Catalogue of named fossil Scarabaeoidea (Lamellicornia)
Catalogue of fossil Scarabaeoidea (Coleoptera: Polyphaga) of the Mesozoic and Tertiary Frank-Thorsten Krell
Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. email: f.krell@nhm.ac.uk
Catalogue of named fossil Scarabaeoidea (Lamellicornia)
The present catalogue is based on the printed version published in 2000 in Invertebrate Taxonomy (Krell, 2000a) but it is updated up to 2003 and includes also some older references that were overlooked. However, it does not include some taxonomic and nomenclatural remarks included in the printed version. All named fossil taxa of Scarabaeoidea are listed according to their current classification. The classification of Scarabaeoidea follows the proposed system of Browne & Scholtz (1998, 1999) and Lawrence & Newton (1995). Species of doubtful identity are in brackets; such instances are only indicated if the fossils definitively lack any diagnostic character of Scarabaeoidea. These specimens are mostly single elytra. A genus name in brackets indicates that the placement of the species in the genus is doubtful. Abbreviations: L: Lower, M: Middle, U: Upper.
The following species were not included in the printed catalogue:
Cretocoma tologoica Nikolaev, 2002
Paralucanus mesozoicus Nikolajev, 2000
Prototrox transbaikalicus Nikolaev, 2000
Onthophagus rugulosus (Heer, 1870), listed by Ponomarenko & Kirejtshuk
(, accessed 30.March 2004) has been described as Elytridium rugulosum and originally classified as a possible carabid (Heer 1870: 78). It is the middle part of one elytron which lacking any character which might reveal its family.
The genus names „Mimaphodius Nikolajev, 1993?, „Protogeotrupes Nikolajev? and „Cretoglaphyrus Nikolajev?
listed by Ponomarenko & Kirejtshuk (l.c.) are not yet described and, therefore, nomina nuda.
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The author would like to update this catalogue regularly. To have your work included, please provide the author with reprints or pdf files of your papers containing new fossil taxa of Scarabaeoidea or mentioning known taxa or only the presence of Scarabaeoidea in a lagerstätte. Thank you for your cooperation.
(Family Glaresidae)
(Genus Aphodiites Heer)
Aphodiites Heer, 1865: 90, pl. 8 (type species by monotypy: Aphodiites protogaeus Heer). – Scudder 1886: 73;
Crowson 1981: 664 (“Possible Scarabaeoids”), 679 (“probably attributable to this superfamily”);
Dellacasa 1988: 361; Paulian 1988: 390; Cambefort 1991: 51 (“undifferenciated Laparosticti”);
Carpenter 1992: 318 (“Little-known genus”, suborder and family uncertain); Scholtz et al. 1994: 275
(“The diagnostic characters [...] could equally apply to a Glaresis-like beetle”); Scholtz & Chown 1995:
358 (“glaresid-like”); Nikolaev 1998b: 30; Browne & Scholtz 1999: 54 (“Glaresidae-like”); Krell 2000a:
874.
(Aphodiites protogaeus Heer)
Aphodiites protogaeus Heer, 1865: 90, pl. 8; (Jurassic, L Lias, Schambelen, Aargau, Switzerland). – Heer 1883:
100, pl. 8; Scudder 1881-1885: 794; Scudder 1891: 190; Handlirsch 1906: 441, pl. 41 (“Es ist auch hier
keinerlei Anhaltspunkt vorhanden, um die Familie mit einiger Sicherheit erkennen zu können. So gut
wie um eine Scarabaeide kann es sich auch um eine Chysomelide, Tenebrionide usw. handeln.”);
Théodoridès 1952: 32 (“très sceptiques sur la position systématique exacte de ce Coléoptère”);
Dellacasa 1983: 28f (“la sua attribuzione agli Scarabaeoidea è estremamente dubitativa”); Dellacasa
1988: 361; Scholtz 1990: 1029; Carpenter 1992: 318; Krell 2000a: 874.
Aphodiites prologaeus. – Balthasar 1963: 78; Iablokoff-Khnzorian 1977: 137 (family identity doubtful); Morón
Ríos 1984: 117; Morón 2003: 12.
Doubful (no appendages present).
Familia Trogidae
Trox antiquus Wickham
Trox antiquus Wickham, 1909: 129 (Oligocene, Florissant, U.S.A.). – Wickham 1920: 358; Krell 2000a: 874.
(Trox oustaleti Scudder)
Trox oustaleti Scudder, 1879a: 178B (Eocene, Nine-mile Creek, British Columbia, Canada). – Scudder 1890:
487, pl. 2; Scudder 1895a: 35; Scudder 1900: 105; Handlirsch 1907: 839; Handlirsch 1910: 99 (“Of this
order of insects, which, as far as the trustworthiness of the identification is concerned, is distinctly a
discredit to paleoentomology, a series of forms from British Columbia has been brought to light, the
interpretation of which lacks adequate support.”); Wickham 1920: 358; Krell 2000a: 875.
Doubful (only one elytron).
Subfamilia Prototroginae Nikolaev, 2000: 63 (transl. p. 426)
Genus Prototrox Nikolaev
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Prototrox Nikolaev, 2000a: 65 (transl. p. 427) (type species by original designation: Prototrox transbaikalicus
Nikolaev). - Nikolajev, 1999: 178 (nomen nudum); Krell 2000a: 890 (nomen nudum); Nikolaev 2002:
54 (transl. p. 281).
Prototrox transbaikalicus Nikolaev
Prototrox transbaikalicus Nikolaev 2000a: 65 (transl. p. 428) (Lower Cretaceous, Argun? formation, Semen
Creek, Chita District, Transbaikalia, Russia).
Familia Passalidae
Passalus (Passalus) indormitus Cockerell
Passalus (?) indormitus Cockerell, 1927a: 65 (Oligocene, John Day Series, Oregon, U.S.A.). Passalus indormitus. – Reyes-Castillo 1970: 73; Morón 2003: 13.
Passalus (Passalus) indormitus. – Reyes-Castillo 1977: 652; Krell 2000a: 875.
Genus Serrulus Hong
Serrulus Hong, 1983: 6 (type species by original designation: Serrulus sinicus Hong).
Serrulus sinicus Hong
Serrulus sinicus Hong, 1983: 6, 13 (Miocene, Shanwang, China). – Zhang 1989: 149, pl. 40 (Passalidae); Krell
2000a: 875.
Familia Lucanidae
Subfamilia Lucaninae
(Lucanus fossilis Wickham)
Lucanus fossilis Wickham, 1913b: 293 (Oligocene, Florissant, U.S.A.). – Wickham 1920: 358; Maes 1992: 17;
Krell 2000a: 875.
Doubtful (only one elytron).
(Platycerus) sepultus Germar
Lucanus. – Goldfuss 1831: 118; Keferstein 1834: 329; Scudder 1886: 74. Platycerus sepultus Germar, 1837a: 7, pl. 7 (Oligocene, “in carbone fossili territorii Rheni prope Bonnam”,
Germany). – Germar 1837b: 423: (“1 Lucanus”, Platycerus sepultus); Malepeyre 1838: 267; Bronn
1848b: 992; Bronn 1849: 623; Giebel 1852b: 653; Giebel 1856: 29; Goss 1878: 331; Scudder 1891: 567;
Handlirsch 1907: 842; Houlbert 1915: 6; Nikolaev 1990: 121 (transl. p. 119) (generic and even family
affiliation doubtful; may belong near Ceruchus); Maes 1992: 14; Krell 2000a: 875. Systenocerus sepultus. – Statz 1952: 1 (generic affiliation doubtful).
Platycerus zherichini Nikolajev
Platycerus zherichini Nikolajev, 1990: 122 (1991: 121) (Oligocene, Pozhar region, Russia). – Krell 2000a: 875.
Genus Succiniplatycerus Nikolajev
Succiniplatycerus Nikolajev, 1990: 122 (1991: 120) (type species by original designation: Platycerus berendti
Zang, 1905). – Jarzembowski 2000: (3); Krell 2000a: 875.
According to Larsson (1978: 152) Zang “describes a total of 5 species of Systenocerus” from Baltic Amber. This is wrong, since Zang described only the following species and mentioned one other stag beetle species
(Dorcasoides bilobus Motsch.).
Succiniplatycerus berendti (Zang) Nikolaev
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Platycerus berendti Zang, 1905: 199 (Eocene, Baltic Amber). – Bibliography: Spahr 1981: 65, additional: Hieke
& Pietrzeniuk 1984: 313.
Platycerus berendtii. – Maes 1992: 12.
Succiniplatycerus berendti. – Nikolaev 1990: 122 (1991: 120); Krell 2000a: 875.
Dorcus (Eurytrachelus) primigenius Deichmüller
Dorcus (Eurytrachelus) primigenius Deichmüller, 1881: 303, pl. 21 (Eocene, Ku?lín [Kutschlin] near Bílina
[Bilin], Czechia). – Leuthner 1885: 482; Scudder 1891: 514; Po?ta 1900: 265; Handlirsch 1907: 842;
Houlbert 1915: 6; ?íha 1979: 26; Maes 1992: 92; Krell 2000a: 875.
Genus Miocenidorcus Riou
Miocenidorcus Riou 1999: 126 (type species by original designation: Miocenidorcus andancensis Riou). – Riou,
1988: 98 (unpublished); Krell 2000a: 875.
Miocenidorcus andancensis Riou
Miocenidorcus andancensis Riou, 1999: 126, 133 (Miocene, Andance, France). – Riou, 1988: 99 (unpublished);
Krell 2000a: 875.
Subfamilia Paralucaninae Nikolajev, 2000b
Genus Paralucanus Nikolajev
Paralucanus Nikolajev, 2000b: S329 (type species by original designation: Paralucanus mesozoicus Nikolajev).
Paralucanus mesozoicus Nikolajev
Paralucanus mesozoicus Nikolajev, 2000b: S330 (Upper Jurassic, Shara-Tag, Mongolia).
Subfamilia Syndesinae
Ceruchus fuchsii Wickham
Ceruchus fuchsii Wickham, 1911: 58 (Oligocene, Florissant, U.S.A.). – Wickham 1920: 358; Rodeck 1938: 293;
Maes 1992: 11; Krell 2000a: 875.
Subfamilia Chiasognathinae
Genus Protognathinus Chalumeau & Brochier
Protognathinus Chalumeau & Brochier, 2001: 595 (type species by monotypy: Protognathinus spielbergi
Chalumeai & Brochier
Protognathinus spielbergi Chalumeau & Brochier
Protognathinus spielbergi Chalumeau & Brochier, 2001: 595 (Eocene, Grube Messel, Germany).
Subfamilia Aesalinae
Genus Cretaesalus Nikolajev
Cretaesalus Nikolajev, 1993: 90 (type species by original designation: Gretaesalus ponomarenkoi Nikolajev
[ lapsus calami for Cretaesalus]). – Nikolaev 1999: 178; Krell 2000a: 875; Nikolaev 2002: 54 (transl. p.
281).
Cretaesalus ponomarenkoi Nikolajev
Cretaesalus ponomarenkoi Nikolajev, 1993: 90 (U Cretaceous, Kzyl-Zhar, Kazakhstan). – Krell 2000a: 875.
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Subfamilia incerta
Genus Dorcasoides Motschulsky
Dorcasoides Motschulsky, 1856: 27, 30 (type species by monotypy: Dorcasoides bilobus Motschulsky). –
Scudder 1881-1885: 795; Scudder 1886: 74; Handlirsch 1925: 245; Maes 1992: 95; Carpenter 1992:
300; Poinar 1992: 147; Krell 2000a: 875; Schweigert, 2003..
Dorcasoides bilobus Motschulsky
Dorcasoides bilobus Motschulsky, 1856: 27 (fossil, without further data; Motschoulsky 1868 listed the species
among “Insectes contenus dans le succin” which was interpreted as from Baltic Amber [Eocene] by
following authors). – Motschoulsky 1868: 103; Scudder 1881-1885: 794 (fig. 1028); Scudder 1891: 513;
Zittel 1895: 505; Zang 1905: 199; Handlirsch 1907: 842; Broili 1921: 675; Broili 1924: 696; Carpenter
1992: 300; Maes 1992: 95; Krell 2000a: 876.
Dorcasoides nigrescens. – Motschulsky 1856: pl. (lapsus calami? – see Scudder 1891: 513).
Dorcasoides bibolus. – Houlbert 1915: 6.
Genus Ceruchites Statz
Ceruchites Statz, 1952: 5 (type species by monotypy: Ceruchites hahnei Statz). – Carpenter 1992: 300; Krell
2000a: 876.
Ceruchites hahnei Statz
Ceruchites hahnei Statz, 1952: 5 (Oligocene, Rott, Germany). – Sphon 1973: 51; Carpenter 1992: 300; Krell
2000a: 876.
Genus Paleognathus Waga
Paleognathus Waga, 1883: 191 (type species by monotypy: Paleognathus succini Waga). – Bibliography: Spahr
1981: 65, additional: Carpenter 1992: 300f; Poinar 1992: 147; Krell 2000a: 876. Palaeognathus. – Bibliography: Spahr 1981: 65, additional: Deville 1991: 16; Maes 1992: 68; Scholtz & Chown
1995: 364; Schweigert 2003: 179.
Paleognathus succini Waga
Paleognathus succini Waga, 1883: 191 (Eocene, Baltic Amber). – Bibliography: Spahr 1981: 65, additional:
Paulian 1988: 394; Carpenter 1992: 300; Krell 2000a: 876; Chalumeai & Brochier 2001: 593
(“probablement […] Lampriminae”).
Palaeognathus succini. – Bibliography: Spahr 1981: 65, additional: Kozur 1984: 594f; Müller 1989: 245f; Maes
1992: 68.
Palaeognathus succinifer. – Leuthner 1885: 482.
Paläognathus succini. – Handlirsch 1908: 1356.
Paleognatius succini. – Laurentiaux 1953: 477.
Familia Bolboceratidae
Bolboceras inermis Piton
Bolboceras inermis Piton, 1940: 175 (Eocene, Menat, France). – Krell 2000a: 876.
Genus Cretobolbus Nikolajev
Cretobolbus Nikolajev, 1996: 95 (transl. p. 221) (type species by original designation: Cretobolbus rohdendorfi
Nikolajev, 1996). – Nikolaev 1999: 178; Krell 2000a: 876; Nikolaev 2002: 54 (transl. p. 281).
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Cretobolbus rohdendorfi Nikolajev
Cretobolbus rohdendorfi Nikolajev, 1996: 96 (transl. p. 222) (L Cretaceous, Baysa, Russia). – Krell 2000a: 876.
Familia Pleocomidae
Genus Proteroscarabaeus Grabau
Proteroscarabaeus Grabau, 1923: 173 (type species by original designation: Proteroscarabaeus yeni Grabau). –
Ping 1928: 18; Crowson 1974: 68 (“suggestive of Hybosoridae. If it is truly Scarabaeid, then it has
possible affinities to [...] Aclopinae”); Nikritin 1977: 124 (1991: 168); Lin 1980: 230; Crowson 1981:
667 (“modern-looking Scarabeoids”); Ponomarenko 1990: 66; Carpenter 1992: 301; Nikolaev 1996: 91
(transl. p. 217), 1999: 178; Krell 2000a: 876 (subfamily affiliation according to Nikolaev 1996);
Nikolaev 2002: 54 (transl. p. 281).
Protoscarabaeus: Scholtz 1990: 1029; Scholtz & Chown 1995: 358.
(Proteroscarabaeus magnus Nikolajev)
Proteroscarabaeus magnus Nikolajev, 1996: 91 (transl. p. 217) (L Cretaceous, Baysa, Russia). – Krell 2000a:
876.
Doubtful (only one elytron).
(Proteroscarabaeus nikritini Nikolajev)
Proteroscarabaeus nikritini Nikolajev, 1996: 93 (transl. p. 217) (L Cretaceous, Baysa, Russia). – Krell 2000a:
876.
Doubtful (only one elytron).
Proteroscarabaeus robustus Zhang
Proteroscarabaeus dalaziensis (nomen nudum). – Lin 1994: 309 (according to Lin, in litt. 2000).
Proteroscarabaeus robustus Zhang, 1997: 90f, 103 (L Cretaceous, Zhixin Basin, China). – Krell 2000a: 876.
Proteroscarabaeus yeni Grabau
Proteroscarabaeus yeni Grabau, 1923: 175 (L Cretaceous, Laiyang, China). – Ping 1928: 19; Handlirsch 1939:
167; Balthasar 1963: 78; Ponomarenko 1977a: 6 (1991a: 2); Nikritin 1977: 126 (1991: 171) (L
Cretaceous, Baysa, and Chita region, Pad Semen site); Iablokoff-Khnzorian 1977: 137 (oldest reliable
Lamellicornia fossil; belonging to an extinct family-group taxon); Morón Ríos 1984: 117; Hong &
Wang 1990: 114f; Carpenter 1992: 301; Zhang 1992b: 134; Nikolaev 1992: 79 (transl. p. 99); Lin 1994:
305; Nikolaev 1996: 94 (transl. p. 219); Krell 2000a: 876; Morón 2003: 12.
Tribus Cretocomini Nikolaev, 2002: 53 (transl. p. 281)
Genus Cretocoma Nikolaev
Cretocoma Nikolaev, 2002: 54 (transl. p. 281) (type species by original designation: Cretocoma tologoica).
