RECORDED FROM TERMITE NESTS badius Motschulsky (Glossacantha),=affinis Kraatz. Ceylon; with termites (Kraatz, 1859, p. 23; Wasmann, 1894, p. 89; Cameron, 1939, p. 520). excisus Kraatz (Glossacantha) . Ceylon; with termites (Kraatz, 1857b, p. 51; Wasmann, 1894, p. 89; Cameron, 1939, p. 524). feae Wasmann (Rhynchodoma) . Burma; with Odontotermes feae K. and N. Holm- gren (Wasmann, 1896a, p. 620; Cameron, 1939, p. 506). flavtis Cameron (Termitodonia). Java (Cameron, 1936). gestroi Wasmann (Rhynchodonia) . Burma; with Coptotermes curvignathus Holm- gren (Wasmann, 1896a, p. 622; Cameron, 1939, p. 505). heimi Wasmann (Glossacantha) . India; with Odontotermes obesus Rambur (Was- mann, 1899a, p. 150; Cameron, 1939, p. 527). hoplonotus Kraatz (Glossacantha). Ceylon; in termite nest (Kraatz, 1857b, p. 49; Wasmann, 1894, p. 89; Cameron, 1939, p. 524). humerosus Motschulsky (Glossacantha). Ceylon; in termite nest (Motschulsky, 1859, p. 90; Wasmann, 1894, p. 90; Cameron, 1939, p. 522). SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 235 inclytus Wasmann (Ctenodonia). Sierra Leone; with Macrotermes bellicosus Smeathman (Wasmann, 1894, p. 208). insignis Cameron (Termilelia) . Ceylon; with termites? (Cameron, 1939, p. 517). leoninus Wasmann (Rhynchodonia) . Sierra Leone; with termites (Wasmann. 1897b, p. 279). motschulskyi Bernhauer and Scheerpeltz, = /ami HO/US Motschulsky (Rhynchodonia). Ceylon; with termites (Motschulsky, 1859, p. 87; Wasmann, 1894, p. 90; Cameron, 1939, p. 508). obscurus Fabricius (Glossacantha),=tridens Wasmann. Ceylon; with Odontotermes obesus Rambur (Wasmann, 1899a, p. 149; Cameron, 1939, p. 527). peringueyi Bernhauer and Scheerpeltz (Euryndonia), = termitobiii8 Peringuey. Rhodesia; in termite nest (Peringuey, 1904, p. 206). puncticollis Peringuey. Rhodesia; with termites (Peringuey, 1904, p. 211). reicherti Wasmann (Rhynchodonia). Orange Free State; with Odontotermes trans- vaalensis Sjostedt (Wasmann, 1912, p. 94). rhodesiae Bernhauer and Scheerpe\tz,=termitophilus Peringuey. Rhodesia; in termite nest (Peringuey, 1904, p. 206). rhodesiana Cameron. Rhodesia; with termites (Cameron, 1919, p. 85). rubricollis Kraatz (Glossacantha) . Ceylon; in termite nest (Kraatz, 1857b, p. 51; Wasmann, 1894, p. 90; Cameron, 1939, p. 522). seminigra Cameron (Zyras). Rhodesia; with termites (Cameron, 1919, p. 86). serraticornis Kraatz (Crateodonia). Ceylon; with termites (Kraatz, 1857b, p. 48; Wasmann, 1894, p. 90; Cameron, 1939, p. 515). termitarius Wasmann (Rhynchodonia). Burma; with termites (Wasmann, 1897a, p. 30; Cameron, 1939, p. 503). termitocolus Gestro (Rhynchodonia), = termitobia Wasmann. Burma; with Copto- termes curvignathus Holmgren (Gestro, 1886, p. 110; Wasmann, 1894, p. 90; Wasmann, 1896a, p. 622; Cameron, 1939, p. 501). termitophilus Wasmann (Rhynchodonia). Burma; with termites (Wasmann, 1897a, p. 28; Cameron, 1939, p. 502). termilovagans Bernhauer (Glossacantha). East Africa (Bernhauer, 1928, p. 38). Irisulcatus Wasmann. Burma; with Odontotermes feae K. and N. Holmgren (Wasmann, 1896a, p. 623; Cameron, 1939, p. 547). DRUSILLA Leach Drusilla Leach, 1819, in Samouelle, Entom. Compend., p. 177. Type species: Drusilla canaliculata Fabricius. Astilbus Dilwyn, 1829, Memoranda . . . Coleopt. Ins. Swansea, p. 63; Cam- eron, 1939, Fauna Brit. India, Staphyl., 4: 460. Myrmedonia Erichson, 1837, Kafer Mark Branden., p. 286. LIST OF SPECIES OF DRUSILLA RECORDED FROM TERMITE NESTS impressicollis Kraatz, = 8culpticollis Wasmann. Ceylon and India; with Odonto- termes wallonensis Wasmann (Kraatz, 1857b, p. 53; Wasmann, 1899a, p. 151; Cameron, 1939, p. 475). incola Fauvel. Burma; with termites (Fauvel, 1904, p. 63; Cameron, 1939, p. 463). planaticollis Kraatz. Ceylon and India; in termites' nests (Kraatz, 1857b, p. 52; Wasmann, 1894, p. 90; Cameron, 1939, p. 467). TERMOZYRAS Cameron Termozyras Cameron, 1930, Rev. Zool. Afr., 19: 420. Type species: Termo- zyras politus Cameron. 236 FIELDIANA: ZOOLOGY, VOLUME 40 Termozyras politus Cameron Termozyras politus Cameron, 1930, Rev. Zool. Afr., 19: 421 (Belgian Congo; with Macrotermes natalensis Haviland). URODONIA Silvestri Urodonia Silvestri, 1946, Boll. Lab. Ent. Agr. Portici, 6: 318. Type species: Urodonia notabilis Silvestri. Urodonia notabilis Silvestri Urodonia notabilis Silvestri, 1946, Boll. Lab. Ent. Agr. Portici, 6: 321, text figs. 5, 6, 7 (Yen-Bay, Tonkino, French Indo-China; Odontotermes malac- censis Holmgren; Silvestri coll.). Subtribe FELDINA As a temporary measure I am assigning three Indomalayan genera to a subtribe of Myrmedoniini. Felda Blackwelder (=Asticta Wasmann), Zunia Blackwelder (=Disticta Wasmann), and Termi- tobiella Wasmann comprise a group of Caprilermes guests. Until very recently these genera have not been available to me and there has not been sufficient time to make detailed studies of them. Felda and Zunia are probably closer than Wasmann implied, and both categories may not be necessary. Wasmann provided excellent illustrations and descriptions, and there should be no great difficulty in recognizing the genera. The three genera are physogastric but not strongly so; their abdomens are strongly sclerotized and do not exhibit large areas of membranous integument. ZUNIA Blackwelder Disticta Wasmann, 1916, Zool. Jahrb. Syst., 39: 184. Type species: Disticta capritermitis Wasmann. Zunia Blackwelder, 1952, Bull. U. S. Nat. Mus., 200: 409 (new name for Disticta Wasmann, preoccupied). Zunia may be incompletely characterized as follows: Head short, transverse; antennae short, compact, basal segment elongated, segments 2 and 3 subquadrate, 4-10 transverse, slightly incrassate, apparently slightly telescoped. Pronotum short, transverse, almost twice as broad as long, broader than head, anterior margin straight, sides rounded, converging toward base, basal margin slightly arcuate, disk deeply, broadly bifoveolate. Abdomen very broad, but apparently feebly physogastric; sclerites large and only small areas of non-sclerotized integu- ment exposed. Inner paratergites very large. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 237 Zunia capritermitis Wasmann Disticta capritermitis Wasmann, 1916, Zool. Jahrb. Syst., 39: 185, pi. 4, fig. 12, pi. 5, fig. 12 (Tandjong Slamat, Sumatra; Capritermes minor Holmgren; Wasmann coll.). Black, anterior part of head flavate, abdomen piceous, shining, impunctate; elytra and lateral margin of abdomen setulose. Length, 2 mm. Wasmann figured antenna, mandible, maxilla, labial palpi, prementum, and anterior and middle legs, but the figures are too sketchy to be of use for comparative purposes. FELDA Blackwelder Asticta Wasmann, 1916, Zool. Jahrb. Syst., 39: 185. Type species: Asticta butteli Wasmann. Felda Blackwelder, 1952, Bull. U. S. Nat. Mus., 200: 165 (new name for Asticta Wasmann, preoccupied). Wasmann distinguished this genus from Zunia by these char- acters: pronotum not foveolate, but indistinctly bi-impressed ; terminal abdominal segments not conically produced; antennae longer, reaching middle of elytra; longer legs; distinctive mandibles, maxillae, and labial palpi. Wasmann's sketches of the mouth parts of this genus do not provide convincing evidence that it is different from Zunia. The impression of the pronotum is probably a matter of degree, and the abdominal character of little value. The antennae of this genus are longer and more loosely organized and the abdomen appears more physogastric. Felda butteli Wasmann Asticta butteli Wasmann, 1916, Zool. Jahrb. Syst., 16: 186, pi. 4, fig. 13, pi. 5, fig. 13 (Gap, Selangor, British Malaya; Capritermes nemorosus Haviland; Wasmann coll.). Head and pronotum piceous to nigro-piceous, shining and impunctate. Elytra piceous, opaque, densely clothed with erect setae; elytral humeri dilated and densely rugulose; disk slightly impressed. Abdomen piceous, semi-shining, punctate, sides and venter testaceous. Antennal segments 3-10 subquadrate. Length, 2 mm. TERMITOBIELLA Wasmann Termitobiella Wasmann, 1916, Zool. Jahrb. Syst., 39: 187. Type species: TermiiobieHa setipes Wasmann. Wasmann states that this genus is distinguished from Felda by its more slender legs, more slender antennae, which reach to the 238 FIELDIANA: ZOOLOGY, VOLUME 40 apices of the elytra, more densely setose legs, different head, and pronotal, elytral, and abdominal forms. These differences are not obvious from Wasmann's photographs. Termitobiella setipes Wasmann Termitobiella setipes Wasmann, 1916, Zool. Jahrb. Syst., 39: 187, pi. 4, fig. 14 (Gap, Selangor, British Malaya; Capritermes nemorosus Haviland; Was- mann coll.). Head, pronotum, and elytra black, very shiny, glabrous; abdomen rufo- piceous, glabrous above, impunctate. Length, 2.3 mm. Subtribe TERMITONDINA This subtribe is proposed for a genus of African termitophiles associated with Macrotermes. The species are limuloid in form. It is very difficult to place this genus accurately in view of the confusion existing in the Myrmedoniini. TERMITONDA, new genus Type species: Termitonda liberiae, new species. Head much broader than long, sides broadly rounded into base; clypeus moderately declivous; antennal fossae opening cephalad from under a slight vertexal arcade extending across front of head. Gula very broad, its base two- fifths as broad as head, its sides converging in front so that apex is a little more than one-half as broad as base. Submentum short and broad, only a little more than one-third as long as broad (maximum width equal to that of gular base); mentum very short, only three-fourths as long as submentum; ligula filiform. Labial palpi long and slender; 3-segmented, first and third segments elongated, second short. Galea membranous, foliaceous; lacinia shorter, with a comb of setae. Antennae moderately long, incrassate. Pronotum one-half broader than long; strongly convex; base bisinuate; apex deeply emarginate; sides arcuate, converging in front; hypomera broad, invisible from side. Elytra broad and short, sutural length about four-fifths the length of pronotum; epipleurae transverse, visible from below. Prosternum and meso- sternum large; mesosternal process extending for about two-thirds of length of coxae; metasternal process much shorter, acute. Metasternum behind middle coxae moderately large. Hind coxae very large; metepimera large, broadly produced apically. Tarsi 4, 5, 5-segmented. Abdomen subconical, narrowing to a slender apex; ninth tergite deeply incised, thus bearing two slender processes; abdomen densely setose, the apex of each tergite and sternite bearing a row of strong, black bristles. Termitonda liberiae, new species. Figure 36, a. Color reddish-brown. Head, pronotum, and elytra sub-impunctate (extremely minute punctules may be present) and without pubescence; shining. Head with SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 239 a pair of clypeal setae near apex, a seta medial to each antennal fossa, and a small group of short setae medial to each eye. Pronotum with a subapical row of 4 setae, a submarginal row of 4 setae, and 4 discal setae in a transverse row behind the middle. Elytral margin with 3 or 4 stout, laterally directed setae and 3 or 4 finer, erect setae; elytral disk with three transverse rows of 3, 1,4 setae, respec- tively; epipleura with a conspicuous row of about 12 pale setae. Tergites with numerous recumbent setae and the following black setae: tergites 3-7 with an apical row of 4, tergite 8 with 6 apical setae and 4 in a semicircle behind the middle; tergite 9 with 5 on each half. Sternites with 12-14 strong black setae in apical row interspersed with finer, pale bristles; in addition, densely clothed with short hairs. Antennae relatively short, scarcely reaching base of pronotum; its segments with the following relative lengths: 14:6:4:3:4:4:4:4:4 :4 :8. Antennae geniculate, segment 2 a trifle less than one-half as long as scape; segment 3 slender, slightly elongated, only two-thirds as long as second; segments 5-10 subequal in length, incrassate; segment 4 smallest, subquadrate. Pronotum one-half broader than long. Length, 1.75-2 mm. Holotype from Bendija, Liberia; collected by W. M. Mann (Firestone-Smithsonian Expedition); with Macrotermes natalensis Haviland. In collection of United States National Museum. Paratypes. Twenty-one specimens, same data as type; in collections of United States National Museum and Chicago Natural History Museum. Other material: Three specimens from Toumodi, Cote d'lvoire, in Cameron collection, British Museum (Natural History). Termitonda tachyporoides, new species Distinguished from liberiae by chaetotaxy of elytra and abdomen and by proportions of antennal segments. Color light reddish-brown. Head and pronotal chaetotaxy as in liberiae. Elytral margin with 4 erect and 4 laterally directed setae, disk with setae in several irregular transverse rows as follows: 4 in sub-basal row, 4 in sub-apical row, one near suture, and 3 near sides. Tergites with moderately numerous, pale, semi- recumbent hairs and with dark pale-tipped setae as follows: tergites 3-6 usually with 8 strong setae in a marginal row (two groups of 4 setae, widely separated); tergite 7 with a transverse row of about 6 dark setae (weaker than on preceding segments); tergite 8 with 8 or more dark setae on each half. Sternites with a marginal row of long, tsrong bristles and moderately numerous semi-recumbent pale hairs. Antennal segments with the following relative lengths: 18 : 8 : 6 : 5 : 7:6:6:5:5:5: 10; segments 3 and 4 relatively larger than in T. liberiae. Length, 2.5-2.8 mm. Holotype. From Rwindi Camp, Belgian Congo, collected May 6, 1948, by Alfred E. Emerson; with Macrotermes natalensis Havi- land. In collection of Chicago Natural History Museum. 240 FIELDIANA: ZOOLOGY, VOLUME 40 Paratype. One specimen, same data as type. Other material: Four specimens, Toumodi, Cote d'lvoire, Cameron collection, British Museum (Natural History). Subtribe TERMITOZYRINA A small group of termitophilous species occurring in the Neo- tropical Region probably form a natural group within the Myr- medoniini. Our knowledge of the tribe is as yet too limited to permit comparison with other tribal groups. TERMITOPHAGUS Silvestri Termitophagus Silvestri, 1945, Comm. Pont. Acad. Sci., 9: 530. Type species: Termitophagus synterminus Silvestri. Head almost one-half broader than long (excluding labrum); inserted in pronotum to a raised occipital margin which is continuous laterally with the infraorbital carina. Eyes large, occupying a large part of the side of the head. Antennae geniculate; scape moderately large, its inner surface a little flattened; segment 2 subcylindrical; segment 3 elongated; segments 4-10 moderately in- crassate; segments 5-10 transverse. Gular sutures subparallel except where diverging near base. Maxillary sinuses large, but not continued as shallow sulci to base of head. Mouth parts typical of tribe. Pronotum twice as broad as long; strongly convex at middle, the sides be- coming less so; broadest about midway between base and apex; side margins converging very strongly in front, much less so behind; apex deeply emarginate (equal to one-eighth length of pronotum); base strongly bisinuate; posterior angles prominent. Elytra a little longer than pronotum, but their sutural length only four-fifths the length of the pronotum; outer apical angles acutely produced. Scutellum large, not carinate. Mesosternal process very short, metasternal process broad and long; metasternum without raised line connecting posterior margins of mesocoxal acetabula. Femora moderately broad and thin, tibiae cylindrical. Middle tarsi with segments of the following relative lengths: 16 : 12 : 10 : 9 : 20; hind tarsi: 22 : 18 : 14 : 10 : 24. Abdomen without noteworthy features. Eighth tergite with an apical row of very minute teeth. Termitophagus synterminus Silvestri. Figure 35, 6. Termitophagus synterminus Silvestri, 1945, Comm. Pont. Acad. Sci., 9: 532, text figs. 7, 8 (Jabaquara, Sao Paulo, Brazil; Syntermes grandis Rambur; Silvestri coll.). Material examined, Sixteen specimens, Sao Miguel Paulista, Sao Paulo, Brazil, October 31, 1950; 21 specimens, Sao Paulo, Brazil, October 31, 1950; specimens from Sao Paulo, June 11, 1951; all collected by R. L. Araujo; with Syntermes wheeleri Emerson (determined by R. L. Araujo). SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 241 Color rufo-testaceous, except head brown and pronotal disk irregularly brownish. Dorsum without erect setae; sternites 3-7 with a few short setae along apical margins; sternite 8 with 14 fine, dark setae. Head, except for impunctate clypeus, moderately densely punctate and pubescent. Pronotum uniformly moderately densely punctate and clothed with a fine, recumbent pubescence. Elytral punctation a little more dense, the slightly coarser pubescence arising from feeble asperities. Tergites, paratergites, and sternites moderately densely and asperately punctulate, the short, fine, pale hairs recumbent. Head and pronotal disk with faint traces of reticulation; sides of pronotum, elytra, and abdomen distinctly so. Length, 4-7 mm. TERMITONUSA Borgmeier Termitonusa Borgmeier, 1950, Rev. Ent., 21 : 660. Type species: Termitonusa sequax Borgmeier. Borgmeier stated that this genus is distinguished from Termito- phagus by its more slender form, position of the eyes, form of the gula, and presence of an apical row of erect setae on the tergites. Although I have not studied this genus, there does not seem to be much justification for separating it from Termitophagus. Termitonusa sequax Borgmeier Termitonusa sequax Borgmeier, 1950, Rev. Ent., 21: 662, text figs. 32-38 (Campinas, Goias, Brazil; Cornitermes similis Hagen; Borgmeier coll.). Borgmeier's description and figures should be consulted for the characteristics of this species. The host of the holotype has been redetermined as Procornitermes araujoi Emerson. IHERINGOCANTHARUS Bernhauer Iheringocantharus Bernhauer, 1912, Verh. Zool. Bot. Ges. Wien, 62: 47. Type species: Iheringocantharus ypiranganus Bernhauer. Distinguished from the other genera of the subtribe Termito- zyrina as well as from all known members of the tribe Myrmedoniini (s. str.) by its large, distinctive pronotum, which completely covers the head. The markedly limuloid body-form is a distinguishing feature of the genus, for it is quite unlike the typical myrmedonine facies. Bernhauer placed Iheringocantharus in the subfamily Tachy- porinae, probably because it bears a superficial resemblance to certain tachyporine genera. It is impossible to judge the correct systematic position of this genus from Bernhauer's description, as he made several errors, including the incorrect tarsal formula. The 242 FIELDIANA: ZOOLOGY, VOLUME 40 mouth parts and other structures clearly show the affinity of this genus to the Termitozyrina. Iheringocantharus has a facies very similar to that of Termitohospes and may be misidentified as a member of that genus unless the mouth parts and tarsi are examined. Body-form limuloid, the head covered by the large, shield-shaped pronotum; pronotum very broad and long, nine-tenths broader than long, its sides and apex deflexed, the lateral and apical margins continuously arcuate; head in a vertical position but without appreciable modification. Iheringocantharus ypiranganus Bernhauer. Figure 35, c. Iheringocantharus ypiranganus Bernhauer, 1912, Verh. Zool. Bot. Ges. Wien, 62: 48 (Ypiranga, Sao Paulo, Brazil; no host given; Chicago Natural History Museum). Material examined. The holotype in the Bernhauer collection. One specimen from Interlagos, Sao Paulo, Brazil, collected June 11, 1951, and one from Sao Miguel, Paulista, Sao Paulo, Brazil, col- lected October 10, 1950, by R. L. Araujo; both with Syntermes wheeleri Emerson (determined by R. Araujo). Color dark testaceous; shining. Pronotum finely and irregularly punctulate, the punctules very minute and sparse medially but more definite and more dense toward the lateral margins. Elytra almost impunctate, with only suggestions of extremely fine punctules. Pronotum with 34 moderately long, curved setae as follows: 9 in a submarginal row along each side, a transverse row of 4 at level of last seta of submarginal row, 4 longitudinal rows of 2 setae near center of disk, and 4 setae on posterior margin. Outer elytral margins with several setae and a number of fine, pale hairs; elytral disk with 1, 3, 2, 1 setae in longitudinal rows. Tergites 3-6 with 6 long setae and several finer hairs on apical margin and one seta near lateral margin; tergite 7 with one seta near each lateral margin and a pair of subapical ones; tergite 8 with 2 setae on each lateral margin, 4 in a subapical row, and 2 on apical margin. Sternites 3-7 with a transverse row of 4 dark setae about the middle, one on each lateral margin, and 8-10 dark setae and a number of pale hairs on posterior margin; eighth sternite with 14 strong setae along the arcuate lateral and apical margins and 6 in a curving subapical row, as well as numerous fine, pale hairs. Antennae with segments of the following relative lengths: 22 : 12 : 12 : 8 : 6 : 6 : 6 : 7 : 7 : 7 : 15. Pronotal base bisinuate, posterior angles rounded. Elytral apices very deeply emarginate, the outer apical angles acutely angulate and very conspicuous; border within emargination almost straight. Eighth tergite with a marginal row of 10 strong teeth. Length, 2.25 mm.; width, 1.3 mm. TERMITOZYRAS, new genus Type species: Termitozyras adamsoni, new species. Distinguished from Termitophagus and Termitonusa by the much smaller size of its species; the presence of shallow sulci, margined by fine lines, along each side of the gula; the trapezoidal, less trans- Fig. 35. (a) Termitozyras adamsoni Seevers. (b) Termitophagus synterminn* Silvestri. (c) Iheringocantharus ypiranganm Bernhauer. (d) Termitosymbia nitida Seevers. 