Search

In the Neotropics, orphnine scarab beetles are represented by the endemic tribe Aegidiini Paulian, 1984 with five genera and over 50 species. Phylogenetic analysis based on morphological characters of all supraspecific taxa of Orphninae showed that Aegidiini is comprised of two lineages. New subtribes, Aegidiina subtr. nov. (Aegidium Westwood, 1845, Paraegidium Vulcano et al., 1966, Aegidiellus Paulian, 1984, and Onorius Frolov & Vaz-de-Mello, 2015, and Aegidinina subtr. nov. (Aegidinus Arrow, 1904) are proposed to better reflect this phylogeny. Two new species of Aegidinus are described: A. alexanderi sp. nov., from the Yungas in Peru and A. elbae sp. nov. from the Caqueta moist forests ecoregion in Colombia. A diagnostic key to Aegidinus species is given.

The third instar larvae of Aphodius (Alocoderus) hydrochaeris (Fabricius, 1798) and A. (Bodilus) ictericus (Laicharting, 1781) are described based on scanning electron microscopy and COI sequences. COI barcode sequence for A. (A.) hydrohaeris is provided for the first time. Two haplotypes are discovered in A. (B.) ictericus.

Aphodius (Alocoderus) hydrochaeris (Fabricius, 1798) Figs. 1–21 Material examined. More than a hundred of the third instar larvae were collected on 7–9.XI. 2019 in cow dung near Dosang and Topal settlements (Astrakhan Province, south of the European part of Russia). All larvae were about the same size and similar exteriorly. Thirty live larvae were taken to the ZIN laboratory and in two months 11 adults were obtained. For two adults and one larva the DNA barcodes were obtained (Genbank accession numbers: MW 672592 – MW 672594, voucher numbers: COI121912, COI121919, COI121920). Description of the third instar larva. Larva of typical C-shape form, relatively slender, covered with sparse, short setae, with whitish, semitransparent integument except for head. Head width: 2.0± 0.05 mm, length (without labrum): 1.43± 0.05 mm. Head capsule color varies from dark-brown with paler base and apex of frons and area around antennal support (Fig. 1) to brown with unclear darker pattern of epicranium (Fig. 4) with intermediate variants (Figs 2, 3). Head surface dull, microreticulate (Figs 9, 11). Frontal sutures are visible as very fine, darker lines. Epicranial suture approximately 4 times shorter than frons height. Each pleural sclerite of epicranuim with 5 long setae: 4 near antennal support and 1 near epicranial suture. A few shorter setae located on pleural sclerite are arranged in irregular rows. Frons with 2 pairs of long setae and 3 pairs of short setae. Stemmata absent. Clypeus trapezoidal, brown. Postclypeus (3/4 length of clypeus) darker than preclypeus. Surface of clypeus without prominent tubercles. First 3 proximal antennomeres about same length, antennomere 1 with whitish stripe, antennomere 3 with small conical process and 5 minute setae distally. Ultimate antennomere short and conical, with 1 smooth elongated sensory area medially near apex and small apical area with a few basiconic sensilla and 1 minute seta. Labrum (Figs 13–15) trilobed apically, with a pair of long setae and 1–2 pairs of short setae dorsally, 24 setae on anterior and lateral margins, and a pair of short robust setae ventrobasally. Clithra present, symmetrical. Phoba (protophoba, dexiophoba, and laeophoba sensu Ritcher (1966)), as a semicircular ridge of inclinate mediad and sclerotized apically spinules; most spinules pectinate apically. Short additional ridge situated anteriorly on right side. Phoba anteriorly flanked with a row of ca. 20 placoid sensilla; 2 larger placoid sensilla between clithra. Epitorma long, parallel-sided, rounded and slightly widened apically. Pternotormae developed, subsymmetrical. Mandibles (Figs 16–19) triangular, asymmetrical, external sides with 3 setae. Left mandible slightly longer than right mandible, its scissorial part wider, S 1 and S 2 teeth widely separated by sinuation, S 2 and S 3 close, separated by incision. Right mandible: S 1 and S 2 teeth fused, S 3 separated from them by incision. Base of mandible light brown, scissorial and molar part dark black. Maxillae symmetrical (Figs 12, 20, 21). Cardo with 4 short setae: 2 on ventral side and 2 on lateral margin near base of stipes. Ventral side of stipes with 1 long proximal and 2 short distal setae, dorsal side with a row of 13–14 stridulatory teeth and 3 setae near base of palpifer (2 longer setae and 1 minute seta). Palpifer without stridulatory teeth, with one short seta ventrally. Maxillary palpus with 4 palpomeres; palpomeres 1 and 4 with 1 seta each, palpomere 3 with 2 setae. Ventral side of galea with longitudinal row of 9 short setae. Dorsal side and apex of galea with 6 longer setae. Dorsal side of lacinia with 5 long and thick setae and 3 shorter and slenderer setae basally, ventral side with 1 short seta basally. Uncus of lacinia tridentate, with 1 or 2 thick setae basad of the apical teeth. Postmentum with 2 pairs of short seta. Proximal part of prementum with a pair of long setae. Distal part of prementum with pair of long setae. Glossa surrounded laterally by rows of thick and ling setae reaching submentum ventroapically; distal setae long and acute, proximal setae shorter and somewhat spatulate, specifically in right row. Glossa with 3 pairs of setae apically, 2 pairs of larger placoid sensilla, a transverse row of ca. 20 smaller placoid sensilla proximad of larger sensilla, and a row of short, triangular spinules immediately proximad of row of placoid sensilla. Middle and hind legs of subequal length, fore legs slightly shorter. Each leg with 36 setae: 3 on coxa, 7 on trochanter, 11 on femur, 13 on tibia, and 2 on tarsus. Tarsal apices not serrate in lateral view, leg segments without tubercles. Thoracic and abdominal segments with short and sparse setae. Folds of abdominal tergites 1-6 with shorter, thick setae arranged in transverse rows medially; setae not situated on conical tubercles. Lateral bulges of abdominal segments with 3 setae. Protoracic spiracles about 2 times larger than abdominal spiracles; abdominal spiracles of subequal size, minute. Raster (Figs 6–8) with pallidia of 8-11 relatively long, strongly sclerotized, depressed spinules (Fig. 10); pallidia curved and converge towards basal margin of sternite. Anal sternite also bears smaller spinules on each side of pallidia. Lower anal lobe not sinuate laterally (Fig. 5). Diagnosis. The larva of A. (A.) hydroсhaeris was described by Martynov (1999), however the differential diagnosis was not provided and specifically the chaetotaxy of the maxilla was not described. In the key to Aphodius larvae (Krell 1997) this species comes out in the couplet with A. (Melinopterus) prodromus (Brahm, 1790) but differs from it in the larger head size, the absence of the red hue in the head capsule coloration, and setation of the lacinia having five longer and thicker setae and three shorter and slenderer ones basally., Published as part of Frolov, Andrey V., Vishnevskaya, Maria S. & Akhmetova, Lilia A., 2021, Identification of larvae of two Aphodius Helwig, 1798 (Coleoptera: Scarabaeidae: Aphodiinae) species using morphology and DNA barcode, pp. 153-164 in Zootaxa 5047 (2) on pages 154-158, DOI: 10.11646/zootaxa.5047.2.4, http://zenodo.org/record/5540644, {"references":["Ritcher, P. O. (1966) Wite Grubs and their Allies. A study of North American scarabaeoid larvae. Oregon State Monographs, Studies in Entomology 4. Oregon State University Press, Corvallis, Oregon, 219 pp.","Martynov, V. V. (1999) Opisanie lichinok zhukov roda Aphodius (Coleoptera, Scarabaeidae). Soobshhenie 2 [Description of the larvae of dung-beetles of the genus Aphodius (Coleoptera, Scarabaeidae). Communication 2]. Vestnik Zoologii, 33 (4 - 5), 35 - 43.","Krell, F. - T. (1997) Uberfamilie: Scarabaeoidea. Nachtrag zur 33. Familie: Scarabaeidae. 14. Gattung: Aphodius Illiger. In: Klausnitzer B. (ed.), Die Larven der Kafer Mitteleuropas. 4. Band. Polyphaga. Pars 3 sowie Erganzungen zum 1. bis 3. Band. Goecke & Evers, Krefeld, pp. 106 - 123."]}

Aphodius (Bodilus) ictericus (Laicharting, 1781) Figs. 22–45 Material examined. More than a hundred of the third instar larvae were collected on 7.–9.XI. 2019 in cow dung near Dosang and Topal settlements (Astrakhan Province, south of the European part of Russia). All larvae were about the same size and similar exteriorly. Thirty live larvae were taken to the ZIN laboratory and in two months 13 adults were obtained. For two adults and three larvae DNA barcodes were obtained (Genbank accession numbers: MW 672587 – MW 672591, voucher numbers: COI121905, COI121906, COI121908, COI121921, COI121922). Description of the third instar larva. Larva of typical C-shape form, relatively slender, covered with sparse, short setae, with whitish, semitransparent integument except for head. Head width: 1.51± 0.04 mm, length (without labrum): 1.13± 0.03 mm. Head surface shiny, smooth (Figs 33, 34). Color pattern of head capsule varies from light brown with unclear lighter spots to dark-brown pleural sclerites with distinct lighter spots and brown frons (Figs 22–25). Frontal sutures are visible as very fine, darker lines. Epicranial suture approximately 4 times shorter than frons height. Each pleural sclerite of epicranuim with 5 long setae: 4 near antennal support and 1 near epicranial suture. A few shorter setae located on pleural sclerite are arranged in irregular rows. Frons with 2 pairs of long setae and 3 pairs of short setae. Stemmata absent. Clypeus trapezoidal, brown. Postclypeus (2/3 length of clypeus) darker than preclypeus. Surface of clypeus without prominent tubercles. First 3 proximal antennomeres about same length, antennomere 1 with whitish stripe, antennomere 3 with small conical process and 5 minute setae distally. Ultimate antennomere short and conical, with 1 smooth elongated sensory area medially near apex and small apical area with a few basiconic sensilla and 1 minute seta. Labrum (Figs 37–39) trilobed apically, with a pair of long setae and 1–2 pairs of short setae dorsally, 24 setae on anterior and lateral margins, and a pair of short robust setae ventrobasally. Clithra present, symmetrical. Phoba (protophoba, dexiophoba, and laeophoba sensu Ritcher (1966)), as a semicircular ridge of inclinate mediad and sclerotized apically spinules; most spinules pectinate apically. Short additional ridge situated anteriorly on right side. Phoba anteriorly flanked with a row of ca. 20 placoid sensilla; 2 larger placoid sensilla between clithra. Epitorma long, parallel-sided, rounded and slightly widened apically. Pternotormae developed, subsymmetrical. Mandibles (Figs 40–43) triangular, asymmetrical, external sides with 3 setae. Left mandible slightly longer than right mandible, its scissorial part wider, S 1 and S 2 teeth widely separated by sinuation, S 2 and S 3 close, separated by incision. Right mandible: S 1 and S 2 teeth fused, S 3 separated from them by incision. Base of mandible light brown, scissorial and molar part dark black. Maxillae symmetrical (Fig. 36, 44, 45). Cardo with 4 short setae: 2 on ventral side and 2 on lateral margin near base of stipes. Ventral side of stipes with 1 long proximal and 2 short distal setae, dorsal side with a row of 8 stridulatory teeth and 3 setae near base of palpifer (2 longer setae and 1 minute seta). Palpifer without stridulatory teeth, with one short seta ventrally. Maxillary palpus with 4 palpomeres; palpomeres 1 and 4 with 1 seta each, palpomeres 3 with 2 setae; seta in 1st palpomere modified (Fig. 36, arrowed). Ventral side of galea with longitudinal row of 9–10 short setae. Dorsal side and apex of galea with 6 longer setae. Dorsal side of lacinia with 5 long and thick setae and 1 shorter and slenderer seta basally, ventral side with 1 short seta basally. Uncus of lacinia tridentate, with 1 thick seta basad of the apical teeth. Postmentum with 2 pairs of short setae. Proximal part of prementum with a pair of long setae. Distal part of prementum with a pair of long setae. Glossa surrounded laterally by rows of thick and ling setae reaching submentum ventroapically; distal setae long and acute, proximal setae shorter and somewhat spatulate, specifically in right row. Glossa with 3 pairs of setae apically, 2 pairs of larger placoid sensilla, a transverse row of ca. 20 smaller placoid sensilla proximad of larger sensilla, and a row of short, triangular spinules immediately proximad of row of placoid sensilla. Middle and hind legs of subequal length, fore legs slightly shorter. Each leg with 36 setae: 3 on coxa, 7 on trochanter, 11 on femur, 13 on tibia, and 2 on tarsus. Tarsal apices not serrate in lateral view, leg segments without tubercles. Thoracic and abdominal segments with short and sparse setae. Folds of abdominal tergites 1–6 with shorter, thick setae arranged in transverse rows medially; setae not situated on conical tubercles. Lateral bulges of abdominal segments with 2 setae. Protoracic spiracles about 2 times larger than abdominal spiracles; abdominal spiracles of subequal size, minute. Raster without pallidia, with 56–60 relatively long, strongly sclerotized spinules. Medial spinules robuster and slightly flattened and widened apically (Fig. 35). Spinules are not arranged in rows or, in some specimens, medial spinules appear to be arranged in irregular longitudinal rows (Figs. 26–31). Lower anal lobe not sinuate laterally (Fig. 32). Diagnosis. The Krell’s (1997) key to Central European Aphodius larvae includes the larva of this species, however it is characterized as having a distinct smooth area of the raster which reaches middle of the raster. Examination of our material shows that this character is not reliable and the smooth area is unclear in most specimens examined. The third instar larvae of A. (B.) ictericus is better diagnosed by having a smooth, shiny head with width ca. 1.5 mm and a characteristic color pattern of the pleural sclerites, raster with ca. 50–60 spinules without distinct pallidia and with unclear or indistinct smooth area basally, and by the modified setae on the first maxillary palpomere. The latter character, however, needs further examination because the larvae of few species of Aphodius were studied with SEM., Published as part of Frolov, Andrey V., Vishnevskaya, Maria S. & Akhmetova, Lilia A., 2021, Identification of larvae of two Aphodius Helwig, 1798 (Coleoptera: Scarabaeidae: Aphodiinae) species using morphology and DNA barcode, pp. 153-164 in Zootaxa 5047 (2) on pages 158-162, DOI: 10.11646/zootaxa.5047.2.4, http://zenodo.org/record/5540644, {"references":["Ritcher, P. O. (1966) Wite Grubs and their Allies. A study of North American scarabaeoid larvae. Oregon State Monographs, Studies in Entomology 4. Oregon State University Press, Corvallis, Oregon, 219 pp.","Krell, F. - T. (1997) Uberfamilie: Scarabaeoidea. Nachtrag zur 33. Familie: Scarabaeidae. 14. Gattung: Aphodius Illiger. In: Klausnitzer B. (ed.), Die Larven der Kafer Mitteleuropas. 4. Band. Polyphaga. Pars 3 sowie Erganzungen zum 1. bis 3. Band. Goecke & Evers, Krefeld, pp. 106 - 123."]}

Orphnus mysoriensis Westwood, 1845 Figs 1–2 Orphnus mysoriensis Westwood, 1845: 176. Orphnus detegens Walker, 1859a: 54 (synonymy by Arrow 1912). Orphnus scitissimus Walker, 1859b: 220 (synonymy by Arrow 1912). Orphnus mysoriensis – Lacordaire 1856: 130 (catalogue). — Gemminger & Harold 1869: 1073 (catalogue). — Arrow 1912: 29 (catalogue, synonymy). — Schmidt 1913: 80 (catalogue). — Mittal 2005: 46 (catalogue). — Frolov 2012: 794 (catalogue). — Khaustov & Frolov 2018: 182 (host of Trochometridium tribulatum Cross, 1965). Orphnus detegens – Preudhomme de Borre 1886: 26 (catalogue). Orphnus scitissimus – Preudhomme de Borre 1886: 26 (catalogue). Differential diagnosis Orphnus mysoriensis is similar to O. medvedevi sp. nov. in having the pronotum with a more or less developed tubercle on the base medially and the endophallus with one group of spinules. It differs from the latter in body length (8.0–10.0 mm, as opposed to 4.6–6.0 mm in O. medvedevi sp. nov.), spinules of endophallus as a rather dispersed group of more than 10 spinules (as opposed to a compact cluster consisting of less than 10 spinules in O. medvedevi sp. nov., Fig. 1G vs Fig. 3G), and tubercle on the base of pronotum normally with two punctures (as opposed to always smooth tubercle in O. medvedevi sp. nov.). Type material of Orphnus mysoriensis Lectotype (here designated, Fig. 1A–E, G) INDIA • ♂; “Mysore / Orphnus mysoriensis Westw / TYPE WESTWOOD Trans. Ent. Soc. 4. 1846. P.176. Coll. Hope Oxon. / TYPE COL: 484 1/3 Orphnus mysoriensis Westw. HOPE DEPT.OXFORD”; OUM. Paralectotypes INDIA • 1 ♂; “ Mysore / TYPE COL: 484 2/3 Aegidium mysoriensis Westw. HOPE DEPT.OXFORD”; OUM • 1 ♀; “ Mysore / TYPE COL: 484 3/3 Aegidium mysoriensis Westw. HOPE DEPT.OXFORD”; OUM. Type material of Orphnus detegens Lectotype (here designated, Fig. 2 E – G) SRI LANKA • ♂; “ Type / Ceylon 59 106 / Orphnus detegens Walker. a.n.H. (type) / detegens Wlk ”; BMNH. Additional material examined SRI LANKA – Central Province • 3 ♂♂, 4 ♀♀; Kandy; 7°18′ N, 80°38′ E; MHNG • 2 ♂♂; same locality data as for preceding; MNHB • 7 ♂♂; same locality data as for preceding; 1–18 Apr. 1991; Jiri Kolibac leg.; NHMB • 1 ♀; same locality data as for preceding; 18 Mar. 1973; G. Zimmermann leg.; MNHB • 2 ♂♂; same locality data as for preceding; 1908; G.B. Longstaff leg.; OUM • 2 ♂♂, 2 ♀♀; Nalanda; 7°40′ N, 80°39′ E; V. De Poll leg.; MNHN • 1 ♀; same locality data as for preceding; 1889; I.Z. Kannegeter leg.; MNHN • 2 ♂♂, 1 ♀; Peradenyia; 7°16′ N, 80°35′ E; MHNG • 3 ♂♂; same locality data as for preceding; Friederichs S.G. leg.; MNHB • 2 ♂♂; same locality data as for preceding; 2 Apr. 1907; O. John leg.; ZIN • 1 ♂; same locality data as for preceding; 3 Apr. 1907; O. John leg.; ZIN • 2 ♂♂, 1 ♀; same locality data as for preceding; Apr. 1901; MNHN • 5 ♂♂; same locality data as for preceding; Apr. 1914; Friederichs S.G. leg.; MNHB • 2 ♂♂; same locality data as for preceding; Apr. 1939; J. Vinson leg.; MNHN • 1 ♀; Punduloya; 7°01′ N, 80°40′ E; OUM • 1 ♂, 1 ♀; Sigiriya; 7°57′ N, 80°45′ E; Oct. 1977; MCSN • 1 ♂, 1 ♀; same locality data as for preceding; Oct. 1977; NMPC • 1 ♂; Weragamtota; 7°19′ N, 80°59′ E; 13 Sep. 1953; F. Keiser leg.; NHMB. – North Central Province • 2 ♀♀; Anuradhapura; 8°19′ N, 80°24′ E; Friederichs S.G. leg.; MNHB • 4 ♂♂; Kala Oya; 8°13′ N, 80°06′ E; 7 Nov. 1983; MNHN • 1 ♂; same locality data as for preceding; 8 Nov. 1983; MNHN • 2 ♂♂; Nat. Park Wilpattu; 8°25′ N, 80°03′ E; 7 Oct. 1982; G.S. Medvedev leg.; ZIN • 1 ♂; Nat. Park Wilpattu, Talawila; 8°25′ N, 80°03′ E; 9 Oct. 1982; G.S. Medvedev leg.; ZIN • 1 ♂, 3 ♀♀; Wilpattu N.P. 17 km WNW entrance Tala Wila; 8°25′ N, 80°03′ E; 8 Oct. 1982; V.F. Zaitzev leg.; ZIN. – North Western Province • 2 ♂♂, 1 ♀; Puttalam; 8°02′ N, 79°50′ E; 1899; W. Horn leg.; MNHN • 1 ♂; Rajakadaluwa; 7°39′ N, 79°50′ E; 22 Aug. 1953; F. Keiser leg.; NHMB. – Sabaragamuwa Province • 2 ♀♀; Sabaragamuwa, Ratnapura; 6°45′ N, 80°30′ E; 20 Jan. 1970; Mussard, Besuchet, Lobl leg.; MCSN. – Southern Province • 3 ♀♀; Bentota; 6°26′ N, 79°60′ E; 14 Aug. 1978; P. Cabella leg.; MCSN • 1 ♂; same locality data as for preceding; 16 Aug. 1978; P. Cabella leg.; MCSN • 1 ♀; same locality data as for preceding; 20 Aug. 1978; P. Cabella leg.; MCSN • 3 ♂♂, 5 ♀♀; same locality data as for preceding; 23 Mar. 1973; G. Zimmermann leg.; MNHB • 1 ♀; Matara; 5°57′ N, 80°33′ E; 20 Oct. 1953; F. Keiser leg.; NHMB. – Western Province • 1 ♀; Colombo; 6°56′ N, 79°51′ E; MNHN • 1 ♂; same locality data as for preceding; MHNG • 1 ♂; same locality data as for preceding; Feb. 1884; MNHB • 1 ♂, 1 ♀; same locality data as for preceding; Mar. 1953; MHNG • 3 ♂♂; Hendala; 6°59′ N, 79°53′ E; 15 Aug. 1979; MNHN • 3 ♀♀; Kalutara; 6°35′ N, 79°58′ E; Mar. 1986; W.U.E. Just leg.; NMPC • 2 ♂♂; Kalutara Distr., Tebuwana; 6°35′ N, 80°03′ E; 13 Mar. 2000; S. Mahunka and L. Mahunka-Papp leg.; HNHM • 1 ♀; Mount Lavinia; 6°50′ N, 79°52′ E; 5 Apr. 1973; G. Benick leg.; MNHB • 5 ♂♂, 4 ♀♀; Wadduwa; 6°40′ N, 79°56′ E; V. De Poll leg.; MNHN. – Sri Lanka (no exact locality) • 9 ♂♂, 1 ♀; MNHN • 3 ♂♂, 2 ♀♀; IRSNB • 1 ♂; OUM • 1 ♂; NMPC • 2 ♂♂; ZIN • 4 ♂♂, 5 ♀♀; MNHB • 1 ♂, 1 ♀; MHNG • 1 ♀; Diener leg.; HNHM • 3 ♀♀; Friederichs S.G. leg.; MNHB • 2 ♀♀; S. Niethner leg.; MNHB • 1 ♂; Templeton leg.; OUM • 1 ♂; Walker leg.; BMNH • 2 ♂♂; Wollaston leg.; OUM • 11 ♂♂; 1871; Thwaites leg.; OUM • 2 ♂♂; 1873; Thwaites leg.; OUM • 5 ♂♂; 1889; H.Fruhstorfer leg.; MNHN • 1 ♂; Mar. 1889; H. Fruhstorfer leg.; MNHN. Variation Body length of the examined specimens varied from 8.0 to 10.5 mm (males) and from 7.0 to 9.0 mm (females). Head and pronotum armature in males varied from well-developed with long lateral pronotal processes and frontoclypeal horn (Fig. 2A) to excavated pronotum with gibbosities aside of the excavation and frontoclypeal tubercle (Fig. 2C), with intermediate variants (Fig. 2B). Distribution The species is distributed rather widely on Sri Lanka except for the eastern part (Fig. 1H)., Published as part of Frolov, Andrey V. & Akhmetova, Lilia A., 2021, Review of the Orphninae (Coleoptera: Scarabaeidae) of Sri Lanka, with description of a new species of genus Orphnus Macleay, 1819, pp. 40-54 in European Journal of Taxonomy 767 (1) on pages 41-45, DOI: 10.5852/ejt.2021.767.1485, http://zenodo.org/record/5514853, {"references":["Westwood J. O. 1845. On the lamellicorn beetles which possess exserted mandibles and labrum, and 10 - jointed antennae. Transactions of the Entomological Society of London 4: 155 - 180. https: // doi. org / 10.1111 / j. 1365 - 2311.1846. tb 01350. x","Walker F. 1859 a. Characters of some apparently undescribed Ceylon insects. The Annals and Magazine of Natural History, Including Zoology, Botany and Geology Series 3 3: 50 - 56. Available from https: // www. biodiversitylibrary. org / page / 2317765 [accessed 25 Aug. 2021].","Arrow G. J. 1912. Pachypodinae, Pleocominae, Aclopinae, Glaphyrinae, Ochodaeinae, Orphninae, Idiostominae, Hybosorinae, Dynamopinae, Acanthocerinae, Troginae. In: Junk W. & Schenkling S. (eds) Coleopterorum Catalogus 43: 1 - 66. W. Junk, Berlin.","Walker F. 1859 b. Characters of some apparently undescribed Ceylon insects. The Annals and Magazine of Natural History, Including Zoology, Botany and Geology Series 3 4: 217 - 224. https: // doi. org / 10.1080 / 00222935908697111","Lacordaire T. 1856. Histoire naturelle des Insectes. Genera des Coleopteres ou expose methodique et critique de tous les genres proposes jusqu'ici dans cet ordre d'insectes. Tome troiseme contenant les familles des pectinicorne et lamellicornes. Roret, Paris. https: // doi. org / 10.5962 / bhl. title. 8864","Gemminger M. & Harold E. 1869. Catalogus Coleopterorum hucusque descriptorum synonymicus et systematicus. Vol. 4 Scarabaeidae: 979 - 1346. E. H. Gummi, Munich. https: // doi. org / 10.5962 / bhl. title. 9089","Schmidt A. 1913. Coleoptera. Lamellicornia. Fam. Scarabaeidae. Subfam. Aegialinae, Chironinae, Dynamopinae, Hybosorinae, Idiostominae, Ochodaeinae, Orphninae. Genera Insectorum 150. V. Verteneuil & L. Desmet, Bruxelles.","Mittal I. 2005. Diversity and conservation status of dung beetles (Laparosticti: Scarabaeidae: Coleoptera) in North India. Bulletin of the National Institute of Ecology 15: 43 - 51.","Frolov A. V. 2012. Diagnosis, classification, and phylogenetic relationships of the orphnine scarab beetles (Coleoptera, Scarabaeidae: Orphninae). Entomological Review 92: 782 - 797. https: // doi. org / 10.1134 / S 0013873812070056","Khaustov A. A. & Frolov A. V. 2018. A new species, new genus and new records of heterostigmatic mites (Acari: Heterostigmata) phoretic on scarab beetles of the subfamily Orphninae (Coleoptera: Scarabaeidae). Zootaxa 4514: 181 - 201. https: // doi. org / 10.11646 / zootaxa. 4514.2.3","Preudhomme de Borre A. 1886. Descriptions de deux especes nouvelles du genre Aegidium Westwood suivies de la liste des Orphnides du Musee royal d'histoire naturelle de Belgique. Annales de la Societe entomologique de Belgique 30: 24 - 26. Available from https: // www. biodiversitylibrary. org / page / 12278747 [accessed 15 Aug. 2021]."]}

