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1 INFORMATION T O USERS This reproduction was made from a copy of a manuscript sent to us for publication and microfilming. While the most advstnced technology has been used to photograph and reproduce this manuscript, the quality of the reproduction is heavily dependent upon the quality of the material submitted. Pages in any manuscript may have indistinct print. In all cases the best available copy has been filmed. The following explanation of techniques is provided to help clarify notations which may appear on this reproduction. 1. Manuscripts may not always be complete. When it is not possible to obtain missing pages, a note appears to indicate this. 2. When copyrighted materials are removed from the manuscript, a note appears to indicate this. 3. Oversize materials (maps, drawings, and charts) are photographed by sectioning the original, beginning at the upper left hand comer and continuing from left to right in equal sections with small overlaps. Each oversize page is also filmed as one exposure and is available, for an additional charge, as a standard 35mm slide or in black and white paper format. * 4. Most photographs reproduce acceptably on positive microfilm or microfiche but lack clarify on xerographic copies made from the microfilm. For an additional charge, all photographs are available in black and white standard 35mm slide format.* *For more information about black and white slides or enlarged paper reproductions, please contact the Dissertations Customer Services Department. T TA /Î-T Dissertation i J l Y l i Information Service University Microfilms International A Bell & Howell Information C om pany 300 N. Z eeb Road, Ann Arbor, Micfiigan 48106

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3 Stribling, James Bentley WORLD GENERIC REViSION OF PTILODACTYLIDAE (COLEOPTERA;DRYOPOIDEA) The Ohio State University Ph.D University Microfilms I nternsti 0 nâl SOO N. zeeb Road, Ann Arbor, Ml 48106

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5 WORLD GENERIC REVISION OF PTILODACTYLIDAE (COLEOPTERA: DRYOPOIDEA) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By James B. Stribling, B.S., M.S. * * * * * The Ohio State University 1986 Dissertation Committee: Approved by C. A. Triplehorn N. F. Johnson D. E. Johnston Û- Adviser Department of Entomology B. D. Valentine

6 Copyright by James Bentley Stribling 1986

7 To My Parents ii

8 ACKNOWLEDGEMENTS There are many people to whom I am very grateful for academic, moral, physical, and financial support during my graduate career at The Ohio State University: my adviser. Dr. Charles A. Triplehorn, for sound advice for the duration and understanding when I needed it most; Dr. Norman F. Johnson for numerous discussions, suggestions, and welcome advice on systematic theory and taxonomic procedure; my officemates, colleagues, and fellow graduate students, especially, Paul S. Cwikla, Michael A. Ivie, Richard S. Miller, and John A. Shuey, for seemingly unending discussions on systematics, for companionship on collecting trips, and for tolerance during these last few months; John F. Lawrence for much advice and comment on this as well as smaller simultaneous projects; W. Calvin Welbourn for many hours spent in setting up the AT&T 3B2 and QTREE and then tutoring me on their use; John F. Lawrence, Charles W. O'Brien, and Arthur J. Gilbert for gifts of specimens; Roland L= Seymour (Department of Botany, The Ohio State University) for fungal identificaiii

9 tions; Ho-Yih Liu (also Department of Botany, 0. S. 0.) for translating some of Nakane's works from Japanese; all of the curators in the "Materials" section for giving me the opportunity to study their specimens (even past expiration dates on loan forms); Niki Lowell and Adam Rubinstein for the habitus illustrations (Figures 3 (N. L«) and 1 and 2 (A. R.); and Helen W. Monk for typing part of the manuscript. Partial funding of museum trips was provided by Sigma Xi (Grant-in-Aid of Research), The Ohio State University Graduate School (Graduate Student Alumni Research Award), and the Knull Fund, Department of Entomology. I am very grateful to them. I owe immense thanks to my wonderful parents. Dr. and Mrs. Loutrelle, for their financial support, endless moral support, encouragement, and for all of those times I needed "a good talkin' to" and got it. Finally, I thank my wife, Sally, for the hours of assistance she has given me in my research, taking time out of her own to process specimens, to type, or to take over household duties when I might be nearing a deadline. My greatest thanks to her are for her unwavering love, encouragement, and companionship. IV

10 VITA October 13, Born - Jackson, Mississippi B. S., The University of Mississippi, Oxford, Mississippi 1982 # «# #M# S #, The Ohio State University, Columbus, Ohio 1980-present...Graduate Research/Teaching Associate, Department of Entomology, The Ohio State University PRESENTATIONS Phylogenetic Implications of Metathoracic Wing Venation in Dryopoidea (Coleoptera). Annual Meeting of Entomological Society of America, Hollywood, Florida. The Biology and Systematics of Anchvtarsus (Coleoptera: Ptilodactylidae). Annual Meeting of the North Central Branch, Entomological Society of America, Lexington, Kentucky. The Biology, Phylogeny, and Zoogeography of the Wing-Toe Beetles (Ptilodactylidae). Departmental seminar. Department of Entomology, The Ohio State University, Columbus The Faces of Systematic Biology. Invited seminar. Department of Biology, The University of Mississippi, Oxford.

11 1981. The New World Genera of the Ptilodactylidae (Coleoptera). Annual Meeting of the Entomological Society of America, San Diego, California. PUBLICATIONS Revision of Anchvtarsus (Coleoptera: Dryopoidea) and a key to the New World genera of Ptilodactylidae. Ann. Entomol. Soc. Amer. 79: Taxonomic and nomenclatorial notes on Caribbean Tropicus Pacheco (Coleoptera: Heteroceridae). Proc. Ent. Soc. Wash. 86 (4): (with M. A. Ivie) Report on the LeConte symposium on the classification and phylogeny of the Coleoptera, Detroit and Columbus, Coleop. Bull. 38 (1): (with M. A. Ivie) FIELDS OF STUDY Major Field: Systematic Entomology with specialization in Coleoptera biosystematics. VI

12 TABLE OF CONTENTS ACKNOWLEDGEMENTS... iii VITA... V LIST OF TABLES... ix LIST OF FIGURES x CHAPTER PAGE I. INTRODUCTION...1 II. METHODS AND MATERIALS...4 Methods Phylogenetic... 4 General... 8 Material III. NATURAL HISTORY...16 IV. CHARACTERS, POLARIZATION, PHYLOGENETIC ANALYSIS, AND CLASSIFICATION Monophyly of the Ptilodactylidae Character discussion H e a d Thorax Abdomen Character polarization Discussion of phylogenetic analysis and resulting classification V. TAXONOMY...38 Key to World Genera of Ptilodactylidae (Adults) The World Genera of Ptilodactylidae Anchvtarsus Guerin-Meneville vii

13 Cladotoma Westwood Paraiichas White Hovactvla Fairmaire Pseudocladotoma Pic Octoqlossa Guérin-Méneville Bradvtoma Guérin-Méneville Aploqlossa Guérin-Méneville Anchycteis Horn Epilichas White Bvrrocrvptus Broun Araeopidius Cockerell Stenafricus NEW GENUS Daemon de Laporte Falsotherius P i c Pseudoepilichas Armstrong and Nakane Ectvphodactvla NEW GENUS Ptiloderes NEW G E N U S Pherocladus Fairmaire Lachnodactvla Champion Ptilodactvla Illiger Stirophora Champion Chelonariomorphus Pic Epiptyqma NEW GENUS Therius Guerin-Mènev i l i e Incertae sedis Exclusions LITERATURE CITED APPEmiX A. Figures Vlll

14 LIST OF TABLES TABLE PAGE 1. Fungal spores and hyphae identified from gut content of some species of Ptilodactlidae Hypothesized polarities of characters used in phylogenetic analysis of the genera of Ptilodactylidae Phylogenetic hypothesis resulting from QTREE analysis of characters in Table 2. Reversals are indicated by dots at the upper right hand side of the character numbers; characters with multiple apomorphic appearances are 2, 3, 4, 5, 7, 9, 11, 12, 13, 14, 15, 16, 18, 20, 23, 25, 27, 29, 31, 35, and » Proposed supraspecific classification of the Ptilodactylidae based on phyletic sequencing 37 ix

15 LIST OF FIGURES FIGURES PAGE 1. Dorsal habitus. Anchvcteis velutina, male Dorsal habitus, Ptilodactvla militaris, male Dorsal habitus. Ptiloderes sp., female.» Serrate antenna; Bradvtoma lineata Ramous antenna, nonarticulated rami; Paralichas q u e r i n i Ramous antenna, nonarticulated rami, Cladotoma m aculicollis Ramous antenna, articulated rami; Ptilodactvla sp S. Tarsus, lateral view showing simple tarsomeres and large onychium; Eulichas sp Tarsomere IV, dorsal view showing large onychium; Eul%cnas sp....o Pectinate ungues; Paralichas querini Bifid ungues; Ptilodactvla sp. The mesal furcations are often more or less truncate Modified tarsus showing reduction of tarsomere IV, ventral lobe of III, simple ungues, and reduced onychium; Cladotoma cucullatus Simple tarsus: Araeopidius monachus Simple tarsus showing minor lobes on tarsomeres II and III, and no reduction of IV; Pseudocladotoma maculata Labium, ventral view, palp and setae omitted from X

16 right half; modified ligula, biacuminate with each half tapered to blunt apex; Ptiloderes sp Labium, ventral view, palp and setae omitted from right half; hexacuminate ligula; Bradvtoma a u r i t a Labium, ventral view, palp and setae omitted from right half; modified ligula, biacuminate with each half tapered to blunt apex, apical palpomere with highlighted area of membranous region showing portion of setal clothing; Ectvphodactvla fiski Labium, ventral view, palp and setae omitted from right half; modified ligula, biacuminate; Falsotherius atricolor Labium, ventral view, palp and setae omitted from right half; modified ligula, quadracuminate with lateral apices reduced; Anchvcteis velutina Labium, ventral view, palp and setae omitted from right half; modified ligula, quadracuminate; B v r rocry ptus sp Labium, ventral view, palp and setae omitted from right half; modified ligula, quadracuminate; Epilichas candezei Labium, ventral view, palp and setae omitted from right half; modified ligula, quadracuminate; Cladotoma sp Labium, ventral view, palp and setae omitted from right half; simple ligula folded ventrally; Stirophora Ivciformis Labium, ventral view, palp and setae omitted from right half; modified ligula, multiacuminate; Paralichas trivittis Labium, ventral view, palp and setae omitted from right half; modified ligula, hexacuminate; Octoqlossa sp Left maxilla, ventral view; multiacuminate galea; Paralichas sp Right maxilla, ventral view, lacinia forming spore xi

17 brush, apical palpomere elongate and largely membranous; Lachnodactvla parviscutum Left maxilla, ventral view, biacuminate galea; Epilichas candezei Left maxilla, ventral view, biacuminate galea with mesal branch absent; Anchvcteis velutina Left maxilla, ventral view, biacuminate galea with mesal branch reduced; Bvrrocrvptus sp Left maxilla, ventral view, lacinia forming spore brush; Aploqlossa aureonotata Left maxilla, ventral view, galea and lacinia forming spore brush; Daemon gigas Left maxilla, ventral view, cylindrical galea with apical cluster of setae; Bradvtoma aurita Left maxilla, ventral view; Ptiloderes sp Right maxilla, ventral view; Falsotherius at r1color Right maxilla, ventral view, apical palpomere with highlighted area of membranous region showing portion of setal clothing; Ectvphodactvla fiski Left mandible, dorsal view; Aploqlossa aureonotata Left mandible, dorsal view, showing concave molar area; Daemon gigas Left mandible, dorsal view; Paralichas querini Right mandible, dorsal view; Octoqlossa cvaneipennis Left mandible, dorsal view; Lachnodactvla parviscutum Left mandible, dorsal view; Anchvcteis velutina Left mandible, dorsal view; Epilichas candezei.. xii

18 Labrum, dorsal view; Falsotherius atricolor Labrum, dorsal view; Aploqlossa aureonotata Labrum, dorsal view; Cladotoma sp Labrum, dorsal view; Ptilodactvla sp Labrum, dorsal view; Anchvcteis velutina Head and prothorax, frontal view, shaded areas = eyes, blackened areas = antennal Insertions; Octoqlossa sp Head and prothorax, frontal view, shaded areas - eyes, blackened areas = antennal insertions; Araeopidius monachus Head and prothorax, frontal view, shaded areas = eyes, blackened areas = antennal insertions; Cladotoma sp Head and prothorax, frontal view, shaded areas = eyes, blackened areas = antennal insertions; Ptiloderes sp Bead and prothorax, frontal view, shaded areas = eyes, blackened areas = antennal insertions; Ectvphodactvla fiski Head and prothorax, frontal view, shaded areas = eyes, blackened areas = antennal insertions; Ptilodactvla sp Tergum and sternum of male visible abdominal segment VI, lateral view; Eoiptvcma rufa Male visible abdominal segment VI, dorsal view of Figure 55; Epiptyqma rufa Tergum and sternum of male visible abdominal segment VI, lateral view; Epiptyqma sp Male visible abdominal segment VI, dorsal view of Figure 57; Epiptyqma sp Tergum and sternum of male visible abdominal segment VI, lateral view; Epiptyqma sp xiii

19 60. Male visible abdominal segment VI, dorsal view of Figure 59; Epiptvoma sp Head and prothorax, lateral view, shaded area = eye; Pherocladus sp Head and prothorax, lateral view, shaded area = eye; Araeopidius m o n a c h u s Head and prothorax, lateral view, shaded area = eye; Octoqlossa sp Head and prothorax, lateral view, shaded area = eye; Paralichas q u e r i n i Head and prothorax, lateral view, shaded area = eye; Bradvtoma lineata Head and prothorax, lateral view, shaded area = eye; Ectvphodactvla fiski Head and prothorax, lateral view, shaded area = eye; Chelonariom orphus sp Head and prothorax, lateral view, shaded area = eye; Ptilodactvla sp Anterior dorsum; Araeopidius monachus Anterior dorsum; Chelonariomorphus subconvexus Anterior dorsum; Ectvphodactvla fiski Anterior dorsum; Paralichas querini Anterior dorsum; Ptiloderes sp Pronotum, dorsal view; Ptiloderes sp Pronotum, dorsal view; Ptiloderes sp Pronotum, dorsal view; Ptiloderes sp Ovipositor, dorsal view; Aploqlossa sp Ovipositor, dorsal view; Araeopidius monachus XIV

20 79. Ovipositor, distal portion, dorsal view; Paralichas trivittis Ovipositor, distal portion, dorsal view; Stirophora sulcipennis Ovipositor, distal portion, dorsal view; Daemon sp Ovipositor, distal portion, dorsal view; Falsotherius atricolor Ovipositor, dorsal view; Cladotoma sp» Ovipositor, right lateral view of Figure 83; Cladotoma sp Aedeagus, ventral view; Bradvtoma aurita Aedeagus, right lateral view of Figure 85; Bradvtoma aurita Aedeagus, dorsal view; Aploqlossa aureonotata., Aedeagus, right lateral view of Figure 87; Aploqlossa aureonotata Aedeagus, dorsal view; Ectvphodactvla fiski Aedeagus, dorsal view; Chelonariomorphus sp Aedeagus, dorsal view; Paralichas querini Aedeagus, dorsal view; Stenafricus sp» Aedeagus, dorsal view; Cladotoma sp Aedeagus, dorsal view; stirophora Ivciformis Aedeagus, dorsal view; Epilichas candezei Aedeagus, dorsal view; Lachnodactvla monticola XV

21 CHAPTER I INTRODUCTION The Ptilodactylidae is a family of semiaquatic and terrestrial beetles which is nearly cosmopolitan in distribution (Stribling, 1986). Species are present in every major biogeographic region except the Palearctic and Chile. The greatest species diversity is in tropical areas with roughly similar numbers in the Old and New Worlds. Characteristics traditionally used to recognize ptilodactylids have been a heart-shaped (= cordiform) mesoscutellum, pseudotetramerous tarsi (with the third tarsomere ventrally lobed and the fourth reduced), and the pectinate antennae of the males. Exclusive use of these characters has allowed recognition of several genera as ptilodactylids, but has also led to confusion regarding phylogenetic relationships among these genera as well as between the Ptilodactylidae and other Dryopoidea (Crowson, 1967, 1978; Lawrence and Newton, 1982; Arnett 1968). An extension of this confusion is a misunderstanding of the generic composition of the family (Arnett, 1968; Borror, ^ al. 1978). Boving and Craighead (1931) recognized affinities between the larvae of Anchvtarsus Guerin-Meneville and Ptilodactvla Illiger and placed them together, elevating

22 the group to familial status. Later, fourteen more genera 2 were transferred from the Dascillidae to the Ptilodactylidae (Crowson, 1967). For a more complete review of the suprageneric taxonomic history see Stribling (1986). There have been few regional studies of the Ptilodactylidae in the Old World. Deleve (1972) investigated the fauna of the Philippines and the Bismarck Archipelago, describing 20 species of Ptilodactvla, one of Pherocladus Fairmaire, and the genus valoka Deleve. Many papers have treated various Oriental species (White, 1859; Nakane, et 1963; Nakane, 1948, 1950, 1952, 1956, 1963, 1977; Sato, 1964, 1968, 1979, 1983; Chûjô and Sato, 1970; Lewis, 1895) usually describing small numbers of Ptilodactvla, Epilichas White, and Paralichas White. Armstrong and Nakane (1956) described Pseudoepilichas from Japan. Even fewer studies have dealt with the New World fauna. Johnson and Freytag (1978, 1982) described species and reviewed the Ptilodactvla of the United States. Champion, in the "Biologia Centrali-Americana" (1897), recognized fifty new species of Ptilodactvla and four new genera from Mexico and Central America. Between 1916 and 1958, Maurice Pic described 206 species of Ptilodactvla (102 in the New World), and 7 monotypic genera. This is the first comprehensive study of the Ptilodactylidae. Although I was unable to examine the

23 types of all described species, I was able to study 3 representatives of nearly all genera and of many species. As a result, I present a phylogenetic hypothesis of the major monophyletic lineages of the Ptilodactylidae of the world. This hypothesis will allow evaluation of phylogenetic relationships of subsequently studied or discovered taxonomic units, and of the significance of new characters and character states.

24 CHAPTER II METHODS AND MATERIALS Phylogenetic Delineation of supraspecific taxonomic categories in classifications has long been highly subjective, often being based on overall similarity rather than on objective statements of phylogenetic relationship. Cladistic methodology (Hennig, 1966; Eldredge and Cracraft, 1980; Wiley, 1981) has provided systematic biology with a method for recognition of groups of organisms sharing unique phylogenetic history. Construction of phylogenetic hypotheses through employment of these methods produces a hierarchically-arranged diagram of operational taxonomic units (OTü's) which may be translated into the hierarchy of the Linnean system of classification. Representation of this common history in classificatory form has been discussed in regard to its information content (Farris, 1980) and application and necessity for non-systematic biological subdisciplines (Dobson, 1985). In this study I use outgroup comparison analysis for postulation of derived, or apomorphic, character states. Joint possession of apomorphic character states among OTü's indicates probable common ancestry and monophyly. I recognize monophyletic taxa (Hennig, 1966) as those

25 composed of: 1) the most recent common ancestor which is 5 inferred to possess a particular apomorphy, and 2) all known descendants of that ancestor. Paraphyletic taxa (Farris, 1974) are converted into monophyletic ones by synonymization and/or splitting of previously recognized taxa. In some cases this requires erection of new taxa. Eldredge and Cracraft (1980: 39) point out that monophyletic higher taxa must consist of two or more species. Monotypic genera, therefore, cause non-monophyly (specifically, paraphyly) in their sister-genera by allowing exclusion of one of the descendants of that ancestor. The only ancestor which a species has in common with a lineage is also ancestral to another part of that lineage. To avoid paraphyly, all descendants of that ancestor must be included in the taxon. If the sister-otu of the species is of preexisting generic status, they are combined. In all cases this tactic produces a monophyletic genus. In other words, a monotypic genus may only be recognized when its species possesses no synapomorphy with a preexisting genus. The most important single component of phylogenetic methodology is the postulation of the relative polarity of character states. Although many criteria have been discussed as potential aids in this inference (Crisci and Stuessy, 1980; de Jong, 1980), outgroup comparison is

26 6 regarded as the most reliable method (Watrous and Wheeler, 1981; Haddison et al., 1984; Donoghue and Cantino, 1984) and I have used this technique exclusively. The Ptilodactylidae are included in the Dryopoidea (Crowson, 1955, 1960, 1978; Lawrence, 1982; Lawrence and Newton, 1982) the latter having been hypothesized to consist of two main lineages (Lawrence and Newton, 1982): 1) Callirhipidae - Eulichadidae - Ptilodactylidae - Chelonariidae; 2) Heteroceridae - Lutrochidae - Limnichidae - Dryopidae - Elmidae - Psephenidae. The Eulichadidae have also been hypothesized to be the sister-group of the Ptilodactylidae (Crowson, 1978). Assuming monophyly of the former lineage, I used the following taxa for outgroup comparison: Eulichadidae Eulichas spp. Stenocolus scutellaris LeConte Callirhipidae Zenoa picea (Beauvois) unidentified sp. from Vietnam Psephenidae Psephenus herricki (DeKay) After demarcation of characters within Ptilodactylidae into binary components, hypotheses of apomorphy through the process of polarization were sought. This process

27 entailed the following steps: 7 1) Check for occurrence of character state in Eulichadidae. Absence indicates probable apomorphy. 2) Check for occurrence in Callirhipidae. Absence corroborates hypothesis of apomorphy. 3) Check for occurrence in Psephenidae. Absence, further corroboration. If the character state is found in Eulichadidae and either Callirhipidae or Psephenidae, it is hypothesized plesiomorphic and its alternative state, apomorphic. In some cases, reference to outgroups did not provide evidence for the polarity of complex variation, such as that of the mouthparts and tarsi. Rather than attempt to align this variation into polarized transformation series I have chosen to follow what I consider to be a more conservative approach. With no evidence as to polarity or of the interrelationships of the character states, any attempt at coding them as a series would produce immediate subjective bias. I experimented with coding each of the states as independent apomorphies; however, this introduced much homoplasy and longer trees. Elimination of these characters from the analysis lowered homoplasy and consequently shortened trees. The phylogenetic computer program used for character matrix analysis was QTREE, written in C by Dr. L. E.

