INSECTA MUNDI, Vol. 11, No.1, March, 1997 51 Classification, natural history, and evolution of the Epiphloeinae (Coleoptera: Cleridae). Part I. The genera ofEpiphloeinae. Weston Opitz Kansas Wesleyan University Department of Biology 100 East Claflin Salina, Kansas 67401 Abstract: The subfamily Epiphloeinae is defined to include fourteen genera as follows: Epiphloeus Spinola; Pilosirus, new genus; Plocamocera Spinola; Iontoclerus, new genus; Arenaria, new genus; Ichnea Laporte; Diapromeces, new genus; Pyticeroides Kuwert; Ellipotoma Spinola; J{atamyurus, new genus; Megatrachys, new genus; Madoniella Pic; Hapsidopteris, new genus; and Teutonia, new genus. The following type-species are described: Pilosirus brunoi, new species; Arenaria chiapas, new species; Diapromeces aclydis, new species; J{atamyurus paxillus, new species; Megatrachys paniculus, new species; Hapsidopteris diastenus, new species; and Teutonia nova, new species. Elloplium humerale Klug is designated as the typespec~es of IOlltoclerus. The genus Madolliella is removed from the subfamily Korynetinae andis declared a senior synonym of Phlogl.stostemus Wolcott. Neiclmea is synonymized with Pyticeroides. This treatise includes a key to the genera of Epiphloeinae, descriptions of the genera and new type-species, and distribution map for each genus. Kcy words: Clericlae, Checkered Beetles, Epiphloeinae, Generic Synopsis Introduction Ichnoides in his Tableau Synoptique des Clairi- Members of Epiphloeinae have been classi- ones. Then, in 1844, in Monographie des Terediles, fied in various subfamilies. Moreover, these Spinola added Plocamocera to the above beetles have not been studied collectively at any men tioned genera and classified the three taxonomic level. The purpose of this paper is to genera under Clerites Hydnoceroides Tableau begin a series of publications intended to clarify Generique des Clerites. At first, this classification epiphloeine relationships of classification, natural was adopted by Lacordaire (1857:421) and followed history, and to speculate about their evolution. by Desmarest (1860). Subsequently, Lacordaire The study begins with a generic synopsis which (1857:422) regrouped the epiphloeine species un- will be followed by revisions of the genera. The der Phyllobenides, a scheme of classification project will end with a proposed evolutionary adopted by Gorham (1860, 1877), Lohde (1900), history of the subfamily. Schenkling (1903, 1906, 1910), Gahan (1910), and Mimicry is extensively ingrained in the struc- Blackwelder (1945). The more modern concept of tural and behavioral evolu tion of the Cleridae. This epiphloeine classification was first introduced by has been indicated by various authors and ably Kuwert (1893), who aligned the species under summarized in a recent important work by Mawd- genera of "Epiphloinen." This classification scheme sley (1994). The mimetic character of clerids has was later refined and published by Wolcott (1947), to some extent influenced the sequence of my Corporaal (1950), Barr (1950,1962), Knull revisionary works, in that as I delved into the (1951), Arnett (1960), Winkler (1961), and taxonomic problems of one group, I invariably found Crowson (1964). Today, it is widely accepted that a preponderance of mimics of other distantly relat- the species under study belong to the subfamily ed genera mistakenly included in my request for Epiphloeinae. unsorted material. This was the circumstance Material and methods that fueled my interest in the Epiphloeinae be- This study is based on several thousand etles some of which are superficially similar to specimens and involved nearly all the nominal the lampyrid and/or lycid-like members of Perily­ pus of the subfamily Clerinae (Ekis, 1977). species now assigned to Epiphloeinae. Many beetles of other subfamilies were also examined. Literature review The specimens were borrowed from various The first published account to bring the institutions or personally field collected. Field col- epiphloeine species to taxonomic order was lected specimens were preserved in Pampel's presented by Spinola (1841), who listed Ichnea fluid (Ekis, 1977) for study of internal organs. As and Epiphloeus under the category Clairiones has been true of my previous revisionary works, 52 Volume 11, No.1, March, 1997, INSECTA MUNDI the results of this study are based in part on an that I have retained for completion of this part extensive outgroup comparison that served to of the study will be deposited in collections indicat- establish the foundation for assessments of charac- edin the text by the following abbreviations: AMNH: ter state phylogeny. Specifically, character state American Museum of Natural History, Entomol- phy logeny was surmised by implementation of the ogy, New York, New