Cretocoma tologoica Nikolaev
Cretocoma tologoica Nikolaev, 2002: 54 (transl. p. 281) (L Cretaceous, Shar Tolgoy, Bayan-Hongor Aymag,
Mongolia)
Familia Glaphyridae
Glaphyrus antiquus Heer
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Glaphyrus antiquus Heer, 1862: 79, pl. 6 (Miocene, Öhningen “Insektenschicht des untern Bruches”, Germany).
– Heer 1865: 380; Heer 1883: 406; Handlirsch 1907: 839; Krell 2000a: 876.
Glaphyrus. – Kolbe 1932: 150.
Amphicoma defuncta Wickham
Amphicoma defuncta Wickham, 1910: 49 (Oligocene, Florissant, U.S.A.). – Wickham 1920: 358; White 1995: 3;
Krell 2000a: 876.
Familia Geotrupidae
Subfamilia Geotrupinae
Geotrupes atavus Oustalet
Geotrupes atavus Oustalet, 1874: 199, pl. 3 (Oligocene, Aix en Provence, France). – Goss 1878: 339; Scudder
1891: 524; Handlirsch 1907: 839; Meunier 1921: 4; Théobald 1937: tabl. 11; Théodoridès 1952: 46;
Krell 2000a: 877.
Gymnopleurus atavus. – Meunier 1921: 12 (“gehört wahrscheinlich auch zur Gattung Gymnopleurus”).
Geotrupes germari Heer
Geotrupes germari Heer, 1862: 71, pl. 6 (Miocene, Öhningen, Kesselstein, Germany). – Heer 1865: 379;
Oustalet 1874: 200; Heer 1883: 405; Scudder 1891: 524; Handlirsch 1907: 839; Meunier 1921: 4; Krell
2000a: 877.
Geotrupes jiaoyanshanensis (Hong) Zhang
Obitiscarabaeus jiaoyanshanense Hong, 1983: 6, 13 (Miocene, Shanwang, China). – Hong 1985: 40, pl. 21;
Zhang 1989: 172.
Geotrupes jiaoyanshanensis. – Zhang 1989: 152, 172; Krell 2000a: 877.
(Geotrupes messelensis Meunier)
Geotrupes messelensis Meunier, 1921: 11, pl. 2 (Eocene, Messel, Germany). – Théodoridès 1952: 46;
Koenigswald 1987: 140; Krell 2000a: 877.
Doubtful (poor preservation).
Geotrupes rottensis Statz
Geotrupes rottensis Statz, 1952: 7 (Oligocene, Rott, Germany). – Sphon 1973: 52; Krell 2000a: 877.
Geotrupes vetustus Germar
Geotrupes vetustus Germar, 1837a: 6, pl. 6 (Oligocene, “e carbone fossili territorii Rheni prope Bonnam”,
Germany). – Malepeyre 1838: 267; Brullé 1839: 20; Germar 1849: 57; Heer 1847: 62; Bronn 1849: 624
(Dynastidae); Giebel 1852b: 653; Giebel 1856: 35; Heer 1862: 72; Goss 1878: 331; Scudder 1891: 525;
Handlirsch 1907: 839; Meunier 1921: 4; Statz 1952: 2; Krell 2000a: 877.
(Genus Geotrupoides Handlirsch)
Geotrupoides Handlirsch, 1906: 545 (type species by monotypy: Geotrupoides lithographicus Deichmüller). –
Nikritin 1977: 119 (1991: 162); Crowson 1981: 667 (“modern-looking Scarabeoids”), 679 (“could well
be an early representative of the Geotrupidae [...] line”); Schlüter 1987: 133; Hong & Wang 1990: 115;
Scholtz 1990: 1029; Carpenter 1992: 300; Scholtz & Chown 1995: 358; Nikolaev 1998b: 30; Krell
2000a: 877.
(Geotrupoides) fortus Ren, Zhu & Lu
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Geotrupoides fortus Ren et al., 1995: 436 (L Cretaceous, Chifeng City, Inner Mongolia, China). – Krell 2000a:
877.
Ren et al. classified this species as being close to G. sulcatus Nikritin which is currently in Holcorobeus. Hence,
the generic affiliation of the G. fortus is doubtful.
Geotrupoides jiaoheensis Hong
Geotrupoides jiaoheense Hong, 1992: 415, pl. 161 (L Cretaceous, Jilin Province, China). – Krell 2000a: 877.
Generic affiliation needs revision since genus is dubious.
(Geotrupoides lithographicus (Deichmüller) Handlirsch)
Geotrupes lithographicus Deichmüller, 1886: 69 (U Jurassic, Eichstädt [Lithographischer Schiefer], Germany). –
Scudder 1891: 206; Iablokoff-Khnzorian 1977: 137 (family identity doubtful); Ponomarenko 1980: 113,
pl. 16 (photo of holotype).
Geotrupoides lithographicus. – Handlirsch, 1906: 545, pl. 45 (“kann man doch nicht sicher sagen, dass die Form
zu den Scarabaeiden gehört, solange weder Fühler noch Beine bekannt sind”); Théodoridès 1952: 32;
Nikritin 1977: 119 (1991: 162); Ponomarenko 1980: 119; Morón Ríos 1984: 117; Hong & Wang 1990:
115f (G. lingraphicus, lapsus calami); Hong 1992: 415f; Krell 2000a: 877; Morón 2003: 12. Geotrupides lithographicus. – Balthasar 1963: 78 (“die Einreihung zu den Scarabaeoidea [ist] nur eine gewagte
Annahme”).
After having studied the holotype Ponomarenko (1980) claimed that this fossil belongs to “Scarabaeidae” (= Scarabaeoidea), but further classification is not possible. However, this interpretation remains questionable since
only traces of the coxae and neither tibiae nor antennae are present.
Geotrupoides nodosus Hong & Wang
Geotrupoides nodosus Hong & Wang, 1990: 116, 180, pl. 21 (L Cretaceous, Laiyang, China). – Krell 2000a: 877.
Generic affiliation needs revision since genus is doubtful.
Geotrupoides saxosus Zhang
Geotrupoides saxosus Zhang, 1997: 90, 103 (U Cretaceous, Zhixin Basin, China). – Lin 1994: 309 (nomen
nudum); Krell 2000a: 877.
Generic affiliation needs revision since genus is doubtful.
Geotrupoides songyingziensis Hong
Geotrupoides songyingziense Hong, 1984: 170, pl. 71 (L Cretaceous, Yixian, China). – Krell 2000a: 877.
Generic affiliation needs revision since genus is doubtful.
Genus Orrhodomala Zhang
Orrhodomala Zhang, 1989: 154, 425 (type species by original designation: Orrhodomala protista Zhang). –
Krell 2000a: 877.
Orrhodomala protista Zhang
Orrhodomala protista Zhang, 1989: 154, 425 (Miocene, Shanwang, China). – Krell 2000a: 877.
Subfamilia Cretogeotrupinae Nikolajev, 1996: 97 (transl. p. 222)
Genus Cretogeotrupes Nikolajev
Cretogeotrupes Nikolajev, 1992: 80 (transl. p. 101), type genus (type species by original designation:
Cretogeotrupes convexus Nikolajev). – Nikolaev 1999: 178; Krell 2000a: 878; Nikolaev 2002: 54
(transl. p. 281).
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Cretogeotrupes convexus Nikolajev
Cretogeotrupes convexus Nikolajev, 1992: 81 (transl. p. 102) (L Cretaceous, Baysa, Russia). – Krell 2000a: 878.
Familia Hybosoridae
Genus Coprologus Heer
Coprologus Heer, 1847: 60 (type species by monotypy: Coprologus gracilis Heer). – Stitzenberger 1851: 100;
Rogg 1852: 22; Giebel 1856: 35; Scudder 1881-1885: 795; Scudder 1886: 74; Handlirsch 1907: 839;
Handlirsch 1925: 246; Balthasar 1963: 79; Carpenter 1992: 330; Krell 2000a: 878.
Caprologus. – Giebel 1852b: 653.
Coprologus gracilis Heer
Coprologus gracilis Heer, 1847: 60, pl. 2 (Miocene, Öhningen, Germany). – Bronn 1849: 624; Giebel 1856: 35;
Heer 1865: 379; Heer 1883: 405; Nikolaev 1996: 96 (transl. p. 222) (doubts the former classification as
belonging to the Bolboceratidae, “presumable belongs to the Hybosoridae”); Krell 2000a: 878.
Caprologus gracilis. – Giebel 1852b: 653.
Genus Procoilodes Ocampo
Procoilodes Ocampo, 2002: 123 (type species by original designation: Procoilodes adrastus Ocampo).
Procoilodes adrastus Ocampo
Procoilodes adrastus Ocampo, 2002: 125 (Miocene, Dominican Amber).
Genus Tyrannasorus Ratcliffe & Ocampo
Tyrannasorus Ratcliffe & Ocampo, 2001: 351 (type species by original designation: Tyrannasorus rex Ratcliffe
& Ocampo).
Tyrannasorus rex Ratcliffe & Ocampo
.Tyrannasorus rex Ratcliffe & Ocampo, 2001: 353 (Miocene, Dominican Amber). – Ocampo 2002: 123.
Subfamilia Anaidinae Nikolajev
(Genus Cretanaides Nikolajev)
Cretanaides Nikolajev, 1996: 94 (transl. p. 219) (type species by original designation: Cretanaides trogopterus
Nikolajev). – Nikolaev 1999: 178; Krell 2000a: 878; Nikolaev 2002: 54 (transl. p. 281).
(Cretanaides trogopterus Nikolajev)
Cretanaides trogopterus Nikolajev, 1996: 95 (transl. p. 220) (L Cretaceous, Baysa, Russia). – Krell 2000a: 878. Doubtful (only one elytron).
Subfamilia Hybosorinae
Genus Cretohybosorus Nikolajev, 1999
Cretohybosorus Nikolajev, 1999: 178 (type species by original designation: Cretohybosorus buryaticus
Nikolajev). – Krell 2000a: 878; Nikolaev 2002: 54 (transl. p. 281).
Cretohybosorus buryaticus Nikolajev
Cretohybosorus buryaticus Nikolajev, 1999: 179 (L Cretaceous, Baysa, Russia). – Krell 2000a: 878.
Cretohybosorus striatulus Nikolajev
Cretohybosorus striatulus Nikolajev, 1999: 179 (L Cretaceous, Baysa, Russia). – Krell 2000a: 878.
Hybosorus lividus Heer
Hybosorus lividus Heer, 1862: 77, pl. 6 (Miocene, Öhningen, Germany). – Scudder 1891: 534; Handlirsch 1907:
838; Krell 2000a: 878.
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Phaeochrous tertiarium (Deichmüller) Nikolaev
Bolboceras tertiarium Deichmüller, 1881: 304 (Eocene, Ku?lín [Kutschlin] near Bílina [Bilin], Czechia). –
Scudder 1891: 474; Po?ta 1900: 265; Handlirsch 1907: 838; ?íha 1979: 26.
Phaeochrous tertiarium (Deichmüller). – Nikolaev, 1996: 96 (transl. p. 222); Krell 2000a: 878.
Familia Ochodaeidae
Subfamilia Cretochodaeinae Nikolaev, 1995a: 78 (transl. p. 120)
Genus Cretochodaeus Nikolajev
Cretochodaeus Nikolajev, 1995a: 79 (type species by original designation: Cretochodaeus mongolicus
Nikolajev). – Nikolaev 1999: 178; Krell 2000a: 878; Nikolaev 2002: 54 (transl. p. 281).
Cretochodaeus mongolicus Nikolajev
Cretochodaeus mongolicus Nikolajev, 1995a: 79 (L Cretaceous, Bon-Tsagan, Mongolia). – Krell 2000a: 878.
(Cretochodaeus striatus Nikolajev)
Cretochodaeus striatus Nikolajev, 1995a: 80 (L Cretaceous, Bon-Tsagan, Mongolia). – Krell 2000a: 878. Doubtful (very incomplete, lacking legs and head appendages).
Genus Mioochodaeus Nikolajev
Mioochodaeus Nikolajev, 1995a: 81 (type species by original designation: Geotrupes proaevus Germar). – Krell
2000a: 878.
Mioochodaeus proaevus (Germar) Nikolaev
Geotrupes proaevus Germar, 1849: 57, pl. 2 (Oligocene, Orsberg near Rott, Germany). – Giebel 1852b: 653;
Giebel 1856: 36; Scudder 1891: 525; Handlirsch 1907: 838.
Geotrupes proavus. – Goss 1878: 331; Meunier 1921: 4.
Mioochodaeus proaevus. – Nikolaev, 1995a: 81; Krell 2000a: 878.
Familia Scarabaeidae
Subfamilia Scarabaeinae (incl. Coprinae)
Anachalcos mfwangani Paulian
Anachalcos mfwangani Paulian, 1976: 1 (Miocene, Lake Victoria, Kenya). – Krell 2000a: 879.
Genus Ateuchites Meunier
Ateuchites Meunier, 1898a: 114 (type species by monotypy: Ateuchites grandis Meunier). – Handlirsch 1925:
246; Théodoridès 1952: 34; Balthasar 1963: 79; Iablokoff-Khnzorian 1977: 137; Carpenter 1992: 300;
Krell 2000a: 879; Paetel 2001: 234.
Ateuchites grandis Meunier
Ateuchites grandis Meunier, 1898a: 114 (Oligocene, Armissan, Aude, France). – Handlirsch 1907: 836;
Carpenter 1992: 300; Krell 2000a: 879.
Choeridium ebenium Horn
Choeridium ebenium Horn, 1876: 245 (Pleistocene, Irvingtonian, Port Kennedy caves, Pennsylvania, U.S.A.). –
Scudder 1890: 490, pl. 1; Wickham 1920: 358; Théodoridès 1952: 36; Krell 2000a: 879.
Choeridium ? ebenium Horn. – Lesley 1889: xiii; Scudder 1891: 490; Scudder 1900: 104; Handlirsch 1908: 1126.
Copris druidum Heer
page 11
Copris druidum Heer, 1862: 73, pl. 6 (Miocene, Öhningen, Kesselstein, Germany). – Heer 1865: 378f; Heer
1883: 404f; Scudder 1891: 500; Handlirsch 1907: 837; Krell 2000a: 879.
Copris (Copris) kartlinus Kabakov
Copris (Copris) kartlinus Kabakov, 1988: 110 (Pliocene, Kisatibi formation, Georgia). – Krell 2000a: 879.
Copris leakeyorum Paulian
Copris leakeyorum Paulian, 1976: 1 (Miocene, Lake Victoria, Kenya). – Krell 2000a: 879.
Copris pristinus Pierce
Copris pristinus Pierce, 1946a: 124; (Pleistocene, Rancho La Brea tar pits, Los Angeles, U.S.A.). – Halffter
1959: 176; Matthews 1961: 35, 67, 69; Matthews & Halffter 1968: 160 (rembuchei-Gruppe); Sphon
1973: 52; Miller et al. 1981: 626; Stock & Harris 1992: 70, 84; Miller 1997: 188; Krell 2000a: 879;
Morón 2003: 12.
Copris subterraneus Heer
Copris subterranea Heer, 1862: 74, pl. 3 (Miocene, Öhningen, Kesselstein, Germany). – Heer 1865: 379; Heer
1883: 405; Scudder 1891: 501; Handlirsch 1907: 837; Krell 2000a: 879.
Gymnopleurus deperditus Heer
Gymnopleurus deperditus Heer, 1862: 73, pl. 6 (Miocene, Öhningen, Kesselstein, Germany). – Handlirsch 1907:
836; Meunier 1921: 11; Krell 2000a: 879.
(Gymnopleurus eocaenicus Meunier)
Gymnopleurus eocaenicus Meunier, 1921: 12, pl. 3 (Eocene, Messel, Germany). – Koenigswald 1987: 140; Krell
2000a: 879; Paetel 2001: 234.
Gymnopleurus eocenicus. – Théodoridès 1952: 46.
Doubtful (poor preservation).
Gymnopleurus rotundatus Heer
Gymnopleurus rotundatus Heer, 1862: 73, pl. 6 (Miocene, Öhningen, Kesselstein, Germany). – Heer 1865: 378f;
Heer 1883: 404f; Scudder 1891: 527; Handlirsch 1907: 837; Meunier 1921: 11; Krell 2000a: 879.
Gymnopleurus sisyphus Heer
Gymnopleurus sisyphus Heer, 1847: 64, pl. 7 (Miocene, Öhningen, Germany). – Bronn 1849: 625; Stitzenberger
1851: 100; Giebel 1852b: 653; Giebel 1856: 38; Heer 1862: 72; Scudder 1891: 527; Handlirsch 1907:
839; Meunier 1921: 11f; Krell 2000a: 879.
Heliocopris antiquus Fujiyama
Heliocopris antiquus Fujiyama, 1968: 203 (Miocene, Noto, Japan). – Harusawa 1994: 23; Krell 2000a: 879.
Metacatharsius rusingae Paulian
Metacatharsius rusingae Paulian, 1976: 2 (Miocene, Lake Victoria, Kenya). – Krell 2000a: 879.
Oniticellus amplicollis Heer
Oniticellus amplicollis Heer, 1862: 76, pl. 6 (Miocene, Öhningen, Kesselstein, Germany). – Heer 1865: 378f;
Heer 1883: 404f; Scudder 1891: 558; Handlirsch 1907: 837; Krell 2000a: 879, 2000b: 177.
(Onitis magus Heyden)
Onitis magus Heyden, 1862: 65, pl. 10 (Oligocene, Rott, Germany). – Krantz 1867: 315; Scudder 1891: 558;
Handlirsch 1907: 837; Krell 2000a: 879.
