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Issue: Vol. 5839 No. 1: 26 Jun. 2026
Type: Article
Published: 2026-06-26
DOI: 10.11646/zootaxa.5839.1.3
Page range: 71-109
Abstract views: 78
PDF downloaded: 10

The Silphinae (Staphylinidae) and Agyrtidae of Kentucky with an illustrated key to the fauna of eastern North America

Department of Entomology; University of Kentucky; S225 Agriculture Science Center North; 1100 South Limestone; Lexington; Kentucky 40546
College of Medicine; University of Kentucky; 230C Multidisciplinary Science Building; 725 Rose St.; Lexington; KY 40536
1308 Lago Drive; Frankfort; KY 40601
Department of Entomology; University of Kentucky; S225 Agriculture Science Center North; 1100 South Limestone; Lexington; Kentucky 40546; College of Medicine; University of Kentucky; 230C Multidisciplinary Science Building; 725 Rose St.; Lexington; KY 40536
Coleoptera Biodiversity checklist regional fauna

Abstract

The carrion beetle fauna (Staphylinidae: Silphinae) of eastern North America has seen some recent changes and additions in the past decade. Two species of Silpha have been introduced and have become established and Nicrophorus vespilloides was discovered to be two species. In this paper, we present a fully illustrated key to the Silphinae of eastern North America (U.S. states entirely east of the Mississippi River and Canadian provinces east of the Ontario-Manitoba border) that reflects these faunal changes. We focus on the silphine fauna of Kentucky and include the single species of Agyrtidae (primitive carrion beetles), as common sampling methods will often pick up members of both groups. We include 22 species of Silphinae and one species of Agyrtidae in our key, and we report one species of Agyrtidae and 11 species of Silphinae from Kentucky, including one new state record (Nicrophorus defodiens). We present the following information for each eastern North American species in an annotated list: diagnostic characters, recent taxonomic changes (if applicable), life history information and more detailed information for Kentucky species including counties, collecting years and months, collections housing the specimens, and collecting methods. Distribution maps for Kentucky specimens are presented. Using this dataset, the bias-corrected iChao2 species richness estimator predicted that 12 species should be found in Kentucky, indicating that our sampling is likely complete.

References

  1. Aleksandrowicz, O. & Komosiński, K. (2005) On the fauna of carrion beetles (Coleoptera, Silphidae) of Mazurian lakeland (north-eastern Poland). In: Skłodowski, J., Huruk, S., Barševskis, A. & Tarasiuk, S. (Eds.), Protection of Coleoptera in the Baltic Sea Region. Warsaw Agricultural University Press, pp. 147–153.
  2. Anderson, R.S. (1982a) Burying beetle larvae: Nearctic Nicrophorus and Oriental Ptomascopus morio (Silphidae). Systematic Entomology, 7 (3), 249–264. https://doi.org/10.1111/j.1365-3113.1982.tb00443.x
  3. Anderson, R.S. (1982b) Resource partitioning in the carrion beetle (Coleoptera: Silphidae) fauna of southern Ontario: Ecological considerations. Canadian Journal of Zoology, 60, 1314–1325. https://doi.org/10.1139/z82-178
  4. Anderson, R.S. & Peck, S.B. (1985) The Carrion Beetles of Canada and Alaska (Coleoptera: Silphidae and Agyrtidae). The Insects and Arachnids of Canada, Part 13. Canadian Government Publishing Centre, Ottawa, 121 pp.
  5. Arnett, R.H. (1946) Coleoptera notes I: Silphidae. The Canadian Entomologist, 78 (7–8), 131–134. https://doi.org/10.4039/Ent78131-7
  6. Backlund, D.C., Marrone, G.M., Williams, C. & Tilmon, K. (2008) Population estimate of the endangered American burying beetle, Nicrophorus americanus Olivier (Coleoptera: Silphidae) in South Dakota. The Coleopterists Bulletin, 62, 9–15. https://doi.org/10.1649/982.1
  7. Baldwin, B. (1975) A Faunal Study of Illinois Silphidae (Coleoptera). Master’s Thesis no. 2487, Eastern Illinois University, 45 pp. [https://thekeep.eiu.edu/theses/2487]
  8. Bedick, J.C., Ratcliffe, B.C., Hoback, W.W. & Higley, L.G. (1999) Distribution, ecology, and population dynamics of the American burying beetle [Nicrophorus americanus Olivier (Coleoptera, Silphidae)] in south-central Nebraska, USA. Journal of Insect Conservation, 3, 171–181. https://doi.org/10.1023/A:1009684926873
  9. Beninger, C.W. (1994) Phenology, reproductive biology and habitat associations of Nicrophorus Fab. (Coleoptera: Silphidae) of the Mer Bleue Ontario area (Ottawa, Canada). Memoirs of the Entomological Society of Canada, 126, 135–143. https://doi.org/10.4039/entm126169135-1
  10. Beninger, C.W. & Peck, S.B. (1992) Temporal and spatial patterns of resource use among Nicrophorus carrion beetles (Coleoptera: Silphidae) in a Sphagnum bog and adjacent forest near Ottawa, Canada. Canadian Entomologist, 124, 79–86. https://doi.org/10.4039/Ent12479-1
  11. Benowitz, K.M., Amukamara, A.U., McKinney, E.C. & Moore, A.J. (2019) Development and the effects of extended parenting in the cold-breeding burying beetle Nicrophorus sayi. Ecological Entomology, 44, 11–16. https://doi.org/10.1111/een.12670
  12. Bergamin, S. & Smits, A. (2015) How do the molluscivorous beetles Carabus granulatus and Phosphuga atrata (Insecta, Coleoptera) deal with sinistral and dextral prey? Vita Malacologica, 13, 49–51.
