identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
45227F36FFB8FFB5B5BA2709FB8EF8FA.text	45227F36FFB8FFB5B5BA2709FB8EF8FA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chalcoderus Erichson 1847	<div><p>Chalcoderus Erichson, 1847, status restored</p><p>Chalcoderus Erichson, 1847: 763 .</p><p>Chalcoderus Erichson (Lacordaire 1855: 109) (synonymy with Onthophagus Latreille, 1802).</p><p>Chalcoderus Erichson (Harold 1867: 23) (citation as synonym of Onthophagus Latreille).</p><p>Chalcoderus Erichs. (Harold 1869: 1024) (citation as synonym of Onthophagus Latreille).</p><p>Chalcoderus Erichson (d’Orbigny 1914: 618) (citation as synonym of Onthophagus Latreille).</p><p>Chalcoderus Erichson (Boucomont 1914: 272) (citation).</p><p>Chalcoderus Er. (Boucomont &amp; Gillet 1927: 118) (citation as synonym of Onthophagus Latreille).</p><p>Chalcoderus Erichson (Balthasar 1963: 153) (citation as synonym of Onthophagus Latreille).</p><p>Chalcoderus Erichson (Palestrini 1982) (suggested classification as nomen oblitum after citation and subsequent designation of the type species).</p><p>Chalcoderus Erichson, 1847 (Branco 2011: 12, 18) (citation).</p><p>Parascatonomus (Morettoeus) Ochi &amp; Kon, 2023: 578 (description) new objective synonymy.</p><p>Type species: Ateuchus maculatus Fabricius, 1801 (cited as “ O. maculatus F.”), subsequent designation by Palestrini (1982: 101) 2 under Article 69.1.1. (Branco 2011: 12).</p><p>Original description (Erichson 1847): “1. Chalcoderus, hier ist das Kinn so lang als die Stämme der Maxillen, Schmal; der Kopf unbewehrt, das Halsschild hoch und gleichmässig gewölbt, hinten ungerandet; die hinteren Füsse etwas breit. Hierher gehört At. maculatus F., Onthoph. signatipennis Dej. Lap. ”</p><p>Redescription</p><p>Body. Not distinctly constricted between pronotum and elytra. Pronotum with very dense, short, fine, transparent and recumbent setae posterior to each granule, elytra with short, fine, translucent and recumbent setae posterior to each granule or puncture (these setae frequently erased—x40 magnification). Ventral surface with short or moderately long, not very dense, pale and recumbent setae on the episterna, the lateral areas of the metasternum and the sides of the ventrites.</p><p>2 Palestrini (1982) was proposed by M. Zunino and redacted under his supervision (C. Palestrini, personal communication). In the MNHN collections, we found a specimen of C. maculatus identified by Zunino and labelled as follows: “ SÉNÉGAL [Mali] / KOULIKORO / J. VUILLET 1910 (light blue) // Museum Paris / ex Coll. / R. Oberthur (light blue) // Onthophagus / ( Parascatonomus) / maculatus (F.) / det. Zunino 1982 (handwritten by Zunino) // WSD00258500 // Chalcoderus maculatus / (Fabricius, 1801) / P. Moretto det. 2025”. The publication date of Palestrini’s paper and the identification date of this specimen are the same (1982). We can confirm that it is a specimen of Ateuchus maculatus Fabricius, 1801 and we retain it as the reference for the species Palestrini (1982) called Parascatonomus maculatus (Fabricius, 1801) .</p><p>Head. Flattened, totally unarmed, narrow, more or less elongate and parabolic in shape, more or less truncated at apex; proportionally narrower and more elongate than all other African and Asian Onthophagini species 3; genae protruding very little or not, clypeo-genal sutures present but very narrow and indistinct.Apical margin of epipharynx strongly emarginate (Ochi &amp; Kon 2023: 531, Fig. 1h). Antennae with 9 articles, scape short, invisible dorsally, segments 3 to 6 of different sizes, the first article of the club cup-shaped, the other two progressively smaller.</p><p>Thorax. Pronotum metallic coloured or black, simple, convex or very convex, covered by convex or very convex, very dense, elongate granules; sometimes with a longitudinal median depression on the posterior part of the disc, distinctly depressed above the erased and absent posterior angles; anterior angles sharp, lateral side of the pronotum bordered, base of the pronotum angulate in the middle, not bordered except sometimes shortly in the median angle; erased posterior angles theoretically located at the termination of the lateral edges of the pronotum; prosternal foveae clear; elevation of metasternum not well defined.</p><p>Elytra. Light coloured with dark spots and/or patches; interstriae with fine and not very dense granules or punctures, or with very dense granulate punctures more or less confluent and vermiculate ( C. maculatus and C. signatipennis).</p><p>Legs. Basal tooth (tooth 1) of protibia strongly reduced, teeth 2 and 3 more elongate, spatulate and strongly developed; apical tooth directed obliquely forward; terminal mobile spur of the protibia spatulate in the male, normal in the female; insertion of the protarsus just behind the articulation of the terminal spur, on the inferior side of the tibia.</p><p>Pygidium. Base of the pygidium bordered, angulate in the middle, with simple punctures, fairly dense to very dense but less dense near apex, denser and more or less raspy near base.</p><p>Aedeagus. See Ochi &amp; Kon (2023: 581, Figs 1 b-d, 2b-d, 3b-d).</p><p>Rationale: Transferring the type species of Chalcoderus Erichson, 1847, to Parascatonomus Paulian, 1932, created a problem with the priority of names (Palestrini 1982). To maintain taxonomic stability, Palestrini (1982) suggested retaining the subgeneric name as Parascatonomus by regarding Chalcoderus as a nomen oblitum under Article 23.9. However, the validity of Chalcoderus is restored once it is considered to belong to a genus or subgenus distinct from Parascatonomus (Branco 2011: 18), particularly as by designating the type species, Palestrini (1982) made Chalcoderus an available name from Erichson (1847). Thus, by virtue of having the same type species, Parascatonomus (Morettoeus) Ochi &amp; Kon, 2023, becomes an objective junior synonym of Chalcoderu s (Article 61.3.3.).