28 Watrous (FMNH); it was executed on an AT&T 3B2/300. The 8 program searches for the most parsimonious trees using the mode of search specified by the options chosen. I used the following options: -bo This specifies the number of pathways to be explored in the search for the shortest tree. The numerical value can be altered from 0-3, each increase causing more trees to be examined. -k2 This specifies that the computer randomly sort the OTD*s; the numerical value, how many times the sort should be repeated. General Relaxing. - Specimens were relaxed in an allpurpose ammonia and detergent household cleanser in a sonic cleaner. They were then vibrated in tap water to remove the ammonia and detergent. Specimens mounted on points or cards were removed before relaxing. Dissecting. - All mouthparts and genitalia were removed from relaxed specimens with jewelers' forceps (no. 3 or no. 5) and/or minuten pins mounted in wooden splints. Dissected parts were dried of excess water and placed in glycerine in small glass dishes in a wooden holder (DeLong and Davidson, 1937) for serial observation. Clearing. - when necessary for viewing detail.

29 dissected structures were cleared in hot KOH and water 9 solution for approximately 3 minutes. The specimen was then removed; rinsed in cold tap water, and examined. If degree of clearing was insufficient, it was placed back in the KOH and heated further. Illustration. - Mouthparts or genitalia were placed on a shallow depression slide with a minimal amount of glycerine. The specimens were covered with a drop of K-Y (registered trademark) Lubricating Jelly (Johnson & Johnson), positioned, and allowed to stand for 2-3 minutes. The specimen was then covered with a drop of glycerine and allowed to stand for another 2-3 minutes until the K- Y/glycerin interface stabilized. Illustration then commenced. The K-Y gel is water soluble and, when allowed to stand uncovered, loses moisture. Thus, it becomes somewhat coagulated, preventing drifting of the specimens. This technique (P. S. Cwikla, pers. comm.) is ideal for illustration purposes. Illustrations were made with a camera lucida on Wild M5A dissecting scope for all except the mouthparts and genitalia which were illustrated with a drawing tube on a Wild H12 compound microscope. Storage of dissections. - Dissected structures are stored in polyethylene microvials with a drop of glycerine. The vials are mounted on the same pins with the specimens. Taxonomic lists. - The taxonomic lists following the

30 descriptions contain all names, specific, subspecific, and varietal, which have been published under the ptilodactylid 10 genera. Asterisks (*) indicate those of which I have seen holotypes, type-compared specimens, or specimens which I feel were reliably determined. The specific epithets are listed in alphabetical order followed by author and date of description; acronyms in parentheses indicate the location of the holotype or type series as known by my examination of the specimen, personal communication with curators, or from the literature. At the end of almost all entries is a geographic locality indicating where the type specimen(s) was (were) collected; in some instances additional information was added from label data. Occasionally, no locality was published with the description and I have seen no determined specimens.

31 Materials 11 This study is based on approximately 25,000 specimens accumulated through loans from the following museum, university, and private collections. Acronyms listed are those used in the text for citation of specimens and for type repositories, AMNH American Museum of Natural History, New York, NY; L. H, Herman, Jr. ANSP Academy of Natural Sciences of Philadelphia, Philadelphia, PA; Donald Azuma. BMNH British Museum (Natural History), London, England; C, M. F. von Hayek, P. M, Hammond, CASC California Academy of Sciences, San Francisco, CA; D. H, Kavanaugh, Jr, CBBC Cheryl B, Barr, private collection. Baton Rouge, LA, CISB California Insect Survey, University of California, Berkeley, CA; J. A, Chemsak, CNCI Canadian National Collection of Insects, Agriculture Canada, Ottawa, Ontario; J, M, Campbell, L. LeSage, A. Smetana, CSIR Commonwealth Scientific and Industrial Research Organization, Canberra, ACT; J, F, Lawrence,

32 CüCC Department of Entomology, Clemson University, Clemson, SC; J. C. Horse. 12 DKYC EEHC Daniel K. Young, personal collection, Madison, WI. Enns Entomological Museum, University of Missouri, Columbia, MO; E. G. Riley. EGRC Edward G. Riley, personal collection. Baton Rouge, LA. FMNH Field Museum of Natural History, Chicago, IL; H. G. Nelson. FSCA Florida State Collection of Arthropods, Gainsville, FL; M. C. Thomas, R. E. Woodruff. GLPC Gary L. Peters, personal collection, Corvallis, OR. HAHC Henry F. and Ann T. Howden, personal collection, Ottawa, Ontario. HUIC Museum fur Naturkunde der Humboldt-Universistat zu Berlin; M. Uhlig. ICCM Carnegie Museum of Natural History, Pittsburgh, PA; G. Ekis. INHS Illinois Natural History Survey, Urbana, IL; D. W. Webb. INPA Institute Nacional de Pesguisas da Amazonia, Manaus, Brasil; V. Py-Daniel. IZWP Institut Zoologique, Académie Polonaise des Sciences, Warszawa, Poland; S. A. Slipinski.

33 JBSC 13 James B. Stribling, personal collection, Columbus, OH. JEWC James E. Wappes, personal collection. Woodlands, TX. KJOC KÜBC Karl Joplin, personal collection, Columbus, OH. Kagoshima University Biological Collections, Kagoshima-Shi, Japan; Takehiko Nakane. LACM Los Angeles County Museum of Natural History, Los Angeles, CA; J. P. Donoghue, R. R. Snelling. LGBZ Larry G. Bezark, personal collection, Sacramento, CA. LSUC Louisiana State University, Baton Rouge, LA; C. B. Barr, J. B. Chapin. MAIC MCZC Michael A. Ivie, personal collection, Bozeman, NT. Museum of Comparative Zoology, Harvard University, Cambridge, MA; A. F. Newton, Jr. MNHP Museum National D'Histoire Naturelle, Paris, France; N. Berti, C. Girard. MRAC Musée Royal de L'Afrique Centrale, Tervuren, Belgium; J. Decelle. MSUC Michigan State University, Lansing, MI; R. L. Fisher. HUIC Mississippi State University, Starkville, MS; R. Brown. MZSP Museu de Zoologia da Universidade de Sao Paulo,

34 Sao Paulo, Brazil; Cleide Costa. 14 NARZ Northern Arizona University, Flagstaff, AZ; C. D, Johnson. NCSÜ North Carolina State University, Raleigh, NC; C. Parron. NDSU North Dakota State University, Fargo, ND; E. U. Balsbaugh, Jr. NMNH National Museum of Natural History, Smithsonian Institution, Washington, D. C.; P. J. Spangler, J. M. Kingsolver. OSUC Ohio State University, Columbus, OH; C. A. Triplehorn. OSUO Oregon State University, Corvallis, OR; G. L. Peters. RHTC Robert H. Turnbow, personal collection. Fort Rucker, AL. RSMC Richard S. Miller, personal collection, Columbus, OH. RSNB Institut Royal des Science Naturelles de Belgique, Bruxelles, Belgium; P. Dessart. RUIC Rutgers University, New Brunswick, NJ; G. W. Wolfe. SMSH Stovall Museum of Science and History, University of Oklahoma, Norman, OK; H. P. Brown. TAMT Texas A & M University, College Station, TX; S.

35 Merritt. ÜART UBCZ University of Arizona, Tuscon, AZ; F. G. Werner. University of British Colombia, Vancouver, BC, Canada; S. G. Cannings. UCDC University of California, Davis, CA; R. 0. Schuster. UICM University of Idaho, Moscow, ID; W. F. Barr, Jr. UMCE Université de Montreal, Quebec, Canada; M. Coulloudon. UMIS University of Mississippi, Oxford, MS; P. R. Lago. UNTC Universidad Nacional de Tucuman, Argentina; A. Willink. VPSU Virginia Polytechnic Institute and state University, Blacksburg, VA; M. Kosztarab. The following acronyms are used only in reference to type repositories. ANMM BANM National Museum of Victoria, Melbourn, Australia. Museu de Ciencias Naturales, Buenos Aires, Argentina. DEIE Institute of Plant Protection Research, Eberswalde, Germany (formerly, Deutsches Entomologisches Institut). HECM Hope Entomological Museum, Oxford University, Cambridge, England. UZHD Universiteits Zoologiske Museum, Copenhagen.

36 CHAPTER III NATURAL HISTORY In general, the biology of ptilodactylids is poorly known. The larvae of Anchvtarsus are aquatic and feed on decaying vegetable material in lotie situations (LeSage and Harper, 1976; Spangler, 1966, 1981, 1982, 1983; Stribling, 1986). Though found in flowing water, individuals are not subjected to the force of the current due to specific location within leaf packs. Anchvtarsus bicolor has also been collected in large numbers from beaver dams in Ontario (L. LeSage, pers. comm.). Other larvae known to be aquatic, Anchvcteis velutina Horn and Araeopidius monachus (LeConte), have been reported to burrow in rocky substrates and to feed on the roots of emergent vegetation (Leech and Chandler, 1956; Brown, 1972), They have also been listed as feeding on plant materials by shredding (Doyen and Ulrich, 1978) and as boring in soft wood (Dudley and Anderson, 1982). Specimens of Anchvcteis have been collected from the vegetation within a bog runoff; and Araeopidius from duff and leaves in creeks (H. P. Brown, pers. comm.). On 3 July, 1982, in Oaxaca (near Vista Hermosa) I collected five species of Ptilodactvla (95 specimens) by 16

37 beating in an area of approximately 20 square meters. Some 17 of the vegetation was spotted with sooty mold; gut content analysis of these specimens has yielded no structures which I could identify as fungal spores or hyphae. However, fungal components were discovered in the guts of other specimens in the course of genitalic dissections (see Table 1). White (1859) described Paralichas querini from specimens which had emerged from structures he proposed were pupal cases of the beetles. When I examined these specimens in the BMNH I found an anonymous note attached to one of them saying that the structure is a tineid larval case (Lepidoptera). A noté on another specimen states that both the tineid larva and beetle have been removed. The largest numbers of adults are collected nocturnally at lights. Diurnally, they are found on vegetation with the aquatic species usually near water.

38 Table 1. Fungal spores and hyphae identified from gut 18 contents of some species of Ptilodactylidae (fungal identifications by R. L. Seymour)..Species. Epjptygma rufa (Champion) Stirophora sulcipennis Champion Ptilodactvla sp, Aploalossa sp. Lachnodactvla monticola Champion Epiptvqma sp. Epiptvqma maculata (Champion) Epiptvqma rufipennis (Pic) Daemon sp. Contents Pteroconium sp. Botrvoderma sp. mixed unknown hyphomycete and coelomycete spores mixed unknown hyphomycete and coelomycete spores Alternaria sp. Capnodendron sp. coelomycetespores Pestalotia sp. mixed unknown hyphomycete and coelomycete spores Coniosporium sp. Pestalotia sp. Alternaria sp. Pestalotia sp, Pestalotia sp. Scleroqraphium sp. possibly, Hvcoenterolobium sp.

39 CHAPTER IV CHARACTERS, POLARITY, PHYLOGENETIC HYPOTHESIS, AND CLASSIFICATION Honophvlv of the Ptilodactylidae The lineage of the Dryopoidea consisting of the Callirhipidae, Eulichadidae, Ptilodactylidae, and Chelonariidae has been hypothesized as monophyletic (Lawrence and Newton, 1982). Crowson (1978) indicated the probable sister-group relationship of the Eulichadidae to the Ptilodactylidae (Crowson, 1978). The Eulichadidae, as well as the Callirhipidae, possesses a large onychium (Figs. 8, 9); in the Ptilodactylidae this structure is absent or very highly reduced (Figs ). I hypothesize this as evidence in support of monophyly of the Ptilodactylidae. Character discussion Suss and Puppin (1976) presented an investigation of the adult and larval anatomy of Ptilodactvla exotica Chapin, Therefore, I will discuss and illustrate only those adult integumental features used in phylogenetic inference and descriptions. The head and thorax have provided the majority of characters used in this study. Abdominal characters are limited to the form of male visible abdominal sterna V and 19

40 20 VI (including tergum of the latter), the ovipositor, and the aedeagus. Head. The 11-articled antennae are often sexually dimorphic with the females usually more conservative. With the exception of some female Cladotoma. ramous antennae are found only in males and have the mesally-directed rami either articulated or nonarticulated. The articulated rami (Figs. 2, 7) occur from antennomeres IV-X and have a narrow membranous base. The rigid nonarticulated rami (Figs. 1, 5, 6) occur on antennomeres III-X and arise proximally, medially, or distally on the antennomeres. This type of ramus is found in Cladotoma and is very broadly dorsoventrally flattened (Fig. 6). In some female Cladotoma, rami are present but are much shorter and narrower than in conspecific males. Serrate antennae vary from extremely weak (Fig. 3) to very strong serrations (Fig. 4). Almost all females possess this type of antennal structure and show the entire range of serrations. Serrate antennae in males are usually more strongly serrate than in conspecific females. The antennomeres are mesally serrate from III-IX. The epistomal sulcus extends across the frons below and approximate to the antennal insertions. It is sometimes deeply impressed with the frons even with or raised above the clypeal level. The sulcus may be weakly impressed with

41 the frons and clypeus equal in level; it is also 21 occasionally absent. Head shape is usually approximately quadrate (Pigs. 51, 52, 54). Octoqlossa. Ectvphodactvla NEW GENUS, and Araeopidius depart from this configuration with a more or less elongate head capsule (Figs. 49, 50, 53). The maxillary variation is large, with the major differences occurring in the galea and lacinia. In many genera, particularly within the Ptilodactylinae, the lacinia has a very well-developed spore brush similar to that described by Ashe (1984) for the subtribe Gyrophaenina (Staphylinidae: Aleocharinae) in which the lacinial apex forms a discoid structure densely beset with short, stout spines (Figs. 27, 31). The spines and setae of the lacinia are sometimes thinner, more elongate, less dense, and not in a discoid structure or arrangement (Figs. 26, 28-30, 33-36). I restrict my definition of spore brush to exclude these types. In Daemon and Stenafricus NEW GENUS, the galea is closely associated with the spore brush, having similar short, very stout spines (Fig. 32). Likewise, the galea of Ptilodactylinae is closely associated with the spore brush disc but possess longer and finer spines or setae (Fig. 27, 34). In this study, the galeae of Daemon and Stenafricus are considered to be part of the spore brush.

42 In other forms, the laclniae may be apically blunt 22 (Fig. 36) or acute, almost always heavily setose, with the galeae exhibiting the greatest variation, being either biacuminate or multiacuminate. Eoilichas and Anchvcteis are described as having biacuminate galeae (Figs. 28, 29); however, the mesad furcation of Anchvcteis is underdeveloped in relation to the laterad (Fig. 29). Multiacuminate galeae are seen in most Cladotominae and possess three or more furcations, each with a number of short, apparently random subbranches (Fig. 26). Mandibular variation includes 1-3 denticles, presence or absence of molar areas, dorsal ridge, and prostheca. In Eoilichas (Fig. 43), the thin dorsomedian portion of the mandibles often possesses an emargination which I do not consider to be homologous to true multidenticulations seen, e. g., in Aploglossa (Fig. 37) and Daemon (Fig. 38). Distinct molar areas (Figs. 37, 38, 41, 42) are often present with that of the left mandible usually concave (molar socket) and the right, more or less convex. The prostheca is located mesally and distally to the molar areas; it is a lightly melanized lobe, with shorter or longer setae (Figs. 37, 38, 40-43). The labrum varies in shape, condition of anterior margin, and presence or absence of anterolateral tufts of setae. In the Cladotominae, the labrum is strongly

43 transverse (basal width approximately 3 1/2 times its greatest length. Fig. 46). All other forms have the basal width no more than approximately twice the median length (Figs. 44, 45, 47, 48). The anterior margin of the labrum is usually slightly rounded or straight; in Falsotherius and Pherocladus it is concave (Fig. 44). The ligula is variable in shape, being anteriorly simple, bi-, guadri-, hex-, or multiacuminate. These conditions consist of each half of the ligula forming one, two, three or more apices which are variable in their acuteness (Figs , 24, 25). In the multiacuminate condition of Paralichas, each half has four or five branches similar in form to the multiacuminate galea, each branch with a number of apparently random subbranches (Fig. 24). In some forms the ligula is simple and is more or less ventrally and transversely folded (Fig. 23). Thorax. The lateral margins of the prothorax are either obsolete or acute. The acute condition is characterized by a smooth carina-like ridge which begins at the posterior margin and extends variable distances anteriorly (Figs. 51, 52, 54, 61, 64, 67, 68). In incompletely acute margins (Figs. 52, 54, 61, 67, 68), the ridge extends anywhere from 1/2 to nearly the entire lateral prothoracic length, the anterior end usually curving somewhat ventrally. Completely acute margins are

44 continuous to the anterior edge (Figs. 51, 64). Lateral 24 margins are either in the same plane as that of the anterior margin (termed, equiplanar. Fig. 67), or approximately mediolateral on the prothorax. Some forms possessing incompletely acute margins appear to be equiplanar, e. g. Stirophora. some Pherocladus. and Ptiloderes NEW GENUS (Fig. 52); however, the anterior end of the lateral ridge curves ventrally, or, in the case of Pherocladus is actually dorsal to the anterior margin (Fig. 61), and are not considered equiplanar. Genera possessing equiplanar margins are Daemon and Chelonariomorohus, the former with the lateral and anterior margins continuous and the latter, continuous or very nearly so (Fig. 67). In genera possessing obsolete lateral margination there is no smooth carina. Occasionally, e.g. in Bradvtoma. Aploglossa. and some Anchvtarsus. there may appear to be an acute lateroposterior margin; however, when closely examined, it is seen that the ridge is not smooth and is thus considered obsolete. The pronotum is anteriorly produced dorsal to the anterior margin in Pherocladus (Fig. 67); Ectvphodactvla has the pronotum smoothly expanded dorsally, laterally, and anteriorly (Fig. 66), the head in repose nearly flush with the margins of anterior prothoracic opening. The posterior pronotal margin is bisinuate and usually

45 more or less crenulate (Pigs. 69, 71, 72), Chelonariomor- 25 phus and Ptiloderes are exceptions to this with a smooth basal margin (Figs. 70, 73-76). Of those genera possessing crenulate basal margins, Aploglossa, Pseudoepilichas, Chaetodactvla, and some Bradvtoma and Anchvtarsus have these crenulations very weakly formed. The mediobasal lobe of the hind margin may be smooth or crenulate and usually has three produced denticles (Figs. 69, 73-76). Even with denticles, the mediobasal lobe is considered smooth when there is lack of one or more interdenticular crenulations. The mediobasal lobe may also be crenulate without produced denticles; this condition is rare and is exhibited only by Pseudocladotoma and some Paralichas (Fig. 72). The notai projection is always present, usually long, ventrally acute or truncate (Figs. 61, 62, 64, 66, 67, 68). In some genera this projection is shorter and usually not acute (Figs. 63, 65). Often present is a posterior and lateral expansion of the prosternum which has been termed 'cowling' (Hlavac, 1975). The cowling often meets the anterior edge of the notai projection causing concealment of the protrochantin. Both conditions, the visibility (Fig ) and concealment (Fig. 61, 66-68) of the protrochantin, are widespread among the genera. In Falsotherius, Ectvphodactvla, and Pherocladus the meeting of the cowling with the notai projection is loose (Pigs.