243 244 FIELDIANA: ZOOLOGY, VOLUME 40 verse pronotum, which has an almost straight apex, scarcely In- sinuate base, and rounded posterior angles; elytral apices not emarginate and their outer apical angles not produced; longer, more slender antennae, which are scarcely incrassate; non-pubescent integuments; and conspicuous macrosetae. Head more than two-fifths broader than long (excluding labrum); inserted in pronotum to a raised occipital line. Eyes less than one-half as long as head. Gula rather broad, its sides converging in front. Maxillary sinuses apparently small but continued to base of head as broad, shallow depressions on each side of gula; these shallow grooves margined by fine lines (best visible in slide mounts). Infraorbital carinae continuous with raised occipital line. Mandibles tapering to very slender apices. Mentum and submentum generalized. Maxillae typical of the Zyrini except that galea is not much longer than lacinia. Maxillary palpi moderate in size; segment 2 slender at base and broader at apex; segment 3 longer than 2, twice as long as broad. Labial palpi 3-segmented, small; basal segment longer and broader than subquadrate second segments. Pronotum two-fifths to two-thirds broader than long, evenly convex, its base broadly arcuate, with just a trace of sinuation laterally, posterior angles broadly rounded, sides feebly arcuate, converging in front, apex almost straight. Elytra about equal in length to pronotum, their apices straight, and their outer apical angles not produced. Wings with basal setose lobe. Mesosternal process broad and short; metasternal process long and slender, but chiefly at a "sub-surface" level, for there is a raised line across the metasternum from the posterior margins of the mesocoxal acetabula. Femora broad and thin. Tibiae slender, cylindrical. Tarsi 4, 5, 5-segmented. Fore-tarsi with segments 1-3 short, subequal, segment 4 equal to 2 and 3 combined. Middle tarsi with segments of the following relative lengths: 8 : 7 : 7 : 6 : 10. Hind tarsi: 12 : 9 : 8 : 7 : 12. Abdomen generalized in structure; tergites without carinae or sulci. Termitozyras adamsoni, new species. Figure 35, a. Color light reddish-brown. Head without setae. Pronotum with an apical row of 6 black setae, each lateral margin with 2, disk with transverse row of 4, and basal border with 4. Elytral margins with 3 and disk with 4 setae. Tergite 2 with a black seta near each apical angle; tergites 3-6 with a subapical row of 6 strong setae; tergite 7 with a pair of very small, apical setae; tergite 8 with 4 sub- apical and 4 strong, apical setae; tergite 9 with a dense clothing of short, pale setae and 6 black setae. Each outer paratergite with one strong seta. Sternites with a sub-basal row and an apical row of strong, black setae. Head with a short, sparse pubescence. Pronotum very minutely punctulate. Antennae with seg- ments of the following relative lengths: 18 : 10 : 12 : 11 : 10 : 10 : 9 : 8 : 8 : 8 : 17; segments 1-5 subequal in width, segments slightly incrassate beyond fifth; seg- ments, except 8-10, longer than broad; 8-10 subequal in length and width. Pro- notum two-thirds broader than long; sides declivous but not strongly deflexed; width obviously greater than that of head; anterior margin not closely embracing head. Length, 3.5 mm. Holotype from Arena Forest, Trinidad, British West Indies; collected February 22, 1945, by R. G. Donald and F. O. Iwenjora SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 245 (ex Adamson collection); with Labiotermes labralis Holmgren. In collection of Chicago Natural History Museum. Paratypes. Thirty-five, same data as type, from two colonies of termites; 21, Arena Forest, May 24, 1944, A. M. Adamson; 20, Mt. Tabor, Trinidad, December 15, 1945, E. McCallan, J. T. Davey, and G. Williams (ex Adamson coll.); all with Labiotermes labralis Holmgren. Termitozyras guianae, new species Closely related to adamsoni, but differing in the chaetotaxy of pronotum and elytra as follows: apical margin of pronotum with 6 setae and lateral margins with 2 but with no setae on disk or on basal margin; elytra with only 2 setae on lateral margins. Color rufo-testaceous. Length, 2.1 mm. Holotype from Kartabo, British Guiana, collected July 24, 1920, by Alfred E. Emerson; with Labiotermes labralis Holmgren. In collection of Chicago Natural History Museum. Paratype. One specimen, same data as holotype. Termitozyras boliviae, new species Distinguished from adamsoni by these differences in chaetotaxy: pronotal setae of same general arrangement but considerably weaker; elytral disk with short, fine setae but no dark macrosetae; head, pronotum, and elytra more minutely and sparsely punctulate and with almost imperceptible hairs. The pronotum is only two-fifths broader than long and although it is as large as in the other two species, the sides are much more strongly deflexed and it appears narrower. The pronotum is more strongly convex than in the other species and the front margin closely embraces the head so that the head is as broad as the pronotum in front. Length, 2.2 mm. Holotype from Cachuela Esperanza, Beni, Bolivia; collected by W. M. Mann (Mulford Expedition); with Labiotermes labralis Holmgren. In collection of United States National Museum. Paratypes. Six specimens, same data as type, in Chicago Natural History Museum and United States National Museum. TERMITOSYMBIA, new genus Type species: Termitosymbia nitida, new species. Distinguished from all other genera of the tribe by its very distinctive physogastric abdomen. In addition, its pronotum has a distinctive form, the elytra are unusually long, and the metepimera are not produced behind the elytra. 246 FIELDIANA: ZOOLOGY, VOLUME 40 Head a little broader than long; vertex strongly depressed in type specimens (probably due to collapse of thinly sclerotized integument). Antennae inserted under a narrow vertexal arcade, the fossae opening cephalad. Clypeus trans- versely convex, strongly declivous. Infraorbital carina absent. Gula long and moderately broad, its sides converging moderately in front. Labium generalized; labial palpi small. Maxillae and maxillary sinuses typically myrmedoniine. Pronotum robust, one-half broader than long, broadest at a point three-fifths the distance from base to apex, from which point sides converge strongly to rounded basal and apical angles; base strongly arcuate, apex less so; disk with a pair of deep pits and a shallow sub-basal impression; pronotum margined. Elytra with moderately deep sulci beginning behind humeri and extending obliquely to middle of each apical margin. Middle coxae separated by a broad, short mesosternal process and a longer, narrower metasternal process. Metepimera not elongated. Abdomen physogastric, twice as broad as pronotum, some sclerites, particu- larly sternites, separated by non-sclerotized integument; somewhat scaphoidal in appearance, tergites very broad, paratergites strongly elevated to form lateral margins; inner paratergites extremely broad, about three-tenths as broad as tergites; sternites strongly convex, their lateral margins curved so as to be visible from above. Termitosymbia nitida, new species. Figure 35, d. Color flavo-testaceous; tergites very pale, except 7-9, which are brown; dorsum very shiny. Head and pronotum with extremely sparse, minute punctu- lation and pubescence (glabrous in appearance) ; head with obsolescent reticulation. Scutellum reticulated, pubescent; elytra with small, darkly pigmented, sparsely distributed punctules, more numerous near base, and with extremely sparse, minute pubescence. Head with a row of about 6 setae on clypeal margin and a seta medial to each antennal fossa. Pronotum with 4 short, black setae on anterior margin, 4 on each lateral margin, 2 at base, and 4 on disk (one lateral to each pronotal fovea, and a pair behind the foveae). Elytra without setae. Tergites 3-6 without punctation or pubescence, obsoletely reticulated; tergites 7-8 with numerous longitudinal, carinulate lines; tergites with an apical row of short, fine setae. Sternites 3-7 with a distinctive vestiture of short, fine, dark, erect hairs, very dense except on sternites 3 and 7 and on sides of sternites 4-6; third sternite with a moderate number of long, black hairs in addition to apical row; sternites 4-7 with numerous long, black setae on the sides; these are es- pecially long near lateral margins. Antennae geniculate; with relative lengths and widths as follows: 28 (16): 12 (8) : 18 (12) : 18 (11) : 16 (11) : 16 (12) : 16 (12) : 15 (12) : 15 (13) : 15 (14) : 22 (13); scape robust, segment 2 small, 3 slightly ovoidal, segments 4-8 subcylindrical, 9-10 slightly obtrapezoidal. Length, 3 mm. Holotype. From Cacheula Esperanza, Beni, Bolivia, collected by W. M. Mann (Mulford Expedition); with Labiotermes labralis Holmgren. In collection of United States National Museum. Paratype. One specimen, same data as type; in collection of Chicago Natural History Museum. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 247 Tribe PSEUDOPERINTHINI Subfamily Pseudoperinthinae, Cameron, 1939, Fauna Brit. India, Staphyl., 4: 1. Wasmann placed Pseudoperinthus in the Aleocharinae, but Cameron transferred it to a new subfamily in the belief that its head structure does not conform to the aleocharine pattern. He noted that the antennae are inserted under an arcade, and that there is a fine, cariniform line across the front. The Pseudoperinthus head is by no means as unusual as Cameron implies and the genus certainly belongs to the Aleocharinae. Was- mann's choice of a generic name seems to be very appropriate, inasmuch as this genus bears a very strong resemblance to Perinthus, even to the point of having a frontal cariniform line (a character which does not, however, appear in all Perinthina). Pseudoperinthus may, indeed, be more closely allied to the Perinthina than has been suggested. Specimens of Pseudoperinthus have not been available until very recently and the necessary comparative studies for its correct placement have not been made. Several characteristics of the genus do not conform to the Perinthina character pattern, the 4, 5, 5-segmented tarsi and the absence of polytrichous coeloconic sensilla from the terminal antennomeres, in particular. Cameron figured the head of fletcheri and Wasmann published photographs of malayanus, but the species were so poorly described that diagnostic features cannot be given. PSEUDOPERINTHUS Wasmann Pseudoperinthus Wasmann, 1916, Zool. Jahrb. Syst., 39: 194; Cameron, 1939, Fauna Brit. India, Staphyl., 4: 1. Type species: Pseudoperinthus malay- anus Wasmann. Body-form limuloid. Head small, hypognathous; inserted in pronotum to eyes. Front with sharp, cariniform line between eyes; antennae inserted under arcade; clypeus depressed. Eyes small. Infraorbital carina present. Antennae 11-segmented. Lacinia longer than galea, ciliate; galea slender, acute, with about 8 spines. Gula broad; mentum trapezoidal, its apex almost straight. Labial palpi 2-segmented. Ligula deeply bifid. Pronotum large, transverse, convex. Meso- sternal process very narrow, elongated; middle coxae contiguous. Tarsi 4, 5, 5- segmented. Abdomen non-physogastric; fusiform; apex acute. Length, 1.75 mm. Pseudoperinthus malayanus Wasmann Pseudoperinthus malayanus Wasmann, 1916, Zool. Jahrb. Syst., 39: 195, pi. 4, fig. 16, a, b, pi. 5, fig. 16, c-h (Tandjong Slamat, Sumatra; Lacessiti- termes laborator Haviland; Wasmann coll.). 248 FIELDIANA: ZOOLOGY, VOLUME 40 Pseudoperinthus fletcheri Cameron Pseudoperinthus fletcheri Cameron, 1939, Fauna Brit. India, Staphyl., 4: 3, text figs. 1, 2 (Coorg, Mercara, India; Eutermes sp.; Cameron coll.). Tribe ATHETINI The large generic complex related to Atheta probably deserves tribal status but it is impossible at present to characterize the Athetini. A comparative study of the genera now assigned to Myrmedoniini (s. lat.) must first be made. Subtribe GOPTOTERMOECIINA COPTOTERMOECIA Oke Coptoternioecia Oke, 1933, Proc. Roy. Soc. Victoria, 45: 135. Type species: Coptotermoecia alutacia Oke. This genus seems to be a member of the Atheta complex of genera but it is impossible at present to place it accurately within the group. A characterization of the genus is difficult and will not be attempted at this time. The head and thorax of Coptotermoecia seem to be rather generalized, and distinctive features are not evident. The abdomen, which is somewhat scaphoidal in form and perhaps incipiently physogastric, will doubtless prove to be of importance in determining the tribe to which Coptotermoecia is ultimately assigned. Oke's figure of the Coptotermoecia abdomen is very misleading, for it was based on specimens that were distorted in the process of drying. Specimens in alcohol, or those that are mounted dry after dehydration, have moderately inflated abdomens. Coptotermoecia alutacia Oke Coptotermoecia alutacia Oke, 1933, Proc. Roy. Soc. Victoria, 45: 136, text figs. 48, 84-88 (Natya and Violet Town, Victoria, Australia; Coptotermes acinaciformis Froggatt; Oke coll.). Material examined. Thirty specimens, Shannons Creek, Capital Territory, Australia; collected July, 1939, by F. J. Gay. One specimen, Mt. Keira, New South Wales, Australia; collected October 7, 1948, by C. E. Chadwick (W. 0. Steel coll.). Host of all specimens: Coptotermes acinaciformis Froggatt (=C. lactens Froggatt). Color pale reddish-brown; antennae, mouth parts, paratergites, and elytral tips paler. Head smooth, with a very few fine punctules and very inconspicuous SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 249 pubescence. Pronotum conspicuously punctate, the moderate-sized punctures irregularly arranged in about ten to twelve rows from base to apex; pronotum obsoletely reticulate. Elytra very smooth, with a few setigerous punctures and a suggestion of minute punctulation. Head with a pair of setae medio-caudad to each eye. Pronotum with 20 moderately long setae as follows: 6 on anterior margin, 6 on basal margin, 2 on each lateral margin, 4 on disk. Elytra with four irregular, longitudinal rows of setae comprised of 2 or 3, 3, 2, 3 setae, respectively, beginning with sutural row. Tergites smooth, glabrous, non-reticulate; tergites 3-6 with 6 very long setae and a few minor ones in apical rows; tergite 7 with about 8 long setae in a sub- apical row; tergite 8 and terminal sclerites with a moderately dense vestiture of medium to long, pale hairs. Sternites almost glabrous, with apical rows of long setae but with only a few scattered hairs elsewhere. Antennae geniculate, with basal segment a little shorter than second and third combined; segment 2 longer than 3, both slender; segment 4 slightly transverse, obtrapezoidal ; segments 6-10 transverse, subequal in length, feebly incrassate. Third segment of labial palpi much shorter than two basal segments combined. Pronotum one-half broader than long. Abdomen only slightly broader than elytra, its sides subparallel. Tergites quite convex; inner paratergites one-sixth as broad as tergites; seventh tergite scarcely, if at all, longer than other tergites. Length, 2.5-2.7 mm. Coptotermoecia gayi, new species. Figure 36, b. Distinguished from C. alutacia Oke by its paler coloration; impunctate pronotum (except for extremely fine punctulation); antennal proportions; somewhat longer third segment of labial palpi; proportionately longer and narrower pronotum; proportion- ately broader and more strongly inflated abdomen (one-third broader than elytra); relatively broad inner paratergites; seventh tergite approximately twice as long as other tergites; distinctive abdominal chaetotaxy. Color rufo-flavate, dorsum of abdomen brownish ; antennae, mouth parts and under side paler rufo-flavate. Head smooth, shining; with a scarcely perceptible reticulation. Pronotum and elytra smooth, shining; with sparse, very fine punc- tules and very fine, stubbly pubescence. Head and pronotal chaetotaxy as in C. alutacia. Tergites 3-7 smooth, almost glabrous; tergite 7 with a faint reticulation; tergites 3-6 with a subapical row of long, pale bristles and with a few shorter setae in addition; tergite 7 with 4 or 5 irregular rows of moderately long, fine setae that become longer apically; tergite 8 densely clothed with moderately long, fine, pale hairs and toward apex with a dense, recumbent pubescence. Para- tergites with a moderate number of long setae. Sternites with a uniform vestiture of medium density, the fine, moderately long, pale hairs erect, semi-erect, or recumbent. Antennae geniculate, segments 2 and 3 slightly elongated; segments 4-10 transverse, increasing in width and length. Basal segment of labial palpi moder- ately large; second segment narrower and a little shorter; third segment long and 250 FIELDIANA: ZOOLOGY, VOLUME 40 slender, almost as long as two basal segments together. Pronotum three-tenths broader than long; pronotal apex almost straight, slightly sinuate; pronotum a little broader in front of middle. Elytra generalized. Wingless. Abdomen moderately inflated; scaphoidal in form; one-third broader than elytra. Inner paratergites relatively broad, about three-tenths as broad as tergites. Seventh tergite approximately twice as long as any of the preceding tergites. Length, 3 mm.; width of abdomen, 1 mm. Holotype from Shannons Creek, Capital Territory, Australia; collected July, 1939, by F. J. Gay; with Coptotermes acinaciformis Froggatt. In collection of Division of Entomology, Commonwealth Scientific and Industrial Research Organization, Canberra. Paratypes. Fifty-four specimens, same data as holotype, in collections of the above institution and Chicago Natural History Museum. Subtribe TERMITOTELINA This category is proposed for a single genus, Termitolelus, recorded from a termite nest in Africa. The relationship of Termi- totelus to the Athetini is yet to be determined. I examined the type of Termitolelus schultzei Wasmann but could come to no decision as to its systematic position, except that it probably does not belong to any of the other termitophile categories of this paper. TERMITOTELUS Wasmann Termitotelus Wasmann, 1908, Denkschr. Med.-Nat. Ges. Jena, 13: 444. Type species: Termitotelus schultzei Wasmann. Termitotelus schultzei Wasmann Termitotelus schultzei Wasmann, 1908, Denkschr. Med.-Nat. Ges. Jena, 13: 444, pi. 22, a, figs. 7, la, 76 (Kookfontein, Klein-Namaland, South West Africa; Microhodotermes viator Latreille; Wasmann coll.). Tribe PHILOTERMITINI, new tribe This tribe is proposed for a few species of New World termito- philes associated with Reticuliiermes and Coptotermes of the Rhino- termitidae. Philotermes Kraatz has always been assigned to the tribe Bolitocharini because of its 4, 4, 5-segmented tarsi. The Bolitocharini are so poorly characterized at present that com- parison of the Philotermitini with them is almost impossible. Tribal diagnosis is deferred until the bolitocharine complex is more care- fully analyzed. The Philotermitini are ecologically distinctive, for most of the Bolitocharini are fungus inhabitants. Philotermes SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 251 I Fig. 36. (a) Termitonda liberiae Seevers. (b) Coptotermoecia gayi Seevers. apparently has no morphological characteristics that we associate with termitophily but inasmuch as they are obligatory guests they must certainly have some degree of physiological adaptation to the colonial life. The nature of their mouth parts suggests that they are predaceous. Six North American species and one Central American species are known. KEY TO GENERA OF PHILOTERMITINI Front of head strongly deflexed, clypeus and labrum vertical; head with a V-shaped frontal cariniform line extending laterally above the antennal fossae; ligula bifid; presternum carinate; lateral lobe of aedeagus not prolonged as a slender process Neophitotermes, new gen. Head slightly declivous in front; clypeus and labrum more or less horizontal; head without complete cariniform line (sometimes with a trace above antennal fossae); ligula simple; presternum not carinate; lateral lobe of aedeagus prolonged as a slender process Philotermes Kraatz PHILOTERMES Kraatz Philotermes Kraatz, 1857, Linn. Ent., 11: 13; Blatchley, 1910, Coleopt. Indiana, p. 343; Fenyes, 1920, Gen. Insect., 173B: 126; Seevers, 1938, Ann. Ent. Soc. Amer., 31 : 430. Type species: Philotermes pilosus Kraatz. 