/60 3&/YI =ttC\~I Qj/ ~c1:,.1f'-1IC ULJ;;Q( .ll".'l.i t1\~..m. UNIVERSITY OF MARYLAND Z: ~E !IJ MARYLAND AGRICULTURAL EXPERIMENT STATION COLLEGE PARK - EASTERN SHORE - RESEARCH FARMS ~ NOl.lVlS ~ The Biology of the Silphidae (Coleoptera):. : A Coded Bibliography : (\ ?? .... : ,, O.P. Young MP 981 July 1983 -----------------~- : 911364 Contents 1 Introduction 1 Summary of Biology 2 Coverage of Bibliography 3 Format of Citations 4 Index 5 Acknowledgements 6 Bibliography 46 Subject Index l The Biology of the Silphidae (Coleoptera): A Coded Bibliography Orrey P. Young 1 Introduction The family limits for the Silphidae have been variously de- fined for many years and only in the last 20 years or so has a more restricted definition become generally accepted. It was de- cided to follow the recent delineation of the family Silphidae by Madge(l980), which includes only the traditional subfamilies Silphinae, Nicrophorinae and Agyrtinae. Excluded from the Sil- phidae are the Bathysciinae, Camiarinae, Catopinae, Coloninae and Leptininae, groups now relegated to the families Leiodidae or Leptinidae. As constituted by Madge (1980), the family Sil- phidae worldwide contains 70 genera and approximately 250 species. Summary of Biology Although members of the Silphidae are usually called "car- rion beetles" (e.g. Dillon and Dillon, 1961), such a designation requires some conditional statements. Species in the subfamily Agyrtinae are poorly known and the meager biological information available suggests that they are found in the leaf litter and/or soil of forested habitats and consume carrion, dung, fungi, de- caying vegetation and/or the organisms associated with such materials. More is known of members of the Silphinae, a group which shows some diversity of food use. Species such as Blitophaga opaca, B. undata and Silpha bituberosa are primarily plant feed- ers, and can cause economically significant damage on crops such as spinach and beets (Martin, 1945). More typical, perhaps, are Silpha americana and S. novaboracensis, which feed on dipterous larvae found on carrion. Species specializei for preying on land 1Faculty Research Associate, Department of Entomology, University of Maryland, College Park, Maryland. Current Address: Southern Grain Insects Research Laboratory, USDA-SE-ARS, Tifton, Georgia; r snails include Ablattaria laevigata and Phosphuga subrotundata. One species, Xylodrepa quadripunctata, is an arboreal predator which feeds primarily on caterpillars. At one time it was intro- duced into the United States as a predator on the Gypsy Moth (Bur- gess and Crossman, 1929). Members of the subfamily Nicrophorinae are the well known "burying" beetles which, besides interring vertebrate carcasses as food for their young, consume dipterous larvae, vertebrate flesh and dead arthropods (Pukowski, 1933). The life history of members of the Silphinae is relatively simple. Mating usually occurs in the spring or early summer; eggs are laid shallowly in soil and hatch in 5 to 7 days. Larvae pass through three instars in 12 to 22 days, consuming the same types of food as adults. Pupation occurs in the soil in a pupal cell, with the prepupal/pupal stages lasting 9 to 30 days. Emer- gence of adults occurs in mid to late summer, usually after the previous year's adults have died, with subsequent hibernation as unmated adults (Balduf, 1935). The life history of members of the Nicrophorinae illustrates the most advanced degree of parental care in Coleoptera and has been described in detail by Pukowski (1933) and discussed by Wil- son (1971). A pair of adult beetles bury a vertebrate carcass or a portion thereof, form a food ball by masticating and manipulat- ing it, lay eggs in a depression on top of the ball and feed the young by regurgitation in the early stages of development. Lar- vae are protected from predators by the adults. The larvae, although able to feed on the food ball when mature, have