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^^i. UNITED STATES AGRICULTURE PREPARED BY ñka DEPARTMENT OF HANDBOOK AGRICULTURAL ^S&?f AGRICULTURE NUMBER 529-28 RESEARCH ^^^^ SERVICE FAMILIES OF COLEóPTERA IN AMERICA NORTH OF MEXICO

Fascicle' Family Year issued Fascicle' Family Year issued Fascicle' Family Year issued

1 Cupedidae 1979 46 Callirhipidae 102 Biphyllidae 2 Micromalthidae 1982 47 Heteroceridae 1978 103 Byturidae 1991 3 Carabidae 48 Limnichidae 1986 104 Mycetophagidae 4 Rhysodidae 1985 49 Dryopidae 1983 105 Ciidae ^. 1982 5 Amphizoidae 1984 50 Elmidae 1983 107 Prostomidae 6 Haliplidae 51 Buprestidae 109 Colydiidae 8 Noteridae 52 Cebrionidae 110 Monommatidae 9 Dytiscidae 53 Elateridae 111 Cephaloidae 10 Gyrinidae 54 Throscidae 112 Zopheridae 13 Sphaeriidae 55 Cerophytidae 115 Tenebrionidae 14 Hydroscaphidae 56 Perothopidae 116 Alleculidae 15 57 Eucnemidae 117 Lagriidae 16 Hydrophilidae 58 Telegeusidae 118 Salpingidae 17 Georyssidae 61 Phengodidae 119 Mycteridae 18 Sphaeritidae 62 Lampyridae 120 Pyrochroidae 1983 20 Histeridae 63 Cantharidae 121 Othniidae 21 Ptiliidae 64 Lycidae 122 Inopeplidae 22 Limulodidae 65 1989 123 Oedemeridae 23 Dasyceridae 66 Nosodendridae 124 Melandryidae 24 Micropeplidae 1984 67 Dermestidae 125 Mordellidae 1986 25 Leptinidae 69 Ptinidae 126 Rhipiphoridae 26 70 Anobiidae 1982 127 Meloidae 27 Scydmaenidae 71 Bostrichidae 128 Anthicidae 28 Silphidae 1993 72 Lyctidae 129 Pedilidae 29 Scaphidiidae 74 Trogositidae 130 Euglenidae 30 Staphylinidae 76 Cleridae 131 Cerambycidae 31 Pselaphidae 78 Melyridae 132 Bruchidae 32 Lucanidae 79 133 Chrysomelidae 33 Passalidae 81 Sphindidae 134 Nemonychidae 34 Scarabaeidae 1984 82 Nitidulidae 135 Anthribidae 35 Eucinetidae 83 Rhizophagidae 138 Allocorynidae 1991 36 Helodidae 86 Cucujidae 140 Brentidae 37 90 Cryptophagidae 141 Platypodidae 1979 38 Dascillidae 92 Languriidae 1983 142 Scolytidae 39 Rhipiceridae 93 Erotylidae 143 Curculionidae 1983 40 Byrrhidae 94 Phalacridae 144 Stylopidae 41 Psephenidae 1983 95 Cerylonidae 1982 145 Fossil Coleóptera 42 Brachypsectridae 96 Corylophidae 146 Ithyceridae 43 Artematopidae 97 Coccinellidae 44 98 Endomychidae 1986 45 Chelonariidae 100 Lathridiidae

^Missing numbers are those assigned in the computer program to families not found in the United States and Canada.

Microfiche or printed copies of this publication may be purchased from the National Technical Information Service, 5285 Port Royal Road, Springfield, VA 22161. For additional information, contact NTIS at the address above.

All programs and services of the U.S. Department of Agriculture are offered on a nondiscriminatory basis without regard to race, color, national origin, religion, sex, age, marital status, or handicap. A CATALOG OF THE COLEÓPTERA OF AMERICA NORTH OF MEXICO FAMILY: SILPHIDAE

By S. B. PECK S.E. MILLER DEPT. OF BIOLOGY BISHOP MUSEUM CARLETON UNIVERSITY P.O. BOX 19000A OTTAWA, ON K1S 5B6 HONOLULU, HI 9681 7-0916 CANADA

£Sß^ UNITED STATES AGRICULTURE P^3F^3^J¿.^X DEPARTMENT OF HANDBOOK AGRICULTURAL i^' AGRICULTURE NUMBER 529-28 RESEARCH w SERVICE

September 1993

FOREWORD

Many species of are important pests of agricultural crops, stored food products, forests, wood products and structures, and fabrics. Many other species, in contrast, are beneficial in the biological sup- pression of pest and weeds, as well as in the of plant , carcasses, and dung. Part of our national responsibility to American agriculture is to provide correct identification of species of American beetles so that appropriate controls can be applied. Most information about animal species, whether agricultural, biological, or experimental, is filed under the species' scientific names. These names are therefore the keys to retrieval of such information. Because some species have been known by several names, a complete listing of these names for each species is neces- sary. For the user of scientific names, an up-to-date taxonomic catalog providing currently accepted names and pertinent bibliographic and distributional data is an indispensable tool. Although taxonomic literature is constantly changing to reflect current work, the traditional published taxonomic catalog remains static with updating left to the individual user until it is revised. Production of catalogs in the past has been laborious with long printing delays resulting in data that are obsolete before being published. However, the computer now provides the capability of storing, updating, and retrieving taxonomic data; rapid publica- tion through computer-driven typesetting machinery; and a greater degree of currentness and flexibility. All the fascicles in this catalog of the beetles of America north of Mexico are produced by an ori- ginal group of computer programs, designed and written during a pilot project by personnel of the Systematic Entomology Laboratory and the Communication and Data Services Division, Agricultural Research Service.

R. D. Plowman Administrator Agricultural Research Service

PREFACE

The Coleóptera, or beetles, are represented in the world by about 220,000 described species, of which about 24,000 occur in the United States and Canada. A comprehensive taxonomic catalog of beetles for this area has not been available except the series of world-based ''Coleopterorum Catalogus" volumes ( 1909-present, Junk, Berlin). The Leng "Catalogue of the Coleóptera of America North of Mexico" (J.D. Sherman, Jr., Mt. Vernon, NY), which was published in 1920 with supplements to the end of 1947, is a checklist. However, it has served professional and amateur alike for nearly 60 years as the principal source of scientific names of beetles. Since 1947, many new taxa have been described and many changes in status and nomenclature have appeared in numerous scattered publications, but little effort has been made to summarize these changes. This catalog will supplant the Leng catalog and supply additional essential information. It is produced by an original suite of storage, retrieval, and printing programs written especially for automated taxonomic catalogs. The catalog for each family is published as a separate fascicle with its introductory text, bibliography, and sequence. The publishing of separate fascicles makes data available shortly after they are assembled. Computer tapes for each fascicle are maintained for updating and necessary reprinting. The information on each family is the responsibility of the respective author or authors. The editors modify it only to correct obvious errors and to make it conform to the requirements of the computer programs. No original proposal for a new name, taxon, status, or classification is given, such data having been previously published, but new host and distributional data are often listed. The rules of "The International Code of Zoological Nomenclature" are followed. The geographic scope of this catalog includes the continental United States, Canada, Greenland, and the associated continental islands. Names of taxa found only in other regions are excluded. If the range of a species extends outside these geographic limits, this fact is indicated. On (or inside oO the back cover is a map of the 12 faunal regions based on historical and faunal criteria to simplify distribution recordings. Two-letter Postal Service style abbreviations are used for States and Provinces, and faunal regions are indicated in each distribution record by a diagonal line between groups of abbreviations. It is not the purpose of this catalog to present a complete scheme of higher classification within the order. The familial makeup is somewhat intermediate between that of R.H. Amett in "The Beetles of the United States" (1960-62, Catholic University Press, Washington, DC) and that of R.A. Crowson in "The Natural Classification of the Families of Coleóptera" (1967, Biddies Ltd., Guildford, England). Modifications of these two systems are largely those advocated by J.F. Lawrence based in part on suggestions by taxonomic specialists for certain families. Generic groups and higher categories within the family are arranged phylogenetically as indicated by the author of the particular fascicle, and species group names with their respective synonyms are arranged alphabeti- cally. Names referable to incertae sedis and nomen dubium are listed separately at the end of the nearest appli- cable taxon with notations as to their status. Each available name is followed by its author, date proposed, and page number referring to the complete bibliographic citation containing the original description. Following each generic name are the type-species and method of its designation, necessary explanatory notes, and pertinent references on immature stages, , redescription, ecology, and keys. After the specific name entry are the original genus (if different from the present placement), type-locality, geographical distribution by State, Province, and broad extralimital units, explanatory notes, pertinent references to immature stages, taxonomy, redescription, and ecology, depository of type-specimen and its sex, and hosts. In addition to the list under the map (on or inside of the back cover) of faunal regions, the following abbreviations are used in this catalog:

ABBREVIATIONS, GENERAL

Amer. Bor.—America Borealis Mus.—Museum Amer. Sept.—America Septentrionalis N. Amer.—North America Automat.—Automatic Orig. des.—Original designation C. Amer.—Central America Preocc.—Preoccupied Co.—County S. Amer.—South America Cosmop.—Cosmopolitan Sp.—Species Design.—Designated Subseq. monot.—Subsequent monotypy F.—Female Subsp.—Subspecies Holarc.—Holarctic Taut.—Tautonymy Isl.—Island Univ.—University M.—Male USA—United States of America Mex.—Mexico Var.—Variety Monot.—Monotypy W. Ind.—West Indies

MUSEUMS IN THE UNITED STATES AND CANADA

AMNH—American Museum of Natural History, FSCA—Florida State Collection, Gainesville New York HAHC—H. & A. Howden Collection, Ottawa, ANSP—Academy of Natural Sciences, Phila- Canada delphia, PA ICCM—Carnegie Museum, Pittsburgh, PA BPBM—Bernice P. Bishop Museum, Honolulu INHS—Illinois Natural History Survey, Urbana BYUC—Brigham Young University, Provo, UT JGEC—J. G. Edwards Collection, San Jose, CA CASC—California Academy of Sciences, San KMFC—K. M. Fender Collection, McMinnville, Francisco OR CISC—University of California, Berkely KSUC—Kansas State University, Manhattan CNCI—Canadian National Collections, Ottawa LACM—Los Angeles County Museum, CA CUIC—Cornell University, Ithaca, NY LSUC—Louisiana State University, Baton Rouge CWOB—C. W. O'Brien Collection, Tallahassee, FL MCZC—Museum of Comparative Zoology, DHKC—D. H. Kistner Collection, Chico State Harvard University, Cambridge, MA College, CA MSUC—Michigan State University, East Lansing ELSC—E. L. Sleeper Collection, Long Beach, CA NCSM—North Carolina State University, Raleigh FMNH—Field Museum of Natural History, Chicago, NYSM—New York State Museum, Albany IL OSEC—Oklahoma State University, Stillwater OSUC—Ohio State University, Columbus OSUO—Oregon State University, Corvallis