252 FIELDIANA: ZOOLOGY, VOLUME 40 Head prognathous; clypeus feebly declivous, transversely convex; frontal cariniform line incomplete (a raised line above antennae may extend medially). Eyes small to medium-sized. Infraorbital carinae present. Antennae robust; segments 4-10 cylindrical, broader than long. Labium generalized, mentum trapezoidal, ligula simple, rather long; labial palpi 3-segmented, two basal seg- ments moderately robust, the third filiform. Mandibles long and strong, falcate. Galea and lacinia subequal in length ; lacinia with a long comblike row of spinose setae. Maxillary palpi 4-segmented; second segment claviform, third fusiform to strongly compressed. Pronotum transverse, one-half or more longer than broad; broadest at middle or slightly in front of middle, base and sides continuously arcuate, basal angles obsolete or very nearly so; sides deflexed, and area near anterior angles usually depressed and densely clothed with long, fine, yellow hairs. Middle coxae con- tiguous; mesocoxal acetabula margined externally and caudally, and broadly confluent medially; mesosternal process short, angulate, and metasternal process very short. Tarsi 4, 4, 5-segmented. Abdomen not physogastric; moderately broad, its sides subparallel. Seventh tergite with a broad, V-shaped emargination; apex of eighth tergite unmodified, emarginate, or dentate. KEY TO SPECIES OF PHILOTERMES KRAATZ 1. Vertex and occiput without erect setae, and with very few fine, recumbent hairs; head without ground sculpture; medial one-half of elytra glabrous. pilosus Kraatz Vertex and occiput with some erect setae, and with a moderately dense vestiture of finer hairs; head reticulated or densely punctate; elytra entirely clothed with hairs 2 2. Eighth tergite emarginate in both sexes 3 Eighth tergite dentate in male, unmodified in female 5 3. Sternites very densely, finely pubescent (appearing tomentose). werneri, new sp. Sternites with fine, pale, erect hairs and scattered bristles, but not pubescent . 4 4. Elytra clothed with pale, recumbent hairs; antennal segments 2-11 conjointly seven-tenths to four-fifths longer than pronotum cubitopilis, new sp. Elytra clothed with pale, erect hairs; antennal segments 2-11 conjointly two-fifths to three-fifths longer than pronotum fnchsii Kraatz 5. Tergites 3-5 with 6 setae in apical row, the second and fifth fine and pale; tergites 6-8 reticulated emersoni Seevers Tergites 3-5 with 8 setae in apical row, the second, third, sixth, and seventh fine and pale; tergites 6-8 obsoletely reticulated pensylvanicus Kraatz Philotermes pilosus Kraatz. Figure 37, a. Philotermes pilosus Kraatz, 1857, Linn. Ent., 11: 14, pi. 1, fig. 11, pi. 2, fig. 13 (Tennessee region; with termites; ?Deuts. Ent. Inst.); Schwarz, 1889, Proc. Wash. Ent. Soc, 1:1; Blatchley, 1910, Coleopt. Indiana, p. 343; Fenyes, 1920, Gen. Insect., 173B: 127, pi. 2, fig. 7; Seevers, 1938, Ann. Ent. Soc. Amer., 31: 432, pi. 2, figs. 5, 7, 10, 15. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 253 Fig. 37. (a) Philotermes pilosus Kraatz. Kraatz. (c) Neophilotermes laxicornis Sharp. (b) Philotermes pensylvanicus Material examined. Framingham (C. A. Frost), Melrose Heights (USNM; R. H. Timberlake), and Tyngsboro (USNM, Hubbard and Schwarz), Massachusetts. Huntington and Forest Hills, Long Island, New York. Washington, D.C. (USNM, Hubbard and Schwarz). Crawford and Posey counties, Indiana (Purdue Univ., Blatchley coll.). Turkey Run State Park, Indiana (Seevers). Olney, Illinois (A. Emerson). Host: Reticulitermes flavipes Kollar. Color yellowish-brown; head, pronotum, elytra, and tergites often darker. Head very minutely and sparsely punctate and with very fine, sparsely arranged yellow pubescence, a little longer behind the eyes. Pronotum densely punctate; clothed with a yellow pubescence except for a narrow median impunctate and nearly glabrous strip. Pronotum with many dark, pale tipped setae; these are absent from the median strip and the region adjacent to anterior angles. Pronotal shoulders with a dense vestiture of long, pale hairs. Lateral half of each elytron densely clothed with fine, white, recumbent pubescence and scattered setae; median areas of elytra almost glabrous and with only one or a few setae. Tergites 254 FIELDIANA: ZOOLOGY, VOLUME 40 with a row of 6 dark setae; sternites 3-5 with fine pubescence and scattered setae, other sternites glabrous, with only two rows of dark setae near apex. Antennae robust, segments 4-10 subequal; eleventh only three-fifths longer than tenth. Pronotum distinctive: about twice as broad as long; broader than either head or elytra; broadest just behind center; surface slightly convex medially then slightly concave laterally before leveling off toward the side margins; anterior border sinuate, anterior angles rounded and slightly depressed; the sides behind broadest point and the base continuously arcuate; posterior angles obsolete. Anterior tibiae having their sides nearly parallel; their distal four-fifths or more with a very dense clothing of short, stiff, yellow hairs except for a strip along outer ventral margins; their outer margins with 5 or 6 slender black setae, apex with a stout spine. Middle tibiae increasing in width distally, their apices some- what swollen; ventral surfaces glabrous but outer surfaces with a clothing of short yellow hairs on swollen distal area and extending almost to base along inner margin. Hind tibiae slender, their outer surfaces with some fine, pale hairs unlike vestiture of other tibiae. Length, 3-4 mm.; width, 1 mm. Philotermes fuchsii Kraatz Philotermes fuchsii Kraatz, 1857, Linn. Ent., 11: 15 (Tennessee region; in termite nest; ?Deuts. Ent. Inst.); Blatchley, 1910, Coleopt. Indiana, p. 343; Seevers, 1938, Ann. Ent. Soc. Amer., 31: 433, pi. 2, fig. 9. Material examined. Hales Bar, Tennessee (A. Emerson); with Reticulitermes virginicus Banks. Chain Bridge, Virginia (USNM, T. E. Snyder); with R. virginicus Banks. Gainesville, Florida (0. Falls) ; with R. virginicus Banks. Crescent City, Florida (USNM, Hubbard and Schwarz). Color light-brown. Head, pronotum, and elytra densely clothed with fine, pale, erect hairs; elytral pubescence particularly conspicuous (best viewed in profile). Head with a transverse row of about 4 dark setae on vertex and a basal row of about 6 on occiput; each half of pronotum and each elytron with about five irregular, longitudinal rows of erect setae. Antennae relatively short; seg- ments 2-11 conjointly two-fifths to three-fifths longer than pronotum; variation of specimens as follows: Crescent City (1.4; 1.5; 1.5; 1.5; 1.5; 1.53; 1.54); Gaines- ville (1.5); Hales Bar (1.63); Chain Bridge (antennae broken). Pronotum two- fifths broader than long, evenly convex, anterior margin almost straight, sides arcuate, broadest about center, sides behind center continuously arcuate with base, basal angles obsolete. Eighth tergite emarginate, the emargination shallowly angulate at center. Eighth sternite slightly angulate medially. Length, 2.1-2.4 mm.; width, 0.7 mm. Remarks. It is now clear that specimens previously identified as fuchsii belong to two species. Inasmuch as I have not seen Kraatz' type of fuchsii, it is difficult to determine with certainty which species it represents. The species characterized above seems to fit the original description somewhat better, especially with SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 255 regard to the following features: pronotum clothed with long, very fine pubescence; elytral pubescence a little longer than on foreparts; antennal segments 4-10 broad and slightly transverse. The species commonly confused with fuchsii is described below. Philotermes cubitopilis, new species Distinguished from fuchsii by the recumbent pubescence of head, pronotum, and elytra, and by the relatively longer antennae. When compared to the pronotal length, which is the same in both species, the antennae are relatively longer in all specimens examined. It is difficult to attribute this increase to particular segments, as each seems to contribute to the greater length. Color light brown. Head, pronotum, and elytra densely pubescent, the pale hairs in a recumbent position, especially on the elytra. Setae of head, pronotum, and elytra as in fuchsii. Antennal segments 2-11 conjointly seven-tenths to four-fifths longer than pronotum; variation of specimens as follows: Tampa (1.76; 1.77; 1.77; 1.8; 1.8); Gainesville (1.76); Daphne (1.74); Arkadelphia (1.7); Washing- ton (1.7; 1.7). Pronotum and abdomen as in fuchsii. Length, 2.1-2.5 mm.; width, 0.7 mm. Holotype from Tampa, Florida, collected November 26, 1936, by Olive Falls; with Reticulitermes virginicus Banks. In collection of Chicago Natural History Museum. Paratypes. Four specimens, same data as type. One specimen, Gainesville, Florida, October 25, 1937, Olive Falls; with R. virginicus Banks. One specimen, Daphne, Alabama, December 29, 1940, B. A. Maina; with R. virginicus Banks. Three specimens, Washing- ton, D.C.; USNM, Hubbard and Schwarz. One specimen, Arka- delphia, Arkansas, April 4, 1937, A. E. Emerson. Philotermes werneri, new species Distinguished from fuchsii and cubitopilis, to which it is most nearly related, by the broader pronotum (three-fifths broader than long), the densely pubescent sternites, the strongly compressed third segment of maxillary palpi, and the differences in chaetotaxy described below. Color yellowish-brown. Head with a moderately dense cover of rather long, erect, pale, very fine hairs; without recumbent pubescence or dark setae. Head finely, closely reticulated. Pronotum with a dense clothing of fine, pale hairs which increase in length laterad to become long and conspicuous on the deflexed sides; in addition, with numerous longer dark setae, with fine, pale tips. Elytra with numerous fine, pale, semi-recumbent hairs, directed in general toward the outer apical angles; in addition, with erect setae, black with pale tips, which are sparse on disk but fairly numerous along the sides. 256 FIELDIANA: ZOOLOGY, VOLUME 40 Tergites 3-6 with short, sparse, pale hairs and an apical row of much longer, semi-recumbent bristles. Third sternite with numerous erect hairs of moderate length but not pubescent; sternites 4-7 densely pubescent (almost tomentose) and with a moderate number of erect setae; sternites 3-7 with an apical row of longer setae; sternites reticulated; eighth sternite without reticulation or pu- bescence but with a moderate number of setigerous tubercles bearing long, fine hairs. Pronotum three-fifths broader than long; broadest a little in front of center, sides behind this point converging at about forty-five degrees and con- tinuously arcuate with the rounded base; apex arcuate and a little sinuate; apical angles slightly obtuse, depressed. Eighth tergite broadly notched at middle; eighth sternite feebly, obtusely angulate at middle. Third segment of maxillary palpi strongly compressed (in contrast to the usual fusiform condition). Length, 2.75 mm.; width, 0.85 mm. Holotype from Murrells Inlet, Georgetown County, South Caro- lina, collected May 10, 1949, by Floyd Werner and W. Nutting; with Reticulitermes virginicus Banks. In collection of Chicago Natural History Museum. Paratypes. One specimen, same data as type. One specimen, Gainesville, Florida; collected October 3, by Hubbard and Schwarz; in United States National Museum. Philotermes pensylvanicus Kraatz. Figure 37, b. Philotermes pensylvanicus Kraatz, 1857, Linn. Ent., 11: 15 (Pennsylvania; in termite nest; ?Deuts. Ent. Inst.); Schwarz, 1889, Proc. Wash. Ent. Soc, 1:1; Seevers, 1938, Ann. Ent. Soc. Amer., 31: 434, pi. 2, fig. 6. Material examined. Oakridge, Pennsylvania (CNHM). Ithaca, New York (USNM, R. C. Shannon). Washington, D.C. (USNM, Hubbard and Schwarz). Clarksville, Tennessee (USNM, S. E. Crumb). Smith Station, LaPorte Co., Indiana (CNHM, A. E. Emerson). Jerseyville, Illinois (USNM, T. E. Snyder). Olney, Illinois (CNHM, A. E. Emerson). Peoria, Illinois (CNHM). Mt. Vernon, Texas (CNHM, A. E. Emerson). In all instances where the host has been collected and determined, it has been Reticulitermes flavipes Kollar. Color reddish-brown. Head, pronotum, and elytra closely and rather coarsely punctate; clothed with short, recumbent pubescence; pronotum with a dense cluster of hairs near apical angles. Head with short dark setae as follows: an irregular, oblique row of 5 setae on each side from between antennal fossae to a point behind the eye; 2 on vertex between the above rows; an occipital row of 6; 2 or 3 on postgenae. Pronotum and elytra with numerous scattered setae. Sternites 3 and 4 with 5 or 6 irregular rows of setae; sternites 5 and 6 with 3 or 4 similar rows; eighth sternite with a "gland opening" surrounded by hairs of a distinctive pattern; sternites 3 and 4 sparsely and irregularly clothed with fine hairs, but not reticulated; sternites 5 and 6 with moderately numerous, rather coarse, setigerous tubercles, which are not restricted to apical half of sternite; SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 257 sternite 7 with numerous setigerous carinules; sternites 7 and 8 usually less heavily sclerotized than the others, sometimes translucent; sternite 8 arcuately produced at middle. Tibiae slender and densely pilose but not as in P. pilosu*. Pronotum three-fifths broader than long. Length, 3 mm. Philotermes emersoni Seevers Philotermes emersoni Seevers, 1938, Ann. Ent. Soc. Amer., 31 : 435, pi. 2, fig. 8 (Indiana Dunes State Park, Indiana; Reticulitermes arenincola Goellner; Chicago Natural History Museum). Closely related to pensylvanicus, from which it may be dis- tinguished by the following characteristics: smaller size (length, 2 mm.); the arrangement of the apical row of setae on tergites 3-6; pronotum about one-half broader than long (three-fifths broader than long in pensylvanicus) . Material examined. Specimens from Beverly Shores, Indiana, Michigan City, Indiana, and Crawford County, near Wyandotte Cave, Indiana; all with R. arenincola Goellner. NEOPHILOTERMES, new genus Type species: Neopkilotermes laxicornis Sharp (= Philotermes laxicornis Sharp). Distinguished from Philotermes by the following characteristics: Head short, transverse, strongly deflexed in front, clypeus and labrum vertical; clypeus rather long, transversely convex; head with a V-shaped cariniform line across front and extending laterally above antennal fossae; antennae slender; their four basal segments elongated; segments 5-10 cylindrical, broader than long; ligula bifid; pronotum strongly transverse, almost twice as broad as long; elytra with apical margin sinuate; prosternum carinate; seventh tergite simple; lateral lobe of aedeagus not prolonged as a slender process. Neophilotermes laxicornis Sharp. Figure 37, c. Philotermes laxicornis Sharp, 1883, Biol. Cent. Amer., Coleopt., 1, part 2, p. 171, pi. 5, fig. 7 (Las Mercedes, Guatemala; no host; British Museum, Natural History). Color testaceous, shining, integuments not reticulated. Body sparsely and coarsely pubescent except for a small area of pronotal disk; the hairs recumbent except on parts of head. Abdomen with a sparse clothing of moderately long, pale setae. Antennae slender, segments 1-4 elongated, 5-10 short, incrassate. Length, 2.25 mm. Material examined. A series of specimens from Hamburg Farm, Guapiles, and Iberia Farm, Costa Rica (USNM, Nevermann coll.); with Coptotermes crassus Snyder. 258 FIELDIANA: ZOOLOGY, VOLUME 40 Tribe TERMITODISCINI Subfamily Termitodiscinae: Wasmann, 1912, Zeitschr. wiss. Zool., 101: 91; 1916, Zool. Jahrb. Syst., 39: 176; Bemhauer and Schubert, 1916, Coleopt. Cat., 67: 443; Cameron, 1932, Fauna Brit. India, Staphyl., 3: 314; Scheerpeltz, 1933, Coleopt. Cat., 129: 1471; Silvestri, 1947, Arch. Zool. Ital., 31 : 125. Wasmann (1899c) originally placed Termitodiscus in the Aleo- charinae but subsequently transferred it and Discoxenns to the Termitodiscinae. Staphylinid specialists have generally accepted this subfamily, placing it near the Tachyporinae or the Pygosteninae. Inasmuch as careful study of Termitodiscus and allied genera has revealed no character or group of characters justifying subfamily status for these species, they are herein returned to the Aleocharinae. The head capsule, mouth parts, thoracic structures, abdomen, and genitalia appear to fall within the range of variation of the Aleo- charinae. The antennae are unusual, but the Aleocharinae are noted for the wide variety of antennal types. Until recently only two genera of Termitodiscini were recognized. Silvestri (1947) elevated a subgenus of Termitodiscus to generic status but chose the name Ldssodiscus Grasse" and Lesperon instead of the prior name Termitogerrus Bernhauer, which he probably overlooked. The three genera are differentiated in the following section by a series of characters not heretofore utilized. All genera of Termitodiscini occur with fungus-growing termites : Termitodiscus in India and Africa with species of Odontotermes; Termitogerrus in Africa with Macrotermes; and Discoxenus in India and Ceylon with Odontotermes. It will be of interest to know if the Indomalayan Macrotermes nest harbor species of this tribe. Grasse" and Poisson (1940) made some interesting observations on the behavior of one species of this tribe, Termitogerrus lepidulus. On a number of occasions they saw these small beetles perched on the minute "mycotetes" of the fungus gardens, and, with the aid of a lens, observed them feeding on conidia, blastospores, and fungus filaments. They also noted on numerous occasions that the beetles groomed their hosts, licking the legs and bodies of workers and nymphs with rapidly moving mouth parts. They suggest that the beetles may feed on debris, such as fungus spores, adhering to the bodies of their hosts. On no occasion did they observe the termites receiving secretions from the beetles. The principal characteristics of the tribe are as follows: Body-form limuloid; head concealed beneath the very large pronotum; hypognathous; antennae in- serted in deep fossae medial to the eyes, their scapes resting in these fossae; SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 259 antennae 10- or 11 -segmented, their second and third segments small, the fourth to terminal segment forming a compact "club"; pronotum approximately twice as broad as long, its sides and apex continuously arcuate; pronotal hypomera very large, but not visible from the side; elytra broad, their epipleura very large; prosternum keeled; mesocoxae set in margined acetabula, narrowly or broadly separated; hind coxae broad, produced laterally to meet the elongated metepimera; tarsi 4, 5, 5-segmented; abdomen acuminate; aedeagus typically aleocharine in structure. KEY TO GENERA OF TERMITODISCINI 1. Mesocoxae widely separated; antennae 10- or 11-segmented, short, not, or very slightly, visible from above; antennal segments 3-10 (or 11), strongly incrassate, usually compressed 2 Mesocoxae narrowly separated; antennae 11-segmented, elongated, easily visible from above; antennal segments 3-11 form a robust, fusiform club. Discoxenus Wasmann 2. Mesocoxae separated by broad meso- and metasternal processes; mesosternum slightly declivous at middle; pronotal margin slightly flared; antennae 10- segmented Termitodiscus Wasmann Mesocoxae separated by a long, broad, metasternal process; mesosternum almost vertical at middle and thus scarcely visible from below; pronotal margin not flared; antennae 11-segmented Termitogerrus Bernhauer TERMITODISCUS Wasmann Termitodiscus Wasmann, 1899, Deuts. Ent. Zeitschr., 43: 147; 1912, Zeitschr. wiss. Zool., 101: 92; 1916, Zool. Jahrb. Syst., 39: 179; Cameron, 1932, Fauna Brit. India, Staphyl., 3: 317; Silvestri, 1947, Arch. Zool. Ital., 31: 125. Type species: Termitodiscus heimi Wasmann. Form of body relatively broad and flat. Head broad and short; the moderately broad antennal fossae widely separated, not continued as genal grooves for reception of small intermediate antennal segments. Genae sharply carinate beneath eyes. Antennae 10-segmented, short, scarcely extending beyond pronotal margin; segments 3-10 forming a compact, incrassate, moderately compressed "club." Pronotum very large, about twice as broad as long, its arcuate margin slightly flared, its base broadly emarginate. Prosternum thinly keeled. Anterior coxae moderately broad, strongly convex. Mesocoxae small, oval, widely sepa- rated; mesosternal process about twice as long as metasternal process; meso- sternum moderately declivous, longer in midline than metasternum. Hind coxae short, about three times as broad as long. Remarks. A species revision has not been attempted, as only a fraction of the species has been available. The African species recorded as belonging to this genus are probably congeneric, if I may judge by one African species of uncertain identity at hand. The following key is based on descriptions and keys in Wasmann (1912, 1916), Cameron (1932), and Silvestri (1947), as well as personal observations. 