Onitis magnus. – Statz 1952: 2.
page 12
Zonitis. – After Janssens (in Balthasar 1963: 79) it is rather a species of Zonitis (Meloidae) and Heyden may
have mixed up the names. Slender tibiae without denticles indicate that it is not an Onitis.
Doubtful (tibiae structure untypical for Scarabaeoidea; poorly preserved).
Onthophagus bisontinus Heer
Onthophagus bisontinus Heer, 1862: 76, pl. 76 (Miocene, Öhningen, “Insektenschicht des unteren Bruches”,
Germany). – Heer 1865: 379; Heer 1883: 405; Scudder 1891: 559; Handlirsch 1907: 837; Krell 2000a:
880.
Onthophagus crassus Heer
Onthophagus crassus Heer, 1862: 75, pl. 6 (Miocene, Öhningen, Kesselstein, Germany). – Heer 1865: 379; Heer
1883: 405; Oustalet 1874: 196; Scudder 1891: 559; Handlirsch 1907: 837; Krell 2000a: 880.
Onthophagus everestae Pierce
Onthophagus everestae Pierce, 1946a: 131 (Pleistocene, Los Angeles. U.S.A.). – Sphon 1973: 52; Miller et al.
1981: 627f; Wilson 1986: 101; Stock & Harris 1992: 70, 84; Miller 1997: 187f; Krell 2000a: 880.
Onthophagus luteus Oustalet
Onthophagus luteus Oustalet, 1874: 194, pl. 2 (Oligocene, Aix en Provence, France). – Goss 1878: 339; Scudder
1891: 559; Handlirsch 1907: 837; Théobald 1937: tabl. 11; Théodoridès 1952: 46; Krell 2000a: 880.
Onthophagus ovatulus Heer
Onthophagus ovatulus Heer, 1847: 64, pl. 7 (Miocene, Öhningen, Germany). – Bronn 1849: 624; Giebel 1852b:
653; Giebel 1856: 39; Heer 1865: 379; Heer 1883: 405; Scudder 1891: 559; Handlirsch 1907: 837;
Krell 2000a: 880.
Onthophagus prodromus Heer
Onthophagus prodromus Heer, 1862: 75, pl. 6 (Miocene, Öhningen, Kesselstein, Germany). – Heer 1865: 378f;
Heer 1883: 404f; Scudder 1881-1885: 795; Oustalet 1874: 196; Scudder 1891: 559; Handlirsch 1907:
837; Krell 2000a: 880.
Onthophagus statzi Krell
Onthophagus muelleri Statz, 1952: 8 (nec Novak, 1921) (Oligocene, Rott, Germany).
Onthophagus mulleri. – Sphon 1973: 52.
Onthophagus statzi Krell, 1990: 187. – Krell 2000a: 880.
Onthophagus urusheeri Krell
Onthophagus urusheeri Krell, 2000a: 880.
Onthophagus urus Heer, 1847: 62, pl. 2 (Miocene, Öhningen, Germany). – Stitzenberger 1851: 100; Giebel
1852b: 653; Giebel 1856: 39; Bronn 1849: 624; Heer 1862: 76; Heer 1865: 379; Heer 1883: 405;
Scudder 1891: 559; Handlirsch 1907: 837; preoccupied by Onthophagus urus Ménétries, 1832: 175.
Phanaeus antiquus Horn
Phanaeus antiquus Horn, 1876: 245 (Pleistocene, Irvingtonian, Port Kennedy caves, Pennsylvania, U.S.A.). –
Goss 1878: 340; Scudder 1890: 489, pl. 1; Scudder 1891: 565; Scudder 1900: 104; Handlirsch 1908:
1126; Wickham 1920: 358; Théodoridès 1952: 36; Krell 2000a: 880.
Phanaeus labreae (Pierce) Miller
Palaeocopris labreae Pierce, 1946a: 130 (Pleistocene, Rancho La Brea tar pits, Los Angeles, U.S.A.). –
Matthews 1961: 35 (“appears to be a composite of two genera”); Sphon 1973: 52; Stock & Harris 1992:
84; Wilson 1986: 101.
page 13 Paleocopris labreae. – Halffter 1959: 176.
Phanaeus labreae. – Miller et al. 1981: 627; Krell 2000a: 880.
Genus Prionocephale Lin
Prionocephale Lin, 1980: 230 (type species by original designation: Prionocephale deplanate Lin). – Krell
2000a: 880; Paetel 2001: 234.
Prionocephale deplanate Lin
Prionocephale deplanate Lin, 1980: 230 (U Cretaceous, Zheijang, China). – Lin 1994: 314; Krell 2000a: 880. Prionocephale deplanae. – Lin 1983: 394.
Genus Scelocopris Zhang
Scelocopris Zhang 1989: 150, 425 (type species by original designation: Scelocopris enertheus Zhang). – Krell
2000a: 880; Paetel 2001: 234.
Scelocopris enertheus Zhang
Scelocopris enertheus Zhang, 1989: 151, 425 (Miocene, Shanwang, China). – Krell 2000a: 880.
Subfamilia Aegialiinae
Aegialia rupta Scudder
Aegialia rupta Scudder, 1890: 489, pl. 8 (Eocene, Green River, Wyoming, U.S.A.). – Scudder 1891: 461;
Scudder 1900: 104; Handlirsch 1907: 838; Wickham 1920: 358; Dellacasa 1988: 359; Krell 2000a: 880.
Genus Cretaegialia Nikolayev
Cretaegialia Nikolayev, 1994: 5 (type species by original designation: Cretaegialia rhypariformis Nikolayev). –
Nikolaev 1999: 178; Krell 2000a: 880; Nikolaev 2002: 54 (transl. p. 281).
Cretaegialia aphodiiformis Nikolayev
Cretaegialia aphodiiformis Nikolayev, 1994: 7 (L Cretaceous, Baysa, Russia). – Krell 2000a: 880.
Cretaegialia rhypariformis Nikolayev
Geotrupoides sulcatus Nikritin, 1977: 120 [part.]. – Nikolayev 1994: 6.
Cretaegialia rhypariformis Nikolayev, 1994: 6 (L Cretaceous, Baysa, Russia). – Krell 2000a: 880.
Subfamilia Aphodiinae
Tribus Aphodiini
Aphodius aboriginalis Wickham
Aphodius aboriginalis Wickham, 1912: 22 (Oligocene, Florissant, U.S.A.). – Wickham 1913a: 17; Wickham
1913c: 360; Wickham 1914b: 455; Wickham 1920: 358; Dellacasa 1988: 400; Krell 2000a: 881.
Aphodius anteactus Krell
Aphodius anteactus Krell, 2000a: 881.
Aphodius antiquus Heer, 1847: 66, pl. 7 (Miocene, Öhningen, Germany). – Bronn 1849: 624; Stitzenberger 1851:
100; Giebel 1852b: 653; Giebel 1856: 40; Scudder 1891: 468; Handlirsch 1907: 838; Schmidt 1910:
140; Dellacasa 1988: 400; preoccupied by Aphodius antiquus Faldermann, 1835: 367.
Aphodius boryslavicus ?omnicki
Aphodius boryslavicus ?omnicki 1894: 83, pl. 7 (Pleistocene, Borys?aw, Poland). – Krell 2000a: 881.
Aphodius bosniaskii Handlirsch
page 14
? Aphodius bosniaskii Handlirsch, 1907: 838 (Miocene, Gabbro, Livorno, Italy) [only length and location given,
no real description, which, however, may be considered as a formal one sensu Art. 12 ICZN]. –
Schmidt 1910: 140 (“Aphod.? Bosniaskii”).
Aphodius bosniaskii. – Dellacasa 1983: 29; Dellacasa 1988: 400; Krell 2000a: 881.
Aphodius brevipennis Heer
Aphodius brevipennis Heer, 1862: 77, pl. 6 (Miocene, Öhningen, Kesselstein, Germany). – Scudder 1891: 468;
Handlirsch 1907: 838; Schmidt 1910: 140; Dellacasa 1988: 400; Krell 2000a: 881.
Aphodius charauxi Piton
Aphodius charauxi Piton, 1940: 176 (Eocene, Menat, France). – Krell 2000a: 881.
Aphodius florissantensis Wickham
Aphodius florissantensis Wickham, 1911: 59 (Oligocene, Florissant, U.S.A.). – Wickham 1912: 23; Wickham
1913a: 17; Wickham 1914b: 456; Wickham 1920: 358; Dellacasa 1988: 400; Krell 2000a: 881.
Aphodius granarioides Wickham
Aphodius granarioides Wickham, 1913a: 17 (Oligocene, Florissant, U.S.A.). – Wickham 1912: 23 (nomen
nudum); Wickham 1913b: 295; Wickham 1920: 358; Théodoridès 1952: 35; Dellacasa 1988: 400; Krell
2000a: 881.
Aphodius helvolus Statz
Aphodius helvolus Statz, 1952: 6 (Oligocene, Rott, Germany). – Sphon 1973: 52; Dellacasa 1991: 36; Krell
2000a: 881.
Aphodius inundatus Wickham
Aphodius inundatus Wickham, 1914a: 262 (Oligocene, Florissant, U.S.A.). – Wickham 1920: 358; Théodoridès
1952: 35; Dellacasa 1988: 400; Krell 2000a: 881.
Aphodius krantzi Heyden & Heyden
Aphodius krantzi Heyden & Heyden, 1866: 138, pl. 22 (Oligocene, Rott, Germany). – Krantz 1867: 315; Goss
1878: 333; Scudder 1891: 468; Handlirsch 1907: 838; Schmidt 1910: 140; Statz 1952: 2; Dellacasa
1988: 400; Krell 2000a: 881.
Aphodius laminicola Wickham
Aphodius laminicola Wickham, 1910: 49 (Oligocene, Florissant, U.S.A.). – Wickham 1911: 59; Wickham 1912:
22f; Wickham 1913a: 17; Wickham 1914b: 456; Wickham 1920: 358; Dellacasa 1988: 400; White
1995: 4; Krell 2000a: 881.
Aphodius mediaevus Wickham
Aphodius mediaevus Wickham, 1914b: 455 (Oligocene, Florissant, U.S.A.). – Wickham 1920: 358; Dellacasa
1988: 400; Krell 2000a: 881.
Aphodius meyeri Heer
Aphodius meyeri Heer, 1847: 67, pl. 7 (Miocene, Öhningen, Germany). – Bronn 1849: 624; Stitzenberger 1851:
100; Giebel 1852b: 653; Giebel 1856: 40; Scudder 1891: 468; Handlirsch 1907: 838; Schmidt 1910:
140; Dellacasa 1988: 400; Krell 2000a: 881.
Aphodius praeemptor Wickham
Aphodius praeemptor Wickham, 1913a: 17, pl. 6 (Oligocene, Florissant, U.S.A.). – Wickham 1914b: 456;
Wickham 1920: 358; Dellacasa 1988: 400; Krell 2000a: 881.
Aphodius precursor Horn
page 15
Aphodius precursor Horn, 1876: 245 (Tertiary/Pleistocene, Bone Caves of Port Kennedy, Pennsylvania, U.S.A.).
– Goss 1878: 340; Scudder 1890: 488, pl. 1; Scudder 1900: 105; Wickham 1920: 358; Krell 2000a: 881.
Aphodius praecursor. – Lesley 1889: v; Scudder 1891: 469; Handlirsch 1908: 1126; Schmidt 1910: 140;
Dellacasa 1988: 400.
Aphodius rhinocerontis ?omnicki
Aphodius rhinocerontis ?omnicki 1894: 81, pl. 7 (Pleistocene, Borys?aw, Poland). – Krell 2000a: 881.
Aphodius ruthenus ?omnicki
Aphodius ruthenus ?omnicki 1894: 82, pl. 7 (Pleistocene, Borys?aw, Poland). – Krell 2000a: 882.
Aphodius schlickumi Statz
Aphodius schlickumi Statz, 1952: 6 (Oligocene, Rott, Germany). – Dellacasa 1991: 36; Lutz 1996: 45; Krell
2000a: 882.
Aphodius senex Wickham
Aphodius senex Wickham, 1914b: 456 (Oligocene, Florissant, U.S.A.). – Wickham 1920: 358; Dellacasa 1988:
400; Krell 2000a: 882.
Aphodius shoshonis Wickham
Aphodius shoshonis Wickham, 1912: 23 (Oligocene, Florissant, U.S.A.). – Wickham 1913a: 17; Wickham
1914b: 455; Wickham 1920: 358; Dellacasa 1988: 400; Krell 2000a: 882.
Aphodius subater ?omnicki
Aphodius subater ?omnicki 1894: 80, pl. 7 (Pleistocene, Borys?aw, Poland). – Krell 2000a: 882.
Aphodius theobaldi Krell
Aphodius theobaldi Krell, 2000a: 882.
Aphodius incertus Théobald, 1937: 126, pl. 10 (Oligocene, Sannoisien du Gard, France). – Théodoridès 1952: 46;
preoccupied by Aphodius incertus Ballion, 1878: 282.
Tribus Eupariini
Ataenius europaeus Quiel
Ataenius europaeus Quiel, 1910: 187 (Eocene, Baltic Amber). – Kolbe 1925: 150; Dellacasa 1988: 412; Krell
2000a: 882.
Ataenius patescens Scudder
Ataenius patescens Scudder, 1893: pl. 1 (not described, only figured; Oligocene, Florissant, U.S.A.). – Scudder
1900: 104, pl. 11 (description); Handlirsch 1907: 838; Schmidt 1910: 140; Wickham 1910: 48;
Wickham 1912: 22; Wickham 1913b: 295; Wickham 1913c: 360; Wickham 1914b: 453; Wickham
1920: 358; Dellacasa 1988: 412; Krell 2000a: 882.
Ataenius restructus (Wickham) Wickham
Aphodius restructus Wickham, 1912: 22 (Oligocene, Florissant, U.S.A.). – Wickham 1913a: 17; Dellacasa 1988:
400.
Ataenius restructus. – Wickham 1914b: 454; Wickham 1920: 358; Krell 2000a: 882.
Oxyomus nearcticus Wickham
Oxyomus nearcticus Wickham, 1914b: 453 (Oligocene, Florissant, U.S.A.). – Wickham 1920: 358; Dellacasa
1988: 403; Krell 2000a: 882.
Saprosites cascus Britton
page 16
Saprosites cascus Britton, 1960: 36 (Eocene, Bognor Regis, Sussex, England). – Jarzembowski 1992: 94; Krell
2000a: 882.
Saprosites succini (Zang) Kolbe
Aphodius succini Zang, 1905: 204 (Eocene, Baltic Amber). – Handlirsch 1908: 1356; Schmidt 1910: 140. Ataenius succini. – Quiel 1910: 187; Dellacasa 1988: 412.
Saprosites succini. – Kolbe 1925: 150; Hieke & Pietrzeniuk 1984: 316; Krell 2000a: 882.
Subfamilia Melolonthinae
(Diplotaxis aurora Wickham)
Diplotaxis aurora Wickham, 1913b: 294 (Oligocene, Florissant, U.S.A.). – Wickham, 1912: 26; Wickham 1920:
358; Vaurie 1960: 414 (doubts generic affiliation because of the described characters); Krell 2000a: 882.
Doubtful (only one elytron).
(Diplotaxis) simplicipes Wickham
Diplotaxis ? simplicipes Wickham, 1912: 25 (Oligocene, Florissant, U.S.A.). – Wickham 1920: 358; Vaurie
1960: 414 (doubts generic affiliation because of the described structures of the legs); Krell 2000a: 882.
Genus Eophyllocerus Haupt
Eophyllocerus Haupt, 1950: 56 (type species by present designation: Eophyllocerus scrobiculatus Haupt [it is
the first mentioned species with most specimens found]). – Crowson 1981: 680; Scholtz & Chown 1995:
364; Krell 2000a: 882.
Eophyllocerus scrobiculatus Haupt
Eophyllocerus scrobiculatus Haupt, 1950: 57 (Eocene, Geiseltal, Germany). – Krumbiegel 1982: 15; Müller
1989: 244; Krell 2000a: 882.
Eophyllocerus glaucinus Haupt
Eophyllocerus glaucinus Haupt, 1950: 58 (Eocene, Geiseltal, Germany). – Krumbiegel 1982: 15; Krell 2000a:
882.
Macrodactylus pluto Wickham
Macrodactylus pluto Wickham, 1912: 24 (Oligocene, Florissant, U.S.A.). – Wickham 1913b: 294; Wickham
1920: 358; Rodeck 1938: 294; Krell 2000a: 882.
Macrodactylus propheticus Wickham
Macrodactylus propheticus Wickham, 1912: 25 (Oligocene, Florissant, U.S.A.). – Wickham 1920: 358; Rodeck
1938: 294; Krell 2000a: 882.
Melolontha greithiana Heer
Melolontha greithiana Heer, 1847: 67 (Oligocene, Greith, Switzerland). – Bronn 1848a: 720; Bronn 1849: 624;
Heer 1865: 378, 380; Heer 1883: 406; Scudder 1891: 551; Handlirsch 1907: 840; Krell 2000a: 882.
Melolontha greithana. – Giebel 1852b: 652; Giebel 1856: 32; Goss 1879: 145.
(Melolontha solitaria Novák)
Melolontha solitaria Novák, 1878: 92, pl. 3 (Miocene, Mok?ina [Krottensee] near Kynšperk n. O., Czechia). –
Scudder 1891: 552; Po?ta 1900: 264; Handlirsch 1907: 840; ?íha 1977: 22; ?íha 1979: 26; Krell 2000a:
883.
Doubtful (only one elytron).
