WUKYUB8ARY *1 WRWtLHAil KBo 71995 UNIVERSITY OF ILLINOIS LIBRARY AT URBANA-CHAMPAIGN BIOLOGY Hi 01 1995 590.5 Fl n.s. A Zoology NEW SI RIES, N;G. 67 Current Classificatio \6 Slaphyliniformia (< F. Newton, Jr. ret K. Thaver K 1992 Publication 1434 PUBLISHED BY FIELD M LSI UM OF NATURAL HISTORY .;-u'Fputer ittcrns in visions, pj ton Publishers. The inches ^22 X 2? arrangement t< ws: Pen arid ink here abbrcv> luntianum a [ED ON ACID-FREE PAPER. FIELDIANA Zoology NEW SERIES, NO. 67 Current Classification and Family-Group Names in Staphyliniformia (Coleoptera) Alfred F. Newton, Jr. Margaret K. Thayer Department of Zoology Field Museum of Natural History Chicago, Illinois 60605-2496 BIOLOGY LIBRARY Accepted April 16, 1991 W BURHUL HALL Published May 29, 1992 Publication 1434 flUG 10 199? PUBLISHED BY HELD MUSEUM OF NATURAL HISTORY ■'VS OJ © 1 992 Field Museum of Natural History Library of Congress Catalog Card Number: 91-77160 ISSN 0015-0754 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents List of Tables Abstract 1 Introduction 2 Family-Group Names and the Code 2 Methods 4 Current Classification of Staphyliniformia 5 Discussion and Summary of Changes .... 14 Diagnoses of Taxa 25 Empelinae Newton & Thayer, subfam. nov 25 Solieriinae Newton & Thayer, nom. nov. . . 27 Catalog 28 Histeridae 29 Hydrophilidae 31 Sphaeritidae 33 Synteliidae 33 Agyrtidae 33 Hydraenidae 33 Leiodidae 34 Pselaphidae 37 Ptiliidae 44 Scydmaenidae 45 Silphidae 46 Staphylinidae 46 Unplaceable Names 67 Acknowledgments 67 Literature Cited 68 Addendum 83 Index 84 1. Synopsis of current classification of Staph- yliniformia 8 2. Summary of changes from recent usage of family-group names in Staphylinifor- mia 16 in Back cover: Megarthrus pictus Motschulsky (Staphylinidae) Current Classification and Family-Group Names in Staphyliniformia (Coleoptera) Alfred F. Newton, Jr. Margaret K. Thayer Abstract The current classification of the beetle series Staphyliniformia is discussed, and a classification is presented that includes 455 taxa above the genus level (ranging in rank from subtribe through superfamily). This classification is used as the framework for a catalog of earliest use and synonyms of the valid names of these taxa, with explanatory discussions as needed. The main provisions of the International Code of Zoological Nomenclature relating to family-group names are reviewed. Application of the Code to such names in Staphyliniformia requires changes (summarized in a table) in about 60 commonly used names. A third of these needed changes have already been pointed out and implemented by others, but 39 are first noted here, required by priority (20), homonymy (2), homonymy of type genus (6), or incorrect stem formation (11, including unjustified emendations). The most significant of the changes adopted here, grouped by family, are Hydrophilidae: Acidocerina (for Helocharina); Leiodidae: Leptodirini (for Bathysciini), Platypsyllinae (for Lep- tininae), Sogdini (for Hydnobiini); Pselaphidae: Iniocyphini (for Tanypleurini), Natypleurina, nom. nov. (for Tanypleurina); Ptiliidae: Cephaloplectinae (for Limulodinae); Scydmaenidae: Cyrtoscydmini (for Euconnini, Neuraphini, or Stenichnini), Mastiginae (for Clidicinae); Staph- ylinidae: Athetini (for Callicerini), Clavilispinina, nom. nov. (for Paralispinina), Dorylophilini (for Deremini), Geostibina (for Callicerina), Glyptomina, nom. nov. (for Calocerina), Homalotini (for Bolitocharini or Gyrophaenini), Hypocyphtini (for Oligotini), Hyptiomina (for Holisina), Leptanillophilini (for Mimecitonini), Lomechusini (for Myrmedoniini or Zyrasini), Miman- ommatini (for Dorylomimini), Mycetoporini (for Bolitobiini), Paglini, nom. nov. (for Pachy- glossini), Solieriinae, nom. nov. (for Physognathinae), Strigotina (for Acrotonina), and Thora- cophorini (for Lispinini). A few required changes are not implemented for reasons discussed in the text, which include pending applications to the ICZN. Ten cases apparently involving homonymy of family-group names (but not their type genera) were discovered and are discussed. Those in which the Staphyliniformia name is junior are Leiodidae: Triarthrini Jeannel, 1962 (not Ulrich, 1930, Trilobita); Pselaphidae: Metopiini Raf- fray, 1904 (not Foerster, 1868, Hymenoptera; senior to Townsend, 1908, Diptera); Staphylin- idae: Callicerini Jacobson, 1908 (not Rondani, 1856, Diptera), Cyphini Lohse, 1974 [nom. nud.?] (not Leng, 1920, Coleoptera: Curculionidae), and Toxoderina Bernhauer & Schubert, 1911 (not Saussure, 1869, Mantodea). Those in which the Staphyliniformia name is senior are Hydrophilidae: Helocharina Orchymont, 1919 (predates Metcalf, 1 965, Homoptera [nom. nud.?]), Hydrobiini Mulsant, 1844 (predates Troschel, 1857, Mollusca); Staphylinidae: Cryptobiina Casey, 1905 (predates Hollande, 1952, Protozoa [nom. nud.?]), Steninae MacLeay, 1825 (pre- dates Fraser & Purves, 1 960, Mammalia [nom. nud.?]), and Tachininae Fleming, 1 82 1 (predates Robineau-Desvoidy, 1830, Diptera). The name Empelinae, subfam. nov. (type genus, Empelus LeConte; Staphylinidae), is made available for the first time; diagnoses are provided for this and for Solieriinae. In Pselaphidae, FIELDIANA: ZOOLOGY, N.S., NO. 67, MAY 29, 1992, PP. 1-92 the replacement name Natypleurus, nom. nov., is proposed for the genus Tanypleurus Raffray (not Steenstrup & Luetken). Many common family-group names in Coleoptera have been attributed to "Leach, 1817"; evidence is presented that they should actually be treated as "Fleming, 1 82 1 ." Relative priority of four works published by Erichson and by Heer in 1839 is discussed. Introduction The beetle series Staphyliniformia is an enor- mous group including over 57,000 named species. This is about a sixth of the order Coleoptera and nearly 4% of named living organisms. These spe- cies are currently placed in nearly 500 named groups above the generic level, ranging in rank from subtribe to superfamily. Existing classifications of this group are in many cases widely divergent from one another, and are inadequate or unsatisfactory in many respects. A fundamental problem is the lack of a clear philo- sophical basis for many of these classifications. Authors of even the most novel classifications have provided little explanation or justification for their systems. There is growing agreement that classi- fications should reflect evolution, using demon- strably monophyletic groups and reflecting a firm understanding of phylogenetic relationships. With some exceptions, existing classifications have evolved without consideration of these factors, and many probably do not represent phylogenetic re- lationships within their groups. Even when in- ferred phylogeny is used in constructing classifi- cations, there is often wide disagreement among different authors on the conclusions and their ap- plication to a classification. Progress is being made steadily in this area for some included groups, but it is not yet possible to propose a comprehensive classification of Staphyliniformia based on thor- ough phylogenetic analysis. Two other limitations of existing classifications of Staphyliniformia, and of phylogenetic studies as well, are more easily dealt with now. One is incompleteness: the last complete worldwide clas- sification is well over half a century old (Coleop- terorum Catalogus, 1910-1934; see "Methods" for full list). The intervening decades have seen a tre- mendous amount of taxonomic activity. Most works during this period have, however, included only higher taxa (i.e., down to family or subfamily level) or have been regional in scope, with many "exotic" groups not mentioned or placed and the impact of studies from other regions overlooked or not fully considered. A second limitation has been inadequate attention to correct use of family- group names by many authors, due to lack of in- formation on the names themselves or miscon- ceptions about the rules governing use of such names. The purpose of the present work is to help overcome these limitations by presenting (1) an updated worldwide classification of Staphylinifor- mia above the generic level, and (2) a review of family-group nomenclature to help determine the correct names to apply to suprageneric taxa. We hope that providing these necessary tools will fa- cilitate progress toward a new and phylogenetically based classification of Staphyliniformia as well as stability in use of names. In addition, the present paper will serve as a contribution toward the prep- aration of an official register of family-group names as recently discussed by the International Com- mission on Zoological Nomenclature (ICZN, 1990c; Savage, 1990). Family-Group Names and the Code Determining the correct names to be applied to family-group taxa is not a simple task, as others have noted (e.g., Madge, 1989; Michener, 1986; Nilssonetal., 1989;Silfverberg, 1990; Watt, 1975). The use of family-group names in zoology was not formally regulated to the degree that genus- and species-group names were until the appearance of the modern International Code of Zoological No- menclature (ICZN, 1961c, 1985a, hereafter re- ferred to as "the Code," or implied by citation of Article numbers). Family-group names under the Code are applied to taxa ranked above genus, up to and including superfamily. Each is based on a generic name, or "type genus," rather than on a taxonomic concept. The lack of clear principles guiding choice of names before 1 96 1 resulted in a certain amount of in- consistency and confusion in usage, much of it arising from the confounding of nomenclatural matters and taxonomic concepts in the application of names. Even when citations of earlier usage of a family-group name have been given, they have often referred to the cited author's concept of the group (i.e., its scope), rather than to nomenclatural proposal (or validation) of a group name; it is usu- FIELDIANA: ZOOLOGY ally impossible to tell from the citation itself which was involved. Also, new names have often been used without being noted as new. Perhaps because of such problems, catalogs such as the Coleopter- orum Catalogus and cataloging services such as the Zoological Record did not (indeed, could not) keep as careful records of family-group names as of generic and specific ones. Hence, these standard references often do not include the earliest use of each family-group name, and some names are omitted. Since the introduction of formal rules in 1961, zoologists have given increased attention to the issue of correct family-group name usage. Within Staphyliniformia, three families have benefited from recent reviews of this subject in conjunction with study at lower taxonomic levels: Histeridae (Mazur, 1984), Pselaphidae (Newton & Chandler, 1989), and Hydrophilidae (Hansen, 1990). No- menclature! review of generic names, an essential foundation for determination of correct family- group names, has also been completed for Staph- ylinidae (Blackwelder, 1952; Tottenham, 1949), European Ptiliidae (Bistrom & Silfverberg, 1979), and Silphidae and Agyrtidae (Madge, 1 980). With- in Staphyliniformia, this leaves the generic no- menclature of only Scydmaenidae, Leiodidae (sen- su lato), Staphylinidae: Scaphidiinae, and non- European Ptiliidae not yet comprehensively re- viewed. Problems in the application of generic names that are type genera of family-group names have resulted in several applications to the Inter- national Commission on Zoological Nomencla- ture (hereafter called the ICZN or the Commis- sion) to conserve or clarify those names (ICZN, 1957-1961b, 1969a-1984, 198 5b- 1991). The ex- istence of this broad foundation of generic-level work and the relative lack of study at the family- group level make an overall review of family-group nomenclature in Staphyliniformia timely. The basic task in family-group nomenclature is determination of the earliest use (author and date) and availability of individual family-group names. These data are independent of the suprageneric classification in use, and the results of such in- vestigations can be presented clearly in a straight- forward alphabetical listing (e.g., Madge, 1989; Nilsson et al., 1989; Watt, 1975). Since the ulti- mate use of such information, however, is the de- termination of the correct names to be applied to particular taxa, it seems to us much more useful to present the results in the context of a classifi- cation (e.g., Michener, 1986) in order to make the ramifications of the nomenclatural data readily apparent. The correct name to apply to a taxon may be only one of many available names that compete in priority for application to that taxon, and the correct name may change according to different concepts of the composition of the taxon (see "Current Classification of Staphyliniformia," below). The definitions of two commonly misunder- stood nomenclatural terms are worth repeating here. An available name is one that meets specified provisions of the Code (Arts. 10-20, and not ex- cluded under Art. lb). Available names can in- clude currently valid names, junior synonyms, un- justified emendations, etc. A valid name is an available name that is the correct name for a taxon at a given point in time. The validity of a name depends on taxonomic judgment (i.e., on the user's concept of a taxon, which determines which avail- able names compete to be the valid name of the taxon) as well as nomenclatural rules, such as pri- ority, that are governed by the Code. Because misconceptions about the application of family-group names are common, it seems worthwhile to highlight the principles most im- portant in determining correct family-group names. The Code should be consulted for general prin- ciples applicable to all scientific names and for additional detail and occasional exceptions to those listed here. The most basic principles are Each family-group name is based on a nominal type genus, and its placement in a classification follows that of the type genus (Art. 35c). The taxonomic concept adopted by the author or subsequent user of a name is not relevant. Each name must, when first published, be a noun in the nominative plural based on the generic name then used as valid for a genus contained in that family-group taxon (Art. llf{i)l), and must end in a latinized suffix (Art. 1 lf{i)3). The type genus need not be the present valid name for the genus (Art. 40), nor need it be the oldest genus included in the taxon (Art. 64), but it cannot be a junior homonym (Art. 39). Each name must be formed from the correct stem of the name of its type genus; if not so spelled originally, it must be corrected (keeping its orig- inal authorship and date). A name based on a misspelling or an unjustified emendation of a generic name must be corrected so as to use the original spelling of the generic name (Arts. 32c(iii), 35d). The endings -idae and -inae are to be used for family and subfamily, respectively (Art. 29a); -oidea and -ini are recommended for NEWTON & THAYER: CLASSIFICATION OF STAPHYLINIFORMIA superfamily and tribe, respectively (Recom- mendation 29A). (Subtribal endings are not mentioned in the Code, but -ina is widely ac- cepted in Coleoptera and is used here.) All names based on a given type genus are coor- dinate, i.e., take the author and date of the first available name based on that generic name, re- gardless of rank (Arts. 36a, 63a). The Principle of Priority is fundamental to appli- cation of family-group names: the correct name for a taxon is the oldest available name based on a genus included in the taxon (Art. 23). In the case of synonymous names of the same date, priority is determined by the first reviser (Art. 24) (not by, e.g., page number if in the same work). The Principle of Homonymy applies to family- group names: the same name (i.e., based on a given stem) cannot be applied to two or more different taxa (Arts. 52, 53a). A family-group name based on a generic name that is a junior homonym must be rejected (even if no family- group name has yet been based upon the senior homonym) (Art. 39). Nonhomonymous generic names differing only in their endings can, how- ever, have identical stems, potentially giving rise to homonymous family-group names. Cases in which such homonymous family-group names have been proposed must be referred to the Commission for arbitration (Art. 55b). A name published before 1 900 and available ex- cept for not being latinized is available with its original authorship and date if it has been sub- sequently latinized, generally accepted, and at- tributed to the author of the non-latinized form (Art. llflni)). A name proposed after 1 930 must be accompanied by a description or definition "that purports to differentiate the taxon," or a reference thereto, or must be a replacement name (Art. 13). For names proposed before 1931, no description or definition is required: for example, a family- group name merely formed from the stem of a generic name and used in a published work (even if only in a list or index) is thereby made avail- able (see Art. 1 2b for this and other validating indications). If a name was rejected before 1 96 1 because its type genus became recognized as a junior syn- onym, and the replacement name (based on the senior synonym) became generally accepted, the replacement name is to be maintained. A name thus adopted retains its own author and date, but for purposes of determining priority takes the precedence of the replaced name (of which it is deemed to be a senior synonym) (Art. 40b). If the author and date of the replacement name are cited, they should be given as: author, date of replacement name (date of replaced name) (Recommendation 40A). After 1960, synonymy of the type genus has no effect on the family- group name based on it. The above discussion reflects the provisions of the current Code (ICZN, 1985a). At a July 1990 meeting (ICZN, 1990c; Savage, 1990), the ICZN began to consider fundamental changes to the Code, two of which would have important implications for determination of some family-group names in Staphyliniformia. The first of these changes "will severely constrain the strict application of priority by giving heaviest weight to the criterion of current usage as the determinant for establishing the va- lidity of names" (Savage, 1990). One option dis- cussed was automatic rejection (without applica- tion to the ICZN) of a forgotten senior synonym when a junior name has been in general use; such a rule would affect about half of the priority-based changes implemented here. The second change in- volves determination of correct spelling of names, which would be "established by current usage rather than Latin grammar" (Savage, 1 990); 1 1 of the name corrections made here apparently would not be required if the proposed new rules were now in effect. We learned of the proposed changes to the Code after completing our manuscript. We are certainly sympathetic with the overall objective of the changes, namely improving the stability and ease of application of nomenclature, and with the gen- eral thrust of the proposals (ICZN, 1990c; Savage, 1 990). We can see no alternative, however, to ap- plying the current Code here, because it represents the rules in effect now and for the immediate fu- ture. Furthermore, many details of the proposed changes are not yet formulated (or at least circu- lated) and, in any case, the final provisions of a new edition of the Code could be substantially different from those now proposed, after the dis- cussion by zoologists that is part of the constitu- tional procedure for modifying the Code. Methods We began the search for earliest use of particular family-group names with the most recent world FIELDIANA: ZOOLOGY catalog, at or above the species level, for each fam- ily. With the exceptions of Histeridae (Mazur, 1984), Pselaphidae (Newton & Chandler, 1989), and Hydrophilidae (sensu lato) (Hansen, 1990), this was the relevant volumes of the Coleoptero- rum Catalogus, which were completed for all Staphyliniformia between 1910 and 1934 (Bern- hauer & Schubert, 1910-1916; Bernhauer & Scheerpeltz, 1 926; Bickhardt, 1910;Csiki, 1910a,b, 1911, 1919; Hatch, 1928, 1 929; Hetschko, 1926a,b Jeannel, 1914; Knisch, 1924; Raffray, 1911 Scheerpeltz, 1933, 1934; Schenkling, 1931a,b Zaitzev, 1910). We also consulted the Genera In- sectorum for the relatively few groups treated in that series (Bickhardt, 1916, 1917, 1918;Desneux, 1906;Fenyes, 1918, 1920, 1921b; Gardner, 1935; Raffray, 1 908). We searched the Zoological Record through Volume 127 (1990/91) for names de- scribed in years subsequent to the catalogs men- tioned above (but found that family-group names were not consistently cited there until about the 1 960s). Other sources of special value were the comprehensive but little-known list of insect fam- ily-group names in Handlirsch (1925), the com- plete list of pre-1816 Coleoptera family-group names of Watt (1975), and the catalogs of Agassiz (1846, 1847) and Lucas (1920). We examined all works cited in the above sources as apparent first uses of particular family-group names for both those names and others included in Staphyliniformia. In addition, we scanned hun- dreds of revisions, older or regional catalogs and checklists, and general handbooks and identifi- cation guides for additional names or earlier ci- tations. With the two exceptions indicated, we have seen all the works listed in "Literature Cited." We included literature available to us through January 1992 in this survey. We constructed a computerized database of all names, including exact original spelling, rank, type genus, author/year/page reference, and present taxonomic position in the classification used here (see "Current Classification of Staphyliniformia," below). We constructed a second database con- taining the full references for all family-group names, including exact publication dates as far as we could determine them. We checked the status of type genera in recent reviews of names in the genus-group for staphy- liniform families (Bistrom & Silfverberg, 1979; Blackwelder, 1952; Hansen, 1990; Madge, 1980; Mazur, 1984; Newton & Chandler, 1989; Totten- ham, 1949). We also checked standard sources (Neave, 1939, 1940, 1950; Edwards & Hopwood, 1 966; Edwards & Vevers, 1975; Zoological Record for 1965 and later) for homonymy of the names of type genera and for potential homonymy of family-group names based on distinct generic names with identical stems. We searched Hand- lirsch (1925) and a variety of recent catalogs for homonymous family-group names in insects and a number of general references for such names in other groups. Finally, we consulted all Opinions of the ICZN concerning genus-group or family- group names in Staphyliniformia (ICZN, 1957— 1961b, 1969a-1984, 1985b-1991). In a further effort to locate overlooked family- group names, as well as solicit comments on the classification used here and on potential problems resulting from implementation of the name changes required by application of the Code, we distributed copies of the first draft of this manuscript to nu- merous systematists now working on all or part of Staphyliniformia (see "Acknowledgments"). Current Classification of Staphyliniformia Classifications are often a matter of considerable disagreement among workers on a group. Staphy- liniformia is no exception, and there is no uni- versally accepted classification at present. The last complete published worldwide classification for most families appeared in the Coleopterorum Cat- alogus (1910-1934) and is now far out of date. More modern classifications exist for some fam- ilies, but most are regional in scope or otherwise incomplete. In this work, we have tried to integrate all work published to date to produce a classifi- cation representing something of a consensus of systematists at present. A true consensus does not appear to exist on many points, or is difficult for us to determine, so the resulting classification un- avoidably includes a certain number of arbitrary decisions. We are aware that no one is likely to be entirely happy with this classification, and in fact we ourselves do not endorse it as the best possible representation of Staphyliniformia at present. Fu- ture work will bring better understanding of phy- logenetic relationships among the included taxa, with resultant changes in classification. Such need- ed changes will, of course, change the application of some family-group names (as would some cur- rent variations in classification), because they will compete in priority with different sets of names from those shown here. The composition of the series Staphyliniformia NEWTON & THAYER: CLASSIFICATION OF STAPHYLINIFORMIA is reasonably well agreed upon (Crowson, 1981; Lawrence & Newton, 1982; Paulian, 1988). There is, however, no such agreement on the higher clas- sification (superfamily and family) within this group. The families are commonly organized into either two superfamilies, Hydrophiloidea and Staphylinoidea (e.g., Lawrence & Newton, 1982), or three superfamilies, with Histeroidea separated from Hydrophiloidea (e.g., Crowson, 1981; Pauli- an, 1988). One family (Hydraenidae) is included in either Hydrophiloidea or Staphylinoidea about equally often. Family concepts vary enormously: Hydrophilidae, Leiodidae, and Staphylinidae in the broadest senses are divided into up to 6, 5, and 10 families, respectively, in some classifica- tions. Similar but generally less drastic disagree- ments may be found at lower taxonomic levels. The choice of a classification to use in a general review such as this is therefore difficult. We are among those presently involved in trying to clarify phylogenetic relationships and develop a new clas- sification of Staphyliniformia, in our case with spe- cial emphasis on Staphylinidae sensu lato (AFN, MKT) and Leiodidae sensu lato (AFN). With rel- atively few exceptions (nearly all being tribes or subtribes included in Pselaphidae or the staphy- linid subfamily Aleocharinae), one or both of us has examined representatives of all of the family- group taxa currently recognized in Staphylinifor- mia. In cases of ambiguity or lack of consensus in the ranking and taxonomic concepts that have been applied to particular taxa, our choice has been guided by our knowledge of the taxa in question. However, the present work is not an appropriate vehicle for introducing changes to existing taxo- nomic concepts or classification, and we refrain from doing so. Our overall framework is the superfamily and family classification of Lawrence & Newton (1982). At lower taxonomic levels, when we have found no clear consensus in current classifications, we have tended to recognize subfamilies and many tribes in a broad rather than narrow sense. How- ever, at the lowest levels (tribe or subtribe), we have tried to be comprehensive in listing as valid all family-group taxa that have been proposed and not subsequently formally synonymized. Within some groups, such as Leptodirini (Leiodidae; = Bathysciini), Euaesthetinae and Scaphidiinae (both Staphylinidae), and Agyrtidae, fairly elaborate classifications have been proposed that are not now in general use. In other cases, new family- group taxa have been used only once and never mentioned again in the literature. It is often not clear to us if these taxa have been explicitly re- jected, simply overlooked, or never restudied. In such cases we have included these taxa as valid groups, in the hope that highlighting their exis- tence will encourage specialists on these groups to deal with them in some way. Specific comments on the classification we have adopted for each family are presented below, in the sequence in which the families appear in our catalog (alphabetically within each superfamily). A synopsis of the classification used here is pre- sented in Table 1 , arranged as in the catalog, sub- taxa being listed alphabetically within each taxon. Hydrophiloidea HlSTERIDAE The recent worldwide classification of Mazur (1984) is used, except that Hetaeriinae are not di- vided into tribes (Helava et al., 1985). Yelamos and Ferrer (1988) modified Mazur's classification by dropping all taxa one rank (without adequate justification, in our view). Hydrophilidae The worldwide classification of Knisch (1924) is adopted here, except for the removal of Hy- draeninae and Limnebiinae to Hydraenidae (e.g., Perkins, 1980) and the addition of Georissidae as a subfamily of Hydrophilidae (Emden, 1956). A few subfamilies of Knisch (1924) have been treat- ed by some later authors (e.g., Crowson, 1955, 1981; Hansen, 1987) as families, and some of his tribes of Hydrophilinae as subfamilies; this treat- ment has not been universally adopted, however (e.g., Lawrence & Newton, 1982), and is not fol- lowed here. Hansen (1990, in prep.) will propose a new classification of this family, but because this is not yet formally published we do not adopt it here. We have, however, made use of data in- cluded in Hansen (1990) regarding published fam- ily-group names. (See Addendum, p. 83.) Staphylinoidea Agyrtidae Separate family status for this group, generally treated as a subgroup of Silphidae (e.g., Hatch, FIELDIANA: ZOOLOGY 1928, 1957; Arnett, 1963, 1985; Madge, 1980). has been advocated recently (Lawrence, 1982; Lawrence & Newton, 1982; Anderson & Peck, 1985; Peck, 1990). Jeannel (1936) placed the gen- era of this family, along with some extraneous genera, in four tribes, one of which (Necrophilini) is evidently a nomen nudum. The internal orga- nization of the group has not been reviewed re- cently, and tribes have not been used in works treating the group as a family (Anderson & Peck, 1985; Peck, 1990). Hydraenidae The classification proposed by Perkins (1980) as extended by Hansen (1991) is used. its relationships, we maintain Pselaphidae here as a separate family. Ptiliidae Seevers and Dybas (1943) advocated removal of the subfamily Cephaloplectinae (as Limulodi- nae) to form a separate family, but the return of this family to Ptiliidae has been adopted in some recent classifications (Crowson, 1981; Lawrence, 1982; Lawrence & Newton, 1982). The internal organization of the family used here is, except for the addition of Cephaloplectinae, that discussed by Dybas (1976), who indicated the artificial na- ture of the existing formal groups but did not pro- pose a new system. Leiodidae This family is used here in the broad sense of Crowson (1981), Lawrence & Newton (1982), and Peck (1990) to include groups often placed in up to five separate families (e.g., Jeannel, 1936) or included in the old broad concept of Silphidae (e.g., Jeannel, 1914; Hatch, 1928). The internal organization has not been completely reviewed re- cently. The classification and ranking of taxa at the subfamilial and tribal levels is adapted from Peck (1973, Cholevinae; 1990, Catopocerinae, Cholevinae, Leiodinae), Wheeler (1984, Leiodi- nae), Newton (1985, Camiarinae), and Perkovsky ( 1 99 1 , Sogdini). Tribal and subtribal groups with- in Cholevinae are also adapted in part from Jean- nel (1936) and Perreau (1989) except for Lepto- dirini, which is modified from Gueorguiev (1974, 1976). SCYDMAENIDAE The classification of this family has not been recently reviewed at the world level. The internal classification adopted here is compiled from the systematic literature, including Csiki (1919), Ar- nett (1963), and recent work of Franz (1985, 1989, 1990). Silphidae The restricted concept and the classification of Anderson and Peck (1985) and Peck (1990) are used, including the removal of some subfamilies or tribes to form a separate family Agyrtidae (qv.) and the removal of other groups to the families Leiodidae or Staphylinidae (Newton, 1975, 1985; Lawrence & Newton, 1982). Pselaphidae The classification follows Newton and Chandler (1989) and Coulon (1989), as modified by Be- suchet (1987, 1991). This family has recently been formally