260 FIELDIANA: ZOOLOGY, VOLUME 40 1 . Pronotum and elytra with numerous semi-recumbent setae of an unusual type (the feebly notched apices broader than the bases) Indian species (2) Pronotum glabrous African species (5) 2. Length, 1.6-1.9 mm 3 Length, 1-1 .4 mm 4 3. Antennal club two and one-half times as long as width of terminal segment, which is one-third shorter than other segments of club conjointly. heimi Wasmann Antennal club very short, only twice as long as width of terminal segment, which is as long as other segments of club conjointly . . escherichi Wasmann 4. Length, 1.4 mm.; coloration brighter; terminal antennal segment a little shorter than remainder of club butteli Wasmann Length, 1 mm.; coloration darker; terminal antennal segment much shorter than remainder of club minutus Cameron 5. Elytra and abdomen with recumbent yellow hairs; antennal club very short and broad, strongly incrassate; terminal antennal segment a little broader than long, nearly triangular; color pitchy brown, abdomen lighter, pronotal margin yellow; length, 1 mm splendidus Wasmann Elytra and abdomen without recumbent yellow hairs but with erect pale hairs, sparsely arranged; antennal club short, strongly tapering; terminal antennal segment as long as broad; color dark brown, abdomen only lighter at tip, pronotum with narrow yellow margin; length, 1.5 mm braunsi Wasmann For characteristics, see Silvestri, 1947, fig. 3 (1-22) . . .transvaalensis Silvestri Termitodiscus heimi Wasmann. Figure 38, a. Termitodiscus heimi Wasmann, 1899, Deuts. Ent. Zeitschr., 43: 147, pi. 1, fig. 1, a-/(Sangamner; Wallon, Ahmednagar District, India; Odontotermes obesus Rambeau and O. wallonensis Wasmann; Wasmann coll.); 1912, Zeitschr. wiss. Zool., 101: 93, pi. 5, fig. 4; 1916, Zool. Jahrb. Syst., 39: 181, pi. 4, fig. 8, a-b, pi. 5, fig. 8, c; Cameron, 1932, Fauna Brit. India, Staphyl., 3: 318; Silvestri, 1947, Arch. Zool. Ital., 31: 127, fig. 1 (1-7). Termitodiscus heimi var. vicinior Silvestri, 1947, Arch. Zool. Ital., 31: 127, pi. 2 (Barkuda Island, India; Odontotermes sp.; Silvestri coll.). Material examined. One cotype, Wallon, Ahmednagar District, India (Heim); 2 specimens, Sangamner, India; 1 specimen, Kendal, India; all in Bernhauer coll. (Chicago Natural History Museum). Termitodiscus escherichi Wasmann Termitodiscus escherichi Wasmann, 1911, in Escherich, Termitenleben auf Ceylon, p. 231 (Peradeniya, Ceylon; Odontotermes redemanni Wasmann; Wasmann coll.); 1912, Zeitschr. wiss. Zool., 101: 94; 1916, Zool. Jahrb. Syst., 39: 181, pi. 4, fig. 9, pi. 5, fig. 9, a; Cameron, 1932, Fauna Brit. India, Staphyl., 3: 318. Termitodiscus escherichi var. piceus Wasmann, 1916, Zool. Jahrb. Syst., 39: 181 (Peradeniya, Ceylon; Odontotermes ceylonicus Wasmann; Wasmann coll.). Material examined. One cotype (Buttel), Peradeniya, Ceylon, in Bernhauer coll. (Chicago Natural History Museum). Fig. 38. (a) Termilodiscus heimi Wasmann. (b) Termitogerrtis bur- geoni Bernhauer. (c) Discoxenus lepisma Wasmann. 261 262 FIELDIANA: ZOOLOGY, VOLUME 40 Termitodiscus butteli Wasmann Termitodiscus butteli Wasmann, 1916, Zool. Jahrb. Syst., 39: 181, pi. 4, fig. 10, pi. 5, fig. 10, a (Peradeniya, Ceylon; Odontotermes obscuriceps Wasmann; Wasmann coll.); Cameron, 1932, Fauna Brit. India, Staphyl., 3: 319. Material examined. One cotype, Peradeniya, Ceylon, in Bern- hauer coll. (Chicago Nat. Hist. Mus.). Termitodiscus minutus Cameron Termitodiscus minutus Cameron, 1926, Trans. Ent. Soc. London, 74: 171 (Dehra Dun, India; "in nest of termites;" Cameron coll.); 1932, Fauna Brit. India, Staphyl., 3: 319. Termitodiscus splendidus Wasmann Termitodiscus splendidus Wasmann, 1899, Deuts. Ent. Zeitschr., 43: 401 (Shivyre, Natal; Odontotermes vulgaris Haviland; Wasmann coll.); 1912, Zeitschr. wiss. Zool., 101 : 94, pi. 5, fig. 5. Termitodiscus braunsi Wasmann Termitodiscus braunsi Wasmann, 1912, Zeitschr. wiss. Zool., 101: 94 (Botha- ville, Orange Free State; Odontotermes transvaalensis Sjostedt; Wasmann coll.). Termitodiscus transvaalensis Silvestri Termitodiscus transvaalensis Silvestri, 1947, Arch. Zool. Ital., 31: 129, text fig. 3 (Transvaal; Termes angustatus Rambur; Silvestri coll.). The host is Odontotermes angustatus Rambur. TERMITOGERRUS Bernhauer Termitogerrus Bernhauer, 1932, Rev. Zool. Afr., 22: 157 (as subgenus of Termitodiscus). Type species: Termitogerrus burgeoni Bernhauer. Lissodiscus GrassS and Poisson, 1940, Bull. Soc. Ent. France, 45: 84 (as sub- genus of Termitodiscus); Silvestri, 1947, Arch. Zool. Ital., 31: 131. Type species: Lissodiscus lepidulus Grasse and Poisson. New synonymy. Coloration piceous to dark brown. Body-form narrower and more convex than in Termitodiscus. Pronotum glabrous, elytra subglabrous. Head broad and moderately short; antennal fossae medium-sized, rather widely separated; genae deeply grooved between eyes and maxillae for reception of second and third antennal segments, thus giving rise to a distinctive plate adjacent to maxillae. Antennae 11-segmented (possibly 10-segmented in some species); basal segment short and stout; second and third short and more slender; segments 4-10 short and incrassate; 5-10 transverse and compressed. Gula broad and moderately long; submentum short, mentum large, trapezoidal, labial palpi 3-segmented. Galea and lacinia subequal in length; maxillary palpi long and slender. Pronotum slightly less than twice as long as broad; large, strongly convex, sides and apex continuously arcuate, margin not flared, basal angles produced. Outer apical angles of elytra very strongly produced; apex of elytra appearing SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 263 deeply emarginate. Prosternum thinly keeled. Anterior coxae very broad and flat, their ventral surfaces feebly convex. Mesosternum almost vertical at middle (only a short triangular area visible); not produced between middle coxae, which are widely separated by a broad convex metasternal process that extends forward to their anterior margins. Hind coxae very large; about twice as broad as long. Remarks. Termitogerrus was proposed by Bernhauer (1932a) as a subgenus of Termitodiscus, a fact that was apparently overlooked by Grass and Poisson (1940), who described Lissodiscus as a sub- genus of Termitodiscus, and by Silvestri (1947), who elevated Lissodiscus to generic rank. The genus is comprised of African species that live with Macrotermes. They are easily distinguished from Termitodiscus by the vertical mesosternum and the very large metasternal process between the widely separated mesocoxae. Additional differences are the narrower, more convex, glabrous pronotum, the larger hind coxae, the very broad, flat anterior coxae, and the 11-segmented antennae. Inasmuch as only two species of Termitogerrus have been studied, no attempt will be made to present species diagnoses. Silvestri's detailed figures should be examined by one attempting to identify species of the genus. Termitogerrus burgeon! Bernhauer. Figure 38, 6. Termitodiscus (Termitogerrus) burgeoni Bernhauer, 1932, Rev. Zool. Afr., 22: 157 (Haut-Uele, Moto Tora, Belgian Congo; Termes natalensis Haviland; Chicago Natural History Museum). Material examined. Type and one cotype (Bernhauer coll.). In addition, 13 specimens, Rwindi Camp, Belgian Congo, collected May 6, 1948, by Alfred E. Emerson, with Macrotermes natalensis Haviland. Termitogerrus lepidulus Grass and Poisson, new comb. Termitodiscus (Lissodiscus) lepidulus Grasse and Poisson, 1940, Bull. Soc. Ent. France, 45: 84, text figs. 1-8 (Kouibli and DananS, C6te d'lvoire, French West Africa; Bellicositermes natalensis Haviland). Lissodiscus lepidulus Grasse and Poisson, Silvestri, 1947, Arch. Zool. Ital., 31: 132. Material examined. Specimens from Bendija, Liberia, collected by W. M. Mann (U.S. Nat. Mus.); with Macrotermes natalensis Haviland. Termitogerrus habilis Silvestri, new comb. Lissodiscus habilis Silvestri, 1947, Arch. Zool. Ital., 31 : 132, text fig. 4 (Con- akry, French Guinea, French West Africa; Bellicositermes bellicosus Smeathman; Silvestri coll.). 264 FIELDIANA: ZOOLOGY, VOLUME 40 Termitogerrus quaerens Silvestri, new comb. Lissodiscus quaerens Silvestri, 1947, Arch. Zool. Ital., 31: 134, text fig. 5 (Thies, Senegal, French West Africa; Bellicositermes bellicosus Smeathman; Silvestri coll.). Termitogerrus bellicosi Silvestri, new comb. Termitodiscus bellicosi Silvestri, 1905, Redia, 3: 348, fig. (Adi Ugri, Eritrea; Bellicositermes bellicosus Smeathman; Silvestri coll.); Wasmann, 1912, Zeitschr. wiss. Zool., 101: 94. Lissodiscus bellicosi Silvestri, 1947, Arch. Zool. Ital., 31 : 135, text fig. 6. DISCOXENUS Wasmann Discoxenus Wasmann, 1904, Zool. Jahrb., Suppl., 7: 655; 1912, Zeitschr. wiss. Zool., 101: 92; 1916, Zool. Jahrb. Syst., 39: 176; Cameron, 1932, Fauna Brit. India, Staphyl., 3: 315. Type species: Discoxenus assmuthi Was- mann. Head longer and narrower than in the other genera. Antennal fossae longer, broader, and deeper than in the other genera to receive the relatively large basal antennal segment. Antennae 11-segmented, relatively long, projecting well beyond front margin of pronotum; scape large, fusiform; segment 2 small, more or less set in apex of scape; segments 3-11 forming a long, compact fusiform "club." Pronotum about twice as broad as long, basal angles broadly rounded, base feebly emarginate, its margin flared. Outer apical angles of elytra not produced, apices a little emarginate medially, straight toward the sides. Mesosternum almost horizontal. Mesocoxae large, narrowly separated by long metasternal process which broadens at apex before meeting the very broad mesosternal process. KEY TO SPECIES OF DISCOXENUS WASMANN (from Wasmann, 1916) 1. Antennae with penultimate segment much longer than broad; slender terminal segment almost awl-shaped, three times as long as broad; elytral disk with transverse rows of numerous long yellow setae 2 Antennae with penultimate segment not longer than broad; terminal segment bluntly cone-shaped, twice as long as broad; segments 4-7 transverse, the visible part of antennae only as long as pronotum; elytra only one-half as long as pronotum; elytral disk very sparingly setose; reddish-brown, elytra darker; 1.8 mm. in length crassicornis Wasmann 2. Larger species, almost 2.5 mm. in length; elytra about two-thirds as long as pronotum; antennae very robust, the visible part longer than pronotum; segments 4-7 transverse; terminal segment very small and slender, four times as long as broad; coloration yellowish-brown lepisma Wasmann Smaller species, less than 2 mm. in length; elytra about one-half as long as pronotum; visible part of antennae not longer than pronotum; terminal segment only three times as long as broad 3 3. Antennae moderately pointed; segments 5-7 not transverse; elytral disk with a few almost recumbent yellow setae; color pitch-brown; length, 1.8-1.9 mm. assmuthi Wasmann Antennae strongly pointed; segments 4-7 transverse; elytral disk thickly covered with erect yellow setae; reddish-brown with pitch-brown elytra; length, 1.8 mm acuticornis Wasmann SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 265 Discoxenus lepisma Wasmann. Figure 38, c. Discoxenus lepisma Wasmann, 1904, Zool. Jahrb., Suppl., 7: 656 (Kendal, Ahmednagar District, India; Odontotermes obesus wallonensis Wasmann; Wasmann coll.); 1916, Zool. Jahrb. Syst., 39: 178, pi. 4, fig. 7; Cameron, 1932, Fauna Brit. India, Staphyl., 3: 316, pi. 3, fig. 6. Material examined. One specimen, Dehra Dun, India (Bern- hauer coll.; Chicago Nat. Hist. Mus.). Discoxenus assmuthi Wasmann Discoxenus assmuthi Wasmann, 1904, Zool. Jahrb., Suppl., 7: 656 (Khandala, Bombay, India; Odontotermes obesus Rambur; Wasmann coll.); 1916, Zool. Jahrb. Syst., 39: 178; Cameron, 1932, Fauna Brit. India, Staphyl., 3:316. Discoxenus crassicornis Wasmann Discoxenus crassicornis Wasmann, 1916, Zool. Jahrb. Syst., 39: 178, pi. 4, fig. 6 (Peradeniya, Ceylon; Odontotermes redemanni Wasmann; Wasmann coll.); Cameron, 1932, Fauna Brit. India, Staphyl., 3: 317. Discoxenus acuticornis Wasmann Discoxenus acuticornis Wasmann, 1916, Zool. Jahrb. Syst., 39: 178 (Pera- deniya, Ceylon; Odontotermes obscuriceps Wasmann; Wasmann coll.); Cameron, 1932, Fauna Brit. India, Staphyl., 3: 317. Subfamily PYGOSTENINAE Pygostenini: Fauvel, 1899, Rev. d'Ent., 18: 5. Pygosteninae: Wasmann, 1902, Verh. Deuts. Zool. Ges., 1902: 91; 1916, Zool. Jahrb. Syst., 39: 170; 1926, Ent. Mitteil., 15: 113; Cameron, 1932, Fauna Brit. India, Staphyl., 3: 320. The subfamily Pygosteninae is probably most noteworthy for including both myrmecophilous and termitophilous species, a respect in which it is perhaps unique among staphylinid groups. The host situation, with minor exceptions, seems to be this: Ants of the subfamily Dorylinae (driver-ants) are hosts of some species through- out the range of the subfamily in the Ethiopian and Indomalayan regions; fungus-growing termites are hosts of most Indomalayan species of Doryloxenus. According to present evidence, African components of the subfamily are much more numerous, and ex- tremely few of these, if indeed any, are truly termitophilous. The Ethiopian species of Doryloxenus, the first of the genus to be dis- covered, are guests of doryline ants, whereas their Asiatic counter- parts in India, Ceylon, and Java occur with Macrotermes and Odontotermes. 266 FIELDIANA: ZOOLOGY, VOLUME 40 Inasmuch as the Pygosteninae should be considered from a much broader viewpoint than is possible in this monograph, no aspect of the problem is included. Mr. David Kistner of Chicago is now engaged in comprehensive studies on the Pygosteninae, and the results will be published in the near future. Fig. 39. Doryloxenus brevicornis Cameron. Species of Pygosteninae recorded from termite nests are in- cluded in lists in this paper. Subfamily TRICHOPSENIINAE Trichopsenii LeConte and Horn (Group II, Tribe Tachyporini), 1883, Smithson. Misc. Coll., no. 507, p. 100. Trichopsenini Eichelbaum (as a tribe of Tachyporinae), 1909, Mem. Soc. Ent. Belg., 17: 196; Wasmann, 1916, Zool. Jahrb. Syst., 39: 196. Schizelythrinae Kemner, 1925, Ent. Tidskr., 46: 122. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 267 Trichopseniinae Seevers, 1941, Ann. Ent. Soc. Amer., 34: 320; Borgmeier, 1950, Rev. Ent., 21 : 642. A discussion of certain aspects of the evolutionary history of the Trichopseniinae is presented in an earlier section of this paper. KEY TO GENERA OF TRICHOPSENIINAE 1. Elytra split for almost their entire length, lateral half of each diverging strongly (Java) Schizelythron Kemner Elytra not split lengthwise 2 2. Pronotum extremely large, twice as broad as long, its anterior and lateral margins continuously arcuate; head and forelegs not visible from above; body-form limuloid 7 Pronotum not as above, its anterior angles distinct (obsolescent in Trichop- senius) 3 3. Antennal segments 3-10 campanulate; abdomen ovoid, strongly inflated (Australia) Mastopsentus Seevers Antennal segments 3-10 not campanulate; abdomen at most feebly inflated . 4 4. Inner margins of elytra beveled, forming a median V-shaped groove (United States) Xenisliisa LeConte Inner margins of elytra not beveled 5 5. Pronotum strongly bi-impressed ; metasternum behind mesocoxae about four- sevenths as long as metasternal width from median line to lateral margin (Neotropical Region) Megaxenistusa Seevers Pronotum not impressed; metasternum behind mesocoxae as long as or longer than metasternal width from median line to lateral margin 6 6. Sides of abdomen parallel (abdomen of physogastric individuals uniformly swollen); metasternum behind mesocoxae about equal to metasternal width from median line to lateral margin (United States) Trichopsenius Horn Sides of abdomen strongly convergent apically (only basal abdominal seg- ments physogastric); metasternum behind mesocoxae one-third greater than metasternal width from medial line to lateral margin (Neotropical Region). Rhinotermopsenius Seevers 7. Anterior margin of pronotum distinctly angulate; pronotum and elytra with- out macrosetae (Malaysia, Micronesia) Hamitopseni us Wasmann Anterior margin of pronotum not angulate; pronotum and elytra conspicu- ously setose (Neotropical Region) Termitopsenius Wasmann TRICHOPSENIUS Horn Trichop8enius Horn, 1877, Trans. Amer. Ent. Soc, 6: 88; Seevers, 1941, Ann. Ent. Soc. Amer., 34: 323. Type series: Trichopsenius depressus LeConte. Head strongly deflexed, partly concealed by pronotum. Antennae inserted medial to eyes; 1 1 -segmented ; segments 2-5 subquadrate, 6-10 incrassate. Prono- tum one-half broader than long; sides slightly deflexed; anterior and lateral margins continuously arcuate; base arcuate, posterior angles rounded. Elytra as broad as pronotum and a little longer. Metasternal length equal to width of metasternum from median line to lateral margin. Metasternal plates covering most of trochanters and proximal part of femora. Sides of abdomen subparallel ; stenogastric to slightly physogastric. Aedeagus of male remarkably large, oc- 268 FIELDIANA: ZOOLOGY, VOLUME 40 cupying almost all of abdomen posterior to third segment; distal sclerite of its lateral lobe long and slender, protruding from abdomen. KEY TO SPECIES OF TRICHOPSENIUS 1. Apical margin of female eighth tergite with a series of setigerous processes. californicus, new sp. Apical margin of female eighth tergite without setigerous processes 2 2. Pronotum two-thirds broader than long; Japan japonicus, new sp. Pronotum one-third to one-half broader than long 3 3. Posterior tibiae with two rows of about ten very long, curved bristles (one- third to one-half as long as tibiae) longipes Seevers Posterior tibiae with scattered, relatively short setae 4 4. Eighth tergite of female not incised xenoflavipes Seevers Eighth tergite of female with apical incision 5 5. Eighth tergite of female carinate laterally; elytral setae about one-fourth as long as elytra depressus LeConte Eighth tergite of female not carinate; elytral setae about one-half as long as elytra frosti Seevers Trichopsenius depressus LeConte. Figure 40, a, b. Hypocyptus depressus LeConte, 1863, Smithson. Misc. Coll., 6: 30 (Louisiana; no host cited; Museum of Comparative Zoology). Trichopsenius depressus LeConte, Horn, 1877, Trans. Amer. Ent. Soc, 6: 88, pi. 1, figs. 1, 2; Seevers, 1941, Ann. Ent. Soc. Amer., 34: 323, pi. 1, figs. 3, 7, 11, pi. 2, figs. 27, 29; 1945, Pan. Pac. Ent., 21: 69, text fig. 4. Material examined. Type from Louisiana; one male and one female from Columbus, Texas; one specimen from Crescent City, Florida; all in LeConte collection (Mus. Comp. Zool.). A series from Gainesville and Jacksonville, Florida, and Memphis, Ten- nessee; all from nests of Reticulitermes virginicus Banks (Chicago Nat. Hist. Mus.). The previously recorded host, Reticulitermes hageni Banks (Seevers, 1945, p. 69) is incorrect. Color testaceous. Pronotum with 5 or 6 setae on lateral margins; disk with a hexagon of 6 setae medially, a group of 3 on each side, an irregular, subapical row of about 6, and a basal row of about 8. Elytra with three irregular transverse rows of about 5 bristles each. Tergites with an apical row of long bristles inter- spersed with short ones. Antennae of both sexes similar; segments 3 and 4 longer than broad, 5 and 6 subquadrate, 7-10 transverse. Anterior border of pronotum feebly emarginate at middle. Female with seventh tergite somewhat produced, angulate at middle; eighth tergite notched medially and elevated laterally to form a strong semicircular carina on each side; lateral plates of ninth segment broad, compressed, rounded at apex. Male with seventh tergite feebly angulate at middle; eighth tergite arcuate, its apex rounded, the converging sides forming a right angle. / ** Fig. 40. Trichopsenius depressus LeConte: (a) male with aedeagus extruded; (b) female, (c) Xenistusa hexagonalis Seevers. (d) Rhinotermopsenius saltatorius Seevers. 