a high mortality in the absence of adults. Pupation occurs in the orig- inal chamber, at which time the adults either die or leave. Coverage of Bibliography An investigator who wishes to survey the literature for information on silphid biology presently has few options. Com- monly used bibliographic files stored in computers, such as AGRICOLA, BIOSIS and CAB, do not contain material prior to 1969. The most recent bibliography of the world Silphidae is the Cole- opterorum Catalogus of 1928 by M. H. Hatch. This work primarily covers taxonomy and distribution and contains few other types of citations. The most recent bibliography of silphids in the United States is contained in C. W. Leng's Catalogue of the Col- eoptera of America, 1920, with supplements covering the period from 1920 to 1947. The Leng catalog also primarily covers tax- onomy and distribution, with some biology references. Unfortun- ately, both the Leng and Hatch citations are abbreviated, without 2 t~tles, and are thus of reduced value. Fortunately a catalogue of American Silphidae north of Mexico is in press (Peck and Miller), which should bring the taxonomy and distribution liter- ature on this topic up to date. Given the coverage of the various catalogues, it was decided to include in this bibliography of Silphidae through 1981: 1. Publications in the United States containing information other than taxonomy and distribution. 2. Non-United States publications after 1927 containing any information on silphids. 3. Non-United States publications prior to 1928 containing information other than taxonomy and distribution. Several exceptions to this coverage have been made to allow for inclusion of major reference works, such as catalogues, manuals for identification, general bibliographies, etc. Because of the author's particular interests in behavior and ecology, certain subject areas may be more completely covered than others. A concerted effort was made to survey the litera- ture involving ground beetles, inhabitants of nests and burrows, disease transmission, stomach analyses of vertebrate predators and medico-legal entomology. Taxonomic works and faunal lists generally were avoided; only those that either had identification keys or information on behavior-ecology were cited. Format of Citations The Harvard system of citation was employed with several refinements. Foreign language references are presented in both the original language and in English translation, with the ex- ception of Russian and Chinese, which are listed only in English. An English summary contained in a foreign language publication is so indicated. If a reference is a book, the pages containing silphid information are listed. Certain parenthetical information appears at the end of each citation. When three or fewer species are mentioned in the ref- erence, they are listed, using the following generic abbrevia- tions: ? A= Ablattaria N = Necrophorus Aca = Acanthopsilus P = Phosphuga Ac= Aclypea Pt= Pteroloma Ag= Agyrtes Pto = Ptomaphila Ap = Apteroloma S = Silpha B = Blitophaga T = Thanatophilus Ne= Necrodes X = Xylodrepa Ne= Necrophilus 3 \.. If more than three species are mentioned, only the number of species is indicated. When species are not identified, they are then indicated as "Silphidae spp.". The information in each reference is additionally coded by subject according to the following scheme: 1. SYSTEMATICS. Taxonomy, geographic distribution, evolu- tion, speciation, diagnostic aids to identification, fossils. lA. Keys for identification lB. Fossils 2. MORPHOLOGY. Descriptions of internal and external struc- tures, histology, cytology, pathology, genetics, internal and external parasites. 2A. Parasites 3. PHYSIOLOGY. Neural, metabolic, sensory processes; organ and system functions; biochemistry and toxicology. 4. LIFE HISTORY AND HABITS. Life cycle, phenology, growth and development, biological rhythms, feeding and oviposition habits. 