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Phylogeny, chaetotaxy, and ecology of the Onthophilinae and (Coleoptera: Histerdae)

Kovarik, Peter William, Ph.D.

The Ohio State University, 1994

UMI 300 N. Zeeb Rd. Ann Arbor, MI 48106

PHYLOGENY, CHAETOTAXY, AND ECOLOGY OF THE ONTHOPHLLINAE AND TRIBALINAE (COLEOPTERA; HISTERDAE)

DISSERTATION

Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University

By

Peter William Kovarik, B.A., M.S.

*****

The Ohio State University 1994

Dissertation Committee: Approv

W. A. Foster D. J. Horn N. F. Johnson Co-Adviser C. A. Triplehom Department of Entomology To Horace R. Burke and Rupert L. Wenzel

“The study o f entomology is one o f the mostfascinating ofpursuits. It takes its votaries into the treasure-houses of Nature, and explains some o f the wonderful series o f links whichform the great chain o f creation. It Ic^s open before us another world, o f which we have been hitherto unconscious, and shows us that the tiniest , so small perhaps that the unaided eye can scarcely see it, has its work to do in the world, and does it. ”

Rev. J.G. W ood

u Acknowledgments

My initial thanks go to my advisors Dr. Norman F. Johnson and Dr. Charles A. Triplehom for their guidance, support, and camaraderie, I wish to also thank close friends and committee members Dr. Woodbridge A. Foster and Dr. David J. Horn for sharing their ideas on science and philosophy. I am grateful for the tutelage of Rupert L. Wenzel and Alfred F. Newton, who were functionary, though unoflScial, committee members. The collective editorial comments of the aforementioned people substantially improved this document. I am particularly indebted to George Keeney for key assistance in rearing. Special thanks are due P. Skelley, K. Stephan, R. W. Jones, A. F. Newton, J. W. Summerlin, M. Archangelsky, W. Godwin, B. Carr, D. Verity, and R. Westcott for providing me with preserved immatures and /or culture stock. Thanks are also due K. Roach, P. Kaufl&nan, P. Lima, and D. Knott (USDA/APHIS/PPQ) for providing me with importation permits. I owe much to the following individuals for financial assistance for the period beyond my allotted departmental support: N. F. Johnson, J. Kovarik, B. Armitage, K. Summers, D. McIntyre and L. Summers. I wish to acknowledge the artistic and technical contributions of R. Hancock, S. Hakola, J. Mitchell, and J. Diaz. I also wish to thank J. Wenzel and C. Marshall for assistance with HENNIG86.

Ill I am extremely grateful to Dr. James W. Kimbrough for fungal spore identifications. I wish to thank the following individuals and lending institutions for the specimens I borrowed: L. H. Herman, American Museum of Natural History, New York; E. DuBoise, British Museum of Natural History, London; E. R. Hoebeke, Cornell University, Ithaca; H. R. Burke and E. G. Riley. Texas A&M University, College Station; H. M. André, Musee Royal De L’Afiique Centrale, Tervuren; J. F. Lawrence, Australian National Insect Collection, Canberra; E. Matthews, South Australian Museum, Adelaide; R. L. Wenzel and A. F. Newton, Field Museum of Natural History, Chicago; N. Berti, Paris Museum, M. Uhlig, Humboldt University Museum, Berlin. Thanks are due Karl Stephan, C. Klopp, and J. Wenzel for translation assistance and Kim Summers for formatting and proofing this document. Finally I wish to thank Miguel Archangelsky, Brian Burrell, Flavia Ejchel, Bob Jones, Bob Hancock, Jolanda Kovarik, John Mitchell, Steve Passoa, Keith Philips, Brady Porter, Paul Skelley, Jim Stoeckel, Kim Summers, Nancy Tomei, and Bruce Weaver for moral support.

IV VTTAE

May 25,1957 Bom - Norwalk, CT

1975-1979 ...... B.A. Biology Quinnipiac College Quinnipiac, CT

1979-1983 ...... M.S. Entomology Texas A&M College Station, TX

MAJOR AREA OF STUDY Entomology Table of Contents

Dedication ...... ii

Acknowledgments ...... iii

Vita...... iv

List of Tables ...... vii

List of Figures...... viii

CHAPTER I Chaetotaxy of Larval (Coleoptera: ) based on a Description of Onthophilus nodatus LeConte ...... 1 Introduction ...... 1 Materials and Methods ...... 4 Terminology and Homology ...... 3 Discussion ...... 16

CHAPTER n Pupal Chaetotaxy of Histeridae (Coleoptera: Hydrophiloidea) with a Description of the Pupa of Onthophilus kimi Ross ...... 39 Introduction ...... 39 Materials and Methods ...... 40 Results...... 40 Discussion ...... 45

CHAPTER m Development of divisus Marseul (Coleoptera: Histeridae) Introduction ...... 50 Materials and Methods ...... 51 Results...... 53 Discussion ...... 56

VI CHAPTER IV A Revision of the Genera of Onthophilinae and Tribalinae (Coleoptera: Histeridae) ...... 61 Introduction ...... 61 Materials and Methods ...... 63 Terminology and Morphology ...... 64 Results...... 75 Phylogentic Analysis ...... 158

CHAPTER V Chaetotaxy of the Immature Stages of the New World Idoliini (Coleoptera: Histeridae) Based on gibba Lewis and Eutribalus americamis (LeConte) ...... 257 Introduction ...... 257 Results...... 258 Discussion ...... 269

APPENDICES Appendix A ...... 307 Appendix B ...... 313 Appendix C ...... 317 Appendix D ...... 318 Appendix E ...... 320 Appendix F ...... 322

REFERENCES...... 323

YU List of Tables

CHAPTER I Table 1. Material Examined ...... 3 Table 2. Examples of setal and pore homologs found in carabid, staphylinied, and histerid larvae ...... 18

CHAPTER n Tables. Material Examined ...... 42

CHAPTER m Table 4. Duration of developmental stages for Epierus divisus under laboratory conditions ...... 54

Table 5. Duration of developmental stages for Euspiloius assimilis under laboratory conditions ...... 54

Table 6. Independent t-test comparing duration of developmental stages of Epierus divisus verus Euspiloius assimilis under laboratory conditions ...... 54

CHAPTER IV Table 7. Material Examined ...... 65

vm Table of Figures

CHAPTER I Figure 1...... 19 Figure 2 ...... 21 Figure 3 ...... 23 Figure 4 ...... 25 Figure 5...... 27 Figure 6...... 29 Figure 7 ...... 31 Figure 8 ...... 33 Figure 9 ...... 35 Figure 10...... 37

CHAPTER n Figure 11...... 46 Figure 12...... 48

CHAPTER IV Figure 13...... 167 Figure 14...... 169 Figure 15...... 171 Figure 16...... 173 Figure 17...... 175 Figure 18 ...... 177 Figure 19 ...... 179 Figure 20...... 181 Figure 21...... 183 Figure 22...... 185 Figure 23...... 187 Figure 24...... 189 Figure 25...... 191 Figure 26...... 193

IX Figure 27...... 195 Figure 28 ...... 197 Figure 29 ...... 199 Figure 30...... 201 Figure 31...... 203 Figure 32...... 205 Figure 33...... 207 Figure 34...... 209 Figure 35...... 211 Figure 36...... 213 Figure 37...... 215 Figure 38 ...... 217 Figure 39 ...... 219 Figure 40...... 221 Figure 41...... 223 Figure 42...... 225 Figure 43 ...... 227 Figure 44 ...... 229 Figure 45...... 231 Figure 46...... 233 Figure 47 ...... 235 Figure 48 ...... 237 Figure 49 ...... 239 Figure 50...... 241 Figure 51...... 243 Figure 52...... 245 Figure 53...... 247 Figure 54...... 249 Figure 55...... 251 Figure 56...... 253 Figure 57...... 255 Figure 58 ...... 256

CHAPTER V Figure 59 ...... 271 Figure 60...... 273 Figure 61...... 275 Figure 62...... 277 Figure 63...... 279 Figure 64...... 281 Figure 65...... 283 Figure 66...... 285 Figure 67...... 287 Figure 68 ...... 289 Figure 69 ...... 291 Figure 70...... 293 Figure 71...... 295 Figure 72...... 297 Figure 73 ...... 299 Figure 74...... 301 Figure 75...... 303 Figure 76...... 305

XI CHAPTER I

CHAETOTAXY OF LARVAL HISTERIDAE (COLEOPTERA: HYDROPHH.OIDEA) BASED ON A DESCRIPTION OF ONTHOPHILUS NODATUS Ju^COmE

Introduction Histeridae is a large family with >300 genera and 3,700 species worldwide (Newton 1991). The family is considered most closely related to Synteliidae, Sphaeritidae, and Hydrophilidae (Lawrence & Newton 1982). A comprehensive cladistic analysis of the Hydrophiloidea or the Histeridae has not yet been attempted. According to Newton (1991), larval histerids are poorly known and >30 genera have been described. These descriptions are based on gross morphological features such as the antennae, mouthparts, legs, and urogomphi. Study of the distribution of setae (mechanoreceptors), other sensilla, and pores has been largely neglected. Immatures provide an array of new characters that supplement those of the adult stage (Goulet 1979). Proper setal nomenclature and homology can be extremely valuable for use and interpretation of this data set. Accurate serial homology permits recognition of variations in setal patterns and sclerite development on segments of different tagmata. It also eliminates the need to generate different nomenclatural systems for the thorax and abdomen respectively (e.g., Bousquet & Goulet 1984). Few attempts have been made to homologize the setae of beetle larvae, and no system exists that can be directly applied to larval histerids for morphological descriptions. In this study I serially homologize the setae of the thorax and abdomen (with the exception of abdominal segment 10) based on a detailed morphological description of

1 2 Onthophilus nodatus J. E. LeConte. The Onthophilus was chosen because it is missing only two primary setae, and the tribe Onthophilini is considered ancestral by Crowson (1974) because of the many plesiomorphic characters found in both larvae and adults.

Materials and Methods Parental stock was obtmned in October and November from Latimer County, OK, using pitfall traps baited vrith swine feces. Voucher specimens were deposited in the larval collections of The Ohio State University, the Field Museum of Natural History, Chicago, and the collection of P.W.K. Descriptions of larvae are based on several specimens cleared in lactic acid and mounted in depression slides with Hoyers’ medium. By warming the slide and moving the coverslip, specimens can be rotated along their longitudinal axis and thus observed at any angle. Specimens in depression slides allow room for setae to orient more or less as they would in a living specimen. Therefore, some setae appear foreshortened. By including dorsal, ventral, and lateral views of each segment, setae that were illustrated foreshortened in one view are likely to be fully extended in another. The specimens were studied at 200x with a Wild M12 compound microscope fitted with a Bausch and Lomb stereo polarizer. Mouthparts and legs, which required high magnification, were removed and mounted on standard glass slides. The proposed setal and pore notation is based on the study of first and second instar Onthophilus nodatus. Larvae of O. kimi Ross and 0. giganteus Helava were also examined for any significant variations in setal and pore distribution within the genus. Exemplars from outgroups (Hydrophilidae and Synteliidae) and all recognized histerid subfamilies (except Niponiinae, Trypeticinae, and Hetaerinae) (Table 1) were examined to Table 1. Material Examined.

Subfamily Species Carabidae Pterostichus sp. Staphylinidae PMonthis sp. Synteliidaie Syntelia histeroides Lewis Hydrophilidae Helophous orientalis Motschulslqr Histeridae Aletes politus LeConte Plegaderus transversus Say Tryponaeinae Trypanaeus sp. fratemus Say Geomysaprimts goffl Ross psyche Casey pensyvcmicus PaykuU Euspiloius assimilus PaykuU sp. pumilio Erichson Xestipyge multistriatum Lewis sp. Anaplaeus marginatus LeConte sp. Platylomalus aequalis Say seeversi Wenzel Onthophilinae Onthophilus nodatus LeConte 0. kimi Ross O. giganteus Helava Tribalinae Epierus divisus Marsuel Idolia gibba Lewis Parepierus sp. bimaculatus L. egregius Casey abbreviatus F. Spilodiscusfloridarms Ross lecontei Marseul Baconia sp. Hololepta sp. Iliotona sp. Phelister panamensis LeConte 4 ensure that my system could be extrapolated to other related taxa. Setal maps of the metathorax and first abdominal segment were prepared for all histerid genera listed in Table 1. This was done to ensure that Onthophilus larvae were not missing a significant number of primary setae. Setae were individually measured and subsequently grouped into four categories based on overall length: minute (<0.033 mm); short (0.033-0.063 mm); medium (0.064-0.141 mm); and long (>0.141 mm). The exact measurements used to delimit size classes may change in subsequent descriptions, depending on the size of the larvae. Mandibular seta 2, reduced to the point of being unrecognizable as a seta, is independently categorized as vestigial. Number and location of setae are generally constant between the two instars; therefore, all setae are regarded as primary.

Terminology and Homology My procedure for determining serial and general homologies was a modification of the approach used by Belkin (1950). I selected the first abdominal segment as a starting point for the setal maps. My intent was to identify the full complement of setae on this segment and homologize them with those of the thorax. However, the discovery of 3 minute setae on the anterior margin of the metathorax, with no apparent serial homologs on the first abdominal segment, required special consideration. I decided not to label these setae as “extras”; instead, I incorporate them into my nomenclatural system. This decision was influenced by the occurrence of apparent homologs of these setae on both the thorax and abdomen of larval staphylinids and carabids. I consider it is probable they are lost fi*om the abdomen of larval histerids. Onthophilus nodatus larvae lack at least one trochanter seta (TR4) and an abdominal pleural seta (PL22). On the basis of outgroup comparison, I interpret the loss of these setae in O. nodatus as a derived feature. I have accordingly left room for these setae in my numbering system. 5 The abdomen and, to a lesser degree, the thorax are modified for crawling via peristaltic waves of contractions. The thoracic legs of histerid larvae are reduced and do not appear to play a significant role in locomotion. The abdominal tergites and stemites are greatly subdivided. Major differences exist in the sclerite and setal distribution of the thorax and most abdominal segments. However, sclerite and setal distribution on the first abdominal segment is somewhat intermediate between those of the rest of the abdomen and thorax, enabling approximate determinations of the serial homologies among abdominal and thoracic setae. Relative size and setal position, as well as certain stable morphological features such as pores, certain sclerites, spiracles, and egg bursters, were used to determine both general and serial homologies. Three setae (TES, 9, and 19), consistently elongated in most histerid genera, are particularly helpful for determining setal homologies on the thorax and abdomen. The nota of the metathorax and abdomen are conspicuous sclerites which generally bear a lateral (TE8, 9) and medial (TEIO, 11) pairs of setae and 2 pores (TEf, g) on each side. Likewise the anterior pleurites of the metathorax and abdomen are easily located and consistently bear three setae (PL23, 24 and 25). The ninth abdominal segment is unlike the other abdominal segments in that it bears paired urogomphi. The sclerites and setal pattern of this segment differ fi*om those of preceding abdominal segments. The setae and pores of the urogomphi are not considered to have serial counterparts on any other segments. The tenth abdominal segment or pygopod articulates ventrally on abdominal segment 9 and is modified into an ambulatory structure. Although it seems probable that setae on this segment have serial counterparts on other abdominal segments, their identities are not yet clear. Therefore, setae on this reduced and highly modified segment are numbered separately as pygopod setae. Similarly, no attempt was made to serially homologize the setae occurring on the head capsule with those elsewhere on the body. 6 Larval sclerites have two distinct textures: smooth and granular. Smooth cuticle is platelike, shiny, and generally darkly pigmented. Granular cuticle appears less rigid and is coarsely textured and lightly pigmented. In histerid larvae, smooth cuticle is generally restricted to the head, certain portions of the thoracic sclerites, legs, and urogomphi. In addition there are also areas of mosaic patterning associated with the protergum. Pores are usually associated with smooth cuticle. The majority of pores appear to be campaniform sensilla, but a minority may be glandular ducts through the sclerotized cuticle. No attempt has been made to ascribe function to any of the sensilla and therefore the term “pore” refers to a minute circular landmark of unknown function. In larval histerids, pores are generally confined to the head, tergal regions of the thorax and abdomen, and urogomphi. Because of the highly dissected nature of the abdominal tergites, many of the pores present on the thorax are presumably lost on the abdomen. The pronotal shield represents the least subdivided tergite present in histerid larvae. Consequently pore homology is based on this region. The notation for setae and pores used in this article closely follows Bousquet & Goulet (1984). Setae are coded by two capital letters and a number. The letters are an abbreviation of the sclerite or appendage on which the seta is located. Pores are similarly coded with a lower case letter distinguishing a specific pore. Setal groups consisting of a variable number of setae are coded by two capital letters preceded by a lower case “g” (for group). The following abbreviations stand for the appendage, part of appendage, structure, etc., that generally bear setae: AN, antenna; AS, anterior stemite; AT, anterior tergite; AP, anterior pleurite; CO, coxa; DH, dorsohumeral tergite; DL, dorsolateral tergite; ES, epistemum; EM, epimeron; FE, femur; PR, fi-ontale; HT, humeral tergite; IS, interstemite; IT, intertergite; LA, labium; LH, laterohumeral tergite; LP, lateral prestemite; LS, lateral stemite; IT, lateral tergite; MN, mandible; MX, maxilla; PC, precoxite; PL, posterior pleurite; PP, pygopod; PR, prestemum; PS, prestemite; PT, protergum; ST, 7 stemite; TA, tarsungulus; TI, tibia; TR, trochanter; UG, urogomphus. Additionally, several other structures are designated by similar notations: EB, egg burster; PE, penicillus; SD, sensillum digitiformium; SE, sensorium; TF, tentorial fossa. The three setal regions on the first abdominal segment of Onthophilus nodatus are tergal, pleural, and sternal. The boundaries of these regions are defined by the relative positions of specific setae and the relationship of some of these setae to the spiracle. For instance, the tergal setal group are those setae which occur above the spiracle. The pleural and sternal setae occur below the spiracle. The pleural setal group are decidedly lateral. The landmarks 1 have used to delimit these regions were easily recognizable in histerid genera I examined (Table 1). There are two types of setae on the head capsule of histerid larvae. The first are typical mechanoreceptors and are rigid, nonplumose, have a socket, and occur singly. The second type are flexible, often plumose, generally lack a socket, and often occur in groups. These modified setae are restricted to the labium, maxilla, and the anterior margin of the fi-ontale. Most other sensilla are not coded but are described and figured. The majority are short and bluntly rounded, those on the antenna of Idolia are long, tapered, and resemble mechanoreceptors. This latter type of sensillum is termed “setiform”. Three types of aspérités occur in histerid larvae: (1) tubercles, minute wartlike prominences lacking microtrichia; (2) chalazae, minute prominences bearing a tapered, often elongate spinule; (3) denticles, minute prominences bearing short, stout, thomlike projections. Denticles are restricted to the tergum and sternum of the thorax and abdomen. They occur on areas of the body which come in contact with the substrate during movement such as the prolegs and ampullae. In Baconia the proleg denticles closely resemble the crochets of caterpillars. Tubercles and chalazae are often widely distributed on the membranous areas of the thorax and abdomen. 8 My interpretation of the morphology of the maxilla differs slightly from that of Newton (1991). I regard the proximal palpal segment as a fusion of sclerites from both the galea and palp. This fusion creates the illusion that the galea (digitiform appendage) arises from the apex of the first palpal segment. A similar modification of the maxilla occurs in larval Sialis. Nasale Morphology. The morphology of the anterior margin of the frontale (nasale) (Fig. 3 A) varies considerably among histerid taxa. Seta FRl may be found on the dorsal surface of this area or at the anterior border as in Onthophilus. Four minute peglike setae are consistently found associated with the frontale teeth. The gFR setal group may arise from sockets or may appear fused to the head capsule. There are additional setae arising from the ventral surface of the nasale, most of which are vestigial or minute and concealed from above. In some histerid genera, some of these setae are prominent, extending well beyond the anterior margin of frontale. I term these subnasal setae (SN). Thoracic Morphology. Sclerites occurring on the thorax can be easily homologized among genera. I therefore considered it informative to apply names to the larger sclerites on the thorax. Although every attempt was made to apply existing terms to the major thoracic sclerites of histerid larvae, the current terminology did not adequately describe some morphological features peculiar to this group. Thus the creation of new terms was sometimes unavoidable. In some histerid larvae there are intersegmental sclerites. I refer to these sclerites collectively as intersclerites and specifically as intertergites (IT) and interstemites (IS). Prothorax (Fig. 9A-C). The protergum (PT) consists of a single sclerotized plate extending ventrad to a membranous pleat which is bordered by setae PL23,24, and 27 (see Fig. 4B). This sclerite is actually a composite of tergal and dorsal pleural elements. The epistemum (ES) is a large sclerite beneath the pronotum and anterior to the procoxa. 9 A very short pleural suture extends dorsad of the coxal articulation. This suture separates the epistemum from the much smaller epimeron (EM), which occurs posteriodorsad of the fore coxa. Below the epistemum lies the prestemum (PR). In larval histerids this area is always subdivided into a medial sclerite (prestemite [PS]) and two lateral sclerites (lateral prestemite [LP]). Anteromedial to the procoxa is a minute sclerite to which I refer as precoxite (PC). Directly posterior to the prestemum and lying between the fore coxae is the prostemite (ST). Flanking the prostemite on each side are two small sclerites which lack setae in all larval histerids examined thus far. I call these lateral stemites (LS). Mesothorax (Fig. 9A-C). Humeral tergites (HT) are paired teigites directly behind the protergal shield. The mesonotum covers most of the dorsal surface of the mesothorax. Directly behind the outer posterior margins of the mesonotum occur a pair of intertergites (IT) which lack setae. A single pair of lateral subdivisions of the notum, termed lateral tergites (LT), occurs below the mesonotum. Directly below the lateral tergites are the pleurites. There are two primaiy subdivisions of the pleural sclerites, anterior (AP) and posterior pleurites (PL). Directly below the posterior pleurites and dorsal to the mesocoxae are the anterior epistemum and the posterior epimeron. Below the epistemum and anterior to the coxae are the precoxites. Anterior to the precoxites is a pair of minute anterior stemites (AS). Behind the precoxites are the lateral stemites. Between the lateral stemites lies the mesostemite. Metathorax (Fig. 9A-C). There are three pairs of humeral tergites, a dorsal pair (DH) and two lateral pairs (LH). Posterior to the dorsal humeral tergites lies the metanotum. An additional tergal subdivision, the dorsolateral tergite (DL), is located between the metanotum and the lateral tergite. Two pairs of anterior stemites are on this segment. The remaining sclerites resemble those found on the mesothorax. Abdominal Morphology. Abdominal sclerite morphology is relatively stable within a genus but is highly variable among genera. The histerid larval abdomen consists 10 of numerous tergal, pleural, and sternal subdivisions. Tergal elements occur above the level of the spiracle. Pleural elements lie below the level of the spiracle. Egg bursters (EB) (Fig. 6A) are associated with the dorsal humeral sclerite of the first abdominal segment of the first instar. Intertergites and interstemites occur in some but not all the examined taxa. Abdominal segments I-VUI in histerid larvae typically bear both dorsal ampullae and ventral prolegs. Retractor muscles are associated with the prolegs of larval histerids. Enlarged asperities are generally found in association with ampullae and prolegs. The morphology and chaetota^qr of abdominal segment IX is considerably different firom the preceding abdominal segments. Consequently, serially homologizing setae on this segment presented a challenge, particularly because the urogomphi and their associated setae are restricted to this segment. My concept of what constitutes segment IX may actually be a composite of the anterior intersegmental region and the segment proper. On segment IX there are two pairs of tergites without setae anterior to TEl. I term these sclerites anterior tergites (AT) (Fig. lOE). Ventrally, a pair of interstemites (IS) (Fig. lOF) occurs anterolaterad of ST30 (see Fig. 8A). Laterad to the interstemites is a small sclerite without setae which appears to be a subdivision of the midsized sclerite bearing ST36 (see Fig. 8B) on abdominal segments n-Vm. Segment X in histerids is a reduced ambulatory structure or pygopod. Results First Instar. (Figs. 1-8). Head. Head capsule width: 0.45 ± 0.01 mm (X ± SD; n = 6). Epicranial suture lyriform with short stem, medial ecdysial line absent. Stemmata absent. Frontale. (Figs. 1 and 10) with 11 setae (FRl-FRl 1), 1 setal group (gFR) and 6 pores (FRa-FRf) on each side. Setal lengths: minute (FRl, 3, 4, 5, 6, 7, 9,10,11); medium (FR2, 8). gFR consisting of 4 short nonplumose setae with distinct sockets. Nasale with 2 prominent asymmetrical truncate teeth. Several vestigial and single short 11 subnasale seta present. Parietal. (Figs. lA-B, 4A) with 19 setae (PA12-PA30) and 13 pores (PAg-PAs) on each side. Setal lengths: minute (PA12, 13, 16, 17, 18,19,24, 28), short (PA15, 20,26,27), medium (PA21,2 9, 30), long (PA14,22, 23, 25). Antenna. (Fig. 2A-C): Segment 1, sclerotized portion with 4 pores (ANa-d); ANb, d dorsolateral, ANa, c mesolateral; distal membrane with ventral sensillum. Segment 2 with single dorsal pore (ANe), distal membrane with 4 papillate sensilla and 2 sensoria (SE1,2). Three sensilla dorsal, one ventral. Dorsal sensilla near base of sensorium 1 <1/2 length of remaining 3, which are about same length as sensorium 1. Segment 4 with 2 terminal and 4 equidistantly spaced papillate sensilla surrounding terminal pair, longer member of terminal pair slightly longer than other distal sensillae. Mandible. (Fig. 2D) with seta of medium length (MNl), a vestigial seta (MN2) and 3 pores (MNa-c); pores MDb dorsolateral; a, dorsal; c, dorsomedial. Apex of mandible pitted, incisor area bidentate with small tooth above retinaculum. Penicillus (PE) setae nonplumose. Maxilla. (Fig. 2E- F ) Stipes with 6 (MXl-6) setae, 4 pores (MXa-d), 2 setal groups (gMXl-2) and 4 short, flexible nonplumose setae on dorsal distal membrane. Setal lengths: minute (MXl, 6), short (MX2, 5), medium (MX4), long (MX3). Pores MXa, c, d dorsal, MXb lateral; gMXl consisting of numerous flexible plumose setae; gMX2 consisting of 4 minute rigid setae with sockets. Proximal palpal segment with 3 setae (MX7-9), 2 pores (MXe, f), 4 flexible nonplumose setae dorsal on membrane. Setal lengths: minute (MX9); short (MX7, 8). Pore MXe dorsal, MXf lateral; 2 setae (MX8, 9) arising terminally from galea. Single minute sensillum mesoventral on membrane. Second palpal segment with single dorsal pore (MXg) and mesoventral sensillum. Segment 3 with single dorsal pore (MXh) and several porelike sensilla. Segment 4 with 10 porelike sensilla. Prominent sensillum digitorum (SD) arising dorsally. Eleven villiform sensilla basiconica of varying lengths arising from palpal apex; all shorter than sensillum digitiformium. Labium. Mentum (not illustrated) reduced, somewhat concealed, with distal portion membranous, bearing 12 numerous flexible plumose setae dorsally and two tapering membranous arms laterally; proximal portion sclerotized and fused to undersurface of frontale. Prementum (Fig. 2G- H) with small obtuse spinose lobe on each side, base of dorsal surface bearing small teeth and tubercles, with 3 setae (LAl-3), 2 pores (LAa-b), and numerous sensilla. Setal lengths: minute (LAI, 3), short (LA2). Sensillum distribution as follows: single minute sensilla on membrane at base of first palpal segment ventrally, 2 short peglike sensilla flanking seta LA3 in distal membrane of prementum dorsally. Labial palp 2 segmented, first palpal segment with sensillum in membrane dorsally. Second palpal segment with prominent sensillum digitiformium dorsally and 10 circular sensillae distributed over surface. Eleven villiform sensilla basiconica of varying lengths arising from apex of palpus, all shorter than sensillum digitiformium. Thorax. Prothorax. (Figs. 4B, 5A-B). Notum with 21 setae (TEl-5, 7-13, 16-21, 23, 24, 27) (shield with subset [TEl-5, 7-13,16,17]), and 13 pores (TEa-m) on each side. Setal lengths: minute (TEl, 3, 7, 10, 16, 18,20, 21,24, 27), short (TES, 11, 12, 17, 23), medium (TE2, 4, 13), long (TE5, 9,19). Epistemum without setae; epimeron subdivided, upper half with single short seta (ST40). Presternum'. both prestemite and lateral prestemite well developed, lateral prestemite extending posteriorly beyond prestemite by 0.25 its length; prestemite roughly quadrate with anterior angles slightly convergent; prestemum with 9 setae (PRl-9) on each side. Setal lengths: minute (PRl-5, 9), short (PR8), medium (PR7), long (PR6). Prostemite diamond-shaped and about as long as wide, central area with smooth cuticle; with 3 setae (ST30,44,46) on each side. Setal lengths: minute (ST30,44), long (ST46). Lateral stemite small and oval: precoxite very small with 3 minute setae (ST31,32, 36). Mesothorax (Figs. 4B, 5A-B). Notum with 8 setae (TE3-5, 7-11) and 5 pores (TEe-i) on each side. Setal lengths: minute (TEIO), short (TE3, 7), medium (TE4, 8, 11), long (TES, 9). Humeral tergites each with 5 minute setae (TEl, 2, 12, 13,14) and 3 pores (TEa-c). Intertergite small and oval. Lateral tergite with 13 7 setae (TEl5-21) and single pore (TEj). Setal lengths: minute (TEl5,20,21), short (TE16,17,18), long (TEl 9). Posterior pleurite large, rectangular, with 2 setae (PL27, 28) arising in posterior 1/3; setal lengths: minute (PL28), medium (PL27). Anterior pleurite 1/6 as large as posterior pleurite and partly surrounding thoracic spiracle, with 2 minute setae (PL23,24). Anterior stemite with single minute seta (ST36). Epistemum with 2 minute setae (PL34,35). Epimeron with single short seta (ST40). Precoxite with 1 short (ST32) and 1 medium-length seta (ST31). Lateral stemites elongate and narrow, slightly shorter than mesostemite. Mesostemite with 4 setae (ST29, 30,44,46) on each side. Setal lengths: minute (ST29), medium (ST30,44), long (ST46). Mesostemite roughly quadrate. Metathorax (Figs. 17-19). Notum with 4 setae (TE8-11) and 2 pores (TEf, g) on each side. Setal lengths: minute (TEIO), medium (TE8, 11), long (TE9). Dorsal humeral tergites each with single minute setae (TE2) and 1 pore (TEb). One minute seta (TEl) just anterior to dorsal humeral tergite. Lateral humeral tergites each with single minute seta (TE12, 13), a single minute humeral seta (TE14) located next to anterior margin of lateral tergite. Intertergite small and oval. Single short seta (TE3) on small sclerite in space between notum and dorsolateral tergite. Dorsolateral tergite with 3 setae (TE4, 5,7) and 2 pores (TEe, i). Setal lengths: minute (TE7), medium (TE4), long (TE5). Lateral tergite nearly identical to homolog on preceding segment. Anterior 1/2 of posterior pleurite slightly tapered, otherwise similar to mesothoracic homolog. Anterior pleurite 2 times as large as mesothoracic homolog and about 1/2 as large as posterior pleurite, with two minute (PL23, 24) and 1 short setae (PL25). Anterior sclerites each with minute seta (ST36) and (ST37). Epistemum similar to preceding homolog. Epimeron with minute (ST39) and short seta (ST40). Precoxite, lateral stemites, and metastemite similar to preceding homologs. Aspérités. Tubercles widely distributed in membranous areas of thorax, especially laterally; denticles restricted to anterior stemum of meso- and metathorax, and a few anterior to metanotum. 14 Leg. Metathoracic Leg (Fïg. 3B-C). Coxa with 19 setae (COl-19) and 3 pores (COa-c). Setal lengths: minute (COl-10,12,13), short (COM, 16-19), medium (CDU, 15). Areas of granular cuticle present. Trochanter with 5 setae (TRl-3, [4 absent], 5, 6) and 7 pores (TRa-g); setal lengths: minute (TRl, 2, 6) short (TR3, 5). Femur with 8 setae (FEl-8) and 2 pores (FEa, b); setal lengths: minute (FEl, 4-7), short (FE2, 3), medium (FES). Sclerotized cuticle reduced proximally, isolating FEl on membrane. Tibia with 16 minute setae (TIl-16) and 2 pores (TIa, b). Tarsungulus with 2 minute setae (TAl, 2). Abdomen. Segment I. Tergum (Figs. 6A, 7A) with 19 setae (TEl, 3,4-11, 13-21), 5 pores (TEb, e, f, g, j), and 14 tergites on each side. Setal lengths: minute (TEl, 14,15), short (TE8,10,17), medium (TE3,4, 6,11), long (TE5, 9, 16, 19). Tergites: TEl on minute sclerite. TE3 on lateral margin of narrow elongate sclerite with egg burster. TE8- 11 sharing large quadrate central sclerite. TE6, 7 sharing oval midsized sclerite. TE4 on midsized sclerite. TE14-16 on midsized sclerite. TE17 on small sclerite. TE5 on midsized sclerite. TE19,20 sharing midsized sclerite. TE13, 18, and 21 each on minute sclerites. Two midsized tergites without setae posterior to spiracle. Three pairs intertergites. Fleuron (Fig. 19) with 6 setae (PL23-28) and 4 pleurites. Setal lengths: minute (PL23, 26), short (PL24), medium (PL28), long (PL25, 27). Pleurites: PL23-25 on anterior pleurite. PL26, 27 sharing midsized pleurite. PL28 on midsized pleurite. Single midsized pleurite lacking setae posterioventral to PL28. Stemum (Fig. 6B) with 17 setae (ST29-36, 38-46) and 13 stemites on each side. Setal lengths: minute (ST29, 34-36, 39,42), short (ST31, 32, 44,45), medium (ST30, 33, 38,41), long (40, 43,46). Stemites: ST29 on minute sclerite. ST30 on midsized sclerite. ST31, 32 sharing midsized sclerite. ST33 on small sclerite. ST34 and ST35 each on minute sclerite. ST36 on small sclerite. ST38-40 sharing large sclerite. ST41 and ST43 each on midsized sclerites. ST42 on minute sclerite. ST44, 45 sharing midsized sclerite. ST46 on large medial stemite, midsized interstemite without setae anterior to sclerite with ST35. Prolegs poorly developed. 15 Aspérités. Distribution of tubercles as in thorax; denticles restricted dorsally to area behind ampulla, ventrally in front of prolegs, and a few on prolegs. Segments II-VIII. Like segment I but without egg bursters. SegmentIX. Tergum (Figs. 7B, 8A) with 9 setae (TEl, 3, 5, 9,16-19,21), 2 pores (TEf, g), and 10 tergites on each side. Setal lengths: minute (TEl, 3,18), short (TE17, 21), medium (TE9,16), long (TES, 19). Tergites: single pair quadrate midsized anterior tergites. TEl and TE3 each on minute sclerite. TEf, g each on minute sclerites. TE9 and 18 each on small sclerite. TE19 on midsized sclerite; TES and 17 and TE16 and 21 each sharing a midsized sclerite. Pleuron (Fig. 7B, 8A) with S setae (PL24-28). Setal lengths: short (PL24,28), medium (PL26), long (PL2S, 27). Pleurites: PL28 on small sclerite, PL24 and 2S and PL26 and 27 each sharing a midsized sclerite. Stemum (Figs. 20, 22) with S setae (ST30, 36, 38,40,46). Setal lengths: minute (ST30, 36), short (ST38), long (ST40,46). Stemites: Single pair midsized interstemites, ST30 and ST36 each on minute sclerites. Midsized sclerite without setae posterior to ST36. ST46 on midsized stemite. ST38,40 sharing midsized sclerite. Aspérités. Tubercles widely distributed in membranous areas of abdomen. Urogomphi (Figs. 7B, 8A-B). Each with 13 setae (UGl-13) and S pores (UGa-e). Setal lengths: minute (UGl, 4, 11), short (UG2,3), medium (UGS), long (ÜG6, 8-10,12, 13). Sclerites: UGl and 2 and UG3 and 4 each sharing midsized sclerite; UGS-7 sharing midsized sclerite. Pygopod (Figs. 7B, 8B). With 7 setae (PPl-7) and 1 pore (PPa) on each side. Setal lengths: minute (PPl, 6), short (PP2, 3), medium (PP7), long (PP2,4). Sclerites: PPl-3 sharing midsized sclerite; PP4 and PPS each on small sclerite; PP6 and PP7 occupying large sclerite; pygopod (including apical membrane) without aspérités. Second Instar (Figs. 9-10). Head capsule width: 0.77 ± .04mm (« = 4). Morphologically nearly identical to first instar. Differences include lack of egg bursters. 16 lack of ecdysial lines on head, and palpi and antennae more elongate in second instar. Some setae appearing shorter in second instar because they have not increased proportionally to growth of body. Mesostemite with additional area of smooth cuticle (Fig. 9B) lacking in first instar.

Discussion The study of beetle larval chaetotaxy is in its infancy. Only four major works on setal and pore nomenclature of larval Coleoptera have appeared recently (Bousquet & Goulet 1984, Ashe & Watrous 1984, Alaire 1990, Wheeler 1990). I chose not to follow exactly any of the prescribed methodologies and setal nomenclatural systems because I disagreed with certain aspects in each of these works. I agree with Ashe & Watrous (1984) in the importance of using serial homology, but I differ in my choice of instar for study as well as my choice of a starting point for serial homology. They base their setal terminology on the third instar because it has the most complete setal pattern. This completeness is because of the presence of subprimary and secondary setae (see Wheeler 1990). Although it may be desirable to name both the primary and secondary setae of later instars, using primary setae alone facilitates determination of serial homology on different segments. Primary setae may also approximate the ancestral pattern of setal distribution in a given taxon. For these reasons, I base my setal terminology on the jBrst instar. Ashe & Watrous (1984) selected the pronotum as the “base” for determining serial homology because it has a complete set of setae. I regard this completeness as a specialization. In other taxa (e.g.. Lepidoptera) the prothorax contains additional setae not found on other segments (XDl, 2) (Stehr 1987). I selected the first abdominal segment as a starting point for two reasons. First, I agree with Belkin (1952) and consider the abdomen to be the least modified tagma (despite modifications for locomotion in larval Histeridae) with a relatively unmodified pattern of setal distribution. I consider the setal 17 distribution patterns of the thorax and modified abdominal segments IX and X to be independently derived fi"om those found in the abdomen. Accordingly, selection of a generalized abdominal segment (I-Vm) as a “base” affords the best opportunity for determining serially homologous setae both posteriorly and anteriorly. Secondly, abdominal segments of histerid larvae bear the most setae. The prothorax (excluding the prestemum) of Onthophilus is missing 18 setae that are present on the first abdominal segment. Bousquet & Goulet (1984) made no attempt to serially homologize setae. Therefore, I chose not to follow their methodology exactly; however, I found their setal and pore nomenclatural system to be very useful. My procedure differed fi*om Bousquet & Goulet’s (1984) in that I consecutively numbered setae on a single tagma rather than renumbering each setal group independently. An obvious advantage of independent numbering is its ease in accommodation of yet undiscovered setae. A disadvantage is the repetition of setal numbers on a segment (e.g ., TE2, PL2, ST2 might all be found on a single segment). My decision to use a single setal number per segment simplified the terminology because I did not have to subdivide the tergal sternal and pleural regions into arbitrary sections. I acknowledge that taxa may be found bearing setae not included in this work, but I feel I exanuned enough genera to minimize the risk of this occurrence. Any “new” setae are likely to be specializations. I have found examples of “extra” setae in some of the saprinine and tryponaeine genera. A strong justification for an independent nomenclatural system for histerid larvae is that many setae occur here which have no apparent homologs in staphylinid or carabid larvae. However, it is interesting that several homologous setae were detected in these groups (Table 2). 18

Table 2. Examples of setal and pore homologs found in carabid, staphylinid, and histerid larvae.

Nomenclatural convention Bousquet & Ashe& Kovarik Goulet Watrous (Chpater 1 (1984) (1984) BODY REGION Head PA7 Ed3 PA14 PA9 Ell PA21 PAA Ec3 PAX

Abdomen TE9 P4 TE19 TEll P2 PE9

I believe that an ancestral distribution pattern of mechanoreceptors can be established for Coleoptera. This ground plan will undoubtedly be a useful tool in phylogenetic studies of . Figure 1. Head capsule of Onthophilus nodatus, first instar (fiexible setae omitted). (A) Dorsal view. (B). Ventral view. Line scale 0.2 mm.

19 20

vil

•n

>28

Figure 1. Figure 2. Mouthparts and antennae of Onthophilus nodatus, first instar. (A) Right antenna, ventral view. (B) Right antenna, dorsal view. (C) Detail, apex antennal segment 3, dorsal view. (D) Left mandible, dorsal view. (E) Right maxilla, dorsal view. (F) Right maxilla, ventral view. (G) Prementum, ventral view. (H) Prementum, ventral view. (Shading indicates membrane). Line scale, 0,1 mm.

21 22

Ob

oe oa

oc ►PE

oa

SO

*c gMXt

•a

H

Figure 2. Figure 3. Frontale and metathoracic leg of Onthophilus nodatus, first instar. (A) Detail of fi'ontale. (B) Metathoracic leg, anterolateral view. (C) Metathoracic leg, posterolateral view. Line scale, 0.1 mm.

23 24

SF gFR oa

«0 A •c

3% ^ CO Tl FE ao

B

FE K

CO c

Figure 3 Figure 4. Head capsule and pro- and mesothorax of Onthophilus nodatus, first instar. (A) Head capsule, lateral view. (Flexible setae omitted). (B) Pro- and mesothorax, lateral view. Line scale, 0.2 mm.

25 26

•m •o

J^o l2®

•m ;>21

Figure 4. Figure 5. Pro- and mesothorax o î Onthophilus nodatus, first instar. (A) Dorsal view. (B) Ventral view. Line scale, 0.2 mm.

27 28

l^a-

ee

og

10?

|PR7

24' Pft( l i p a A#

146

Figure 5. Figure 6. Metathorax and first abdominal segment of Onthophilus nodatus, first instar. (A) Dorsal view. (B) Ventral view. Line scale, 0.2 mm.

29 30

'' 1

5

mm #

Mh. r4=»^ ; |//

I

Figure 6. Figure 7. Onthophilus nodatus, first instar. (A) Metathorax and first abdominal segment, lateral view. (B) Abdominal segments IX and 7^ lateral view. Line scale, 0.2 mm.

31 32

m 39/,

Figure 7. Figure 8. Onthophilus nodatus, first instar. (A) Abdominal segments IX and X, dorsal view. (B) Abdominal segments IX and X, ventral view. Line scale, 0.2 mm.

33 34

Figure 8. Figure 9. Onthophilus nodatus, second instar. (A) Thorax, dorsal view. (B) Thorax, ventral view. (C) Head and thorax, lateral view. (D) Head, dorsal wew. (E) Head, ventral view. Shading indicates sclerotized cuticle; darkest shading indicates darkly pigmented sclerotized cuticle; middle tone indicates smooth sclerotized cuticle; lightest shading indicates granular cuticle; mottled shading indicates mosaic patterning. Line scale, 0.5 mm.

35 36

I J

Figure 9. Figure 10. Abdomen of Onthophilus nodaius, second instar. (A) Abdominal segment H, dorsal view. (B) Abdominal segment H, ventral view. (C) Abdominal segment H, lateral view. (D) Abdominal segment IX and X, lateral. (E) Abdominal segments IX and X, dorsal view. (F) Abdominal segments IX and X, ventral view. Shading as described for Fig. 9. Line scale, 0.5 mm.

37 38

S

4Q " ÔÔ

Figure 10. CHAPTER n

PUPAL CHAETOTAXY OF HISTERD)AE (COLEOPTERA; HYDROPEHjOmEA) w r r a A DESCRIPTION OF THE PUPA OF ONTHOPHILUS KIRNIROSS

Introduction Pupal morphology of histerid beetles has been all but neglected. Pupal descriptions are available for only ten of the 300+ world genera (Newton, pers, comm.). Several of the available pupal descriptions are insufficiently detailed to be useful in a comparative study. In none of these works was any attempt made to identify setae or setal groups. The duration of the pupal stage in the Histeridae is relatively short (ca. 13 days for those species for which the developmental rate has been determined) (Linder 1967, Summerlin e/a/. 1981, Morgan, etal. 1983, Summerlin e/ar/. 1984, Summerlin and Fincher 1987, Summerlin and Fincher 1988, Summerlin etal. 1989). In the larval stage, setae function as a tactile network which enables a sightless histerid larva to interact vrith a complex environment. This has resulted in a relatively stable pattern of setal distribution (see Chapter 1). Histerid pupae are always concealed in a cell fashioned in the developmental substrate, which may or may not be reinforced with a proteinaceous cement. Here, the need for a sensory network is likely diminished. This may account for the greater plasticity in the pattern of setal distribution among histerid pupae found in this study. The pupal body form is essentially that of the adult beetle, although structures are not always sharply demarcated. Therefore, the same terms used to describe the morphology of adult beetles are applied to the pupa. Most pupal setae can be easily

39 40 homologized among different subfamilies. At lower taxonomic levels it is possible to determine setal homology with greater precision due to stable sub-patterns of setal distribution, A description of every sub-pattern is beyond the scope of this work. This study is designed to identify features peculiar to the pupal stage of histerid beetles and also to provide a working template for describing histerid pupae.

Materials and Methods Several second instar larvae of Onthophilus kimi Ross were collected from a Geomys sp. (Rodentia: Geomyidae) nest/dung chamber on April 4,1992 near Mneola, Wood County, Texas. These larvae were reared to the pupal stage in the laboratory. Voucher specimens are deposited in the larval collections of the Field Museum of Natural History, Chicago, and the author. Descriptions of pupae are based on three specimens. The specimens were studied at SO X with a Wild MS dissecting microscope. The description and setal notation are based on the study of pupae of O. kimi. Pupae of O. nodatus LeConte were also examined for any significant variations in morphology or setal distribution within the genus. Pupal exemplars fi*om Hydrophilidae and all recognized histerid subfamilies (except Niponiinae, Trypeticinae, and Hetaerinae) (Table 3) were examined to ensure that the setal nomenclature was applicable to other related taxa.

Results Terminology and Morphology. Setal homologies were determined by the relative position occupied on the body of the pupa. Pupal setae in Histeridae, although apparently socketed, lack flexibility at their bases. The terminology used in this paper is a modification of that of May (1978). Both individual setae and setal groups bear the name of the body region firom where they originate. For example, elytral setae would 41

Table 3. Material Examined.