  13. Bivand, R.S., Pebesma, E.J. & Gomez-Rubio, V. (2013) Applied spatial data analysis with R, Second edition. Springer, New York, XVIII + 405 pp. https://doi.org/10.1007/978-1-4614-7618–4
  14. Blouin-Demers, G. & Weatherhead, P.J. (2000) A novel association between a beetle and a snake: parasitism of Elaphe obsoleta by Nicrophorus pustulatus. Ecoscience, 7, 395–397. https://doi.org/10.1080/11956860.2000.11682609
  15. Blouin-Demers, G., Weatherhead, P.J. & Row, J.R. (2004) Phenotypic consequences of nest-site selection in black rat snakes (Elaphe obsoleta). Canadian Journal of Zoology, 82, 449–456. https://doi.org/10.1139/z04-014
  16. Brewer, J.W. & Bacon, T.R. (1975) Biology of the carrion beetle, Silpha ramosa Say. Annals of the Entomological Society of America, 68, 786–790. https://doi.org/10.1093/aesa/68.5.786
  17. BugGuide (2021) Available from: https://bugguide.net/node/view/15740 (accessed 16 December 2025)
  18. Burke, K.W., Groulx, A.F. & Martin, P.R. (2023) The competitive exclusion–tolerance rule explains habitat partitioning among co-occurring species of burying beetles. Ecology, 105, e4208. https://doi.org/10.1002/ecy.4208
  19. Burke, K.W., Wettlaufer, J.D., Beresford, D.V. & Martin, P.R. (2020) Habitat use of co-occurring burying beetles (genus Nicrophorus) in southeastern Ontario, Canada. Canadian Journal of Zoology, 98, 591–602. https://doi.org/10.1139/cjz-2019-0287
  20. Cai, C., Tihelka, E., Giacomelli, M., Lawrence, J.F., Ślipiński, A., Kundrata, R., Yamamoto, S., Thayer, M.K., Newton, A.F., Leschen, R.A.B., Gimmel, M.L., Lu, L., Engel, M.S., Bouchard, P., Huang, D., Pisani, D. & Donoghue, P.C.J. (2022) Integrated phylogenomics and fossil data illuminate the evolution of beetles. Royal Society Open Science, 9, 211771. https://doi.org/10.1098/rsos.211771
  21. Canitz, J., Sikes, D.S., Knee, W., Baumann, J., Haftaro, P., Steinmetz, N., Nave, M., Eggert, A.-K., Hwang, W. & Nehring, V. (2021) Cryptic diversity within the Poecilochirus carabi mite species complex phoretic on Nicrophorus burying beetles: phylogeny, biogeography, and host specificity. Molecular Ecology, 31, 658–674. https://doi.org/10.1111/mec.16248
  22. Chao, A., Ma, K.H., Hseih, T.C. & Chiu, C.H. (2016) SpadeR: Species-Richness Prediction and Diversity Estimation with R. R package Version 0.1.1. Available from: https://CRAN.R-project.org/package=SpadeR (accessed 24 December 2022) https://doi.org/10.32614/cran.package.spader
  23. Chapman, E.G., Leavengood, J.M., Jr. & Dupuis, J.R. (2025) The Cleridae and Thanerocleridae of eastern North America, with illustrated keys, updated distributions, and special emphasis on the Kentucky fauna. Zootaxa, 5639 (1), 001–088. https://doi.org/10.11646/zootaxa.5639.1.1
  24. Chapman, E.G., Richards, A.B. & Dupuis, J.R. (2023) The longhorn beetles (Coleoptera: Cerambycidae) of Kentucky with notes on larval hosts, adult nectar use, and semiochemical attraction. Zootaxa, 5229 (1), 001–089. https://doi.org/10.11646/zootaxa.5229.1.1
  25. Chiu, C.-H., Wang, Y.-T., Walther, B.A. & Chao, A. (2014) An improved nonparametric lower bound of species richness via a modified good-turing frequency formula. Biometrics, 70 (3), 671–682. https://doi.org/10.1111/biom.12200
  26. Clark, C.U. (1895) On the food habits of certain dung and carrion beetles. Journal of the New York Entomological Society, 3 (2), 3–61. https://www.jstor.org/stable/25002650
  27. Cole, A.C. (1942) Observations of three species of Silpha. American Midland Naturalist, 28, 161–163. https://doi.org/10.2307/2420698
  28. Conrad, T., Roberts, L., Steiger, S. & Ringlein, M. (2024) Vibrations from the crypt: Investigating the possibility of vibrational communication in burying beetles. Entomologia Experimentalis et Applicata, 172, 1154–1165. https://doi.org/10.1111/eea.13519
  29. Cook, L.M., Smith, A.N., Meyers, P.J., Creighton, J.C. & Belk, M.C. (2019) Evidence for differential diel activity patterns in two co-occurring species of burying beetles (Coleoptera: Silphidae: Nicrophorinae). Western North American Naturalist, 79 (2), 270–274. https://doi.org/10.3398/064.079.0212
  30. Creighton, J.C., Astarache, R.B., Lomolino, M.V. & Belk, M.C. (2009) Effect of forest removal on the abundance of the endangered American burying beetle, Nicrophorus americanus (Coleoptera: Silphidae). Journal of Insect Conservation, 13, 37–43. https://doi.org/10.1007/s10841-007-9115-4
  31. Davis, L.R. (1980) Notes on beetle distributions with a discussion of Nicrophorus americanus Olivier and its abundance in collections (Coleoptera: Scarabaeidae, Lampyridae, and Silphidae). The Coleopterists Bulletin, 34 (2), 245–252. https://www.jstor.org/stable/4000138
  32. DeMarco, K.V. & Martin, P.R. (2020) A case of a pustulated carrion beetle (Nicrophorus pustulatus, Coleoptera: Silphidae) burying live tree swallow (Tachycineta bicolor, Passeriformes: Hirundinidae) nestlings under the nest. The Canadian Field-Naturalist, 134, 217–221. https://doi.org/10.22621/cfn.v134i3.2369
  33. Dorsey, C.K. (1940) A comparative study of the larvae of six species of Silpha (Coleoptera: Silphidae). Annals of the Entomological Society of America, 33, 120–139. https://doi.org/10.1093/aesa/33.1.120
  34. Eggert, A.K. & Müller, J.K. (1992) Joint breeding in female burying beetles. Behavioral Ecology and Sociobiology, 31 (4), 237–242. https://doi.org/10.1007/BF00171678
  35. Eisner, T. & Meinwald, J. (1982) Defensive spray mechanism of a silphid beetle (Necrodes surinamensis). Psyche, 89, 357–367. https://doi.org/10.1155/1982/41643
  36. Engasser, E.L., Stone, R.L. & Jameson, M.L. (2021) Habitat associations of carrion beetles (Coleoptera: Silphidae) across a full annual cycle. Environmental Entomology, 50 (3), 605–614. https://doi.org/10.1093/ee/nvab011
