identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03E287E2FFFAFFD157C5066E6144FAC6.text	03E287E2FFFAFFD157C5066E6144FAC6.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Geotrupidae	<div><p>Geotrupidae: CRETOGEOTRUPINAE Nikolajev 1996 (L Cretaceous)</p> <p>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.</p> </div>	https://treatment.plazi.org/id/03E287E2FFFAFFD157C5066E6144FAC6	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Krell, Frank- Thorsten	Krell, Frank- Thorsten (2006): Fossil Record And Evolution Of Scarabaeoidea (Coleoptera: Polyphaga). The Coleopterists Bulletin (mo 5) 60: 120-143, DOI: 10.1649/0010-065x(2006)60[120:fraeos]2.0.co;2
03E287E2FFFAFFD157C50192630BFB8A.text	03E287E2FFFAFFD157C50192630BFB8A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lucanidae	<div><p>Lucanidae: PARALUCANINAE Nikolajev 2000 b (U Jurassic)</p> <p>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.</p> </div>	https://treatment.plazi.org/id/03E287E2FFFAFFD157C50192630BFB8A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Krell, Frank- Thorsten	Krell, Frank- Thorsten (2006): Fossil Record And Evolution Of Scarabaeoidea (Coleoptera: Polyphaga). The Coleopterists Bulletin (mo 5) 60: 120-143, DOI: 10.1649/0010-065x(2006)60[120:fraeos]2.0.co;2
03E287E2FFFAFFD157C507296366F9E3.text	03E287E2FFFAFFD157C507296366F9E3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Pleocomidae Le Conte 1861	<div><p>Pleocomidae: CRETOCOMINAE Nikolajev 2002, stat. nov. (L Cretaceous)</p> <p>Diagnosis: Pronotum without leathery anterior border; vein R3(5 RA4) 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.</p> </div>	https://treatment.plazi.org/id/03E287E2FFFAFFD157C507296366F9E3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Krell, Frank- Thorsten	Krell, Frank- Thorsten (2006): Fossil Record And Evolution Of Scarabaeoidea (Coleoptera: Polyphaga). The Coleopterists Bulletin (mo 5) 60: 120-143, DOI: 10.1649/0010-065x(2006)60[120:fraeos]2.0.co;2
03E287E2FFFBFFD057F803B06141FDAD.text	03E287E2FFFBFFD057F803B06141FDAD.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Glaphyridae	<div><p>Glaphyridae: CRETOGLAPHYRINAE Nikolajev 2005 c, stat. nov. (L Cretaceous)</p> <p>Diagnosis: Mandibles and labrum exposed; eyes partly separated by canthus; mesocoxae contiguous; radius at anterior wing margin; vein R3(5 RA4) 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.</p> </div>	https://treatment.plazi.org/id/03E287E2FFFBFFD057F803B06141FDAD	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Krell, Frank- Thorsten	Krell, Frank- Thorsten (2006): Fossil Record And Evolution Of Scarabaeoidea (Coleoptera: Polyphaga). The Coleopterists Bulletin (mo 5) 60: 120-143, DOI: 10.1649/0010-065x(2006)60[120:fraeos]2.0.co;2
03E287E2FFFBFFD057F802F16114FEA5.text	03E287E2FFFBFFD057F802F16114FEA5.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Ochodaeidae	<div><p>Ochodaeidae: CRETOCHODAEINAE Nikolajev 1995 a (L Cretaceous–Oligocene)</p> <p>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 (5 RA4) 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.</p> </div>	https://treatment.plazi.org/id/03E287E2FFFBFFD057F802F16114FEA5	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Krell, Frank- Thorsten	Krell, Frank- Thorsten (2006): Fossil Record And Evolution Of Scarabaeoidea (Coleoptera: Polyphaga). The Coleopterists Bulletin (mo 5) 60: 120-143, DOI: 10.1649/0010-065x(2006)60[120:fraeos]2.0.co;2
03E287E2FFFBFFD057F800B76282FBCB.text	03E287E2FFFBFFD057F800B76282FBCB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Scarabaeidae Latreille 1802	<div><p>Scarabaeidae: CRETOSCARABAEINAE Nikolajev 1995 b (L Cretaceous)</p> <p>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.</p> </div>	https://treatment.plazi.org/id/03E287E2FFFBFFD057F800B76282FBCB	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Krell, Frank- Thorsten	Krell, Frank- Thorsten (2006): Fossil Record And Evolution Of Scarabaeoidea (Coleoptera: Polyphaga). The Coleopterists Bulletin (mo 5) 60: 120-143, DOI: 10.1649/0010-065x(2006)60[120:fraeos]2.0.co;2
03E287E2FFFBFFD057F806126169FB22.text	03E287E2FFFBFFD057F806126169FB22.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Scarabaeidae Latreille 1802	<div><p>Scarabaeidae: LITHOSCARABAEINAE Nikolajev 1992 (L Cretaceous)</p> <p>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.</p> </div>	https://treatment.plazi.org/id/03E287E2FFFBFFD057F806126169FB22	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Krell, Frank- Thorsten	Krell, Frank- Thorsten (2006): Fossil Record And Evolution Of Scarabaeoidea (Coleoptera: Polyphaga). The Coleopterists Bulletin (mo 5) 60: 120-143, DOI: 10.1649/0010-065x(2006)60[120:fraeos]2.0.co;2
03E287E2FFFBFFDF57F806C96244FD6C.text	03E287E2FFFBFFDF57F806C96244FD6C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Scarabaeidae Latreille 1802	<div><p>Scarabaeidae: PROTOTROGINAE Nikolajev 2000 a (L Cretaceous)</p> <p>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 2000 a) 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.</p> <p>Scarabaeidae: Melolonthinae: CRETOMELOLONTHINI Nikolajev 1998 (L Cretaceous) Diagnosis: Labrum not covered by clypeus; clypeus without anterior border; anterior border of pronotum leathery; Radius 1 (5 RA3) and R3 (5 RA4) apically approximated (but not fused); meso- and metatibia with single transverse carina; apical spurs of meso- and 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.</p> <p>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 (5 RA4) curving smoothly without joining radius (5 RA3). 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.</p> </div>	https://treatment.plazi.org/id/03E287E2FFFBFFDF57F806C96244FD6C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Krell, Frank- Thorsten	Krell, Frank- Thorsten (2006): Fossil Record And Evolution Of Scarabaeoidea (Coleoptera: Polyphaga). The Coleopterists Bulletin (mo 5) 60: 120-143, DOI: 10.1649/0010-065x(2006)60[120:fraeos]2.0.co;2
03E287E2FFF6FFDD56C3037960C9FBCF.text	03E287E2FFF6FFDD56C3037960C9FBCF.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Scarabaeoidea sensu Crowson 1981	<div><p>Paleoecology of Scarabaeoidea</p> <p>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.</p> <p>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 (1997 b) 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.</p> </div>	https://treatment.plazi.org/id/03E287E2FFF6FFDD56C3037960C9FBCF	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Krell, Frank- Thorsten	Krell, Frank- Thorsten (2006): Fossil Record And Evolution Of Scarabaeoidea (Coleoptera: Polyphaga). The Coleopterists Bulletin (mo 5) 60: 120-143, DOI: 10.1649/0010-065x(2006)60[120:fraeos]2.0.co;2