Subfamily Species Agyritidae Necrophilus hydrophiloides Guérin-Méneviile Hydrophilidae Helophorus orientalis Motschulsky Histeridae Abraeinae Aeletes politus LeConte Plegaderus transversus Say Trypanaeinae Tiypanaeus sp, Saprininae Hypocaccus fratemus Say gofft Ross Xerosaprinus orbiculatus MarseuI Sapritms pensylvcmicus Paykull Euspiloius assimilis Paykull Gnathoncus sp. Dendrophilinae Carcinops pumilio Erichson Xestipyge multistriatum Lewis X. conjunctum Say Dendrophilus sp. Platylomalus aequalis Say Paromalus seeversi Wenzel Onthophilinae Onthophilus nodatus LeConte O. kimi Ross Tribalinae Epierus divisus MarseuI Idolia gibba Lewis Parepierus sp. Histerinae Atholus bimaculatus L. Hister abbreviatus F. Spilodiscusfloridarms Ross Platysoma lecontei MarseuI Baconia sp. Hololeptayucateca MarseuI Phelister panamensis LeConte 42 encompass all setae found on the pupal elytra. Individual setae are coded with two capital letters (e.g., LA for labral setae). A lower case g appears in front of setal groups (e.g., gOC for ocular setal group) and a lower case b appears in front of setal bands. Setae may or may not arise from tubercles. In a few taxa, setae expand slightly rather than taper distally. Setae were grouped into three size categories based on overall length: minute, short and long. Setae were not measured; therefore size classes are imprecise. The pupal integument is uniformly covered with minute asperities which are not resolvable under low magnification. Head Morphology (Fig. 11A-D). All examined non-histerine genera bear a pair of distal protuberances arising from the dorsal surface of the labrum. These are referred to as labral papillae (LP). The pupal eyes, not readily apparent early in development, eventually take on a red hue. The antennal scape in all known saprinine genera bears a pair of protuberances or extensions distally. These are termed scape papillae (SP). All head setae except for those on the labroclypeal region are termed epicranial setae. These are further subdivided into three setal groups: the frontal epicranial (FE), ocular epicranial (OE) and vertical epicranial (VE). There are two pairs of frontal epicranial setae, a medial pair (MF) and a ventral pair (VF). The vertical setae arise medially on the vertex, generally above the eyes. The ocular setae arise laterally on the epicranium, usually close to the mesal margin of the eye. The frontal setae arise from the frons. Those setae arising at or below the level of the anterior articulation of the mandible are termed clypeal (CL) setae. Setae arising from the area of the labrum are termed labral setae (LA). Labral setae are minute. Thoracic Morphology. Prothoracic setae are generally restricted to the notum. The setae are divisible into four major groups: the anterior pronotal (AN), lateral pronotal (LN), posterior pronotal (PN), and interior pronotal (IN). The anterior pronotal group generally consists of three setae. In some taxa these setae border the anterior margin of 43 the pronotum. In other taxa these setae are more centrally located on the pronotal disc. Interior pronotal setae are those which are isolated on the disc of the pronotum away from other prothoracic setal groups. The exact number of notai setae is variable. An average number of setae per group is reported in the description. A conspicuous feature of the pupal mesonotum is a raised central area which corresponds to the adult scutellum. Setae may arise from the pupal scutum (termed scutal [SC] setae) or scutellum (termed scutellar [ST] setae). A functional spiracle is located on the mesopleuron. The metanotum is essentially featureless and generally lacks setae. Setae arising here are termed metanotal (MN). The pupal elytra have a similar configuration to those of the adult. Ridges or costae are present in some taxa. Two rather distinct patterns of elytral setation were encountered. The first type consists of three usually discrete setal bands arranged obliquely on each elytron. Each band consists of numerous, generally short setae arranged linearly. Setal bands on the elytra are coded as bEL. The second condition consists of a group of relatively few setae (less than 10) which were not arranged in bands. This configuration is coded as gEL. The hindwings in some histerid genera bear a protuberance on the dorsal surface of the wing apex; these are termed wing tubercles (WT). Setae occur on the pupal hind wings in certain saprinine genera as well as in Plegaderus Erichson. In saprinines a cluster of minute setae occurs along the midline of the hindwing where it joins the elytra. In Plegaderus, a single seta protrudes in approximately the same area of the hindwing. Setae arising on the hindwing are termed hind wing setae (HW). The pupal legs of certain genera bear a single seta on the dorsal surface of the femora distally. These setae are termed femoral (FE). Abdominal morphology. The pupal abdomen generally resembles the adult abdomen. Functional spiracles are present only on the first four abdominal segments. 44 This phenomenon was first reported by Verhoeff (1918) in staphylinid pupae and is widespread in the staphyliniform beetles (Newton pers. comm.). In most cases the full complement of spiracles is re-established in the adult stage (Newton pers. comm ). Urogomphi with hardened and darkened apices persist on the dorsal surface of the ninth abdominal segment in most histerid pupae. In some taxa, the pupal pygidium (T7) bears a groove along the midline. Setae originating on the pupal tergites are termed tergal (TE). Tergal setae may occur singly or in groups. Setae arising in the area between the tergites and pleurites or lateral-most on the tergites are termed laterotergal (LT). This setal position is generally discernable even in cases where the tergites bear a continuous row of setae. Setae arising on the stemites dorsolaterally slightly below the juncture of the tergites and stemites are termed laterostemal (LS) and any setae arising ventrad to these on the stemites are termed sternal (ST). In saprinine pupae, tergal setae on T7 have apparently shifted to the posterior margin of the tergite. Description of Pupa of Onthophilus kirni. (Fig. 12A-B) All setae tapered. Head. Four long epicranial setae on each side; single mediofrontal seta (MF), 2 in ocular epicranial group (gOE) and single vertical epicranial (VE). Pair of minute labroclypeal setae (LA) on each side, each pair arising directly above prominent labral papilla (LP). Thorax. Pronotum with 10-15 long setae on each side, 3 in anterior pronotal group (gAN), 4-5 in lateral pronotal group (gLN), and 3-7 in posterior pronotal group (gPN). Mesonotum with single long scutal seta (SC) on each side. Metanotum lacking setae. Elytron costate and with 3 long setae (gEL). Hind Wings. Hind wings with tapered wing tubercle (WT) and no setae. Abdomen. Tergites 1-6 each with single long tergal setae (TE) on each side. Laterotergal seta (LT) on single specimen only, absent in others; located on segments 2-6. Laterotergal setae on segments 2-4 mostly short; those on segments 5, 6 long. Single laterostemal seta (LS) on abdominal segments 2-7; 45 laterostemal setae on segments 2-6 long, on segment 7 short. Pygidium lon^tudinally ridged and lacking setae. Urogomphi well-developed.

Discussion This study establishes a nomenclature to describe chaetotaxy and morphological peculiarities of histerid pupae. It also provides a means to homologize setae and setal groups among histerid genera. The pattern of setal distribution in histerid pupae varies considerably among genera. Although the pattern of setal distribution appears to be relatively stable within a species, a fair amount of interspecific variability was also detected, especially on the pronotum. For example, the pupa of Onthophilus nodatus LeConte differs fi’om O. kimi in having up to six lateral pronotal setae on each side, an additional scutal seta, and single scutellar seta on each side of the mesonotum. The single specimen available for study also lacked laterotergal setae. Therefore, caution must be used in generalizing character information above the species level, especially when using the exemplar approach. Setal distribution was found to be more stable in histerid larvae (see Chapter 1). Figure 11. Head-on views of head and prothorax of histerid pupae. (A) Hister abbreviatus. (B) Xestipyge conjunctum (C) Epierus divisus (D) Geomysaprinus goffl. Line scale, 0.5 mm.

46 47

i-i- U

•g06

SP' iC l,

Figure 11. Figure 12. Setal maps of Onthophilus kimi pupa. (A) Dorsal and lateral view composite (B) Lateral view. Line scale, 1.0 mm.

48 49

SAN

,SLN qPN ‘goe / ^ 11 LA

LSI

Figure 12, CHAPTER m

DEVELOPMENT OF EPIERUS DIVISUS MARSEUL (COLEOPTERA: HISTERTOAE).

Introduction Little is known about the biology of the histerid genus Epierus. The 52 described species occur mainly in the Neotropics, with a few species in the Nearctic, Oriental, Palearctic, and Australasian zoogeographic regions. Adult Epierus are usually found beneath bark and appear to be specialists on fungal spores (Moser 1971, unpublished data). Adult Epierus are apparently long-lived as evidenced by some species surviving more than a year in laboratory culture (unpublished data). Larval Epierus have been collected under loose bark of dead trees (including Quercus) where a layer of moist organic debris has accumulated. The larvae likely burrow through this material in search of food. Larvae of one species, Epieruspulicarius Erichson, have been extracted from accumulated organic debris within tree and stump cavities. Epierus larvae have been collected in the field from mid-June to late August. The developmental rates of only a few species of histerids are known. These taxa include several histerine genera; Hister (Linder 1967, Summerlin etal. 1981, Summerlin etal. 1984, Summerlin and Fincher 1 9 8 8 ) , (Linder 1967), (Bomemissza 1968), Atholus (Summerlin and Fincher 1987), and Pleasius (Jepson 1914); three saprinine genera: Saprinus (Linder 1967, Mochizuki 1985), Gnathoncus (Linder 1967), dXiàEuspilotus (Summerlin etal. 1989); a single dendrophiline genus, Carcinops (Morgan et al. 1983) and a single abraeine genus, Teretriosoma (Rees 1985). Reporting of developmental data was generally inconsistent. In some cases the developmental rates 50 51 of the two larval instars were pooled while in other cases they were reported separately. Some authors recognized and included the developmental times for pre-pupae while others did not, indicating a need for a standard methodology in order to make more precise comparisons of developmental rates among histerid taxa. An objective of this study was to compare the duration of each developmental stage of the subcortical jÿ/en/.s divisus to a histerid associating with an ephemeral system to see how they differed. I selected the dung/carrion-associated species Euspiloius assimilis (Paykull) to compare vAi)! Epierus divisus. The development of this species had been determined by Summerlin et al. (1989), but these authors failed to delimit prepupal development. To remedy this, I studied the development of Euspiloius assimilis and included duration of the pre-pupa.

Materials and Methods Adult parental stock of Epierus divisus was obtained from beneath the bark of a fallen tree in CaAon La Libertad, Tamaulipas, Mexico. Parental stock of Euspiloius assimilis was collected from Latimer Co., Oklahoma. Voucher specimens were deposited in the larval collections of The Ohio State University; the Field Museum of Natural History, Chicago; and the collection of PWK. Laboratory cultures o îEpierus divisus were started with adults confined in clear plastic rearing containers (6.6 x 9.0 cm). The bottom of each container was covered with a layer of moist sand approximately 2 mm deep, to which was added a 4 mm layer of sifred organic debris from an oak tree cavity. Blotting paper was placed on top of the organic debris, and oak bark was placed on top of the blotting paper. Both the sand and the blotting paper were kept moist (but not saturated) at all times and the containers were kept in darkness. Adults were provided with hoXh Musca eggs and early instar Sarcophaga larvae from laboratory cultures. All rearing took place at about 25° C. Rearing containers were inspected daily for the 52 presence of eggs. Eggs deposited in the moist sand were clearly visible through the bottom of the container and were transferred to petri dishes containing moist filter paper and inspected daily for eclosion. Initial attempts to maintmn larvae in petri dishes with moist filter paper fmled because larvae were trapped and drowned in water condensation. An alternate method of rearing was developed using plaster-filled watch glasses provided with a central depression 2.5 cm wide x 0.8 cm deep. A thin layer of sifted organic debris was added to the bottom of the depression and covered by a circle of filter paper cut to a diameter slightly less than that of the depression in the plaster. Larvae were placed in the depression and confined there by covering the plaster with a 5.5 cm circle of filter paper and a 4 cm diameter circular cardboard pillbox cover centered above the depression. The cover was kept tightly appressed to the watch glass with rubber bands. Relatively high humidity was maintained in these cages by adding water to the plaster and placing the rearing cage within a sealed Ziploc* bag with moist paper toweling. Larvae were provided with hoXhMusca eggs and early instar Sarcophaga larvae and examined daily for eclosion. Pre-pupae were transferred from the rearing cages to small petri dishes partly filled with slightly dampened sifted organic debris. To prevent escape of the pre-pupa, the lid of the petri dish was held in place with rubber bands. The container was placed within a sealed Ziploc® bag with moist paper toweling and kept in darkness. Occasionally, the pre­ pupa would construct its pupal cell against the bottom or side of the dish. This would permit observation of pre-pupa with minimal disturbance. Laboratory culture oî Euspiloius assimilis followed the methodology of Summerlin (1989). Egg and larval isolation and maintenance was similar to that used for Epierus divisus. The only major difference in larval rearing procedures was the substitution of moist sand for sifted organic material in the plaster rearing containers. 53 Histerids develop through 5 discrete stages: egg, first instar, second instar, pre-pupa, pupa, and adult. An improved resolution of the developmental history of a species can be achieved if each stage is measured; if a standardized method of rearing and data recording are adopted, direct comparison of developmental rates among species would be possible. The starting and stopping point of the egg, larval, and pupal stages are obvious. Each stage begins and ends with ecdysis with the exception of the egg stage, which begins with deposition and ends with eclosion of the first instar. The beginning of the prepupal stage is perhaps not as obvious, but this is an important and distinct part of development; the prepupal stage marks the transition of an actively feeding larva to one that stops feeding and begins constructing a pupal cell or cocoon. As noted by Linder (1967) this stage is marked by a color change. The feeding larva is translucent yellow with the gut easily visible beneath the integument. The pre-pupa is opaque white with the gut no longer visible. A completely white larva signals the end of larval development and the commencement of prepupal development. Paired samples t-tests were performed with Systat® to compare sample means of developmental rates of Epierus divisus versus Euspiloius assimilis.

Results Egg, larval, prepupal, and pupal development Eggs oî Epierus divisus were ovoid and pale cinnamon. Length of 14 eggs was 1.24 ± 0.04 mm (X ± SD) and width 0.44 ± 0.03 mm. The surface of the chorion was dull, apparently due to microscopic texture. Unlike the membranous chorion found in most histerid eggs, the chorion of Epierus divisus is thick and partially retains its shape after larval eclosion. Just before eclosion fi-om the egg, the mandibles, terminal palpal segments, and setae darken and can be clearly seen through the chorion. Most sclerotized larval structures harden and darken soon after emergence fi-om the egg. 54 The duration of the developmental stages of Epierus divisus and Euspiloius assimilis are summarized in Tables 4 and 5 respectively. Results of a t-test are summarized in Table 6, The egg stage oîEpierus divisus lasted 10.75 ± 0.71 days (n=14) and was significantly longer than that of Euspiloius assimilis (t = 26.46; dfi= 17; P = 0.00).

Table 4. Duration of developmental stages ïor Epierus divisus under laboratory conditions.

Duration Life stage n X(days) Range SD Egg 8 10.75 10-12 0.71 Instar 1 6 10.50 8-15 2.74 Instar 2 6 6.33 6-8 0.82 Pre-pupa 5 7.83 7-8 0.41 Pupa 4 10.50 10-12 1.00

Table 5. Duration of developmental stages for Euspiloius assimilis under laboratory conditions.

Duration Life stage n X(days) Range SD Egg 11 2.09 1-3 0.70 Instar 1 13 3.15 2-4 0.89 Instar 2 13 3.69 2-4 0.63 Pre-pupa 9 5.11 4-6 0.78 Pupa 7 7.57 7-8 0.53

Table 6. Independent t-test comparing duration of developmental stages of Epierus divisus versus Euspiloius assimilis and under laboratory conditions.

Life stage T DF P Egg 26.46 17 0.00 Instar 1 8.93 17 0.00 Instar 2 7.75 17 0.00 Pre-pupa 7.00 12 0.00 Pupa 6.46 9 0.00 55 First instar ^/en/jr divisus were capable of killing and feeding on soft-bodied , including conspeciftcs, soon after eclosion. First instar larvae completed development in 8-15 days. The development of first instar J^/e/t/5r divisus was significantly longer than that of Euspiloius assimilis (t = 8.93; df= 17; P = 0.00). Before molting, first msXzx Epierus divisus larvae sought a sheltered place and became quiescent until the onset of the molting process. Second instar larvae required 6-8 days to complete development. Development of second instar i^/en/5 divisus was significantly longer than that 0Î Euspiloius assimilis (t = 7.75; df= 17; P = 0.00). Pre-pupae oîEpierus divisus constructed ovoid cells in the organic substrate provided for them. Little or no structural protein was used by Epierus divisus pre-pupae during pupal cell construction as evidenced by the fragile walls of pupal cells. Before pupation, pre-pupae became quiescent and assumed a “C”-shape. The prepupal stage required 7 to 8 days. Prepupal development oîEpierus divisus was significantly longer than that oî Euspiloius assimilis (t = 7.00; df= 12; P = 0.00). Newly eclosed pupae were entirely transparent. As development proceeds, pupae gradually became an opaque white with red eyes. Toward the end of pupal development, the teneral adult was clearly visible through the pupal integument. Before eclosion, the adult had already begun the process of hardening and darkening. Eventually, the movements of the head and legs freed the adult from the membranous pupal integument. The duration of the pupal stage for this species was 10-12 days. Pupal development of Epierus divisus was significantly longer than that of Euspiloius assimilis (t = 6.46; df= 9; P = 0.00). Larval Feeding Behavior. The following observations on larval feeding behavior apply not only to Epierus but to other members of the family reared by the author. Laboratory-reared Sarcophaginal larva were provided as prey. When histerid larvae contacted a prey item with their palpi, they lunged forward to seize the prey with their 56 mandibles. If successful, larvae moved their mandibles alternately back and forth until the prey’s cuticle is pierced. Shortly after piercing the integument, larvae would often release their prey and proceed to hunt for other items. The bitten and released maggots thrashed about and attempted to crawl away, but even those that appeared to have only a minor injury had great diflBculty moving and were soon still, possibly indicating the use of venom. Several prey were generally wounded or killed before feeding commenced. The mobile nature of maggots and other insect larvae preyed upon by histerid larvae may have resulted in this predatory behavior. Histerid larvae are liquid feeders. Digestion is extraoral and histerid larvae are equipped with a pharyngeal pump. During feeding, the anterior gut, clearly visible through the larval integument, expanded and contracted in rapid succession as the liquid contents of the prey were ingested. All larval mouthparts as well as the antennae were used to manipulate a prey item for fluid extraction. Discussion The lack of a standardized approach in presenting developmental data and, to a lesser degree, significant differences in rearing temperatures, hampered comparisons of developmental rates of Epierus with those of other taxa. It should be noted that the histerids for which data are available develop in warm weather in association with ephemeral systems (i.e. dung or carrion) and feed as adults and larvae primarily on eggs and larvae of flies that develop on these substrates. This is probably the most common trophic association for members of this family (Baulduf 1935, Reichardt 1941, Hinton 1945, Wenzel 1962, Vienna 1980). Epierus divisus is the first histerid studied whose life cycle is not tied to a highly ephemeral resource. The average egg development oîEpierus divisus (ca. 11 days) was considerably longer than that oîEu^iîotus assimilis (ca. 2 days) and all other histerids previously studied. The maximum reported duration of embryogenesis for both histerine and 57 saprinine histerids studied was 6 days at 20-25® C (Linder 1967). The average duration of the egg stage for C pumilio (Erichson), the only dendrophiline studied, was 6.3 days at 25® C. (Morgan et al. 1983). The average developmental rate oîEpierus divisus first instars (ca. 10 days) was also much longer than that oî Euspiloius assimilis (ca. 3 days). First instar development of some histerine species took at most 7 days to complete at 20- 25® C (Linder 1967). The average developmental rate oîEpierus divisus second instars was ca. 7 days. This was nearly twice as long as that of Euspiloius assimilis. Unfortunately, most developmental rates determined for the second instar do not distinguish between the feeding second instar from the pre-pupa. Exceptions include Hisier unicolor L. and Margarinoius siriola (Sahlberg). The development of second instar larvae for these species took respectively 6 and 9-10 days to complete at 20-25® C (Linder 1967). Prepupal development took an average of ca. 8 days îov Epierus divisus and an average of ca. 5 days îot Euspiloius assimilis. Pre-pupae oîH. unicolor developed in 2 days while those ofM. siriola took 5-6 days 20-25® C (Linder 1967). The average pupal development for Epierus divisus took ca. 10 days. This is similar to pupal development of Euspiloius assimilis, which took ca. 8 days to complete. Pupal development of some histerine species was lengthy. Pupal development o î Hisier nomas Erichson averaged 17 days at 25-28® C (Summerlin and Fincher 1988) while that

0ÎMargarinoius carbonarius (Hofiman) took 17-18 days to complete at 20-25® C (Linder 1967). The species with the longest known pupal duration is Pactolinus chinensis (Quensel). Pupae of this species required 3-5 weeks to complete development at 25- 30® C (Bomemissza 1968). Differences in the developmental rate of histerid larvae associated with ephemeral systems versus those that are not may reflect prey availability. Histerids associated with ephemeral systems feed as both adults and larvae on soft-bodied insects. At carrion, histerids prey mainly on fly larvae, particularly larval blow flies (Bomemissza 1957, Jirôn 58 and Cartin 1981, Braack 1987). At dung, histerids also are reported to feed on fly larvae, particularly larval muscids (Hafez 1939, Hammer 1941, Mohr 1943), and also on scarab larvae (Hafez 1939, Mohr 1943). At optimal temperature and moisture levels, dung and carrion will foster development of large numbers of fly larvae. A single jfresh cattle dropping can contain well over a hundred fly larvae (Mohr 1943). This translates into an abundant and concentrated source of prey for predators. However, the rapid degradation of non- structural components of ephemeral systems promotes rapid development of associated fly larvae. Independent fleld studies on carrion decomposition by Bomemissza (1957) and Braack (1987) found that blow fly larvae colonize carcasses almost immediately and begin declining after about 14 days. In the field, muscid and sarcophagid flies developing on bovine dung colonize this substrate almost immediately upon deposition and complete their development in about 6 days. Scarab larvae first appear in 3-day-old bovine dung and complete their development after about 16 days. The arrival of adult histerids at ephemeral systems tends to lag behind prey colonization by several days (Hammmer 1941, Kessler and Balsbaugh, 1972, Braack 1987), reducing the window for foraging by larval histerids but increasing the likelihood of prey being plentiful. Maggots may also escape predation because they rapidly become too large for predators to subdue (Valiela 1974). At carrion, allowing for a 2-day lag time for colonization and 2 days for egg development, larval Euspiloius assimilis preying on blow fly larvae would have about 10 days before their prey became scarce. The foraging window for predation of muscid and/or sarcophagid larvae developing on bovine dung would be approximately 2 days. If scarab larvae are included as potential prey items at bovine dung, the foraging window would extend to ca. 12 days. Both of these scenarios assume immediate oviposition by the adult histerids. Under laboratory conditions the foraging of Euspiloius assimilis larvae is completed in about 7 days, well within typical foraging windows afforded to them in the 59 field. Egg and first instar development oîEpierus divisus were considerably longer than that of Euspiloius assimilis, underscoring ecological differences. If development of Epierus divisus is superimposed on foraging windows of ephemeral systems, it becomes apparent that prey would be gone long before the normal period of larval foraging is completed. Many subcortical species of histerids colonize trees in their early stages of decay and feed on flies coloniring decaying phloem and cambium, or bast (unpublished data). According to Hamilton (1978), decaying bast offers a rich and well balanced diet but this material is rapidly consumed; hence, histerids associated with flies developing on bast are apt to be subjected to the same pressures of other ephemeral systems. The developmental substrate oîEpierus, typically accumulated organic material beneath loose bark of trees dead for at least a year, is not nearly as ephemeral as carrion, dung, or bast. In older dead trees, the only remaining primary source of nutriment is cellulose. Relatively few organisms (mainly bacteria, fungi, protists, and termites) are able to digest cellulose. Furthermore, breakdown of wood by these organisms requires ample moisture and generally warm temperatures (Subramanian 1983). During winter in temperate regions, the process of degradation invariably slows, extending the duration of the wood in the field. Hence, decaying trees provide a relatively long-lived and stable habitat for TQ^lophagous and mycophagous insects. Predators of these primary consumers, like larval Epierus divisus, are provided with a less restricted window for development than their counterparts associated with ephemeral resources. Loose bark adhering to wood tends to modify the microclimate beneath it by trapping heat and moisture providing a suitable habitat for growth and sporulation of fungi on which adult Epierus specialize. Larval Epierus probably feed on soft-bodied insects that also occur beneath bark. Specific prey items of larval Epierus have not been identified; however, I have examined extracted firom microhabitats supporting these larvae. Potential prey items include larval 60 Coleoptera, termites, ant brood, Collembola, and mites, Diptera larvae were rarely encountered. If prey is scarcer under bark than ephemeral systems, which is apparently the case, a lengthened larval development may be advantageous. The relative temporal stability of a rotten log would allow a longer period for egg development. Ecological information on histerid beetles is rather scare. Histerids seem to be preadapted for exploiting ephemeral systems; there are only two larval instars and their large eggs produce precocious larvae capable of capturing prey shortly after eclosion. Developmental data are unknown for histerids inhabiting tree holes or those which occur in association with flies developing on rotting bast. Data are also lacking ftom species inhabiting the nests of rodents, ants, or termites. CHAPTER IV A REVISION OF THE GENERA OF ONTHOPHILINAE AND TRIBALINAE (COLEOPTERA: HISTERIDAE).

Introduction Histerid beetles belong to the staphylinlform lineage. Until recently, they were placed in the superfamily Histeroidea along wdth the related families Synteliidae and Sphaeritidae. However, recent work on the higher classification of the Coleoptera by Lawrence and Newton (1982) placed histerids in the superfamily Hydrophiloidea which includes the families Hydrophilidae, Synteliidae, and Sphaeritidae. Except for Helava’s (1985) revision of the Hetaeriinae, suprageneric revisions of histerid beetles are essentially nonexistent. Members of this family have been assumed to be predators both as adults and larvae. Wenzel (pers. comm.) was aware of the presence of fungal spores in the gut of adult Epierus. This observation, published as a personal communication in a paper by Moser et al. (1971), has largely gone unnoticed. An examination of the mouthpart structure and gut contents of examples of all onthophUine genera has revealed most to be microphagous and most mycophagous. Adult onthophiline and tribaline histerids are long-lived. I have maintained specimens from several genera in captivity for more than a year. While developmental data are unavailable for most onthophilines, the developmental period fi-om egg to adult in Epierus divisus Marseul took ca. 48 days (see Chapter 3).

61 62

Before Wenzel’s (1944) classification of histerids, the Tribalinae were regarded by Bickhardt (1916, 1917) to constitute a tribe of the , Bickhardt’s Tribalini included the genera ^/en/s, Parepierus, Stictostix, Tribalister, Triballodes, Idolia, Sphaericosoma, and Caerostemus. The genera Onthophilus, Epiechinus, Gïymma, and Peplogfyptus were considered to belong to the Abraeinae primarily on the basis of a single character involving antennal insertion in a fovea between the eyes and next to the medial edge of the eyes. Wenzel (1944) redefined the Abraeinae, removing the aforementioned genera without placing them elsewhere, indicating in a footnote that “There is reason to suspect that these genera may belong to the Tribalinae”. In this work, he also elevated the Tribalini to subfamilial rank. Wenzel (1944) established two large divisions; Saprinomorphae (including the Abraeinae) and Histeromorphae, which respectively lack or have a prosternai lobe. He was the first to mention and ascribe importance to the modifications of the prothorax for antennal protection. He described expansions extending laterally in fi-ont of the prosternai carinae which he termed alae. The alae form a ventral shield covering antennal cavities situated in the anterior prothoracic angles. Wenzel (1962) included the presence of labral setae as a key character for the Tribalinae. He placed the genus Onthophilus in the Tribalini and moved Tribalister (which lacks labral setae) to the Histerinae. Mazur (1972) placed Glymma candezei Marseul in the Onthophilini of the Tribalinae. Mazur (1973), included in the Onthophilini the genera Onthophilus, Epiechinus, Glymma, Vuattouxirms, Peploglyptus, and Sculptura. Vienna (1974) elevated the tribe Onthophilini to the rank of subfamily. This change in status was justified by the presence of strong carinae on the elytra as well as contrasting differences in aedeagus morphology. He noted that the aedeagus of abraeines lacks an 63 articulated phallobase which is present in Onthophilus and Epiechinus. Vienna (1974) also excluded the genus Vuattouxinus from the Onthophilini because of the apparent lack of an articulated phallobase. Mazur (1984) recognized two subfamilies, the Onthophilinae and Tribalinae, in his world catalogue ofHsteridae. In the Onthophilinae he included the genera Onthophilus, Epiechinus, Sculptura, Sigillum, Vuattouxinus, Peploglyptus, and Glymma. In the Tribalinae he included the Epierus (including the subgenus ), Pseudepierus, Stictostix, Parepierus, Scaphidister, Trihalus, Tribalasia, Idolia, Sphaericosoma, Caerostemus. He also moved the genus Triballodes from the Tribalinae to the Dendrophilinae. Following Wenzel’s suggestion, Mazur (1988) elevated the subgenus Plagiogramma to the level of genus. This decision was based primarily on the presence of a both a meso-metastemal suture and a marginal mesostemal suture (see Fig. 25C and D) and the typically elongate aedeagal parameres.

Materials and Methods Specimens of nearly all taxa in Table 7 were dissected for the purpose of examining fine structure of the antenna, mouthparts, hind wings, and terminalia. For antennal and mouthpart dissections, specimens were relaxed in near boiling water so that the head could be removed. The head was placed in a vessel of lactic acid and heated on a slide warming tray. Antennae and mouthparts were mounted on standard slides in Hoyers’ medium. Several shards of glass were used to suspend the cover slip above the subject to prevent breakage and distortion from flattening. Preparation of hind wings involved first softening and expanding them in near-boiling soapy water. The wings were then placed in lactic acid for a few minutes and then mounted on standard slides using Hoyers’ medium. Terminalia were cleared in lactic acid, dissected apart and mounted in depression slides 64 Table 7. Material Examined

Family Species Hydrophilidae Helophorus orientalis Motschulsky Syntelüdae Syntelia histeroides Lewis Syntelia westwoodi Sallé Sphaeritidae Spherites politus Mannerheim Ifisteridae Dendrophilus opacus Ross Ampleus marginatus LeConte Spaericosoma ovum Marseul Onthophilus Idmi Ross Onthophilus nodatus LeConte Onthophilus punctatus Müller Vuattouxinus borassicola Thérond Epiechinus planistemus Bickhardt Epiechinus rappi Bickhardt Epiechinus kivuensis (Thérond) Epiechinus leleupi (Thérond) Epiechinus candezii (Marseul) Australepierus australis (Helava & Howden) Australepierus foderatus (Lewis) Australepierus phyllohius (Broun) Australepierus punctulipennis (Broun) Australepierus spinellus (Broun) Plagiogramma gentilis (Horn) Plagiogramma subtropica (Casey) Plagiogramma frontale (Kirsch) Plagiogramma pubifrons (Hinton) Plagiogramma rhinocera (Marseul) Epierus regularis (Beauvois) Epierus pulicarius Erichson Epierus beccari Marseul Epierus sp. Stictostix califomica (Horn) Stictostix frontalis (MacLeay) Stictostix parra (Marseul) 6 5 Table?, (continued)

Family Species Stictostix ectatommae Lea Peploglyptus helfragei LeConte Peploglyptus n. sp. Tribalus ascaphus Marseul Tribalus capensis (Paykuil) Tribalus minimus ^ossi) Tribalus scaphidiformis (Hliger) Eutribalus americanus (LeConte) Eutribalus marseuli (Gomy) Eutribalus opimus (Lewis) Eutribalus agrestis (Marseul) Eutribalus puncticeps (Lewis) Eutribalus koenigius (Marseul) Parepierus amandus (Schmidt) Parepierus salvazai (Desbordes) Parepierus velox (Cooman) Parepierus vitalisi (Desbordes) Paridolia laevidorsis (Lewis) Paridolia sp. Idolia gibba Lewis Idolia laevissima (LeConte) Idolia scitua Lewis 66 with glycerine jelly. The removal of gut contents required initial clearing of the body cavity in warm lactic acid for several days. The gut was then removed and mounted on a standard slide with a relatively small amount of Hoyers’ medium and a large cover slip. It was possible to spread out the gut contents by gently moving the coverslip back and forth before the medium had set. Specimens were studied with a Wild M12 compound microscope fitted with a Bausch and Lomb stereo polarizer. Dissected parts and whole specimens intended for SEM were cleaned by sonication in Windex*. Specimens were then transferred to 100% ETOH and later critical point or air dried. Specimens to be air dried were placed in 100% ETOH and transferred to filter paper. Specimens were mounted on aluminum stubs with double-stick tape, amber coated and studied with a JOEL JSM-820 operating at 20kV.

Terminology and Morphology Seta!, Pore, and Cuticular Terminology: Patterns of setation on the mouthparts, antennae and terminalia were found to vary considerably among taxa. However, on the maxilla and labia predictable patterns of setation were detected among both in- and outgroup members. These are setae are referred to as primary setae (rs) (Fig. 43). All other setae are termed secondary setae. The term seta denotes hair or spinelike socketed structures. The term setula is used to describe seta-like structures without a detectable socket. The term pore generally refers to a minute, circular cuticular landmark of unknown function. However, one pore found on the prosternai keel termed the cctriml pore (cp) (Fig. 13C), may be secretory. These pores, located between the lateral marginal prosternai and carinal striae, were found to be universally present in Tribalinae and Onthophilinae. The pores were also present in Anaplecms but absent in Dendrophilus. The function of these pores is unknown but they may be gland openings, hficrographs of these structures find them often encrusted with material which presumably had emanated 67 from these structures. Areas adjacent to the pores are imbricately textured in some species of Idolia and Epierus (Fig. 36D). This type of texture is found nowhere else on the surface of these beetles and may have evolved to facilitate dissemination of emitted volatiles. In some taxa, the cuticle was shagreened, or densely covered with microscopic tubercles (Fig. 4 IB, D) which diffract light to produce turquoise iridescence. Antennal Morphology: The scape morphology is variable within the Onthophilinae and Tribalinae. In some taxa the scape is slender and elongate and in others short and robust. In some members of the Tribalus genus group, a short, blunt medioapical projection termed scape tooth (to) (Fig. 27C) is present. The dorsal surface of the scape may or may not be expanded and pitted. Armature commonly consists of minute and short setae. Surface texture is generally imbricate. The Oagellum consists of 7 segments, each bearing a whorl of sensilla chaetica. Surface texture of the flagellar segments is also generally imbricate. The compact antennal club is formed from the frision of three flagellomeres segments. In the out Anapleus and Dendrophilus, the club segments (Fig. 27A) are clearly divided by 2 deep sulci. These are termed the distal sulcus (ds) and proximal sulcus (ps). In Anapleus and Dendrophilus the sulci contain a row of sensilla basiconica. In most tribaline taxa these sutures are either absent or evanescent. In the genus Tribalus, there is a deep pit associated with the proximal sulcus termed the sulcal p it (sp) (Fig. 32A). Surface sculpture when present is imbricate. Three of the four types of sensilla present on the club segment of onthophiline and tribaline histerids corresponded to the descriptions and illustrations of sensilla found on the antenna of Ips (Coleoptera: Scolytidae) (Borden 1968). They include sensilla trichodea, sensilla basaconica, and sensilla chaetica. Sensilla basaconica were typically arranged in transverse whorls, distinct plaques, or were irregularly distributed. An additional sensillum type found on the club of 68 most species oîEpiechinus, consists of short nipple-like structures termed sensilla mammillae. Labrum/Epipharynx Morphology: The dorsal labral surface in all onthophilines bears at least 2 pairs of long robust setae. The inner surface of the labrum or epipharynx is divided into a pair of outer lateral areas (la) (Fig. 28B) and a central medial area (ma) (Fig. 28B) by two oblique rows of setulae, the inner brushes (ib) (Fig. 28A) The lateral and dorsal margin of the lateral area are bordered with a fiinge of regularly spaced elongate setae termed lateral fringe (IQ (Fig. 28A). Lateral fringe setae that extend onto the medial area are often peg-like and are termed medial peg setae (mp) (Fig. 28B). The lateral area is uniformly covered with scales which may be denticulate and non­ overlapping or broadly rounded and overlapping. In some taxa, some lateral area scales are elongate and hairlike. When numerous, these elongate scales form an outer brush (ob) (Fig. 28A). Vestiges of an outer brush are found in several onthophiline genera. A dorsomedial brush (db) (Fig. 28A) composed of short stiff setulae generally borders the medial area dorsally. Several robust, often peg-like setae originate directly above the dorsomedial brush and are termed dorsomedial peg setae (dp) (Fig. 28A). A longitudinal ridge usually bisects the medial area. The medial area usually bears numerous sensilla. Mandible Morphology: The upper surface of the mandible is divisible into four parts; a dorsolateral area (dl) (Fig. 29A), a dorsomesal area (dm) (Fig. 29A), the prostheca (pr) (Fig. 29A), and the mola (mo) (Fig. 29A). The surface of the dorsolateral area is commonly textured, setose, and with several robust apical setae that appear to be contact chemoreceptors (cc) (Fig. 20A) (D. Johnston pers. comm.). Multiple pits, depressions, or cavities thought to be mycangia (my) (Fig. 20A) (M. Thomas, pers. comm ), are present on the dorsolateral areas of some tribaline and onthophiline taxa. Internally several pores are visible apically via transmitted light. The apex of the dorsolateral area is commonly bidentate. 69 The surface of the dorsomesal area is smooth. Cuticular cavities are present mainly along the mesal margin of the dorsomesal area. An articulated setose appendage (aa) (Fig. 29 A) arises from the basomesal margin of the mesal area. This structure and the area to which it attaches directly above the mola is hyaline via transmitted light. The prostheca is either broadly attached to the mesal area or apically detached. The dorsal prosthecal setulae vary in length, organization, and structure. The mola may or may not project mesally. The inner molar surface is generally equipped with a series of linear channels which expand slightly beneath the surface. Lawrence (1989) described this structural arrangement as a food press. The upper surface of the mola bears a field of microvilli termed the dorsal microtrichialfield (df) (Fig. 29A). The microtrichia may or may not be arranged in small packets forming distinct patterns. Much of the lower surface of the mandible is quite smooth. The mesal margin basally bears another field of microvilli termed the ventral microtrichialfield (vf) (Fig. 21C). Setulae of the ventral surface of the prostheca vary in length, structure, and organization. In Onthophilus and Vuattouxirms specialized comb setulae (Fig. 2 IB) are present. In the Idolia and Epierus genus groups, a complex file-like structure termed radula (ra) (Figs. 35B-D, 38D) is present on ventral prosthecal surface. One or two small openings termed ventral pores (vp) (Fig. 21 A) (possible gland ducts) may be present on the ventral mandibular surface. Maxilla Morphology: Terminology for the gross morphology of the maxillae follows that of Crampton (1923) and Williams (1938). Two major types of galeal (ga) (Fig. 43B) modification occur in the Onthophilinae and Tribalinae. An “unmodified” type is found in most Tribalini (Fig. 39A, B) and is similar to that found in outgroups Anapleus, Dendrophilus and all predatory forms examined thus far. It consists of an apically flaring, sclerotized base armed mesally and apically with robust, elongate, setulae. 70 In Onthophilus, Vuattouxinus, and mosX Australepierus the dorsal surface of the galea is densely covered with relatively short, stiff tapered setulae (Figs, 23C-D, 39C). In these taxa sclerotization of the galeal base is reduced. The dorsal surface of the galea in the Idoliini, Epierus, most Plagiogramma, some Stictostix and Australepierus is armed with elongate spatulate setulae. The galea oî Epiechinus appears to be somewhat intermediate between the two afore mentioned types. The lacinia (Ic) (Fig. 43B) is usually broadly attached to the mediostipes. In the Idoliini, the lacinia is largely detached from the mediostipes. The lacinial fringe mainly consists of slender tapered setulae, but in a number of taxa distinct flattened and broad setulae termed blade-like setulae also occur near the lacinial apex. An armed outgrowth of the dorsoapical lacinial margin is present in some onthophilines. This unusual structure is termed the strobilus (Fig. 40C). The strobilus is armed along its length with minute setulae and may terminate in a cluster of elongate blade-like setulae. The ventral lacinial disc is distally armed with a variably modified lacinial tooth. Also on the ventral lacinial disc are one or more slender setae termed lacinial discal setae. The mediostipes (me) (Fig. 43 A) commonly bears internal cuticular cavities. The surface of the mediostipes is generally rugose and bears numerous small setae and usually a single pore. The central ventral sclerite or basistipes (bs) (Fig. 43A) commonly bears 2 pores and is armed with 2-3 primary setae (l®s) (Fig. 43A) which are generally more elongate and robust than secondary setae. The basitipes is often textured, sometimes strongly so. The cardo (ca) (Fig. 43 A) bears a dorsal condyle articulated proximally with the basostypes. The cardo armature is variable and the surface is generally smooth. The lateral-most sclerite, the palpifer (pf) (Fig. 43B) is typically strongly textured and always bears an elongate, sometimes robust ventroapicalprimary seta. At least one pore occurs on the ventral surface. A small sclerite, the dististypes (di) (Fig. 43 A) occurs just above the ventral apex of the palpifer. The dorsal surface of the palpifer is variably armed and often bears a carina running basolaterally to mesoapically. 71 The dorsal apex of the palpifer commonly bears one to several usually elongate, robust dorsoapical primary setae. In the Idoliini, the dorsal surface of the palpifer bears an unusual extension termed the palpifer condyle (pc) (Figs. 40A, 52A). The dorsal membrane may be covered with minute to short vestiture termed dbrsal membrane setulae. The labial palp is four segmented. The proximal palpal segment always bears a pair of minute setae on the lateral margin and a cluster of pores in the apical membrane. The second palpal segment is commonly armed with short setae laterally and lateroapically. The third palpal segment is plesiomorphicly armed with a whorl of 5 short apical primary setae. The terminal palpal segment is variably armed and plesiomorphicly bears a lateral cluster of sensilla digitiformia termed the palpal organ (Fig. 40B), several apical pores, and several isolated sensilla digitiformia near the apex. The apical membrane usually bears relatively few minute sensilla basaconica. Labium Morphology: Terminology for the gross morphology of the prementum follows that of Williams (1938). The paraglossae (pa) (Fig. 43E) and glossae (gl) (Fig. S5E) in histerids consist of membranous lobes. Paraglossal lobes are always present whereas distinct glossal lobes may or may not be apparent. The dorsal labial surface is densly covered with microtrichiae which are variably organized among genera. In Onthophilus and Tribalus the pile is relatively long and dense, whereas in and Eutribalus the pile is much shorter. The apical ligular margin is fringed with setulae. The mesoventral ligular disc is armed with relatively long and often robust setae termed ligular disc setae. A generally elongate and acuminate structure termed the ligular sclerite (Is) (Fig. 43E) articulates with the paraglossae basolaterally. The palpiger (pg) (Fig. 43E) commonly bears a single minute seta and at least one pore on the ventral surface; it may or may not be armed with setae laterally. The variable basal appendix is termed the basal extension (be) (Fig. 43E). The basal palpal segment commonly bears a single minute lateral seta. The second palpal segment commonly bears at least a single pore and is 72 armed with an apical whorl of 4 short to medium- length primary setae and a variable number of lateral setae. The terminal palpal segment is variably armed and expanded and dorsoventrally flattened in some taxa (Fig. 40D). Apical pores and sensilla digitiformia are often present. Sensilla basaconica in the distal membrane are either short and few in number or relatively long and numerous. The base or substructure of the prementum is internal and underlies the mentum. The substructure has apparently not been studied elsewhere in the Coleoptera and consequently terminology presented here is new. A small, variably shaped sclerite, the dorsomedial sclerite (do) (Fig. 43E), lies between the basal extensions. A pair of partially hyaline structures termed lateral arms (ar) (Fig. 43E) extend from the mesal base of the glossa to the apical mesal border of the basal substructure. A small medial sclerite is sometimes present in the center of the apical substructure. The basal substructure consists of a body which bears anteriorly a pair of produced, elevated, and laterally flattened processes termed alae (sa) (Fig. 43E). The alae in certain groups are ventromesally closed by thin cuticle, forming a cupule. A much shorter process projecting anteriorly process termed the median lobe (ml) (Fig. 43E) is generally present between the alae. The substructure body may be laterally lobed. Projecting posteriorly from the base of the substructure body are a pair of variable, dorsoventrally flattened structures termed substructure legs (si) (Fig. 43E). The base of these legs are often expanded into plates which may have lateral extensions with sclerotized apices. Antennal Protection: The antennal club of histerid beetles is replete with olfactory sensilla. It is likely that without these structures, histerid beetles would not be able to locate volatiles emitted from decaying materials and fermenting liquids, and possibly fungal spores (J. Kimbrough, pers. comm.) and. Several different methods of antennal protection have evolved within the Histeridae. In the least elaborate of these methods, characteristic oî Anapleus and Dendrophilus, the antennae are shielded by the fore tibiae 73 and femora when the beetle withdraws its appendages. Antennal grooves (ag) (Fig. 13C) are present alongside the prosternai keel. Depressions for receiving the enlarged antennal club are present in some taxa (eg. Sphaericosoma and Geocolus). Saprinines similarly use their withdrawn forelegs to protect their antenna but in nearly all taxa there are distinct cavities for receiving the antennal club in front of the procoxae and next to the prosternai keel. In onthophiline and tribaline histerids the anterior angles of the prothorax are equipped with cavities (ac) (Fig. 13B, D) into which the antennae are drawn when the beetles are disturbed. Hind Wings; Hind wing venation follows that of Kukalovâ-Peck and Lawrence (1993). The hind wing venation of Dendrophilus strongly resembling that found in Sphaerites and Syntelia. Shared features include the presence of J and AP3+4, KP2 and RP3+4 complete and extending to the wing margin as veins, and CUA2+3+4 lining up to the right of CuA. In Sphaerites and Syntelia, MP3+4 and CUA2+3+4 are true veins whereas in the Histeridae these veins are present as sclerotized membrane. A distinct pigmented area occurring immediately distad to the pinch in histerids is termed RA3+4 stigma (Fig. 42A). Carinal, stria!, and sutural terminology: Terminology used to describe external sutures, striae, and carinae follows that used by Wenzel and Dybas (1941) and Helava (1978). The 2 lateral metastemal striae are differentiated as lateral and dorsolateral metastemal striae. The lateral metastemal striae extend from the meso-metastemal margin to the metacoxae. The dorsolateral metastemal striae extend from the mesocoxae to the metastemal-metepistemal suture. The pair of striae extending from the metacoxae to the posterior margin of the first apparent abdominal stema are termed lateral abdominal striae (las) (Fig. 15B). Terminalia: The terminology applied to the normally concealed last three abdominal segments for both males and females varies and is occasionally conflicting (Helava 1978, 74 1985; Vienna 1980). In an effort to resolve these discrepancies, I studied the last three abdominal segments of exemplars from all histerid subfamilies except for the Niponiinae, Trypanaeinae, and Tiypecticinae. I examined and compared component structures in each of the three abdominal segments in both sexes to identify serially homologous structures. I then compared terminalia morphology between sexes. The terminal abdominal segments in the male essentially form a telescoping syringe. Those of the female form an extensible ovipositor bearing terminal coxites. Despite their functional differences, the component parts of the male copulatory apparatus and the female ovipositor are derivable from differentially modified tergal, pleural, and sternal elements. The un-modified eighth segment in males (Fig. 56A, B, C) is composed of a large tergite (te) and smaller paired articulating distal processes. The morphology of the male tergite in onthophilines is variable. The posterior margin is often medially emarginate. In some cases the emargination is extreme, producing lateral tergal extensions or tails. The tergite often bears minute setae and may not bear cuticular cavities. I decided to term the articulated processes coxites (cx). This term has traditionally been applied to the terminal articulated structures of the female ovipositor which I regard as homologous to those found in males. The coxites of females each bear an articulated apically setose structure termed a stylus. The coxites of male Dendrophilus, Anapleus and apparently some species oî Stictostix also bear styli nearly identical to those found in females. The coxites of male onthophilines are armed apically with setae which usually include some which are short to medium length. In addition, the internal apical surface in some taxa is lined with a pubescent membrane. The un-modified ninth segment (Fig. 56D, E, F) resembles the eighth in terms of structure and the relative proportions of the two elements. However, in this segment, the coxites have apparently fused, forming a distal coxal carapace (cc). The dorsal position of the coxal carapace has apparently led to some confusion, as this structure is generally 7 5 considered to be the tergum of the 10th segment. However evidence suggests that the coxal carapace is likely a serial homolog of the coxites of the previous segment. This structure, besides occupying approxi\mately the same position as the coxites of the previous segment is sometimes divided by a longitudinal suture and apparently the product of fusion of two separate structures. Like the coxites of the eighth segment, the coxal carapace is armed apically with relatively long setae and is lined with an internal pubescent membrane in some taxa. The ninth segment has a ventral sclerite associated with it not found in the eighth which is generally referred to as the spiculum gastrale (sg). This structure may be the eight stemite. The terminology of the aedeagus (Fig. 56G, H, Ï) follows that of Lawrence and Brittain (1991). The morphology of the preceding segments and that of the 10th segment of the female shed some light on possible serial correspondence to some of the component structures of the aedeagus. The phallobase (ph) is probably homologous with the terete of the 8th and 9th segments while the parameres (pm) appear to be homologous with the coxites of the same segments. The origin of the median lobe remains obscure.