  37. Essig, E.O. (1926) Insects of Western North America. MacMillian, New York, 1035 pp.
  38. Ferreira, R.N. (2017) First record of Silpha puncticollis Lucas, 1846 (Coleoptera: Silphidae, Silphinae) for North America. Arquivos Entomolóxicos, 17, 101–103. https://doi.org/10.5281/zenodo.12765131
  39. Fisher, R.M. & Tuckerman, R.D. (1986) Mimicry of bumble bees and cuckoo bumble bees by carrion beetles (Coleoptera: Silphidae). Journal of the Kansas Entomological Society, 50, 20–25. https://www.jstor.org/stable/25084730
  40. Gissler, C.F. (1880) Biological studies on Silpha ramosa. American Entomologist, 3, 265–267.
  41. Goe, M.T. (1919) Life history and habits of Silpha inaequalis Fab (Coleoptera). Entomological News, 30, 253–255.
  42. Grappone, B.A. (2025) Ecology and Taxonomy of Necrophilous Arthropods in Central Alberta. Master’s thesis, University of Alberta, 270 pp.
  43. Gries, C., Gilbert, E.E. & Franz, N.M. (2014) Symbiota—A virtual platform for creating oucher-based biodiversity information communities. Biodiversity Information Journal, 2014, e1114. https://doi.org/10.3897/BDJ.2.e1114
  44. Haitlinger, R. (2025) Mites (Acari) associated with Silphinae (Coleoptera: Staphylinidae): a worldwide review. Acarological Studies, 7 (2), 72–100. https://doi.org/10.47121/acarolstud.1724127
  45. Hall, C.L., Howard, D.R., Smith, R.J. & Mason, A.C. (2015) Marking by elytral clip changes stridulatory characteristics and reduces reproduction in the American burying beetle, Nicrophorus americanus. Journal of Insect Conservation, 19, 155–162. https://doi.org/10.1007/s10841-015-9755-8
  46. Hall, C.L., Mason, A.C., Howard, D.R., Padhi, A. & Smith, R.J. (2013) Description of acoustic characters and stridulatory pars stridens of Nicrophorus (Coleoptera: Silphidae): A comparison of eight North American species. Annals of the Entomological Society of America, 106 (5), 661–669. https://doi.org/10.1603/AN13001
  47. Hanley, G.A. & Cuthrell, D.L. (2008) The Carrion beetles of North Dakota, Including Descriptions and Identification Keys for the Entire North American Silphid Fauna. Cyril Moore Science Center Science Monograph #4, Minot State University, Minot, ND, 52 pp.
  48. Harrison, L. (2021) Thermal ecology and plasticity in the burying beetle, Nicrophorus orbicollis: An observational field study. Master’s thesis, Queen’s University, Kingston, Ontario, Canada, 56 p.
  49. Hatch, M.H. (1927) Studies on the Silphinae. Journal of the New York Entomological Society, 35, 331–71. https://www.jstor.org/stable/25004219
  50. Hatch, M.H. & Reuter, W., Jr. (1934) Coleoptera of Washington: Silphidae. University of Washington Publications in Biology, 1 (3), 147–162.
  51. Heinrich, B. (2012) A heretofore unreported instant color change in a beetle, Nicrophorus tomentosus Weber (Coleoptera: Silphidae). Northeastern Naturalist, 19, 345–352. https://doi.org/10.2307/41684241
  52. Hocking, M.D., Ring, R.A. & Reimchen, T.E. (2006) Burying beetle Nicrophorus investigator reproduction on Pacific salmon carcasses. Ecological Entomology, 31, 5–12. https://doi.org/10.1111/j.0307-6946.2006.00747.x
  53. Howden, A.T. (1950) The succession of beetles on carrion. Master’s Thesis, University of North Carolina, 83 p.
  54. Ikeda, H., Kagaya, T., Kubota, K. & Abe, T. (2008) Evolutionary relationships among food habit, loss of flight, and reproductive traits: Life-history evolution in the Silphinae (Coleoptera: Silphidae). Evolution, 62 (8), 2065–2079. https://doi.org/10.1111/j.1558-5646.2008.00432.x
  55. Ikeda, H., Kubota, K., Kagaya, T. & Abe, T. (2007) Flight capabilities and feeding habits of silphine beetles: Are flightless species really “carrion beetles”? Ecological Research, 22, 237–241. https://doi.org/10.1007/s11284-006-0012-1
  56. iNaturalist (2021) Available from: https://www.inaturalist.org (accessed 16 December 2025)
  57. Katakura, H. & Fukuda, H. (1975) Faunal makeup of ground and carrion beetles in Komiotoineppu, Hokkaido University Nakagawa Experimental Forest, northern Japan, with some notes on research problems. Research bulletin of the College of Experimental Forestry, Hokkaido University, 32, 75–92. https://hdl.handle.net/2115/20952
  58. Katovich, K., Kriska, N.L., Williams, A.H. & Young, D.K. (2005) Carrion Beetles (Coleoptera: Silphidae) of Wisconsin. The Great Lakes Entomologist, 38 (1), 30–41. https://doi.org/10.22543/0090-0222.2121
  59. Keller, M.L., Howard, D.R. & Hall, C.L. (2019) Spatiotemporal niche partitioning in a specious silphid community (Coleoptera: Silphidae Nicrophorus). The Science of Nature, 106, 57. https://doi.org/10.1007/s00114-019-1653-6
  60. Keller, W.L. & Heske, E.J. (2001) An observation of parasitism of black rat snake (Elaphe obsoleta) eggs by a beetle (Nicrophorus pustulatus) in Illinois. Transactions of the Illinois State Academy of Science, 94, 167–169.