46 61, 66). The anterior margin of the mesoscutellum is associated with the mediobasal lobe of the pronotum in what has been called an interlocking mechanism or system (Hlavac, 1975; Lawrence, 1982, and personal communication). It takes three forms: smooth (Fig. 70), crenulate (Fig. 72), and cordiform (Pig. 69, 71, 73). The interlocking mechanism is best developed in forms having cordiform mesoscutella in which the median denticle of the basal lobe corresponds with the scutellar notch. Elytra may be strongly or obsoletely striate; either form may possess deeply impressed sutural striae which are usually most evident in the basal one-fifth or one-quarter bounding the mesoscutellum (e. g., Fig. 69). The prosternai process extends posteriorly between the mesocoxae and is usually received in a groove formed by the mesad ridges of the mesocoxal cavities and is not a true excavation. Sometimes a mesosternal excavation is present, it is a depression of the mesosternum into which the prosternai process fits, and may be shallow or relatively deep. A mesosternal keel is best developed in Stirophora and Chelonariomorphus. It is consistently present but weakly developed in Epiptvqma; rarely, a minor keel is present in Ptilodactvla. The metasternal transverse suture is visible

47 just anterior to the metacoxae, usually most distinct on 27 either side of the discrimen; it less distinct laterally. In some cases, e. g. Chelonariomorphus, Aploglossa. and Epiptvqma, the suture is not visible. The metacoxal plates form the ventral wall of the coxal grooves which serve for reception of the metafemora, and may be very weakly or relatively well-developed. The tarsi provide characters involving tarsomere IV, the ventral surface of the tarsomeres, and the ungual form. Tarsomere IV is often reduced, sometimes being inconspicuous. When not reduced, IV may be simple and approximately the same size as III (Fig. 13, 14), or it may have a ventral lobe. Frequently, when tarsomere IV is reduced. III has an expanded ventral lobe (Fig. 12). All ventral tarsomere lobes in Ptilodactylidae are entire and not bilobed as in some Dascillidae and Rhipiceridae. Tarsi are termed simple if tarsomere IV is not reduced or lobed. Occasionally, as in Epilichas, Anchvcteis, and Therius, IV may be lobed. In these and Falsotherius, the remaining tarsomeres have ventral fleshy pads which occasionally are produced into weak lobes. In Pseudocladotoma there is no reduction in IV and II - III have short, narrow lobes (Fig. 14). Ongues occur in three forms: simple (Fig ), bifid (Fig. 11), and pectinate (Fig. 10). Bifid ungues have two

48 28 acute points (Fig. 11) or the inner furcation shortened and more or less truncate. In the pectinate ungues of Paralichas (Fig. 10) each unguis has a mesally-directed row of fine teeth similar to those of alleculines (Tenebrionidae). Abdomen. The apex of male visible abdominal sternum V is variable in its shape and may be bisinuate, emarginate or simple. In male Epiptvqma NEW GENUS, visible tergum and sternum VI are sometimes very heavily sclerotized with variously shaped and ventrally-directed tergal flanges (Figs ). This segment is very rarely extruded and is usually visible in caudal view. The aedeagus is of the trilobed type and exhibits much variation. Consistent variation is found in the presence of subapical accessory projections of the median lobe (Figs. 94). Parameres are movable and may have apical or subapical nonmelanized projections (Figs. 85, 86, 90, 92), and in some cases, e. g. Aploglossa, are very short in relation to the basal piece (Fig. 87, 88). The ovipositor is usually very long with the bacula + valvulae 3 to 4x the length of the coxites. In some genera the ovipositor is shortened, the length of the bacula + valvulae less than twice that of the coxites (Figs. 77, 78). This condition is most pronounced in Araeopidius (Fig. 78). The coxites may (Figs. 77, 78, 81) or may not

49 (Fig. 19, 80, 82-84) be divided into proximal and distal 29 sections; styli may be present or absent, and, when present, apical (Fig. 78) or subapical (Figs. 77, 81). In the Cladotominae, the coxites are completely or almost completely fused with the bacula and valvulae (Fig. 79, 83, 84). The coxites in Cladotoma are apparently fused with heavily sclerotized paraprocts, forming a very rigid structure (Fig. 83, 84). Character polarization In this section, the character numbers, including those in parentheses, refer to Table 2. Char Antennal rami. Articulated rami are postulated to be apomorphic due to unique occurrence within the Ptilodactylidae; antennal rami of the Callirhipidae (as well as all others known to me) are nonarticulated. Char Epistomal sulcus. Reference to Eulichadidae and Callirhipidae showing absence of this sulcus indicates probable apomorphy in its presence. Char. 3, - Head shape. Occurrence of the basically quadrate condition in the examined outgroups requires postulation of its plesiomorphy; thus, elongation is considered apomorphic. Chars Mandibles. Absence of prostheca (4) and molar areas (5) are coded as apomorphic as a result of

50 presence of the former in Eulichadidae and Callirhipidae 30 and the latter in the Eulichadidae. Char Ligula. In Eulichadidae, the ligula is simple; thus, modification of this structure from the simple condition is coded as apomorphic. Chars. 6, Labrum. The short, transverse configuration of the labrum (6) is postulated as apomorphic with the alternative state occurring in Eulichadidae and Callirhipidae. Other labral modifications, anterior margin concave (26) and presence of anterolateral setal tufts (27) are likewise considered apomorphic due to their absence in the outgroup. Chars. 23, Maxilla. Presence of distinct lacinial (23) and galeal (24) spore brushes is hypothesized as apomorphic with the absence of these structures in the outgroup. Coded as separate apomorphies are the biacumination of the galea (31) and the reduction of the mesad branch of this condition (33). A further modification is the apomorphic polyacuminate galea (32). The apical palpomere being elongate and largely membranous (48) is also considered apomorphic. None of these conditions are found in the Eulichadidae or Callirhipidae. Chars Pronotum, Presence of incompletely acute lateral margination (8) is considered apomorphic due to its absence in the Eulichadidae and Psephenidae; thus.

51 the obsolete and completely acute conditions are 31 hypothesized plesiomorphic. The equiplanar margins (9), shortened notai projections (29), concealment of the protrochantins (10), and the anterior pronotal production (10), not found in any component of the outgroup, are postulated apomorphic» Chars. 11, 28, Thoracic sterna. Presence of the metasternal transverse suture (11) in eulichadids and callirhipids indicates probable plesiomorphy; therefore, the alternative absence is postulated apomorphic. Reduction or absence of a mesosternal excavation (28) is considered apomorphic due to the broad and deep condition of that feature in the Eulichadidae. The mesosternal keel (34) is not found in the outgroup, thus, its presence is apomorphic. Chars , Tarsi. Eulichadidae and Callirhipidae have simple tarsi and ungues. Any modification is thus hypothesized as apomorphic (17, 20). The various tarsal modifications, tarsomere IV reduced (19) or Icbed (35), presence of ventral, fleshy pads on two or more tarsomeres (18), and ungues pectinate (21) and basal tooth/bifid (36), are considered apomorphic. Char Sixth visible abdominal segment, males. The unique occurrence of this heavily sclerotized abdominal segment with a ventrally directed tergal flange and its

52 absence in the outgroup indicates an autapomorphy for 32 Epiptyqma. Chars , Aedeagus. The subapical accessory projections (25) of the median lobe are postulated apomorphic with their absence in the outgroup; presence of nonmelanized parameral projections (12) and short parameres (30) are also considered apomorphic due to absence of the projections and presence of long parameres in Eulichadidae and Callirhipidae. Chars Ovipositor. In the Eulichadidae, the ovipositor is long, the coxites transversely divided and articulated with the bacula, and styli apical; therefore, a shortened ovipositor (16), coxites not divided (13) or articulated with the bacula (fused; 39), and subapical (15) or absent (14) styli are hypothesized apomorphic. Discussion of phvlooenetic analysis and resulting classification Previous classifications of Ptilodactylidae (possibly only implied) (Hlavac, 1975; Lawrence and Newton, 1982) have recognized two or three subfamilies ([Anchytarsinae + Ptilodactylinae], or [Araeopidiinae + Anchytarsinae + PtilodactylinaeJ). Crowson (1967) suggested that several subfamilies exist within the Ptilodactylidae. The

53 phylogenetic hypothesis presented (Table 3) supports his 33 statement. Basing groupings only on synapomorphy and by phyletic sequencing (Wiley, 1979), I propose recognition of six subfamilies (Table 4). The numbers in parentheses below refer to characters listed in Table 2 and the phylogenetic hypothesis (Table 3), The proposed Ptilodactylinae is similar to that formerly recognized and with the addition of Daemonini and Pseudoepilichini. The former is included on the basis of ungual modification (bifid) (20, 36) and the latter of ovipositor simplification (13, 14) and concealment of protrochantins (10). The Epilichinae are hypothesized monophyletic on the synapomorphic biacuminate galea (31), a character appearing twice in the proposed phylogeny, also exhibited by the basal Anchytarsinae. The apomorphic shortening of parameres (with a corresponding lengthening of the basal piece) (30) suggests monophyly of Aploglossinae; the Octoglossinae and Cladotominae are each hypothesized monophyletic on a number of synapomorphiies. Two unresolved multichotomies occurred in the analysis. Both consistently appeared in multiple runs of QTREE and could probably be resolved by addition of further characters, especially larval. Some genera were not included in the phylogenetic

54 analysis due to lack of information, i. e., only having 34 knowledge of one sex (in some cases, that of the type of the generic type species) or only a few specimens in poor condition. In constructing the key I have been able to use characters which I have never observed as sexually dimorphic. Therefore, I have placed those genera in the key and provided descriptions. I will discuss the probable subfamilial or tribal placement for two of them in the phylogenetic diagram, and thus, in the classification. The third Therius is placed incertae sedis due to lack of males and the poor condition of the examined material. Hovactvla possesses a transverse labrum (6), no molar areas (5) or prostheca (4), and a polyacuroinate galea (32), and is placed in the Cladotominae. pseudocladotoma possesses a transverse labrum (6), no molar areas (5) or prostheca (4), and coxites not articulated with bacula (39), and is also placed in the Cladotominae.

55 35 Table 2. Hypothesized polarities of characters used in phylogenetic analysis of the genera of Ptilodactylidae. Characters Rami Epistomal sulcus Head shape Prostheca Molar area Labral length States Plesiomorphic non-articulated absent quadrate present present long Apomorphic articulated present elongate absent absent short ant.'ly produced incompl. acute equiplanar concealed absent with not divided absent subapical Pronotum not ant.'ly produced Pronotum: lat, marg. not Acute margins not equiplanar Protrochantins visible Metasternal trans. suture present Parameres w/o non-mel. projs. Ovipositor: coxites divided Ovipositor: styli present Styli present apical Valvifers + baculae:length much greater than coxites less than 2x Tarsi simple not simple Tarsi: ventral aspect w/o fleshy pads w/pads Tarsomere IV not reduced reduced Ungues simple not simple Ungues: not simple not pectinate pectinate Abd. tergum VI (male) w/o vertical flange with flange Lacinia not forming spore brush Spore brush lacinia only lacinia + galea Aedeagus:median lobe w/o access, proj. w/access. proj. Labrum:ant. angles w/o setal tufts w/tufts Labrum not anteriorly concave Mesosternal excavation deep Notai projection long shallow short Parameres long short Galea Galea not biacuminate not multiacuminate biacuminate multiacuminate Galea:biacuminate Mesosternal keel mesai furc, well-dev. not absent present Tarsomere 4:not reduced not lobed lobed Ungues:not simple not bifid or toothed Ligula simple not simple Apical max. palpomere (male) not elong./memb. elong./memb. Coxite/baculum/valvifer fusion no yes

56 36 Table 3. Phylogenetic hypothesis resulting from QTREE analysis of characters in Table 2. Reversals are indicated by dots at the upper right hand side of the character numbers; characters with multiple apomorphic appearances are 2, 3, 4, 5, 7, 9, 11, 12, 13, 14, 15, 16, 18, 20, 23, 25, 27, 29, 31, 35, and 38. 'i r / d 30 / 23 /2 5 I/ " * " 3 8 " 37' J 37

57 37 Table 4. Proposed supraspecific classification of the Ptilodactylidae ranked by phyletic sequencing. PTILODACTYLIDAE Anchytarsinae ^ ^ Anchvtarsus Guerin-Meneville Cladotominae Cladotoma Westwood Paralichas White Pseudocladotoma Pic Hovactvla Pairmaire Octoglossinae ^ ^ Octoqlossa Guerin-Meneville Aploglossinae Bradvtoma Guerin-Meneville Aploqlossa Gulrin-Meneville Epilichinae Anchvcteis Horn Epilichas White Bvrrocrvptus Broun Araeopidius Cockerell Ptilodactylinae Daemonini Stenafricus NEW GENÜS Daemon de Laporte Pseudoepilichini Falsotherius Pic Pseudoepilichas Armstrong and Nakane Ectvphodactvla NEW GENUS Ptilodactylini Ptiloderes NEW GENUS Pherocladus Pairmaire Lachnodactvla Champion Ptilodactvla Illiger Stirophora Champion Chelonaricmorppus Pic Epiptvqma NEW GENUS

58 CHAPTER V TAXONOMY KEY TO WORLD GENERA OF PTILODACTYLIDAE (ADULTS) 1. Protrochantins exposed (Figs, 62-65) *. Protrochantins concealed (Figs. 61, ). c, Tarsi simple (Fig. 13) '. Tarsi not simple, tarsomere IV reduced (Fig. 12) or lobed, II and III lobed (Fig. 14), or 1 or more tarsomeres with ventral fleshy pads Labrum transverse (Fig. 46), basal width more than 3x greatest length '. Labrum not transverse (Figs. 44, 45, 47, 48), basal width approx. 2x greatest length Ungues pectinate (Fig. 10)....Paralichas 4*. Ungues simple (Figs )...Hovactvla 5. Mesoscutellum cordiform (Figs. 1-3, 69, 71, 73),. 6 5'. Mesoscutellum not cordiform, basally smooth (Fig. 70), or crenulate (Fig. 7 2 )...Anchvtarsus 6. Lateral prothoracic margination incompletely acute (Figs. 52, 54, 61, 67, 6 8 )..Bvrrocrvptus 6 *. Lateral prothoracic margination obsolete (Figs. 49, 50, 53, 62, 63, 65, 6 6 ).....Anchvcteis 7. Tarsomere IV reduced (Fig. 12) '. Tarsomere IV not reduced, usually lobed

59 8. Mesoscutellum basally smooth (Fig. 70) or crenulate 39 (Fig. 72) '. Mesoscutellum cordiform (Figs. 1-3, 69, 71, 73) Tarsomeres II and III ventrally lobed '. Tarsomere II not lobed (Figs. 12, 13) Mandibles bidentate; Oriental region pseudoepilichas 10'. Mandibles tridentate; Neotropical region bradvtoma 11. Labrum transverse (Pig. 46), lateral prothoracic margination complete (Figs. 51, 64) Cladotoma 11'. Labrum not transverse (Pigs. 44, 45, 47, 48), lateral margination obsolete (Figs. 49, 50, 53, 63, 65, 66).....Aploqlossa 12. Prothoracic margination equiplanar (Figs. 67), complete; head quadrate (as in Figs. 52, 54); Madagascar...Daemon 12'. Prothoracic margination not equiplanar, obsolete (Fig. 63); head usually elongate (Fig. 49); Neotropics....Octoqlossa 13. Epistomal sulcus absent (Figs. 49, 51, 53) '. Epistomal sulcus present (Figs. 50, 52, 54)., Tarsomere IV ventrally lobed... Epilichas 14'. Tarsomere IV not lobed (Fig. 14).. Pseudocladotoma

60 Lateral prothoracic margination obsolete (as in Figs. 63r 65).....Anchvcteis 15'. Lateral prothoracic margins acute Ungues simple (Figs ); southern Africa Therius 16'. Ungues with basal teeth (as in Fig. 11 with mesal furcations more or less truncate); central Africa....Stenafricus NEW GENUS 17. Mesoscutellum basally smooth (Fig. 70) or crenulate (Fig. 72) '. Mesoscutellum cordiform (Figs. 2, 3, 71, 73) Prothoracic margination equiplanar (Fig. 67); Neotropics.....Chelonariomorphus 18'. Prothoracic margination not equiplanar; Indonesia..... Falsotherius 19. Prothorax bulbous, lateral margination obsolete, cowling loosely meeting notai projection (Fig. 53, 6 6 )....Ectvphodactvla NEW GENUS 19'. Prothorax not bulbous, lateral margination incompletely acute, cowling tightly fitting against notai projection (Figs. 61, 6 8 ) With mesosternal keel '. Without mesosternal k e e l Dorsum usually glabrous, very shining, median mesoscutellary notch usually continuing as

61 distinct stride; males, tergum of visible 41 abdominal segment VI with vent rally-directed flange (Pigs )...Epiptvqma NEW GENUS 21*. Dorsum covered with dense, recumbant setae, or, elytra very strongly striate; males, tergal flange absent; aedeagus, median lobe with subapical accessory projections (Fig. 94}... Stirophora 22. LateropeSterior pronotal angles broadly explanate (Figs. 3, 52, 73-76); galeae not forming spore brush (Fig. 34).Ptiloderes NEW GENUS 22. Lateroposterior pronotal angles not broadly explanate (Pigs. 2, 54); galeae forming spore brush (Fig. 27, and as in Fig. 31) Pronotum with strong anterior production (Fig. 61); Indonesia, Philippines..Pherocladus 23. Pronotum without strong anterior production (Fig. 6 8 ); cosmopolitan Apical maxillary palpomere, males, enlarged, elongate, and largely membranous (Fig. 27), females, usually somewhat elongate and mostly sclerotized......lachnodactvla 24. Apical maxillary palpomere simple, securiform (as in Pig. 31)...Ptilodactvla

62 THE WORLD GENERA OF PTILODACTYLIDAE 42 Anchvtarsus Anchvtarsus Guerin-Meneville, 1843a: 194 (type species, by monotypy: Atopa bicolor Melsheimer, 1845: 221); Blanchard, 1845: 56; Guerin-Meneville, 1849: 1; Lacordaire, 1857: 264; LeConte, 1853: 229, 1861: 179, (1865): 50; Horn, 1880: 8 6, 1881: 87; LeConte and Horn, 1883: 170; Champion, 1897: 593; Blatchley, 1910: 689; Bertrand, 1935: 138, 1939: 307, 1956: 275, 1966: 143, 1972: 391; Arnett, 1968: 443; Spangler, 1966: 397, 1981: 208, 1982: 386, 1983: 161; Brown, 1972: 24; 1975; 149; Hlavac, 1975: 182; Doyen and Ulrich, 1978: 229; White, 1980: 94; Stribling, 1986: 232. Tetraalossa Champion, 1897: 593 (type species, by monotypy: T, palpalis Champion, 1897: 594); Bertrand, 1956: 278, 1972: 393; Brown, 1975: 150; Stribling, 1986: 220. I have recently redescribed this genus (Stribling, 1986) and will here give a synopsis of characters not pointed out in that paper as well as a diagnosis. Molar areas and dorsal ridge are present and well-developed. The apical maxillary palpomeres are simple and largely sclerotized. The pronotal mediobasal lobe and mesoscutellum are either crenulate or smooth (as in Figs.