combined or linked with certain parts of the family Staphylinidae (Naomi, 1985; Thayer, 1987). Inclusion of Pselaphidae in Staphylinidae will require some modification of internal orga- nization of the group, because two "extra" ranks are in use in Pselaphidae (between family and sub- family and between subfamily and tribe). For this reason, and pending more conclusive evidence of Staphylinidae There is very wide disagreement on the limits of this family as well as on the composition and ranking of subordinate taxa (e.g., Lohse, 1964; Moore, 1964; Coiffait, 1972; Tikhomirova, 1973; Lawrence & Newton, 1982; Naomi, 1985; Newton & Thayer, 1988). The traditional family Staphy- linidae has been broken into 3-10 families by some authors (Coiffait, 1972; Naomi, 1985) but is still recognized in the broad sense by the others cited above. Three relatively small taxa frequently re- garded as separate families are treated as staphy- linid subfamilies here because strong arguments NEWTON & THAYER: CLASSIFICATION OF STAPHYLINIFORMIA Table 1. Synopsis of current classification of Staphyliniformia. Subtaxa are listed alphabetically within each taxon. STAPHYLINIFORMIA Latreille, 1802 HYDROPHILOIDEA Latreille, 1802 HISTERIDAE Gyllenhal, 1808 HISTEROMORPHAE Gyllenhal, 1808 DENDROPHILINAE Reitter, 1909 ANAPLEINI Olexa, 1982 BACANIINI Kryzhanovskij, 1976 DENDROPHILINI Reitter, 1909 PAROMALINI Reitter, 1909 HETAERIINAE Marseul, 1857 HISTERINAE Gyllenhal, 1808 EXOSTERNINI Bickhardt, 1914 HISTERINI Gyllenhal, 1808 HOLOLEPTINI Hope, 1840 OMALODINI Kryzhanovskij, 1972 PLATYSOMATINI Bickhardt, 1914 ONTHOPHILINAE MacLeay, 1819 TRIBALINAE Bickhardt, 1914 SAPRINOMORPHAE Blanchard, 1845 ABRAEINAE MacLeay, 1819 ABRAEINI MacLeay, 1819 ACRITINI Wenzel, 1944 ACRITOMORPHINI Wenzel, 1944 PLEGADERINI Portevin, 1929 TERETRIINI Bickhardt, 1914 CHLAMYDOPSINAE Bickhardt, 1914 NIPONIINAE Fowler, 1912 SAPRININAE Blanchard, 1845 TRYPANAEINAE Marseul, 1857 TRYPETICINAE Bickhardt, 1914 HYDROPHILIDAE Latreille, 1802 EPIMETOPINAE Zaitzev, 1908 GEORISSINAE Laporte, 1840 HELOPHORINAE Leach, 1815 HYDROCHINAE Thomson, 1859 HYDROPHILINAE Latreille, 1802 AMPHIOPINI Kuwert, 1890 BEROSINI Mulsant, 1844 CHAETARTHRIINI Bedel, 1881 (1844) HYDROBIINI Mulsant, 1844 ACIDOCERINA Zaitzev, 1908 HYDROBIINA Mulsant, 1844 HYDROPHILINI Latreille, 1802 SPERCHEINAE Erichson, 1837 SPHAERIDIINAE Latreille, 1802 CERCYONINI Horn, 1890 MEGASTERNINI Mulsant, 1844 OMICRINI Smetana, 1975 RYGMODINI Orchymont, 1916 SPHAERIDIINI Latreille, 1802 SPHAERITIDAE Shuckard, 1839 SYNTELIIDAE Lewis, 1882 STAPHYLINOIDEA Latreille, 1802 AGYRTIDAE Thomson, 1859 AGYRTINI Thomson, 1859 LYROSOMATINI Horn, 1880 PTEROLOMATINI Thomson, 1862 HYDRAENIDAE Mulsant, 1844 HYDRAENINAE Mulsant, 1844 HYDRAENIDINI Perkins, 1980 HYDRAENINI Mulsant, 1844 HYDRAENINA Mulsant, 1844 LIMNEBIINA Mulsant, 1844 OCHTHEBIINAE Thomson, 1859 LEIODIDAE Fleming, 1821 CAMIARINAE Jeannel, 1911 AGYRTODINI Jeannel, 1936 CAMIARINI Jeannel, 1911 NEOPELATOPINI Jeannel, 1962 CATOPOCERINAE Hatch, 1927 (1880) CATOPOCERINI Hatch, 1927 (1880) GLACICAVICOLINI Westcott, 1968 CHOLEVINAE Kirby, 1837 ANEMADINI Hatch, 1928 ANEMADINA Hatch, 1928 EOCATOPINA Jeannel, 1936 NEMADINA Jeannel, 1936 PARACATOPINA Jeannel, 1936 CHOLEVINI Kirby, 1837 CATOPINA Chaudoir, 1845 CHOLE VINA Kirby, 1837 EUCATOPINI Jeannel, 1921 LEPTODIRINI Lacordaire, 1854 (1849) ANTROHERPONINA Jeannel, 1910 BATHYSCIINA Horn, 1880 BATHYSCIOTINA Gueorguiev, 1974 GHIDINIINA Gueorguiev, 1974 LEPTODIRINA Lacordaire, 1854 (1849) PHOLEUINA Reitter, 1886 PLATYCHOLEINA Horn, 1880 SPELAEOBATINA Gueorguiev, 1974 ORITOCATOPINI Jeannel, 1936 PTOMAPHAGINI Jeannel, 1911 PTOMAPHAGIN A Jeannel, 1911 PTOMAPHAGININA Szymczakowski, 1964 COLONINAE Horn, 1880 (1859) FIELDIANA: ZOOLOGY Table 1. Continued. LEIODINAE Fleming, 1821 AGATHIDIINI Westwood, 1838 ESTADIINI Portevin, 1914 LEIODINI Fleming, 1821 PSEUDOLIODINI Portevin, 1926 SCOTOCRYPTINI Reitter, 1884 SOGDINI Lopatin, 1961 SOGDINA Lopatin, 1961 TRIARTHRINA Jeannel, 1962 PLATYPSYLLINAE Ritsema, 1869 PSELAPHIDAE Latreille, 1802 BRACHYSCELIA Raffray, 1890 BATRISINAE Reitter, 1882 AMAUROPINI Jeannel, 1948 BATRISINI Reitter, 1882 AMBICOCERINA Leleup, 1970 BATRISINA Reitter, 1882 LEUPELIIN A Jeannel, 1954 STILIPALPINA Jeannel, 1954 METOPIASINI Raffray, 1904 EUPLECTINAE LeConte, 1861 EUPLECTOMORPHI LeConte, 1861 EUPLECTINI LeConte, 1861 ACETALIINA Jeannel, 1958 BIBLOPLECTIN A Jeannel, 1959 BIBLOPORELLINA Jeannel, 1952 BIBLOPORINA Park, 1951 CHRESTOMERINA Jeannel, 1962 EUPLECTINA LeConte, 1861 PANAPHANTINA Jeannel, 1950 RHINOSCEPSINA Bowman, 1934 TRIMIINA Bowman, 1934 TRIMIODYTINA Jeannel, 1964 PTERACMINI Jeannel, 1962 RAFFRAYIINI Jeannel, 1949 TROGASTRINI Jeannel, 1949 MITRAMETOPINA Park, 1952 PHTEGNOMINA Park, 1951 RHEXIINA Park, 1951 TRISIGNINA Park & Schuster, 1955 TROGASTRINA Jeannel, 1949 JUBOMORPHI Raffray, 1904 JUBINI Raffray, 1904 FARONINAE Reitter, 1882 BYTHINOPLECTINI Schaufuss, 1890 BYTHINOPLECTINA Schaufuss, 1890 PYXIDICERIN A Raffray, 1904 DIMERINI Raffray, 1908 FARONINI Reitter, 1882 MAYETIINI Winkler, 1925 GONIACERINAE Reitter, 1882 (1872) BRACHYGLUTOMORPHI Raffray, 1904 ARNYLLIINI Jeannel, 1952 BRACHYGLUTINI Raffray, 1904 BARADINA Park, 1951 BRACHYGLUTINA Raffray, 1904 DECARTHRINA Park, 1951 EUPSENIINA Park, 1951 HALORABYXINA Leleup, 1969 PSELAPTINA Park, 1976 GONIACEROMORPHI Reitter, 1882 (1872) BYTHININI Raffray, 1890 BYTHININA Raffray, 1890 MACHAERITINA Jeannel, 1950 XENOBYTHINA Jeannel, 1950 GONIACERINI Reitter, 1882 (1872) INIOCYPHINI Park, 1951 DALMODINA Park, 1951 GLOBINA Jeannel, 1959 INIOCYPHINA Park, 1951 NATYPLEURINA Newton & Thayer, nom. nov. PYGOXYINI Reitter, 1909 TRICHONYCHINI Reitter, 1882 VALDINI Park, 1953 PROTEROMORPHI Jeannel, 1949 IMIRINI Jeannel, 1949 PROTERINI Jeannel, 1949 TYCHOMORPHI Raffray, 1904 SPELEOBAMINI Park, 1951 TYCHINI Raffray, 1904 MACROSCELIA Raffray, 1890 CLAVIGERINAE Leach, 1815 CLAVIGERINI Leach, 1815 COLILODIONINI Besuchet, 1991 TIRACERINI Besuchet, 1986 PSELAPHINAE Latreille, 1802 CTENISTOMORPHI Blanchard, 1845 ATTAPSENIINI Bruch, 1933 CEOPHYLLINI Park, 1951 CHALCOPLECTINI Oke, 1925 CTENISTINI Blanchard, 1845 ODONTALGINI Jeannel, 1949 PACHYGASTRODINI Leleup, 1969 PET ANOPINI Jeannel, 1954 SCHISTODACTYLINI Raffray, 1890 SOMATIPIONINI Jeannel, 1949 TMESIPHORINI Jeannel, 1949 TYRINI Reitter, 1882 CENTROPHTHALMINA Jeannel, 1949 HAMOTINA Park, 1951 JANUSCULINA Cerruti, 1970 TYRINA Reitter, 1882 NEWTON & THAYER: CLASSIFICATION OF STAPHYLINIFORMIA Table 1. Continued. CYATHIGERIMORPHI Schaufuss, 1872 BARROSELLINI Leleup, 1973 CYATHIGERINI Schaufuss, 1872 HYBOCEPHALINI Raffray, 1890 MACHADOINI Jeannel, 1951 PSELAPHOMORPHI Latreille, 1802 ARHYTODINI Raffray, 1890 PHALEPSINI Jeannel, 1949 PSELAPHINI Latreille, 1802 PTILIIDAE Erichson, 1845 ACROTRICHINAE Reitter, 1909 (1856) CEPHALOPLECTINAE Sharp, 1883 NANOSELLINAE Barber, 1924 PTILIINAE Erichson, 1845 SCYDMAENIDAE Leach, 1815 MASTIGINAE Fleming, 1821 CLIDICINI Casey, 1897 LEPTOMAST ACINI Casey, 1897 MASTIGINI Fleming, 1821 SCYDMAENINAE Leach, 1815 ASCYDMINI Casey, 1897 CEPHENNIINI Reitter, 1882 CHEVROLATIINI Reitter, 1882 CYRTOSCYDMINI Schaufuss, 1889 EUTHEIINI Casey, 1897 LEPTOSCYDMINI Casey, 1897 PLAUMANNIOLINI Costa Lima, 1962 SCYDMAENINI Leach, 1815 SIAMITINI Franz, 1989 SYNDICINI Csiki, 1919 SILPHIDAE Latreille, 1807 NICROPHORINAE Kirby, 1837 SILPHINAE Latreille, 1807 ST APHYLINIDAE Latreille, 1802 ALEOCHARINAE Fleming, 1821 ACTOCHARINI Bernhauer & Schubert, 1911 ALEOCHARINI Fleming, 1821 ALEOCHARINA Fleming, 1821 COMPACTOPEDIINA Kistner, 1970 HODOXENINA Kistner, 1970 ATHETINI Casey, 1910 ATHETINA Casey, 1910 COPTOTERMOECIINA Kistner & Pasteels, 1970 GEOSTIBINA Seevers, 1978 MICROCEROXENINA Kistner, 1970 NASUTIPHILINA Kistner, 1970 SCHISTOGENIINA Fenyes, 1918 STRIGOTINA Casey, 1910 TAXICERINA Lohse, 1989 TERMITOTELINA Kistner, 1970 THAMIARAEINA Fenyes, 1921 AUTALIINI Thomson, 1859 COROTOCINI Fenyes, 1918 ABROTELINA Seevers, 1957 COROTOCINA Fenyes, 1918 EBURNIOGASTRINA Jacobson et al., 1986 NASUTITELLINA Jacobson et al., 1986 SPHURIDAETHINA Pace, 1988 TERMITOCHARIN A Seevers, 1957 TERMITOCUPIDINA Jacobson et al., 1986 TERMITOGASTRINA Bernhauer & Scheerpeltz, 1926 TERMITOICEINA Jacobson et al., 1986 TERMITOPITHINA Jacobson et al., 1986 TERMITOPTOCHINA Fenyes, 1921 TIMEPARTHENINA Fenyes, 1921 CREMATOXENINI Mann, 1921 DEINOPSINI Sharp, 1883 DIESTOTINI Mulsant & Rey, 1871 DIGLOTTINI Jacobson, 1909 DIGRAMMINI Fauvel, 1900 DIMONOMERINI Cameron, 1933 DORYLOPHILINI Fenyes, 1921 DREPANOXENINI Kistner & Watson, 1972 ECITOCHARINI Seevers, 1965 ECITOGASTRINI Fenyes, 1918 EUSTENIAMORPHINI Bernhauer & Scheer- peltz, 1926 FALAGRIINI Mulsant & Rey, 1873 FELDINI Kistner, 1972 GYMNUSINI Heer, 1839 HETEROTAXINI Fenyes, 1921 HOMALOTINI Heer, 1839 BOLITOCHARINA Thomson, 1859 DINARDOPSINA Bernhauer & Scheerpeltz, 1926 GYROPHAENINA Kraatz, 1856 HOMALOTINA Heer, 1839 OXYPODININA Fenyes, 1918 SILUSINA Fenyes, 1918 HOPLANDRIINI Casey, 1910 HYGRONOMINI Thomson, 1859 HYGRONOMINA Thomson, 1859 SAPHOGLOSSINA Bernhauer & Scheer- peltz, 1926 HYPOCYPHTINI Laporte, 1835 LEPTANILLOPHILINI Fenyes, 1918 LABIDOPULLINA Jacobson & Kistner, 1991 10 FIELDIANA: ZOOLOGY Table 1. Continued. LEPTANILLOPHILINA Fenyes, 1918 MIMECITINA Bernhauer & Scheerpellz, 1926 MIMONILLINA Bemhauer & Scheerpeltz, 1926 LEUCOCRASPEDINI Fenyes, 1921 LOMECHUSINI Fleming, 1821 LOMECHUSINA Fleming, 1867 MYRMEDONIINA Thomson, 1867 TERMITONDINA Seevers, 1957 TERMITOZYRINA Seevers, 1957 MASURIINI Cameron, 1939 MESOPORINI Cameron, 1959 MIMANOMMATINI Wasmann, 1912 MYLLAENINI Ganglbauer, 1895 OXYPODIN1 Thomson, 1859 APHYTOPODINA Bernhauer & Scheer- peltz, 1926 BLEPHARHYMENINA Klimaszewski & Peck, 1986 DINARDINA Mulsant & Rey, 1873 MEOTICINA Seevers, 1978 OCYUSINA Mulsant & Rey, 1874 OXYPODINA Thomson, 1859 PHLOEOPORINA Thomson, 1859 TACHYUSINA Thomson, 1859 PAGLINI Newton & Thayer, nom. nov. PARADOXENUSINI Bruch, 1937 PEDICULOTINI Adam, 1987 PHILOTERMITINI Seevers, 1957 PLACUSINI Mulsant & Rey, 1871 PHYLLODINARDINI Wasmann, 1916 PHYTOSINI Thomson, 1867 PRONOMAEINI Mulsant & Rey, 1873 PSEUDOPERINTHINI Cameron, 1939 PYGOSTENINI Fauvel, 1899 SCEPTOBIINI Seevers, 1978 SKATITOXENINI Kistner & Pasteels, 1969 TERMITODISCINI Wasmann, 1904 TERMITOHOSPITINI Seevers, 1941 HETA1ROTERMITINA Seevers, 1957 TERMITOHOSPITINA Seevers, 1941 TERMITONANNINI Fenyes, 1918 PERINTHINA Bernhauer & Scheerpeltz, 1926 TERMITONANNINA Fenyes, 1918 TERMITOPAEDIINI Seevers, 1957 TERMITUSINI Fenyes, 1918 TERMITOSPECTRINA Seevers, 1957 TERMITUSINA Fenyes, 1918 TRICHOPSENIINI LeConte & Horn, 1883 TRILOB1TIDEINI Fauvel. 1899 APATET1CINAE Fauvel, 1895 APHAENOSTEMM1NAE Peyerimhoff, 1914 DASYCERINAE Reiner, 1887 EMPELINAE Newton & Thayer, subfam. nov. EUAESTHETINAE Thomson, 1859 ALZADAESTHETINI Scheerpeltz, 1974 AUSTROESTHETINI Cameron, 1944 EUAESTHETINI Thomson, 1859 FENDERIINI Scheerpeltz, 1974 NORDENSKIOLDIINI Bernhauer & Schu- bert, 1911 STENAESTHETINI Bernhauer & Schubert, 1911 HABROCERINAE Mulsant & Rey, 1877 LEPTOTYPHLINAE Fauvel, 1874 CEPHALOTYPHLINI Coiffait, 1963 ENTOMOCULIINI Coiffait, 1957 LEPTOTYPHLINI Fauvel, 1874 METROTYPHLINI Coiffait, 1963 NEOTYPHLINI Coiffait, 1963 MEGALOPSIDIINAE Leng, 1920 MICROPEPLINAE Leach, 1815 NEOPHONINAE Fauvel, 1905 OLISTHAERINAE Thomson, 1859 OMALIINAE MacLeay, 1825 ANTHOPHAGINI Thomson, 1859 ARPEDIOMIMINI Cameron, 1917 CORNEOLABIINI Steel, 1950 CORYPHIINI Jacobson, 1908 BOREAPHILINA Zerche, 1990 CORYPHIINA Jacobson, 1908 EUSPHALERINI Hatch, 1957 GLYPHOLOMATINI Jeannel, 1962 HADROGNATHINI Portevin, 1929 MICRALYMMINI Mulsant & Rey, 1880 MICROSILPHINI Crowson, 1950 OMALIINI MacLeay, 1825 TETRADELINI Fauvel, 1904 OSORIINAE Erichson, 1839 ELEUSININI Sharp, 1887 LEPTOCHIRINI Sharp, 1887 OSORIINI Erichson, 1839 OSORIINA Erichson, 1839 PAROSORIINA Bernhauer & Schubert, 1911 THORACOPHORINI Reiner, 1909 CLAVILISPININA Newton & Thayer, nom. nov. GLYPTOMINA Newton & Thayer, nom. nov. LISPININA Bernhauer & Schubert, 1910 THORACOPHORIN A Reiner, 1909 NEWTON & THAYER: CLASSIFICATION OF STAPHYLINIFORMIA 11 Table 1. Continued. OXYPORINAE Fleming, 1821 OXYTELINAE Fleming, 1821 COPROPHILINI Heer, 1839 OXYTELINI Fleming, 1821 PAEDERINAE Fleming, 1821 PAEDERINI Fleming, 1821 ACANTHOGLOSSINA Coiffait, 1982 ASTENINA Hatch, 1957 CRYPTOBIINA Casey, 1905 CYLINDROXYSTINA Bierig, 1943 DOLICAONINA Casey, 1905 ECHIASTERINA Casey, 1905 LATHROBIINA Laporte, 1835 LITHOCHARINA Casey, 1905 MEDONINA Casey, 1905 PAEDERINA Fleming, 1821 SCOPAEINA Mulsant & Rey, 1878 STILICINA Casey, 1905 STILICOPSINA Casey, 1905 PINOPHILINI Nordmann, 1837 PINOPHILINA Nordmann, 1837 PROCIRRINA Bernhauer & Schubert, 1912 PHLOEOCHARINAE Erichson, 1839 PIESTINAE Erichson, 1839 PROTEININAE Erichson, 1839 ANEPIINI Steel, 1966 NESONEINI Steel, 1966 PROTEININI Erichson, 1839 PSEUDOPSINAE Ganglbauer, 1895 SCAPHIDIINAE Latreille, 1807 CYPARIINI Achard, 1924 HETEROSCAPHINI Achard, 1914 SCAPHIDIINI Latreille, 1807 CERAMBYCISCAPHINA Pic, 1915 DIATELIINA Achard, 1924 SCAPHIDIINA Latreille, 1807 SCAPHIINA Achard, 1924 SCAPHISOMATINI Casey, 1894 BAEOCERIDIINA Achard, 1924 BAEOCERINA Achard, 1924 SCAPHISOMATINA Casey, 1894 TOXIDIINI Achard, 1924 SOLIERIINAE Newton & Thayer, nom. nov. ST APHYLININAE Latreille, 1802 DIOCHINI Casey, 1906 OTHIINI Thomson, 1859 PLATYPROSOPINI Lynch, 1884 STAPHYLININI Latreille, 1802 AMBLYOPININA Seevers, 1944 CRASPEDOMERINA Bernhauer, 1911 HYPTIOMINA Casey, 1906 PHILONTHINA Kirby, 1837 QUEDIINA Kraatz, 1857 STAPHYLININA Latreille, 1802 TANYGNATHININA Reitter, 1909 TRIACRINA Bernhauer, 1931 XANTHOPYGINA Sharp, 1884 XANTHOLININI Erichson, 1839 STENINAE MacLeay, 1825 TACHYPORINAE MacLeay, 1825 CORDOBANINI Bernhauer, 1910 DEROPINI Smetana, 1983 MEGARTHROPSINI Cameron, 1919 MYCETOPORINI Thomson, 1859 SYMMIXINI Bernhauer, 1915 TACHYPORINI MacLeay, 1825 VATESINI Seevers, 1958 TRICHOPHYINAE Thomson, 1859 TRIGONURINAE Reiche, 1865 relating them to particular groups of staphylinids have been made recently and they have been for- mally included in Staphylinidae by some authors: Scaphidiidae (Kasule, 1966; Crowson, 1981; Law- rence & Newton, 1982), Dasyceridae (Thayer, 1 987), and Micropeplidae (e.g., Ganglbauer, 1 895; Bernhauer & Schubert, 1910; Arnett, 1963, 1985; Lohse, 1964; Campbell, 1968; Thayer, 1987). Pse- laphidae have also been combined with a part of Staphylinidae sensu lato in one recent study (Na- omi, 1985), but the disputed relationships, large size, and complex hierarchical classification of this group have led us to retain pselaphids as a separate family for the time being. The system of staphylinid subfamilies recog- nized here, we feel, approaches a consensus among those recent specialists using a relatively broad traditional concept of this family, and differs most from the deliberately artificial system of Moore ( 1 964) used in some North American works (Moore & Legner, 1979; Arnett, 1985). Two monotypic subfamilies, Empelinae and Solieriinae, have ap- peared in the literature but lack available names; these are treated in a later section. The larger sub- families with much internal classification, and a few others, are discussed individually below. Aleocharinae— Used here in the broad sense of Hammond (1975) to include those staphylinids that share a uniquely derived complex aedeagal structure. Some of the included taxa are com- 12 FIELDIANA: ZOOLOGY monly treated as separate subfamilies, notably Trichopseniini (e.g., Pasteels & Kistner, 1971; Seevers, 1978) and Hypocyphtini (e.g., Arnett, 1 963; Lohse, 1 974). Seevers (1978) made the most comprehensive recent attempt to deal with the internal classification of the subfamily, and we have generally followed his arrangement. However, his classification is problematic in several ways, in- cluding the absence of numerous non-Nearctic taxa and the use of many new subtribal names that are nomina nuda. Also, he organized the Nearctic tribes into a new system of supertribes, most of which appear only as headings in a checklist and cannot be applied readily to a world classification; these supertribes are