Genus Melolonthites Heer (collective group)
page 17
Melolonthites Heer, 1847: 71 (as a collective group, this taxon has no type species [Art. 13.3.2 ICZN], though a
the type species was designated by Carpenter 1992: Melolonthites aciculatus Heer.). – Bronn 1848a:
720; Bronn 1849: 624; Rogg 1852: 22; Giebel 1856: 33; Scudder 1886: 74; Scudder 1895: 120;
Handlirsch 1925: 246; Théodoridès 1952: 36; Carpenter 1992: 323 (suborder and family uncertain);
Krell 2000a: 883.
(Melolonthites aciculatus Heer)
Melolonthites aciculata Heer, 1847: 71, pl. 2 (Miocene, Öhningen, Germany). – Stitzenberger 1851: 100; Giebel
1852b: 652; Giebel 1856: 33.
Melolonthites aciculatus. – Bronn 1848a: 720; Heer 1849: 624; Scudder 1891: 552; Handlirsch 1907: 840;
Carpenter 1992: 323; Krell 2000a: 883.
Doubtful (only one abdomen [5 sternites]).
(Melolonthites collinsi Wickham)
Melolonthites collinsi Wickham, 1929: 149 (Eocene, Holcomb, Tennessee, U.S.A.). – Wickham 1933: 104;
Krell 2000a: 883.
Doubtful (only one elytron).
Melolonthites deperditus Heer
Melolonthites deperdita Heer, 1847: 71, pl. 2 (Miocene, Öhningen, Germany). – Stitzenberger 1851: 100; Giebel
1852b: 652; Giebel 1856: 33.
Melolonthites deperditus. - Bronn 1848a: 720; Bronn 1849: 624; Scudder 1891: 552; Handlirsch 1907: 840;
Krell 2000a: 883.
Melolonthites interemtus Cockerell
Melolonthites interemtus Cockerell, 1926: 319 (Oligocene, Kudia River, Siberia, Russia) (only part of an ala;
“placed in Heer?s blanket-genus Melolonthites merely to avoid proposing a new generic name”). – Krell
2000a: 883.
(Melolonthites kollari Heer)
Melolonthites kollari Heer, 1847: 72, pl. 7 (Miocene, “in den Kohlen zu Parschlug in Steiermark”, Austria). –
Bronn 1848a: 720; Bronn 1849: 624; Giebel 1856: 33; Scudder 1891: 552; Handlirsch 1907: 840; Beier
1952: 131; Krell 2000a: 883.
Melolonthites kolleri. – Giebel 1852b: 652.
Doubtful (described after only a drawing of one elytron).
(Melolonthites laterosinuatus Piton in Piton & Théobald)
Melolonthites laterosinuatus Piton in Piton & Théobald, 1935: 78, pl. 3 (Mio/Pliocene, Cinérites de Varennes,
France). – Piton 1936: 18 (“latero-sinuatus”); Krell 2000a: 883.
Doubtful (only one elytron).
(Melolonthites lavateri Heer)
Melolonthites lavateri Heer, 1847: 73, pl. 7 (Miocene, Öhningen, Germany). – Bronn 1848a: 720; Bronn 1849:
624; Stitzenberger 1851: 100; Giebel 1852b: 652; Giebel 1856: 34; Scudder 1891: 552; Handlirsch
1907: 840; Krell 2000a: 883.
Doubtful (only one elytron).
(Melolonthites obsoletus Heer)
page 18
Melolonthites obsoleta Heer, 1847: 73, pl. 2 (Miocene, Öhningen, Germany). – Stitzenberger 1851: 100; Giebel
1852b: 652; Giebel 1856: 34.
Melolonthites obsoletus. – Bronn 1848a: 720; Bronn 1849: 624; Scudder 1891: 552; Handlirsch 1907: 840; Krell
2000a: 883.
Doubtful (only one elytron, abdomen and a part of the thorax indistinctly visible).
(Melolonthites parschlugianus Heer)
Melolonthites parschlugiana Heer, 1847: 72 (Miocene, “in den Kohlen zu Parschlug in Steiermark”, Austria). –
Unger 1870: 3, pl. 1.
Melolonthites parschlugianus. – Bronn 1848a: 720; Scudder 1891: 552; Handlirsch 1907: 840; Beier 1952: 131;
Krell 2000a: 883.
Melolonthites parschluganus. – Bronn 1849: 624.
Melolonthites parschlugana. – Giebel 1852b: 652; Giebel 1856: 33.
Doubtful (described after only a drawing of one elytron).
Phyllophaga avus (Cockerell) Cockerell
Melolonthites avus Cockerell, 1921: 36, pl. 8 (Eocene, White River, Green River Formation, Colorado, U.S.A.).
– Morón 2003: 13.
Phyllophaga avus. – Cockerell 1925: 10 (Eocene, Roan Mountains, Green River Formation, Colorado, U.S.A.);
Wickham 1927: 55; Krell 2000a: 883.
Phyllophaga disrupta Cockerell
Phyllophaga disrupta Cockerell, 1927b: 587 (Oligocene, Florissant, U.S.A.). – Wickham 1933: 104; Krell 2000a:
883; Morón 2003: 13.
(Phyllophaga extincta (Wickham) Wickham)
Lachnosterna ? extincta Wickham, 1916: 9 (Oligocene, Florissant, U.S.A.). – Rodeck 1938: 294.
Phyllophaga extincta. – Wickham 1920: 358; Krell 2000a: 883.
Doubtful (only one elytron).
Rhizotrogus longimanus Heer
Rhizotrogus longimanus Heer, 1847: 69, pl. 7 (Miocene, Öhningen, Germany). – Bronn 1849: 624; Stitzenberger
1851: 100; Giebel 1852b: 652; Giebel 1856: 32; Heer 1865: 380; Heer 1883: 406; Scudder 1891: 576;
Handlirsch 1907: 839; Krell 2000a: 883.
Listrochelus puerilis Wickham
Listrochelus puerilis Wickham, 1914b: 459, pl. 6 (Oligocene, Florissant, U.S.A.). – Wickham 1920: 358; Krell
2000a: 884; Morón 2003: 12.
Tribus Cretomelolonthini Nikolajev 1998
Cretomelolonthini Nikolajev, 1998: 80 (transl. p. 515).
Genus Cretomelolontha Nikolajev
Cretomelolontha Nikolajev, 1998a: 81 (transl. p. 516) type genus (type species by original designation:
Cretomelolontha transbaikalica Nikolajev). – Nikolaev 1999: 178; Krell 2000a: 884; Nikolaev 2002:
54 (transl. p. 281).
Cretomelolontha transbaikalica Nikolajev
page 19
Cretomelolontha transbaikalica Nikolajev, 1998a: 82 (transl. p. 517) (L Cretaceous, Baysa, Russia). – Krell
2000a: 884.
Subfamilia Dynastinae
Ligyrus compositus Wickham
Ligyrus compositus Wickham, 1911: 59 (Oligocene, Florissant, U.S.A.). – Wickham 1914b: 461; Wickham 1920:
358; Rodeck 1938: 294; Krell 2000a: 884.
(Ligyrus) effetus Wickham
Ligyrus effetus Wickham, 1914b: 461 (Oligocene, Florissant, U.S.A.). – Wickham 1920: 358; Krell 2000a: 884.
The generic classification was founded “by the facies”, that means not by characters. Therefore, the
generic affiliation is doubtful.
Pentodon bellerophon Heyden & Heyden
Pentodon bellerophon Heyden & Heyden, 1866: 139, pl. 23 (Oligocene, Rott, Germany). – Krantz 1867: 315;
Scudder 1891: 564; Handlirsch 1907: 841; Houlbert 1915: 64; Kolbe 1932: 150; Statz 1952: 3; Krell
2000a: 884.
(Pentodon proserpinae (Heer) Heer)
Scarabaeus proserpinae Heer, 1862: 78, pl. 6 (Miocene, Öhningen, Kesselstein, Germany). – Scudder 1891: 578. Pentodon proserpinae. – Heer 1865: 379; Heer 1883: 405; Scudder 1891: 564; Handlirsch 1907: 841; Houlbert
1915: 64; Kolbe 1932: 150; Krell 2000a: 884.
Doubtful (only one elytron).
Strategus cessatus Wickham
Strategus cessatus Wickham, 1914b: 461 (Oligocene, Florissant, U.S.A.). – Wickham 1920: 358; Ratcliffe 1976:
123, 169 [only elytron, but revised by Ratcliffe]; Krell 2000a: 884; Morón 2003: 13.
Subfamilia Rutelinae
Adoretus recticlypeus Zhang
Adoretus recticlypeus Zhang, 1989: 167, pl. 47 (Miocene, Shanwang, China). – Zhang et al. 1994: 110; Krell
2000a: 884.
Adoretus rhinus Zhang, Sun & Zhang
Adoretus rhinus Zhang, Sun & Zhang, 1994: 109, pl. 16 (Miocene, Shanwang, China). – Krell 2000a: 884.
Anomala amblobelia Zhang
Magniscarabaeus furvus part. – Hong 1985: 83.
Anomala amblobelia Zhang, 1989: 158, 160, 164, 174, pl. 42 (Miocene, Shandong, China). – Krell 2000a: 884.
Anomala brachytarsia Zhang, Sun & Zhang
Anomala brachytarsia Zhang, Sun & Zhang, 1994: 104, 107f, pl. 16 (Miocene, Shanwang, China). – Krell 2000a:
884.
Anomala endoxa Zhang, Sun & Zhang
Anomala endoxa Zhang, Sun & Zhang, 1994: 103, 106f, pl. 16 (Miocene, Shanwang, China). – Krell 2000a: 884.
Anomala eversa Zhang
Anomala eversa Zhang, 1989: 158, 165, 167, pl. 44 (Miocene, Shanwang, China). – Krell 2000a: 884.
Anomala exterranea Wickham
page 20
Anomala exterranea Wickham, 1914b: 459 (Oligocene, Florissant, U.S.A.). – Wickham 1920: 358; Krell 2000a:
884.
Anomala fugax Heer
Anomala fugax Heer, 1862: 80, pl. 6 (Miocene, Öhningen, Kesselstein, Germany). – Heer 1865: 380; Heer 1883;
406; Scudder 1891: 465; Handlirsch 1907: 841; Krell 2000a: 884.
Anomala furva (Hong & Wang) Zhang
Magniscarabaeus furvus Hong & Wang in Hong, 1983: 5, 13 (Miocene, Shanwang, China). – Hong 1985: 38, pl.
17; Hong & Wang 1986: 7; Hong & Wang 1987: 120. Anomala furva. – Zhang 1989: 158, 162, 164, 166f, 173, pl. 43; Zhang et al. 1994: 104, pl. 15; Krell 2000a: 884. Anomala turua. – Sun 1995: 38.
Anomala lochmocola Zhang
Anomala lochmocola Zhang, 1989: 158, 164, pl. 45 (Miocene, Shanwang, China). – Krell 2000a: 885.
Anomala orcina Zhang, Sun & Zhang
Anomala orcina Zhang, Sun & Zhang, 1994: 103, 105, pl. 15 (Miocene, Shanwang, China). – Krell 2000a: 885.
Anomala palaeobrunnea Krell
Anomala palaeobrunnea Krell, 2000a: 885.
Magniscarabaeus brunneus Hong, 1985: 39, pl. 42 (Miocene, Shanwang, China). – Hong & Wang 1987: 119ff. Anomala brunnea. – Zhang 1989: 158, 160f; junior secondary homonym of Anomala brunnea Klug, 1855
(extant species from Moçambique).
(Anomala) primigenia Heyden & Heyden
Anomala primigenia Heyden & Heyden, 1866: 138, pl. 22 (Oligocene, Rott, Germany). – Krantz 1867: 315;
Goss 1878: 333; Scudder 1891: 465; Handlirsch 1907: 840; Ohaus 1940: 74; Statz 1952: 2 (generic
affiliation doubtful); Krell 2000a: 885.
Anomala punctulata Zhang
Anomala punctulata Zhang, 1989: 158, 165, pl. 44 (Miocene, Shanwang, China). – Krell 2000a: 885.
Anomala scia Zhang
Anomala scia Zhang, 1989: 158, 166, pl. 45 (Miocene, Shanwang, China). – Krell 2000a: 885.
Anomala scudderi Wickham
Anomala scudderi Wickham, 1914b: 460 (Oligocene, Florissant, U.S.A.). – Wickham 1920: 358; Krell 2000a:
885.
Anomala synemosyna Zhang
Anomala synemosyna Zhang, 1989: 158, 161, pl. 42 (Miocene, Shanwang, China). – Krell 2000a: 885.
(Anomala) thetis Heyden & Heyden
Anomala thetis Heyden & Heyden, 1866: 139, pl. 24 (Oligocene, Rott, Germany). – Krantz 1867: 315; Goss
1878: 333; Scudder 1891: 465; Handlirsch 1907: 840; Ohaus 1940: 74; Statz 1952: 2 (generic affiliation
doubtful); Krell 2000a: 885.
(Anomala) tumulata Heyden & Heyden
Anomala tumulata Heyden & Heyden, 1866: 138, pl. 23 (Oligocene, Rott, Germany). – Krantz 1867: 315; Goss
1878: 333; Scudder 1891: 465; Handlirsch 1907: 841; Meunier 1920: 48 (no Lamellicornia); Ohaus
1940: 74; Statz 1952: 2 (Menier?s specimen: no Lamellicornia; Heydens? specimen may be one; generic
affiliation doubtful); Krell 2000a: 885.
page 21
Anomala ursa Zhang, Sun & Zhang.
Anomala ursa Zhang, Sun & Zhang, 1994: 104, 108f, pl. 16 (Miocene, Shanwang, China). – Krell 2000a: 885.
Genus Anomalites Fri?
Anomalites Fri?, 1885: 163 (type species by monotypy: Anomalites fugitivus Fri?). – Scudder 1881-1885: 794;
Scudder 1886: 74; Handlirsch 1925: 246; Carpenter 1992: 300, 330; Krell 2000a: 885.
Anomalites fugitivus Fri?
Anomalites fugitivus Fri?, 1885: 163 (Tertiary, Süßwasserquarz Nogent le Rotrou, France). – Handlirsch 1907:
841; Carpenter 1992: 300; Krell 2000a: 885.
Genus Zhangsunia Krell
Zhangsunia Krell, 2000a: 885.
Dolichopoda Zhang, Sun & Zhang, 1994: 101, 279 (type species by original designation: Dolichopoda extumida
Zhang, Sun & Zhang). – Jarzembowski 2000: (6); preoccupied by Dolichopoda Bolivar, 1880: 72
(Ensifera: Rhaphidophoridae).
Zhangsunia extumida (Zhang, Sun & Zhang) Krell Dolichopoda extumida Zhang, Sun & Zhang, 1994: 101f (Miocene, Shanwang, China).
Zhangsunia extumida. – Krell 2000a: 885.
Holotrichia cressona Zhang, Sun & Zhang
Holotrichia cressona Zhang, Sun & Zhang, 1994: 102f (Miocene, Shanwang, China). – Krell 2000a: 885.
Holotrichia spatha Zhang
Holotrichia spatha Zhang, 1989: 156 (Miocene, Shanwang, China). – Krell 2000a: 885.
Mimela rhenana (Heyden) Ohaus in Statz Anoplognathus rhenanus Heyden, 1862: 65, pl. 10 (Oligocene, Rott, Germany). – Krantz 1867: 315; Scudder
1891: 465; Handlirsch 1907: 841; Ohaus 1940: 74.
Mimela rhenana (Heyden). – Ohaus in Statz, 1952: 2; Krell 2000a: 885.
Genus Miolachnosterna Wickham
Miolachnosterna Wickham, 1914b: 458 (type species by original designation: Miolachnosterna tristoides
Wickham). – Théodoridès 1952: 36; Carpenter 1992: 301; Krell 2000a: 885.
Miolachnosterna tristoides Wickham
Miolachnosterna tristoides Wickham, 1914b: 458, pl. 8 (Oligocene, Florissant, U.S.A.). – Wickham 1920: 358;
Carpenter 1992: 301; Krell 2000a: 886; Morón 2003: 13.
Genus Pelidnotites Cockerell
Pelidnotites Cockerell, 1920: 463 (type species by monotypy: Pelidnotites atavus Cockerell). – Carpenter 1992:
301; Krell 2000a: 886.
Pelidnotites atavus Cockerell
Pelidnotites atavus Cockerell, 1920: 462 (Eocene, U Ypresian - Lutetian, Bournemouth, England). – Carpenter
1992: 301; Jarzembowski 1996: 210; Krell 2000a: 886.
Genus Petraeianus Zhang
Petraeianus Zhang, 1989: 168, 426 (type species by original designation: Petraeianus ruderatus Zhang). – Krell
2000a: 886.
Petraeianus ruderatus Zhang
Petraeianus ruderatus Zhang, 1989: 168, 426 (Miocene, Shanwang, China). – Krell 2000a: 886.
page 22
Genus Petulantis Zhang, Sun & Zhang
Petulantis Zhang, Sun & Zhang, 1994: 110, 279 (type species by original designation: Petulantis yimengensis
Zhang, Sun & Zhang). – Krell 2000a: 886.
Petulantis yimengensis Zhang, Sun & Zhang
Petulantis yimengensis Zhang, Sun & Zhang, 1994: 110, pl. 17 (Miocene, Shanwang, China). – Krell 2000a: 886.
Subfamilia Hopliinae
Hoplia striatipennis Wickham
Hoplia striatipennis Wickham, 1914b: 457 (Oligocene, Florissant, U.S.A.). – Wickham 1920: 358; Krell 2000a:
886.