Orphnus bicolor (Fabricius, 1801) Fig. 4 Scarabaeus bicolor Fabricius, 1801: 9. Orphnus bicolor – MacLeay 1819: 119. — Westwood 1845: 176. — Lacordaire 1856: 130 (catalogue). — Gemminger & Harold 1869: 1072 (catalogue). — Arrow 1912: 29 (catalogue). — Schmidt 1913: 79 (catalogue). — Frolov 2012: 793 (catalogue). Differential diagnosis Orphnus bicolor is similar to O. parvus in having the pronotum without tubercle on base medially and endophallus with a few groups of spinules. It differs from the latter in having the parameres in dorsal view with small but distinct notches basad of the lateral teeth (Fig. 4F), longer parameres (about 0.7 length of phallobase [Fig. 4E] as opposed to 0.5 length of phallobase in O. parvus [Fig. 5E]), and abdominal sternite 8 medially longer than sternites 4–7 combined (as opposed to sternite 8 medially as long as sternites 4–7 combined in O. parvus). Type material Lectotype (here designated, Fig. 4 A – F) INDIA • ♂; “ TYPE / Ind. orient. Daldorff Mus. T. Lund Geotrupes bicolor F. / Lectotype Geotrupes bicolor F. A.Frolov des. 2016”; ZMUKG. Paralectotype INDIA • 1 ♂; “ TYPE / Paralectotype Geotrupes bicolor F. A.Frolov des. 2016”; ZMUKG. Additional material examined SRI LANKA – Central Province • 1 ♂; Weragamtota; 7°19′ N, 80°59′ E; 13 Sep. 1953; F. Keiser leg.; NHMB. – Southern Province • 1 ♂, 1 ♀; Tissamaharana; 6°17′ N, 81°17′ E; 30 Oct. 1983; MNHN. Variation Body length of the examined specimens varies from 7.5 to 9.0 mm (males) and 8.6 mm (female). Distribution The species was described form “East India ”. In Sri Lanka, it is known from two rather distant localities in the centre and on the southern coast (Fig. 4I)., Published as part of Frolov, Andrey V. & Akhmetova, Lilia A., 2021, Review of the Orphninae (Coleoptera: Scarabaeidae) of Sri Lanka, with description of a new species of genus Orphnus Macleay, 1819, pp. 40-54 in European Journal of Taxonomy 767 (1) on page 48, DOI: 10.5852/ejt.2021.767.1485, http://zenodo.org/record/5514853, {"references":["Fabricius J. C. 1801. Systema Eleutheratorum secundum ordines, genera, species: adiectis synonymis, locis, observationibus, descriptionibus. Tomus I. Impensis Bibliopolii Academici novi. Kiel.","MacLeay W. S. 1819. Horae Entomologicae: or Essays on the Annulose Animals. Vol. 1 Part 1. S. Bagster, London. https: // doi. org / 10.5962 / bhl. title. 48636","Westwood J. O. 1845. On the lamellicorn beetles which possess exserted mandibles and labrum, and 10 - jointed antennae. Transactions of the Entomological Society of London 4: 155 - 180. https: // doi. org / 10.1111 / j. 1365 - 2311.1846. tb 01350. x","Lacordaire T. 1856. Histoire naturelle des Insectes. Genera des Coleopteres ou expose methodique et critique de tous les genres proposes jusqu'ici dans cet ordre d'insectes. Tome troiseme contenant les familles des pectinicorne et lamellicornes. Roret, Paris. https: // doi. org / 10.5962 / bhl. title. 8864","Gemminger M. & Harold E. 1869. Catalogus Coleopterorum hucusque descriptorum synonymicus et systematicus. Vol. 4 Scarabaeidae: 979 - 1346. E. H. Gummi, Munich. https: // doi. org / 10.5962 / bhl. title. 9089","Arrow G. J. 1912. Pachypodinae, Pleocominae, Aclopinae, Glaphyrinae, Ochodaeinae, Orphninae, Idiostominae, Hybosorinae, Dynamopinae, Acanthocerinae, Troginae. In: Junk W. & Schenkling S. (eds) Coleopterorum Catalogus 43: 1 - 66. W. Junk, Berlin.","Schmidt A. 1913. Coleoptera. Lamellicornia. Fam. Scarabaeidae. Subfam. Aegialinae, Chironinae, Dynamopinae, Hybosorinae, Idiostominae, Ochodaeinae, Orphninae. Genera Insectorum 150. V. Verteneuil & L. Desmet, Bruxelles.","Frolov A. V. 2012. Diagnosis, classification, and phylogenetic relationships of the orphnine scarab beetles (Coleoptera, Scarabaeidae: Orphninae). Entomological Review 92: 782 - 797. https: // doi. org / 10.1134 / S 0013873812070056"]}

Orphnus parvus (Wiedemann, 1823) Fig. 5 Geotrupes parvus Wiedemann, 1823: 6. Orphnus nanus Westwood, 1845: 177 (synonymy by Arrow 1912). Orphnus parvus ��� Gemminger & Harold 1869: 1073 (catalogue). ��� Arrow 1912: 30 (catalogue). ��� Schmidt 1913: 80 (catalogue). ��� Frolov 2012: 794 (catalogue). Differential diagnosis Orphnus parvus is similar to O. bicolor in having the pronotum without a tubercle on base medially and endophallus with a few groups of spinules. It differs from the latter in having the parameres in dorsal view without notches basad of lateral teeth (Fig. 5G), shorter parameres (about 0.5 length of phallobase [Fig. 5E] as opposed to 0.7 length of phallobase in O. bicolor [Fig. 4E]), and abdominal sternite 8 medially as long as sternites 4���7 combined (as opposed to sternite 8 medially distinctly longer than sternites 4���7 combined in O. bicolor). Type material of Geotrupes parvus Lectotype (here designated, Fig. 5 A ��� C) INDIA ��� ♂; ���Bengal Mai 1808 Parvus ♂ Wied. / Mus. Westerm. / TYPE ���; ZMUKK. Paralectotype INDIA ��� 1 ♂; ���Mus. Westerm. / TYPE ���; ZMUKK. Type material of Orphnus nanus Lectotype (here designated, Fig. 5 D ��� H) INDIA ��� 1 ♂; ��� TYPE COL: 482 1/2 Orphnus nanus Westw. HOPE DEPT.OXFORD / Orphnus nanus Westw East Ind [...] Hearsey / TYPE WESTWOOD Trans. Ent. Soc. 4.1846.P.177 Coll. Hope Oxon. / [���] / W / LECTOTYPUS Orphnus nanus Westw. Frolov et al. 2016 ���; ZMUKK. Paralectotype INDIA ��� 1♂;��� TYPE COL: 482 2/2 Orphnus nanus Westw. HOPE DEPT.OXFORD / TYPE WESTWOOD Trans. Ent. Soc. 4.1846.P.177 Coll. Hope Oxon. / Prof. Westwood���s private collection purchased from Miss Swann 1895 Collected by Gen. Hearsey in India / [9���H] / Orphnus nanus Westw ���; ZMUKK. Additional material examined SRI LANKA ��� Central Province ��� 2 ♂♂; Kandy; 7��18��� N, 80��38��� E; MHNG ��� 1 ♀; Peradenyia; 7��16��� N, 80��35��� E; MHNG ��� 1 ♂, 1 ♀; same locality data as for preceding; Friederichs S.G. leg.; MNHB ��� 1 ♂; same locality data as for preceding; 3 Apr. 1907; O. John leg.; ZIN ��� 1 ♀; Sigiriya; 7��57��� N, 80��45��� E; Oct. 1977; MHNG ��� 2 ♀♀; Weragamtota; 7��19��� N, 80��59��� E; 13 Sep. 1953; F. Keiser leg.; NHMB. ��� North Central Province ��� 1 ♀; Anuradhapura; 8��19��� N, 80��24��� E; Friederichs S.G. leg.; MNHB ��� 1 ♂; Kala Oya; 8��13��� N, 80��06��� E; 7 Nov. 1983; MNHN ��� 1 ♀; same locality data as for preceding; 8 Nov. 1983; MNHN ��� 1 ♀; Vilpattu Nat. Park, Talawila; 8��25��� N, 80��03��� E; 9 Nov. 1983; G.S. Medvedev leg.; ZIN. ��� Southern Province ��� 1 ♀; 9 km SW of Yala; 6��22��� N, 81��31��� E; 21 Oct. 1982; V.F. Zaitzev leg.; ZIN ��� 8 ♂♂; Southern Province, Bentota; 6��26��� N, 79��60��� E; 23 Mar. 1973; G. Zimmermann leg.; MNHB ��� 1 ♂; Point de Galle; 6��01��� N, 80��13��� E; MHNG ��� 1 ♂; Tissamaharama; 6��17��� N, 81��17��� E; 8 Aug. 1978; P. Cabella leg.; MCSN ��� 2 ♂♂, 2 ♀♀; same locality data as for preceding; 30 Oct. 1983; MNHN ��� 2 ♂♂; Yala; 6��22��� N, 81��31��� E; 21 Oct. 1982; G.S. Medvedev leg.; ZIN. ��� Western Province ��� 1 ♂; Colombo; 6��56��� N, 79��51��� E; Friederichs S.G. leg.; MNHB ��� 1 ♂; same locality data as for preceding; Mar. 1953; G. Frey leg.; MHNG ��� 1 ♂, 1 ♀; Negombo; 7��12��� N, 79��50��� E; 23 Mar. 1973; B. Huttler leg.; NMPC ��� 1 ♀; Wadduwa; 6��40��� N, 79��56��� E; V. De Poll leg.; MNHN. ��� Sri Lanka (no exact locality) ��� 2 ♂♂, 1 ♀; OUM ��� 1 ♀; MHNG ��� 1 ♂; MNHB ��� 1 ♂; Friederichs S.G. leg.; MNHB ��� 1 ♂; Parry leg.; IRSNB ��� 1 ♂; S. Niethner leg.; MNHB. Variation Body length of the examined specimens varies from 6.1 to 8.2 mm (males) and 6.0 to 7.5 mm (females). Distribution The species is distributed rather widely throughout Sri Lanka except for the north-eastern part (Fig. 5J)., Published as part of Frolov, Andrey V. & Akhmetova, Lilia A., 2021, Review of the Orphninae (Coleoptera: Scarabaeidae) of Sri Lanka, with description of a new species of genus Orphnus Macleay, 1819, pp. 40-54 in European Journal of Taxonomy 767 (1) on pages 48-52, DOI: 10.5852/ejt.2021.767.1485, http://zenodo.org/record/5514853, {"references":["Wiedemann C. R. W. 1823. Zweihundert neue Kafer von Java, Bengalen, und dem Vorgebirge der guten Hoffnung. Zoologisches Magazin Kiel 2: 1 - 133. Available from https: // www. biodiversitylibrary. org / page / 14928532 [accessed 25 Aug. 2021].","Westwood J. O. 1845. On the lamellicorn beetles which possess exserted mandibles and labrum, and 10 - jointed antennae. Transactions of the Entomological Society of London 4: 155 - 180. https: // doi. org / 10.1111 / j. 1365 - 2311.1846. tb 01350. x","Arrow G. J. 1912. Pachypodinae, Pleocominae, Aclopinae, Glaphyrinae, Ochodaeinae, Orphninae, Idiostominae, Hybosorinae, Dynamopinae, Acanthocerinae, Troginae. In: Junk W. & Schenkling S. (eds) Coleopterorum Catalogus 43: 1 - 66. W. Junk, Berlin.","Gemminger M. & Harold E. 1869. Catalogus Coleopterorum hucusque descriptorum synonymicus et systematicus. Vol. 4 Scarabaeidae: 979 - 1346. E. H. Gummi, Munich. https: // doi. org / 10.5962 / bhl. title. 9089","Schmidt A. 1913. Coleoptera. Lamellicornia. Fam. Scarabaeidae. Subfam. Aegialinae, Chironinae, Dynamopinae, Hybosorinae, Idiostominae, Ochodaeinae, Orphninae. Genera Insectorum 150. V. Verteneuil & L. Desmet, Bruxelles.","Frolov A. V. 2012. Diagnosis, classification, and phylogenetic relationships of the orphnine scarab beetles (Coleoptera, Scarabaeidae: Orphninae). Entomological Review 92: 782 - 797. https: // doi. org / 10.1134 / S 0013873812070056","Fabricius J. C. 1801. Systema Eleutheratorum secundum ordines, genera, species: adiectis synonymis, locis, observationibus, descriptionibus. Tomus I. Impensis Bibliopolii Academici novi. Kiel.","Frolov A. V., Montreuil O. & Akhmetova L. A. 2016. Review of the Madagascan Orphninae (Coleoptera: Scarabaeidae) with a revision of the genus Triodontus Westwood. Zootaxa 4207: 1 - 93. https: // doi. org / 10.11646 / zootaxa. 4207.1.1"]}

Orphnus medvedevi sp. nov. urn:lsid:zoobank.org:act: 3F99F2E5-0FD6-4CE9-A3AC-53A7FBA4241D Fig. 3 Differential diagnosis Orphnus medvedevi sp. nov. is similar to O. mysoriensis in having the pronotum with more or less developed tubercle on base medially and endophallus with one group of spinules. It differs from the latter in shorter body (4.6–6.0 mm, as opposed to 8.0–10.0 mm. in O. mysoriensis), spinules of endophallus as a compact cluster consisting of less than 10 spinules (as opposed to a rather dispersed group of more than 10 spinules in O. mysoriensis, Fig. 3G vs Fig. 1G), and tubercle on the base of pronotum always smooth (as opposed to having normally two punctures in O. mysoriensis). Etymology The new species is named after Gleb Sergeevich Medvedev (1931–2009), a Russian coleopterist. Type material Holotype SRI LANKA • ♂; “ Shri Lanka Vilpattu [Sri Lanka, Wilpattu] 7.10.1982 G. Medvedev ”; ZIN. Paratypes SRI LANKA • 2 ♀♀; “ Shri Lanka Vilpattu [Sri Lanka, Wilpattu] 8.10.1982 G. Medvedev ”; ZIN • 1 ♂, 6 ♀♀; “ Sri Lanka. Nat. Park Wilpattu. Talawila 9.X. 982. Medvedev”; ZIN • 13 ♂♂, 13 ♀♀; “ Sri Lanka, Wilpattu N.P. 17 km WNW entrance Tala Wila, 8.X.1982 V.F. Zaitzev leg.”; ZIN. Description Male, holotype (Fig. 3A, D–G) BODY. Length 6.0 mm. Colour uniformly brown. CLYPEUS. Wide, with convex anterior margin, rounded laterally, finely crenulate. Genae small, not protruding past eyes. Frontal suture indistinct. Clypeus with long, slender horn directed upwards and slightly curved backwards apically. Dorsal surface of head finely punctate. Labrum deeply sinuate in the middle, distinctly protruding past clypeus. PRONOTUM. With rounded sides, about 1.5 times as wide as long, with deeply concave disc, conical lateral processes aside excavation, and smooth, rounded tubercle medially near base. Anterior angles acute; posterior angles rounded, indistinct in dorsal view. Pronotum bordered on anterior margin and base. Lateral margins with long, sparse, brown setae. Sides irregularly punctate with round punctures separated by 3–5 puncture diameters. SCUTELLUM. Subtriangular, narrowly rounded apically, about 1/10 length of elytra. ELYTRA. About as long as wide, with distinct humeral humps, widest in middle, lateral margins slightly rounded in basal half. First (sutural) stria distinct, as feebly impressed groove with row of punctures. Other stria before humeral humps as rows of round setiferous punctures, separated by more than 3 their diameters except for base of elytra. Elytral intervals covered with minute punctures. WINGS. Macropterous. LEGS. Protibiae with 3 outer teeth. Lateral margin basad of outer teeth not crenulate. Apical spur of protibia absent. Left protarsus absent. Middle and hind legs similar in shape; metafemora and metatibiae about ⅛ as long as the mesofemora and mesotibiae. Mesotibia and metatibiae somewhat triangular with 2 apical spurs, inner margin almost straight, with 1 transverse keel. Upper spur of tibiae as long as two basal tarsomeres. Claws ⅓ length of apical tarsomere. Femora almost impunctate. ABDOMEN. Ventrally irregularly punctate, pubescent, with sparse, long setae. Abdominal sternite 8 medially shorter than sternites 4–7 combined. Pygidium invisible from above, with slightly truncate apex in caudal view. Plectrum triangular with rounded apex, wider than long. AEDEAGUS. Parameres relatively short (0.5 length of phallobase), curved downwards, tapering apically, spear-shaped in dorsal view, with feebly visible lateral teeth (Fig. 3D–E). Endophallus with a compact cluster of 6 spinules (Fig. 3G). Female Female (Fig. 3H) differs from the male in having a relatively smaller pronotum without armature, frontoclypeus without process, and prothoracic spur present. Variation Body length of the paratypes varies from 5.9 to 4.6 mm (males) and 4.7 to 6.0 mm (females). Some of the paratypes have darker elytra. Head and pronotum armature in males paratypes varies from relatively well-developed, similar to that of the holotype, to excavated pronotum with gibbosities beside the excavation and frontoclypeal tubercle (Fig. 3C), with intermediate variants (Fig. 3B). Distribution The species is known from one locality in north-western Sri Lanka (Fig. 3I)., Published as part of Frolov, Andrey V. & Akhmetova, Lilia A., 2021, Review of the Orphninae (Coleoptera: Scarabaeidae) of Sri Lanka, with description of a new species of genus Orphnus Macleay, 1819, pp. 40-54 in European Journal of Taxonomy 767 (1) on pages 45-48, DOI: 10.5852/ejt.2021.767.1485, http://zenodo.org/record/5514853