Results Key to world genera of Onthophilinae and TVibalinae.

1 Pronotum with at least 2 costae (Figs. 14A, 15A,C,26B)...... 2 1' Pronotum lacking costae ...... 6

2(1) Antennae < width of head capsule ...... 3 2' Antennae much longer than head capsule width ...... 5

3(2) Frons and at least pronotal bead with erect amber scales (Fig. 41C); conspicuous fovea generally present on meso-, meta-, first apparent abdominal sternum; (mainly Old World tropics) ...... 4 3' Erect amber scales absent, elytra costate, sternum punctate; Australia...... Australepierus (in part) ^4. australis\ 7 6 4(3) Sternal fovea present (Fig. 15D), elytra with raised costate; cuticle shagreened, iridescent ...... Epiechnus 4* Sternal fovea absent (Fig. 15B), elytra with impressed striae; cuticle not shagreened ...... Vuattouxinus

5(2') Pair of trichome-lined foveae anterolaterally on pronotal disc (Fig. 26B, C) and prosternai keel; prosternai lobe produced; carinal striae present; umbiculately punctate; (New World) ...... Peploglyptus 5* Pronotum and prosternai keel lacking fovea; prosternai lobe not produced; carinal striae absent; sternum punctate; Holarctic and Oriental distribution ...... Onthophilus

6(1') Antennae much longer than head capsule width (western Nearctic and Australia)...... Stictostix 6' Antennae < head capsule width ...... 7

7(6') Lateral abdominal striae absent; antennal club externally divided by apical and basal sutures (Figs. 27D, 32A); whorls of sensilla basiconica originating from sutures accentuating divisions; elytra lacking distinct and complete dorsal and sutural striae; hind wing membrane pigment unapparent; dorsal body outline somewhat circular (Fig. 26D); generally somewhat flattened dorsoventrally (Old World) ...... Tribalus 7' Lateral abdominal striae present; antennal club lacking external sutures, although transverse whorls of sensilla basiconica originating from sutural areas give a divided appearance; elytral striae variable; hind wing membrane pigment distinct ...... 8

8(7') Laterofrontal sutures very long, V-shaped; body form globose (Fig. 30C- D) or lentil-shaped; elytra with at most with single subhumeral stria, terminal segment of labial palpi expanded and flattened dorsoventrally; small beetles generally 2 mm or less ...... 9 8' Laterofrontal sutures usually short; if long they are angular and not V- shaped; body form never globose or lentil shaped; elytral striation and terminal labial palpal segment, and size variable ...... 10

9(8) Pronotum and elytra margined; meso-metastemal stria superficially indicated (Oriental) ...... Paridolia 9' Pronotum and elytra not margined; meso-metastemal stria lacking (Fig. 36C); (Neotropical) ...... Idolia 77 10(8') Antennal club interrupted by 1 or 2 transverse whorls of sensilla basiconica (Fig. 17C, D); elytra with at least one complete dorsal stria, dorsal body outline generally oval (Fig. 14B, C, D); mandibles bidentate; terminal labial palpal segment acuminate ...... 13 10' Antennal club not transversely interrupted; other characters variable 11

11(10') Mandibles bidentate (Fig. 20D); prosternai lobe not produced; antennal club ovoid in outline (Fig. 17B); terminal segment of labial palpi acuminate; galea expanded laterally, dorsal surface densely covered with short setulae; elytra often with a complete set of striae; dorsal body outline generally oval; elytra rarely costate (Australia and New Zealand) Australepierus 11 ' Mandibles unidentate (Fig. 34A, C); prosternai lobe produced; antennal club hatchet shaped in outline (Fig. 32B); terminal segment of labial palpi expanded and flattened dorsoventrally; dorsal body outline always circular, elytra never costate ...... 12

12(11') Meso-metastemal stria transverse or feebly arcuate anteriorly (Fig. 3 6A); mentum generally densely covered with flexible, plumose setae of uniform length; elytra with suturé striae only; generally not dorsoventrally flattened (Old World tropics; one Nearctic species) ...... Eutribalus 12' Meso-metastemal stria strongly anteriorly arcuate (Fig. 36B); mentum with mixture of short and long stiff setae; elytra usually with 4 complete dorsal striae in addition to suturais often dorsoventrally flattened (Oriental) ...... Parepierus

13(10) Meso-metastemal stria present in addition to marginal metastemal stria (Fig. 25D); marginal metastemal stria often evanescent medially (Neotropical with 2 Nearctic and 1 Palearctic species) Plagiogramma 13' Meso-metastemal stria absent (Fig. 25C); marginal metastemal stria not evanescent medially (Neotropical with some Nearctic; Oriental Palearctic, and Australasian species) ...... Epierus 78 Onthophilinae Thompson 1862 Included in the subfamily Onthophinae Thompson, are the genera Onthophilus Leach, Epiechinus Lewis, Vuattouxinus Thérond, Australepierus^ new genus, Epierus Erichson and Plagiogramma Tarsia.

T ype genus : Onthophilus Leach. Adult Redescription: Body outline (dorsal) commonly circular or ovoid. Head. Frons with laterofrontal carina; verticofrontal carinae present or absent. Antenna. Surface texture generally imbricate. Eighth flagellar segment about half as long as wide. Club completely fused, sulci reduced or absent. Labrum setose. Epipharynx. Lateral area margin usually with regularly spaced fringe of marginal setae; lateral area disc bearing mainly relatively small, tooth like non-overlapping scales; vestiges of outer brush present or absent; inner brush well developed and sometimes bordered apically by setular comb. Mesal area with dorsomedial brush present. Mandibles commonly with dorsolateral area constricted and covered with regularly spaced mostly minute setae; longer robust contact chemoreceptors may be present dorsoapically. Mesal area apically acuminate or not. Mola jutting mesad; microtrichia of dorsal microtrichial field well-developed or somewhat reduced. Prostheca with dorsal setulae short; some setulae may be more elongate distally but never extending beyond incisor; inner prosthecal setulae variably. Maxilla. Galea variously modified for microphagy. Lacinia with imbricate texture dorsally, ventral rugosity and with or without strobilus; lacinial tooth unmodified; lacinial spines present; lacinial fiinge with variable blade-like setulae. Proximal palpal segment laterally armed with a pmr of minute primary setae. Third palpal segment generally armed with apical whorl of 5 primary setae. Terminal palpal segment variably armed. Labium. Mentum with anterior margin outwardly arcuate or unidentate and with several internal cuticular cavities (possible gland ducts) located near anterior margin. Palpal segments un-textured. Palpiger generally with an elongate basal extension and with single minute seta on ventral disc. 79

Proximal palpal segment armed with single minute basolateral seta and bearing several pores in apical membrane. Second palpal segment generally with 4 ventral primary setae. Terminal palpal segment variably armed and with short sensilla basiconica at palpal apex. Ligular sclerites relatively long and broad; premental substructure otherwise variable. Thorax. Prothorax. Notum with or without costae; sternum with alae and with anterior margin of lobe straight or produced; carinal stria present or lacking. Fore-tibia generally slender and occasionally toothed. Elytra striate or costate. Hind wings with variable venation; wing membrane nearly always with dark maculations. Meso-Metastemal surfaces variably punctate; marginal mesostemal stria generally distinct; other striae variable. Abdomen. Propygidium and pygidium variably punctate and occasionally costate. First apparent stemite with lateral abdominal striae. Diagnosis: Members of the Onthophilinae can be distinguished from those of the Tribalinae and other histerids by the following combination of characters: Antennal club fused with sulci vestigial or absent; labrum setose; epipharynx lateral areas with reduced, denticulate and generally non-overlapping scales; mandibles with prostheca apically detached (except in Onthophilus)', lacinia with strobilus (secondarily lost va. Plagiogramma and Epierus)', prosternai alae present; hind wings generally with dark maculations.

Onthophilus genus group The Onthophilius genus group is an informal group of related genera including Onthophilus Leach, Vuattatouxinus Thérond, and Epiechinus Lewis. Adult description: Head. Frons generally with vertico-frontal carina. Antenna. Length variable. Club with at most single transverse whorl of sensilla basiconica; sensory plaque present or absent. Epipharynx. Lateral fringe and inner brush complete. Mandibles with contact chemoreceptors at apex of lateral area. Medial area apically acuminate. Mola jutting mesad. Microtrichia of dorsal microtrichial field well-developed. Prostheca with 80 dorsal setulae short; inner prosthecal setulae with or without comb setulae. Maxilla. Galea with numerous flexible setulae on ventral surface; dorsal surface densely covered with short, slender setulae. Lacinia with or without strobilus; flinge complete. Palpifer carina abbreviated basally. Terminal palpal segment armed with several minute setae or appearing unarmed. Labium. Mentum with anterior margin outwardly arcuate. Palpiger with elongate basal extension. Terminal palpal segment appearing unarmed. Substructure alae elongate, strongly elevated, and open ventromesally. Thorax. Prothorax. Notum with 1-3 costae per side; sternum with anterior margin of prosternai lobe straight or feebly outwardly arcuate; carinal stria present or lacking. Fore-tibia slender with anterior margin toothed. Hind wings with wing vein RP2 complete; otherwise venation variable. Meso- Metastemal surfaces punctate and occasionally with fossulate fovea. Abdomen. Propygidium and pygidium punctate and oflen costate. First apparent stemite occasionally with fossulate fovea. Diagnosis: Members of Onthophilus species group can be distinguished fi*om those of the Epierus species group by the following characters: Frons with vertico-fi'ontal carina; mandibular medial area apically acuminate; palpifer carina complete; pronotum with at least one pair of costae; anterior margin of fore-tibia toothed; hind wings with wing vein RP2 complete.

Genus Onthophilus Leach (Figs. 14A, 16A, ISA, 20A, 21A-C, 39C, 40B-C, 42A, 43, 56).

T ype Species Hister sulcatus Moll Adult redescription: Body outline (dorsal) (Fig. 14A) oval to circular. Head. Frons. Laterofrontal carina (when distinct) relatively long and joining at the level of the clypeus. Verticofrontal carinae occasionally indistinct. Antenna (Fig. 16A) longer than width of head capsule. Scape petiolate basally, gradually expanded distally; dorsal surface 8 1 smooth or punctate; uniform covering of minute setae with several scattered short robust setae. Club somewhat flattened dorsoventrally and densely covered mainly with sensilla trichodea; prominent elongate sensilla chaetica originate from areas adjacent to former club division and toward antennal apex; some short transverse bands of sensilla basaconica originating from former club divisions; distal sulcus lacking; weak proximal sulcus remnant superficially discernible and well indicated with transmitted light. Epipharynx (Fig. ISA). Broadly ovoid in outline. Outer margin of lateral area with tight fiinge of ca. IS regularly spaced elongate setae. Lateral area narrow and densely covered with small, mostly non-overlapping acuminate scales; outer brush remnant present centrally. Inner brush composed of medium length setulae. Medial area with 3 short peg setae on each side; sensilla relatively large, tuberculate, and uniformly distributed; dorsomedial brush setulae relatively long and robust, brush expanded centrally and narrow laterally; keel basally abbreviated; dorsomedial peg setulae relatively slender and elongate. Mandibles (Figs. 20A, 21A-C). Dorsolateral area expanded basally and with large basal concavity; disc and lateral margin armed with minute setae and commonly 3 short, robust, apical contact chemoreceptors; some taxa with one or more relatively large depressions or cavities (putative mycangia) (Fig. 20A [my]) present midway along dorsolateral area. Mesal area with articulated setose appendage sparsely armed with minute setulae. Dorsal microtrichial field extensive, completely concealing molar cavities when viewed dorsally; microtrichia well-developed. Ventral microtrichial field (Fig. 21 A, C) very broad and extending to base of mandible. Prostheca attached apically; some dorsal setulae apically branched; ventral prostheca with comb setulae (Fig. 2 IB). Maxilla (Figs. 39C, 40B-C, 43A-C). Galea flaring distally; ventral surface largely membranous and covered with relatively long, flexible setulae; basal 1/2 of dorsal surface of galea sclerotized and armed laterally with setulae; apical 1/2 of dorsal surface membranous and very densely covered with slender stiff setulae with mesally curved tips (Fig. 39C, 43B). Lacinial strobilus (Fig. 82 40C) detached, sometimes elongate and slender, and without terminal blade-like setulae; lacinial tooth (Fig. 43C) robust, with basal portion more than twice as long as toothlets; several discal setae clustered directly below tooth, fiinge setulae short and slender; distal 1/3 of dorsal lacinial disc armed with short setulae; blade-like setulae numerous, short, and slender. Mediostipes broad, untextured, and with numerous minute setae, a pore, and several small internal cuticular cavities. Basistipes untextured, and bearing three long, robust, primary setae and 2 pores. Cardo untextured and with several short and minute setae. Palpifer without surface texture and laterally armed with numerous minute to medium length setae; ventroapical primary seta long and robust; dorsolateral surface with cluster of minute-short setae basally; 3 short-long dorsoapical primary setae present; other generally minute setae sometimes present dorsoapically on palpifer. Dorsal membrane with numerous minute denticulate setulae basally and increasing in length slightly toward apex. Proximal palpal segment untextured, relatively long (longer than 3rd segment), centrally constricted and outwardly bent. Second palpal segment long, about twice as long as wide, untextured, and armed with short setae laterally, and 2 pores ventroapically. Third palpal segment with imbricate texture basally; commonly armed with a few minute lateral setae. Terminal palpal segment with basal imbricate texture; palpal organ (Fig. 40B) covering 2/3 of the distal lateral margin; several short and minute setae distributed mainly on ventral surface; several pores and sensilla digitiformia present apically. Labium (Fig. 43D-F). Mentum (Fig. 43D) rectangular; generally with mixture of 8-10 long robust setae and numerous short and minute setae on ventral surface; lateral margins fiinged with short and minute setae. Prementum (Fig. 43E-F) with lobes of paraglossae broad, long, and with minute denticulate setulae basolaterad; paraglossal appendix lacking; ligular fiinge consisting of mixture of slender and robust medium length setae; robust setae sometimes apically expanded and forked; glossal lobes undifferentiated; ligular disc setae robust, medium length and -15 in number, several pores near setal bases. Palpiger with 83 several pores on dorsal surface and several medium length setae on lateral margin. Proximal palpal segment relatively long, ca. twice as long as wide. Second palpal segment with variable number of short setae laterally and with medium length primary setae and pore at apicax. Terminal palpal segment long and narrow, dorsal surface with mainly minute setae, some short setae commonly present on mesal margin; palpal organ at middle of segment; several pores and longitudinal sensilla digitiformia at apex. Substructure with dorsomedial sclerite with anterior face small and oval and posterior extension greatly expanded laterally. Hyaline section of central arms slender and elongate; pigmented section short and cylindrical. Medial sclerite apparently absent. Median lobe slender and elongate, slightly swollen at middle. Body bordering alae with broad lateral lobe on each side. Legs relatively short, divergent, mesal border arcuate. Leg bases laterally expanded and with short lateral extensions with sclerotized apices. Thorax. Prothorax. Notum with 2-6 costae; sternum with anterior margin of lobe straight; carinal stria lacking. Fore­ tibia elongate, slender and variably armed. Elytra with eight costae; deep pits sometimes found in intervals along anterior margin. Hind wings (Fig. 42A) with R4 and RP separate; RP relatively long; RA3+4 stigma more than 2x longer than wide; RA3 absent; RA4 and RPl dark and distinct; RP connected to but not fused with MAl+2; RP2 and RP3+4 complete, extending to wing margin as veins; Mp3+4 distinct and long, nearly reaching wing margin; CUA2+3+4 lined up to right of CuA; CUA and AAl+2 joining beyond end of jugal lobe and terminating in characteristic eye; AP3+4 present; J indicated; pigmented patterns relatively dark and distinct. Meso- and metastema with surfaces punctate, often coarsely so; marginal mesostemal suture indistinct; meso-metastemal suture and longitudinal metastemum suture either faintly indicated or obliterated by punctures. Abdomen. Stemites with varying degrees of punctation. Abdomen. Propygidium and pygidium punctate and sometimes costate. Male Terminalia (Fig. 56). Eighth segment (Fig. 56A-C). Tergite dorsum about as long as broad and obtusely emarginate proximally 84 and acutely emarginate distally; armed with minute setae dorsolaterally and laterally. Coxites relatively short, conical and with minute setae on dorsal and ventral surface and several short setae towards apex; numerous minute pore-like cuticular cavities present ventrally and mesally; apical pubescent membrane present. Ninth segment (Fig. 56D-F) distinctly capsular. Tergite subdivided into 2 laterally broad valves armed dorsoapically and ventroapically with minute setae. Coxal carapace elongate, ca. 3 x longer than wide and with distinct longitudinal suture; armed distally with several short and minute setae; pore-like cuticular cavities present distolaterally; apical pubescent membrane present. Spiculum gastrale elongate, relatively broad, somewhat expanded distally and posteriorly bisected for about 1/2 of length of sclerite. Aedeagus (Fig. 56G-I) relatively elongate with phallobase globular or tubular; parameres commonly parallel-sided, divided distally, and armed with minute-short setae on ventroapical surface. Diagnosis: The following combination of characters will distinguish adult Onthophilus from all other histerid genera; Antennae longer than head capsule width; ventral prosthecal setulae pectinate; ventral galeal surface densely covered with flexible setae; dorsoapical galeal surface very densely covered with stiff, slender setulae; strobilus detached, appendicular, and without terminal blade-like setulae; dorsal lacinial disc armed; proximal maxillary palpal segment relatively long, centrally constricted and outwardly bent; second palpal segment slender and elongate; ligular disc setae robust medium length and numerous; palpiger laterally armed with several medium length setae; pronotum with at least 2 costae; prostemum with anterior margin strmght and carinal striae absent; elytra with 8 costae and punctate intervals; hind wings RP2 and RP3+4 complete; Mp3+4 nearly complete; AAl+2 joining beyond end of jugal lobe and terminating in characteristic eye; AP3+4 present; propygidium and pygidium costate and punctate; teminalia with 8th and 9th segments capsulate; parameres of aedeagus distally divided. 85 Distribution. Over 90% of the described species of Onthophilus occur in the Holarctic. The remainder are Oriental. Immature Stages. The larva of Onthophilus nodatus LeConte is described in Chapter 1. The pupa of O. kim i is described in chapter 2. A single second instar of an undetermined species of Onthophilus was extracted by P. Skelley from soil beneath rabbit dung in January. Larvae of O. kim i have been collected from soil within the nest and/or fecal chambers of pocket gophers, Geomys sp., (Geomjddae: Rodentia) from several localities. P. Skelley collected larvae of this species in Alachua Co., Florida during January and February ; P. Skelley, R. Tumbow, and M. Thomas collected others during February in Dale Co., Alabama; W. Godwin collected O. kim i larvae during April in Wood Co. Texas; and K. Stephan took many larvae of this species during November and March in Latimer Co. Oklahoma. A first instar larva of O. giganteus was collected by P. Skelley in a Geomys burrow in January in Alachua Co., Florida. Biology/Ecology. Adult and larval Onthophilus have been found associating with dung, carrion, tree wounds, mammal nests, rotting fungi and plant material (Helava 1978, Ohara and Nakane 1986, unpublished label data). Several of these species are attracted to proprionic acid (Helava 1978). Adults and larvae maintained in the laboratory required relatively high humidity (Kovarik, unpublished data). Unlike most histerids, adult periodicity and reproduction for members of this genus apparently coincides with cooler months of the year. Adult periodicity and flight activity was determined for Onthophilus nodatus via automated pitfall traps in Burleson County Texas, by Summerlin et al. (1993). This species was found to be active mainly in the winter/spring (February-April) and autumn/winter (September-December). Specimens were captured between 1200 and 2200 h.. Peak flight activity occurred between 1500 and 1600 h.. Adult activity for two obligate nidicophilic species of Onthophilus associated with pocket gophers was determined by P. Skelley (unpublished data) in Alachua County, 86 Florida via pitfall traps placed within gopher burrows. Onthophilus kimi Ross and O. gigcmteus Helava, both associated with pocket gophers. Adults of O. kim i were active within the burrow systems from late December to mid-March. The activity period for Onthophilus gigcmteus is more restricted than that of O. kimi. Adults of O. gigcmteus were active from late November to early February. Very few records are avmlable for field collected larvae. Helava (1978) erroneously concluded that adult Onthophilus were predators on the basis of their mouthpart structure. The mouthparts of Onthophilus are adapted for filter feeding. The comb hairs on the inner surface of the prostheca of adult Onthophilus are nearly identical to those found in larval biphyllids (Coleoptera) which feed on fermenting liquid. According to Lawrence (1989) these hairs act as filter feeding devices. The dorsal surface of the galea (Fig. 39C, 43B) is a modified filtering device, the structure of which is unique to the Onthophilus genus group and probably the Hsteridae. Another elaborate filter is present on the inner surface of the ligula and substructure of the prementum. It is composed of overlapping layers of fine setulae. I have observed adults of O. kim i feeding on fresh guinea pig dung. The beetles appeared not to ingest any solid material, but rather use their mouthparts to gather liquid on the surface of the dung. The gut contents of a specimen of Onthophilus nodatus from Alachua County, Florida contained spores of the hyphomycetes Trichocladium pyriforrme Dixon and Cladosporium sp. The ephemeral nature of dung, carrion, and other rotting substrates in warm weather may explain why adult periodicity of Onthophilus generally coincides with cooler temperatures. During the summer in temperate regions, dung is rapidly eliminated by fly larvae and scarab beetles (Mohr 1943). In cooler weather when most dung-associated Diptera and Coleoptera are inactive dung remains relatively undisturbed for longer periods of time (Mohr 1943, Merritt and Anderson 1977, Laurence 1954). Cooler temperatures 87 may provide the best window of opportunity to obtain liquid nutriment from dung and other ephemeral rotting substrates. Some species of Onthophilus associate with ants. Hirano (1986) reported an association of O. silvae Lewis with Lasiusfuliginosus (Latreille). W. P. MacKay discovered a specimen of O. lecontei in the nest of Pogonomyrmex rugosus Emery and an undetermined species of Onthophilus in the nest of P. subnitidus Emery. A number of species of Onthophilus are associated with mammal nests or burrows. A few appear to be nonspecific in their selection of host nest. The Nearctic O. deflectus has been found in “mouse” and wood rat, Neotoma sp., nests (Helava 1978 and unpublished label data). The Palearctic O. punctatus has been found in nests of several mammal species including ground squirrels, (Citellus spp.), voles, (Microtus spp.), mice, (Mus spp.), old world rabbits, (Orctolagus spp.), old world moles, (Talpa spp.). Hamsters, (Cricetus sp.), and the old world badger me les Storr. Twenty-five percent of the described species of Onthophilus inhabit the burrows systems of certain species of rodents. These species complete their life cycle within the nest and fecal chambers of the rodent burrow and apparently only rarely leave them. According to Helava (1978) the Nearctic O. cynomysi inhabits open burrows of prairie dogs, Cynomys sp. (Rodentia: Sciuridae). Onthophilus lecontei Horn and O. thomomysi Helava (and probably 0 . soltaui Casey) inhabit the closed burrows of Western pocket gophers, Thomomys sp. (Rodentia: Geomyidae). Onthophilus kimi Ross, O. gigcmteus Helava, and O. wenzeli Helava inhabit the closed burrows of Eastern pocket gophers, Geomys sp. (Rodentia: Geomyidae). O. convictor inhabits burrows of the long clawed mole-vole Prometheomys schaposchnikawski (Rodentia: Muridae) (Kryanovskij 1977). Nidicophilus species typically have more elongate legs and antennae than their free living counterparts. With a lengthened stride, these beetles may be able to more quickly traverse distances within burrows which for some species of pocket gopher average 53 meters in 88 length (Brown and Hickman 1973). Some of the nidicolous species have pits or depressions in the mandibles (Fig. 20A [my]) that appear to be mycangia. These beetles may inoculate fresh sources of dung with fungal spores and other microbes via material contained within these structures.

Genus Vuattowdnus Thérond (Figs. 15A-B, 16B, 19A, 21D, 44)

T ype Species Vuattatouxinus borrasicola Thérond Adult redescription: Body outline (dorsal) (Fig. ISA-B) oval. Head. Frons with short laterofrontal carina; frontal carinae and vertex sparsely covered with minute, erect, amber scales. Antenna (Fig. 16B) shorter than width of head capsule. Scape short, slightly longer than broad, and swollen; surface with strong imbricate texturing and armed dorsally with mixture of minute slender and short stout setae. Club somewhat flattened, oval in outline, and densely covered with sensilla trichodea; group of sensilla basaconia arranged in large, shallow depression, forming distinct plaque on anterodorsal club surface; elongate, regularly spaced sensilla chaetica originating near former sutures and toward antennal apex; all vestiges of antennal sulci completely lacking. Epipharynx (Fig. 19A) very broad and trapezoidal in outline. Outer margin of lateral area with loose fringe of ca. 10 robust regularly spaced medium-length setae; lateral area with outer brush remnant near dorsal margin; most scales on lateral area reduced and denticulate; some larger rounded scales present mesally. Inner brush apically abbreviated and composed of relatively long setulae. Medial area with three peg setae on each side; medial area sensilla minute and distributed mainly laterally; dorsomedial brush setulae short; brush expanded centrally and narrow laterally; dorsomedial peg setae very short and robust. Mandibles (Figs. 2GB, 2 ID). Dorsolateral area expanded basally and with large basal concavity; disc armed vdth minute setae and 3-4 short, robust, apical contact chemoreceptors. 89 Dorsomedial area with articulated setose appendage armed with relatively long, robust setulae. Dorsal microtrichial field slightly smaller than that of Onthophilus; ventral microtrichial field not extending to mandible base; prostheca apically detached dorsal setulae short and unbranched; ventral setulae (Fig. 2 ID) arranged in transverse rows progressively decreasing in length; comb setulae present at apex. Maxilla (Fig. 44A-C). Galea flaring distally; ventral surface densely covered with medium-length flexible setulae; apical margin of galea flinged with medium length spatulate setulae; apical 1/2 of dorsal surface of galea densely covered with slender stiff setulae with mesally curved apices. Lacinia with strobilus absent, lacinial tooth (Fig. 44C) with basal portion about as long as toothless; several discal setulae arising below tooth; flinge setulae short and slender; blade-like setulae numerous, short, and slender. Mediostipes relatively narrow and bearing numerous minute setae, a pore, and a relatively large cuticular cavity. Basistipes with some surface texture and with three robust, medium length, apically barbed primary setae; some minute secondary setae and 2 pores also present. Cardo untextured and with several minute setae. Palpifer with surface texture, two pores and several short basolateral setae; palpifer ventoapical primary seta long, robust, and apically barbed; dorsolateral surface with long row of short regularly spaced setae; dorsal apex with 1 long and 2 medium- length primary setae, a peg-like sensillum, and single pore. Dorsal membrane with minute denticulate setulae. Proximal palpal segment untextured and short. Second palpal segment ca. twice as long as wide, with some proximal texture, 2 basal pores, and armed laterally and apically with short setae. Third palpal segment with basolateral surface texture and armed laterally with several minute setae and single ventral pore; apical primary setae short. Terminal palpal segment with mesal and basal surface texture, palpal organ restricted to basal 1/2 of segment; single ventral pore present; armed apically with vestigial setae; several pores and sensilla digitiformia at apex. Labium (Fig. 44D-F). Mentum (Fig. 44D) trapezoidal and with 5 long robust barbed setae, several medium length setae, and 90 numerous short and minute setae on ventral surface. Prementum (Fig. 44E-F) with lobes of paraglossae short, broadly rounded, and lacking appendix; weakly produced lobule located basolaterally; ligular fiinge composed of short-medium length setulae; glossal lobe undifferentiated; ligular disc setae long, robust, and numbering 2 per side; several pores present at glossal base mesally. Palpiger with single pore on dorsal surface. Proximal palpal segment short, slightly wider than long. Second palpal segment with single ventral pore and armed with 2 minute lateral setae and short-medium length primary setae. Terminal palpal segment elongate, ovoid, and bearing a few vestigial setae; palpal organ basal and several pores and sensilla digitiformia at palpal apex. Substructure with dorsomedial sclerite wedge-shaped. Hyaline section of central arms slender and elongate; pigmented section long and cylindrical. Medial sclerite absent. Median lobe long and broad. Legs very short, about 1/3 length of entire substructure; mesal border arcuate, leg base with lateral hyaline extension terminating in sclerotized knob. Thorax. Prothorax. Notum with bead and with weakly developed pronotal costa 1; disc uniformly covered with minute erect amber scales; costae and bead with larger scales; sternum with anterior margin of lobe straight; carinal stria present. Fore-tibia slightly expanded distally. Elytra with a complete set of striae and relatively uniformly covering of small scale-bearing punctures. Hind wings with R4 and RP separate; RP relatively short; RA3+4 stigma more than 2x longer than wide; RA3 very short; RA4 and RPl dark and distinct; RP fused with MAI+2 along most of its length; RP relatively short; RP2 complete; RP3+4 apically abbreviated, ending as vein about half way to wing margin and extending to wing margin as sclerotized membrane; Mp3+4 distinct and apically abbreviated, ending well short of wing margin; AAl+2 joining at end of jugal lobe, AA3+4 fusing with CUA2+3+4 shortly beyond end jugal lobe; CU2+3+4 represented apically by slender vein; AP3+4 absent; J indicated basally; pigmented patterns relatively dark and distinct. Meso- and metastuma surfaces (including legs) with small scale-bearing punctures; marginal metastemal suture 91 absent; meso-metastemal suture faintly impressed only; longitudinal metastemal suture absent. Abdomen. Stemites with small, shallow punctures with minute scales; lateral abdominal sutures carinate. Propygidium and pygidium uniformly covered with small, shallow punctures with minute scales. Male Terminalia. (not figured) Eighth segment. Tergite dorsum about as long as broad and deeply emarginate proximally; remaining disc medially hyaline giving tergite subdivided appearance; tergite produced lateroposteriorly into rounded tails on each side and armed dorsoapically with few minute setae; cuticular cavities present dorsoapically. Coxites about as long as tergite, relatively broad, somewhat dorsoventrally flattened, and armed distally with 2 short stout setae; several minute, inconspicuous setae present on doroapical and dorsolateral surfaces; numerous irregular cuticular cavities located dorsoapically and dorsolaterally; ventral surface of coxite largely membranous and with apical pubescent membrane. Ninth segment non-capsular and travois shaped. Tergite subdivided into 2 proximally tailed, unarmed valves with numerous cuticular cavities. Coxal carapace short, broad, and composed of sclerotized and granular cuticle; carapace without longitudinal suture and armed distally with 2 short conspicuous setae; cuticular channels present distally; apical pubescent membrane lacking. Spiculum gastrale missing firom the single male specimen available for study. Aedeagus short and stout with phallobase asymmetrical and fused to paramered; parameres united distally and armed with short, stout setae at ventral apex. Diagnosis: The following combination of characters will distinguish adult Vuattouximis firom all other histerid genera; Frontal carinae and vertex with erect amber scales; antennal scape short, swollen, and armed with short, stout setae; club with sensilla basiconica plaque and lacking sensilla mammillae; mandibles bearing some pectinate ventral prosthecal setulae; ventral galeal surface densely covered with flexible setulae; dorsoapical galeal surface densely covered with slender stiff setulae; strobilus lacking; terminal maxillary palpal segment unarmed; mentum with some setae barbed; paraglossae 92 with small basolateral lobule; terminal labial palpal segment unarmed; labial substructure legs very short; leg base with lateral extensions; pronotum with bead and weak pronotal costa 1, bead, costae, and disc with erect amber scales; prostemum anterior margin straight; with carinal stria present; elytra striate uniformly punctate; each puncture with minute scale; meso- and metastemum un-sculptured and with scaled punctures; propygidium and pygidium with scaled punctures; cuticle not shagreened. Distribution. Vuattoitxinus borassicola is known only from the environs of Pakobo, Ivory Coast. Immature Stages. Immature stages of Vuaitowcinus are unknown. Biology/Ecology. Comb hairs on the prostheca and the filter-like arrangement of setulae on the dorsal surfaceof the galea suggests that V. borassicola probably filter feeds on fermenting liquids. R. Vuattoux reported that this species is found in association with dying Rônier palm trees (Borassus flabellifer L.). The gut contents of a specimen of this species contained unidentifiable organic material.

Genus Epiechinus Lewis (Figs. 15C-D, 16C-D, 19B, 20C, 41C,D, 42C, 45)

T ype Species Onthophilus costipennis Fahraeus in Boheman Adult redescription: Body shape (Fig. 15C-D) circular to oval in outline. Cuticle (Fig. 4 ID) shagreened. Head. Frons with moderately long laterofrontal carinae; Frontal carina and vertex with minute, erect, amber scales. Antenna (Fig. 15C-D) shorter than width of head capsule; scape short, about as broad as long, and swollen; surface occasionally pitted and covered mainly with short, stout setae. Club somewhat rounded and densely covered with blunt sensilla trichodea and usually sensilla mammillae; sensilla basaconia oriented longitudinally, with one large sensilla basiconica grouping in large shallow depression, forming a distinct plaque on anterodorsal club surface (Fig. 15D); 93 sensilla chaetica apparently absent; all vestiges of antennal sulci lacking. Epipharynx (Fig. 19B). Trapezoidal in outline. Outer margin of lateral area with loose fringe of ca. 10 regularly spaced elongate setae; scales on lateral area reduced and denticulate and with some larger scales located mesally. Inner brush complete and composed of relatively short setulae. Medial area with three peg setae on each side; medial area sensilla minute and distributed basally and laterally; dorsomedial brush setulae short; brush wide centrally and narrowing laterally; dorsomedial peg setae short and robust. Mandibles (Fig. 20C). Dorsolateral area basally expanded and with large proximal concavity; disc armed with minute setae and 4 short, robust apical contact chemoreceptors. Articulated setose appendage armed with relatively long, robust setulae. Dorsal microtrichial field more restricted than that found in other Onthophilus genus group; ventral microtrichial field not extending to mandible base; prostheca detached apically; dorsal setulae short to medium length; ventral setulae arranged in transverse rows progressively decreasing in length. Maxilla (Fig. 45A-C). Galea (Fig. 45A-B) flaring distally; ventral surface generally sparsely covered with short flexible setulae; distal margin flinged with short spatulate setulae; apical 1/2 of dorsal surface bearing mainly short spatulate setulae with some short, tapered setulae laterobasally and short, stiff setulae with curved apicesmesobasally; dorsal sclerotized galeal base unarmed. Lacinia with strobilus ending in several mesally bent blade-like setulae; lacinial tooth (Fig. 45C) wth basal portion about as long as toothless; few discal setulae present below tooth; flinge blade-like setulae numerous, short and narrow. Mediostipes very broad, laterally textured and bearing numerous minute setae, a pore and relatively large cuticular cavity. Basistipes partly textured and bearing several minute setae, 3 apically barbed, robust, medium length primary setae, and 2 pores. Cardo untextured and armed with several minute setae. Palpifer with strong, imbricate texturing laterally; lateral disc generally armed with a few minute setae; ventroapical primary seta long, robust and apically barbed; dorsolateral surface armed with row of minute and short 94 setae; dorsal apex with one to several generally medium-length primary setae. Dorsal membrane with minute denticulate setulae basally, becoming progressively longer towards apex. Proximal palpal segment untextured and short. Second palpal segment about as long as wide and with some poximal texture; typically armed ventroapically with 3 medium-length setae and ventrally with a single seta. Third palpal segment untextured and usually armed with short lateral and apical primary setae; pores present or absent. Terminal palpal segment untextured and armed with vestigial setae only; palpal organ restricted to basal 1/2 of segment; several pores and sensilla digitiformia located at apex. Labium (Figs. 45D-F). Mentum (Fig. 45D) trapezoidal and commonly with 6 long robust barbed setae, several medium-length setae, and numerous short and minute setae located on ventral surface. Prementum (Fig. 45E-F) with lobes of paraglossae narrow, very short, and divergent; appendix lacking; small basolateral lobule present on paraglossae; ligular fringe consisting of medium-length setulae; weakly projecting; ligular disc setae of medium length, robust, apically blunt, and commonly 3 in number; several pores located at glossal base mesally. Palpiger commonly with single pore on dorsal surface. Proximal palpal segment short, slightly wider than long. Second palpal segment with primary setae medium-length and barbed; several pores commonly present. Terminal palpal segment short, oval, and bearing a few vestigial setae; palpal organ basal; several pores and sensilla digitiformia at apex. Substructure with dorsomedial sclerite wedge-shaped. Hyaline section of central arms slender and elongate; pigmented section long and cylindrical. Medial sclerite absent. Median lobe slender and fairly elongate. Legs very short; about 1/ 3 length of entire substructure; mesal border arcuate, leg bases mesally expanded. Thorax. Prothorax. Notum with bead and at least pronotal costa 1; costae 2 & 3 present in some taxa; disc, bead and costae usually with relatively large, erect amber scales; sternum with anterior margin of lobe straight or feebly outwardly arcuate; carinal stria present and carinate. Fore-tibia apically expanded and armed anteriorly with relatively 95 long, sharp teeth. Elytra with at least 4 costae; costa generally armed with large, erect amber scales; intervals generally with rows of deep punctures. Hind wings (Fig. 42C). R4 and RP separate; RA3+4 stigma elongate more than 7x longer than wide; RA3 absent; RA4 and RPl light but distinct; RP fused with MAl+2 along most of its length; RP relatively short; RP2 complete; RP3+4 apically abbreviated, ending as vein about 1/2 way to wing margin and extending to wing margin as sclerotized membrane; Mp3+4 distinct and apically abbreviated, ending well short of wing margin; CUA and AAl+2 joining at end of jugal lobe; CU2+3+4 short; AA3+4 long, nearly reaching wing margin; AP3+4 absent; J indicated basally; pigmented patterns relatively light but distinct. Meso- and metastema with fossulate fovea; fovea varying in shape; median sulcus sometimes present on metastemum; meso-metastemal suture and sections of metastermal longitudinal suture generally indicated; surface (including legs) commonly scaly. Abdomen. Stemites. First apparent stemite usually with lateral fossulate foveae. Posterior margin of remaining abdominal stemites punctate. Propygidium and pygidium punctate, typically carinate, and scaly. Male Terminalia. (not figured) Eighth segment. Tergite dorsum about as long as broad and deeply emarginate proximally; remaining disc medially hyaline giving tergite subdivided appearance; tergite produced lateroposteriorly into rounded tails on each side; tergite generally with several cuticular cavities dorsoapically. Coxites about as long as tergite and somewhat dorsoventrally flattened; coxite ventral surface largely membranous and armed distally with 1 to several short, stout setae; other minute setae may be present on dorsoapical and dorsolateral surfaces; cuticular cavities present dorsoapically and dorsolaterally; apical pubescent membrane present. Ninth segment non-capsular and travois shaped. Tergite subdivided into 2 tailed unarmed valves commonly with numerous anterior cuticular cavities. Coxal carapace short and broad, longitudinal suture lacking and typically armed distally with 2-4 short conspicuous setae; apical pubescent membrane lacking. Spiculum gastrale usually broad posteriorly and narrowing anteriorly and 96 expanding into an apical crescent partially encircling the aedeagus. Aedeagus short, stout, and generally with an asymmetrical phallobase (phallobase reduced and fused to parameres in some taxa); parameres united distally and variably armed apically; commonly with pore­ like cuticular cavities. Diagnosis: The costate elytra, fossulate fovea on the meso-, meta- and first abdominal sternum, cuticular shagreening, and erect amber scales will distinguish adult Epiechinus firom all other genera of histerids. Distribution. Over 60% of described species of^/ecA W f are Afi^otropical. The remaining species are distributed in the Oriental, Australasian, and Palearctic zoogeographic regions. Immature Stages. Immature stages oîEpiechinus are unknown. Biology/Ecology. Adults oî Epiechinus are microphagus and most taxa appear to be fungal spore specialists. Label data indicates that most Epiechinus are associated with rotting wood or forest litter. Thérond (1973) mentioned the association oîE. desbordesi Burgeon with recently fallen tree branches. Kanaar (1992) reported the same species associated with defunct termite nests. The gut contents of a specimen o î Epiechinus rcqppi Bickhardt fi’om the Shimba Hills, Kenya, contained spores of the hyphomycete Spadicodes obovata (Cooke & Ellis) and a coelomycete (Diplodia or Botryodiplodia). The sternal depressions, characteristic of most Epiechinus, may be mycangia. Some of the cavities on the bodies of Scolytini used for fimgal transport have associated glands producing an oily secretion to hold spores (Crowson 1981). Most sternal fovea and the elytra bear deep cuticular pores which secrete substances that keep the beetle encrusted with debris. This debris may contain and assist in the transport of spores of protists or fungi. One Afi'otropical species oî Epiechinus is myrmecophilious. Epiechinus kivuensis Therond inhabits the nests oîMyrmicaria sp.. 97

Epierus species group The Epierus genus group is an informal group of related genera including Australepierus, new genus, Plagiogramma Tarsia., and EpierusEnchson. Adult Description: Head. Frons with verticofrontal carinae lacking. Antennae shorter than width of head capsule. Scape variable and armed with minute setae. Club with sensilla basiconica distribution patchy or linear. Epipharynx with lateral fiinge basally abbreviated; inner brush complete or incomplete and bordered distally by comb. Mandibles with contact chemoreceptors present or absent. Medial area somewhat broad apically. Microtrichia of dorsal microtrichial field either well-developed or somewhat reduced. Prostheca detached apically; setulae generally short but occasionally more elongate distally; inner prosthecal setulae variably reduced and modified. Maxilla. Galea with an obtuse or acute mesal extension and variably armed. Lacinial strobilus present or absent; lacinial fiinge sometimes basally abbreviated . Palpifer carina apically abbreviated or absent. Labium. Mentum with anterior margin outwardly arcuate or unidentate. Palpiger with an elongate or reduced basal extension. Thorax. Prothorax. Notum generally lacking costae; sternum with anterior margin of prosternai lobe straight or produced; carinal stria present or absent. Fore-tibia lacking teeth. Elytra usually stiate and rarely costate. Hind wings with wing vein RP2 apically abbreviated. Meso- Metastemal surfaces generally finely punctate but rarely coarsely punctate. Abdomen. Propygidium and pygidium generally finely punctate but rarely coarsely punctate and costate. Diagnosis: Members o î Epierus species group can be distinguished fi’om those of Onthophilius species group by the following characters: verticofrontal carina lacking; epipharyngeal lateral fiinge reduced; mandibular medial area apically broad; galea extending mesally; palpifer carina apically abbreviated or absent; anterior margin of fore­ tibia lacking teeth; hind wings with wing vein RP2 apically abbreviated. 98 Genus Australepierus, new genus. Syn.: Parepierus Bickhardt (in part) (Figs. 14B, 17A-B, 19C-D, 20D, 23C-D, 37B, 38C, 42E, 46).