4A. Nests and burrows of vertebrates 4B. Stomachs of vertebrate consumers 4C. Human cadavers (medico-legal entomology) 4D. Disease transmission 5. POPULATION PARAMETERS. Population dynamics and modeling; effects of biotic factors on natality, mortality, survivorship and fecundity. 6. BEHAVIOR. Descriptive and experimental; foraging, feeding, reproductive, antipredator; locomotory, orientation, communication .. 7. ECOLOGY. Effect of physical environmental factors on species survival parameters. 8. METHODS. Sampling, trapping, rearing techniques, equipment. A particular reference may be coded in several subject areas. Only those publications that have been personally examined are coded. Index Each citation is assigned a unique number. A subject index is presented at the end of the bibliographic listings with cita- tions listed by number under the appropriate subject heading. Included in the bibliography are citations that were not obtain- able but are listed because their titles indicated silphid in- formation. These titles have.not been assigned a number and thus have not been coded or included in the index. 4 Acknowledgements The author's introduction to the silphid literature occurred while performing his dissertation research with the support of the Smithsonian Institution (1974 to 1976). The literature search largely was conducted while the author was employed as a faculty research associate in the Department of Entomology, University of Maryland, College Park (1979-1980). Personnel at the National Agricultural Library (USDA) and the University of Georgia Science Library have been of considerable value in the final phases of preparation (1980-1981). J. A. Payne (USDA, Byron, GA) and S. B. Peck (Carleton Univ., Ottawa) made available their extensive personal reprint files and were of assistance in other ways. P. P. Shubeck (Montclair St. College, NJ) reviewed several early drafts of the bibliography and was a source of encouragement. The difficult task of typing the manuscript was performed by J. Garrick, P. J. Young and L. King. Financial support for publication was provided by the Department of Ento- mology, University of Maryland, and the Maryland Agricultural Experiment Station. 5 Bibliography ABBOTT, C. E. 1927. An aberrant specimen of Necrophorus 001 tornentosus (Weber) ( Coleop. : Silphidae). Entornol. News 38:39. [2]. ABBOTT, C. E. 1927. Experimental data on the olfactory 002 sense of Coleoptera, with special reference to the Necrophori. Ann. Entornol. Soc. Arner. 20:207-216. [ 4 spp. ; 3 , 4, 6 J . ABBOTT, C. E. 1927. Further observations on the olfactory 003 powers of the Necrophori. Ann. Entornol. Soc. Arner. 20:550-553. [N. orbicollis; 3,6]. ABBOTT, C. E. 1936. The toxicity of Trirnethylarnine for 004 Necrophorus orbicollis (Say). Psyche 43:37-39. [3]. ABBOTT, C. E. 1937. The necrophilous habit in Coleoptera. 005 Bull. Brooklyn Entornol. Soc. 32:202-204. [N. orbicollis; 4]. AKOPYAN, M. M. 1953. 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Zur biologie des Necrophorus vestigator 024 Herschel nebst beschreibung der larve und nymphe. 7 The biology of Necrophorus vestigator Hers. with descriptions of larva and pupa. Entomol. Blat. 8:197-203. [2,4]. BICKHARDT, H. 1909. Die varietaten unserer schwarzen 025 totengraberarten. / The varieties of black burying beetle species. Entomol. Blat. 5:73-76. [N. germanicus, humator; l]. BILY, S., and J. PAVLICEK. 1970. A comparison of the soil 026 coleopterous fauna in three types of meadow in Bohemia. Acta Entomol. Bohemoslov. 67:287-303. [S. obscura, A. laevigata; 4,5,7]. BJORKMAN, F., and M. H. HATCH. 1939. Note on Silpha (Than- 027 atophilus) coloradensis Wickham. Pan-Pacif. Entomol.-- 15:96. [4]. BLAIR, K. G. 1931. A note on the arthropod fauna of 028 rooks' nests. Entomol. Mon. Mag. 67:35-36. [P. atrata; 4A]. BLAISDELL, F. E. 1901. Pupa of Necrophilus hydrophiloides. 