</p><p>Similarities: Chalcoderus and Parascatonomus share some morphological characters that are unique in the Onthophagini. Mentum usually longer than wide. Epipharynx not strongly transverse, chaetopariae not arising from antero-lateral corner and both almost parallel with fine bristles. Clypeo-genal sutures present but very narrow and indistinct. Angular postero-medial edge of the pronotum projected posteriorly. Basal tooth (tooth 1) of protibia more or less reduced or absent, teeth 2 and 3 more elongate, spatulate and strongly developed, apical tooth directed obliquely forward; insertion of the protarsi positioned behind or just beside the articulation of the terminal spur, on the inferior side of the tibia. Base of the pygidium strongly angulate in the middle. Shared shape of the aedeagus, parameres with a baso-lateral elevation, medio-lateral and apical teeth present, shared shape of the copulatory lamella (Ochi &amp; Kon 2017, 2023).</p><p>Differences: Parascatonomus subgenera show a range of clear differences from one another and from Chalcoderus . Given the impossibility of clearly describing the external morphological differences between the genus Chalcoderus and the genus Parascatonomus taken as a whole, it was decided to provide diagnostic characters for each of the six accepted subgenera of Parascatonomus, emphasizing how each differs from the genus Chalcoderus . As presented below, the morphological external characters have been described following the published descriptions (Ochi &amp; Kon 2017; Ochi et al. 2021) and after examination of specimens belonging to several different species of each subgenus (except for Parascatonomus (Necramator), klapperichi group (Ochi &amp; Kon 2017)). All are distributed within Asia with some ranges extending into the eastern Palaearctic. To the contrary, the known distribution of Chalcoderus is restricted to Africa. For the shape of the epipharynx, the parameres and the copulatory sclerites, see Ochi &amp; Kon (2017), Ochi et al. (2021) and Ochi &amp; Kon (2023).</p></div>	https://treatment.plazi.org/id/45227F36FFB8FFB5B5BA2709FB8EF8FA	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFBEFFB2B5BA268CFD15FAC0.text	45227F36FFBEFFB2B5BA268CFD15FAC0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Parascatonomus (Laonthophagus) Frey 1971	<div><p>Parascatonomus (Laonthophagus) Frey, 1971 (based on simultaneously collected males and females)</p><p>Body. General shape similar to P. ( Parascatonomus), strongly convex, dark brown. Fine medium recumbent setae on the sides and close to the base of elytra.</p><p>Head. Large, genae strongly protruding, clypeus comprising two long, strong and acute upturned teeth in male (shorter in female); frontal carina straight, well developed, protruding angularly in the middle; vertex extended backward in a very long and very large, parallel-sided lamina (as large as the head at the level of the eyes in males, shorter and trapezoidal in female), strongly toothed at the posterior angles.</p><p>Thorax. Pronotum depressed behind lamina of the vertex but very convex with a strong tubercle protruding forward in the middle, tubercle followed by a large, shallow, densely granular depression in the middle of the disc, the granules elongate inside the depression. A large area extending on each side of the depression to the posterior border of the pronotum, showing scaly-shaped granules or simple fine punctures on the largest specimens. No depression above posterior angles. Elevation of metasternum well developed.</p><p>Elytra. Striae fine, interstriae convex, shagreened on the disc of elytra, smooth on the sides, with very fine and sparse granules on the disc and simple, fine, not very dense punctation on the lateral interstriae.</p><p>Legs. Protibia enlarged, basal tooth short, teeth 2 to 4 strong, long and acute; insertion of pro-tarsi beside the articulation of the terminal spur; inner border of the protibia extended in a tooth at apex in male; terminal mobile spur of the protibia not modified.</p><p>Pygidium. With dense, medium-sized punctures.</p></div>	https://treatment.plazi.org/id/45227F36FFBEFFB2B5BA268CFD15FAC0	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFBEFFB2B5BA24A8FD1EFD74.text	45227F36FFBEFFB2B5BA24A8FD1EFD74.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Parascatonomus (Parascatonomus) Paulian 1932	<div><p>Parascatonomus (Parascatonomus) Paulian, 1932</p><p>Body. Elongate, distinctly constricted between pronotum and elytra, sides parallel, mostly smooth and without setae dorsally (except laterally on elytra in P. fujiokai), entirely black. Long, erect, orange setae on the prosternum, the mesepisternes, the anterior declivity and the lateral areas of the metasternum</p><p>Head. Wider than long, clypeus deeply emarginate with a finger-like tooth in the middle of the emargination, genae very protruding.</p><p>Thorax.Anterior angles of pronotum not protruding, square or rounded, more or less largely granulate (flattened granules), or only near the anterior angles, or totally smooth; not depressed above posterior angles; elevation of metasternum well developed.</p><p>Elytra. Striae quite fine, interstriae with very fine punctures or totally smooth.</p><p>Legs. Pro-tibiae very enlarged, basal tooth proportionally small, teeth 2 and 3 very large and strong, terminal tooth narrower and acute, terminal mobile spur of the protibia strongly curved downwards and acute in both sexes. Insertion of pro-tarsi before the articulation of the terminal spur, on the inferior side of the tibia.</p><p>Pygidium. With dense to sparse, fine punctures.</p></div>	https://treatment.plazi.org/id/45227F36FFBEFFB2B5BA24A8FD1EFD74	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFBEFFB2B5BA21D0FC2EF800.text	45227F36FFBEFFB2B5BA21D0FC2EF800.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Parascatonomus (Peculiaricerus) Ochi & Kon 2017	<div><p>Parascatonomus (Peculiaricerus) Ochi &amp; Kon, 2017</p><p>Body. General shape similar to P. ( Parascatonomus). Head and pronotum with a metallic lustre, elytra black. Pronotum and elytra with recumbent setae.</p><p>Head. Wider than long, polygonal; genae strongly protruding; clypeus with a large upturned process, or, deeply and widely notched in the middle, with a large upturned process at the middle of the notch; a strong tubercle in place of the clypeal suture; vertex with a blunt keel. Segments of the club of the antennae showing a very complicated shape.</p><p>Thorax. Pronotum entirely and densely granular except for a small finely punctate area near base in the middle; anterior part with an elongate smooth tubercle or a fine longitudinal carina in the middle; indistinctly depressed above the posterior angles; elevation of metasternum well developed.