'Abbreviations for U. S. and Canadian museums abridged from Arnett, R. H., Jr., and Samuelson, G. A., 1969, "Directory of Coleóptera Collections of North America (Canada Through Panama)," Cushing-Malloy, Ann Arbor, MI, 123 pp. PMNH—Peabody Museum, Yale University, SMSH—Stovall Collection, University of New Haven, CT Oklahoma, Norman PSUC—Pennsylvania State Museum, University TAMU—Texas A. & M. University, College Park Station PURC—Purdue University, West Lafayette, IN UCDC—University of California, Davis RUIC—Rutgers University, New Brunswick, NJ UICM—University of Idaho, Moscow SEMC—Snow Museum, University of Kansas, UMMZ—University of Michigan, Ann Arbor Lawrence UMRM—University of Missouri, Columbia SJSC—San Jose State College, CA USNM—U.S. National Museum of Natural SLWC—S. L. Wood Collection, Provo, UT History, Washington, DC WSUC—Washington State University, Pullman

MUSEUMS IN FOREIGN COUNTRIES

BMNH—British Museum (Natural History), NMPC—Narodni Museum, Prague, London Czechoslovakia GUHC—Glasgow University, Hunterian SCUT—Spinola College, University of Turin, College, Scotland Italy HMOX—Hope Museum, Oxford, England SMTD—Staatliches Museum für Tierkunde, IPZE—Institut Pflanzenschutzforschung Dresden, East Germany Zweigstelle, Eberswald, East Germany UNAM—Universidad Nacional Autónoma, IRSB—Institut Royal Sciences Belgique, Mexico City Brussels UZMC—University Zoological Museum, MFNB—Museum für Naturkunde (Humboldt), Copenhagen, Denmark Berlin UZMH—University Zoological Museum, MGFT—Museum G. Frey, Tutzing, Munich, Helsinki, Finland West Germany ZMAS—Zoological Museum, Academy of MHNL—Museum d'Histoire Naturelle, Lyon, Sciences, Leningrad France ZMPA—Zoological Museum, Polish Academy of MNHP—Museum National d'Histoire Naturelle, Sciences, Warsaw Paris ZMUL—Zoological Museum, University of MNSL—Museum of Natural Sciences, Leipzig, Lund, Sweden East Germany ZMUM—Zoological Museum, University of MZBS—Museum Zoologia, Barcelona, Spain Moscow NHRS—Naturhistoriske Riksmuseet, Stockholm ZSBS—Zoologische Sammlung Bayerischen Staates, Munich, West Germany ACKNOWLEDGMENTS

We are indebted to many individuals who contributed to the planning and development of this catalog. We are especially grateful to the following specialists who helped to make it as complete and accurate as possible: Richard H. Foote, Systematic Entomology Laboratory (SEL), Agricultural Research Service (ARS), for his sug- gestions, guidance, and encouragement; C. W. Sabrosky, SEL, for valuable counsel on nomenclatural problems; J. F. Lawrence, Division of Entomology, Commonwealth Scientific and Industrial Research Organization, Can- berra, Australia, for his recommendations on higher categories; and more than 50 coleopterists in Canada, the United States, and Mexico for voluntarily contributing information about their specialty groups. We wish to acknowledge the extensive computer-programming and editing-system support provided by members of the former Communications and Data Services Division of ARS during the eariy years of this project. We also thank Elaine Jamison for the data entry necessary for each fascicle.

R. D. Gordon, editor in chief J. C. Barch, associate editor F. C. Thompson, computer program advisor Systematic Entomology Laboratory, Agricultural Research Service Washington, DC

Editorial Board

J. M. Campbell, Biosystematics Research Institute, Agriculture Canada, Ottawa, ON T. L. Erwin, Department of Entomology, Smithsonian Institution, Washington, DC H. F. Howden, Department of Biology, Carieton University, Ottawa, ON J. M. Kingsolver, Florida State Collection of Arthropods, Gainesville, FL P. J. Spangler, Department of Entomology, Smithsonian Institution, Washington, DC T. J. Spilman, Systematic Entomology Laboratory, ARS, Washington, DC R. E. White, Systematic Entomology Laboratory, ARS, Washington, DC

CONTENTS

Family Silphidae 1 Tribe Agyrtini 2 Tribe Silphini 3 Tribe Nicrophorini 5 Bibliography IQ Index 23 Family SILPHIDAE By S. B. Peck and S. E. Miller Silphidae, the so called beetles, are from 5 to 35 mm long, are often brightly col- ored, and usually are found feeding on dead or sometimes larvae. Some, however, seem more associated with decaying fungi or vegetable matter, and one genus is strictly phy- tophagous. The attractive red or orange and black Nicrophorus species are called burying bee- tles because they can rapidly excavate the soil from under a dead mouse or other small animal until it sinks into the ground. Parental care of the larvae makes Nicrophorus one of the most social of all beetles (work of Pukowski, abstracted in Balduf, 1935; Milne and Milne, 1944, 1976). The Silphini do not show such parental care. are carrion feeders, Lyrosoma are littoral, and and Apteroloma are montane and probably predatory. Little is known of the biology of the Agyrtini. Limited aspects of silphid biology in restricted areas have been studied, but some reports are unreliable because of confused nomenclature. Olfactory sensitivity, of primary importance in locating carrion, is acute, and in electrophysiological studies, the sensory receptors and spe- cific decay compounds that activate them have been identified. Many species that live in one area can ''divide'' the carrion resource by differing in seasonality, diel periodicity, or habitat preferences (Anderson, 1982c; Shubeck, 1971; Wilson and Fudge, 1984; Wilson and Knollenberg, 1984; Wilson, Knollenberg, and Fudge, 1984). Carrion-baited pitfall traps are best for collecting silphids. They often can be used as experimental systems to answer specific ecological questions (Scott, et al., 1987; Shubeck, 1976a, 1976b). Silphids, biologically important in maintaining ecosystem health and productivity, aid in the recycling of organic matter. While engaged in this process, they reduce the frequency and suitability of carrion as a site of fly breeding and development. Their feeding may destroy po- tential infection foci of pathogenic bacteria or parasitic worms. Rabies virus, but not anthrax bacteria, is inactivated in the gut of Nicrophorus. Silphids phoretically carry many species of and nonpathogenic as commensals. There has been much confusion in the use of generic and specific names for the silphids of North America. For instance, the names Nicrophorus and Necrophorus have interchangeably been used by authors, but Nicrophorus is considered correct. For the 15 species of Nicrophorus of the United States and Canada, over 110 names have been applied, including about 65 used as variations and aberrations with no taxonomic status. Reviews and revisions (Madge, 1980; and Miller and Peck, 1979), in combination with this catalog, should bring the fauna into - ary perspective with the world fauna and should help establish nomenclatorial stability. A de- tailed study of adult and larval characters of Agyrtini, with about 45 world species (Lawrence, 1982; Lawrence and Newton, 1982; and Newton, unpublished), shows that this tribe should be elevated to family status and placed near the Leiodidae. Lawrence and Newton (1982) also indi- cate that the remaining two tribes-Silphini with about 90 world species and Nicrophorini with about 68 world species-could be united as a subfamily of the Staphylinidae. Madge (1980) uses a different system (, Nicrophorinae, Trigonurinae, and Agyrtinae) for the Silphidae, which he regards as a valid family. Other important general works on North American silphids are by Anderson (1982a), Anderson and Peck (1985), Arnett (1944), Hatch (1927a, 1927b, 1928, 1957), Madge (1980), Miller and Peck (1979), Peck and Anderson (1985), Peck and Kaulbars (1988), and Peck and Miller (1982). Keys to identify the genera and species are in Anderson and Peck (1985) and Peck (1989). Several species recorded in the literature as occurring in North America represent unes- tablished introductions of palaearctic species or mislabelings. These species, which are not in- cluded in the catalog, are quadrimaculara (Scopoli, 1772), Nicrophorus germanicus (Linnaeus, 1758), N, humator (Gleditsch, 1767), N vespillo (Linnaeus, 1758), tristis ill- liger, 1798), and S. tyrolensis (Laicharting, 1781). Additional information may be found in Arnett (1944), Hatch (1927b), and Miller and Peck (1979). Nicrophorus grandior Angelí (1912) was proposed for a specimen of N germanicus (lectotype by Peck and Miller, 1982) and for a specimen of N humator, both mistakenly labeled ''California.'' Eurasian synon5niis of holarctic genera and species (not used in the North American litera- ture and not included here) may be found in Hatch (1928). We have not included aberrations and varieties named by Hatch (1957) because he stated that these ''do not refer to discrete pop- 2 COLEÓPTERA CATALOG ulations but to segregates of the same/' Varietal names proposed by Portevin are included, however, as they may have been meant in a subspecific sense. R. S. Anderson, R. B. Madge, and A. F. Newton contributed to the preparation of this cata- log. This manuscript was received October 1981, modified September 1986, and amended No- vember 1992. Tribe AGYRTINI Genus NECROPHILUS Latreille Necrophilus Latreille, 1829: 500. Type-species: Silpha subterránea Dahl (monot.). Necrobius Gistel, 1834: 147. Type-species: Silpha subterránea Dahl (monot.). hydrophiloides Guerin-Meneville, 1835: pi. 17, fig. 12. "File de Norfolk, dans TAmerique boréale''; AK/ BC WA OR ID/ CA. First published as a named figure in the text (Guerin-Meneville, 1844: 61). ater Motschulsky, 1845: 363. AK. IMMATURE STAGES: Blaisdell, 1901: 263. pettiti Horn, 1880: 243. ON: Grimsby; MI ON/ MO IL IN OH KY/ NY VA/ LA AL TN GA SO NC FL. The lectotype was designated in Peck, 1981: 181. TYPE DEPOSITORY: MCZC. SEX OF TYPE: F. ECOLOGY: Peck, 1981: 182. Genus IPELATES Reitter Ipelates Reitter, 1885: 89, 90. Type-species: Hadrambe latissimus Reitter (monot.). Pelâtes Horn, 1880: 244 (preoccupied by Cuvier and Valenciennes, 1829). Type-species: Necrophilus latus Mannerheim (monot.). Sphaeroloma Portevin, 1905: 422. Type-species: Sphaeroloma sikkimensis Portevin (orig. des.). Pelatines Cockerell, 1906: 240 (replacement name for Pelâtes Horn). Type-species: Necrophilus latus Mannerheim (automat.). Brachyloma Portevin, 1914: 214. Type-species: Brachyloma curtum Portevin (design, by Hatch, 1928: 70) = striatipennis (Lewis). Necrophilodes Champion, 1923: 49. Type-species: Necrophilus castaneicolor Champion (orig. des.). Misspelled Necrophiloides by Hatch, 1927b, p. 333 and Hlisnikovsky, 1963, p. 314. TAXONOMY: Schawaller, 1983: 101. latus Mannerheim, 1852: 331 {Necrophilus), AK: Sitkha; AK NT/ BC WA OR/ AB/ CA. Genus Froelich Agyrtes Froelich, 1799: 15 (nominate subgenus extra-limital). Type-species: Mycetophagus castaneus Fabricius (design, by Latreille, 1810: 427). TAXONOMY: Hlisnikovsky, 1964b: 275; Peck, 1974: 501. Subgenus AGYRTECANUS Reitter Agyrtecanus Reitter, 1901: 102. Type-species: Agyrtes bicolor Laporte (monot.). Lendomus Casey, 1924: 184. Type-species: Lendomus politus Casey (monot.) = longulus (LeConte). longulus LeConte, 1859a: 282 {Necrophilusl CA: San Francisco, Table Mt.; AK/ BC WA OR ID/ CA. The lectotype was designated by Peck, 1974: 501. TYPE DEPOSITORY: MCZC. politus Casey, 1924: 185 {Lendomus). PQ: Anticosti Isl. Error of nomenclature and distribution from erroneous specimen data (Peck, 1974: 504). TYPE DEPOSITORY: USNM. TAXONOMY: Peck, 1974: 501. SILPHIDAE 3