therefore not used here. The tribal and subtribal classification used here is thus based on Seevers (1978) but heavily modified and aug- mented by reference to Ashe ( 1 99 1 ); Fenyes (1918, 1920, 1921a,b); Hammond (1975); numerous works of Kistner, Pasteels, and coworkers (see "Literature Cited," plus others); Jacobson et al. (1986); Jacobson and Kistner (1991); Klima- szewski (1982); Klimaszewski and Peck (1986); Lohse (1974, 1989); Lohse et al. (1990); Muona (1979); Pace (1989); and Seevers (1957, 1965), as well as the Coleopterorum Catalogus (Bernhauer & Scheerpeltz, 1926; Scheerpeltz, 1934). A num- ber of subtribes recently proposed by Seevers ( 1 978) and Muona ( 1 979) are nomina nuda. We list these as synonyms under the appropriate tribes; in the case of coordinate names subsequently made available, we list the nomina nuda under the avail- able names. It is possible that we have overlooked additional family-group taxa, as well as some syn- onymies and changes in rank, in this most difficult and poorly known staphylinid subfamily. Euaesthetinae— After Scheerpeltz ( 1 974). This tribal system, which did not place all known gen- era, has not been discussed or adopted by others, and needs critical review. Coiffait (1984) charac- terized a tribe Tamotini by the supposedly 3-seg- mented tarsi of Tamotus, but Puthz (1973) had already shown that Tamotus has 4-segmented tarsi and placed it in Euaesthetini. Leptotyphlinae— After Coiffait (1963, 1972). Omaliinae— No complete recent review; tribes are not universally used and many genera have never been formally placed in tribes. The system of tribes and subtribes used here is adapted from Bernhauer and Schubert (1910), Hatch (1957), Newton (1975, 1985), Portevin (1929), Scheer- peltz (1933), Thayer and Newton (1979), and Zerche (1990). We follow Hatch (1957), Ham- mond (1971), and Thayer (1985) rather than Ar- nett (1963, 1985) or Watanabe (1990) in place- ment of Brathinus. Osorhnae— Modified from Blackwelder ( 1 942) by elevation to tribal status of Eleusinini (treated as the subfamily Eleusiinae [sic] by Watt & McColl, 1982). Ox ytelinae— After Herman's (1970) world re- view, as slightly modified by Newton (1982b). In an overlooked work, Scheerpeltz ( 1 944) proposed a new tribe for the new genus Trigonobregma; the genus appears to belong in Coprophilini of Her- man (1970). Paederinae— No complete recent review. The classification is adapted from Arnett (1963), Blackwelder (1944), Bordoni (1975), Hatch (1957), and Herman (1981, 1991). Phloeocharinae— Concept of Herman (1972); excludes Olisthaerus (to subfamily Olisthaerinae; e.g., Lohse, 1964) and Derops (to Tachyporinae, Smetana, 1983). Piestinae— Used in a restricted sense, excluding the former tribes Trigonurini and Apateticini (re- moved to Silphidae by some authors, including Madge, 1980; here treated as staphylinid subfam- ilies), other tribes or subtribes removed to Oso- riinae by Blackwelder ( 1 942), and genera removed to Phloeocharinae (Herman, 1972) and Pseudop- sinae (Newton, 1982a). Proteininae— After Steel (1966). Pseudopsinae— Concept of Newton (1982a). Scaphidiinae— After Achard (1924), modified by generic placements of Lobl (1971) and later generic synonymies (Lobl, 1977, 1987). Achard's system has not been fully used or reviewed by Lobl or other recent workers on this group. Staphylininae— Used in the broad sense (Blackwelder, 1943;Kasule, 1966), including Xan- tholininae of many authors and six subfamilies of Moore (1964). The tribes grouped as Xantholi- ninae by Coiffait (1972) and Smetana (1982) are here treated as tribes of Staphylininae. The re- maining staphylinine family-group taxa are in need of comprehensive review; they are treated here as subtribes of Staphylinini (Blackwelder, 1944; Hatch, 1957; Arnett, 1963) rather than separate tribes (many other authors) or in some cases sub- families (Moore, 1964; Arnett, 1985). Smetana (1984) has made recent changes to this group. In- clusion of Creophilus and Thinopinus in the sub- tribe Staphylinina rather than Xanthopygina fol- lows Sharp (1884), Coiffait (1956), Lohse (1964), and Newton (in prep.). If these genera are placed in Xanthopygina, as done by Bernhauer and Schu- bert (1914), Cameron (1932), Hatch (1957), Coif- NEWTON & THAYER: CLASSIFICATION OF STAPHYLINIFORMIA 13 fait (1974), and Outerelo and Gamarra (1985), then Creophilina Kirby, 1837, becomes the valid name of that group. Tachyportnae— After Smetana (1983), with the addition of two tribes (Cordobanini and Symmix- ini) not discussed in the recent literature. Several tribes formerly included in Tachyporinae have been moved to Aleocharinae (Hammond, 1975). Xantholininae— See Staphylininae. Discussion and Summary of Changes Even before the imposition of formal rules cul- minating in the current Code (ICZN, 1961c, 1985a), most systematists working on Staphylin- iformia generally followed widely accepted (though informal) guidelines for use of family-group names, including the Principle of Priority. This is evident, for example, in the extensive published discus- sions covering the proper name to apply to Bathy- sciini (see {5}, below). Thus, full application of the Code to the use of family-group names in this group requires relatively few changes in current usage of the nearly 500 names concerned. Table 2 summarizes all such changes that we and others have found recently and the reasons for them. (The examples of recent implementation given may not include the first use of the change indicated, but we have found no previous implementation of those labeled only "present work.") In the following catalog we have implemented the current Code, making a deliberate effort to justify existing usage if possible, in the interest of stability. We have adopted all but six required changes of which we are aware. Five are the subject of current applications to the Commission (Hy- drophilidae: Helophorinae, see {3}, below; Leiod- idae: Coloninae, see {3}, below; Staphylinidae: Tachyporinae, see {16}, below; and Staphylinidae: Quediina and Xantholinini, see {23}, below). The sixth is a case where an imminent taxonomic change will make the question of priority irrele- vant (Hydrophilidae: Rygmodini and Cylomini; see Addendum). For the most part, we feel that implementing the remaining changes will not cause serious problems or confusion. Specialists on the groups involved may disagree and wish to apply to the Commission to conserve some other well- known names, such as Bolitocharini, Myrmedon- iini, and Bolitobiini in Staphylinidae; Limulodi- nae in Ptiliidae; or Leptininae and Bathysciini in Leiodidae. The last-mentioned is perhaps one es- pecially deserving of conservation because of the extensive ecological and physiological literature on the group. There are a few cases involving hom- onymy of family-group names in which we simply maintain existing usage for the time being (see {21}, below). The reasons for some changes require more ex- tensive comment than can be readily indicated in Table 2 and the catalog. These are discussed here and referred to by number in Table 2 and the catalog. { 1 } Acceptance of Non-latinized Names— Arti- cle 1 1 ffjii) provides for the availability of a pre- 1 900 non-latinized name, if it has been sub- sequently latinized, "generally accepted," and at- tributed to the author of the non-latinized name. The last two conditions are often difficult to de- termine, as Madge (1989) and others have pointed out, given the casual way in which family-group names have generally been used. Madge (1989) urged the acceptance of all non-latinized names based on type genera, at least when the names are from well-known publications. In our review of family-group names, we have accepted as available all non-latinized names that have been subsequently latinized, if we could find at least one subsequent citation of the non-latin- ized name as the origin of the name. We indicated other non-latinized names in the catalog as "una- vail. ?" if we could find no subsequent latinized use, or no subsequent citation of that particular non-latinized use. We have found only two cases in which name changes would result from accep- tance of non-latinized names that evidently have not been subsequently latinized or otherwise cited: Coryphiina Jacobson, 1908, is preceded by Bo- reaphilaires Mulsant & Rey, 1 880, for the name of the present tribe Coryphiini; Zerche (1990) used Boreaphilina as a new subtribal name with- out reference to Mulsant & Rey's earlier name. Dolicaones Casey, 1905, is preceded by Gnati- menitos Solier, 1849, and Gnathymenites La- cordaire, 1854, for the name of the present sub- tribe Dolicaonina. {2} Correction of Stem To Be Used in Forming Family-Group Name-The Code (Arts. 29, 35d(i)) provides that a family-group name must be cor- rectly formed from the stem of the name of its type genus, and if not so formed originally is to be corrected. The proper stem for family-group name formation for generic names that are or end 14 FIELDIANA: ZOOLOGY in Latin or Greek words is the genitive singular form with the case ending removed (Art. 29b). The family-group name stems of most generic names that are, or can be regarded as, Latin nouns are determined easily by dropping the nominative case ending (most commonly -us, -a, -um, -es, and many -is). The Code (Appendices B, D; Examples) and numerous lexicons (e.g., in English, Jaeger, 1944; Woods, 1944, 1966; Brown, 1956; also see Steyskal, 1 980) offer guidance in determining fam- ily-group stems. Several names requiring a stem change to form their genitive have been used cor- rectly in Staphyliniformia (corresponding generic endings and stems given): -gaster, -gastr-; -hospes, -hospit-; -onyx, -onych-; and -termes, -termit-. We have, however, found numerous names in Staphy- liniformia that have not been formed correctly, either at the time of their proposal or in subsequent use. In the catalog, all of these are indicated as "(incorrect original [or subsequent] spelling)," but are not otherwise discussed there. Because these misspellings fall into relatively few categories, it seems worthwhile to discuss each category here once, with cross-references by number in Table 2 and the catalog. {2a} A relatively trivial but common error in family group name formation is inappropriately adding or eliminating an -/- at the end of the stem of the name of the type genus, for example, Val- diini instead of the correct Valdini (from Valdd) or Rhexini instead of the correct Rhexiini (from Rhexius). Many of these have already been cor- rected in the literature, and such errors are nu- merous and obvious enough from the information given in the catalog that we do not list them here. On the other hand, in names such as Hydrophilii, Pselaphii, and Staphyliniae of Latreille (1802), it is difficult to determine what was meant as stem and what as ending (-/'/-//', -aeAiae). The endings clearly are not ones currently in use, but whether the original spellings were correct in terms of stem formation cannot be determined. Somewhat less obvious errors are those based on stems incorrectly formed through misinterpre- tation of Latin or Greek case endings and stems. We discuss these here grouped by ending, and list the names newly affected. One incorrectly formed family-group name of each type is given as an example. In the interest of stability, we have tried to keep existing usage if it can be justified (see especially {2f}). {2b} -on. This is a Greek ending; the genitive stem of the noun depends on its gender. Masculine and feminine nouns ending in -on keep the -on as the end of their stems; neuter nouns drop it. (A few masculine nouns with long o in the ending have genitive stems ending in -ont-\ see Steyskal, 1 980.) Four names in Staphyliniformia based on neuter generic names of this type must be cor- rected: Decarthrina (not Decarthronina), Pho- leuina, Mimecitini, and Elachistarthrini. The last two of these are currently junior synonyms. {2c} -soma, -loma, and -stoma. These are all endings of Greek neuter nouns (meaning body, edge, and mouth) whose genitive stems end in -somat-, -lomat-, and -stomal-, respectively. Su- perficially, they resemble, and can be mistaken for, Latin nouns ending in -a (which are normally fem- inine, but occasionally masculine). Type genera of family-group names in Staphyliniformia having the three endings listed have been uniformly treat- ed as neuter, and thus must be regarded as Greek nouns. The family-group names based on these genera that have not been corrected previously are Coelostomatini (not Coelostomini; a junior syn- onym), Lyrosomatini, Pterolomatini, Glypholo- matini, Scaphisomatini, and Conosomatini (a ju- nior synonym). The similar-looking name Cyloma was indicated by Sharp at the time of its descrip- tion as having "no classical derivation"; the fam- ily-group stem of such a name is determined (Art. 29b(ii)) by the author first basing a family-group name upon it (Cyllomina Zaitzev, 1908, the sec- ond "1" added erroneously), so Cylom- is the re- sulting family group stem. {2d} -ops. These are Greek nouns, originally either masculine or feminine, but to be treated as masculine under the Code (Art. 30a(ii)). Their gen- itive stems end in -op-. Most such names in Staph- yliniformia have been formed correctly, but three require correction: Amauropini (not Amauropsi- ni), Deropini, and Heterothopini (a junior syn- onym). {2e} -as. This is a Greek ending, occasionally adopted in Latin (Code: Appendix D); the genitive stem of the noun depends on its gender. Masculine nouns ending in -as drop the -as to form their stems; neuter nouns replace -as with -at-, and fem- inine nouns replace -as with -ad- . The only needed correction of this type that we found in Staphy- liniformia is Zyrini (not Zyrasini; a junior syn- onym); this correct spelling was used by Bradley ( 1 930). Seevers (1957) also seems to have followed this interpretation in describing the genus Ter- mitozyras (evidently derived from Zyras) and bas- ing the family-group name Termitozyrina on it. {2f} -opsis, -scepsis, -scelis, and -charis. These are endings of somewhat ambiguous standing. All NEWTON & THAYER: CLASSIFICATION OF STAPHYLINIFORMIA 15 Table 2. Summary of changes from recent usage of family-group names in Staphyliniformia required by the Code (ICZN, 1985a). Numbers in braces < } refer to items in "Discussion and Summary of Changes." Name change required From: To: Reason Examples of recent implementation HISTERIDAE Platysomini HYDROPHILIDAE Georyssinae Helocharina Helophorinae Rygmodini AGYRTIDAE Lyrosomini Pterolomini Platysomatini Georissinae Acidocerina Elophorinae Cylomini Lyrosomatini Pterolomatini LEIODIDAE Anisotomidae, -inae Leiodidae, -inae Anisotomini Bathysciini Catopinae, -ini Coloninae Hydnobiini Leptininae Pholeuonina PSELAPHIDAE Amauropsini Articerini Bythininae Bythinomorphi Decarthronina Fustigerini Metopiini Octomicrini Pyxidicerini Tanypleurina Tanypleurini Zethopsina PTILIIDAE Limulodinae Agathidiini Leptodirini Cholevinae, -ini Koloninae Sogdini Platypsyllinae Pholeuina Amauropini Tiracerini Goniacerinae Goniaceromorphi Decarthrina Clavigerodini Metopiasini Dimerini Bythinoplectini Natypleurina, nom. nov. Iniocyphini Bythinoplectina Cephaloplectinae SCYDMAENIDAE Clidicinae Mastiginae Euconnini Cyrtoscydmini Neuraphini Cyrtoscydmini Stenichnini Cyrtoscydmini correction of stem {2c} original spelling of stem {3) priority original spelling of stem priority correction of stem {2c} correction of stem {2c} priority {6,19} Art. 1 1 f(i)( 1 ); priority {6} priority {5} priority original spelling of stem priority {7} priority {8} correction of stem {2b} correction of stem {2d} misidentified type genus priority priority {4} correction of stem {2b} priority group name homonymy {9} priority priority preoccupied type genus {10} preoccupied type genus {10} priority priority priority {19} priority priority priority Mazur, 1973, 1984 Pope, 