</p><p>Elytra. Striae fine, interstriae flat, shagreened, with not very dense granules or rough punctures.</p><p>Legs. Protibiae moderately enlarged, distinctly incurved in apical third, basal tooth small, teeth 2 and 3 large and fairly strong, terminal tooth narrower and acute; insertion of pro-tarsi just beside or just before the articulation of the terminal spur, which is slightly curved backwards and acute in both sexes.</p><p>Pygidium. Shagreened, with dense or very dense, medium punctures.</p></div>	https://treatment.plazi.org/id/45227F36FFBEFFB2B5BA21D0FC2EF800	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFBFFFB3B5BA27A4FD3CFA26.text	45227F36FFBFFFB3B5BA27A4FD3CFA26.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Parascatonomus (Granulidorsum) Ochi & Kon 2017	<div><p>Parascatonomus (Granulidorsum) Ochi &amp; Kon, 2017</p><p>Body. Not elongate, strongly convex, small to medium sized. Dorsum usually dull with a metallic lustre, elytra rarely testaceous. Pronotum, elytra and pygidium with recumbent setae. In agreement with Ochi &amp; Kon (2017) this “subgenus is unique in having the pronotum and elytra wholly covered with very aggregated coarse granules, the elytral granules mostly forming several longitudinal sinuous rows, and the pronotal granules also partly arranged so”.</p><p>Head. Normal size, frequently totally flattened, elongate and parabolic in shape, sometimes truncated or emarginate at apex, sometimes arcuate with carinae present on frons and vertex.</p><p>Thorax. A distinct depression above posterior angles of pronotum present or not, sometimes a depression near the base, in the middle. Elevation of metasternum present.</p><p>Legs. Basal tooth of the fore tibia totally reduced or normally sized, teeth 2 and 3 normal, apical tooth frequently very narrow and acute. Terminal mobile spur of the protibia not modified in both sexes.</p><p>Pygidium. Covered with dense fine granules.</p></div>	https://treatment.plazi.org/id/45227F36FFBFFFB3B5BA27A4FD3CFA26	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFBFFFB3B5BA21BCFC95F82F.text	45227F36FFBFFFB3B5BA21BCFC95F82F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Parascatonomus (Necramator) Ochi & Kon 2017	<div><p>Parascatonomus (Necramator) Ochi &amp; Kon, 2017</p><p>Parascatonomus (Necramator) nitidus group (4–14 mm)</p><p>Body. Elongate and oval, distinctly constricted between pronotum and elytra. Entirely black, black with metallic lustre, or only the pronotum with a strong metallic colour.</p><p>Head. Usually large with angularly protruding cheeks. Clypeus variable, pointed, sometimes with a median tooth, or rounded; frontal carina present or absent; sometimes a more or less elevated carina on the vertex.</p><p>Thorax. Pronotum simply formed, regularly convex, usually glabrous with simple punctation (rarely hairy, entirely covered with very large, dense granules and / or punctures, or, large vermiculated punctures).</p><p>Elytra. Usually glabrous with simple punctation on interstriae. (rarely hairy with fine granules on interstriae).</p><p>Legs. Protibia with four well developed teeth.Terminal mobile spur of the protibia slightly spatulate or thickened in male, or not.</p><p>Pygidium. Glabrous with simple, medium to fine punctures.</p></div>	https://treatment.plazi.org/id/45227F36FFBFFFB3B5BA21BCFC95F82F	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFBFFFB3B5BA24E0FCB9FC1F.text	45227F36FFBFFFB3B5BA24E0FCB9FC1F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Parascatonomus (Pseudonthophagus) Balthasar 1959	<div><p>Parascatonomus (Pseudonthophagus) Balthasar, 1959</p><p>Body. Elongate, moderately ( P. penicillatus) to strongly convex, distinctly constricted between pronotum and elytra. Head, pronotum and pygidium metallic coloured. Elytra light brown with two curved rows of more or less well developed and coalescing dark brown patches, a more or less developed transverse spot or darkened area just before the apex of the elytra and a spot at the base of interstria 5 (sometimes missing); juxta-sutural interstriae with a weak metallic lustre. Pronotum and elytra with long, dense, white recumbent setae.</p><p>Head. Normally large, not narrow, flattened, elongate and parabolic in shape, truncated at apex; genae moderately protruding. In males, clypeus flat, simply punctured, “forehead” moderately depressed (effaced area between frontal suture and vertex), vertex with a blunt keel. In females, clypeus flat, almost entirely densely rugose, “forehead” with a deep, smooth depression (effaced area between frontal suture and vertex), vertex elevated in a triangular or rounded lamina or a horn just behind the depression.</p><p>Thorax. Pronotum very densely covered in contiguous, strongly-flattened granules, scaly on the disc, each granule followed by a very fine, simple, setose puncture; pronotum simply convex in male, depressed behind the lamina in females; disc with a more or less longitudinal shallow depression near the base in the middle; also a distinct depression above posterior angles, topped with a fine “paintbrush” of erected bristles; scutellum visible, even if very minute; elevation of metasternum present.</p><p>Elytra. Striae fine, interstriae convex, shagreened, with fairly dense, fine raspy punctures on top, becoming simple, less fine and denser on the sides.</p><p>Legs. Protibia not enlarged, basal tooth strongly reduced, second tooth somewhat reduced, teeth 3 and 4 fairly strong and spatulate; inner border extended in a small tooth at apex in male (at least in P. penicillatus); terminal mobile spur not modified in both sexes.</p><p>Pygidium. With fine, dense to not very dense punctures.</p></div>	https://treatment.plazi.org/id/45227F36FFBFFFB3B5BA24E0FCB9FC1F	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFBCFFB0B5BA24E0FF2CFDA3.text	45227F36FFBCFFB0B5BA24E0FF2CFDA3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Parascatonomus (Necramator) tricornis (Wiedemann 1823)	<div><p>Parascatonomus (Necramator) tricornis group (7.5–17 mm)</p><p>Body. Not elongate.