Department of Entomology The Natural History ...

Coleopterists Society Monograph Number 5:120–143. 2006.

FOSSIL RECORD AND EVOLUTION OF SCARABAEOIDEA (COLEOPTERA: POLYPHAGA) FRANK-THORSTEN KRELL Department of Entomology The Natural History Museum Cromwell Road, London SW7 5BD, UK [email protected] Abstract The fossil record of scarab beetles (Coleoptera: Scarabaeoidea) is reviewed and its relevance for the reconstruction of the evolutionary history of the taxon is explored. After a discussion of the different kinds of preservation of scarab fossils, including preservation of color, the richest scarab deposits are identified. From the fossil record, the minimum age of Scarabaeoidea is determined as 152 myr. To develop an idea about the appearance of the first scarab, the ground-pattern of Scarabaeoidea is reconstructed for the first time on the basis of published phylogenetic analyses. Extinct scarabaeoid family-group taxa are described and discussed. Cretocomini and Cretoglaphyrini are upgraded to subfamily rank for reasons of consistency. Most of their diagnostic character states are controverted or not polarized. The minimum age of extant scarabaeoid family-group taxa is deduced from the fossil record. Ancient feeding habits, particularly the development of coprophagy, are discussed. The youngest extinct species described are from the Pleistocene, whereas extant species have been recorded since the Pliocene.

It is obvious, however, that we need to examine as objectively as possible any evidence, recent or fossil, that will shed light on the movement and development of different groups of organisms. —Henry F. Howden, 1972 Scarab beetles (Scarabaeoidea or Lamellicornia) are well-represented in the fossil record. Around 1,000 fossil specimens are known, with the first having been discovered in Linneán times (Bertrand 1763). There have been 244 species formally described, 204 of which show characters indicating that they belong to the Scarabaeoidea, beyond body or elytron shape. All species are compiled in a recently published catalog (Krell 2000) which is updated in the Appendix of this paper. Fossil insects continue to be described in large numbers. New scarabaeoid fossils have been recorded or formally named every year since the publication of the catalog (Chalumeau and Brochier 2001; Nikolajev 2000a, b, 2002, 2004, 2005a–c; Ocampo 2002; Ratcliffe and Ocampo 2001; Ratcliffe et al. 2005; Schweigert 2003; Wappler 2003). In this review I present and discuss major aspects of the fossil record of Scarabaeoidea and its applicability and drawbacks for reconstructing the evolutionary history of this taxon. Preservation of Scarab Fossils The usefulness of fossils for scientific study and the feasibility of classification depend on the kind of preservation. Grimaldi and Engel (2005) described the various preservation types in detail. Most scarabaeoid fossils show detailed structures because without them they would not be identifiable as scarabaeoids. In current paleoentomology, single elytra or body silhouettes without appendages 120

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or hind wings are rarely assigned to a beetle family (with some exceptions, e.g., Nikolajev 1996, 2004). Three-dimensional Fossils The most impressive fossils are the three-dimensional, almost undistorted amber inclusions, the best of which show an almost full set of external characters (although the original chitin has been diagenetically altered and is no longer present; Stankiewicz et al. 1998). Often, however, the inclusions are partly covered by dirt, verlumung (milky impurities), other specimens, or optical distortions caused by the amber itself. Scarabaeoidea in amber are known from the Cretaceous (Lebanon–Crowson 1981; Myanmar–Krell, Ballerio, Azar, Buckley, unpubl.), Eocene (Baltics–Spahr 1981), and Miocene (Dominican Republic– Ratcliffe and Ocampo 2001; Ocampo 2002). I am currently studying the known Lebanese, Myanmar, and Baltic specimens. More recent inclusions of scarabaeoids have become known from copal which can be of recent to Pleistocene or Miocene age (Schlu¨ter and Gnielinski 1987). This material has rarely been published, probably only by Giebel (1862), who considered an undescribed ‘Serica sp.’ to be an amber inclusion but seems more likely to be Apogonia sp. (Melolonthinae) (Quiel 1911). Three-dimensional ‘petrified’ scarabaeoids in numbers are known only from the Miocene of Rusinga and Mfangano Island in Lake Victoria, Kenya (Leakey 1952). The high quality of these calcified specimens allows assignment to a genus, and three of them were described later by Paulian (1976) as dung beetle species of Anachalcos Hope, Copris Mu¨ller, and Metacatharsius Paulian. Single, threedimensional scarabs were found in the Eocene London Clay of Bognor Regis, England (the pyritized Saprosites cascus Britton (Aphodiinae); Britton 1960), in the Eocene-Oligocene phosphorites of Quercy in France (possible dynastine and questionable aphodiine; Handschin 1950), and in the Tertiary fresh-water quartz of Nogent le Rotrou in France (Anomalites fugitivus Fricˇ, 1885; a possible melolonthine or ruteline). Recently, a calcified specimen of Dynastinae was discovered in Pliocene Laetoli in Tanzania (Krell, unpublished). The youngest calcified 3-D scarabaeoid is probably a geotrupid from the Holocene travertine of Weimar, Germany (Puhlmann et al. 1991). Tar pits have revealed a wealth of insects, although they are mostly in fragments. The most famous are Rancho La Brea in Los Angeles County, where the preservation of Pleistocene chitin has even been proven (Stankiewicz et al. 1997). Five extant, two extinct, and two unidentifiable (though named) Scarabaeidae species were found there (Miller et al. 1981; Stock and Harris 1992). Scarabaeoidea are also known from the tar pits of Talara, Peru (Pleistocene–Churcher 1966), Las Breas de San Felipe, Cuba (MiocenePleistocene–Valde´s 1999), and Fyzabad, Trinidad (Upper Pleistocene-Holocene–Blair 1927; Kugler 1927). Similar quality of preservation is found in numerous Quaternary sites up to the present (see below). Two-dimensional Fossils Most insect fossils are flat compressions or impressions (Engel and Grimaldi 2005). If the embedding sediment is fine-grained, structures are preserved in great detail, often in higher resolution than in three dimensional petrified fossils. The best examples are the Eocene oil shales of Messel (Germany), the Oligocene sapropel/diatomite couplets of Florissant (see below), the Lower Cretaceous Jehol

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Biota in China (Zhang and Zhang 2003; Zhang et al. 2003), and the fine-grained limestone of the Brazilian Cretaceous lagersta¨tte of Santana (Grimaldi 1991), the scarabaeoids of which have not yet been studied. In Florissant and Santana, microbial mats protected sediment and deceased biota from the effects of hydrodynamic and biological disruption (Harding and Chant 2000) leading to excellent preservation. Flat (or flattened) beetles often changed their relative dimensions or the topographic relations of body parts, either through external pressure by the embedding of sediment or before by internal bloating and disintegrating during decay (Martıńez-Delclo`s and Martinell 1993; Lutz 1990; Duncan et al. 2003; Smith et al. 2006). Color Preservation Fossil beetles with colors preserved, although by far not the oldest colored fossils (Parker 2005), are among the most impressive invertebrate fossils. Such specimens are known from the German lagersta¨tten Rott (Oligocene–Hellmund 1988; not scarabaeoid), Geiseltal (Eocene–Haupt 1950; the melolonthine Eophyllocerus is grey to dark blue, with green and violet in E. glaucinus Haupt), and thousands of specimens (including many Scarabaeoidea) from Eocene Messel (Parker and McKenzie 2003; Krell, unpublished). Many Messel beetles show bright metallic colors that have been identified as original and are caused by multilayer reflectors of the preserved cuticle (Parker and McKenzie 2003). The only colorful scarabaeoids published so far are Protognathinus spielbergi Chalumeau and Brochier (Schaal and Ziegler 1992; Chalumeau and Brochier 2001) and another unidentified stag beetle (Lausch et al. 2000). The Richest Scarab Lagersta¨tten Scarabaeoidea fossils and ichnofossils have been described or recorded from 100 Mesozoic and Tertiary (Krell 2000 and Appendix) and countless Quaternary localities, most of which either yielded only a few species, or their biota have not yet been comprehensively studied. The overwhelming majority of fossil scarabaeoids are from deposits of the northern hemisphere. The most diverse Mesozoic fauna has been described from Russia, China, Kazakhstan, and Mongolia, but Santana in Brazil (Grimaldi 1991) will take a strong position as soon as the scarabaeoids are studied because of the high quality of preservation. South America is the center of scarabaeoid ichnofossils (fossil dung balls) (Genise et al. 2000; Krell 2000), but Santana remains the only locality known for body fossils. Africa is gaining in importance with ongoing studies of Tertiary deposits. Mesozoic and Tertiary Scarabaeoidea are missing from Australia (Jell 2004), but Porch (in press) found extant Aphodiinae species in late Pleistocene deposits in Victoria and Tasmania. A ‘hemi-fossil’ reported from Australia by Karube (2002) is a degraded elytra, probably of the extant native stag beetle Phalacrognathus muelleri (MacLeay), found in a swampy area near Cairns at a depth of 30 cm. Antarctica has yielded only Eocene ichnofossils of coprophagans (Laza and Roguero 1990). Speciose fossil scarabaeoid fauna has been recorded from only a few lagersta¨tten, all of which have an exceptional quality of fossil preservation: Baissa/Baysa, Zaza Formation (Lower Cretaceous), Transbaikalia, Buryat Republic, Russia Age: Probably pre-Barremian (.130 myr) (Zherikhin et al. 1999). General description: Zherikhin et al. 1999. Remarks: Fine-grained sediments deposited in an intermontane lake. A very rich site. More than 20,000 specimens of the 250,000 fossil insects housed in the Palaeontological Institute in Moscow are from here (Zherikhin 1999; Zherikhin et al.