Key to species of Orphnus from Sri Lanka (males) 1. Pronotum with more or less developed tubercle on base medially (Figs 1A–B, 2A–C, 3A–C). Endophallus with 1 group of spinules (Figs 1G, 3G)....................................................................... 2 – Pronotum without tubercle on base medially (Figs 4A, H, 5A, H). Endophallus with more than 1 distinct group of spinules (Figs 4D, 5D).......................................................................................... 3 2. Larger: body length 8.0–10.0 mm. Spinules of endophallus as a rather dispersed group of more than 10 spinules (Fig. 1G). Tubercle on base of pronotum normally with 2 punctures; punctures may be close to each other or indistinct in some specimens............. Orphnus mysoriensis Westwood, 1845 – Smaller: body length 4.6–6.0 mm. Spinules of endophallus in a compact cluster consisting of less than 10 spinules (Fig. 3G). Tubercle on base of pronotum smooth...... Orphnus medvedevi sp. nov. 3. Parameres in dorsal view with small but distinct notches basad of lateral teeth (Fig. 4F). Parameres longer, about 0.7 length of phallobase (Fig. 4E). Abdominal sternite 8 medially longer than sternites 4–7 combined............................................................................... Orphnus bicolor (Fabricius, 1801) – Parameres in dorsal view without notches basad of lateral teeth (Fig. 5G). Parameres shorter, about 0.5 length of phallobase (Fig. 5F). Abdominal sternite 8 medially as long as sternites 4–7 combined.................................................................................. Orphnus parvus (Wiedemann, 1823), Published as part of Frolov, Andrey V. & Akhmetova, Lilia A., 2021, Review of the Orphninae (Coleoptera: Scarabaeidae) of Sri Lanka, with description of a new species of genus Orphnus Macleay, 1819, pp. 40-54 in European Journal of Taxonomy 767 (1) on page 52, DOI: 10.5852/ejt.2021.767.1485, http://zenodo.org/record/5514853, {"references":["Westwood J. O. 1845. On the lamellicorn beetles which possess exserted mandibles and labrum, and 10 - jointed antennae. Transactions of the Entomological Society of London 4: 155 - 180. https: // doi. org / 10.1111 / j. 1365 - 2311.1846. tb 01350. x","Fabricius J. C. 1801. Systema Eleutheratorum secundum ordines, genera, species: adiectis synonymis, locis, observationibus, descriptionibus. Tomus I. Impensis Bibliopolii Academici novi. Kiel.","Wiedemann C. R. W. 1823. Zweihundert neue Kafer von Java, Bengalen, und dem Vorgebirge der guten Hoffnung. Zoologisches Magazin Kiel 2: 1 - 133. Available from https: // www. biodiversitylibrary. org / page / 14928532 [accessed 25 Aug. 2021]."]}

Cerhomalus absconditus (Petrovitz, 1971) comb. nov. Figs 2, 5A, 6 Orphnus (Cerhomalus) absconditus Petrovitz, 1971: 3. Differential diagnosis Cerhomalus absconditus can be easily separated from congeners by having the apices of the parameres rounded (in apical view), without semi-circular excavations (Fig. 2G, J) and without keels or fossae on ventral side (Fig. 2K), and the base of elytra with somewhat tile-shaped punctures (Fig. 2 A���B). Material examined Holotype DEMOCRATIC REPUBLIC OF THE CONGO ��� ♂; ���COLL. MUS. CONGO Tshuapa: Yokamba /1953 J. Stevenard / HOLOTYPUS / O. (Cerhomalus) absconditus n.sp. Petrovitz���; RMCA. Paratypes DEMOCRATIC REPUBLIC OF THE CONGO ��� 1 ♀ ���COLL. MUS. CONGO Kasai: Ipamu -1937 R. Soeur Imelda / PARATYPUS / O. (Cerhomalus) absconditus n.sp. Petrovitz���; RMCA ��� 5 ♀♀; ���COLL. MUS. CONGO Kwango: Djuma 1953 /R.P.G. Leta/ PARATYPUS / O. (Cerhomalus) absconditus n.sp. Petrovitz / coll. Petrovitz ���; RMCA ��� 1 ♂; ���COLL. MUS. CONGO Lulua: Kapanga ��� XI-1933 F.G. Overlaet / PARATYPUS / O. (Cerhomalus) absconditus n.sp. Petrovitz���; RMCA ��� 1 ♂; ���MUS. ROY. AFR. CENTR. Kasai: terr. Dekese I.1960 F. Francois / PARATYPUS / O. (Cerhomalus) absconditus n.sp. Petrovitz���; RMCA ��� 1 ♀; ���MUSEE DU CONGO Kasai: Don de la Cie du Kasai / PARATYPUS / R. DET. 1939 F / O. (Cerhomalus) absconditus n.sp. Petrovitz���; RMCA ��� 1 ♂; ���MUSEE DU CONGO Kasai: Ipamu - VIII -1922 P. Vanderijst / PARATYPUS / R. DET. 1939 F / O. (Cerhomalus) absconditus n.sp. Petrovitz���; RMCA ��� 2 ♂♂; ���Musee du Congo Kinda (Katanga) 1927 (Don M. Muller.) / Kinda Katanga 1926 Congo / O. (Cerhomalus) absconditus n.sp. Petrovitz / PARATYPUS / R. DET. 1939 F / 139���; MHNG ��� 1 ♀; ���MUSEE DU CONGO Murlambongo (Idiofa) - 1930 (Les Soeurs de la Mission) / PARATYPUS / O. (Cerhomalus) absconditus n.sp. Petrovitz���; RMCA ��� 1 ♂; ���Musee du Congo Sankuru: Komi (Lodja) 27-I-1930 J. Ghesquiere / R. DET 1939 F / Cerhomalus mechowi Qued. / O. (Cerhomalus) absconditus n.sp. Petrovitz / PARATYPUS ���; RMCA ��� 1 ♂; ���COLL. MUS. CONGO Sankuru: Katako-Kombe VIII - 1952 Dr M. Fontaine / O. (Cerhomalus) absconditus n.sp. Petrovitz / PARATYPUS / PARATYPUS���; MCSNG ��� 1 ♂, 1 ♀; ���COLL. MUS. CONGO Tshuapa: Ikela 1956 R. Deguide / PARATYPUS / O. (Cerhomalus) absconditus n. sp. Petrovitz���; RMCA ��� 1 ♀; ���COLL. MUS. CONGO Tshuapa: Ikela 1956 R.P. Lootens / PARATYPUS / O. (Cerhomalus) absconditus n. sp. Petrovitz���; RMCA ��� 1 ♀; ���COLL. MUS. CONGO Tshuapa: Bosekele VI-1952 J. Van Vynckt / O. (Cerhomalus) absconditus n.sp. Petrovitz / PARATYPUS ���; RMCA ��� 1 ♀; ���COLL. MUS. CONGO Tshuapa: Yokamba /1953 J. Stevenard / ALLOTYPUS / O. (Cerhomalus) absconditus n.sp. Petrovitz���; RMCA ��� 1 ♂; ���COLL. MUS. CONGO Terr. de Dibaya Kamponde 1945 Rev. Fr. ALLAER 1945 / PARATYPUS / O. (Cerhomalus) absconditus n. sp. Petrovitz���; RMCA ��� 1 ♀; ���COLL. MUS. CONGO Katoka-Luluabourg / -1938 R. P. VANKERCKHOVEN / PARATYPUS / O. (Cerhomalus) absconditus n.sp. Petrovitz / coll. Petrovitz ���; RMCA ��� 1 ♂; ���MUSEE DU CONGO Luebo - III-1931. 1 ex. J.P. Colin / O. (Cerhomalus) absconditus n.sp. Petrovitz / PARATYPUS���; RMCA. Other material DEMOCRATIC REPUBLIC OF THE CONGO ��� 1 ♀; ��quateur, Tshuapa, Etata; Aug.���Sep. 1969; J. Hauwaert leg.; RMCA. ANGOLA ��� 1 ♂, 3 ♀♀; Bi��, Catabola; 15���27 Sep. 2012; T. Lackner leg.; BMNH ��� 4 ♀♀; Lunda Norte, Andrada; 7��41��� S, 21��17��� E; Aug.���Sep. 1952; M. Fetchkowsky leg.; RMCA ��� 2 ♀♀; Dundo; Sep. 1948; Machado leg.; RMCA ��� 1 ♂; Cuando Cubango Province, N of Menongue City; 1600 m a.s.l.; 22 Nov. 2019; S. Dementev leg.; SDCMR. CAMEROON ��� 1 ♂; 1912; BMNH. Distribution This is the most widely distributed species of the genus. The majority of records are from the Congo basin, but the species occurs up to Cameroon in the north (no precise locality is recorded) and to the Ancient Plateau in Angola in the south (Fig. 6)., Published as part of Frolov, Andrey V. & Akhmetova, Lilia A., 2021, Taxonomic revision of the Afrotropical scarab beetle genus Cerhomalus Quedenfeldt, 1884 (Coleoptera: Scarabaeidae: Orphninae), pp. 36-50 in European Journal of Taxonomy 739 (1) on pages 40-42, DOI: 10.5852/ejt.2021.739.1265, http://zenodo.org/record/4604232, {"references":["Petrovitz R. 1971. Beitrag zur Kenntnis der Gattung Orphnus M'Leay (Orphninae, Scarabaeidae, Coleoptera). Revue de zoologie et de botanique africaines 84: 1 - 46."]}

Cerhomalus petrovitzi sp. nov. urn:lsid:zoobank.org:act: A04DF237-5C78-4B76-BEDB-7453CCEAB827 Figs 3, 5C, 6 Differential diagnosis Cerhomalus petrovitzi sp. nov. is most similar to C. mechowi in having the ventral fossae on the apices of the parameres completely bordered (Figs 1F, 3G) and the apices right angled in lateral view (Figs 1C, 3D) but can be separated from it in having the widened part of a paramere (in lateral view) about two thirds the length of the paramere (Fig. 3D) and ventral fossa of a paramere longer and narrower (Fig. 3G). Etymology The new species is named after Rudolf Petrovitz (1906���1974), a German coleopterist. Type material Holotype DEMOCRATIC REPUBLIC OF THE CONGO ��� ♂; ���MUSEE DU CONGO BELGE KISANTU P. Goossens ���; MNHN. Paratypes DEMOCRATIC REPUBLIC OF THE CONGO ��� 2 ♂♂, 1 ♀; same collection data as for holotype; RMCA ��� 1 ♂; ���MUSEE DU CONGO Belge [Kasa��-Occidental] (Don M. Corpentier)���; MHNG. Other material NIGERIA ��� 1 ♂; ��� Nigeria W. Afr. / (Staudinger coll.) 1914-412.���; BMNH. Description Male, holotype (Fig. 3 A���B, D���G) Body length 20.1 mm. Colour uniformly brown to black. Anterior margin of frontoclypeus somewhat rectangular, slightly convex in middle, bordered and slightly serrate in dorsal view (Fig. 3A). Frontoclypeus with a keel-shaped transverse process near anterior margin; height of keel about half its width. Surface of frontoclypeus almost smooth, with minute punctures. Eye tubercles feebly developed. Eyes rather large: width about 1/6 distance between eyes in dorsal view. Antennae 10-segmented. Pronotum widely rounded laterally, as wide as elytra. Anterior border wide. Basal border narrow, keelshaped, separated from pronotal disc by deep groove with row of longitudinally elongated punctures. Pronotal disc with a rounded depression occupying more than half of the surface, somewhat rugose anteriorly. Most of pronotum surface covered with minute punctures separated by more than 3 puncture diameters. Anterolateral angles with much larger and denser punctures; posteriolateral angles with same punctures but in smaller numbers. Scutellum subtriangular, narrowly rounded apically, about 1/10 length of elytra. Elytra 1.1 times longer than wide, with distinct humeral humps. Elytra widest in middle, lateral margins almost parallel in basal half. First (sutural) stria distinct, as groove with row of punctures. Other stria before humeral humps as rows of round setiferous punctures. Striae laterad of stria 5 feebly distinct. Elytral intervals covered with minute punctures, somewhat sparser than those on pronotum. Macropterous. Abdominal sternite 8 medially longer than sternites 6 and 7 combined; sternite 6 about as long as sternite 7. Pygidium invisible from above, with slightly truncate apex. Plectrum triangular with rounded apex, wider than long. Aedeagus. Ventral fossae on the apices of the parameres completely bordered, long and narrow (Fig. 3G); apices of parameres right angled in lateral view, its widened part (in lateral view) about two thirds the length of paramere (Fig. 3D). Female Female (Fig. 3C) differs from male in having a relatively smaller, convex pronotum without tubercles and only with a small depression medially, frontoclypeus with much smaller, low keel not bimodal apically, protibial spur, and pygidium with rounded apex. Body length 21.0 mm. Variation Body length of examined male paratypes varies from 17.0 to 22.5 mm. Distribution The species is known from western Democratic Republic of the Congo (Fig. 6). The record from Nigeria is doubtful and need confirmation, therefore the specimen is not included in the type series., Published as part of Frolov, Andrey V. & Akhmetova, Lilia A., 2021, Taxonomic revision of the Afrotropical scarab beetle genus Cerhomalus Quedenfeldt, 1884 (Coleoptera: Scarabaeidae: Orphninae), pp. 36-50 in European Journal of Taxonomy 739 (1) on pages 42-43, DOI: 10.5852/ejt.2021.739.1265, http://zenodo.org/record/4604232

Cerhomalus Quedenfeldt, 1884 Figs 1���6 Cerhomalus Quedenfeldt, 1884: 289, pl. VIII, fig. 7. Cerhomalus ��� Karsch 1887: 5. ��� Quedenfeldt 1888: 165. ��� Arrow 1912: 30. ��� Schmidt 1913: 71. ��� Paulian, 1948: 11. Orphnus (Cerhomalus) Quedenfeldt ��� Petrovitz 1971: 2. Type species Cerhomalus mechowi Quedenfeldt, 1884, by monotypy. Diagnosis The largest orphnines (body length 15���22 mm). Colour uniformly dark brown to black. Clypeus of both sexes with a transverse keel, higher and apically bilobate in males (Fig. 1A) and lower but distinct in females (Fig. 1G). Pronotal disc of males more or less depressed, sometimes slightly excavated, but distinct pronotal lateral processes always absent. Elytra convex, without ridges, with humeral umbones. Elytral surface sparsely covered with rounded punctures; punctures arranged in longitudinal striae, which are more distinct on disc. Protibiae with 3 strong outer teeth similar in both sexes. Apical protibial spur absent in males. Middle and posterior legs similar in shape; posterior femora and tibiae about �� longer than middle ones. Parameres symmetrical, not setose apically, very wide in lateral view; inner sides of parameres with longitudinal excavations in 3 species (Fig. 1E). Ventroapical part of phallobase strongly sclerotized, in shape of 2 incompletely separated sclerites (Figs 1C, 5A, va. phlb. sc.). Species composition The genus comprises four species including two new ones described below. Distribution The genus occurs in Central and West Africa with the majority of records from the Congo Basin (Fig. 6). Key to species of Cerhomalus (males) 1. Apices of parameres rounded in apical view, without semi-circular excavations (Fig. 2G, J) and without keels or fossae on ventral side (Fig. 2K); base of elytra with somewhat tile-shaped punctures (Fig. 2 A���B).................................................. Cerhomalus absconditus (Petrovitz, 1971) comb. nov. ��� Apices of parameres with semi-circular excavations in apical view (Figs 1E, 3F, 4F, J) and with more or less developed fossae on ventral sides bordered with keel (Fig. 1F); base of elytra smooth (Figs 1A, G, I, 3A, C, 4A, C)............................................................................................................ 2 2. Ventral fossae on apices of parameres not bordered proximally (Fig. 4G, K, arrowed); apices of parameres in lateral view acute-angled (Fig. 4D, H)................... Cerhomalus quedenfeldti sp. nov. ��� Ventral fossae on apices of parameres completely bordered (Figs 1F, 3G); apices of parameres in lateral view right-angled (Figs 1C, 3D)............................................................................................ 3 3. Widened part of paramere (in lateral view) about half length of paramere (Fig. 1C); ventral fossa of paramere shorter and wider (Fig. 1F)............................... Cerhomalus mechowi Quedenfeldt, 1884 ��� Widened part of paramere (in lateral view) about two thirds length of paramere (Fig. 3D); ventral fossa of paramere longer and narrower (Fig. 3G).............................. Cerhomalus petrovitzi sp. nov., Published as part of Frolov, Andrey V. & Akhmetova, Lilia A., 2021, Taxonomic revision of the Afrotropical scarab beetle genus Cerhomalus Quedenfeldt, 1884 (Coleoptera: Scarabaeidae: Orphninae), pp. 36-50 in European Journal of Taxonomy 739 (1) on pages 37-38, DOI: 10.5852/ejt.2021.739.1265, http://zenodo.org/record/4604232, {"references":["Quedenfeldt G. 1884. Verzeichniss der von Herrn Major a. D. von Mechow in Angola und am Quango- Strom 1878 - 1881 gesammelten Pectinicornen und Lamellicornen. Berliner Entomologische Zeitschrift 28: 265 - 340. https: // doi. org / 10.1002 / mmnd. 18840280207","Karsch F. 1887. Altes und Neues uber Koleopteren. Berliner Entomologische Zeitschrift 31: 1 - 8. https: // doi. org / 10.1002 / mmnd. 18870310105","Quedenfeldt G. 1888. Beitrage zur Kenntniss der Koleopteren-Fauna von Central-Afrika nach den Ergebnissen der Lieutenant Wissman'schen Kassai-Expedition 1883 bis 1886. Berliner Entomologische Zeitschrift 32: 155 - 219.","Arrow G. J. 1912. Pachypodinae, Pleocominae, Aclopinae, Glaphyrinae, Ochodaeinae, Orphninae, Idiostominae, Hybosorinae, Dynamopinae, Acanthocerinae, Troginae. In: Junk W. & Schenkling S. (eds) Coleopterorum Catalogus 43: 1 - 66. W. Junk, Berlin.","Schmidt A. 1913. Coleoptera. Lamellicornia. Fam. Scarabaeidae. Subfam. Aegialinae, Chironinae, Dynamopinae, Hybosorinae, Idiostominae, Ochodaeinae, Orphninae. Genera Insectorum 150: 1 - 87. V. Verteneuil & L. Desmet, Brussels.","Paulian R. 1948. Revision des Orphnus africains (Coleoptera, Scarabaeidae). Annales de la Societe entomologique de France 117: 1 - 75.","Petrovitz R. 1971. Beitrag zur Kenntnis der Gattung Orphnus M'Leay (Orphninae, Scarabaeidae, Coleoptera). Revue de zoologie et de botanique africaines 84: 1 - 46."]}

Cerhomalus quedenfeldti sp. nov. urn:lsid:zoobank.org:act: AC2F5EC0-10F3-4262-A85E-F06B7B699139 Figs 4, 5D, 6 Differential diagnosis Cerhomalus quedenfeldti sp. nov. is similar to C. mechowi and C. petrovitzi sp. nov. in having the apices of the parameres with semi-circular excavations in apical view and with more or less developed fossae on ventral sides bordered with a keel, but can be separated from them in having the ventral fossae on the apices of the parameres not bordered proximally (Fig. 4 G–K, arrowed) and the apices of the parameres in lateral view acute-angled (Fig. 4D, H). Etymology The new species is named after Friedrich Otto Gustav Quedenfeldt (1817–1891), a German coleopterist. Type material Holotype REPUBLIC OF THE CONGO • ♂; “Congo Brazzaville O.R.S.T.O.M. XI-1965 R. Paulian ”; MNHN. Paratypes REPUBLIC OF THE CONGO • 1 ♂; “ Mai 1978 Voka Congo Coll. Th. Porion ”; MNHN; 6 ♀♀; “Congo Brazzaville O.R.S.T.O.M. XI-1965 R. Paulian”; MNHN • 1 ♀; “ CONGO BRAZZAVILLE XI-57 / ORSOM PARIS MOUCHET 1957”; MNHN • 1 ♀; “ Soil-Zoological Exp. Congo-Brazzaville Brazzaville ORSTOM / 18.10.1963 No 3 singled in park leg. Endrody-Younga”; HNHM • 1 ♂; “ Soil-Zoological Exp. Congo-Brazzaville Brazzaville ORSTOM park / 16.1.1964 No 695 soil trap in forest leg. Balogh & Zicsi ”; HNHM • 1 ♀; “ Soil-Zoological Exp. Congo-Brazzaville Brazzaville ORSTOM park / 22.12.1963 No 492 soil trap leg. Balogh & Zicsi ”; MHNG • 2 ♂♂, 3 ♀♀; “ Brazzaville Congo XI-1963 / MUSEUM PARIS Mission A. Descarpentiers et A. Villiers 1963-1964”; MNHN • 1 ♂; “ Soil-Zoological Exp. Congo-Brazzaville Kindamba, Kimboukou river / 28.10.1963. No 38 singled on river side leg. Endrody-Younga ”; MHNG. Description Male, holotype (Fig. 4 A–B, D–G) Body length 19.8 mm. Colour uniformly brown to black. Anterior margin of frontoclypeus somewhat rectangular, slightly convex in middle, bordered and slightly serrate in dorsal view (Fig. 4A). Frontoclypeus with a keel-shaped transverse process near anterior margin; height of keel about ¾ its width. Surface of frontoclypeus almost smooth, with minute punctures. Eye tubercles feebly developed. Eyes rather large: width about 1/6 distance between eyes in dorsal view. Antennae 10-segmented. Pronotum widely rounded laterally, as wide as elytra. Anterior border wide. Basal border narrow, keelshaped, separated from pronotal disc by deep groove with row of longitudinally elongated punctures. Pronotal disc depressed, somewhat rugose anteriorly. Most of pronotum surface covered with minute punctures separated by more than 3 puncture diameters. Anterolateral angles with much larger and denser punctures; posteriolateral angles with same punctures but in smaller numbers. Scutellum subtriangular, narrowly rounded apically, about 1/10 length of elytra. Elytra 1.1 times longer than wide, with distinct humeral humps. Elytra widest in middle, lateral margins almost parallel in basal half. First (sutural) stria distinct, as groove with row of punctures. Other stria before humeral humps as rows of round setiferous punctures. Striae laterad of stria 5 feebly distinct. Elytral intervals covered with minute punctures, somewhat sparser than those on pronotum. Macropterous. Abdominal sternite 8 medially longer than sternites 6 and 7 combined; sternite 6 about as long as sternite 7. Pygidium invisible from above, with slightly truncate apex. Plectrum triangular with rounded apex, wider than long. Aedeagus. Apices of parameres acute angled in lateral view (Figs 4D, H, 5D), with semi-circular excavations in apical view (Fig. 4F, J) and with more or less developed fossae on ventral sides not bordered proximally (Fig. 4G, K). Female Female (Fig. 4C) differs from male in having a relatively smaller, convex pronotum without tubercles, frontoclypeus with much smaller, low keel not bimodal apically, protibial spur, and pygidium with rounded apex. Variation Body length of examined male paratypes varies from 18.5 to 22.5 mm (males) and from 16.0 to 19.0 mm (females). Distribution The species is known from a few localities in the Republic of the Congo (Fig. 6).