similis Fall, 1937: 29. CA: Pasadena; CA. TYPE DEPOSITORY: MCZC. TAXONOMY: Peck, 1974: 504. Genus LYROSOMA Mannerheim Lyrosoma Mannerheinl, 1853: 174. Type-species: Pteroloma pallidum Eschscholtz (design, by Hatch, 1928: 71). TAXONOMY: Hlisnikovsky, 1964a: 39. opacum Mannerheim, 1853: 175. AK: Atkha, Aleutian Isl.; St. Paul, Pribilof Isl.; AK; Other: Siberia. HOST: Supposedly living in rotting kelp, at times completely submerged by tides (Van Dyke, 1921: 161). Also recorded from nests of maritime birds (Ferris, 1918: 125). Genus PTEROLOMA Gyllenhal Pteroloma Gyllenhal, 1827: 418. Type-species: Harpalus forsstromii Gyllenhal (monot.). Adolus Fischer von Waldheim, 1828: 242. Type-species: Adolus brunneus Fischer von Waldheim (monot.) = forsstromii (Gyllenhal). Holocnemis Schilling, 1829: 93. Type-species: Holocnemis grauenhorstii Schilling (monot.) = forsstromii (Gyllenhal). TAXONOMY: Madge, 1980: 354. nebrioides Brown, 1933: 213. AB: Waterton; BC ID/ AB MT. TYPE DEPOSITORY: CNCI. SEX OF TYPE: F. ECOLOGY: Johnson, 1987: 381. Genus APTEROLOMA Hatch Apteroloma Hatch, 1927a: 12 (as subgenus). Type-species: Necrophilus tenuicorne LeConte (orig. des.). TAXONOMY: Van Dyke, 1928: 19; Bolivar y Pieltain and Hendrichs, 1972: 207; Miller and Peck, 1979: 89. KEYS: Van Dyke, 1928: 21; Bolivar y Pieltain and Hendrichs, 1972: 210; Miller and Peck, 1979: 98. arizonicum Van Dyke, 1928: 24 (Pteroloma). AZ: near Prescott; AZ NM TX. TYPE DEPOSITORY: CASC. caraboides Fall, 1907: 235 (Pteroloma). WA: Wenatchee; CA: Mt. San Antonio. TYPE DEPOSITORY: MCZC. tahoecum Fall, 1927: 136 (Pteroloma). CA: Lake Tahoe; OR/ CA NV. TYPE DEPOSITORY: MCZC. SEX OF TYPE: F. tenuicornis LeConte, 1859b: 84 (Necrophilus). WA: Puget Sound. TYPE DEPOSITORY: MCZC. SEX OF TYPE: F. TAXONOMY: Peck and Miller, 1982: 154. Tribe SILPHINI Genus Leach Necrodes Leach, 1815: 88. Type-species: Silpha littoralis Linnaeus (design, by Curtis, 1830: pi. 334). Asbolus Bergroth, 1884: 229 (unnecessary replacement name for Necrodes Leach). Type- species: Silpha littoralis Linnaeus (automat.). Protonecrodes Portevin, 1922: 508. Type-species: Silpha surinamensis Fabricius (design, by Hatch, 1928: 123). TAXONOMY: Madge, 1980: 354. surinamensis Fabricius, 1775: 72 (Silpha). southern (North) America; BC WA OR ID/ AB SK MB MT SD/ MN WI MI ON PQ/ NK PE NS NF/ UT/ NE KS lA MO IL IN OH 4 COLEÓPTERA CATALOG

KY/ NY PA NJ DE MD DC WV VA/ ME NH VT MA RI CT/ TX OK/ AR LA MS AL TN GA SC NC FL. TYPE DEPOSITORY: GUHC. IMMATURE STAGES: Dorsey, 1940: 130; Ratcliffe, 1972: 366. TAXONOMY: Ratcliffe, 1972: 364. ECOLOGY: Ratcliffe, 1972: 380; Eisner and Meinwald, 1983: 358. Genus Gistel Oxelytrum Gistel, 1848: 190. Type-species: Oxelytrum aequinoctiale Gistel (orig. des.) = cayennense (Sturm). Oxelytrium (error). {In Gistel, 1857: 545, 566). Oxelytron (error). {In Gistel, 1857: 546). Hyponecrodes Kraatz, 1876: 357, Type-species: Silpha cayennensis Sturm (design, by Madge, 1980: 355). Katanecrodes Schouteden, 1905: 195. Type-species: Silpha apicalis BruUe (design, by Hatch, 1928: 113). Paranecrodes Portevin, 1921b: 81. Type-species: Necrodes biguttatus Philippi (monot.). TAXONOMY: Madge, 1980: 357. discicolle BruUe, 1836: 75, pi. 5, fig. 4 {Silpha). Bolivia: near Cochabamba; TX; Mex., C. Amer., S. Amer. analis Chevrolat, 1843: 26 {Necrodes). Mex.: ''Orixaba''. discretas Portevin, 1921b: 84 {Hyponecrodes) (as variety). '^Guatemala, Colombie, Venezuela, Brezil, Paraguay''. The stated type-locality also includes ''Argentine et Pérou'' (Portevin, 1921b: 84). TAXONOMY: Davis, 1980: 249; Peck and Anderson, 1985: 263. Genus Leach Thanatophilus Leach, 1815: 89. Type-species: Silpha sinuata Fabricius (monot.). Pseudopelta Bergroth, 1884: 229 (unnecessary nomen nudum). Type-species: Silpha sinuata Fabricius (monot.). Philas Portevin, 1903: 331. Type-species: Silpha truncata Say (monot.). Silphosoma Portevin, 1903: 333. Type-species: Silpha metallescens Fairmaire (monot.). Chalcosilpha Portevin, 1926: 31. Type-species: Silpha micans Fabricius (design, by Hatch, 1928: 79). TAXONOMY: Schawaller, 1981: 3. coloradensis Wickham, 1902: 180 {Silpha). CO: Argentine Pass, nr. Georgetown, over 12,000 ft; AK/ BC/ MT/ WY UT CO/ NM. TYPE DEPOSITORY: USNM. SEX OF TYPE: M. obalskii Portevin, 1920b: 507 {Silpha). BC: Lake Teslin. ECOLOGY: Bjorkman and Hatch, 1939: 96; Peck and Anderson, 1982: 112. lapponicus Herbst, 1793: 209, pi. 52, fig. 4 {Silpha). Lappland; AK YT NT GL/ BC WA OR ID/ AB SK MB MT ND SD/ MN WI MI ON PQ/ NS NF/ CA/ WY UT CO/ NE KS lA IL/ PA DC/ NH RI/ AZ NM; Mex., Holarc. See Hatch (1928: 80) for Eurasian synonyms. caudata Say, 1823: 192 {Silpha). ^'Missouri Territory". The neotype was not designated (Peck and Miller, 1982: 154). tuberculata Germar, 1824: 81 {Silpha). "North America". californica Mannerheim, 1843: 253 {Silpha). CA. IMMATURE STAGES: Dorsey, 1940: 126. ECOLOGY: Korobeinikov and Esyunin, 1984: 40. sagax Mannerheim, 1853: 173 {Silpha). AK: Kaktnu River, Kenai Peninsula; AK YT NT/ BC WA OR ID/ AB SK MB ND SD/ ON PQ/ WY CO. The species was long considered a junior synonym of Thanatophilus trituberculatus (Kirby, 1830). TAXONOMY: Miller and Peck, 1979: 91. SILPHIDAE 5 trituberculatus Kirby, 1837: 101 (). '^Boreal America''; AK NT/ AB SK MB/ ON. The specific type-locality is between New York and Cumberland House (SK). IMMATURE STAGES: Anderson, 1987: 34. TAXONOMY: Miller and Peck, 1979: 91. truncatus Say, 1823: 193 (Silpha). "eastern Colorado''; CO/ NE KS/ AZ NM TX; Mex. The neotype was designated by Peck and Miller, 1982: 154. TYPE DEPOSITORY: MCZC. SEX OF TYPE: M. Genus OICEOPTOMA Leach Oiceoptoma Leach, 1815: 89. Type-species: Silpha thoracica Linnaeus (monot.). Oeceoptoma Agassiz, 1847: 256 (unjustified emendation). Type-species: Silpha thoracica Linnaeus (monot.). Isosilpha Portevin, 1920a: 398. Type-species: Eusilpha hypocrita Portevin (monot.). IMMATURE STAGES: Dorsey, 1940: 123. inaequale Fabricius, 1781: 87 (Silphal "North America"; ND SD/ MN WI MI ON PQ/ CO/ KS MO IL IN OH KY/ NY PA NJ MD/ MA RI CT/ TX OK/ AR LA AL TN GA NC FL noveboracense Forster, 1771: 17 (Silphal ^^Northern North America"; AB SK MB MT ND SD/ MN WI MI ON PQ/ NS/ WY/ lA MO IL IN OH KY/ NY PA NJ DC WV VA/ ME NH VT MA RI CT/ TX OK/ AR MS TN GA NC. marginalis Fabricius, 1777: 215 (Silpha). "Boreal America". ECOLOGY: Anderson, 1989: 18. rugulosum Portevin, 1903: 333 (Silpha). GA: Savannah; IL IN/ NY/ TX OK/ AL GA SC NC FL. This name was erroneously put into synon5any under O. inaequale by Hatch, 1927b: 343. The lectotype was designated by Peck and Kaulbars, 1988: 50. TYPE DEPOSITORY: MNHP. Genus Kirby and Spence Necrophila Kirby and Spence, 1828: 509. Type-species: Silpha americana Linnaeus (orig. des.). Necrobora Hope, 1840: 151 (unnecessary name). Tjrpe-species: Silpha americana Linnaeus (orig. des.). Necrotropha Gistel, 1848: 121. Type-species: Silpha americana Linnaeus (monot.). Eusilpha Semenov-Tian-Shanskij, 1891: 299. Type-species: Silpha japónica Motschulsky (design, by Hatch, 1928: 111). Calosilpha Portevin, 1920a: 396. Type-species: Silpha ioptera KoUar and Redtenbacher (design, by Hatch, 1928: 113). Deutosilpha Portevin, 1920a: 396. Type-species: Silpha rufithorax Wiedemann (monot.). Chrysosilpha Portevin, 1921a: 538. Type-species: Silpha formosa Laporte (design, by Hatch, 1928: 112). Deuterosilpha (error). (In Hatch, 1928: 112). americana Linnaeus, 1758: 360 (Silpha). '^America" (north); MB SD/ MN WI MI ON PQ/ NS/ lA MO IL IN OH KY/ NY PA NJ DE MD WV VA/ NH MA RI CT/ TX OK/ AR AL TN GA SC FL. peltata LeConte, 1853: 279 (Silpha). PA. Name first published as Scarabaeus peltatus by Catesby (1748: pi. 10, fig. 7) and repeated by Edwards (1771), but suppressed by Int. Comm. Zool. Nomen. Opinion 259. Name first validated by LeConte (1853: 279). affine Kirby, 1837: 103 (Oiceoptoma). NS. canadense Kirby, 1837: 104 (Oiceoptoma). ON: Lake Huron. terminatum Kirby, 1837: 103 (Oiceoptoma). NS. IMMATURE STAGES: Dorsey, 1940: 126. ECOLOGY: Fisher and Tuckerman, 1986: 21; Anderson, 1989: 18. Genus HETEROSILPHA Portevin Heterosilpha Portevin, 1926: 83. Type-species: Silpha ramosa Say (design, by Hatch, 1928: 100). 6 COLEÓPTERA CATALOG