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1999). 30 described scarabaeoid species (Krell 2000 and Appendix). Oldest records for Bolboceratidae, Geotrupidae, and Scarabaeidae: Aegialiinae and Melolonthinae. Messel (Middle Eocene), Germany Age: 47 myr (Franzen 2005). General description: Schaal and Ziegler 1992. Remarks: Oil shales deposited in a maar lake (Harms et al. 2004). Just over 3% of ca. 11,500 preserved Messel insect fossils in Senckenberg are Scarabaeoidea. Some 420 scarabaeoid specimens have been preserved, deposited mainly in the Senckenberg research station at Messel itself, rendering Messel the richest scarabaeoid lagersta¨tte. At least 20 species can be reliably differentiated (Krell, unpublished), and one was described from a photograph: Protognathinus spielbergi Chalumeau and Brochier (Lucanidae: Chiasognathinae). Oldest record for Scarabaeidae: Hopliinae (together with a specimen from Baltic amber, both unpublished). I am currently working on a monograph on the scarabaeoid fauna of Messel. Eckfelder Maar (Middle Eocene), Germany Age: 44 myr (Mertz et al. 2000). General description: Lutz 2003. Remarks: Oil shales deposited in a maar lake; 196 scarabaeoid specimens, assigned to 8+ unnamed species of Lucanidae, Geotrupidae, and Scarabaeidae: Aclopinae, Aphodiinae, Melolonthinae, Rutelinae, and Cetoniinae (Wappler 2003). Oldest record for Cetoniinae. Florissant (Lower Oligocene), Colorado, U.S.A. Age: 34 myr (Evanoff et al. 2001). General description: Meyer 2003. Remarks: Lakedeposited shales and tuffs (volcanic ash with self-sedimented diatom mats, and pumice); 32 described scarabaeoid species (see Meyer 2003). Rott (Upper Oligocene), Germany Age: 25 myr (Mo¨rs 1995; Koenigswald 1996). General description: Koenigswald 1996. Remarks: Oil shales deposited in a lake, possibly a caldera; 16 described, one unnamed scarabaeoid species (see Krell 2000). ¨ hningen (Middle Miocene), Germany O Age: 13.6–14.8 myr (Berger et al. 2005). Remarks: Molasse (detritus shed from the rising Alpine mountains); 33 described scarabaeoid species (see Krell 2000). All species were described by Oswald Heer between 1847 and 1865 and haven’t been revised since except for Frentzen’s (1927) cursory review of the fauna. Shanwang (Middle Miocene), Shandong, China Age: ,12–15 myr (Zhao et al. 2002). General description: Yang and Yang 1994; Sun 1995. Remarks: Diatomaceous shale deposited in a volcanically related lake basin; 27 described, two unnamed scarabaeoid species (see Krell 2000).

Minimum Age of Scarabaeoidea The fossil record can only define the minimum age of a taxon, because it is highly unlikely that first individuals of the stem species of a taxon were fossilized and that this stem species, which initially is likely to look similar to its sister species, has developed an autapomorphy that allows us to diagnose the species as stem species (i.e., an autapomorphy that is still diagnosable as autapomorphy of the extant taxon and preserved in the fossil). To reliably identify a scarabaeoid fossil, we need at least one of the two unequivocal scarabaeoid synapomorphies: (1) an antenna with lamellate club and (2) an intrinsic spring mechanism for unfolding the hind wings, recognizable in many fossils by a triangular, oval, or oblique sclerotized field proximal to the pinch between Radius Anterior (1 + 2) and the apical part of Radius Anterior 3 (or RA 3 + 4) (terminology after Kukalova´-Peck and Lawrence 1993) (Fig. 1). The meso- and metatibia of most Scarabaeoidea have transverse, setose or spinose keels on the outer side. This character has never been recognized as an

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autapomorphy of Scarabaeoidea, but it is lacking in virtually all other Coleoptera (Lawrence et al. 2000) and thus might serve as another relatively reliable diagnostic character. The oldest fossil apparently showing a scarabaeoid autamomorphy is Juraclopus rohdendorfi Nikolajev (Fig. 2) (Nikolajev 2004, 2005b) from the Upper Jurassic of Karatau, Kazakhstan (152–158 myr; Grimaldi and Engel 2005). The photograph of the fossil shows a structure on the left side of the head that is certainly a lamellate antenna. Two other species from the same lagersta¨tte, Antemnacrassa nigrimontana (Nikolajev 1992) and A. albosulcata Nikolajev 2004, show scarabaeoid tibiae with transverse keels. Reliable Scarabaeoidea from the Cretaceous are numerous (Krell 2000), but none of the pre-Karatau supposed scarabaeoid fossils show any diagnostic characters of the Scarabaeoidea (e.g., Aphodiites protogaeus Heer, Lower Jurassic, Fig. 3). In 1996, when checking the rich beetle collection from the Upper Triassic Molteno Formation, South Africa, deposited in the South African National Biodiversity Institute, Pretoria, I could not find a single scarab. In all other known Triassic lagersta¨tten of fossil insects, Scarabaeoidea are yet to be discovered (Krell 2000; Papier et al. 2005). Therefore, the minimum age of Scarabaeoidea is currently 152 myr. The Basal Scarab–Still a Mystery Since the ground-pattern (grundplan) of Scarabaeoidea has not been reconstructed in the phylogenetic literature, and Mesozoic scarabs already are morphologically diverse, we have to compile possible ground pattern character states to get an idea of what the first scarab might have looked like. Since sister species look very similar just after speciation, the habitus of members of the sister group of Scarabaeoidea might help. However, the sister group of Scarabaeoidea has not been reliably identified. The clade Hydrophiloidea + Histeroidea is a likely candidate, but its sister group relationship to Scarabaeoidea is weakly supported morphologically (Beutel and Leschen 2005) and molecular analyses are equivocal (Caterino et al. 2005). The hypothesis of a sister group relationship of Dascillidae and Scarabaeoidea as discussed by Crowson (1995) is not supported by the latest cladistic analyses (Grebennikov and Scholtz 2004; Beutel and Leschen 2005). Using the data from Beutel and Leschen (2005) and Hansen (1997a), I hypothesize the adult ground-pattern of the crown group Scarabaeoidea (below). It comprises unambiguous autapomorphies of superordinated groups (retained as plesiomorphies in Scarabaeoidea) and autapomorphies of Scarabaeoidea (marked by asterisks in the description below). To this, I added diagnostic characters given by Lawrence and Britton (1991) and extracted additional information from the internal phylogeny of Scarabaeoidea as presented by Scholtz et al. (1994) and Browne and Scholtz (1999) who consider Glaresidae to be the sister of the remaining Scarabaeoidea. Glaresidae possess three unique and seven controverted apomorphies: Radius Posterior 1 and RP3 + 4 reduced, 5 abdominal sternites; 10 antennomeres, ocular canthus present, mandibles with molar lobe and prostheca, true epipharynx present, foraminated tentorium, protrochantin concealed, and procoxae closed. The respective plesiomorphic states are included in the scarabaeoid ground-pattern. However, it remains incomplete and preliminary, since the data used were originally not defined and selected to reconstruct comprehensive ground-patterns.


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Fig. 1. Scarabaeoid wing from the Eocene oil shales of Messel (Melolonthinae). Arrow indicates the sclerotized field proximal to the pinch between Radius Anterior (1 + 2) and the apical part of Radius Anterior 3. (Inv. No. MeI5984 [Senckenberg]). Fig. 2. Juraclopus rohdendorfi Nikolajev from Upper Jurassic Karatau, Kazakhstan, the oldest scarabaeoid with a preserved lamellicorn antenna. Photo: Georgy Nikolajev. Fig. 3. Aphodiites protogaeus Heer from Lower Lias (Jurassic) of Schambelen, Switzerland. Long considered to be the oldest scarabaeoid fossil despite no diagnostic scarabaeoid characters present. From Heer (1865, pl. 8).

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Scarabaeoid Ground-Pattern. Clypeus enlarged (lateral margin typically more than 1.53 as long as frons anterior to eyes); labium enlarged, plate-like and strongly sclerotized; posterior labiohypopharyngeal complex strongly narrowed between mandibles and maxillae, with steeply ascending posterior part; no true epipharynx; mandibles without prostheca and molar area; 11 antennomeres; *antenna with a lamellate club; club with three pubescent segments, basal segments almost glabrous; ocular canthus absent; tentorium invaginated; accessory posterior pronotal ridge present; prothorax enlarged, modified for burrowing with large coxae; procoxal cavities open; protrochantin exposed; metacoxae without plates; *anterior border of hind wings with sclerotized field proximal to a pinch, as part of a spring folding mechanism; Radius Posterior 1 and 3 + 4 present; six or seven abdominal sternites; tergite 8 free, forming a pygidium. This is the baseline, and further reconstruction is currently unfounded. In any case, the ground-pattern of the first scarabaeoid stem-group species, i.e., the sister species of the first hydrophiloid/histeroid, would not have developed most of the scarabaeoid apomorphies but would be similar to ancestral hydrophiloid-histeroids and probably not distinguishable from them in the fossil record. A glaresid-like, burrowing beetle with full wing venation (thus probably larger than extant glaresids), primitive mandibles, six or seven abdominal sternites, and without an ocular canthus might be a possible approximation of the first scarab, but only if the stem species of the Glaresidae crown group existed shortly after the stem species of Scarabaeoidea. In case of extensive morphological evolution in a long stem line of Glaresidae before the last common ancestor of extant Glaresidae evolved, the first scarab could have looked considerably different from extant glaresids.

Extinct Higher Taxa in the Scarabaeoidea Nikolajev (1992, 1995a, b, 1996, 1998, 2000a, b, 2002, 2005c) is the only author to have introduced extinct higher taxa into the scarabaeoid classification. He described ten family-group taxa—the original diagnoses of which, however, reveal only a weak justification in some cases. Lucanidae: PARALUCANINAE Nikolajev 2000b (U Jurassic) Diagnosis: Body elongate; mandibles and labrum exposed, labrum not fused with clypeus; 10-segmented antennae weakly geniculate; ocular canthus present; pro- and mesocoxae contiguous; metatibiae broadened to apex; six visible sternites. Remarks: Six clearly visible sternites in combination with an ocular canthus differentiate this taxon from any extant Lucanidae. Paralucaninae was erected on the basis of a unique combination of characters rather than phylogenetic considerations. Geotrupidae: CRETOGEOTRUPINAE Nikolajev 1996 (L Cretaceous) Diagnosis: Relatively small body size; abdomen completely concealed by elytra; elytra with more than 11 striae each; protibia tridentate; mesocoxae contiguous; meso- and metatibiae transversely bicarinate. Remarks: The number of elytral striae and the venation of the hind wings (Nikritin 1977) indicate affiliation with Geotrupidae, but Nikolajev (1996) does not give a single diagnostic character separating Cretogeotrupinae from Geotrupinae. Pleocomidae: CRETOCOMINAE Nikolajev 2002, stat. nov. (L Cretaceous) Diagnosis: Pronotum without leathery anterior border; vein R3(5RA4) not developed in apical part of ala; mesocoxae contiguous or close; six visible sternites; female genitalia with ‘large styles’. Remarks: According to the drawing in Nikolajev (2002), the first two characters are hardly traceable. Based on the fossil as shown, Nikolajev’s reconstruction is rather speculative since he uses the absence of structures in incompletely preserved fossils as characters for classification. The characters presented are shared with other scarabaeoid taxa, including the allegedly diagnostic reduced RA4 (also in Glaresidae; Scholtz et al. 1994, but see Balthasar 1943). Nikolajev treated the group as a tribe (Cretocomini), and, because he considered Pleocomidae a subfamily, the Cretocomini must be upgraded to subfamily rank to be consistent with the current system.