1977; Hansen, 1987, 1990; Lohse, 1989 present work (ex Hansen, 1990) McCorkle, 1965; Hansen, 1990; NOT IMPLEMENT- ED {3} NOT IMPLEMENTED (see Catalog; Hansen, 1990) present work present work Arnett, 1963, 1985; Peck, 1973, 1990; Lohse, 1989 Arnett, 1963; Lohse, 1989; Peck, 1990 Silfverberg, 1990; present work Zwick, 1979; Perreau, 1989; Peck, 1990 NOT IMPLEMENTED {3} Perkovsky, 1991 present work present work present work Besuchet, 1986; Newton & Chandler, 1989 Newton & Chandler, 1989 present work present work Newton & Chandler, 1989 present work Newton & Chandler, 1989; Coulon, 1989 Newton & Chandler, 1989; Coulon, 1989 present work present work Newton & Chandler, 1989; Coulon, 1989 Lawrence & Newton, 1982 present work present work present work present work 16 FIELDIANA: ZOOLOGY Table 2. Continued. Name change required Reason Examples of recent From: To: implementation STAPHYLINIDAE Acrotonina Strigotina priority present work Anthobiini Eusphalerini misidentified type genus Hatch, 1957; Arnett, 1963; Muona, 1979; Watanabe, 1990 Aphytopina Aphytopodina correction of stem {2g} present work Atanygnathina Tanygnathinina priority Burakowski et al., 1980 Bolitobiini Mycetoporini priority {17} present work Bolitocharini Homalotini priority {12} present work Callicerina Geostibina group name homonymy {11} present work Callicerini Athetini group name homonymy {11} Klimaszewski & Peck, 1986; Lohse, 1989; Lohse et al., 1990 Calocerina Glyptomina, nom. nov. preoccupied type genus present work Deremini Dorylophilini priority present work Deropsini Deropini correction of stem {2d} present work Dorylomimini Mimanommatini priority present work Glypholomini Glypholomatini correction of stem {2c} present work Gyrophaenini Homalotini priority {12} present work Holisina Hyptiomina priority {15} present work Lispinini Thoracophorini priority present work Mimecitonini Leptanillophilini priority {13} present work Myrmedoniini Lomechusini priority {14,19} present work Oligotini Hypocyphtini priority present work Pachyglossini Paglini, nom. nov. preoccupied type genus present work Paralispinina Clavilispinina, nom. nov. preoccupied type genus present work Physognathinae Solieriinae, nom. nov. preoccupied type genus present work Quediina Platycnemina priority NOT IMPLEMENTED {23} Scaphisomini, -ina Scaphisomatini, -ina correction of stem {2c} present work Suniina Astenina misidentified type genus Hatch, 1957; Arnett, 1963 Tachyporinae Tachininae priority {19} NOT IMPLEMENTED {16} Xantholinini Agrodini or Gyrohypnini priority NOT IMPLEMENTED {23} Zyrasina Myrmedoniina priority {14} Seevers, 1978; Lohse, 1989 Zyrasini Lomechusini priority {14,19} present work are originally Greek, but can also be interpreted as latinized Greek endings. The two interpreta- tions result in different genitive stems (and thus differently spelled family-group names). The al- ternative forms are Genitive stem Genitive stem Nominative ending if Greek end ing if Latin -ops is -opse- -ops- -charis -charit- -char- -scepsis -scepse- -sceps- -scelis -scelid- -scel- In Staphyliniformia, all the family-group names in use that are based on generic names having these endings use the Latin stems shown above, thus implicitly interpreting the generic names as latin- ized Greek nouns. Because this universal inter- pretation can be justified linguistically, it seems preferable for stability to maintain existing usage by treating these generic names in Staphylinifor- mia as Latin nouns. There seems little reason to overturn established and justifiable usage merely because an alternative interpretation of the stems (often with more complex spelling!) is possible. {2g} -pus. When this termination is derived from the Greek pous, meaning foot, the genitive stem ends in -pod-. Staphyliniform names requiring correction on this basis are Aphytopodina (not Aphytopina) and Ocypodina (a junior synonym). Several other names ending in -pus that have given rise to family-group names actually are derived from unrelated words ending in -pus (three ending in -metopus; also Platyprosopus and Stilipalpus) and thus have stems ending in -p-. {2h} -odus. When this termination is derived from the Greek odous, meaning tooth, the genitive stem ends in -odont-. None of the basonyms of family-group names in Staphyliniformia have this NEWTON & THAYER: CLASSIFICATION OF STAPHYLINIFORMIA 17 termination, although a few seem to: Holozodus appears to be derived instead from holo-, whole, and zodion, a small carved figure; Rygmodus is evidently derived from another generic name ("near Amarygmus,''\ White, 1846); and Stylopo- dus actually ends in -podus. {2i} Finally, there are two type genera requiring individual discussion. The first is Eleusis Laporte, upon which Sharp (1 887b) based the family-group name Eleusinina, with the stem Eleusin-. Laporte gave no indication of its derivation. Agassiz ( 1 846) indicated the generic name as a proper noun (oth- erwise unexplained) from the Greek. Sharp clearly regarded the generic name as being the Greek name Eleusis (genitive, Eleusinos), the name of a town in Attica where the goddess Ceres was worshiped (also applied to Ceres herself). Some later workers have used a name coordinate with Sharp's but based on the stem Eleusi-. There seems to be no justification for that spelling; if the generic name were regarded as Latin or latinized instead of Greek, its genitive stem would be Eleus-. {2j} The second unique case is that of Hister, upon which the family name Histeridae and sev- eral nominotypical subtaxa are based. This ap- pears to be simply a (second declension) Latin noun, whose stem should be Histr- {hister, -tri, — histrio, an actor; Simpson, 1968). This stem has been used in only three of the works cited by, for instance, Agassiz (1846) and Mazur (1984). Lin- naeus, in describing the genus, gave no indication of its derivation. Marseul (1854) gave its origin as "mot etrusque, [meaning] histrion." Jaeger (1944) also said: "hister- . . . Etruscan hister, an actor, giving rise to L. histrio, . . . -onis . . ." (emphasis added here). Hister is thus a non-Latin, non-Greek noun. For such generic names, the stem "for the purposes of the Code is that used by the author who establishes a family-group name based on that generic name" (Art. 29b(ii)). Gyllenhal ( 1 808) used Hister-, so that is the correct stem. {3} Confusion Concerning Original Spelling of Type Genus— In four cases, family -group names in common use have been based on emendations (only one justified) of the original spelling of the type genus. In two of these cases, Elophorinae and Georis- sinae of the family Hydrophilidae, the family-group names have already been corrected and used by several authors (see Table 2). In the case of the former, however, R. B. Angus (letter of 1 8 October 1990 to A. F. Newton) has submitted an appli- cation to the ICZN (Case No. 2796) requesting conservation of the more common usage Helo- phorus. Approval of this request would make Hel- ophorinae the correct spelling, and we maintain this prevailing usage here. The leiodid genus commonly called Colon was originally spelled Kolon by Herbst (1797). There is no doubt that this is Herbst's intended spelling; it is used as a generic heading and for two species in the text, as well as in the "Verzeichnis" on page vi and in the legend to figures of both species on plate 109. The name Colon is an evident emen- dation of Kolon Herbst by Illiger (1801), and is the base for the currently accepted family-group name Coloninae Horn, 1 880. The original spelling Kolon has been pointed out by (e.g.) Hatch (1928) and Pope (1977), but never adopted as the valid spelling for the genus or the family-group name based on it. Correcting the generic and family- group names to Kolon and Koloninae would cause a substantial alphabetical shift in this name, prob- ably creating considerable confusion. In view of the wide distribution of the genus and widespread usage of the names, Dr. H. Silfverberg (letter of 11 July 1990 to M. K. Thayer) is preparing an application to the Commission requesting conser- vation of the current usage, Colon and Coloninae. With this application pending, we maintain exist- ing usage here. The staphylinid genus commonly called Thora- cophorus was originally spelled Thoraxophorus by Motschulsky (1837). Both Erichson (1840a) and Motschulsky (1840) emended this to Thoracopho- rus, in which form the name has been subsequently used by all authors except Burakowski et al. (1979). Blackwelder (1952) considered Thoracophorus to be a justified emendation (of an error of transcrip- tion), but such emendations are not justified ac- cording to the Code (Art. 32c(ii)). It appears, in fact, that Motschulsky's original spelling was much more than an error of transcription. He gave the meaning of the name as "un homme ou un cheval arme d'une cuirasse [a man or horse armed with a breastplate]." The Greek word he gave as the supposed base of the name was incorrectly spelled, however. The correct spelling of the Greek word he obviously intended (see, e.g., Liddell & Scott, 1869, p. 7 19; Brown, 1956, p. 165) is indeed trans- literated as Thoracophorus. We regard Motschul- sky's (1837) French designation of the meaning of the name as indicating his true intent, and pro- viding internal evidence of an inadvertent error in the original (Greek) spelling of the name (as required by Art. 32c(ii) of the Code and illustrated by the Examples associated with it). The original 18 FIELDIANA: ZOOLOGY spelling, Thoraxophorus, must therefore be cor- rected to Thoracophorus, as it has been by virtually all authors since. {4} Nonstandard Ranks and Endings— In cer- tain families, formal genus-based names have been used between the ranks of family and subfamily (e.g., Histeridae: names ending in -morphae), or between subfamily and tribe (e.g., Pselaphidae: names ending in -morphi). These have generally been treated as independent of the coordinate set of family-group names regulated by the Code, and therefore have been cited with their own authors and dates (e.g., Mazur, 1984; Newton & Chandler, 1989). Because such names are based on a type genus, and are within the range of ranks covered by the Code, we see no reason why these names should not be treated the same as all other coor- dinate family-group names, taking the author and date of the oldest name based on the type genus. With one exception, this interpretation requires only a change in citation of author and date for such names in our catalog; in all cases, the first use of the name in its nonstandard form is also I cited. The one change is the pselaphid name By- thinomorphi, which becomes Goniaceromorphi. {5} Bathysciini/Leptodirini— The first family- group name to be applied to this taxon was Stagobiinae Schiodte (1849), based on the genus Stagobins Schiodte, but Stagobius was soon rec- ognized as a junior synonym of Leptodirus Schmidt. Lacordaire (1854) pointed out this synonymy and replaced the name Stagobiinae with "Leptoder- ides," a non-latinized name based on Leptoderus Schmidt, 1852 (an unjustified emendation of Lep- todirus Schmidt, 1832). Latinized versions of La- cordaire's name had appeared by 1859 (Lepto- derini Gutfleisch & Bose, 1859; Leptoderidae Kraatz, 1859a), and were used extensively in cat- alogs and lists published during the following 20 years (e.g., the 1862, 1868, and 1877 editions of Schaum's Catalogus Coleopterorum Europae, 1 863 and later editions of Marseul's Catalogue des Co- leopteres d 'Europe et du Bassin de la Mediter ranee en Afrique et en Asie) and intermittently thereafter (e.g., Reitter, 1891, 1906; Handlirsch, 1925; Hatch, 1933). Abeille de Perrin (1878) was evidently the first to use a name "Leptodirites" based on the correct original spelling Leptodirus, but his name is not a formal group name according to Jeannel (1936); Hatch (1933) was probably the first to use the correct spelling in a formal latinized name. Hatch (1933) and Arnett (1963, 1985) also used the name at a higher level, as Leptodiridae, for the group generally known as Catopidae (treated as Cholevinae in the catalog, below). Horn (1880) was evidently the first to use the name Bathysciae, based on Bathyscia Schiodte, for this group; he gave no reason for not using a name based on Leptoderus or Leptodirus, which he in- cluded in the group. Most subsequent authors, in- cluding virtually all systematists and ecologists working on this taxon, have followed Horn, using a name based on Bathyscia as the valid name for the group (e.g., Jeannel, 1910, 1911, 1914, 1936; Hatch, 1957; Laneyrie, 1967, 1978; Gueorguiev, 1974, 1976; Peck, 1973, 1990). Reitter (1884, 1886), using group names based on both Bathyscia and Leptoderus, adopted one based on Bathyscia for the larger group including both, but later (Reit- ter, 1891, 1 906) chose Leptoderini for the larger group, and still later ( 1 909) returned to Bathysci- ini. We have not been able to find discussions regarding the replacement of Leptoderini with Bathysciini until Hatch (1933), Jeannel (1936), and Zwick ( 1 979), who were unaware of the latinized names based on Leptoderus used prior to Horn ( 1 880). Lacordaire's ( 1 854) name Leptoderides was accepted (with correction) by Hatch (1933), re- jected by Jeannel (1936) because it was not latin- ized, and rejected by Zwick (1979) because it was based on an unjustified emendation. Silfverberg ( 1 990) independently arrived at conclusions sim- ilar to ours (next paragraph) regarding the priority of Leptodirini Lacordaire over Bathysciini Horn. Schiodte's ( 1 849) name based on the junior syn- onym Stagobius should be rejected under Article 40b, due to universal rejection of this name by authors from 1854 on. Names based on the un- justified emendation Leptoderus are available with original author and date, with correction of the stem to the original spelling of the genus, Lepto- dirus (Art. 35d), and with date of precedence of 1849 (Art. 40b). We accept Lacordaire's non-la- tinized name as providing availability of Lepto- dirini by provision of Article 1 1 ftjii), because his name was subsequently latinized and widely ac- cepted with attribution to him. If one wishes to question whether Leptodirini has been "generally accepted," because of more widespread use of the later name Bathysciini, then the first latinized use of the name (Kraatz, 1 859a, or Gutfleisch & Bose, 1859) would provide availability. In either case, present evidence indicates that Leptodirini is the correct name to apply to this taxon. Handlirsch (1925) and Arnett (1963) cited "So- lier, 1834" as author and date for Leptodiridae, but Zwick (1979) and Silfverberg (1990) correctly NEWTON & THAYER: CLASSIFICATION OF STAPHYLINIFORMIA 19 pointed out that Solier's (1834) name "Leptoder- ides" was not based on a genus and was applied to a section of Coleoptera: Heteromera! {6} Anisotomini/Agathidiini— The first use of the name Anisotomidae was by Stephens (1829a, p. 157, published February 1), who, however, did not mention Anisotoma Panzer as a valid genus in this group. His list of species belonging to the group included under Leiodes Latreille all those species normally included in both Leiodes and An- isotoma, so that Anisotoma must be considered an implied junior synonym of Leiodes. Later in the same year, Stephens repeated this treatment ( 1 829b, published June 1) and finally (1 829c, pub- lished July 15) mentioned Anisotoma explicitly, but only as a junior synonym. Anisotomidae Ste- phens is therefore not available (Art. 1 lf(i)l). Lat- er, the genera Leiodes and Anisotoma were mis- identified by Schmidt (1841), with their identities reversed, leading to much subsequent confusion. Erichson (1 845) used a name Anisotomidae based on Anisotoma of Schmidt, not Panzer, and there- fore Erichson's name is also unavailable. Reitter (1884) straightened out the misidentifications of Schmidt (1841) and was apparently the first to use a name Anisotomidae