63 70 or 72). Sutural striae are deeply impressed and more 43 evident than other striae near the mesoscutellum. Diagnostic combination. This genus may be recognized by the following combination of characters: simple tarsi (as in Fig. 13), protrochantins visible (as in Pig. 65), epistomal sulcus absent, lateral pronotal margination obsolete, male antennae serrate (as in Fig. 4), parameres with apical nonmelanized projections, and ovipositor with styli. MATERIAL EXAMINED: 104 specimens (AMNH, MNHP, BMNH, CASC, CNCI, EGRC, FMNH, ICCM, MSDC, MCZC, NMNH, NCSD, OSDC, SMSH, JBSC, DMCE, OMIS). Taxonomic list - Anchvtarsus *bicolor (Melsheimer), 1845 (Atopa) (MCZC); eastern North America *folliculipalpus Stribling, 1986 (CNCI); Central, S. America *instriatus Pic, 1931b (MNHP); Costa Rica palpalis (Champion), 1897a (Tetraalossa) (BMNH); Mex., Cent., S. Amer. substriatus Champion, 1897a (BMNH); Nicaragua

64 Cladotoma 44 Cladotoma Westwood, 1837: 254 (type species: Ç. ovalis Westwood 1837); Guerin-Meneville, 1843b: 194; 1843a: 1; Blanchard, 1845: 56; Lacordaire, 1857: 276; Champion, 1897a: 626; Stribling, Telon Champion, 1897a: 626 (type species: T. cucullatus Champion 1897a: 626); Stribling, 1986: 232. NEW SYNONYMY. These two genera are synonymized due to joint possession of apomorphic pseudotetramery (Fig. 12) within the Cladotominae and the coxites of the ovipositor nonarticulated with the bacula (Figs. 83, 84). Diagnostic combination. This genus may be recognized by the following combination of characters: lateral pronotal margination complete (as in Fig. 64), labrum transverse (Fig. 46), tarsi (Fig. 12) tarsomere IV reduced, III ventrally lobed, and simple ungues. Description. Body Size. Length mm, width at humeral angles mm. Head. Quadrate; antennae, males with nonarticulated, broad, dorsoventrally flattened rami, antennomeres III-X (Fig. 6 ), females usually strongly serrate, sometimes with much shorter rami formed as in males; epistomal sulcus

65 45 absent; labrum strongly transverse (Fig. 46); mandibles (as in Fig. 39) bidentate, without prostheca, molar area, or dorsal ridge; maxillae (as in Fig. 26), lacinia acute, galea polyacuminate; labium, ligula (Fig. 22) quadracuminate. Thorax. Prothorax (as in Fig. 64) somewhat anteriorly produced, with complete, acute mediolateral margination (Fig. 51, as in 64), strong basal crenulation, mediobasal lobe crenulate, with produced denticles, mesoscutellum crenulate (as in Fig. 72); notai projection very long, thin, trochantins visible (as in Fig. 64); mesosternal keel absent; elytra obsoletely striate, sutural striae not more deeply impressed; metasternal transverse suture visible near"discrimen, laterally obsolete; metacoxal plates welldeveloped; tarsi (Fig. 12), article IV reduced. III ventrally lobed, no fleshy pads, ungues simple. Abdomen. Male sternum V modified, sinuate; aedeagus (Fig. 93), median lobe without mediolateral flanges or subapical accessory projections, parameres long, without nonmelanized apical projections; ovipositor (Pigs. 83, 84) long, coxites not divided, not articulated with bacula, fused with heavily sclerotized paraprocts, styli absent. Species excluded: russula Pic (Cerophytidae), vittata Pic fpseudolichas: Dascillidae).

66 MATERIAL EXAMINED: 26 specimens (OSDC, EGRC, KJOC, CNCI, 46 MNHP, FMNH, INHS, BMNH). Taxonomic list - Cladotoma bruchii Pairmaire, 1904 (BANM); Argentina *maculicollis Pairmaire, 1904 (MNHP); Argentina *heptosa (NEW NAME FOR maculicollis Pic, 1911 [MNHP]); Brazil (Rio Verde) *marqinata Pic, 1933 (MNHP); Brazil ovalis Westwood, 1837 (HEMC); Brazil subvittata Guerin-Meneville, 1861; Brazil thoracica Guerin-Meneville, 1843a; French Guiana

67 Paralichas 4 7 Paralichas White, 1859; 287 (type species, by monotypy: P. querini White 1859: 287); Nakane, 1948: 5, 1956: 52. Eucteis Guerin-Meneville, 1861: 539 (type species, by monotypy: E. bimaculata Guérin-Meneville, 1861: 539); Pairmaire, 1886: 395; Lewis, 1895: 99. Paralvchus Pairmaire, 1886: 395 (unjustified emendation). Odontonvx Guérin-Meneville, 1843b: 194 (not Stephens, 1827 in Carabidae; type species, by monotypy: Atopa ornata Melsheimer, 1845: 220 (transferred by Guérin-Meneville (1843b) [ =Dasvtes trivittis Germar, 1824: 76, Lacordaire (1857: 267)1; Blanchard, 1845: 56; Guérin- Meneville, 1849: 1; LeConte, 1861: 179; Horn, 1880: 8 6 ; LeConte and Horn, 1883: 170; Blatchley, 1910; 689; Arnett, 1968: 443; Hlavac, 1975: 182; Stribling, 1986: 232. NEW SYNONYMY. This synonymy is based on joint possession of the apomorphic pectinate ungues (Pig. 10). Diagnostic combination. This genus may be recognized by the autapomorphic pectinate ungues. Description. Body Size. Length mm, width at humeral angles 3o5-5o0 mm. Head. Quadrate; antennae, males often with

68 nonarticulated rami, antennomeres III-X (Fig. 5), females, 48 sometimes males, serrate; epistomal sulcus absent; labrum (as in Fig. 46) strongly transverse, anteriorly truncate or slightly convex; mandibles (Fig. 39) bidentate, without prostheca, molar areas, or dorsal ridge; maxillae (Fig. 26), galea multiacuminate, apical palpomere mostly sclerotized and melanized; labium (Fig. 24), ligula multiacuminate. Thorax. Prothorax (Fig. 64) somewhat anteriorly produced with complete, acute mediolateral margination, strong basal crenulation, mediobasal lobe crenulate, with or without produced denticles (Fig. 72); mesoscutellum crenulate (Fig, 72); notai projection very long, thin, trochantins visible (Fig. 64); mesosternal keel absent; elytra strongly striate, sutural striae not more deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete; metacoxal plates welldeveloped; tarsi (as in Fig. 13) simple, without ventral fleshy pads; ungues pectinate (Fig. 10). Abdomen. Male sternum V simple or modified, when modified, sinuate; aedeagus (Fig. 91), median lobe without mediolateral flanges and subapical accessory projections, parameres long, without nonmelanized apical projections; ovipositor (Fig. 79) long, coxites not divided, not articulated with bacula; styli absent.

69 Species excluded: pjceiceps Pic (Pseudocladotoma\ 49 MATERIAL EXAMINED: 136 specimens (ICCM, RHTC, MCZC, RÜIC, OSDC, KOBC, MNHP, NMNH, INHS, CASC). There are two undescribed Neotropical species, one specimen each from Mexico and El Salvador. Taxonomic list - Paralichas *bicolorpes Pic, 1911 (MNHP); Taiwan *querini White, 1859 (BMNH); China Eucteis bimaculata Guerin-Meneville, 1861 Lewis, 1895 (synonymized with querini) V. V. V. *disconiqer Pic, 1939 (MNHP); China *diverselineatus Pic, 1939 (MNHP); China *iunctus Pic, 1939 (MNHP); China *qriseolineatus Pic, 1939 (MNHP); no locality *hiqoniae Lewis, 1895 (MNHP); Japan ssp. qvotokui Nakane, 1963; Japan niqer Pic, 1931a (MNHP); Madagascar *pectinata (Kiesenwetter), 1874 (Odontonvx); China, Japan rufolimbata (Pairmaire), 1886 (Eucteis); Kiang-si, China V. *apicornis Pic, 1939 (MNHP); China *striolatus Pairmaire, 1897 (MNHP); Sze-tchouen *subnitidus Pic, 1911 (MNHP); China *trivittus (Germar), 1824 (Dasvtes); eastern N. America Atopa ornata Melsheimer, 1845 Guerin-Meneville, 1849 (transferred ornata to Odontonvx)

70 Hovactvla Hovactvla Pairmaire, 1901: 174 (type species, by monotypy: H. dermestoides Pairmaire, 1901: 175). Diagnostic combination. This genus may be recognized by the following combination of characters: labrum transverse (as in Pig. 46) and tarsi and ungues simple (as in Pig. 13). Description. Body size. Length mm, width at humeral angles mm. Head. Quadrate; antennae serrate (as in Pig. 4); epistomal sulcus absent, labrum transverse (as in Pig. 46), basal width approx. 4x greatest length, anteriorly convex; mandibles (as in Pig. 39) bidentate without prostheca or molar areas, with dorsal ridge; maxillae, lacinia acute not forming spore brush, galea multiacuminate (as in Pig. 26); apical maxillary palpomere mostly sclerotized; labium, ligula quadracuminate. Thorax. Pronotum somewhat anteriorly produced, with mediolateral margins completely acute (as in Figs. 51, 64), strong basal crenulation, mediobasal lobe crenulate, with produced denticles (as in Pig. 72), notai projection long, protrochantins visible (as in Pig. 64); mesoscutellum basally crenulate (as in Pig. 72), mesosternal excavation

71 deep, keel absent; elytra strongly striate, sutural striae 51 not deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete, coxal plates underdeveloped; tarsi simple (as in Fig. 13), without fleshy ventral pads, ungues simple. Abdomen. Male sternum V slightly sinuate; aedeagus, median lobe without mediolateral spines or subapical accessory projections, parameres without nonmelanized projections. Females unknown. MATERIAL EXAMINED: 2 specimens (MNHP, CNCI). Specimens from the type locality and South Africa. The latter represents an undescribed species. Species excluded: rufescens Pic (incertae sedis). Taxonomic list - Hovactvla *dermestoides Pairmaire, 1901 (MNHP); Madagascar

72 52 Pseudocladotoma Pseudocladotoma Pic, 1918: 3 (type species, by monotypy: P. maculata Pic, 1918: 4). Diagnostic combination. This genus may be recognized by the following combination of characters: labrum transverse, complete lateral margination, serrate antennae, simple ungues and, tarsomere IV not reduced. Description. Body size. Length S mm, width at humeral angles mm. Head. Quadrate; epistomal suture absent; labrum transverse (as in Fig. 46), basal width approx. 3x greatest length, anteriorly convex; mandibles monodentate (as in Fig. 40), without prostheca or molar areas (as in Fig. 39), with dorsal ridge; maxillae, lacinia acute, not forming spore brush, galea blunt; apical palpomere mostly sclerotized; labium, ligula biacuminate (as in Fig. 18). Thorax. Lateral margins completely acute (as in Fig. 64), somewhat anteriorly produced; basally crenulate, mediobasal lobe crenulate without produced teeth; notai projection long, protrochantins visible (as in Fig. 64); mesosternal excavation, keel absent, mesoscutellum basally crenulate (as in Fig. 72); elytra strongly striate, sutural striae not deeply impressed; metasternal transverse suture

73 53 visible near discrimen, laterally obsolete; metacoxal plates underdeveloped; tarsi (Fig. 14} robust, article IV not reduced, II-III sometimes with very narrow ventral lobes, without ventral fleshy pads, ungues simple. Abdomen. Ovipositor long, coxites not divided, not articulated with bacula, styli absent. Males unknown. MATERIAL EXAMINED: 4 specimens (MNHP). Taxonomic List - Pseudocladotoma maculata Pic, 1918 (MNHP); Java piceiceps (Pic), 1930a (Paralichas) NEW COMBINATION (MNHP); China rufonotatus (Pic), 1931b (Epilichas) NEW COMBINATION (MNHP); Borneo vitaticollis (Pic), 1930a (Epilichas) NEW COMBINATION (MNHP); China

74 Octoqlossa 54 Octoqlossa Guerin-Meneville, 1843a: 1 (type species, by monotypy: Q» femoralis Guerin-Meneville, 1843a: 2); Guerin-Meneville, 1843b: 194; Lacordaire, 1857: 268; Stribling, 1986: 232. Astvloqlossa Pic, 1919: 6 (type species, by monotypy: A. coerulipennis Pic, 1919: 7); Stribling, NEW SYNONYMY. These two genera are synonymized due to the synapomorphic elongation of the head, lack of molar areas, ovipositors with subapical styli and, shortened notai projections. Diagnostic combination. This genus may be recognized by the following combination of characters: lateral pronotal margination obsolete, head elongate, trochantins visible (Figs. 49, 63), tarsomere IV reduced (as in Fig. 12 but somewhat distally expanded), II-III lobed ventrally, mesoscutellum cordiform. These species often have dorsal elytral patterns of metallic blue and green. Description. Body size. Length mm, width at humeral angles mm. Head. Elongate (Fig. 49; less so in 0. rufa): antennae serrate (as in Fig. 4), epistomal sulcus absent, labrum

75 long, basal width less than 2x greatest length, anteriorly convex; mandibles (Fig. 40) long, monodentate, with prostheca, without dorsal ridge or molar areas; maxillae, galea acute, sometimes multiacuminate, lacinia acute, not forming spore brush; apical palpomere mostly sclerotized; labium, ligula hexacuminate (Fig. 25). Thorax. Prothorax with obsolete lateral margins (Fig. 63); basal margins strongly crenulate, mediobasal lobe crenulate, with produced denticles (as in Fig. 71); notai projection short, protrochantins visible (Fig. 63); mesoscutellum cordiform; mesosternal excavation shallow, keel absent; elytra strongly striate, sutural striae not deeply impressed; metasternal transverse suture visible near discrimen, laterlly obsolete, coxal plates underdeveloped; tarsi, article IV reduced (as in Fig. 12 but somewhat distally expanded), II-III lobed, without ventral fleshy pads, ungues simple. Abdomen. Male sternum V emarginate; aedeagus, median lobe without mediolateral flanges or subapical accessory projections; ovipositor (as in Fig. 77) short, bacula and valvulae less than 2 x coxites, coxites divided, articulated with bacula, styli subapical. MATERIAL EXAMINED: 20 specimens (MCZC, BMNH, ANSP, MNHP, NMNH, CNCI). Specimens were examined from Panama, Costa Rica,

76 Colombia, Ecuador, and Venezuela. Taxonomie list - Octoqlossa *coerulipennis Pic, 1919 (Astvloqlossa) NEW COMBINATION (MNHP); Colombia *cvaneipennis Pic, 1924a (MNHP); Costa Rica femoralis Guérin-Meneville, 1843b; Panama, Colombia *rufa Pic, 1924b (MNHP); Colombia

77 Bradvtoma Bradvtoma Guerin-Meneville, 1843a: 1 (type species: B. aurita Guérin-Meneville, 1843a: 1, by monotypy); Guerin-Meneville, 1843b: 194; Blanchard, 1845: 56; Lacordaire, 1857: 277; Stribling, 1986: 232. Brithvcera Erichson, 1847: 175 (type species: B. Ivciformis Erichson, 1847: 175, by monotypy), 1848: 102; Lacordaire, 1857: 276. Diagnostic combination. This genus may be recognized by the following combination of characters: tarsomere IV reduced, II-III ventrally lobed, without ventral fleshy pads, labrum not anteriorly concave, lateral pronotal margination obsolete, male antennae serrate (Fig. 4), mesoscutellum basally smooth (as in Fig. 70) or crenulate (as in Fig. 72), and ligula not simple, with variable acumination (Fig. 16). Description. Body size. Length mm, width at humeral angles mm. Head. Quadrate; antennae serrate (Fig. 4); epistomal sulcus present or absent; labrum long, basal width 2 x or less greatest length (as in Fig. 47), anteriorly truncate or slightly convex; mandibles tridentate, with prostheca,

78 molar area, and dorsal ridge (as in Fig. 37); maxillae 58 (Fig. 33), lacinia blunt, not forming spore brush, galea cylindrical, with apical cluster of setae, not closely associated with lacinia; apical palpomere mostly sclerotized; labium, ligula modified (Fig. 16), hexacuminate with varying degrees of reduction. Thorax. Prothoracic margins (Pig. 65) obsolete, occasionally appearing posteriorly acute but margins without anterior, ventral curvature or smooth carina; basally crenulate, mediobasal lobe crenulate or not, with or without produced denticles; notai projection long, trochantins visible; mesosternal keel, excavation absent; mesoscutellum basally smooth, elytra strongly striate, sutural striae deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete; metacoxal plates weakly developed; tarsi, article IV reduced, II-III lobed, without ventral fleshy pads; ungues simple (as in Fig. 12). Abdomen. Male sternum V simple or modified, when modified, sinuate; aedeagus (Figs. 85, 8 6 ), median lobe without mediolateral spines, parameres with nonmelanized projections, ovipositor (as in Fig. 81) long, coxites not divided, articulated with bacula, styli subapical. MATERIAL EXAMINED: 12 specimens (MNHP, CNCI, RSN^ ).

79 Taxonomic list - Bradvtoma 55 *aurita Guerin-Meneville, 1843; Brasil *lineata Pic, 1921 (MNHP); Brasil Ivciformis (Erichson), 1847 (Brithvcera); Peru

80 Aploalossa 60 Aploqlossa Guerin-Meneville, 1849; 1 (type species, here designated: A. sallei Guerin-Meneville, 1849: 2 [ =Aploqlossa marqinata Guerin-Meneville, 1849: 2]); Lacordaire, 1857: 276; Kasap and Crowson, 1975; 442; Stribling 1986: 232. Haploqlossa Gemminger and von Harold, 1869: 1623 (unjustified emendation); Champion, 1897: 623; Kasap and Crowson, 1975: 442. Diagnostic combination. This genus may be recognized by the following combination of characters: tarsi (as in Pig. 12), tarsomere IV reduced. III lobed ventrally, epistomal sulcus present, protrochantins visible, obsolete lateral margination (as in Fig. 65), male antennae serrate (as in Fig. 4), mesoscutellum basally smooth (as in Fig. 70), and simple ligula. Description. Body size. Length mm, width at humeral angles mm. Head. Quadrate; antennae (as in Fig. 4), serrate; epistomal sulcus present; labrum (Fig. 45), not transverse, anterior margin more or less concave, with tufts cf setae at anterolateral angles; mandibles tridentate, with prostheca, molar areas, and without dorsal ridge (Fig. 37);

81 maxillae (Fig. 31), lacinia forming spore brush, galea Cl closely associated with spore brush, but, with longer, thinner, and less dense setae, apical palpomere mostly sclerotized; labium (as in Fig. 23), ligula simple. Thorax. Prothorax (as in Fig. 65) with obsolete lateral margins, sometimes appearing posteriorly acute, but lacking smooth ridge, usually more or less angulate at posterior one-fifth; weak basal crenulation, mediobasal lobe crenulate, without produced denticles; mesoscutellum basally smooth (as in Fig. 70); notai projection short, trochantins visible (as in Fig. 65); mesosternal excavation shallow, keel absent; elytra obsoletely striate, sutural striae not deeply impressed; metasternal transverse suture absent; metacoxal plates underdeveloped; tarsi (as in Fig. 12), article IV reduced. III ventrally lobed, without ventral fleshy pads, ungues simple. Abdomen. Male visible sternum V modified, emarginate; aedeagus, median lobe without mediolateral flanges, sometimes with subapical accessory projections (as in Fig. 94), parameres short, without nonmelanized apical projections (Pigs. 87, 88); ovipositor (Fig. 77) short, valvifers less than 2x coxites, coxites articulated with bacula; divided, with subapical styli. MATERIAL EXAMINED: 145 specimens (NMNH, AMNH, CASC, BMNH, CNCI, MAIC, FMNH, MNHP, JBSC, UNTC).

82 Taxonomie list - Aploqlossa 69 *anqustata Champion, 1897a (BMNE); Panama *aureonotata Pic, 1913d (MNHP); Peru V. qermaini Pic, 1927b (DEIE); Bolivia *baeri Pic, 1913d (MNHP); Peru *boliviensis Pic, 1913d (MNHP); Bolivia *collaris Guérin-Meneville, 1849; Izabal, Guatemala Champion, 1897a (redescription) V. V. *reducta Pic, 1934e (MNHP); no locality *testaceicollis Pic, 1913d (MNHP); no locality *ferruqinea Champion, 1897a (BMNH); Guatemala *laiovei Pic, 1913d (MNHP); Venezuela sallei Guerin-Meneville, 1849; Caracas, Venezuela marqinata Guérin-Meneville, 1849 *scutellaris Pic, 1913d (MNHP); Neotropics V. V. *niqricollis Pic, 1913d (MNHP); Brazil *obscuricolor Pic, 1913d (MNHP); Amazon *suturalis Pic, 1934c (MNHP); Bolivia V. V, V. *diversipes Pic, 1934c (MNHP); no locality *humeralis Pic, 1934c (MNHP); no locality *notaticollis Pic, 1934c (MNHP); no locality *testaceipes Pic, 1934c (MNHP); Peru

83 Anchvcteis Anchvcteis Horn, 1880: 87 (type species, by monotypy: A. velutina Horn, 1880: 87); LeConte and Horn, 1883: 170; Leech and Chandler, 1956: 367; Arnett, 1968: 443; Hlavac, 1975: 182; Doyen and Ulrich, 1978: 229; Stribling, 1986: 232. Amphicteis Bertrand, (1966: 144, 1972: 392; incorrect subsequent spelling). Diagnostic combination. This genus may be recognized by the following combination of characters: male antennae with nonarticulated rami (Pig. 1), epistomal sulcus present but shallow, lateral pronotal margins obsolete, protrochantins visible (as in Fig. 65), tarsomeres I-IV with ventral fleshy pads, IV sometimes lobed, and molar areas present (Fig. 42). Description. Body size. Length mm, width at humeral angles mm. Head. Quadrate; antennae, males with non-articulated rami, antennomeres III-X (Figs. 1, 5), females serrate (as in Fig. 4); epistomal sulcus present, shallow; labrum (Fig. 48), not transverse, anteriorly convex; mandibles (Fig. 42) monodentate, with prostheca and molar areas, with or without dorsal ridge; maxillae (Fig, 29), lacinia blunt.