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Ochodaeidae: CRETOCHODAEINAE Nikolajev 1995a (L Cretaceous–Oligocene) Diagnosis: Mandibles and labrum exposed; clypeus with ‘stripe’ on anterior margin; eyes partly separated by canthus; procoxae ‘appear contiguous’; mesotibiae without carinae; radius extending along anterior margin; radius sector (5RA4) at angle to radius and interrupted near base. Remarks: Nikolajev’s drawing resembles the hind wings of Glaphyridae (see Balthasar 1943) and the other diagnostic characters also would fit this family. Glaphyridae: CRETOGLAPHYRINAE Nikolajev 2005c, stat. nov. (L Cretaceous) Diagnosis: Mandibles and labrum exposed; eyes partly separated by canthus; mesocoxae contiguous; radius at anterior wing margin; vein R3(5RA4) is directed at an angle to radius and inserts distant from wing apex. Remarks: Because Nikolajev (2005c) considers the Glaphyridae a subfamily of Scarabaeidae, his tribe Cretoglaphyrini should be upgraded to subfamily rank to be consistent with the current system (Scholtz and Gebennikov 2005). However, Nikolajev’s differentiation from the remaining Glaphyrinae is incorrect: R3 inserts distant from the wing apex in other Glaphyrinae (Balthasar 1943), and in Anthypna Latreille, the mesocoxae are close or contiguous also. Scarabaeidae: CRETOSCARABAEINAE Nikolajev 1995b (L Cretaceous) Diagnosis: Mandibles and labrum exposed; antennal club with more than three segments; outer margin of protibia with three denticles; mesocoxae contiguous (Cretoscarabaeus Nikolajev) or narrowly separated (Cretorabaeus Nikolajev); meso- and metatibiae with transverse keels and with apical spurs located at one side of tarsus. Remarks: The documentation of the fossils only as line drawings does not allow Nikolajev’s interpretations to be reassessed, e.g., the number of articles of the clavus which is not identifiable, and antennae are only present in Cretoscarabaeus. All character states apart from three protibial denticles are considered plesiomorphic for the Scarabaeoidea by Nikolajev himself. Three protibial denticles, however, also are present in other Scarabaeoidea such as Glaresidae, Hybosoridae, Ceratocanthidae, Ochodaeidae, and several subfamilies of Scarabaeidae. Since Nikolajev uses the family rank Scarabaeidae for Scarabaeoidea sensu Crowson (1981), Cretoscarabaeinae should be upgraded to family level to adjust the author’s intention to the current system. However, because of the possible paraphyly of this group I refrain from upgrading, since the family level is subjectively more important owing to wider usage. Scarabaeidae: LITHOSCARABAEINAE Nikolajev 1992 (L Cretaceous) Diagnosis: Body size large; mandibles and labrum exposed; outer margin of protibiae with three denticles; meso- and metatibiae with two transverse keels; mesocoxae separated by mesothoracal process; two free veins between cubitus and first basally linked vein (Media Posterior 1 + 2). Remarks: Two free veins distal to the cubitus are present in Glaresidae, Lucanidae, Trogidae, Hybosoridae, and some Geotrupidae (Kempers 1923; Balthasar 1943). The other diagnostic characters also are widespread in the Scarabaeoidea. Scarabaeidae: PROTOTROGINAE Nikolajev 2000a (L Cretaceous) Diagnosis: Mesocoxae elongate, contiguous; mesotibia with two transverse keels; elytra covering pygidium; five visible sternites; lateral margins of sternites forming sharp edges. Remarks: According to Nikolajev, the taxon is diagnosed by symplesiomorphies only (in respect to Trogidae). Since Nikolajev classifies Trogidae, Geotrupidae, and Scarabaeidae as subfamilies (of Scarabaeidae 5 Scarabaeoidea without Lucanidae and Passalidae), his Prototroginae should be upgraded to family level to be consistent with the current classification (Lawrence et al. 2000; Scholtz and Gebennikov 2005). However, since Prototroginae are diagnosed solely by symplesiomorphies (Nikolajev 2000a) and are likely to be paraphyletic, I refrain from formally giving it the new status of a family. Nevertheless, there are no extant scarabaeoids with five visible sternites and tibiae with two transverse keels.

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Scarabaeidae: Melolonthinae: CRETOMELOLONTHINI Nikolajev 1998 (L Cretaceous) Diagnosis: Labrum not covered by clypeus; clypeus without anterior border; anterior border of pronotum leathery; Radius 1 (5RA3) and R3 (5RA4) apically approximated (but not fused); meso- and metatibia with single transverse carina; apical spurs of mesoand metatibia close-set; pygidium free; two last abdominal spiracles on sternites; six visible sternites. Remarks: All diagnostic characters seem to be plesiomorphic with respect to Melolonthinae (wing venation) or even Scarabaeidae. Cretomelolonthinae might be paraphyletic as defined. Scarabaeidae: Aclopinae?: HOLCOROBEINI Nikolajev 1992 (U Jurassic–L Cretaceous) Diagnosis: Medium-sized body; mandibles and labrum exposed; outer margin of protibia with three denticles; meso- and metatibia with two transverse keels; mesocoxae contiguous; each elytron with ten ‘dot-like grooves’; radius gradually thinning toward apex and running along frontal margin of hind wing; radius sector (5RA4) curving smoothly without joining radius (5RA3). Remarks: Nikolajev (2004) integrated the Holcorobeini into Aclopinae although two characters do not fit his own diagnosis of this subfamily: the tarsi of Holcorobeini are not or only slightly longer than the tibia (much longer in Aclopinae with the exception of Xenaclopus Arrow), and the meso- and metatibiae can have more than one keel on the outer side (one keel in Aclopinae with a rudimentary second in Xenaclopus). Moreover, the metaventrite and coxae are much smaller in the fossils than in extant Aclopinae.

The Systematic Status of Extinct Higher Scarabaeoid Taxa Although Nikolajev did a remarkable job in presenting and interpreting the Mesozoic scarabs of Russia, Kazakhstan, and Mongolia, the mostly insufficient documentation and inconsistent phylogenetic argumentation (often explicitly based on symplesiomorphies) reduce the value of his classification. Neither does he consider any possible displacement of body parts nor changes in size due to gaseous inflation during decay (Lutz 1990; Mendes 1999; Duncan et al. 2003) and fossilization. For example, in a flattened scarab that has the abdomen protruding beyond the elytra, it is impossible to reconstruct the original shape, e.g., whether the pygidium originally was covered by the elytra or not. The distance of the fossilized traces of coxae might have changed slightly during fossilization due to flattening and displacement. Mouthparts can be squeezed out of the head by gaseous pressure or simply be disarticulated and displaced in a decaying beetle. Apart from Prototroginae and Cretogeotrupinae, which do not have the head preserved, all other extinct higher taxa have the labrum and mandibles exposed. The characters mostly visible in fossils are coxae and wing venation, which have been neglected in phylogenetic analyses. Likewise, the evolution and possible homoplasy of the transverse tibial keels, typical for many groups of Scarabaeoidea and rare in other groups, is yet to be reconstructed. Most of the characters Nikolajev used to define extinct higher taxa are widely distributed across Scarabaeoidea. Therefore, a cladistic analysis including more wing venation and leg characters seems to be the best procedure for evaluating and fitting these proposed extinct taxa into the scarabaeoid classification system. As currently defined, most of them are likely to be either paraphyletic or synonyms of extant taxa, but without having seen the original material, I refrain from proposing formal synonymies. The Minimum Age of Extant Higher Taxa To determine the mimimum age of scarabaeoid families and subfamilies, I will consider the current classification of specimens even if the evidence is poor. For example, Nikolajev’s arguments for placing Cretaclopus Nikolajev, Juraclopus


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Fig. 4. Fossil record of Scarabaeoidea based on the phylogenetic system, corrected and updated from Krell (2000). Thick lines begin with the first fossil record of a clade. The phylogenetic system is adapted from Browne and Scholtz (1998, 1999) and Ahrens (2006), combined. Either Hybosoridae or Ceratocanthidae are paraphyletic with respect to the other (Grebennikov and Scholtz 2004; Grebennikov et al. 2004), but because of contradicting evidence, they are kept as sister groups for the time being. Abbreviations: Cretac.– Cretaceous; Copr.–Coprinae; L–Lower; Scarab.–Scarabaeidae; U–Upper.

Nikolajev, and Holcorobeini in Aclopinae are not well supported. Since the phylogenetic position of Aclopinae within the Scarabaeoidea has never been analysed (see Browne and Scholtz 1999; Scholtz and Grebennikov 2005), apomorphies of the taxon are unknown. Nikolajev considered the long tarsi of Cretaclopus, combined with exposed mouthparts, evidence enough to justify the assignment. However, the first character state is likely to be plesiomorphic in Scarabaeoidea and the second is homoplastic (e.g., present in Aphodiinae, Melolonthinae). One of the oldest lineages (Fig. 2) according to Nikolajev’s (1996, 2002) assignment of fossils is the Pleocominae, which is based on the interpretation of Grabau’s (1923) original description and drawing of Proteroscarabaeus yeni. He assigned it to Pleocomidae because of six denticles on the outer edge of the left protibia (which could be a result of degradation), and an antennal club consisting of seven segments (which could be a displaced, damaged antenna or maxilla). The other two species of Proteroscarabaeus Grabau he described were represented by single elytra, and P. robustus Zhang from China has neither antennae nor tibiae preserved (Zhang 1997). As doubtful as these and other similar assignments are, we cannot falsify them on the basis of the available evidence. I therefore considered them in (Fig. 4) as an indicator for the minimum age of extant higher scarabaeoid taxa, but they are weak hypotheses. If we follow these interpretations, all family-level radiations

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occurred in the Mesozoic. Also, half of the Scarabaeidae subfamilies of the socalled ‘Cenozoic Clade’ (Scholtz and Chown 1995) originated in the Mesozoic. Only the Rutelinae-Trichiinae clade, Ahrens’s (2006) ‘melolonthine group I’, diversified in the Lower Tertiary (Fig. 4). Paleoecology of Scarabaeoidea Assigning feeding or other ecological traits to fossil beetles is notoriously difficult. Narrative scenarios have some appeal and may be intrinsically consistent (e.g., Ponomarenko 2003), but rely on extensive interpretations of sparse evidence. We have only two sources of evidence: fossilized traces of feeding activities and optimizing feeding habits of extant taxa onto cladograms (e.g., Scholtz and Chown 1995). Fossilized evidence of feeding activity of insects refers mainly to phytophagy and coprophagy. Traces of leaf feeding are frequently documented in the fossil record (Scott et al. 1992), but identifying the tracemakers is virtually impossible. The fossil evidence for coprophagy is large, and assigning it to the Scarabaeoidea is straightforward if we consider scarabs to be the only dung feeders producing dung balls and tunnels. Beutel and Leschen (2005) identified saprophagy as an apomorphy for all Polyphaga taxa included in their analysis. It was hypothesized to be the ancestral feeding habit of the hydrophiloid lineage (Hydrophiloidea, Histeroidea, and Scarabaeoidea) by Hansen (1997b) and of ‘laparostict’ scarabs by Cambefort (1991). However, by mapping extant feeding habits onto the then latest phylogenetic tree, Scholtz and Chown (1995) identified mycetophagy as plesiomorphic conditions for the adults and humus feeding for the larvae of scarabaeoids. The topology of the scarabaeoid tree has changed since, and some extant feeding habits were not considered in the analysis, e.g., coprophagy in adult Trogidae (Krell et al. 2003 and references therein), predatory Hybosoridae and Cetoniinae, sap-feeding Dynastinae and Lucanidae, and necrophagy in Cetoniinae. A new analysis is beyond this review but necessary to test the hypothesis of a fungus-feeding scarabaeoid stem species with humus-feeding larvae. Coprophagy The currently widely accepted hypothesis is that coprophagy developed mainly as a consequence of Tertiary mammal radiations in grasslands (see Scholtz and Chown 1995), though it is likely to have originated before that. Halffter and Matthews (1966:58) proposed this hypothesis on the following grounds: 1. 2. 3.