correctly based on Aniso- toma Panzer. However, the name Leiodidae Flem- ing (1821) has priority over Anisotomidae Reitter (1 884) for the name of this family, and Agathidiini Westwood (1838) has priority over Anisotomini Reitter (1884) for the tribe that includes both Agathidium and Anisotoma. {7} Hydnobiini/Sogdini-Perkovsky (1988) es- tablished that the genus Sogda Lopatin, 1961, type genus of the monotypic family Sogdidae Lopatin ( 1 96 1 ), is a senior subjective synonym of the genus Trichohydnobius Vogt, 1961, of the leiodid tribe Hydnobiini Jeannel, 1962. As noted later by Per- kovsky (1991), the name Sogdini has priority over Hydnobiini and Triarthrini (= Hydnobiini) Jean- nel (1962b). Triarthrini is also a junior homonym (see {21}), however, so in light of Perkovsky's (1991) use of Triarthrina as a subtribal name, the case should be referred to the Commission. {8} Leptininae/Platypsyllinae— The name Lep- tinidae has been attributed by virtually all authors, including LeConte (1872), to LeConte (1866). In that work, however, no family-group name based on the genus Leptinus was actually used. LeConte (1866, p. 368) merely said, after comparing the genus to members of several other families, "I therefore infer that Leptinus is a highly specialized type, representing a distinct family, having less affinity with Silphidae than with Hydrophilidae." The name Leptinidae was apparently first made available by LeConte (1872), subsequent to pro- posal of the name Platypsyllidae by Ritsema (1 869). {9} Metopiini/Metopiasini— The name Meto- piini Raffray ( 1 904b) (type genus, Metopias Gory), in current use for a tribe of Pselaphidae, is a junior homonym of the name Metopioidae Foerster (1868) (type genus, Metopius Panzer), in current use for a subfamily of Ichneumonidae (Insecta: Hymenoptera). There is also a third homonym, Metopiini Townsend (1908) (type genus, Metopia Meigen) in common use for a tribe of Sarcophag- idae (Insecta: Diptera). In cases such as this, where the names of type genera have identical stems and result in homonymous family-group names, the Code (Art. 55b) requires that the case be referred to the Commission for a ruling. We have submit- ted an application to the Commission (with T. Pape; Case No. 2793) to resolve this situation, including fixation of Metopias- as the stem of Me- topias Gory (giving Metopiasini) and Metopia- as the stem of Metopia Meigen (giving Metopiaini). Pending a decision by the Commission (and an- ticipating approval of our proposal), we here use Metopiasini as the valid name for this group; Jean- nel (1949) used this incorrect spelling of Raffray's name. {10} Tanypleurini, -ina/Iniocyphini, Natypleu- rina— In a recent review of the genera and higher taxa of Pselaphidae, Newton and Chandler (1989) overlooked the fact that Tanypleurus Raffray (1890) was preoccupied by Tanypleurus Steenstrup & Luetken, 1861. The currently used family-group names Tanypleurini and Tanypleurina Jeannel (1949), based on Tanypleurus Raffray, are there- fore no longer available and must be replaced. The names Iniocyphini and Dalmodini, both of Park (1951) and therefore of equivalent priority, are available for use as a tribal name; we choose the former to replace Tanypleurini, because of doubts about the identity of the type species of Dalmodes, type genus of Dalmodini (see Newton & Chandler, 1989, p. 7). The subtribe Tanypleurina has no available re- placement name. Because there are no other avail- able names for the genus Tanypleurus Raffray, we propose the replacement name Natypleurus, nom. nov. (gender masculine; type species, Tanypleurus 20 FIELDIANA: ZOOLOGY malaianus Raffray). We also propose the replace- ment name Natypleurina, nom. nov. (type genus, Natypleurus, nom. nov.) for the unavailable name Tanypleurina Jeannel ( 1 949). {11} Callicerini, -ina/ Athetini, -ina— Horion (1967) and Lohse ( 1 969) used the name Callicerini instead of Athetini for this tribe because Callicerus Gravenhorst was the oldest generic name in the group, without referring to Jacobson's ( 1 908) much earlier use of Callicerini. This usage continued for a time (e.g., Lohse, 1974; Muona, 1979) and then changed back to Athetini (Lohse, 1 989; Lohse et al., 1990) without explanation. Muona (1979) and Lohse et al. (1990) listed Callicerina and Athetina as separate subtribes within Callicerini/ Athetini, thus placing the two type genera in separate groups at a lower level. Although Callicerini Jacobson (1908) has priority over Athetini Casey (1910b), it is a junior homonym of Callicerini Rondani (1856) (based on Callicera Panzer), in use for a tribe of Syrphidae (Insecta: Diptera). This case of family-group homonymy must be referred to the Commission (Art. 55b), which can either suppress the junior name or change the stem of one hom- onym. The relative merit of these alternatives as far as usage within Staphylinidae depends on the taxonomic situation within the group. Athetini is available (and is in fact better known) as a name for any group including Atheta (with or without Callicerus), but a subordinate taxon including Cal- licerus and not Atheta would need a different name. Muona (1979) placed the genus Geostiba in the subtribe Callicerina; the available name Geosti- bina Seevers can be used for a taxon including Geostiba and Callicerus. The key question is, if Geostiba and Callicerus were to be separated, would Callicerus likely be combined with another genus already the basis of a family-group name? If not, a family-group name based on Callicerus would still be needed, and a stem change (for it or Cal- licera) would therefore be the most desirable course. Lacking the knowledge of this group necessary to answer the question, we will leave it to specialists in the group to submit an appropriate application to the Commission. In the meantime, we maintain existing usage of Athetini (the next oldest name) for the tribe and we use Geostibina Seevers for Callicerina of recent authors. If the name Calli- cerini Jacobson is made available by changing its stem (or that of the older Callicerini Rondani) through exercise of the plenary powers of the Com- mission, Jacobson's name would become the old- est family-group name in the present Athetini and would take precedence over any other name for taxa that include Callicerus. {12} Bolitocharini/Gyrophaenini/Homaloti- ni— Seevers (1978, p. 5) pointed out that the name Gyrophaenini Kraatz (1856) had priority over Bolitocharides Thomson (1859), but was incon- sistent in his use: in the main entry in his text (Seevers, 1978, p. 160) he used Bolitocharini, but in the checklist (Seevers, 1978, p. 266) he used Gyrophaenini. However, the still older name Homalotida Heer (1839a; see {20}) has priority over both names. Heer (1839a,b) used the genus Homalota Mannerheim in a very broad sense, ev- idently following Erichson (1837), rather than the restricted (monotypic) original sense of Manner- heim, and did not specifically mention the type species of Homalota, H. plana (Gyllenhal). Erich- son (1837) included Homalota plana in his con- cept of the genus; although he indicated later (Er- ichson, 1839a, pp. 689, 700, 1839b) that he had misidentified H. plana in his 1837 work, he still included the true H. plana (Gyllenhal) in his broad concept of the genus, and compared this species to H. linearis (Gravenhorst), included in Homa- lota by Heer (1839a,b) as well as Erichson (1837). Thus, although both Erichson (1837,1 839a,b) and Heer ( 1 839a,b) used the genus Homalota in a sense that included many species now placed in different tribes, and Erichson's (1837) original use of this expanded generic concept included a misidentifi- cation of the type species, we see no basis for con- sidering Homalota of Erichson and Heer to be a misidentified type genus, and thus no reason to reject Heer's family-group name. {13} Mimecitonini/Leptaniilophilini— Seevers (1965) used the name Mimecitonina (should be Mimecitina; see {2b}, above) for this group. In a list (1965, p. 188) he included the name "Mime- citonini, Wasmann, 1916" without further refer- ence. We have searched the publications of Was- mann for 1916 (including the one in Seevers's bibliography) and other publications in which Wasmann discussed his genus Mimeciton, but we cannot find any evidence that Wasmann used a family-group name based on this genus. The first use of the name Mimecitonini appears to be by Bernhauer and Schubert (1926), who gave no ref- erence to an earlier use of this name. The name Leptanillophilini Fenyes (1918) precedes Mime- citonini and Mimonillae, both of Bernhauer and Scheerpeltz(1926). NEWTON & THAYER: CLASSIFICATION OF STAPHYLINIFORMIA 21 {14} Zyrasini/Myrmedoniini/Lomechusini — This tribe was long known by the name Myrme- doniini, until Jeannel and Jarrige ( 1 949) proposed the new name "Zyrasini" (correct spelling would be Zyrini; see {2e}, above) because Myrmedonia had been synonymized with Zyras; Bradley ( 1 930) had earlier proposed "Zyrini" for the same reason. Usage since then has not stabilized: Lohse (1969, 1974), Kistner (1975), Klimaszewski (1982), Lohse et al. ( 1 990), and other authors have used Zyrasini, but Arnett (1963), Palm (1968), Seevers (1978), Muona (1979), Klimaszewski and Peck (1986), Lohse ( 1 989), and others have used Myrmedoniini for this group. Because the criterion of Article 40b of "general acceptance" of such a replacement name has not been met, Myrmedoniini would seem to be the appropriate name to use. However, Lo- mechusidae Fleming, 1821, has priority over both Myrmedoniini and Zyrasini, and should become the tribal name. At the subtribal level, the argu- ment above for use of Myrmedoniini over Zyrasini remains relevant, and the subtribe including the genus Zyras (= Myrmedonia) should be known as Myrmedoniina. {15} Holisina/Hyptiomina— Casey (1906) first used the name "Hyptiomae" for a group based on his new genus Hyptioma. Shortly thereafter (Cas- ey, 1910a), he pointed out that Hyptioma was a junior synonym of Holisus Erichson, but did not mention a family-group name. The next use of a family-group name involving this genus was ap- parently by Blackwelder ( 1 944), who used the sub- tribal heading "Holisi" for Holisus and another genus, without comment. Newton (1988), acting in the belief that Blackwelder's name was a re- placement name for "Hyptiomae" and therefore acceptable under Article 40b, used "Holisina" as a subtribal name for this group when redefining it. We can find no direct evidence, however, that Blackwelder intended his name as a replacement name rather than a new name (several of which appear in the same catalog). If considered a new name, "Holisi" is not available because it was not accompanied by a description or indication ac- ceptable for that time (Art. 1 3a). If considered a replacement name, it is still doubtful if it can be considered available because we can find no evi- dence that it was used at all, much less generally accepted before 1 96 1 (Art. 40b; the only other use of this name we can find is Newton, 1988). Thus "Hyptiomina" is the valid name for this group, and "Holisina," inadvertently made available by Newton (1988), is a junior synonym. {16} Tachyporinae and Tachininae— Tachini- dae Fleming (1821) (based on Tachinus Graven- horst, 1802 [Coleoptera: Staphylinidae]) is older than Tachinariae Robineau-Desvoidy (1830) (based on Tachina Meigen, 1803 [Diptera: Ta- chinidae]). Family-group names based on Tach- inus were frequently used in Staphylinidae until about 1839 (e.g., Mannerheim, 1830; Westwood, 1838; Heer, 1839a). Since then, however, the name Tachyporinae (MacLeay, 1 825) has been used uni- formly for the staphylinid group in question, and the name Tachinidae has been virtually univer- sally used in Diptera. Deliberate replacement of Tachininae by Tachyporinae in Staphylinidae ap- parently began with Erichson (1839a,b), who used Tachyporinae to avoid conflict with Tachinidae in Diptera (according to Erichson, 1840b, p. 230). His regarding Tachininae (Staphylinidae) as the junior name apparently resulted from attribution of Tachininae to Mannerheim (1830) (e.g., Erich- son, 1839b, p. 25), making it appear to be a junior synonym of MacLeay's Tachyporinae. We have found no discussion of the use of the junior hom- onym Tachinidae Robineau-Desvoidy in Diptera, although it was listed alongside the Coleoptera names (including Tachinidae Fleming, as "Leach 1817"; see {19} below) by Agassiz (1847). In re- cent years a group Tachinini, subordinate to Tach- yporinae, has sometimes been used in the staphy- linids (e.g., Coiflait, 1954). Because Tachyporinae (Staphylinidae) and Tachinidae (Diptera) are both well-established and frequently used names, we have continued existing usage of these names here, and have submitted an application to the Com- mission (with C. W. Sabrosky; Case No. 2786) requesting suppression of family-group names based on Tachinus before Mannerheim ( 1 830) and change of the stem of Tachinus to Tachinus- (for taxa including this genus and not Tachyporus). If approved, this would validate existing usage and leave (1) Tachinidae Robineau-Desvoidy avail- able in Diptera, (2) Tachyporinae MacLeay as the correct name for the staphylinid taxon including both Tachyporus and Tachinus, and (3) an avail- able name based on Tachinus for taxa including Tachinus but not Tachyporus. {17} Mycetoporini/Bolitobiini — Mycetoporini Thomson (1859) has priority over Bolitobiini Horn (1877). Mycetoporini was used extensively for a few decades (e.g., Thomson, 1867; Seidlitz, 1874, 1 889; Sahlberg, 1 876) but then was rather abruptly replaced by Bolitobiini, which has been in uni- versal use since (e.g., Ganglbauer, 1895; Bern- 22 FIELDIANA: ZOOLOGY hauer & Schubert, 1916; Cameron, 1932; Black- welder, 1944; Arnett, 1963; Lohse, 1964; Tikhomirova, 1973). We have found no discus- sion of this change, but it may have arisen from Ganglbauer's (1895) adoption of Bolitobiini with- out citation of Mycetoporini. Recently, Outerelo and Gamarra (1985) used the name "Mycetopo- riini" for a tribe of "Bolitobiinae," without ref- erence to an earlier use of the name. Thomson ostensibly based his name on Myce- toporus Mannerheim, 1830, but in fact included the previously designated type species of that ge- nus in a separate genus, Ischnosoma Stephens, 1829 (Tottenham, 1949; Blackwelder, 1952). A recent application submitted to the Commission by J. M. Campbell (Case No. 2733) discusses the complexities of this situation at the generic level and requests conservation of Mycetoporus in the sense of Thomson and nearly all subsequent au- thors. Mycetoporini Thomson ( 1 859) is the proper name for a family-group taxon including both My- cetoporus of Thomson (= Schinomosa Tottenham if the application is denied) and Ischnosoma. If the request for conservation is denied (leaving My- cetoporini Thomson based on a mistaken type- species designation) and a future author wishes to place Ischnosoma (= Mycetoporus) and Schino- mosa in separate family-group taxa, then appli- cation of the family-group name would need to be referred to the Commission (Art. 65b). {18} Necrophilidae Gistel— This family name appeared between the families Peltidae (= Tro- gossitidae) and Silphidae in a checklist of Bavarian insects (Gistel, 1856), with the only included spe- cies being ""Thymallus Lat. limbatus" (= Thy- malus limbatus (Fabricius), now in the family Tro- gossitidae). Gistel's (1856) family names, many of them new, were evidently all based on generic names, although the implied type genus was not always included in the checklist. In the case of Necrophilidae, there are no generic names usually associated with Thymalus or other Trogossitidae that would give rise to such a family name. How- ever, the adjacent family Silphidae (in which Thy- malus and some other trogossitids were included by some early authors) included at that time two such names, neither of which was