269 270 FIELDIANA: ZOOLOGY, VOLUME 40 Length, 1-1.5 mm. Legs relatively short: total lengths 0.55 mm., 0.72 mm., and 0.85mm.; femora 0.25 mm., 0.28 mm., and 0.3 mm.; tibiae 0.17 mm., 0.22 mm., and 0.26 mm.; tarsi 0.14 mm., 0.23 mm., and 0.35 mm. Trichopsenius xenoflavipes Seevers Trichopsenius xenoflavipes Seevers, 1945, Pan. Pac. Ent., 21: 70, text fig. 3 (Crawford County, near Wyandotte Cave, Indiana; Reticulitermes flavipes Kollar; Chicago Natural History Museum). Material examined. A series from Olney, Urbana, and Spring- field, Illinois; all with Reticulitermes flavipes Kollar (Chicago Nat. Hist. Mus.). Testaceous. Chaetotaxy as in depressus except that the setae of elytra and abdomen are one-third longer (elytral bristles about one-half as long as elytra). Antennal segments 3-7 of male longer than broad, 8-10 transverse; segments 3-6 of female longer than broad, 7-10 transverse, a little broader than in male. An- terior margin of pronotum not emarginate at middle. Female abdomen with seventh tergite evenly arcuate; eighth arcuate, not notched; lateral plates of ninth segment slender, tapering to a very acute apex. Legs as in depressus. Length, 1-1.5 mm. Trichopsenius frosti Seevers Trichopsenius frosti Seevers, 1945, Pan. Pac. Ent., 21 : 70, text fig. 5 (Sherborn, Massachusetts; Reticulitermes flavipes Kollar; Chicago Natural History Museum). Material examined. A series from Boston, Forest Hills, and Wellesley, Massachusetts (U. S. Nat. Mus.). Color testaceous. Chaetotaxy as in xenoflavipes. Female abdomen with seventh tergite very feebly arcuate, faintly sinuate on each side of middle; eighth tergite with a broad, deep incision, but otherwise unmodified. Male seventh tergite moderately and uniformly arcuate; eighth tergite broadly and obtusely arcuate; the blade-like distal end of lateral lobes of aedeagus rather stout. Length, 1.25 mm. Trichopsenius longipes Seevers Trichopsenius longipes Seevers, 1945, Pan. Pac. Ent., 21: 71 (Skull Valley, Tooele County, Utah; Reticulitermes sp.; United States National Museum). Material examined. A series from San Antonio Canyon, near Camp Baldy, San Bernardino County, California ; with Reticulitermes hesperus Banks (Chicago Nat. Hist. Mus.). Color flavo-testaceous. Chaetotaxy and antennae as in T. depressus. An- terior margin of pronotum bisinuate. Abdomen moderately physogastric, uni- formly dilated, its sides scarcely converging posteriorly. Seventh and eighth tergites of female simple; lateral plates of ninth segment rather slender, apex SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 271 acute. Tibiae with two rows of long, fine, curved bristles, those of hind legs much longer. Length, 1.9 mm. (stenogastric specimens) and 2.8 mm. (physogastric speci- mens). Legs relatively long: total lengths 0.65 mm., 0.92 mm., and 1.1 mm.; femora 0.27 mm., 0.35 mm., and 0.35 mm.; tibiae 0.21 mm., 0.3 mm., and 0.35 mm.; tarsi 0.17 mm., 0.29 mm., and 0.44 mm. Trichopsenius californicus, new species Distinguished from other species by its distinctive eighth female tergite. Testaceous. Chaetotaxy as in depressus. Pronotum a little more than one- half broader than long (77 : 50); broadest one-fifth the distance from base to apex; sides in front of broadest point converging rather strongly and continuously arcuate with apex; basal angles obtusely rounded; base almost straight. Hind tibiae with one row of about 10 fine setae that are one-third as long as the tibia. Legs long: hind leg 1 mm., hind femur 0.3 mm., hind tibia 0.3 mm., hind tarsus 0.4 mm. Apex of eighth female tergite with a series of setigerous processes: a long, median process, bifid at apex, flanked on each side by a slightly shorter process, bifid or not; lateral to these processes are two very short, tubercle-like processes. Male abdomen not seen. Abdomen stenogastric. Length, 1.4-1.5 mm. Holotype a female from Stanford University, California; collected April 5, 1902; with Reticulitermes hesperus Banks. In collection of Chicago Natural History Museum. Paratypes. Two females, same data as type. Trichopsenius japonicus, new species This is the first palaearctic species of this genus to be recorded. It is probably more closely related to longipes than to any other American species. It resembles the latter in having two rows of long, fine setae on the lateral margins of the hind tibiae, but the setae are less than one-third as long as the tibia, and the rows consist of only 4 macrosetae. The pronotum is two-thirds broader than long, differing from all American species in this respect. The lateral plates of the ninth abdominal segment are unusually short, broad, and stout. Color rufo-testaceous. Chaetotaxy as in depressus. Pronotum two-thirds broader than long (70 : 43); broadest two-fifths of distance from base to apex; sides strongly converging in front of broadest point; anterior angles obsolete; sides behind broadest point continuously arcuate with base; apex almost straight. Hind leg 0.9 mm., hind femur 0.3 mm., hind tibia 0.3 mm., hind tarsus 0.31 mm. Eighth female tergite unmodified. Length, 1.35 mm.; width, 0.6 mm. 272 FIELDIANA: ZOOLOGY, VOLUME 40 Holotype. A female from Kinschin, Japan, collected by M. Yano; with Reticulitermes speratus Kolbe. In Wasmann collection, Natuurhistorisch Museum, Maastricht, The Netherlands. Paratype. One female, same data as type; in Chicago Natural History Museum. XENISTUSA LeConte Xenistusa LeConte, 1880, Trans. Amer. Ent. Soc, 8: 167; Seevers, 1941, Ann. Ent. Soc. Amer., 34: 323. Type species: Xenistusa cavernosa LeConte. Head and thorax slender; mesothorax, metathorax, and abdomen inflated slightly; sides of thorax and abdomen with some non-sclerotized integument. Head prognathous, only base of occiput concealed by pronotum; clypeus very short; labrum broadly and moderately emarginate. Gula broad, trapezoidal, gular sutures apparently terminating at posterior tentorial pits, as sides of sub- mentum are not delimited from postgenae. Segment 2 of labial palpi globose, segment 3 without terminal spicule. Pronotum one-half broader than long, not much broader than head. Elytra with a very narrow, slit-like groove near each lateral margin, and with the inner margins beveled, giving rise to a distinctive V-shaped groove between them. Elytral epipleura bent at right angles to elytra a short distance from the sides, which thus appear carinate. Anterior coxae slender at base, clavate distally, the ventral surfaces somewhat convex and the upper surfaces nearly flat. Meso- sternum and its process very short; metasternal process long and slender. Meta- sternum long, strongly convex, its sides almost vertical; metasternal plates very narrow in front and not covering the femora, but expanding posteriorly to cover the medial half of the trochanters in repose. Male with terminal abdominal segments as follows: tergite 8 with strongly arcuate apex; sternite 8 with deeply emarginate base and arcuate apex; tergite 9 smaller than 8 and concealed by it, articulating at sides with large, elongated, lateral plates of ninth segment which protrude from apex of abdomen; these plates articulate at base with long, slender apodemes that extend anteriorly as far as segment 4 and are visible through the translucent sternites. Sternite 9 small, concealed by 8. Bulbous median lobe of aedeagus with several sclerites; lateral aedeagal lobes also with several sclerites, the apical one shorter and broader than in Trichopsenius and not protruding from abdomen. Female with sternite 8 and tergite 8 shorter than those of male. Lateral plates of ninth segment very broad at base, the apical one-half tapered. Xenistusa cavernosa LeConte Xenistusa cavernosa LeConte, 1880, Trans. Amer. Ent. Soc, 8: 167 (Co- lumbus, Texas; no host given; Museum of Comparative Zoology); Schwarz, 1889, Proc. Wash. Ent. Soc, 1: 1 (host, Termes flavipes). Xenistusa fossata LeConte, 1880, Trans. Amer. Ent. Soc, 8: 167 (Columbus, Texas; no host given; Museum of Comparative Zoology); Schwarz, 1889, Proc. Wash. Ent. Soc, 1: 1 (host, Termes flavipes). New synonymy. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 273 Xenistusa pressa LeConte, 1880, Trans. Amer. Ent. Soc, 8: 167 (Columbus, Texas; no host given; Museum of Comparative Zoology); Schwarz, 1889, Proc. Wash. Ent. Soc, 1: 1 (host, Termes flavipes). New synonymy. Material examined. The types of LeConte's three species. The three species of Xenistusa proposed by LeConte were based on three specimens collected by E. A. Schwarz from a colony of Reticulitermes in a log near Columbus, Texas. Schwarz (1889) later expressed doubt, after examining the mounted specimens, that they represented three species. There have been no subsequent reports on this problem, probably because no more material has become available. It has seemed improbable that three specimens of a genus from the same termite nest would belong to three species, yet LeConte described appreciable differences in pronotal structure. When I had an opportunity to examine briefly LeConte's specimens, I was surprised to note how different the pronota actually appear. The differences very likely resulted from distortion of the thin integu- ments when the specimens were dried and probably do not represent species characters. The pronotum of the cavernosa type specimen, described by LeConte as profoundly bilobed, has a deep, median, V-shaped sulcus. The pronotum of the fossata type specimen, described as having a large, very deep, triangular excavation from base to apex, has a very broad, V-shaped sulcus. The pronotum of the pressa type specimen is very feebly depressed. It is my conclusion that the three conditions resulted from different degrees of distortion, but as the medial depressions are remarkably sym- metrical, one cannot state with finality that they are artifacts. The ultimate conclusion must be based on a study of additional specimens, preferably preserved in alcohol. Minor differences in the LeConte specimens, such as variations in lengths of elytral bristles, may be sexual differences such as were noted in the case of hexagonalis. Schwarz (1889) made the interesting comment that the living beetles, with their cylindrical bodies, greatly extended abdomens, and peculiar mode of locomotion, resembled the termites much more than one would suspect from dried specimens. Xenistusa hexagonalis Seevers. Figure 40, c. Xenistusa hexagonalis Seevers, 1941, Ann. Ent. Soc. Amer., 34: 324, pi. 1, figs. 6, 10, pi. 2, figs. 19-26 (Gainesville, Florida; Reticulitermes hageni Banks; Chicago Natural History Museum). The host has been re- determined as Reticulitermes virginicus Banks. 274 FIELDIANA: ZOOLOGY, VOLUME 40 Male with light reddish-brown head and thorax and paler abdominal sclerites (segments 3-6 testaceous, the others darker). Female generally darker than male, with head and thorax brown and abdominal sclerites light reddish-brown. Vertex of head broadly and shallowly concave; with 2 setae between antennae and 2 on margin of clypeus. Pronotum with 6 setae in form of hexagon at center of disk, 3 on each side forming a large triangle, 6 on anterior border, 4 on lateral borders, and 4 on basal border. Elytra rather densely setose, the moderately long, pale hairs arranged in 6-8 irregular, longitudinal rows of 8-10 hairs. Antennal scape somewhat compressed; segment 2 short, moderately globose; segments 3-10 increasing in width; segment 11 short, broadly transverse. Pronotum broadest a short distance behind front margin; apex bisinuate, anterior angles broadly rounded; sides converging posteriorly, forming a continuous arc with base; pronotal disk not impressed. Elytra slightly broader than pronotum and nearly twice as long. Segments 1-4 of front tarsi subequal, segment 5 longer; basal segment of middle tarsi subequal to segments 2-5 combined, segment 5 as long as segments 2-4 combined; basal segment of hind tarsi longer than segments 2-5 combined, 2-4 decreasing in length, segment 5 equal to segments 3-4 combined. Males and females about equal in length but the latter more robust, and the head, pronotum, and elytra larger. Length, 1.9-2.1 mm. Female head 0.47 mm. broad; pronotum 0.54 mm. broad and 0.35 mm. long; elytra 0.6 mm. broad and 0.65 mm. long. Male head 0.42 mm. wide; pronotum 0.48 mm. broad and 0.33 mm. long; elytra 0.52 mm. broad and 0.55 mm. long. MEGAXENISTUSA Seevers Megaxenistusa Seevers, 1945, Pan-Pac. Ent., 21: 66. Type species: Mega- xenistusa rhinotermitis Seevers. The closest relative of this genus may be Xenistusa, to which it bears a resemblance. It is distinguished from that genus by the form and proportions of the metasternum and metepisternum, and by its unmodified elytra. Body-form cylindrical. Head strongly compressed, deflexed. Antennal scape flat, its medial surface concave; segments 2-10 sub cylindrical. Gula sclerotized; mentum trapezoidal. Labial palpi with segment 2 asymmetrical, strongly com- pressed. Maxillary palpi long and slender, segment 2 arcuate, segment 3 fusiform, segment 4 conical. Pronotum robust, transverse, not margined; its disk bi- impressed. Elytra as broad as pronotum but longer; their surfaces somewhat irregular, with broad, shallow impressions; sutural margins not beveled. Mesosternum short; mesocoxae slightly separated by carinate processes of mesosternum and metasternum. Metasternum relatively short and broad; its length behind mesocoxae about seven-tenths the width of metasternum from median line to metepisternum along posterior margin; each half of metasternum strongly oblique (a cross section of metasternum would be markedly V-shaped); metasternal plates moderate in size, not much larger than posterior trochanters; metepisternum large, four-fifths as broad at caudal margin as long. Femora broad, compressed; tibiae somewhat less so. Tarsi 5-segmented, basal segment of middle and hind legs extremely long, exceeding the other segments combined. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 275 Megaxenistusa rhinotermitis Seevers Megaxenistusa rhinotermitis Seevers, 1945, Pan-Pac. Ent., 21: 68 (Kartabo, British Guiana; Rhinotermes marginalia Linnaeus; Chicago Natural History Museum). Color reddish-brown; elytra and areas of pronotum darker. Head with 2 frontal and 4 clypeal setae. Pronotum with 4 setae on anterior margin, one sub- humeral seta, and 4 longitudinal rows of setae on each half^a discal row of 4, a row of 3 in impressed area, a pair lateral to impression, and a marginal row of 3. Elytra with 5 or 6 longitudinal rows of hairs. Abdomen sparsely and regularly setose, with scattered long hairs among short ones and marginal rows of long bristles. Second antennal segment short; segments 3-6 subequal, longer than broad; 7-10 shorter, incrassate; segment 11 as long as 10 but narrower. Pronotum with two large oval impressions on the disk, extending forward from base about two-thirds the length of pronotum; anterior margin arcuate, sides arcuate, con- verging moderately in front; base bisinuate. Length, 3 mm. TERMITOPSENIUS Wasmann Termitopsenius Wasmann, 1902, Boll. Mus. Zool. Torino, 17, no. 427, p. 4; Silvestri, 1903, Redia, 1: 201; 1947, Arch. Zool. Ital., 31: 144. Type species: Termitopsenius limulus Wasmann. Pulicipsenius Seevers, 1941, Ann. Ent. Soc. Amer., 34: 327. Type species: Pulicipsenius acanthoscelis Seevers. New synonymy. Body-form limuloid. Pronotum very large, completely covering the head; strongly deflexed anteriorly and laterally, the lateral and apical margins con- tinuously arcuate. Head broad and short, dorso-ventrally compressed; so oriented with respect to pronotum that its dorsum is ventral and its mouth parts are directed caudad. Eyes prominent. Vertex broad, antennae inserted near eyes some distance from anterior tentorial pits; clypeus extremely short; genae in front of eyes deeply excavated to receive basal antennal segment. Antennae 11-seg- mented; scape compressed, its inner surface concave; segments 3-10 incrassate; terminal segment membranaceous, usually reduced in size. Mandibles with a medial tooth on inner margin, but not toothed externally. Gula and submentum very short due to shortening of head and its position with respect to pronotum. Mesosternum very short, its process between middle coxae very slender, slightly carinate. Metasternum extremely large, its metasternal plates large. Abdomen acuminate. Termitopsenius limulus Wasmann. Figure 41, b. Termitopsenius limulus Wasmann, 1902, Boll. Mus. Zool. Torino, 17, no. 427, p. 5 (Posadas, Misiones, Argentina; Capritermes opacus Hagen; Wasmann coll.); Silvestri, 1903, Redia, 1: 201, pi. 6, figs. 283, 284; Bruch, 1938, Notas Mus. La Plata, 3: 155; Silvestri, 1947, Arch. Zool. Ital., 31: 146, fig. 9. Material examined. One specimen from Rio Negro, Parana, Brazil; with Neocapritermes opacus Hagen (ex Reichensperger coll.). 276 FIELDIANA: ZOOLOGY, VOLUME 40 Color rufo- testaceous, pronotal disk darker, head reddish-brown. Head with 2 vertexal setae and 2 on clypeus. Pronotum with two transverse subapical rows of 4 setae each, 2 or more setae on lateral margins, and disk with two longitudinal rows of 3 and 2 setae, respectively. Pronotal disk with an extensive non-setose area. Elytra with four rows of 2, 3, 3, 4 long, fine setae. Tergites 3-7 and sternites 3-8 with 6 long setae on apical margin. Third tergite pubescent, third sternite with short pale semi-erect hairs; other sclerites glabrous. Antennal scape two- thirds as broad as long, its inner surface concave; scape too broad to fit in genal groove. Antennal segments with the following relative lengths: 18 : 6 : 6 : 4 : 5 : 6:8:8:8:8:6; terminal segment small, membranaceous; segments 3-5 sub- equal in width, segment 6 much broader than 5, segments 6-10 scarcely incrassate. Hind tibiae with 3 large spines on outer margin and 3 at apex, as well as several short spines. Termitopsenius acanthoscelis Seevers, new comb. Figure 42. Termitopsenius acanthoscelis Seevers, 1941, Ann. Ent. Soc. Amer., 34: 329, pi. 1, figs. 2, 4, 5, 9, 14, pi. 2, figs. 15-18, 28 (Kartabo, British Guiana; Rhinotermes marginalis Linnaeus; Chicago Natural History Museum). Distinguished from limulus by smaller size; differences in chaeto- taxy of pronotum, elytra, and abdomen; stouter spines of hind tibiae; and proportions of antennal segments. Color testaceous. Head with 2 vertexal setae and 2 on clypeus. Pronotum with 4 rows of. very long, slender, pale setae on each half comprised of 5, 3, 2, 2 setae, respectively, beginning with medial row. Elytra with four rows of 2, 3, 3, 3 setae. Tergites 3-7 with 4 very long setae in apical row; tergite 8 with 4 long setae and 2 shorter ones medially. Sternites 3-8 with 6 very long apical setae, sternite 8 with a few shorter ones in addition. Tergites 3 and 4 with a few short, recumbent hairs, sternite 3 pubescent (especially near middle), sternite 4 slightly so; other abdominal sclerites glabrous. Lateral plates of ninth segment with a few long setae. Hind tibiae with 4 strong spines on outer margin and 3 stout ones at apex; these spines are considerably stouter than those of limulus. Antennae differing from those of limulus in that segments 3-10 are gradually incrassate and the apical segment is not as small or as membranaceous. Length, 1.75 mm. HAMITOPSENIUS Wasmann Hamitopsenius Wasmann, 1916, Zool. Jahrb. Syst., 39: 198. Type species: Hamitopsenius caudatus Wasmann. Hamitopsenius is very closely allied to the neotropical Termitopsenius, from which it may be distinguished by the following characters: anterior margin of pronotum distinctively angulate in front of and below eyes; mandibles with an unusual tooth on outer margin; mesosternal process broader between mesocoxae and more conspicuously elevated as a carina; metasternal process broader and with a rounded apex; antennal segments 3-6 considerably smaller than those of Termi- topsenius; pronotum and elytra without macrosetae. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 277 Fig. 41. (a) Mastopsenius aus- tralis Seevers. (b) Termilopsenius limulm Wasmann. (c) Hamilo- psenius micronesiae Seevers. Wasmann stated that the antennae are 10-segmented in this genus but I have not been able to examine the antennae of caudatus to determine the number of segments. The antennae of micronesiae are 11-segmented. Hamitopsenius caudatus Wasmann HamilopseniM caudatus Wasmann, 1916, Zool. Jahrb. Syst., 39: 201, pi. 4, fig. 17, a-c, pi. 5, fig. 17, d-k (Tandjong Slamat, Sumatra; Amitermea dentatw Haviland; Wasmann coll.). Material examined. One topotype (cotype; U. S. Nat. Mus.). 