Virtual absence of necrophagy in grassland scarabs, but the African savanna fauna has an abundant and speciose scarabaeoid-dominated carrion community (Cambefort 1984; Krell and Linder 1996; Krell et al. 2003). Dung ball rolling could have evolved only in open habitats. However, rollers are frequent and speciose in Old World tropical forests. Subterranean nidification is more advantageous in the open than in buffered habitats. However, subterranean nidification does not only protect from climatic adversities but also from predators and provides a mechanism for avoiding competition within the feces irrespective of the habitat.

The rich African dung beetle fauna indicates a radiation of dung beetles in Africa’s mammal-rich grasslands, but the rich dung beetle fauna in, e.g., Bornean


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rainforest (Davis 2000), clearly shows that open grassland is not a necessary condition for the development of a speciose coprocenosis. Is mammal dung necessary? Most extant dung beetles feed on mammal dung since in present terrestrial ecosystems, it is the most abundant dung type occurring in usable portions. However, in the Mesozoic dinosaur dung undoubtedly occurred in larger quantities. The extant Archosauria-dung fauna is either barely studied (crocodile dung), or the dung portions are generally small and less suitable for dung beetles (bird droppings). However, the latter have been shown to attract dung beetles (Vinson 1951; Halffter and Matthews 1966; Oppenheimer and Begum 1978; Tanaka 1985; pers. obs.), and crocodile dung has been seen to attract Nesosisyphus vicinus (Vinson) in Mauritius (Motala and Krell, unpublished). Extant Squamata and amphibian dung has been proven to be attractive to at least a few dung beetles (Young 1981). However, as long as large ‘reptilian’ feces are not thoroughly studied (e.g., Komodo dragon, crocodiles) and found to be generally unattractive, we have no hint to rule out dinosaur dung as having triggered the evolution of coprophagy. If Cretaceous dinosaurs also fed on grass as suggested by Prasad et al. (2005), their dung might have been more similar to the dung of extant grass-feeding mammals than to the dung of extant predatory reptiles. Intuitively, we may expect dinosaur feces to have occurred in large portions as it was impressively visualized in the movie ‘Jurassic Park’. However, evidence is scarce. It is difficult to definitely assign coprolites to dinosaurs, and most fossilized dinosaur feces are rather small, less then 10 cm long and less than 5 cm in diameter (Thulborn 1991), but still a potentially useful resource for smaller coprophagans and, in piles, for the largest species. Karen Chin and colleagues found a ‘king-sized’ tyrannosaur coprolite of 20 cm 3 10 cm in Saskatchewan (Chin et al. 1998) that would satisfy the needs of many modern dung beetles and Trogidae for feeding and reproduction. Feeding traces of coprophagous invertebrates have been found on coprolites from as early as the Lower (Northwood 2005) and Upper Triassic (Wahl et al. 1998) and Upper Cretaceous (Matley 1941), but any hypothesis about the responsible organisms remains speculation. Wahl et al. (1998) proposed Diptera larvae as possible trace-makers of the tiny tunnels (less than 1 mm diameter) because dung beetles were too large, but the Upper Cretaceous amber scarabs or the smallest extant Aphodiinae are small enough to have produced such traces. From the Upper Cretaceous onwards, fossil dung balls similar to the ones made by extant dung beetles are known from South America. In Tertiary paleosols, they have been recorded from North America, Antarctica, Asia, and Africa (Krell 2000). Additionally, Chin and Gill (1996) found a coprolite of a Cretaceous herbivorous dinosaur with burrows backfilled with sediment, and, in the surrounding sediment, burrows backfilled with fecal material. Only dung beetles are known to produce such structures. This has been taken as proof for dung beetle activity in the Mesozoic. Most Mesozoic scarab fossils, however, have a different body shape than extant dung beetles. Prionocephale deplanate Lin from the Upper Cretaceous of the Lanxi formation, Zhejiang (92–83.5 myr, Lin 1994) is an exception. It resembles members of the Onthophagini or even a telecoprid (roller), and, if it is a scarabaeine, with a body length of about 2 cm, it could have processed a substantial amount of dung. Although dung feeding is unlikely to be the ancient feeding habit of Scarabaeoidea, it may have developed early in the Mesozoic.

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Quaternary Scarabs–The Present in the Past The oldest fossil Scarabaeoidea tentatively assigned to extant species are those from the Pliocene of Willershausen, Germany: Aphodius rufipes (L.) (Aphodiinae), Oryctes nasicornis (L.) (Dynastinae) (Gersdorf 1971), and Lucanus cervus (L.) (Lucanidae) (Schweigert 2003), whereas the youngest extinct fossil species have been described from three Pleistocene deposits: 1.

2.

3.

Port Kennedy Caves: A sinkhole in the Valley Forge National Historical Park, Pennsylvania (Daeschler et al. 1993; Bechtel et al. 2005): Choeridium ebenium Horn, Phanaeus antiquus Horn, Aphodius precursor Horn (Horn 1876). Rancho La Brea: The famous tar pits in Los Angeles, California (Stock and Harris 1992): Copris pristinus Pierce, Onthophagus everestae Pierce, Phanaeus labreae (Pierce), Serica kanakoffi Pierce, the last two species being unidentifiable at the species level and, therefore, possible synonyms of extant species (Miller et al. 1981). Borislav, W Ukraine: Aphodius boryslavicus Łomnicki, A. rhinocerontis Łomnicki, A. ruthenus Łomnicki, and A. subater Łomnicki (Łomnicki 1894).

Most Pleistocene specimens can be assigned to extant species, though their past and present ranges might differ. The most famous case for such a range change is Aphodius holdereri Reitter, a Tibetan dung beetle species that was found in Late Pleistocene deposits in England (Coope 1973). Another dung beetle, Aphodius bonvouloiri Harold, today a Spanish endemic, was one of the most abundant beetles in Great Britain in the warm interstadial in the middle of the last glaciation (Coope 1990). As late as the Holocene, species were found well outside their present ranges, e.g., Valgus hemipterus (L.) (Valginae) in southern Sweden in an archeological site 7,000–5,000 BP (Lemdahl 1990) or Protaetia aeruginosa (Drury) (Cetoniinae) from peat of the Subboreal Period (ca. 5,000–2,500 BP) in Denmark (Johnsen and Krog 1948); both were north of their present distributions. Such range shifts render identifications difficult because not only the local fauna but the fauna of the whole continent or even hemisphere has to be considered to find extant matches or to decide whether a Pleistocene fossil is a new species. Angus (1973) and Kus´ka (1991) identified all four Pleistocene Helophorus (Hydrophilidae) and three of the five weevil species described by Łomnicki (1894) from Borislav (Ukraine) as extant species that do no longer occur in this area, but in Siberia or higher up in the Carpathians. The same may happen to Łomnicki’s four fossil Aphodius species in a future revision. Miller et al. (1981) considered it possible that the fossil species from La Brea will be discovered in the extant Mexican fauna, and the fossils from Port Kennedy have never been revised. Although morphological evolution (Brower 1994; Huey et al. 2000) and speciation in insects can be rapid (Mendelson and Shaw 2005), such evidence in the fossil record of Scarabaeoidea is scarce and needs to be re-evaluated. In this respect, the scarabaeoid fossil record doesn’t differ from that of other insect groups. As stated by Buckland and Coope (1991): ‘‘Contrary to initial expectations, Quaternary entomology provides no evidence for any evolution of insects during the Middle and Late Pleistocene’’. However, Quaternary entomology is a rich source for reconstruction of climate and habitat change and for changing ranges of extant species (Ponel 1993; Ashworth et al. 1997; Porch and Elias 2000), and Scarabaeoidea are among the taxa most commonly used (Porch and Elias 2000). The literature is vast (Buckland and Coope 1991) and growing, and cannot be summarized here.

John Day Series, Oregon Shara-Teg, Mongolia Baltic Amber Laiyang, China Baissa, Russia Baissa, Russia Baissa, Russia Bayan-Teg, Mongolia Dominican Amber Bon-Tsagan, Mongolia Karatau, Kazakhstan Menat, France* Baissa, Russia Lanxi, China Leskovo, Russia Messel, Germany /Baltic Amber Baissa, Russia Tottori, Japan Bournemouth, England Clarno Formation, Oregon Eckfelder Maar, Germany Clarkia Fossil Beds, Idaho ¨ hningen, Germany O ¨ hningen, Germany O

U Oligocene [c. 33–39 myr] U Jurassic [161–145 myr] Eocene [ca. 50 myr] L Cretaceous [ca. 130 myr?] L Cretaceous [.130 myr] L Cretaceous [.130 myr] L Cretaceous [.130 myr] U Jurassic Miocene [ca. 15–20 myr] L Cretaceous [130–125 myr]

U Jurassic [152–158 myr] U Paleocene [56 myr] L Cretaceous [.130 myr] U Cretaceous [92–83.5 myr] L Cretaceous [145–140 myr] L/M Eocene [47/50 myr] L Cretaceous [.130 myr] M Miocene L/M Eocene [50–42 myr]) M Eocene [44.6–46.8 myr] M Eocene [44 myr] L Miocene [17–20 myr] M Miocene [13.6–14.8 myr] M Miocene [13.6–14.8 myr]

Passalidae Lucanidae Trogidae Pleocomidae Bolboceratidae Glaphyridae Geotrupidae Hybosoridae Ceratocanthidae Ochodaeidae Scarabaeidae Aclopinae Aphodiinae Aegialiinae Scarabaeinae/Coprinae Sericinae Hopliinae Melolonthinae Euchirinae Rutelinae Dynastinae Cetoniinae Osmodermatinae Valginae Trichiinae