T ype Species: Epierus^imllus"&xo\xa. Adult description: Body outline (dorsal) (Fig. 14B) oval to circular. Head. Frons with short laterofrontal carinae. Antenna (Fig. 17A-B) shorter than width of head capsule; scape morphology variable; generally armed with uniform covering of minute setae. Club somewhat flattened dorsoventrally and densely covered with sensilla trichodea; sensilla basaconica distribution irregular; elongate, regularly spaced sensilla chaetica originating near former antennal divisions and towards antennal apex; all vestiges of antennal sulci lacking. Epipharynx (Fig. 19C-D) generally broadly rounded in outline; marginal setal fringe confined to dorsal and upper mesal margin of epipharynx. Lateral area with outer brush remnant near dorsal margin; most scales on lateral area reduced and denticulate; some larger scales present mesally and basally. Inner brush complete and composed of relatively short setulae. Medial area with one conspicuous and two less conspicuous peg setae on each side; medial area sensilla numerous, relatively large, and uniformly distributed; dorsomedial brush setulae elongate; brush centrally dense with reduced, recumbent setulae extending laterad; dorsomedial peg setae short and less than 10 in number. Mandibles (Figs. 20D, 38C). Dorsolateral area weakly expanded basally; disc and armed with minute setae and 3 short, robust apical contact chemoreceptors. Dorsomedial area broad; articulated setose appendage densely armed with minute setulae. Microtrichia (Fig. 37B) well-developed; ventral microtrichial field ending short of mandibular base; prostheca slightly detached apically and with some dorsal setulae barbed, ventral setulae (Fig. 38C) arranged in transverse rows progressively decreasing in length. Maxilla (Figs. 23C-D, 46A-C) with galea (Figs. 23C-D, 46A-B) transversely expanded; ventral surface with variable covering of short setulae; apical margin flinged with short 99 setulae; apical 1/2 of dorsal surface commonly with concentric transverse rows of short narrowly spaced setulae (Fig. 23C-D); at least one species with some spatulate setulae on dorsal surface; cluster of short, stiff setulae present on dorsomesal margin. Lacinia (Fig. 46A-B) with dorsal imbricate texture and some ventral rugosity; strobilus terminating in several blade-like setulae; lacinial tooth (Fig. 46C) with basal portion more than twice as long as toothless; some discal setulae directly below tooth; ffinge setulae short and slender; blade-like setulae numerous, short and slender. Mediostipes generally very broad and bearing many, mainly minute setae, a pore, and several small internal cuticular cavities. Basistipes untextured, commonly bearing at least three short-medium-length primary setae and 2 pores. Cardo untextured and armed with some minute setae. Palpifer untextured; lateral disc bearing a pore and some short to medium-length setae; ventroapical primary seta long and robust; dorsolateral disc with cluster of minute-short setae basally; carina apically abbreviated; dorsal apex with one or more generally medium-length dorsoapical primary setae. Dorsal membrane basally denticulate. Proximal palpal segment untextured and relatively long, slightly longer than wide. Second palpal segment long, about twice as long as wide, untextured, and generally bearing 2 ventroapical pores; armed laterally and ventroapically with short setae. Third palpal segment sometimes with vestigial basal texture and armed laterally with several minute setae; pores present or absent. Terminal palpal segment sometimes with faint basal texturing; palpal organ restricted to basal 1/2 of segment; armed with numerous mostly vestigial setae; several pores and sensilla digitiformia at apex. Labium (Fig. 46D-F). Mentum (Fig. 46D) trapezoidal with 6-8 long robust setae and numerous short and minute setae on ventral surface. Prementum (Fig. 46E-F) with lobes of paraglossae often strongly projecting, very narrow, and without appendix; ligular fiinge consisting of short slender setulae which may extend obliquely behind glossae; glossal lobes (when present) narrowly acuminate, strongly projecting, and apically tufted with medium-length setulae; ligular disc setae generally slender, medium- 100

length, and 3-6 in number; several pores at glossal base mesally. Palpiger with single pore on dorsal surface. Proximal palpal segment slightly longer than wide. Second palpal segment primary setae short; pores present or absent. Terminal palpal segment elongate, ovoid; and armed with a few vestigial setae; palpal organ situated basally; several pores and sensilla digitiformia at apex. Substructure with dorsomedial sclerite roughly triangular. Hyaline section of central arms short and robust; pigmented section short and generally cylindrical. Medial sclerite present or absent. Alae elongate, strongly elevated and open ventromesally. Median lobe reduced and acuminate; substructure body with lateral lobe on each side; legs usually relatively short; about 1/2 entire length of substructure; mesal border arcuate, leg base only slightly expanded and with short, dorsolateral projections. Thorax. Prothorax. Notum lacking bead; pronotal costae 3 present in A. australis; disc variably punctate; sternum with anterior margin of prostemd lobe straight; carinal striae present. Fore-tibia usually slender, only slightly expanded distally, and anteriorly armed with short spines. Elytra generally with at least 4 complete outer dorsal striae and variably punctate or sometimes costate. Hind wings (Fig. 42E) with R4 and RP separate; RA3+4 stigma relatively short, ca. 2x longer than wide; RA3 absent; RA4 dark and distinct; RPl faint and indistinct; RP fused with MAl+2 along most of its length; RP relatively short; RP2 apically abbreviated, ending as vein about midway to wing margin, continuing to wing margin as sclerotized membrane; RP3+4 apically abbreviated, ending as vein about half way to wing margin and extending to wing margin as sclerotized membrane; Mp3+4 distinct and apically abbreviated, ending well short of wing margin; CUA and AAl+2 joining slightly beyond at end of jugal lobe; beyond juncture sclerotized membrane extends nearly to wing margin; AP3+4 absent; J indicated and relatively elongate; pigmented patterns relatively light but distinct. Meso- and metastema with marginal metastemal and meso-metastemal suture; metastemal longitudinal suture typically present; surface punctate coarsely so in .<4. australis. 101 Abdomen. Stemites variably punctate. Propygidium and pygidium usually unsculptured and variably punctate but occasionally carinate and coarsely punctate. Male Terminalia (not figured). Eighth segment capsular. Tergite dorsum entire, commonly obtusely emaiginate proximally; bearing numerous cuticular cavities dorsolaterally. Coxites relatively large, conical and armed with single short setae towards apex; numerous minute pore-like cuticular cavities present ventrally and mesally; apical pubescent membrane present. Ninth segment distinctly capsular. Tergite subdivided into 2 laterally broad valves armed dorsoapically and ventroapically with minute setae. Coxal carapace elongate, ca. 3x longer than wide and with distinct longitudinal suture; armed distally with several short and minute setae; pore-like cuticular cavities present distolaterally; apical pubescent membrane present. Spiculum gastrale elongate, relatively broad, somewhat expanded distally and posteriorly bisected for about 1/2 of length of sclerite. Aedeagus typically with an elongate tubular phalobase; parameres divided distally and armed with minute-short setae on ventroapical surface. Diagnosis: The following combination of characters will distinguish adult Australepierus firom other histerid genera: antennal club with sensilla basiconica distribution irregular; epipharynx with marginal setal fiinge confined to dorsal and upper mesal margin of lateral area; galea with acute mesal extension; ventral galeal surface variably covered with short flexible setae; dorsoapical galeal surface typically bearing concentric rows of short setulae; strobilus ending in blade-like setulae; mediostipes often very broad; first maxillary palpal segment longer than wide; second maxillary palpal segment more than twice as long as wide; prementum with lobes of paraglossae usually extremely short and narrow; glossal lobes generally short, narrowly acuminate, and apically tufted with setulae; labial substructure legs commonly relatively short. Distribution. Australepierus is restricted to New Zealand and Australia. 102 Immature Stages. A. F. Newton of the Field Museum, Chicago provided me with larvae and a pupa found in association with adult A. punctulipemis (Broun) and also larvae found in association ynXh Australepierus phylobius (Broun) that he and M, Thayer collected in New Zealand. I am describing the larvae and pupae of^4. punciulipennis as part of a larger work on larvae of the Epierus genus group. Biology/Ecology. Adults o îAustralepierus are microphagous and apparently fungal spore specialists. The gut contents of specimens of/I. punctulipennis collected by Newton from beneath the bark of Nothofagus south of Turangi, New Zealand contained spores of several hyphomycete species including Exosporium fungorum (Fries), Dichotomophthora sp., Dendryphilopsis atra (Corda), Bactrodesmium transversianum (Peyrolel), Dreschlera sp., and Cladiosorium sp.. The most abundant spore type was Cladosporium. The structure of the dorsal surface of the galea of most Australepierus is unique within the subfamily. The concentric rows of short setulae may act as a harvesting device to gather mainly smaller spores such as those of Cladiosporium. In addition to living under bark, some species o î Australepierus have been extracted from organic debris and leaf litter. One species oî Australepierus, A. australis (Helava), was attracted to pitfall traps baited with decaying fungi and dung. The dorsoapical galeal surface setulation oîA.foederatus (Lewis) is différent from other species o î Australepierus. This species has apical spatulate setae resembling those found in Plagiogramma and Epierus and a basal remnant of the concentric transverse rows of short setulae typical of the genus.

Genus Plagiogramma Tarsia in Curia Syn.: Pseudepierus Casey (Figs. 14C, 17D, 18B, 22A, 24A-B, 25A-B, D, 47)

T ype Species: Plagiogramma brasiliense Tarsia in Curia 103 Adult redescription: Body outline (dorsal) circular to ovoid (Fig. 14C). Head. Frons with short latero-frontal carinae. Antenna (Figs. 17D) shorter than head capsule width; scape slender and expanded dorsoapically; surface generally textured and armed with minute setae. Club usually apically tapered and lacking external sulci; former antennal divisions indicated as dark bands encircling club with transmitted light; surface densely covered with sensilla trichodea; transverse whorl of sensilla basaconica completely encircling club at former distal division; some short transverse bands of sensilla basaconia arising at level of former proximal division; elongate, regularly spaced sensilla chaetica originating near former sulci and toward antennal apex. Epipharynx (Fig. 18B) trapezoidal in outline and slightly detached from inner surface of labrum distally. Marginal setal fringe subdivided; one portion confined to base of outer lateral areas; other portion apparently forming elaborate comb along medial margin of lateral area. Lateral area disc with outer brush remnant medioapically; most scales reduced and denticulate; some rounded near mesal margin. Inner brush short and incomplete; brush setulae relatively short. Medial area with two peg setae on each side; medial area sensilla numerous, minute, and evenly distributed; dorsomedial brush setulae narrow, only slightly expanded mesally and extend laterally as thin band; dorsomedial peg setae elongate and slender. Mandibles (Fig. 22A) sexually dimorphic in some taxa. Dorsolateral area moderately expanded basally; armed with vestigial setae only; apical contact chemoreceptors absent. Dorsomedial area apically accuminate; articulated setose appendage with minute pubescence. Microtrichia somewhat reduced; ventral microtrichial field not reaching mandibular base. Prostheca strongly detached apically; apical dorsal setulae longer and more robust than remainder; ventral prosthecal setulae forming radula. Maxilla (Fig. 47A-C) with galea (Fig. 47A-B) expanded and rounded; mesal margin bending ventrally and densely covered with tapered setulae; apical 1/2 of dorsal surface generally bearing elongate, spatulate setulae; basolateral margin of galea with distinct fiinge of tapered setulae. Lacinia dorsal outline 104 broadly oval; strobilus absent; lacinial tooth (Fig. 47C) with basal portion more than twice as long as toothless; single robust spine below tooth; fiinge typically extending posteriorly to about the middle of structure; blade-like setulae numerous, long and relatively broad. Mediostipes with some rugosity, and armed with a few short and several minute setae; structure also bearing several pores, and multiple, small, internal cuticular cavities. Basistipes textured and usually with 3 short to medium-length setae, several minute setae, and 2 pores. Cardo generally with some texturing and with a few short and minute setae. Palpifer with some lateral texturing; lateral disc generally bearing a few minute setae and pore; ventroapical primary seta robust and short to medium-length; dorsolateral surface with cluster of short setae basally, several short discal setae, several short primary setae and single minute peg seta at dorsal apex; minute denticulate setulae in membrane basally; proximal palpal segment untextured, very short and band-like, second palpal segment stout, about 1.5 times as wide as long, generally with some texture, and with short setae and generally 2 pores laterally; third palpal segment traces of imbricate patterning mesally and armed with minute lateral setae; terminal palpal segment sometimes with faint patterning basomesally, palpal organ either basal or extending apically; numerous minute and several short setae present; single pore on ventral surface basally; several pores apically. Labium (Fig. 47D-F). Mentum (Fig. 47D) trapezoidal, with dorsomedian projection and textured surface; generally 4 long robust plumose seta and several minute, short, and medium-length setae on ventral surface; lateral margins with minute setae. Prementum (Fig. 47E-F) with lobes of paraglossae short and moderately broad, divergent and with appendbc, ligular fiinge composed of medium-length robust, apically flared and barbed setulae, glossal lobes narrow and apically rounded; ligular disc setae variable; pair of distinctive long and extremely robust disc setulae present in some taxa. Palpiger with several pores on dorsal surface. Proximal palpal segment short. Second palpal segment with several apical pores and short or medium-length primary setae. Terminal palpal 105 segment more than twice as long as broad and armed with several short to medium-length setae; palpal organ situated basally; several pores and sensilla digitiformia at apex. Substructure with dorsomedial sclerite generally triangular. Hyaline section of central arms slender and elongate; pigmented section very long and cylindrical. Medial sclerite present. Alae moderately elongate and elevated and closed ventromesally; cupule emarginate. Median lobe short and broad. Substructure body bordering alae with small lateral lobe on each side; substructure legs long, about twice as long as body of substructure; mesal border acuminate, leg base broadly expanded and with moderately long lateral extensions with sclerotized apices. Thorax. Prothorax. Notum unsculptured; disc usually with sparse, fine punctation. Sternum with anterior margin of prosternai lobe produced; carinal striae present. Fore-tibia somewhat expanded and armed anteriorly with short spines. Elytra generally with sparse, fine punctation and a reduced number of striae; at least 3 complete outer dorsal striae typically present. Hind wings (Fig. 42B). Wing veins R4 and RP proximally united; RA3+4 stigma relatively short, ca. 2x longer than wide; RA3 faint and elongate, about as long as RA4; RA4 and RPl dark and distinct; RP fused with MAl+2 along ca. 1/2 its length; RP relatively short; RP2 apically abbreviated, ending as vein about half way to wing margin and continuing nearly to wing margin as faint sclerotized membrane; RP3+4 apically abbreviated, ending as vein slightly more than half way to wing margin and extending nearly to wing margin as sclerotized membrane (RP3+4 slerotized membrane occasionally very dark, lending appearance of completeness); Mp3+4 dark, distinct and complete; CUA2+3+4 on line with CuA and long, extending to wing margin; CUA and AAl+2 remaining discrete for their entire length; AP3+4 absent; J indicated as basal vestige; pigmented patterns relatively dark and distinct. Meso- and metastema typically with sparse, fine punctation; marginal metastemal and meso- metastemal sutures present (Fig. 25D); metastemal longitudinal suture evanescent. Abdomen. Stemites, propygidium and pygidium typically with sparse, fine punctation. 106 Male Terminalia (not figured) variable. Eighth segment with tergite typically reduced; proximal border may be thickly margined. Coxites often relatively large (in some taxa nearly twice as long as tergite), but occasionally reduced; typically armed with several minute seta mesally and one to several short apical setae; cuticular cavities typically present; apical pubescent membrane absent. Ninth segment loosely capsular or reduced. Tergite usually subdivided into 2 laterally broad valves bearing vestigial setae and cuticular cavities. Coxal carapace variably reduced; lacking longitudinal suture, cuticular cavities and setae; apical pubescent membrane absent. Spiculum gastrale elongate, and generally broad. Aedeagus with tubular phallobase nearly as long as or longer than parameres; parameres typically broad basally and tapering apically; distally divided and usually armed with minute-short setae mainly on ventroapical surface. Sexual Dimorphism. Plagiogramma is one of 2 onthophiline genera of that includes taxa exhibiting pronounced sexual dimorphism. Male Plagiogramma have either a setigerous tubercle (Fig. 25 A) or a setigerous patch (Fig. 25B) in middle of anterior margin of clypeus. In some taxa the mandibles may be asymmetrically developed with the dorsolateral area of right mandible bearing a large, anteriorly projecting digitiform lobe. The head and body shape is may also be sexually dimorphic (Fig. 24A-B). The fi’ons in males of certain species otPlagiogramma is enlarged and the area above the antennal insertion project acutely. In at least one species, P. frontalis, dimorphic traits are more pronounced in some males than others. Diagnosis: The following combination of characters will distinguish adult Plagiogramma fi’om all other histerid genera: Males with setigerous tubercle or setigerous patch in middle of anterior margin of clypeus; antennal club usually acuminate apically and with transverse whorl of sensilla basiconica completely encircling club at level of former distal division; epipharynx slightly detached from inner surface of labrum; dorsal portion of m a ria i fringe of epipharynx modified into an elaborate comb along mesal 107 margin of lateral area; all mandibular setae vestigial; dorsoapical galeal surface with slender, elongate, usually spatulate setulae; lateral margin of galea with distinct fiinge of tapered setulae; lacinia oval in outline dorsally and broadly attached to mediostipes ventrally; strobilus absent; lacinea with single robust spine; fiinge generally extending posteriorly to about the middle of lacinia; palpifer ventroapical primary seta robust and short to medium-length; dorsoapical primary setae several in number and short; apical margin of mentum unidentate; ligular fiinge setulae distally expanded and apically barbed; terminal segment of labial palp with numerous short setae; leg base with moderately long lateral extensions; meso-metastemal stria present in addition to marginal metastemal stria. Distribution. Over 90% of the described species oî Plagiogramma are distributed in the Neotropics. Two species occur in the Nearctic and one in the Palearctic. Immature Stages. I have reared two species o f Plagiogramma, P. subtropica (Casey) and P. puhifrom (Hinton). I am describing the larvae and pupae of the former species. I extracted a single larva of P. puhifrons fi-om a pile of Atta detritus in Chiapas, Mexico in July. Larvae of an unidentified species of Plagiogramma was extracted fi'om litter under an old tree on Barro Colorado Island in Panama by A. Newton in February. Biology/Ecology. Adults o fPlagiogramma are microphagous and fungal spore specialists. Some species are leaf litter inhabitants while others inhabit cavities in soft rotting wood. Adults of some wood inhabiting species are noctumally active. I have observed adult Plagiogramma grazing on fungi on the surface of deadfall in Chiapas, and P. Skelley recently collected several species of this genus on log surfaces at night in Loreto Province, Peru. One species of Plagiogramma, P. puhifrons, is apparently restricted to accumulated detritus produced by leafcutter antSi4//a sp. (Hymenoptera: Formicidae). The gut contents of P. subtropica extracted fi’om cypress litter in Northern Florida yielded unidentifiable organic material and spores of the hyphomycete species Bactrodesmium cerdricola Ellis, Spadicoides grovei Ellis, and spores of the coelomycete species 108 Coryneum casteneicola Berkeley & Curtiss. Also found were protist endospores. Gut contents of the litter-inhabiting Plagiogramma gentilis from Lawrence County, Arkansas contained spores of a discomycete {Sclerotinum sp.) as well as those of a species of hyphomycete (Dendryphiopsis sp). The gut contents of another specimen of P. gentilis collected in Montgomery County, Maryland contained unidentifiable organic material and spores of the hyphomycete species Coniosporium olivaceum Link ex. Fries, B. transversianum, and an undetermined Cardona. Protist endospores and spores of the hyphomycete species S. bina (Corda) were found in the gut of P. frontalis, a rotting wood inhabiting species collected in Aragua, Venezuela.

Genus Epierus Erichson (Figs. 14D, 17C, 18C-D, 22B-D, 23A-B, 25C, 37C, 38D, 48).

T ype species: Histerfitlvicomis Fabricius. Adult redescription: Body outline (dorsal) ovoid (Fig. 14D); some species dorsoventrally flattened. Head. Frons with short latero-frontal carinae; males of sexually dimorphic species with setigerous tubercle in middle of anterior margin of clypeus Antenna (Fig. 17C). Scape slender, dorsally expanded and with strong surface texture and minute setae. Club somewhat flattened dorsoventrally, apically truncate and lacking external sutures; former antennal divisions superficially absent but indicated with transmitted light; surface very densely covered with sensilla trichodea; transverse whorl of sensilla basaconica completely encircling club at level of former apical division; some short transverse bands of sensilla basaconia arising at level of proximal sulcus; elongate, regularly spaced sensilla chaetica originating near former sulci and towards antennal apex. Epipharynx (Fig. 18C-D). Trapezoidal in outline and largely detached from the inner surface of labrum. Portion of marginal setal flinge confined to basolateral margin of lateral areas; another portion apparently modified into an elaborate comb along mesal margin of 109 lateral area; outer brush apparently lacking; most scales reduced and denticulate. Inner brush short, covering basal 1/3 of epipharynx basally; brush setulae relatively short. Medial area with two peg setae on each side and numerous minute sensilla; dorsomedial brush narrow through out and with short setulae; dorsomedial peg setae elongate and slender. Mandibles (Figs. 22B-D, 37C, 38D). Dorsolateral area moderately expanded basally; setae vestigial apically and minute basally, apical contact chemoreceptors absent. Dorsomedial area broad; articulated setose appendage with minute pubescence. Microtrichia (Fig. 37C) somewhat reduced; ventral microtrichial field not reaching mandibular base. Prostheca strongly detached apically; apical dorsal setulae longer and more robust than remainder; ventral prosthecal setulae forming radula (Fig. 38D). Maxilla (Figs. 23A-B, 48A-C) with galea (Figs. 23A-B, 48A-B) expanded and apically rounded; mesal margin produced, ventrally bent and densely covered with spatulate setulae; apical 1/2 of dorsal surface with elongate spatulate setulae; lateral margin of galea devoid of setulae. Lacinia narrowly rectangular in dorsal outline and broadly attached to mediostipes; strobilus absent; lacinial tooth (Fig. 48C) with basal portion more than twice as long as toothless; single robust lacineal spine below tooth; fiinge with blade-like setulae numerous, long, slender, and concentrated apically; fiinge very abbreviated, not extending below level of insertion of lacinial tooth. Mediostipes partly ruge and typically bearing a few short and minute setae, a single pore, and a few small internal cuticular cavities. Basistipes textured and commonly with 3 short to medium-length primary setae, several minute setae and 2 pores. Cardo generally with some texturing and with several short and minute setae. Palpifer with some lateral texturing; lateral disc generally with a few minute setae and single pore; ventroapical primary seta long and extremely robust; dorsolateral surface lacking basal setae and bearing several short discal setae, two long primary setae and single minute peg at dorsal apex; minute denticulate setulae in membrane basally; proximal palpal segment untextured very short and band-like, second palpal segment 110 stout, about 1.5 times as wide as long, generally with some texture, and with short setae and lateral pore; third palpal segment with some patterning and armed with short lateral setae; terminal palpal segment sometimes with faint patterning basally, palpal organ restricted to basal 1/2 of segment; numerous short setae present on dorsal surface; a few short lateroapical and numerous minute setae on ventral surface; single pore on ventral surface basally; several pores and very short sensilla digitiformia apically. Labium (Fig. 48D-F). Mentum (Fig. 48D) trapezoidal, with textured surface, and with dorsomedian and a pair of small dorsolateral projections; ventraal disc with several long robust plumose seta and several minute, short, and medium-length setae on ventral surface. Prementum (Fig. 48E-F) with lobes of paraglossae short and narrow, and with minute denticulate setulae on lateral margins and with appendbc; ligular fiinge composed of medium-length robust, apically flared and barbed setulae, glossal lobes weakly produced, broad and apically rounded; ligular disc setae commonly numbering 4, including long and extremely robust pair. Palpiger with several pores on dorsal surface. Proximal palpal segment short and band-like. Second palpal segment with short or medium-length setae apical primary setae and a pore. Terminal palpal segment long, more than twice as long as broad; dorsal and lateroventral surface vith many short to medium-length setae; palpal organ basal; several pores at apex. Substructure with dorsomedial sclerite wedge-shaped. Hyaline portion of central arms slender and elongate; sclerotized section short and cylindrical. Medial sclerite apparently absent. Alae short and weakly elevated. Median lobe short and broad. Body of substructure bordering alae un-lobed; legs long, about twice the entire length of substructure; mesal border acuminate, leg base somewhat mesally expanded. Thorax. Prothorax. Notum unsculptured; disc commonly with sparse, fine punctation. Sternum with prosternai lobe produced and occasionally with imbricate texture; carinal striae present. Fore-tihia somewhat expanded and armed anteriorly with short spines. Elytra generally with sparse, fine punctation and full set of complete dorsal striae. Hind Ill wings. R4 and RP separate; RA3+4 stigma relatively short, ca, 2x longer than wide; RA3 absent; RA4 and RPl dark and distinct; RP not fused with MAl+2; RP2 apically abbreviated, ending as vein about half way to wing margin and continuing nearly to wing margin as faint sclerotized membrane; RP3+4 apically abbreviated, ending as vein ca. 3/4 of the way to wing margin and extending nearly to wing margin as sclerotized membrane; Mp3+4 dark, distinct and complete; CUA2+3+4 on line with CuA; CUA and AAl+2 uniting beyond end of jugal lobe and forming a difiuse eye; AP3+4 absent; J indicated as basal vestige; pigmented patterns relatively dark and distinct. Meso- and melastema commonly with sparse, fine punctation; marginal metastemal suture present, meso- metastemal sutures absent (Fig. 2SC); metastemal longitudinal suture evanescent. Abdomen. Stemites, propygidium and pygidium commonly with sparse, fine punctation. Male Terminalia (not figured) somewhat variable. Eighth segment with tergite reduced and unarmed; dorsum occasionally emarginate and bisected; proximal border may be thickly margined. Coxites usually about as long as tergite, broad basally and apically rounded; often armed with setal rows; a single short apical seta commonly present; cuticular cavities often found laterally; apical pubescent membrane absent. Ninth segment reduced. Tergite commonly subdivided into 2 tailed unarmed valves with cuticular cavities anteriorly. Coxal carapace reduced and entirely membranous; apical pubescent membrane absent. Spiculum gastrale commonly elongate, relatively narrow, and parallel sided. Aedeagus with tubular phallobase commonly much longer than parameres. Parameres variable but often very short and divided apically. Sexual Dimorphism. Epierus is the other onthophiline genus with pronounced sexually dimorphic taxa. In relatively few species of Epierus, males possess a setigerous tubercle in middle of anterior margin of clypeus not found in females. As in Plagiogramma, some taxa have asymmetrically developed mandibles (Fig. 22C-D) with 112 the dorsolateral area of right mandible (Fig. 22C) bearing a large, anteriorly projecting digitiform lobe. Diagnosis: The following combination of characters will distinguish adult Epierus from other histerid genera; Frons in males of some taxa with setigerous tubercle; Antenna with transverse whorl of sensilla basaconica completely encircling club at level of former distal division; epipharyngeal inner brush apically bordered by setular comb; mandibular setae mostly vestigial; ventral surface of prostheca with radula; dorsoapical galeal surface with slender, elongate spatulate setulae; lateralobasal margin of galea lacking tapered setal fringe; strobilus absent; lacinial fringe basally abbreviated, not extending below level of insertion of lacinial tooth; palpifer ventroapical primary seta long and extremely robust; two long dorsoapical primary setae and single minute peg at dorsal apex of palpifer; mentum tridentate; ligular fringe setae distally expanded and apically barbed; pair of ligular disc setae long and extremely robust; terminal segment of labial palp with dense covering of short setae; meso-metastemal stria absent. Distribution. Over 60% of the described species o îEpierus are Neotropical. The remdning taxa are distributed in the Oriental, Nearctic, Australasian, and Palearctic zoogeographic regions. Immature Stages. The larva and pupa of E. lucidulus were described by Costa et al. (1988). I have reared five species o ïEpierus: E. divisus Marseul, E. incultus Marseul, E. pulicarius Erichson, E. regularis (Beauvois), and an undescribed species of Epierus from Mexico. I have extracted larvae of E. incultus from organic debris beneath oak bark from Chiapas, Mexico, in August and those of another from organic material beneath bark in Chiapas, Mexico in August. Larvae of E. pulicarius were extracted from organic material within an oak tree cavity in Ottawa County, Ohio in August and from organic debris within hollow beech stumps in Franklin County, Ohio during July-August. 113 Biology/Ecology. kàw\i Epierus are both macrophagous predators and microphagous spore specialists. R. Wenzel recorded E. pulicarius (Erichson) as a general predator and a mycophagist in Moser et al. (1971). Adults of most species appear to inhabit humic material that accumulates beneath the bark of alien tree trunks. They actively tunnel through this material creating a network of stable runways. I have extracted adult E. pulicarius from accumulated organic debris within a tree cavity. The gut contents of a specimen oîE. beccari from Banguay Island, Borneo contained spores of the following species of hyphomyceted: Bactrodesmium betulicola Ellis, and S. hina. The gut contents of a specimen oîE. regularis from Alachua County, Florida contained spores of the hyphomycete species Dictyosporium heptasporium (Garvaglio), Chalaropsis theilavioides Peyrolel and Cordana sp.. The gut of a specimen oîE. heterognathus from Chiapas, Mexico contained protist spores as well as those of the hyphomycete S. bina. Courtship: A rather peculiar form of courtship exists for Epierus. Prior to copulation, males stay in close contact with females by gathering in their mouthparts a highly elastic substance adhering to the female’s pygidium. Since there are no obvious glands or gland ducts associated with the female pygidium or propygidium, the source of this substance is likely the male. This material might be delivered through the frontal tubercle in those species that have it. During courtship the frons of the male generally remains in close contact with the female’s pygidium. When a gap between the pair develops, it is bridged by an elastic strand of this substance. The male is able to close the gap by gathering in this strand with his mouthparts. 114 IHbalinae Bickhardt, 1917

T ype G enus : Tribalus Erichson Included in the subfamily Tribalinae Bickhardt are the genera Tribalus Erichson, Stictostix Marseul and Peploglyptus LeConte, Idolia Lewis and the related genera Eutribalus Marseul, Parepierus Schmidt, and ParidoUa Kovarik, new genus. Redescription: Body outline (dorsal) commonly circular or ovoid. Head. Frons commonly with latero-frontal carinae only. Antenna with club completely fused, sulci reduced or absent. Labrum with setigerous punctures. Epipharynx with lateral area margin with regularly spaced fringe of marginal setae, lateral area disc generally with relatively large, broad, overlapping scales; vestiges of outer brush absent; inner brush variably developed. Medial area with dorsomedial brush present or absent. Mandibles with dorsolateral area generally armed with minute to short inconspicuous setae, several longer robust contact chemoreceptive setae may be present dorsoapically. Dorsomedial area with articulated setose appendage. Mola produced or retracted; microtrichia of dorsal microtrichial field either well-developed or atrophied. Maxilla with galea either relatively unmodified or modified for mycophagy. Lacinia with or without blade-like setulae. Proximal palpal segment short, band-like, and with 2 minute laterobasal primary setae and pore cluster in apical membrane. Penultimate palpal segment with a reduced number of primary setae. Terminal segment bearing palpal organ and commonly with apical sensilla digitiformia and pores. Labium with mentum bearing internal cuticular cavities near apical margin. Palpiger commonly with single minute primary seta and with a tapered or reduced basal extension. Proximal palpal segment short, band-like, and armed with single minute basolateral primary seta and with several pores in apical membrane. Penultimate palpal segment plesiomorphicly with 4 setae encircling apex. Terminal palpal segment with or without palpal organ. Ligular sclerites proximally convergent in most taxa. Thorax. Prothorax. Notum costate in some taxa; sternum with alae and with anterior margin of 115 lobe produced; lateral marginal prosternai striae present; lateral prosternai stria absent; carinal stria present or lacking. Fore-tibia lacking teeth. Elytra with or without costae. Hind wings with wing veins RP2 and RP3+4 apically abbreviated and wing vein AP3+4 missing; maculations reduced or absent. Meso-Metastemal surfaces variably punctate; marginal mesostemal suture and dorsolateral metastemal suture distinct; meso- metastemal, longitudinal metastemal, and lateral metastemal sutures present or absent. Abdomen. Propygidium and pygidium variably punctate and never costate. Diagnosis: Members of the Tribalinae can be distinguished from those of the Onthophilinae and other histerids by the following combination of characters: Antennal club completely fused with sulci vestigial or absent; labmm setose; epipharynx lateral areas with well developed, generally broad, overlapping scales; lacinia with strobilus lacking; third maxillary palpal segment with reduced number of primary setae; well developed prostemal alae present; hind wings generally with faint or no maculations.

Tribalus genus group The Tribalus genus group is an informal group of related genera including Tribalus Erichson, Stictostix Marseul and Peploglyptus LeConte. Adult description: Body outline (dorsal) circular to ovoid. Head. Antenna generally as long or longer than head capsule width. Scape occasionally with ventroapical tooth. Eighth antennal segment variable; club with or without vestigial sutures. Epipharynx trapezoidal in outline. Iimer bmsh well developed; dorsomedial brush present; medial area with relatively few sensilla restricted to area’s lateral margins. Mandibles with dorsolateral area uniformly broad, imbricately textured and armed with minute setae; apical contact chemoreceptive setae present or absent. Mola protracted or somewhat retracted; microtrichia of dorsal microtrichial field well-developed or extremely long, especially near mesal border. Prostheca setulae generally short. Maxilla with galea 116 relatively unmodified and commonly armed with robust tapered setulae. Lacinia with rugose dorsal surface texture; lacinial tooth short and stout; variable number of discal setae; fiinge variable; blade-like setulae present or absent. Mediostipes broad and bearing minute setae and single pore. Basistypes untextured, with 2 pores, and variably armed. Cardo untextured and variably armed. Palpifer untextured, variably armed, and bearing single lateroapical pore and complete carina. Dorsal membrane with variable armature. Second palpal segment stout; about as long as broad and variably armed. Third palpal segment armed only laterally. Terminal palpal segment variably armed and with elongate apical sensilla digitiformia; sensilla basiconica at palpal apex short. Labium with trapezoidal mentum; anterior margin strdght or tridentate. Prementum with ligular fiinge setae medium-length and tapered; ligular disc with at least 2 pores near setal bases. Palpiger bearing 1 or 2 pores and with long or short basal extension. Proximal palpal segment short and band-like. Second palpal segment commonly armed laterally with 1 or 2 short secondary setae and 4 short-medium-length apical primary setae. Terminal palpal segment variably armed and bearing several pores and short sensilla basaconica at apex. Substructure with dorsomedial sclerite wedge-shaped. Central arms variable. Medial sclerite present or absent. Ligular sclerites variably oriented. Alae closed anteriorly forming an emarginate cupule. Median lobe, body and legs of substructure variable. Thorax. Prothorax. Notum with or without costae and or raised bead; sternum with anterior margin of lobe produced; carinal striae striate or carinate. Hind wings. R4 and RP proximally united; RP relatively long; CUA2+3+4 in line with CUA and extremely faint and short, about as long as CUA; CUA and AAl+2 uniting beyond end of jugal lobe; AP3+4 absent; pigmented patterns extremely faint and indistinct. Meso- and metastemum with marginal metastemal, longitudinal metastemal, and dorsolateral metastemal stria present and distinct; meso-metastemal suture present or absent; lateral metastemal stria absent. Abdomen. Stemites. First apparent stemite with or without lateral abdominal 117 striae; remaining stemites variably punctate, Male Terminalia variable but with apical pubescent membrane of eighth and ninth segment absent and with coxites of abdominal segment Vm armed with relatively long spines. Diagnosis: Members of the Tribalus genus group can be distinguished from those of the Idolia genus group by the following combination of characters: Antennae usually as long as or longer than head capsule width; epipharyngeal dorsomedial brush present; epipharyngeal medial area with the relatively few sensilla present restricted to area’s lateral margins; mandibular dorsolateral area uniformly broad. Pigmented pattern of hind wings always extremely faint and with R4 and RP proximally united. Coxites of abdominal segment Vm armed with relatively long spines.

Genus Stictostix Marseul (Figs. 26A, 27B, 28B, 29B, 38A, 39B, 41A, 49)

T ype Species Epierusparra Marseul Adult Redescription: Body outline (dorsal) (Fig. 26A) oval to circular. Areas of cuticle including pronotal disc and thoracic stemites of one species, S. frontalis, with dense umbiculate punctures (Fig. 41 A). Head. Frons with relatively long laterofrontal carinae. Antenna (Fig. 27B) longer than head capsule width; scape elongate, angulate, and with or without ventroapical tooth; dorsal surface imbricately textured and uniformly armed with minute setae. Eighth antennal segment wider than long. Club somewhat flattened dorsoventrally and densely covered with sensilla trichodea; some short transverse bands of elongate sensilla basaconica originating from area of former proximal sulcus; transverse whorl of sensilla basaconica completely encircling club at area of former distal sulcus; prominent elongate sensilla chaetica originate from areas adjacent to former club division and towards antennal apex; former antennal divisions superficially absent but indicated in some taxa as incomplete dark bands on ventral surface with transmitted light. 118

Epipharynx (Fig. 28B). Broadly trapezoidal in outline. Outer margin of lateral area with tight fiinge of ca. IS regularly spaced elongate setae. Inner brush apically abbreviated and composed of relatively broad band of short, slender setulae; row of several minute sensilla bordering lateral area mesoapicaly. Lateral area with relatively large and apically rounded overlapping scales; outer brush absent. Dorsomedial brush setulae forming transverse, mesally expanded band of short, slender setulae; medial area with two prominent dorsomedial peg setae on each side. Mandibles (Figs. 29B, 3 8 A). Dorsolateral area generally with short robust apical contact chemoreceptors; apical 1/2 of dorsolateral area with several small, deep pits. Dorsomedial area broad; articulated setose appendage with short pubescence. Mola projecting strongly mesad; microtrichia well-developed; ventral microtrichial field approaching mandible base. Prostheca with dorsal setulae robust, and of medium-length distally, progressively decreasing in length proximally; some distal setae apically forked; ventral setulae (Fig. 3 8 A) loosely organized into several rows of mesally curved, stiff setulae, each row progressively decreasing in length. Maxilla (Figs. 39B, 49A-C). Galeal base sclerotized and with or without pores; ventromesal and ventroapical surface bearing robust, short to medium-length tapered setulae. Lacinia with lacinial tooth (Fig. 49C) moderately elongate and robust; with basal portion generally more than twice as long as toothlets; one or two discal setae present near tooth base; fiinge setulae relatively elongate; blade-like setulae relatively broad and elongate. Mediostipes partially rugose, and several small internal cuticular cavities. Basistipes armed with some short setae and three or less relatively long, robust primary setae. Cardo generally armed ventrally and laterally with several short and minute setae. Palpifer lateral disc commonly armed with numerous minute to short setae; ventoapical primary seta robust and medium- length; dorsolateral surface usually armed with row of minute-short setae along carina; dorsal apex with two long and generally one short primary setae. Dorsal membrane with cluster of short setulae basally. Second palpal segment with mainly ventral surface texture 119 and typically armed with minute and short setae laterally and bearing 1-2 pores ventrally. Third palpal segment with mainly dorsal surface texture and commonly armed laterally with single short seta and distally with maximum of 3 minute-short dorsoapically primary setae. Terminal palpal segment with mesobasal texture; palpal organ situated basally; individual sensilla digitiformia relatively elongate; armature variable and with several pores and elongate apical sensilla digitiformia. Labium (Fig. 49D-F). Mentum (Fig. 49D) anteriorly tridentate, with acute dorsomedial and blunt lateral projections; disc surface textured and with mixture of stout, medium-length, sometimes apically barbed setae and slender short and minute setae. Prementum (Fig. 49E-F) with lobes of paraglossae broad and with short, narrow appendix; ligular fringe consisting of long, robust, tapered setulae. Glossal lobes indistinct; ligular disc setae commonly numbering 3 per side; basal pair much shorter than remainder; with several pores near setal bases. Palpiger bearing single pore and elongate basal extension. Terminal palpal segment elongate and mesally and laterally armed mainly with short setae; palpal organ extending nearly to apex of segment; several apical pores present. Substructure with dorsomedial sclerite generally elongate and distally expanded. Hyaline section of central arms slender and elongate; pigmented section long and cylindrical. Medial sclerite usually present. Ligular sclerites distally convergent and slender. Alae relatively elongate and closed basomesally. Median lobe relatively elongate. Substructure body very short, much shorter than legs. Legs very elongate with mesal border arcuate. Leg bases narrow and with short lateral extensions. Thorax. Prothorax. Notum with an elevated bead surrounding disc; disc generally strongly punctate. Sternum with carinal striae striate. Fore-tibia elongate and slender; foretibia variably spinose. Elytra with elevated bead and with subhumeral and at least 2 dorsal costae; costae feebly elevated in some taxa. Hind wings. RA3+4 stigma relatively long, ca. Sx longer than wide; RA3 short; RA4 dark and distinct; RPl extremely light and indistinct; RP partially fused with MAl+2; RP2 apically abbreviated and consisting 120

entirely of sclerotized membrane ending about half way to wing margin; RP3+4 apically abbreviated, ending as vein ca. 2/3 of way to wing margin and extending beyond this for a short distance as sclerotized membrane; Mp3+4 extremely faint and short, about as long as CUA2+3+4; J faintly indicated, Meso- and metastema with surfaces strongly punctate and meso-metastemal stria present. Abdomen. Propygidium and pygidium typically densely and deeply punctate. Stemites. First apparent stemite with lateral abdominal striae; remaining stemites usually with strong punctures. Male Terminalia (not figured) Eighth segment commonly capsular. Tergite dorsum typically longer than broad and generally emarginate proximally and unarmed; dorsal disk sometimes bearing minute, pore-like cuticular cavities laterally. Coxites relatively long, conical and armed with minute, short and medium-length setae on dorsal lateral and ventral surfaces; coxal apex with or without setose styli; numerous cuticular cavities present mainly ventrally and mesally. Ninth segment reduced. Tergite subdivided into 2 variably shaped distally bilobed valves. Coxal carapace absent. Spiculum gastrale elongate, expanded posteriorly, narrowing for most of length then expanding laterally at apex. Aedeagus relatively elongate with tubular phallobase; parameres commonly parallel-sided and divided distally, ventrally bending and armed with minute-short setae on ventroapical surface. Diagnosis: The following combination of characters will distinguish adult Stictostix from other genera of Histeridae: Antennal scape shorter than combined length of flagellum and club; labrum /epipharynx relatively long; mandibles with apical contact chemoreceptors and several small pits; galeal setulae at most medium-length and much shorter than length of galea; mediostipes with multiple cuticular cavities; anterior border of mentum tridentate; glossal lobes of prementum lacking; palpiger with elongate basal extension; labial substructure central arms hyaline portion slender and elongate; pigmented section long and cylindrical; medial sclerite present; substructure body very short; legs very long; pronotum with an elevated bead surrounding disc; elytra with elevated bead and 121 with subhumeral and at least 2 dorsal costae; hind wing vein RP2 apically abbreviated; ending about half way to wing margin; male terminalia with eighth segment coxites relatively long and armed with at least some medium-length setae; ninth segment with coxal carapace absent; aedeagus with tubular phallobase and apically bifid and ventrally setose parameres. Distribution. Seventy-five percent of the described species of Stictostix occur in Australia. The remmnder occur in the western Nearctic. Biology and Ecology. Little biological information is available for Stictostix. The preferred microhabitat of some species appears to be flood debris or leaf litter along creeks and rivers, particularly in xeric areas. S. califomica has been collected at blacklight indicating that this species is nocturnal. While the Nearctic S. califomica is apparently sometimes found in association with ants, this symbiosis is probably not obligatory. Two MCZ specimens of S. califomica from Olympia, Washington were mounted with specimens of the m i Monica mutica (Emery). A specimen fi’om the CASC was mounted with an unidentified ant species. The Australian species S. ectatomae and S. biserata have been collected firom nests of the ant Ectatoma metallicum. Although the mouthparts of most Stictostix do not appear not to be modified for spore gathering, the galea of S. frontalis is equipped with spatulate setulae. The gut contents of a specimen of S. califomica collected in California contained unidentifiable organic material and spores of the hyphomycete species C. olivacium. At least 2 species of Stictostix, S. parra and S. califomica apparently have female biased sex ratios. Both of these species are associated with water courses mainly in xeric or seasonally dry areas. Immature Stages. The immature stages of Stictostix are unknown. 122 Genus Peploglyptus J. L. LeConte (Figs. 26B-C, 27C, 28C, 29C, 37A, 41B, 50)

T ype Species Peploglyptus belfragei J. L. LeConte. Adult Redescription: Body outline (dorsal) (Fig. 26B) almost rectangular. Cuticle shagreened (Fig. 4 IB). Head. Frons with relatively long laterofrontal carinae. Antenna (Fig. 27C) longer than width head capsule. Scape elongate, angulate and with ventroapical tooth; dorsal surface with imbricate textured and fairly uniform covering of minute setae. Eighth antennal segment about as long as vdde. Club apically acuminate and densely covered with relatively elongate sensilla trichodea; some short transverse bands of elongate sensilla basaconica originating from regions of former proximal sulcus; transverse and nearly complete whorl of sensilla basaconica encircling club at former distal division; prominent elongate sensilla chaetica arising from areas adjacent to former club division and towards antennal apex; all vestiges of former antennal divisions absent. Epipharynx (Fig. 28C). Trapezoidal in outline. Outer margin of lateral area with fringe of ca. 10 regularly spaced elongate setae. Inner brush apically abbreviated; setulae short and relatively robust; row of several minute sensilla bordering lateral area mesoapicaly; Lateral area with relatively large, well-developed overlapping scales; some scales with toothed margins; outer brush absent. Dorsomedial brush setulae flattened and apically bifurcate; forming thick mesal cluster. Medial area with two prominent dorsomedial peg setae on each side. Mandibles (Figs. 29C) Dorsolateral area with minute setae arising from minute, shallow pits; apical contact chemoreceptors absent. Medial area broad; articulated setose appendage with short pubescence. Mola extending strongly mesad; microtrichia (Fig. 37A) well-developed; ventral microtrichial field approaching mandible base. Prostheca with dorsal setulae robust, and of medium-length distally, progressively decreasing in length proximally; some distal setae apically forked; ventral setulae loosely organized into several rows of stiff, mesally curved, slender setulae; each row progressively decreasing in 123

length. Maxilla (Figs. 50A-C). Galeal base sclerotized and with one to several ventral pores; ventromesal and ventroapical surface with short to medium length robust tapered setulae. Lacinia with rugose surface texture dorsally; lacinial tooth (Fig. SOC) moderately elongate and robust; with basal portion about as long as toothlets; one discal setae present near tooth base; fiinge setulae short; blade-like setulae, relatively short and very broad. Mediostipes broad, with some surface rugosity, and several small internal cuticular cavities. Basistipes with some minute and three long robust primary setae. Cardo armed ventrally and ventrolaterally with mostly minute setae. Palpifer lateral disc armed with minute setae; ventoapical primary seta robust and medium-length; dorsalateral surface with cluster of mainly short setae basally; dorsal apex with at least one long and medium- length primary seta; peg-like sensillum also present. Dorsal membrane with patch of minute setulae. Second palpal segment with dorsal and ventral surface texture and with minute setae and single pore present ventrally. Third palpal segment with mainly basal surface texture and armed with single 2 short apical primary setae; single pore located ventrally. Terminal palpal segment mesobasally textured; palpal organ basal; ventroapically armed with a few vestigial setae and with single ventrolateral pore; apical 1/2 of ventral surface armed with several minute setae; palpal apex bearing several pores and very elongate sensilla digitiformia. Labium (Fig. 50D-F). Mentum (Fig. 50D) anteriorly tridentate with acute lateral and dorsomedial projections; surface of disc textured and with mixture of short to medium-length stout setae and slender minute setae. Prementum (Fig. 50E-F ) with paraglossal lobes and appendbc broad; ligular fiinge setulae long, robust and apically blunt. Glossal lobes absent; ligular disc setae medium-length and 2 per side; some pores present near setal bases. Palpiger with single pore and with elongate basal extension. Terminal palpal segment armed mesally and dorsally with short setae; palpal organ basal; palpal apex bearing several pores and numerous short sensilla basaconica. Substructure with dorsomedial sclerite wedge-shaped. Central arms with 124 hyaline section slender and elongate; pigmented section long and cylindrical. Medial sclerite present. Ligular sclerite parallel and relatively broad. Alae and median lobe relatively elongate. Substructure body short, slightly shorter than legs and with narrow lateral lobe on each side. Legs elongate with mesal border acuminate. Leg bases narrow and with short lateral extensions. Thorax. Prothorax. Notum with elevated bead surrounding disc and with pronotal costa 1 on each side; costae terminating distally into pair of trichome lined fossae (Fig. 26C ); disc with some umbiculate punctures (Fig. 4 IB) mesally and basally; bead with some minute erect scales. Sternum with carinal striae carinate and a pair of small trichome lined fossae encroaching on carina distally. Fore-tibia elongate; anterior margin somewhat expanded distally and variably spinose. Elytra with elevated bead and with subhumeral and at least 2 dorsal costae; intervals densely covered with relatively large umbiculate punctures (Fig. 4 IB) costae with minute erect scales. Hind wings. RA3+4 stigma long, ca. 3x longer than wide; RA3 absent; RA4 and RPl extremely faint and indistinct; RP partially fused with MAl+2; RP2 apically abbreviated and consisting entirely of sclerotized membrane ending about half way to wing margin; RP3+4 apically abbreviated, ending as vein ca. 1/2 of way to wing margin; Mp3+4 extremely faint and short, about as long as CUA2+3+4; J absent. Meso- and Metastema with large umbiculate punctures; meso-metastemal stria indistinct. Abdomen. Propygidium and pygidium with large umbiculate punctures. Stemites. First apparent stemite with lateral abdominal striae and umbiculate punctures; remaining stemites generally with umbiculate punctures. Male Terminalia (not figured) Eighth segment capsular. Tergite dorsum longer than broad and proximally emarginate and unarmed; dorsal disk bearing minute, pore-like cuticular cavities laterally. Coxites relatively long, conical and armed with minute, short and medium-length setae on dorsal lateral and ventral surfaces; coxal apex bearing veiy reduced setose styli; numerous cuticular cawties present mainly ventrally and mesally. Ninth segment reduced. Tergite subdivided into 2 125 relatively narrow and distally bilobed valves. Coxal carapace absent. Spiculum gastrale elongate, broad, and laterally expanding at distal apex. Aedeagus relatively elongate with tubular phallobase; parameres parallel-sided and divided distally, apex ventrally bent and armed with setal fiinge ventroapically. Diagnosis: The pronotal and prostemal trichome-lined fovea, shagreened cuticle and apically acuminate antennal club will distinguish adult Peploglyptus firom other genera of Histeridae. Distribution. Peploglyptus occurs in the Nearctic and Neotropical regions. Biology and Ecology. On the basis of label data, the preferred microhabitat for members of this genus appears to be flood debris. The trichome lined fossae on the pronotum and prostemum suggest an association with ants (Kanaar 1981). One specimen was collected firom a rotten log inhabited by ants in a creek bottom in Latimer Co., Oklahoma (K. Stephan, pers. comm). The maxillae are relatively unmodified and resemble those of its sister genus Stictostix. The gut contents of an undescribed species of Peploglyptus fi"om flood debris in San Luis Potosi, Mexico, were examined. They contained unidentified organic material and spores of the following hyphomycete species: D. heptasporium, C. casteneicola, and Asterosporium betulinum Peck. Immature Stages. The immature stages oî Peploglyptus are unknown.