Cerhomalus quedenfeldti sp. nov. urn:lsid:zoobank.org:act: AC2F5EC0-10F3-4262-A85E-F06B7B699139 Figs 4, 5D, 6 Differential diagnosis Cerhomalus quedenfeldti sp. nov. is similar to C. mechowi and C. petrovitzi sp. nov. in having the apices of the parameres with semi-circular excavations in apical view and with more or less developed fossae on ventral sides bordered with a keel, but can be separated from them in having the ventral fossae on the apices of the parameres not bordered proximally (Fig. 4 G���K, arrowed) and the apices of the parameres in lateral view acute-angled (Fig. 4D, H). Etymology The new species is named after Friedrich Otto Gustav Quedenfeldt (1817���1891), a German coleopterist. Type material Holotype REPUBLIC OF THE CONGO ��� ♂; ���Congo Brazzaville O.R.S.T.O.M. XI-1965 R. Paulian ���; MNHN. Paratypes REPUBLIC OF THE CONGO ��� 1 ♂; ��� Mai 1978 Voka Congo Coll. Th. Porion ���; MNHN; 6 ♀♀; ���Congo Brazzaville O.R.S.T.O.M. XI-1965 R. Paulian���; MNHN ��� 1 ♀; ��� CONGO BRAZZAVILLE XI-57 / ORSOM PARIS MOUCHET 1957���; MNHN ��� 1 ♀; ��� Soil-Zoological Exp. Congo-Brazzaville Brazzaville ORSTOM / 18.10.1963 No 3 singled in park leg. Endrody-Younga���; HNHM ��� 1 ♂; ��� Soil-Zoological Exp. Congo-Brazzaville Brazzaville ORSTOM park / 16.1.1964 No 695 soil trap in forest leg. Balogh & Zicsi ���; HNHM ��� 1 ♀; ��� Soil-Zoological Exp. Congo-Brazzaville Brazzaville ORSTOM park / 22.12.1963 No 492 soil trap leg. Balogh & Zicsi ���; MHNG ��� 2 ♂♂, 3 ♀♀; ��� Brazzaville Congo XI-1963 / MUSEUM PARIS Mission A. Descarpentiers et A. Villiers 1963-1964���; MNHN ��� 1 ♂; ��� Soil-Zoological Exp. Congo-Brazzaville Kindamba, Kimboukou river / 28.10.1963. No 38 singled on river side leg. Endrody-Younga ���; MHNG. Description Male, holotype (Fig. 4 A���B, D���G) Body length 19.8 mm. Colour uniformly brown to black. Anterior margin of frontoclypeus somewhat rectangular, slightly convex in middle, bordered and slightly serrate in dorsal view (Fig. 4A). Frontoclypeus with a keel-shaped transverse process near anterior margin; height of keel about �� its width. Surface of frontoclypeus almost smooth, with minute punctures. Eye tubercles feebly developed. Eyes rather large: width about 1/6 distance between eyes in dorsal view. Antennae 10-segmented. Pronotum widely rounded laterally, as wide as elytra. Anterior border wide. Basal border narrow, keelshaped, separated from pronotal disc by deep groove with row of longitudinally elongated punctures. Pronotal disc depressed, somewhat rugose anteriorly. Most of pronotum surface covered with minute punctures separated by more than 3 puncture diameters. Anterolateral angles with much larger and denser punctures; posteriolateral angles with same punctures but in smaller numbers. Scutellum subtriangular, narrowly rounded apically, about 1/10 length of elytra. Elytra 1.1 times longer than wide, with distinct humeral humps. Elytra widest in middle, lateral margins almost parallel in basal half. First (sutural) stria distinct, as groove with row of punctures. Other stria before humeral humps as rows of round setiferous punctures. Striae laterad of stria 5 feebly distinct. Elytral intervals covered with minute punctures, somewhat sparser than those on pronotum. Macropterous. Abdominal sternite 8 medially longer than sternites 6 and 7 combined; sternite 6 about as long as sternite 7. Pygidium invisible from above, with slightly truncate apex. Plectrum triangular with rounded apex, wider than long. Aedeagus. Apices of parameres acute angled in lateral view (Figs 4D, H, 5D), with semi-circular excavations in apical view (Fig. 4F, J) and with more or less developed fossae on ventral sides not bordered proximally (Fig. 4G, K). Female Female (Fig. 4C) differs from male in having a relatively smaller, convex pronotum without tubercles, frontoclypeus with much smaller, low keel not bimodal apically, protibial spur, and pygidium with rounded apex. Variation Body length of examined male paratypes varies from 18.5 to 22.5 mm (males) and from 16.0 to 19.0 mm (females). Distribution The species is known from a few localities in the Republic of the Congo (Fig. 6)., Published as part of Frolov, Andrey V. & Akhmetova, Lilia A., 2021, Taxonomic revision of the Afrotropical scarab beetle genus Cerhomalus Quedenfeldt, 1884 (Coleoptera: Scarabaeidae: Orphninae), pp. 36-50 in European Journal of Taxonomy 739 (1) on pages 45-47, DOI: 10.5852/ejt.2021.739.1265, http://zenodo.org/record/4604232

Cerhomalus absconditus (Petrovitz, 1971) comb. nov. Figs 2, 5A, 6 Orphnus (Cerhomalus) absconditus Petrovitz, 1971: 3. Differential diagnosis Cerhomalus absconditus can be easily separated from congeners by having the apices of the parameres rounded (in apical view), without semi-circular excavations (Fig. 2G, J) and without keels or fossae on ventral side (Fig. 2K), and the base of elytra with somewhat tile-shaped punctures (Fig. 2 A–B). Material examined Holotype DEMOCRATIC REPUBLIC OF THE CONGO • ♂; “COLL. MUS. CONGO Tshuapa: Yokamba /1953 J. Stevenard / HOLOTYPUS / O. (Cerhomalus) absconditus n.sp. Petrovitz”; RMCA. Paratypes DEMOCRATIC REPUBLIC OF THE CONGO • 1 ♀ “COLL. MUS. CONGO Kasai: Ipamu -1937 R. Soeur Imelda / PARATYPUS / O. (Cerhomalus) absconditus n.sp. Petrovitz”; RMCA • 5 ♀♀; “COLL. MUS. CONGO Kwango: Djuma 1953 /R.P.G. Leta/ PARATYPUS / O. (Cerhomalus) absconditus n.sp. Petrovitz / coll. Petrovitz ”; RMCA • 1 ♂; “COLL. MUS. CONGO Lulua: Kapanga – XI-1933 F.G. Overlaet / PARATYPUS / O. (Cerhomalus) absconditus n.sp. Petrovitz”; RMCA • 1 ♂; “MUS. ROY. AFR. CENTR. Kasai: terr. Dekese I.1960 F. Francois / PARATYPUS / O. (Cerhomalus) absconditus n.sp. Petrovitz”; RMCA • 1 ♀; “MUSEE DU CONGO Kasai: Don de la Cie du Kasai / PARATYPUS / R. DET. 1939 F / O. (Cerhomalus) absconditus n.sp. Petrovitz”; RMCA • 1 ♂; “MUSEE DU CONGO Kasai: Ipamu - VIII -1922 P. Vanderijst / PARATYPUS / R. DET. 1939 F / O. (Cerhomalus) absconditus n.sp. Petrovitz”; RMCA • 2 ♂♂; “Musee du Congo Kinda (Katanga) 1927 (Don M. Muller.) / Kinda Katanga 1926 Congo / O. (Cerhomalus) absconditus n.sp. Petrovitz / PARATYPUS / R. DET. 1939 F / 139”; MHNG • 1 ♀; “MUSEE DU CONGO Murlambongo (Idiofa) - 1930 (Les Soeurs de la Mission) / PARATYPUS / O. (Cerhomalus) absconditus n.sp. Petrovitz”; RMCA • 1 ♂; “Musee du Congo Sankuru: Komi (Lodja) 27-I-1930 J. Ghesquiere / R. DET 1939 F / Cerhomalus mechowi Qued. / O. (Cerhomalus) absconditus n.sp. Petrovitz / PARATYPUS ”; RMCA • 1 ♂; “COLL. MUS. CONGO Sankuru: Katako-Kombe VIII - 1952 Dr M. Fontaine / O. (Cerhomalus) absconditus n.sp. Petrovitz / PARATYPUS / PARATYPUS”; MCSNG • 1 ♂, 1 ♀; “COLL. MUS. CONGO Tshuapa: Ikela 1956 R. Deguide / PARATYPUS / O. (Cerhomalus) absconditus n. sp. Petrovitz”; RMCA • 1 ♀; “COLL. MUS. CONGO Tshuapa: Ikela 1956 R.P. Lootens / PARATYPUS / O. (Cerhomalus) absconditus n. sp. Petrovitz”; RMCA • 1 ♀; “COLL. MUS. CONGO Tshuapa: Bosekele VI-1952 J. Van Vynckt / O. (Cerhomalus) absconditus n.sp. Petrovitz / PARATYPUS ”; RMCA • 1 ♀; “COLL. MUS. CONGO Tshuapa: Yokamba /1953 J. Stevenard / ALLOTYPUS / O. (Cerhomalus) absconditus n.sp. Petrovitz”; RMCA • 1 ♂; “COLL. MUS. CONGO Terr. de Dibaya Kamponde 1945 Rev. Fr. ALLAER 1945 / PARATYPUS / O. (Cerhomalus) absconditus n. sp. Petrovitz”; RMCA • 1 ♀; “COLL. MUS. CONGO Katoka-Luluabourg / -1938 R. P. VANKERCKHOVEN / PARATYPUS / O. (Cerhomalus) absconditus n.sp. Petrovitz / coll. Petrovitz ”; RMCA • 1 ♂; “MUSEE DU CONGO Luebo - III-1931. 1 ex. J.P. Colin / O. (Cerhomalus) absconditus n.sp. Petrovitz / PARATYPUS”; RMCA. Other material DEMOCRATIC REPUBLIC OF THE CONGO • 1 ♀; Équateur, Tshuapa, Etata; Aug.–Sep. 1969; J. Hauwaert leg.; RMCA. ANGOLA • 1 ♂, 3 ♀♀; Bié, Catabola; 15–27 Sep. 2012; T. Lackner leg.; BMNH • 4 ♀♀; Lunda Norte, Andrada; 7°41′ S, 21°17′ E; Aug.–Sep. 1952; M. Fetchkowsky leg.; RMCA • 2 ♀♀; Dundo; Sep. 1948; Machado leg.; RMCA • 1 ♂; Cuando Cubango Province, N of Menongue City; 1600 m a.s.l.; 22 Nov. 2019; S. Dementev leg.; SDCMR. CAMEROON • 1 ♂; 1912; BMNH. Distribution This is the most widely distributed species of the genus. The majority of records are from the Congo basin, but the species occurs up to Cameroon in the north (no precise locality is recorded) and to the Ancient Plateau in Angola in the south (Fig. 6).

Aegidium simplex Frolov & Akhmetova, new species Figs. 1���9. Type material. Holotype, male: ��� Columbia Dr. Will. ��� (ZMHUB). Paratype, female with the same data as for holotype (ZMHUB). Description. Holotype, male (Fig. 1, 3, 5���9). Body length 10.9 mm, width of elytra 5.2 mm, width of pronotum 4.8 mm. Upper side of body shiny, color uniform brown, head and pronotum slightly darker. Clypeus symmetrical with slightly convex anterior margin and rounded anterior angles. Head surface densely punctate with round punctures separated by less than a puncture diameter on clypeus, becoming denser and coarser on frons near eyes. Head without traces of medial horn or tubercle. Mandibles symmetrical, protruding past anterior margin of clypeus. Labrum rounded, small, slightly protruding past clypeus (in dorsal view). Pronotum 1.5 times wider than long, widest at middle. Anterior margin with a border interrupted medially by a tubercle. Lateral margins crenulate. Base bordered. Disc of pronotum slightly depressed in middle. Surface densely punctate, punctures separated by about a diameter on disc, becoming denser towards sides. Scutellum elongate, rounded apically, about 1/12 length of elytra, smooth. Elytra moderately convex on disc with maximum width approximately at middle. Humeral and apical humps distinct. Elytra with 1 feebly distinct longitudinal ridge on disc between suture and humeral hump. First (sutural) elytral interval almost not elevated. Striae indistinct. Elytra densely punctate with coarse, irregularly shaped punctures not arranged in distinct longitudinal rows on disc. Base of elytra feebly bordered. Wings fully developed. Protibiae with 3 outer teeth and a smaller inner tooth. Lateral margin basad of outer teeth slightly crenulate. Apex and internal margin of tibia with a few slender setae. Protarsi about 1/2 length of protibiae. Claws 1/3 length of tarsomere 5, which is somewhat longer than tarsomeres 3 and 4 combined, and somewhat thicker than other tarsomeres. Tarsomere 1 as long as tarsomeres 2���4 combined. Ventral surface of femora punctate with rounded punctures. Mesothoracic legs and metathoracic legs similar in shape; metafemora and metatibiae about 1.2 times longer than mesofemora and mesotibiae. Femora sparsely punctate with rounded punctures. Tibiae somewhat triangular, with two apical spurs. Mesotibiae without tuft of setae ventroapically (Fig. 3). Upper tibial spur shorter than tarsomeres 1���3 combined; lower spur shorter than tarsomeres 1���2 combined. Claws 1/3 length of tarsomere 5, which is relatively slender, as long as tarsomere 2 and twice as short as tarsomere 1. Abdominal sternites punctate with coarse irregularly shaped punctures. Sternite 8 medially about 2 times wider than sternites 4���7. Pygidium triangular, convex, partly hidden under elytra, irregularly punctate with transverse punctures. Aedeagus. Phallobase 2.1 times longer than parameres, tube shaped (Fig. 5). Parameres reasonably wider than apical part of phallobase in dorsal view (Fig. 6). Ventrobasal transverse plate of parameres absent (Fig. 7). Endophallus without symmetrical strongly sclerotized sclerites (Fig. 9). Cranial part of spiculum gastrale narrow, tapering and rounded at very apex (Fig. 8). Female. Female (Fig. 2) differs from male in having a distinct protibial spur and relatively smaller pronotum without depression and anterior tubercle. Body length 12.0 mm, width of elytra 5.5 mm, width of pronotum 4.7 mm. Differential diagnosis. In the key to the South American mainland Aegidium species (Frolov et al. 2017b), the new species comes out in the couplet with A. reichei Preudhomme de Borre, A. bolivianum Frolov & Akhmetova, and A. varians Frolov, Akhmetova & Vaz-de-Mello. All these species are similar in having the mesotibiae without a tuft of setae ventroapically and in the lack of ventrobasal transverse plate of parameres. Aegidium simplex can be separated from these species by the parameres being wider in lateral view and the length/width ratio is approximately 2.0 (versus being narrower in lateral view and the length/width ratio being 3.5���4.0). Etymology. From Latin, ��� simplex ��� for an ordinary appearance of the beetles., Published as part of Frolov, Andrey V. & Akhmetova, Lilia A., 2021, A new species of Aegidium Arrow (Coleoptera: Scarabaeidae: Orphninae) from Colombia, pp. 583-586 in Zootaxa 4927 (4) on pages 583-585, DOI: 10.11646/zootaxa.4927.4.9, http://zenodo.org/record/4543187, {"references":["Frolov, A. V., Akhmetova, L. A. & Vaz-de-Mello, F. Z. (2017 b) Revision of the mainland species of the Neotropical genus Aegidium Westwood (Coleoptera: Scarabaeidae: Orphninae). Journal of Natural History, 51, 1035 - 1090. https: // doi. org / 10.1080 / 00222933.2017.1319519"]}

Genus Madecorphnus Paulian, 1992 Madecorphnus Paulian, 1992: 171; Frolov 2010a, 2010c. Drepanognathus Lansberge, 1886: 92 (in part: Paulian 1977; synonym of Orphnus MacLeay: Frolov 2005). Sissantobius Ritsema, 1888: 217 (in part: Paulian 1937, 1977; synonym of Orphnus MacLeay: Frolov 2005). Type species. Drepanognathus falciger Lansberge, 1886, designated by Paulian (1977). Diagnosis. Madecorphnus are small-sized (4.5–7.0 mm) beetles with mostly uniform, brown to black coloration. Upper side of the body has a few setae with distinct locations: 1 seta on the elytral base near epipleuron, 1 seta on the posterior angle of the pronotum, 1 seta approximately in the middle of the lateral margin of the pronotum, 2 setae on the anterior angle of the pronotum, 2 setae on the lateral sides and 2 setae on the anterior margin of the frontoclypeus. Mandibles asymmetrical, subequal in length in females and subequal to strongly unequal in males. Frontoclypeus symmetrical to asymmetrical in males, wide, bordered anteriorly, smooth, without tubercles or ridges. Pronotum wider than long, smooth, similar in both sexes, without any depressions, tubercles, or ridges. Propleurae smooth, convex, without carinae separating anterolateral areas from basal area. Scutellum triangular, narrowly rounded apically, about 1/10 the length of the elytra. Elytra convex, with feebly marked humeral humps, smooth, with only first stria distinct. Wings fully developed. Metepisternon triangular, tapering caudally, with slightly widened anterodorsal angle slightly overlapping epypleuron. Protibiae with 3 outer teeth. Phallobase strongly sclerotized dorsally and with a thin membrane ventrally. Parameres relatively long, their apices tapering or curved downwards, without setae. Internal sac of aedeagus has species-specific armature. Sexual dimorphism in Madecorphnus is weaker than in the other Orphninae taxa. The sexes mostly differ in the shape of the mandibles. Males with the relatively feebly-developed mandibles are similar to females. In Madecorphnus, the sexual dimorphism character common for all Orphninae genera, the absence of distinct protibial spur in males, is less expressed. Often, the apical seta is longer and more robust than the others and is similar to the spur (Fig. 10 H). However, examination of this seta at higher magnification shows that it bears minute, feebly visible setae while the true spurs are always smooth. Madecorphnus males also lack any frontoclypeal horns and prothoracic ridges, tubercles or excavations, which are found in the majority of species of almost all genera of the Orphninae. Diagnostic characters. The shape of the mandibles is species specific at least in some species, but it is subject to allometric variability and therefore cannot always be a reliable diagnostic character. The most useful characters to separate species proved to be the number and shape of the internal sac sclerites and the shape of the parameres. Species composition, distribution, and habitat. Madecorphnus is endemic to Madagascar and the most speciose genus of the Madagascan Orphninae with 19 described species. The species are distributed throughout the island but mostly in its eastern part. The notable exception is M. falculoides, which was found in the western part. The majority of the localities agree well with the current distribution of the remnants of indigenous forests. Little is known about bionomy of Madecorphnus. Some specimens were collected in carrion baited pitfall traps. However, it is unknown if they were attracted to carrion or captured accidently. A few specimens were collected in flight intercept traps or sifted out of forest litter. Madecorphnus species are probably generalist saprophagous litter dwellers but the labels of most of the collected specimens lack any data about the way the beetles were collected. Nesting behavior and preimaginal stages are so far unknown. Remarks. The first known species of the genus Madecorphnus and its type species, M. falciger (Lansberge, 1886), was originally described in the genus Drepanognathus Lansberge, 1886, along with D. mandibularis Lansberge, 1886 (currently Orphnus mandibularis). Since the generic name Drepanognathus was preoccupied, the replacement name Sissantobius Ritsema, 1888 was used for the two species. The name Madecorphnus was proposed by Paulian (1977) to accommodate M. falciger and S. falculoides Paulian from Madagascar and to separate these two species from African Sissantobius mandibularis (sensu Paulian), the type species of the genus Sissantobius. Synonymy of the names Drepanognathus, Sissantobius, and Madecorphnus is discussed in more detail in Paulian (1992) and Frolov (2005). Paulian (1977: 1204) illustrated the peculiar aedeagus and evaginated internal sac of M. falciger (sensu Paulian), but these illustrations do not agree with the aedeagus of the type of M. falciger. The shape of the parameres is similar to that of M. niger Frolov, 2010, but the internal sac with numerous short spinules having wide round bases is unknown in any described Madecorphnus species. We failed to find this specimen, apparently belonging to an undescribed species, in the collection of MNHN., Published as part of Frolov, Andrey V., Montreuil, Olivier & Akhmetova, Lilia A., 2016, Review of the Madagascan Orphninae (Coleoptera: Scarabaeidae) with a revision of the genus Triodontus Westwood, pp. 1-93 in Zootaxa 4207 (1) on pages 22-23, DOI: 10.11646/zootaxa.4207.1.1, http://zenodo.org/record/200444, {"references":["Paulian, R. (1992) Un nouveau genre et une nouvelle espece d'Orphnidae de Madagascar (Coleoptera, Scarabaeoidea). Revue Francaise d'Entomologie (N. S.), 14, 169 - 171.","Frolov, A. V. (2010 a) A new species of scarab beetle, Madecorphnus cuccodoroi (Coleoptera: Scarabaeidae: Orphninae), from northern Madagascar. Zootaxa, 2531, 64 - 68.","Frolov, A. V. (2010 c) Revision of the Madagascan genus Madecorphnus Paulian (Coleoptera, Scarabaeidae, Orphninae). Journal of Natural History, 44, 1095 - 1111.","Lansberge, J. W. van (1886) Scarabaeides, Bupretides et Cerambycides de l'Afrique occidentale envoyes au Musee de Leyden par M. M. Veth et van der Kellen. Notes from the Leyden Museum, 8, 69 - 120.","Paulian, R. (1977) Les Orphnidae de Madagascar. Bulletin du Museum National d'histoire naturelle, Paris. Series 3, 411, 1199 - 1223.","Frolov, A. V. (2005) Taxonomic position of Sissantobius mandibularis (Lansberge) and synonymy of Orphnus over Sissantobius (Coleoptera, Scarabaeidae, Orphninae). Journal of Afrotropical Zoology, 2, 69 - 72.","Ritsema, C. (1888) Lijst der Entomologische Geschriften van Mr J. W. van Lansberge. Tijdschrift voor Entomologie, 31, 201 - 34.","Paulian, R. (1937) Faune entomologique de Madagascar. Coleoptera Lamellicornia, Scarabaeidae Acanthocerini, Trogini, Aulonocnemis, Hybosorini, Orphnini et Ochodaeini. Bulletin de l'Academie malgache (nouvelle serie), 19, 129 - 143."]}