84 not forming spore brush, galea apically acute, biacuminate 64 with mesal branch highly reduced or absent; apical palpomere mostly sclerotized and melanized, not exceptionally flattened or expanded; labium (Fig. 19), ligula quadracuminate with lateral branches highly reduced. Thorax. Prothorax (as in Fig. 65) with lateral margination obsolete, basal margin strongly crenulate, mediobasal lobe crenulate, with produced denticles (as in Fig. 71); mesoscutellum cordiform; notai projection long, trochantins visible (as in Fig. 65); mesosternal keel absent, excavation present, shallow; elytra strongly striate, sutural striae not more deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete; metacoxal plates well-developed; tarsomere IV not reduced, I-IV with ventral fleshy pads sometimes being drawn into small lobes; ungues simple. Abdomen. Male visible sternum V modified, sinuate; aedeagus with mediolateral flanges, without subapical accessory projections; parameres long, without nonmelanized apical projections; ovipositor long, coxites articulated with bacula, divided or not, styli present or absent, when present, subapical. MATERIAL EXAMINED: 56 specimens (UCDC, FMNH, OSUO, MCZC, OSUC, UMCE, MNHP, BMNE, KÜBC, CASC, NMNH, CNCI, G L PC).

85 Taxonomie list - Anchvcteis. 65 *brunneicornis (Lewis), 1895 (Epilichas) NEW COMBINATION (BMNH); Japan ssp. usori (Nakane), 1958; Japan *monticola (Nakane), 1952 (Epilichas) NEW COMBINATION; Japan *velutina Horn, 1880 (MCZC); northwestern ü. S. A,

86 Epilichas Epilichas White, 1859: 290 (type species, by monotypy: E. candezei White, 1859; 289); Nakane, 1948: 5, 1956: 52; Crowson, 1967: 50. Diagnostic combination. This genus may be recognized by the following combination of characters: male antennae with nonarticulated rami (as in Figs. 1, 5), epistomal sulcus absent, lateral margination obsolete, protrochantin visible (as in Fig. 65), tarsomere IV lobed, I-III with ventral fleshy pads, and molar areas absent (Fig. 43). Description. Body size. Length ram, width at humeral angles mm. Head. Quadrate; antennae, males with non-articulated rami on antennomeres III-X (as in Figs. 1, 5), females serrate (as in Fig. 4); epistomal sulcus absent; labrum (as in Fig. 48), not transverse, anteriorly truncate or convex; mandibles (Fig. 43) monodentate, with prostheca, without molar areas, with or without dorsal ridge; maxillae (Fig. 28), lacinia blunt, not forming spore brush, galea apically acute, biacuminate; apical palpomere mostly sclerotized, usually flattened and laterally expanded; labium (Fig. 21), ligula quadracuminate. Thorax. Prothorax (as in Fig. 63) with lateral

87 margination obsolete, basal margin strongly crenulate, mediobasal lobe crenulate, with produced teeth (as in Fig. 71); mesoscutellum cordiform; notai projection long, trochantins visible (as in Fig, 64); mesosternal keel absent, excavation shallow; elytra obsoletely striate, sutural striae not deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete; metacoxal plates well-developed; tarsi, article IV lobed, articles I-III with ventral fleshy pads occasionally being drawn into small lobes; ungues simple. Abdomen. Male visible sternum V modified, sinuate; aedeagus (Fig. 95), median lobe without mediolateral spines or subapical accessory projections; parameres, long, without apical flange or nonmelanized apical projections; ovipositor, long, coxites articulated with bacula, divided, with apical styli (as in Fig. 78 but bacula 4-5x coxital length). Species excluded: brunneicornis Lewis (Anchvcteis), monticola Nakane (Anchvcteis), rufonotatus Pic (Pseudocladotoma), varieqatus Carter (Bvrrocrvptus), vittaticollis Pic (Pseudocladotoma), apicicornis Pic (Falsotherius). MATERIAL EXAMINED: 46 specimens (MNHP, BMNH, NMNH, CNCI, KUBC).

88 Taxonomie list - Epilichas 68 *atricolor Lewis, 1895 (BMNE); Japan basipes Pic, 1927d (MNHP); Vietnam (Tonkin) *candezei White, 1859 (BMNH); China *flabellatus (Kiesenwetter), 1874 (Octoqlossa)r Japan *testaceipes Pic, 1927a (MNHP) Nakane, 1952 (synonymized with flabellatus) V. amamianus Nakane, 1963; Japan ssp. mutsuensis Nakane, 1958; Japan ssp. vakushimensis Nakane, 1952; Japan f, oshiïïianus Nakane, 1952; Japan V. rubricollis Nakane, 1952; Japan ssp. tamaii Sato, 1964; Iriomote-jima, Sakishima mivatakei Nakane, 1952; Japan niqrinis Carter, 1936; Australia *niaripes Pic, 1914b; Vietnam (Tonkin) V. testaceicollis Pic, 1914b; Vietnam oblonqus (Carter), 1935 (Dascillus) (ANMM); Australia obscurus (Pic), 1923a (Eulichas): Vietnam (Tonkin) Deleve (redescription) rufescens Pic, 1927c; Vietnam (Tonkin) serraticornis Carter, 1935; Australia

89 Bvrrocrvptus Bvrrocrvptus Broun, 1893: 1136 (type species, by monotypy: Bvrrocrvptus urouharti Broun, 1893: 1137); Kasap and Crowson, 1975: 442. Diagnostic combination. This genus may be recognized by the following combination of characters: antennae serrate (as in Fig. 4), epistomal sulcus present, lateral pronotal margination incompletely acute, protrochantins visible (as in Pig. 65), mandibles bidentate (as in Fig. 41), and simple tarsi (as in Fig. 13). Description. Body size. Length mm, width at humeral angles mm. Head. Quadrate; antennae serrate (as in Fig. 4); epistomal sulcus present; labrum (as in Fig. 47), basal width approx. 2x greatest length, anterior margin truncate or slightly'convex; mandibles (as in Fig. 41) bidentate, prostheca, molar areas, and dorsal ridge present; maxillae, lacinia blunt, not forming spore brush, galea (Fig. 30) biacuminate, mesal branch sometimes highly reduced, apical palpomere mostly sclerotized and melanized; labium (as in Fig. 20), ligula quadracuminate. Thorax. Prothorax with acute lateral margination, incomplete; basal margin strongly crenulate, mediobasal

90 lobe crenulate, with produced denticles (as in Fig. 71), 70 notai projection long, trochantins visible (as in Fig. 64); mesoscutellum cordiform; sternal excavation shallow, keel absent; elytra strongly striate, pronotum and elytra sometimes with broad, irregular punctation, sutural striae usually deeply impressed basally; metasternal transverse suture visible near discrimen, laterally obsolete; metacoxal plates well-developed; tarsi (as in Fig. 13) simple without fleshy pads, ungues simple. Abdomen. Male sternum V simple or modified, when modified, sinuate; aedeagus, median lobe without mediolateral flanges or subapical accessory projections; parameres long, wtihout nonmelanized apical projections; ovipositor, long, articulated with bacula, coxites divided, with apical styli (as in Fig. 78 with length of bacula 4-5x that of coxites). MATERIAL EXAMINED: 136 specimens (CSIR, CNCI, BMNH, JBSC). Taxonomic list - Bvrrocrvptus *urauharti Broun, 1893 (BMNH); New Zealand *varieaatus (Carter), 1935 NEW COMBINATION (Epilichas) (ANMM); N. Queensland There are about 6 species with additional records from Papua, New Guinea and New South Wales. It is probable that

91 71 the other species of Epilichas described by Carter (1935) belong here; however, examination of the types is necessary.

92 Araeopidius 72 Araeopus LeConte, 1874: 56, not Spinola (1839) in Hemiptera; Horn, 1880: 88; LeConte and Horn, 1883: 170. Araeopidius Cockerell, 1906: 241 (type species: by monotypy: Araeopus monachus LeConte, 1874: 56); Hlavac, 1975: 182; Lawrence and Newton, 1982: 278; Stribling, 1986: 232. Diagnostic combination. This genus may be recognized by the following combination of characters; head somewhat elongate (Fig. 50), lateral pronotal margins obsolete, protrochantins visible (Fig. 62), mesoscutellum cordiform (Fig. 69), tarsi simple (Fig. 13), and sutural striae deeply impressed (Fig. 69). Description. Body size. Length mm, width at humeral angles mm. Head. Elongate (Fig. 50); antennae serrate (Fig. 4); epistomal sulcus shallow, labrum long, anteriorly truncate or slightly concave; mandibles (as in Pig. 42) monodentate, with prostheca, molar areas, and dorsal ridge; maxillae, lacinia acute (as in Fig. 29), not forming spore brush, galea biacuminate (as in Fig. 28); apical palpomere mostly sclerotized, somewhat elongate; labium, ligula quadracuminate (as in Fig. 21).

93 Thorax. Pronotum with obsolete lateral margins (Fig )/ strong basal crenulation, mediobasal lobe crenulate, with produced denticles (Fig. 69); notai projection long, protrochantins visible (Fig. 62); mesoscutellum cordiform, mesosternal excavation deep, keel absent, elytra obsoletely striate, sutural striae deeply impressed (Fig. 69); metasternal transverse suture visible near discrimen, laterally obsolete, coxal plates well-developed; tarsi (Fig. 13) simple, with reduced onychium, without ventral fleshy pads, ungues simple. Abdomen. Male sternum V simple; aedeagus, median lobe without mediolateral flanges and subapical accessory projections, parameres long, without nonmelanized apical projections; ovipositor (Fig. 78) short, bacula and valvulae slightly longer than coxites, coxites articulated with bacula, divided, styli apical. MATERIAL EXAMINED: 175 specimens (GLPC, OSUO, COBC, INKS, FMNH, MCZC, CASC, OSUC). Taxonomic list - Araeopidius monachus (LeConte), 1874 (Araeopus); northwestern North America

94 7 4 Stenafricus NEW GENUS Stenafricus NEW GENUS (type species, present designation: Stenactvla auberti Pic, 1925b: 14). Stenactvla Fairmaire, 1904: 346, in part. Diagnostic combination. This genus may be recognized by the following combination of characters: lateral pronotal margins acute, nearly complete, notai projection short (as in Fig. 63), protrochantins visible (as in Fig. 63), tarsomere IV lobed, no ventral fleshy pads, ungues with basal tooth, and lacinia and galea forming spore brush (Fig. 32). Description. Body size. Length mm, width at humeral angles mm. ' Head. Quadrate; antennae serrate (as in Fig. 4); epistomal sulcus present; labrum (as in Fig. 47) long, basal width 2x or less greatest length, anteriorly truncate or slightly convex; mandibles tridentate, with prostheca and molar areas (as in Fig. 37), without dorsal ridge; maxillae (Fig. 32), galea and lacinia forming spore brush, apical palpomere mostly sclerotized; labium, ligula simple, often somewhat ventrally, transversely (as in Fig. 23 though not as distinctly) folded. Thorax. Prothorax with lateral margins acute, usually

95 -s nearly complete, occasionally complete, margination more or less mediolateral, not equiplanar; basal margin crenulate, mediobasal lobe crenulate, with produced teeth (as in Fig. 72), notai projection short, protrochantins visible (as in Fig. 63); mesoscutellum cordiform or crenulate; mesosternal excavation, keel absent; elytra obsoletely striate, sutural striae not deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete, metacoxal plates well-developed; tarsi, article IV lobed, without ventral fleshy pads, ungues with basal tooth. Abdomen. Male sternum V more or less emarginate; median lobe with mediolateral flanges; parameres long, with nonmelanized subapical projections (Fig. 92); ovipositor (as in Fig. 81} long, coxites divided, articulated with bacula, styli subapical. MATERIAL EXAMINED: 37 specimens (MNHP, MRAC). Seven or eight species from the type localities, Nigeria, Angola, and Zaire. Further type studies are necessary to determine how many are described. Taxonomic list - Stenafricus *auberti (Pic), 1925b (Stenactvla) NEW COMBINATION (MNHP); Gabon *stranqulata (Pic), 1928a (Stenactvla) NEW

96 76 COMBINATION (MNHP); French Congo *nitidissimus (Pic), 1914a (Therius) NEW COMBINATION (MNHP); west central Africa

97 D a e m o n 7 7 Daemon de Laporte, 1836; 24 [type species, by monotypy: Ptilodactvla gioas de Laporte, 1836: 23 (= Colobodera ovatus Klug, 1838: 59)]; Lacordaire, 1857: 278; Kasap and Crowson, 1975: 442, Colobodera Klug, 1838: 67 (type species, present designation: Ç. ovatus Klug, 1838: 67); Lacordaire, 1857: 278; Fairmaire, 1896; 341. Diagnostic combination. This genus may be recognized by the equiplanar prothoracic margination (as in Fig. 67 with lateral margins continuous with anterior) and visible protrochantins (as in Fig. 63). Description. Body size. Length mm, width at humeral angles mm. Head. Quadrate; antennae very weakly serrate, not sexually dimorphic; epistomal sulcus present; labrum (as in Fig. 47), not transverse, basal width approx. 2x basal width, anteriorly truncate or slightly convex; mandibles (Fig. 38) bidentate, with prostheca, molar areas, and dorsal ridge, molar socket exaggerated; maxillae (as in Fig. 32), lacinia forming spore brush, galea closely associated and integrated with spore brush, possessing similar spines; apical palpomere mostly sclerotized and

98 melanized; labium (as in Pig, 23), ligula simple, often 78 somewhat ventrally, transversely folded. Thorax, Prothorax with acute lateral margins, complete, equiplanar (as in Fig, 67), and continuous with anterior margin; strong basal crenulation, mediobasal lobe crenulate, with three produced denticles; mesoscutellum cordiform (as in Fig, 71); notai projection long, trochantins visible (as in Pig, 64); mesosternal excavation and keel absent; elytra obsoletely striate, sutural striae not deeply depressed; metasternal transverse sutrue visible near discrimen, laterally obsolete; metacoxal plates present, underdeveloped; tarsi (as in Fig, 12), article IV reduced. III ventrally lobed, without ventral fleshy pads, ungues simple. Abdomen, Male visible sternum V modified, emarginate; aedeagus, median lobe without mediolateral flanges, with or without subapical accessory projections; parameres without nonmelanized projections, often with long apical setae; ovipositor (Fig, 81), long, coxites divided, articulated with bacula, with subapicla styli. Species excluded: subiunctus Pic (Ptilodactvla), MATERIAL EXAMINED: 302 specimens (MNHP, MRAC, IZWP, NMNH),

99 Taxonomie list - Daemon 79 aegualis Fairmaire, 1896 (MNHP); Madagascar *apicalis Pic, 1924b (MNHP); no locality V. donckieri Pic, 1924b atopoides Fairmaire, 1896 (MNHP); Madagascar V. *notaticollis Pic, 1913b (MNHP); Madagascar *atricornis Pic, 1924b (MNHP); Madagascar *basalis Fairmaire, 1896 (MNHP); Madagascar *berqrothi Pic, 1924b (MNHP); Madagascar cebrionoides Fairmaire, 1896 (MNHP); Madagascar colossus Fairmaire, 1896 (MNHP); Madagascar distinquendus Fairmaire, 1896 (MNHP); Madagascar *diversipes Pic, 1924b (MNHP); Madagascar *elateroides Fairmaire, 1896 (MNHP); Madagascar elonqatus (Klug), 1838 (Colobodera); Madagascar Lacordaire, 1857 (transferred to Daemon) Fairmaire, 1896 (redescription) fasciolatus Fairmaire, 1896 (MNHP); Madagascar *qeniculatus Pic, 1924b (MNHP); Madagascar qiqas de Laporte, 1836 (MNHP); Madagascar ovatus (Klug), 1838 (Colobodera) Lacordaire, 1857 (synonymized with qiqas) Fairmaire 1896 (redescription) (Daemoniminus) grandis Pic, 1953b (MNHP); Madagascar linearis de Laporte, 1836; Madagascar

100 Fairmaire, 1896 {redescription) 80 lineellus Fairmaire, 1896 (KNHP); Madagascar *lonqulus Fairmaire, 1896 (MNHP); Madagascar *minutus Fairmaire, 1896 (MNHP); Madagascar mucronatus (Klug), 1838 (Colobodera); Madagascar Lacordaire, 1857 (transferred to Daemon) Fairmaire, 1896 (redescription) *niqer Pic, 1953b (MNHP); Madagascar *niariceps Fairmaire, 1896 (MNHP); Madagascar *niqrithorax Pic, 1913b (MNHP); no locality nitidus (Klug), 1838 (Colobodera); Madagascar Lacordaire, 1857 (transferred to Daemon) Fairmaire, 1896 (redescription) notaticollis Pic, 1913b; no locality *pallidipes Pic, 1924b (MNHP);.Madagascar *quadrinotatus Pic, 1913b (MNHP); no locality striatus (Klug), 1838 (Colobodera); Madagascar Lacordaire, 1857 (transferred to Daemon) Fairmaire, 1896 (redescription) *subelonqatus Pic, 1913b (MNHP); no locality *trifasciatus Fairmaire, 1896 (MNHP); Madagascar *truncatulus Fairmaire, 1896 (MNHP); Madagascar *vittatus Pic, 1949 (MNHP); Madagascar

101 Falsotherius 81 Falsotherius Pic, 1913a: 165 (type species, by subsequent designation [Deleve, 1971: 355]): Falsotherius atricolor Pic, 1913a: 165); Diagnostic combination. This genus may be recognized by the following combination of characters: antennae slightly serrate, epistomal sulcus absent, protrochantins loosely concealed, tarsomere IV reduced, I-III with ventral, fleshy pads, II-III lobed, and males with apical labial (Fig. 18) and maxillary (Fig. 35) palpomeres ovoid and half-membranous. Description. Body size. Length mm, width at humeral angles mm. Head, Quadrate; antennae slightly serrate, nearly filiform; epistomal sulcus absent; labrum (Fig. 44) not transverse, basal width 2x or less greatest length, anterior margin concave, mandibles tridentate (as in Fig. 37), without prostheca, molar areas, or dorsal ridge; maxillae (Fig. 18), lacinia blunt, not forming spore brush, galea blunt with small production on lateral sub-apex; labium (Fig. 35), ligula biacuminate; labial and maxillary palpi shortened, in males, apical palpomeres ovoid, 1/2 membranous.

102 Thorax; Prothorax somewhat anteriorly produced, with 82 acute mediolateral margins very short, strong basal crenulation, mediobasal lobe crenulate, with produced denticles (as in Fig. 71); mesoscutellum cordiform; notai projection long, cowling loosely meeting anterior edge, trochantins loosely concealed (as in Fig. 66); mesosternal keel, excavation absent; elytra strongly striate, sutural striae not more deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete; metacoxal plates underdeveloped; tarsi, article IV reduced (as in Fig. 12), I-III with ventral, fleshy pads, II-III drawn into lobes; ungues simple. Abdomen. Male sternum V modified, emarginate; aedeagus without mediolateral spines or flanges, or subapical accessory projections; parameres long, without nonmelanized apical projections; ovipositor (Fig. 82) long, coxites not divided, articulated with bacula, styli absent. MATERIAL EXAMINED: 62 specimens (BMNH, NMNH, ÜCDC, MNHP, IZWP). Taxonomic list - Falsotherius *apicicornis (Pic), 1934a (Eoilichas) NEW COMBINATION (MNHP); Selangor (Malaysia) *atricolor Pic 1913a (MNHP); Sumatra *dohertvi Pic 1913a (MNHP); Malacca (Malaysia)

103 Pseudoepilichas 83 Pseudoepilichas Armstrong and Nakane, 1956: 212 (type species, by monotypy: Epilichas niponicus Lewis 1895: 101). Diagnostic combination. This genus may be recognized by the following combinatin of characters: male antennae with nonarticulated rami (as in Fig. 5), epistomal sulcus present, protrochantins visible (as in Fig. 65), tarsomere IV reduced (as in Fig. 12), II-III ventrally lobed, without fleshy pads, and apical palpomeres simple, largely sclerotized. Description. Body size. Length mm, width at humeral angles mm. Head. Quadrate; antennae, males with nonarticulated rami, antennomeres III-X (as in Fig. 5), females serrate; epistomal sulcus present; labrum, not transverse, basal width approx. 2x greatest length, anterior margin truncate or slightly convex (as in Fig. 47); mandibles bidentate, with prostheca, dorsal ridge (as in Fig. 41), without molar areas; maxillae, lacinia blunt, not forming spore brush, galea biacuminate (as in Fig. 28); labium, ligula quadracuminate (as in Fig. 21); apical palpomeres simple, mostly sclerotized.