Genus Tribalus Erichson (Figs. 26D, 27D, 28D, 29D, 32A, 38B, 39A, 42D, 51)

T ype Species: Hister capensis PaykuU. Designated by Bickhardt. Adult Redescription: Body outline (dorsal) (Fig. 26D) generally circular. Head. Frons with moderately long laterofi’ontal carinae. Antenna (Figs. 27D, 32A) about as long as or shorter than width head capsule. Scape variable but generally expanded dorsally; dorsal surface with imbricate texture and uniformly armed with minute setae. Eighth 126 antennal segment about as wide as long. Club (Fig. 32A) abruptly expanded proximally, distally truncate, and densely covered with sensilla trichodea; former antennal divisions superficially evident; transverse bands of relatively short sensilla basaconica originating fi'om proximal sulcus and almost completely encircling club; proximal sulcus shallow but with deep ventromesal sulcal pit (sp) concealing some sensilla basiconica; a second transverse nearly complete whorl of sensilla basaconica originating fi’om shallow distal sulcus; a third incomplete transverse row of sensilla basiconica located near antennal apex; prominent elongate sensilla chaetica originating immediately below basal and apical sutures and towards antennal apex. Epipharynx (Fig. 28D). Trapezoidal in outline and extremely broad and narrow. Outer margin of lateral area with tight fiinge of ca. 10 regularly spaced elongate setae. Inner brush setulae short and slender. Some minute sensilla bordering lateral area mesoapicaly. Lateral area with mixture of relatively large and smaller scales with those laterad apically acuminate and remainder apically rounded. Dorsomedial brush setulae forming transverse and mesally expanded band of short, slender setulae. Medial area with two minute dorsomedial peg setae on each side. Mandibles (Figs. 29D, 38B). Dorsolateral area with uniform covering of minute setae arising firom minute, shallow pits. Dorsomedial area broad and with articulated setose appendage densely covered with relatively long pubescence. Mola retracted and nearly even with inner mandible margin; microtrichia very elongate along mesal margin; moderately elongate elsewhere; ventral microtrichial field relatively narrow and extending to mandibular base. Prostheca with dorsal setulae short and slender, progressively increasing in length and thickness, forming relatively dense mat of setulae; ventral setulae loosely organized into several rows of very slender setulae, each row progressively decreasing in length. Maxilla (Figs. 39A, S1Â-C) Galeal base largely sclerotized and with lateral pores; ventromesal and ventroapical surface armed with long robust tapered setulae; dorsolateral margin of galea with 3 medium-length tapered setulae. Lacinia with some rugosity 127 dorsally; lacinial tooth (Fig. 51C) moderately elongate and robust; with basal portion ca. twice as long as toothlets; numerous setulae on lacinial disc partially obscuring tooth; fiinge setulae relatively elongate; blade-like setulae absent. Mediostipes broad, untextured and with single internal cuticular cavity. Basistipes with several short and three longer robust primary setae. Cardo armed ventrally and ventrolaterally with short and medium- length setae. Palpifer lateral disc armed with short setae; ventroapical primary seta medium-length and slender; dorsal surface with pores and mainly minute setae oriented along oblique carina; dorsal apex with 2 long primary setae and minute peg-like sensillum; dorsal membrane with small patch of short setulae. Second palpal segment with dorsal and ventral surface texture; bearing minute and short lateral setae and several ventral and dorsolateral pores. Third palpal segment with mainly mesal and dorsal surface texture; bearing minute lateral setae and pores. Terminal palpal segment mesobasally textured and laterally armed with minute setae; numerous widely spaced vestigial setae on dorsal and ventral surface; palpal organ basal; single ventrolateral pore present near palpal organ; palpal apex with several pores and very elongate sensilla digitiformia. Labium (Fig. 51D- E). Mentum (Fig. 5 ID) trapezoidal and with straight anterior margin; disc surface lacking texture and with mixture of stout, medium-length setae and slender minute setae. Prementum (Fig. 51E-F) with paraglossal lobes relatively narrow; appendix broad, bluntly rounded and not well differentiated; ligular fiinge consisting of medium-length to long, robust, tapered setulae. Glossal lobes short; ligular disc setae long, apically blunt and numbering 2 per side; two pores commonly present near setal bases; palpiger with 2-3 pores and with short, straight basal extension. Second palpal segment sometimes with dorsal surface texture. Terminal palpal segment of variable dimensions, with some basomesal texture, and generally 2 ventral pores; armed mesally, dorsally, and laterally with minute-medium-length robust, setae; palpal organ basal; palpal apex bearing several pores, and numerous short sensilla basaconica. Substructure with dorsomedial sclerite 128 elongate. Hyaline portion of central arms robust and relatively short; pigmented portion short, broad, and platelike. Medial sclerite absent. Ligular sclerites slender and posteriorly convergent. Alae relatively short and closed mesally. Median lobe short. Body of substructure elongate, about as long as legs. Legs relatively short, with mesal border rounded. Leg bases narrow and extremely elongate sclerotized lateral extensions. Thorax. Prothorax. Notum without costae; marginal stria abbreviated posteriorly margin slightly elevated anteriorly and anterolaterally; disc densely and finely punctate. Sternum with striate carinal striae. Fore-tibia short anteriorly expanded, and anteriorly armed with short spines. Elytra densely covered with fine punctures; sutural and internal subhumeral striae absent; apical remnants of dorsal striae 1-3 typically present; external subhumeral, marginal elytral and marginal epiplural stria complete. Hind wings (Fig. 42D). RA3+4 stigma relatively long, ca. 4x longer than wide; RA3 moderately long, about a third as long as RA4; RA4 dark and distinct; RPl light but distinct; RP not fused with MAl+2; RP2 apically abbreviated, ending about halfway to wing margin; RP3+4 apically abbreviated, ending as vein ca. 2/3 of way to wing margin and extending to wing margin as sclerotized membrane; Mp3+4 extremely faint and very short, about as long as CUA; J faintly indicated. Meso- and metastema with surfaces generally densely punctate; meso- metastemal stria present. Abdomen. Propygidium and pygidium generally densely and finely punctate. Stemites. First apparent stemite lacking lateral abdominal striae; remaining stemites punctate along posterior margin. Male Terminalia (not figured) Eighth segment reduced. Tergite dorsum largely hyaline, wider than long, and proximally tailed; dorsal disk with some lateral cuticular cavities. Coxites short and very broad; armed with mixture of minute, short, and medium length setae on dorsal lateral and ventral surfaces; styli absent; numerous cuticular cavities present mainly ventrally and mesally. Ninth segment reduced. Tergite subdivided into 2 rather broad, distally bilobed valves. Coxal carapace longer than broad, medially hyaline, and bearing pore-like cuticular cavities 129 laterally. Spiculum gastrale elongate, narrow and distally expanded and bifurcate. Aedeagus generally with relatively elongate and asymmetrically tubular phallobase; parameres distally divided with ventroapical surface unarmed. Diagnosis: The following combination of characters will distinguish adult Tribalus from other histerid genera: Eighth antennal segment about as long as wide; club with former antennal divisions superficially evident; basal suture with deep ventromesal pit containing some sensilla basiconica; epipharynx extremely broad and narrow; epipharyngeal lateral area with mixture of large and small scales; mandibular mola nearly even with irmer mandible margin; dorsal microtrichia elongate, especially along mesal margin; prostheca dorsal setulae forming relatively dense mat; galea with long robust tapered setulae and dorsolateral margined with 3 medium-length tapered setulae; ventral lacinial disc densely armed with setulae; blade-like lacinial setulae apparently absent; mediostipes with single internal cuticular cavity; palpiger dorsal apex with 2 long primary setae and minute peg-like sensillum; anterior margin of mentum straight; prementum with short glossal lobes; palpiger with 2-3 pores and relatively short basal extension; substructure dorsomedial sclerite elongate; pigmented portion of substructure central arms short, broad, and platelike; substructure medial sclerite absent; substructure ligular sclerites slender and distally convergent; alae of substructure relatively short; body of substructure elongate; substructure legs relatively short with leg bases with elongate, sclerotized lateral extensions; pronotal and elytral costae lacking; hind wing vein RP2 apically abbreviated vein; lateral abdominal striae absent; male eighth tergite with dorsum largely hyaline and proximally tailed; ninth segment with coxal carapace present; aedeagus generally with asymmetrically tubular phalobase and unarmed parameres. Distribution. Tribalus occurs in the Palaearctic, Afrotropical, and Oriental regions. Biology and Ecology. Vienna (1980) listed several microhabitats where T. scaphidiformis and T. minimus are known to occur. They include beneath stones; in leaf 130 litter; including litter at the base of larch trees {Tamarix sp ); and under poplar bark (Populus sp.). The species is also common in sand dunes along the coast. Tribalus minimus is found in similar ecological settings (except for costal dunes). Both species have been found in association with numerous ant genera. Vienna (1993) included ecological information recorded on labels of the material he examined for his revision of the Afrotropical species of Wbalus. Several species were taken in traps baited with meat, excrement, or rotting fruit while others have been extracted from forest litter and compost. Specimens of T. foveolatus Vienna were found beneath stones, while those of T. subfasciatus opacinotus Vienna were collected from marshy ground near a stream. A few species were collected beneath bark and one was extracted from flood debris. Some species inhabit open grasslands and one species, T. cavemicola (Lewis) is cave-dwelling. T. kochi Thérond has been taken from a wide variety of ecological settings including ant nests. Prinz (1984) found T. capensis to be prevalent in semi-dry and dry bovine dung. He observed adults of this species feeding on small scarab larvae and small fly larvae associated with the dung. The gut contents of several species of Tribalus were examined. All contained unidentifiable organic material and no spores of any kind. Immature Stages. The immature stages of Tribalus are unknown.

Idolia genus group The Tribalus genus group is an informal group of related genera including Idolia Lewis, Eutribalus Marseul, Parepierus Schmidt, and ParidoUa, new genus. Description: Body outline (dorsal) generally circular. Head. Frons laterofrontal carinae length variable. Antenna not longer than width head capsule. Scape relatively short and dorsally expanded; surface strongly textured, occasionally pitted, and armed with minute setae. Flagellum with 8th segment about as long as wide. Club imbricately patterned, somewhat flattened dorsoventrally and lacking distal and proximal sulci. 131 Epipharynx with lateral area with relatively large, well-developed overlapping scales; outer brush absent. Inner brush setulae atrophied in some taxa. Dorsomedial brush absent. Dorsomedial peg setae present or absent. Mandibles. Dorsolateral area usually constricted basally and uniformly covered with minute or short setae and lacking contact chemoreceptors; apex uni- or bidentate. Dorsomedial area broad; articulated setose appendage either reduced or broadly flattened. Mola usually flush with inner mandibular margin and not extending mesad; microtrichia of microtrichial field atrophied (Fig. 37D). Prostheca with two distinct layers of dorsal setulae: outer layer elongate distally, extending beyond incisor; inner layer variable; ventral prosthecal setulae forming radula. Ventral microtrichial field extremely reduced (Fig. 35B). Maxilla with galea expanded and apically truncate; surface scaled or setose in some taxa; armed dorsoapically with short tapered setulae proximally and spatulate setulae distally; doromesal galeal base with setulae cluster; dorsolateral galeal base generally bearing elongate setulae cluster. Lacinia with venose texture dorsally and ventrally rugose; detached fi'om and oriented nearly perpendicular to mesal margin of mediostipes. Mediostipes rugosely textured and with single pore and several generally minute setae. Basistipes variably textured, with two pores, and with maximum of 2 primary setae. Cardo commonly un-textured and with variable setation. Palpifer ventral surface textured ventrally and laterally and armed with medium-length ventroapical primary seta and minute secondary setae; dorsal surface with variably shaped condyle; dorsal apex with single long primary setae. Dorsal membrane setulae absent; second palpal segment stout, about as long as wide and laterally armed with short setae. Third palpal segment about as long as or slightly longer than wide; textured, and laterally armed with short setae. Terminal palpal segment elongate, dorsoventrally flattened, and armed laterally with short setae; palpal organ extending distally 2/3 length of palpal segment; distal 1/3 of segment with uniform covering of extremely minute setae; palpal apex with several pores and relatively elongate sensilla 132 basiconica and lacking sensilla digitiformia. Labium. Mentum trapezoidal and often basolaterally produced; anterior shelf present or lacking; lateral margins fiinged with mmnly minute setae; discal setation variable. Prementum paraglossal lobes with lateral surface texture; appendix present or lacking; glossal lobes distinct or lacking; ligular disc lacking pores. Palpiger with very short basal extension. Second palpal segment with variable setation. Terminal palpal segment (Fig. 40D) apically expanded and dorsoventrally flattened; mesally armed with stout, usually blunt setae; with numerous relatively long sensilla basiconica apically; palpal organ and apical sensilla digitiformia absent. Hyaline section of central arms generally unapparent; pigmented portion variable. Dorsomedial sclerite usually wedge-shaped with lateroapical projections. Medial sclerite absent; ligular sclerites slender and distally convergent; alae of substructure closed forming cupule. Median lobe of substructure present or absent; legs and leg bases variable. Thorax. Prothorax. Notum lacking costae; disc variably punctate. Sternum with anterior margin of lobe produced; carinal stria carinate or striate. Fore-tibia short, anteriorly expanded, and anteriorly armed with short spines. Elytra rarely with marginal bead; striation and punctation variable. Hind wings. BAl+2 reduced in size; RA3 dark, distinct and relatively elongate; RA4 dark and distinct; RP relatively short; AP3+4 absent; pigmented patterns but distinct. Meso- and metastema with marginal mesostemal mesally evanescent; meso-metastemal, longitudinal metastemal and lateral metastemal stria variable; dorsolateral metastemal stria present. Abdomen. Propygidium and pygdium variably punctate. Stemites. First apparent stemite with lateral abdominal striae; remaining stemites variably punctate along posterior margin. Male Terminalia. Eighth segment reduced. Tergite variable. Coxites short and very broad; and variably armed. Ninth segment reduced. Tergite subdivided into 2 very narrow distally bilobed valves. Coxal carapace variable. Spiculum gastrale rodlike. Aedeagus variable. 133 Diagnosis: Members of the Idoîia genus group can be distinguished from those of the Tribalus genus group by the following combination of characters: Epipharyngeal dorsomedial brush absent; epipharyngeal inner brush reduced or atrophied; mandibular lateral area basally constricted; mola retracted; dorsal microtrhichia atrophied; prostheca with setulae extending beyond mandibular teeth; mandibular radula present; spatulate setae of galea very thin; lacinia largely detached from mediostipes; palpiger with dorsal condyle; maxillary dorsal membrane unarmed; ventral surface of paraglossal lobe with surface texture; palpiger with short basal extension; terminal labial palpal segment distally expanded and dorsoventrally flattened and with relatively long sensilla basiconica.

Genus Eutribalus Bickhardt (n. stat.) (Figs. 30A, 31C, 32B, 33D, 34C, 35A-C, 35A, 39D, 40A, D, 42F, 52) Syn.: Tribalus Mais&x\ Caerostemus J. L. LeConte. New synonymy Tribalasia Cooman. New synonymy

T ype Species Tribalus agrestis Marseul. Adult redescription: Body outline (dorsal) (Fig. 30A) ovoid to circular. Head. Frons with short laterofrontal carinae. Antenna (Fig. 31C) Club securiform (Fig. 32B) and with surface densely covered with sensilla trichodea; sensilla basaconica distribution broad and irregular; sensilla chaetica short, rather unorganized and few in number. Epipharynx OFig. 33D) somewhat triangular in outline and apically truncate. Outer margin of lateral area with loose fringe of about 20 elongate and apically plumose setae; becoming abruptly short and spinelike along dorsal and dorsomesal margin of lateral area. Inner brush apically abbreviated and composed of vestigial setulae. Medial area generally with three minute peg setae on each side; surface densely covered with minute sensilla. Dorsomedial peg setae apparently absent. Mandibles (Figs. 34C, 35A-C) with dorsolateral area apically 134 unidentate and basally abbreviated; densely armed with minute, setae or shallow setigerous punctures. Mola flush with inner mandibular margin. Prostheca with some outer dorsal setulae (Fig. 35A) apically forked; inner prosthecal setulae (Fig. 35B) short, distally flattened, and apically bent apically; radular setulae (Fig. 35C) slender and with apex bending mesad at 90° angle; radular setulae arranged in densely and slightly curving rows. Maxilla (Figs. 39D, S2Â-C) with ventral galeal disc scaled and unarmed; ventroapical galeal margin fiinged with spatulate setulae; ventromesal m a r^ with fiinge of tapered setulae; dorsoapical galeal disc armed with narrow proximal band of short tapered setulae followed by band of elongate extremely thin spatulate setulae; distal dorsomesal margin armed with short tapered setulae; lateral margin with cluster of several elongate slender setulae apically. Lacinia narrowly rectangular in dorsal outline; lacinial tooth (Fig. S2C) veiy elongate and extremely slender; single slender spine present below tooth; fiinge setulae medium-length, slender, and usually confined to lateral margin; blade-like setulae lacking. Mediostipes armed with minute and short setae. Basistipes strongly, reticulately textured; armed with short apical setae and generally 1-2 medium length primary setae. Cardo with several short setae laterally. Palpifer strongly reticulate laterally; dorsolateral surface with broadly triangular condyle (Figs. 40A) and usually armed with minute setae. Second palpal segment with some texture basally and with several pores ventrolaterally. Third palpal segment strongly textured and commonly with two lateroapical pores on dorsal surface; terminal palpal segment strongly textured. Labium (Figs.). Mentum (Fig. 52D) with anterior border deeply emarginate and basolaterally produced; anterior shelf sometimes present and filling emargination; lateral and anterolateral margins fiinged with short and minute setae; disc untextured and generally uniformly covered with long flexible plumose setae. Prementum (Fig. S2E-F) with lobes of paraglossae broad, and armed laterally with minute denticulate setulae; appendix present or lacking; ligular fiinge consisting of long slender tapered setulae; glossal generally weakly produced; ligular disc 135 setae generally numbering 4 per side. Palpiger commonly with several pores on dorsal surface. Second palpal segment textured and armed mesally and laterally; mesal armarture consisting of cluster of several short and single medium-length setae. Terminal palpal segment with dorsal and ventral surface texture; disc variably setose; lateral margin armed with several short slender setae; several extremely minute setae near apex of segment. Substructure with hyaline portion of central arms unapparent; pigmented portion large and variably shaped. Median lobe short and broad. Legs generally long, about twice the entire length of substructure; mesal border acuminate; leg base large, somewhat quadrate and with short mesal extension. Thorax. Prothorax. Notum generally with complete marginal stria; disc variably punctate. Sternum; carinal stria carinate. Elytra unmargined in most taxa; striation and punctation variable; dorsal striae evanescent; marginal epipleural and marginal elytral striae generally present; humeral stria present or not. Hind wings (Fig. 42F). RA3+4 stigma elongate, about 3 x longer than wide; RPl light but distinct; RP not fused with MAl+2; RP2 apically abbreviated, extending ca. halfway to wing margin; RP3+4 apically abbreviated, ending as vein slightly less than half way to wing margin and extending to wing margin as relatively dark sclerotized membrane; Mp3+4 light and elongate; CUA2+3+4 short, in line with CUA and about half its length; CUA and AAl+2 joining beyond end of jugal lobe and forming eye; J only faintly indicated. Meso- and metastema with meso-metastemal stria (Fig. 36A) distinct and straight; longitudinal metastemal generally distinct; lateral metastemal stria abbreviated. Abdomen. Propygidium and pygidium generally with at least some relatively large punctures. Stemites generally with some relatively large punctures along posterior margin. Male Terminalia (not figured). Tergite of eighth segment variably shaped, lightly sclerotized, and bearing few cuticular cavities. Coxites short and very broad; and apically armed with several relatively long setae. Ninth segment reduced. Coxal carapace lightly sclerotized and armed apically with a few short and minute setae. Tergite subdivided into 2 very 136 narrow distally bilobed valves. Aedeagal phalobase generally short and fused to parameres. Parameres somewhat flattened and expanded laterally. Diagnosis: The following combination of characters will distinguish adult Eutribalus from all other histerid genera: Frons with short laterofrontal carinae; antennal club securiform with sensilla basiconica irregularly distributed; epipharyngeal fringe plumose; inner brush setulae vestigial; mandibles unidentate; galeal disc scaled and unarmed; lacinial fringe lacking blade-like setulae; palpifer condyle broadly triangular; anterior margin of mentum emarginate; disc nearly always with uniform covering of long flexible plumose setae; prementum with glossal lobes rarely produced; ligular setulae non- plumose; elytra lacking any complete dorsal stria; meso-metastemal stria distinct and straight; longitudinal metastemal stria generally distinct; lateral metastemal stria abbreviated. Distribution. Seventy percent of the described species oîEutribalus occur in the Oriental realm. Three species are Afrotropical, two Australasian, and one Nearctic. Biology and Ecology. Members of this genus are associated with decaying wood. The single new world species, Eutribalus americanus is known to associate with both decaying hardwoods and pines. This species also frequents lumber mill sawdust piles (R. Wenzel, pers. comm.). Eutribalus americanus is attracted to UV light and I have observed specimens active at night on fallen beech tree trunks. The adult mouthparts in members of this genus are specialized for gathering and processing spores. Spores from the gut of the Nearctic Eutribalus americanus and the Oriental E. marseuli were remarkably similar. The gut of a specimen oîE. americanus from Columbus, Ohio, contained the following species of hyphomycete spores: B. microleucorum Spegazzini, bina, Corynespora smithii (Berkeley & Broome). Also found were some protist spores. The hyphomycete spores species found in the gut of a specimen oîE. marseuli from Irian Jaya, Indonesia included B. traversianum (Peyrolel) and S. bina. 137 Immature Stages. I made several an attempts to rearÆ americanus in the laboratory and all were unsuccessful. However, I was able to collect larvae of this species from the field. Two of these larvae were reared to the pupal stage in the laboratory. These larva were found in relatively dry wood shavings (produced mainly by ants) in and around rotting beech stumps. The cuticle of Eutribalus larvae is remarkably transparent and the Malpighian tubules and hind gut appear white, probably due to accumulated uric acid. This condition seems to be indicative of larvae that inhabit relatively dry microhabitats, as it also occurs in sand dwelling saprinine larvae. I was not able to determine what E. americanus larvae were consuming in the field. Larvae m^tmned in the laboratory readily accepted house fly eggs. Like Idolia, pre-pupae construct sturdy, silk-lined circular cocoons. The larvae and pupae of E. americanus are described in the following chapter.

Genus Parepierus Marseul (Figs. 30D, 31 A, 33B, 34A, 36B, Syn.: Scqphidister Cooman Tribalus Maiseul 1855.

T ype S?Ecm Epierus amandus Schmidt 1892 Adult redescription: Body outline (dorsal) (Fig. 30D) generally circular. Body form generally dorsoventrally flattened. Head. Frons with short laterofrontal carinae. Antenna (Figs. 31 A) with sensilla basaconica irregularly distributed on securiform club; sensilla chaetica short, rather unorganized, and few in number. Epipharynx (Fig. 33B) somewhat triangular in outline. Outer margin of lateral area with fiinge of less than 10 elongate setae; becoming abruptly short and spinelike along dorsal margin of lateral area. Inner brush complete and composed of well-developed setulae. Medial area commonly with 2 small and 2 minute peg setae on each side and numerous minute sensilla. 138

Dorsomedial peg setae small and 2 in number. Mandibles (Figs. 34A). Dorsolateral area apically unidentate, basally constricted, and bearing punctures with minute but conspicuous setae. Mola of one species, P. amcmdus, is unusual in that it is mesally produced, whereas those of other species examined were not. Prostheca with inner dorsal setulae short, robust, and tapered; radular setulae relatively robust with only proximal setulae apically bent; distal setulae gradually curving mesad; setulae arranged in dense and slightly curving rows. Maxilla (Fig. 53 A-C) with ventral galeal disc variably scaled and unarmed; ventroapical galeal margin fringed with elongate slender setulae; ventromesal margin with commonly incomplete fiinge of tapered setulae; doroapical galeal disc with proximal band of short to medium-length tapered setulae followed by band of elongate, spatulate setulae; dorsomesal galeal margin armed distally with cluster of medium-length tapered setulae; lateroapical galeal margin with cluster of several elongate slender setulae. Lacinia narrowly elliptical in dorsal outline; lacinial tooth (Fig. 53C) short and slender; discal spines generally absent; fiinge setulae slender, medium-length proximally, short distally; blade-like setulae lacking. Mediostipes armed with several minute setae. Basistipes reticulately textured and generally 2 short primary setae. Cardo with several short and medium-length setae. Palpifer with strong ventral and lateral reticulate texture; palpifer condyle generally narrowly triangular. Second palpal segment textured ventrally and mesally and with several ventrolateral pores; lateral armature on this and remaining palpal segments consisting of short, slender setae. Third palpal segment with mesal and dorsal surface texturing. Terminal palpal segment with strong surface texture mesally; texture evanescent laterally; segment generally bearing several pores. Labium (Figs. 53D- E). Mentum (Fig. 53D) with anterior border deeply emarginate and basolaterally produced; typically with anterior shelf produced and largely filling emargination; lateral and anterolateral margins fiinged with short and minute setae; disc laterally textured and armed with mixture of minute setae, and short to medium-length robust setae. Prementum 139 (Fig. 53E-F) with paraglossal lobes relatively narrow and with appendix; ligular fiinge consisting of long, slender, tapered and apically plumose setulae. Glossal lobes strongly produced, broad and apically rounded; ligular disc setae commonly numbering 3 per side. Palpiger commonly with several pores on ventral surface. Second palpal segment with mesal and dorsal surface texture and variably armed. Terminal palpal segment with dorsal and mesal surface texture and generally several pores. Substructure with pigmented section of central arms broad, flattened and roughly triangular; median lobe absent; substructure legs long, about twice the entire length of substructure; mesal border somewhat; leg base moderately expanded. Thorax. Prothorax. Notum usually with marginal stria; disc generally finely punctate. Sternum with striate carinal striae. Elytra un-margined and finely punctate; striation variable but most species with some complete dorsal striae present; marginal epipleural and marginal elytral striae present; humeral stria present usually absent. Hnd wings. RA3+4 stigma short, less than 3 x longer than wide; RPl light but distinct; RP partially fused with MAl+2; RP2 apically abbreviated, extending ca. 2/3 of way to wing margin; RP3+4 apically abbreviated, ending as vein ca. half way to wing margin and extending to wing margin as sclerotized membrane; Mp3+4 absent; CUA2+3+4 absent; AAl+2 absent or vestigial; J indicated as distinct basal vestige. Meso- and metastema with meso-metastemal stria (Fig 36B) distinct and outwardly arcuate; longitudinal metastemal evanescent; lateral metastemal stria absent. Abdomen. Propygidium and pygidium finely punctate. Stemites with fine punctures along posterior margin. Male Terminalia. (not figured) Tergite of eighth segment variably shaped, lightly sclerotized, and bearing few cuticular cavities. Coxites short and very broad; and apically armed with several relatively long setae. Ninth segment reduced. Coxal carapace lightly sclerotized and armed apically with a few short and minute setae. Tergite subdivided into 2 very narrow distally bilobed valves. Aedeagal phalobase generally short and fused to parameres. Parameres somewhat flattened and expanded laterally. 140 Diagnosis: The following combination of characters will distinguish adult Parepierus from all other idoliini genera: Frons with short laterofrontal carinae; antennal club securiform and with sensilla basiconica irregularly distributed; epipharynx with lateral area fringe non-plumose setae; inner brush setulae well-developed; mandibles unidentate; dorsal galeal disc scaled and unarmed; lacinia fringe lacking blade-like setulae; anterior margin of mentum emarginate and with disc setae not flexible or plumose; prementum with distinct glossal lobes; ligular fringe setulae plumose; Mp3+4 and CUA2+3+4 of hind wing absent; meso metastemal stria outwardly arcuate; lateral metastemal stria absent. Distribution. Parepierus is restricted to the Oriental realm. Biology and Ecology. Members of this genus are associated with decaying wood. The gut contents of an undescribed species oî Parepierus collected under bark in Mindanao, Philippine Islands contained spores of an undetermined species of Endophragmia (Hyphomycetes). Cooman (1933) made some observations on live adults of Parepierus velox in Vietnam. When disturbed this species does not play dead as is common for members of the family. Instead it takes flight. Its local movement is rapid and interrupted by short sudden stops. Immature stages. The immature stages o î Parepierus are unknown.

Genus ParidoUaf new genus Syn. : Tribalus Marseul (Figs. 30C, 3 ID. 32D, 33C, 34D, 54)

T ype Species: Tribalus comes (Cooman) Adult description: Body outline (dorsal) (Fig. 30C) circular. Body form generally globose. Head. Frons with laterofrontal carinae very long, V-shaped, and meeting at clypeus. Antenna. (Fig. 3 ID) with club (Fig. 32D) apically rounded; surface sparsely covered with sensilla trichodea; elongate sensilla basiconica arising along lateral and mesal 141 margins and forming two transverse whorls completely encircling club; sensilla chaetica elongate, a few present along basomesal and basolateral margin; others forming whorl near former apical division and an apical tuft. Epipharynx (Fig. 33C) somewhat trapezoidal in outline and apically truncate. Outer margin of lateral area with loose fiinge of less than 10 short, apically plumose setae. Inner brush setulae well-developed. Mesal area generally with 2 small peg setae on each side; surface densely covered with minute sensilla. Two minute dorsomedial peg setae present. Mandibles (Fig. 34D) with dorso lateral area apically bidentate and basally abbreviated and armed with setigerous punctures; setae minute, basally and short apically. Prostheca with inner dorsal setulae short. Radula extremely narrow. Maxilla (Figs. 54A-C) with ventral galeal disc untextured and unarmed; ventromesal margin with nearly complete fiinge of progressively shortened, tapered setulae; ventroapical galeal margin generally fiinged with stiff apically truncate spatulate setulae; lateral margin with or without setulae cluster; dorsoapical setulae composition variable and often complex. Lacinia somewhat distally acuminate; lacinial tooth (Fig. S4C) robust and short; fiinge setulae medium-length and slender, and with extremely elongate apical blade-like setulae; Mediostipes armed with some short and minute setae. Basistipes vith reticulate surface texture and 2 medium-length primary setae. Cardo with several short setae laterally. Palpifer strongly reticulately textured ventrolaterally; palpifer condyle broadly rounded and concave. Second palpal segment with dorsal and ventrolaterally surface texture and bearing and single ventral pore. Third palpal segment with dorsal and ventrolateral surface texture and single ventral pore; dorsal surface of terminal palpal segment strongly textured and with several lateroventral pores. Labium (Figs. 54D-E). Mentum (Fig. 54D) with anterior border slightly emarginate and basolaterally produced; with short anterior shelf; lateral margins fiinged with minute setae; disc untextured and with a mixture of medium-length and short setae; medium-length setae branched in some taxa. Prementum (Fig. S4E-F) with lobes of paraglossae broad. 142

and fiinged laterally with minute setulae and bearing appendix; ligular fiinge consisting of a mixture of short mmnly lateral setulae and long slender, sometimes branched setulae. Glossal lobes weakly produced; ligular disc setae commonly numbering 2 per side. Palpiger generally with single pore on dorsal surface. Second palpal segment untextured and mesally armed with single medium-length seta and laterally with minute and short setae. Terminal palpal segment with some dorsal surface texture; disc armed with several short setae ventrally and dorsolaterally; lateral margin armed with robust medium-length setae. Substructure with hyaline portion of central arms unapparent; pigmented portion small and ovoid. Median lobe apparently absent. Legs relatively short, only slightly longer than body of substructure; mesal border acuminate; leg base only slightly expanded. Thorax. Prothorax. Notum with marginal stria; disc generally finely punctate. Sternum with carinal stria stiate. Elytra not margined and finely punctate; striation variable but generally with some complete dorsal stria present; marginal epipleural and marinai elytral striae present; humeral striae usually absent. Hind wings. Hind wings. R4 and RP proximally united; RA3+4 stigma short, less than 2 x longer than wide; RPl extremely faint and indistinct; RP partially fused with MAl+2; RP2 apically abbreviated, extending ca. 2/3 of way to wing margin; RP3+4 apically abbreviated, ending as vein slightly less than half way to wing margin and extending to wing margin as relatively light sclerotized membrane; Mp3+4 absent; CUA2+3+4 faint and short; CUA and AAl+2 joining beyond end of jugal lobe; J indicated as short, distinct basal piece. Meso-' and metastema with meso-metastemal stria present and generally outwardly arcuate; longitudinal metastemal absent; lateral metastemal stria present. Abdomen. Propygidium and pygidium finely punctate. Stemites with fine punctures along posterior margin. Male Terminalia (not figured). Tergite of eighth segment variably shaped, lightly sclerotized, and bearing cuticular cavities. Coxites short and apically armed with several short setae. Ninth segment variably reduced. Aedeagal phalobase moderately long. Parameres variable. 143 Diagnosis: The following combination of characters will distinguish dÀvXiParidolia from all other histerid genera: Body form globose; laterofrontal carinae very long, V- shaped and meeting at clypeus; antenna club apically rounded and sparsely covered with sensilla trichodea; sensilla basiconica elongate and forming two transverse whorls encircling club; epipharynx trapezoidal in outline, apically truncate and fringed laterally with plumose setae; epipharyngeal inner brush setulae well-developed; mandibles bidentate; ventral galeal disc smooth and unarmed; galeal ventroapical margin with stiff spatulate setulae; lacinial short and robust; palpifer condyle broadly rounded and concave; mentum with anterior border slightly emarginate and with short anterior shelf; disc with a mixture of medium-length and short setae; glossal lobes indistinct; second palpal segment armed mesally with single medium-length seta; lateral metastemal stria present. Distribution. Paridolia is restricted to the Oriental realm. Biology and Ecology. Members of this genus are associated with decaying wood. The gut contents of a P. laevidorsis Lewis collected from a tree stump on Mindanao Island, Philippines, contained spores of the hyphomycete species P. Iraversianunt and P. betulicola. Immature Stages. The immature stages of Paridolia are unknown.

Genus Idolia Lewis 1885 (Figs. BOB, BIB, B2C, BBA, B4B, BSD, B6C-D, B7D, 55)

T ype Species Idolia gibba Lewis Adult redescription: Adult description: Body outline (dorsal) (Fig. BOB) circular. Body form globose. Head. Frons with laterofrontal carinae very long, V-shaped and meeting at clypeus. Antenna (Fig. B IB) with club (Fig. B2C) apically truncate; surface relatively sparsely covered with sensilla trichodea; transverse whorls of sensilla basaconica encircle club at level of both former proximal and distal divisions; some relatively short 144 sensilla chaetica originating near areas of former sutures and towards antennal apex. Epipharynx (Fig, 33 A) somewhat trapezoidal in outline. Outer margin of lateral area with loose fiinge of less than 10 regularly spaced short, apically plumose setae; fiinge setae abruptly decreasing in length on dorsal margn of lateral area. Distal portion of inner brush diverging at ca. 45° angle. Medial area commonly with 3 relatively large and 2 minute peg setae on each side and numerous minute sensilla. Dorsomedial peg setae short and numbering 2. Mandibles (Figs. 34B, 35D, 37D). Dorsolateral area apically bidentate and basally abbreviated and armed with short, conspicuous setae. Prostheca with inner dorsal setulae slender and relatively elongate and forming a distinct comb; radular setulae (Fig. 35D) slender and arranged in dense, oblique rows; proximal setulae apically bent at 90° degree angle, distal setulae gradually curving mesad. Maxilla (Figs. 55A-C) with ventral galeal disc untextured and armed with some apical setulae; ventromesal margin with nearly complete fiinge of slender setulae; ventroapical margin fiinged with stiff spatulate setulae; dorsal surface of galea armed proximally with relatively broad band of minute and short setae and followed by row of short to medium-length very stout, tapered setulae and relatively broad band of elongate, apically truncate spatulate setulae; an isolated cluster of medium-length tapered setulae present on dorsal surface near galea base; dorsal and ventrolateral membranous galeal base armed with minute denticulate setulae. Lacinia somewhat elongate oval in dorsal outline; some short fiinge setulae present dorsoapically; ventral surface extremely broad; surface divided into mesal base and broad membranous extension; lacinial tooth (Fig. S5C) elongate and slender; fiinge with extremely elongate apical blade-like setulae; fiinge setulae of medium-length and slender. Mediostipes armed with several minute setae. Basistipes with reticulate surface texture and armed with single medium length apical primary seta and some minute secondary setae. Cardo armed with several minute setae on ventral surface and a few longer setae laterally and mesally. Palpifer with some strong reticulate texturing ventrolaterally; 145 palpifer condyle acuminate with sharply bent apex. Second palpal segment with limited texture ventrally and with two ventral pores. Third palpal segment with nearly uniform texturing and 2 ventrolateral pores. Terminal palpal segment with strong, m ^ y ventral surface texture and lateral pores. Labium (Figs. 55D-F). Mentum (Fig S5D) not produced basolaterally and with anterior border weakly emarginate; anterior shelf absent; lateral margins fiinged with slender minute setae; disc uniformly textured and bearing a mixture of minute, short, and medium-length slender setae. Prementum (Fig. 55E-F) with lobes of paraglossae relatively narrow and with appendix; ligular fiinge consisting of long, slender, tapered setulae. Glossal lobes strongly produced, somewhat apically acuminate; ligular disc setae commonly numbering 4 per side. Palpiger bearing two pores and armed with at least 2 minute setae. Second palpal segment without surface texture; armed lateroapically with minute setae and apically with 5 short and single long lateral seta. Terminal palpal segment relatively elongate and only slightly expanded apically; uniformly textured and with single ventral pore. Ventral surface sparsely armed with mainly short setae; dorsolateral surface with dense covering of minute slender setae. Substructure with hyaline section of central arms elongate and extending to mesal glossal base; pigmented portion short. Medial sclerite absent. Median lobe short and broad. Legs short, about as long as substructure body; mesal border acuminate; leg base mesally expanded. Thorax. Prothorax. Notum with marginal stria; very sparsely and finely punctate. Sternum sometimes with imbricate texture (Fig. 36D); carinal stria stiate. Elytra not margined and very sparsely and finely punctate; with marginal epipleural, elytral, and subhumeral striae only. Hind wings. R4 not joined with RP; RA3+4 stigma relatively long, about 4x longer than wide; RPl dark and distinct; RP partially fused with MAl+2; RP2 apically abbreviated, extending less than half way to wing margin and continuing a short distance as sclerotized membrane; RP3+4 apically abbreviated, ending as vein about 2/3 of way to wing margin and extending to wing margin as sclerotized membrane; Mp3+4 light and 146 apically abbreviated; CUA2+3+4 short, in line with CUA and about half its length; CUA and AAl+2 joining beyond end of jugal lobe and forming eye; J short and distinct. Meso- and metastema (Fig. 36D) with meso-metastemal stria absent; longitudinal metastemal evanescent; lateral metastemal stria absent. Abdomen. Propygidium and pygidium veiy sparsely and finely punctate. Stemites with fine punctures along posterior margin. Male Terminalia (not figured). Tergite of eighth segment variably shaped, lightly sclerotized, and bearing cuticular cavities. Coxites short and very broad; and apically armed with several relatively long setae. Ninth segment reduced. Tergite subdivided into 2 very narrow distally bilobed valves. Aedeagal phalobase generally short and fused to parameres. Parameres united distally. Diagnosis: The following combination of characters will distinguish adult Idolia from other genera of Histeridae: Body form globose with sparse, fine punctures; laterofrontal carinae very long, V-shaped, and meeting at clypeus; antennal club apically tmncate and with pair of transverse whorls of sensilla basaconica encircling club; epipharynx with inner bmsh setulae atrophied; mandibles bidentae; inner dorsal prosthecal setulae relatively long and strmght; ventral galeal disc smooth and unarmed; galeal ventroapical margin with stiff spatulate setulae; isolated cluster of medium-length tapered setulae on dorsal galeal base; dorsal lacineal surface divided into a mgose mesal base and a broad membranous extension; lacinial spines absent; lacinial fiinge with extremely elongate distal blade-like setulae; palpifer condyle acuminate with sharply bent apex; mentum not produced basolaterally and lacking anterior shelf; glossal lobes distinct; palpiger with more than one setae on ventral surface; second palpal segment with long lateroapical seta; terminal palpal segment relatively elongate and only slightly expanded apically; elytra not margined and with marginal epipleural, elytral, and subhumeral striae only; meso- and metastema with distinct lacking meso-metastemal stria and lateral metastemal stria. 147 Distribution. Idolia is mainly Neotropical with one species barely extending into the Nearctic realm. Biology and Ecology. Members of this genus are associated with decaying wood. At least one species is commonly found beneath loose bark. The gut contents of a specimen of Idolia gibba Lewis collected under the bark of a large dead tree in CWapas, Mexico contained spores of the hyphomycete species S. obovata and S. groveri as well as spores of an unidentified coelomycete (Diplodea ox Botryodiplodid). The gut contents of a specimen of/, scitula Lewis from Aragua, Venezuela contained spores of an unidentified coelomycete (Diplodea ox Botryodiplodid). I collected several larvae of/, gibba from accumulated organic material beneath the loose bark of a dead trees in Chiapas, Mexico in August, 1991. Immature Stages. Larvae of /. scitula were collected in February by A. Newton on Barro Colorado Island from litter beneath an old tree. Larvae of /. gibba were collected in August from humic material beneath loose bark of a fallen tree in Chiapas, Mexico. Idolia larvae, like larvae of Eutribalus, have transparent cuticles and accumulated what appears to be uric acid in their hind gut and Malpighian tubules. What these larvae feed on in nature is unknown. Larvae maintained in the laboratory readily accepted house fly eggs. Pre-pupae constructed circular cocoons in which to pupate using silk secreted from the anus. The cocoons were very sturdy and required considerable effort to break into them to remove the pupa for study. The larva and pupa of /. gibba are described in chapter S. Species List The following is a list of species names for the Onthophilinae. The source of this information was Mazur (1984, 1988), Gomy (1984), Yelamos (1991), and Johnson et al (1992) and Vienna (1993). Rather than provide detailed locality data for each species, each name is followed by the biogeographic realm(s) where it occurs: AA = Australasian; AT = Afrotropical; NA =Nearctic; NT = Neotropical; OR = Oriental; PA = Palearctic. I did not examine material for all of the species listed. Those that were 148 examined are notated in the following manner; 1 = primary type was seen; 2 = secondary type was seen; * = authoritatively determined specimen(s) were seen. I made no attempt to synonymize names at the species level. This can best be done when onthophiline genera are revised at the species level. My main objective in examining material was to ensure that the material conformed to the generic concepts proposed in this revision.