Orphnus (Orphnus) brevialatus Frolov & Akhmetova, new species (Figs. 1 A–D, 2A–D) Type material. Holotype, male with the label “ TANZANIA 16.XII.2006 Chunya to Mbeya, 2500 m Werner leg.” (ZIN). Paratype, female with the same data as the holotype (ZIN). Description. Holotype, male (Figs. 1 A–C, 2A–C). Body elongate, convex, shiny; length 11.8 mm, maximum width 5.8 mm. Color uniformly dark brown. Head: Clypeus wide, with convex anterior margin, rounded laterally, finely crenulate. Genae small, not protruding past eyes. Frontal suture indistinct. Clypeus with long, slender horn, slightly curved posteriad (Figs. 1B, C). Dorsal surface of head almost impunctate. Labrum deeply sinuate in the middle, distinctly protruding past clypeus. Pronotum: with rounded sides, about 1.2 times wider than long, with deeply concave disc and large lateral ridges almost triangular in lateral view (Fig 1C). Anterior angles acute; posterior angles rounded, indistinct in dorsal view. Pronotum bordered on anterior margin and base. Base of pronotum rugose. Sides with coarse, rounded punctures separated by about 0.5–1.0 puncture diameters. Lateral margins with long, sparse, brown setae. Scutellum: wide, semicircular, about 1/16 the length of elytra. Elytra: convex, about as wide as long, widest in the middle, with almost indistinct humeral umbones. Striae indistinct. Elytra with coarse, ocellate punctation; punctures denser near base and sparser on disc and in apical part. Wings: reduced, about 1/3 length of elytra. Legs: Protibiae with 3 outer teeth. Lateral margin basad of outer teeth not crenulate. Apical spur of protibia absent. Protarsi of the holotype broken. Middle legs and hind legs similar in shape; metafemora and metatibiae about 1/8 longer than the mesofemora and mesotibiae. Meso- and metatibiae somewhat triangular with 2 apical spurs, inner margin almost straight, with 1 transverse keel. Upper spur of tibiae as long as two basal tarsomeres. Claws 1/3 length of apical tarsomere. Femora almost impunctate. Abdomen: Abdominal sternites irregularly punctate, pubescent, with sparse, long setae. Visible sternite 6 medially about 2 times longer than sternites 2–5 combined. Pygidium: Surface irregularly punctate with transverse punctures. Aedeagus: with relatively long (0.6 length of phallobase), curved downwards parameres tapering apically (Fig. 2B). Parameres spear-shaped in dorsal view, with angulate lateral teeth. Endophallus with 2 groups of microspinules and a cluster of larger spinules (Fig. 2C). Female (Fig. 1D) differs from the male in having a relatively smaller pronotum without armature, frontoclypeus without process, presence of prothoracic spur. Body length of the female paratype 10.5 mm. Diagnosis. From the other flightless Orphnus species, O. brevialatus new species differs in having a welldeveloped pronotal armature with two long, acute lateral processes. In the shape of the head, pronotum, and male genitalia it is similar to O. compactilis Quedenfeldt, 1884 but differs from it, in addition to the reduced wings and shorter body, in the shape of the endophallic armature. Distribution. The new species is known from a single locality in Kipengere Range (Livingstone Mountains), the eastern escarpment of the East African Rift in East Africa. The label does not contain exact locality but suggests that the specimens were collected on the road between towns of Chunya and Mbeya (Fig. 2D). Etymology. From Latin adjectives in the nominative singular brevis (= short), and alatus (= winged), referring to the reduced wings.

Orphnus (Orphnus) brevialatus Frolov & Akhmetova, new species (Figs. 1 A���D, 2A���D) Type material. Holotype, male with the label ��� TANZANIA 16.XII.2006 Chunya to Mbeya, 2500 m Werner leg.��� (ZIN). Paratype, female with the same data as the holotype (ZIN). Description. Holotype, male (Figs. 1 A���C, 2A���C). Body elongate, convex, shiny; length 11.8 mm, maximum width 5.8 mm. Color uniformly dark brown. Head: Clypeus wide, with convex anterior margin, rounded laterally, finely crenulate. Genae small, not protruding past eyes. Frontal suture indistinct. Clypeus with long, slender horn, slightly curved posteriad (Figs. 1B, C). Dorsal surface of head almost impunctate. Labrum deeply sinuate in the middle, distinctly protruding past clypeus. Pronotum: with rounded sides, about 1.2 times wider than long, with deeply concave disc and large lateral ridges almost triangular in lateral view (Fig 1C). Anterior angles acute; posterior angles rounded, indistinct in dorsal view. Pronotum bordered on anterior margin and base. Base of pronotum rugose. Sides with coarse, rounded punctures separated by about 0.5���1.0 puncture diameters. Lateral margins with long, sparse, brown setae. Scutellum: wide, semicircular, about 1/16 the length of elytra. Elytra: convex, about as wide as long, widest in the middle, with almost indistinct humeral umbones. Striae indistinct. Elytra with coarse, ocellate punctation; punctures denser near base and sparser on disc and in apical part. Wings: reduced, about 1/3 length of elytra. Legs: Protibiae with 3 outer teeth. Lateral margin basad of outer teeth not crenulate. Apical spur of protibia absent. Protarsi of the holotype broken. Middle legs and hind legs similar in shape; metafemora and metatibiae about 1/8 longer than the mesofemora and mesotibiae. Meso- and metatibiae somewhat triangular with 2 apical spurs, inner margin almost straight, with 1 transverse keel. Upper spur of tibiae as long as two basal tarsomeres. Claws 1/3 length of apical tarsomere. Femora almost impunctate. Abdomen: Abdominal sternites irregularly punctate, pubescent, with sparse, long setae. Visible sternite 6 medially about 2 times longer than sternites 2���5 combined. Pygidium: Surface irregularly punctate with transverse punctures. Aedeagus: with relatively long (0.6 length of phallobase), curved downwards parameres tapering apically (Fig. 2B). Parameres spear-shaped in dorsal view, with angulate lateral teeth. Endophallus with 2 groups of microspinules and a cluster of larger spinules (Fig. 2C). Female (Fig. 1D) differs from the male in having a relatively smaller pronotum without armature, frontoclypeus without process, presence of prothoracic spur. Body length of the female paratype 10.5 mm. Diagnosis. From the other flightless Orphnus species, O. brevialatus new species differs in having a welldeveloped pronotal armature with two long, acute lateral processes. In the shape of the head, pronotum, and male genitalia it is similar to O. compactilis Quedenfeldt, 1884 but differs from it, in addition to the reduced wings and shorter body, in the shape of the endophallic armature. Distribution. The new species is known from a single locality in Kipengere Range (Livingstone Mountains), the eastern escarpment of the East African Rift in East Africa. The label does not contain exact locality but suggests that the specimens were collected on the road between towns of Chunya and Mbeya (Fig. 2D). Etymology. From Latin adjectives in the nominative singular brevis (= short), and alatus (= winged), referring to the reduced wings., Published as part of Frolov, Andrey V. & Akhmetova, Lilia A., 2020, Description of a new brachypterous scarab species, Orphnus brevialatus (Coleoptera: Scarabaeidae: Orphninae) from East Africa, with notes on flightlessness in the orphnines, pp. 425-431 in Zootaxa 4750 (3) on page 426, DOI: 10.11646/zootaxa.4750.3.8, http://zenodo.org/record/3707541

Frolov, Andrey V., Akhmetova, Lilia A. (2020): Description of a new brachypterous scarab species, Orphnus brevialatus (Coleoptera: Scarabaeidae: Orphninae) from East Africa, with notes on flightlessness in the orphnines. Zootaxa 4750 (3): 425-431, DOI: https://doi.org/10.11646/zootaxa.4750.3.8

Genus Aphodius Helwig, 1798 Type species: Scarabaeus fimetarius Linnaeus, 1758, Published as part of Frolov, Andrey V., Akhmetova, Lilia A. & Kr��l, David, 2020, The taxonomic status of Aphodius insperatus Petrovitz, A. longipennis Rakovi�� and A. inylchekensis Frolov (Coleoptera: Scarabaeidae: Aphodiinae), pp. 390-394 in Zootaxa 4728 (3) on page 390, DOI: 10.11646/zootaxa.4728.3.9, http://zenodo.org/record/3626486

Hybalus punicus Baraud, 1991 Hybalus punicus Baraud, 1991: 22. The species is only known from the type series collected in Fondouk-Djedid (Fig. 2)., Published as part of Frolov, Andrey, Akhmetova, Lilia & Montreuil, Olivier, 2019, A review of Hybalus Dejean from Tunisia (Coleoptera: Scarabaeidae: Orphninae) with description of a new species, pp. 290-294 in Zootaxa 4671 (2) on page 293, DOI: 10.11646/zootaxa.4671.2.10, http://zenodo.org/record/3442338, {"references":["Baraud, J. (1991) Revision du genre Hybalus Brulle, 1834 (Coleoptera, Scarabaeoidea, Orphninae). Annales de la Societe Entomologique de France, New Series, 27, 3 - 28."]}

Frolov, Andrey, Akhmetova, Lilia, Montreuil, Olivier (2019): A review of Hybalus Dejean from Tunisia (Coleoptera: Scarabaeidae: Orphninae) with description of a new species. Zootaxa 4671 (2): 290-294, DOI: https://doi.org/10.11646/zootaxa.4671.2.10

[Hybalus bigibber Reitter, 1892] Hybalus bigibber Reitter, 1892e: 260; Normand 1936: 376; Baraud 1991: 14. This species was described from Sicily and recorded from Tunisia by Normand (1936) (Ghardimaou and Souk El Arba [Jendouba]), although the later author was unsure about the identification of the Hybalus specimens he studied. The record from Jendouba probably belongs to H. algiricus. We don���t know any specimen to confirm the presence of this species on Tunisian territory., Published as part of Frolov, Andrey, Akhmetova, Lilia & Montreuil, Olivier, 2019, A review of Hybalus Dejean from Tunisia (Coleoptera: Scarabaeidae: Orphninae) with description of a new species, pp. 290-294 in Zootaxa 4671 (2) on page 293, DOI: 10.11646/zootaxa.4671.2.10, http://zenodo.org/record/3442338, {"references":["Normand, H. (1936) Contribution au Catalogue des Coleopteres de la Tunisie. Fascicule 10. Bulletin de la Societe d'Histoire naturelle de l'Afrique du Nord, 27, 355 - 383.","Baraud, J. (1991) Revision du genre Hybalus Brulle, 1834 (Coleoptera, Scarabaeoidea, Orphninae). Annales de la Societe Entomologique de France, New Series, 27, 3 - 28."]}

Hybalus normandi Baraud, 1980 Hybalus normandi Baraud, 1980: 60; 1991: 20. This species was described from Tamera (Baraud 1980) and later recorded also from Cap Bon (Baraud 1991) (Fig. 2)., Published as part of Frolov, Andrey, Akhmetova, Lilia & Montreuil, Olivier, 2019, A review of Hybalus Dejean from Tunisia (Coleoptera: Scarabaeidae: Orphninae) with description of a new species, pp. 290-294 in Zootaxa 4671 (2) on page 293, DOI: 10.11646/zootaxa.4671.2.10, http://zenodo.org/record/3442338, {"references":["Baraud, J. (1980) Les Hybalus Brulle de Tunisie. Nouvelle Revue d'Entomologie, 10, 59 - 62.","Baraud, J. (1991) Revision du genre Hybalus Brulle, 1834 (Coleoptera, Scarabaeoidea, Orphninae). Annales de la Societe Entomologique de France, New Series, 27, 3 - 28."]}

[Hybalus parvicornis Lucas, 1855] Hybalus parvicornis Lucas, 1855: 556; Normand 1936: 376; Baraud 1991: 21. This species is known from a few localities in Algeria. It was recorded from Tunisia by Normand (1936) but he mentioned that the identification was doubtful. We don���t know any specimens to confirm the presence of this species on Tunisian territory., Published as part of Frolov, Andrey, Akhmetova, Lilia & Montreuil, Olivier, 2019, A review of Hybalus Dejean from Tunisia (Coleoptera: Scarabaeidae: Orphninae) with description of a new species, pp. 290-294 in Zootaxa 4671 (2) on page 293, DOI: 10.11646/zootaxa.4671.2.10, http://zenodo.org/record/3442338, {"references":["Normand, H. (1936) Contribution au Catalogue des Coleopteres de la Tunisie. Fascicule 10. Bulletin de la Societe d'Histoire naturelle de l'Afrique du Nord, 27, 355 - 383.","Baraud, J. (1991) Revision du genre Hybalus Brulle, 1834 (Coleoptera, Scarabaeoidea, Orphninae). Annales de la Societe Entomologique de France, New Series, 27, 3 - 28."]}

Hybalus algiricus Petrovitz, 1968 Hybalus algiricus Petrovitz, 1968: 6; Baraud, 1991: 11. Material examined. Tunisia: Jendouba, El Feija National Park, N36��27��48�� E08 14��11��, 590 m, 3.III.2012, W. Ziegler leg., 1 male (ZIN), 2 males (CWZ); Sillana Province, N Kesra, 8.III.2012, N35��51��20�� E9��23��41��, 1000m, W. Ziegler leg., 1 male (CWZ); Beja, 11 km of Teboursouk Pass, N36��29��48�� E09��09��12��, 1.III.2012, Meybohm leg., 1 male (CWZ); 2 males MNHN La Kesra Tunisie De Vauloger. This species was described from Algeria (without more precise locality) and known from the single type specimen. It is recorded from Tunisia for the first time (Fig. 2)., Published as part of Frolov, Andrey, Akhmetova, Lilia & Montreuil, Olivier, 2019, A review of Hybalus Dejean from Tunisia (Coleoptera: Scarabaeidae: Orphninae) with description of a new species, pp. 290-294 in Zootaxa 4671 (2) on page 293, DOI: 10.11646/zootaxa.4671.2.10, http://zenodo.org/record/3442338, {"references":["Baraud, J. (1991) Revision du genre Hybalus Brulle, 1834 (Coleoptera, Scarabaeoidea, Orphninae). Annales de la Societe Entomologique de France, New Series, 27, 3 - 28."]}

Frolov, Andrey V., Akhmetova, Lilia A., Vaz-de-Mello, Fernando Z. (2019): Contribution to the knowledge of the Neotropical canopy dwelling genus Onorius (Coleopteraı Scarabaeidaeı Orphninae): another species from southern Ecuador and notes on the hind gut morphology in the Orphninae. Journal of Natural History 53 (17): 1071-1077, DOI: 10.1080/00222933.2019.1632952

Onorius sp. Material examined Female. ‘ Zamora Chinchipe Paquisha Alto Marzo 2008 Col. V. Carvajal ’ (CEMT). Diagnosis This species differs from O. inexpectatus in the sculpture of the body (Figure 1a): head is punctate with transversely elongate, sometimes adjoining punctures (punctures are round and relatively sparse in O. inexpectatus); pronotum with coarser ocellate punctures (punctures are smaller and not ocellate in O. inexpectatus); elytra with characteristic U- to V-shaped punctures each bearing a distinct yellowish seta posteriorly (punctures are round in O. inexpectatus; setae in the punctures are smaller, barely visible at the same magnification); proximal ileum has two large, symmetrical diverticula (Figure 2b). Description Female (Figures 1 and 2). Body length 9.0 mm, width of elytra 4.6 mm, width of pronotum 3.5 mm. Colour uniform brown with pronounced bronze tint. Head densely punctate with transversely elongate punctures; some punctures fused near anterior margin of clypeus. Head without traces of medial horn or tubercle. Left mandible slightly protruding past clypeus in dorsal view; labrum not protruding past clypeus. Clypeus rounded anteriorly. Pronotum 1.5 times wider than long, widest at basal 0.3 of its length. Anterior margin without border, basal and lateral margins bordered. Disc with shallow impression anteriorly and near base, without tubercles or deep fossae. Surface punctate with round ocellate punctures separated by 1 – 3 puncture diameters. Scutellum triangular, narrowly rounded apically, about 1/15 length of elytra, coarsely punctate. Elytra convex, with marked humeral and apical humps, without ridges or other elevated areas.Maximum width approximately at the middle. Striae indistinct. Elytra punctate with U- or V-shaped punctures; each puncture bears a short, yellow seta posteriorly. Base of elytra not bordered. Wings fully developed. Protibiae with 3 outer teeth, without smaller inner tooth. Lateral margin basad of outer teeth not crenulate. Apex and internal margin of tibia with a few slender setae. Protarsi well developed, about 2/3 length of protibiae. Claws 1/3 length of tarsomere 5. Tarsomere 5 as long as tarsomeres 2 – 4 combined. Tarsomere 1 as long as tarsomeres 2 – 4. Hind and middle legs similar in shape; metafemora and metatibiae about 1.2 times longer than mesofemora and mesotibiae. Femora sparsely punctate with elongate punctures. Tibiae somewhat triangular, with 2 apical spurs. Upper tibial spur as long as tarsomeres 1 – 3; lower spur as long as or slightly longer than tarsomere 1. Claws 1/3 length of tarsomere 5. Tarsomere 5 relatively robust, as wide as the other tarsomeres, as long as tarsomeres 2 – 4 combined and as long as tarsomere 1. Abdominal sternites punctate with elongate punctures. Sternite 8 about 2 times wider than sternites 4 – 7 medially. Pygidium transverse, irregularly punctate, partly hidden under elytra. Male unknown. Distribution and habitat The new species is known from only one locality, Alto Marzo in Zamora-Chinchipe province of Ecuador (Figure 1f), a place on the western side of the Cordillera del Cóndor, an elevated (over 2000m) enclave in the Amazonian hylaea that appears to present high endemicity. The exact habitat and collecting circumstances of the specimen are unknown. Considering the characters of the genus and complete absence of the soil contamination, normally found in all Aegidiini taxa except Onorius, we think that the new species is a canopy dweller similar to O. inexpectatus.