TAXONOMY: Miller and Peck, 1979: 92. aenescens Casey, 1886: 171 (Silpha), CA: San Francisco; OR/ CA; Mex. The lectotype was designated by Miller and Peck, 1979: 93. TYPE DEPOSITORY: USNM. SEX OF TYPE: M. REDESCRIPTION: Miller and Peck, 1979: 93. ramosa Say, 1823: 193 {Silphal "the upper Missouri'* (River); NT/ BC WA OR ID/ AB SK MB MT ND SD/ MN WI MI ON/ CA NV/ WY UT CO/ NE/ AZ NM; Mex. The neotype was designated by Miller and Peck, 1979: 93. TYPE DEPOSITORY: MCZC. SEX OF TYPE: M. cervaria Mannerheim, 1843: 252 (Silpha). CA. IMMATURE STAGES: Dorsey, 1940: 129; Brewer and Bacon, 1975: 787. REDESCRIPTION: Miller and Peck, 1979: 93. ECOLOGY: Brewer and Bacon, 1975: 786.

Genus Reitter Aclypea Reitter, 1885: 81. Type-species: Peltis undata Mueller (design, by Hatch, 1928: 93). This name is used in reference to Blitophaga following Seidlitz (1888: 311), who is regarded as first reviser in accordance with Article 24 (a) (i) of the International Code of Zoological Nomenclature (Miller and Peck, 1979: 90). Blitophaga Reitter, 1885: 82. Type-species: Silpha opaca Linnaeus (design, by Hatch, 1928: 95). TAXONOMY: Anderson and Peck, 1984: 248. bituberosa LeConte, 1859c: 6 (Silpha), WY: near Ft. Bridger; NT/ BC WA OR ID/ AB SK MB MT ND/ CA/ WY UT CO/ NE KS. TYPE DEPOSITORY: MCZC. SEX OF TYPE: F. IMMATURE STAGES: Cooley, 1917: 97; Anderson and Peck, 1984: 254. TAXONOMY: Peck and Miller, 1982: 154. ECOLOGY: Cooley, 1917: 95; Anderson and Peck, 1984: 251. HOST: Adults and larvae feed on Monolepis nuttalliana (Chenopodiaceae) and related native and introduced plants, such as beet, sugarbeet, spinach, pumpkin, squash, etc. (Hatch, 1957: 10). opaca Linnaeus, 1758: 361 (Silpha). Europe; AK YK NT; Other: Eurasia. Older records of ^4. bituberosa are often erroneously under this name (Miller and Peck, 1979: 90). ECOLOGY: Anderson, 1989.

Tribe NICROPHORINI Gtfenus NICROPHORUS Fabricius Nicrophorus Fabricius, 1775: 71. Type-species: Silpha uespillo Linnaeus (design, by Latreille, 1810: 427). Many species of Nicrophorus were described under the unjustified emendation Necrophorus (Herman, 1964: 5; Madge, 1980: 357). Designation of the lectotype by Latrielle was for the emendation Necrophorus, Necrophorus Thunberg, 1789: 7 (unjustified emendation). Type-species: Silpha vespillo Linnaeus (design, by Latreille, 1810: 427). Necrophagus (error) Leach. 1815: 88. Cyrtoscelis Hope, 1840: 149. Type-species: Silpha vespillo Linnaeus (orig. des.). Acanthopsilus Portevin, 1914: 223. Type-species: Necrophorus concolor Kraatz (monot.). Necrocharis Portevin, 1923: 68. Tjrpe-species: Silpha Carolina Linnaeus (monot.). Eunecrophorus Semenov-Tian-Shanskij, 1933: 152. Type-species: Olivier (orig. des.). Necrocleptes Semenov-Tian-Shanskij, 1933: 153. Type-species: Silpha humator Gleditsch (orig. des.). Necropter Semenov-Tian-Shanskij, 1933: 154. Type-species: Necrophorus investigator Zetterstedt (orig. des.). SILPHIDAE 7

Stictonecropter Semenov-Tian-Shanskij, 1933: 154. Type-species: Necrophorus pustulatus Herschel (orig. des.). Neonicrophorus Hatch, 1946: 99. Type-species: Silpha germánica Linnaeus (orig. des.). IMMATURE STAGES: Anderson, 1982a: 256. TAXONOMY: Portevin, 1926; Hatch, 1927a; Herman, 1964: 6; Madge, 1980: 357. ECOLOGY: Milne and Milne, 1944: 312, and 1976: 87; Wilson, 1983: 57; Anderson, 1982a: 249, and 1982c: 1315. americanus Olivier, 1790: 6, pi. 1, fig. 3. '^North America'^; MN WI MI ON PQ/ NS/ NE KS lA MO IL IN OH KY/ NY PA NJ MD DC VA/ ME NH MA RI CT/ AR MS TN GA SCNC. grandis Fabricius, 1792: 247. ''North America''. virginicus Froelich, 1792: 123. VA. ECOLOGY: Davis, 1980: 248; Anderson, 1982b: 363; Hecht, 1988: 39618; Kozol et al., 1989: 167. carolinus Linnaeus, 1771: 530 (Silpha). "Carolina"; AB/ WY CO/ NE KS IL/ NY MD VA/ AZ NM TX OK/ LA MS AL TN GA SC NC FL. mediatus Fabricius, 1801: 334 (Nicrophorus). "Carolina". mysticallis Angelí, 1912: 307. AZ: Tuba. TYPE DEPOSITORY: USNM. defodiens Mannerheim, 1846: 513. AK: Sitka; AK YK NT/ BC WA OR ID/ AB SK MB MT SD/ MN WI MI ON QB/ NK PE NS NF/ CA NV/ WY CO/ NY NJ WV/ ME NH VT MA/ TN NC. pollinctor LeConte, 1854: 19 (junior homonym of Nicrophorus pollinctor Mannerheim, 1853). OR, WA. The lectotype was designated by Peck and Miller, 1982: 152. TYPE DEPOSITORY: MCZC. SEX OF TYPE: M. conversator Walker, 1866: 320. BC: ^'Slecy Station'\ plagiatus Motschulsky, 1869: 352. CA. lateralis Portevin, 1903: 330 (as variety of vespilloides), CA. IMMATURE STAGES: Leech, 1934: 37. TAXONOMY: Leech, 1937: 156; Miller and Peck, 1979: 95; Peck and Miller, 1982: 152. ECOLOGY: Leech, 1934: 36. guttula Motschulsky, 1845: 53. AK: Sitka; BC WA OR ID/ AB SK MT ND SD/ MN/ CA NV/ WY UT CO/ NE/ AZ NM; Mex. hecate Bland, 1865: 382. ''Colorado Territory''. The lectotype was designated by Miller and Peck, 1979: 97. TYPE DEPOSITORY: ANSP. punctostriatus Pierce, 1949: 66 (as subspecies of guttula). CA: Los Angeles Co., Rancho La Brea. The Rancho La Brea specimens are of the Pleistocene Age (Pierce, 1949: 55). TYPE DEPOSITORY: LACM. immaculosis Hatch, 1957: 15. CA. This name was used as an aberration by Hatch, 1927b: 362 but as a subspecies by Hatch, 1957: 15; it takes the authorship and date of use as a subspecies (International Code of Zoological Nomenclature, Art. 10b: p. 10). TYPE DEPOSITORY: IPZE. TAXONOMY: Miller and Peck, 1979: 97; Peck and Miller, 1982: 153; Anderson and Peck, 1986: 286. hybridus Hatch and Angelí, 1925: 216. MB: Aweme; BC WA OR ID/ AB MB MT ND/ MN/ WY UT CO/ NE lA/ AZ NM. TYPE DEPOSITORY: USNM. investigator Zetterstedt, 1824: 151. Sweden; AK YT/ BC WA OR ID/ AB MT/ MN ON PQ/ NS NF/ WY UT CO/ ME/ AZ NM; Helare. See Hatch (1928: 139) for Eurasian synonyms. maritimus Guerin-Meneville, 1835:, pi. 17, fig. 8. AK: Sitcha Isl. This was first published by Guerin-Meneville in 1835 as a named figure; text (p. 60) published in 1844. melsheimeri Kirby, 1837: 97. 'ïrom New York to Cumberland-House" (SK). TYPE DEPOSITORY: BMNH. 8 COLEÓPTERA CATALOG