York, 10024; BMNH: six criteria for character analysis developed by me British Museum (Natural History), Entomology, in an earlier paper (Ekis, 1977: 117). Lundberg SW 5BD, London, England; CASC: California Acad- (1972), Ross (1974), and Watrous and Wheeler emy of Science, Entomology, San Francisco, Cal- (1981) have detailed the methods of outgroup ifornia, 94118; CNCI: Canadian National Collec- comparisons. tion of Insects, Entomology Research Institute, The assessment of a character state disconti- Ottawa, Ontario, Canada; FMNH: Field Museum nuity as being generic in magnitude is a highly of Natural History, Entomology, Chicago, Illinois, subjective matter. It is the character state that 60605; JNRC: Jacques Rifkind Collection, 11322 makes the genus, not the genus that makes the Camarillo St., #304, North Hollywood, California, character state. Mayr (1969) clearly summarized 91602; MCZC: Museum of Comparative Zoology this concept of the genus. Herein, I have Harvard University, Entomology, Cambridge, attempted to achieve a balance among the charac- Massachusetts,02138; MCMC: Museo de Historia ter state discontinuities judged to be generic in Natural de la Ciudad de Mexico, Apartado 18845, rank. That is, when a particular characteristic Mexico, D.F.; MNHN:Museum Nationalcl'Histoire gap (discontinuity) among the species was consid- Naturelle, Entomologie, 45 bis, Rue de Buffon, ered potentially a generic level discontinuity, I Paris (Ve), France; MZSP: Museu de Zoologia da compared the magnitude of observed difference U niversidade de Sao Paulo, Caixa postal with the character state difference among other 7172,01.05, Sao Paulo, Brazil; USNM: National genera. Specifically, to establish subfamily rank I Museum of Natural History, Smithsonian Institu- used the presence of pro notal tactile organs (Figs. tion, Entomology, Washington D.C., 20560; OSUC: 3,6), serrulate protibia (Fig. 5), and geographic The Ohio State University, Museum of Biological distribution. For defining generic rank I relied on Diversity, 1315 Kinnear Road, Columbus, Ohio, structural differences of the metatibia, antenna, 43212; WFBC: William F. Barr Collection, 1415 and male genitalia. Borah Avenue, Moscow, Idaho, 83843; WFBM: Number of articles of the antenna is an William F. Barr Museum, Department of important diagnostic characteristic for placement Entomology, University of Idaho, Moscow, Idaho, of specimens in their appropriate genera. 83844; WOPC: Weston Opitz Collection, Kansas Unfortunately, some of the antennal articles, Wesleyan University, Department of Biology, 100 especially those of the funicle (Fig. 44), are difficult East Claflin, Salina, Kansas, 67401. I am indebted to discern. These articles are usually small and to the curators of these collections who entrusted profusely setose in some species to an extent that me with material in their charge. I am particularly their anatomical limits are indistinguishable. To greatful to William F. Barr and to Charles A. solve this problem I subjected an antenna to a hot Triplehorn for various courtesies including the solution of potassium hydroxide for some 15 review of this manuscript. This research was sup- minutes, then observed the antenna under tap ported by a National Science Foundation Grant water. This treatment expands the integument (DEB 7910 962). between the antennal articles thus clearly indicating their anatomical limits. In this publica- Subfamily Diagnosis tion, I have illustrated all the important character- Species of Epiphloeinae are readily distin- istics that identify the genera of Epiphloeinae. guished from other Cleridae by the presence of 2 Illustration and dissection techniques, and the discal and 2 paralateral punctiferous and setifer- use of descriptive terms, essentially follow those ous depressions (Figs. 3, 6) on the pronotum. The used in my earlier work with Perilypus (Ekis, anterior margin of the protibia is serrulate (Fig. 1977). 5) and the fourth tarsal article of the metatarsus Most of the borrowed specimens on which this is cryptic (Fig. 129). The members of this study is based have been returned to their owners subfamily are found only in the New World. Their identified and labeled as species to be described or range extends from the United States to Central as species already described. The few specimens Argentina. INSECTA MUNDI, Vol. 11, No.1, March, 1997 53 Key to the Genera of Epiphloeinae 1. Antenna composed of 11 articles ............................. 2 12(10).Elytral surface corrugated, densely set with tuber- - Antenna composed of less than 11 articles ........ 3 c'les and setose pencils (Fig. 105) .................... .. ................................... !l1egatrachys, new genus 2(1). Antennal funicular articles approximately equal - Elytral