Lepitrix germanica Heer
Lepitrix germanica Heer, 1862: 80, pl. 6 (Miocene, Öhningen, Germany). – Heer 1865: 378, 380; Heer 1883:
406; Scudder 1891: 546; Handlirsch 1907: 839; Krell 2000a: 886.
Subfamilia Sericinae
Genus Lithanomala Nikolajev
Lithanomala Nikolaev, 1992: 79 (transl. p. 100) (type species by original designation: Proteroscarabaeus
oblongus Ponomarenko). – Nikolaev 1998a: 82 (transl. p. 519); Nikolaev 1999: 178; Krell 2000a: 886.
Lithanomala crassa (Ponomarenko) Nikolaev
Proteroscarabaeus crassus Ponomarenko, 1990: 66, pl. 6 (L Cretaceous, Leskovo, Russia). Lithanomala crassa. – Nikolaev, 1992: 80 (transl. p. 100); Krell 2000a: 886.
Lithanomala oblonga (Ponomarenko) Nikolaev
Proteroscarabaeus oblongus Ponomarenko, 1990: 66, pl. 6 (L Cretaceous, Leskovo, Russia).
Lithanomala oblonga. – Nikolaev, 1992: 79 (transl. p. 100); Krell 2000a: 886.
Lithanomala sibirica (Ponomarenko) Nikolaev
Proteroscarabaeus sibiricus Ponomarenko, 1990: 68, pl. 6 (L Cretaceous, Leskovo, Russia). Lithanomala sibirica. – Nikolaev, 1992: 80 (transl. p. 100); Krell 2000a: 886.
Maladera spinitibialis Statz
Maladera spinitibialis Statz, 1952: 9 (Oligocene, Rott, Germany). – Krell 2000a: 886.
Maladera ? spinitibialis. – Sphon 1973: 52.
Genus Mioserica Zhang
Mioserica Zhang, 1989: 157, 426 (type species by original designation: Mioserica margelis Zhang). – Krell
2000a: 886.
Mioserica margelis Zhang
Mioserica margelis Zhang, 1989: 157, 426 (Miocene, Shanwang, China). – Krell 2000a: 886.
(Serica) antediluviana Wickham
Serica antediluviana Wickham, 1912: 23 (Oligocene, Florissant, U.S.A.). – Wickham 1914a: 263; Wickham
1920: 358; Rodeck 1938: 294; Krell 2000a: 886.
According to the description, the specimen shows no character allowing to classify it as belonging to the
Sericinae.
(Serica) cockerelli Wickham
page 23
Serica cockerelli Wickham, 1914a: 262 (Oligocene, Florissant, U.S.A.). – Wickham 1920: 358; Théodoridès
1952: 35; Krell 2000a: 886.
According to the description, the specimen shows no character allowing to classify it as belonging to the
Sericinae.
Serica kanakoffi Pierce
Serica kanakoffi Pierce, 1946b: 132 (Pleistocene, Rancho La Brea tar pits, Los Angeles, U.S.A.). – Sphon 1973:
53; Miller et al. 1981: 627; Stock & Harris 1992: 84; Krell 2000a: 886.
Serica minutula Heer
Serica minutula Heer, 1862: 79, pl. 6 (Miocene, Öhningen, Kesselstein, Germany). – Heer 1865: 380; Heer 1883:
406; Scudder 1891: 579; Handlirsch 1907: 839; Krell 2000a: 886.
Subfamilia Cetoniinae
(Cetonia defossa Weyenbergh)
Cetonia (?) defossa Weyenbergh, 1869a: 282, pl. 37 (Amphicoma ?) (U Jurassic, Solnhofen, Germany). –
Weijenbergh 1869b*: [232]; Weijenbergh 1874: 103 [“defosfa”, lapsus calami] or 109 [“defossa”];
Goss 1879: 148; Winkler 1878: 96; Altena 1958.
Cetonia defossa. – Weijenbergh 1869b: [232]; Scudder 1891: 197; Meunier 1897: 235, pl. 10 (“indéterminable
empreinte de coléoptère”); Krell 2000a: 887.
? Ditomoptera defossa (Coleoptera incert. sed.). – Handlirsch 1906: 546 (classification doubted by Altena 1958)
Genus Genonota Zhang
Genonota Zhang, 1989: 170, 426 (type species by original designation: Genonota mochthera Zhang). – Krell
2000a: 887.
Genonota mochthera Zhang
Genonota mochthera Zhang, 1989: 170f, 426, pl. 46 (Miocene, Shanwang, China). – Krell 2000a: 887.
Genonota scola Zhang
Genonota scola Zhang, 1989: 170f, pl. 47 (Miocene, Shawang, China). – Krell 2000a: 887.
Macronota shandongiana Zhang
Macronota shandongiana Zhang, 1989: 169, pl. 45 (Miocene, Shandong, China). – Krell 2000a: 887.
Subfamilia Trichiinae
Gnorimus aedilis (Heer) Frentzen
Trichius (Gnorimus) aedilis Heer, 1862: 81, pl. 6 (Miocene, Öhningen, Kesselstein, Salamanderschicht,
Germany). – Heer 1865: 378, 380; Scudder 1891: 591.
Trichius aedilis. – Heer 1883: 404f; Scudder 1891: 591; Handlirsch 1907: 841. Gnorimus aedilis. – Frentzen 1927: 141; Kolbe 1931: 117; Kolbe 1932: 150; Krell 2000a: 887.
Gnorimus lugubris (Heer) Kolbe
Trichius (Gnorimus) lugubris Heer, 1862: 81, pl. 6 (Miocene, Öhningen, Germany). – Heer 1865: 380; Scudder
1891: 591.
Trichius lugubris. – Heer 1883: 405; Scudder 1891: 591; Handlirsch 1907: 841; Frentzen 1927: 141; Krell 2000a:
887.
Gnorimus lugubris. – Kolbe 1931: 117; Kolbe 1932: 150
page 24
Trichius amoenus Heer
Trichius amoenus Heer, 1847: 74, pl. 7 (unknown locality). – Bronn 1849: 624; Stitzenberger 1851: 100; Giebel
1852b: 652; Giebel 1856: 30; Heer 1865: 380 (deducing: Switzerland); Heer 1883: 405; Scudder 1891:
591; Handlirsch 1907: 841; Frentzen 1927: 141; Krell 2000a: 887.
Trichius rotundatus Heer
Trichius rotundatus Heer, 1862: 82, pl. 6 (Miocene, Öhningen, Kesselstein, Germany). – Scudder 1891: 591;
Handlirsch 1907: 841; Krell 2000a: 887.
Trichius unifasciatus Heer
Trichius (Trichius) unifasciatus Heer, 1862: 82, pl. 6 (Miocene, Öhningen, Kesselstein, Germany). – Scudder
1891: 592; Handlirsch 1907: 842; Kolbe 1931: 117; Krell 2000a: 887.
Subfamilia Valginae
Valgus oeningensis Heer
Valgus oeningensis Heer, 1862: 82, pl. 6 (Miocene, Öhningen, Kesselstein, Germany). – Heer 1865: 378, 380;
Heer 1883: 404-406; Scudder 1891: 594; Handlirsch 1907: 842; Frentzen 1927: 141; Kolbe 1931: 117;
Krell 2000a: 887.
Subfamilia Euchirinae
Cheirotonus otai Ueda
Cheirotonus otai Ueda, 1989: 107 (Miocene, Tottori, Japan). – Krell 2000a: 887.
Subfamilia Cretoscarabaeinae Nikolajev, 1995b: 147 (transl. p. 185)
Genus Cretoscarabaeus Nikolajev
Cretoscarabaeus Nikolajev, 1995b: 147 (transl. p. 186), type genus (type species by original designation:
Cretoscarabaeus gibbosus Nikolajev). – Nikolaev 1999: 178; Krell 2000a: 887; Nikolaev 2002: 54
(transl. p. 281).
Cretoscarabaeus gibbosus Nikolajev
Cretoscarabaeus gibbosus Nikolajev, 1995b: 147 (transl. p. 186) (L Cretaceous, Baysa, Russia). – Krell 2000a:
887.
Cretoscarabaeus lentiginosus Nikolajev
Cretoscarabaeus lentiginosus Nikolajev, 1995b: 148 (transl. p. 188) (L Cretaceous, Baysa, Russia). – Krell
2000a: 887.
Genus Cretorabaeus Nikolajev
Cretorabaeus Nikolajev, 1995b: 149 (transl. p. 189) (type species by original designation: Cretorabaeus
elongatus Nikolajev). – Nikolaev 1999: 178; Krell 2000a: 887.
Cretorabaeus elongatus Nikolajev
Cretorabaeus elongatus Nikolajev, 1995b: 149 (transl. p. 189) (L Cretaceous, Baysa, Russia). – Krell 2000a:
887.
Subfamilia Lithoscarabaeinae Nikolajev, 1992: 76 (transl. p. 96)
Genus Lithoscarabaeus Nikolajev
page 25
Lithoscarabaeus Nikolaev, 1992: 78 (transl. p. 97), type genus (type species by original designation:
Proteroscarabaeus baissensis Nikritin). – Nikolaev 1999: 178; Krell 2000a: 888; Nikolaev 2002: 54
(transl. p. 281).
Lithoscarabaeus baissensis (Nikritin) Nikolaev
Proteroscarabaeus baissensis Nikritin, 1977: 124 (1991: 168) (L Cretaceous, Baysa, Russia). – Hong & Wang
1990: 115.
Proteroscarabaeus daissensis. – Morón 2003: 12
Lithoscarabaeus baissensis. – Nikolaev 1992: 78 (transl. p. 97); Krell 2000a: 888.
Familiae Incertae Sedis
Genus Aliscarabaeus Hong
Aliscarabaeus Hong, 1983: 5 (type species by original designation: Aliscarabaeus granulatus Hong). – Hong
1985: 37 (Alscarabaeus, lapsus calami); Krell 2000a: 888.
Aliscarabaeus granulatus Hong
Aliscarabaeus granulatus Hong, 1983: 5, 13 (Miocene, Shanwang, China). – Hong 1985: 37f, pl. 16; Krell
2000a: 888.
Genus Avitortor Ponomarenko
Avitortor Ponomarenko, 1977b: 42 (1991b: 54) (originally classified as belonging to Gyrinidae) (type species by
original designation: Avitortor primitivus Ponomarenko). – Schlüter 1987: 132; Nikolaev 1992: 80
(transl. p. 101) (Scarabaeidae); Nikolaev 1999: 178; Krell 2000a: 888; Nikolaev 2002: 54 (transl. p.
281).
Avitortor primitivus Ponomarenko
Avitortor primitivus Ponomarenko, 1977b: 42 (1991b: 55) (L Cretaceous, Baysa, Russia). – Nikolaev 1992: 80
(transl. p. 101) (Scarabaeidae); Krell 2000a: 888.
Avitortor leptoscelis (Nikritin) Nikolaev
Geotrupoides leptoscelis Nikritin, 1977: 122 (1991: 165) (L Cretaceous, Baysa, Russia). – Hong 1984: 171;
Hong & Wang 1990: 116 (G. ceptoscelis, lapsus calami); Hong 1992: 416. Avitortor leptoscelis. – Nikolaev 1992: 80 (transl. p. 101); Krell 2000a: 888.
Genus Cretoserica Nikolajev
Cretoserica Nikolajev, 1998a: 83 (transl. p. 519) (type species by original designation: Cretoserica latitibialis
Nikolajev), incertae subfamiliae. – Nikolaev 1999: 178; Krell 2000a: 888; Nikolaev 2002: 54 (transl. p.
281).
Cretoserica latitibialis Nikolajev
Cretoserica latitibialis Nikolajev, 1998a: 83 (transl. p. 520) (L Cretaceous, Baysa, Russia). – Krell 2000a: 888.
Genus Hongscarabaeus Krell
Hongscarabaeus Krell, 2000a: 888
Proscarabaeus Hong, 1982: 126 (type species by original designation: Proscarabaeus brunneus Hong). – Xiao
et al. 1994: 81; Nikolaev 1998b: 30; preoccupied by Proscarabaeus Schrank, 1781 (Coleoptera
Meloidae).
Hongscarabaeus brunneus (Hong) Krell
Proscarabaeus brunneus Hong, 1982: 126 (L Cretaceous, Jiuquan, China).
page 26
Hongscarabaeus brunneus. – Krell 2000a: 888.
(Genus Hybosorites Nikolajev)
Hybosorites Nikolajev, 1996: 98 (type species by original designation: Hybosorites fissuratus Nikolajev). –
Nikolaev 1999: 178; Krell 2000a: 888.
(Hybosorites fissuratus Nikolajev)
Hybosorites fissuratus Nikolajev, 1996: 98 (L Cretaceous, Baysa, Russia). – Krell 2000a: 888.
Doubtful (only one elytron).
Genus Mesoscarabaeus Hong
Mesoscarabaeus Hong, 1982: 123 (type species by original designation: Mesoscarabaeus corneus Hong). –
Nikolaev 1998b: 30; Krell 2000a: 888.
Mesoscarabaeus corneus Hong
Mesoscarabaeus corneus Hong, 1982: 124 (L Cretaceous, Jiuquan, China). – Krell 2000a: 888.
Mesoscarabaeus morulosus Hong
Mesoscarabaeus morulosus Hong, 1982: 125 (L Cretaceous, Jiuquan, China). – Krell 2000a: 888.
(Genus Opiselleipon Bode)
(Opiselleipon Bode, 1953: 230; pl. 11 (type species by monotypy: Opiselleipon gravis Bode); not assigned to a
family by the author. – Crowson 1981: 664 (“Possible Scarabaeoids”), 679 (“Scarabaeoid-like
appearance”); Carpenter 1992: 324 (“Little-known genus”, suborder and family uncertain); Scholtz &
Chown 1995a: 358; Krell 2000a: 888.
(Opiselleipon grave Bode)
Opiselleipon gravis Bode, 1953: 230, pl. 11 (L Jurassic, Hondelage near Braunschweig, Germany). – Scholtz
1990: 1029; Carpenter 1992: 324; Krell 2000a: 888.
Tribus Holcorobeini Nikolajev, 1992: 81 (transl. p. 102)
Genus Holcorobeus Nikritin
Holcorobeus Nikritin, 1977: 127 (1991: 172) [“Holcoribeus”, lapsus calami, since all species are named
Holcorobeus], type genus (type species: Holcorobeus vittatus Nikritin). – Crowson 1981: 667
(“modern-looking Scarabeoids”); Schlüter 1987: 133; Nikolaev 1998b (classified as Aclopinae, but
without giving an explanation); Nikolaev 1999: 178; Jarzembowski 2000: (8); Krell 2000a: 889;
Nikolaev 2002: 54 (transl. p. 281).
Holcoribeus Nikritin (see above). – Ponomarenko 1990: 68; Carpenter 1992: 300 (“Family assignment
doubtful”).
Antemnacrassa Gomez Pallerola 1979: 340 (syn. Nikolaev 1992: 82 who called it Antemnocrassa).
Since the differences between Holcorobeus and Geotrupoides (sensu Nikolaev) are fluid, the undiscussed transference of some species from the latter to the first genus by Nikolaev (1992) is questionable.
Subgenus Holcorobeus Nikritin
Holcorobeus Nikritin, 1977: 127 (1991: 172) (“Holcoribeus”; see above); subgenus introduced by Nikolaev
1992: 82 (transl. p. 103); Krell 2000a: 889.
Holcorobeus (Holcorobeus) atrosulcatus Nikolajev
Holcorobeus (Holcorobeus) atrosulcatus Nikolajev, 1992: 84 (transl. p. 106) (L Cretaceous, Baysa, Russia). –
Krell 2000a: 889.
page 27
Holcorobeus (Holcorobeus) evittatus Zhang
Holcoribeus evittatus Zhang, 1992a: 334 (L Cretaceous, Laiyang Formation, China). – Krell 2000a: 889.
Legs lacking exept for hind coxae and femora. On this basis, the classification is not reliable.
Holcorobeus (Holcorobeus) incertus Ponomarenko
? Holcoribeus incertus Ponomarenko 1990: 68, pl. 6 (L Cretaceous, Turga Formation, Russia; Gidari Formation:
Karabon, Pavlovka, Russia).
Holcorobeus (Holcorobeus) incertus. – Nikolaev, 1992: 83 (transl. p. 104); Krell 2000a: 889.
Holcorobeus (Holcorobeus) latitibialis Nikolajev
Holcorobeus (Holcorobeus) latitibialis Nikolajev, 1992: 84 (transl. p. 106) (L Cretaceous, Baysa, Russia). –
Krell 2000a: 889.
Holcorobeus (Holcorobeus) longipes (Ponomarenko) Nikolaev
Geotrupoides longipes Ponomarenko, 1986: 98, pl. 8 (L Cretaceous, Myangad, Mongolia). Holcorobeus (Holcorobeus) longipes. – Nikolaev, 1992: 83 (transl. p. 104); Krell 2000a: 889.
Holcorobeus (Holcorobeus) maculatus Nikolajev
Holcorobeus (Holcorobeus) maculatus Nikolajev, 1992: 85 (transl. p. 107) (L Cretaceous, Baysa, Russia). –
Krell 2000a: 889.
Holcorobeus (Holcorobeus) monreali (Gomez Pallerola) Nikolaev
Antemnacrassa monreali Gomez Pallerola 1979: 340 (L Cretaceous, Montsec, Spain). – Gomez Pallerola 1986:
720f.
Holcorobeus (Holcorobeus) monreali. – Nikolaev, 1992: 83 (transl. p. 104); Krell 2000a: 889.