particeps Fischer von Waldheim, 1844: 139 (as variety of Nicrophorus lunatus). AK: Sitka. Heller (1896: 23) shows Turcoman, the locality cited by Fischer von Waldheim (1844: 139), to be in error. aleuticus Gistel, 1848: 190. AK: Sitka. infodiens Mannerheim, 1853: 170. AK: Unalaschka Isl. pollinctor Mannerheim, 1853: 169. AK: Atkha Isl. Peck and Miller (1982:152) designated the lectotype. confossor LeConte, 1854: 20. WA: Prairie Paso. TYPE DEPOSITORY: MCZC. SEX OF TYPE: M. TAXONOMY: Arnett, 1947: 110; Peck and Miller, 1982: 152. marginatus Fabricius, 1801: 334. ^^America"; BC WA OR ID/ AB MT ND SD/ MN WI MI ON I PQ / CA/ WY UT CO/ NE KS lA MO IL IN KY/ NY PA NJ MD VA/ ME MA RI ' CT/ AZ NM TX OK/ AR LA TN NC FL; Mex. ] requiescator Gistel, 1848: 190 (synon3miy following Hatch, 1928: 147). ''North America*'. montezumae Matthews, 1888: 92. Mex. TYPE DEPOSITORY: BMNH. labreae Pierce, 1949: 63, figs. 4-10 (as subspecies of guttula). CA: Los Angeles C!o., Rancho La Brea. The Rancho La Brea specimens are of the Pleistocene Age (Pierce, 1949: 55). Miller and Peck (1979: 96) designated the lectotype. TYPE DEPOSITORY: LACM. latifrons Pierce, 1949: 67, fig. 14 (as subspecies of investigator). CA: Los Angeles Co., Rancho La Brea. The Rancho La Brea specimens are of the Pleistocene Age (Pierce, 1949: 55). TYPE DEPOSITORY: LACM. mckittricki Pierce, 1949: 66, fig. 11. CA: Kern Co., McKittrick. The McKittrick specimens are of the Pleistocene Age (Pierce, 1949: 55). TYPE DEPOSITORY: LACM. obtusiscutellum Pierce, 1949: 67, fig. 12. CA: Los Angeles Co., Rancho La Brea. The Rancho La Brea specimens are of the Pleistocene Age (Pierce, 1949: 55). TYPE DEPOSITORY: LACM. TAXONOMY: Miller and Peck, 1979: 96. mexicanas Matthews, 1888: 91. Mex.; UT CO/ KS lA/ AZ NM; Mex. ECOLOGY: Halffter et al., 1982: 51, and 1983: 649. nigrita Mannerheim, 1843: 251. CA; BC WA OR/ CA. The name is used as a noun in apposition; it is masculine singular of Nigritae, an African people living along the Niger River. ruficornis Motschulsky, 1869: 352 (Necrophorus). CA. alpha Pierce, 1949: 67, fig. 13 (as subspecies of investigator). CA: Los Angeles Co., Rancho La Brea. The Rancho La Brea specimens are of the Pleistocene Age (Pierce, 1949: 55). Miller and Peck (1979: 95) designated the lectotype. TYPE DEPOSITORY: LACM. TAXONOMY: Miller and Peck, 1979: 95-6. obscurus Kirby, 1837: 97. 'ïrom New York to Cumberland-House'' (SK); ID/ AB SK MB MT ND SD/ MN/ UT/ NE lA MO. melsheimeri LeConte, 1853: 275. NB, UT. orbicoilis Say, 1825: 177. ^^United States''; SD/ MN WI MI ON/ NS/ NE KS lA MO IL IN OH KY/ NY PA NJ MD DC WV VA/ ME NH MA RI CT/ OK/ AR AL TN GA SC NC FL. Peck and Miller (1982: 153) designated a neotype. TYPE DEPOSITORY: MCZC. SEX OF TYPE: M. hallii Kirby, 1837: 98. NS, MA. SILPHIDAE 9

quadrisignatus Laporte, 1840: 1. "Amérique du Nord*\ TAXONOMY: Peck and Miller, 1982: 153. ECOLOGY: Scott, 1989: 133; Trumbo. 1991: 82. pustulatus Herschel, 1807: 271. "North America''; MN WI MI ON PQ/ KS lA MO IL IN OH/ NY PA NJ MD DC WV VA/ ME NH RI CT/ TX OK/ AR GA NC FL. bicolon Newman, 1838: 385. "North America". tardus Mannerheim, 1853: 170. AK: Sitka Isl. sayi Laporte, 1840: 2. "North America''; AB SK MB/ MN WI MI ON PQ/ NF/ lA IL IN/ NY PA NJ MD WV/ ME NH MA CT/ OK/ AR AL TN NC. lunulatus Gistel, 1848: 189 (synonymy following Hatch, 1928: 129). "North America". lunatus LeConte, 1853: 277 (preoccupied Fischer von Waldheim, 1842). NY, GA. Peck and Miller (1982: 152) designated the lectotype. TYPE DEPOSITORY: MCZC. SEX OF TYPE: M. Inniger Harold, 1868: 104 (replacement name for lunatus LeConte). NY. TAXONOMY: Peck and Miller, 1982: 152. tomentosus Weber, 1801: 47. "America"; MB MT SD/ MN WI MI ON PQ/ NB NS/ CO/ NE KS lA MO IL IN OH/ NY PA NJ MD DC VA/ ME NH MA CT/ NM TX OK/ AR AL TN GA SC NC FL. velutinus Fabricius, 1801: 334. "Carolina". aurigaster Portevin, 1925: 170 (as variety of velutinus), "United States". ECOLOGY: Scott and Traniello, 1988: 693; Trumbo, 1990: 491. vespilloides Herbst, 1783: 32. Germany: Berlin; AK YT NT/ BC/ AB SK MB ND/ MN MI ON PQ/ NB NS NF/ IL IN/ NY PA/ ME NH MA; Holarc. See Hatch (1928: 136) for Eurasian synonyms. hebes Kirby, 1837: 96. NS. TYPE DEPOSITORY: BMNH. pygmaeus Kirby, 1837: 98, pi. II, fig. 3. "Rocky Mountains". TYPE DEPOSITORY: BMNH. TAXONOMY: Leech, 1937: 158. ECOLOGY: Bartlett, 1987a: 471, 1987b: 179, and 1988: 297. Unplaced Species pulsator Gistel, 1848: 190 (Necrophorus) (incertae sedis). Canada. NOMEN NUDUM brasiliensis Dejean, 1833: 118 (Necrodes) (not available). Brasilia, nomen nudum; this name weis published without a diagnosis or indication. 10 BIBLIOGRAPHY

Agassíz, L. 1847 Nomenciatoris zoologici index universalis, continens nomina systematica classium, ordinum, familiarum et generum animalium omnium, tam viventium quam fos- silium, secundum ordinem alphabeticum unicum disposita, adjectis homonymiis plantarum, nee non variis adnotationibus et emendationibus. Jent and Gass- mann, Soloduri [= Solothurn, Switzerland], viii and 1-393 pp. Date from Fletcher, 1979. Anderson, R. S. 1982a Burying larvae: Nearctic Nicrophorus and Oriental Ptomascopus morio (Sil- phidae). Systematic Entomology, vol. 7, no. 3, pp. 249-264. Anderson, R. S. 1982b On the decreasing abundance of Nicrophorus americanus Olivier (Coleóptera: Sil- phidae) in Eastern North America. The Coleopterists Bulletin, vol. 36, no. 2, pp. 362-365. Anderson, R. S. 1982c Resource partitioning in the carrion beetle (Coleóptera: Silphidae) fauna of south- ern Ontario: ecological and evolutionary considerations. Canadian Journal of Zo- ology, vol. 60, no. 6, pp. 1314-1325. Anderson, R. S. 1987 The of Thanatophilus trituberculatus (Kirby) (Coleóptera: Silphidae). The Co- leopterists Bulletin, vol. 41, no. 1, p. 34. Anderson, R. S. 1989 Potential phylogenetic utility of mating behavior in some carrion beetles (Coleóp- tera: Silphidae: Silphinae). The Coleopterists Bulletin, vol. 43, no. 1, p. 18. Anderson, R. S. and S. B. Peck 1984 Bionomics of Nearctic species of Aclypea Reitter: Phjrtophagous **carrion'' beetles (Coleóptera: Silphidae). Pan-Pacific Entomologist, vol. 60, no. 3, pp. 248-255. Anderson, R. S. and S. B. Peck 1985 The and arachnids of Canada. Part 13. The carrion beetles of Canada and Alaska (Coleóptera: Silphidae and ). Agriculture Canada, Ottawa, publ. 1778, 121 pp. Anderson, R. S. and S. B. Peck 1986 Geographic patterns of colour variation in North American Nicrophorus burying beetles (Coleóptera: Silphidae). Journal of Natural History, vol. 20, no. 3, pp. 283- 297. Angelí, J. W. 1912 Two new North American species of Necrophorus (Coleop.). Entomological News, vol. 23, no. 7, 307 pp. Arnett, R. H., Jr. 1944 A revision of the Nearctic Silphini and Nicrophorini based upon the female genita- lia (Coleóptera, Silphidae). Journal of the New York Entomological Society, vol. 52, no. 1, pp. 1-25. Arnett, R. H., Jr. 1947 Coleóptera notes II: Silphidae. The Canadian Entomologist, vol. 79, no. 6, pp. 110- 113. Balduf, W. V. 1935 The bionomics of entomophagous Coleóptera. J. S. Swift Co., Inc., St. Louis, pt. 1, 220 pp. Reprinted in 1969 by E. W. Classey, Hampton, England. SILPHIDAE 11

Bartlett, J. 1987a Evidence for a sex attractant in burying beetles. Ecological Entomology, vol. 12, no. 4, pp. 471-472. Bartlett, J. 1987b Filial cannibalism in burying beetles. Behavioral Ecology and Sociobiology, vol. 21, no. 3, pp. 179-183. Bartlett, J. 1988 Male mating success and parental care in (Coleóptera: Sil- phidae). Behavioral Ecology and Sociobiology, vol. 23, no. 5, pp. 297-303. Bergroth, E. 1884 Bemerkungen zur dritten Auflage des Catalogus Coleopterorum Europae auctori- bus L. V. Heyden, E. Reitter et J. Weise. Berliner Entomologische Zeitschrift, vol. 28, pt. 2, pp. 225-230. Bjorkman, F. and M. H. Hatch 1939 Note on Silpha (Thanatophilus) coloradensis Wickham. The Pan-Paciñc Entomolo- gist, vol. 15, no. 2, p. 96. Blackwelder, R. E. 1949 Studies on the dates of books on Coleóptera. I. The Coleopterists' Bulletin, vol.3, pp.42-46. Blaisdell, F. E., Sr. 1901 of Necrophilus hydrophiloides. Entomological News, vol. 12, no. 9, pp. 263- 264. Bland, J. H. B. 1865 Descriptions of several new species of North American Coleóptera. Proceedings of the Entomological Society of Philadelphia, vol. 4, pp. 381-384. Bolivar y Pieltain, C. and J. Hendrichs 1972 Distribución en Norteamérica del genero holartico Pteroloma Gyllenhal, 1827 y es- tudio de tres formas mexicanas (Col. Silph.). Ciencia, Mexico, vol. 27, no. 6, pp. 207-216. Brewer, J. W. and T. R. Bacon 1975 Biology of the carrion beetle Silpha ramosa Say. Annals of the Entomological Soci- ety of America, vol. 68, no. 5, pp. 786-790. Brown, W. J. 1933 Two undescribed species of the old family Silphidae with notes on some characters that have been used to divide the group. The Canadian Entomologist, vol. 65, no. 9, pp. 213-215. Brülle, A. 1836 Insectes de TAmerique méridionale. In d'Orbigny, A. D., Voyage dans V Amérique méridionale (le Brésil, la Republique Orientale de V Uruguay, la Republique Ar- gentine, la Patagonie, la Republique du Chili, la Republique de Bolivia, la Re- publique du Pérou), execute penda les annes 1826, 1827, 1828, 1829, 1830, 1831, 1832, et 1833. Librairie de la Société géologique de France, Paris, vol. 6, pt. 2: Insectes, pp. 1-222. Dates from Sherborn and Griffin, 1934. Brülle was the author to page 60. After page 60, the author was Blanchard. Casey, T. L. 1886 Descriptive notices of North American Coleóptera. I. Bulletin of the California Academy of Sciences, vol. 2, no. 6, pp. 157-264, illus. Casey, T. L. 1924 Additions to the known Coleóptera of North America. Memoirs on the Coleóptera. New Era Printing Co., Lancaster, PA, vol. 11, pp. 1-347. 12 COLEÓPTERA CATALOG