278 FIELDIANA: ZOOLOGY, VOLUME 40 Coloration brown. Pronotum and elytra glabrous and without macrosetae. Tergites with a row of very long macrosetae on apical margin and a moderate number of pale, recumbent setae elsewhere. Pronotum three-fifths broader than long; its base strongly bisinuate. Sutural length of elytra more than one-third as long as pronotum. Lateral plates of ninth segment stout and long, nearly three-fourths as long as entire abdomen. Hamitopsenius micronesiae, new species. Figure 41, c. Distinguished from caudatus by its paler coloration, distinctive chaetotaxy of tergites, broader pronotum, longer elytra, and much shorter lateral plates of ninth abdominal segments. Color light rufo-testaceous. Pronotum and elytra glabrous in appearance but with extremely fine, sparse punctulation and pubescence; without macrosetae, the elytra with a few short hairs on lateral margins. Tergites 3-6 with a row of 8-10 very long, fine macrosetae about midway between base and apex; basal half of each tergite with sparse vestiture of recumbent and semi-erect hairs, apical half glabrous; seventh tergite with 6 macrosetae; eighth tergite with 4. Sternites with numerous pale, semi-erect hairs; sternites 3-7 with a pair of long, dark, closely placed bristles near each lateral margin. Pronotum three-fourths broader than long; its base feebly bisinuate, basal angles produced. Sutural length of elytra almost three-fourths as long as pronotum; elytral apices almost straight. Lateral plates of ninth abdominal segment only one-fourth as long as abdomen. Hind tibiae with two spines on lateral margin and three at apex. Length, 1.5 mm. Holotype from Koror Island, Palau Islands, Micronesia; collected November 22, 1947, by Henry S. Dybas (Pacific Science Board Expedition); with Prorhinotermes, n. sp. In collection of United States National Museum. Paratypes. Ten specimens, Koror Island, from three colonies of Prorhinotermes, n. sp.; collected November 22, 1947, January 17, 1948, and February 10, 1948, by Henry S. Dybas. In Chicago Natural History Museum, United States National Museum, and Bishop Museum. RHINOTERMOPSENIUS Seevers Rhinotermopsenius Seevers, 1941, Ann. Ent. Soc. Amer., 34: 329. Type species: Rhinotermopsenius saltatorius Seevers. Distinguished from Termitopsenius by its non-limuloid form; smaller pronotum and elytra; physogastric abdomen; longer, more slender, feebly spinose hind legs; labial palpi; and absence of genal grooves. Head small, deflexed, covered in part by pronotum; thorax and basal segments of abdomen somewhat swollen and with areas of white, non-sclerotized integument; SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 279 abdomen acuminate posterior to fourth segment. Head and mouth parts very similar to those of Termitopsenius but genae not grooved in front of eyes. Gula and submentum very short, the former feebly sclerotized. Segment 2 of labial palpi asymmetrical (very similar to that of Trichopsenius). Antennal segments 4-7 longer than those of Termitopsenius. Pronotum moderate in size, trapezoidal in shape, broadest at base, its sides not appreciably deflexed. Elytra moderate Fig. 42. Termitopsenius acanthoscelis Seevers. (a) Ventral view of meta- sternum, illustrating articulation of hind leg (right metasternal plate removed), (b) Diagrammatic section showing head of trochanter, (c) Dorsal view of tro- chanter head in articulation with two processes. in size, its sides nearly straight; epipleura very small. Metasternal lobes rela- tively narrow. Hind legs one-half longer than those of Termitopsenius. Abdomen distinctive; basal segments somewhat physogastric, terminal segments acuminate. Rhinotermopsenius saltatorius Seevers. Figure 40, d. Rhinotermopsenius saltatorius Seevers, 1941, Ann. Ent. Soc. Amer., 34: 330, pi. 1, figs. 1, 8, 12, 13 (Kartabo, British Guiana; Rhinotermes marginalis Linnaeus; Chicago Natural History Museum). Color light reddish-brown, abdomen lighter than foreparts. Head sparsely and microscopically punctate, the punctures usually arranged in small groups; with 2 vertexal setae medial to antennae. Pronotal disk with four longitudinal 280 FIELDIANA: ZOOLOGY, VOLUME 40 rows of 5 setae each, and with several small setae on lateral margins. Tergites and sternites 3-7 with several rows of moderately long, very pale, erect setae; tergite 8 with apical setae and a subapical row of 8 pale setae. Lateral plates of ninth segment very stout and strongly setose. Length, 1.75-2 mm. MASTOPSENIUS Seevers Mastopsenius Seevers, 1945, Pan-Pac. Ent., 21: 64. Type species: Masto- psenius australis Seevers. A very distinctive genus, characterized especially by its large, conical, physogastric abdomen. The antennae, with their cam- panulate intermediate segments, and the elytra, with their con- tinuously arcuate lateral and apical margins, are distinctive. Body robust, fusiform. Head dorso-ventrally compressed; deflexed a little, not covered by pronotum. Antennae elongated, segments 3-9 campanulate. Eyes large. Gula well sclerotized; mentum and submentum strongly transverse; ligula bifid. Pronotum robust, strongly transverse, margined except for medial one-half of base, which is strongly deflexed. Elytra narrower than pronotum, their lateral and apical margins continuously arcuate, the outer apical angles obsolete. Winged. Prosternum broadly tuberculate. Mesosternum large. Meso- and metasternal processes slender, carinate. Metasternum relatively short and narrow behind middle coxae, its width from median line to side margin scarcely greater than width of the large metepimeron. Metasternal plates moderately large, about equal to posterior trochanters. Mesothorax, metathorax, and basal abdominal segments greatly inflated, exposing considerable white, non-sclerotized integument. Abdomen conical. Mastopsenius australis Seevers. Figure 41, a. Mastopsenius australis Seevers, 1945, Pan-Pac. Ent., 21: 65, text figs. 1, 2 (Townsville, Queensland, Australia; Mastotermes darwiniensis Froggatt; Chicago Natural History Museum). Head brown; antennal segments 1-3 flavo-testaceous, segments 4-10 with basal one-third black and apex brown; pronotum dull to shining black, with a reddish median area; elytra piceous to dark brown; abdominal sclerites bright reddish-brown; legs brown, coxae, trochanters, and femora with dusky margins. Head with 2 vertexal setae between antennae. Pronotum with four irregular, transverse rows each with about 10 fine hairs. Elytra with four very irregular rows of fine hairs. Abdominal sclerites sparsely and uniformly clothed with fine, erect hairs. Basal antennal segment strongly clavate, its medial surface concave; segment 2 one-third as long; segments 3 and 4 subequal in length, 5-7 a trifle longer, 8-10 decreasing in length. Length, 5.5-6.2 mm. SCHIZELYTHRON Kemner Schizelythron Kemner, 1925, Ent. Tidskr., 46: 110. Type species: Schizely- thron javanicum Kemner. SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 281 The position of this genus within the subfamily is not apparent at present but it certainly belongs here. Kemner, recognizing its remarkable qualities, proposed a subfamily to receive it, but in so doing ignored the existence of its closest relatives. The genus is well illustrated and described by Kemner and may be recognized at once by its unique elytra, which are split lengthwise from a point near the base. Inasmuch as I have not examined specimens of Schizelylhron, I shall not attempt to characterize it in detail. The mouth parts seem to be typical of the subfamily. The head is moderately deflexed but not covered by the pronotum. The pronotum has a deep pit on each side of the disk and a broad deep impression near the base. The abdomen is more or less cylindrical, not unlike that of Megaxenislusa, and appears to be moderately physogastric. Schizelythron javanicum Kemner Schizelylhron javanicum Kemner, 1925, Ent. Tidskr., 46: 119, text figs. 1-3, pi. 3 (Buitenzorg, Java; Schedorhinotermes javanicus Kemner; ?Kemner coll.). Subfamily TACHYPORINAE Termitophily is an insignificant feature of the Tachyporinae, yet brief mention must be made of the genus Termitoplus Silvestri. A few years ago Silvestri, although recording this genus with Syn- termes, placed it near Wasmannotherium Bernhauer, a neotropical genus associated with Eciton (army ants). It is apparent from Silvestri's excellent illustrations that Termitoplus is not closely allied to Wasmannotherium (in fact does not belong to the same tribe), and an examination of the genus confirmed this viewpoint. Wasmannotherium and Vatesus comprise a small group of distinctive tachyporine species that are always found in the company of Eciton, and it would be a serious mistake to distort the host pattern by linking them with the termitophilous genus Termitoplus. Termitoplus is in fact related to Erchomus Motschulsky (=Copro- porus Kraatz), a large genus of free-living tachyporines, and my preliminary comparison of the two genera did not reveal a significant basis for separation. However, I am not familiar enough with Erchomus to state that Termitoplus is a synonym of it. More than seventy species of Erchomus, mostly collected from logs, under bark, and from vegetable debris, have been recorded from the Neotropical Region alone. Termitoplus grandis Silvestri, the only known species, is ap- parently a regular inhabitant of Syntermes mounds, but whether or not it has an obligatory relationship to the Syntermes societies is yet to be determined. For a description and illustrations of this species, the interested reader is referred to Silvestri's paper. 282 FIELDIANA: ZOOLOGY, VOLUME 40 TERMITOPLUS Silvestri Termitoplus Silvestri, 1945, Comm. Pont. Acad. Sci., 9: 547. Type species: Termitoplus grandis Silvestri. Termitoplus grandis Silvestri Termitoplus grandis Silvestri, 1945, Comm. Pont. Acad. Sci., 9: 551, text figs. 13, 14 (Jabaquara, Sao Paulo, Brazil; and Jaguariahiva, Parana, Brazil; Syntermes grandis Rambur; Silvestri coll.). Record. Silvestri also recorded this species from Loreto, Misi- ones, Argentina, with Syntermes obtusus Holmgren. Material examined. Brazil: Sao Paulo (Interlagos), Sao Paulo, June 5, 1951, with Syntermes wheeleri Emerson (determined by R. Araujo); and Sao Paulo (Congohas), Sao Paulo, July 3, 1951, with Syntermes dims Burmeister (determined by R. Araujo); both col- lections by R. L. Araujo. Subfamily HYPOCYPHTINAE This is a small subfamily which, although sometimes classed as a tribe of the Tachyporinae, is almost certainly more closely related to the Aleocharinae. A detailed comparative study is needed to establish the validity of the subfamily category or to place this group in its proper subfamily. At present the subfamily includes Cypha Leach (= Hypocyphthus Gyllenhal) and a few related genera. Only a few species are associ- ated in any way with termites, and it is doubtful if these have an obligatory relationship to termite societies. It is probable, there- fore, that they do not conform to the definition of termitophiles used in this revision. Some species have been recorded so frequently with termites that they cannot be ignored, even if at other times they occur under bark of logs or even in ant colonies. Species of Anacyptus are extremely minute, being less than one millimeter in length. They are probably the smallest staphylinids, except for the subterranean Leptotyphlinae. Anacyptus has a very wide distribution in the Americas, where it has been recorded with Reticulitermes, Coptotermes, and Diversitermes as well as in logs and under bark. I have material that was collected with Eciton in North Carolina. The host-guest relationship is apparently so weak that Anacyptus is extremely polyxenous. I recently examined, in the Cameron collection of the British Museum (Natural History), an undescribed genus that was collected SEEVERS: TERMITOPHILOUS STAPHYLINIDAE 283 with termites in Angola. This genus (Mesoporus Cameron, MS) will soon be described in a posthumous paper by Cameron. Although believed by Cameron to represent a new subfamily, it doubtless has a position close to Anacyptus in the Hypocyphtinae. ANACYPTUS Horn Anacyptus Horn, 1877, Trans. Amer. Ent. Soc, 6: 87. Type species: Ana- cyptus testaceus LeConte. Microcyptus Horn, 1883, Trans. Amer. Ent. Soc, 10, proc, p. 1. Anacyptus testaceus LeConte Hypocyptus testaceus LeConte, 1853, Smithson. Misc. Coll., 6: 30 (Athens, Georgia; under pine bark; Museum of Comparative Zoology). Anacyptus testaceus LeConte, Horn, 1877, Trans. Amer. Ent. Soc, 6: 87, 125; Schwarz, 1878, Proc. Amer. Phil. Soc, 17: 440; Blackwelder, 1943, Bull. U. S. Nat. Mus., 182: 532. Microcyptus testaceus LeConte, Schwarz, 1889, Proc Ent. Soc. Wash., 1: 160; Blatchley, 1910, Coleopt. Indiana, p. 442. Records. Blackwelder records this species from Arizona, Texas, and Florida (U. S. Nat. Mus. coll.), Georgia (LeConte coll.), Georgia and Arizona (Horn, 1877), Lake Superior (Casey coll.), Cayamas, Cuba (Schwarz coll., U. S. Nat. Mus.), and Mont- serrat (Hubbard coll., U. S. Nat. Mus.). Material examined. Florida: North Miami, December 4, 1940, E. M. Miller, with Prorhinotermes simplex Hagen; Royal Palm State Park, March 23, 1941, V. Dropkin and A. Emerson, with Neotermes castaneus Burmeister; Tampa, November 15, 1936, O. Falls, with Reticulitermes hageni Banks. North Carolina: Southern Pines, April 16, 1949, M. Wing, with Ecilon (Neivamyrmex) nigrescens Cresson. Arkansas: Hot Springs, July 1, 1935, C. Seevers, with Reticulitermes tibialis Banks. Other material, possibly not belonging to this species, was col- lected in Mexico: Penuela, Vera Cruz, August 17, 1941, H. Dybas, under bark; and Huichihuayan, San Luis Potosi, June 18, 1941, H. Dybas, under bark. CALLOPSENIUS Wasmann Eupsenius Wasmann, 1902, Boll. Mus. Zool. Torino, 17: 5. Type species: Eupsenius clavicornis Wasmann. Callopsenius Wasmann, 1903, Deuts. Ent. Zeitschr., 1903: 236 (for Eupsenius, preoccupied, LeConte, 1850). 284 FIELDIANA: ZOOLOGY, VOLUME 40 Callopsenius was placed by Wasmann in the subfamily Cephalo- plectinae, which has since been shown (Seevers and Dybas, 1943) to be extremely artificial. Cephaloplectus and related genera were transferred to the family Limulodidae (more closely allied to the Ptiliidae than to the Staphylinidae), while other genera were assigned to existing staphylinid subfamilies. I have since examined the type of Wasmann's Callopsenius clavicornis, which unfortunately lacks a head and part of the thorax, and can only conclude that it belongs to the Hypocyphtinae. The hind coxae are particularly indicative of relationship to this group; they bear large lamellar plates that partially conceal the hind femora. Callopsenius clavicornis Wasmann Eupsenius clavicornis Wasmann, 1902, Boll. Mus. Zool. Torino, 17, no. 427, p. 6 (Tacurupucu, Paraguay; Heterotermes longiceps Snyder; Wasmann coll.); Silvestri, 1903, Redia, I: 202, pi. 6, figs. 285, 286. IV. Systematic Catalogues < Q >- K Pu m O K o w Eh w K fa o O I I H S3 i. - x> bfi.* flj z. 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W V. a* ftj to ^ ft*fc ^a, 301 A LIST OF TERMITES AND THEIR STAPHYLINID GUESTS Family Mastotermitidae Mastotermes darwiniensis Froggatt Mastopsenius australis Seevers Australia Family Rhinotermitidae Coptotermes crassus Snyder Neophilotermes laxicornis Sharp Coptotermes acinaciformis Froggatt Coptotermoecia alutacia Oke Coptotermoecia gayi Seevers Prorhinotermes, new sp. Hamitopsenius micronesiae Seevers Reticulitermes arenincola Goellner Philotermes emersoni Seevers Reticulitermes flavipes Kollar Philotermes pensylvanicus Kraatz Philotermes pilosus Kraatz Trichopsenius frosti Seevers Trichopsenius xenoflavipes Seevers Reticulitermes hesperus Banks Trichopsenius californicus Seevers Trichopsenius longipes Seevers Reticulitermes speratus Kolbe Trichopsenius japonicus Seevers Reticulitermes virginicus Banks Philotermes cubitopilis Seevers Philotermes fuchsii Kraatz Philotermes werneri Seevers Trichopsenius depressus LeConte Xenistusa hexagonalis Seevers Reticulitermes sp. indet. Xenistusa cavernosa LeConte Rhinotermes marginalis Linnaeus Megaxenistusa rhinotermitis Seevers Rhinotermopsenius saltatorius Seevers Termitopsenius acanthoscelis Seevers Schedorhinotermes intermedius Brauer Termitoecia fabulosa Bernhauer Schedorhinotermes javanicus Kemner Schizelythron javanicum Kemner Guatemala, Costa Rica Australia Australia Palau Islands United States United States United States United States United States United States United States Japan United States United States United States United States United States United States British Guiana British Guiana British Guiana Australia Java 302 Family Termitidae; Nasutitermitinae Bulbitermes constrictoides Holmgren Affinoptochus exclusus Kemner Bulbitermes singaporiensis Haviland Termitoptochus indicia Silvestri Constrictotermes cavifrons Holmgren Corotoca guyanae Mann Spirachtha schioedtei Mann Spirachtha mirabilis Mann Constrictotermes cyphergaster Silvestri Corotoca araujoi Seevers Corotoca melantho Schi0dte Corotoca phylo Schi0dte Spirachtha eurymedusa Schi0dte Termitocola silvestrii Wasmann Cornitermes cumulans Kollar Macrotrichurus brasiliensis Silvestri Termitohospes unicolor Silvestri Termitozophilus laetus Silvestri Cornitermes pugnax Emerson Termitozophilus mirandus Mann Cornitermes, sp. indet. Termitohospes limulus Seevers Termitohospes tachyporoides Seevers Diversitermes diversimilis Silvestri Perlinctus quaesitus Silvestri Termitonicus uroclaviger Silvestri Termitosodalis fasciatus Silvestri Diversitermes, new sp. Perlinctus foveicollis Seevers Grallatotermes africanus Harris Idioptochus lehmensicki Seevers Millotoca tanganyikae Seevers Hospitalitermes butteli Holmgren Termitoptochus sumatranus Silvestri (larva) Labiotermes labralis Holmgren Termitosymbia nitida Seevers Termitozyras boliviae Seevers Labiotermes labralis Holmgren, new subsp. Termitocomes trinidadensis Seevers Termitozyras adamsoni Seevers Termitozyras guianae Seevers Lacessititermes laborator Haviland Pseudoperinthus malayanus Wasmann Nasulitermes aquilinus Holmgren Termitohospes brasiliana Seevers Termitohospes silvestrii Seevers Termilopithus crassiusculus Seevers Termitosyne platygastra Seevers Xenogaster inflata Wasmann Nasulitermes ? arenarius Silvestri Thyreoxenus autuorii Silvestri Java Malaya British Guiana British Guiana British Guiana Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil, Argentina, Paraguay British Guiana Brazil Brazil Brazil Brazil Brazil Brazil Tanganyika Tanganyika Sumatra Bolivia Bolivia Trinidad Trinidad British Guiana Sumatra Brazil Brazil Brazil Brazil Brazil Brazil 303 Nasutitermes bivalens Holmgren Perinthus bolivari Seevers Termitogaster chavantinae Seevers Termitogaster nigricollis Silvestri Termitogaster wenzeli Seevers Thyreoxenus pulchelltcs Mann Nasutitermes brevirostris Oshima Lauella palauensis Seevers Nasutitermes ceylonicus Holmgren Termitoptochus ceylonicus Silvestri (larva) Nasutitermes chaquimayensis Holmgren Abroteles rurrenabaquensis Seevers Perinthus bolivari Seevers Termitogaster beniensis Seevers Termitophya holmgreni Wasmann Thyreoxenus convexinotus Seevers Nasutitermes columbicus Holmgren Perinthus dudleyanus Casey Termitohospes panamensis Seevers Termitophya piliventris Seevers Thyreoxenus major Mann Xenopelta cornuta Mann Nasutitermes corniger Motschulsky Abroteles beaumonti Casey Perinthus dudleyanus Casey Perinthus hageni Seevers Perinthus mayae Seevers Termitogaster insolens Casey Termitogaster testaceus Seevers Termitomorpha fissipennis Casey Termitomorpha costaricensis Seevers Nasutitermes costalis Holmgren Perinthus dudleyanus wasmanni Mann Perinthus xenocostalis Seevers Termitogaster brevis Mann Termitohospes miricorniger Seevers Termitomorpha meinerti Wasmann Termitomorpha fissipennis Casey Termitophya flaviventris Mann Thyreoxenus brevitibialis Seevers Thyreoxenus parviceps Mann Nasutitermes coxipoensis Holmgren Abroteles badia Seevers Nasutitermes dendrophilus Holmgren Abroteles bisetosus Seevers Perinthus dudleyanus major Seevers Termitogaster bicolor Seevers Termitophya inornata Seevers Nasutitermes ehrhardti Holmgren Xenogaster pilosula Seevers Nasutitermes ephratae Holmgren Abroteles beaumonti Casey Eburniola gastrovittata Seevers Perinthus bolivari Seevers Brazil Brazil Brazil Brazil Brazil Palau Islands Ceylon Bolivia Bolivia Bolivia Bolivia, Peru Bolivia Panama Panama Panama Panama Panama Panama Panama Panama, Costa Rica Honduras, Guatemala Panama, Honduras Costa Rica Panama Costa Rica British Guiana British Guiana, Trinidad British Guiana, Trinidad British Guiana, Trinidad British Guiana, Grenada British Guiana British Guiana, Trinidad Colombia British Guiana, Trinidad Brazil Ecuador Ecuador Ecuador Ecuador Brazil Panama Panama Trinidad 304 Perinthus dudleyanus Casey Perinthus dudleyanus wasmanni Mann Perinthus guatemalae Seevers Perinthus hageni Seevers Termitogaster diversicollis Seevers Termitogaster emersoni Mann Termitogaster magdalenae Seevers Termitohospes miricorniger Seevers Termitophya emersoni Seevers Thyreoxenus albidus Seevers Thyreoxenus cucullatus Seevers Thyreoxenus pulchellus Mann Nasutitermes fulviceps Silvestri Abroteles philetaerus Silvestri Termitophya heyeri Wasmann Xenogaster inflata