  61. Knee, W. (2017) New Macrocheles species (Acari, Mesostigmata, Macrochelidae) associated with burying beetles (Silphidae, Nicrophorus) in North America. ZooKeys, 721, 1–32. https://doi.org/10.3897/zookeys.721.21747
  62. Kočárek, P. (2003) Decomposition and Coleoptera succession on exposed carrion of small mammal in Opava, the Czech Republic. European Journal of Soil Biology, 39 (1), 31–45. https://doi.org/10.1016/S1164-5563(02)00007-9
  63. Kozol, A.J., Scott, M.P. & Traniello, J.F.A. (1988) The American burying beetle, Nicrophorus americanus: studies on the natural history of a declining species. Psyche, 95, 167–176. https://doi.org/10.1155/1988/79403
  64. Lampert, L.L., Jr. (1977) Notes on Silpha inaequalis Fabricius (Coleoptera: Silphidae). The Coleopterists Bulletin, 31, 132. https://doi.org/10.5962/p.371951
  65. Lane, C. & Rothschild, M. (1965) A case of Müllerian mimicry of sound. Proceedings of the Royal Entomological Society of London. Series A, General Entomology, 40 (10-12), 156–158. https://doi.org/10.1111/j.1365-3032.1965.tb00305.x
  66. LaPlante, S. (1997) Premieres evidences de l’etablissement d’une population de Silpha tristis Illiger (Coleoptera: Silphidae) en Amerique du Nord. Fabreries, 22, 85–93.
  67. Lawrence, J.F. & Newton, A.F. (1982) Evolution and classification of beetles. Annual Review of Ecology and Systematics, 13, 261–290. https://doi.org/10.1146/annurev.es.13.110182.001401
  68. Leclerc, L., Bedard, N., Bloin, P., Djoumad, A. & Picq, S. (2025) First Records of Silpha (Phosphuga) atrata Linnaeus, 1758 (Coleoptera: Staphylinidae: Silphinae) in North America. The Coleopterists Bulletin, 79 (4), 651–658. https://doi.org/10.1649/0010-065X-79.4.651
  69. Leech, H.B. (1934) The family history of Nicrophorus conversator Walker. Proceedings of the Entomological Society of British Columbia, 31 (1934), 36–40.
  70. LeGros, D.L. & Beresford, D.V. (2010) Aerial foraging and sexual dimorphism in burying beetles (Silphidae: Coleoptera) in a central Ontario forest. Journal of the Entomological Society of Ontario, 141, 3–10.
  71. Lomolino, M.V., Creighton, J.C., Schnell, G.D. & Certain, D.L. (1995) Ecology and conservation of the endangered American burying beetle (Nicrophorus americanus). Conservation Biology, 9, 605–614. https://doi.org/10.1046/j.1523-1739.1995.09030605.x
  72. Lowe, A.J. & Lauff, R.F. (2012) Arboreal Burials in Nicrophorus spp. (Coleoptera: Silphidae). Psyche, 2012, 6 p. https://doi.org/10.1155/2012/578650
  73. MacRae, T.C. (2017) Why is this male carrion beetle “biting” the female’s antennae? Available from: https://beetlesinthebush.com/wp-content/uploads/2017/07/136_macrae_2017_male-carrion-beetle.pdf
  74. Mahlerová, K., Jakubec, P., Novák, M. & Růžička, Y. (2021) Description of larval morphology and phylogenetic relationships of Heterotemna tenuicornis (Silphidae). Scientific Reports, 11, 16973. https://doi.org/10.1038/s41598-021-94744-x
  75. Majka, C.G. (2011) The Silphidae (Coleoptera) of the Maritime Provinces of Canada. Journal of the Acadian Entomological Society, 7 (2011), 83–101.
  76. McCain, C.M., Monk, E., Hinson, K., Szewcyzk, T. & D’Oench, H. (2021) Key to the carrion beetles (Silphidae) of Colorado and neighboring states, Version 3: posted online December 2021. Available from: http://spot.colorado.edu/~mccainc/PDFs/Key to Silphidae of Colorado.pdf
  77. Mckenna, D.D., Shin, S., Ahrns, D., Balke, M., Beza-Beza, C., Clarke, D.J., Donath, A., Escalona, H.E., Friedrich, F., Letschi, H., Liu, S., Maddison, D., Mayer, C., Misof, B., Murin, P.J., Niehuis, O., Peters, R.S., Podsiadlowski, L., Pohl, H., Scully, E.D., Yan, E.V., Zhou, X., Slipinski, A. & Beutel, R.G. (2019) The evolution and genomic basis of beetle diversity. Proceedings of the National Academy of Sciences, 116 (49), 24729–24737. https://doi.org/10.1073/pnas.1909655116
  78. Mckenna, D.D., Wild, A.L., Kanda, K., Bellamy, C.L., Buetel, R.G., Caterino, M.S., Farnum, C.W., Hawks, D.C., Ivie, M.A., Jameson, M.L., Leschen, R.A.B., Marvaldi, A.E., McHugh, J.V., Newton, A.F., Robertson, J.A., Thayer, M.K., Whiting, M.F., Lawrence, J.F., Slipinski, A., Maddison, D.R. & Farrel, B.D. (2015) The beetle tree of life reveals that Coleoptera survived end‐Permian mass extinction to diversify during the Cretaceous terrestrial revolution. Systematic Entomology, 40 (4), 835–880. https://doi.org/10.1111/syen.12132
  79. Merrick, M.J. & Smith, R.J. (2004) Temperature regulation in burying beetles (Nicrophorus spp.: Coleoptera: Silphidae): Effects of body size, morphology and environmental temperature. Journal of Experimental Biology, 207, 723–733. https://doi.org/10.1242/jeb.00807
  80. Milne, L.J. & Milne, M.J. (1944) Notes on the behavior of burying beetles (Nicrophorus spp.). Journal of the New York Entomological Society, 52, 311–327. https://www.jstor.org/stable/25005075
  81. Mroczkowski, M. (1949) Notes on the successive appearance of some species of the genera Nicrophorus and Neonicrophorus. Polskie Pismo Entomologiczne, 19, 196–199.
  82. Mullins, P.L., Riley, E.G. & Oswald, J.D. (2013) Identification, distribution, and adult phenology of the carrion beetles (Coleoptera: Silphidae) of Texas. Zootaxa, 3666, 221–251. https://doi.org/10.11646/zootaxa.3666.2.7
  83. Murgia, G., Loi, F., Cappai, S. & Cogoni, M.P. (2022) Entomological investigation of the main entomatic adversities for terrestrial gastropods Helix aspersa Müller (Mollusca Gastropoda Pulmonata): A preliminary study in Sardinian heliciculture farms. Insects, 13 (7), 660. https://doi.org/10.3390/insects13070660
  84. Newton, A.F. (1997) Review of Agyrtidae (Coleoptera), with a new genus and species from New Zealand. Annales Zoologici (Warszawa), 47 (1/2), 111–156.