</p><p>Head. Differs from the nitidus group by the diverse ornaments of the head: frontal carina absent or present or replaced by a tubercle in the middle; vertex with a transverse carina, a horn or a tubercle, or a lamina extended into a horn at its tip; rarely, antennae club segments strongly elongate.</p><p>Thorax. Pronotum with at least a simple depression anteriorly, or with a thoracic protrusion overlying a more or less large and deep depression, frequently a more or less distinct longitudinal furrow on the disc and at least the main part covered with very dense contiguous flattened granules; lateral fovea present.</p><p>Elytra. With simple, rough or granulose punctation.</p><p>Pygidium. Usually glabrous with simple, large to medium punctures, sometimes with sparse, pale recumbent setae.</p></div>	https://treatment.plazi.org/id/45227F36FFBCFFB0B5BA24E0FF2CFDA3	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFBCFFB1B5BA2630FE5BFCC7.text	45227F36FFBCFFB1B5BA2630FE5BFCC7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Parascatonomus (Necramator) klapperichi (Balthasar 1952)	<div><p>Parascatonomus (Necramator) klapperichi group (4.3–8 mm)</p><p>Diagnosis following Ochi &amp; Kon (2017). This small group of two species differs from the others by the small size, the compact, convex body not constricted between pronotum and elytra; the head and pronotum black; the elytra bicoloured, black with testaceous design, or testaceous with black design. Head totally unarmed. Protibia with the basal tooth reduced (at least in P. takahashii Masumoto, Ochi &amp; Hanboonsong, 2013).</p><p>The description of the genus Uenoeus Ochi &amp; Kon, 2017, and the formal division of Parascatonomus into five subgenera by Ochi &amp; Kon (2017) were largely inspired by the preliminary work of Huijbregts &amp; Krikken (2009) who simply defined functional groups for the species. Following the transfer of a sixth subgenus to Parascatonomus (Ochi et al. 2021), it seemed logical to describe Morettoeus Ochi &amp; Kon, 2023 as a seventh subgenus, particularly since one member of group 15 of Afrotropical Onthophagus (d’Orbigny 1913) had already been transferred to the genus Parascatonomus by Palestrini (1982). It is probable that this material examined by Palestrini (1982) was also examined for the morphological phylogeny of Tarasov &amp; Solodovnikov (2011: Fig. 11, clade F, X) in which the transferred species (“ O. (Parascatonomus) maculatus ”) was nested in the Parascatonomus clade. However, the congeneric status of Chalcoderus and Parascatonomus species is questionable since this phylogeny showed poor resolution of cladistic relationships between Chalcoderus, Uenoeus and the subgenera of Parascatonomus in clades F and X (all cited as species of Onthophagus). For instance, a species of the genus Uenoeus was nested within species of the genus and subgenus Parascatonomus, whereas species of the subgenus Necramator were grouped with those of other subgenera within three different subclades, one containing “ O [nthophagus] maculatus ”. Furthermore, there are strong, overall, morphological differences between Afrotropical Chalcoderus and Oriental Parascatonomus subgenera.</p><p>Nevertheless, it should be noted that a further two species, O. coeruleicollis Arrow, 1907, from India and Onthophagus regalis Arrow, 1907, from Sri Lanka (Arrow 1931: 185–186, pl. 1: fig. 8–9), appear close to Chalcoderus species. Concerning O. coeruleicollis, Arrow (1907: 430) noted that “This species and the following one [ O. regalis] belong to a group of which several have been described from tropical Africa, and of which O. maculatus, F., is the type.” However, they exhibit characters that differentiate them from species of the genus Chalcoderus within which, they will not currently be included pending detailed examination. Both O. regalis and O. coeruleicollis have long, pale, oblique setae on the head, the pronotum and the underside. Setae are also long and dense on the elytra of O. regalis but shorter and less dense on the elytra of O. coeruleicollis . They have a brush of long, erected setae on the anterior border of the postero-lateral depressions of the pronotum. The head is proportionally larger and Arrow (1907) notes that in O. coeruleicollis, “the posterior margin of the head forms a sharp edge but is not produced”. They also exhibit differences in the mobile protibial spur. Arrow further indicates a possible sexual character for O. coeruleicollis: “a very minute tubercle sometimes traceable near the middle of the head probably indicates the male”. Bacchus (1978) states that each type series contains a male, which we have not had the opportunity to study.</p><p>Although it is possible that a revision that includes these two species might expand the known distribution of Chalcoderus to the western Oriental region, the possibility of convergent morphology due to similarities in habits cannot currently be discounted. A molecular phylogeny might help to resolve the status of relationships between these two species, Parascatonomus subgenera, Uenoeus and Chalcoderus . It would also assist in ensuring there are no discrepancies between phylogeny and classification based on the phenetic characters described above.</p><p>Taxonomy—species</p><p>We are confident that all of the species formerly included in Afrotropical Onthophagus group 15 of d’Orbigny (1913, 1915) belong in Chalcoderus . Therefore, we formally transfer the following nine currently recognised species to the genus, Chalcoderus Erichson, 1847: Onthophagus (s. l.) foliiceps Quedenfeldt, 1884, Onthophagus (s. l.) graphicus Wallengren, 1881, Onthophagus (s. l.) inermiceps d’Orbigny, 1905, Onthophagus (s. l.) longiceps d’Orbigny, 1904, Onthophagus (s. l.) planaticeps d’Orbigny, 1913, Onthophagus (s. l.) semicroceus d’Orbigny, 1915, Onthophagus (s. l.) simpliciceps (d’Orbigny, 1905), Parascatonomus (Morettoeus) imbellis (d’Orbigny, 1905), Parascatonomus (Morettoeus) maculatus (Fabricius, 1801) . A tenth species ( Onthophagus signatipennis de Castelnau, 1840) is revalidated from synonymy with Ateuchus maculatus Fabricius, 1801, and its placement in Chalcoderus is restored. Onthophagus chloroderus d’Orbigny, 1905, is removed from the list of valid group 15 species and is, here, synonymized with Ateuchus maculatus Fabricius, 1801 (currently Chalcoderus maculatus (Fabricius, 1801)) .