Locality

Minimum age (evidence)

Taxon

Juraclopus rohdendorfi Nikolajev 2005b Aphodius charauxi Piton 1940 Cretaegialia spp. (Nikolajev 1994) Prionocephale deplanate Lin (Krell 2000) Lithanomala spp. (Nikolajev 1992) undescribed (Krell unpublished) Cretomelolontha transbaikalica Nikolajev 1998a Cheirotonus otai Ueda 1989 Pelidnotites atavus Cockerell 1920 Oryctoantiquus borealis Ratcliffe and Smith 2005 undescribed (Wappler 2003) Osmoderma sp., undescribed (Lewis 1985) Valgus oeningensis Heer 1862 Trichius spp. (Heer 1862)

Passalus indormitus Cockerell (Reyes Castillo 1977) Paralucanus mesozoicus Nikolajev 2000b undescribed (Krell unpublished) Proteroscarabaeus yeni Grabau (Nikolajev 2002) Cretobolbus rohdendorfi Nikolajev 1996 Cretoglaphyrus spp. (Nikolajev 2005c) Cretogeotrupes convexus Nikolajev 1992 Jurahybosorus mongolicus (Nikolajev 2005d) undescribed (Poinar and Poinar 1999) Cretochodaeus mongolicus Nikolajev 1995a

Species (reference)

Table 1. Minimum age of scarabaeoid family-group taxa based on fossil evidence. Their minimum age based on the age of the sister group is shown in (Fig. 4). (*Menat was originally thought to be Eocene and was listed as such in my catalog (Krell 2000), but Vincent et al. (1977) dated it at ca. 56 myr [Palaeocene]).

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Scarabaeoid fossils are recorded continuously up to historical times, e.g., from Roman and Medieval times in Rhenania (Koch 1970) and from a 15th century site in Sweden (Lemdahl and Thelaus 1989). Outlook Grimaldi (2001) noted that ‘Despite Hennig’s influence and background, paleoentomology remains largely descriptive and taxonomic.’ Whereas Quaternary fossils are widely used to reconstruct and model paleoclimates and distribution patterns, with Mesozoic and Tertiary beetle fossils a descriptive level is mostly as far as we can get. However, the number of exceptionally wellpreserved scarabaeoid fossils being published is constantly increasing and their consideration in more comprehensive cladistic analyses may only be a matter of time. Only then can their classification and the reconstruction of the evolutionary history of Scarabaeoidea be improved. Acknowledgments I am deeply indebted to Georgy Nikolajev (Al-Faraby University Almaty, Kazakhstan) for updating me about his latest finds and allowing me to use his picture of Juraclopus in this paper. John Anderson, then National Botanical Institute, Pretoria, allowed me to look through the Molteno beetles. Svetlana Nikolajeva and Max Barclay (NHM London) helped with translating Russian literature. Haruki Karube (Kanagawa Prefectural Museum of Natural History, Japan) gave further information about his Australian fossil. Nick Porch (Monash University, Australia) sent me his unpublished manuscript on the first Australian scarab fossils. My postdoc, Dorothy Newman, improved the manuscript considerably.

Literature Cited Ahrens, D. 2006. The phylogeny of Sericini and their position within the Scarabaeidae based on morphological characters (Coleoptera: Scarabaeidae). Systematic Entomology 31:113–144. Angus, R. B. 1973. Pleistocene Helophorus (Coleoptera, Hydrophilidae) from Borislav and Starunia in the western Ukraine, with a reinterpretation of M. Łomnicki’s species, description of a new Siberian species, and comparison with British Weichselian faunas. Philosophical Transactions of the Royal Society of London (B) 265:299– 326. Ashworth, A. C., P. C. Buckland, and J. P. Sadler (eds.). 1997. Studies in Quaternary Entomology. An Inordinate Fondness for Insects. Quaternary Proceedings 5:i–viii, 1–303. ¨ ber die Nervatur des Hinterflu¨gels der Lamellicornien. Veˇstnik Balthasar, V. 1943. U ˇ eske´ Spolecˇnosti Nauk Trˇ´ıda Matematicko-Prˇ´ırodoveˇdecka´ 1942, Kra´lovske´ C 11:1–44. Bechtel, T. D., J. L. Hojdila, S. H. Baughman II, T. DeMayo, and E. Doheny. 2005. Relost and refound: Detection of a paleontologically, historically, cinematically(?), and environmentally important solution feature in the carbonate belt of southeastern Pennsylvania. The Leading Edge 24:537–540. Berger, J. P., B. Reichenbacher, D. Becker, M. Grimm, K. Grimm, L. Picot, A. Storni, C. Pirkenseer, and A. Schaefer. 2005. Eocene-Pliocene time scale and stratigraphy of the Upper Rhine Graben (URG) and the Swiss Molasse Basin (SMB). International Journal of Earth Sciences 94:711–731. Bertrand, E. 1763. Dictionnaire Universel des Fossiles Propres, et des Fossiles Accidentels […]. Tome premier. Gosse et Pinet, La Hate. Beutel, R. G., and R. A. B. Leschen. 2005. Phylogenetic analysis of Staphyliniformia (Coleoptera) based on characters of larvae and adults. Systematic Entomology 30:510–548.


135

Blair, K. G. 1927. Insect remains from oil sand in Trinidad. Transactions of the Entomological Society London 75:137–141. Britton, E. B. 1960. Beetles from the London Clay (Eocene) of Bognor Regis, Sussex. Bulletin of the British Museum (Natural History), Geology 4:27–50, pls 2-7. Brower, A. V. Z. 1994. Rapid morphological radiation and convergence among races of the butterfly Heliconius erato inferred from patterns of mitochondrial DNA evolution. Proceedings of the National Academy of Science U.S.A. 91:6491–6495. Browne, J., and C. H. Scholtz. 1998. Evolution of the scarab hindwing articulation and wing base: a contribution toward the phylogeny of the Scarabaeidae (Scarabaeoidea: Coleoptera). Systematic Entomology 23:307–326. Browne, J., and C. H. Scholtz. 1999. A phylogeny of the families of Scarabaeoidea. Systematic Entomology 24:51–84. Buckland, P. C., and G. R. Coope. 1991. A Bibliography and Literature Review of Quaternary Entomology. J. R. Collins Publications, University of Sheffield. ´ tude ećologique des coleópte`res Scarabaeidae de Coˆte d’Ivoire. Cambefort, Y. 1984. E Travaux des Chercheurs de la Station de Lamto 3:1–294. Cambefort, Y. 1991. From saprophagy to coprophagy [pp. 22–35]. In: Dung Beetle Ecology. I. Hanski and Y. Cambefort (eds.). Princeton: Princeton University Press. Caterino, M. S., T. Hunt, and A. P. Vogler. 2005. On the constitution and phylogeny of Staphyliniformia (Insecta: Coleoptera). Molecular Phylogenetics and Evolution 34:655–672. Chalumeau, F., and B. Brochier. 2001. Une forme fossile nouvelle de Chiasognathinae : Protognathinus spielbergi (Coleoptera, Lucanidae). Lambillionea 101:593–595. Chin, K., and B. D. Gill. 1996. Dinosaurs, dung beetles, and conifers: Participants in a Cretaceous food web. Palaios 11:280–285. Chin, K., T. T. Tokaryk, G. M. Erickson, and L. C. Calk. 1998. A king-sized theropod coprolite. Nature 393:680–682. Churcher, C. S. 1966. The insect fauna from the Talara tar-seeps, Peru. Canadian Journal of Zoology 44:985–993. Cockerell, T. D. A. 1920. Fossil Arthropods in the British Museum.—II. Annals and Magazine of Natural History, Series 9 5:454–463, pl. 16. Coope, G. R. 1973. Tibetan species of dung beetle from Late Pleistocene deposits in England. Nature 245:335–336. Coope, G. R. 1990. The invasion of Northern Europe during the Pleistocene by Mediterranean species of Coleoptera [pp. 203–215]. In: Biological Invasions in Europe and the Mediterranean Basin. F. di Castri, A. J. Hansen, and M. Debusche (eds.). Kluwer, Dordrecht, Netherlands. Crowson, R. A. 1981. The Biology of the Coleoptera. Academic Press, London, U.K. Crowson, R. A. 1995. Some interesting evolutionary parallels in Coleoptera [pp. 63–85]. In: Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. J. Pakaluk and S. A. S´lipin´ski (eds.). Muzeum i Instytut Zoologii PAN, Warszawa. Daeschler, E., E. E. Spamer, and D. C. Parris. 1993. Review and new data on the Port Kennedy local fauna and flora (Late Irvingtonian), Valley Forge National Historical Park, Montgomery County, Pennsylvania. The Mosasaur 5:23–41. Davis, A. J. 2000. Species richness of dung-feeding beetles (Coleoptera: Aphodiidae. Scarabaeidae, Hybosoridae) in tropical rainforest at Danum Valley, Sabah, Malaysia. Coleopterists Bulletin 54:221–231. Duncan, I. J., F. Titchener, and D. E. G. Briggs. 2003. Decay and disarticulation of the cockroach: implications for preservation of the blattoids of Writhlington (Upper Carboniferous), UK. Palaios 18:256–265. Evanoff, E., W. C. McIntosh, and P. C. Murphey. 2001. Stratigraphic summary and 40 Ar/39Ar geochronology of the Florissant Formation, Colorado. Proceedings of the Denver Museum of Nature and Science series 4(1):1–16. Franzen, J. L. 2005. The implications of the numerical dating of the Messel fossil deposit (Eocene, Germany) for mammalian biochronology. Annales de Paleóntologie 91:329–335.