Onorius Frolov & Vaz-de-Mello, 2015 Type species Onorius inexpectatus Frolov & Vaz-de-Mello, 2015, by original designation., Published as part of Frolov, Andrey V., Akhmetova, Lilia A. & Vaz-de-Mello, Fernando Z., 2019, Contribution to the knowledge of the Neotropical canopy dwelling genus Onorius (Coleoptera�� Scarabaeidae�� Orphninae): another species from southern Ecuador and notes on the hind gut morphology in the Orphninae, pp. 1071-1077 in Journal of Natural History 53 (17) on page 1072, DOI: 10.1080/00222933.2019.1632952, http://zenodo.org/record/3673335, {"references":["Frolov AV, Vaz-de-Mello FZ. 2015. A new genus and species of Orphninae (Coleoptera: Scarabaeidae) associated with epiphytes in an Andean cloud forest in Ecuador. Zootaxa. 4007: 433 - 436. doi: 10.11646 / zootaxa. 4007.3.10."]}

Onorius sp. Material examined Female. ��� Zamora Chinchipe Paquisha Alto Marzo 2008 Col. V. Carvajal ��� (CEMT). Diagnosis This species differs from O. inexpectatus in the sculpture of the body (Figure 1a): head is punctate with transversely elongate, sometimes adjoining punctures (punctures are round and relatively sparse in O. inexpectatus); pronotum with coarser ocellate punctures (punctures are smaller and not ocellate in O. inexpectatus); elytra with characteristic U- to V-shaped punctures each bearing a distinct yellowish seta posteriorly (punctures are round in O. inexpectatus; setae in the punctures are smaller, barely visible at the same magnification); proximal ileum has two large, symmetrical diverticula (Figure 2b). Description Female (Figures 1 and 2). Body length 9.0 mm, width of elytra 4.6 mm, width of pronotum 3.5 mm. Colour uniform brown with pronounced bronze tint. Head densely punctate with transversely elongate punctures; some punctures fused near anterior margin of clypeus. Head without traces of medial horn or tubercle. Left mandible slightly protruding past clypeus in dorsal view; labrum not protruding past clypeus. Clypeus rounded anteriorly. Pronotum 1.5 times wider than long, widest at basal 0.3 of its length. Anterior margin without border, basal and lateral margins bordered. Disc with shallow impression anteriorly and near base, without tubercles or deep fossae. Surface punctate with round ocellate punctures separated by 1 ��� 3 puncture diameters. Scutellum triangular, narrowly rounded apically, about 1/15 length of elytra, coarsely punctate. Elytra convex, with marked humeral and apical humps, without ridges or other elevated areas.Maximum width approximately at the middle. Striae indistinct. Elytra punctate with U- or V-shaped punctures; each puncture bears a short, yellow seta posteriorly. Base of elytra not bordered. Wings fully developed. Protibiae with 3 outer teeth, without smaller inner tooth. Lateral margin basad of outer teeth not crenulate. Apex and internal margin of tibia with a few slender setae. Protarsi well developed, about 2/3 length of protibiae. Claws 1/3 length of tarsomere 5. Tarsomere 5 as long as tarsomeres 2 ��� 4 combined. Tarsomere 1 as long as tarsomeres 2 ��� 4. Hind and middle legs similar in shape; metafemora and metatibiae about 1.2 times longer than mesofemora and mesotibiae. Femora sparsely punctate with elongate punctures. Tibiae somewhat triangular, with 2 apical spurs. Upper tibial spur as long as tarsomeres 1 ��� 3; lower spur as long as or slightly longer than tarsomere 1. Claws 1/3 length of tarsomere 5. Tarsomere 5 relatively robust, as wide as the other tarsomeres, as long as tarsomeres 2 ��� 4 combined and as long as tarsomere 1. Abdominal sternites punctate with elongate punctures. Sternite 8 about 2 times wider than sternites 4 ��� 7 medially. Pygidium transverse, irregularly punctate, partly hidden under elytra. Male unknown. Distribution and habitat The new species is known from only one locality, Alto Marzo in Zamora-Chinchipe province of Ecuador (Figure 1f), a place on the western side of the Cordillera del C��ndor, an elevated (over 2000m) enclave in the Amazonian hylaea that appears to present high endemicity. The exact habitat and collecting circumstances of the specimen are unknown. Considering the characters of the genus and complete absence of the soil contamination, normally found in all Aegidiini taxa except Onorius, we think that the new species is a canopy dweller similar to O. inexpectatus., Published as part of Frolov, Andrey V., Akhmetova, Lilia A. & Vaz-de-Mello, Fernando Z., 2019, Contribution to the knowledge of the Neotropical canopy dwelling genus Onorius (Coleoptera�� Scarabaeidae�� Orphninae): another species from southern Ecuador and notes on the hind gut morphology in the Orphninae, pp. 1071-1077 in Journal of Natural History 53 (17) on pages 1072-1075, DOI: 10.1080/00222933.2019.1632952, http://zenodo.org/record/3673335

Frolov, Andrey V., Akhmetova, Lilia A., Vaz-de-Mello, Fernando Z. (2019): Contribution to the knowledge of Aegidinus Arrow (Coleoptera: Scarabaeidae: Orphninae): new species and comments on the classification and nomenclature. Journal of Natural History 53 (11): 725-747, DOI: 10.1080/00222933.2019.1606953

Aegidinus simulatus Colby, 2009 (Figure 4 (e���h)) Aegidinus simulatus Colby, 2009: 32 Material examined PERU. Loreto: Curaray River, Gaseoducto Barrett, 25 February 2008, W. Yawarcani leg., two males and three females (UNMSM). COLOMBIA. Meta: San Mart��n, Tocancipa Ranch, 520 m, forest, pitfall, April 2006, J. Noriega leg., one male (CEMT). Diagnosis Aegidinus simulatus is similar to Ae. brasiliensis, Ae. howeae and Ae. colbyae sp. nov. in having mediobasal margins of the dorsomedial lobes of the parameres strongly sclerotised, but can be distinguished from them by having the dorsal sides of the parameres strongly overlapping and separated by a slit (Figure 4 (f), arrowed). Description of male Body length 9.5 ��� 10.0 mm. Colour uniformly blackish brown. Anterior margin of frontoclypeus with a horn rounded apically (Figure 4 (e)). Pronotum with widely rounded lateral margins, narrower than elytra, 1.6 times wider than length. Posterior angles widely rounded. Anterior margin bordered, border narrowly interrupted medially, with short, somewhat bimodal tubercle. Base of pronotum not bordered, with obsolete punctures laterally. Pronotal disc feebly excavated anteromedially, with two gibbosities in centre. Pronotum punctate with a few large rounded punctures laterally and transversely elongated punctures anteromedially and with minute, feebly visible punctures throughout. Elytra almost as long as wide, widest medially and rounded apically, with humeral and apical humps. First elytral stria as continuous line not connected basally with undulate line from scutellum to humeral hump. Other striae marked with rows of sparse elongate punctures. Protibia without medioapical tooth. Aedeagus relatively long and slender, ratio of phallobase length/paramere width 2.6 (Figure 4 (f)). Ventroapical plate of phallobase absent (Figure 4 (h)). Parameres symmetrical, mediobasal margins of dorsomedial lobes strongly sclerotised (4G), dorsal sides of the parameres strongly overlapping and separated by a slit (Figure 4 (f), arrowed). Distribution The species was described from Ali��ahui (Ecuador, Napo Province) and Gigante (Huila, Colombia). The new records are from Curaray River (Peru, Loreto) and San Mart��n (Colombia, Meta) (Figure 7)., Published as part of Frolov, Andrey V., Akhmetova, Lilia A. & Vaz-de-Mello, Fernando Z., 2019, Contribution to the knowledge of Aegidinus Arrow (Coleoptera: Scarabaeidae: Orphninae): new species and comments on the classification and nomenclature, pp. 725-747 in Journal of Natural History 53 (11) on page 743, DOI: 10.1080/00222933.2019.1606953, http://zenodo.org/record/3675571

Aegidinus guianensis (Westwood, 1845) (Figure 1 (a���i)) Aegidium guianense Westwood, 1845: 175 = Aegidinus oreibates Colby, 2009: 26, syn. nov. = Aegidinus crypticus Colby, 2009: 20, syn. nov. Type locality. Guyana, Upper Takutu-Upper Essequibo. Type material examined Aegidium guianensis: holotype (Figure 1 (a)), female at OUMNH labelled ��� Guiana / Aegidium? Guianense Westw./TYPE WESTWOOD Trans. Ent. Soc L.1846.P.175 T. 12. 8. 5. Coll. Hope Oxon./TYPE COL: 488 Aegidium guianensis Westw. HOPE DEPT. OXFORD/ W/WB ���. Aegidium oreibates: holotype (Figure 1 (f)), male at CMN labelled ��� TRINIDAD: ST. GEORGE 8 km N. Arima, Simla Res. Sta., 260 m, 14 ��� 24.VI.1993 S. & J. Peck, lower mont. rainforest, f.i.t., 93 ��� 48/ Aegidinus oreibates Colby HOLOTYPE ���. Paratypes: GUYANA. Potaro-Siparuni: one female at CMN labelled ��� Guyana: Iwokrama F. R. 4��40 �� 19N 58�� 41 �� 04W 100 ��� 200 m, V-VI.2001 R.Brooks & Z.Falin in FIT/ Aegidinus oreibates J. Colby PARATYPE ���. TRINIDAD AND TOBAGO. One female at IRSNB labelled ��� ♀ /Coll. R. I. Sc. N. B. Caparo Trinidad A. Heyne, Berlin-Wilm./ Aegidinus oreibates Colby PARATYPE / Paratype / Aegidinus guianense Westw ♀ E. Benderitter, det ���. Arima: one male at CMN labelled ��� TRINIDAD, 16 km N Arima Andrews Trace, 620m up. montane rainforest FIT, 7 ��� 24.VI.1993 S&J Peck, 93 ��� 14/ PARATYPE Aegidinus oreibates J. Colby ���; one male at CMN labelled ��� TRINIDAD, 8 km N Arima Simla Res. Sta. 260m 24.VI ��� 8.VII.93, lower montane rainforest FIT S&J Peck, 93 ��� 57/ Aegidinus oreibates J. Colby PARATYPE ���; one female at CMN labelled ��� TRINIDAD: ST. GEORGE 8 km N. Arima, Simla Res. Sta., 260 m, 14 ��� 24. VI.1993 S. & J. Peck, lower mont. rainforest, f.i.t., 93 ��� 48/ Aegidinus oreibates Colby ALLOTYPE ���; one male and two females at CMN labelled ��� TRINIDAD: ST. GEORGE 8 km N. Arima, Simla Res. Sta., 260 m, 14 ��� 24.VI.1993 S. & J. Peck, lower mont. rainforest, f.i.t., 93 ��� 48/ Aegidinus oreibates Colby PARATYPE ���; two males and one female at CMN labelled ��� TRINIDAD, 8 km N Arima Simla Res. Sta. 260m 6 ��� 14.VI.93 trop. forest FIT S&J Peck, 93 ��� 9/ PARATYPE Aegidinus oreibates J. Colby ���. Couva-Tabaquite-Talparo: one male at CMN labelled ��� TRINIDAD, 11 km SE Arima, Arena For. Res. 13 ��� 22.VI.93, 80m rainforest FIT S&J Peck, 93 ��� 30/ PARATYPE Aegidinus oreibates J. Colby ���; one male at CMN labelled ��� TRINIDAD, 13 km S Arima 2km N Talparo, Quesnell Farm, rainforest FIT 12 ��� 22.VI.93, 50m S&J Peck, 93 ��� 28/ PARATYPE Aegidinus oreibates J. Colby ���. VENEZUELA. Bol��var: one female at CMN labelled ��� VENEZUELA: BOLIVAR Guri, 14.VI.1996 H.&A. Howden Dry forest L +/ Aegidinus oreibates J. Colby PARATYPE ���. Additional material examined VENEZUELA. Bol��var: Guri, dry forest, 14 June 1996, H. Howden and A. Howden leg., one female (CEMT); Guri, wet forest, 17 June 1996, B.D. Gill leg., one female (BGCO). TRINIDAD AND TOBAGO. 11 km SE Arima, Arena Forest Reserve, 80 m, rainforest, FIT, 13 ��� 22 June 1993, S. Peck, J. Peck leg., one male (BGCO); 8 km N Arima, Simla Research Station, 260 m, lower mountane rainforest FIT, 6 ��� 14 June 1993, S. Peck, J. Peck leg., one male (BGCO); 16 km N Arima, Andrews Trace, 620 m, upper montane rainforest, FIT, 24 June ��� 7 July 1993, S. Peck, J. Peck leg., one male (BGCO); 8 km N Arima, Simla Research Station, 260 m, lower montane rainforest, FIT, 24 June ��� 8 July 1993, S. Peck, J. Peck leg., one female (BGCO). GUYANA. Potaro-Siparuni: Pataro-Siparuni Iwokara Rain Forest, 70 m, human faeces baited, 13 April ��� 31 May 2009, S. Phelps leg., one female (OUMNH). SURINAME. Road to Amotopo, km 39, unbaited pitfalls, secondary forest, 29 May ��� 2 June 1981, J. Huijbregts leg., one female (RMNHL); Sipaliwini W-bank Suriname River opposite Pingpe 4.041667�� N, 55��45 W, FIT in forest, 25 January ��� 1 February 2012, M.A. Hielkema leg., one female (RMNHL); road to Amotopo, km 34, primary forest, pitfalls baited with human dung, 28 May ��� 1 June 1981, J. Huijbregts leg., one female (RMNHL). FRENCH GUIANA. Mont Grand Matoury, 24 January 2013, SEAG leg., two females (ZIN); 7 January 2013, SEAG leg., one male (ZIN); Gourdonville, 1905, E. Le Moult leg., one male and one female (MNHN). BRAZIL. Roraima: Cariri, Vila Napole��o, 4 May 1972, J. Celso leg., one male (CEMT); 10 April 1972, J. Celso leg., one female (CEMT). Par��: Bel��m, one female (MNHN). Diagnosis Aegidinus guianensis is most similar to Ae. howdenorum but differs from it in having the ventrolateral lobe of the parameres without a subapical tooth (Figure 1 (h)). From other Aegidinus species it differs in having the phallobase with a ventroapical plate and symmetrical parameres (Figure 1 (g,i)). Remarks Paulian (1984) and Colby (2009) drew male characters of Ae. guianensis from the type of Ae. steinheili, which was considered a synonym of Ae. guianensis after Arrow (1904). Colby (2009) erroneously considered the female specimen with the label ��� Cayenne ���, housed in BMNH, to be the holotype of Ae. guianensis and drew the female characters from it. This specimen, in our opinion, belongs to Ae. howeae Colby or Ae. brasiliensis, but it lacks the abdomen and genitalia. Ae. guianensis was described from British Guiana (nowadays Guyana) and the holotype of it is housed in OUMNH (Figure 1 (a,b)). It agrees with the original description of the species, and its mouthparts were dissected and illustrated by Westwood (1845, pl. XII, fig. 5). The specimen was likely collected by Robert Schomburgk in Pirara on the shores of Lake Amucu (Darren Mann, pers. comm.). By the shape of the female genitalia and external morphological characters, the type of Ae. guianensis (Figure 1 (a,d)) cannot be distinguished from Ae. oreibates Colby (Figure 1 (c,e)), described from a long series mostly from the Guiana Shield and the island of Trinidad. For this reason, we propose that Aegidinus oreibates Colby is a synonym of Aegidium guianense Westwood. Because this long series includes both sexes, the male characters of Ae. guianensis can be reliably based on the males from this series. Ae. crypticus Colby was described from Guyana from a single male specimen. The difference between Ae. oreibates and Ae. crypticus is, according to the author, that in the latter species the median lobes of the parameres are longer and thinner and the shape of the ventral plate of phallobase at the apex is much rounder (Colby 2009). However, the detailed photographs of the parameres provided in the original descriptions (Colby 2009, figs 46 ��� 49, 59 ��� 60) show that these characters are not diagnostic: the median lobes of the parameres are of about the same length and width and the ventral plates are equally rounded. The differences between the photographs are due to the slightly different angle of view and because the illustrated parameres of Ae. oreibates are opened while those of Ae. crypticus are close to each other. Therefore, we propose that Aegidinus crypticus Colby is also a synonym of Aegidium guianense Westwood., Published as part of Frolov, Andrey V., Akhmetova, Lilia A. & Vaz-de-Mello, Fernando Z., 2019, Contribution to the knowledge of Aegidinus Arrow (Coleoptera: Scarabaeidae: Orphninae): new species and comments on the classification and nomenclature, pp. 725-747 in Journal of Natural History 53 (11) on pages 733-734, DOI: 10.1080/00222933.2019.1606953, http://zenodo.org/record/3675571, {"references":["Paulian R. 1984. Les Orphnidae Americains (Coleopteresi Scarabaeoidea). Annales de la Societe Entomologique de France (N S). 20: 65 - 92.","Colby J. 2009. Monographic revision of the genus Aegidinus Arrow (1904) and generic phylogeny of the world Orphninae (Coleoptera: Scarabaeidae: Orphninae). Insecta Mundi. 76: 1 - 41.","Arrow GJ. 1904. XXVII. Sound-production in the Lamellicorn Beetles. T Roy Ent Soc London. 52: 709 - 750. doi: 10.1111 / j. 1365 - 2311.1904. tb 02761. x","Westwood JO. 1845. On the lamellicorn beetles which possess exserted mandibles and labrumi and 10 - jointed antennae. Trans Entomol Soc London. 4: 155 - 180. doi: 10.1111 / j. 1365 - 2311.1846. tb 01350. x"]}

Aegidinus noriegai Frolov, Akhmetova and Vaz-de-Mello sp. nov. (Figure 4 (a ��� d)) Type locality. Colombia, Antioquia, Medell��n. Type material examined Holotype. Male (Figure 4 (a)) at CEMT labelled ��� COLOMBIA: Antioquia. Porce. Picardia V- 1998. pastizal, pitfall. J. Noriega ���. Diagnosis Aegidinus noriegai sp. nov. is most similar to Ae. candezei but can be distinguished from it by the longer and more asymmetrical parameres, ventroapical plate of the phallobase longer than wide (Figure 4 (b ��� d)), and the protibia without medioapical tooth. Description Male (Figure 4 (a)). Body length 8.5 mm. Colour uniformly blackish brown. Anterior margin of frontoclypeus with a horn rounded apically. Pronotum with widely rounded lateral margins, narrower than elytra, 1.6 times wider than length. Posterior angles widely rounded. Anterior margin bordered, border interrupted medially, with feeble gibbosity. Base of pronotum not bordered, with a few large rounded punctures laterally and a few small medially. Pronotal disc feebly excavated anteromedially, with two gibbosities in centre. Pronotum punctate with a few large rounded punctures laterally and anteromedially and with minute, feebly visible punctures throughout. Elytra almost as long as wide, widest medially and rounded apically, with humeral and apical humps. First elytral stria as continuous line, connected basally with undulate line from scutellum to humeral hump. Other striae marked with rows of sparse punctures; punctures somewhat V- and comma-shaped on basal part of elytra, becoming smaller towards apices. Protibia without medioapical tooth. Aedeagus relatively short and wide, ratio of phallobase length/paramere width 1.8 (Figure 4 (b)). Ventroapical plate of phallobase narrow and long, 1/3 as wide as phallobase and longer than wide (Figure 4 (d)). Parameres strongly asymmetrical (Figure 4 (c,d)): right paramere with relatively long medial and lateral processes, medial process relatively straight; left paramere with vestigial lateral process; medial process curved outwards, with small tooth near apex. Distribution The species is known from a single locality in Antioquia, Colombian Andes (Figure 7). Etymology The new species is named after Jorge Noriega (University of Los Andes, Bogot��, Colombia)., Published as part of Frolov, Andrey V., Akhmetova, Lilia A. & Vaz-de-Mello, Fernando Z., 2019, Contribution to the knowledge of Aegidinus Arrow (Coleoptera: Scarabaeidae: Orphninae): new species and comments on the classification and nomenclature, pp. 725-747 in Journal of Natural History 53 (11) on pages 738-739, DOI: 10.1080/00222933.2019.1606953, http://zenodo.org/record/3675571

Aegidinus steinheili (Harold, 1880), distinct species (Figure 2 (a���g)) Aegidium steinheili Harold, 1880: 42 Aegidinus guianensis nec Westwood, 1845: 175, auct.: Arrow 1904: 739, 1912: 31; Paulian 1984, 98; Colby 2009: 20 Type locality. Colombia, Antioquia, San Carlos. Type material examined Lectotype. Male (Figure 2 (a)) at MNHN labelled ��� S. Carlos/Columbia Aegidinus Steinheili Harold /Col. Benderitter ae. guianense/ Aegidinus guganesis (Westw.) R. Paulian det./ Aegidinus steinheili (Har.) R. Paulian det./Museum Paris/Museum Paris Coll. M. Pic/ HOLOTYPE / Aegidinus guianensis ���. The labels of this specimen do not agree completely with the data provided by Paulian (1984), but we believe that this is the specimen that Paulian intended to designate as the lectotype. There are cases in which Paulian did not properly label his material with type labels (Olivier Montreuil, Antoine Mantilleri, pers. comm.). However, this specimen is the only one bearing Paulian ��� s handwritten identification labels and it definitely belongs to the type series from San Carlos. Paralectotypes. COLOMBIA. Antioquia: one female at MNHN labelled ��� San Carlos /G. J. Arrow Vidit 1903./ Ex. Musaeo E. Steinheil / MNHN ���; two males at MNHN labelled ��� San Carlos /G. J. Arrow Vidit 1903./sp. nov./ Ex. Musaeo E. Steinheil / Museum Paris / Aegidinus candezei (Preudhomme de Borre) det. J.B. Colby 2008/ MNHN ���; one male at MNHN labelled ��� Nare (Wallis)/G. J. Arrow Vidit 1903./ Ex. Musaeo E. Steinheil / Museum Paris / MNHN 03 ��� 2005 ADSmith/ MNHN / Aegidinus guianensis (Westwood) det. J.B. Colby 2008 ���; one female at BMNH labelled ��� S. Carlos / Aegidinus guianensis, Westw. Agrees with type (♀)/ Ex. Musaeo E. Steinheil /ex Coll. Oberthur. 1904 ��� 175./ Cotype / BMNH / Aegidinus guianensis Westwood INVALID TYPE det. J.B. Colby 2007 ���; one female at MNHN labelled ��� San Carlos /G. J. Arrow Vidit 1903./ Ex. Musaeo E. Steinheil / Brit. Mus. Accepit./ MNHN ���; one female at MNHN labelled ��� San Carlos /G. J. Arrow Vidit 1903./ Ex. Musaeo E. Steinheil / MNHN ���; one female at MNHN labelled ��� San Carlos /G. J. Arrow Vidit 1903./ Ex. Musaeo E. Steinheil / Museum Paris / MNHN 03 ��� 2005 ADSmith/ MNHN / Aegidinus guianensis (Westwood) det. J.B. Colby 2008 ���; one male at MNHN labelled ��� San Carlos /G. J. Arrow Vidit 1903./ Steinheili Harold / Ex. Musaeo E. Steinheil / Brit. Mus. Accepit./ Museum Paris / MNHN / Aegidinus guianensis (Westwood) det. J.B. Colby 2008 ���; one male at BMNH labelled ��� San Carlos / Steinheili Har. [...]/ Ex. Musaeo E. Steinheil /ex Coll. Oberthur. 1904 ��� 175./ Cotype /BMHN/ Aegidinus guianensis Westwood INVALID TYPE det. J.B.Colby 2007 ���. Additional material examined COLOMBIA. One male at MNHN, Canoas [not traced locality]. One male at IRSNB labelled ��� Coll. R. I. Sc. N. B. Bresil Steinheil. ex coll. Cand��ze/ Aegidium Alatum Cast. Br. / Aegidinus guianensis (Westwood) det. J.B. Colby 2008 ���; a mislabelled specimen (see below) most probably collected in San Carlos. Diagnosis Aegidinus steinheili is most similar to Ae. petrovi Colby and Ae. teamscaraborum Colby but differs from them by having ventrolateral lobes of parameres long and slender (in lateral view), noticeably longer than dorsomedial lobes (Figure 2 (d ��� f)). The female has a punctation of the body similar to that of the male (Figure 2 (c)); the external female genitalia are as in Figure 2 (g). Remarks Arrow (1904) considered Ae. steinheili (Harold 1880) to be a synonym of Ae. guianensis (Westwood 1845) although he did not study the genitalia of the specimens. Paulian (1984) and Colby (2009) followed Arrow, apparently because they did not study the type of Ae. guianensis and thus misinterpreted this species. Examination of the female type of Ae. guianensis and conspecific males (see above) showed that Ae. steinheili is distinct from it and all other Aegidinus species. Paulian (1984) studied most of the Aegidinus specimens from San Carlos and one from Nare and, based on the external characters, identified them as two species, Ae. steinheili (= Ae. guianensis sensu Paulian) and Ae. candezei. He designated the larger male from San Carlos, with the denser punctation of the body, as the lectotype of Ae. steinheili (Figure 2 (a)). We dissected all specimens of this series and found that characters of the male and female genitalia are congruent with the characters of the external morphology. In this material we found two male morphotypes and two female morphotypes and we agree with Paulian that they belong to the two species mentioned above. In the collection of IRSNB there is a male specimen of this species with the labels ��� Coll. R.I.Sc.N.B. Bresil Steinheil. ex coll. Cand��ze/ Aegidium Alatum Cast. Br. / Aegidinus guianensis (Westwood) det. J.B.Colby 2008 ���. It was among a small series of Aegidiellus alatus Laporte de Castelnau having the same printed labels ��� Coll. R.I.Sc.N.B. Bresil [...] ��� but with varying additional data. These labels were placed by IRSNB technical staff under previously unlabelled specimens that were in the collection drawer next to the bottom label ��� Aegidiellus alatus ���. The record of Ae. steinheili from Brazil (Colby 2009, as Ae. guianensis) is apparently based on this mislabelled specimen which is very similar to the larger males of the type series (Figure 2 (b))., Published as part of Frolov, Andrey V., Akhmetova, Lilia A. & Vaz-de-Mello, Fernando Z., 2019, Contribution to the knowledge of Aegidinus Arrow (Coleoptera: Scarabaeidae: Orphninae): new species and comments on the classification and nomenclature, pp. 725-747 in Journal of Natural History 53 (11) on pages 735-736, DOI: 10.1080/00222933.2019.1606953, http://zenodo.org/record/3675571, {"references":["Arrow GJ. 1904. XXVII. Sound-production in the Lamellicorn Beetles. T Roy Ent Soc London. 52: 709 - 750. doi: 10.1111 / j. 1365 - 2311.1904. tb 02761. x","Arrow GJ. 1912. Pachypodinaei Pleocominaei Aclopinaei Glaphyrinaei Ochodaeinaei Orphninaei Idiostominaei Hybosorinaei Dynamopinaei Acanthocerinaei Troginae. In: Coleopterorum Catalogus. Berlin: Junki W.; p. 1 - 66.","Paulian R. 1984. Les Orphnidae Americains (Coleopteresi Scarabaeoidea). Annales de la Societe Entomologique de France (N S). 20: 65 - 92.","Colby J. 2009. Monographic revision of the genus Aegidinus Arrow (1904) and generic phylogeny of the world Orphninae (Coleoptera: Scarabaeidae: Orphninae). Insecta Mundi. 76: 1 - 41.","Harold E. 1880. Verzeichniss der von E. Steinheil in Neu-Granada gesammlten coprophagen Lamellicornien. Stettiner entomologische Zeitung. 41: 13 - 46.","Westwood JO. 1845. On the lamellicorn beetles which possess exserted mandibles and labrumi and 10 - jointed antennae. Trans Entomol Soc London. 4: 155 - 180. doi: 10.1111 / j. 1365 - 2311.1846. tb 01350. x"]}