Catesby, M. 1748 The natural history of Carolina, Florida, and the Bahama Islands. (Edition III, Geo. Edwards, editor). London, vol. 2, (1743) 100 pp. and appendix (1748), 20 pp. Champion, G. C. 1923 Some Indian Coleóptera (10). Entomologist's Monthly Magazine, vol. 59, no. 707, pp. 43-53, 77-80. Chevrolat, L. A. A. 1843 Coléoptères du Mexique (226) (Pentameres: Hydrocanthares, sternoxes, terediles, necrophages, lamellicornes). Magasin de Zoologie, (series 2), vol. 13, pp. 1-37, illus. Cockerell, T. D. A. 1906 Preoccupied generic names of Coleóptera. Entomological News, vol. 17, no. 7, pp. 240-244. Cooley, R. A. 1917 The spinach carrion beetle Silpha bituberosa Lee. Journal of Economic Entomolo- gy, vol. 10, no. 1, pp. 94-102. Cowan, C. F. 1971 On Guerin's Iconographie: Particularly the insects. Journal of the Society for the Bibliography of Natural History, vol. 6, pt. 1, pp. 18-29. Curtis, J. 1830 British entomology; being illustrations and descriptions of the genera of insects found in Great Britain and Ireland: Containing coloured figures from nature of the most rare and beautiful species, and in many instances of the plants upon which they are found. London, vol. 7, pis. 290-337. Citation for first edition, numerical arrangement (Blackwelder, 1949). Dahl, G. 1807 Ein merkwürdiger neuer Kaefer Silpha subterránea. Magazin fuer Insektenkunde, vol. 6, pp. 362-364. Davis, L. R., Jr. 1980 Notes on beetle distributions, with a discussion of Nicrophorus americanus Olivier and its abundance in collections (Coleóptera: Scarabaeidae, Lampyridae, and Sil- phidae). The Coleopterists Bulletin, vol. 34, no. 2, pp. 245-251. Dejean, P. F. M. A. 1833 Catalogue des coléoptères de la Collection de M. le comte Dejean. Mequignon - Marvis, Paris, Livr. 2, pp. 97-176. Dorsey, C. K. 1940 A comparative study of the larvae of six species of Silpha (Coleóptera, Silphidae). Annals of the Entomological Society of America, vol. 33, no. 1, pp. 120-139. Edwards, G. 1771 A catalogue of the animals and plants represented in Catesby's natural history of Carolina: With the Linnean names. Appended to Edwards' 1971 edition of Ca- tesby's "The natural history of Carolina, Florida, and the Bahama Islands.'' London, 2 volumes. Reprinted in 1974 by Beehive Press, Savannah, Georgia. Eisner, T. and J. Meinwald 1983 Defensive spray mechanism of a silphid beetle {). Psyche, vol. 89, nos. 3-4, pp. 357-367. Fabricius, J. C. 1775 Systema entomologiae sistens insectorum classes, ordines, genera, species, adiectis synonymis, locis, descriptionibus, observationibus. Kortii, Flensburgi et Lipsiae [= Flensburg and Leipzig], 832 pp. SILPHIDAE 13

Fabricius, J. C. 1777 Genera insectorum eorumque characteres naturales secundum numerum, figuram, situm et proportionem omnium partium oris adiecta mantissa specierum nuper detectarum. Bartsch, Chilonii [= Kiel], xiv and 310 pp. Date from Hagen, 1862. Fabricius, J. C. 1781 Species insectorum exhibentes eorum differentias specifícas, synonyma auctorum, loca natalia, metamorphosin adiectis observationbus, describtionibus. C. E. Bohnii, Hamburgi et Kilonii [= Hamburg and Cologne], vol. 1, iii-viii and 552 pp. Fabricius, J. C. 1792 Entomología systematica emendata et aucta. Secundum classes, ordines, genera, species adjectis synonimis, locis observationibus, descriptionibus. Proft, Hafniae [= Copenhagen], vol. 1, pt. 1, xx and 330 pp.; pt. 2, 538 pp. Fabricius, J. C. 1801 Systema eleutheratorum secundum ordines, genera, species adiectis synonymis locis, observationibus, descriptionibus. Bibliopolii Academici Novi. Kiliae [= Kiel], vol. 1, 506 pp. Reprinted in 1970 by Asher and Co., Vaals. Fall, H. C. 1907 New Coleóptera from the southwest. III. The Canadian Entomologist, vol. 34, no. 7, pp. 235-243. Fall, H. C. 1927 New Coleóptera XII. The Canadian Entomologist, vol. 59, no. 6, pp. 136-141. Fall, H. C. 1937 Miscellaneous notes and descriptions (Coleóptera). The Canadian Entomologist, vol. 69, no. 2, pp. 29-32. Ferris, G. F. 1918 An apparently new species of Leptinillus (Coleóptera, Leptinidae). The Canadian Entomologist, vol. 50, no. 4, pp. 125-128. Fischer von Waldheim, G. 1828 Entomographia Imperil Rossici. A. Semen, Mosquae [= Moscow], vol. 3, pp. viii and 3-314, illus. Fischer von Waldheim, G. 1842 Catalogus Coleopterorum in Sibiria Orientali a Cel. Gregorio Silide Karelin CoUec- torum. Moscow, pp. 1-28. Fischer von Waldheim, G. 1844 Spicilegium entomographiae rossicae. Bulletin de la Société Imperiale des Natura- listes de Moscou, vol. 17, pp. 3-144. Fisher, R. M. and R. D. Tuckerman 1986 Mimicry of bumble bees and cuckoo bumble bees by carrion beetles (Coleóptera: Silphidae). Journal of the Kansas Entomological Society, vol. 59, no. 1, pp. 20-25. Fletcher, D. S. 1979 Geometroidea. In The generic names of moths of the world. Volume 3. I. W. B. Nye, editor. British Museum (Natural History), London, xx and 243 pp. Forster, J. R. 1771 Novae species insectorum. Centuria I. Davies and White, London, viii and 100 pp. Froelich, J. A. 1792 Bemerkungen ueber einige seltene Kaefer aus der Insectensammlung des Herrn. Hofr. und Prof. Rudolph in Erlangen. Der Naturforscher, vol. 26, pp. 68-165. 14 COLEÓPTERA CATALOG

Froelich, J. A. 1799 Einige neue Gattungen und Arten von Kaefern. Der Naturforscher, voL 28, pp. 1- 65. Germar, E. F. 1824 Insectorum species novae aut minus cognitae, descriptionibus illustratae. I. C. Hendelii and Filii, Halae [= Halle], vol. 1, xxiv and 624 pp., illus. Gistel, J. 1834 Entomologische Notizen. Zeitschrift fuer Zoologie und vergleichende Anatomie, vol. 1, pt. 3, pp. 128-151. Gistel, J. 1848 Naturgeschichte des Thierreichs fuer hoehere Schulen. 2nd ed. Scheitlin & Krais, Stuttgart, xvi and 216 pp., 4 unnumbered pp., illus. Gistel, J. 1857 Achthundert und zwanzig neue oder unbeschriebene wirbellose Thiere. Straubing, vol. 2, pp. 513-606. Also published as a separate work, pp. 1-94. Gleditsch, J. G. 1767 Vermischte Physicalisch-Botanisch-Oeconomische Abhandlungen. Halle, vol. 3, 24 unnumbered pp., 397 pp., and 11 unnumbered pp., illus. Guerin-Meneville, F. E. 1835 Iconographie du Regne animal de G. Cuvier, ou representation d'après nature de Tune des espèces les plus remarquables, et souvent non encore figurées, de chaque genre d'animaux. Insectes. Bailliere, Paris, (1829-1844) vol. 7, pp. 1-576, illus. Title page date, 1829-1838; plate 17 publ. 1835; text publ. 1844; dates from Cowan, 1971. Gyllenhal, L. 1810 Insecta Svecica descripta. Classis I. Coleóptera sive Eleutherata. Leveretz, Scaris, vol. 1, pt. 2, iv-xx and 660 pp. Gyllenhal, L. 1827 Insecta Svecica descripta. Classis I. Coleóptera sive Eleuterata. Fleischer, Lipsiae [= Leipzig], vol. 1, pt. 4, viii and 761 pp. Hagen, H. A. 1862 Bibliotheca entomológica. Die litteratur ueber das ganze Gebiet der Entomologie bis zum Jahre 1862. W. Engelmann, Leipzig, vol. 1, A-M, 556 pp. Halffter, G., S. Anduaga, and C. Huerta 1982 Nidificacion en Nicrophorus (Coleóptera, Silphidae). Folia Entomológica Mexicana, no. 54, pp. 50-52. Halffter, G., S. Anduaga, and C. Huerta 1983 Nidification des Nicrophorus (Col. Silphidae). Bulletin de la Société Entomologique de France, vol. 88, nos. 7 and 8, pp. 648-666. Harold, E. von 1868 Miscellen [= Miscellaneous notes]. In Coleopterologische Hefte. Carl Merhoffs Verlag, vol. 4, 104 pp. Hatch, M. H. 1927a Studies on the carrion beetles of Minnesota, including new species. University of Minnesota Agricultural Experiment Station, St. Paul, Technical Bulletin, no. 48, 19 pp. Hatch, M. H. 1927b Studies on the Silphinae. Journal of the New York Entomological Society, vol. 35, no. 4, pp. 331-371. SILPHIDAE 15