  85. Newton, A.F. (2025) StaphBase: Staphyliniformia world catalog database (version Aug 2022). In: Bánki, O., Roskov, Y., Döring, M., Ower, G., Hernández Robles, D. R., Plata Corredor, C. A., Stjernegaard Jeppesen, T., Örn, A., Vandepitte, L., Hobern, D., Schalk, P., DeWalt, R. E., Ma, K., Miller, J., Orrell, T., Aalbu, R., Abbott, J., Adlard, R., Aedo, C., Aescht, E., Alexander, S., Alonso-Zarazaga, M. A., Alvarez, B., Andrella, G. C., Antonietto, L. S., Arango, C., Artois, T., Atahuachi Burgos, M., Atkinson, S., Atwood, J. J., Bagnatori Sartori, Â. L., Bailly, N., Baixeras, J., Baker, E., Balan, A., Bamber, R., Bandyopadhyay, S., Barber-James, H., Barbosa Pinto, R., Barrett, R., Bartolozzi, L., Bartsch, I., Beccaloni, G., Bellamy, C. L., Bellan-Santini, D., Bellinger, P. F., Ben-Dov, Y., Blasco-Costa, I., Boatwright, J. S., Bock, P., Borges, L. M., Bossard, R. L., Bota-Sierra, C., Bouchard, P., Bourgoin, T., Boury-Esnault, N., Boxshall, G., Boyko, C., Brandão, S., Braun, H., Bray, R., Brehm, G., Brinda, J. C., Brock, P. D., Broich, S. L., Brown, J., Brown, S., Bruce, N., Brullo, S., Bruneau, A., Bush, L., Büscher, T., Błażewicz-Paszkowycz, M., Cabras, A., Cairns, S., Calonje, M., Cardinal-McTeague, W., Cardoso, D., Cardoso, L., Castilho, R. C., Castro Silva, I. C., Cervantes, A., Chevillotte, H., Choo, L. M., Christiansen, K. A., Cianferoni, F., Cigliano, M. M., Clarke, R., Cobra e Monteiro, T., Collins, A., Compton, J., Copilaș-Ciocianu, D., Corbari, L., Cordeiro, R., Cortés-Hernández, K., Costello, M., Crameri, S., Cruz-López, J. A., & Cárdenas, P., Daly, M., Daneliya, M., Dauvin, J.-C., Davie, P., De Broyer, C., De Grave, S., De Lima, H. C., De Prins, J., De Prins, W., De la Estrella, M., DeSalle, R., Decker, P., Decock, W., Delgado-Salinas, A., Deliry, C., Dellapé, P. M., Den Heyer, J., Dijkstra, K.-D., Dmitriev, D. A., Dohrmann, M., Dorado, Ó., Dorkeld, F., Downey, R., Duan, L., Díaz, M.-C., Eades, D. C., Egan, A. N., Eitel, M., El Nagar, A., Emig, C. C., Engel, M. S., Esquete Garrote, P., Evans, G. A., Evenhuis, N. L., Falcão, M., Farruggia, F., Fauchald, K., Fautin, D., Favret, C., Fišer, C., Forró, L., Fortuna-Perez, A. P., Fortune-Hopkins, H., Fritsch, P., Froese, R., Fuchs, A., Fujimoto, S., Furuya, H., Gagnon, E., Garic, R., Garnett, S., Gasca, R., Gattolliat, J.-L., Gerken, S., Gibson, D., Gibson, R., Gielis, C., Gilligan, T., Giribet, G., Girón Duque, J. C., Gittenberger, A., Giusso del Galdo, G., Gofas, S., Goncharov, M., Gondim, A. I., Goodwin, C., Govaerts, R., Grabowski, M., Granado, A. de A., Gray, A., Gregório, B. de S., Grehan, J. R., Grether, R., Grimaldi, D. A., Gross, O., Guerra-García, J. M., Guglielmone, A., Guilbert, E., Gusenleitner, J., Haas, F., Hadfield, K. A., Hajdu, E., Hassler, M., Hastriter, M. W., Hausmann, A., Hayward, B. W., Hendrycks, E., Henry, T. J., Hernandes, F. A., Hernandez, F., Hernández-Crespo, J. C., Herrera Bachiller, A., Hine, A., Hodson, A., Hoeksema, B., Hoenemann, M., Holstein, J., Hooge, M., Hooper, J., Hopkins, H., Horak, I., Horton, T., Hošek, J., Hughes, C., Hughes, L., Huys, R., Häuser, C., Janssens, F., Jaume, D., Javadi, F., Jazdzewski, K., Johnson, K. P., Jordão, L., Jóźwiak, P., Kakui, K., Kallies, A., Kamiński, M. J., Kanda, K., Karanovic, I., Kathirithamby, J., Kelly, M., Kim, Y.-H., King, R., Kirk, P., Kitching, I., Klautau, M., Klitgaard, B. B., Koenemann, S., Korovchinsky, N. M., Kotov, A., Kramina, T., Krapp-Schickel, T., Kremenetskaia, A., Krishna, K., Krishna, V., Kroh, A., Kroupa, A. S., Kury, A. B., Kury, M. S., Kvaček, J., Lachenaud, O., Lado, C., Lambert, G., Lana C. Atunes, L., Lazarus, D., Le Coze, F., Le Roux, M. M., LeCroy, S., Ledis Linares, J., Lee, S., Lewis, G. P., Li, S.-J., Li-Qiang, J., Lichtwardt, R. (†), Lim, S.-C., Lohrmann, V., Longhorn, S. J., Lorenz, W., Lowry, J., Lozano, F., Lumen, R., Lyal, C. H. C., Lörz, A.-N., Madin, L., Magnien, P., Mah, C., Mal, N., Mamos, T., Manconi, R., Mansano, V., Markello, K., Martin, J. H., Martin, P., Mashego, K. S., Maslin, B., Mattapha, S., McFadden, C., McKamey, S., McMurtry, J. A., Medrano, M. A., Mees, J., Mendes, A. C., Merrin, K., Mesa, N. C., Messing, C., Mielke, C. G. C., Migeon, A., Miller, D. R., Mills, C., Minelli, A., Mitchell, D., MolluscaBase eds., Molodtsova, T., Montenegro Valls, J. F., Mooi, R., Morandini, A., Moreira da Rocha, R., Morrow, C., Moteetee, A., Murillo-Ramos, L., Murphy, B., Narita, J. P. Z., Nemys eds., Nery, D. G., Neu-Becker, U., Neuhaus, B., Newton, A., Ng Kee Lin, P., Nicolson, D., Nielsen, J. E., Nijhof, A., Nishikawa, T., Norenburg, J., O’Hara, T., Ochoa, R., Ohashi, H., Ohashi, K., Ollerenshaw, J., Oosterbroek, P., Opresko, D., Osborne, R., Osigus, H.