Wasmann Nasutitermes fumipennis Walker Termitoptocinus australiensis Silvestri Nasutitermes gaigei Emerson Poduroides bovingi Mann Termitospeclrum thoracicum Mann Nasutitermes globiceps Holmgren Abroteles lobata Seevers Termitogaster emarginatus Seevers Nasutitermes guayanae Holmgren Eburniola leucogaster Mann Termitophya arnica Mann Thyreoxenus major Mann Xenopelta cornuta Mann Nasutitermes, new sp. Xenogaster reichenspergeri Seevers Nasutitermes incurvus Sjostedt Paraperinthus paucisela Seevers Perinthodes africanus Seevers Nasutitermes intermedium Banks Xenogaster fossulatus Mann Nasutitermes itapocuensis Holmgren Termitophya heyeri Wasmann Xenogaster inflata Wasmann Xenogaster inquilina Borgmeier Xenogaster subnuda Seevers Nasutitermes jaraguae Holmgren Abroteles compact icornis Borgmeier Abroteles pubicollis Seevers Eburniola lujae Seevers Termitophya heyeri Wasmann Xenogaster nana Seevers Xenogaster glabriventris Seevers Xenogaster reichenspergeri Seevers Nasutitermes latifrons Sjostedt Perinthodes africanus Seevers Nasutitermes luzonicus Oshima Termitoptochus luzonicus Silvestri (larva) Termitoptochus philippinus Silvestri Panama British Guiana, Trinidad Guatemala Panama, Costa Rica Colombia British Guiana, Panama Colombia British Guiana, Trinidad British Guiana, Panama Colombia Colombia British Guiana, Trinidad Argentina Brazil Brazil, Uruguay Australia British Guiana British Guiana Brazil Brazil British Guiana British Guiana, Trinidad British Guiana British Guiana Brazil Belgian Congo Belgian Congo British Guiana Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Belgian Congo Philippines Philippines 305 Nasutitermes maculiventris Sjostedt Termitella lujae Wasmann Termitomimus latipes Seevers Nasutitermes matangensis Haviland Affinoptochus exclusus Kemner Nasutitermes meinerli Wasmann Abroteles beaumonti Casey Eburniola gastrovittata Seevers Perinthus dudleyanus Casey Termitomorpha meinerti Wasmann Nasutitermes minimus Holmgren Termilophya holmgreni Wasmann Nasutitermes mojosensis Holmgren Termitogaster wasmanni Holmgren Termitophya holmgreni Wasmann Nasutitermes nigriceps Haldeman Termitosynodes williamsi Seevers Trachopeplus setosus Mann Trachopeplus disjunctipennis Seevers Nasutitermes novarumhebridarum Holmgren and Holmgren Thyreoxenus solomonensis Seevers Nasutitermes octopilis Banks Paratermitosocius vestitus Mann Termitosocius microps Seevers Nasutitermes olidus Hill Lauella vitiensis Mann Nasutitermes peruanus Holmgren Abroteles rurrenabaquensis Seevers Perinthus dudleyanus major Seevers Perinthus hageni Seevers Termitoides marginatus Seevers Neotermitogaster colonus Seevers Termitogaster impressicollis Seevers Termitogaster puncticeps Seevers Termitomorpha manni Seevers Termitophya ecuadoriensis Seevers Nasutitermes pilifrons Holmgren Termitocola cylindricornis Seevers Nasutitermes pilosus Snyder Abroteles rurrenabaquensis Seevers Perinthus xenocostalis Seevers Termitomorpha manni Seevers Thyreoxenus pulchellus Mann Nasutitermes pluriarticulatus Silvestri Termitogaster nigricollis Silvestri Nasutitermes proximus Silvestri Abroteles compacticornis Borgmeier Termitophya araujoi Borgmeier Xenogaster inquilina Borgmeier Nasutitermes rufirostris Hill Hetairotermes insulanus Seevers Termitobra perinthoides Seevers Belgian Congo Belgian Congo Java Colombia Colombia Colombia Venezuela Bolivia Bolivia Bolivia Ecuador British Guiana Ecuador Solomon Islands British Guiana British Guiana Fiji Islands Bolivia Ecuador Ecuador Ecuador Ecuador Ecuador Bolivia Bolivia Ecuador Panama Bolivia Bolivia Bolivia Bolivia Brazil Brazil Brazil Brazil Palau Islands Palau Islands 306 Nasutitermes sanctaeanae Holmgren Termitogaster nigricollis Silvestri Argentina Nasutitermes similis Emerson Termitophya punctata Mann British Guiana Xenopelta tricornis Mann British Guiana Nasutitermes surinamensis Holmgren Perinthus tarsatus Mann British Guiana Termitohospes guianae Seevers British Guiana Nasutitermes torqualus Sjostedt Termitella foveolata Reichensperger Belgian Congo Termitella lujae Wasmann Belgian Congo Termitella rubricollis Seevers Belgian Congo Nasutitermes usambarensis Sjostedt Termitellodes lativentris Seevers Belgian Congo Termitomimus emersoni Seevers Belgian Congo Termitopullus sociusculus Reichensperger Belgian Congo Nasutitermes, sp. indet. Termitomorpha huachii Seevers Bolivia Thyreoxenus boliviae Seevers Bolivia Nasutitermes (s. lat.) spp., Silvestri Oecidiophilus mimellus Silvestri Argentina Fonsechellus bicolor Silvestri Brazil Fonsechellus diversicolor Silvestri Brazil Macrognathellus crassicornis Wasmann Paraguay Paracornitermes emersoni Araujo Autuoria elegantulum Silvestri Brazil Procornitermes araujoi Emerson Termitonusa sequax Borgmeier Brazil Termituncula gracilipes Borgmeier Brazil Procornitermes striatus Hagen Termitonannus domunculi Silvestri Argentina Subulitermes microsoma Silvestri Mormellus bicolor Silvestri Brazil Parvidolum microsomatis Silvestri Brazil Termitonannus microsomatis Silvestri Brazil Subulitermes, new sp. Fonsechellus fragilis Seevers Brazil Syntermes dirus Burmeister Termitoplus grandis Silvestri Brazil Syntermes grandis Rambur Termitonannus validus Silvestri Brazil Termitophagus synterminus Silvestri Brazil Syntermes molestus Burmeister Termitopelta fulgens Borgmeier Brazil Syntermes wheeleri Emerson Iheringocantharus ypiranganus Bernhauer Brazil Termitophagus synterminus Silvestri Brazil Tenuirostritermes cinereus Buckley Eburniogaster texana Brues United States Tenuirostritermes, new sp. Termitonidia jaliscensis Seevers Mexico Tenuirostritermes tenuirostris Desneux Eburniogaster termitocola Seevers United States Termitonidia lunata Seevers United States 307 Tenuirostritermes, new sp. Eburniogaster anahuaci Seevers Termitonidia michoacani Seevers Termitonidia tarascani Seevers Trinervitermes biformis Wasmann Doryloxenus eutermitis Wasmann Eutermitophila fletcheri Cameron Termitotima assmuthi Wasmann Trinervitermes heimi Wasmann Termitoptochus peninsularis Silvestri (larva) Trinervitermes rapulum Sjostedt Idiogaster escherichi Wasmann Trinervitermes trinerviformis Holmgren Paracorotoca akermani Warren Termitomimus entendveniensis Tragardh Trinervitermes trinervius Rambeau Oideprosoma mirandum Silvestri Velocitermes beebei Emerson Blapticoxenus brunneus Mann Termitonicus mahout Mann Termitosodalis barticae Seevers Velocitermes heteropterus Silvestri Termitoiceus anastrephoproctus Silvestri Termitoiceus simulans Silvestri Termitosius pauciseta Silvestri Mexico Mexico Mexico India India India India Eritrea Natal Natal French Guinea British Guiana British Guiana British Guiana Brazil Brazil Paraguay Family Termitidae; Macrotermitinae Macrotermes bellicosiis Smeathman Termitobia physogastra Wasmann Termitobia paolii Gridelli Termitogerrus bellicosi Silvestri Termitogerrus habilis Silvestri Termitogerrus quaerens Silvestri Termitopidex grandicornis Fauvel Macrotermes gilvm Hagen Dioxeuta microps Sharp Neodioxeuta oudemansi Franssen Macrotermes goliath Sjostedt Termitobia sp. Macrotermes malaccensis Haviland Dioxeuta indosinensis Silvestri Macrotermes natalensis Haviland Termitonda liberiae Seevers Termitonda tachyporoides Seevers Termitobia burgeoni Cameron Termitobia rhodesiae Seevers Termitogerrus burgeoni Bernhauer Termitogerrus lepidulus G. and L. Termitopulex natalensis Wasmann Macrotermes swaziae Fuller Termitobia sp. Gold Coast Italian Somaliland Eritrea French Guinea Senegal Ethiopia Sarawak, Java, Sumatra, Malaya, Philippines Java Tanganyika Indo-China Liberia, Ivory Coast Belgian Congo, Ivory Coast Belgian Congo, Liberia Rhodesia Belgian Congo Ivory Coast Natal Transvaal 308 Macrotermes, sp. indet. Dioxeuta flaveseens Cameron Neodioxeuta burmae Seevers Microtermes vadsehaggae Sjostedt Termitopulex sjoestedli Eichelbaum Odontotermes angustatus Ram bur Termitodiseus transraalensis Silvestri Odontotermes ceylonieus Wasmann Termitodiseus escherichi Wasmann Odontotermes jaranieus Holmgren Doryloxenus triartieulatus Kemner Odontotermes obesus Rambur Diseoxenus assmuthi Wasmann Doryloxenus termitophilus Wasmann Termitodiseus heimi Wasmann Odontotermes obseuriceps Wasmann Discoxenus acuticornis Wasmann Doryloxenus butteli Wasmann Doryloxenus splendidus Wasmann Termitodiseus butteli Wasmann Odontotermes redemanni Wasmann Discoxenus erassicornis Wasmann Doryloxenus eeylonieus Wasmann Doryloxenus peradenyiae Wasmann Termitodiseus escherichi Wasmann Odontotermes transvaalensis Sjostedt Termitodiseus braunsi Wasmann Termitotecna braunsi Wasmann Odontotermes vulgaris Haviland Termitodiseus splendidus Wasmann Odontotermes wallonensis Wasmann Discoxenus lepisma Wasmann Doryloxenus transfuga Wasmann Termitodiseus heimi Wasmann Pseudacanthotermes spiniger Sjostedt Termitopaedia kohli Wasmann Burma Burma Tanganyika Transvaal Ceylon Java India India India Ceylon Ceylon Ceylon Ceylon Ceylon Ceylon Ceylon Ceylon Orange Free State Orange Free State Natal India India India Belgian Congo Family Termitidae; Amitermitinae Amitermes dentatus Haviland flamitopsenius caudatus Wasmann Sumatra Amitermes unidentatus Sjostedt Termitotropha o'neili Wasmann South Africa Anoplotermes ater Hagen Termitonannus schmalzi Wasmann Brazil Termitonilla luteola Borgmeier Brazil Anoplotermes bequaerti Snyder and Emerson Termitonannus major Wasmann Paraguay Timeparthenus oglobini Silvestri Argentina Anoplotermes hageni Snyder and Emerson Timeparthenus regius Silvestri Brazil Anoplotermes meridianus Emerson Termitonannus silvestrii Wasmann Argentina 309 Anoplotermes pacificus Miiller Macrotrichuris brasiliensis Silvestri Macrotrichuris notabilis Silvestri Termitonannus schmalzi Wasmann Anoplotermes turricola Silvestri Ptocholellu8 mimus Silvestri Anoplotermes, new sp. Termitonannus gatuni Seevers Termitonannus setosus Seevers Anoplotermes, sp. indet. Eunannodes reconditi Silvestri Nannellus anoplotermitis Silvestri Termitonannus brachycerus Silvestri Termitonannus echinoides Seevers Termitonannus parvulus Silvestri Termitonannus proximatus Silvestri Speculitermes reconditus Silvestri Termitonannus vagans Silvestri Speculitermes silvestrii Emerson Termitocomes wasmanni Seevers Brazil Brazil Brazil Brazil Panama Panama Brazil Brazil Brazil Bolivia Brazil Brazil Brazil British Guiana Family Termitidae; Termitinae Capritermes capricornis Wasmann Termitochara kraatzi Wasmann Capritermes nemorosus Haviland Felda butteli Wasmann Termitobiella setipes Wasmann Zunia capritermitis Wasmann Cubitermes fungifaber Sjostedt Termitusa hystrix Wasmann Termitusa lujae Wasmann Termitusa sjoestedti Wasmann Cubitermes glebae Sjostedt Termitusa cameroni Reichensperger Cubitermes sankurensis Wasmann Termitusa lujae Wasmann Cubitermes subcrenulatus Silvestri Protermitobia comes Seevers Cubitermes, sp. indet. Termitusa escalerae Fauvel Termitusa mundus Bernhauer Termitusa quadricollis Cameron Microcerotermes sikorae Wasmann Termitochara kraatzi Wasmann Neocapritermes opacus Hagen Termitopsenius limulus Wasmann Noditermes cristifrons Wasmann Termitusa lativentris Seevers Termes baculiformis Holmgren Millotoca mirotermitidis Paulian Madagascar Malaya Malaya Sumatra Belgian Congo Belgian Congo Cameroons Tanganyika Belgian Congo Sierra Leone Rio Muni Cameroons Belgian Congo Madagascar Argentina French Equatorial Africa Madagascar 310 REGIONAL LIST OF TERMITOPHILOUS STAPHYLINIDAE /. Neotropical Region ARGENTINA Abroteles philetaerus Silvestri Oecidiophilus mimellus Silvestri Termitogaster nigricollis Silvestri Termitonannus silvestri i Wasmann Termitonannus domunculi Wasmann Termitoniciis uroclaviger Silvestri Termitopsenius limidus Wasmann Termitozophilus laetus Silvestri Timeparthenus oglobini Silvestri BOLIVIA Abroteles rurrenabaquensis Seevers Perinthiis bolivari Seevers Perinthus xenocostalis Seevers Termitogaster beniensis Seevers Termitogaster puncticeps Seevers Termitogaster wasmanni Holmgren Termitomorpha huachii Seevers Termitomorpha manni Seevers Termitonannus echinoides Seevers Termitophya holmgreni Wasmann Termitosymbia nitida Seevers Termitozyras boliviae Seevers Thyreoxenus boliviae Seevers Thyreoxenus convexinotus Seevers Thyreoxenus pulchellus Mann BRAZIL Abroteles badia Seevers Abroteles compacticornis Borgmeier Abroteles lobata Seevers Abroteles pubicollis Seevers Autuoria elegantula Silvestri Corotoca araujoi Seevers Corotoca melantho Schi0dte Corotoca phylo Schi0dte Eburniola lujae Seevers Euannodes reconditi Silvestri Fonsechellus bicolor Silvestri Fonsechellus diversicolor Silvestri Fonsechellus fragilis Seevers Iheringocantharus ypiranganus Bern- hauer Macrotrichurus brasiliensis Silvestri Macrotrichurus notabilis Silvestri Mormellus bicolor Silvestri Nannellus anoplotermitis Silvestri Parvidolum microsomatis Silvestri Perinthus bolivari Seevers Perlinctus quaesitus Silvestri Perlinctus foveicollis Seevers Ptocholellus mimus Silvestri Spirachtha eurymedusa Schi0dte Termitocola silvestrii Wasmann Termitogaster chavantinae Seevers Termitogaster emarginatus Seevers Termitogaster nigricollis Silvestri Termitogaster wenzeli Seevers Termitohospes brasiliana Seevers Termitohospes limultis Seevers Termitohospes silvestrii Seevers Termitohospes tachyporoides Seevers Termitohospes unicolor Silvestri Termitoiceus anastrephroproctus Silves- tri Termitoiceus simulans Silvestri Termitonannus brachycerus Silvestri Termitonannus microsomatis Silvestri Termitonannus parvulus Silvestri Termitonannus proximatus Silvestri Termitonannus schmalzi Wasmann Termitonannus vagans Silvestri Termitonannus validus Silvestri Termitonicus uroclaviger Silvestri Termitonilla luieola Borgmeier Termitonusa sequax Borgmeier 311 Termitopelta fulgens Borgmeier Termitophagus synterminus Silvestri Termitophya araujoi Borgmeier Termitophya heyeri Wasmann Termitopithus crassiusculus Seevers Termitoplus grandis Silvestri Termitosodalis fasciatus Silvestri Termitosyne platygastra Seevers Termitozophilus laetus Silvestri Termituncula gracilipes Borgmeier Timeparthenus regius Silvestri Thyreoxenus autuorii Silvestri Thyreoxenus pulchellus Mann Xenogaster glabriventris Seevers Xenogaster inflata Wasmann Xenogaster inquilina Borgmeier Xenogaster nana Seevers Xenogaster pilosula Seevers Xenogaster reichenspergeri Seevers Xenogaster subnuda Seevers BRITISH GUIANA Blapticoxenus brunneus Mann Corotoca guyanae Mann Eburniola leucogaster Mann Megaxenistusa rhinotermitis Seevers Paratermitosocius vestitus Mann Perinthus dudleyanus wasmanni Mann Perinthus tarsatus Mann Perinthus xenocostalis Seevers Poduroides bovingi Mann Rhinotermopsenius saltatorius Seevers Spirachtha mirabilis Mann Spirachtha schioedtei Mann Termitocomes wasmanni Seevers Termitogaster brevis Mann Termitogaster emersoni Mann Termitohospes miricorniger Seevers Termitohospes guianae Seevers Termitomorpha meinerti Wasmann Termitomorpha fissipennis Casey Termitonicus mahout Mann Termitophya arnica Mann Termitophya emersoni Seevers Termitophya flaviventris Mann Termitophya punctata Mann Termitopsenius acanthoscelis Seevers Termitosocius microps Seevers Termitosodalis barticae Seevers Termitospectrum thoracicum Mann Termitozophilus mirandus Mann Termitozyras guianae Seevers Thyreoxenus major Mann Thyreoxenus parviceps Mann Thyreoxenus pulchellus Mann Trachopeplus setosus Mann Xenogaster fossulatus Mann Xenopelta cornuta Mann Xenopelta tricornis Mann COLOMBIA Abroteles beaumonti Casey Eburniola gastrovittata Seevers Perinthus dudleyanus Casey Termitogaster diversicollis Seevers Termitogaster magdalenae Seevers Thyreoxenus albidus Seevers Thyreoxenus cucullatus Seevers Thyreoxenus brevitibialis Seevers COSTA RICA Abroteles beaumonti Casey Neophilotermes laxicornis Sharp Perinthus hageni Seevers Termitogaster testaceus Seevers Termitomorpha costaricensis Seevers ECUADOR Abroteles bisetosus Seevers Neotermitogaster colonus Seevers Perinthus dudleyanus major Seevers Perinthus hageni Seevers Termitogaster bicolor Seevers Termitogaster impressicollis Seevers Termitoides marginatum Seevers Termitophya ecuadoriensis Seevers Termitophya inornata Seevers Termitosyne williamsi Seevers Trachopeplus disjunctipennis Seevers 312 GUATEMALA Neophilotermes laxicornis Sharp Perinthus guatemalae Seevers Perinthus mayae Seevers Perinthus mayae Seevers HONDURAS Termitogaster insolens Casey Eburniogaster anahuaci Seevers Termitonidia jaliscensis Seevers MEXICO Termitonidia michoacani Seevers Termitonidia tarascani Seevers PANAMA Abroteles beaumonti Casey Eburniola gastrovittata Seevers Perinthus dudleyanus Casey Perinthus hageni Seevers Termitocola cylindricornis Seevers Termitogaster insolens Casey Termitohospes panamensis Seevers Termitomorpha fissipennis Casey Termitonannus gatuni Seevers Termitonannus setosus Seevers Termitophya emersoni Seevers Termitophya piliventris Seevers Thyreoxenus major Mann PARAGUAY Macrognathellus crassicornis Wasmann Termitonannus major Wasmann Termitosius pauciseta Silvestri Termitozophilus laetus Silvestri PERU Termitophya holmgreni Wasmann TRINIDAD, TOBAGO, GRENADA Eburniola leucogaster Mann Perinthus dudleyanus wasmanni Mann Perinthus bolivari Seevers Perinthus xenocostalis Seevers Termitocomes trinidadensis Seevers Termitogaster brevis Mann Termitogaster emersoni Mann Termitohospes mirieorniger Seevers Termitomorpha meinerti Wasmann Termitophya arnica Mann Termitophya flaviventris Mann Termitozyras adamsoni Seevers Thyreoxenus parviceps Mann Thyreoxenus pulchellus Mann URUGUAY Xenogaster inftala Wasmann VENEZUELA Termitomorpha meinerti Wasmann 313 II. Nearctic Region UNITED STATES Eburniogaster termitocola Seevers Eburniogaster texana Brues Philotermes cubitopilis Seevers Philotermes emersoni Seevers Philotermes fuchsii Kraatz Philotermes pilosus Kraatz Philotermes pensylvanicus Kraatz Philotermes werneri Seevers Termitonidia lunata Seevers Trichopsenius californicus Seevers Trichopsenius depressus LeConte Trichopsenius frosti Seevers Trichopsenius longipes Seevers Trichopsenius xenoflavipes Seevers Xenistusa cavernosa LeConte Xenistusa hexagonalis Seevers 777. Palaearctic Region JAPAN Trichopsenius japonicus Seevers IV. Indomalayan Region Dioxeuta microps Sharp BORNEO Termitobaena bryanti Bernhauer Dioxeuta flavescens Cameron BURMA Neodioxeuta burmae Seevers CEYLON Discoxenus acuticornis Wasmann Discoxenus crassicornis Wasmann Doryloxenus butteli Wasmann Doryloxenus ceylonicus Wasmann Doryloxenus peradenyiae Wasmann Doryloxenus splendidus Wasmann Termitodiscus butteli Wasmann Termitodiscus escherichi Wasmann Termitoptochus ceylonicus Silvestri INDIA Discoxenus assmuthi Wasmann Discoxenus lepisma Wasmann Doryloxenus brevicornis Cameron Doryloxenus eutermitis Wasmann Doryloxenus longesetosus Cameron Doryloxenus termitophilus Wasmann Doryloxenus transfuga Wasmann Eutermitophila fletcheri Cameron Pseudoperinthus fletcheri Cameron Termitodiscus heimi Wasmann Termitodiscus minutus Cameron Termitoptochus peninsularis Silvestri Termitotima assmuthi Wasmann INDO-CHINA Dioxeuta indosinensis Silvestri 314 JAVA Affinoptochus exclusus Kemner Lauella javana Seevers Dioxeuta microps Sharp Neodioxeuta oudemansi Franssen Doryloxenus triarticulafus Kemner Schizelythron jaranicum Kemner MALAYA Dioxeuta microps Sharp Hetairotermes piceus Cameron Felda butteli Wasmann Termitobiella setipes Wasmann Hetairotermes agilis Cameron Termitoptochus indicus Silvestri Hetairotermes bryanti Cameron PHILIPPINE ISLANDS Dioxeuta microps Sharp Termitoptochus luzonicus Silvestri Neodioxeuta sp. Termitoptochus philippinus Silvestri SUMATRA Dioxeuta microps Sharp Termitoptochus sumatranus Silvestri Hamitopsenius caudatus Wasmann Zunia capritermitis Wasmann Pseudoperinthus malayanus Wasmann V. Australian and Papuan Regions AUSTRALIA Coptotermoecia alutacia Oke Mastopsenius australis Seevers Coptotermoecia gayi Seevers Termitoecia fabulosa Bernhauer Hetairotermes latebricola Lea Termitoecia wilsoni Cameron Hetairotermes punctiventris Lea Termitoptocinus australiensis Silvestri FIJI ISLANDS Lauella vitiensis Mann PALAU ISLANDS Hamitopsenius micronesiae Seevers Lauella palauensis Seevers Hetairotermes insulanus Seevers Termitobra perinthoides Seevers SOLOMON ISLANDS Thyreoxenus solomonensis Seevers VI. Ethiopian Region BELGIAN CONGO Doryloxenus wasmanni Cameron Perinthodes africanus Seevers Paraperinthus pauciseta Seevers Pygostenus infimus Wasmann 315 Pygostenus termitophilus Wasmann Termitella foveolata Reichensperger Termitella gridellii Bernhauer Termitella lujae Wasmann Termitella rubricollis Seevers Termitellodes lativentris Seevers Termitonda tachyporoides Seevers Termitobia burgeoni Cameron Termitogerrus burgeoni Bernhauer Termitomimus emersoni Seevers Termitomimus latipes Seevers Termitopaedia kohli Wasmann Termitopullus sociusculus Reichensper- ger Termitusa hystrix Wasmann Termitusa lujae Wasmann Termitusa quadricollis Cameron THE CAMEROONS Termitusa mundus Bernhauer Termitusa sjoestedti Wasmann Idiogaster