</p><p>The following is a complete and updated list of all known species that were former members of Afrotropical Onthophagus group 15 but are now placed in the genus Chalcoderus Erichson, 1847 . High resolution photographs of type material are provided for all of the valid species. Notes are provided for each species concerning its validity, status and synonyms. Lectotypes are designated where necessary. Special attention is paid to Ateuchus maculatus Fabricius, 1801, Onthophagus signatipennis de Castelnau, 1840, and Onthophagus chloroderus d’Orbigny, 1905, which are fully revised in order to correct confusion in their taxonomy and distributions as well as to precisely establish the type species of the genus Chalcoderus . The material available for the other species is insufficient to allow for their revision.</p></div>	https://treatment.plazi.org/id/45227F36FFBCFFB1B5BA2630FE5BFCC7	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFBDFFBFB5BA207CFDA6FEFB.text	45227F36FFBDFFBFB5BA207CFDA6FEFB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chalcoderus maculatus (Fabricius 1801) Fabricius (Erichson 1847	<div><p>Chalcoderus maculatus (Fabricius, 1801), restored combination</p><p>(Fig. 1)</p><p>Ateuchus maculatus Fabricius, 1801: 64 .</p><p>“ At. Maculatus F.” (Erichson 1847: 763) (citation).</p><p>Chalcoderus maculatus Fabricius (Erichson 1847: 763) (change of genus).</p><p>O [nthophagus] maculatus F. (Lacordaire 1855: 109 (note 3)) (citation).</p><p>Onthophagus maculatus Fabr. (Harold 1869: 1032) (citation).</p><p>O [nthophagus] maculatus Fabricius (d’Orbigny 1902: 114) (citation).</p><p>Onthophagus chloroderus d’Orbigny, 1905a: 449 (description) (new subjective synonymy) (Fig. 2)</p><p>Onthophagus chloroderus d’Orbigny (d’Orbigny 1913: 257–258) (key).</p><p>Onthophagus chloroderus d’Orbigny (d’Orbigny 1914: 657) (catalogue).</p><p>Onthophagus chloroderus d’Orb. (Boucomont &amp; Gillet 1927: 161) (citation sensu d’Orbigny 1913).</p><p>“ O. maculatus F.” (Palestrini 1982: 101) (subsequent designation as type species of Chalcoderus Erichson, 1847).</p><p>Onthophagus (Parascatonomus) maculatus F. (Palestrini 1982: 98) (transfer to the subgenus Parascatonomus).</p><p>Ateuchus maculatus (Fabricius, 1801) (Branco 2011: 18) (citation as type species of Chalcoderus Erichson, 1847)</p><p>Parascatonomus (Morettoeus) chloroderus (d’Orbigny, 1905a) (Ochi &amp; Kon 2023: 580) (transfer to Parascatonomus ( Morettoeus)).</p><p>All citations of Onthophagus chloroderus published subsequent to d’Orbigny (1905a) relate to Chalcoderus maculatus (Fabricius, 1801) (see summary below in section for Chalcoderus signatipennis (de Castelnau, 1840)) .</p><p>Type material examined: Ateuchus maculatus Fabricius, 1801 . Lectotype (not sexed): The type series of this species comprises three specimens according to Zimsen (1984: 31). To stabilize the nomenclature, here, we designate a lectotype (Article 74), which carries the following labels “ Guinea. [Ghana] / Meier. / Mus: T. Lund / Ateuchus / maculatus. F. {handwritten on square label} // small green square // TYPE {black print on red label} // ZMUC / 00508558 // square code ZMUC Fabricius 001127 // square code WORLD SCARAB DATABASE WSD00255537 // Lectotype Ateuchus maculatus / Fabricius, 1801 Moretto, / Deschodt &amp; Davis des. 2025 {red print} // Chalcoderus maculatus / (Fabricius, 1801) / Moretto, Deschodt &amp; Davis / det. 2025” ZMUC. Paralectotypes (2): 1 ex. “small green square // TYPE {black print on red label} // ZMUC / 00508559 // square code ZMUCFabricius 001128 // square code WORLD SCARAB DATABASE WSD00255538 // Onthophagus / maculatus / (Fab.) / P.Moretto det. 2008 {half handwritten, half printed} // Paralectotype Ateuchus / maculatus Fabricius, 1801 / Moretto, Deschodt &amp; Davis / det. 2025 {red print} // Chalcoderus / maculatus / (Fabricius, 1801) / Moretto, Deschodt / &amp; Davis det. 2025”; 1 ex. “small green square // TYPE {black print on red label} // ZMUC / 00508560 // square code ZMUC Fabricius 001129 // square code WORLD SCARAB DATABASE WSD00255539 // Onthophagus / maculatus / (Fab.) / P.Moretto det. 2008 {half handwritten, half printed} // Paralectotype Ateuchus / maculatus Fabricius, 1801 / Moretto, Deschodt &amp; Davis / det. 2025 {red print} // Chalcoderus maculatus / (Fabricius, 1801) / Moretto, Deschodt &amp; Davis / det. 2025” (ZMUC).</p><p>Type locality: “Guinea” [Ghana] (Moretto &amp; Génier 2015: 4)</p><p>Original description (Fabricius 1801): “maculatus . 45. A. clypeo integro, capite thoraceque cupreis, elytris testaceis nigro-maculatis. Habitat in Guinea. Mus. Dom. Lund. Statura et magnitude praecedentium. Capitis clypeus rotundatus, integer, cupreus. Thorax gibbosus, laevis, cupreus, nitidus. Elytra substriata, Testacea, strigis duabus e maculis quadratis nigris. Corpus atrum.”</p></div>	https://treatment.plazi.org/id/45227F36FFBDFFBFB5BA207CFDA6FEFB	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFB3FFBFB5BA25F2FE3CFC63.text	45227F36FFB3FFBFB5BA25F2FE3CFC63.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Onthophagus chloroderus d'Orbigny 1905	<div><p>Onthophagus chloroderus d’Orbigny, 1905</p><p>(Fig. 2)</p><p>Holotype by monotypy “ Sierra / Leone {handwritten, black bordered} // Ex-Musaeo / H.W.Bates / 1892 {black bordered} // maculatus F. / v. chloroderus / var. nov. d’Orb. {handwritten by d’Orbigny} // H. d’Orbigny / Onth. Afr. 1902 // Chloroderus / d’Orb. {handwritten by d’Orbigny} // H. d’Orbigny / vidit 1906 // HOLOTYPE {black print on red label} // Muséum Paris / ex. Coll. / R.Oberthür // Chalcoderus / maculatus / (Fabricius, 1801) / Moretto, Deschodt / &amp; Davis det. 2025” (MNHN).</p><p>Type locality: “ Sierra Leone ” ( d’Orbigny (1905) clearly states that this specimen is the type).</p><p>Original description (d’Orbigny, 1905a): “… vert, avec le dessous du corps et les pattes plus foncés, ou d’un bleu foncé... les élytres d’un testacé clair, y compris les épipleures, avec le bord basiliaire très étroitement noir et deux rangées transverses de petites taches noires…”</p><p>Comparison between the types of Onthophagus chloroderus d’Orbigny, 1905, and Ateuchus maculatus Fabricius, 1801, shows there are no differences, except in the colour of the pronotum. Aedeagi extracted from non-type green (Guinea) and red (Côte d’Ivoire) individuals also show no differences. Following the principle of priority, we propose Onthophagus chloroderus d’Orbigny, 1905 as a junior subjective synonym of Ateuchus maculatus Fabricius, 1801 .