104 Thorax. Prothorax with acute lateral margination, Q ^ incomplete, anterior end of margin not curving ventrally; basal margin weakly crenulate, mediobasal lobe smooth, with three produced denticles; notai projection long, trochantin visible (as in Pig. 64); mesoscutellum basally smooth (as in Fig. 70); mesosternal excavation, keel, absent; elytra obsoletely striate, sutural striae not deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete; metacoxal plates well-developed; tarsi, article IV reduced (as in Fig. 12), II-III lobed, without ventral fleshy pads; ungues simple. Abdomen. Male sternum V simple; aedeagus, median lobe with mediolateral flanges, without subapical accessory processes; parameres long, without nonmelanized apical projections; ovipositor (as in Fig, 80) long, coxites not divided, articulated with bacula, styli absent. MATERIAL EXAMINED; 8 specimens (KUBC, BMNH). Taxonomic - Pseudoepilichas *niponicus (Lewis), 1895 (Epilichas) (BMNH); Japan robustior Nakane, 1963; Japan

105 EctvPhodactvla NEW GENUS 85 Ectvphodactvla NEW GENUS (type species, by monotypy: E. fiski NEW SPECIES). Diagnostic combination. This genus may be recognized by the following combination of characters: lateral pronotal margination obsolete, protrochantins loosely concealed (Fig. 66), tarsomere IV reduced. III ventrally lobed, simple ungues (as in Fig. 12), head somewhat elongate, clypeus mostly covering labrum (Fig. 53), and males with apical maxillary and labial palpomeres elongate and mostly membranous (Figs. 17, 36). Description. Body size. Length mm, width at humeral angles 3-4 mm. Head. Rather elongate; antennae serrate; epistomal sulcus absent, labrum (Fig. 53) long, basal width 2x or less greatest length, anteriorly convex; mandibles tridentate, with prostheca and dorsal ridge (as in Fig. 37), without molar areas (as in Fig. 39); maxillae (Fig. 36), lacinia blunt, not forming spore brush, galea blunt; in males, apical maxillary and labial palpomeres elongate and largely membranous, membranous areas covered with short, evenly-spaced sensilla (Figs. 17, 36), in females, elongate and mostly sclerotized; labium (Fig. 17), ligula

106 biacuminate, each half tapered to blunt apex. 86 Thorax. Pronotum (Figs. 53, 66, 71) strongly dorsoanteriorly rounded, with obsolete lateral margination (Fig. 66), strong basal crenulation, mediobasal lobe crenulate, with produced denticles (Fig. 71); notai projection long, protrochantins concealed, cowlings loosely meeting anterior edge of notai projection (Pig. 66); mesoscutellum cordiform (Fig. 71); mesosternal excavation, keel absent; elytra obsoletely striate, sutural striae not deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete; coxal plates welldeveloped; tarsi (as in Fig. 12), article IV reduced. III ventrally lobed, without ventral fleshy pads, ungues simple. Abdomen, Male sternum V emarginate; aedeagus (Fig. 89), median lobe without mediolateral flanges or subapical accessory projections, parameres long, without nonmelanized accessory projections; ovipositor (as in Fig. 80) long, coxites not divided, articulated with bacula, styli absent. The following species description does not include characters discussed in the generic description. Ectvphodactvla fiski NEW SPECIES Type material. Holotype male, label data: COSTA RICA:

107 Heredia, Prov,, Pto. Viejo, Finca la Silva, II , 87 FWFisk (OSÜC). Paratype males, 8 specimens: Mexico, V. C. Volcan San Martin SE. slope, B & B Valentine/ beating at camp, el. 4000', VII-59 (OSDC); Coyame, Lake Catemaco, Veracruz, MEXICO, VII.63-Black, Its. D. R. Whitehead (NMNH); COSTA RICA, Turrialba, VII , P. J. Spangler (2 spec., NMNH); COSTA RICA, Turrialba, Cartage, III-11-67, W.R. Enns, coll./u Mo, C (EEMC); Costa Rica: Heredia, pr. LaSelva nr. Puerto Viejo 50m ,80, RS Anderson (CNCI); COSTA RICA Cart. Pr., 20 Km. E Turrialba, June 9, , J.E. Wappes (OSUC); Honduras, 1940/ W. von Hagen (AMNH). Paratype females, 4 specimens: MEX, Yerba Buena, 20 mi. N. Bochil, Chis. VI , D.E. Bright (CNCI); MEX, Veracruz, El Bastanal, nr., Coyame, ca. 2500', San Andres Mts., IX,19.65/ George E. Ball, D. R. Whitehead, collectors (NMNH); Mexico, V. C. Volcan San Martin SE. slope, B & B Valentine/ beating el,, ', 23-VII-59 (OSÜC); COSTA RICA: Puntarenas, Monteverde area, 6 June-14 June 1973, meters/ Erwin & Hevel, Central American Expedition, 1973 (NMNH). Description. Antennae very weakly serrate; clypeus lengthened, nearly covering labrum (Fig. 53); dorsal mandibular ridge prominent; anterolateral angles of ligula more produced than inner (Fig. 17); prominent groove dorsomesally to each eye; pronotum and elytra minutely

108 88 punctate, sparsely covered with fine setae; aedeagus (Fig. 89), median lobe very thin, parameres dorsoventrally compressed, broad, blade-like, apex and subapex covered with short setae. Larvae unknown. Etymology. Ectvphodactvla is from a combination of the greek words "ektyphos" (puffed up, or swollen) and "dactylos" (finger). "Ektyphos" is in reference to the bulbous pronotum; "dactylos" is taken from the family name, Ptilodactylidae; and the specific epithet "fiski" is in honor of Dr. Frank W. Fisk, Emeritus Professor, Department of Entomology, The Ohio State University, collector of the holotype specimen. MATERIAL EXAMINED: 13 specimens (NMNH, OSUC, CNCI, AMNH). Specimens are from Mexico (Chiapas, Veracruz), Costa Rica, and Honduras. Taxonomic list - Ectvphodactvla *fiski NEW SPECIES (OSUC); Mexico, Central America

109 Ptiloderes NEW GENUS Ptiloderes NEW GENUS (type species, by monotypy: P. humeralifer Pic, 1928c: 8). Diagnostic combination. This genus may be recognized by the following combination of characters: protrochantins concealed, lateral pronotal margination incompletely acute (as in Fig. 68), posterolateral pronotal angles broadly explanate (Pigs. 3, 52, 73-76), tarsi (as in Fig. 12), tarsomere IV reduced. III ventrally lobed, mesoscutellum cordiform (Figs ), and lacinia not forming spore brush (Fig. 34). Body size. Length mm, width at humeral angles mm. Head. Quadrate; male antennae with articulated rami (as in Fig. 7), female serrate; epistomal sulcus present, labrum (as in Fig. 47) long, basal width 2x or less greatest length, anteriorly truncate or slightly concave; mandibles bidentate, with prostheca and dorsal ridge (as in Fig. 41), without molar areas (as in Fig. 39); maxillae (Fig. 34), lacinia blunt, not forming spore brush, galea acute, apical palpomere mostly sclerotized; labium (Fig. 15), ligula with each half tapered to blunt apex. Thorax. Pronotum with incompletely acute lateral margins (Fig. 52, as in 58), hind angles broadly explanate

110 (Fig. 3, 52, 73-76), basal margin weakly crenulate, 90 mediobasal lobe smooth, with produced denticles (Figs ); notai projection long, protrochantins concealed (as in Fig. 68); mesoscutellum cordiform (Fig. 73); mesosternal excavation, keel absent; elytra obsoletely striate, sutural striae deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete; coxal plates well-developed; tarsi (as in Fig. 12), article IV reduced, III lobed, without ventral fleshy pads, ungues bifid. Abdomen. Male sternum V emarginate; aedeagus, median lobe without mediolateral flanges or subapical accessory projections, parameres long, witout nonmelanized apical projections; ovipositor (as in Fig. 80) long, coxites not divided, articulated with bacula, styli absent. Etymology. Ptiloderes is a combination of "ptilon" (Gr., feather, wing, leaf) and "dere" (Gr., fneck, throat) and is in reference to the broadly explanate posterolateral angles of the pronotum. MATERIAL EXAMINED: 45 specimens (IZWP, INPA, CNCI, EGRC, BMNH, LACM, MZSP, MNHB, NMNH, MNHP, CASC). There are 6 or 7 species in this genus from Panama, Ecuador, and Brazil. Further type studies are necessary to determine which species are described. Taxonomic list - Ptiloderes *humeralifer (Pic), 1928c (Ptilodactvla) NEW COMBINATION (MNHP); Brazil

111 Pherocladus 91 Pherocladus Fairmaire 1881: 372 (type species, by monotypy: P. dermestoides Fairmaire, 1881: 372); Deleve, 1972: 264. Diagnostic combination. This genus may be recognized by the following combination of characters: lateral prothoracic margination incompletely acute, protrochantins concealed and extreme anterior pronotal production (Fig. 61). Description. Body size. Length mm, width at humeral angles mm. Head, Quadrate; male antennae (as in Figs. 2, 7) with articulated rami, antennomeres IV-X, females serrate (as in Fig. 4); epistomal sulcus present; labrum (as in Fig. 47) not transverse, basal width 2x or more greatest length; mandibles (as in Fig. 41) bidentate, with prostheca, molar areas, and dorsal ridge; maxillae (as in Fig. 31), lacinia forming spore brush, galea acute, closely associated with spore brush and with longer, finer setae; apical maxillary palpomere mostly sclerotized; labium (as in Fig. 23), ligula simple, ventrally, transversely folded. Thorax. Prothoracic margins incompletely acute, extreme anterior pronotal production, dorsal to anterior

112 margin (Fig. 61); basal margin strongly crenulate, 92 mediobasal lobe crenulate, with three produced denticles (as in Fig. 71); mesoscutellum cordiform; notai projection long, protrochantins concealed (Fig. 61); mesosternal keel, excavation absent; elytra obsoletely striate, sutural striae variable, may or may not be deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete; metacoxal plates well-developed; tarsi (as in Fig. 12), article IV reduced. III lobed, without ventral fleshy pads; ungues with basal teeth. Abdomen. Male sternum V emarginate; aedeagus, median lobe without mediolateral spines or subapical accessory projections, parameres long, without apical nonmelanized projections; ovipositor (as in Pig. 80) long, coxites not divided, articulated with bacula, without styli. Species excluded; andrewesi Bourgeois (Ptilodactvla) MATERIAL EXAMINED: 9 specimens (MNHP, NMNH, BMNH, MCZC, IZWP).

113 Taxonomie list - Pherocladus 93 atriceps Pic, 1923c; Sumatra *capucina Pic, 1923c (MNHP); Sumatra *castanescens Pic, 1923c (MNHP); Borneo dermestoides Fairmaire, 1881; Insel Viti (Fiji Isl.) *inlatefalis Pic, 1923c (MNHP); Sumatra palawanus Deleve, 1972 (ÜZMD); Palawan, Philippines *plicata Pic, 1923c (MNHP); Borneo *sinqularicollis Pic, 1935 (BMNH); Borneo *testaceomaculata Pic, 1923c (MNHP); Sumatra

114 Lachnodactvla 94 Lachnodactvla Champion, 1897a; 658 (type species, here designated: L. monticola Champion, 1897a: 659); Schaeffer, 1906: 115; Hlavac, 1975: 182; Stribling, 1986; 232. Diagnostic combination. This genus may be recognized by the following combination of characters: protrochantins concealed (as in Fig. 68), lateral pronotal margination incompletely acute (as in Fig. 68), tarsomere IV reduced, III ventrally lobed (as in Fig. 12), mesoscutellum usually crenulate (as in Fig. 72), males with apical maxillary palpomeres thickened, elongate, and largely membranous (Fig. 27), and females usually with apical maxillary palpomere slender, elongate, and mostly sclerotized. Description. Body size. Length mm, width at humeral angles mm. Head. Quadrate; males, antennae with articulated rami (as in Figs. 2 and 7), females serrate (as in Pig. 4, usually more weakly serrate); epistomal sulcus present, labrum (as in Fig. 47) long, basal width 2x or less greatest length; mandibles (Fig. 41) bidentate, with prostheca, molar areas, and acute dorsal ridge; maxillae (Fig. 27), lacinia forming spore brush, galea acute.

115 closely associated with spore brush, with longer, thinner, 95 and less dense setae; in males, apical palpomere elongate, mesal 1/2 of more membranous, covered with evenly spaced, very short sensilla, in dried specimens often appearing as longitudinal furrow, in females usually somewhat elongate (less so in L. arizonica Schaeffer), mostly sclerotized. Thorax, Pronotum (as in Fig, 68) with incompletely acute lateral margins, strong basal crenulation, mediobasal lobe crenulate, with produced denticles; notai projection long, protrochantins concealed; mesoscutellum cordiform or crenulate; mesosternal excavation, keel absent; elytra strongly striate, sutural striae not deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete or absent; coxal plates well-developed; tarsi (as in Fig. 12), article IV reduced. III lobed, without ventral fleshy pads; ungues with basal tooth. Abdomen, Male sternum V emarginate; aedeagus (Fig, 96) without mediolateral flanges or subapical accessory projections, parameres long, without nonmelanized apical projections; ovipositor (as in Fig, 80) long, coxites not divided, articulated with bacula, styli absent, MATERIAL EXAMINED: 303 specimens (CNCI, HUIC, FMNH, CISB, FSCA, BMNH, UCDC, OSUC, AMNH, MCZC, COBC, JBSC, UCRC, RSNB, NMNH, EGRC, MSUC, TAMT).

116 96 Specimens are from Arizona, Texas, Florida, Mexico, C. America, and Colombia; those from Florida and Colombia represent 2 undescribed species. Taxonomic list - Lachnodactvla *arizonica Schaeffer, 1906 (NMNH); southern Arizona *monticola Champion, 1897 (BMNH); Mexico, Guatemala *parviscutum Champion, 1897 (BMNH); Mexico, Guatemala, Costa Rica *texana Schaeffer, 1906 (NMNH); Texas, Mexico

117 97 Ptilodactvla Ptilodactvla llliger, 1807: 342 (type species, by monotypy: Ptilodactvla elaterina Illiger, 1807: 342 [= Pvrochroa nitida DeGeer, 1775: 27]; Latreille, 1929: 461; de Laporte, 1836: 21, 1840; 258; Blanchard, 1845; 57; Lacordaire, 1857: 279; LeConte, 1861: 179; Chenu, 1870: 16; Kirsch, 1874: 371; Horn, 1880: 90; LeConte and Horn, 1883: 171; Champion, 1897: 627; Blatchley, 1910: 690; Legros, 1947: 96; Nakane, 1956: 51; Arnett, 1968: 461; Deleve, 1972: 264; Stribling, 1986: 232. Hvpselothorax Kirsch, 1866: 188 (I have never seen this publication, thus I cannot here record the type species). Ptvlodactvla de Laporte, 1836: 21 (incorrect subsequent spelling), Stenactvla Fairmaire, 1896: 346 (type species, by monotypy: Stenactvla pallioes Fairmaire, 1896: 346). NEW SYNONYMY. Falsodaemon Pic, 1913c: 4 (type species, by monotypy: Falsodaemon madecassus Pic, 1913c: 4). NEW SYNONYMY. Theriomorphus Pic, 1913c: 1 (type species, by monotypy: Theriomorphus niasensis Pic, 1913c: 1). NEW SYNONYMY. Due to the large number of species included in Ptilodactvla it is premature to propose hypothetical

118 synapomorphy. Therefore, since the type species of 98 Stenactvla, Falsodaemon, and Theriomorphus cannot be distinguished from Ptilodactvla, the genera are synonymized. Diagnostic combination. This genus may be recognized by the following combination of characters; lateral pronotal margination incompletely acute, trochantins concealed (Fig. 68), tarsi (as in Fig. 12) tarsomere IV reduced. III lobed ventrally, ungues with basal tooth, mesoscutellum usually cordiform, apical palpomeres mostly sclerotized and securiform, and male antennae with articulated rami (Fig. 2, 7). Description. Body size. Length mm, width at humeral angles < mm. Head. Quadrate; male antennae with articulated rami (Figs. 2, 7), female serrate; epistomal sulcus present, labrum long (Fig. 47), basal width 2x or less greatest length, anteriorly truncate or slightly convex; mandibles (as in Fig. 41) bidentate, with prostheca, molar areas, and dorsal ridge; maxillae (as in Fig. 31), lacinia forming spore brush, galea acute, closely associated with spore brush, with longer, thinner, and less dense setae, apical palpomere mostly sclerotized; labium (as in Fig. 23),.

119 ligula simple, usually somewhat ventrally, transversely 99 folded. Thorax. Pronotum with lateral margins incompletely acute (Fig. 68), basal margin strongly crenulate, mediobasal lobe crenulate, with produced denticles (as in Fig. 71); notai projection long, protrochantins concealed (Fig. 68); mesoscutellum cordiform, rarely, crenulate; mesosternal excavation absent, keel, rarely, slightly developed; elytra strongly or obsoletely striate, sutural striae not deeply impressed; metasternal transverse suture usually not visible, coxal plates well-developed; tarsi (as in Fig. 12), article IV reduced. III lobed, without ventral fleshy pads, ungues bifid or with basal tooth. Abdomen, Male sternum V usually emarginate; aedeagus, median lobe without mediolateral flanges or subapical accessory projections, parameres long, without nonmelanized projections; ovipositor (as in Fig. 80) long, coxites not divided, articulated with bacula, styli absent. SPECIES EXCLUDED: humeralifer Pic (Ptiloderes); all following species are transferred to Epiotvoma: rufa Champion, rufipennis Pic, corvina Champion, ebenina Champion, maculate Champion, nigricornis Champion. More than 20,000 specimens are available for examination from the repositories in the "Material" section.