-J., Oswald, J. D., Ota, Y., Otte, D., Ouvrard, D., Pandey, A., Pape, T., Paulay, G., Paulson, D., Pauly, D., Pennington, R. T., Pereira, J. da S., Perez-Gelabert, D., Petrusek, A., Phillipson, P., Pinheiro, U., Pires Morim, M., Pisera, A., Pitkin, B., Plotkin, D., Poatskievick Pierezan, B., Poore, G., Povydysh, M., Praxedes, R. A., Pulawski, W. J., Pyle, R., Pühringer, F., Rajaei, H., Rakotonirina, N., Ramos, G., Rando, J., Ranzato Filardi, F., Raz, L., Read, G., Rees, T., Reich, M., Reimer, J. D., Rein, J. O., Reynolds, J., Rincón, J., Rius, M., Robertson, T., Robinson, G., Robinson, G. S. (†), Rodríguez, E., Ruggiero, M., Ríos, P., Rützler, K., Sanborn, A., Sanjappa, M., Santos, S. G., Santos-Guerra, A., Sartori, M., Sattler, K., Schierwater, B., Schilling, S., Schley, R., Schmid-Egger, C., Schmidt-Rhaesa, A., Schoolmeesters, P., Schorr, M., Schrire, B., Schuchert, P., Schuh, R. T., Schönberg, C., Schütz Rodrigues, R., Scoble, M., Segers, H., Seijo, G., Seleme, E. P., Senna, A., Serejo, C., Sforzi, A., Shenkar, N., Siegel, V., Sierwald, P., Sihvonen, P., Silva Flores, A., Silva de Carvalho, C., Simon, M. F., Simonsen, T., Simpson, C. E., Sinniger, F., Sirichamorn, Y., Skvarla, M., Smith, A. D., Smith, V. S., Soares Gissi, D., Sokoloff, D., Sotuyo, S., Soulier-Perkins, A., South, E. J., Souza-Filho, J. F., Spearman, L., Spelda, J., Steiner, A., Stemme, T., Sterrer, W., Stevenson, D., Stiewe, M. B. D., Stirton, C. H., Strand, M., Straub, S., Stueber, G., Stöhr, S., Subramaniam, S., Swalla, B., Swedo, J., Sánchez-Ruiz, M., Sørensen, M. V., Taiti, S., Takiya, D. M., Tandberg, A. H., Tavakilian, G., Taylor, K., Thessen, A., Thomas, J. D., Thomas, P., ThripsWiki, Thuesen, E., Thulin, M., Thurston, M., Thuy, B., Todaro, A., Torke, B. M., Turiault, M., Turner, J. R. G., Turner, T., Turon, X., Tyler, S., UCD Community, Uetz, P., Ulmer, J. M., Vacelet, J., Vachard, D., Vader, W., Valls Domedel, G., Van der Burgt, X., Vanhoorne, B., Vatanparast, M., Verhoeff, T., Vonk, R., Väinölä, R., Walker-Smith, G., Walter, T. C., Wambiji, N., Wanke, D., Watling, L., Weaver, H., Webb, J., Welbourn, W. C., Whipps, C., White, K., Wilding, N., Williams, G., Wilson, A. J. G., Wing, P., Winitsky, S., Wirth, C. C., Wojciechowski, M., Woodman, S., Xavier, J., Yi, T., Yoder, M., Yu, D. S. K., Yunakov, N., Zahniser, J., Zeidler, W., Zhang, R., Zhang, Z. Q., Zinetti, F., d’Hondt, J.-L., de Moraes, G. J., de Oliveira, A. B. R., de Voogd, N., del Río, M. G., van Haaren, T., van Nieukerken, E. J., van Ofwegen, L., van Soest, R., Şentürk, O., ITIS, ICTV, LPWG, PaleoBioDB & World Flora Online (Creators) (Eds.), Catalogue of Life (Version 2025-03-14). Catalogue of Life, Amsterdam. https://doi.org/10.48580/dg9ld-3gk
  86. Oksanen, J., Blanchet, F.G., Friendly, M., Kindt, R., Legendre, P., McGlinn, D., Minchin, P.R., O’Hara, R.B., Simpson, G.L., Solymos, P., Stevens, M.H.H., Szoecs, E. & Wagner, H. (2020) vegan: Community Ecology Package. R package Version 2.5-7. Available from: https://CRAN.R-project.org/package=vegan (accessed 24 December 2022)
  87. Pebesma, E.J. (2018) Simple Features for R: Standardized Support for Spatial Vector Data. The R Journal, 10, 439–446.
  88. Pebesma, E.J. & Bivand, R.S. (2005) Classes and methods for spatial data in R. R News, 5, 9–13.
  89. Peck, S.B. (1981) Distribution and biology of flightless carrion beetle Necrophilus pettitii in eastern North America. Entomological News, 92, 181–185.
  90. Peck, S.B. (2001a) 18. Agyrtidae C.G. Thomson, 1859. In: Arnett, R.H., Jr. & Thomas, M.C. (Eds.), American Beetles, Volume 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC Press, Boca Raton, pp. 247–249.
  91. Peck, S.B. (2001b) 21. Silphidae Latreille, 1807. In: Arnett, R.H., Jr. & Thomas, M.C. (Eds.), American Beetles, Volume 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC Press, Boca Raton, pp. 268–271.
  92. Peck, S.B. & Kaulbars, R.S. (1987) A synopsis of the distribution of the carrion beetles (Coleoptera: Silphidae) of the conterminous United States. Proceedings of the Entomological Society of Ontario, 188, 47–81.
  93. Pirone, D.J. (1974) Ecology of necrophilous and carpophilous Coleoptera in a southern New York woodland (phenology, aspection, trophic and habitat preferences). Ph.D. thesis, Fordham University, 769 p.