mentioned by Gistel: Necrophila Kirby & Spence, 1 828 (still a genus of Silphidae), and Necrophilus Latreille, 1 829 (now a genus of Agyrtidae). Because of uncertainty about which, if either, of these names might have been meant by Gistel, we conclude that Necro- philidae Gistel is unavailable, as it does not meet the criterion of availability set out in Article HfliXD. {19} "Leach 1817'VFleming, 1821, Names-A number of early family-group names in Staphy- liniformia were cited by Agassiz (1846) as "Leach 1817," with the reference "Encycl. Brit." Subse- quent workers (including Agassiz, 1847) have at most credited these names to Leach, 1817 (without specific reference). They are not, however, among the new family-group names in Leach's Zoological Miscellany, nor are they in his other 1817 works. According to R. B. Madge (letter of 14 May 1 990), Leach wrote the articles on " Annulosa" and "Entomology" in the Supplement to the 4th, 5th, and 6th editions of the Encyclopedia Britannica, but neither contains family-group names. The names attributed by Agassiz ( 1 846, 1 847) to Leach, 1817, appear instead in the "Insecta" article of the Supplement, an article written by John Fleming. Authorship of articles in the Supplement is indi- cated by a code given at the end of each article. "Q.Q." at the end of the "Insecta" article meant J. Fleming, and "V." following the "Annulosa" and "Entomology" articles meant Leach; Volume 1 contains a list of contributors and their corre- sponding codes (R. B. Madge, letter of 14 May 1990). Publication dates for the parts of the Sup- plement are given in Volume 6, Part 2; Volume 5, Part 1, containing the "Insecta" article, is there dated as July 1821. The peculiar spelling of many names in Fleming (1821), reproduced in Agassiz's lists and there at- tributed to Leach, supports the idea that Fleming's "Insecta" article was Agassiz's source for them. For example, some names are formed upon entire generic names instead of generic stems (e.g., Leio- desidae and, outside Staphyliniformia, Clerusidae, Ptinusidae, Erotylusidae, etc.); others have the ter- mination -adae instead of -idae (e.g., Spheridi- adae and, outside Staphyliniformia, Melolontha- dae); and some have nonstandard stems (e.g., Mastigoidae). MacLeay ( 1 830), in a caustic review of Fleming's overall work including the article in the Supplement, criticized these spellings and con- cluded "Luckily, however, no great harm is done; for few naturalists place [Fleming's] names even in their list of synonyms." This statement may explain why Fleming's names were generally over- looked or ignored by contemporary authors. Flem- ing's names are fully available, however, and most of them have been cited at least occasionally fol- lowing their appearance in Agassiz (1846, 1847). The attribution of these names to "Leach 1817" NEWTON & THAYER: CLASSIFICATION OF STAPHYLINIFORMIA 23 (Encyclopaedia Britannica) is apparently a mistake on the part of Agassiz (1846), and should be cor- rected to Fleming (1821). Wheeler (1986, p. 135) independently arrived at the conclusion that an- other "Leach 1817" name, Lymexylonidae, was first published by Fleming (1821). {20} Priority of Erichson, 1839/Heer, 1839, Names— Several family-group names in Staphy- linidae were first published in 1839, in up to four places by two authors: Erichson (1 839a, Die Kdfer der Mark Brandenburg, 1(2); 1839b, Genera et Species Staphylinorum, pt. 1) and Heer (1839a, Fauna coleopterorum Helvetica, 1(2); 1839b, Die Kdfer der Schweiz, 1(2)). (Heer (1839b) was a sep- arate printing of a paper that appeared in Neue Denkschriften der allgemeinen schweizerischen Gesellschaft fur die gesammten Naturwissenschaf- ten, Volume 3 (1839), according to the title page of the separate, but Volume 4 (1840) according to Hagen (1862) and the title page of the journal it- self.) The title pages and other standard biblio- graphic sources give only "1839" as the date of publication for all four of these works. Both Erich- son (1839b) and Heer (1839a) cited (with page numbers) new names published by Erichson (1839a). Heer (1839b), in a checklist that gave at most authors' names with the taxa listed, included names of many species described by Erichson (1839a) and Heer (1839a). The names published as new in Heer (1839a) were listed as "Heer" in Heer (1839b). Heer (1839a,b) did not refer to Erichson (1839b), and vice versa. Later, Heer (1841a, p. 553) expressed the opinion that Erich- son (1839b) appeared in 1840, the year after Heer (1839a), but other evidence (e.g., publication no- tice in Entomologische Zeitung [Stettin]: 1 : 12) shows that Erichson (1839b) did indeed appear in 1839. Erichson (1840b), in a review of 1839 en- tomological literature, discussed all four 1839 works in the apparently chronological sequence Erichson (1839a), Heer (1839a), Heer (1839b), and Erichson (1839b), although without explicit men- tion of absolute or relative dates of publication. Although we have no exact publication dates for any of the 1839 works (so technically they should be considered simultaneously published), it is clear that Erichson (1839a) appeared before the other works; Heer (1839a) probably preceded Heer (1839b); and the priority of Erichson (1839b) rel- ative to Heer (1839a) and Heer (1839b) is uncer- tain. We accept Erichson's (1840b) own sequence of discussion of these works as setting their order of precedence and treat Erichson (1839b) as the last published of the four. {21} Apparently Homonymous Family-Group Names— Under Article 55b of the Code, if "hom- onymy between family-group names results from similarity but not identity of the names of their type genera, the case is to be referred to the Com- mission for a ruling. . . ." The Commission may either reject the junior homonym or amend the stem of one name, thus removing the homonymy. If rejected, a junior homonym must be replaced unless it is rejected as a junior synonym (Art. 60). We have found 10 instances of apparently hom- onymous family-group names in Staphyliniformia and other taxa. Strict application of the Code would require that most of these be submitted to the Commission. We are submitting applications con- cerning two of these, but for the reasons indicated below we are not pursuing the other cases. Finding and tracking family-group names in other groups is generally as difficult as in Staphyliniformia, and there may be others we have not found. Full ci- tations and type genera for all names are given in the catalog. Callicerini Jacobson (1908), Staphylinidae, not Callicerini Rondani (1856), Diptera. See {11}, above. Cryptobiina Casey (1905) Staphylinidae, predates Cryptobiinae Hollande (1952), Protozoa. The junior name as cited is a nomen nudum; if it has been validated elsewhere, the case needs to be referred to the Commission. Cyphinae Lohse (1974), Staphylinidae, not Cy- phini Leng (1920), Coleoptera: Curculionidae. We can find no earlier use of Cyphinae (Staph- ylinidae) than Lohse (1974), but it was given there as a synonym of Hypocyphtinae. If this was its first use, it is therefore not available from that date (Art. 1 le). Adam (1987) used the in- correct spelling Cyphainae as the valid name for a group including the same staphylinid genus Cypha, but without making it available or giving any reference to an earlier use. If the staphylinid name has been validated elsewhere, the case needs to be referred to the Commission. Helocharina Orchymont (1919), Hydrophilidae, predates Helocharini Metcalf (1965), Homop- tera. The junior name as cited is a nomen nudum; if it has been validated elsewhere, the case needs to be referred to the Commission. 24 FIELDIANA: ZOOLOGY Hydrobiini Mulsant (1844), Hydrophilidae, pre- dates Hydrobiidae Troschel (1857), Mollusca. Recently, in an application to the Commis- sion dealing with other problems in Mollusca, Rosenberg and Davis ( 1 990) requested that Hy- drobiidae Troschel be placed on the Official List of Family-Group Names, apparently unaware of the homonymy indicated above. We have submitted a comment on that application rec- ommending against such action until the hom- onymy has been dealt with (Newton & Thayer, 1990). The case needs to be submitted to the Commission. Metopiini Raffray ( 1 904b), Pselaphidae, not Me- topiinae Foerster (1868), Hymenoptera; both predate Metopiini Townsend ( 1 908), Diptera. See {9}, above. Steninae MacLeay ( 1 825), Staphylinidae, predates Stenidae Fraser & Purves (1960), Mammalia. The junior name as cited is a nomen nudum; if it has been validated elsewhere, the case needs to be referred to the Commission. It has been treated by at least some workers as a synonym of Delphinidae. Tachininae Fleming (1821), Staphylinidae, pre- dates Tachinidae Robineau-Desvoidy (1830), Diptera. See {16}, above. Toxoderina Bernhauer & Schubert (1911), Staph- ylinidae, not Toxoderinae Saussure (1869), Mantodea. The junior name has seldom, if ever, been used except by Scheerpeltz ( 1 933). It is currently treated as a junior synonym of Coprophilini, the type genus having been synonymized with Homalotrichus (Herman, 1970). An application requesting rejection of Toxoderina Bernhauer & Schubert should be submitted to the Commis- sion. Triarthrini Jeannel ( 1 962b), Leiodidae, not Triar- thrinae Ulrich (1930), Trilobita. The junior name has recently been considered either a junior synonym (Daffner, 1983; Peck, 1 990) or a subtribe of Sogdini (Perkovsky, 1 99 1 ). 1910, 1916; Blackwelder, 1944; -o-: Erichson, 1834; Agassiz, 1847; Dohrn, 1865; Bickhardt, 1914; Mazur, 1 984). Although there has been some discussion of the discrepancy (e.g., Dohrn, 1865, 1870), there has been no satisfactory resolution of the conflict. We Anally discovered its source, how- ever: there were two separately typeset printings of the original description, one of which (Esch- scholtz, 1829a, p. 1 1) used Trypanaeus, the other (Eschscholtz, 1829b, p. 10) Tryponaeusl We can find no evidence regarding relative or exact pub- lication dates of the two editions, both of which bear the date 1829. The two editions must thus be considered as simultaneously published, with the correct name to be determined by the first reviser (Art. 24). The first reviser appears to have been Agassiz (1846), who cited both spellings as "Eschscholtz 1829," and used Trypanaeus as the correct form. We therefore choose to use Trypan- aeus (also the more common spelling) and Try- panaeinae. {23} Quediina/Platycnemina and Xantholinini/ Agrodini/Gyrohypnini — Nordmann (1837) es- tablished two group names, Platycnemidiformes and Agraeformes, for new genera now placed in Quediina Kraatz, 1857 and Xantholinini Erich- son, 1839, respectively. In each case, Nordmann's group name has not been used or even cited sub- sequently, and his type genus has long been treated as a synonym or subgenus of another genus (Pla- tycnemus of Haematodes Laporte, Agrodes of Plo- chionocerus Dejean). Kirby (1 837) based the name Gyrohypnidae on the genus Gyrohypnus "Kirb. Steph.," but his name has not been used or cited since, and the genus Gyrohypnus has long been placed in Xantholinini. Hatch (1957) indepen- dently proposed Gyrohypnini (based on a now- rejected concept of Gyrohypnus [see ICZN, 1983]) as a new name for Xantholinini. Since the names Quediina and Xantholinini (and their coordinate forms) have been in universal use for well over a century, one of us (AFN) is submitting a proposal to the ICZN requesting their conservation over the never-used names of Nordmann and Kirby. { 22 } Try ponaeinae/Trypanaeinae — Marseul (1 857) based the name Trypaneens (later latinized by Jacobson, 1910) on Trypanaeus Eschscholtz, citing Erichson's (1834) spelling Tryponaeus as a typographical error. Both spellings of the generic (and resultant family-group) name have been used (-a-: Agassiz, 1846; Marseul, 1857; Bickhardt, Diagnoses of Taxa Empelinae Newton & Thayer, subfam. nov. Type genus: Empelus LeConte, 1 86 1 . NEWTON & THAYER: CLASSIFICATION OF STAPHYLINIFORMIA 25 The genus Empelns LeConte includes the single rare species E. brunnipennis (Mannerheim), known from southern Alaska to California. The genus was originally described in the tribe Clambini of the family Silphidae (LeConte, 1861), and was re- tained as a rather aberrant member of that tribe (e.g., Horn, 1880), even as the group was later elevated to family status as Clambidae (e.g., Hatch, 1929, 1957). Crowson (1955) and Endrody-Youn- ga (1959) removed Empelus from that family and placed it in Anisotomidae (now Leiodidae), with- out further comment. Later, Crowson (1 960) char- acterized the genus as one of the most primitive members of Staphylinoidea, and discussed how other staphylinoid families might be derived from it, but did not assign the genus to a family-group taxon. Hammond (1971) commented that Em- pelus "... clearly does not belong [in Leiodidae], but has distinct affinities with the Proteininae [Staphylinidae] and may eventually be assigned to that subfamily." Perhaps with these discussions of Crowson (1960) or Hammond (1971) in mind, but without citing them, many authors providing checklists or discussions of higher taxa of Coleoptera have men- tioned a staphylinoid family Empelidae (e.g., Ab- dullah, 1969, and later works; Crowson, 1981; Paulian, 1988), or a staphylinid subfamily Em- pelinae (e.g., Hlavac, 1975; Lawrence & Newton, 1982; Thayer, 1987). However, none of these pub- lications using a family-group name based (ex- plicitly or by inference) on Empelus LeConte pro- vides a differential diagnosis or description of the family-group taxon, or reference thereto, that would make the name available (Art. 1 3a). Because a family-group name based on Empelus is being widely used in the systematic literature, and because of our conviction that such a name will be needed regardless of the final consensus on ranking and exact placement of the group, we here provide a differential diagnosis and discussion to make the name available. Descriptions or mention of significant characteristics of Empelus brunni- pennis have been published by LeConte (1861), Horn (1880, including figures of habitus and an- tenna), Hatch (1957, including habitus figure), Crowson (1960), Hammond (1971), Hlavac (1975, including figure of prothorax), and Thayer (1987). Our diagnosis and discussion are based on those sources and on examination of cleared and slide- mounted specimens of adults of both sexes of the species. We have also examined a probable syn- type in the Museum of Comparative Zoology, Harvard University. The immature stages are un- known. Diagnostic description— Small (under 2 mm long), compact, and dorsally convex beetles with: head hypognathous, strongly flattened, without ocelli, with antennal groove ventrally between eye and maxillary foramen; antenna 1 1 -segmented with loose but very distinct 3 -segmented club; man- dible with membranous prostheca, large molar lobe, without preapical teeth; maxillary palp 4- segmented, segment 3 subquadrate, segment 4 about 4 times as long as 3; pronotum short and broad, evenly convex, with rounded antero- and posterolateral corners; presternum a very narrow transverse strip anterior to procoxae, without in- tercoxal process; elytra without striae, nearly cov- ering abdomen (no more than 3 segments ex- posed), elytral epipleural fold only half as long as elytron; abdominal sternites 3-8 visible, sternite 3 with carina-delimited coxal cavities; abdominal sternite 8 with basal projection associated with glandular structures (projection more than one third as wide as sternite); functional spiracles on abdominal segments 1-8; abdominal interseg- mental membranes long, apically attached, with brick-wall pattern; wing-folding patches on ab- dominal terga 2-4; pro- and mesocoxa oblique, very elongate; mesosternum very short, leaving procoxa and mesocoxa separated from each other by less than width of either; hind coxa transverse, excavate posteriorly, covering short femur in