</p></div>	https://treatment.plazi.org/id/45227F36FFB3FFBFB5BA25F2FE3CFC63	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFB0FFBDB5BA224CFCA5FB4A.text	45227F36FFB0FFBDB5BA224CFCA5FB4A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chalcoderus signatipennis (de Castelnau 1840) MNHN	<div><p>Chalcoderus signatipennis (de Castelnau, 1840), restored status, restored combination</p><p>(Fig. 3)</p><p>Onthophagus signatipennis de Castelnau, 1840: 85 (described from Senegal).</p><p>“Onthoph. signatipennis Dej. Lap. ” (Erichson 1847: 763) (citation).</p><p>Chalcoderus signatipennis Laporte (Erichson 1847: 763) (change of genus).</p><p>O[nthophagus] signatipennis Casteln. (Lacordaire 1855: 109 (note 3)) (citation).</p><p>Onthophagus signatipennis Casteln. (Harold 1869: 1032) (synonymy with O. maculatus Fabr.).</p><p>Onthophagus maculatus Fabricius (d’Orbigny 1905a: 450) (comparison with “ O. chloroderus n. sp. ”—Starting point of the confusion).</p><p>Onthophagus maculatus Fabricius, 1801 (d’Orbigny 1913: 257–258) (citation consistent with earlier synonymy of O. signatipennis).</p><p>Onthophagus maculatus Fabricius (d’Orbigny 1914: 656) (catalogue).</p><p>Onthophagus signatipennis Castelnau (d’Orbigny 1914: 657) (citation as synonym of O. maculatus Fabr.).</p><p>Onthophagus maculatus F. 1801 (Boucomont &amp; Gillet 1927: 180) (citation sensu d’Orbigny 1913).</p><p>Onthophagus signatipennis Cast. (Boucomont &amp; Gillet 1927: 180) (citation as synonym of O. maculatus F. sensu d’Orbigny).</p><p>Parascatonomus (Morettoeus) maculatus (Fabricius, 1801) (Ochi &amp; Kon 2023: 579 (transfer to Parascatonomus ( Morettoeus)).</p><p>All citations of Onthophagus maculatus published subsequent to Onthophagus maculatus sensu d’Orbigny (1905a: 450) relate to Chalcoderus signatipennis de Castelnau, 1840 (see summary below).</p><p>The type material is presumed destroyed (Evenhuis 2012). Here, we designate a neotype ♂ selected from specimens held in the MNHN: “ SENEGAL / FORET DE BANDIA // Museum Paris / Collection générale // Neotype ♂ / Onthophagus / signatipennis / de Castelnau, 1840 / Moretto, Deschodt / &amp; Davis des. 2025 {handwritten on red label} // Chalcoderus signatipennis / (de Castelnau 1840) / Moretto, Deschodt &amp; Davis / det. 2025” (MNHN).</p><p>Type locality (Article 76.3): Sénégal, Région de Thies, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-16.95&amp;materialsCitation.latitude=14.6" title="Search Plazi for locations around (long -16.95/lat 14.6)">Forêt de Bandia. The</a> “Forêt de Bandia” corresponds to the current private “Réserve de Bandia” (14° 36′ N, 16° 57′ W).</p><p>Original description (de Castelnau 1840): “17. Onthophagus signatipennis . Long. 4 lig. ½. Larg. 2 lig. ½. — D’un rouge cuivreux; corselet très-bombé, entièrement granuleux; élytres d’un brun jaune, striées, avec deux bandes transversales irrégulières, noires, formées de taches qui se réunissent ordinairement: elles offrent aussi chacune une tache noire vers l’extrémité; dessous du corps et pattes d’un vert cuivreux. — Sénégal.”</p><p>From this description, it is clear that Onthophagus signatipennis de Castelnau, 1840, corresponds to what d’Orbigny (1913) believed to be Ateuchus maculatus Fabricius, 1801 . It is here removed from synonymy with Ateuchus maculatus Fabricius, 1801 and its valid species status is restored. As Erichson (1847) described the genus Chalcoderus for both Ateuchus maculatus Fabricius, 1801, and Onthophagus signatipennis de Castelnau, 1840 (cited as At. maculatus F., Onthoph. signatipennis Dej. Lap.), the combination Chalcoderus signatipennis (de Castelnau, 1840) is, therefore, restored.</p><p>Distribution of C. signatipennis (Fig. 4) according to studied specimens: Gambia, Guinea Bissau and Senegal west of the Senegal River, plus mostly on Tertiary sediments of the Senegalese Basin (Thieblemont et al. 2016); Western Mali (“Haut Sénégal, Khayes” PMOC).</p><p>To summarize, Onthophagus maculatus sensu d’Orbigny (1913: 257–258) is actually Chalcoderus signatipennis (de Castelnau, 1840) whereas Onthophagus chloroderus d’Orbigny, 1905 (1905a: 450; 1913: 258) is actually Chalcoderus maculatus (Fabricius, 1801) . The descriptions given by d’Orbigny (1913: 257–258) are precise and allow the two species to be easily separated. Subsequent erroneous citations with respect to these inversions have extended to the present day and it is appropriate to interpret nomenclature in these subsequent publications accordingly, including Ochi &amp; Kon (2023, Figs 1a–j and 2a–d).</p></div>	https://treatment.plazi.org/id/45227F36FFB0FFBDB5BA224CFCA5FB4A	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFB1FFBDB5BA21D9FCEAF801.text	45227F36FFB1FFBDB5BA21D9FCEAF801.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chalcoderus longiceps (d'Orbigny 1904) Moretto & Deschodt & Davis 2026	<div><p>Chalcoderus longiceps (d’Orbigny, 1904), new combination</p><p>(Fig. 6)</p><p>Onthophagus longiceps d’Orbigny, 1904: 285 .</p><p>Type material examined (photos only): Lectotype ♂, present designation “afr. or. / allem. {handwritten} // longiceps / n. sp. d’Orb. {handwritten by d’Orbigny} // Ex-Typis {printed in red, framed in black} // cf. Ann. Mus. Civ. Genov. / XLI.1904, p. 285 {handwritten} // J.J. Gillet det. vend. / O.( Onthophagus) / longiceps d’Orb. / R.M.H.N. Belg. 10.640 // Lectotype Onthophagus / longiceps d’Orbigny, 1904 / Moretto, Deschodt &amp; Davis / des. 2025 {red} // Chalcoderus longiceps / (d’Orbigny, 1904) / Moretto, Deschodt &amp; Davis / det. 2025” (IRSNB). Paralectotype ♀ “afr. or. / allem. {handwritten} // Ex-Typis {printed in red, framed in black} // cf. Ann. Mus. Civ. Genov. / XLI.1904, p. 285 {handwritten} // J.J. Gillet det. vend. / O.( Onthophagus) / longiceps d’Orb. / R.M.H.N. Belg. 10.640 // Paralectotype Onthophagus / longiceps d’Orbigny, 1904 / Moretto, Deschodt &amp; Davis det. 2025 {red} // Chalcoderus longiceps (d’Orbigny, 1904) / Moretto, Deschodt &amp; Davis det. 2025” (IRSNB) .</p><p>Type locality: “Afrique orientale allemande” [Tanzania].