  94. Potticary, A.L., Belk, M.C., Creighton, J.C., Ito, M., Kilner, R., Komdeur, J., Royle, N.J., Rubenstein, D.R., Schrader, M., Shen, S.-F., Sikes, D.S., Smiseth, P.T., Smith, R., Steiger, S., Trumbo, S.T. & Moore, A.J. (2024) Revisiting the ecology and evolution of burying beetle behavior (Staphylinidae: Silphinae). Ecology and Evolution, 14, e70175. https://doi.org/10.1002/ece3.70175
  95. Pukowski, E. (1933) Oekologische Untersuchungen an Necrophorus F. Zeitschrift Fur Morphologie und Okologie der Tiere, 27, 518–586. https://doi.org/10.1007/bf00403155
  96. Purrington, F.F. & Davidson, R.L. (2000) New southerly distribution records for the boreal carrion beetle, Nicrophorus vespilloides (Coleoptera: Silphidae). Entomological news, 111 (5), 355–358.
  97. R Core Team (2021) R: A language and environment for statistical computing. In: R Foundation for Statistical Computing, Vienna. Available from: https://www.R-project.org/ (accessed 21 January 2025)
  98. Raithel, C. (1991) American burying beetle (Nicrophorus americanus) recovery plan. U.S. Fish & Wildlife Service, Newton Corner, 80 p.
  99. Ratcliffe, B.C. (1972) The natural history of Necrodes surinamensis (Fabr.) (Coleoptera: Silphidae). Transactions of the American Entomological Society, 98, 359–410. https://www.jstor.org/stable/25078118
  100. Ratcliffe, B.C. (1996) The Carrion Beetles (Coleoptera: Silphidae) of Nebraska. Bulletin of the University of Nebraska State Museum, Volume 13, 100 pp.
  101. Reed, H.B., Jr. (1958) A study of dog carcass communities in Tennessee, with special reference to the insects. American Midland Naturalist, 59 (1), 213–245. https://doi.org/10.2307/2422385
  102. Rigsby, S.N., Cavallaro, M.C., Fuhlendorf, S.D. & Hoback, W.W. (2025) Use of charred vertebrate carcasses by carrion beetles: Implications of controlled burns and wildfires for the American burying beetle, Nicrophorus americanus. Insect Conservation and Diversity, 2025, 1–11. https://doi.org/10.1111/icad.12844
  103. Ruzicka, J. (1994) Seasonal activity and habitat associations of Silphidae and Leiodidae: Cholevinae (Coleoptera) in central Bohemia. Acta Societatis Zoologicae Bohemicae, 58, 67–78.
  104. Schawaller, W. (1981) Taxonomie und Faunistik der Gattung Thanatophilus (Coleoptera: Silphidae). Stuttgarter Beitrage zur Naturkunde, Series A (Biologie), 351, 1–21.
  105. Scott, M.P. (1994a) Competition with flies promotes communal breeding in the burying beetle Nicrophloruts tomentosis. Behavioral Ecology and Sociobiology, 34, 367–373. https://doi.org/10.1007/s002650050052 https://www.jstor.org/stable/4600954
  106. Scott, M.P. (1994b) The benefit of paternal assistance in intra-and interspecific competition for the burying beetle, Nicrophorus defodiens. Ethology ecology & evolution, 6 (4), 537–543. https://doi.org/10.1080/08927014.1994.9522978
  107. Scott, M.P. (1998) The ecology and behavior of burying beetles. Annual review of entomology, 43 (1), 595–618. https://doi.org/10.1146/annurev.ento.43.1.595
  108. Scott, M.P. & Traniello, J.F. (1990) Behavioural and ecological correlates of male and female parental care and reproductive success in burying beetles (Nicrophorus spp.). Animal Behaviour, 39 (2), 274–283. https://doi.org/10.1016/S0003-3472(05)80871-1
  109. Scott, M.P., Traniello, J.F.A. & Fetherston, I.A. (1987) Competition for prey between ants and burying beetles (Nicrophorus spp.): differences between northern and southern temperature sites. Psyche, 94, 325–332. https://doi.org/10.1155/1987/56594
  110. Schrader, M. & Galanek, J. (2022) Stridulation is unimportant for effective parental care in two species of burying beetle. Ecological Entomology, 47 (1), 18–24. https://doi.org/10.1111/een.13086
  111. Shubeck, P.P. (1970) Silphidae attraction to carrion-baited air cans versus carrion-baited ground cans. The Coleopterists Bulletin, 24, 66–70. https://doi.org/10.5962/p.372627 https://www.jstor.org/stable/3999393
  112. Shubeck, P.P. (1971) Diel periodicities of certain carrion beetles. The Coleopterists Bulletin, 25, 41–46. https://doi.org/10.5962/p.372563
  113. Shubeck, P.P. (1976) Carrion beetle responses to poikilotherm and homoiotherm carrion. Entomological News, 89, 265–269.
  114. Shubeck, P.P. (1983) Habitat preferences of carrion beetles in the Great Swamp National Wildlife Refuge, New Jersey (Coleoptera: Silphidae, Dermestidae, Nitidulidae, Histeridae, Scarabaeidae). Journal of the New York Entomological Society, 91, 333–341. https://www.jstor.org/stable/25009379
  115. Sikes, D.S., Madge, R.B. & Newton, A.F. (2002) A catalog of the Nicrophorinae (Coleoptera: Silphidae) of the world. Zootaxa, 65, 1–304. https://doi.org/10.11646/zootaxa.65.1.1
  116. Sikes, D.S., Thayer, M.K. & Newton, A.F. (2024) Large carrion and burying beetles evolved from Staphylinidae (Coleoptera, Staphylinidae, Silphinae): a review of the evidence. ZooKeys, 1200, 159–182. https://doi.org/10.3897/zookeys.1200.122835
  117. Sikes, D.S. & Rathiel, C.J. (2002) A review of hypotheses of decline of the endangered American burying beetle (Silphidae: Nicrophorus americanus Olivier). Journal of Insect Conservation, 6, 103–113. https://doi.org/10.1023/A:1020947610028
  118. Sikes, D.S., Trumbo, S.T. & Peck, S.B. (2016) Cryptic diversity in the New World burying beetle fauna: Nicrophorus hebes Kirby; new status as a resurrected name (Coleoptera: Silphidae: Nicrophorinae). Arthropod Systematics and Phylogeny, 74 (3), 299–309. https://doi.org/10.3897/asp.74.e31872
  119. Sikes, D.S. & Venables, C. (2013) Molecular phylogeny of the burying beetles (Coleoptera: Silphidae: Nicrophorinae). Molecular Phylogenetics and Evolution, 69 (3), 552–565. https://doi.org/10.1016/j.ympev.2013.07.022
  120. Smith, R.J., Bonilla, M., Calahan, C. & Mann, J. (2000) Comparison of reproductive success of in-situ burial versus the use of abandoned burrows for carcass interment by Nicrophorus investigator (Coleoptera: Silphidae). Journal of the Kansas Entomological Society, 73, 148–154.