029 Entomol. News 12:263-264. [2]. BLATCHLEY, W. S. 1910. An illustrated descriptive cata- 030 logue of the Coleoptera or beetles (exclusive of the Rhynchophora) known to occur in Indiana. (Silphidae, p. 272-291). Indiana Dept. Geol. Nat. Resour., Bull. No. 1. 1386 pp. [l,lA,4]. BLAZEJEWSKI, F. 1956. Chrzaszcze trupozerne rezerwatu 031 cisowego Wierzchlas. / Necrophorus beetles of the Wierzchlas Yew-tree Preserve. (Eng. summary.) Zesz. Naukowa Univ. Torun. Biol. 1:63-90. [4,5,7]. BLISS, R. Q. 1949. Necrophorus secondary sexual dif- 032 ferences. Entomol. News 60:197-204. [2]. BOCKELER, W. 1977. Fortfuhrende untersuchungen an kafern 033 als experimentellen transitwirten von Trichinella spiralis (Owen 1835) / Further examinations of beetles as transit-hosts of Trichinella spiralis (Owen 1835). (Eng. summary.) Zool. Anz., Jena 199:42-76. [B. transcaspica, T. ferrugatus, N. humator; 2A,3,4D]. 8 I BOECKH, J. 1962. Elektrophysiologische untersuchungen an 034 einzelnen geruchsrezeptoren auf den antennen des toten- grabers (N~crophorus, Coleoptera). / Electrophysiological investigations on single odor receptors of the antenna of carrion beetles (Necrophorus, Coleoptera). Z. Vergl. Physiol. 46:212-248. L2,3,6]. BOLIVAR Y PIELTAIN, C. 1940. Sohre algunos insectos 035 alpinos de los volcanes de Mexico. / On some alpine insects of Mexican volcanoes. Rev. Soc. Mexicana Hist. Nat. 1:175-177. [Ap. sp.; l]. BORCHERS, H. A. 1968. Distribution and composition of populations of mites symbiotic on Decrophilous beetles. Ph.D. thesis, Iowa State University. 126 pp. BORNEMISSZA, G. F. 1957. An analysis of arthropod succes- 036 sion in carrion and the effect of its decomposition on the soil fauna. Aust. J. Zool. 5:1-12. [Pto. lachrymosa; 4 , 5 , 6 , 7 ] ? BOVING, A. G., and F. C. CRAIGHEAD. 1930. An illustrated 037 synopsis of the principal larval forms of the order Coleoptera. Entomol. Amer. 11:1-351. [S. noveboracensis, Ne. littoralis; 1,2]. BRADLEY, J. C. 1930. A manual of the genera of beetles of 038 America north of Mexico. (Silphidae, pp. 61-62). Ithaca, New York: Daw, Illston and Co. 360 pp. [l,lA]. BRAGG, A. N. 1957. Use of carrion by the beetle, Canthon 039 laevis (Coleoptera, Scarabaeidae). Southwestern Natur. 2:173. [4]. BRASS, P. 1914. Abdominal segment der kaferlarven als 040 bewegungsorgan / Abdominal segments of beetle larvae as mobility organs. Zool. Jahrb. Abt. Syst. Geog. Biol. Tiere 37:65-122. [S. rugosa; 2]. BREWER, J. W., and T. R. BACON. 1975. Biology of the 041 carrion beetle Silpha ramosa Say. Ann. Entomol. Soc. Amer. 68:786-790. [2,4,5]. BRIVIO, C. 1954. Studi sui Necroohorini (Coleoptera, Silphidae) / Studies in the Necrophorini. Natura 44:85-88. BRUNETTI, B. 1931. Ricerche sul proventricolo degli 042 insetti. / Research on the proventriculus of insects. Atti Soc. Toscana Sci. Nat. Mem. 41:110-146. [S. carinata; 2]. 9 ,......- BUNING, J. 1979. 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Eine neue gattung und art, Pseudo- 296 silpha n. gen. roderi n. sp., aus dem Bhutan-Himalaya. Coleoptera, Silphidae, Pterolomini. /Anew genus and species, Pseudosilpha roderi, from the Bhutan-Himalaya. Coleoptera: Silphidae: Pterolomini. (Eng. summary.) Entomol. Basil. 3:103-108. [l]. SCHAWALLER, W. 1978. Zwei neue Pteroloma-arten aus dem 297 Nepal-Himalaya, erganzende bemerkungen zu und Katalog der Pterolomini. / Two new species of Pteroloma from Nepal-Himalaya, with supplementary annotations to the Catalogue of the Pterolomini. Senckenbergiana Biol. 58:171-184. [10 spp.; l]. SCHAWALLER, W. 1979. Morphologische variation bei Silpha 301 tristis und synonymie von Silpha franzi (Coleoptera, Silphidae). / Morphological variation of Silph~ t~istis and synonomy of Silpha franzi. (Eng. summary.) Stutt- garter Beitr. Naturk., Ser. A., 328:1-8. [1,2]. SCHAWALLER, W. 1980. 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SYSTEMATICS: 008, 012, 013, 014, 015, 016, 017? 025, 030, 035, 037, 038, 043, 044, 058, 059, 064, 066, 074, 075, 078, 079, 082, 087, 097, 099, 102? 