Holcorobeus (Holcorobeus) nigrimontanus Nikolajev
Holcorobeus nigrimontanus Nikolajev, 1992: 83 (transl. p. 105) (U Jurassic, Karatau, Kazakhstan). – Krell
2000a: 889.
(Holcorobeus (Holcorobeus) nigrovittatus Nikolajev)
Holcorobeus nigrovittatus Nikolajev, 1992: 86 (transl. p. 108) (L Cretaceous, Baysa, Russia). – Krell 2000a: 889. Doubtful (only one elytron).
Holcorobeus (Holcorobeus) picturatus Nikritin
Holcorobeus picturatus Nikritin, 1977: 129 (1991: 175) (L Cretaceous, Baysa, Russia). Holcorobeus (Holcorobeus) picturatus. – Nikolaev 1992: 83 (transl. p. 104); Krell 2000a: 889.
Holcorobeus (Holcorobeus) punctatus (Ponomarenko) Nikolaev
Proteroscarabaeus punctatus Ponomarenko, 1986: 97f, pl. 8 (L Cretaceous, Gurvan-Erenii-Nuru, Mongolia).
Holcorobeus (Holcorobeus) punctatus. – Nikolaev, 1992: 83 (transl. p. 104); Krell 2000a: 889.
Holcorobeus (Holcorobeus) sulcatus (Nikritin) Nikolaev
Geotrupoides sulcatus Nikritin, 1977:120 (1991: 163) (L Cretaceous, Baysa, Russia). – Hong 1984: 170; Hong
& Wang 1990: 116 (G. sdulcatus, lapsus calami); Hong 1992: 416; Ren et al. 1995: 436; Zhang 1997:
90.
Holcorobeus (Holcorobeus) sulcatus. – Nikolaev 1992: 83 (transl. p. 104); Krell 2000a: 889.
Holcorobeus (Holcorobeus) vitimensis (Nikritin) Nikolaev
Geotrupoides vitimensis Nikritin, 1977: 123 (1991: 167) (L Cretaceous, Baysa, Russia). – Hong & Wang 1990:
116; Hong 1992: 416.
Holcorobeus vitimensis. – Nikolaev, 1992: 83 (transl. p. 104); Krell 2000a: 889.
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Holcorobeus (Holcorobeus) vittatus Nikritin
Holcorobeus vittatus Nikritin, 1977: 128 (1991: 173) (L Cretaceous, Baysa, Russia). – Nikolaev 1992: 83 (transl.
p. 104); Krell 2000a: 890.
Holcoribeus vittatus. – Carpenter 1992: 301; Zhang 1992a: 334f.
Subgenus Mesaclopus Nikolajev
Mesaclopus Nikolajev, 1992: 86 (transl. p. 108) (type species by original designation: Holcorobeus mongolicus
Nikolajev). – Krell 2000a: 890.
Holcorobeus (Mesaclopus) mongolicus Nikolajev
Holcorobeus (Mesaclopus) mongolicus Nikolajev, 1992: 86 (transl. p. 108) (U Jurassic-L Cretaceous, Khutel-
Khara, Mongolia). – Krell 2000a: 890.
presumed Scarabaeoidea:
Genus Tetragonides Bode
Tetragonides Bode, 1953: 225, pl. 11 (type species by monotypy: Tetragonides magnus Bode) – Carpenter 1992:
326 (“Little-known genus.”, suborder and family uncertain); Krell 2000a: 890.
Tetragonides magnus Bode
Tetragonides magnus Bode, 1953: 225 (L Jurassic, Hondelage, Germany) (“Die Gestalt erinnert an die der (?)
Lucaniden.”). – Carpenter 1992: 326; Krell 2000a: 890.
Fossil taxa later synonymised with extant taxa Genus Magniscarabaeus Hong & Wang
Magniscarabaeus Hong & Wang in Hong, 1983: 5 (type species by original designation: Magniscarabaeus
furvus Hong & Wang). – Hong & Wang 1986: 6.
= Anomala Samouelle, 1819. – Zhang 1989: 162; Krell 2000a: 890. Genus Obitiscarabaeus Hong
Obitiscarabaeus Hong, 1983: 6 (type species by original designation: Obitiscarabaeus jiaoyanshanensis Hong).
– Hong 1985: 40.
= Geotrupes Latreille, 1796. – Zhang 1989: 152; Krell 2000a: 890. Genus Palaeocopris Pierce
Palaeocopris Pierce, 1946a: 130 (type species by monotypy: Palaeocopris labreae Pierce, 1946). – Edmonds
1972: 855 (“morphological resemblance to this genus with Phanaeus”).
Paleocopris. – Halffter 1959: 176.
= Phanaeus MacLeay, 1819. – Miller et al. 1981: 627; Krell 2000a: 890. Canthon simplex antiquus Pierce, 1946a: 120 (Pleistocene, Rancho La Brea tar pits, Los Angeles, U.S.A.). –
Spohn 1973: 52.
Boreocanthon simplex antiquus. – Halffter 1959: 176.
= Canthon (Boreocanthon) simplex LeConte. – Miller et al. 1981: 626; Stock & Harris 1992: 84; Krell 2000a:
890.
Canthon praticolus vetustus Pierce, 1946a: 122 (Pleistocene, Rancho La Brea tar pits, Los Angeles, U.S.A.). –
Sphon 1973: 52.
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Boreocanthon praticola vetustus. – Halffter 1959: 176.
= Canthon (Boreocanthon) praticola LeConte. – Miller et al. 1981: 626; Stock & Harris 1992: 84; Krell 2000a:
890.
Nomina nuda and unpublished names
Genus Melolonthidium Phillips, 1871: 174; M Jurassic, Stonesfield Slate, England; without indication. – Brodie
1873: 23; Goss 1879: 143; Handlirsch 1907: 572; Krell 2000a: 890. Genus Pseudopentodon Filhol, 1892: 2 (without indication). – Krell 2000a: 890.
Pseudopentodon blanchardi Filhol, 1892: 2 (Eocene-Oligocene, Quercy, France; without indication). –
Handschin 1944: 2, 19, 21; Handschin 1950: 138 (Pseudopantodon); Théodoridès 1952: 47; Krell
2000a: 890.
Ichnotaxa, fossil nests and broodballs of Scarabaeoidea
Ichnogenus Coprinisphaera Sauer
Coprinisphaera Sauer, 1955: 123 (type species by monotypy: Coprinisphaera ecuadoriensis Sauer). –
Häntzschel 1962: W189; Häntzschel 1975: W52; Martinez 1982: 48; Genise 1993: 50; Donovan 1994: 209;
Genise & Cladera 1995: 78; Buatois et al. 1998: 226; Genise & Laza 1998: 220; Genise 1999: 110; Genise et al.
2000: 49ff; Krell 2000a: 890.
Cangabola Lengerken, 1955: 937 (type species by monotypy: Cangabola ecuadoriensis Sauer)
Coprinsphaera. – Sauer 1956: 550; Halffter & Matthews 1966: 154; Retallack 1991: 182, 296; Duringer et al.
2000: 264.
Devincenzia Roselli, 1939: 81 (type species by monotypy: Devincenzia murguiai Roselli; preoccupied by
Devincencia Kraglievich, 1932 – see Genise & Bown 1994: 109). (syn.) Devicenzichnus Roselli, 1976: 167 (type species by monotypy: Devicenzichnus murguiai). – Martinez 1982: 48; Buatois et al. 1998: 226. (syn.)
Devinzenzichnus. – Genise 1993: 50 (synonym of Coprinisphaera); Genise & Laza 1998: 213. (syn.)
Coprinisphaera ecuadoriensis Sauer
„„bolas”. – Bruet 1950.
Coprinisphaera ecuadoriensis Sauer, 1955: 123 (U Pleistocene, northern Ecuador). – Sauer 1956: 555; Sauer
1959: 119ff; Häntzschel 1962: W189; Häntzschel 1975: W52; Martinez 1982: 48; Genise & Bown 1994:
109; Genise 1999: 110; Krell 2000a: 890.
Cangabola ecuadoriensis. – Lengerken 1955: 937.
Coprinsphaera ecuadoriensis. – Sauer 1956: 550.
Coprinsphaera ecuadorensis. – Halffter & Matthews 1966: 154.
Coprinisphaera ecuadorensis. – Genise 1993: 50.
Coprinisphaera frenguellii Genise & Bown
Coprinisphaera frenguellii Genise & Bown, 1994: 110 (Miocene, Estancia Ana Maria, Prov. Santa Cruz,
Argentina). – Buatois et al. 1998: 226; Krell 2000a: 891.
Coprinisphaera murguiai (Roselli) Genise & Bown
Devincenzia murguiai Roselli, 1939: 81 (U Cretaceous, Colonia, Uruguay). – Genise 1983: 50.
Devincenzichnus murguiai. – Roselli 1976: 166f; Genise 1993: 50; Genise & Bown 1994: 109.
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Devicenzichnus murguiai. – Roselli 1976: 167; Martinez 1982: 48.
Devincenzichnus murguiae. – Retallack 1990: 219.
Devinzenzichnus murgiiae. – Donovan 1994: 209.
Coprinisphaera murguiai. – Genise & Bown 1994: 109; Krell 2000a: 891.
Ichnogenus Eatonichnus Bown, Hasiotis, Genise, Maldonaldo & Browers Eatonichnus Bown, Hasiotis, Genise, Maldonaldo & Browers, 1997: 52 (type species by original designation:
Xenohelix utahensis Gilliland & LaRocque; “might be constructions of dung beetles”). – Genise &
Bown 1998: 222; Genise & Laza 1998: 214, 218; Genise 1999: 110; Genise et al. 2000: 53f, 57
(“possible dung beetle nest”); Krell 2000a: 891.
Eatonichnus claronensis Bown, Hasiotis, Genise, Maldonaldo & Browers Eatonichnus claronensis Bown, Hasiotis, Genise, Maldonaldo & Browers, 1997: 54 (Eocene, J.G. Eaton locality,
Utah, U.S.A.). – Genise & Bown 1998: 222; Krell 2000a: 891.
Eatonichnus utahensis (Gilliland & La Rocque) Bown, Hasiotis, Genise, Maldonaldo & Browers
Xenohelix? utahensis Gilliland & La Rocque, 1952: 502, pl. 59 (Eocene, Bald Knoll Canyon, Utah, U.S.A.). –
Kilpper 1962: 56.
Eatonichnus utahensis. – Bown et al. 1997: 52; Genise & Laza 1998: 222; Krell 2000a: 891.
Ichnogenus Fontanai Roselli
Fontanai Roselli 1939: 79 (type species by monotypy: Fontanai kraglievichi Roselli). – Genise & Laza 1998: 213 (“is a dung beetle brood mass”); Genise et al. 2000: 49, 51, 53f, 58f; Krell 2000a: 891. Fontanaichnus Roselli 1976: 167 (type species by monotypy: Fontanaichnus kraglievichi). – Retallack 1990: 219; Donovan 1994: 209; Buatois et al. 1998: 226. (syn.)
Fontanai kraglievichi Roselli
Fontanai kraglievichi Roselli, 1939: 79 (U Cretaceous-L Tertiary, Asencio Formation, Uruguay). – Genise 1993: 53 (classification impossible); Genise & Bown 1994: 112 (“insufficiently known to meaningfully interpret
them”); Krell 2000a: 891.
Fontanaichnus kraglievichi. – Roselli 1976: 167; Retallack 1990: 219 (“Likely fossil nests of dung beetles”);
Donovan 1994: 209 (“Possible fossil nests of dung beetles”).
Ichnogenus Madinaichnus Roselli
Madinaichnus Roselli, 1987: 23 (type species by monotypy: Madinaichnus larranagai). – Genise & Laza 1998: 213; Buatois et al. 1998: 226; Genise 1999: 110; Genise et al. 2000: 54; Krell 2000a: 891. Genise (1999) supposed Madinaichnus, Martinezichnus, and Microicoichnus to be synonymous to Coprinisphaera.
Madinaichnus larranagai Roselli
Madinaichnus larranagai Roselli, 1987: 23 (U Cretaceous – L Tertiary, Uruguay). – Krell 2000a: 891.
Ichnogenus Martinezichnus Roselli
Martinezichnus Roselli, 1987: 22 (type species by monotypy: Martinezichnus francisi Roselli). – Genise &
Cladera 1995: 78; Genise & Laza 1998: 213; Buatois et al. 1998: 226; Genise 1999: 110; Genise et al. 2000: 54; Krell 2000a: 891.
Martinezichnus francisi Roselli
Martinezichnus francisi Roselli, 1987: 22 (Cretaceous, Asencio Formation, Uruguay?). – Genise & Cladera 1995: 78f; Krell 2000a: 891.
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Ichnogenus Microicoichnus Roselli
Microicoichnus Roselli, 1987: 49 (type species by monotypy: Microicoichnus lafurcadai Roselli). – Buatois et al.
1998: 226; Genise & Laza 1998: 213; Genise 1999: 110; Genise et al. 2000: 54; Krell 2000a: 891.
Microicoichnus lafurcadai Roselli
Microicoichnus lafurcadai Roselli, 1987: 49 (U Cretaceous – L Tertiary, Uruguay). – Krell 2000a: 891.
Ichnogenus Monesichnus Roselli
Monesichnus Roselli, 1987: 39 (type species by monotypy: Monesichnus ameghinoi Roselli). – Laza et al. 1994;
Bown et al. 1997: 55f (“might be constructions of dung beetles”); Buatois et al. 1998: 226; Genise & Laza 1998: 218; Genise et al. 2000: 49, 51, 53f, 58f; Krell 2000a: 891.
Monesichnus ameghinoi Roselli
Monesichnus ameghinoi Roselli, 1987: 39 (U Cretaceous – L Tertiary, Asencio Formation, Uruguay). – Laza et
al. 1994; Bown et al. 1997: 56; Genise & Laza 1998; Genise 1999: 110; Krell 2000a: 891.
Ichnogenus Pallichnus Retallack
Pallichnus Retallack, 1984: 580 (type species by original designation: Pallichnus dakotensis Retallack). –
Genise 1993: 50; Genise & Bown 1994: 109; Buatois et al. 1998: 226; Genise 1999: 110; Genise et al. 2000: 53;
Krell 2000a: 891.
Pallichnus dakotensis Retallack
Pallichnus dakotensis Retallack, 1984: 581 (Oligocene, South Dakota, U.S.A.). – Genise 1993: 50; Krell 2000a: 891.
Ichnogenus Scaphichnium Bown & Kraus
Scaphichnium Bown & Kraus, 1983: 106 (type species by original designation: Scaphichnium hamatum Bown &
Kraus). – Buatois et al. 1998: 226; Genise & Laza 1998: 214, 218; Krell 2000a: 892.
Scaphichnium hamatum Bown & Kraus
Scaphichnium hamatum Bown & Kraus, 1983: 106 (L Eocene, Willwood formation, Wyoming, U.S.A.). –
Hasiotis et al. 1993 (classified as traces of Scarabaeoidea); Bown et al. 1997: 56; Genise & Laza 1998: 222; Genise 1999: 110; Krell 2000a: 892.
1unnamed trace fossils:
Africa: Pleistocene, Rutana, Burundi: “loges fossiles des coprophages” (Basilewsky 1951).
Pliocene (2-3 Myr), Makapansgat Limeworks, South Africa (Kitching 1980). Pliocene (3,46-3,76 Myr), Laetoli, Tanzania: structure resembling a dung ball of Heliocopris (Sands 1987: 423) (= Coprinisphaera sp. according to Genise et al. 2000).
Asia: U Miocene, Dhok Patan Formation, Pakistan: Coprinisphaera sp. (Retallack 1991: 182, 296). North America: Pliocene, Jalisco, México: “bola-nido” (Morón Ríos 1984: 27).
L Eocene, Golden Valley Formation, North Dakota, U.S.A.: striae in coprolites “appear to have been made by
scatophagous beetles” (Jepsen 1963: 680).
U Cretaceous, Two Medicine Formation, Montana, U.S.A.: coprolites (Chin & Gill 1996).
South America: U Pleistocene, Buenos Aires and Lujan; Santa Fé, Tezanos Pinto; Tucumán, Tafí del
Valle: Coprinisphaera sp. (Genise et al. 2000).
1 more possible traces of dung beetles are compiled by Retallack (1990: 223)
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U Pleistocene, Sopas, Uruguay: Coprinisphaera sp. (Genise et al. 2000).
L Pleistocene, Los Galpones, Argentina: “bolas” of cf. Canthon (Frenguelli 1938a, b; Coprinisphaera sp.
according to Genise et al. 2000).
U Pliocene (- L Pleistocene), Chapadmalal and San Andrés Formation, Buenos Aires, Argentina:
Coprinisphaera sp. (Genise et al. 2000).
U Pliocene, Piquete Formation, Sierra de Vaqueros, Salta, Argentina: cf. Phanaeus, Megathopa; Coprinisphaera sp. (Alonso et al. 1982; Genise et al. 2000).
U Miocene, Las Flores, San Juan, Argentina: “nidos de escarabeidos” (Contreras 1996; Coprinisphaera sp.
according to Genise et al. 2000).
U Miocene, Collón-Curá Formation, Río Negro and Neuquén, Argentina: “peras” of cf. Megathopa, cf.
Onthophagus, etc. (Frenguelli 1938a, 1939; Laza 1986a; Coprinisphaera sp. according to Genise et al. 2000).
U Miocene, Paso de las Carretas, San Luis, Argentina: “?nidos? de escarabideos” (Pascual & Bondesio 1981:125; Coprinisphaera sp. according to Genise et al. 2000).