  121. Smith, G., Trumbo, S.T., Sikes, D.S., Scott, M.P. & Smith, R.L. (2007) Host shift by the burying beetle, Nicrophorus pustulatus, a parasitoid of snake eggs. Journal of Evolutionary Biology, 20, 2389–2399. https://doi.org/10.1111/j.1420-9101.2007.01404.x
  122. Smith, R.J. & Heese, B. (1995) Carcass selection in a high altitude population of the burying beetle, Nicrophorus investigator (Silphidae). The Southwestern Naturalist, 40, 50–55. https://www.jstor.org/stable/30054393
  123. Smith, R.J. & Merrick, M.J. (2001) Resource availability and population dynamics of Nicrophorus investigator, an obligate carrion breeder. Ecological Entomology, 26, 173–180. https://doi.org/10.1046/j.1365-2311.2001.00307.x
  124. Steele, B.F. (1927) Notes on the feeding habits of carrion beetles. Journal of the New York Entomological Society, 35, 77–81. https://www.jstor.org/stable/25004181
  125. Trumbo, S.T. (1990a) Interference competition among burying beetles (Silphidae, Nicrophorus). Ecological Entomology, 15, 347–355. https://doi.org/10.1111/j.1365-2311.1990.tb00816.x
  126. Trumbo, S.T. (1990b) Reproductive success, phenology and biogeography of burying beetles (Silphidae: Nicrophorus). American Midland Naturalist, 124, 1–11. https://doi.org/10.2307/2426074
  127. Trumbo, S.T. (1991) Reproductive benefits and the duration of parental care in a biparental burying beetle, Necrophorus orbicollis. Behavior, 117 (1-2), 82–105. https://doi.org/10.1163/156853991X00139
  128. Trumbo, S.T. (1992) Monogamy to communal breeding: exploitation of a broad resource base by burying beetles (Nicrophorus). Ecological Entomology, 17, 289–298. https://doi.org/10.1111/j.1365-2311.1992.tb01060.x
  129. Trumbo, S.T. (1994) Interspecific competition, brood parasitism, and the evolution of biparental cooperation in burying beetles. Oikos, 69, 241–249. https://doi.org/10.2307/3546144
  130. Trumbo, S.T. & Bloch, P.L. (2000) Habitat fragmentation and burying beetle abundance and success. Journal of Insect Conservation, 4, 245–252. https://doi.org/10.1023/A:101
  131. Ulyshen, M., Hanula, J.L. & Horn, S. (2007) Burying beetles (Coleoptera: Silphidae) in the forest canopy: The unusual case of Nicrophorus pustulatus Herschel. The Coleopterists Bulletin, 61, 121–123. https://doi.org/10.1649/924.1
  132. [USFWS] United States Fish & Wildlife Service (1989) Endangered and threatened wildlife and plants; determination of endangered status for the American burying beetle. Federal Register, 54, 29652–29655.
  133. [USFWS] United States Fish & Wildlife Service (2020) Endangered and threatened wildlife and plants; reclassification of the American burying beetle from endangered to threatened with a section 4(d) rule. Federal Register, 85, 65241–65261.
  134. Walker, T.L. & Hoback, W.W. (2007) Effects of invasive eastern red cedar on capture rates of Nicrophorus americanus and other Silphidae. Environmental Entomology, 36 (2), 297–307. https://doi.org/10.1093/ee/36.2.297
  135. Welch, J.F., Windmiller, B. & Berkholtz, J. (2023) Observation of the uncommon burying beetle Nicrophorus hebes (Coleoptera: Staphylinidae: Silphinae), feeding on a dead Blanding’s turtle (Emydoidea blandingii) in Massachusetts. Northeastern Naturalist, 30 (3), N44–N47. https://doi.org/10.1656/045.030.0309
  136. Wettlaufer, J.D. (2019) Partitioning resources through the seasons: A test of the competitive ability – cold tolerance trade-off hypothesis in seasonally breeding beetles. Master’s thesis, Queen’s University, Kingston, Ontario, Canada, 98 p.
  137. Wickham, H. (2016) ggplot2: Elegant Graphics for Data Analysis. Springer-Verlag, New York. [program]
  138. Wilhelm, S.I., Larson, D.J. & Storey, A.E. (2001) Habitat preference of two burying beetles (Coleoptera: Silphidae: Nicrophorus) living among seabirds. Northeastern Naturalist, 8 (4), 435–442. https://doi.org/10.2307/3858447
  139. Wilson, D.S. & Fudge, J. (1984) Burying beetles: intraspecific interactions and reproductive success in the field. Ecological Entomology, 9, 195–203. https://doi.org/10.1111/j.1365-2311.1984.tb00715.x
  140. Wilson, D.S. & Knollenberg, W.G. (1987) Adaptive indirect effects: the fitness of burying beetles with and without their phoretic mites. Evolutionary Ecology, 1987 (1), 139–159. https://doi.org/10.1007/BF02067397
  141. Wilson, D.S., Knollenberg, W.G. & Fudge, J. (1984) Species packing and temperature dependent competition among burying beetles (Silphidae: Nicrophorus). Ecological Entomology, 9, 205–216. https://doi.org/10.1111/j.1365-2311.1984.tb00716.x
  142. Zachariassen, K.E. (1980) The role of polyols and nucleating agents in cold-hardy beetles. Journal of Comparative Physiology, 140, 227–234. https://doi.org/10.1007/BF00690407

How to Cite

Chapman, E.G., Clarkson, M.D., Wright, C.E. & Dupuis, J.R. (2026) The Silphinae (Staphylinidae) and Agyrtidae of Kentucky with an illustrated key to the fauna of eastern North America. Zootaxa, 5839 (1), 71–109. https://doi.org/10.11646/zootaxa.5839.1.3