116, 117, 118, 119, 120, 121, 122, 123, 124, 138, 140, 141, 142, 143, 147, 167, 168, 169, 181, 183, 186, 188, 190, 192, 194, 202, 203, 207, 209, 210, 216, 228, 238, 300, 301, 302, 303, 314, 316, 318, 319, 320, 331, 343, 355, 356, 363, 364, 366, 370, 379, 382 lA: 012, 016, 017, 030, 038, 061, 063, 097, 116, 117, 122, 124, 139, 163, 170, 178, 246, 255, 280, 316, 318 lB: 018, 056, 057, 208, 217, 222, 223, 263 2. MORPHOLOGY: 001, 012, 013, 016, 024, 029, 032, 034, 037, 040, 041, 042, 043, 044, 055, 062, 063, 067, 074, 080, 081, 092, 093, 105, 107, 171, 184, 188, 189, 200, 206, 237, 246, 248, 276, 284, 293, 294, 301, 302, 310, 317, 333, 334, 335, 342, 345, 347, 349, 350, 351, 353, 356, 358, 368, 370, 376, 379, 381, 383, 384, 385 2A: 033, 051, 065, 077, 115, 173, 215, 231, 233, 358, 367 3. PHYSIOLOGY: 002, 003, 004, 033, 034, 080, 081, 206, 237, 291, 292, 368 4. LIFE HISTORY AND HABITS: 002, 005, 006, 008, 011, 013, 015, 017, 019, 021, 023, 024, 026, 027, 030, 031, 036, 039, 041, 045, 047, 049, 050, 052, 053, 054, 055, 059, 060, 063, 064, 066, 068, 069, 070, 071, 072, 075, 076, 082, 083, 084, 085, 086, 088, 090, 098, 099, 100, 102, 103, 104, 106, 107, 108, 109, 110, 111, 117, 122, 124, 125, 126, 127, 128, 129, 130, 131, 132, 133, 134, 135, 136, 137, 144, 145, 146, 147, 148, 150, 151, 152, 154, 155, 156, 157, 164, 165, 166, 167, 168, 172, 173, 174, 175, 180, 182, 184, 185, 188, 190, 192, 194, 195, 199, 201, 206, 209, 213, 215, 216, 218, 219, 221, 225, 227, 229, 230, 231, 232, 240, 243, 246, 247, 248, 249, 250, 251, 252, 253, 254, 255, 256, 257, 258, 260, 264, 265, 266, 267, 268, 270, 271, 272, 273, 274, 276, 278, 279, 282, 283, 285, 286, 287, 289, 290, 291, 292, 293, 294, 304, 305, 307, 309, 312, 313, 315, 321, 322, 323, 324, 325, 3 2 6, 328, 329, 330, 331, 332, 336, 338, 339, 340, 341, 342, 344, 346, 349, 354, 356, 357, 359, 360, 361, 362, 364, 369, 371, 372, 373, 374, 375, 376, 377, 378, 379, 387 46 4A: 028, 076, 089, 094, 113, 114, 153, 197, 198, 205, 211, 212, 239, 248 4B: 007, 091, 101, 160, 161, 162, 196, 311, 337, 380 4C: 009, 010, 095, 158, 176, 177, 191, 204, 214 4D: 022, 033, 073, 149, 244 5 . POPULATION PARAMETERS: 020, 021, 026, 031, 036, 041, ? 046, 049, 068, 069, 070, 076, 096, 104, 110, 111, 112, 146, 148, 152, 154, 155, 156, 164, 165, 166, 194, 195, 201, 206, 229, 241, 242, 245, 247, 249, 258, 260, 275, 304, 306, 3 0 9, 322, 357, 369, 387 6. BEHAVIOR: 002, 003, 021, 034, 036, 062, 076, 084, 085, 088, 098, 104, 128, 129, 130, 131, 132, 133, 134, 135, 136, 137, 159, 175, 184, 185, 188, 200, 206, 219, 220, 224, 226, 232, 235, 236, 237, 265, 268, 270, 271, 276, 288, 321, 325, 365, 368,. 386 7 . ECOLOGY: 008, 018, 020, 021, 026, 031, 036, 046, 049, 053, 057, 068, 069, 070, 076, 084, 085, 096, 104, 110, 111, 112, 146, 148, 152, 154, 155, 156, 164, 165, 166, 172, 186, 188, 194, 195, 201, 2 0 6, 217, 229, 245, 247, 249, 251, 258, 260, 265, 268, 272, 275, 278, 279, 281, 304, 306, 3 0 9, 322, 326, 352, 357, 359, 360, 361, 369, 387 8. METHODS: 048, 069, 075, 098, 179, 193, 204, 234, 259, 277, 308, 323, 327 47 Contribution number 6593 and miscellaneous publications number 981 of the Maryland Agricultural Experiment Station, Depart- ment of Entomology. The University of Maryland is an equal opportunity institution with respect to both education and employment. The university's policies, programs and activities are in conformance with pertinent Federal and state laws and regulations on nondis- crimination regarding race, color, religion, age, national origin, sex and handicap. Inquiries regarding compliance with Title VI of the Civil Rights Act of 1964, as amended; Title IX of the 1972 Education Amendments: Section 504 of the Rehabil- itation Act of 1973; or related legal requirements should be directed to the Assistant to the Director for Affirmative Action, Symons Hall, University of Maryland, College Park, MD 20742. Tel. (3Cl) 454-5981.