U Miocene, Andalhualá, Catamarca, Argentina: Coprinisphaera sp. (Genise et al. 2000).
U Miocene, Cerro Azul, La Pampa/Buenos Aires, Argentina: Coprinisphaera sp. (Genise et al. 2000).
U Miocene-L Pliocene, Monte Hermoso, Buenos Aires, Argentina: Coprinisphaera sp. (ichnofossil) (Genise et
al. 2000).
U Oligocene, Colhue-Huapi Formation, Argentina (U): Coprinisphaera sp. (Laza 1986b, Genise et al. 2000).
Oligocene, Ludados lustrosos, Mendoza, Argentina: Coprinisphaera sp. (Genise et al. 2000).
U Eocene, Musters Formation, Chubut Province, Argentina: Coprinisphaera sp. (Genise et al. 2000).
M Eocene, Laguna del Mate, Chubut Province, Argentina: “similar fossils of Coprinisphaera” (Retallack 1990:
218).
L Eocene, Santa Cruz, Punta Casamayor, Los Leones, and Chubut, Comodoro Rivadavia, Argentina: “peras” of
cf. Megathopa and cf. Phanaeus (Frenguelli 1938a, b; Laza 1986b; Coprinisphaera sp. according to Genise et
al. 2000).
Palaeocene, Rio Chico, Chubut, Argentina: Coprinisphaera sp. (Genise et al. 2000).
Palaeocene (?), Comallo, Rio Negro, Uruguay: fossil nests (Retallack 1990: 217).
Tertiary (Oligocene?), Chubut, Cañadón del Loco, Argentina: “nidos de escarabeidos” (Frenguelli 1941).
Antarctica: Eocene, Isla Marambio (Seymour), La Meseta formation, Peninsula Antartica: “un molde
interno de nido de escarabeido integrante de la subtribu Phanaeina (Scarabaeinae)” (Laza & Reguero 1990;
Coprinisphaera sp. according to Genise et al. 2000).
Other fossils originally described as Lamellicornia
Ichnogenus Fictovichnus Johnston, Eberth & Anderson
Fictovichnus Johnston, Eberth & Anderson, 1996: 521 (type species by original designation: Fictovichnus
gobiensis Johnston, Eberth & Anderson) (pupal chambers of Scarabaeidae, Tenebrionidae, or Curculionidae),
containing the ichnospecies F. gobiensis and F. parvus Johnston, Eberth & Anderson, 1996. - Genise et al.
2002a: (probably [junior] synonym of Rebuffoichnus Roselli, 1987 [Curculionidae]); Genise et al. 2002b (pupal chamber of Curculionidae).
Genus Oryctites Oppenheim (not valid)
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Oryctites Oppenheim, 1888: 238. – Ponomarenko 1971: 72 (transl. p. 67): Adephaga incertae sedis; Krell 2000a:
892.
Since Oppenheim only indicates a fossil species of Oryctes Illiger, 1798 (“Für Oryctes spricht noch der
Umstand...”), the name Oryctites Oppenheim is not valid according Art. 20 ICZN.
(Oryctes) fossilis (Oppenheim) Houlbert
Oryctites fossilis Oppenheim, 1888: 238, pl. 31, fig. 2 (Jurassic, U Malm, Solnhofen, Germany). – Scudder 1891:
216; Meunier 1898b: 112, 133, pl. 23; Ponomarenko 1971: 67: Adephaga incertae sedis.
Oryctes fossilis. – Houlbert 1915: 64 (“Handlirsch assure que cette identification est inexacte”); Krell 2000a: 892.
? Cerambycinus fossilis (Cerambycidae). – Handlirsch 1906: 547 (“gewiss kein „Oryctes?”).
(Oryctes pluto Weyenbergh)
Oryctes pluto Weyenbergh 1869a: 282, pl. 27 (Jurassic, U Malm, Solnhofen, Germany). – Weijenbergh 1869b*:
2[232]; Weijenbergh 1874: 102 or 109; Goss 1879: 148; Winkler 1878: 95; Scudder 1891: 216; Winkler
1896: 313 (“Coléoptère sp. Meunier”); Meunier 1897, pl. 10 (“Aucun organe de ce fossile n?étant
suffisamment conservé, on doit se borner à dire que cette empreinte est celle d?un grand coléoptère.”);
Houlbert 1915: 64 (“n?est pas déterminable”); Krell 2000a: 884.
Oryctes grandis Weijenb. – Weijenbergh 1869b [nomen nudum].
Pseudohydrophilus avitus (Heyden, 1847) Handlirsch, 1906 (Blabera avita Heyden, 1847: 100). – Handlirsch
1906: 544; belongs to Hydrophilidae.
Genus Progeotrupes Oppenheim
Progeotrupes Oppenheim, 1888: 239 (type species by monotypy: Progeotrupes jurassicus Oppenheim). – Broili
1921: 675; Broili 1924: 697; Laurentiaux 1953: 476; Krell 2000a: 892. – belongs to Blattodea (see below).
Progeotrupes jurassicus Oppenheim
Progeotrupes jurassicus Oppenheim, 1888: 239, pl. 31 (Jurassic, U Malm, Solnhofen, Germany). – Scudder
1891: 217; Meunier 1898b: 109 (“C?est avec le plus grand doute, que je place ce coléoptère près des
Lamellicornes du genre Geotrupes”); Handlirsch 1906: 549, pl. 45 [1907] (“Hat sicher nichts mit Geotrupes zu
tun.”); Ponomarenko 1971: 80 (transl. p. 75) (Blattodea); Krell 2000a: 893.
Geotrupes jurassicus. – Meunier 1898b: 133.
Genus Prosynactus Bode
Prosynactus Bode, 1953: 224 (type species by monotypy: Prosynactus scissus Bode) (“Ein Typ, der an manche (?) Lucaniden erinnert.”); Krell 2000a: 893. – belongs to Tachypachidae (see below).
Prosynactus scissus Bode
Prosynactus scissus Bode, 1953: 224, pl. 11 (Lias, Beienrode, Germany). – Ponomarenko 1992: 180 (belongs to Coleoptera Tachypachidae); Krell 2000a: 893.
Ichnogenus Rebuffoichnus Roselli
Rebuffoichnus Roselli, 1987: 24 (type species by monotypy: Rebuffoichnus casamiquelai). – Buatois et al. 1998:
226; Genise & Laza 1998: 213; Genise et al. 2000: 54; Krell 2000a: 892; Genise et al. 2002a: (probably [senior] synonym of Fictovichnus Johnston, Eberth & Anderson, 1996 [Curculionidae]); Genise et al. 2002b (probably
pupal chamber of Curculionidae, but separate from Fictovichnus).
Rebuffoichnus casamiquelai Roselli
2 There are two printings with slightly different arrangement of the text.
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Rebuffoichnus casamiquelai Roselli 1987: 24 (U Cretaceous – L Tertiary, Asencio Formation, Uruguay). –
Genise et al. 1999: 29 (pupal chamber of Coleoptera – Curculionidae, Scarabaeidae or Tenebrionidae); Krell 2000a: 892; Genise et al. 2002a: 230f; Genise et al. 2002b.
Genus Scarabaeides Germar
Scarabaeides Germar, 1839: 218 (type species by monotypy: Scarabaeides deperditus Germar). – Giebel 1846:
148; Bronn 1848b: 1119; Bronn 1849: 625; Weyenbergh 1873: 240; Krell 2000a: 893. Mesobelostomum Haase, 1890: 21 (Heteroptera: Belostomatidae); synonym of Scarabaeides because of identical
type species.
Scarabaeides deperditus Germar
“Scarabaeus (der Gattung Phileurus nahe stehend)”. – Germar 1837b: 422.
Scarabaeides deperditus Germar, 1839: 218 (Malm, Solnhofen, Germany). – Brodie 1845: 109; Bronn 1848b:
1119; Bronn 1849: 625; Giebel 1852a: 207; Quenstedt 1852a: 313 (“Wäre es ein Käfer, so sollte man doch Reste
von Flügeldecken erwarten.”), 786 (index: “Scarabaeoides depertitus” [lapsus calami]; Quenstedt 1852b: pl. 5;
Giebel 1856: 36 (“Die Stellung des Thieres bleibt völlig zweifelhaft.”); Hagen 1862: 110 (“grosse Aehnlichkeit
mit Belostoma; ein Käfer ist es bestimmt nicht.”), 113 (ad Hemiptera); Weyenbergh 1869a: 249 (ad Hemiptera); Goss 1879: 147; Scudder 1886: 73 (“Belostomidae”); Scudder 1891: 176 (“Belostomatidae”); Zittel 1895: 504; Krell 2000a: 893.
Scarabaeoides deperditus. – Weijenbergh 1869b: [233] (Heteroptera, Geocorises).
Scaraboeides deperditus, Scarabeides deperditus. – Brodie 1873: 21.
Scarabaeus deperditus. – Weyenbergh 1874: 88f, 103 or 109; Winkler 1878: 96; Scudder 1891: 176. Hydrophilus deperditus. – Weyenbergh 1873: 239; Weyenbergh 1874: 84 or 89, 103 or 109; Winkler 1878: 96; Winkler 1896: 309.
Belostomum deperditus. – Assmann 1877: 192.
Belostoma deperditum. – Deichmüller 1886: 61; Meunier 1896: 93; Winkler 1896: 309.
Belostoma deperdita. – Scudder 1891: 176.
Mesobelostomum deperditum. – Haase 1890: 21; Handlirsch 1906: 637, pl. 51 (1907); Handlirsch 1925: 210; Müller 1989: 247.
Genus Tetragonotrachelus Bode
Tetragonotrachelus Bode, 1953: 224 (type species by monotypy: Tetragonotrachelus gracilis Bode) (“Äußerlich
etwas an den Lucanidentyp erinnernd.”); Krell 2000a: 893.
synonym of Prosynactus Bode, 1953: 224 (Coleoptera Tachypachidae) (Ponomarenko 1992: 181).
Tetragonotrachelus gracilis Bode
Tetragonotrachelus gracilis Bode, 1953: 224, pl. 11 (Lias, Beidenrode, Germany); Krell 2000a: 893. Prosynactus gracilis. – Ponomarenko 1992: 181 (belongs to Coleoptera: Tachypachidae).
Genus Troxites Goldenberg
Troxites Goldenberg, 1854: 36 (type species by monotypy: Troxites germari Goldenberg). – Giebel 1856: 36;
Goldenberg 1867: 7; Gerstaecker 1868: 289 (“Zugehörigkeit zu der Ordnung der Coleopteren zumindest sehr
zweifelhaft”); Scudder 1879b: 19 (“curculionid”); Scudder 1881-1885: 794 (“dürfte eine fossile Frucht sein”); Scudder 1886: 73 (“probably a fruit”); Krell 2000a: 893.
Troxites germari Goldenberg
page 35
Troxites germari Goldenberg, 1854: 36 (Carboniferous; Steinkohlen, Altenwald near Saarbrücken). – Giebel
1856: 36; Goldenberg 1867: 7; Brodie 1873: 27; Goldenberg 1873: 7, pl. 2; Goldenberg 1877: 50; Scudder 1879b: 17; Goss 1880: 297; Scudder 1891: 97f (“Probably not an insect.”); Handlirsch 1906: 342 (refers to the next author?s interpretation); Schlechtendal 1912: 159ff, pl. 1 (“unzweifelhaft das Bruchstück einer Crustacee etwa ? Arthropleura armata, vielleicht haben wir in ihm ein Stückchen Fühler oder Bein”); Handlirsch 1919: 81 (“Arthropleura-Fühlerglied”); Handlirsch 1922: 221 (“Arthropleurarest”); Waterlot 1934: 262 (“fossile à affinité douteuse”); Krell 2000a: 893.
Xyloryctes planus Fri? (ichnospecies)
Xyloryctes planus Fri?, 1877: 16, pl. 3 (Carboniferous, Ny?an near Plzen, Czechia; Oberhohndorf, Germany). –
Scudder 1879b: 17; Scudder 1891: 98 (“coleopteroid”); Handlirsch 1905: 328 (“Die Deutung dieser
Gebilde als Bohrlöcher von Coleopteren erscheint mir gewagt.”); Handlirsch 1919: 590 (arthropod
origin questionable); Handlirsch 1922: 219 (no insect); Krell 2000a: 893.
Xyloryctes septarius Fri? (ichnospecies)
Xyloryctes septarius Fri?, 1877: 15, pl. 3 (U Carboniferous, Swinná near Radnitz, Bohemia, Czechia). – Scudder
1879b: 17; Scudder 1891: 98 (“coleopteroid”); Handlirsch 1905: 337 (not from an insect); Handlirsch
1919: 590 (arthropod origin questionable); Handlirsch 1922: 219 (no insect); Krell 2000a: 893. Fri? never claimed that these ichnofossils were lebensspuren of Scarabaeoidea. However, he gave them the ichnogeneric name Xyloryctes. I suppose that he did not realize that this name was already proposed for extant Dynastinae from America (Xyloryctes Hope, 1837).
Tertiary and Mesozoic lagerstätten with fossils of Scarabaeoidea
Pliocene (> 1,8 Myr)
Europe: Willershausen, Harz, northern Germany (U): Aphodius (Acrossus) rufipes (L.), Aphodiinae spec.,
Mimela spec., Oryctes nasicornis L. (Gersdorf 1971), Coprinae sp., Cetoniinae sp. (reliable) (Gersdorf 1976); Lucanus cervicalis (= cervus (L.)) (Schweigert 2003).
Boulder-Formation, Mundesley, England: cfr. Copris lunaris (Lyell 1840: 175).
Kisatibi formation, Georgia (L?): Copris (Copris) kartlinus (Kabakov 1988).
Africa: Makapansgat, South Africa: ichnofossils of coprophagans (Kitching 1980). Kouba Olanga, palaeo-lake Chad system, Chad (5 Myr): fossil brood-balls of dung beetles (Duringer et al. 2000).
Pliocene (3,46-3,76 Myr), Laetoli, Tanzania: Coprinisphaera sp, (Sands 1987: 423; Genise et al. 2000: 55).
North America: Jalisco, México: ichnofossils of coprophagans (Morón Ríos 1984). South America: Piquete Formation, Sierra de Vaqueros, Salta, Argentina (U): ichnofossils of coprophagans (“bolas”) (Alonso et al. 1982; Coprinisphaera sp. according to Genise et al. 2000: 55).
Chapadmalal and San Andrés Formation (transition to Pleistocene), Buenos Aires, Argentina: Coprinisphaera sp.
(Genise et al. 2000).
Transition Mio/Pliocene: Europe: Simetite (Sicilian resin): (Scarabaeus bilobus L.) (=
Caelosis biloba) (Ferrara 1805: 136 [wrong determination; Krell 1996: 18]; Kohring & Schlüter 1989: 45). Cinérites de Varennes, France: (Melolonthites laterosinuatus) (Piton & Théobald 1935).
North America: Beaufort Formation, Prince Patrick Island, Canada (3-5,7 Myr): Aegialia sp.
(Matthews 1976, Matthews et al. 1990).
page 36
South America: Monte Hermoso, Buenos Aires, Argentina (U Miocene-L Pliocene): Coprinisphaera
sp. (ichnofossil) (Genise et al. 2000: 55).
Miocene (> 5 Myr)
Europe: Montagne d?Andance, Ardeche, France (U, Turolian): Miocenidorcus andancensis, cf. Geotrupes
stercorarius, Thyphoeus sp. [= Typhaeus sp.], cf. Amphimallon ater (Riou 1988: 103ff, 1999).
Habichtswald near Kassel, Germany (L): cfr. Aphodius (Landgrebe 1843); doubtful (legs and antennae absent). Öhningen, Badenia, Germany (M, Sarmat, Serravallian, 14,5 Myr): Coprologus gracilis, Geotrupes germari, Gymnopleurus deperditus, G. rotundatus, G. sisyphus, Copris druidum, C. subterraneus, Onthophagus
bisontinus, O. crassus, O. ovatulus, O. prodromus, O. urusheeri, Oniticellus amplicollis, Aphodius anteactus,
A. brevipennis, A. meyeri, Hybosorus lividus, Rhizotrogus longimanus, (Melolonthites aciculatus), M.
3deperditus, (M. lavateri), (M. obsoletus), Lepitrix germanica, Anomala fugax, Serica minutula, Gnorimus aedilis, G. lugubris, Trichius amoenus, T. rotundatus, T. unifasciatus, Valgus oeningensis, Glaphyrus
antiquus, Pentodon proserpinae (Heer 1847, 1862, 1865, Frentzen 1927), Lucanus sp. (Serres 1829: 235),
Geotrupes sp., cf. Trichius sp. (obviously not considered by Heer) (Scudder 1895b: 120). Parschlug, Styria, Austria (U): (Melolonthites kollari), (M. parschlugianus) (Heer 1847).
Mok?ina (Krottensee) near Kynšperk n. O., Cypris Shale, Cheb Basin, Bohemia, Czechia (L, Burdigalian):
(Melolontha solitaria) (Novák, 1878).
Gabbro, Livorno, Italy (L, Messinian): Aphodius bosniaskii (Handlirsch 1908).
Euboea (Evia), Greece (L): Geotrupes spec. (Bachmayer et al. 1971).
Asia: Dhok Patan Formation, Pakistan (U): Coprinisphaera sp. (Retallack 1991: 182, 296).