escherichi Wasmann Termitopulex grandicornis Fauvel ERITREA Termitogerrus bellicosi Silvestri ETHIOPIA Termitopulex grandicornis Fauvel FRENCH EQUATORIAL AFRICA Termitusa lativentris Seevers FRENCH WEST AFRICA Oideprosoma miranda Silvestri (French Guinea) Termitonda liberiae Seevers (Ivory Coast) Termitonda tachyporoides Seevers (Ivory Coast) Termitogerrus habilis Silvestri (French Guinea) Termitogerrus lepidulus Grasse and Poisson (Ivory Coast) Termitogerrus quaerens Silvestri (Senegal) GOLD COAST Termitobia physogastra Wasmann ITALIAN SOMALILAND Termitobia paolii Gridelli Termitobia burgeoni Cameron Termitogerrus lepidulus Grasse and Lesperon LIBERIA Termitonda liberiae Seevers RIO MUNI Termitusa escalerae Fauvel 316 SIERRA LEONE Protermitobia comes Seevers TANGANYIKA TERRITORY Idioptochus lehmensicki Seevers Termitopulex sjoestedti Eichelbaum Millotoca tanganyikae Seevers Termilusa cameroni Reichensperger Protermitobia kirbyi Seevers UNION OF SOUTH AFRICA Paracorotoca akermani Warren Termitomimus entendveniensis Tragardh Termitobia rhodesiae Seevers Termitopulex natalensis Wasmann Termitodiscus braunsi Wasmann Termitotecna braunsi Wasmann Termitodiscus transvaalensis Silvestri Termitotropho o'neili Wasmann Termitodiscus splendidus Wasmann VII. Malagasy Region MADAGASCAR Millotoca mirotermitidis Paulian Termitochara kraatzi Wasmann Termitana perrieri Fairmaire 317 A LIST OF DOUBTFULLY TERMITOPHILOUS SPECIES Subfamily STAPH YLININAE Belonuchus penetrans Silvestri, 1945, Comm. Pont. Acad. Sci., 9: 522, text figs. 3, 4 (Brazil; Syntermes silvestrii Holmgren, S. molestus Burmeister). Quediosoma sericoilius Cameron, 1920, Ent. Month. Mag., 56: 216 ( = termito- phagum Cameron, 1926, Trans. Ent. Soc. London, 1926: 367) (India; after exposing combs of Odontotermes obesus). Termitoquedius iheringi Bernhauer, 1912, Verh. Zool. Bot. Ges. Wien, 62: 46 (Brazil; termitophilous or myrmecophilous). Termitoquedius is a genus of ecitophilous Staphylinidae. Subfamily PAEDERINAE Acalophaena basalis Lynch, 1884, Bol. Acad. Nac. Cienc. Cordoba, 7: 270 (Ar- gentina; Procornitermes lespesii Fr. Mtiller). Astenus paranensis Lynch, 1884, Bol. Acad. Nac. Cienc. Cordoba, 7: 299 (Ar- gentina; Procornitermes lespesii Fr. Muller). Paederus termitophilus Wasmann, 1911, Rev. Zool. Afr., 1: 101 (Africa; Odonto- termes monodon var. lujana Wasmann). Termitosaurus insinuatus Silvestri, 1945, Comm. Pont. Acad. Sci., 9: 525, figs. 5, 6 (Brazil; Syntermes molestus Burmeister). Subfamily OXYTELINAE Oxytelus simulans Cameron, 1930, Rev. Zool. Bot. Afr., 19: 405 (Belgian Congo; Macrotermes natalensis Haviland). Oxytelus termitophilus Cameron, 1928, Rev. Zool. Bot. Afr., 16: 18 (Belgian Congo; Macrotermes natalensis Haviland). Subfamily OSORIINAE Osorius frater Lynch, 1884, Bol. Acad. Nac. Cienc. Cordoba, 7: 345 (Argentina; Procornitermes lespesii Fr. Muller). Thoracophorus heyeri Wasmann, 1902, Tijd. Ent., 45: 101 (Brazil; Nasutitermes fulviceps Silvestri). Subfamily TACHYPORINAE Conosomus heathi Wasmann, 1902, Tijd. Ent., 45: 99 (California; Zootermopsis angusticollis Hagen). Conosomus termitophilus Wasmann, 1902, Tijd. Ent., 45: 99 (Brazil; Nasutitermes fulviceps Silvestri). Conosomus convexiusculus Wasmann, 1902, Tijd. Ent., 45: 99 (Brazil; Nasuti- termes fulviceps Silvestri). 318 Subfamily EUAESTHETINAE Edaphus termitophilus Bernhauer, 1916, Ark. Zool., 10, no. 5, p. 2 (Australia; no host given). Subfamily ALEOCHARINAE Atheta convivens Silvestri, 1945, Comm. Pont. Acad. Sci., 9: 541 (Brazil; Syntermes grandis Rambur). Atheta silvestrii (new name for Atheta termitaria Silvestri, 1945, Comm. Pont. Acad. Sci., 9: 538; preoccupied by A. termitaria Bernhauer, 1932) (Argentina; Anoplotermes bequaerti Snyder and Emerson). Atheta syntermitis Silvestri, 1945, Comm. Pont. Acad. Sci., 9: 535 (Brazil; Syn- termes grandis Rambur). Atheta termitaria Bernhauer, 1932, Boll. Lab. Zool. Portici, 26: 13 (French Guinea). Atheta termitobia Wasmann, 1894, Krit. Verz., p. 211 (Brazil; Syntermes dims Burmeister). Atheta termitophila Motschulsky, 1859, Etudes Ent., 8: 61 (Ceylon). Aleochara croceipennis Motschulsky, 1858, Bull. Soc. Nat. Moscou, .31: 238 (Ceylon; in termite nest). Chledophila annularis Cameron, 1920, Trans. Ent. Soc. London, 1920: 231 (Fiji Islands). Coenonica puncticollis Kraatz, Linn. Ent., 11: 46 (Ceylon). Dabra termitophila Lea, 1906, Proc. Linn. Soc. N. S. Wales, 31: 215 (Australia; Coptotermes raffrayi Wasmann. Also with Iridomyrmex in Victoria (Lea, 1910, p. 134]). Demerinda termitophila Cameron, 1927, Ent. Month. Mag., 63: 224 (India; frequents exposed combs of Odontotermes |Cameron, 1939, p. 240)). Fenyesia nigra Cameron, 1920, Trans. Ent. Soc. London, 1920: 270 (Malaya; with termites in rotting log). Myrmecopora termitophila Cameron, 1948, Mem. Mus. Nat. d'Hist. Nat., (2), 20: 237 (Ivory Coast; with a termite). Orphnebius termitis Motschulsky (= Hygroptera termitis), 1859, Etudes Ent., 8: 87 (Ceylon; in termite nest). Oxypoda termitophila Bernhauer, 1901, Deuts. Ent. Zeitschr., 1901: 252 (Mada- gascar; no host given). Philotermimus setiger Reichensperger, 1915, Medd. Goteborgs Mus. Zool., 16: 3 (Natal; Odontotermes tragardhi Holmgren). Pelioptera micans Kraatz, 1857, Linn. Ent., 11: 56 (=Termitopora adustipennis Motschulsky, 1859, Etudes Ent., 8: 93) (Ceylon; in termite nest). Pelioptera opaca Kraatz, 1857, Linn. Ent., 11: 56 (Ceylon; in termite nest). Philastilbus opulentus Bernhauer, 1929, Rev. Zool. Bot. Afr., 18: 248 (German East Africa; possibly termitophilous). Porus ferrugineus Kraatz, 1857, Linn. Ent., 22: 48 (Sudan; probably termito- philous). Porus ochraceus Westwood, 1839, in Royle, Illust. Bot. Himal. Mts., 1 : 55 (India). Recorded as termitophilous (Kraatz, 1857a, p. 21; Wasmann, 1894, p. 89). 319 Rhopalinda termitophila Cameron, 1927, Ent. Month. Mag., 63: 223 (India; frequents the exposed combs of "Termes"). Termitolara opacella Bernhauer, 1927, Rev. Zool. Bot. Afr., 15: 336 (Cameroons; with Eutermes sp.). Termitolara reichenspergeri Bernhauer, 1927. Rev. Zool. Bot. Afr., 15: 240 (Cam- eroons; with Nasutitermes latifrons Sjostedt). 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Subfamilies, Tribes, and Subtribes Abrotelina (subtribe), 41, 121 Aleocharinae (subfamily), 61, 319 Athetini (tribe), 248 Coptotermoeciina (subtribe), 248 Corotocina (subtribe), 27, 35, 140 Corotocini (tribe), 25, 62, 63 Euaesthetinae (subfamily), 319 Feldina (subtribe), 236 Hetairotermitina (subtribe), 191, 201 Hypocyphtinae (subfamily), 282 Myrmedoniina (subtribe), 233 Myrmedoniini (tribe), 62, 232 Osoriinae (subfamily), 318 Oxytelinae (subfamily), 318 Paederinae (subfamily), 318 Perinthina (subtribe), 45, 62, 63, 164 Philotermitini (tribe), 54, 63, 250 Pseudoperinthini (tribe), 63, 247 Pygosteninae (subfamily), 265 Staphylininae (subfamily), 318 Tachyporinae (subfamily), 281, 318 Termitocharina (subtribe), 139 Termitodiscini (tribe), 54, 63, 258 Termitogastrina (subtribe), 29, 32, 70 Termitohospitina (subtribe), 50, 191, 192 Termitohospitini (tribe), 48, 63, 191 Termitonannina (subtribe), 46, 63, 181 Termitonannini (tribe), 42, 164 Termitondina (subtribe), 238 Termitopaediini (tribe), 52, 214 Termitospectrina (subtribe), 191 Termitotelina (subtribe), 250 Termitozyrina (subtribe), 240 Termitusina (subtribe), 51, 192 Timeparthenina (subtribe), 29, 30, 64 Trichopseniinae (subfamily), 266 77. Genera and Subgenera Abroteles Casey, 122 Acalophaena Sharp, 318 Affinoptochus Kemner, 33, 115 Aleochara Gravenhorst, 319 Anacyptus Horn, 283 Astenus Dejean, 318 Asticia Wasmann, 237 Astilbus Dilwyn, 235 Atheta Thomson, 319 Autuoria Silvestri, 30, 68 Belonuchu8 Nordmann, 318 Blapticoxenus Mann, 199 Callopsenius Wasmann, 283 Ceratoxenus Mann, 106 Chledophila Cameron, 319 Cono8omu8 Motschulsky, 318 Coptotermoecia Oke, 248 Corotoca Schi0dte, 38, 141 Corymbogaster Mann, 65 Dabra Olliff, 319 Demerinda Cameron, 319 Dioxeuta Sharp, 217 Di8Coxenti8 Wasmann, 264 Dislicta Wasmann, 236 Doryloxenus Wasmann, 265 Drusilla Leach, 235 Eburniogaster Seevers, 41, 135 Eburniola Mann, 39, 150 Edaphus Motschulsky, 319 Eunannodes Silvestri, 46, 182 Eupsenius Wasmann, 283 E titer mitophila Cameron, 45, 179 Eutermitoptochus Silvestri, 150 Felda Blackwelder, 237 Fenye8ia Cameron, 319 Fon8echellti8 Silvestri, 130 Glaphyrinus Bernhauer, 209 329 330 FIELDIANA: ZOOLOGY, VOLUME 40 Hamitopsenius Wasmann, 276 Hetairotermes Cameron, 50, 201 Idiogaster Wasmann, 33, 109 Idioptochus Seevers, 33, 115 Iheringocantharus Bernhauer, 241 Jacobsonella Silvestri, 217 Lauella Mann, 27, 45, 177 Lissodiscus Grasse and Poisson, 262 Macrognathellus Silvestri, 45, 174 Macrotrichurus Silvestri, 189 Mastopsenius Seevers, 280 Megaxenistusa Seevers, 274 Microcyptus Horn, 283 Millotoca Paulian, 35, 119 M or melius Silvestri, 134 Myrmecopora Saulcy, 319 Myrmedonia Erichson, 11, 235 Nannellus Silvestri, 46, 181 Neodioxeuta Seevers, 220 Neophilotermes Seevers, 257 Neotermitogaster Seevers, 98 Oecidiophilus Silvestri, 130 Oideprosoma Silvestri, 40, 162 Orphnebius Motschulsky, 319 Osorius Latreille, 318 Oxypoda Mannerheim, 319 Oxytelus Gravenhorst, 318 Paederus Fabricius, 318 Paracorotoca Warren, 35, 119 Paraperinthus Seevers, 45, 180 Paratermitosocius Seevers, 199 Parvidolum Silvestri, 133 Pelioptera Kraatz, 319 Perinthodes Seevers, 45, 179 Perinthus Casey, 45, 167 Perlinctus Silvestri, 132 Philastilbus Bernhauer, 319 Philotermes Kraatz, 251 Philotermimus Reichensperger, 319 Poduroides Mann, 45, 175 Porus Westwood, 319 Protermitobia Seevers, 230 Pseudoperinthus Wasmann, 247 Ptocholellus Silvestri, 30, 69 Quediosoma Cameron, 318 Rhinotermopsenius Seevers, 278 Rhopalinda Cameron, 319 Schizelythron Kemner, 280 Silvestrinus Bernhauer, 222 Spirachtha Schi0dte, 11, 36, 39, 144 Termitana Fairmaire, 50, 208 Termitella Wasmann, 33, 110 Termitellodes Seevers, 114 Termitissa Reichensperger, 110 Termitobaena Bernhauer, 27, 225 Termitobia Wasmann, 226 Termitobiella Wasmann, 237 Termitobra Seevers, 50, 205 Termitochara Wasmann, 41, 139 Termitocola Seevers, 45, 172 Termitocolonus Seevers, 199 Termitocomes Seevers, 188 Termitoctesis Bernhauer, 110 Termitodiscus Wasmann, 259 Termitoecia Bernhauer, 50, 207 Termitogaster Casey, 89 Termitogerrus Bernhauer, 262 Termitohospes Seevers, 50, 192 Termitoiceus Silvestri, 127, 129 Termitoides Seevers, 99 Termitolara Bernhauer, 320 Termitolinus Wasmann, 224 Termitomimus Tragardh, 11, 36, 39, 153 Termitomorpha Wasmann, 84 Termitomus Silvestri, 200 Termitonannus Wasmann, 46, 182 Termitonda Seevers, 238 Termitonicus Mann, 45, 175 Termitonidia Seevers, 41, 136 Termitonilla Borgmeier, 20, 42, 190 Termitonusa Borgmeier, 241 Termitopaedia Wasmann, 216 Termitopelta Borgmeier, 45, 166 Termitophagus Silvestri, 240 Termitophya Wasmann, 28, 32, 71 Termitopithus Seevers, 80 Termiloplus Silvestri, 282 Termitopora Motschulsky, 319 Termitopsenius Wasmann, 275 Termitoptochus Silvestri, 11, 35, 116 Termitoptocinns Silvestri, 39, 148 Termitopulex Fauvel, 222 Termitopullus Reichensperger, 39, 147 Termiloquedhis Bernhauer, 318 Termitosaurus Silvestri, 318 Termitoscapha Bernhauer, 147 Termitosius Silvestri, 134 Termitosocius Seevers, 198 Termitosodalis Seevers, 200 Termitosomus Seevers, 84 Termitospectrum Mann, 50, 206 Termitosymbia Seevers, 245 Termitosyne Seevers, 81 Termitosy nodes Seevers, 83 Termitotecna Wasmann, 229 Termitotelus Wasmann, 250 Termitothymus Silvestri, 122 Termitotima Wasmann, 33, 108 Termitotropha Wasmann, 226 Termitozophilus Silvestri, 65 Termitozyras Seevers, 242 Termituncula Borgmeier, 28, 30, 64 Termitusa Wasmann, 51, 209 Termophila Lea, 201 INDEX 331 Termozyras Cameron, 235 Tetraphilus Silvestri, 182 Tetrasticta Kraatz, 320 Thaxteria Fenyes, 84 Thoracophorus Motschulsky, 318 Thyreoxenus Mann, 36, 40, 156 Timeparthenus Silvestri, 30, 69 Trachopeplus Mann, 88 Trianellus Silvestri, 130 Trichopsenius Horn, 267 Urodonia Silvestri, 236 Urolitus Silvestri, 320 Xenistusa LeConte, 272 Xenogaster Wasmann, 100 Xenopelta Mann, 106 Zunia Blackwelder, 236 Zyras Stephens, 233 III. Species and Subspecies acanthoscelis Seevers, 276 acuticornis Wasmann, 265 adamsoni Seevers, 244 africanus Seevers, 180 agilis Cameron, 202 akermani Warren, 119 albidus Seevers, 160 alutacia Oke, 248 arnica Mann, 78 anahuaci Seevers, 136 anastrephoproctus Silvestri, 129 annularis Cameron, 319 anoplotermitis Silvestri, 182 araujoi Borgmeier, 73 araujoi Seevers, 143 assmuthi Wasmann (Discoxenus) , 265 assmuthi Wasmann (Termitotima) , 108 australiensis Silvestri, 148 australis Seevers, 280 autuorii Silvestri, 158 badia Seevers, 127 barticae Seevers, 200 basalis Lynch, 318 beaumonti Casey, 123 bellicosi Silvestri, 264 beniensis Seevers, 96 bicolor Seevers, 93 bicolor Silvestri (Fonsechellus) , 131 bicolor Silvestri (Mormellus), 134 bi8etosns Seevers, 124 bolivari Seevers, 170 boliviae Seevers (Termitozyras), 245 boliviae Seevers (Thyreoxenus), 158 bovingi Mann, 175 brachycerus Silvestri, 186 brasiliana Seevers, 193 brasiliensis Silvestri, 189 braunsi Wasmann (Termitodiscus), 262 braunsi Wasmann (Termilotecna) , 230 brevis Mann, 97 brevitibialis Seevers, 160 brunneus Mann, 199 bryanti Bernhauer, 225 bryanti Cameron, 203 burgeoni Bernhauer, 263 burgeoni Cameron, 229 burmae Seevers, 221 butteli Wasmann (Doryloxenus), 300 butteli Wasmann (Felda), 237 butteli Wasmann (Termitodiscus), 262 californicus Seevers, 271 cameroni Reichensperger, 212 capritermitis Wasmann, 237 caudatus Wasmann, 277 cavernosa LeConte, 272 ceylonicus Silvestri, 118 ceylonicus Wasmann, 300 chavantinae Seevers, 94 clavicornis Wasmann, 284 colonus Seevers, 99 comes Seevers, 232 compacticornis Borgmeier, 124 convexinotus Seevers, 161 convexiusculus Wasmann, 318 convivens Silvestri, 319 cornuta Mann, 107 costaricensis Seevers, 86 crassicornis Wasmann (Discoxenus), 265 cras8icorni8 Wasmann (Macrognathel- lus), 174 crassiusculus Seevers, 81 croceipennis Motschulsky, 319 cubitopili8 Seevers, 255 cucullalu8 Seevers, 158 cylindricorni8 Seevers, 173 depressus LeConte, 268 disjunctipennis Seevers, 89 diversicollis Seevers, 98 diver sicolor Silvestri, 131 domuncidi Silvestri, 184 dudleyanus Casey, 167 echinoides Seevers, 187 ecuadoriensis Seevers, 74 elegantulum Silvestri, 68 emarginatu8 Seevers, 97 emer8oni Mann, 94 emer8oni Seevers (Philotermes) , 257 emersoni Seevers (Termitomimus), 38, 154 emersoni Seevers (Termitophya), 74 332 FIELDIANA: ZOOLOGY, VOLUME 40 entendveniensis Tragardh, 154 ericiogaster See vers, 199 erythraeanus Bernhauer, 222 escalerae Fauvel, 211 escherichi Wasmann (Idiogaster), 109 escherichi Wasmann (Termitodiscus), 260 eurymedusa Schi0dte, 146 eutermitis Wasmann, 300 exclusus Kemner, 115 fabulosa Bernhauer, 207 fasciatus Silvestri, 200 ferrugineus Kraatz, 319 fissipennis Casey, 86 flavescens Cameron, 218 flaviventris Mann, 76 fletcheri Cameron (Eutermitophila), 179 fletcheri Cameron (Pseudoperinthus), 248 formicicola Oke, 204 fossata LeConte, 272 fossulata Mann, 105 foveicollis Seevers, 132 foveolata Reichensperger, 111 fragilis, Seevers, 131 f rater Lynch, 318 frosti Seevers, 270 fuchsii Kraatz, 254 fulgens Borgmeier, 166 gastrophysa Eichelbaum, 227 gastrovittata Seevers, 152 gatuni Seevers, 187 gayi Seevers, 249 gestroi Bernhauer, 148 glabriventris Seevers, 105 gracilipes Borgmeier, 65 grandicornis Fauvel, 222 grandis Silvestri, 282 gridellii Bernhauer, 112 guatemalae Seevers, 172 guianae Seevers (Termitohospes), 194 guianae Seevers (Termitozyras), 245 guyanae Mann, 144 habilis Silvestri, 263 hageni Seevers, 172 heathi Wasmann, 318 heimi Wasmann, 260 hexagonalis Seevers, 273 heyeri Wasmann (Termitophya), 73 heyeri Wasmann (Thoracophorus), 318 holmgreni Wasmann, 76 huachii Seevers, 85 hystrix Wasmann, 211 iheringi Bernhauer, 318 impressicollis Seevers, 93 indicus Silvestri, 117 indosinensis Silvestri, 220 inflata Wasmann, 101 inornata Seevers, 74 inquilina Borgmeier, 106 insinuatus Silvestri, 318 insolens Casey, 91 insulanus Seevers, 204 insular is Fenyes, 85 jaliscensis Seevers, 138 japonicus Seevers, 271 javana Seevers, 178 javanicum Kemner, 281 kirbyi Seevers, 231 kohli Wasmann, 216 kraatzi Wasmann, 140 laetus Silvestri, 66 latebricola Lea, 201 latipes Seevers, 154 lativentris Seevers (Termitellodes) , 114 lativentris Seevers (Termitusa), 213 laxicornis Sharp, 257 leai Cameron, 177 lehmensicki Seevers, 116 lepidulus Grasse and Lesperon, 263 lepisma Wasmann, 265 leucogaster Mann, 151 liberiae Seevers, 238 limulus Seevers, 196 limulus Wasmann, 275 lobata Seevers, 126 longipes Seevers, 270 lujae Seevers, 152 lujae Wasmann (Termitella), 110 lujae Wasmann (Termitusa), 211 lunata Seevers, 137 luteola Borgmeier, 190 luzonicus Silvestri, 118 magdalenae Seevers, 95 mahout Mann, 176 major Mann, 38, 161 major Seevers, 170 major Wasmann, 184 malayanus Wasmann, 247 manni Seevers, 88 marginatus Seevers, 99 mayae Seevers, 170 meinerti Wasmann, 85 melantho Schi0dte, 142 micans Kraatz, 319 michoacani Seevers, 138 micronesiae Seevers, 278 microps Seevers, 198 microps Sharp, 217 microsomatis Silvestri (Parvidolum), 133 microsomatis Silvestri (Termitonannus), 186 mimellus Silvestri, 130 mimus Silvestri, 70 minutus Cameron, 262 mirabilis Mann, 147 INDEX 333 mirandum Silvestri, 164 mirandus Mann, 66 miricorniger Seevers, 193 mirotermitidis Paulian, 120 mundus Bernhauer, 212 nana Seevers, 103 natalensis Wasmann (Termitolinus), 225 natalensis Wasmann (Termitopulex), 223 nigeriensis Silvestri, 320 nigra Cameron, 319 nigricollis Silvestri, 95 nitida Seevers, 246 notabilis Silvestri (Macrotrichurus), 190 nolabilis Silvestri (Urodonia), 236 novaehollandiae Silvestri, 150 ochraceus West wood, 319 oglobini Silvestri, 69 o'neili Wasmann, 226 opaca Kraatz, 319 opacella Bernhauer, 320 opulentus Bernhauer, 319 oudemansi Franssen, 221 palauensis Seevers, 178 panamanensis Seevers, 194 paolii Gridelli, 229 paranensis Lynch, 318 parviceps Mann, 157 parvulus Silvestri, 186 pauciseta Seevers, 181 pauciseta Silvestri, 134 penetrans Silvestri, 318 peninsularis Silvestri, 118 pensylvanicus Kraatz, 256 peradenyiae Wasmann, 300 perinthoides Seevers, 205 perrieri Fairmaire, 209 philetaerus Silvestri, 127 philippinus Silvestri, 118 phyla Schi0dte, 143 physogastra Wasmann, 227 piceus Cameron, 202 piceus Wasmann, 260 pilivenlris Seevers, 78 pilosula Seevers, 102 pilosus Kraatz, 252 platygastra Seevers, 82 polita Kraatz, 320 politus Cameron, 236 pressa LeConte, 273 proximatus Silvestri, 186 pubicollis Seevers, 126 pulchellus Mann, 157 punctata Mann, 80 puncliceps Seevers, 92 punctiventri8 Lea, 203 quadricollis Cameron, 213 quaerens Silvestri, 264 quaesitus Silvestri, 132 recondili Silvestri, 182 regius Silvestri, 69 reichenspergeri Bernhauer, 320 reichenspergeri Seevers, 104 rhinotermiti8 Seevers, 275 rhodesiae Seevers, 228 rubricollis Seevers, 112 rurrenabaquensis Seevers, 124 saltatoriu8 Seevers, 279 schioedtei Mann, 146 schmalzi Wasmann, 183 schultzei Wasmann, 250 sequax Borgmeier, 241 8ericoiliu8 Cameron, 318 setiger Reichensperger, 319 setipes Wasmann, 238 setosus Mann, 88 setosus Seevers, 187 silvestrii Seevers (Atheta), 319 silvestrii Seevers (Termitohospes) , 197 silvestrii Wasmann (Termitocola), 173 silvestrii Wasmann (Termitonannus), 183 8imopelta Mann, 86 simulans Cameron, 318 simulans Mann, 85 simulans Silvestri, 129 sjoestedti Eichelbaum, 223 sjoestedti Wasmann, 210 sociusculus Reichensperger, 148 solomonen8is Seevers, 162 splendidus Wasmann (Doryloxenus) , 300 splendidus Wasmann (Termitodi8cu8) , 262 subnuda Seevers, 103 sumatranus Silvestri, 118 synterminus Silvestri, 240 8yntermiti8 Silvestri, 319 tachyporoides Seevers (Termitohospes), 197 tachyporoides Seevers (Termitonda), 239 langanyikae Seevers, 120 tarascani Seevers, 139 tarsatus Mann, 172 termitaria Bernhauer (Atheta), 319 termitaria Silvestri (Atheta), 319 termitis Motschulsky, 319 termitobia Silvestri, 217 termitobia Wasmann, 319 termitocola Seevers, 135 termitophila Bernhauer, 319 termilophila Cameron (Demerinda), 319 termitophila Cameron (M yrmecopora) , 319 termitophila Cameron (Rhopalinda), 320 termitophila Lea, 319 termitophila Motschulsky, 319 termitophilus Bernhauer, 319 termitophilus Cameron, 318 termitophilus Wasmann (Conosomus), 318 334 FIELDIANA: ZOOLOGY, VOLUME 40 termitophilus Wasmann (Doryloxenus) , 300 termitophilus Wasmann (Paederus) , 318 testaceus LeConte, 283 testaceus Seevers, 92 texana Brues, 135 thoracicum Mann, 206 transfuga Wasmann, 300 transvaalensis Silvestri, 262 triarticulatus Kemner, 300 tricornis Mann, 107 trinidadensis Seevers, 188 unicolor Silvestri, 194 uroclaviger Silvestri, 176 vagans Silvestri, 184 validus Silvestri, 294 vestitus Mann, 199 vicinior Silvestri, 260 vitiensis Mann, 177 wasmanni Holmgren (Termitogaster), 92 wasmanni Holmgren (Termitophya), 76 wasmanni Mann, 168 wasmanni Seevers, 188 wenzeli Seevers, 96 werneri Seevers, 255 williamsi Seevers, 83 wilsoni Cameron, 208 xenocostalis Seevers, 170 xenoflavipes Seevers, 270 ypiranganus Bernhauer, 242 Publication 815