Hatch, M. H. 1928 Silphidae IL In Coleopterorum Catalogus auspiciis et auxilio W. Junk editus A. S. Schenkung. W. Junk, Berlin, vol. 7, pt. 95, pp. 63-244. Hatch, M. H. 1946 Mr. Ross H. Arnett's ^'Revision of the Nearctic Silphini and Nicrophorini.'' Jour- nal of the New York Entomological Society, vol. 54, no. 2, pp. 99-103. Hatch, M. H. 1957 The beetles of the Pacific Northwest. Part II: . University of Washington Publications in Biology, University of Washington Press, Seattle, vol. 16, 384 pp. Hatch, M. H. and J. W. Angelí 1925 A new North American Necrophorus, Journal of the New York Entomological So- ciety, vol. 33, no. 4, p. 216. Hecht, A. 1988 Endangered and threatened wildlife and plants: Proposed endangered status for the American . The Federal Register, vol. 53, no. 196, pp. 39617- 39621. Heller, K. M. 1896 Necrophorus lunatus und particeps Fischer. Entomologische Nachrichten, vol. 22, no. 2, pp. 23-24. Herbst, J. F. W. 1783 Kritisches Verzeichniss meiner Insektensammlung. 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Mannerheim, C. G. von 1853 Dritter Nachtrag zur Kaefer-Fauna der Nord-Amerikanischen Laender des Russis- chen Reiches. Bulletin de la Société Imperiale des Naturalistes de Moscou, vol. 26, no. 3, pp. 95-273. Matthews, A. 1888 In D. Sharp et al., Family Silphidae. Biología Centrali-Americana. Insecta. Coleóp- tera. Printed by Taylor and Francis, London, vol. 2, pt. 1, pp. 72-101, illus. Miller, S. E. and S. B. Peck 1979 Fossil carrion beetles of Pleistocene California asphalt deposits, with a synopsis of Holocene California Silphidae (Insecta: Coleóptera: Silphidae). Transactions of the San Diego Society of Natural History, vol. 19, no. 8, pp. 85-106. Milne, L. J. and M. J. Milne 1944 Notes on the behavior of burying beetles (Nicrophorus spp.). Journal of the New York Entomological Society, vol. 52, no. 4, pp. 311-327. Milne, L. J. and M. J. Milne 1976 The social behavior of burying beetles. Scientific American, vol. 235, no. 2, pp. 84- 89. Motschulsky, V. von 1845 Remargues sur la collection de Coléoptères russes de Victor de Motchulsky. Article 1. Bulletin de la Société Imperial des Naturalistes de Moscou, vol. 18, no. 1, pp. 1-127, illus. Motschulsky, V. von 1869 Enumeration des nouvelles espèces de Coléoptères rapportes de ses voyages. VIII. Bulletin de la Société Imperial des Naturalistes de Moscou, vol. 42, pt. 2, pp. 348- 410. Mueller, O. F. 1776 Zoologiae Danicae prodromus, seu animalium Daniae et Norvegiae Indigenarum Characteres, nomina, et synonyma imprimis popularium. Hallageri, Havniae [= Copenhagen], xxxii and 282 pp. Newman, E. 1838 Entomological notes. Entomological Magazine, vol. 5, pp. 372-402. Olivier, A. G. 1790 Entomologie, ou histoire naturelle des insects, avec leurs caracteres génériques et spécifiques, leur description, leur synon3niiie, et leur figure enluminée. Coléop- tères. Baudoin, Paris, vol. 2, no. 10, 8 pp., illus. Peck, S. B. 1974 A review of the Agyrtes (Silphidae) of North America. Psyche, vol. 81, nos. 3-4, pp. 501-506. Peck, S. B. 1981 Distribution and biology of the flightless carrion beetle Necrophilus pettitii in east- ern North America (Coleóptera: Silphidae). Entomological News, vol. 92, no. 5, pp. 181-185. Peck, S. B. 1989 Insecta: Coleóptera Silphidae and the associated families Agyrtidae and Leiodidae. In Dindal, D. (ed.). Soil biology guide. John Wiley and Sons, New York, ch. 37. Peck, S. B. and R. S. Anderson 1982 The distribution and biology of the alpine-tundra carrion beetle Thanatophilus co- loradensis (Wickham) in North America (Coleóptera: Silphidae). The Coleopter- ists Bulletin, vol. 36, no. 1, pp. 112-115. SILPHIDAE 19

Peck, S. B. and R. S. Anderson 1985 Taxonomy, phylogeny and biogeography of the carrion beetles of Latin America (Coleóptera: Silphidae). Quaestiones Entomologicae, vol. 21, no. 3, pp. 247-317. Peck, S. B. and M. M. Kaulbars 1988 A synopsis of the distribution and bionomics of the carrion beetles (Coleóptera: Sil- phidae) of the conterminous United States. Proceedings of the Entomological So- ciety of Ontario, (1987) vol. 118, pp. 47-81. Peck, S. B. and S. E. Miller 1982 Type designations and synonymies for North American Silphidae (Coleóptera). Psyche, vol. 89, nos. 1-2, pp. 151-156. Pierce, W. D. 1949 Fossil arthropods of California. 17. The silphid bur3dng beetles in the asphalt de- posits. Bulletin of the Southern California Academy of Sciences, vol. 48, pt. 2, pp. 55-70. Portevin, G. 1903 Remarques sur les necrophages du museum et description d'espèces nouvelles. Bul- letin du Museum National d'Histoire Naturelle, Paris, vol. 9, no. 7, pp. 329-336. Portevin, G. 1905 Troisième note sur les Silphides du Museum. Bulletin du Museum National d'His- toire Naturelle, Paris, vol. 11, no. 6, pp. 418-424. Portevin, G. 1914 Revision des Silphides, Liodides et Clambides du Japon. Annales de la Société En- tomologique de Belgique, vol. 58, pp. 212-236. Portevin, G. 1920a Revision des Silphini et Necrophorini de la region Indo-Malaise. Bulletin du Museum National d'Histoire Naturelle, Paris, vol. 26, no. 5, pp. 395-401. Portevin, G. 1920b Silphides nouveaux de la collection du museum. Bulletin du Museum National d'Histoire Naturelle, Paris, vol. 26, no. 6, pp. 505-508. Portevin, G. 1921a Note sur quelques Silphides et Liodides de la collection Grouvelle. Bulletin du Museum National d'Histoire Naturelle, Paris, vol. 27, no. 7, pp. 535-538. Portevin, G. 1921b Revision du genre Hyponecrodes Kraatz. Miscellanea Entomologicia, vol. 25, no. 11, pp. 81-85. Portevin, G. 1922 Note sur quelque silphides des collections du Museum. Bulletin du Museum Na- tional d'Histoire Naturelle, Paris, vol. 28, no. 7, pp. 506-508. Portevin, G. 1923 Revision des Necrophorini du globe. Bulletin du Museum National d'Histoire Na- turelle, Paris, vol. 29, no. 1, pp. 64-71; no. 2, 141-146; no. 3, 226-233; no. 4, 303-309. Portevin, G. 1925 Revision des Necrophorini du globe. Bulletin du Museum National d'Histoire Na- turelle, Paris, vol. 31, no. 2, pp. 165-170. Portevin, G. 1926 Les grands necrophages du globe. Encyclopédie Entomologique, vol. 6, pp. 1-270. Ratcliffe, B. C. 1972 The natural history of Necrodes surinamensis (Fabr.) (Coleóptera: Silphidae). Transactions of the American Entomological Society, vol. 98, no. 4, pp. 359-410. 20 COLEÓPTERA CATALOG Reitter, E. 1885 Bestimmungs-Tabellen der europaeischen Coleopteren. XII. Necrophaga. (Platyp- syllidae, Leptinidae, Silphidae, Anisotomidae und Clambidae). Verhandlungen Naturforschen Vereines, W. Burkart, Bruenn [= Br'no], vol. 23, pp. 3-122. Reitter, E. 1901 Uebersicht der bekannten Agyrtes-Arten. Wiener Entomologische Zeitung, vol. 20, p. 102. Say, T. 1823 Descriptions of coleopterous insects collected in the late expedition to the Rocky Mountains, performed by order of Mr. Calhoun, Secretary of War, under the command of Major Long. Journal of the Academy of Natural Sciences of Phila- delphia, vol. 3, pt. 1, pp. 139-216. Reprinted in LeConte, 1859d. Say, T. 1825 Descriptions of new species of coleopterous insects inhabiting the United States. Journal of the Academy of Natural Sciences of Philadelphia, vol. 5, pt. 1, pp. 160-204. Reprinted in LeConte, 1859d. Schawaller, W. 1981 Taxonomie und Faunistik der Gattung Thanatophilus (Coleóptera: Silphidae). Stuttgarter Beitrage zur Naturkunde, Serie A (Biologie), no. 351, pp. 1-21. Schawaller, W. 1983 Die Arten der Gattung Ipelates (sensu lato) (Coleóptera, Silphidae, Agyrtinae). Revue Suisse de Zoologie, vol. 90, fase. 1, pp. 101-110. Schilling, P. S. 1829 Holocnemis. Beitraege zur Entomologie, besonders in bezug auf die schlesische Fauna , vol. 1, pp. 93-94. Schouteden, H. 1905 Les Hyponecrodes sud-americains. Annales de Société Entomologique de Belgique, vol. 49, pp. 195-199. Scopoli, J. A. 1772 Annus histórico naturalis Annus, Lipsiae [= Leipzig], vol. 5, 128 pp. Scott, M. P. 1989 Male parental care and reproductive success in the burying beetle Nicrophorus or- bicollis. Journal of Behavior, vol. 2, no. 1, pp. 133-137. Scott, M. P. and J. F. A. Traniello 1988 Behavioural cues trigger ovarian development in the burying beetle Nicrophorus tomentosus. Journal of Insect Physiology, vol. 33, no. 10, pp. 693-696. Scott, M. P., J. F. A. Traniello, and I. A. Fetherston 1987 Competition for prey between ants and burying beetles (Nicrophorus spp.): differ- ences between northern and southern temperature sites. Psyche, vol. 94, nos. 3- 4, pp. 325-332. Seidlitz, G. 1888 See Seidlitz, 1891 . Seidlitz, G. 1891 Fauna Báltica. Die Kaefer (Coleóptera) der deutschen Ostseeprovinzen Russlands, ed. 2 (1887-1891). Koenigsberg, Familien LVI, Gattungen 192 and Arten 818 pp. Fauna Báltica was initially issued in 6 fascicles that were combined in 1891 as a complete work: fase. 1, 1887, Familien pp. I-XL, Gattungen pp. 1-16, Arten pp. 1-96; fase. 2, 1888 (spring), Gattun- gen pp. 17-48, Arten pp. 97-224; fase. 3, 1888 (autumn), Familien pp. XLI-XLVII, Gattungen pp. 49- 80, Arten pp. 225-336; fase. 4, 1889, Gattungen pp. 81-128, Arten pp. 337-512; fase. 5, 1890, Gattun- gen pp. 129-160, Arten pp. 513-608; fase. 6, 1891, Familien pp. XLIX-LVI, Gattungen pp. 161-192, Arten pp. 609-818. SILPHIDAE 21