Key to Aegidinus species (males) 1. Parameres separated into dorsomedial and ventrolateral lobes (Figures 1 (h,i), 2(e,f) and 3(b ��� d))....................................................................................................................................................... 2 - Parameres not separated into dorsomedial and ventrolateral lobes (Figure 6 (c ��� e))................................................................................................................................... Aegidinus cornutus 2. Phallobase with ventroapical plate (Figures 1 (g), 3(c) and 4(d))......................................... 3 - Phallobase without ventroapical plate (Figures 2 (d), 4(h) and 5(d)).............................. 6 3. Parameres symmetrical.......................................................................................................................... 4 - Parameres asymmetrical.................................................................................................................... 5 4. Ventrolateral lobe of paramere with subapical tooth (Colby 2009, fig. 55)....................................................................................................................................................... Aegidinus howdenorum - Ventrolateral lobe of paramere without subapical tooth (Figure 1 (h)).......................................................................................................................................................... Aegidinus guianensis 5. Parameres longer, more asymmetrical (Figure 4 (b,c)); ventroapical plate of phallobase longer than wide (Figure 4 (d)); protibia without medioapical tooth.................................................................................................................................................................... Aegidinus noriegai sp. nov. - Parameres shorter, less asymmetrical (Figure 3 (b,d)); ventroapical plate of phallobase wider than long (Figure 3 (c)); protibia with medioapical tooth Aegidinus candezei 6. Mediobasal margins of dorsomedial lobes of parameres feebly sclerotised, membranous (Figure 2 (e)); protibia with medioapical tooth...................................................................... 7 - Mediobasal margins of dorsomedial lobes of parameres strongly sclerotised (4G, 5E); protibia without medioapical tooth............................................................................................. 9 7. Ventrolateral lobes of parameres long and slender (in lateral view), noticeably longer than dorsomedial lobes (Figure 2 (f)).................................................................................... steinheili - Ventrolateral lobes of parameres triangular and obtuse in lateral view, not longer than dorsomedial lobes..................................................................................................................... 8 8. Ventrolateral lobes of parameres as long as dorsomedial lobes...... Aegidinus petrovi - Ventrolateral lobes of parameres noticeably shorter than dorsomedial lobes.............................................................................................................................. Aegidinus teamscaraborum 9. Dorsal sides of parameres strongly overlapping and separated by slit (Figure 4 (f), arrowed).................................................................................................................... Aegidinus simulatus - Dorsal sides of parameres less overlapping and not separated by slit (Figure 5 (f)) 10 10. Dorsal processes of parameres carina shaped (Figure 5 (e), arrowed)............................................................................................................................................................ Aegidinus colbyae sp. nov. - Dorsal processes of parameres tooth or spur shaped........................................................ 11 11. Dorsal processes of parameres long, spur shaped...................... Aegidinus brasiliensis - Dorsal processes of parameres short, tooth shaped........................ Aegidinus howeae, Published as part of Frolov, Andrey V., Akhmetova, Lilia A. & Vaz-de-Mello, Fernando Z., 2019, Contribution to the knowledge of Aegidinus Arrow (Coleoptera: Scarabaeidae: Orphninae): new species and comments on the classification and nomenclature, pp. 725-747 in Journal of Natural History 53 (11) on pages 727-731, DOI: 10.1080/00222933.2019.1606953, http://zenodo.org/record/3675571, {"references":["Colby J. 2009. Monographic revision of the genus Aegidinus Arrow (1904) and generic phylogeny of the world Orphninae (Coleoptera: Scarabaeidae: Orphninae). Insecta Mundi. 76: 1 - 41."]}

Aegidinus venezuelensis Colby, 2009 Aegidinus venezuelensis Colby, 2009: 38 Type locality. Venezuela, Distrito Capital, Caracas. Type material examined Holotype. Female at MNHN labelled ��� V��n��zu��la Caracas/EX MUSEO N VAN DE POLL/ MUS��UM PARIS 1936 COLL. A. BOUCOMONT/ Aegidinus brasiliensis R. Paulian det. Arr./ Aegidinus venezuelensis Colby HOLOTYPE ���. Remarks The type of this species cannot be reliably distinguished from Ae. guianensis and Ae. howdenorum females. Distribution suggests that it may belong rather to Ae. howdenorum, but additional material is needed. It should be noted that Ae. howdenorum is known from a pair of specimens; the third female from ��� Colombia ��� was tentatively identified as Ae. howdenorum and cannot be distinguished from females of Ae. guianensis. Considering the similarity of the two species ��� guianensis and howdenorum differ only in a slightly different shape of the parameres, the difference smaller than in other Aegidinus species ��� additional material is also needed to clarify whether this difference is not due to the geographic variability among populations of one species., Published as part of Frolov, Andrey V., Akhmetova, Lilia A. & Vaz-de-Mello, Fernando Z., 2019, Contribution to the knowledge of Aegidinus Arrow (Coleoptera: Scarabaeidae: Orphninae): new species and comments on the classification and nomenclature, pp. 725-747 in Journal of Natural History 53 (11) on pages 741-742, DOI: 10.1080/00222933.2019.1606953, http://zenodo.org/record/3675571

Aegidinus candezei (Preudhomme de Borre, 1886) (Figure 3 (a���e,g,h,j,k)) Orphnus candezei Preudhomme de Borre, 1886: 116 Aegidinus candezei (Preudhomme de Borre); Arrow, 1912: 31; Schmidt, 1913: 76; Paulian 184: 89; Colby 2009: 17 Aegidinus sunidigea Colby, 2009, syn. nov. Type locality. Colombia, Choc��. Type material examined Lectotype (designated here, Figure 3 (a,e)), male at IRSNB labelled ��� Coll. R. I. Sc. N. B. Colombie ex coll Candeze Choco Wallis /det. P. de Borre 1886 Orphnus Candezei ♂ Type/ Holotype / Lectotypus Orphnus candezei P.d.B. Frolov des. 2018 ���. Paralectotype (Figure 3 (f,i,l)), female at IRSNB labelled ��� Coll. R. I. Sc. N. B. Colombie: Choco ex Wallis ex coll. Candeze /det. P. de Borre 1886 Orphnus Candezei ♀ Type/ Allotype / Aegidinus candezei (Preudhomme de Borre) det. J.B. Colby 2009 / Paralectotypus Orphnus candezei P.d.B. Frolov des. 2018 ���. Additional material examined COLOMBIA. Boyac��: Otanche, 5 May 1995, Arri��gada leg., five females (CEMT). Santander: Puerto Ara��jo, Los Manantiales Ranch, May 2000, J. Noriega leg., one male (CEMT); Puerto Ara��jo, Las Marias, 130 m, May 2000, J. Noriega leg., one female (CEMT). Tolima: Mariquita, Municipal forest, 680 m, April 2000, J. Noriega leg., one male (CEMT). Antioquia: Medell��n, forest, pitfall, April 1998, J. Noriega leg., one male (CEMT). Diagnosis Ae. candezei is most similar to Ae. noriegai sp. nov. but can be distinguished from it by the shorter and less asymmetrical parameres, ventroapical plate of the phallobase wider than long (Figure 3 (b ��� d)), and protibia with medioapical tooth. Remarks Aegidinus candezei was described with no explicit information about the number of the examined specimens although it was indicated that both sexes were studied. There are two specimens (Figure 3 (a,i)) in the IRSNB collection which agree with the original description and might be the only specimens studied by Preudhomme de Borre. These specimens should be considered syntypes since no lectotype had been designated so far. The male specimen bears a red printed label that reads ��� Holotype ��� (Figure 3 (e)) and the female specimen the label ��� Allotype ��� (Figure 3 (f)). These labels were placed by the IRSNB technical staff in 1980 ��� 2000 (Alain Drumont, pers. comm.). Paulian (1984) mentioned the ��� holotype ��� apparently based on these labels. The identity and conspecificity of the two specimens were not questioned by Paulian (1984) and Colby (2009), and the latter author drew the female characters of Ae. candezei from the female syntype. However, we think that the two syntypes are not conspecific. The body punctation and the shape of the genitalia of the female syntype (Figure 3 (i,l)) do not agree with the female specimens from San Carlos (Figure 3 (h,k)) belonging to the series of both sexes with trustworthy locality data. Only the male syntype bears the original locality label ��� Choco Wallis ���. The locality label of the female syntype was added by IRSNB technical staff later and put under an unlabelled specimen. Although it seems probable that both specimens originate from Candeze ��� s collection and might have been collected by Wallis, the locality of the female syntype is unclear since Wallis made extensive expeditions throughout South America (Veitch 1906). Aegidinus sunidigea Colby was described from a female (Figure 3 (g,j)) from San Carlos. As discussed above, the series from San Carlos includes two species represented by both sexes, so the female characters of Ae. candezei can be inferred from these specimens with sparse punctation, including the type specimen of Ae. sunidigea. To ensure stability of the nomenclature, we here designate the male syntype (Figure 3 (a)) as the lectotype of Orphnus candezei and propose that Aegidinus sunidigea is a synonym of Orphnus candezei., Published as part of Frolov, Andrey V., Akhmetova, Lilia A. & Vaz-de-Mello, Fernando Z., 2019, Contribution to the knowledge of Aegidinus Arrow (Coleoptera: Scarabaeidae: Orphninae): new species and comments on the classification and nomenclature, pp. 725-747 in Journal of Natural History 53 (11) on pages 736-737, DOI: 10.1080/00222933.2019.1606953, http://zenodo.org/record/3675571, {"references":["Colby J. 2009. Monographic revision of the genus Aegidinus Arrow (1904) and generic phylogeny of the world Orphninae (Coleoptera: Scarabaeidae: Orphninae). Insecta Mundi. 76: 1 - 41.","Preudhomme de Borre A. 1886. Liste des Lamellicornes Laparostictiques recueillis par feu Camille Van Volxem pendant son voyage au Bresil et a la Plata en 1872 i suivie de la description de dix huit especes nouvelles et un genre nouveau. Annales de la Societe entomologique de Belgique. 30: 103 - 120.","Paulian R. 1984. Les Orphnidae Americains (Coleopteresi Scarabaeoidea). Annales de la Societe Entomologique de France (N S). 20: 65 - 92.","Veitch JH. 1906. Hortus Veitchii: A history of the rise and progress of the nurseries of Messrs. James Veitch and Sonsi together with an account of the botanical collectors and hybridists employed by them and a list of the most remarkable of their introductions. London: J. Veitch and Sons; p. 684 p."]}

Aegidinus cornutus Colby, 2009 (Figure 6 (a���f)) Aegidinus cornutus Colby, 2009: 18 = Aegidinus unicus Colby, 2009: 37, syn. nov. Type locality. Brazil, Mato Grosso, Sinop. Type material examined Aegidinus cornutus: Holotype. Male at CMN labelled ���SINOP 12��31�� S, 55��37�� W BR 163 km 500 a 600 Mato Grosso, BRASIL 350 m IX.1974 Alvarenga & Roppa col./Cole����o M. Alvarenga/H. & A. HOWDEN COLLECTION ex. A. Martinez coll./ Aegidinus cornutus Colby HOLOTYPE ���. Paratype. Male at CMN labelled ���Oct. 976 BRASIL Minas Gerais Sin��polis Alvarenga ��� leg. Coll. Martinez /H. & A. HOWDEN COLLECTION ex. A. Martinez coll./ Aegidinus cornutus Colby PARATYPE ���. Additional material examined BRAZIL. Mato Grosso: Sinop, 12��31�� S, 55��37�� W, BR 163 km 500���600, 350 m, September 1974, Alvarenga, Roppa leg., two males and one female (CEMT). Par��: Reden��ao, Pinkaiti ��� Aik, October 1999, P.Y. Scheffler leg., 18 males and nine females (CEMT); Reden��ao, Pinkaiti ��� Aik, FIT, 2 November 1998, P.Y. Scheffler leg., one male (CEMT). Diagnosis Aegidinus cornutus can be easily distinguished by the parameres not separated into dorsomedial and ventrolateral lobes (Figure 6 (c ��� e)). External female genitalia as in Figure 6 (f). Remarks Aegidinus cornutus and Ae. unicus were both described from Sinop, Mato Grosso, Brazil, from a male and a female, respectively. Additional specimens of both sexes with the same locality labels as the holotype and a rather long series of both sexes from Reden��ao (Figure 6 (a,b)) show that the the two names belong to the same species. Therefore, a new synonymy is here proposed. The label of the paratype of Ae. cornutus is wrong. There is no locality ��� Sin��polis ��� in Minas Gerais, and Sinop has been probably misinterpreted as an abbreviation; if in the original data the state was noted as MGr or MG, it could be confused with Minas Gerais (instead of Mato Grosso). M. Alvarenga went to Sinop (Mato Grosso), then a just founded town, more than once in the 1970s and he was proud of his extensive collecting done there (personal comments to FZVM on several occasions, 1992 ��� 2008)., Published as part of Frolov, Andrey V., Akhmetova, Lilia A. & Vaz-de-Mello, Fernando Z., 2019, Contribution to the knowledge of Aegidinus Arrow (Coleoptera: Scarabaeidae: Orphninae): new species and comments on the classification and nomenclature, pp. 725-747 in Journal of Natural History 53 (11) on page 740, DOI: 10.1080/00222933.2019.1606953, http://zenodo.org/record/3675571

Madecorphnus grandis Frolov, Smith, and Akhmetova, new species (Figures 1���5.) Type material. Holotype: male at FMNH labeled ��� MADAGASCAR: Befin- / gotra (11.0 km WSW), / R��s Anjanaharibe-Sud, / 14��45���S, 49��27���E, / 1565m, montane rain- / forest, 16.XI.1994 // FMHD #94-58, Winkler / extraction of sifted litter / (leaf mold, rotten wood) / B. L. Fisher #1232 (1-50) / FIELD MUS. NAT.���. Description. Holotype, male (Figs. 1���5). Body length 7.5 mm (without mandibles). Color uniformly dark brown, almost black, legs and underside of body somewhat paler. Right mandible 2 times longer than left, without tooth behind apex (Fig. 3). Labrum subtrapezoidal, with slightly rounded sides, length about 1/8 width (in dorsal view). Clypeus slightly asymmetrical, apically obtuse, with 2 long and 3 shorter setae on the apical margin. Canthus and frontal suture indistinct. Clypeus slightly depressed apicomedially. Head without traces of frontoclypeal suture, finely punctate with minute punctures separated by greater than 4 times their diameter. Pronotum approximately 1.5 times wider than long, widest medially. Disc of pronotum convex, without any depressions, tubercles, or ridges. Punctation on pronotum similar to that on head. Margins with relatively wide border, lateral margins with 4 long setae: 1 seta on basal angle, 1 seta approximately in the middle of lateral margin, and 2 setae on the apical angle. Scutellum triangular, angulate apically, about 1/12 length of elytra. Elytra convex, with distinct humeral and apical umbones, widest at basal third. First stria distinct and reaching the apex of elytron, other striae indistinct. Elytra with double punctation: entire surface with minute punctures similar to those on head and pronotum; disc with larger, sparse, elongate, setigerous punctures. Epipleura with long, sparse, brown setae. Base of elytron with border from scutellum to humeral callus. Wings fully developed. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex with robust, spur-like seta and a few smaller setae basally. Mesothoracic and metathoracic legs similar in shape to each other. Longer tibial spur shorter than mesotarsomeres 1���2 in mesothoracic legs and as long as metatarsomeres 1���2 in metathoracic legs. Apices of parameres rounded in lateral view (Fig. 4). Lateral teeth of parameres poorly developed, situated approximately in middle of lateral paramere margin. Endophallus with 2 large, robust spinules with a sclerotization near base of one of spinules, and 2 smaller, comma-shaped sclerites (Fig. 5). Female unknown. Diagnosis. This species is similar to M. peyrierasi Frolov, 2010 and, especially, M. perinetensis Frolov, 2010 in the shape of the endophallus armature but differs in having comma-shaped sclerites longer than in other species, wider and rounded in lateral view apices of parameres, and in less developed lateral teeth of parameres. The new species is also 1.5���2.0 mm longer than M. peyrierasi and M. perinetensis specimens and somewhat longer than the holotype of M. falcatus Paulian, 1992, the largest Madecorphnus species known so far. Distribution. The new species is known from a tropical rainforest in the Anjanaharibe-Sud Reserve, in mid-altitude escarpments of northern Madagascar (Fig. 6). The nearest Madecorphnus locality is that of M. barclayi Frolov, 2012 in the northern part of the Masoala Peninsula (Frolov 2012), some 100 km southeast of Anjanaharibe-Sud. The similar species M. peyrierasi and M. perinetensis occur some 500 km to the south (Fig. 6). Etymology. Adjective in the nominative singular. From the Latin grandis for large, denoting the relatively large size of the species., Published as part of Frolov, Andrey V., Smith, Andrew B. T. & Akhmetova, Lilia A., 2017, A new species of the genus Madecorphnus (Coleoptera: Scarabaeidae: Orphninae) from Madagascar in Zootaxa 4341 (2), DOI: 10.11646/zootaxa.4341.2.7, http://zenodo.org/record/1039495, {"references":["Paulian, R. (1992) Un nouveau genre et une nouvelle espece d'Orphnidae de Madagascar (Coleoptera, Scarabaeoidea). Revue Francaise d'Entomologie, New Series, 14, 169 - 171.","Frolov, A. V. (2012) Three new species of scarab beetle genus Madecorphnus Paulian, 1992 (Coleoptera: Scarabaeidae: Orphninae) from Madagascar. Proceedings of the Zoological Institute RAS, 316, 159 - 165."]}