136


¨ ningen am Frentzen, K. 1927. Die fossile Insektenfauna der obermioza¨nen Kalkmergel von O Bodensee. Archiv fu¨r Insektenkunde des Oberrheingebietes und der angrenzenden La¨nder 2:137–147. Fricˇ, A. 1885. Uiber einen fossilen Maika¨fer (Anomalites fugitivus Fr.) aus dem tertia¨ren Su¨sswasserquarz von Nogent le Rotrou in Frankreich. Sitzungsberichte der Ko¨nigl. Bo¨hm. Gesellschaft der Wissenschaften in Prag 1884, Vortra¨ge in den Sitzungen der mathematisch-naturwissenschaftlichen Classe: 163–165. Genise, J. F., M. G. Mańgano, L. A. Buatois, J. H. Laza, and M. Verde. 2000. Insect trace fossil associations in paleosols: The Coprinisphaera ichnofacies. Palaios 15:49–64. Gersdorf, E. 1971. Weitere Ka¨fer (Coleoptera) aus dem Jungtertia¨r Norddeutschlands. Geologisches Jahrbuch 88:629–670. Giebel, C. 1862. Wirbelthier und Insektenreste im Bernstein. Zeitschrift fu¨r die Gesammten Naturwissenschaften 20:311–321. Grabau, A. W. 1923. Cretaceous fossils from Shantung. Bulletin of the Geological Survey of China 5:148–181, 2 pls. Grebennikov, V. V., A. Ballerio, F. C. Ocampo, and C. H. Scholtz. 2004. Larvae of Ceratocanthidae and Hybosoridae (Coleoptera: Scarabaeoidea): study of morphology, phylogenetic analysis and evidence of paraphyly of Hybosoridae. Systematic Entomology 29:524–543. Grebennikov, V. V., and C. H. Scholtz. 2004. The basal phylogeny of Scarabaeoidea (Insecta: Coleoptera) inferred from larval morphology. Invertebrate Taxonomy 18:321–348. Grimaldi, D. A. 1991. The Santana formation insects [pp. 379–405]. In: Santana Fossils: An Illustrated Atlas. J. G. Maisey (ed.). T.F.H. Publications, Neptune City, NJ. Grimaldi, D. 2001. Insect evolutionary history from Handlirsch to Hennig, and beyond. Journal of Paleontology 75:1152–1160. Grimaldi, D., and M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press, Cambridge, U.K. Halffter, G., and E. G. Matthews. 1966. The natural history of dung beetles of the subfamily Scarabaeinae (Coleoptera, Scarabaeidae). Folia Entomologica Mexicana 12– 14:1–312. Handschin, E. 1950. Zur Entstehung der Quercyfossilien. Proceedings, Eighth International Congress of Entomology: 137–140. Hansen, M. 1997a. Phylogeny and classification of the staphyliniform beetle families (Coleoptera). Biologiske Skrifter, Kongelige Danske Videnskabernes Selskab 48:1–340. Hansen, M. 1997b. Evolutionary trends in ‘‘staphyliniform’’ beetles (Coleoptera). Steenstrupia 23:43–86. Harding, I. C., and L. S. Chant. 2000. Self-sedimented diatom mats as agents of exceptional fossil preservation in the Oligocene Florissant lake beds, Colorado. Geology 28:195–198. Harms, F.-J., H. Buness, M. Felder, and T. Nix. 2004. Neue Vorstellungen zur Entstehung der Welterbesta¨tte Grube Messel in aktuellen geologischen und seismischen Schnitten. Bericht, Naturwissenschaftlicher Verein Darmstadt, N.F. 27:71–86. Haupt, H. 1950. Die Ka¨fer (Coleoptera) aus der eoza¨nen Braunkohle des Geiseltales. Geologica 6:i–viii,1–168. Heer, O. 1862. Beitra¨ge zur Insektenfauna Oeningens. Coleoptera. Geodaphagen, Hydrocanthariden, Gyriniden, Brachelytren, Clavicornen, Lamellicornen und Buprestiden. Natuurkundige Verhandelingen van de Hollandsche Maatschappij der Wetenschappen te Haarlem 16(2):1–90, pls. Hellmund, M. 1988. Eine Ka¨ferelytre mit Farberhaltung aus der oberoligoza¨nen Bla¨tterkohle von Rott im Siebengebirge. Decheniana 141:343. Horn, G. H. 1876. Notes on some coleopterous remains from the bone cave at Port Kennedy, Penna. Transactions of the American Entomological Society 5:241–245. Howden, H. F. 1972. Systematics and zoogeography: science or politics? Systematic Zoology 21:129–131.


137

Huey, R. B., G. W. Gilchrist, M. L. Carlson, D. Berrigan, and L. Serra. 2000. Rapid evolution of a geographic cline in size in an introduced fly. Science 287:308– 309. Jell, P. A. 2004. The fossil insects of Australia. Memoirs from the Queensland Museum 50:1–124. Johnsen, P., and H. Krog. 1948. Potosia speciosissima Scop. subfossil in Denmark (Coleoptera Lamellicornia). Entomologiske Meddelelser 25:252–262. Karube, H. 2002. A hemi-fossil of Phalacrognathus muelleri (Coleoptera, Lucanidae) found near Cairns, Queensland, Australia. Gekkan-Mushi 378:34 [in Japanese]. Kempers, K. J. W. B. 1923. Abbildungen von Flu¨gelgea¨der der Coleopteren. Entomologische Mitteilungen 12:71–115, 162–163. Koch, K. 1970. Subfossile Ka¨ferreste aus ro¨merzeitlichen und mittelalterlichen Ausgrabungen im Rheinland. Entomologische Bla¨tter 66:41–56. Koenigswald, W. von (ed.). 1996. Fossillagersta¨tte Rott bei Hennef im Siebengebirge. 2nd ed. Rheinlandia-Verlag, Siegburg, Germany. Krell, F.-T. 2000. The fossil record of Mesozoic and Tertiary Scarabaeoidea (Coleoptera: Polyphaga). Invertebrate Taxonomy 14:871–905. Krell, F.-T., J. Korb, and P. Walter. 2003. The beetle fauna of hyaena latrines: Coprocenoses consisting of necrophagous beetles (Coleoptera: Trogidae, Scarabaeidae). Tropical Zoology 16:145–152. Krell, F.-T., and C. Linder. 1996. Preliminary study on necrocenoses in the Guinea savanna in the Ivory Coast (Coleoptera). Proceedings, XX International Congress of Entomology, Firenze, Italy, August 25–31, 1996: 547. Kugler, H. G. 1927. Report on a fossil locality for arthropods and vertebrates in Trinidad (B.W.I.). Transactions of the Entomological Society London 75:141–142. Kukalova´-Peck, J., and J. F. Lawrence. 1993. Evolution of the hind wing in Coleoptera. Canadian Entomologist 125:181–258. Kus´ka, A. 1991. Revision of fossil Curculionidae (Coleoptera) from Borysław and Starunia near Lwo´w. Polskie Pismo Entomologiczne 61:89–95. Lausch, E., T. Ernsting, and M. Mu¨ller. 2000. Aus der Tiefe der Zeit. Geo 4/2000: 148–170. Lawrence, J. F., E. B. Britton. 1991. Coleoptera (Beetles) [pp. 543–683]. In: The Insects of Australia. A Textbook for Students and Research Workers. Division of Entomology, Commonwealth Scientific and Industrial Research Organisation (ed.). Melbourne University Press, Carlton, Australia. Lawrence, J. F., A. M. Hastings, M. J. Dallwitz, T. A. Paine, and E. J. Zurcher. 2000. Beetles of the World: A Key and Information System for Families and Subfamilies. CSIRO, Collingwood, Australia [CD-ROM]. Laza, J. H., and M. A. Reguero. 1990. Extensioń faunistica de la antigua regioń Neotropical en la penıńsula anta´rtica durante el Eoceno. Ameghiniana 26:245. Leakey, L. S. B. 1952. Lower Miocene invertebrates from Kenya. Nature 169:624–625. Lemdahl, G. 1990. Ett subfossilfynd av bladhorningen Valgus hemipterus fra˚n Ska˚ne. Entomologisk Tidskrift 111:99–100. Lemdahl, G., and Thelaus, M. 1989. Subfossila skalbaggsfynd fra˚n medeltida Halmstad. Entomologisk Tidskrift 110:39–41. Lewis, S. E. 1985. Miocene insects from the Clarkia deposit of northern Idaho [pp. 245–264]. In: Late Cenozoic History of the Pacific Northwest. C. J. Smiley, A. E. Leviton, and M. Berson (eds.). Interdisciplinary Studies on the Clarkia Fossil Beds of Northern Idaho, San Francisco, CA. Lin, Q.-b. 1980. [Fossil Insects] [pp. 211–244]. In: Zhe-Wan Zhongshengdai huoshan chenji yan diceng de huafen ji duibi. Zhongguo Kexueyuan. Nanjing Dizhi Gushengwu Yanjiusuo [Academia Sinica. Nanjing Institute of Geology and Palaeontology] (ed.). [‘Fossils of Mesozoic Deposits of Volcanic Origin in Zhejiang and Anhui.’] Science Press, Peking, China [in Chinese]. Lin, Q.-b. 1994. Cretaceous insects of China. Cretaceous Research 15:305–316. Łomnicki, A. M. 1894. Pleistocen´skie owady z Borysławia. Muzeum Imienia Dzieduszyckich we Lwowie 4:1–116, 9 pls.