Subfamily Onthophilinae Thomson Genus Onthophilus Leach 1. Onthophilus qffinis Redtenbacher [PA] 2. Onthophilus altematus (Say) [NA] * 3. Onthophilus annoi Ohara [PA] 4. Onthophilus bickhardti Reitter [PA] 5. Onthophilus convictor Normand [PA] * 6. Onthophilus cynomysi Helava [NA] 7. Onthophilus deflectus Helava [NA] * 8. Onthophilus extrordinarus Reichardt [PA] 9. Onthophilusflavicomis Lewis [PA] 10. Onthophilusflohri Lewis [NA- Mexican plateau] * 11. Onthophilusfoveipennisl^ms [PA] 12. Onthophilus giganteuslA&XdLVd. [NA] * 13. Onthophilusglobosus (Oliver) [PA] 14. Onthophilus intermixus Helava [NA] * 15. Onthophilusjulii Lewis [NA- Mexican plateau] 16. Onthophilus hmiyai Adachi [PA] 17. Onthophilus kimi'Ro^s [NA] * 18. Onthophilus lecontei Horn [NA] * 19. Onthophilus melampusRéichaidt [PA] 20. Onthophilus niponensis Lewis [PA] 21. OnthophilusnodatushçConiQ [NA] * 22. Onthophilus ordinarius Lewis [PA] * 23. Onthophilusostreatus'LQms [PA, OR] * 24. Onthophiluspluricostatus’LeConie [NA] * 25. OnthophiluspunctatusMàWQx [PA] 26. Onthophilus sculptilis Lewis [OR] 27. Onthophilus silvae Lewis [PA] 28. Onthophilus soltaui Casey [NA] * 29. Onthophilus striatus (Forster) [PA] * 30. Onthophilus thonomysi Helava [NA] 1 31. Onthophilus tuberculatusl^tms [PA] 32. Onthophilus wenzeli Helava [NA] * 149 Genus Vuattouxînus Thérond 1. Vuattouxinus borassicola Thérond [AT] 2

Genus Epîechinus Lewis Syn.: G/ymma Marseul Sculptura Thérond Sigillum Thérond 1. Epiechinus arboreus Lewis [PA] 2. Epiechinus bipartitus Lewis [AT] * 3. Epiechinus candezii (Marseul) [AT] n. comb. 2 4. Epiechinus caucasicola Bickhardt [PA] 5. Epiechinus cavistemsus Bickhardt [AA] 6. Epiechinus dementi Thérond [AT] 7. Epiechinus commersoni Gomy [AT] 8. Epiechinus costatus (MacLeay) [AA] 9. Epiechinus costipennis (Fahraeus) [AT] 10. Epiechinus desboresi Burgeon [AT] 11. Epiechinus julvosetosus (Sahlberg) [PA] 12. Epiechinus hispidus (Paykull) [OR] 13. Epiechinus hova (Lewis) [AT] 14. Epiechinus kivuensius (Thérond) [AT] n. comb. 1 15. Epiechinus kuntzeni Bickhardt [AT] 16. Epiechinus laceratus Schmidt [AT] 17. Epiechinus lagunae Heller [OR] 18. Epiechinus leleupi (Thérond) [AT] n. comb. 1 19. Epiechinusmalayieus'Q\c\ihaxdX [OR?] 20. Epiechinus marseuli Lewis [OR] * 21. EpiechinusnotogaeusBic)âméi [AA] 22. Epiechinus novemcostatus (Marseul) [AT] 23. Epiechinus perrieri Fairmaire [AT] 24. Epiechinus planistemus Bickhardt [AA] * 25. Epiechinuspumilus Thérond [AT] 26. Epiechinuspunciistemus (Lewis) [AT] 27. Epiechinus rappi Bickhardt [AT] * 28. Epiechinus resinus Schmidt [AT] 29. Epiechinus sculptus Gomy [AT] 30. Epiechinus seriepunctatus Bickhardt [AT] 31. Epiechinus sulcistemus Bickhardt [AT] 32. Epiechinus taprobanae Lewis [OR] 33. Epiechinus Iherondi Gomy [AT] 34. Epiechinus tuberculistemus (Lewis) [AT] 150

Genus Austraiepierus Kovarik, n. genus Syn.: Bickhardt Onthophilus Leach 1. Australepierus abrogatus (Broun) [AA] 1 2. Australepierus australis (Helava & Howden) [AA] n. comb, * 3. Australepierus crenulatus (Broun) [AA] n. comb. * 4. Australepierusfoederatus (Lewis) [AA] n. comb. * 5. Australepierus phyllobius ^roun) [AA] n. comb. * 6. Australepierusplaniceps (Broun) [AA] n. comb. 1 7. Australepierus punctulipennis (Broun) [AA] n. comb 1 8. Australepieruspurus (Broun) [AA] - n. comb. 1 9. Australepierus rufescens (Reitter) [AA] - n. comb * \QAustralepierus rusticus (Broun) [AA] \\ Australepierus simplex ^roun) [AA] n. comb. 2 llAustralepierus spinellus (Broun) [AA] n. comb. * Vi Australepierus sylvanus(^Qmi) [AA] n. comb. 2

Genus Plagiogramma Tarsia Syn.: Pseudepierus CzsQy 1. Plagiogramma arciger (Marseul) [NT] 2. Plagiogramma brasiliense Tarsia [NT] * 3. Plagiogramma caviscuta (Marseul) [NT] 4. Plagiogramma coproides (Marseul) [NT] * 5. Plagiogramma darlingtoni (Wenzel) [NT] 6. Plagiogramma epulo (Marseul) [NT] 7. Plagiogrammafissus (Marseul) [NT] * 8. Plagiogrammafomicata (Marseul) [NT] 9. Plagiogrammafrater (Marseul) [NT] 10. Plagiogrammafrontale (Kirsch) [NT] * 11. Plagiogramma hastata (Marseul) [NT] 12. Plagiogramma gentilis OHom) [NA] n. comb. * 13. Plagiogramma glabra (Bickhardt) [NT] * 14. Plagio^amma guyanense (Mazur) [NT] * 15. Plagiogramma incas (Marseul) [NT] 16. Plagiogramma intermedia (Marseul) [NT] * 17. Plagiogramma italica (Paykull) [PA] n. comb. * 18. Plagiogramma leleupae (Wenzel) [NT] 19. Plagiogramma lucens (Marseul) [NT] 20. Plagiogramma marseulii (Kirsch) [NT] 21. Plagiogramma mundas (Erichson) [NT] * 22. Plagiogramma nitiscens (Marseul) [NT] 23. Plagiogramma paradoxa (Mazur) [NT] 151 24. Plagiogramma patrueîis (Lewis) [NT] 25. Plagiogramma peruana (Schmidt) [NT] ♦ 26. Plagioff'ammapubifrons (Hinton) [NT] * 27. Plagiogramma rhinocera (Marseul) [NT] * 28. Plagiogramma schmidti (Wenzel & Dybas) [NT] * 29. Plagiogramma spaerula (Marseul) [NT] 30. Plagiogramma stratipyga (Wenzel) [NT] 31. Plagiogramma subtropica (Casey) [NA] * 32. Plagiogramma tersus (Erichson) [NT] 33. Plagiogramma trux (Marseul) [NT] *

Genus Epierus Erichson 1. Epierus alberti Marseul [NT] 2. Epierus alutaceus Marseul [NT] * 3. Epierus angularis Schmidt [NT] 4. Epierus antillarum Marseul [NT] 5. Epierus beccarii Marseul [OR, AA] * 6. Epierus binasutus Cooman [OR] 7. Epierus bisbistriatus Marseul [NT] * 8. Epierus biscissus Marseul [AA] 9. Epierus brurmipennis Marseul [NT] 10. Epierus complus Erichson [PA] * 11. Epierus comutus Casey [NA] * 12. Epierus cylindricus Wenzel [NT] 13. Epierus dalaunayi Lewis [OR] 14. Epierus devius LeConte [NT] 15. Epierus divisus Marseul [NT] * 16. Epierus ellipticus LeConte [NA] * 17. Epierus floridanus Casey 18. Epierus foveolatus Hinton [NT] 19. Epierus fulvicomis ÇFabricius) [NT] * 20. Epierus humeristrius Schmidt [NT] * 21. Epierus Marseul [NT] * 22. Epierus inscriptus Schmidt [NT] 23. Epierus insularis Schmidt [OR] 24. Epierus /«v/ûft/j Marseul [NT] * 25. Epierus kraatzi Schmidt [NT] 26. Epierus laevistrius Marseul [NT] 27. Epierus latior Bickhardt [NT] 28. Epierus longulus Marseul [NT] 29. Epierus lucidulus Erichson [NT] * 30. Epierus lucus Erichson [PA] 31. Epierus mariae Marseul [NT] 152 32. Epierus mehicanus LeConte [NT] * 33. Epierus mundus Erichson [NT] 34. Epierus nasicomis Bickhardt [OR] 35. Epierus nemoralis Lewis [OR] 37. Epierus notius Marseul [NT] * 38. Epierus novellus Zimmermann [NA] 39. Epierus ohlongus Casey [NA] 40. Epierus obsolescens Casey [NT] 41. Epierus planulus Lewis [OT, NA] * 42. Epierus pulicarius Erichson [NA] * 43. Epierus regularis (Benuvois) [NA] * 44. Epierus sauteri Bickhardt [OR] 45. Epierus scitus Lewis [NT] 46. Epierus singulistriatus Hinton [NT] 47. Epierus smaragdinus Maiseul [NT] 48. Epierus toxopei Desbordes [AA] 49. Epierus -vagcms Marseul [NT] 50. Epierus vandepoHi Schmidt [NT] * 51. Epierus vethi (Bickhardt) [NT] 52. Epierus villiersi Cooman [OR] 53. Epierus waterhousii Marseul [NT] *

Subfamily Tribalinae Wenzel Genus Stictostîx Marseul 1. Stictostix biseriata (Schmidt) [AA] 1 2. Stictostix califomica (Horn) [NA] * 3. Stictostix cognata Lea [AA] 1 4. Stictostix ectatommae Lea [AA] 1 5. Stictostix frontalis (MacLeay) [AA] 1 6. Stictostix leae Lewis [AA] 7. Stictostix mormon Lewis [NA] 1 8. Stictostix parra (Marseul) [AA] *

Genus Peplogfyptus LeConte 1. Peploglyptus belfragei LeConte [PA] 1 2. Peploglyptus golbachi Kanaar [NT] *

Genus Tribalus Erichson 1. Tribalus algericus Olexa [PA] 2. Tribalus amnicola Lewis [AT, PT] 3. Tribalus anatolicus Olexa [PA] 153 4. Tribalus andreinii Müller [AT] 5. Tribalus ascaphus Marseul [AT] * 6. Tribalus atlantis Yélamos [PA] 7. Tribalus brevistemus Vienna [AT] 8. Tribalus capensis (Paykull) [AT] * 9. Tribalus cavemicola Lewis [AT] * 10. Tribalus crypticus Vienna [AT] PT IL Tribalus distinguendus Müller [AT] 12. Tribalus elapsus Vienna [AT] 13. Tribalus endroedyi Vienna [AT] 14. Tribalus excellens Vienna [AT] 15. li-ibalus exilis (Paykull) [OR] 16. Tribalusfastigiatus Marseul [AT] * 17. Tribalusfastigatus lucidus "Vienna [AT] 18. Tribalusfloridus Vienna [AT] 19. Tribalusfoveolatus Vienna [AT] 20. Tribalus freyi Müller [PA] DETFM 21. Tribalus gioiellae Vienna [AT] 22. Tribalus gracilipes'Wi&nra. [AT] 23. Tribalus homi Lewis [OR] PT 24. Tribalus impressibasis Bickhardt [AT] 25. Tribalus impinatus Vienna [AT] 26. Tribalus interruptus Vienna [AT] 27. Tribalus kanaari Vienna [AT] 28. Tribalus kochi Thérond [AT] ETV 29. Tribalus kovariki Vienna [AT] 30. Tribalus lelupi Thérond [AT] 31. Tribalus margiventer Mazur [OR] 32. Tribalus maroccanus Olexa [PA] 33. Tribalus longipes Venna [AT] 34. Tribalus micros Mazur [AT] 35. Tribalus minimus (Rossi) [PA] * 36. Tribalus minutulusTYiéïonà [AT] 37. Tribalus modicus Cooman [OR] 38. Tribalus nitens Vienna [AT] 39. Tribalus oblongus Vienna [AT] 40. Tribalus politus Vienna [AT] 41. Tribaluspunctillatus'B\c)ùi 2iidX [OR] * 42. Tribalus ratti Vienna [AT] 43. Tribalus rubiculus Schmidt [AT] * 44. Tribalus scaphidiformis (Illiger) [PA] * 45. Tribalus similis Vienna [AT] 154 46. Tribalus subdolus Menna [AT] 47. Tribalus subfasciatus Vienna [AT] 48. Tribalus subfasciatus opacinotus Vienna [AT] 1 49. Tribalus tibialis Vienna [AT]

New synonymy: Tribalus bicarinatus Lewis =Anapleus bicarinatus Lewis n. comb. 1 Tribalus unistriatus Lewis =Anapleus bicarinatus Lewis n. comb. R. Wenzel, pers. comm.

Genus Eutribalus Bickhardt, n. stat. Syn.: Tribalasia Coovtan Tribalus LeConte Caerostemus 1. Eutribalus acceptus (Marseul) * 2. Eutribalus agrestis (Marseul) [AT] * 5. Eutribalus americanus (LeConte) [NA] comb. * 4. Eutribalus australis (MacLeay) [AA] * 5. Eutribalus bomba (Marseul) [OR] * 6. Eutribalus catenarius (Lewis) [OR] 7. Eutribalus colombius (Marseul) [OR] * 8. Eutribalus doriae (Marseul) [OR] * 9. Eutribalus eggersi (Bickhardt) [AT] 11. Eutribalus koenigius (Marseul) [OR] 12. Eutribalus marseuli (Gomy) [OR] * 13. Eutribalus ogieri (Marseul) [OR] * 14. Eutribalus onustus (Lewis) [AT] 15. Eutribalus opimus ^ewis) [OR] * 16. Eutribalus pseudostrialis (Mazur) [OR] 17. Eutribalus puncticeps (Lewis) [OR] * 18. Eutribalus suturalis (Lewis) [OR] * 19. Eutribalus tropicus (Lewis) [OR] 20. Eutriblaus vallata Cooman [OR] 1

Genus Parepierus Bickhardt Syn.: Scaphidister Coomm 1. Parepierus alutaceus Cooman [OR] 2. Parepierus amandus (Schmidt) [OR] 2 3. Parepierus arcuatus Bickhardt [OR] 4. Parepierus chaostrius Cooman [OR] 155 5. Parepierus corticola (Bickhardt) [OR] 6. Parepierus indiicola (Mazur) [OR] 7. Parepierus lewisi Bickhardt [OR] 8. Parepierus monticola (Schmidt) [OR] 9. Parepierus musicola (Cooman) [OR] n. comb. 10. Parepierus opacipermis Bickhardt [OR] 11. Parepierus orphanus (Lewis) [OR] 12.ParepierusovulatusBviMmdi [OR] * 13. Parepierus salvazai (Desbordes) [OR] * 14. Parepierus silvaticus Cooman [OR] 15. Parepierus subhutneralis Cooman [OR] * 16. Parepierus tumidifrons Cooman [OR] * 17. Parepierus velox (Cooman) [OR] n. comb. * 18. Parepierus vitalisi (Desbordes) [OR] n. comb. *

Genus Paridolia Kovarik, n. genus Syn.: ThrWf/f Erichson 1. Paridolia cornes {(Zoovcax^[OK\ n. comb. * 2. Paridolia corylophoides (Lewis) [OR] n. comb. 1 3. Paridolia laevidorsis (Lewis) [OR] n. comb. 2 4. Paridolia modicus (Cooman) 5. Paridolia montanus (Lewis) [OR] n. comb. 2 6. Paridolia pumilio (Schmidt) [OR] n. comb. *

Genus Idolia Lewis 1. Idolia antermata Lewis [NT] 2. Idolia gibba Lewis [NA, NT] * 3. /(5fo//a/w/egra Lewis [NT] 4. Idolia laevissima (LeConte) * 5. Idolia nucleola (Marseul) [NT] 6. Idolia punctistemum Lewis [NT] 7. Idolia scitula Lewis *

Spore feeding. Spores of fungi and, to a lesser extent, those of protists were commonly found in the gut of adult onthophilines and tribalines. The only taxa in which no spores were found were Tribalus and Vuattouxirms. Spore feeding îor Epierus, although reported by Wenzel (see Moser et. al. 1971), has largely been overlooked. All fungal spores found to date belong to the form class Deuteromycontia. According to Kimbrough (pers. comm.) this is an artificial grouping of the asexual stage of fungi. The 156 majority of the deuteromycontian genera have been linked to the ascomycetes while a minority are linked to the basidiomycetes. Most adult histerids and adults of the closely related families Synteliidae and Spaeritidae are macrophagous predators. This habit is likely ancestral in the Histeridae. Most histerid adults are trophically dependent on insect protein. Since fungal spores are essentially protein surrounded by chitin (Kimbrough pers. comm.), spore feeding would not represent a change in food quality. However, gathering spores in sufficient quantities to satisfy nutritional requirements presents a problem. According to Lawrence (1989) two major problems in the switching from macrophagy to microphagy are the gathering of often scattered food items and concentrating and moving them to a processing area. The solution to these problems involve modification of the mouthparts. The principal gathering device appears to be the galea of the maxilla. Some of the Tribaline genera with a relatively unmodified galea (eg. Stictostix, and Peploglyptus) do gather and consume some fungal spores as evidenced by examination of gut contents. However in all cases the number of spores in the gut was relatively low when compared to those groups with more specialized mouthparts, suggesting that these taxa are facultative spore feeders. Conversely, the gut of onthophiline and tribaline genera with the galea modified for spore gathering (eg. members of the Epierus and Idolia genus groups) was often packed with spores. Moving gathered spores to the mandibular mola for grinding is likely accomplished via the maxillary lacinia and mandibular prostheca. While the exact function of the prosthecal radula remains undetermined, it may function as the file it resembles, scraping chitin sheathing from spores. It is likely that most onthophiline taxa with mouthparts specialized for microphagy are opportunistic predators. Evidence for this is the acceptance of housefly eggs and early instar larvae by species of Onthophilus, Epierus, Plagiogramma, and Idolia maintained in the laboratory. A possible exception is Eutribalus and perhaps its sister genus Parepierus. Specimens of E. americanus 157 maintained in the laboratory had difficulty feeding on fly eggs and would not feed on early instar housefly larvae. The mandibles of these taxa may be so modified for spore processing as to be inefficient for predation. Some genera of the Onthophilus genus group and most members of the Epierus and Idolia genus groups are found in association with moist rotting wood and feed on fungal spores developing on this substrate. According to Kimbrough (pers. comm.) it is likely that all of the Deuteromycontia are able to break down cellulose via enzymes (cellulase). According to Subramanian (1981), many species of hyphomycetes colonize oak wood, a substrate with which several species o î Epierus and some New World Idoliini are known to associate (Kovarik, unpublished data). These fungi are likely to persist on a rotten log for quite some time. Most fungi need both moisture and warmth for major vegetative growth and firuiting (Subramanian 1983, Kimbrough pers. comm ). Coincidentally, reproduction of the wood-associated histerids appears to be correlated with identical conditions. In those taxa that specialized on gathering fungal and protist spores, three major types galea modifications were encountered. The first consisted of spatulate dorsal galear setulae typical of Epierus (Fig 23 A-B), Plagiogramma, and genera of Idoliini. A very different but apparently equally effective type of setular arrangement is characteristic of the genus Australepierus (Fig. 23C-D). A third type of galeal modified for spore gathering is found in Epiechinus. Although these modified galea differed markedly in terms of their structure, the spore-packed guts of dissected specimens attest to the spore gathering capabilites of each. Mouthpart sturcture of species of Onthophilus and Vuattouxirms borrasicola appear to be modified for liquid filter feeding. The mandibular comb hmrs (Fig. 2 IB) of Onthophilus and V. borrasicola and the galeal filters (Figs. 39C) of all these taxa and gut contents indicate that these beetles are gathering extremely small food particles. According to Lawrence (1989) food gathering in a liquid medium, problems of food gathering are increased because smaller food items are in suspension. 158 These taxa appear to obtain nutriment from fermenting liquids associated with tree wounds, or liquids on the surface of dung or decaying phloem and sapwood. This liquid contains bacteria and fungi that subsist on non-structural carbohydrates (Lawrence 1989, Subramanian 1981). The galea of species of Tribalus (Fig. 39A) is generalized, resembling that found in other histerids known to be predatory. The long robust tapered setulae probably aid in fluid conduction. Several features of the mandibles of Tribalus (Fig. 29D) including a reduced mola, elongate pubescence along mesobasal margin, and long mandibular teeth are typical of predatory Coleoptera. Biological observations by Prinz (1984) combined with a lack of spores in the gut contents suggest that species of Tribalus may be strictly macrophagous predators.

Phylogenetic Analysis Character evaluation: Component structures of male terminalia were generally found to be highly variable within a genus and therefore were of limited importance in determining phylogenetic relationships among genera. In general, surface sculpture and texture and elytral striation were often found to be variable within a genus and likewise were of limited importance. The genus Australepierus is a good example of a genus which contains taxa which are strikingly different in appearance. A. phyllobius (Broun) has an essentially featureless pronotum and elytra and finely punctate meso- and metathoracic sterna. Australepierus australis (Helava) has a carinate pronotum and elytra, and a coarsely punctate meso- and metastemum. Although most species of the genus Parepierus have a full complement of dorsal striae, P. vitalisi (Desbordes) was exceptional in having none. Conversely most ventral thoracic and abdominal sutures and striae were found to be relatively stable characters. Mouthpart and antennal structure proved to be much more stable and diagnostic. Perhaps the microphagous mode of feeding in the Onthophilinae has 159 much to do with the overall stability and usefulness of mouthpart characters. Likewise hind wing venation proved to be very informative and an important character in this analysis. Relationships and out group selection. This analysis was intended to test the putative monophyly of the taxa included in the Onthophilinae and Tribalinae. However the lack of any phylogenetic treatment of taxa above the level of genus in the Hsteridae left me with no designated outgroups to choose for my analysis. My initial strategy in searching for outgroups of my ingroup was to compare prospective outgroup candidates with outgoups of the Histeridae. Hydrophilids, synteliids and sphaeritids were all used to hypothesize character polarity within the Histeridae. Both adults and larvae were examined. One of the most obvious features of histerid larvae are the many setae and pores present on the head and body. Since no system existed for describing and homologizing setae and pores for histerid larvae, an initial priority was to develop such a system in order to maximize the information content of the larvae. This study was important in that it provided coroborating evidence of the relatively unspecialized nature of two taxa, Dendrophilus and Anapleus, which were under consederation for use as potential outgroups of onthophilinae and tribaline taxa. The terminal segment of the urogomphi in histerids always bears two long mechanoreceptors setae, UG12 and 13. Both of these setae commonly arise from the apex of the urogomphus. However in Dendrophilus and Anapleus, seta UG13 arises subapically. This same condition occurs in larval Hydrophiloidea other than Histeridae, including Helophorus, Syntelia, and Sphaerites, indicating that this condition is plesiomorphic. Several adult characters o î Anapleus and Dendrophilus were also found to be plesiomorphic. These include the externally subdivided antennal club, and the presence of a longitudinal antennal groove and and the lack of depressions for receiving the retracted antennal club and no prostemal.alae The hind wing venation of Dendrophilus was 160 decidedly more plesiomorphic than that found in Anapleus. The hind wing venation of Dendrophilus is remarkably complete and closely resembles the hind wings of Sphaerites. Plesiomorphic features include the presence of wing vein AP3+4 and the “off-line” position of the CUA2+3+4. The plesiomorphic positioning of the CUA2+3+4 is. Anapleus hind wings lack these plesiomorphic features. Despite the wing venational differences, I was fairly confident that because of these and other plesiomorphic characters, Dendrophilus Anapleus are indeed, relatively underived histerids, and good candidates for outgoup comparison with the genera of Tribalinae and Onthophilinae. Validitation of established genera. The first step in my actual analysis was to examine adults of all described genera of the Onthophilinae and Tribalinae to determine which among them were valid. To accomplish this, I relied mainly on antennal structure, mouthpart structure, hind wing venation, male genitalic structure, and some external features. I then compiled data matrices for larval, pupal and adult characters. I was unable to obtain larval or pupal material for the following genera: Vuattouxirms, Epiechinus, Stictostix, Peploglyptus, Tribalus, Parepierus and Paridolia. The outgroups selected for my phylogenetic analysis of larval and adult Tribaline and Onthophiline genera were Dendrophilus and Anapleus. Since no pupae oî Anapleus were available, Dendrophilus alone was used as the outgroup for the pupal analysis. Shaericosoma is excluded fi'om the Tribalinae because it lacks developed prosternai alae and because its antennal club is divided by both proximal and distal sulci. However, Shaericosoma does share in common certain derived characters of the hind wings which are found in Anapleus and the Tribalinae. These include apically abbreviated wing veins RP2 and RP3+4, and a lack of maculations and wing vein AP3+4. Shaericosoma also shares in common with Anapleus and Stictostix the presence of pits on the dorsolateral area of the mandibles. Anapleus, Shaericosoma relies its retracted foreleg to protect its antennae. However Shaericosoma differs îcovciAncq)leus in having depressions in the anterior angles of the 161 pronotum for receiving the antennal club. The antenna! grooves of Shaericosoma are nearly perpendicular to the longitudinal axis of the body and are posteriorly bordered by presumptive alae. These characters hint at a possible sister group relationship of Sphaericosoma and the Tribalinae. Character analysis. Initial data analysis of 111 adult characters (the most complete data set) via HENNIG86 phylogenetic program produced only two trees 'mth relatively high retention and consistency indicates, but one of the out groups, Anapleus, was pulled into the ingroup as the sister of Stictostix, Peploglyptus, Tribalus, Eutribalus, Parepierus, Paridolia, and Idolia. This was the first indication of possible paraphyly among these taxa. Initially I considered this to be a problem of working with basal taxa of the Histeridae. Â relatively insignificant amount of homoplasy might be sufiGcient to move an outgroup such as Anapleus into the ingroup. I began re-examining adult characters in an attempt to determine the source of this suggested paraphyly. One character which argued against paraphyly was the presence of prosternai alae (Fig. 13D [al] ) in all members of the Onthophilinae and Tribalinae. Alae are not present in Anapleus or Dendrophilus leaving little doubt that alae are derived structures. The question then becomes, did these structures evolve in a recent common ancestor of tiibaline and onthophiline genera, or did they evolve independently. New evidence suggests the latter conclusion. As mentioned, the hind wing venation oîDendrophilus is remarkably plesiomorphic. It shares a number hind wing synplesiomorphies with Onthophilus (Fig. 42A) not found among other onthophiline or tribaline genera. These include the presence of AP3+4, a complete RP3+4, and dark wing maculations. These and several other unique features, including relatively long lateral palpiger setae in adults, and socketed setae in the fi*ontale group in larvae. However, prosternai alae are present in Onthophilus adults while absent in Dendrophilus. Anapleus adults lack alae but they have derived wing venation characters including the loss of AP3+4 an apically abbreviated 162 RP3+4 and un-maculated wings. These same derived features are found in the genera grouped by HENNIG86 with Anapleus. In order to envision a sister group relationship \iQtwQQn Anapleus and all genera of onthophilinae and tribalinae, one would have to assume the wing vein AP3+4 re-evolviing and RP3+4 re-establishing contact with the wing margin.in Onthopilus. Wing vein reduction tends to be unidirectional in holometabolous insects and it is unlikely that histerid beetles are an exception. A more parsimonious argument would be to accept that the ingroup is paraphyletic and assume that alae have evolved independently in the two groups. The presence of alae in other apparently unrelated groups of histerids (i.e. Histerinae, Chlamydopsinae, and Haeterinae) lends support to an independent origin of these structures. Additional evidence supporting a sister group relationship oî Anapleus with some but not all ingroup taxa are the presence of pits (Fig. 29 A, B) on the lateral area of mandibles va Anapleus and Stictostix and the presence of broad overlapping scales on the epipharyngeal lateral areas la Anapleus and all taxa grouped with it by HENNIG86. Accepting paraphyly among my ingroup genera, I ran two separate analyses combining 355 adult, larval and pupal characters (see appendix 1) for each subset. The first group consisted of the outgroup Dendrophilus and all the taxa grouped with it fi’om my initial analysis, including Onthophilus, Vuattouxinus, Epiechinus, Australepierus, Plagiogramma, and Epierus, or the Onthophilinae. The second group combined the out group Anapleus and the in group taxa formerly grouped with and listed above, or the Tribalinae. The larval and pupal data set for the Onthophilinae was the most complete; data was missing for only two of the ingroup genera, Vuattouxinus and Epiechinus. The larval and pupal data set for the Tribalinae is, unfortunately, far less complete. Larval data was available for the out group Anapleus and larval and pupal data were available for only two taxa of my in group, Eutribalus and Idolia. An analysis of the Onthophilinae via the 163 implicit enumeration option of HENNIG yielded a single cladogram (Fig. 57) with a length of 127, a consistency index of 0.71 and a retention index of 0.60. Because of the incomplete larval and pupal data set for the Tribalinae, I ran two separate analyses; one using adult data only and the other combining the data sets. The analysis of adult characters, again using the implicit enumeration option, produced two equally parsimonious trees each with a length of 151, a consistency index of 0.73 and a retention index of 0.76. A Nelson consensus tree retained from this analysis presented in Fig. 58 is accepted as the probable phylogenetic relationship of tribaline genera. Of the two trees produced by implicit enumeration, the tree not retmned by the HENNIG86 Nelson analysis differs from the retained tree only in placement of the genera of Idoliini. Whereas the Nelson tree placed Idolia as the sister oîParidolia, Eutribalus and Parepierus, the rejected tree indicates a sister relationship for Idolia and Paridolia. The combined character analysis of the Tribalinae produced a single tree in which the only two ingroup genera for which larval and pupal data were available {Eutribalus and Idolia) were grouped together as sister genera. Because a sister group relationship îor Eutribalus and Idolia is unsupported by adult characters, the tree resulting from the analysis of combined data is regarded as the product of missing larval data for other Idoliini genera rather than a valid relationship. An improved resolution of relationships among genera of Idoliini is anticipated once the immature stages of Paridolia and Parepierus are discovered. The Onthophilinae constitutes a monophyletic group which is well-supported by adult and larval characters. Adult characters include: epipharyngeal lateral with reduced, denticulate and generally non-overlapping scales; mandibles with prostheca apically detatched (except Onthophilus) and the presence of a lacineal strobilus. Larval characters include a reduction of asperities (except Onthophilus) and the presence of 2 anterior tergites on each side of abdominal segment IX. 164 Adults of the Onthophilus genus group share a number of synapomorphies including a vertico-frontal carina, an apically acuminate mandibular medial area, prosthecal comb setulae Epiechims), deep punctures along the elytral intervals, and teeth on the anterior margin of the fore-tibia. The hypothesized sister group relationship of Vuattouxinus and Epiechinus is supported by the erect scale vestiture, the antennal club sensory plaque, a swollen antennal scape armed with short setae, an unarmed terminal maxillary palpal segment, lateral lobules on the paraglossae, male abdominal segment Vm with very large and dorsoventrally flattened coxites, male abdominal segment IX travois shaped, and apically fused parameres. Adult synapomorphies supporting the Epierus genus group include a mesally extending galea, an apically abbreviated or absent palpifer carina, and apically abbreviated KP2 hind wing veins. Larval characters supporting this clade include metathoracic setae TEl, 2, and pore TEb sharing a sclerite, the absence of metathoracic seta TE3, metathoracic setae ST36 and 37 sharing a sclerite, abdominal segments I-VUI with a single medial interstemite, setae ST29 and 30 sharing a tergite, abdominal tergal sclerites TE5 and TE17 fused, abdominal segment IX with seta TE9 and pore TEg sharing a sclerite, urogomphal setae UGl-7 sharing a common sclerite, and a cuticle largely free of asperities. The hypothesized sister group relationship of Epierus and Plagiogramma is very well supported by adult and immature stages. Adult synapomorphies include: sexually dimorphic characters e.g., males with a setigerous tubercle on the clypeus and with a digitiform projection on the lateral area of the left mandible; non-dimorphic characters such as a detached epipharynx, a comb along the mesal margin of the lateral area, the loss of mandibular contact chemoreceptors, the presence of a mandibular radula, reduction of mandibular microtrichia, spatulate setulae on the galea, the palpifer with a very robust ventroapical primary seta, ligular fringe setulae apically expanded and barbed, and a pair 165 of extremely robust glossal disc setae. Larval characters in support of this clade include the absence of frontale seta FRl, the loss of abdominal segment TE17, abdominal segments I-Vm with ST33 isolated on a separate sclerite while ST31 and 32 share a sclerite, and digitiform paragomphi in the second instar. The Tribalinae constitutes a monophyletic group which is defined by the following adult characters: epipharyngeal lateral with well developed, and generally broad overlapping scales; third maxillary palpal segment with a reduced number of primary setae; and hind wings with faint or no maculations. Adults of the Tribalus genus group share the following synapomorphies: antennae generally at least as long as the head capsule width, an apical tooth on the antennal; a uniformly broad dorsolateral area on mandible, no maculations on the hind wings, wing veins KP4 and RP proximally united, and coxites of male abdominal segment Vm armed with relatively long setae. The hypothesized sister group relationship of Stictostix and Peploglyptus is supported by the following adult characters: antennae longer than the head capsule width, umbiculate punctures and elytral costae in some taxa, a tridentate mentum, premental medial sclerite present, and relatively long prementum substructure legs. The Idolia species group is particularly well-supported by adult, larval and pupal characters. Adult characters include: an absence of the epipharyngeal dorsal brush, reduction of the inner brush, constriction of the lateral area of the mandibles, retraction of the mola, reduction of the mandibular microtrichia, prostheca with extremely long apical setulae, mandibular radula is present, lacinea is distally detatched from mediostipes, a dorsal condyle is present on the palpiger, the maxillary dorsal membrane is unarmed, and the terminal labial palpal segment is dorsoventrally flattened and has relatively long sensilla basiconica. Larval synapomorphies include setiform sensilla on the antennae, an unarmed dorsal surface of the prementum, the presence of a unique pore (LAc) on the labial palpi, a unique seta, ST45, on the meso- and metastemites; vestigial metathoracic lateral humeral 166 tergites; tibiae with some short setae; a lack of metathoracic intertergites or thoracic interstemites; abdominal segments I-Vni with setae TE14,15,16 on separate sclerites;

abdominal segment DC lacking pores and with seta TE16 and 21 each on separate sclerites, and with an apical urogomphal seta UGll. Pupal synapomorphies include a protuberant galeal scaffold and the mesonotum with at least 4 scutal setae per side. The hypothesized sister group relationship of Eutribalus and Parepierus within the Idoliini is supported by the following adult characters: antennal club securiform and with sensilla basiconica patchily distributed, unidentate mandibles, and with the ventral surface of galea imbricately textured. This analysis is an important first step towards bringing phylogenetic methodology to bear on the relationships of genera of histerid beetles. It also offers insight into character polarity for the family as a whole. While establishing monophyly for the Onthophilinae and Tribalinae, this analysis has suggested the probable paraphyly of the Dendrophilinae. A great deal of work remains to be done on the higher classification of the Histeridae. Characters which have been historically overlooked, such as mouthpart structure, hind wing venation, and larval and pupal chaetotaxy proved to be pivotal in this analysis. These same characters will undoubtedly prove to be equally important in further works on phylogenetic relationship within the Histeridae. Figure 13. Prothoracic modifications for antennal protection: Anapleus vs. Stictostix. (A) Anapleus marginatus^ head on view. (B) Stictostix califomica, head on view. (C)A. marginatus, ventral view. (D) S. califomica, ventral view. Line scale, 100.0 pm.

167 168

Figure 13. Figure 14. Exemplars of Onthophilinae. (A) Onthophilus sp. (B) Australepierus spinellus. (C) Plagiogramma frontale. (D) Epierus lucidulus ÎEnchson. Line scale, 1.0 mm.

169 170

Figure 14. Figure 15. Exemplars of Onthophilinae. (A) Vuattouxinus borassicola, dorsal habitus, (B) V. borassicola, ventral habitus. (C) Epiechinus kivuensis, dorsal habitus. E. kivuensis, ventral habitus. Line scale, 1 mm.

171 172

• . / /'V- . ,. ^■^• \ & 11® ' - ÆÊ

Figure 15. Figure 16. Antennae of Onthophilinae, dorsal view. (A) Onthophilus striatus (Forster), (B) Vuattouxinus borassicola. (C) Epiechinus rappi. (D) E. rappi^ detail of club sensory plaque. Line scale, 100.0 pm (A, B, C); 10.0 pm (D).

173 174

Figure 16. Figure 17. Antennae of Onthophilinae, dorsal view. (A) Australepieruspunctulipermis. (B) A. punctulipermis club, ventral view. (C) Epierus sp. (D) Plagiogramma schmidti club, ventral view. Line scale, 100.0 pm (A, C); 10.0 pm (B, D).

175 176

Figure 17. Figure 18. Epipharynx of Onthophilinae. (A) Onthophilus kimi. (B) Plagiogramma frontale. (C) Epierus sp. detail of inner brush base and medial area. (D) E. lucidulus Erichson. Line scale, 10.0 pm (A); 100.0 pm (B, C, D).

177 178

Figure 18. Figure 19. Epipharynx of Onthophilinae. (A) Vuattouxinus borassicola. (B) Epiechinus rappi. (C) Australepierus punctulipennis. (D) Australepieruspunctulipennis detail of inner brush base and medial area. Line scale, 10.0 pm.

179 180

s

.Mm

Figure 19. Figure 20. Mandibles of Onthophilinae, dorsal view. (A) Onthophilus kimi, (B) Vuattouxinus borassicola. (C) Epiechinus kivuensis. (D) Australepierus punctulipennis. Line scale, 100,0 pm.

181 182

I

I

Figure 20. Figure 21. Mandibles of Onthophilinae, ventral view. (A) Onthophilus nodatus. (B) O. nodatus ventral prosthecal comb setulae. (C) 0 . nodatus ventral microtrichia. Vuattouxinus ftorosj/co/a ventral prosthecal setulae. Line scale, 100.0 pm (A); 10.0 pm (B, C, D).

183 184

Figure 21. Figure 22. Mandibles of Onthophilinae, dorsal view. (A) Plagiogrcmima frontale, female. (B) Epierus sp. female. (C) Epierus sp. male, right mandible. (D) Same, left mandible. Line scale, 100.0 pm.

185 186

Figure 22. Figure 23. Galea of Onthophillnae, dorsal view. (A) Epierus sp, (B) Epierus. sp., detail of spatulate setae. (C) Australepieruspunctulipemis. (D) A. punctulipennis detail of galea setulae. Line scale, 10.0 pm.

187 188

VI I

Figure 23. Figure 24. Sexual dimorphism in Plagiogramma. (A) P. frontale^ male, (B) P. frontale^ female. Line scale, 1.0 mm.

189 190

» i

Figure 24. Figure 25. Diagnostic features of Plagiogramma and Epierus. (A) Head of P. subtropica, female, showing setigerous tubercle. (B) Head of P. puhifrons, male, showing setigerous patch. (C) E. pulicarius meso-metastemum. (D) P. schmidti meso-metastemum. Line scale, 100.0 pm.

191 192

Figure 25. Figure 26. Exemplars of Tribalinae. (A) Stictostix mormon Lewis. (B) Peploglyptus belfragei (C) P. belfragei, detail of pronotal fossa. (D) Tribalus capensis. Line scale, 1.0 mm (A, B, D); 100.0 pm (C).

193 194

1 r « fe«

Figure 26. Figure 27. Antennae oîAnapleus and Tribalinae, dorsal view. (A) Anapleus marginatus. (B) Stictostix califomica. (fS) Peploglyptus belfragei. (D) Tribalus scaphidiformis. Line scale, 100.0 pm.

195 196

Figure 27. Figure 28. Epipharynx of Dendrophilus and Tribalinae. (A) Dendrophilus opacus. (B) Stictostix califomica. (C) Peploglyptus beljragei. (D) Tribalus scaphidiformis. Line scale, 10.0 pm.

197 198

Figure 28. Figure 29. Mandibles of Anapleus and Tribalinae, dorsal view. (A) Anapleus marginatus. (B) Stictostix califomica. (C) Peploglyptus belfragei. (D) Tribalus scaphidiformis. Line scale, 100.0 pm.

199 200

Figure 29. Figure 30. Exemplars of Tribalinae. (A) Eutribalus agrestis. (B) Idoliapunctistemum Lewis. (C) Paridolia comes (Cooman). (D) Parepierus salvazai. Line scale, 1.0 mm.

201 202

.m a

Figure 30. Figure 31. Antennae of Tribalinae, dorsal \dew, (A) Parepierus sp. (B) Idolia gibba. (C) Eutribalus americanus. (p ) Paridolia Line sp.. scale, 100,0 pm.

203 204

Figure 31. Figure 32. Antennal club of Tribalinae, ventral view. (A) Tribalus scaphidiformis. (B) Eutribalus americanus. (P) Idolia gibba. (p) Paridolia Line scale, sp. 10.0 pm.

205 206

Figure 32. Figure 33. Epipharynx of Tribalinae. (A) Icblia gibba. (B) Parepierus salvazai. (C) Paridolia sp. Eutribalus americanus. Line scale, 10.0 pm.

207 208

Figure 33. Figure 34. Mandibles of Tribalinae, dorsal view. (A) Pctrepierus salvazai (B) Idolia gibba. {C) Eutribalus americanus. (p ) Paridolia Line sp..scale, 100 nm.

209 210

Figure 34. Figure 35. Mandibles of Tribalinae. (A) Eutribalus americanus dorsal detail of prostheca. (B) E. canericanus, ventral view. (C) E. americanus radula detail. (D) I. gibba radula detail. Line scale, 10.0 pm (A, C, D); 100.0 pm (B).

211 212

Figure 35. Figure 36. Diagnostic features of Tribalinae. (A) Eutribalus americanus meso- metathorax. (B) Parepierus sp. meso-metathorax. (C) Idolia gibha meso- metathorax. (D) I. gibba prosternai sculpture. Line scale, A-C = 100.0 pm; D = 10.0 pm.

213 214

Figure 36. Figure 37. Dorsal mandibular microtrichiae of Onthophilinae and Tribalinae. (A) Peploglypius belfragei. (B) Australepieruspunctulipemis. (C) Epierus sp. (D) Idolia gibba. Line scale, 10.0 pm.

215 216

•rWt>-rSG5

Figure 37. Figure 38. Ventral prostheca of Onthophilinae and Tribalinae. (A) Stictostix califomica (B) Tribalus scctphidrformis (C) Australepierus australis (D) Epierus heterognathus. Line scale, 10.0 pm.

217 218

%

Figure 38. Figure 39. Galea of Onthophilinae and Tribalinae. (A) Tribalus scaphidiformis, dorsal view. (B) Stictostix califomica, ventral view. (C) Onthophilus kimi, ventral view. (D) Eutribalus americanus, ventral view. Line scale, 10.0 pm.

219 220

;

m vy-:i

Figure 39. Figure 40. Miscelaneous moutpart detail of Onthophilinae and Tribalinae. (A) Eutribalus americanus palpiter condyle, lateral view. (B) Onthophilus kimi maxillary palpal organ, lateral view. (C) O. nodatus strobilus ventral view. (D) E. americanus teminal labial palpal segment. Line scale, 10.0 pm.

221 2 2 2

I

Figure 40. Figure 41. Cuticular features of Onthophilinae and Tribalinae. (A) Stictostix frontalis prothoracic umbiculate punctures and scales. (B) Peploglyptus helfragei elytral umbiculate punctures, minute scales, and cuticular shagreening. (C) Epiechinus rappi, detail of large elytral scales. (D) E. rappi, cuticular detail showing shagreening. Line scale, 10.0 pm.

223 224

Figure 41. Figure 42. Hind wings of Onthophilinae and Tribalinae. (A) Onthophilus giganteus. (B) Plagiogramma frontale. (C) Epiechinusplanistemus (D) Tribalus scaphidiformis (E) Australepieruspunctulipennis. (F) Eutribalus americanus. Line scale, 0.5 mm.

225 226

B4 R P s t k u u

Figure 42. Figure 43. Mouthparts of Onthophilus kimi. (A) Right maxilla, ventral view. (B) Right maxilla, dorsal view. (C) Maxillary tooth, dorsal view. (D) Mentum, ventral view. (E) Prementum, ventral view (left ligular fringe and right labial palp omitted for clarity). (F) Right labial palp, dorsal view. Line scale, 0.2 mm.

227 228

' me

ce

/

Figure 43. Figure 44. Mouthparts of VuaUotacinus borassicola. (A) Right maxilla, ventral view. (B) Right maxilla, dorsal view. (C) Maxillary tooth, dorsal view. (D) Mentum, ventral view. (E) Prementum, ventral view (left ligular fiinge and right labial palp omitted for clarity). Right labial palp, dorsal view. Line scale, 0.2 mm.

229 230

1

/■ f //

Figure 44. Figure 45. Mouthparts oîEpiechinus rappi. (A) Right maxilla, ventral view. (B) Right maxilla, dorsal view. (C) Maxillary tooth, dorsal view. (D) Mentum, ventral view. (E) Prementum, ventral view (left ligular fnnge and right labial palp omitted for clarity). (F) Right labial palp, dorsal view. Line scale, 0.2 mm.

231 232

Oo!

Figure 45. Figure 46. Mouthparts oî Australepierus punctulipennis. (A) Right maxilla, ventral view. (B) Right maxilla, dorsal view. (C) Maxillary tooth, dorsal view. (D) Mentum, ventral view. (E) Prementum, ventral view (left ligular fiinge and right labial palp omitted for clarityÿ (F) Right labial palp, dorsal view. Line scale, 0.2 mm.

233 234

f

V

Figure 46. Figure 47. Mouthparts of Plagiogramma gentilis. (A) Right maxilla, ventral view. (B) Right maxilla, dorsal view. (C) Maxillary tooth, dorsal view. (D) Mentum, ventral view. (E) Prementum, ventral view (left ligular fnnge and right labial palp omitted for clarity). (F) Right labial palp, dorsal view. Line scale, 0.2 mm.

235 236

Figure 47. Figure 48. Mouthparts of Epierus pulicarius. (A) Right maxilla, ventral view. (B) Right maxilla, dorsal view. (C) Maxillaiy tooth, dorsal view. (D) Mentum, ventral view. (E) Prementum, ventral view (left ligular fiinge and right labial palp omitted for clarity). (F) Right labial palp, dorsal view. Line scale, 0.2 mm.

237 238

&

Figure 48. Figure 49. Mouthparts of Stictostix californica. (A) Right maxilla, ventral view, (B) Right maxilla, dorsal view. (C) Maxillary tooth, dorsal view. (D) Mentum, ventral view. (E) Prementum, ventral view (left ligular fringe and right labial palp omitted for clarity). (F) Right labial palp, dorsal view. Line scale, 0.2 mm.

239 240

^0

I

Figure 49. Figure 50. Mouthparts of Pepîoglypius belfragei. (A) Right maxilla, ventral view. (B) Right maxilla, dorsal view. (C) Maxillary tooth, dorsal view. (D) Mentum, ventral view. (E) Prementum, ventral view (left ligular fringe and right labial palp omitted for clarityÿ (F) Right labial palp, dorsal view. Line scale, 0.2 mm.

241 242

o œ

Figure 50. Figure 51. Mouthparts of Tribalus scaphidiformis. (A) Right maxilla, ventral view. (B) Right maxilla, dorsal view. (C) Maxillary tooth, dorsal view. (D) Mentum, ventral view. (E) Prementum, ventral view (left ligular fiinge and right labial palp omitted for clarity). (F) Right labial palp, dorsal view. Line scale, 0.2 mm.

243 244

y i

Figure 51. Figure 52. Mouthparts oîEutribalus americanus. (A) Right maxilla, ventral view. (B) Right maxilla, dorsal view. (C) Maxillary tooth, dorsal view. (D) Mentum, ventral view. (E) Prementum, ventral view (left ligular fiinge and right labial palp omitted for clarity). (F) Right labial palp, dorsal view. Line scale, 0.2 mm.

245 246

i r

Figure 52. Figure 53. Mouthparts oîParepierus amandus. (A) Right maxilla, ventral view. (B) Right maxilla, dorsal view. (C) Maxillaiy tooth^ dorsal view. (D) Mentum, ventral view. (E) Prementum, ventral view (left ligular fiinge and right labial palp omitted for clarity). (F) Right labial palp, dorsal view. Line scale, 0.2 mm.

247 248

$

î/J.

Figure S3. Figure 54. Mouthparts o f Paridolia laevidorsis. (A) Right maxilla, ventral view. (B) Right maxilla, dorsal view. (C) Maxillary tooth, dorsal view. (D) Mentum, ventral view. (E) Prementum, ventral view (left li^lar fringe and right labial palp omitted for clanty). (F) Right labial palp, dorsal view. Line scale, 0.2 mm.

249 250

/ f

Figure 54. Figure 55. Mouthparts oîldolia gibha. (A) Right maxilla, ventral view. (B) Right maxilla, dorsal view. (C) Maxillary tooth, dorsal view. (D) Mentum, ventral view. QE) Prementum, ventral view (left ligular fiinge and right labial palp omitted for clarity). (F) Right labial palp, dorsal view. Line scale, 0.2 mm.

251 252

Figure 55. Figure 56. Male genitalia of Onthophilus sp. (A) Eighth abdominal segmant, dorsal view. (B) Same, ventral view, (C) Same, lateral view. (D) Ninth abdominal segment, dorsal view. (E) Same, ventral view. (F) Same, lateral view. (G) Aedeagus, dorsal view. (H) Same, ventral view. (H) Same, lateral view. Line scale, mm.

253 254

ex te V /

ex

M

Figure 56. 255

Figure 57. Cladogram of onthophiline genera, based on both adult and larval characters.

Dendrophilus

Onthophilus

Viiattouxinus

Epicchinus

Australepinis

Plagiogranuna

Epierus

Figure 57. 256

Figure 58. Cladogram of tribaline genera, based exclusively on adult characters.

Anapleus

Stictostix

Peploglyptus

Tribulus

Idolia

Paridolia

Parepierus

Eutribulus Figure 58. CHAPTER V

CHAETOTAXY OF THE IMMATURE STAGES OF THE NEW WORLD IDOLLi GENUS GROUP (COLEOPTERA; HISTERIDAE) BASED ON IDOLU GIBBA LEWIS AND EÜTRIBALÜS AMERICANUS (LECONTE).