Semenov-Tian-Shanskij, A. P. 1891 Diagnoses Coleopterorum novorum ex Asia Centrali et Orientali. Horae Societatis Entomologicae Rossicae, (1890) vol. 25, pp. 262-382. On the table of contents page the author's name is cited as Andreas Semenow. Semenov-Tian-Shansky, A. P. 1933 De tribu Necrophorini (Coleóptera, Silphidae) classificanda et de ejua distributione geographica. Travaux de Tlnstitut Zoologique de TAcademie des Sciences de rURSS (Trudy Zoologicheskogo Institua Leningrad), vol. 1, pp. 149-160. Title in French and Russian; text in Latin and Russian. Sherborn, D. D. and F. J. Griffin 1934 On the dates of publication of the natural history portions of Alcide d'Orbigny's ^'Voyage Amérique Méridionale.*' Annals and Magazine of Natural History, ser. 10, vol. 13, no. 73, pp. 130-134. Shubeck, P. P. 1971 Diel periodicities of certain carrion beetles (Coleóptera: Silphidae). The Coleopter- ists Bulletin, vol. 25, no. 2, pp. 41-46. Shubeck, P. P. 1976a An alternative to pitfall traps in carrion beetle studies (Coleóptera). Entomological News, vol. 87, nos. 5 & 6, pp. 176-178. Shubeck, P. P. 1976b Carrion beetle réponses to poikilotherm and homoiotherm carrion (Coleóptera: Sil- phidae). Entomological News, vol. 87, nos. 9 & 10, pp. 265-269. Sturm, J. 1826 Catalog meiner Insecten-Sammlung. Nuernberg [= Nuremberg], pt. 1, Kaefer, viii and 207 pp., illus. Thomson, C. G. 1862 Skandinaviens Coleóptera, synoptiskt bearbetade. Lund, vol. 4, 269 pp. Thunberg, K. P. 1789 Periculum entomologicum quo characteres generum insectorum. J. Edman, Upsa- liae [= Uppsala], 16 pp. Trumbo, S. T. 1990 Regulation of brood size in a burying beetle, Nicrophorus tomentosus (Silphidae). Journal of Insect Behavior, vol. 3, pp. 491-500. Trumbo, S. T. 1991 Reproductive benefits and the duration of paternal care in a biparental burying beetle, Necrophorus [sic] oribicollis. Behaviour, vol. 117, pp. 82-105. Van Dyke, E. C. 1921 Coleóptera from the Pribilof Islands, Alaska. Proceedings of the California Acade- my of Sciences, ser. 4, vol. 11, no. 4, pp. 156-166. Van Dyke, E. C. 1928 The American species of Pteroloma (Coleoptera-Silphidae) and a new Japanese spe- cies. Bulletin of the Brooklyn Entomological Society, vol. 23, no. 1, pp. 19-26, illus. Walker, F. 1866 List of Coleóptera. In J. K. Lord, The naturalist in Vancouver Island and British Columbia. R. Bentley, London, vol. 2, pp. 309-334. Weber, F. 1801 Observationes entomologicae, continentes novorum quae condidit generum charac- teres, et nuper dectectarum specierum descriptiones. Bibliopolii Academici novi, Kiliae [= Kiel], pp. iv-xii and 4-116. 22 COLEÓPTERA CATALOG

Wickham, H. F. 1902 Two new Silphidae from Colorado. The Canadian Entomologist, vol. 34, no. 7, pp. 180-181. Wilson, D. S. 1983 The effect of population structure on the evolution of mutualism: A field test in- volving burying beetles and their phoretic mites. The American Naturalist, vol. 121, no. 6, pp. 851-870. Wilson, D. S. and J. Fudge 1984 Burying beetles: Intraspecific interactions and reproductive success in the field. Ecological Entomology, vol. 9, no. 2, pp. 195-203. Wilson, D. S. and W. G. Knollenberg 1984 Food discrimination and ovarian development in burying beetles (Coleóptera: Sil- phidae: Nicrophorus). Annals of the Entomological Society of America, vol. 77, no. 2, pp. 165-170. Wilson, D. S., W. G. Knollenberg, and J. Fudge 1984 Species packing and temperature dependent competition among burying beetles (Silphidae, Nicrophorus). Ecological Entomology, vol. 9, no. 2, pp. 205-216. Zetterstedt, J. W. 1824 Nya Svenska Insect-Arter. Konglifa Vetenskaps Akademiens Handlingar, 1824, pp. 149-159. INDEX 23

Names are indexed as follows: longulus LeConte, Agyrtes 2 CAPITALS: All names for taxa above the generic lunatus LeConte, Nicrophorus 9 level; luniger Harold, Nicrophorus 9 Boldface: Valid generic and subgeneric names; lunulatus Gistel, Nicrophorus 9 Roman: Valid specific and subspecific names; Lyrosoma Mannerheim 3 Italics: All invalid names such as synonyms, nomina marginalis Fabricius, Oiceoptoma 5 nuda, and extra-limital taxa even though valid. marginatus Fabricius, Nicrophorus 8 The generic name following the author's name indi- maritimus Guerin-Meneville, Nicrophorus 7 cates the present placement of the species. Synonyms of mckittricki Pierce, Nicrophorus 8 species-group names are listed with the original spelling. mediatas Fabricius, Nicrophorus 7 melsheimeri Kirby, Nicrophorus 7 Acanthopsilus Portevin 6 melsheimeri LeConte, Nicrophorus 8 Aclypea Reitter 6 mexicanus Matthews, Nicrophorus 8 Adolus Fischer von Waldheim 3 montezamae Matthews, Nicrophorus 8 aenescens Casey, Heterosilpha 6 mysticallis Angelí, Nicrophorus 7 affine Kirby, Necrophila 5 nebrioides Brown, Pteroloma 3 Agyrtecanus Reitter 2 Necrobias Gistel 2 Agyrtes Froelich 2 Necrobora Hope 5 AGYRTINI 2 Necrocharis Portevin 6 aleuticus Gistel, Nicrophorus 8 Necrocleptes Semenov-Tian-Shanskij 6 alpha Pierce, Nicrophorus 8 Necrodes Leach 3 americana Linnaeus, Necrophila 5 NecrophagaSy error 6 americanus Olivier, Nicrophorus 7 Necrophila Kirby and Spence 5 analis Chevrolat, Oxelytrum 4 Necrophilodes Champion 2 Apteroloma Hatch 3 arizonicum Van Dyke, Apteroloma 3 Necrophilus Latreille 2 Asbolus Bergroth 3 Necrophorus Thunberg 6 ater Motschulsky, Necrophilus 2 Necropter Semenov-Tian-Shanskij 6 aurigaster Portevin, Nicrophorus 9 Necrotropha Gistel 5 bicoTon Newman, Nicrophorus 9 Neonicrophoras Hatch 7 bituberosa LeConte, Aclypea 6 NICROPHORINI 6 Blitophaga Reitter 6 Nicrophorus Fabricius 6 Bracnyloma Portevin 2 nigrita Mannerheim, Nicrophorus 8 brasiliensis Dejean 9 NOMEN NUDUM 9 californica Mannerheim, Thanatophilus 4 noveboracense Forster, Oiceoptoma 5 Calosilpha Portevin 5 obalskii Portevin, Thanatophilus 4 canadense Kirby, Necrophila 5 obscurus Kirby, Nicrophorus 8 caraboides Fall, Apteroloma 3 obtasiscatellum Pierce, Nicrophorus 8 carolinus Linnaeus, Nicrophorus 7 Oeceoptoma Agassiz 5 caudata Say, Thanatophilus 4 Oiceoptoma Leach 5 cervaria Mannerheim, Heterosilpha 6 opaca Linnaeus, Aclypea 6 Chalcosilpha Portevin 4 opacum Mannerheim, Lyrosoma 3 Chrysositpha Portevin 5 orbicoUis Say, Nicrophorus 8 coloradensis Wickham, Thanatophilus 4 Oxelytriam, error 4 confossor LeConte, Nicrophorus 8 Oxelytron, error 4 conversator Walker, Nicrophorus 7 Oxelytrum Gistel 4 Cyrtoscelis Hope 6 Paranecrodes Portevin 4 defodiens Mannerheim, Nicrophorus 7 particeps Fischer von Waldheim, Nicrophorus 8 Deuterosilpha, error 5 Pelâtes Horn 2 Deutosilpha Portevin 5 Pelatines Cockerell 2 discicolle Brülle, Oxelytrum 4 peltata LeConte, Necrophila 5 discretas Portevin, Oxelytrum 4 pettiti Horn, Necrophilus 2 Eunecrophorus Semenov-Tian-Shanskij 6 Philas Portevin 4 Eusilpha Semenov-Tian-Shanskij 5 plagiatas Motschulsky, Nicrophorus 7 grandis Fabricius, Nicrophorus 7 politus Casey, Agyrtes 2 guttula Motschulsky, Nicrophorus 7 pollinctor LeConte, Nicrophorus 7 hallii Kirby, Nicrophorus 8 pollinctor Mannerheim, Nicrophorus 8 hebes Kirby, Nicrophorus 9 Protonecrodes Portevin 3 hecate Bland, Nicrophorus 7 Pseadopelta Bergroth 4 Heterosilpha Portevin 5 Pteroloma Gyllenhal 3 Holocnemis Schilling 3 pulsator Gistel, Nicrophorus 9 hybridus Hatch and Angelí, Nicrophorus 7 panctostriatus Pierce, Nicrophorus 7 hydrophiloides Guerin-Meneville, Necrophilus 2 pustulatus Herschel, Nicrophorus 9 Hyponecrodes Kraatz 4 pygmaeas Kirby, Nicrophorus 9 immaculosis Hatch, Nicrophorus 7 quadrisignatas Laporte, Nicrophorus 9 inaequale Fabricius, Oiceoptoma 5 ramosa Say, Heterosilpha 6 infodiens Mannerheim, Nicrophorus 8 reqaiescator Gistel, Nicrophorus 8 Investigator Zetterstedt, Nicrophorus 7 ruficornis Motschulsky, Nicrophorus 8 Ipelates Reitter 2 rugulosum Portevin, Oiceoptoma 5 Isosilpha Portevin 5 sagax Mannerheim, Thanatophilus 4 Katanecrodes Schouteden 4 sayi Laporte, Nicrophorus 9 labreae Pierce, Nicrophorus 8 SILPHIDAE 1 lapponicus Herbst, Thanatophilus 4 SILPHINI 3 lateralis Portevin, Nicrophorus 7 Silphosoma Portevin 4 latifrons Pierce, Nicrophorus 8 similis Fall, Agyrtes 3 latus Mannerheim, Ipelates 2 Sphaeroloma Portevin 2 Lendomus Casey 2 Stictonecropter Semenov-Tian-Shanskij 7 24 COLEÓPTERA CATALOG surinamensis Fabricius, Necrodes 3 tomentosus Weber, Nicrophorus 9 tahoecum Fall, Apteroloma 3 trituberculatus Kirby, Thanatophilus 5 tardus Mannerheim, Nicrophorus 9 truncatus Say, Thanatophilus 5 tenuicornis LeConte, Apteroloma 3 tuberculata Germar, Thanatophilus 4 term^natum Kirby, Necrophila 5 ^^Í^^^^^^S N¡!^^ I Thanatophilus Leach 4 virginicus Froelich, Nicrophorus 7 ^oi ft%'-^'^

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