138


Lutz, H. 1990. Systematische und palo¨kologische Untersuchungen an Insekten aus dem Mittel-Eoza¨n der Grube Messel bei Darmstadt. Courier Forschungsinstitut Senckenberg 124:1–165. Lutz, H. 2003. The Eckfeld Maar (Vulkaneifel, Germany)—a key locality for the understanding of the Eocene paleoenvironment of the western Rhenish shield. Geólogie de la France 2003(1):63–67. Martıńez-Delclo`s, X., and J. Martinell. 1993. Insect taphonomy experiments. Their application to the Cretaceous outcrops of lithographic limestones from Spain. Kaupia 2:133–144. Matley, C. A. 1941. The coprolites of Pijdura, Central Provinces. Records of the Geological Survey of India 74:535–547, pls. 34-37. Mendelson, T. C., and K. L. Shaw. 2005. Rapid speciation in an arthropod. Nature 433:375–376. Mendes, M. 1999. An attempt at taphonomic simulation in blattopteras from Araripe (Lower Cretacean) from the NE of Brasil. Proceedings of the First International Palaeoentomological Conference, Moscow 1998:99–101. Mertz, D. F., C. C. Swisher, J. L. Franzen, F.-O. Neuffer, and H. Lutz. 2000. Numerical dating of the Eckfeld maar fossil site, Eifel, Germany: a calibration mark for the Eocene time scale. Naturwissenschaften 87:270–274. Meyer, H. W. 2003. The Fossils of Florissant. Smithsonian, Washington, D.C. Miller, S. E., R. D. Gordon, and H. F. Howden. 1981. Reevaluation of Pleistocene scarab beetles from Rancho La Brea, California (Coleoptera: Scarabaeidae). Proceedings of the Entomological Society Washington 83:625–630. Mo¨rs, T. 1995. Die Sedimentationsgeschichte der Fossillagersta¨tte Rott und ihre Alterseinstufung anhand neuer Saügetierfunde (Oberoligoza¨n, Rheinland). Courier Forschungsinstitut Senckenberg 187:1–129. Nikolajev, G. V. 1992. Taksonomicheskie priznaki i sostav rodov mezozoiskikh plastinchatousykh zhukov (Coleoptera, Scarabaeidae). Paleontologicheskiy Zhurnal 1992(1):76–88. [Translation: Nikolajev, G. V. 1993. Taxonomic criteria and generic composition of Mesozoic lamellicorn beetles (Coleoptera, Scarabaeidae). Paleontological Journal 26:96–111.]. Nikolajev, G. V. 1995a. Materialy k sistematike podsemeistva Ochodaeinae (Coleoptera, Scarabaeidae). Zoologicheskiy Zhurnal 74:72–82. [Translation: Nikolajev, G. V. 1996. New Data on the Systematics of the Subfamily Ochodaeinae (Coleoptera, Scarabaeidae). Entomological Review 75:113–123.]. Nikolajev, G. V. 1995b. Novoe podsemeistvo plastinchatousykh zhukov (Coleoptera, Scarabaeidae) iz nizhnego mela Zabaikal’ya i ego filogeneticheskiye svyazi. Paleontologicheskiy Zhurnal 1995(2):147–151. [Translation: Nikolajev, G. V. 1995. A new subfamily of lamellicorn beetles (Coleoptera, Scarabaeidae) from the Lower Cretaceous of Transbaykalia and its phylogenetic relationships. Paleontological Journal 29:185–192.]. Nikolajev, G. V. 1996. Vidy plastinchatousykh zhukov (Coleoptera, Scarabaeidae) s bol’shim kolichestvom bopozdok na nadkryl’iakh iz nizhemelovogo mestonakhozhdeniia Baisa v Zabaikalie. Paleontologicheskiy Zhurnal 1996(2):91–99. [Translation: Nikolajev, G. V. 1996. Lamellicorn Beetle Species (Coleoptera, Scarabaeidae) with Multistriate Elytra from the Lower Cretaceous Baisa Locality, Transbaikalia. Paleontological Journal 30:217–224.]. Nikolajev, G. V. 1998. Vidy plastinchatousykh zhukov gruppy Pleurosticti (Coleoptera, Scarabaeidae) iz nizhnego mela Zabaikaliya. Paleontologicheskiy Zhurnal 1998(5):77–84, pl. 10. [Translation: Nikolajev, G. V. 1998. Pleurostict Lamellicorn Beetles (Coleoptera, Scarabaeidae) from the Lower Cretaceous of Transbaikalia. Paleontological Journal 32:513–521.]. Nikolajev, G. V. 2000a. Novoe podsemeystvo plastinchatousykh zhukov (Coleoptera, Scarabaeidae) iz nizhiego mela Zavaykal’ya i Mongolii i ego polozhenie v sisteme nadsemeystva. Paleontologicheskiy Zhurnal 2000(4):63–66. [Translation: Nikolajev, G. V. 2000: A new subfamily of lamellicorn beetles (Coleoptera, Scarabaeoidae [sic]) from the Lower Cretaceous of Transbaikalia and Mon-


139

golia, and its position within the superfamily. Paleontological Journal 34:426– 428.]. Nikolajev, G. V. 2000b. New subfamily of the stag beetles (Coleoptera: Scarabaeoidea: Lucanidae) from the Mesozoic of Mongolia, and its position in the system of the Superfamily. Paleontological Journal 34, Suppl. 3:S327–S330. Nikolajev, G. V. 2002. O veroyatioy prinadlezhiosti k podsemeystvu Pleocominae novogo roda plastinchatoysykh zhukov iz nizhiego mela Mongolii. Paleontologicheskiy Zhurnal 2002(3):51–55. [Translation: Nikolajev, G. V. 2002. To possible assignment of a new genus of lamellicorn beetles from the Lower Cretaceous of Mongolia to the subfamily Pleocominae (Coleoptera, Scarabaeidae). Paleontological Journal 36:279– 282.]. Nikolajev, G. V. 2004. Mezozoyskiy etap v evolyutsii podsemeystva Aclopinae (Coleoptera, Scarabaeidae). Tethys Entomological Research 10:33–46. Nikolajev, G. V. 2005a. Novyy rod plastinchatousykh zhukov (Coleoptera, Scarabaeidae) iz nizhiemelovogo mestonakhozhdeniya Baysa v Zabaykal’e. Biologicheskie Nauki Kazakhstana 1:117–120. Nikolajev, G. V. 2005b. Novyy rod podsemeystva Aclopinae (Coleoptera, Scarabaeidae) iz verkhney yury Kazakhstana. Trudy Instituta Zoologii, Ministerstvo Obrazovaniia i Nauki Respubliki, Zoologo-ekologicheskie Issledovaniya 49:112–114. Nikolajev, G. V. 2005c. Plastinchatousye zhuki podsemeystva Glaphyrinae (Coleoptera, Scarabaeidae) v nizhnem mele Zabaykal’ya. Zhivotnyy mir Dal’nego Vostoka 5:69–78. Nikolajev, G. V. 2005d. Novyy rod triby Hybosorini (Coleoptera, Scarabaeidae) iz mezozoya Azii. Tethys Entomological Research 11:27–28. Nikritin, L. M. 1977. Infraotryad Scarabaeiformia. Trudy Paleontologicheskogo Instituta Akademiya Nauk SSSR 161 (Arnol’di, L. V., Zherikhin, V. V., Nikritin, L. M, and A. G. Ponomarenko: Mezozoiskie zhestkokrylye):119–130. [Translation: Nikritin, L. M. 1991. Infraorder Scarabaeiformia. [pp. 162–176]. In: Mesozoic Coleoptera. L. V. Arnol’di, V. V. Zherikhin, L. M. Nikritin, and A. G. Ponomarenko (eds.). Oxonian Press, New Delhi.]. Northwood, C. 2005. Early Triassic coprolites from Australia and their palaeobiological significance. Palaeontology 48:49–68. Ocampo, F. C. 2002. Procoilodes adrastus Ocampo, a new genus and species of Miocene hybosorid in amber from the Dominican Republic (Coleoptera: Scarabaeoidea: Hybosoridae). Coleopterists Bulletin 56:123–126. Oppenheimer, J. R., and J. Begum. 1978. Ecology of some dung beetles (Scarabaeidae and Aphodiidae) in Dacca District. Bangladesh Journal of Zoology 6:23–29. Papier, F., A. Nel, L. Grauvogel-Stamm, and J.-C. Gall. 2005. La diversite´ des Coleoptera (Insecta) du Trias dans le nord-est de la France. Geodiversitas 27:181–199. Parker, A. R. 2005. A geological history of reflecting optics. Journal of the Royal Society Inferface 2:1–17. Parker, A. R., and D. R. McKenzie. 2003. The cause of 50 million-year-old colour. Proceedings of the Royal Society London (B), Supplement (Biology Letters) 270:S151–153. Paulian, R. 1976. Three fossil dung beetles (Coleoptera: Scarabaeidae) from the Kenya Miocene. Journal of the East Africa Natural History Society and National Museum 31(158):1–4. Piton, L. 1940. Paleóntologie du gisement eóce`ne de Menat (Puy-de-Doˆme) (Flore et Faune). Lechevalier, Paris, France. 306 pp. Poinar, G., and R. Poinar. 1999. The Amber Forest. A Reconstruction of a Vanished World. Princeton University Press, Princeton, NJ. Ponel, P. 1993. Les coleópte`res du Quaternaire: leur roˆle dans la reconstruction des ´ cologie 24:5–16. paleóclimats et des paleóećosyste`mes. Bulletin d’E Ponomarenko, A. G. 2003. Ecological evolution of beetles (Insecta: Coleoptera). Acta Zoologica Cracoviensia 46(Suppl.):319–328. Porch, N. (in press). Late Pleistocene beetle records from Australia. In: Encyclopedia of Quaternary Science. S. A. Elias (ed.). Elsevier, Amsterdam, Netherlands.

140


Porch, N., and S. Elias. 2000. Quaternary beetles: A review and issues for Australian studies. Australian Journal of Entomology 39:1–9. Puhlmann, P. W., F. W. Sander, and W. Steiner. 1991. Ein Ka¨ferfund aus dem holoza¨nen Travertin von Weimar/Thu¨ringen. Zeitschrift fu¨r Geologische Wissenschaften 19:591–592. Prasad, V., C. A. E. Stro¨mberg, H. Alimohammadian, and A. Sahni. 2005. Dinosaur coprolites and the early evolution of grasses and grazers. Science 310:1177–1180. Quiel, G. 1911. Bemerkungen u¨ber Coleopteren aud dem baltischen Bernstein. Berliner Entomologische Zeitschrift 55:181–192. Ratcliffe, B. C., and F. C. Ocampo. 2001. Tyrannasorus rex Ratcliffe and Ocampo, a new genus and species of Miocene hybosorid in amber from the Dominican Republic (Coleoptera: Scarabaeoidea: Hybosoridae). Coleopterists Bulletin 55:351–355. Ratcliffe, B. C., D. M. Smith, and D. Erwin. 2005. Oryctoantiquus borealis, new genus and species from the Eocene of Oregon, U.S.A., the world’s oldest fossil dynastine and largest fossil scarabaeid (Coleoptera: Scarabaeidae: Dynastinae). Coleopterists Bulletin 59:127–135. Schaal, S., and W. Ziegler (eds.). 1992. Messel. An Insight into History of Life and of the Earth. Claredon Press, Oxford, U.K. Schlu¨ter, T., and F. von Gnielinski. 1987. The East African Copal its geologic, stratigraphic, palaeontologic significance and comparison with fossil resins of similar age. Occasional Paper, National Museums of Tanzania 8:1–32. Scholtz, C. H., D. J. Browne, and J. Kukalova´-Peck. 1994. Glaresidae, archaeopteryx of the Scarabaeoidea (Coleoptera). Systematic Entomology 19:259–277. Scholtz, C. H., and S. L. Chown. 1995. The evolution of habitat use and diet in the Scarabaeoidea: a phylogenetic approach [pp. 355–374]. In: Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. J. Pakaluk and S. A. S´lipin´ski (eds.). Muzeum i Instytut Zoologii PAN, Warszawa, Poland. Scholtz, C. H., and V. V. Grebennikov. 2005. Scarabaeiformia Crowson, 1960 [pp. 344–425]. In: Coleoptera, Beetles, 1: Morphology and Systematics (Archostemata, Adaphaga, Myxophaga, Polyphaga partim). Handbuch der Zoologie IV (38). R. G. Beutel and R. A. B. Leschen (eds.). Walter de Gruyter, Berlin, Germany. Schweigert, G. 2003. Alpenbock und Hirschka¨fer im Plioza¨n von Willershausen. Fossilien 20:178–182. Scott, A. C., J. Stephenson, and W. G. Chaloner. 1992. Interaction and coevolution of plants and arthropods during the Palaeozoic and Mesozoic. Philosophical Transactions of the Royal Society London (B) 335:129–165. Smith, D. M., A. Cook, and C. R. Nufio. 2006. How physical characteristics of beetles affect their fossil preservation. Palaios 21:305–310. Spahr, U. 1981. Systematischer Katalog der Bernstein- und Kopal-Ka¨fer (Coleoptera). Stuttgarter Beitra¨ge zur Naturkunde, Serie B (Geologie und Palaöntologie) 80:1– 107. Stankiewicz, B. A., D. E. G. Briggs, R. P. Evershed, and I. J. Duncan. 1997. Chemical preservation of insect cuticle from the Pleistocene asphalt deposits of California, U.S.A. Geochimica et Cosmochimica Acta 61:2247–2252. Stenkiewicz, B. A., H. N. Poinar, D. E. G. Briggs, R. P. Evershed, and G. O. Poinar. 1998. Chemical preservation of plants and insects in natural resins. Proceedings of the Royal Society London (B) 256:641–647. Stock, C., and J. M. Harris. 1992. Rancho La Brea. A record of Pleistocene life in California. 7th Ed. Natural History Museum of Los Angeles County, Los Angeles. Sun, B. 1995. Shanwang Fossils. Science Press, Beijing, China. Tanaka, H. 1985. Dung beetles found on raptor pellets. Lamellicornia 1:28–29 [in Japanese]. Thulborn, R. A. 1991. Morphology, preservation and palaeobiological significance of dinosaur coprolites. Palaeogeography, Palaeoclimatology, Palaeoecology 83:341– 366. Ueda, K. 1989. A Miocene fossil of long-armed scarabaeid beetle from Tottori, Japan. Bulletin of the Kitakyushu Museum of Natural History 9:105–110, pl. 1.