</p></div>	https://treatment.plazi.org/id/45227F36FFB1FFBDB5BA21D9FCEAF801	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFB6FFBAB5BA25C1FDACFC89.text	45227F36FFB6FFBAB5BA25C1FDACFC89.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chalcoderus foliiceps (Quedenfeldt 1884) Moretto & Deschodt & Davis 2026	<div><p>Chalcoderus foliiceps (Quedenfeldt, 1884), new combination</p><p>(Fig. 7)</p><p>Onthophagus foliiceps Quedenfeldt, 1884: 277 .</p><p>Type material examined (photo only): Holotype by monotypy “Quango Strom / Major v. Mechow {printed and framed in black} // Ex-Musaeo / Quedenfeldt {printed and framed in black} // o. foliiceps / n. sp. Qdflt {handwritten by Quedenfeldt} // H. d’Orbigny / Ontho. Afr. 1902 // HOLOTYPE {printed in black on red label} // Muséum Paris / ex. Coll. / R.Oberthür // Chalcoderus foliiceps / (Quedenfeldt, 1884) / Moretto, Deschodt &amp; Davis / det. 2025” (MNHN).</p><p>Type locality: “ Malange ” [Angola].</p></div>	https://treatment.plazi.org/id/45227F36FFB6FFBAB5BA25C1FDACFC89	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFB7FFBBB5BA24E0FE5EFD82.text	45227F36FFB7FFBBB5BA24E0FE5EFD82.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chalcoderus graphicus (Wallengren 1881) Moretto & Deschodt & Davis 2026	<div><p>Chalcoderus graphicus (Wallengren, 1881) new combination</p><p>(Fig. 8)</p><p>Onthophagus graphicus Wallengren, 1881: 21 .</p><p>Type material examined: (photo only) Holotype by monotypy “Christiana {gothic characters} // Copris pumili /onis. Wallengr. {handwritten} // S. Afr. Transvaal / Christiana. / Leg. N. Persson. /Coll. Malmö Museum. // Typus {printed and framed in black on red label} // 1963 / 210 {{pale blue} // Zool. Mus. Lund Sweden / Scarabaeidae / Scarabaeinae / Type No. 2769:1 {half handwritten, half printed} // Chalcoderus graphicus / (Wallengren, 1881) / Moretto, Deschodt &amp; Davis / det. 2025” (MZLU).</p><p>Type locality: “Christiana” [South Africa, North West Prov. Dr Ruth Segomotsi Mompati District, Lekwa-Teemane Municipality].</p></div>	https://treatment.plazi.org/id/45227F36FFB7FFBBB5BA24E0FE5EFD82	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFB7FFBBB5BA26FEFC8CFAD3.text	45227F36FFB7FFBBB5BA26FEFC8CFAD3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chalcoderus imbellis (d'Orbigny 1905) Moretto & Deschodt & Davis 2026	<div><p>Chalcoderus imbellis (d’Orbigny, 1905), new combination</p><p>(Fig. 9)</p><p>Onthophagus imbellis d’Orbigny, 1905b: 434 .</p><p>Parascatonomus (Morettoeus) imbellis (d’Orbigny, 1905) (Ochi &amp; Kon 2023: 578) .</p><p>Type material examined (photo only): Lectotype (not sexed), present designation “ GUINEA PORTOGHESE / Bolama / VI-XII 1899. L.Fea // imbellis / n.sp. d’Orb. {handwritten by d’Orbigny} // TYPUS {printed and framed in red} // imbellis / d’Orb. {handwritten and framed in black} // Syntypus / Onthophagus / imbellis / d’Orbigny, 1905 // Museo Civico / di Genova // Lectotype Onthophagus / imbellis d’Orbigny, 1905, / Moretto, Deschodt &amp; Davis / des. 2025 {red} // Chalcoderus imbellis (d’Orbigny, 1905) / Moretto, Deschodt &amp; Davis det. 2025” (MSNG) . Paralectotypes (4). 2 ex. (photos), same labels than the lectotype except “// Paralectotype Onthophagus / imbellis d’Orbigny, 1905 / Moretto, Deschodt &amp; Davis det. 2025 {red} // Chalcoderus imbellis (d’Orbigny, 1905) / Moretto, Deschodt / &amp; Davis det. 2025” (MSNG) . 1 ♂ “ GUINEA PORTOGHESE / Bolama / VI-XII 1899. L.Fea // MUSEUM PARIS / Coll. H. d’Orbigny / 1915 // {aedeagus in a vial} // Paralectotype Onthophagus imbellis d’Orbigny, 1905 / Moretto, Deschodt &amp; Davis det. 2025 {red} // Chalcoderus imbellis (d’Orbigny, 1905) / Moretto, Deschodt &amp; Davis det. 2025”; 1 ♂ “ GUINEA PORTOGHESE / Bolama / VI-XII 1899. L.Fea // MUSEUM PARIS / Coll. H. d’Orbigny / 1915 // imbellis / d’Orb. {handwritten by d’Orbigny} // TYPE {black on red label} // Paralectotype Onthophagus imbellis d’Orbigny, 1905 / Moretto, Deschodt &amp; Davis det. 2025 {red} // Chalcoderus imbellis (d’Orbigny, 1905) / Moretto, Deschodt &amp; Davis det. 2025” (MNHN) .</p><p>Type locality: “ Guinée portugaise: Bolama ” [Guinea Bissau].</p></div>	https://treatment.plazi.org/id/45227F36FFB7FFBBB5BA26FEFC8CFAD3	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFB4FFB8B5BA24E0FBE9FDAE.text	45227F36FFB4FFB8B5BA24E0FBE9FDAE.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chalcoderus inermiceps (d'Orbigny 1905) Moretto & Deschodt & Davis 2026	<div><p>Chalcoderus inermiceps (d’Orbigny, 1905), new combination</p><p>(Fig. 10)</p><p>Onthophagus inermiceps d’Orbigny, 1905a: 450 .</p><p>Type material examined (photo only): Holotype by monotypy “ Samburu / 30. X. to / 20. XI.[18]96. {Printed with top and bottom lines in black} // B. E. Africa / C.S. Betton. / 98—12 {separated by blue line with black basal line} // Determined by / M. d’Orbigny. {printed} / inermiceps / n.sp. d’Orb. {handwritten by d’Orbigny} // Type {black print within red circle} // square code NHMUK015529648 // Chalcoderus inermiceps / (d’Orbigny, 1905) / Moretto, Deschodt &amp; Davis / det. 2025” (NHML).</p><p>Type locality: “ Afrique orientale anglaise: Sambourou ” [Kenya: Samburu].</p></div>	https://treatment.plazi.org/id/45227F36FFB4FFB8B5BA24E0FBE9FDAE	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFB4FFB8B5BA2612FDC0FC59.text	45227F36FFB4FFB8B5BA2612FDC0FC59.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chalcoderus planaticeps (d'Orbigny 1913) Moretto & Deschodt & Davis 2026	<div><p>Chalcoderus planaticeps (d’Orbigny, 1913), new combination</p><p>(Fig. 11)</p><p>Onthophagus planaticeps d’Orbigny, 1913: 259 .</p><p>Type material examined (photo only): Holotype by monotypy “ Congo {handwritten} // planaticeps / n. sp. D’Orb. {handwritten by d’Orbigny} // TYPE {printed and framed in black on red label} // Cf. Ann. Soc. Ent. Fr. / T.82 (1913) p.259 {handwritten} // J.J. Gillet det. vend.: / O.( Onthophagus) / planaticeps d’Orb. / R.M.H.N. Belg. 10.640 {half handwritten, half printed} // Chalcoderus planaticeps / (d’Orbigny, 1913) / Moretto, Deschodt &amp; Davis / det. 2025” (IRSNB).</p><p>Type locality: “ Congo ” [D.R.C.].