Shanwang, Shandong, China (M, Serravallian-Langhian; 15,5-17 Myr): Geotrupes jiaoyanshanensis,
Orrhodomala protista, Onthophagus sp., Phalops sp., Scelocopris enertheus, Adoretus recticlypeus, A. rhinus, Aliscarabaeus granulatus, Anomala amblobelia, A. brachytarsia, A. endoxa, A. eversa, A. furva, A.
lochmocola, A. orcina, A. palaeobrunnea, A. punctulata, A. scia, A. ursa, Zhangsunia extumida, Genonota
mochthera, G. scola, Holotrichia cressona, H. spatha, Macronota shandongiana, Mioserica margelis,
Petraeianus ruderatus, Petulantis yimengensis, Serrulus sinicus (Hong 1983, 1985, Zhang et al. 1994, Zhang
1989).
Noto, Takaya, Yanagida Formation, Japan (M): Heliocopris antiquus, Phyllopertha? sp. (Fujiyama 1968).
Seki, Masaragawa Formation, Sado Island, Japan (L): Aphodius ? sp. (doubtful, only one elytron) (Fujiyama 1985).
Tottori Group, Kokufu Town, Japan (M): Cheirotonus otai (Ueda 1989).
Africa: Lake Victoria, Kenya: Anachalcos mfwangani (Mfwangano Island), Copris leakeyorum (Rusinga
Island), Metacatharsius rusingae (Rusinga I.), Rutelinae, Cetoniinae (Paulian 1976). North-America: Ruby River, Montana, U.S.A. (U): Rutelinae (tentatively) (Zuidema 1950: 121). Clarkia, Idaho, U.S.A. (L, 17-20 Myr): Lucanus sp., Osmoderma sp., Geotrupes sp. (Lewis 1985: 251ff).
Meighen Island, Canada (U): Aegialia sp. (Matthews 1977).
Dominican Amber: Tyrannasorus rex (Ratcliffe & Ocampo 2001), Procoilodes adrastus (Ocampo 2002);
Ceratocanthidae, Lucanidae, Canthidium sp., Rhyparus, Termitodius (Poinar 1992: 151, 285; Poinar & Poinar 1999: 165, 205; Wu without date: 187).
3 The Melolontha listed by Keferstein (1834: 331) likely refers to one of these Melolonthites species.
page 37 South-America: Río Negro, Collón-Curá Formation, and Neuquén, Argentina (U): ichnofossils of
coprophagans (Megathopa and Onthophagus), Coprinisphaera sp. (Frenguelli 1938a; Genise et al. 2000: 55). San Luis, Paso de las Carretas, Argentina (U): ichnofossils of scarabs, Coprinisphaera sp. (Pascual & Bondesio 1981: 125; Genise et al. 2000: 55).
Las Flores, San Juan, Argentina (U): “nidos de escarabeidos” (Contreras 1996; Coprinisphaera sp. according to Genise et al. 2000: 55).
Andalhualá, Catamarca and Cerro Azul, La Pampa/Buenos Aires, Argentina (U): Coprinisphaera sp.
(ichnofossil) (Genise et al. 2000: 55).
Santacrucian, Pinturas, Patagonia, Argentinia (L): Coprinisphaera frenguellii (ichnofossil) (Genise & Bown 1994).
Oligocene (> 24 Myr)
Europe: Rott, Siebengebirge, Germany (U, Chattian, 25 Myr): Geotrupes rottensis, (Onitis) magus,
Mioochodaeus proaevus, Onthophagus statzi, Aphodius helvolus, A. krantzi, A.. schlickumi, Melolontha spec., Mimela rhenana, Anomala primigenia, A. thetis, A. tumulata, Maladera spinitibialis, Pentodon bellerophon, Ceruchites hahnei (Germar 1849, Heyden 1862, Heyden & Heyden 1866, Statz 1952, Krell 1990);
presumably: Geotrupes vetustus, (Platycerus) sepultus (Germar 1837a).
Enspel, Westerwald, Germany (U): Scarabaeoidea (Wedmann 1998, 2000: 47).
Greith (Kohlengrube), Hohenrhone, Switzerland: Melolontha greithiana (Heer 1847).
Brunstatt, Alsatia, France (M): Ammoecius ? sp. [= Aphodius (Ammoecius) sp.] (Förster 1890: 102). Aix en Provence, France (U, Stampian): Onthophagus luteus, Geotrupes atavus (Oustalet 1874), Sisyphus sp., Melolontha sp., Pachypus sp. (Serres 1829: 221f), Cetonia sp. (“resembling C. hirtellus” and “like C. stictica, Fab.”) (Curtis 1829: 295); ? Geotrupes, Melolontha (Rhizotrogus) [single elytron] (Hope 1847). Célas, Gard, France (L, Sannoisian): Aphodius theobaldi (Théobald 1937).
Puy Saint-Jean, Auvergne, France (U; ca. 22 Myr [-L Miocene?]): “Helicopris sp. (?)” [= Heliocopris] (Rudel 1940: 15).
Armissan, Aude, France: Ateuchites grandis (Meunier 1898a).
Barachek Creek, Pozhar Region, Russia: Platycerus zherichini (Nikolaev 1991).
Asia: Kudia River, Primorye Province, Russia (L): Melolonthites interemtus (Cockerell 1926). North America: John Day Series (lower division), Oregon, U.S.A. (U): Passalus indormitus
(Cockerell 1927a).
Ruby Paper Shales, Montana, U.S.A. (U): “scarab” (Becker 1961: 38, pl. 31).
Florissant, Colorado, U.S.A. (L): Trox antiquus, Ataenius patescens, A. restructus, Oxyomus nearcticus,
Aphodius aboriginalis, A. florissantensis, A. granarioides, A. inundatus, A. laminicola, A. mediaevus, A.
praeemptor, A. senex, A. shoshonis, Phyllophaga disrupta, (P. extincta), Macrodactylus pluto, M. propheticus, (Diplotaxis aurora), (Diplotaxis) simplicipes, Listrochelus puerilis, Anomala exterranea, A. scudderi,
Miolachnosterna tristoides, Hoplia striatipennis, (Serica) antediluviana, (S.) cockerelli, Ligyrus compositus, (L.) effetus, Strategus cessatus, Amphicoma defuncta, Ceruchites fuchsii, (Lucanus fossilis) (Cockerell 1927b, Wickham 1909, 1910, 1911, 1912, 1913a, b, 1914a, b). South Dakota: Pallichnus dakotensis (ichnofossil) (Retallack 1984). South America: Colhue-Huapi Formation, Argentina (U): Coprinisphaera sp. (ichnofossil) (Laza 1986b, Genise et al. 2000: 54).
page 38
Santa Cruz and Chubut, Argentina: ichnofossils of coprophagans, Coprinisphaera sp. (ichnofossil) (Frenguelli 1938a, b; Genise et al. 2000: 54).
Ludados lustrosos, Mendoza, Argentina: Coprinisphaera sp. (ichnofossils) (Genise et al. 2000: 54).
Eocene – Oligocene: Europe: Quercy near Toulouse, France (U Eocene to U Oligocene):
Aphodius sp., Pseudopentodon blanchardi nom. nud. (Filhol 1892; Handschin 1944, 1950). Glarus shales, Switzerland (U Eocene - L Oligocene): “HANNETON. Scarabaeus” [= Melolontha s.l. spec.] (Bertrand 1763: 259).
Eocene (> 37 Myr)
Europe: Gurnet Bay, Bembridge Limestone, Isle of Wight (U): Dorcus sp. (Brodie 1845: 47; Goss 1878: 325; Woodward 1878, 1879).
Ku?lín (Kutschlin) near Bílina (Bilin), Bohemia, Czechia (U, U Priabonian): Dorcus (Eurytrachelus) primigenius, Phaeochrous tertiarium (Deichmüller 1881).
Bournemouth, Bagshot Beds, England (M, U Ypresian - Lutetian): Pelidnotites atavus (Cockerell 1920). Geiseltal near Halle, Germany (M, M - U Geiseltalian): Eophyllocerus scrobiculatus, E. glaucinus (Haupt 1950). Messel near Darmstadt, Germany (M, L Geiseltalian): (Geotrupes messelensis), (Gymnopleurus eocaenicus)
(Meunier 1921), Lucanidae (Lutz 1988b), Protognathus spielbergi (Chalumeau & Brochier 2001), many more species (Krell, in prep.).
Eckfelder Maar, Eifel, Germany (M., U Geiseltalian): Scarabaeidae, Melolonthinae (Lutz 1988a).
Baltic Amber: Paleognathus succini (Waga 1883); Succiniplatycerus berendti (Zang 1905), Dorcasoides bilobus (Motschulsky 1856); Ataenius europaeus (Quiel 1910); Saprosites succini (Zang 1905); Aphodius, Trox (Helm
1886: 277), some more species (Krell, in prep.).
Bognor Regis, London Clay, England (L): Saprosites cascus, Onthophagus spec. (Britton 1960: 36f). North America: British Columbia, Canada (M): Ninemile creek: (Trox oustaleti) (Scudder 1879a, Handlirsch 1910); Princeton: Scarabaeidae (Wilson 1977). Green River, Colorado/Wyoming, U.S.A. (L – M): Aegialia rupta (Scudder 1890), Phyllophaga avus (Cockerell 1921, 1925); unidentified specimen of Scarabaeidae (Grande 1984: 249). Willwood Formation, Slick Creek and Fifteenmile Creek, Bighorn Basin, Wyoming, U.S.A. (L): Scaphichnium hamatum (Bown & Kraus 1983: 106; Hasiotis et al. 1993).
Holcomb Property, Henry County, Tennessee, U.S.A. (L): (Melolonthites collinsi) (Wickham 1929).
J.G. Eaton Locality, Claron Formation, Kane County, Utah, U.S.A. (L): Eatonichnus claronensis, E. sp.
(ichnofossils) (Bown et al. 1997).
Golden Valley Formation, North Dakota, U.S.A. (L): striae in coprolites “appear to have been made by
scatophagous beetles” (Jepsen 1963: 680).
Bald Knoll Canyon, Colter and Claron Formation, Sevier and Garfield County, Utah, U.S.A.: Eatonichnus
utahensis (Bown et al. 1997).
South America: Musters Formation, Chubut Province, Argentina (U): Coprinisphaera sp. (Genise et
al. 2000: 54).
Laguna del Mate, Chubut Province, Argentina (M): “similar fossils of Coprinisphaera” (Retallack 1990: 218).
Santa Cruz, Punta Casamayor, Los Leones, and Chubut, Comodoro Rivadavia, Argentina (L): “peras” of cf.
Megathopa and cf. Phanaeus (Frenguelli 1938a, b; Laza 1986b; Coprinisphaera sp. according to Genise et al.
2000: 54).
page 39
Antarctica: Isla Marambio: ichnofossils of coprophagans (Laza & Roguero 1990). Palaeocene (> 54 Myr)
Europe: Menat, Puy-de-Dôme, France (U, Thanetian, 56 Myr): Bolboceras inermis, Aphodius charauxi
(Piton 1940).
South America: Rio Chico, Chubut, Argentina: Coprinisphaera (Genise et al. 2000: 54).
Tertiary (without specification): Europe: Nogent le Rotrou (“Süßwasserquarz”), France: Anomalites fugitivus (Fri? 1885).
MESOZOICUM
Upper Cretaceous (> 65 Myr)
Asia: Kzyl-Zhar, Southern Kazakhstan (Turonian): Cretaesalus ponomarenkoi (Nikolajev 1993).
Zheijang Province, Ho Jia, Zhu Xi, China (Turonian?-Santonian): Prionocephale deplanate (Lin 1980).
Africa: Orapa, Botswana (Cenomanian-Coniacian): Scarabaeidae (Photo) (McKay & Rayner 1986: 11f). North America: Two Medicine Formation, Montana, U.S.A. (Campanian): Coprolithes and fossil soil with nesting traces (Chin & Gill 1996).
South America: (U Cretaceous – L Tertiary) Asencio Formation, Cantera Maldonado, Nueva Palmira, Departamento Colonia, Uruguay: Coprinisphaera murguiai, Fontanai kraglievichi, Madinaichnus sp.,
Martinezichnus francisi, Microicoichnus sp., Monesichnus ameghinoi (ichnotaxa) (Roselli 1939, Genise & Cladera 1995, Genise & Laza 1998).
Lower Cretaceous (> 98 Myr)
Europe: Las Hoyas, Cuenca, Spain (Barremian): Scarabaeidae (Martínez-Delclós 1989: 76). Montsec, Spain (U Berriasian - L Valanginian): Scarabaeidae (Martínez-Delclós & Ruiz de Loizaga 1993);
Holcorobeus monreali (Gomez Pallerola 1986, Nikolaev 1992).
Asia: Lebanon (Amber, Aptian): termitophilous species of Scarabaeoidea (Crowson 1981: 669, 673); “Corythodérine” (= Corythoderini) (Paulian 1988: 394).
Baysa, Zazin series, Transbaykalia, Russia (Valanginian-Hauterivian): Avitortor leptoscelis, A. primitivus,
Cretogeotrupes convexus, Cretohybosorus buryaticus, C. striatulus, Holcorobeus atrosulcatus, H. latitibialis,
H. maculatus, H. nigrovittatus, H. picturatus, H. sulcatus, H. vitimensis, H. vittatus, Proteroscarabaeus yeni, (P. magnus), (P. nikritini), Lithoscarabaeus baissensis, Cretaegialia aphodiiformis, C. rhypariformis,
(Cretanaides trogopterus), Cretobolbus rohdendorfi, (Hybosorites fissuratus), Cretomelolontha
transbaikalica, Cretoserica latitibialis, Cretoscarabaeus gibbosus, C. lentiginosus, Cretorabaeus elongatus
(Ponomarenko 1977b, Nikritin 1977, Nikolaev 1992, 1994, 1995b, 1996, 1998a, 1999). Pad Semen, Argun series, Transbaykalia, Russia: Proteroscarbaeus yeni, Lithoscarabaeus baissensis (Nikritin
1977), Prototrox transbaikalicus (Nikolaev 2000a).
Leskovo, Transbaykalia, Russia: Lithanomala crassa, L. oblonga, L. sibirica (Ponomarenko 1990, Nikolaev
1992).
Turga, Transbaykalia, Russia: Holcorobeus incertus (Ponomarenko 1990).
Gurvan-Erenii-Nuru, Mongolia: Holcorobeus punctatus (Ponomarenko 1986).
Shar Tolgoy, Bayan-Hongor Aymag, Mongolia: Cretocoma tologoica (Nikolaev 2002)
Myangad, Mongolia: Holcorobeus longipes (Ponomarenko 1986).
Bon-Tsagan, Mongolia (U Neocomian – L Aptian): Cretochodaeus mongolicus, (C. striatus) (Nikolaev 1995a).
Chifeng, Inner Mongolia, China: (Geotrupoides) fortus (Ren et al. 1995).
page 40
Laiyang Formation, Shandong Province, China (Barremian-Hauterivian): Geotrupoides nodosus (Hong & Wang
1990), Holcorobeus evittatus (Zhang 1992a), Proteroscarabaeus yeni, P. baissensis (Grabau 1923; Hong &
Wang 1990). There has been some discussion about whether this formation belongs to Lower Cretaceous, to Upper Jurassic (Zhang 1992b), or to both (see Lin 1994). Here I adopt the view of Lin (1994) and Hong (1998).
Jiuquan Basin, Chijinpu Formation, Gansu Province, China (Valanginian; according to Hong 1998): Mesoscarabaeus corneus, M. morulosus, Hongscarabaeus brunneus (Hong 1982).
4Yixian, Hebei Province, China (Tithonian?-Berriasian): Geotrupoides songyingziensis (Hong 1984).
Jilin Province, China: Geotrupoides jiaoheensis (Hong 1992).
South America: Santana, Brazil: Scarabaeidae (Grimaldi & Maisey 1990: 7; Grimaldi 1991: 400 [colour photo]), some more specimens (Krell, in prep.).
Transition Upper Jurassic/Lower Cretaceous: Khutel-Khara; Mongolia:
Holcorobeus (Mesaclopus) mongolicus (Nikolaev 1992).
Upper Jurassic (Malm, > 144 Myr)
Europe: Lithographischer Schiefer, Bavaria, Germany (L Tithonian): (Geotrupoides lithographicus)
(Eichstätt) (Deichmüller 1886), (Cetonia defossa), (Oryctes pluto) (Solnhofen) (Weyenbergh 1869a).
Asia: Karatau, Kazakhstan (Oxfordian – Kimmeridgian): Holcorobeus nigrimontanus (Nikolaev 1992).
Shara-Teg Mountain, Gov?-Altai? Aymag, Mongolia: Paralucanus mesozoicus (Nikolajev 2000b).
Middle Jurassic (Dogger, > 163 Myr)
No reliable records: “Melolonthidium” of Philips (1871) from the Stonesfield Slate, England, was
not described and has to be confirmed. It might refer to the doubtful Melolontha (or “Melolontha ?”) from the
same lagerstätte classified and figured by Murchison (1845: 68, 107, pl. 4). Lower Jurassic (Lias, > 188 Myr)
No reliable records: The family identity of Opiselleipon grave Bode and Tetragonides magnus
Bode (Lias ε from Hondelage, Germany), Aphodiites protogaeus Heer (Lower Lias, Schambelen, Switzerland) and Melolontha ? sp. (Lower Lias, Cracombe, England; Brodie 1845: 101, pl. 9) is untraceable. Carboniferous (> 286 Myr)
No records: Troxites germari Goldenberg from the hard coals of Saarbrücken, Germany, and the ichnotaxa Xyloryctes planus Fri? and X. septarius Fri? from the Upper Carboniferous of Germany and Bohemia, are no fossils or traces of Scarabaeoidea.
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