Introduction Adults members of the Idolia genus group are fungal spore specialists and are associated with rotting tree trunks. Little ecological or biological information is av^able for members of this tribe (see Chapter 4). The 7 described species o f Idolia are mainly Neotropical with 1 species, I. gibba, extending into Nearctic Mexico. The 18 described species oîEutribalus are mainly Oriental and Afrotropical in distribution with a single Nearctic species, Eutribalus americanus. Nomenclatural systems for describing and homologizing setae and pores of larval (Chapter 1) and pupal Histeridae (Chapter 2) were recently developed and are applied to Idolia and Eutribalus for the first time. Parental stock for Idolia gibba was obtained in March, 4 km. east of Palo Seco, Chiapas, Mexico, beneath the loose bark of a fallen tree. Larvae of Eutribalus americanus were extracted fi’om debris within rotting beech stumps on the West Campus of The Ohio State University, Columbus, OH. Voucher specimens were deposited in the larval collections of The Ohio State University, the Field Museum of Natural History, Chicago, and the collection of PWK. Methods for studying larvae, and terminology for naming and homologizing setae and pores, follow those used in Chapter 1. Exact measurements used to delimit setal size classes correspond to those used for Onthophilus nodatus in Chapter 1. Larvae o f I. 257 258 scitua Lewis were examined for any significant variations in setal and pore distribution within 7db//a. Pupal setal nomenclature follows that used in Chapter 2. Drawings are based on live specimens because of the distortion that accompanies ethanol preservation. Laboratory rearing methods follow that described for Epierus divisus Marseul in Chapters.

Results Description of Idolia genus group larvae. Epicranial suture lyriform with short stem, medial ecdysial line absent; antennal segment 2 with 1 dorsal and 2 lateral setiform sensilla and 2 sensoria (SEl, 2) on distal membrane; sensorium SEl larger than SE2; antennal segment 3 with setiform terminal sensilla; mandible lacking mola and with a small tooth below retinaculum; stipes with 4 short flexible setae on dorsal distal membrane; proximal maxillary palpal segment with 2 flexible setae dorsally on membrane; second maxillary palpal segment with pore MXg enlarged; apical membrane of mentum without lobes; dorsal labial surface and lateral lobes unarmed; LAI seta vestigial; pore LAc present dorsally on labial palp; nasale with 4 flexible non-socketed gFR setae; mesothoracic seta TE 10 short; seta ST45 present on meso- and metastemite; meso- and metathoracic seta ST46 of medium length; metathoracic lateral humeral tergites vestigial; minute metathoracic seta ST29 off of metastemite; with some short setae present; metathoracic intertergites absent; thoracic interstemites absent; abdominal segments I-Vni with setae TE14,15, and 16 on separate sclerites; abdominal segments I-VŒ with seta STS 8 on separate sclerite; abdominal segments I-Vm with seta PL25 and ST40 of medium length; abdominal segments IX with seta TEl and pores TEf,g absent; abdominal segment IX with seta TE16 and 21 on separate sclerites; seta UGll borne at apex of urogomphus; pygopod seta PP2 of medium length; all abdominal segments with intertergites and 259 interstemites lacking; membranous cuticle of thorax and all abdominal segments with extensive covering of chalazae, denticles absent; malpighian tubules and hind gut appear white, probably due to accumulated uric acid; sturdy silk-lined cocoons produced by pre­ pupa. Diagnosis of Idolia genus group larvae. Known larvae of Idoliini can be distinguished from known histerid larvae by the following combination of characters: antennal segment 2 with 3 setiform sensilla; pore LAc present dorsally on labial palp; seta ST45 present on meso- and metastemite; and seta UGll bome at apex of urogomphus.

Key to the larvae of New World Idolia genus group. 1 Apical sensilla of maxilla nearly as long as last palpal segment; thoracic posterior pleurites absent; lateral stemites absent ...... Idolia r Apical sensilla of maxilla much shorter than last palpal segment; thoracic posterior pleurites well developed; lateral stemites present ...... Eutribalus

Description of Idolia genus group pupae. Head with galeal scaffold (GS) enlarged and protuberant; labral setae absent; pronotum with lateral pronotal group restricted to posterior half of lateral margin; posterior pronotal group with 5-7 setae in on each side; interior pronotal setae lacking; mesonotum generally with 5 scutal setae on each side and lacking scutellar setae; hind wings lacking tubercle; urogomphi relatively short. Diagnosis of Idolia genus group pupae. Pupae of Idoliini can be distinguished from known tribaline pupae by the protuberant galeal scaffold

Key to the pupae of New World Idolia genus group. 1 Head with ventrofrontal and vertical epicranial setae; pronotum lacking anterior pronotal setae; pygidial setae present ...... Idolia 1* Head with ventrofrontal and vertical epicranial setae lacking; pronotum with anterior pronotal setae; pygidial setae absent ...... Eutribalus 260

Description First Instar Idolia gibba. (Figs. 59-66) Head. Head capsule width: 0.27 ± 0.01 mm. (X ± SD; n = 6). One stemmata on each side. Frontale (Figs. 59A, 61 A) with 10 setae (FR2-FR11) on each side. Setal lengths: vestigial (FR3-6), minute (FR7, 9,11), short (FRIO), medium (FR2, 8). Nasale with 2 asymmetrical weakly protuberant teeth. Three short subnasale setae (SF) present. Parietal (Figs. 59, 62A) setal lengths: minute (PA16,17,18,19), short (PA12, 13, 20, 24, 26,28), medium (PA15,27, 28, 29) long (PA14,21,22, 23,25,30). Antenna. (Fig. 60A-B) Segment 1 about twice as long as wide. Segment 2 about as long as wide; sensillum on distal membrane near base of SEl extremely short; remaining sensilla about as long as terminal antennal segment. Segment 3 with longest member of terminal pair of sensilla longer than last antennal segment, shorter member slightly shorter than peripherals which are nearly as long as last antennal segment. Mandible (Fig. 60C) with medium length laterobasal seta (MNl). Penicillus (PE) setae plumose. Maxilla (Fig. 60D-E). Stipes setal lengths: minute (MXl, 6), short (MX5), medium (MX2, 4), long (MX3). gMXl composed of numerous flexible barbed setae; gMX2 composed of 5-6 minute rigid setae with sockets. Proximal palpal segment setal lengths: minute (MX9), medium (MX7, 8). Segment 4 with 10 pore-like sensilla and apical sensilla basiconica of varying lengths; terminal pair of sensilla basiconica very long, nearly as long as terminal palpal segment. Labium. Prementum (Fig. 60F-G) about as long as wide. Setal lengths: vestigial (LAI), minute (LA3), short (LA2). LA3 arising laterad of 2 short peg-like sensilla in dorsoapical premental membrane. Second palpal segment with 5 circular sensilla distributed over surface. Villiform sensilla basiconica of varying lengths arising from apex of palpus; two are longer than sensillum digitiformium, remainder shorter. Thorax. Prothorax (Figs. 62B, 63A). Notum with 23 setae (TEl-13, 14, 16,17, 18-21, 23,24, 27) (shield with subset [TEl-14, 16, 17, 19]), and 13 pores (TEa-i, k-m) 261

(shield with subset [TEa-i, 1, m]) on each side. Setal lengths: minute (TEIO, 18,20,21, 24, 27), short (TEl, 6, 7, 8, 12, 14, 16, 17, 23), medium (TE2, 3,4,11, 13), long (TE5, 9, 19). Posterior pleurite absent; single minute pleural seta PL28 present. Epistemum and epimeron without setae. Prestemum: Lateral prestemite extending posteriorly beyond prestemite by 30% its length; prestemite roughly triangular; prestemite setal lengths: minute (PRl-5, 8,9), short (PR7), medium (PR6). Prostemite oblong ovate and about twice as long as wide, with smooth cuticle centrally. Setal lengths: short (ST30, 44), medium (ST46). Precoxite very small with 3 minute setae (ST31,32,36), seta ST31, ST32 and ST36 subequal. Mesothorax (Figs. 62B, 63). Notum setal lengths: short (TE6, 7, 8,10), medium (TE4, 5, 9,11), long (TE5). Intertergites elongate-ovoid. Lateral tergite mid-sized, ovoid with 5 setae (TE16-20) Setal lengths: minute (TEl5, 18, 21), short (TE16, 17), medium (TE20), long (TE19). Two isolated, minute laterotergal seta, TE15 and TE21 present. TEIS located just dorsad lateral tergite and TE21 located just anterior to lateral tergite. Posterior pleurite absent; posteropleural setae PL27 and 28 arising from cuticle of pleural region in close proximity to each other. Setal lengths: minute (PL28), short (PL27). Anterior pleurite minute and bearing 2 minute setae (PL23,24). Anterior stemite small with single minute seta (ST36). Epistemum with 2 minute setae (ST34, 35). Epimeron with single minute seta (ST40). Precoxite absent; both precoxal setae ST32 and ST31 minute and arising from cuticle in close proximity to each other. Lateral stemite absent. Mesostemite large and triangular; mesostemite setal lengths: minute (ST29), short (ST30,44), medium (ST45,46). Metathorax (Figs. 64, 65A). Notum setal lengths: short (TE6,10), medium (TEll), long (TE9). Dorsal humeral setae TEl and TE2, arising from humeral region in close proximity to each other and to pore TEb. Minute lateral humeral setae TE12, TE13 and TE14 arising anterior to lateral tergite in close proximity to each other. Dorsolateral 262 tergite mid-sized and with 4 setae (TE4, 5,7, 8). Setal lengths: short (TE7, 8), medium (TE4), long (TE5). Lateral tergite and isolated laterotergal setae nearly identical to homologs on mesothorax. Posterior pleurite absent; posteropleural setae similar to homologs on mesothorax. Anterior pleurite absent; three minute anteropleural setae (PL23,24, 25). Anterior sclerites absent; two isolated, minute anterostemal setae (ST36) and (ST37) present. Epistemum and epimeron similar to homolog on mesothorax. Precoxite absent; precoxal setae similar to homologs on mesothorax. Metastemite shorter and more quadrate than preceding homolog and with 4 setae (ST30,44-46) per side. ST29 on vestigial stemite anteriorad to metastemite on each side. Metastemal setal lengths identical to those of homologs on mesothorax. Metathoracic Leg 61B-C). Coxa setal lengths: minute (COl-10,12) and short (COll, 13-19). Trochanter with 6 setae (TRl-6); setal lengths: minute (TRl- 6). Femur setal lengths: minute (FBI, 2,4-6), short (FE3, 7), medium (FE8). Tibia with 17 setae (TIl-17) and 2 pores (TIa, b). Setal lengths: vestigial (TI7, 9,14,16), minute (TIl- 6,10,12) short (TI8,11,13,15,17). Tarsungulus with 2 short apically barbed setae (TAl, 2). Abdomen. Segment I. Tergum (Figs. 64A, 65 A) with 19 setae (TEl, 3,4-11, 13- 21) and 15 tergtes (many reduced) on each side. Setal lengths: minute (TEl, 8,10,13- 15, 17, 18, 21), short (TE4, 6, 7, 11), medium (TE3, 16,20), long (TE5, 9, 19). Tergites: TEl on vestigial sclerite. TE3 on minute sclerite. Egg burster on rectangular mid-sized sclerite. TE8-11 sharing mid-sized, circular central sclerite. TE6, 7 sharing small circular sclerite. TE4 on minute sclerite. TE14,15 each on vestigial sclerites. TE16 on minute sclerite. TE5 on small circular sclerite. TE19, 20 sharing small sclerite. TE13, and 21 each on vestigial sclerites. TE 18 on minute sclerite. Ampulla poorly developed. Pleuron (Fig. 65A) with 7 setae (PL22-28) and 4 pleurites. Setal lengths: minute (PL22, 23,26, 28), short (PL24, 27), medium (PL25). Pleurites: PL22-25 on mid-sized anterior 263 pleurite. PL26,27 sharing small pleurite. PL28 on vestigial pleurite. Small dorsal pleurite present; lateral and ventral pleurites absent. Sternum (Fig. 62B) with 16 setae (ST29-36, 38-41,43-46) and 13 stemites (many reduced) on each side. Setal lengths: minute (ST29, 32, 34-36, 39), short (ST30,31,33, 38,41,44,45), medium (ST40,46), long (ST43). ST29, 30, 32, 34-36 each on vestigial sclerites. ST31, 33,38 each on minute sclerites. ST39-40 sharing small sclerite. ST41,43 each on small sclerites. ST44, 45 sharing mid-sized sclerite. ST46 on large ovoid medial stemite. Prolegs poorly developed. Segments II-VIII similar to segment I but with better-developed prolegs. SegmentlX. Tergum (Figs. 65B, 66A) with 8 setae (TE3, 5, 9, 16-19,21) and 8 sclerites per side. Setal lengths: minute (TE3,17), short (TE18,21), medium (TE9,16), long (TE5,19). Tergites: TE3 on vestigial sclerite. TE9 on minute sclerite. TE18 on minute sclerite. TE19 and PL28 share mid-sized sclerite representing the fusion of a tergite and pleurite. TB5,16, 17, and 21 each on small circular sclerites. Pleuron (Figs. 65b, 66A-B) with 2 sclerites per side. Setal lengths: short (PL24,28), medium (PL27) long (PL25, 26). Pleurites: PL24,25 sharing mid-sized sclerite. PL26, 27 borne at apex of weakly protuberant paragomphus. Sternum (Figs. 65A, 66B) with 7 setae (ST29,30, 34,36, 38,40,46) and 6 sclerites per side. Setal lengths: minute (ST29, 34, 36), short (ST30, 38), medium (ST46), long (ST40). Stemites: ST30 and 36 each on minute sclerites. ST29 and 34 each on vestigial sclerites. ST46 on small round sclerite. ST38 and 40 sharing small round sclerite. Urogomphi (Figs. 65B, 66A-B). Setal lengths: minute (UGl), short (ÜG2,4, 5, 7), medium (UG3, 6, 9,11), long (UG8,10,12,13). Sclerites: UGl-7 each sharing mid­ sized sclerite. Pygopod (Figs. 65B, 66B). Setal lengths: short (PPl, 3, 4), medium (PP2, 5-7). Sclerites: PPa on vestigial sclerite. PPl-3 sharing circular, mid-sized, sclerite. PP4 and 5 264 sharing mid-sized sclerite. PP6 and PP7 each on small circular sclerites. PPa on minute sclerite dorsad to PP7. Second Instar Head capsule width: 0.41 ± 0.01 mm (n = 3). Structurally quite similar to first instar. Differences include a slightly different head capsule shape, more elongate palpi and antennae, and sclerotized cuticle on mesothorax. Pupa [based on 2 specimens] (Fig. 67). All setae tapered. Head. Four short epicranial setae on each side; 2 mediofiontal setae (MF), 2 in ocular epicranial group (gOE) and single vertical epicranial (VE). Single short ventral fi*ontal seta (VF) on each side. Labral papillae distinct. Thorax. Pronotum with 8-13 short setae on each side, 2-5 in lateral pronotal group (gLN), 5-7 in posterior pronotal group (gPN). Anterior pronotal setae absent. Mesonotum generally with 5 short setae on each side, all belonging to scutal group (gSC). Metanotum lacking setae. Elytron relatively smooth and with 2-3 short setae (gEL). Abdomen. Tergites 1-6 each with 2-3 short tergal setae (TE) and single short laterotergal seta (LT) on each side. Single short laterostemal seta (LS) on abdominal segments 1-6. Pygidium generally with 2 setae on each side. Description First Instar Eutribalus americanus. (Figs. 68-75) Head. Head capsule width: 0.27 ± 0.01 mm. (n = 6). Stemmata absent. Frontale (Figs. 68A, 70A) with 11 setae (FRl-FRll). Setal lengths: vestigial (FR3,4, 6, 7, 9,11), minute (FRl, 5,10); short (FR2); medium (FR8). Nasale with 2 indistinct asymmetrical teeth. Short and minute subnasale setae (SF) present. Parietal (Figs. 68, 70A) setal lengths: minute (PA16-19,24, 28), short (PA12, 15,20, 26), medium (PA13,23,29, 30), long (PA14, 21, 22,25,27). Antenna (Fig. 69A-B) Segment 1 about 4 times longer than wide. Segment 2 about twice as long as wide and with sensillum on distal membrane near base of SEl extremely short; remaining sensilla about half as long as terminal antennal segment. Segment 3 with longer member of terminal pair nearly as long as terminal antennal segment, shorter member of terminal pair slightly shorter than peripheral sensilla which are 265 about as long as width of terminal antennal segment. Mandible. (Fig, 69C ) with short laterobasal seta (MNl). Penicillus setae (PE) apically branched. Maxilla (Figs. 69D-E). Stipes setal lengths; minute (MXl, 5, 6), short (MX2), and medium (MX4,7). gMXl consisting of numerous flexible mostly un-branched setae; gMX2 consisting of 6 minute rigid setae with sockets. Proximal palpal segment setal lengths: minute (MX9); short (MX8), medium (MX7). Segment 4 with 18 pore-like sensilla and apical villiform sensilla basiconica of varying lengths; with one slightly longer than sensillum digitiformium. Labium. Prementum (Figs. 69F-G) about twice as long as wide and with 3 setae (LAI-3), 3 pores (LAa-c), and numerous sensilla. Setal lengths: minute (LAI, 3), short (LA2). Two short peglike sensilla flanking seta LA3 in dorsodistal membrane. Second palpal segment with 10 circular sensilla distributed over surface. Villiform sensilla basiconica of varying lengths arising from palpal apex, 2 slightly longer than sensillum digitiformium; remainder shorter. Thorax. Prothorax (Figs. 71B, 72). Notum with 20 setae (TEl, 2,4-13, 16,17, 19-21,23, 24, 27) (shield with subset [TEl, 2,4-13, 16, 17, 19]), and 13 pores (TEa-j, k- m) (shield with subset [TEa-j]) on each side. Setal lengths: vestigial (TE21), minute (TEl, 6-8, 10, 12, 16,24, 27), short (TE4, 11, 17,20, 23), and medium (TE2, 5, 9,13, 19). Epistemum and epimeron without setae. Posterior pleurite mid-sized and with single minute seta (PL28). Prestemum: both prestemite and lateral prestemite well-developed, lateral prestemite extending posteriorly beyond prestemite by 20% its length; prestemite roughly quadrate with sides slightly convergent. Prestemite setal lengths: minute (PRl-4, 7-9), short (PR5, 6). Prostemite diamond-shaped, about as long as wide, with smooth cuticle centrally. Prostemite setal lengths: minute (ST30), short (ST44), medium (ST46). Lateral stemite small and circular: precoxite very small with 3 minute setae (ST31, 32, 33), seta ST31, longer than ST32 and ST33 subequal. 266

Mesothorax (Figs. 7 IB, 72), Notum Setal lengths: minute (TE6), short (TE4,7, 8, 10,11), medium (TE5, 9). Intertergite broadly oval. Lateral tergite large, oval, with 7 setae (TE15-21) and single pore (TE)). Setal lengths: minute (TEl5,21), short (TE16, 17,18,20), medium (TE19). Posterior pleurite large, narrowly rectangular, with 2 setae (PL27,28); both setae arising from posterior half of sclerite. Setal lengths: minute (PL28), short (PL27). Anterior pleurite half as large as posterior pleurite and surrounding thoracic spiracle, with 2 minute setae (PL23,24). Anterior stemite small with single minute seta (ST36). Epistemum with 2 minute setae (ST34, 35). Epimeron with single minute seta (ST40). Precoxite subdivided; mesal section mid-sized, each with 1 short seta (ST31); dorsal section minute, each with minute seta (ST32). Lateral stemites ovoid and slightly less than half as long as mesostemite. Mesostemite large and triangular. Mesostemite setal lengths: minute (ST29), short (ST 30, 44), medium (ST45, 46). Metathorax (Figs. 73, 74B). Notum setal lengths: short (TE6,10), medium (TEll), long (TE9). Dorsal humeral tergites small; each with 1 minute setae (TE2) and 1 pore (TEb). TEl setae minute; each on vestigial tergite anterior to dorsal humeral tergite. Lateral humeral setae (TE12, 13, 14) minute. Dorsolateral tergite with 3 setae (TE4, 5, 8) and 2 pores (TEe, i). Setal lengths: short (TE8), medium (TE4), long (TES). Lateral tergite nearly identical to homolog on preceding segment. Posterior pleurite somewhat shorter but otherwise similar to mesothoracic homolog. Anterior pleurite smaller but otherwise similar to anterior pleurite on mesothorax; each bearing minute (PL23) and short (PL24) setae. Anterior sclerites each with minute seta (ST36) and (ST37). Epistemum similar to that of the mesothorax. Epimeron with minute setae (ST40). Precoxite, and lateral stemites similar to preceding homologs. Metastemite shorter and more quadrate than mesothoracic metastemite. ST29 on minute stemite anteriorad to metastemite on each side. Metastemal setal lengths identical to those of mesothorax. 267 MetathoracicLeg(B\g.lO'Q-C). Coxa setal lengths: minute (COl-10,12) and short (COll, 13-19). Trochanter with 6 setae (TRl-6); setal lengths: minute (TRl-3, 5, 6), short (TR4). Femur; setal lengths: minute (FEl, 2,4, 6, 7), short (FE3, 5, 8). Ubia with 16 setae (TIl-11, 13-17) and 2 pores (TIa, b). Setal lengths: minute (TI2,4, 5, 7, 9, 15,17) short (Til, 3,6, 8, 11,13,14,16). Tarsungulus with 2 short setae (TAl, 2). Abdomen. Segment I. Tergum (Figs. 73 A, 74A) with 19 setae (TEl, 3,4-11,13- 21), 5 pores (TEb, e, f, g, j), and 15 tergtes on each side. Setal lengths: minute (TEl, 13-15, 18), short (TE3, 6, 8, 21), medium (TE4,7, 10, 11, 16, 17, 20), long (TE5, 9,19). Tergites: TEl on minute sclerite. TE3 on small sclerite. Egg burster on ovoid mid-sized sclerite. TE8-11 sharing large rectangular central sclerite. TE6, 7 sharing elongate-ovoid mid-sized sclerite. TE4 on circular, mid-sized sclerite. TE 14 and 15 each on vestigial sclerites. TE 16 on small circular sclerite. TE5 on mid sized circular sclerite. TE 19, 20 sharing mid-sized circular sclerite. TE13 on vestigial tergite. TE18, and 21 each on minute sclerites. Ampulla weakly developed. Pleuron (Fig. 74A with 7 setae (PL22-28) and 4 pleurites. Setal lengths: minute (PL22,23,26), short (PL24), medium (PL25, 27, 28). Pleurites: PL22-25 on mid-sized, circular anterior pleurite. PL26, 27 sharing mid­ sized, circular pleurite. PL28 on mid-sized ovoid pleurite. Mid-sized dorsal and lateral pleurite present; ventral pleurites absent. Sternum (Fig. 73B, 74A) with 16 setae (ST29- 36,38-41,43-46) and 10 stemites on each side. Setal lengths: minute (ST29, 32, 34-36), short (ST30,31, 33, 38, 39,41,44,45), medium (ST40,43,46). ST29 on minute sclerite. ST30, 31 each on small, ovoid sclerites. ST32,33 sharing small ovoid sclerite. ST34, 35 each on minute sclerite. ST36 on small, ovoid sclerite. ST38 on small ovoid sclerite. ST39-40 sharing small, ovoid sclerite. ST41,43 each on small, somewhat rounded sclerites. ST44,45 sharing mid-sized, ovoid sclerite. ST46 on large medial stemite. Prolegs fairly welldeveloped. 268 Segments II-VIII like segment I but with better-developed prolegs and well- developed, protuberant ampullae. SegmentlX. Teigum (Figs. 74B, 75A) with 8 setae (TE3, 5, 9,16-19,21), and 7 tergites on each side. Setal lengths: minute (TE3,17), short (TE18), medium (TE5,9, 16, 21), long (TE19). Tergites: TE3 on minute sclerite. TE9 on ovoid mid-sized sclerite. TË16, 19, and 21 each on rounded mid-sized sclerites. TE17 on small round sclerite. TE5 and 18 sharing large, ovoid sclerite. Pleuron (Fig. 74B, 75) with 3 sclerites. Setal lengths: short (PL24,28), medium (PL26) long (PL25,27). Pleurites: PL28 on small, circular sclerite. PL24,25 sharing round, mid-sized sclerite. PL26,27 bom at apex of weakly protuberant paragomphus. Sternum (Figs. 74B, 75B) with 7 setae (ST29, 30,34, 36, 38,40,46) and 5 sclerites. Setal lengths: minute (ST29, 30,34, 36), short (ST38, 46), medium (ST40). Stemites: ST36 on small, round sclerite. ST34 on vestigial sclerite. ST30,46 each on mid-sized, somewhat quadrate sclerites. ST38,40 sharing mid-sized sclerite. ST39 on small sclerite. Urogomphi (Figs. 74B, 75). Setal lengths: minute (UGl, 4, 11), short (UG2, 3, 5, 7), medium (UG6, 9, 10), long (UG8,12, 13). Sclerites: UGl-4 each sharing mid-sized sclerite; UG5-7 sharing mid-sized sclerite. Pygopod (Figs. 74B, 75B). Setal lengths: minute (PPl), short (PP4, 6), medium (PP2, 3, 5, 7). Sclerites: PPl-3 sharing mid-sized sclerite. PP4,5 sharing mid-sized sclerite. PP6 and PP7 each on small, round sclerites. PPa on minute sclerite dorsad to PP7. Second Instar. Head capsule width: 0.64 + 0.03 mm (n = 5). Structurally quite similar to first instar. Differences include a slightly different head capsule shape, more elongate palpi and antennae and darker pigmentation in head capsule and some other sclerotized cuticle. 269 Pupa [based on 2 specimens] (Fig. 76). All setae tapered. Head. Three short epicranial setae on each side; single mediofrontal seta (MF), and 2 ocular epicranial group (gOE). Vertical epicranial setae absent. Ventral frontal setae absent. Labral papillae distinct. Thorax. Pronotum with 10-14 short setae on each side, 1-3 in anterior pronotal group, 4 in lateral pronotal group (gLN), and 5-7 in posterior pronotal group (gPN). Interior pronotal setae absent. Mesonotum generally with 5 short setae on each side, all belonging to scutal group (gSC). Metanotum lacking setae. Elytron relatively smooth and with 2 short setae (gEL). Abdomen. Tergites 4-6 each with 3 short tergal setae (TE) and tergites 3 and 4 with single short laterotergal seta (LT) on each side. Single short laterostemal seta (LS) on abdominal segments 2-6. Pygidial setae absent.

Discussion These are the first larval descriptions for the Idolia genus group. Larvae of the two other genera included in this tribe in Chapter 4, Parepierus and Paridolia, remain unknown. Numerous synapomorphies found in the larvae and pupae o îIdolia and Eutribalus support both the monophyly of this genus group and its derived position in the Tribalinae (see Chapter 4). Larval synapomorphies include: setiform sensilla of the antennae; mentum membrane without lacking lobe; a glabrous prementum dorsum; LAI seta vestigial; a unique pore (LAc) on the labial palpi; an apparently unique seta ST45 present on meso- and metastemite; vestigial metathoracic lateral humeral tergites; tibiae with short apical setae; no metathoracic intertergites or thoracic interstemites; abdominal segments I-Vm with setae TE14,15, and 16 on separate sclerites; the same segments with seta ST38 isolated on a separate sclerite; abdominal segment IX lacking pores and with seta TE16 and 21 each on separate sclerites; urogomphal seta UGll apical; no intertergites or interstemites on any abdominal segments and membranous cuticle of thorax; and all abdominal segments with extensive covering of chalazae. Pupal 270 synapomorphies include: the presence of a protuberant galeal scaffold, pronotal lateral pronotal setae restricted to posterior half of lateral margin; and mesonotum with at least 4 scutal setae per side. Idolia larvae possess a number of apparent autapomorphies that readily distinguish them from larval Eutribalus and known larvae of Onthophilinae. Most notable among these is a reduction of most larval sclerites and an nearly complete elimination of others. Another distinctive autapomorphy o îIdolia are the very long terminal sensilla basiconica of the last maxillary palpal segment. Idolia larvae have more thoracic setae than those of Eutribalus including a pronotal seta (TE14) not known from other histerid larvae. Another thoracic seta present in Idolia and absent in Eutribalus (mesothoracic TE3) also occurs in Onthophilus and Dertdrophilus. An interesting autapomorphic setal arrangement in larval Idolia not yet discovered elsewhere in the Histeridae is the occurrence of 4 setae on the metathoracic dorsolateral tergite. Eutribalus larvae have relatively few autapomorphies; among them was the absence of a pronotal seta (TE3) and extreme reduction of the majority of frontale setae. Descriptions of larvae and pupae o î Idolia dxià Eutribalus have provided an important first look at the immature stages of the subfamily Tribalinae. Like the adults, larval and pupal Idoliini have many derived characters. The phylogenetic analysis of the Tribalinae based exclusively on adult characters (see Chapter IV) hypothesized a sister group relationhip for Eutribalus and Parepierus. Therefore I would predict that the undiscovered larvae and pupae of Parepierus will share more characters with Eutribalus than larvae of other genera of the Idolia genus group. The ultimate discovery of immature stages of the more basal tribaline taxa, Stictostix, Peploglyptus, and Tribalus will undoubtedly provide critical information to further test the hypothesized relationships established via adult character analysis. 271

Figure 59. Head capsule of Idolia gibba, first instar larva. (A) Dorsal view. (B) Ventral view, (“flexible” setae omitted). Line scale 0.2 mm.

271 272

Figure 59. 273

Figure 60. Mouthparts and antennae of Idolia gibba, first instar larva. (A) Left antenna, dorsal view, (B) Left antenna, ventral view, (C) Left mandible, dorsal view, (D) Right maxilla, dorsal view, (E) Right maxilla, ventral view, (F) Prementum, ventral view, (G) Prementum dorsal view. Line scale, 0,1 mm. Shading indicates membrane.

273 274

1M 1 •2

O"

lOO

o«

'o

OMXI

>gMX2

Figure 60. 275

Figure 61. Frontale and metathoracic leg of Idolia gibba, first instar larva. Detail of fi"ontale. (A) Metathoracic leg, anterolateral view. (B) Metathoracic leg, posterolateral view. Line scale, 0.1 mm.

275 276

oe

oc

ao

bo

Figure 61. 277

Figure 62. Head capsule and pro- and mesothorax o î Idolia gibba, first instar larva. (A) Head capsule, lateral view. (Flexible setae omitted). (B) Pro- and mesothorax, lateral view. Line scale, 0.2 mm.

277 278

m#

• 8

' 2 7 '2 8 •;

ri •

Figure 62. 279

Figure 63. Pro- and mesothorax of Idolia gibba, first instar larva. (A) Dorsal view. (B) Ventral view. Line scale, 0.2 mm.

279 îÆfjé. a /

(g 3 O 8

k.t

« y

K) 00 o 281

Figure 64. Metathorax and first abdominal segment oî Idolia gibha, first instar larva. (A) Dorsal view. (B) Ventral view. Line scale, 0.2 mm.

281 282

V--S-^ïï

Figure 64. 283

Figure 65. Idolia gibba, first instar iarva. (A) Metathorax and first abdominal segment, lateral view, (B) Abdominal segments IX and X, lateral view. Line scale, 0.2 mm.

283 284

*»* t4

m m m •I-.'.-. i m m »

J ' ' '

B

Figure 65. 285

Figure 66. Idolia gibba, first instar larva. (A) Abdominal segments IX and X, dorsal view. (B) Abdominal segments IX and X, ventral view. Line scale, 0.2 mm.

285 286

Figure 66. 287

Figure 67. Setal maps of IdoUa gibba pupae. (A) Dorsal and lateral view composite. (B) Lateral view. Line scale, 1.0 mm.

287 288

Figure 67. 289

Figure 68. Head capsule of Eutribalus americanus, first instar larva. (A) Dorsal view. (B) Ventral view, (“flexible” setae omitted). Line scale 0.2 mm.

289 290

d" f,

•c

Vl2

og

\28

Figure 68, 291

Figure 69. Mouthparts and antennae oîEutrihalus americanus, first instar larva. (A) Left antenna, dorsal view. (B) Left antenna, ventral view. (C) Left mandible, dorsal view. (D) Right maxilla, dorsal view. (E) Right maxilla, ventral view. (E) Prementum, ventral view. (G) Prementum dorsal view. (Shading as in Figure 58). Line scale, 0.1 mm.

291 292

r

Ob ®Q> o#

oc A

o#

OS gMXl

O b

Figure 69. 293

Figure 70. Frontale and metathoracic leg oîEutribalus americanus, first instar larva. Detail of fi’ontale. (A) Metathoracic leg, anterolateral view, (B) Metathoracic leg, posterolateral view. Line scale, 0.1 mm.

293 294

ta

Ob

od 06 I

h oc

17J<

Figure 70. 295

Figure 71. Head capsule and pro- and mesothorax oî Eutribalus americanus, first instar larva. (A) Head capsule, lateral view. (Flexible setae omitted). (B) Pro- and mesothorax, lateral view. Line scale, 0.2 mm.

295 296

2 9 os

m M M

m s m I

Figure 71. 297

Figure 72. Pro- and mesothorax oîEutrihalus americanus, first instar larva. (A) Dorsal view. (B) Ventral view. Line scale, 0.2 mm.

297 298

— în-y-f—

m

'3 0 ill

Figure 72. 299

Figure 73. Metathorax and first abdominal segment o f Eutribalus americanus, first instar larva. (A) Dorsal view. (B) Ventral view. Line scale, 0.2 mm.

299 300 m g mmm. r n

X

5*4 P46

Av-Ÿ

Figure 73. 301

Figure 74. Eutribalus americanus, first instar larva. (A) Metathorax and first abdominal segment, lateral view. (B) Abdominal segments IX and X, lateral view. Line scale, 0.2 mm.

301 302

t S

Figure 74. 303

Figure 75. Eutribalus americanus, first instar larva. (A) Abdominal segments IX and X, dorsal view. (B) Abdominal segments IX and X, ventral view. Line scale, 0.2 mm.

303 304 il

U012. [UGll 1)013

iSS

CUG9

Figure 75. 305

Figure 76. Setal maps of Eutribalus americanus pupae. (A) Dorsal and lateral view composite. (B) Lateral view. Line scale, 1.0 mm.

305 306

Figure 76. APENDK A - LARVAL CHARACTERS

1) S tw th tfk n a ln la ------H) 0) Ibniâul MnsDum long M 3td antennal legmant; a*het i|ik«l Hmilk 0) Pieniontun eboul M irids u long. notdy u long M SBI.(F) 1) Piementummderlhenlong. 1) Ttamiu] MtMSIum mudi ibocter tfaBi M aitannil wgioont; ottur apiol muillimuchihottoclhinSBl arulcngu3idantennilMgmait;odicr 14) AaaaataraafdaaaalpraananWanaOacn. •pint iemilli KtUbon Old longer Hun SBt.(A) 0) Donalbbialaui&oa toothed and or tubcrculate.(P) 1) Dotaal labial authoe with mmnbranoua patch aamad with mmuteahort 1») Mil >) branched and unbiancfaed aetulae or dcraal U ia l aui6oa arnooth-CA) OINottpplkebk I) T sm iul iennllum much ihcfter than 3rd entennil Mgment; ottier iptcil 144 Aa«aatar»afdaaaal| riniinh lanrfcea.(aanii4llllii) eennUe imKh thorter then SB!.(A) 0) Not applicable. 3) Itamaul iemaiuni u long •* M tntannil Mgment; odwt epicil Mndte 1) Doiaal labial aui&oe with membtanoua patch aoned widi minute or aboat Mta&nn and longer (hen SB UA) branched and unbranched aetiJan 2) Doraal labial ne&ceamooth. 2) Semaerlaefaeeenda Waagnnmt 0) SenaonaZnnallertbanienaona 1^10 15) Lateral labaa ad pranaaadnna. 1) Semotia I & 2 nibequaL(A) 0) Lateral lobea qmoae d). 1) Lateral lobea unarmed or abaent (A). 3) a«nalBaafaacandaat— aleepninr, 0) IVro wnsilla on 2nd antemal Kginent mudi longer lhan o(lien.(IO ISa) Laterallabaaafpiian ntnm (nanaddMra) 1) SenaaBa of 2nd antennal aegment an ahort & aubequal or fiwr aendia on 2nd 0) NotappBoable. antennal aegment long and aetaCxm^A) 1) Lateral lobea unarmed. 2) Lateral lobea abaent 3a) Sanaflh afaeeand an«ani l aagmat (neattddMr») 0} Not applicable. IS) Labial aata LAI length. 1) Sendla of2nd antennal aegment aD abort & aubequaL 0) LAI tninuleXP) 2) Fouraena3laon2ndanlennal aegment long and aelalbanXA) 1) LAIvestigiaL(A)

4) Mandlbolarmala. 12) Labial para LAc 0) Mandible wilfa defined mola.(P) 0) LAeabaent(F) 1) Mandible laddng defined m(ja.(A) 1) LAcprerenl(A)

5) Mandibular teeth. II) gPR arrnatm (aanaddltlra) 0) Mandible with aauB tooth d)ove tetiueiiIum.(P) 0)gFRaetaeaodceted 1) Mandible without aauHtoodi above letinacuhmi,(A) 1) gPRaetae norr-aoolceted

0 Mamdlbadar taelh. (nanaddWva) II) Nualataath(nanaddtdva) 0) Mandible without tooth below letinaculuni. 0)Naaale with2 atronÿy protruding, generally aaymmetrical. teeth. 1) Mandible with tooth bdow letmacuhim. 1) Naaale with weaUy protnrding teeth or teedi laddng.

7) S P aredltoaanaflla antacnahaal— iglary palpal aegnaint (nanaddMra) 21) PrantalaaataMUatatica. 0) Itaminal maxBlaiy palpal aegment whh > 10 poredika aenaBIa. 0) FRI preaent(P) 1) Troûial nuxBlaiy p a ^ aegmcnt with < 10 pon-Bks acaiailla. 1) FRI abaent(A)

I) ParaMXgafaaaxma. 21) Frantala aata FRI length. 0) Pore MXg not enlatged-CP) 0)FRIIongXP) 1) Pore MXg cnlaiged.(A) 1) FRSmcdiumXA)

3) MaxmaapaataMXSIaaglk 22) Stanaaaataatatna. 0) MXSmedium.(IO 0) Stearunata abaent 1) MXSahcctorniinuteXA) 1) Single atemmata preaent

*a) MaiBlaapaaraMXS length, (nanaddltira) 23) Parietal aata PA12 length. 0) NotappBoable. 0) PAI2ahort(P) 1)MX3ahoit 1) PAI2minute(A) 2) MX3 minuta. 24) Parietal ar4aPA13 length. (nanaddUIra) 10) Maatnarp aata MXC length 0) PA13 minute. 0) MX«ahoit(P) DPA Ilahort 1) MXS minute or medhim.(A) 2) PAI3 medium.

10a) Maamarp aata MX* length, (nanaddltlva) 25) Parietal ar«aPA15 length. 0) Not applicable. 0) PAI3medium.(P) 1) MXS minute. 1) PAI3ahorl(A) 2) MXS medium. 24) Parietal aetaPA23 length. II) MaxBlaap aata MX7 length. 0) PA23long.(F) 0) MX7 medium.(P) 1)PA23tnedium.(A) 1)MX7ahoit(A) 27) Parietal aetaPA24 length. 11) Maafflaap aata MXS latagtk 0) PA24ahort(P) 0) MX8medium.(P) 1) PA24minute.(A) 1)MX8ahofl(A) 307 308 21) PWMilMtoPA27lMftk. 49 Pna*nde#MaTE2llMith. 0)PA27nie

2Sa) ParMii Mta PA17 iMgtk. 429 Pr*(ktnck#tliTElll«cll>-(MB*ddWn) (QNotqiplicibls. 0) NotippliGiUe. 1)PAZ7ihoil 1 )m iv m lW # l 2)PA27la«. 2)1B21ibait 2f) P«WMM(mPA2#hmgaL 43) rndMradcMt#TE23h#) 1) PA28imnuleariiwdium.(A) l)TE23mednunXA) 2ta) UaOw) 44) Pn«h*tacke*taST40lwflh.(MwddMT*) QNotqjpBcaUc. 0) ST28 minute. 1)PA28œiniite. 1)8128 ihoct 2) PA29 medium. 43) Pnbact(P) 1) PMiboctCA) 1)TBlminute.(A) 40) PiM tM V ilM tiPR7l«clh. 31) Pi«dMncicMfieeoxiteiet#e. 0)’re6ihoft(P) 1)TE6minule.(A) 51) PrWermlWdlm uirlenr (neMOdHhr.) 0) Proetemite longer dun breed. 34) PrcOMndeaetaTETtaclh. 1) Proetemite is long •* broed. 0) TE7 nmuite.(P) 1) TB7 veitigial or ibort.(A) SI) Mweriegum Olmenrlini (neeudOWtre) 0) Meioteigum mudi wider dun long. 34.) Pr#dMr#ck»et»TE7l«*lfc.(#Mttd4ttlT.) 1) Mesoteigum sbout IS long u wide. 0) NotippScalile. DTETvcftigU. S3 MesedutackietaTE3lM«lh.(Mu4dtliTt) 2)TO7ihoit 0)TE3abecnL 1) 183 minute. 37) Pimhende#i<>TE7lM|lh. 2)TE3elutt 0)TB91oi*(10 1)TB9medium.(A) S4) Meeedwewck set. TE4 length. 0) TE6 ihait or medium.(F) 30) PrMkeradCMdTEniMClk. 1)184minute.(A) 0) TE12 ibcrt-medium.(P) 1)TEI2minute.(A) 53) Meselhsmde,eemTE7 length. 0)1B7ibsent(P) 39) PiedHradCMlaTE14lMtlh. 1) TE7 minute or medium(A) IQIPtdihort# I) TE16 •beentarminute.CA) SSs) Me**ther*dc#e«aTE7 length. (nsnsddltlTs) 0)No(epplicsble. 39*) Pre«fa*tKic(t(aTEI4ba|lh.(Hnddttivc) 1)187 minute. 0)Noti|>plical>le. 2) 187 medium. 1)TEI4*bient 2) TB14 minute. 54) Meeelheradc set* TEO length. (nenaddWre) 0)188 minute. 40) PiedMndcMtaTEUtaclk. 1)188 short 0)TBI8#Uent(P) DTOlSvettwiUA) S7) M eselhsndc set# TEIO length. (nenaddWre) 9TB18minute.(A) 0)18l0minute.(P) 1) 1810 vestigial or thoti(A) 41) PntterKk«rt#TX20lMitL(i 0)TB20 minute. S7a) MesethendcsetaTK101ength.

M) M(Mboft 0) TES minute. 1)1B17minute. DIES aboat 2) TBI7medium. %TES medium.

(1) Meeedwende erne TEH Wgl&.(memmddlUee) I t) MalathanckaelaTElllaa«h. 0)TEI8mimiie. 0)TE10minuteXP) D TBlSihaft 1) TEIO veatigial at aboft(A) % TEH medium. 7N) Matalhetadeee«aTEieiengdB.(eaatad«IHe) <2) Meeeaee»deeelmPL27lem0h. 0) Not applicable. QPL27medium. DTElOveadgiaL 1)PL27ihmt 2)1E10aboat

tXi MeeetWmdepMeexEeeWue. M) MalathatackaadalEKIaagtfa. 0) Meiothancio pieemqle entûe.(P) ll)TE16abaitatmedium.(P) 1) Meiolhoti gc pcecoadle iubdivided.(A) l)TEl$minuta.(A) 2) Maotlianciop(eooxitee|)peKnllyibKnt(A) 11) MaMhatadc tala TE17 length. «4) MeielharackeetaSTMlMClk 0)TB17minute. 0) ST30minute.(P) 1)TE17ahait 1)5130 (bixt(A) 2)TE17medium. ^5I30medium.(A) 12) MalathatackaelaTEItlangth.(nanaddlthra) IS) MeeetWeckmtmSTSl hcgÜL(meueddllhr#) 0)TElSminute. 0) 5131 medium. DTElSahcat 1) 5131 du et 2)TE18medium. 2) 5131 minute. 13) MaMhetack tala PL24 length. f t) M#ee«lw«nde e««eSTJ2 length. («e—ddUlT») 0)PL24mmute.(P) 0)5132 minute. 1)PL24abait(A). 1)5I32iboit 84) MalathatadcaalaPL25atalaa. <7) Meee-HHeHtalkenckee(nSr44ieiVlk(MuddltlTc) 0)PL23pieaent(P) 0)5T44ibort. 1)PL23abaent(A). 1) 5T44 medium. 85) Malathaaatic aala PL17 length. O ) M«e»mmde»ehlheee d c eet»ST4SeWu«. 0) PL27 medium 0)5r45ibMnL(P) 1)PL27 aboat 1) 5T4SpRMnt(A) 84) Malathataalc aala PL28 length. (nanaddWte) W) Mete-end ■ntithereckeeteSTUliitt. 0)PL28 minute. 0) 5T4«long.(P) 1)PL28ibait 1) 5T46medium.(A) 2) PL28 medium

71) MditkendcTEl,TSl,andTEkplecenient 87) MalalhaaackST34andST37placaaaaaat 0) TB1 iiolated, T ) ^ end pa n TEb ihiie sneO •clente.(P) 0) 5134 on minute adeaite; 5137 abaentOO 1) TE 1, TE2, end pore TEb lim e imill ideriteXA) 1) 5134 and 5137 each on veatigial at minute ateanitea or abating amaD ^ TE 1, TE2, end pen TEb eech on vectigiil 0( minute >deritef.(A) ateanite^A)

71) M eW W edcTESW nW n (nw i i dWhr») 87a) MaMhatadc 5T34aaUST37placaanaaat 0) TE3 teigite veitigiel at minute. 0)Nol applicable. 1)TB3teiSite abaent 1) 5134 n d 5T37 eacb on veatigial at minute ateanite. 9 5134 4b 5137 abating aanaO ateanite. 72) M etatbetickTEiaandTEUflifeniir t (neawddWre) 0) TE12 on minute idcdte; TB13 on emtll togite. IS) MaMhatanlepieeexllealalna. 1) TE12 end TE13 ihaiing aman togite. 0) Metatbondc paecoxite enliae.(P) 2) TE12 andTE13 each on watigial at minute teigitet. 1) Metilbnaidc pieooxite aubdivided.(A) I) Metatfaondo pieccnte apptaeait]}'abaent(A) 73) B(ntnaldlllra) 0) Metatexgum wida than long. 83) MaMhatadcST2*| 1) Metatogum about u loaig u wide at longet than wide. 0) 5129 on metaatetnite.(P) 1) 5129 offmetaateanite.(A) 74) Matatharack aata TE3 length. (aeaaddtllTi) 0)TE3 abaent N ) MaMhatadc aala ST38 length, (n addHtva) 1)TE3 minute. 0)5130 minute 2)TE3ahoat 1)5130 aboat

75) Metatketacfc aata TE4 length, (nanaddlllta) 91) MaMhaaadeaala5T3l length. 0)TE4ahait 0)5niaboit{P) 1)TE4 medium 1) 5131 minute at medium.(A) 310 »U) Mt(alkMadCMtoSniiia(lSk(MnUilln) 189) Metelteirmdctegm e te n i4 tea # (m ia iddllUi) 0)No*«pplk*hk. l»1116minute. 1) ST31 minute. 1)1116 vemdgiaL ^ S n i medium. 3 ) H 1 6 ih o it ÿnidmedium. M) MetelhtncteMtiSnilMglh.(MmiMlttee) Q SI32 minute. 118) Mmtetemrmde teg meteTAlAl t e a # (mmamddtelM) D S m ih o it l)XAlft2 minute. DlAlftlmhoit >3) M etetbendeamcatfcte ÿ lA ljU medium. 0) Metethondo 00X1 culide endidy imoo(h.(IO 1) Meteduncio coxi Gudcte botti imoodi ind gnnuhi.(A) lU ) AtedmmlamI megmmad 1 T O 41TE17 mmtel gteraa a te 0) TB3 mndlBlI on eeiante iderite.(I) M) MteteWnetetegMteCOUtemgllLCmeaiddNtee) 1) 1E5 end TE17 ih in teige ovoid icledte.(A) 0)COI3minute. 3) TB3 on luge ovoid iderite; TB17 ibmentCA) 1)C013ihoit 111) Atedmmhu l mmgmmal ITE14,15* * 14 ph n a ieat 35) MteteWmte teg Item C034 te a # . 0)TEK13*&16mhmmmiddzedteqite.(P) 0)COMihoit(P) 1)1E K 15, ernch on minute teqite,TB16an 1) C014minute.(A) until or midsized teqpteXA)

94) MteteWmte teg Item COM t e a # 113) Akdtadamlme^Madl mete TE3 t e a # 0) COI3modium.(P) 0)TB3 medium. 1)COI3ihait(A) 2) TB3 ihort or long.