</p></div>	https://treatment.plazi.org/id/45227F36FFB4FFB8B5BA2612FDC0FC59	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFB4FFB8B5BA2043FC7BFA6F.text	45227F36FFB4FFB8B5BA2043FC7BFA6F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chalcoderus semicroceus (d'Orbigny 1915) Moretto & Deschodt & Davis 2026	<div><p>Chalcoderus semicroceus (d’Orbigny, 1915), new combination</p><p>(Fig. 12)</p><p>Onthophagus semicroceus d’Orbigny, 1915: 386 .</p><p>Type material: d’Orbigny (1915) indicates a size range “Long. 5–5,5 mm” from which we can deduce that there are at least two specimens in the type series, but only one was found (examined from photo) and is here designated as the Lectotype, present designation “MUSÉE DU CONGO / Kapiri / IX.1912 / Miss. Agric. // R. DÉT. / S / 182 // semicroceus / n. sp. D’Orb. {handwritten by d’Orbigny} // TYPE {printed and framed in black on red label} // Lectotype Onthophagus / semicroceus d’Orbigny, 1915, / Moretto, Deschodt and Davis / des. 2025 {red} // Chalcoderus semicroceus / (d’Orbigny, 1915) / Moretto, Deschodt &amp; Davis / det. 2025” (MRAC) .</p><p>Type locality: “Kapiri dans le Katanga ” [ Kapiri, Lualaba, D.R.C.].</p></div>	https://treatment.plazi.org/id/45227F36FFB4FFB8B5BA2043FC7BFA6F	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFB4FFA6B5BA2250FBF2FD3F.text	45227F36FFB4FFA6B5BA2250FBF2FD3F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chalcoderus simpliciceps (d'Orbigny 1905) Moretto & Deschodt & Davis 2026	<div><p>Chalcoderus simpliciceps (d’Orbigny, 1905), new combination</p><p>(Fig. 13)</p><p>Onthophagus simpliciceps d’Orbigny, 1905a: 452 .</p><p>Type material examined (photo only): Holotype by monotypy “simpliciceps / n.sp. d’Orb. {handwritten by d’Orbigny} // D.O.Africa / Tanga 5.4.[18]99 {printed in black on blue label, date handwritten} // Coll. C. Felsche / Kauf 20, 1918 {printed and framed in black on blue label} // Chalcoderus simpliciceps / (d’Orbigny, 1905) / Moretto, Deschodt &amp; Davis / det. 2025” (ZHMB).</p><p>Type locality: “Afrique orientale allemande: Tanga ” [Tanzania].</p><p>Distributional notes: Photographs of additional material in Ochi &amp; Kon (2023) show clear differences in habitus between the three, known, closely-related, West African species of Chalcoderus: C. signatipennis (cited as Parascatonomus (Morettoeus) maculatus), C. maculatus (cited as P. (M.) chloroderus) and C. imbellis (cited as P. (M.) imbellis) despite some apparent overlap in distribution (d’Orbigny 1913). A record for C. imbellis from Ethiopia (d’Orbigny 1913) requires validation.</p><p>Three species have been described from the Democratic Republic of the Congo (DRC) and Angola in central Africa with the single type specimen for C. planaticeps from an unspecified locality in the Congo (DRC) cited as very close to C. semicroceus (d’Orbigny, 1915) . Three species have also been described from Kenya and Tanzania in East Africa with one cited as occurring southwards to Zimbabwe (d’Orbigny 1913). In addition to these three species, C. graphicus has also been recorded from East Africa (Laikipia) (Krell 2005, cited as Onthophagus graphicus) although it was described from South Africa where it shows a wide distribution, extending into neighbouring countries.</p><p>The comparison of types and recently collected specimens from different African countries shows that the genus Chalcoderus is much more diverse than expected. Although d’Orbigny (1913, 1915) recognised all 10 species listed, here, as valid, seven small species show close similarities differing primarily in punctation and patterns of melanic patches or spots on the elytra. Therefore, it would be useful to conduct a full revision of described Chalcoderus species and their distributions that includes further, possibly undescribed, African species and Indian relatives. However, such a revision awaits the collection of further individuals.</p></div>	https://treatment.plazi.org/id/45227F36FFB4FFA6B5BA2250FBF2FD3F	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFAAFFA6B5BA2769FD3AFB59.text	45227F36FFAAFFA6B5BA2769FD3AFB59.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Onthophagus coeruleicollis , Arrow 1907	<div><p>Onthophagus (s. l.) coeruleicollis Arrow, 1907</p><p>(Fig. 14)</p><p>Onthophagus coeruleicollis Arrow, 1907: 430 .</p><p>Type material examined (photos): Lectotype ♀ (Bacchus 1978) “Dharwar, / N. India / 4473 {handwritten} // Onthophagus / coeruleicollis, Arrow / type a.a.m., 1907. {handwritten by Arrow} // Onthophagus ♀ / coeruleicollis, Arr / M.E.Bacchus det 1975 / LECTOTYPE {half printed, half handwritten} // LECTO. / TYPE {printed in black within blue circle}” (NHML) . Paralectotype ♀ “ India // Bowring . / 63.47* // Onthophagus / coeruleicollis / co-type Arrow {handwritten by Arrow} // Onthophagus ♀ / coeruleicollis,Arr / M.E.Bacchus det 1975 / PARALECTOTYPE {half printed, half handwritten} // PARA. / LECTO. / TYPE {printed in black within blue circle}” (NHML) . Type material not examined: Paralectotypes 1 ♂, 1 ♀, Belgaum, Bombay (Bacchus 1978) (NHML) .</p><p>Type locality: “S. India, Dharwar, Belgaum.”</p></div>	https://treatment.plazi.org/id/45227F36FFAAFFA6B5BA2769FD3AFB59	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
45227F36FFAAFFA6B5BA2142FD9AF934.text	45227F36FFAAFFA6B5BA2142FD9AF934.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Onthophagus regalis , Arrow 1907	<div><p>Onthophagus (s. l.) regalis Arrow, 1907</p><p>(Fig. 15)</p><p>Onthophagus regalis Arrow, 1907: 431 .</p><p>Type material examined (photo): Paralectotype ♀ “ Ceylon. / H. P. Green. / 1916 – 157 // Onthophagus / regalis, Arrow {handwritten by Arrow} // MUSÉUM PARIS / 1938 / COLL. A. BOUCOMONT” (MNHN). Type material not examined: Lectotype (Bacchus 1978) ♂, Sri Lanka; Paralectotype ♀, Sri Lanka, Yerbury (Bacchus 1978) (NHML) .</p><p>Type locality: “Ceylon” [Sri Lanka].</p></div>	https://treatment.plazi.org/id/45227F36FFAAFFA6B5BA2142FD9AF934	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.		MagnoliaPress via Plazi	Moretto, Philippe;Deschodt, Christian M.;Davis, Adrian L. V.	Moretto, Philippe, Deschodt, Christian M., Davis, Adrian L. V. (2026): The genus Chalcoderus Erichson, 1847: synonymy of Parascatonomus (Morettoeus) Ochi & Kon, 2023, and transfer of group 15 (d’Orbigny 1913, 1915) Afrotropical Onthophagus Latreille, 1802 (Coleoptera: Scarabaeinae). Zootaxa 5752 (1): 63-84, DOI: 10.11646/zootaxa.5752.1.3, URL: https://doi.org/10.11646/zootaxa.5752.1.3