120 Taxonomie list - Ptilodactvla 100 *acuta Johnson and Freytag, 1978 (NMNH); Georgia *aequinoctialis Champion, 1897a (BMNH); Panama, Costa Rica africana Pic, 1942 (MNHP); Cameroon amamioshimana Nakane, 1963; Japan amazonica Pic, 1947b; Amazon (Brazil) *andrewesi (Bourgeois), 1896 (Pherocladus) NEW COMBINATION; India anqusta Kirsch, 1874; Peru *anqustata Horn, 1880 (MCZC); eastern North America anqustatipennis Pic, 1916b (MNHP); India Sato, 1979 (diagnosis) annulata Pic, 1924 (1923c) (MNHP); South America annulicornis Chevrolat, 1870 (MHCC); Cuba *antennalis Champion, 1897a (BMNH); Honduras *antillarum Champion, 1897b (BMNH); Antilles *apicalis (Pic), 1929 (Stenactvla) NEW COMBINATION (MNHP) ; Africa V. *robustior Pic, 1929 (MNHP); Africa apiceniqra Pic, 1947b; Brazil apicicornis Pic, 1916a (MNHP); Ceylon ardua Sato, 1979 (NMBE); India atra de Laporte, 1836; Cayenne (French Guiana) atricollis Pic, 1917a; Vietnam *atricolor Pic, 1947 (MNHP); Peru

121 *atricornis Pic, 1924 (1923c) (MNHP); Malay Archipelago ICI atrosuturalis Pic, 1916a (MNHP); America meridionalis *australis Bourgeois, 1884 (RSNB); New Caledonia, New Hebrides baeri Pic, 1916a (MNHP); Peru *bakeri Pic, 1923c (MNHP); Borneo, Mindanao Deleve, 1972 (redescription) V. *curtithorax Pic, 1923c (MNHP) V. *sexualis Pic, 1923c (MNHP) V. *mindanaosa Pic, 1923c (MNHP) *banatama Deleve, 1972 (üzmd); Lavongai, New Guinea baroniurbanii Sato, 1979 (NMBE); Nepal benhana Pic, 1929; Vietnam (Tonkin) bhutanica Sato, 1979 (NMBE); Bhutan bicolor Deieve, 1972 (ÜZMD); Palawan, Philippines bicoloriceps Pic, 1923a (NMNH); Trinidad boqorensis Pic, 1916a (MNHP); Malacca (Indonesia) boootensis Pic, 1916a (MNHP); America meridionalis borneensis Pic, 1916a (MNHP); New Guinea boucardi Pic, 1923c; Sumatra brancuccii Sato, 1979 (NMBE); Nepal ssp. darieelinaiana Sato, 1979 (NMBE); India bremensis Pic, 1947b; Brazil breveimpressa Pic, 1923a (MNHP); Brazil brevenotata Pic, 1947b; no locality V. innotata Pic, 1947b; no locality

122 brevepubens Pic, 1947b; Colombia 209 *brevicollis Champion, 1897 (BMNH); Mexico brevipennis Pic, 1916a (MNHP); Sumatra *brevipubescens Deleve, 1972 (UZMD); Yalom, New Britian, New Guinea *breviramus Pic, 1923c (MNHP); Borneo, Philippines Deleve, 1972 (redescription) *breviscutum Champion, 1897a (BMNH); Guatemala *bruneiensis Pic, 1924 (1923c) (MNHP); Borneo brunneipennis Pic, 1923c; Borneo, Philippines *brunneomarqinata Pic, 1924 (1923c) (MNHP); South America brunneonotata Pic, 1916a (MNHP); Mexico burqeoni Pic, 1934d (MRAC); Belgian Congo (Zaire) calcuttana Pic, 1924 (1923c) (MNHP); India calwelli Pic, 1934b; New Guinea *canaliculata Champion 1897a (BMNH); Mexico, Guatemala caracaensis Pic, 1947b; Venezuela carbonaria Chevrolat, 1870 (MHCC); CUba *carinata Johnson and Freytag, 1978 (NMNH); Kentucky castanea de Laporte, 1836 ; Cayenne (French Guiana) castaneonotata Pic, 1924 (1923c) (MNHP); South America catharinae Pic, 1947b; Brazil *caudata Champion, 1897a (BMNH), Central America celebensis Pic, 1916a (MNHP); Malacca (Indonesia) chiriquensis Champion, 1897a (BMNH); Panama

123 cisteloides Kirsch, 1874; Peru 103 compressa Pic, 1947c (MNHP); Peru *confinis Champion, 1897a (BMNH); Central America confusa Deleve, 1972 (ÜZMD); Yalom, New Britian, New Guinea conradti Pic, 1947b; Guatemala convexa Champion, 1897a (BMNH); Mexico, Nicaragua convexicollis Champion, 1897a (BMNH); Central America *corporaali Pic, 1923b (MNHP); Sumatra costaricana Pic, 1947b (MNHP); Costa Rica *costaricensis Champion, 1897a (BMNH); Costa Rica Costata Pic, 1923c (MNHP); South America crenuatostriata Redtenbacher, 1868; Brazil *cruciata Kirsch, 1866; Colombia cucullata Champion, 1897a (BMNH); Central America curta Pic, 1923c; Sumatra *curtipennis Pic, 1923c; Borneo; Philippines curtiramous Pic, 1929; Bolivia daemoniformis Pic, 1917b (MNHP); Banguey dascilliformis Pic, 1923c; Sumatra *debilis Champion, 1897a (BMNH); Mexico, Guatemala decumana Erichson, 1847; Peru *deleta Champion, 1897a (BMNH); Central America delevi (NEW NAME for reducta Deleve, 1972) (UZMD); Tawi Tawi, Philippines

124 104 densepunctata Pic, 1923c (MNHP); South America *dentaticornis Pic, 1924 (1923c) (MNHP); South America *denticollis Champion, 1897a (BMNH); Costa Rica *denticulata Champion, 1897a (BMNH); Central America deplanata Champion, 1897a (BMNH); Panama deplanaticeps Pic, 1929; Bolivia dermestoides Pic, 1923c (MNHP); Celebes (Sulawesi) dilatata Pic, 1929 (MNHP); Bolivia dilaticollis Champion, 1897a (BMNH); Panama dilatithorax Pic, 1947b; Brazil *discedens Deleve, 1972 (UZMD); Lorengau, Manus, N.Guinea discicollis Pic, 1930a; Bolivia discoidalis Pic, 1924 (1923c) (MNHP); South America diversa Pic, 1923c (MNHP); Borneo; Philippines Deleve 1972 (redescription) diversecostata Pic, 1935; Borneo diversepunctata Pic, 1924 (1923c) (MNHP); China diversipubens Pic, 1924 (1923c) (MNHP); South America *dohertvi Pic, 1916a (MNHP); Jobi donckieri (see note) Note; I do not know if this name is published as a full species or is merely a manuscript name. I did not find this type in the Pic collection; he (1952, 1955) published the two varieties under it.

125 105 V. V. ruandana Pic, 1955 (MRAC); Ruanda flandriana Pic, 1952 (MRAC); Belgian Congo (Zaire) elonqata de Laporte, 1836; Colombia emarqinata Chevrolat, 1870 (MHCC); Cuba *equilobata Chapin, 1927 (NMNH); United States *exotica Chapin, 1927 (NMNH); United States Suss and Puppin, 1977 (redescription) favrei Pic, 1924 (1923c) (MNHP); Travancore, India ferruqinea Boheman, 1858; Brazil *forcioata Champion, 1897a (BMNH); Mexico forteimpressa Pic, 1947c (MNHP); Brazil *forticornis Champion, 1897a; Mexico fruhstorferi Pic, 1947c; Brazil frvi Pic, 1924 (1923c) (MNHP); South America furcata Pic, 1954; China fuscicornis Champion, 1897a (BMNH); Guatemala qermaini Pic, 1916a (MNHP); Neotropics *qermana Champion, 1897a (BMNH) ; Panama *qibbicollis Champion, 1897a (BMNH); Panama *qlabrata Champion, 1897a (BMNH); Costa Rica qodavarensis Sato, 1979 (NMBE); Nepal qorhami Pic, 1924 (1923c) (MNHP); South America qounelli Pic, 1947b; Brazil *qracilicornis Deleve, 1972 (UZMD); Yalom, New Britian, New

126 Guinea -0^ *qracilis Champion, 1897a (BMNH); Guatemala qranulata Pic, 1917b (MNHP); Borneo *qranulicollis Champion, 1897a (BMNH); Mexico qriseosuturalis Pic, 1930b; Argentina V, rufescens Pic, 1930b. qriseoundulata Pic, 1928d; Brazil quadalupensis Legros, 1947 (MNHP); Guadeloupe quatemalensis Champion, 1897a; Guatemala hektosa (NEW NAME for rufithorax Pic, 1923c); Sumatra *heterophva Kirsch, 1873; Peru hickeri Pic, 1947b; Brazil hirsuta Pic, 1923c; Borneo, Philippines humeralis Motschulsky, 1863; Ceylon V, picea Motschulsky, 1863; Ceylon humeridens Pic, 1947c; Colombia humerosa Champion, 1897a (BMNH); Mexico, Antilles hyperqlotta Johnson and Freytag, 1982 (NMNH); Texas, 0. S. A. impressa Pic, 1924 (1923c) (MNHP); Ganee, Malay Archipelago impressipennis Pic, 1916a (MNHP); Neotropics incisa Pic, 1924 (1923c) (MNHP); South America inhursuta Pic, 1923c; Sumatra inlateralis Pic, 1924 (1923c) (MNHP); Sumatra

127 instriata Pic, 1916a (MNHP); Malacca (Indonesia) 107 *inteara Champion, 1897a (BMNH); Mexico, Guatemala irregularis Pic, 1928e; Buenos Aires, Argentina ishiqakiana Sato, 1968; Ryukyu Archipelago isoloba Johnson and Freytag, 1982 (NMNH); Kentucky, Ü.S.A. iaponensis (=macrophthalma Nakane) iiriensis Sato, 1979 (NMBE); Nepal *iokoensis (Pic), 1931b NEW COMBINATION (Stenactvla) (MNHP); Cameroon klapperichi Pic, 1954 (DEIE, MNHP); China lacorderei de Laporte, 1836 ; Cayenne (French Guiana) laiovei Pic, 1917b (MNHP); Ho Chi Minh City, Vietnam lamellifera Kirsch, 1873; Peru laoensis Pic, 1926a; Laos lata Pic, 1926b (MNHP); Bolivia lateralis Pic, 1923c; Sumatra laticollis Pic, 1923c; Borneo, Philippines latior Pic, 1923a (MNHP); Trinidad latithorax Pic, 1924 (1923c) (MNHP); Malay Archipelago Delève, 1972 (redescription) leopoldi Pic, 1952 (MRAC); Belgian Congo (Zaire) lerovi Pic, 1952 (MRAC); Belgian Congo (Zaire) loiaensis Pic, 1947c; Ecuador lonqicornis Pic, 1923c (MNHP); Borneo, Philippines Delève, 1972 (redescription)

128 lonqiramus Pic, 1923c (MNHP); Borneo, Philippines Delève, 1972 (redescription) lovasatoi Pic, 1947b; Uruguay luteipes Pic, 1923c (MNHP); Sumatra Delève, 1972 (redescription) luteitarsus Pic, 1947a; Peru *lutescens Champion, 1897a (BMNH); Mexico; Guatemala luzonica Pic, 1923c (MNHP); Borneo; Philippines Delève 1972 (redescription) macrophtalma Legros, 1947 (MNHP); Guadeloupe macrophthalma Nakane, 1963; Japan iaponensis Nakane 1977 UNJUSTIFIED NAME CHANGE madecassus (Pic), 1913c NEW COMBINATION (Palsodaemon) (MNHP); Madagascar madurensis Pic, 1917b (MNHP); India maindroni Pic, 1924 (1923c) (MNHP); India malaccana Pic, 1923a (MNHP); Malacca (Idonesia) malabarensis Pic, 1938b (DEIE); Malabar (SW India) manuanesis Delève, 1972 (UZMD); Manuan, Duke of York, New Guinea *marcida Champion, 1897a (BMNH); Guatemala *marqinata Champion, 1897a (BMNH); Mexico, Guatemala martapurana Pic, 1916a (MNHP); Malacca (Indonesia) maxima Pic, 1923c (MNHP); Bogor (Java) medanensis Pic, 1917b (MNHP); Sumatra

129 *mexicana Champion, 1897a (BMNH); Mexico 109 *militaris Chevrolet, 1870 (MHCC); Cuba minacensis Pic, 1947b; Brazil minima Pic, 1924 (1923c) (MNHP); New Guinea minor Pic, 1923c (MNHP); Borneo, Philippines V. diversicolor Pic, 1923c (MNHP); Borneo, Philippines Delève, 1972 (redescription) miniscula Pic, 1923c (MNHP); Malacca (Indonesia) *minuta Kirsch, 1873; Peru miscella Sato, 1979 (NMBE); Bhutan modesta Pic, 1924 (1923c) (MNHP); South America modiqlianii Pic, 1924 (1923c) (MNHP); Malay Archipelago *montana Champion, 1897a (BMNH); Panama mrazai Pic, 1928d (Prague); Brazil multicostata Pic, 1947c; Brazil multifaria Sato, 1979 (NMBE); Nepal multiimoressa Pic, 1947c; Brazil mussauiana Delève, 1972 (ÜZMD); Talumalaus, Mussau, New Guinea mutata Pic; Bolivia (=scutellaris Pic 1928b, not Kirsch) nanoderma Johnson and Freytag, 1982 (NMNH); Indiana, U.S.A. nigra Kirsch, 1873; Peru nigronotata Pic, 1916a (MNHP); America meridionalis nitens de Laporte, 1836 ; Cayenne (French Guiana)

130 110 nitida (Degeer), 1775 (Pvrochroa); Cayenne (French Guiana) elaterina Illiger 1807 Guerin-Meneville, (illustrated elaterina) nitidissima Pic, 1927b; Formosa nitidor Pic, 1916a (MNHP); Sumatra nodieri Pic, 1947b; Peru notata Pic, 1929; Bolivia notaticollis. Pic 1923b; Sumatra novabritannica Delève, 1972 (UZMD); Yalom, New Britian, NewGuinea *novahibernica Delève, 1972 (UZMD); Danu, New Ireland, NewGuinea obesa Erichson, 1847; Peru obiiauefasciata Pic, 1947b; Bolivia *obovata Champion, 1897a (BMNH); Mexico obscura Kirsch, 1873; Peru obscuriceps Pic, 1923c; Sumatra V. separata Pic, 1923c; Sumatra obscuricollis Pic, 1923c; Borneo, Philippines ocularis Pic, 1916a (MNHP); New Guinea omospila Kirsch, 1873; Peru *opacicollis Pic, 1924 (1923c) (MNHP); South America opacithorax Pic, 1923c (MNHP); Colombia *opima Champion, 1897a (BMNH); Guatemala, Panama

131 *ornata de Laporte, 1836; Brazil 111 *ovalis Delève, 1972 (UZMD); Palawan, Philippines pallescens Kirsch, 1873; Peru pallida Steinheil, 1874; Argentina pallidicolor Pic, 1923c; Sumatra pallidior Pic, 1929; Bolivia pallidipes Pic, 1916a (MNHP); Vietnam (Tonkin) pallidomarqinata Pic, 1924 (1923c) (MNHP); Java *pallipes (Pairmaire), 1896 NEW COMBINATION (Stenactvla) (MNHP); Madagascar *parallela Champion, 1897a (BMNH); Costa Rica, Panama paranana Pic, 1928c; Argentina particularicornis Pic, 1924 (1923c)- (MNHP); Mexico pauli Pic, 1924 (1923c) (MNHP); Brazil V. V, iuneta Pic, 1924 (1923c); South America elonqatior Pic, 1928c; Brazil pendleburvi Pic, 1934a; Malaysia penta (NEW NAME for minuta Pic, 1947b); Brazil peruviana Pic, 1923a (MNHP); Peru peruviensis Pic, 1947b; Peru *pici (NEW COMBINATION, NEW NAME for Stenactvla anqustata Pic, 1929 [MNHP]; Africa) piniqisana Delève, 1972 (UMZD); Palawan, Philippines planatipennis Delève, 1972 (UMZD); Yalom, New Britian, New Guinea

132 praecellens Kirsch, 1873; Peru 11 ') prescutellaris Pic, 1952 (MRAC); Belgian Congo (Zaire) prima (NEW NAME for atricollis Pic, 1947b); Colombia probanda Kirsch, 1873; Peru *pruinosa Champion, 1897a (BMNH); Mexico puberula Pic, 1928c; Brazil pubescens Pic, 1923c; Borneo, Philippines *punctatissima Champion, 1897a (BMNH); Mexico, Guatemala punctatostriata Murray, 1868 (BMNH); Old Calabar punosa Pic, 1947b; Peru *ramea Lewis, 1895 (BMNH); Japan ramicornis Chevrolat, 1870 (MHCC); Cuba rectiramus Pic, 1929; Bolivia V. notaticollis Pic, 1929; Bolivia reducta Pic, 1924 (1923c) (MNHP); Argentina reitteri Pic, 1924 (1923c) (MNHP); South America robusta Pic, 1917b (MNHP); Sumatra roonensis Pic, 1924 (1923c) (MNHP); Malay Archipelago *rotundicollis Champion, 1897a (BMNH); Guatemala rouveri Pic, 1917b (MNHP), Java V. V, notaticollis Pic, 1923b; Sumatra brunneosuturalis Pic, 1938a; Pahang (Malaysia) rubricollis Pic, 1952 (MRAC); Belgian Congo (Zaire) rufescens Pic, 1947b; Colombia ruficolor Pic, 1916a (MNHP); American meridionalis

133 rufipes Pic, 1924 (1923c) (MNHP); South America 113 rufithorax Pic, 1917a; Bolivia rufocincta Pic, 1916a; America meridionalis rufohumeralis Pic, 1924 (1923c) (MNHP); Palawan, Philippines rufosuturalis Pic, 1924 (1923c) (MNHP); Bolivia *rufonotata Pic, 1923a (MNHP); Brazil rufotestacea Champion, 1897a (BMNH); Guatemala rufula Pic, 1947b; Bolivia *ruqulosa Champion, 1897a (BMNH); Mexico saiqonensis Pic, 1929; Ho Chi Minh City, Vietnam sallei Pic, 1924 (1923c) (MNHP); Mexico salti Pic, 1953a; Kilimanjaro (Tanzania) *sanctisvincentis Champion, 1897b (BMNH); Antilles sanqirensis Pic, 1924 (1923c) (MNHP); Malay Archipelago sapitensis Pic, 1924 (1923c) (MNHP); Lombok (Indonesia) satoi (NEW NAME for P. bicolor Sato, 1979 [NMBE]); Bhutan *scabrosa Champion, 1923 (BMNH); Seychelles *scapularis Champion, 1897a (BMNH); Panama scrutata Kirsch, 1866; Colombia *scutata Champion, 1897a (BMNH); Panama scutellaris Kirsch, 1873; Peru scutellaris Pic ( =mutata Pic) secedens Kirsch, 1873; Peru secunda (NEW COMBINATION, NEW NAME for Stenactvla bicolor

134 Pic, 1925b [MNHP]); Madagascar 114 V. V. testaceicornis Pic, 1925b (MNHP); Madagascar testaceipennis Pic, 1925b (MNHP); Madagascar semiobscura Pic, 1923c; Borneo, Philippines sericea de Laporte, 1836 ; Cayenne (French Guiana) *serrata Champion, 1897a (BMNH); Costa Rica, Panama *serricollis (Say), 1823 (Ptilinus); ü. S. A. fusca Melsheimer, 1845 Johnson and Freytag, 1982 (neotype designation) *sexualis Delève, 1972 (ÜMZD); Palawan, Philippines simplex Chevrolat, 1870 (MHCC); Cuba simulans Kirsch, 1873; Peru sinensis Pic, 1923a; China V. sauteri Pic, 1927b; Formosa solidicornis Delève, 1972 (üzmd); Balabac, Philippines sparsepunctata Pic, 1916a (MNHP); Borneo strangulate Pic, 1924c; Sumatra striatella Delève, 1972 (ÜZMD); Palawan, Philippines striatipennis Pic, 1923c (MNHP); Borneo, Phillipines Delève, 1972 (redescription) *strictifrons Delève, 1972 (ÜZMD); Banatam, Lavongai, NewGuinea suapeinsis Pic, 1947b; Bolivia subacuminata Pic, 1916a (MNHP); Java subcastanea Motschulsky, 1863; Java

135 subconvexa Pic, 1923c; Sumatra subdepressa Pic, 1924 (1923c) (MNHP); Malacca (Malaysia) subelonqata Pic, 1924 (1923c) (MNHP); Malacca (Malaysia) subiunctus Pic, 1953b NEW COMBINATION (Daemon) (MNHP); Madagascar *submaculata Champion, 1897a (BMNH); Honduras, Nicaragua subovata Pic, 1924 (1923c) (MNHP); South America *subparallela Champion, 1897a (BMNH); Mexico *substriata Champion, 1897a (BMNH); Costa Rica substriatipennis Pic, 1928c; Brazil subvittata Pic, 1924 (1923c) (MNHP); Malay Archipelago sulcata Champion, 1897a (BMNH); Mexico suturalifer Pic, 1929; Bolivia suturalis Pic, 1916a (MNHP); America meridionalis tabascoana Champion, 1897a (BMNH); Mexico takahashii Sato, 1968; Ryukyu Archipelago tanqana (Pic), 1914 NEW COMBINATION (Stenactvla) (MNHP); no locality tenanarivana (Pic), 1925 NEW COMBINATION (Stenactvla) (MNHP); Madagascar tenuipunctata Delève 1972 (ÜZMD); Yalom, New Britian, New Guinea tenuis Champion, 1897a (BMNH); Guatemala testaceicollis Pic, 1916a (MNHP); America meridionalis V. inhumeralis Pic, 1947b; Bolivia

136 testaceicornis Pic, 1924 (1923c) (MNHP); Malay Archipelago testaceimembris Pic, 1924 (1923c) (MNHP); Madura (Java) testaceipes Pic, 1924 (1923c) (MNHP); South America *testaceohumeralis Pic, 1952 (MRAC); Belgian Congo (Zaire) testaceonotata Pic, 1924 (1923c) (MNHP); Borneo, Philippines Delève 1972 (redescription) *tetra (NEW COMBINATION, NEW NAME for Stenactvla donkieri Pic, 1913c [MNHP]); Congo theresae Pic, 1947a; Brazil thoracica de Laporte, 1840 ; Cayenne (French Guiana) thri (NEW NAME for curta Pic, 1925); Borneo toliana Pic, 1924 (1923c) (MNHP); Celebes (Sulawesi) tonkinea Pic, 1916a (MNHP); Vietnam (Tonkin) transversicollis Pic, 1923a (MNHP); Brazil tricoloricornis Pic, 1923a (MNHP); Peru triimpressa Pic, 1924 (1923c) (MNHP); South America trinotata Lacordaire, 1857; Brazil *tropicalis Champion, 1897a; Central America truncata Pic, 1928c; Brazil truncaticollis Pic, 1947a; Peru tucumana Pic, 1928c; Argentina turrialbana Pic, 1947b; Costa Rica ueleensis Pic, 1952 (MRAC); Belgian Congo (Zaire)