97) Mtetehwadc teg eten r o l l t e a # (BMiddWrt) 113m) Ahdtmhulergmatel m tteTE3tea#(ataiddlHvi) 0)CO18minute. 0)Notmppliemhle. 1)C018iha(t 1)1E3 ihort 2)lB31ong. 98) Mteteteemtec teg Item TR3 t e a # (OTIOiboftOO 114) AhdmaJamI i i gmiaU I mete TE4 t e a # (meaiddldve) l)TR3minute.(A) 0)154 ihort 1)154 minute 99) MtetetewacktegiteiTIUteteax. 2) 154 medium 0)TR4ixeunt(P) 1)Hl4ibMnl(A) 115) AhdeailamliiparatiwteTEdtea# 0)156fnedhun.(10 188) M tetetewKktegNtmTRSteo# 1)156 ihort(A) 0) 1113 ihcrt Of medium. 1)TR5 minute. 114) Ahdmailatlitgmiat l mite TE7 te a # 2) TR3 medium. 0)TE7ihort(P) 1) 157 medium or ibmentCA) 181) MtetehetadctegeteiFEltectetea. 0)FBIonianur.(F) 114)0 Abdmrmla i l i mfmiat l mete TE7 t e a # (amamddlUvm) 1) FBI on membmne.(A) 0)Notmppliembte. 1)157 mbmentCA) 181) Mtemlteeende teg Item FEZ t e a # (aeamddUtee) 2)TE7medium.(A) 0)FB2 minute. DFEZiboiL 117) Abdmmhml mmgnwat I mmteT514 t e a # (amai ddWvm) 0)T516medium.(P) 183) M tetebttadctegiteaT13tea#(Maiddtthrt) 1)TE161ong.(A) 0) TO minute. 1)TOihoft 118) Abdmmhimlmmgawatl mmte TE18 t e a # 0)1518 ihortdO 184) Metelteeeiete teg mete TI4 t e a # (aeamddtetee) 1)TE18minute.(A) 0)TI4 minute. OTIdibofL 119) AbdmmlailimgnMatl mmte TE18 t e a # 0)1520 minute or ihortCF) 185) MeWteeemdc teg mete Til t e a # 1)1520mediumXA) 0)TI9minute.(F) 1) TI9 ihoft cr veitigiiL(A) 111) Abdmmlml mmgmmatl mmteTEll t e a # 0)1521 minute.(tO 185m) MiMboracfctegMtaTUtea#(aiaaddttin) 1)1521 ihort(A) 0)Notepi>lica>)k. 1)119 ihort 111) latmrter# mlmtam D T O vadgiil 0) Intcrtergitem prcient(IO 1) Interteigitem mbment(A) 184) Metelteetadc teg item nil te a # (aeauddUn) 0)TI11 minute. 111) Abdmralai l tmgmart 1 aamraii dgtearlte mtmtam 1)1111 ihoit 0) Donil, Imtenl, end ventnl pleuiitem (ireicnL(F) 1) Vmtnl pleuiite mteung or both litenl md veritnl pleuiitem miiiiiv.(A) 181) Mtemlteermcte teg mete ni3 t e a # 0)T113minute.(P) Him) Abdmailaml mmgrnmal I aamnamdgteamlte mlalam(mmaiddhlrm) 1)T113ihoit(A) 0)Notipplicmble. 2)T113medium.(A) 1) Vaitnl pleuite mteang. 2) U tenl uid vaihml pkunlem miuing. 188) Mtemlteermrte teg meteni4 t e a # (aearnddtetem) 8)H14minute. 113) Abdmralail r at I mite PLH mtmtam. 1)1114 ihort 0)FL22ibient(P) 1)PL22proient(A) 311 124) A»mitwilM|wiwH>t<»PL25iMtflL 141) Abdemlaal eagaant i aata 3T44 langtk (nanaddMra) 0)PL25loi«.(I0 0)5T46 medium. 1)PL25niedium.(A). 1)ST461ong.

115) AliwtaslwgmitflwtePLl?!**». 142) Aaaaalaae abdamhial pralega. (aeaaddHhra) 0)PL27long.(P) 0) Ptolega anned with denbddee. 1) PL37 •boct ot mednnnXA) 1) Pnlesa not aoned with dentridea

124) A btarinlM (M N (IW arttm atiiM u.(M w ldtlh«) 143) Akdamlaal aagmaad n-Vin batmaiaeidia atataa. (aeaaddUre) 0) Piii of Uo il intoitantM . 0) Pair of lateial intentemitea. 1) Singie médiat Bitentemita. 1)Sinÿ e medial and pail of latent interalemitee. SO tntoitanita itmnt ^ Anterior and poatoiac medial and pair of tateial intenteniilea. ^InterteqpteaabacnL 127) AliimlialMffmrttanfS T M phw iMif 0) SI29 on nmuite iteniüe & CT30 on miileized ateo iite^ 144) Abdambwl eepaaadlXlalarOmgMe rtalaa, (aaaaddHKra) 1) ST29 * S130 etdi cn vndgiit ttenâea or ibiing a miiMzad togite^A) 0) S in ^ medial tntertagiie. 1) Pair of round latent « d angle medial intertergitea. 1274 AM imkat w pe ■ » 18T27 A 9T3# f law mt , (mimeddWh,) ^P air ofquadntelalenl and angle medial Interlenplea. 0)Natq)pScabie. ÿlnterlngiteaabaent. 1) s n p ft SI30 eedi on vciligiil stemite*. Q ST29 ft ST30 aliaramiil^ized togSe. 1«S) K ealae TE3, n O i TE17, TEH, ft T E » I (aaaaddHtn) 124) A iiim hn! aigmml l 8T11,8T12 ft STM piaw n n*. IQ TE 19 on aepaiate aderite; 153 ft TEl 8 abare aderite; TES ft TB17 abare (0 sni on aqiaiite aianüe; 5132 and ST33 ilian itaniite^ aderile. I) 5133 cnaepaiata atonite; 5131 and 5132 ibarastemite cr 5T31,5132 ft 1) TB3, TES, ft TB17 abare adaide; TE18 ft TE19 eadi on aepante aderitea. 5133 cadi on aqitnte iteniite.(A) QTE3,TE17, ftTE19 eadi on aeimate aderitea;TES ft 18 abare aderite. 3) TE3, TES^ TE17, TE18; TEI9 each on aqMiateaderitea. lST 31 ,ST12 ftST3 3vl12Sa) Aki lST31,ST12ftST33vl12Sa) (manaddbhe) 4TE3 on aepanie adeiite;TES and TE17 abare aderite;TE18 and 19 ibare (ONolapplkahle. aderile. 1) 5133 on aepaiate atemitr, 5131 and 5132 iharaiteniite 2) 5I31,5132 ft ST33 cacti on aepaiata atemite. 144) Abdanriaalaapnawl IXaataTEPftperaTEgplareaniart.(aanaddblra) 0) TE9 ft TEg (when preaent) eadi on aepante aderitea. 123) AbdamlnlaafaaantlST3ftSTMftST4«placaaMnt 1) TE9 ft TEg abare aderile. 0) 5138,5139 ft 5T40 abare atanite.(P) t) 5138 on aqnrate atendte; 5139 ft 5T40 atian atemite.(A) 147) Abdemlaal aagmant IX aataeTEH ft TEllplacamant. 0)TEI6 ftTE21 each on aqtaiate aderitea. 138) Abdcaalmalaegaaan#l5T43,ST44ftST43,lacamaaft 1)TE16 ft TE21 abare aderile. 0) 5T43 on aqniate atemite; 5T44 ft 5T43 abara atanite.(F) 1) 5T43,5T44. ft 5T4S abare akmile.(A) 148) Abdemlaal aegmeat IX TEOedmbealaeL 0) TE9 teigite minute or amaH and circular. 131) AbdaaataaiaatnaaatlSTMIaacdi 1) TE9 teigite middzed and dongite. 0)5r29minutc.(I0 t)5T29veatigiaL(A) 149) AbdemlaalaagmatlXeataTElatataa. 0)TE1 preaent(P) 132) AbiaiadMtaagaaadlaa«aSTHIanida 1)TE1 ahaentXA) (05I30ahoat(P) t) 5130 medium.W 150) Abdemlaal aepnawt K eata TE17 length. (neandJHtre) 0)TE17abort. 133) Ak

135) Abdemlaal eegaaaW I aebi 5T38 langlb. (aanaddWYa) 152) Abdemlaal aegmaallX eata TE2HaagUL 0)5T38 medium. 0)TE21 abort or medium.(P) 1)5138 abort. 1)TE21 minuteXA)

134) A b ia m lil aapnnrt I ea«a5T4# langlk (nanaddMw) 153) Abdemlaal regaaaaat IX peree TEf and TEg atataa. 0)5T401ong. 0)TEfandTEgpreeent.(0 1) ST40 medium. 1)TEfandTEgabaent(A)

137) Abdamlnel eegma#H ea*aST41lamglb. 154) Paragampbl atataa. 0)ST41abort.(P) 0) IXgitilbem pangoanpbi abaent in aeoond inatar.(F) 1)ST41medium.(A) 1) Paiagoanphi preaent in aecond inatarXA)

138) Abdemtnaleap aaartIaaaaST42rtanm.(# addbhre) 155) Abdemlaal aegmaatlX eata PL24 length. 0)5T42preaenL 0) PL24 abort4nednun.(P) 1)5r42abaent 1)PL2dlong.(A).

139) Akdeaalaalae(meatIaa*aST431an(tlL(naaaddmva) 154) Abdamhial aegmeal IX PL34adertte atataa. 0) 5T43 medium. 0) PL36 aderile not aubdhrided into 2 paita.(P) 1)ST431ong. 1) PL36 aderite subdivided into 2 partt.(A)

140) Abiaaalnnl aapamrt I eata 8T45 lam #. 157) AbdemlaalaegaaaatIXcata8T»,8T30,ftST44phremait 0)5145 aboitOO 0) 5T29 ft 5130 abare aderile; ST4d on ae|»nte adeaite.(IO 1)5r4Sminule.(A) 1) ST29,5130, ft 8T44 each on aqiante aderitea or STT29,5130, 2)5r45veadêal(A) and ST46 sharing adeaite or 5T29, ft 5T46 abare adeaite; 5130 abaenl.(A) 312 157») A k ta ln lM p M rtK M tiS T 2 f,S n i^ 4 S T 4 < p l (■aamddUhre) 173) fygaged m f PM eteriM. (eamaddlUw) 0)No(^Tlk»bk. 0)FFIpR»enL 1) SI79, SmO, & ST16 etdi on sqxnts iderita. 1)FF2abaent 1) SrT29, SI30k & ST4< i h i n id o S e . ^ ST29, ft ST46 ilure idente; sn o iU cnt 174) Pnar»d»a(»PF2tava.(MMddWTe) 0)FF2long. ISQ AkteUi (lX»d»STMiMa«. 1)FF2 medium. 0)SI30picMnt(P) 1)ST30ibMnt(A) I7Q Pygepad aal» PP3 length. (eamadftUva) 0)FF3 ahort. 15f) A >iw l»»lmgn n t K u t» STJ> «Mm». (m iii

1(1) Ak(»adMl»»pMatIXi»(»ST4(l«|lk 177) P7ga,»dee«»PP3l»»#ft 0)ST46meifiuniXF) 0)FF5ahort.(P) 1) ST4$ kog K iboit(A) 1) FF5 minute 04 medium.(A)

1(1») AH i i hwl»«lw w lK»«t»ST((l«il>.(»»M((WlT») 177») P7ger»d»a«aPPStaaf|k(HwddM4e) 0)Notapi&»bk. 0) Not applicable. 1)Sr461ang. 1) FF5 minute. 2)ST4(ihait OFFS medium.

1(2) »»>»» UCl-7 ph w iir t (««M iftttTi) 170 Pyg»pod»et»PP7l»ilk(»amii(lthB) 0) UOl ft U02 than adoits; U03 ft U04 than identa; 0)FF7ehott U05^Uafi,ftU07ahareadaile. 1) FF7 medium. 1) lX]|>7ahan aderile. 1 70 Aapaeltea, (mamaddlllaa) 1(3) UmpmvMaabUClkmglh. 0) Membcanoua eutide laigdy coveied with tuberdea; pteaent on moat teqp 0) UG2 ahort armedhini.(IO andatema. 1)U02ininute.(A) 1) Membfinou» cutide higdy wiSiaut aaperitea; dentide» preaent on moat toga and atcoia; pttdiea of chal» 2»e htenSy. 1(4) Ur»CM«hlM<»UC4laaflh.(MM(dUT») 2) Membfanou» cutide laigdy covered with diahzae; dentidea abeent 0)UQ2 ahort. 1)U02 minute. 1*0) M»lplgM»n tnbul»» and hind gut 2) U02 medium. 0) Malpighian tubulea and hind gut de»t(0 1) Malpighian tubulea and hind gut sued nÛte.(A) 1(5) UregaigM art» UC3 haglh. 0)UO5 ahort oamedium.(P) 1)Ua5minule.(A)

ItS) VngemUMtaUCdiaglk 0)UO6longXP) 1)UO(modium.(A)

1(7) Ur»g»»apUaa<»IIC7laaglh. 0)UO7ahort 04 modium.(P) t)UG7mimite.(A)

1(1) Ur»gaaiBphlaalaUG*l«glh. 0)tX39long

1(f) UretaBgyaatiUCllpwWam 0) u a i l in diatal t/3 of lenninal urogocnpilaegmenL(F) 1) UOl I C». nûdwiy akng lenninal urogomphalaegmenloeUOIIayical l(f») llregarepMaetnUCll 0) Not applicable. 1)UOIIc*. midway along tenninal Ulogompbal aegment 2)U 0II apical

170) Uregompbla»t»IICl%»mdllC13paaltlan. 0) 1X312 apical; U0I3 aul>apicaL(P) 1) U0I2 ft U0I3 both apical(A)

171) Pygepad act»» PP4 and PP3 pla ramirt. 0) FF4 and FFS each on aepante aderitct(F) 1) FF4 and FFS aharing aderite.(A)

172) Pygepad aeta# PP( and PP7 pliieaaamt (nanaddtttve) 0) FF( and FF7 eadi on aepante aderitea 1) FF6 and FF7 ahaiing aderile. APENDK B - ADXJLT CHARACTERS

1) r n a fc ir ia a lilitu . 14a) Eplpbaiyngaalbaaarbraabdavalapeaaut, (waneddlttva) 0) R o m with htooSantil canni only. 0) Not applicable. 1) n o m wilfa hteo&ontal A votieofitmtal c u m 1) Inner bruab narrow. Q Inner bruab veatigiaL 1) Latmfrtatalorinlliaflh. ^ Inner bruib incomplete; bordered diftally by comb. 0) ProM «idi ihoct blaoflnntal cadna. 1) F n n i with laiaofianlal cnint voy long&joins!8 t t dypoH. IS) Eptpbarygeal drraaaaa Aal bruab atatue. 0)Doraomedial bruab preaenb(IO 3) AmfmmêliÊmftK 1) Donomedial bruab ebaentfA) 0) Antoinie M long or longer dun bead op n ik width.(F) 1) Antoinaa ihorter tban bead capaule wkÜ>.(A) 34) Epipharyngeal madtal ara» mmM------. ------0) Seruilb relatively finv in number; Kneady arranged and reatricted to lateral 4) ABtawnl acapa maepbalagy. (mamaddbhra) margina of medial area. (P) 0) Scape idalivdy dongata and gnduaOy expanded. 1) Senailbnwmeroua Anot bncarly arranged; eoemadm* on or moatly 3) ScaiM abort A awoScn. covering medial area ditc.(A)

9) AotaMalacapataadk 17) Maaiillbnlar caartact chaanarareptara aWna. 0) Scape apicaBy unannedOO 0) Contact chemorccepton preaent at apex oflaleral arca.(P) 1) Scape aritfa vcntrcapical tootfa.(A) 1) Contact chemoreceptora abaent fiom lateral area.(A)

d) Aeiaeealacapeeetadei II) Maadlhirlar latarai area pit atatrra. 0) Setae moady miniite, alendet A incanqiieuoue.(IO 0) Lateral area lacking deep pita or depreaaione. 1) Setae moatiy abcft atout, ccnapKuoua.CA) 1) Lateral area with pita or dqxeariona.

7) A1a—al dub ■ aapbalagy. lie ) Marrdlbular lateral area pit atatrra. (rranaddMva) 0) d u b divided into 3 aeotiotta by 2 genoiDy deq> ftdci that 0) Not applicable. cam piet^ encifde dub.(I9 1) Lateral area with aeveral amafi pita. 1) Cbib eoinpletdy fitted; aigMificial tiacea of Ibimer entonnai 2) Lateral area with dqxeaiion or aingie large pit diviaiona oocaaionally paabtant(A) 19) Marrdtkulartaath. 0) Mandibular apex with two teetb(F) 0) Senailla ohactica veqt dongate A ptominentOO 1) Mandibular apex with tingle toodL(A) 1) Senailla diartica abort(A) 21) CaardtUaraaaaadlhalar latarai ararr apex. Saaitflh baifcaalra caudlllau 0) Lateral area unifixmly broad. 0)Anlennal chib wifit two complete oa needy ooinpletettanaveoewhoale 1) Lateral area narrowed aptcally. o f aeniilla baatconica; aeniilla may be concealed within tuki(F) 1) Antennal dub with a in ^ compléta banavene wfaod of aenaSIa batkonica; 21) Caadttlea aaaadtbafcw latarai area baaa. (aaaraidMva) or aensillt baaicanica diatnbution non.lineai.(A) 0) Lateral area broad basaOy. 1) Lateral area baially conrtricted. 9a) SaaitHla baakaatka candWen, (■aaaddWtva) 0) Non applicable. 21) Mandibular aaimrl dirraarphlaaa atatrra. 1) Senailla baatconica finning aingie compléta tianavaie wfaod(A) 0) Mandibka not aexually dimorphio.(IO 2) Scnailla badoonica ananged into btge oonapicuoue plaque.CA) 1) Mandibles in maka aiymmebically devdoped; 3) SataaOla baateanlm Aatrtballam patchy^) dorsolateral area of right mandibis bearing buge lobe.(A)

I t ) Cÿpeal dlaairphliM atataa; 23) Mandibular articulated appendage vaatltara; 0) Maka belong aetigeroua tubcrde or pubeaccnt on dypeua^lO 0) Articulated appendage with rdativeiy long, denae pubeacence. (P) 1) Make with aetigaaua tubetde or pubcacent on dypeua.(A) 1) Articulated appendage with very abort pubeaeence (A).

U) Eplphaayagaalamrglnal ratal Mage caadMam. 24) Mandibular malar davelopnwnt 0) Fdnge complete A finned of mom than d aetae.(F) 0) Mola extending beyond meaal margin of meaal area. 00 1) Piinge leatiictcd A finned of leaa than 6 actae.(A) 1) Mola even with metal matgbi ofmcaal area. (A)

12) Epiphaiyagaal latarai area attartunaart 25) Mandibular aukratrlehlal Held atataa. 0) Latcaal maigina of lateni area attached to inner labial aur&ce.(P) 0) Microtrichia areO developed. 00 1) Lateral maigina of lateni area detached fixxn inner labial aur£aoe.(A) 1) Microtrichia reduced. (A)

II) gplphaiygaal latarai area aeaia aaarpbalagy. 2d) Maadlbular praathaca attarhmaart 0) Diao with broad overlapping acalea.(^ 0) Proatheca not detached apicaDy.0O 1) Diac with reduced or narrow acaka.(A) 1) Proatheca detached apicaIly.(A)

13a) Ep^baayngial latarai area acale marp balagy. (mawaddlUva) 27) Maadlbular deraalpraethaealaatalaa length. 0)Notappbeable. 0) Apical proatbecal actulae relatively abort A not extending beyond mandibular 1) Diac with amall, elongate, and narrow overlapping acalea. teeth.00 2) Diac with email, abort, arid broad ovedapping acalea. 1) Apical proathccal actulae long; extending beyond mandiht Jar teefii(A) 3) Diao with amall, dcniieulate mainly non-overlapping acalea. 11) Maadlbular deraalpraathacalaatalaaeandltlan 14) Epipharyngaal hrraar hrrrah davalapn 0) Dorsal actulae apicaBy fixked.(F) 0) Inner bruab broad rveOsleveloped A coniplcte. 1) Dorsal actulae simple.(A) 1) bmer bruab narrow, incomplete or veatigiaL

313 314 tt) MiattatarlHMrpndiMadpetabMCMittka. 45) Pakplgarderaal ridge atstea. (D Peetinite idle ilwaitOO 0) Donal ridge oomplels.(P) 1) Pedinite Mbe present^ 1) Dona) ridge apkaSy abteeviataiL(A) 2) Oblique ridge ahtenL(A) M) 0)Riduhibsent(P) 4f) Pslÿlgtreaedirle atstea. 1) lUduk pnaent on imur tuifioe of im

31e) Celeel metplielep . (ne—ddMre) 47s) Derasla •) (QNofepplieehle^ to Not applicable. I) Oelee modified (te liquid fitteting beding^A) 1) Donal membrane armed with minute actulae. 9 Oolee modified fix qxxe gellieniigXA) 9 Dorsal membrane unarmed.

311) Celeel merfMogy. (nenmddMve) 41) Prartreal martllaty psipsi aepeset lariglh. (ONoteppliceUe. 0) Pnndmal and aeoond palpal segments wider fiian lo n g ^ 1) Setiilio medhim length i . epehilete 1) Proximal and second palpal segment longer flianwide.(A) ^Sehilioihnrt end tiperedptoidmilly.veo^ thin end ^li)litediitilly.(A) 4f) Third eraxmsry palpal asp rarrt araatesae, 31) CeleeTialraleartHtanBetanL ()) Whorl of at lent S apical prhntries preaent(P) 0) >feiitnl gileel diso leigel/wdfaout eetulee.([) 1) Palpal aegment hterally armed latenly on|y.(A) 1) Ifentnl luifice ofgilet «itfa numetout flocfide Mb!lae.(A) 50) Trrsahral marin try palpal aagmawt saarphalagy. 3Q CileeTentrelwrfecateitmei 0) Ttaminal palpal segment not flattened donoventnlly.0’) 0) \fcnlnl euiboe ofgilee gnooflL(P) 1) Ttaminal paÿal segment flattened dorsoventnlly.(A) 1) Vnbil eut&oe of gilee icaly.(A) 51) Taf h ral sasifllary palpal aagsaasdaassstsrw 34) Csleedenelteenl Steel dedirdilei. 0) Tbrminal palpal segment armed with a rrsxtue of vestigial and usually 0) Geieel hddng iioieted dontl lelit duteer. short setae; most short aetae randomly or ventrsBy distributed(IO QOtleilinneddonmeielly or doooletoiDywilhitoliledcluito of medium- 1) Ikrminal palpal aegment armed wifii mixture of vestigial or minute and short length eelis. setae; short; mainly laterxllydistribuled aetse or ^ipesringunarmod.(A)

33) MeiBiiy hteeeel mirf helegy. Sla) Trrsrteal martllary palpal aagsrant srsaature. (sanaddlHva) I» Lecinee hotdly tttsdted to medioteqiee.00 0)NorHpplicable. 1) Udnes higdy deteched fiom medioteipee,(A) 1) Tbrminal palpal appearing unarmed; with vestigial aetae only. 2) Tbrminal p a ^ aegment with a mixture of vestigial or minute and short 34) Ierlmeel etrehllne testes. setae; 0) Strobiha abeentOO short setae mainly laterally distributed. 1) Strotntui (xeientCA) SI) Tsrsrisal sasilllaey palpal aegsasnt spiral aanteUs dIgllKarsils atatrra. 3‘T) Isfle eslMegesetelML 0) Apical sensilladigllifntmiapresent(P) (9 Fdnge irilb bladeiltem eetuIse.(F) 1) Apical senailla digitifixmia abaent(A) 1) Hledeiltem ediilto Uddng.(A) 53) Mrslssasiarphrlrg^ 31) LedeesIM egeisedltlee, 0) Anterior margin ofmentum straight or outwardly arcuate.(P) 0) Fdnge oomplete or nesdy i o ^ 1) Anterior maqpn uni-or tridentate or emaiginate.(A) 1) Fdnge mbteevisted |saxhnml|y.(A) S3s) Mentssa saarphalagy. (saraaddltlva) 33) LsdessderislinrtMS testae. 0)Non.applicable. 0) Oonel im&oe texture imbDcsteXIO 1) Anterior margin uni-or tridentate. 1) Dooel eurboe texture iugoae.(A) 2) Anterior margn or emarginate; cuticuler shrtf may urukrheemarginatioo.

41) MeHeetlgee cetlcsler csrlUee rsmdhlem 54) Msratnm latarai saarghtarsastsra. (saraaddltlva) 0) Medioteipet with tevetel cuticuler csvitiee.(i’) <9 Lateral margin ofmenbrmfirnged with minute aetae. 1) Medioteipet with lingular cevity Of cutkulir cevities lscldng.(A) 1) lateral margin ofmentum unarmed.

40s) Medleetlpee estkslsr estedee ceedltlen. (seuaddltlre) 55) Paragleasallahevsertralarerfaeatailses. 0)Noteppliceble. 0) Tfentral sur&oe of paragloeaal lobes largdy untextured(l9 1) Mediostÿea with tingle outiculet oavity. 1) Vmtral surfioe of paragloaaal lobes wifii hnbcicate lexture.(A) 2) Medioitipet lacking cuficular oavitiea. S4) Paraglaaaslappasdlx atataa. 41) BaalatlpeaprlsexryatesI testes. 0) Para^oasal appendix present(P) 0) Baiiftipea generally aimed with 2-3 1) Paragloasal appendix abscnt(A) prinaiy letae, aome arising teaally.(P) 1) Baaisripes armed with 1 or 2 ilender ST) ParaglaaaalWsgaaataBlaafandltlaeei ahort to medium length prhnaiy setae 0) Fringe setulae relatively long and unmodified.(F) reatricted upper l/2ofadaite.(A) 1) Fringe setulae of short-medium length artd vsrioualy modified.(A)

42) Palplgarsaetisl prteaaay sets atstea. S7a) MadMledparagleeaalfirhrgaaetalaa types, (sen^addltlvae) 0 Primary aeta medhrmdong and alender.(10 0) Not applicable. 1) Primary sets medium lenÿh and t«iyrolxiat.(A) 1) Fringe setulae medium length and apicaDy branched. ^Fringe aetulae relatively ahort. atoul and occasionally brandied or apically 43) Pslplgardersalprhearyatese testes. fixked. 0) At least two piimary setae pcesentCp) 3) Ringe setulae medium length and biimt 1) Single primary aeta onlyW ^Fringe setulae medhan length, ^acaBy expended and barbed. ^Fringe setulae rdatrvdylorrg and phimoee. 44) Psiplgir peg ates states. 0) Peg setae aheenl(P) SX) Classai lehe devalsf mrrte, (aasaddltira) 1) Peg setae present at donal apex.(A) 0) CHoasal lobes poorly developed or appûcntiy absenL 1) Glossal lobes weB developed and disteict 315 Sf) POTthtwUtcwhlr—mWM 75) 0) Diio idM numbcdng 2-< pet ada, medium-long m d uncx)dified.(F) B)Aoatemum lacking aheXP) 1)Diio iemovmnoudymodi5ed.(A) I) Pioetemum with alaeXA)

Sfà) Pongleeeeldbeeemi tempeiMem (uemeddlllte) 74) Feewtlblal eratielar aaergbr eraaeture. (niuai tin) 0)Notepplicible. 0)AriteriorrDraginoffbte4ibiaatrai|dit Onôngeielaeiuiniberiiigc*. lSpeiiide;iDbuttniediuni length, and tapered. 1) Anterior margin of fixe-tibia toothecL 2) DiiCiataa a mixtun of lavent ilender tapered letulaa and a pair of axtremdy atout bhint aetae. 77) Etytnlaaaapbelegy, B) Elytra with at least 2 complete dorsal striae end lading coaiaoXP) ( t ) PaJplgarnatflielap . 1) Bbrtra with leae than 2 ooeqriets dorad striae or dytra costate.(A) 0) Pa lp i^ wim elongate baaat extenaica(F) 1) Faln p t wilfa a relatively ahoit baial extenaioa(A) 77a) Elytral merpbalegy. (uaraeddNIra) 0) Not applicable. «) SecaadhhUpa^ala 1) Elytra wilh leaa Bran 2 dorad striae. 0) Second palpal aegment ahort and hand kke; wider than long.(P) 2) Elytra with coatee. 1) Second p a ^ aegment longer dianwideXA) 71) Elytnlpn O) Seeaad laMal pa^ al aegaaeat vraetralelaral prlaaaiy aetae la i# k B) Elytra lading deep punctureeXP) I» Ifailioieteial primaiy abort; aubequal to 0^ pdmanea end ahorter dun 1) E ^ with dc 9 puicturaaXA) tenninal palpal aep n entOT 1) Vntiolateial prhnaiy toeiK longer than other pnmanea and longer than 75) B M w lM R A m ebm ae, terminal palpal aeginentCA) 0) RAl+2notreduoedXP0 1)RAI+2reducedXA) O) Tiraimal inhlal palpal aegn int n arphalep . 0) Terminal palpal aegment diataHy acuminate; not datteneddoraoventraHyXF) St) BbadwlngltdailPeendblam. 1) Ttenmal palpal aegment expanded diatally and doraoventrally llattened.(A) B)R4ARPaeparate.(P0 I) R4 & HP meaally united.(A) (4) Tmalaal laMal palpal aagiiial aatal caadtflan. 0) Palpal aegment adth veitigial aetae only, appearing unanned.(P) t l ) Hind wingRA2Hellgaaale«glb.(ai aaidlllri) 1)Pah)al aegment with aome ahort aetae.(A) 0)RA2»4 stigma ahort ca.2x longer than wide. 1) RA244 adgma medium-lengrii ca. 3x longer than wide. C4a) Teeaahaal labial palpal eepaenleetaleamdltlem. (nanaddtttve) 2) RA2+4 stigma long, more then 3x longer than wide. QNotapidicahle. 1) Palpal aegment armed mainly meaally with mhuile and abort aetae. U ) HbadwingRASelatae. 2) P a ^ aegment armed meaaBy, doraaOy, and laterally with ahort to medium B)RA3pteaent(P) ler«thtqiered aetae. l)RA3abeenl(A) 3) Palpal aegment anned meaoapically with lotiuat, epicaOy bhmt aetae and laterally and doraolaterally with aleoderahcrtirredhim length aetae. 53) HhidwlagllPlceadUen. B)RPI dark and diatinctXF) <5) Tamilnal lebh l palpal aagmawt em ana barleenlea atatna, l)RPIl*tbutmatinclXA) 0) Senailla beaioonica abort and aomewhat inoonapicuoua. Q RPl Bunt and indiatinct(A) 1) Senailla baaioonica relatively long and oon^icuoua. 54) Hind wing RP and MAl+2eendltlem. ( f ) Labial anbatmctnre central eanaa niarphalagy, 0)RP&MAI+2aqierale.(P) 0) Central arms fayabne aecdon alender and abort; pigmented aection dongate 1) RP partially fiiaed to MA1+2.(A) andcytindricaL(F) 2) RP completely Breed to MA l+2.(A) 1) Central arms hyabne aection slender and dongate; pigmented aection ahort andcyiindriGsL(A) 15) Hind wing RPloendttlen. 2) Central arms hyalhie aection ahort and robust or unapparent; pigmented 0)RP2oomplele.(P) aection broad and platdike.(A) 1) RP2 abbreviated epicdly 4k continuing to wing margin as aderotized 3) Central arms hyaline aection variable; pigmented aection vestigial.(A) membtane.(A) 2) RP2 abbrevhted apically 4t not continuing to wing margin as aderotized *7) Labial snbannctnt e ggnhr adeaite erlantatlen, membrane.(A) 0) Ugular aderitea parallel, not posteriorly convergent(P) 3) RP2oonaiatingofapically abbreviated aderotized membrane.(A) 1) Ligulsr aderhes poaterioily oonvetgent(A) 14) HlndwtngRPlHmndllian. (5) LabM anbati nrtuie alee marpbd agy. B)RP2oomplete.(P) 0) Also dongate and atrongdydevated and open ventromesallyXF) 1) RP2 abbreviated apically 4k continumg to wing margin as aderotized 1) Alae ahort end weakly devated and largely closed vcntromesally.(A) membtaneXA)

(f ) Labial srdbatriactw e body marpb alagy S7) HbadwbagMPltdrewdltlen, B) Body ofaubetructure bordering alee without lobe on each aide (IQ B) MP2f4 aderotized membrane complete 4k dark.(P) I) Body ofaubstructure bordering alae wtdi lateral lobe on eadiaife(A) 1)MP3M long but apically abbreviated 4k derk.(A) Q MP344 abort and &inL(A) 71) Labial subatiiialuaebedy la legprepertla«.(aansd dltlve) 0) Body elongate; about ulong as or longer than legs. 55) Hind wing CUA2+2M caadtUML 1) Legs dongate; mudt longer than body. 0) CUA2f3t4 oom^ete.(P) 1) aiA 2+2f 4 inoomplete.(A) 71) LAhd aamatrwdrare leg base aaerpbelegy. (nenaddltlre) B) Leg base with lateral extension lacking. SS) Hind wbigAP2t4 atatna. to Leg base extension hyaline ending in aderotized knob. 0)AP3t4 présenta») I) Leg base extension enthdy aderotized. 1)AP344absenL(A)

72) PraraetaldiaeerElbeeeculptarai SB) Marginal aaetaataanalantme atatna, to Pronotd diao earinateXP) 0) Suture indiatinct(P) I) Pronotd diao hddng cerina(A) 1)SutauediatinctXA)

72) Preetaraallabamarpbalagy. SI) M taemataitemalantnre atatna, to Anterior margin of proatemd lobe produoed.(P) 0) Suture atraiÿrt(P) 1) Anterior margin of proatemal lone attaight(A) 1) Suture outwardly arcuate or suture ebaenl(A)

74) Lataaabmarghuil preateawal atria atataa. Sla) Mete nretaatemal anture atatna. (nenaddltha) B) Suture carinate.00 0) Not applicable. 1) Suture not carhute.(A) 1) Suture outwardly arcuate. 2) Suture absent 316 n) UtnlM tutm nlratanf 0)SubB«abMnt(F) 1)SutunpieMnt(A)

0) Suture pce«enL(P) 1) Suture iliMntCA)

M) PnfTiUtBiamMhreiraMank 0) CtiinM tb ie n t^ 1) Creinre preient(A)

95) S l i re I t VIII fut w iw trei i n M itiftn 0) Pufaooent memlifins ibrenLCA) 1) PubcMoit membnne piaenLCIO

K) TreilrevmnwTffcBlip . (9 Itasits entire ot poBtoiody oii»giiut8.(F) 2) 1bgite Itikd pottenodyXA)

97) TertBeVni Jeci iifhilip . 0) Diio unUboniy idaotiad.(P) 1) Diu incdiaOy hyiline.(A)

M) CedteVnielylueelmnre. 19 Coxite with itylUL(P) l)St^leddng.(A)

99) Cexire VIII Mitphiin y. 0) Coxiies longa then conirel, end gaioiQir apically blunt 1) Coxitei about i> long at broad, donovaitnly flattened, and ipcilly bhmt

100) C axtoV m a 0) Coxita anned wifli mixture of minute A abort setae. 1) Coxitet aimed «idi numerous relativdy long setae.

101) Sip iW K peOaaraart eaanabeane rtahii 0) Pubescent membtane absent(A) 1) Pubescent membnne pcesent.(I9

102) TargtteKmeevhelecy. 0) Ibgite irtath i^ bcoad.(F) 1)1bigite vansbly reduced; oflen strep-li&e.(A)

103) Caxalcatagace stains. 0) Coxal caiaiMoe longer than broad.(P) 1) Coxaloaiapaoe broader than long or abacnl(A)

103a) Caial caragace status. (nan aOdWee) l9Notappbcable. 1) Coxal caiapaoe broader tiian long. 2) Coxal carapace absent

104) SfIcatuaa gastrale marp balagy. 0) ^acuhan gastrale elongate, broad A ptateIike.(F) 1)^iiculum gastrale elongate,nano* ArodbkeXA)

103) Pbalabaae maergbalegy. 0) Pbslobaae globuln or tubular A rdativeiy sboftCP) 1) Phalobaie tubular A rdativeiy long(A)

104) Paraaaaeva aasegbalegy. (uana iflltTi) 0) Panmeres divided apically tor more than lA tbeff length. 1) Parameres divided apically Ibr less than 1/2 their length. 2} Parameres apically fined.

107) 0) Parameres armed ventraBy. !) Parameres unanned ventnOy. leg) 0) Cutide not dragreenedOO 1) Cuticle dngrecned.(A)

103) Specialised eutfcalarpauderee. 19 Cutid e Iacldng spni6lirndpunctures.(l9 I) Cutide with umbiculate puncturesXA)

110) Vartttwe. 0) Body without acales.(F) 1) Body with scales.(A)

lU ) Badyfanu. 0) Bexly ibnn not globose.(F) 1) Body ibim globoseXA) APENDK C - PUPAL CHARACTERS

1) C ca«dM |riM *lM alM 4 0) G tktl KtObld not enluied or protnilnnLOO 17) Ahdssiilas! gPY stitas. 1) fiileil «ciflblil enlnged md protnitnnt(A) 0) gP Y setie present(IO 1) gPY setie ibsentCA) Hi Baa

5) BM4ediVriM«.(MMMIin) 0)VPielieptei

() Beeg Mix CL itatu. (D1 CL ictie pieienLOO I)CLidieibienL(A)

Cl) H iiC ietiC L itii» i.( i »-eCCWlTi) 0)Notipp)icibie. ÔlCLidiepieiaü ^ICLidiepteienL

7) B eaC ütiL A ditai. (OLAieUepRienLOO 1) LA ictie ibient(A)

I) PreHilgAWgrfgititM. 0) gAN wiUi 1 minimum of 3 id le per nle.(!0 1) gAN genaiDy with leu Hun 3 Mde pci ûie.(A)

S) PieedilgLNgreegiiWmi. 0) gLN with I miximum ofS id le per sd&CF) 1) gLN genenOy with 1 minimum of 7 id le per iide.(A)

10) Prenetilgm gieQ gpeiWeu, 0) gLK id le pruent ilong interior 1/2 of htenl niiigin.(IO 1) gLN letie restricted to posterior 1/3 oflstenl nuigia(A)

II) Prenetil gIN p u # stiriu. (mewCdlllri) 0) gIN with of 2-3 setie per side. 1) gIN genetiBy «ôth it leut 1 setie p a side. 2) gIN with 4 id le p a side. 3)gIN idieliddng.

12) M esethencki^ stU ae. 0) gSC with ■ k u then 3 setie p a iide.(P) 1) gSC with 3 ietie p a Bde.(A)

11) Meeethende^ST dates. 0) gST id le presentfF) 1) gSr setie ibeent(A)

14) M etU hsnclesdiM T ddas. 0) MT setie presenL(F) 1) MT setie sbient(A)

13) EhrtnlgELcsadittea. 0) gEL setie with 1 nuximum of 3 setu p a side.(IO 1) gEL oonsiiling of 3 sets! bends, esch consisting of 4-20 setie.(A)

14) Abdeadail gTE stitas, (asa iddlllii) 0) gfTE with 1 seti p a side. 1) ETE with 2-3 setie p a side. 2) ETE with 3 setie p a side.

317 APENDK D - ADULT DATA

Anapleus 0000000000 0000000000 1101000000 0000000000 0000000000 0000000000 0000000110 0000000000 0000000000 1100011121 3210000000 0000000000 0000000

Stictostix 0000101011 0000000000 1100000000 0100000000 0000010000 0000000100 0000000130 0000001200 0101000110 1200100031 2111000000 0000101120 0010100

Peploglyptus 0000101013 0000000001 0000000000 0100000000 0000010000 0100000100 0000000130 0000001200 0101000110 1200101031 2111120000 0010101120 0011110

TVibalus 0000101000 0000200001 0000000100 0100000000 0001011100 0100000101 2011101131 0010001312 1100200110 1100100021 2111000000 1010100001 0100000

Eutribalus 0010001113 0000012111 0011100110 1101012201 1101011210 1021120111 2112111000 0010011312 1101000010 1101010021 1110001000 0110001121 0100000

Parepierus 0010001113 0000011111 0011100110 1101012201 1101011210 1021120111 2112110151 0010111312 1101000110 0000120021 3210110000 0110001121 0100000

Paridolia 0110001100 0000011111 0001100110 1101012200 1101011210 1021120111 2112110010 0010011313 1100000110 1100110021 3110111000 0110001121 1100001

318 319 APPENDIX D (cont)

Idolia 0110001000 0000012111 0001100110 1101012200 1101011210 1021120111 2112110001 0010111313 1100000110 1101000021 1111120001 0110001121 0200001

Dendrophilus 0000000000 0001100001 0000000000 0000000000 0000000000 0000000001 1011101001 0000000000 0000000000 0000000000 0000000000 0000000000 0000000

Onthophilus 1000001011 0001300010 1201000000 0010011010 0010000000 0000111000 0011101120 1101001101 0010111011 1210001000 0000120011 0010010000 0010000

Vuattouxinus 1011011012 0001300010 0001000001 0110011010 0000001100 0100111001 1011101120 0000000001 0010011111 0000000001 1010001001 1011101110 0200010

Epiechinus 1011011112 0001311010 0001000001 0100011010 0010001100 1000001001 1011001120 0000000001 0000011011 1210001001 1010001111 1011111110 0201010

Australepierus 0010001013 0101311010 0001001001 0100111010 0010001100 0010111000 0011001121 0001000001 0010001110 0000021011 1011001101 0010001000 1010000

Plagiogramma 0010001011 1111313011 0001011011 0101112100 0000100001 0120110000 0000100141 1200001200 0111100110 0000020010 0011110000 0010000110 1010000

Epierus 0010001011 1111313011 0001011011 0101112100 0000100001 0120110000 0000100140 1210001201 0101000110 0000021010 0011120000 0010000120 1100000 APENDK E - LARVAL DATA

Dendrophilus 1101111010 0000000110 00001000010 0000000102 0000000000 112110110010 10200010000 22012010000 1001120200 0221002001 100011100 1002111100 121220012000 0100011100 020012000 0011000101 133300002 20000000101 0000200000 0000002000 12?

Onthophilus 1101100000 1111110000 00000010101 0111110111 1200000120 000011101210 10200010000 01001011001 0002120000 0110001110 011211000 1010110000 000000000000 0100100000 000000000 1010010101 001000001 10000112111 1100100101 1101000000 100

Australepierus 0000000110 0011010001 10010011010 1000111010 1210000120 000100001101 01010000011 00000001000 1110110000 1100110120 001200011 0010000000 000011000111 1010000011 011200010 1100001010 011310101 00001112011 0010111101 1100001100 010

Plagiogramma 0010011010 0011010001 10010101100 0100111010 1111111001 100001001101 01011101121 10100100000 1110001011 1000100120 000000011 0101000000 000012000101 1010000010 011211011 1110001120 012110100 11011013101 0011111111 1110100111 110

Epierus 0010011110 1100110001 10011101111 1011000011 1110001000 000000001101 01001100011 00000001000 1110111011 1100101120 001200100 1000000000 000012000101 1010000010 011211001 1010011110 022110100 01011012000 1111111101 1110100111 010

320 321 APPENDIX E (cont)

Anapleus 0000000000 0000000001 20012000000 0000000000 0000000110 000000000000 01000000000 10000101011 0110000000 0000000000 110000000 0000000011 000000000201 2000000000 000000000 0000101000 000400010 00100000010 0010000000 0001110100 010

Eutribalus 1201211111 1212011121 11112010020 1112110010 0010011120 111000011101 10011211210 01111100111 1200100100 0111110111 110000101 1101001112 111210111011 1011111111 120011100 0101101000 133201010 00100011010 1200101011 2110012212 121

Idolia 1201211101 1111001121 11111111010 0000120002 1201200002 000000000000 00101211200 01121200112 0201102112 0100010112 111100101 1101000012 110120100110 0011011211 121112100 0101101100 133301010 00101000011 0010001011 2110010212 121 APENDK F - PUPAL DATA

Dendrophilus 0001101000 0000001000

Onthophilus 0110110000 0301100000

Australepierus 0000002101 000011211?

Plagiogramma 0010001100 0200101000

Epierus 0010001100 0100101000

Idolia 1010010110 1111101101

Eutribalus 1110110110 1311101001

322 REFERENCES

Alaire, Y. 1990. Primaiy setae and pores on the cephalic capsule and head appendages of larval Hydroporinae (Coleoptera: Dytiscidae: Hydroporinae). Can. J. Zool. 69: 2255- 2265.

Ashe, J. S. & L. E. Watrous. 1984. Larval chaetotaxy of Aleocharinae (Staphylinidae) based on a description oîAtheta coriaria Kraatz. Coleop. Bull. 38: 165-179.

Baulduf, W. V. 1935. The bionomics of entomophagous Coleoptera. Part 1. John Swift Co., St. Louis. 220 p.

Belkin, J. N. 1950. A revised nomenclature for the chaetotaxy of the mosquito larva (Diptera: Culicidae). Am. Midi. Nat. 44: 678-698.

Bickhardt, H. 1916. Ksteridae. Wytsman, Genera insectorum. 166a. pp. 1-125.

Bickhardt, H. 1917. Histeridae. Wytsman, Genera insectorum. 166b. pp 113-302.

Borden, J. H. 1968. Antennal morphology QÎIps confusus (Coleoptera: Scolytidae). Ann. Entomol. Soc. Am. 61:10-13.

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