re- pose; each femur with tibial groove ventrally; all tarsi 5 -segmented, empodia unisetose; wing- fold- ing asymmetrical; wing with costal hinge proximal to radial sector; aedeagus composed of basally en- larged median lobe with small foramen, pair of free parameres, and internal sac with well-devel- oped flagellum. Discussion— The wing- folding pattern, wing hinge placement, structure of the median lobe of the aedeagus, and abdominal intersegmental pat- tern are derived or apomorphous characteristics of the family Staphylinidae sensu lato, while the presence of the unique glandular structure at the base of abdominal sternite 8 clearly places Em- pelinae in the "Omaliine Group" of staphylinid subfamilies (Hammond, 1971; Lawrence & New- ton, 1982; Thayer, 1987). Within this group, the presence of a full set of functional abdominal spir- acles excludes Empelinae from the "Proteinine Subgroup" of Thayer (1987), in which the inter- mediate abdominal spiracles are atrophied. Em- pelinae also differ from Proteininae (to which 26 FIELDIANA: ZOOLOGY Hammond (1971) suggested they might belong) in having long abdominal intersegmental mem- branes with a brick-wall pattern. This leaves Em- pelinae together with Omaliinae and Aphaeno- stemminae as primitive members of the Omaliine Group, with unresolved relationships to one an- other. Empelinae differ from both Omaliinae and Aphaenostemminae in lacking a postcoxal process of the pronotum. Some of the following features of Empelinae occur in some Omaliinae, but not all of them occur there in combination: lack of ocelli; strong 3-segmented antennal club; antennal groove; long elytra almost completely covering the abdomen dorsally; excavate metacoxae. Solieriinae Newton & Thayer, nom. nov. Type genus: Solierius Bernhauer, 1 92 1 (new name for Physognathus Solier, 1849, not Agassiz, 1847). The genus Solierius Bernhauer (replacement name for Physognathus Solier, preoccupied) is known from a single rare species, S. obscurus (So- lier), found in southern Chile and adjacent parts of Argentina. Solier ( 1 849) originally placed the genus in a new staphylinid subtribe, "Fisognati- tos," and compared it to genera now placed in Omaliinae, with a comment on its pselaphid-like appearance. Lacordaire (1854) repeated Solier's treatment of the genus in French, using the group name "Physognathites." Kraatz (1859b) reviewed the characteristics and placement of the genus, agreed that it did not belong in any of the estab- lished groups of Staphylinidae, and concluded that ". . . die Gruppe der Physognathites am besten neben den Omalini einzuschalten sein. . . ." Evi- dently there has been no subsequent use of a fam- ily-group name for this genus based on either of its generic names. Most subsequent authors and all catalogs have placed Solierius in the tribe Om- aliini (of Oxytelinae) or the equivalent subfamily Omaliinae (e.g., Gemminger & Harold, 1868; Ei- chelbaum, 1909; Bernhauer & Schubert, 1910; Blackwelder, 1944; Coiffait & Saiz, 1968; Puthz, 1974; Shibata, 1970: in "Coryphium complex"). However, Fauvel (1889) noted that the genus ". . . est aberrant et doit former une tribu speciale re- liant les Staphylinides aux Pselaphides . . . ," and Newton (1985), listing the genus as "subfamily uncertain," indicated that it did not belong in Om- aliinae and ". . . cannot be related definitely to any other higher taxon of staphylinids, or to . . . pse- laphids. . . ." We interpret Kraatz's (1859b) name "Physog- nathites" as latinized, from the context of his com- ments quoted above. If this interpretation is ac- cepted, then technically we are simply proposing a new name to replace one based on a preoccupied type genus. If, however, Kraatz's name is inter- preted as a subsequent use of Lacordaire's name of the same spelling, which is clearly not latinized (an accent appears in names of some other taxa of the same rank and appearing in the same format), then our treatment here will constitute the de- scription of a new taxon. In either case, this group currently lacks an available family-group name, and in our opinion requires one (regardless of the rank and placement that eventually will be decided upon). Thus, we provide a diagnosis and discus- sion comparable to that for the new taxon Em- pelinae. Descriptions of Solierius obscurus have been presented by Solier (1849, including an in- accurate habitus figure and seven more accurate detail figures), Kraatz (1859b), Coiffait and Saiz (1968, including a good habitus figure), and Puthz (1974, including figures of maxillary palp and ae- deagus). Our diagnosis and discussion are based on those sources and on examination of cleared and slide-mounted specimens of adults of both sexes of the species. We have also examined syn- types in the Museum National d'Histoire Natu- relle (Paris) and Deutsches Entomologisches In- stitut (Eberswalde). The immature stages are unknown. Diagnostic Description— Slender, pubescent beetles (about 3.5 mm long) with: head narrow, with distinct neck constriction, dorsum of head medially with triangular impression at base grad- ing into carina between antennal bases, eyes strongly protruding; antennal insertions hidden in dorsal view, distinctly separated from anterior margin of head capsule; antennae 1 1 -segmented, not clubbed; mandible with molar lobe and bifid apex; maxillary palp subequal in length to head width including eyes, 4-segmented, segment 3 swollen and distinctly longer and wider than acute- ly conical segment 4; pronotum constricted ba- sally, with 2 pairs of furrows (one paramedian and longitudinal, other more lateral and somewhat oblique) and shallow median pit near base; elytra short, covering part of tergite 3, epipleural fold lacking; abdominal sternites 3-8 visible, function- al spiracles on segments 1-8, segments 3-7 each with two pairs of paratergites; abdominal inter- NEWTON & THAYER: CLASSIFICATION OF STAPHYLINIFORMIA 27 segmental membranes long, apically attached, with brick-wall pattern; sternite 3 with carina-delimited coxal cavities; anterior margin of tergite 7 bisin- uate, projecting further than usual into segment 6; tergal wing-folding patches and sternite 8 gland absent; protrochantin concealed, procoxa project- ing, lacking external keel; all tarsi 5-segmented, em podia bisetose, setae truncate and slightly ex- panded apically; wing-folding asymmetrical; wing with costal hinge proximal to radial sector; male with curled laminar structure projecting ventrally from gula, median cluster of stout setae at apex of prosternum, and apically bifid sternite 9; aedeagus very elongate, basal bulb of median lobe small, parameres long and narrow, slightly asymmetrical; female genital segment with single pair of apically well-sclerotized and densely setose gonocoxites, without styli. Discussion— The wing-folding pattern, wing hinge placement, structure of the aedeagus, and abdominal intersegmental pattern are derived or apomorphic characteristics of the family Staphy- linidae sensu lato (Lawrence & Newton, 1982). The lack of the sternite 8 gland and the presence of two pairs of abdominal paratergites rule out placement of Solierius in the Omaliinae, where it has usually been placed, or anywhere in the "Oma- liine Group" of staphylinid subfamilies (Ham- mond, 1971; Lawrence & Newton, 1982; Thayer, 1987). As yet, we have not found any characters that link it convincingly with any other subgroups of Staphylinidae, and the position of Solieriinae within this large assemblage remains ambiguous. Catalog With two exceptions, we have chosen to limit the taxonomic ranks used in this list to the gen- erally accepted ones of superfamily, family, sub- family, tribe, and subtribe, using the Code-man- dated or recommended endings of -oidea, -idae, -inae, -ini (Art. 29a, Rec. 29A), and -ina, respec- tively (the last not mentioned in the Code, but widely accepted in Coleoptera, at least). In certain families, formal or informal groups have been used between the ranks of family and subfamily (e.g., Histeridae: genus-based names ending in -morphae; Pselaphidae: informal names), or be- tween subfamily and tribe (e.g., Pselaphidae: ge- nus-based names ending in -morphi). In Histeri- dae and Pselaphidae, where such taxa are in general use, we have listed them as valid names and placed them appropriately in the classification. In all oth- er cases, we have cited such names with other names based on the same type genus, or (informal names) as synonyms of the next higher taxon. It has been occasional practice to use informal or formal names for groups between family and superfamily, or of indeterminate status above the family level, as in Reitter (1909), Naomi (1985) and Paulian ( 1 988). Such groups are not in general use and are not included in the classification here. Because these names also are either not formal ones or, if based on a type genus, are established family-group names previously used at the family level or below, we have not included them in our list at all. The rank of infratribe, used recently in Staphyliniformia only within one subtribe (Jacob- son et al., 1986), is not formally shown in the list: the two names concerned are given as synonyms of their subtribe. We have not tried to include all existing uses of family-group names. Citations are given only for the first use of a family-group name based on a given type genus, for independent proposals of names based on the same genus, and for a few subsequent uses of names using different orthog- raphies or unusual suffixes or when it is not clear whether a use is based on an earlier name or in- dependently proposed. Clearly subsequent uses are indicated by a semicolon (;) between the name and reference. We have also attempted to determine and cite the first latinized use of family-group names originally proposed in the vernacular. We have included emendations, but have made no attempt to list all variant spellings, incorporating only those that have attained widespread use. For the sake of brevity, each citation (i.e., name with reference) is listed only once. Because of the Prin- ciple of Coordination, uses of a name at different ranks or for different concepts of a group are not separate nomenclatural acts, and therefore we do not cite such multiple uses. For each taxon recognized here, the correct spelling of the valid name is given as a heading in all upper case letters, followed by its author, date of availability, and (if applicable) currently used or well-known synonyms. Names coordinate with (i.e., based on the same type genus as) a valid name have their full citations given only under the high- est ranked use of that name (up to the family level). Citations for synonymous (or otherwise invalid) names based on other type genera are placed under the lowest ranked taxa to which their type genera are assigned (Art. 35c). As an example, family-group names based on 28 FIELDIANA: ZOOLOGY the genus Staphylinus Linnaeus are used at six different hierarchical levels (series through sub- tribe). Full citations for names based on Staphy- linus are listed only under the heading for the family Staphylinidae. At the other levels (Staphy- liniformia, Staphylinoidea, Staphylininae, Staph- ylinini, and Staphylinina), the respective taxon names appear as headings with author and date, but without full citations. Names based on the genera Creophilus, Thinopinus, and Ocypus, placed in the same subtribe as Staphylinus, are listed only under the heading for the subtribe Staphylinina (i.e., under the lowest taxon to which they can be assigned). The full citation for any name may be found by looking up its type genus in the index. Within each taxon, subtaxa of the next lowest rank are listed in alphabetical order. Within each taxon, the citation for the valid name is first (unless given at a higher taxonomic level), followed by other citations in chronological order. Each citation of a name includes the stated or implied type genus (with its author and date). If the author of a particular citation of a name gave the type genus differently, did not indicate one, or gave a different author for the type genus, this is so indicated (in parentheses). The notation "see Discussion {#}" refers to the numbered items in the section "Discussion and Summary of Changes" above. References for all family-group name ci- tations are listed in "Literature Cited," but those for type genera are not (unless they happen to con- tain cited family-group names). All family-group names and type genera included in the catalog are listed in the index. STAPHYLINIFORMIA Latreille, 1802 HYDROPHILOIDEA Latreille, 1802 HISTERIDAE Gyllenhal, 1808 Histeroides Gyllenhal, 1808: 74 (see Discussion {2j}). Type genus: Hister Linnaeus, 1758. Cyrthisterinae Houlbert & Monnot, 1922: 12 (unavail., not based on genus). Plathisterinae Houlbert & Monnot, 1922: 12 (unavail., not based on genus). Histeromorphae; Wenzel, 1944: 53 (group between family and subfamily; see Discussion {4}). Type genus: Hister Linnaeus, 1758 (not cited). HISTEROMORPHAE Gyllenhal, 1808 DENDROPHILINAE Reitter, 1909 Dendrophilini Reitter, 1909: 288. Type genus: Dendrophilus Leach, 1817. ANAPLEINI Olexa, 1982 Anapleini Olexa, 1982: 38. Type genus: Anapleus Horn, 1873. BACANIINI Kryzhanovskij, 1976 Bacaniini Kryzhanovskij, 1976: 266. Type genus: Bacanius LeConte, 1853. Bacaniini Vienna, 1974: 273 (unavail., no description). Type genus: Bacanius LeConte, 1853. DENDROPHILINI Reitter, 1909 PAROMALINI Reitter, 1909 Paromalini Reitter, 1909: 287. Type genus: Paromalus Erichson, 1834. HETAERIINAE Marseul, 1857 Heteriens Marseul, 1857: 148 (not latinized; avail., Art. 1 lfl[iii); see Discussion {1}). Type genus: Hetaerius Erichson, 1839. Hetaeriini Schmidt, 1 885: 28 1 . Type genus: Hetaerius Erichson, 1 839. Note: First latinized use? Hetaeriomorphini Bickhardt, 1914: 308 (avail., Art. 12b(4)). Type genus: Hetaeriomorphus Schmidt, 1893. Note: Treated incorrectly as nomen nudum by Mazur (1984). HISTERINAE Gyllenhal, 1808 NEWTON & THAYER: CLASSIFICATION OF STAPHYLINIFORMIA 29 EXOSTERNINI Bickhardt, 1914 Exosternini Bickhardt, 1914: 308 (avail., Art. 12b(4)). Type genus: Exosternus Lewis, 1902. Note: Treated incorrectly as nomen nudum by Mazur (1984). HISTERINI Gyllenhal, 1808 HOLOLEPTINI Hope, 1840 Hololeptidae Hope, 1840: 106. Type genus: Hololepta Paykull, 1811. OMALODINI Kryzhanovskij, 1972 Omalodini Kryzhanovskij, 1972: 19. Type genus: Omalodes Erichson, 1834. Omalodini Reichardt, 1941: 37 (unavail., no description). Type genus: Omalodes Erichson, 1834 (not cited). PLATYSOMATINI Bickhardt, 1914 (= Platysomini) Platysomini Bickhardt, 1914: 307 (avail., Art. 12b(4); incorrect original spelling; see Dis- cussion {2c}). Type genus: Platysoma Leach, 1817. Note: Treated incorrectly as nomen nudum by Mazur (1984). Althanini Cooman, 1939: 138 (as tribe of Teretriinae). Type genus: Althanus Lewis, 1903. Platysomatini; Mazur, 1973: 51 (correction of original spelling). Type genus: Platysoma Leach, 1817. ONTHOPHILINAE MacLeay, 1819 Onthophilidae MacLeay, 1819: 25 (with "?"). Type genus: Onthophilus Leach, 1817 (not cited). Onthophilidae Gistel, 1856: 363. Type genus: Onthophilus Leach, 1817. Onthophilina Thomson, 1862: 247. Type genus: Onthophilus Leach, 1817. Scolytini Jacobson, 19 1 1 : 652 (new name for Abraeini (unnecessary); based on preoccupied type genus). Type genus: Scolytus Miiller, 1 764 (not Geoffrey, 1 762; = Onthophilus Leach, 1817). Onthophilinae; Vienna, 1974: 280 (as new subfamily, elevated from tribe). Type genus: Onthophilus Leach, 1817. TRIBALINAE Bickhardt, 1914 Tribalini Bickhardt, 1914: 307 (avail., Art. 12b(4)). Type genus: Tribalus Erichson, 1834. Note: Treated incorrectly as nomen nudum by Mazur (1984). SAPRINOMORPHAE Blanchard, 1845 Saprinites Blanchard, 1 845: 276 (not latinized; avail., Art. 1 lf^iii); see Discussion { 1 }). Type genus: Saprinus Erichson, 1834. Saprinides Lacordaire, 1854: 273 (not latinized; avail., Art. 1 1 f