137 undulata Pic, 1923c (MNHP); Borneo, Philippines Deleve 1972 (redescription) V. V. V, obscurior Pic, 1923c; Borneo, Philippines subreqularis Pic, 1923c; Borneo, Philippines proxima Pic, 1923c; Borneo, Philippines ssp. iavanica Delève, 1972 (UZMD); Java *varicornis Champion, 1897a (BMNH); Panama varieqata Kirsch, 1889; Colombia *venusta Delève 1972 (UZMD); Talumalaus, Mussau, New Guinea vicina Pic, 1923b; Sumatra vilis Kirsch, 1873; Peru *vittata-pic, 1916a (MNHP); America meridionalis V. invittata Pic, 1926b (MNHP); Brazil vuilleti Pic, 1917a; Vietnam (Tonkin) waqneri Pic, 1924 (1923c) (MNHP); South America waterstradti Pic, 1924 (1923c) (MNHP); Malay Archipelago wittei Pic, 1950; Belgian Congo (Zaire) wittmeri Sato, 1979 (NMBE); Bhutan

138 Stirophora 115 Stirophora Champion, 1897a; 660 (type species, by monotypy: S. sulcipennis Champion, 1897a: 660); Stribling, 1986: 232. Chaetodactvla Champion, 1897a: 660 (type species, by monotypy: Ç. Ivciformis Champion, 1897a: 661); Hlavac, 1975: 182; Stribling, 1986: 232. NEW SYNONYMY. This synonymy is on the basis of joint possession of the apomorphic mesosternal keel and the subapical accessory projections of the median aedeagal lobe (Fig. 94). Diagnostic combination. This genus may be recognized by the following combination of characters: tarsomere IV reduced. III ventrally lobed (as in Fig. 12), protrochantins concealed (as in Fig. 68), epistomal sulcus present, lateral pronotal margins incompletely acute (as in Fig. 68), male antennae with articulated rami (as in Fig. 2, 7), mesosternal keel present, usually very prominent, and median aedeagal lobe with subapical accessory projections (Fig. 94). Description. Body size. Length mm; width at humeral angles mm. Head. Quadrate; antennae, males with articulated rami (as in Fig. 2, 7), females serrate (as in Fig. 4); labrum

139 (as in Fig. 47), not transverse, basal width approx. 2x 119 greatest length, anteriorly truncate or slightly convex; epistomal sulcus present; mandibles (as in Fig. 41) bidentate, with prostheca, molar areas, and distinct dorsal ridge; maxillae (as in Fig. 31), lacinia forming welldeveloped spore brush, galea closely associated with spore brush, with longer, less dense, and thinner setae, apical palpomere mostly sclerotized and melanized; labium (Fig. 23), ligula simple, usually ventrally, transversely, folded. Thorax. Prothorax with acute, incomplete lateral margins, anteriorly curved toward venter (as in Fig. 68); weak basal crenulation, mediobasal lobe smooth, with three produced denticles; mesoscutellum cordiform; notai projection long, met anteriorly by cowling, protrochantin concealed (as in Fig. 68); mesosternal excavation absent, keel usually present; elytra strongly striate, striae sometimes obscured by setation, sutural stria not more evident; metasternal transverse suture present near discrimen, laterally obsolete; coxal plates well-developed; tarsi (as in Fig. 12), article IV reduced. III lobed, ungues with basal tooth (as in Fig. 11 with mesal furcation truncate). Abdomen. Male visible sternum V modified, emarginate; aedeagus (Fig. 94), median lobe with subapical accessory

140 120 projections, without mediolateral flanges, parameres long, without nonmelanized apical projections; ovipositor (Fig. 80), coxites not divided, without styli, articulated with bacula. MATERIAL EXAMINED: 104 specimens (CNCI, HAHC, NDSÜ, EEMC, EGRC, PMNH, NMNH, BMNH, ÜICM, AMNH). Taxonomic list - Stirophora *lvciformis (Champion), 1897a (Chaetodactyla) NEW COMBINATION (BMNH); Panama, Costa Rica, Nicaragua, Colombia *sulcipennis Champion, 1897a (BMNH); Panama There are four species; the two undescribed are from Costa Rica.

141 Chelonar iomorphus 121 Chelonariomorphus Pic, 1916a; 2 (type species: Ç. subconvexus Pic, 1916a: 2); Stribling, 1986: 232. Diagnostic combination. This genus may be recognized by the following combination of characters: prothoracic margins equiplanar and protrochantins concealed (Fig. 67). Description. Body size. Length mm, width at humeral angles mm. Head. Quadrate; antennae (as in Figs. 2, 7), males with articulated rami on articles IV-X, females serrate; epistomal sulcus present; labrum (as in Fig. 47) not transverse, basal width approx, 2x greatest length, anterior margin truncate or slightly concave; mandibles (as in Fig. 41) bidentate, with prostheca, molar areas, and dorsal ridge; maxillae (as in Fig, 31), lacinia forming spore brush, galea acute, closely associated with spore brush and with longer, finer setae; apical palpomere mostly sclerotized; ligula (as in Fig. 23) simple, ventrally, transversely folded. Thorax, Prothoracic margins equiplanar, lateral margins continuous, or very nearly so, with anterior (Fig. 67); basal margin smooth, mediobasal lobe smooth, without produced denticles (Fig. 70); mesoscutellum basally smooth

142 (Fig. 70); notai projection long, trochantins concealed 122 (Fig.-67); mesosternal keel present, excavation shallow; elytra moderately or obsoletely striate, sutural striae not deeply impressed; metasternal transverse suture absent; metacoxal plates well-developed; tarsi, artilce IV reduced, III ventrally lobed (as in Fig. 12), without fleshy pads; ungues with basal teeth. Abdomen. Male sternum V emarginate; aedeagus with mediolateral flanges, without accessory projections, parameres long, without apical nonmelanized projections; ovipositor (as in Fig. 80) long, coxites not divided, articulated with bacula; styli absent. MATERIAL EXAMINED; 18 specimens (BMNH, MNHP, INPA, LACM, CNCI, FMNH, NMNH, MZSP). Taxonomic list - Chelonariomorphus *subconvexus Pic, 1916 (MNHP); Bolivia I have examined three species (two undescribed) with specimens from Suriname, British Guiana, Peru, and Ecuador.

143 Epjptyqma NEW GENUS 123 Epjptygma NEW GENUS (type species, present designation: Ptilodactyla rufa Champion, 1897a: 630). Diagnostic combination. This genus may be recognized by the following combination of characters: protrochantins concealed, lateral pronotal margination incompletely acute (as in Fig. 68), tarsi (as in Fig. 12), tarsomere IV reduced. III yentrally lobed, mesosternal keel present, mesoscutellum cordiform with median notch usually continuing as striole to or nearly to apex, and lacinia forming spore brush (as in Fig. 31), and male visible abdominal tergum VI with yentrally-directed flange (Figs ). Description. Body size. Length mm, width at humeral angles mm. Head. Quadrate; male antennae with articulated rami (as in Fig. 7), female serrate (as in Fig. 4); epistomal suture present, labrum (as in Fig. 47) long, basal width 2x or less greatest length, anteriorly truncate or slightly conyex; mandibles (as in Fig. 41) bidentate, with prostheca, molar areas, and dorsal ridge; maxillae (as in Fig. 31), lacinia forming spore brush, with longer, thinner, and less dense setae; apical palpomere mostly

144 124 sclerotized; labium (as in Pig. 23), ligula simple, often somewhat ventrally, transversely folded. Thorax, Pronotum with incompletely acute lateral margins (as in Fig, 68), strong basal crenulations, occasionally smooth on either side of mediobasal lobe, mediobasal lobe crenulate, with produced denticles (as in Fig 71); notai projection long, protrochantins concealed (as in Fig, 68); mesoscutellum cordiform (as in Fig, 71); mesosternal excavation absent, keel present; elytra obsoletely striate, sutural striae usually deeply impressed; metasternal transverse suture absent; coxal plates well-developed; tarsi (as in Fig, 12), article IV reduced. III ventrally lobed, without ventral fleshy pads, ungues with basal tooth. Abdomen, Male sternum V emarginate; segment VI usually heavily sclerotized and with ventrally-directed, posterior tergal flange (Figs, 55-60); aedeagus, median lobe without mediolateral flanges or subapical accessory projections, parameres long, without nonmelanized apical projections; ovipositor (as in Fig, 80) long, coxites not divided, articulated with bacula, styli absent. Etymology, Epiptvcma is greek for "over-fold" and "flap" and is in reference to the modification of the visible abdominal tergum VI in males of this genus, that of

145 the ventrally-directed flanges (Figs ). 125 MATERIAL EXAMINED: 317 specimens (PMNH, MCZC, CASC, EGRC, KJOC, HAHC, IZWP, AMNH, CNCI, JBSC, MZSP, EEMC, NMNH, OSUC, NDSÜ, BMNH, MNHP). There are approximately 20 species from Suriname, Bolivia, Panama, Brazil, Costa Rica, Mexico, Belize, Venezuela, Ecuador, Peru, Nicaragua, and Guatemala. Further type studies are necessary to determine which of these species are described. Taxonomic list - Epi-ptvoma *corvina (Champion), 1897a (Ptilodactvla) NEW COMBINATION (BMNH); Mexico *ebenina (Champion), 1897a (Ptilodactyla) NEW COMBINATION (BMNH); Central America *maculata (Champion), 1897a (Ptilodactyla) NEW COMBINATION (BMNH); Central America *niqricornis (Champion), 1897a (Ptilodactyla) NEW COMBINATION (BMNH); Central America *rufa (Champion), 1897a (Ptilodactyla) NEW COMBINATION (BMNH); Central America *rufjpennis (Pic), 1916a (Ptilodactyla) NEW COMBINATION (MNHP); Neotropics

146 Therius Therius Guerin-Meneville, 1849: 1 (type species, here designated: T. suturalis Guerin-Meneville, 1849: 2); Lacordaire, 1857: 270; Kasap and Crowson, 1975: 442; Hlavac, 1975: 182. Thervus Pic, 1932: 20 (incorrect subsequent spelling). Diagnostic combination. This genus may be recognized by the following combination of characters: tarsomere IV lobed, II-III with ventral fleshy pads, lateral pronotal marginations acute, very nearly reaching anterior margins, protrochantins visible (as in Fig. 64), and mandibles tridentate (as in Fig. 37). Description. Body size. Length mm, width at humeral angles mm. Head. Quadrate; female antennae serrate (as in Fig. 4), epistomal sulcus present; labrum (as in Fig. 47) long, basal width 2x or less greatest length, anteriorly truncate or slightly convex; mandibles (as in Fig. 37) tridentate, with prostheca, molar areas, and dorsal ridge; maxillae, lacinia blunt, not forming spore brush;'galea blunt; apical palpomere mostly sclerotized; labium, ligula with each half bluntly apexed (as in Fig. 15). Thorax. Pronotal margins incompletely acute.

147 margination very nearly meeting anterior; basal margin 127 crenulate, mediobasal lobe crenulate, with produced denticles (as in Fig. 72); notai projection short, protrochantins visible (as in Fig. 63); mesosternal excavation deep, keel absent; mesoscutellum cordiform; elytra strongly striate, sutural striae not deeply impressed; metasternal transverse suture visible near discrimen, laterally obsolete; metacoxal plates welldeveloped; tarsi, article IV not reduced, lobed, with ventral fleshy pads; ungues simple. Abdomen. Ovipositor (as in Fig. 81) long, coxites divided, articulated with bacula, styli subapical. Males unknown. MATERIAL EXAMINED: 10 specimens (MNHP), all in poor condition. Species excluded: nitidissimus Pic (Stenafricus) Taxonomic list - Therius fulvipes Guerin-Meneville, 1849; S. Africa iaspideus Fairmaire, 1878; China *luridipennis Guerin-Meneville, 1849; S. Africa perrieri Pic, 1931a; Madagascar V. bicolor Pic, 1931a; Madagascar *suturalis Guerin-Meneville, 1849; S. Africa *uniformis Pic, 1913a (MNHP); no locality

148 128 COMBINATION (MNHP); French Congo *nitidissimus (Pic)/ 1914a (Therius) NEW COMBINATION (MNHP); west central Africa

149 .29 INCERTAE SEDIS Falsoptilodactvla Pic 1958 Valoka Deleve 1972 Podabrocephalus Pic 1913 Brounia Sharp 1878 Hovactvla rufescens Pic 1913 Stenactvla basicornis Fairmaire 1901 Stenactvla lutea Pic 1946 Stenactvla ruficeps 1952 Stenactvla striata Pic 1930 EXCLUSIONS Cladotoma russula Fairmaire 1904a Cladotoma vittata Pic 1914 Drupeus Lewis 1895 Cerophytidae Pseudolichas Psephenidae; Eubriinae Pseudolichas Fairmaire 1878 Pseudodactvlus Hampe 1866 Singularodaemon Pic 1953b Dascillidae Eurypogonidae (?) Dascilloidea

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156 136 Latreiile, P. A Les crustacés, les arachnides et les insectes, distribues en famille naturelles, ouvrage formant les tomes 4 et 5 de celui de M, le Baron Cuvier sur le Régne animal (loth edition), pp. Paris. Lawrence, J. F Coleoptera, pp In, Sybil P. Parker, Synopsis and classification of living organisms, 2. and A. P. Newton Evolution and classification of beetles. Ann. Rev. Ecol. Syst. 13: LeConte, J. L Synopsis of the Atopidae, Rhipiceridae and Cyphonidae of the United States. Proc, Acad. Nat. Sci. Philadelphia 6: Classification of the Coleoptera of North America, 1. Smithson. Misc. Coll. 136: 1-208, List of the Coleoptera of North America. Smithson. Misc. Coll. 140: Descriptions of new Coleoptera chiefly from the Pacific slope of North America. Trans. Amer. Ent. Soc. 5; and G. H. Horn Classification of the Coleoptera of North America. Washington: Smith. Instit. pp Reprinted 1978 by Arno Press. A N. y. Times Co., New York. Leech, H. B. and H. G. Chandler Aquatic Coleoptera, Ch. 13, pp In, R. L. Usinger (ed.). Aquatic Insects of California. University of California. University of California, Berkeley. Legros, Clement ^Dascilloidea. Faune de l'empire Français. VII. Coléoptères des Antilles. 1: LeSage, L. and P. P. Harper Notes on the life history of the toed-winged beetle Anchvtarsus bicolor (Melsheimer) (Coleoptera: Ptilodactylidae). Coleop. Bull. 30 (3): Lewis, G On the Dascillidae and malacoderm Coleoptera of Japan. Ann. Mag. Nat. Hist. 6 (16):

157 137 Maddison, W. P., M. J, Donoghue, and D. R. Maddison Outgroup analysis and parsimony, Syst, Zool, 33(1): Melsheimer, F, E, 1845, Descriptions of new species of Coleoptera of the United States, Proc, Acad, Nat, Sci, Phil. 2(8): , von Motschulsky, ViLctor, Essai d'un catalogue des insectes de l'ile Ceylon, Bull, Soc, Imp, Nat, Moscou 36(1): Murray, A, 1868, List of Coleoptera received from Old Calabar, on the west coast of Africa, Ann, Mag, Nat, Hist, (4) 2: , Nakane, T, On the Japanese Dascillidae (Coleoptera), Bull, Takarazuka Insectarium 45: 1-16, 1950, Iconographia Insectorum Japonicorum, editio secunda, reformata, Hokuryukan, Tokyo: ,, 1952, New or little known Coleoptera from Japan and its adjacent regions, VII.,.Dascillidae,,, Sci, Rep, Saikyo Univ. (Nat, Sci, and Liv, Sci.) 1 (1): 35-41,, 1956, The beetles of Japan, 31, Shin Konchu 9 (2): 51-55,, 1963, New or little known Coleoptera from Japan and its adjacent regions, 22, Frag, Coleopt., (10-12): 42-48,, 1977, New or little known Coleoptera from Japan and its adjacent regions, 28, Fragmenta Coleopterologica 22/24: 87-98, K. Ohbayashi, S, Nomura and Y, Kurosawa, Iconographia Insectorum Japonicum, Colore naturale édita, 2, (Coleoptera), Hokuryukan, Tokyo, 443 pp. Pic, M, 1911, Coleopteres exotiques nouveaux ou peu connus (Suite,) Echange 27: , 1913a, /Coléoptères exotiques eu partie nouveaux (Suite,), Echange 29: , 1913b, Diagnoses de Dascillides et Cyphonides

158 138 / nouveaux. Echange 29: c. Nouveaux genres et nouvelles espèces de Cyphonides. Mel. Exot.- Ent. 7: d. / Coléoptères exotiques en partie nouveaux (Suite.). Echange 29: a. ^ Coléoptères exotiques en partie nouveaux (Suite.). Echange 30: b. Coléoptères divers da Tonkin et de l'indo- China. Mel. Exot.- Ent. 9: c. Dascillidae, Helodidae, Scraptiidae et Salpingidae. In, Voyage de Ch. Alluaud et R. Jeannel en Afrique orientale ( ). Résultats scientifiques Mémoire 32. Coleoptera. 9: a. Descriptions abrégées diverses. Mél. Exot.- Ent. 20: b. Descriptions abrégées diverses. Mel. Exot.- Ent. 21: a. Trois nouveaux Ptilodactvla. Bull, soc. ent. France 1916: b. Descriptions abrégées diverses. Mel. Exot.- Ent. 22: Courtes descriptions diverses. Mel. Exot.- Ent. 27: Nouveautés diverses. Mel. Exot.-Ent. 30: Nouveautés diverses. Mel. Exot.-Ent. 33: a. Nouveautés diverses. Mél. Exot.-Ent. 40: b. Nouveaux Coléoptères Dascillides de Sumatra. Tijdschr. Ent. 66: c ( %. Contribution a l'étude des Helodidae [Col.]. Échange 39 ( ): 1-12, hors texte.

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162 und Indien-Nepal-Expeditionen des Naturhistorischen Museums in Basel. Coleoptera: Pam. Georissidae, Eydraenidae, Hydrophilidae und Ptilodactylidae. Ent. Basiliensa 4: , Ptilodactylidae. Checklist of the Coleoptera of Japan. 20: 1-5. Say, T, Descriptions of coleopterous insects collected in the late expedition to the Rocky Mountains, performed by order of Mr. Calhoun, Secretary of War, under the command of Major Long. J. Acad. Nat. Sci. Phil. 3(1): Schaeffer, C New Dascyllidae. Psyche 13: Spangler, P. J The Catherwood Foundation Peruvian- Amazon Expedition XIII. Aquatic Coleoptera (Dytiscidae, Noteridae, Gyrinidae, Hydrophilidae, Dascillidae, Helodidae, Psephenidae, Elmidae). Monogr. Acad, Nat. Sci. Phil. 14: , 13 pis Coleoptera. In, Aquatic Biota of Tropical South America, Part I: Arthropoda. S, H. Hurlbert, G. Rodriguez, and N. D. Santos, eds. San Diego State University, San Diego, Cal. xiit 323 pp Coleoptera. In, Aquatic Biota of Mexico, Central America and the West Indies, S. H. Hurlbert and A. Villalobos-Figueroa, eds., San Diego State University, San Diego, California Immature stages and biology of Tetraalossa palpalis Champion (Coleoptera: Ptilodactylidae). Ent. News 94(5): Steinheil, E Symbolae ad historiam coleopterorum Argentiniae meridionalis, ossia enumerazione dei coleotteri raccolti dal prof. P. Strobel nell' Argentinia méridionale, e descrizione delle specie nuove. Attn. Soc. Italians Sci. Nat. 15: Stribling, J. B Revision of Anchvtarsus (Coleoptera: Dryopoidea) and a key to the New World genera of Ptilodactylidae. Ann. Ent. Soc. Amer. 79: Suss, L. and 0. Puppin ( ) Osservazioni sulla morfologia e sulla biologia di Ptilodactvla exotica Chapin (Coleoptera, Ptilodactylidae) nelle

163 serre della Lombardia e contribute bibliografico alio studio del gruppo (*). Bolletino Di Zoologia, Agrararia Di Bachiocultura. Milan. (Ser. 2) 13: Watrous, L. E. and Q. D. Wheeler The outgroup comparison method of character analysis, Syst. Zool. 30(1): Westwood, J Description of some new species of exotic coleopterous insects from the collection of Sir Patrick Walker. Mag. Zool. Bot. 1: White, A Note on the pupa-case of a coleopterous insect from northern China. Ann. Mag. Nat. Hist. (3) 3: White, D. S Coleoptera (Dryopoidea). pp In, Aquatic Insects of upper Three Runs Creek, Savannah River Plant, South Carolina. Pt. I: orders other than Diptera. Eds.: Morse, J. C., J. W. Chapin, D. D. Herlong, and R.S. Harvey. J. Georgia Entomol. Soc. 15: Wiley, E Phylogenetics. The Theory and Practice of Phylogenetic Systematics. John Wiley and Sons, New York. 439 pp.

164 APPENDIX A FIGURES 144

165 Fig. 1. Dorsal habitus. Anchvcteis velutina, male. 145

166 146

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