141

Valde´s, P. 1999. Primeros insectos fo´siles cubanos (Coleoptera: Scarabaeidae: Dytiscidae). Cocuyo 9:17–18. Vincent, P., M. Aubert, P. Boivin, J.-M. Cantagrel, and J.-F. Lenat. 1977. Dećouverte d’un volcanisme paleóce`ne en Auvergne: les maars de Menat et leurs annexes; e´tude geólogique et geóphysique. Bulletin de la Socie´te´ Geólogique de France series 7(19): 1057–1070. Vinson, J. 1951. Le cas des Sisyphes Mauriciens (Insecta Coleoptera). Proceedings of the Royal Society of Arts and Sciences of Mauritius 1:105–122, pl. 5. Wahl, A. M., A. J. M. Martin, and S. T. Hasiotis. 1998. Vertebrate coprolites and coprophagy traces, Chincle Formation (Late Triassic), Petrified Forest National Park, Arizona [pp. 144–148]. In: National Park Service Paleontological Research, Technical Report NPS/NRGRD/GRDTR-98/01. V. L. Santucci and L. McClelland (eds.). National Park Service, Geological Resources Division, Lakewood, CO. Wappler, T. 2003. Die Insekten aus dem Mittel-Eoza¨n des Eckfelder Maares, Vulkaneifel. Mainzer Naturwissenschaftliches Archiv Beiheft 27:i–viii,1–234, 18 pls. Yang, H., and S. Yang. 1994. The Shanwang fossil biota in eastern China—a Miocene Konservat-Lagersta¨tte in lacustrine deposits. Lethaia 27:345–354. Young, O. P. 1981. The attraction of Neotropical Scarabaeinae (Coleoptera: Scarabaeidae) to reptile and amphibian fecal material. Coleopterists Bulletin 35:345–348. Zhang, H.-C. 1997. Early Cretaceous Insects from the Dalazi Formation of the Zhixin Basin, Jilin Province, China. Palaeoworld 7:75–103. Zhang, J.-F., and H.-C. Zhang. 2003. Insects and spiders [pp. 58–67]. In: The Jehol Biota. Chang Mee-mann et al. (eds.). Shanghai Scientific and Technical Publishers, China. Zhang, Z., P. M. Barrett, and J. Hilton. 2003. An exceptionally preserved Lower Cretaceous ecosystem. Nature 421:807–814. Zhao, R., B. Han, and J. G. Liou. 2002. Timing of basalt flows in the Shanwang Basin, Shandong Peninsula, eastern China. Eos, Transactions American Geophysical Union 83(47), Fall Meeting, Suppl.:Abstract V62B-1406. Zherikhin, V. V. 1999. Preface. Proceedings of the First International Palaeoentomological Conference, Moscow 1998:1. Zherikhin, V. V., M. B. Mostovski, P. Vrsansky, V. A. Blagoderov, and E. D. Lukashevich. 1999. The unique Lower Cretaceous locality Baissa and other contemporaneous fossil insect sites in North and West Transbaikalia. Proceedings of the First International Palaeoentomological Conference, Moscow 1998:185–191. .

Appendix. List of fossil Scarabaeoidea described since Krell’s (2000) catalog. Species based on single elytra in brackets. Familia TROGIDAE Subfamilia? Prototroginae Nikolajev 2000a: 63 (transl. p. 426) Prototrox Nikolajev 2000a: 65 (transl. p. 427) (type species by original designation: Prototrox transbaikalicus Nikolajev, 2000) Prototrox transbaikalicus Nikolajev 2000a: 65 (transl. p. 428) (L Cretaceous, holotype: Argun Formation, Semen Creek, Olengui River basin, Chita Region, Transbaikalia. Paratype: Barremian-Aptian, Bon-Tsagaan series, Bayan-Hongor Aymag, Mongolia) Familia LUCANIDAE Subfamilia Paralucaninae Nikolajev 2000b: S328 Paralucanus Nikolajev 2000b: S329 (type species by original designation: Paralucanus mesozoicus Nikolajev) Paralucanus mesozoicus Nikolajev 2000b: S330 (U Jurassic, Shara-Teg, Gov9-Altai9 Aymag, Mongolia)

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Subfamilia Lucaninae Protognathinus Chalumeau and Brochier 2001: 595 (type species by monotypy: Protognathinus spielbergi Chalumeau and Brochier) Protognathinus spielbergi Chalumeau and Brochier 2001: 595 (M Eocene, Messel, Germany) Familia PLEOCOMIDAE Subfamilia Cretocominae Nikolajev stat. nov. Cretocomini Nikolajev 2002: 53 (transl. p. 281) Cretocoma Nikolajev 2002: 54 (transl. p. 281) (type species by original designation: Cretocoma tologoica Nikolajev) Cretocoma tologoica Nikolajev 2002: 54 (transl. p. 281) (L Cretaceous, Shar-Tolgoy, IhBogd Mountain, Bayan-Hongor Aymag, Mongolia) Familia GLAPHYRIDAE Subfamilia Cretoglaphyrinae Nikolajev stat. nov. Cretoglaphyrini Nikolajev 2005c: 70 Cretoglaphyrus Nikolajev 2005c: 70 (type species by original designation: Cretoglaphyrus rohdendorfi Nikolajev, 2005) Cretoglaphyrus rhodendorfi Nikolajev 2005c: 72 (L Cretaceous, Baissa, Russia) [Cretoglaphyrus leptopterus Nikolajev 2005c: 73] (L Cretaceous, Semen, Russia) [Cretoglaphyrus transbaikalicus Nikolajev 2005c: 73] (L Cretaceous, Semen, Russia) [Cretoglaphyrus calvescens Nikolajev 2005c: 73] (L Cretaceous, Semen, Russia) Cretoglaphyrus olenguicus Nikolajev 2005c: 75 (L Cretaceous, Semen, Russia) Cretoglaphyrus zherikhini Nikolajev 2005c: 74 (L Cretaceous, Semen, Russia) Familia HYBOSORIDAE Jurahybosorus Nikolajev 2005d: 27 (type species by original designation: Jurahybosorus mongolicus Nikolajev) Jurahybosorus mongolicus Nikolajev 2005d: 27 (U Jurassic, Bayan-Teg, Mongolia) Procoilodes Ocampo 2002: 123 (type species by original designation: Procoilodes adrastus Ocampo). Procoilodes adrastus Ocampo 2002: 125 (Miocene, Dominican Amber) Tyrannasorus Ratcliffe and Ocampo 2001: 351 (type species by original designation: Tyrannasorus rex Ratcliffe and Ocampo) Tyrannasorus rex Ratcliffe and Ocampo 2001: 253 (Miocene, Dominican Amber) Familia SCARABAEIDAE Subfamilia Aclopinae Tribus Aclopini Cretaclopus Nikolajev 2004: 35 (type species by original designation: Geotrupoides longipes Ponomarenko, 1986) Juraclopus Nikolajev 2005b: 112 (type species: Juraclopus rohdendorfi Nikolajev) Juraclopus rohdendorfi Nikolajev 2005b: 113 (Fig. 2) (U Jurassic, Karatau, Kazakhstan). Nikolajev (2004) [as ‘in litt.’ and figured]. Tribus Holcorobeini (classified as tribe of Aclopinae by Nikolajev 2004) Mongolrobeus Nikolajev 2004: 37 (type species by original designation: Mongolrobeus zherikhini Nikolajev) Mongolrobeus zherikhini Nikolajev 2004: 37 (L Cretaceous; Dund-Uul, Bon-Tsagaan, Bayan-Hongor Aymag, Mongolia) Antemnacrassa Gomez Pallerola has been removed from synonymy with Holcorobeus Nikritin by Nikolajev (2004) who transferred the following species into this genus: A. nigrimontana (Nikolajev), A. picturata (Nikritin), A. vitimensis (Nikritin), A. maculata (Nikolajev), A. incerta (Ponomarenko), and A. punctata (Ponomarenko).


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[Antemnacrassa albomaculata Nikolajev 2004: 42] (L Cretaceous, Shar-Tologoy, Bayan-Leg, Mongolia). Antemnacrassa albosulcata Nikolajev 2004: 39 (U Jurassic, Karatau, Kazakhstan) [Antemnacrassa geminata Nikolajev 2004: 41] (L Cretaceous, Pad Semen, Russia). [Antemnacrassa magna Nikolajev 2004: 41] (L Cretaceous, Baissa, Russia). [Antemnacrassa nebulosa Nikolajev 2004: 41] (L Cretaceous, Baissa, Russia). Baisarabaeus Nikolajev 2005a: 117 (type species by original designation: Baisarabaeus rugosus Nikolajev) Baisarabaeus rugosus Nikolajev 2005a: 119 (L Cretaceous, Baissa, Russia) Subfamilia Dynastinae Oryctoantiquus Ratcliffe and Smith in Ratcliffe, Smith, and Erwin 2005: 128 (type species by original designation: Oryctoantiquus borealis Ratcliffe and Smith) Oryctoantiquus borealis Ratcliffe and Smith in Ratcliffe, Smith, and Erwin 2005: 130 (M Eocene, Clarno Formation, West Branch Creek, Oregon, U.S.A.)