Aegidiellus Paulian, 1984 Type species: Phileurus alatus Castelnau, 1840, by original designation. Aegidiellus: Paulian 1984: 86; Colby 2009: 5; Frolov 2012: 796; Frolov and Vaz-de-Mello 2015: 136; Frolov 2013: 42; Rojkoff and Frolov 2017: 355. Description Aegidiellus comprises small to medium-sized beetles (body length 7–11 mm), brown to black coloured, without pattern. Mandibles subsymmetrical, small, without lateral processes, protruding past anterior margin of frontoclypeus in dorsal view. Frontoclypeus without tubercles, horns or ridges. Propleurae with carinae separating anterolateral areas from basal area. Pronotum of ‘major’ males with deep excavation in the middle and two horn-shaped, lateral pronotal processes directed somewhat anteriorly. In ‘minor’ males, pronotal and head armature less developed: lateral pronotal processes short, tubercle-shaped and anterior frontoclypeal process carina-shaped. In females, pronotum impressed anteriorly on disc and with a tubercle medially on anterior margin. Scutellum narrowly rounded apically, about 1/ 10 length of elytra. Elytra convex, with marked humeral and apical umbones. Elytral striae as longitudinal rows of punctures; sutural stria normally distinct as a depressed furrow in apical half of elytron; elytra without elevated, longitudinal keels or smooth stripes on disc. Wings fully developed in all species. Metepisternon elongated, its posterior angle slightly triangular and situated in distinct concavity of epipleuron. Mesocoxal cavities connected by a hole. Protibiae with three strong outer teeth in both sexes, with a smaller, medial tooth in majority of males. Mesotibiae without tuft of setae ventroapically. Plectrum widely triangular, rounded apically, with a few sensory setae. Stridulatory file with 100–110 fine, slightly curved carinae (carinae counted according to Frolov and Akhmetova (2016)). Phallobase tubeshaped with strongly sclerotized ventral side but without differentiation of ventral and dorsal sclerites; ventroapical plate absent. Parameres symmetrical, apices with short, dense setae. Spiculum gastrale wide, triangular. Endophallus without armature. Subcoxites with sparse, long setae medioapically. Coxites triangular, with row of short spinules mediobasally and long, sparse setae apically. Stili not separated from coxites. Diagnosis Aegidiellus is a member of the putative Aegidiini clade, which also includes Aegidium, Paraegidium and Onorius. These genera share the following characters: mandibles without lateral processes, mesocoxal cavities connected by a hole; and metatibiae without transverse carina (Paulian 1984; Frolov 2012). Aegidiellus can be separated from Paraegidium by the sculpture of the dorsum of the body, which is almost glabrous and sparsely punctate as opposed to being densely, irregularly punctate and densely pubescent in the latter; from Onorius it can be separated by the prominent mandibles (well visible in dorsal view), and relatively slender tarsomeres. Aegidiellus is externally most similar to Aegidium but can be separated from it by the elytral striae as longitudinal rows of punctures, sutural stria normally distinct as a depressed line on apical half of elytron, and elytra without elevated longitudinal keels or smooth stripes on disc. From all Aegidiini, Aegidiellus differs in having setose apices of parameres and female with a tubercle on the anterior margin of the pronotum. Diagnostic characters of species Aegidiellus species are similar externally and the only character that reliably diagnoses the species is the shape of the aedeagus and especially the parameres. Females of Aegidiellus cannot be separated at present. Species composition and distribution Aegidiellus includes three species distributed in the low coastal parts of the Atlantic forest ecoregion. Key to Aegidiellus species (males) 1. Phallobase wider, its anterodorsal margin not protruding (Figure 1 (d)). Parameres wider, their lateral margin with deep sinuation forming a tooth basally (Figure 1 (e,f)).............................................................................................................................. A. dentipenis sp. nov. – Phallobase narrower, its anterodorsal margin membranous, protruding (Figures 2 (d) and 4(c)). Parameres narrower, their lateral margin without sinuation and tooth (Figures 2 (e) and 4(d))...................................................................................................................... 2 2. Parameres with apices wider (Figures 2 (e) and 3(d)). Dorsal side of parameres straight to slightly sinuate in lateral view (Figures 2 (d) and 3(c)).................. A. alatus – Parameres with apices narrower (Figure 4 (d)). Dorsal side of parameres convex in lateral view (Figure (4c))................................................................ Aegidiellus zezaoi sp. nov., Published as part of Frolov, Andrey V., Akhmetova, Lilia A. & Vaz-de-Mello, Fernando Z., 2017, Revision of the Neotropical scarab beetle genus Aegidiellus Paulian (Coleoptera: Scarabaeidae: Orphninae) with description of two new species, pp. 1767-1779 in Journal of Natural History (J. Nat. Hist.) (J. Nat. Hist.) 51 (29 - 30) on pages 1768-1770, DOI: 10.1080/00222933.2017.1353153, http://zenodo.org/record/5182397, {"references":["Paulian R. 1984. Les Orphnidae Americains (Coleopteres, Scarabaeoidea). Ann Soc Ent Fr (N S). 20: 65 - 92.","Castelnau FL. 1840. Histoire Naturelle des Insectes Coleopteres. Avec une introduction renfermant L' Anatomie et la Physiologie des Animaux Articules, par M. Brulle. Paris: P. Dumenil.","Colby J. 2009. Monographic revision of the genus Aegidinus Arrow (1904) and generic phylogeny of the world Orphninae (Coleoptera: Scarabaeidae: Orphninae). Ins Mundi. 76: 1 - 41.","Frolov AV. 2012. Diagnosis, classification, and phylogenetic relationships of the orphnine scarab beetles (Coleoptera, Scarabaeidae: Orphninae). Ent Rev. 92: 782 - 797.","Frolov AV, Vaz-de-Mello FZ. 2015. A new genus and species of Orphninae (Coleoptera: Scarabaeidae) associated with epiphytes in an Andean cloud forest in Ecuador. Zootaxa. 4007: 433 - 436.","Frolov AV. 2013. Stenosternus Karsch, a possible link between Neotropical and Afrotropical Orphninae (Coleoptera, Scarabaeidae). Zookeys. 335: 33 - 46.","Rojkoff S, Frolov AV. 2017. Revision du genre Aegidium Westwood, 1845 des Antilles (Coleoptera, Scarabaeidae, Orphninae, Aegidiini). Ann Soc Ent Fr. 52: 354 - 368.","Frolov AV, Akhmetova LA. 2016. Revision of the subgenus Orphnus (Phornus) (Coleoptera, Scarabaeidae, Orphninae). Euro J Taxon. 241: 1 - 20."]}

Aegidiellus zezaoi sp. nov. (Figure 4 (a–e)) Type locality Brazil, Linhares. Type material examined Holotype. Male at CEMT labelled ‘RFCVRD LINHARES ES BRASIL Data: 01/11/88 Col. JSS’. Paratypes. BRAZIL. Espírito Santo: one female with the same data as the holotype; four males at CEMT labelled ‘ BRASIL: ES Linhares – RFCVRD X.1987 JS Santos’; one male at CMNC labelled ‘ BRASIL Est. E. Santo Mun. Linhares P.N. Sooretama Coll. Martinez Nov.962 /H. & A. HOWDEN COLLECTION ex. A. Martinez coll. / Aegidiellus alatus (La Porte) det. J.B. Colby 2008’; three males at CMNC labelled ‘ BRASIL E° E. Santo Mun. Linhares P.N. Sooretama Coll. Martinez Nov.962 /H. & A. HOWDEN COLLECTION ex. A. Martinez coll. / Aegidiellus alatus (La Porte) det. J.B. Colby 2008’; 33 males and 55 females at CEMT, ZIN, EPGC, BMNH, IRSNB, MHNG, MNHN, NMPC and ZMHUB labelled ‘ BRASIL: Espirito Santo Linhares. Reserva Natural Vale. 19°8ʹ10ʺS 40°3ʹ17ʺW 40m IV-1988 J.S. Santos leg.’. Rio de Janeiro: one female at CMNC labelled ‘ BRASIL E. Santo Linhares Sooretama NOV.62 A. Martinez /H. & A. HOWDEN COLLECTION ex. A. Martinez coll. / Aegidiellus alatus (La Porte det. J.B.Colby 2008’. Diagnosis Aegidiellus zezaoi sp. nov. is most similar to A. alatus but can be separated from it by the parameres more tapering apically and having narrower apices (Figure 4 (d)); dorsal side of parameres slightly convex in lateral view (Figure 4 (c)). Description Holotype, male (Figure 4 (a,c–f)). Body length 8.0 mm. Colour uniformly dark brown. Anterior margin of frontoclypeus slightly convex in middle, slightly sinuate each side of medial convexity, rounded laterally, with a narrow border. Frontoclypeus shiny, coarsely punctate anteriorly and somewhat shagreened, sparsely punctate with smaller punctures posteriorly. Pronotum with widely rounded lateral margins, as wide as elytra, 1.5 times wider than length, 0.6 times length of elytra. Posterior angles sinuate in dorsal view. Anterior margin bordered laterally, convex, not protruding anteriorly, with a conical tubercle-like anterior pronotal process. Base of pronotum without furrow, punctate with irregular row of round to strongly elongated punctures. Pronotum deeply excavated in middle, almost flat on disc. Disc punctate with coarse, rounded punctures separated by one to two puncture anteriorly. Elytra 1.08 times longer than wide, with feebly distinct humeral and apical humps. Elytra widest in basal one-third, rounded apically in dorsal view. Elytral carinae marked with rows of elongated punctures separated by one to two times their length on disc, becoming denser laterally and apically. Sutural stria as furrow in apical one-third. Elytral intervals smooth. Aedeagus with relatively slender phallobase, separated into smaller basal and larger apical parts by constriction (Figure 4 (c)). Apical part of phallobase with protruding anterodorsal margin and without convexity anteroventrally. Lateral margin of parameres without sinuation. Apices relatively narrow and tapering apically (Figure 4 (d)). Spiculum gastrale triangular (Figure 4 (e)). Female (Figure 4 (b)). differs from male in having a long protibial spur, relatively smaller pronotum feebly sinuate medially on disc, anterior margin of pronotum not protruding, and anterior pronotal process small and tubercle-shaped. Variation Body length of males 8.0–9.0, females 7.1–9.3. Most males have less developed lateral pronotal processes and shallower excavation on pronotum than holotype. Distribution The species is known from a few close localities in vicinity of Linhares, Espírito Santo State, Brazil (Figure 1 (b)). Etymology The new species is named after Mr José Simplício dos Santos (Zezão), insect enthusiast and collector of the most part the type series., Published as part of Frolov, Andrey V., Akhmetova, Lilia A. & Vaz-de-Mello, Fernando Z., 2017, Revision of the Neotropical scarab beetle genus Aegidiellus Paulian (Coleoptera: Scarabaeidae: Orphninae) with description of two new species, pp. 1767-1779 in Journal of Natural History (J. Nat. Hist.) (J. Nat. Hist.) 51 (29 - 30) on pages 1776-1778, DOI: 10.1080/00222933.2017.1353153, http://zenodo.org/record/5182397

Aegidiellus dentipenis Frolov, Akhmetova & Vaz-de-Mello, sp. nov. (Figure 1 (a–f)) Type locality Brazil, Rio de Janeiro, near Juturnaiba Lake. Type material examined Holotype. male at CEMT labelled ‘L. Juturnaiba. Araruama, RJ BRASIL, XI.1981 M. Alvarenga //coleçao M. Alvarenga’. Diagnosis Aegidiellus dentipenis sp. nov. can be easily separated from the other species of the genus by the shape of the aedeagus: phallobase wider, its anterodorsal margin not protruding; parameres wider, their lateral margin with deep sinuation basally forming a tooth (Figure 1 (d–f)). It also differs in having a spiculum gastrale of intermediate form between T-shaped (similar to that in most Aegidium and in Paraegidium) and triangular (as in other Aegidiellus species). Description Holotype, male (Figure 1 (a–f)). Body length 8.5 mm. Colour uniformly dark brown. Anterior margin of frontoclypeus slightly convex in middle, slightly sinuate each side of medial convexity, rounded laterally, with a narrow border. Frontoclypeus relatively sparsely punctate with punctures having indistinct margins anteriorly, slightly rugose between eyes. Pronotum with widely rounded lateral margins, as wide as elytra, 1.5 times wider than length, 0.6 times length of elytra. Posterior angles sinuate in dorsal view. Anterior margin bordered laterally, with a conical tubercle-like anterior pronotal process. Base of pronotum without furrow, punctate with irregular row of large, elongated punctures. Pronotum deeply excavated in middle, almost flat on disc. Disc punctate with coarse, rounded punctures separated by one to two puncture diameters except for sparsely punctate area in near base. Elytra 1.08 times longer than wide, with feebly distinct humeral and apical humps. Elytra widest in basal one-third, rounded apically in dorsal view. Elytral carinae marked with rows of round to elongated punctures separated by one ot three diameters. Elytral intervals smooth. Aedeagus with robust phallobase, separated into smaller basal and larger apical parts by constriction (Figure 1 (d)). Apical part of phallobase without protruding anterodorsal margin and with convexity anteroventrally. Lateral margin of parameres with deep sinuation forming a tooth basally (Figure 1 (e,f)). Spiculum gastrale with wide cranial part (Figure 1 (c)). Female. Unknown. Distribution The species is known from one locality near Juturnaiba Lake, Rio de Janeiro State, Araruama Municipality, Brazil (Figure 1 (b)). Etymology From the Latin dentis for ‘tooth’ and penis., Published as part of Frolov, Andrey V., Akhmetova, Lilia A. & Vaz-de-Mello, Fernando Z., 2017, Revision of the Neotropical scarab beetle genus Aegidiellus Paulian (Coleoptera: Scarabaeidae: Orphninae) with description of two new species, pp. 1767-1779 in Journal of Natural History (J. Nat. Hist.) (J. Nat. Hist.) 51 (29 - 30) on pages 1771-1774, DOI: 10.1080/00222933.2017.1353153, http://zenodo.org/record/5182397

Aegidiellus zezaoi sp. nov. (Figure 4 (a–e)) Type locality Brazil, Linhares. Type material examined Holotype. Male at CEMT labelled ‘RFCVRD LINHARES ES BRASIL Data: 01/11/88 Col. JSS’. Paratypes. BRAZIL. Espírito Santo: one female with the same data as the holotype; four males at CEMT labelled ‘ BRASIL: ES Linhares – RFCVRD X.1987 JS Santos’; one male at CMNC labelled ‘ BRASIL Est. E. Santo Mun. Linhares P.N. Sooretama Coll. Martinez Nov.962 /H. & A. HOWDEN COLLECTION ex. A. Martinez coll. / Aegidiellus alatus (La Porte) det. J.B. Colby 2008’; three males at CMNC labelled ‘ BRASIL E° E. Santo Mun. Linhares P.N. Sooretama Coll. Martinez Nov.962 /H. & A. HOWDEN COLLECTION ex. A. Martinez coll. / Aegidiellus alatus (La Porte) det. J.B. Colby 2008’; 33 males and 55 females at CEMT, ZIN, EPGC, BMNH, IRSNB, MHNG, MNHN, NMPC and ZMHUB labelled ‘ BRASIL: Espirito Santo Linhares. Reserva Natural Vale. 19°8ʹ10ʺS 40°3ʹ17ʺW 40m IV-1988 J.S. Santos leg.’. Rio de Janeiro: one female at CMNC labelled ‘ BRASIL E. Santo Linhares Sooretama NOV.62 A. Martinez /H. & A. HOWDEN COLLECTION ex. A. Martinez coll. / Aegidiellus alatus (La Porte det. J.B.Colby 2008’. Diagnosis Aegidiellus zezaoi sp. nov. is most similar to A. alatus but can be separated from it by the parameres more tapering apically and having narrower apices (Figure 4 (d)); dorsal side of parameres slightly convex in lateral view (Figure 4 (c)). Description Holotype, male (Figure 4 (a,c–f)). Body length 8.0 mm. Colour uniformly dark brown. Anterior margin of frontoclypeus slightly convex in middle, slightly sinuate each side of medial convexity, rounded laterally, with a narrow border. Frontoclypeus shiny, coarsely punctate anteriorly and somewhat shagreened, sparsely punctate with smaller punctures posteriorly. Pronotum with widely rounded lateral margins, as wide as elytra, 1.5 times wider than length, 0.6 times length of elytra. Posterior angles sinuate in dorsal view. Anterior margin bordered laterally, convex, not protruding anteriorly, with a conical tubercle-like anterior pronotal process. Base of pronotum without furrow, punctate with irregular row of round to strongly elongated punctures. Pronotum deeply excavated in middle, almost flat on disc. Disc punctate with coarse, rounded punctures separated by one to two puncture anteriorly. Elytra 1.08 times longer than wide, with feebly distinct humeral and apical humps. Elytra widest in basal one-third, rounded apically in dorsal view. Elytral carinae marked with rows of elongated punctures separated by one to two times their length on disc, becoming denser laterally and apically. Sutural stria as furrow in apical one-third. Elytral intervals smooth. Aedeagus with relatively slender phallobase, separated into smaller basal and larger apical parts by constriction (Figure 4 (c)). Apical part of phallobase with protruding anterodorsal margin and without convexity anteroventrally. Lateral margin of parameres without sinuation. Apices relatively narrow and tapering apically (Figure 4 (d)). Spiculum gastrale triangular (Figure 4 (e)). Female (Figure 4 (b)). differs from male in having a long protibial spur, relatively smaller pronotum feebly sinuate medially on disc, anterior margin of pronotum not protruding, and anterior pronotal process small and tubercle-shaped. Variation Body length of males 8.0–9.0, females 7.1–9.3. Most males have less developed lateral pronotal processes and shallower excavation on pronotum than holotype. Distribution The species is known from a few close localities in vicinity of Linhares, Espírito Santo State, Brazil (Figure 1 (b)). Etymology The new species is named after Mr José Simplício dos Santos (Zezão), insect enthusiast and collector of the most part the type series.

Aegidiellus alatus (Castelnau) (Figures 2 (a–f) and 3(a–e)) Phileurus alatus Castelnau 1840: 116. Aegidium alatum: Arrow 1912: 31; Gemminger and Harold 1869: 1073; Preudhomme de Borre 1886: 24; Schmidt 1913: 70; Vulcano et al. 1966: 255; Aegidiellus alatus: Paulian 1984: 86; Colby 2009: 5; Frolov 2012: 796. Aegidium haedulus Westwood 1845: 174, synonymy by Gemminger & Harold 1869: 1073; Preudhomme de Borre 1886: 24 (as synonym of Aegidium alatus). Aegidium haedulum Westwood: Arrow 1912: 31 (as synonym of Aegidium alatum); Schmidt 1913: 70 (as synonym of Aegidium alatum). Type locality Brazil. Type material examined Neotype of Phileurus alatus (here designated according to ICZN article 75, see remarks below) and lectotype of Aegidium haedulum (here designated according to ICZN article 74, see remarks below): male at OUMNH labelled ‘ Aegidium haedulus Dej. Brasil /type of Aegidium haedulum haedulum Westw. /Type Westwood Trans. Ent. Soc. 1846. p. 174. Coll. Hope Oxon /TYPE COL: 485 Aegidium haedulum Westw. HOPE DEPT. OXFORD /LECTOTYPUS Aegidium haedulus West. Frolov et al. 2016 /NEOTYPUS Phileurus alatus Cast. Frolov et al. 2016 ’. Additional material examined BRAZIL. Espírito Santo: Córrego Itá, I.1956, W. Grossmann leg., one male (CMNC). Bahia: Santo Antonio da Barra, Salvador, XI-XII.1888, Gounelle leg., one male (MNHN). Minas Gerais: Doce River Valley, XII.1957, P. Coelho leg., one male (CMNC). Rio de Janeiro: Floresta da Tijuca, 6.I.1977, J. Celso leg., one male and one female (CEMT); Araruama, III.1984, P. Arnaud leg., four males and four females (CEMT); Ponta da Trindade, I.2004, E. D.F. Grossi leg., one male and one female (CEMT); Pico do Corcovado, Guanabara, XII.1965, Alvarenga and Seabra leg., one male (CEMT); Rio de Janeiro, one female (BMNH); Newton Santos leg., one male (MNRJ); Rio de Janeiro, Corcovado, XI.1962, Martinez leg., one male (CMNC); XII.1958, Martinez, Alvarenga and Seabra leg., one female (CMNC); 19.XII.1958, D. Zajciw leg., one female (MNRJ). Brazil (without precise locality), one male (ZIN), one male and one female (MHNG), one male and two females (BMNH), two males and one female (MNHN), one male and one female (NMPC), one male (IRSNB); Steinheil leg., one female (IRSNB); Deyrolle leg., one male and one female (IRSNB). No locality, two males (BMNH), two males and one female (MNHN). Diagnosis Aegidiellus alatus is most similar to A. zezaoi sp. nov. but can be separated from it by the parameres less tapering apically and with wider apices (Figures 2 (e) and 3(d)); dorsal side of parameres straight to slightly sinuate in lateral view (Figures 2 (d) and 3(c)). Description Male (Figure 3 (a)). Anterior margin of frontoclypeus slightly convex in middle, slightly sinuate each side of medial convexity, rounded laterally, with a narrow border. Frontoclypeus relatively densely punctate anteriorly and sparsely punctate to smooth posteriorly; posterior part of frontoclypeus mostly hidden under protruding anterior margin of pronotum. Pronotum with widely rounded lateral margins, as wide as elytra, 1.5 times wider than length, 0.6 times length of elytra. Posterior angles slightly sinuate in dorsal view. Anterior margin bordered laterally, trapezoidal, protruding anteriorly to cover most of posterior part of frontoclypeus, with a conical tubercle-like anterior pronotal process. Base of pronotum without furrow, punctate with irregular row of round to strongly elongated punctures, interrupted medially. Pronotum deeply excavated in middle, almost flat on disc. Disc punctate with coarse, rounded punctures separated by one to two puncture diameters except for sparsely punctate area in near base. Elytra 1.08 times longer than wide, with feebly distinct humeral and apical humps. Elytra widest in basal one-third, rounded apically in dorsal view. Elytral carinae marked with rows of round to elongated punctures separated by three to five times their diameters on disc, becoming denser laterally and apically. Sutural stria as furrow in apical half. Elytral intervals smooth. Aedeagus with relatively slender phallobase, separated into smaller basal and larger apical parts by constriction (Figures 2 (d)and 3(c)). Apical part of phallobase with protruding anterodorsal margin and without convexity anteroventrally. Lateral margin of parameres without sinuation. Apices wide and feebly tapering apically (Figures 2 (e) and 3(d)). Spiculum gastrale triangular (Figure 3 (e)). Female (Figure 3 (b)). differs from male in having a long protibial spur, relatively smaller pronotum feebly sinuate medially on disc, anterior margin of pronotum not protruding, and anterior pronotal process small and tubercle-shaped. Variation Body length of males 9.1–11.0, females 8.5–10.0. Pronotal armature of males varies: some specimens have less developed lateral pronotal processes, shallower excavation on disc, and rounded anterior margin with smaller anterior pronotal process, similar to that in females. Distribution The species is known from a number of localities spanning from Ponta da Trindade in the south to Salvador in the north (Figure 2 (f)). Most of the localities are situated in coastal areas. Remarks Phileurus alatus was described from ‘Brazil’ without more exact locality and indication of the number of specimens examined (Castelnau 1840). A few years later Westwood (1845) described Aegidium haedulus. In the catalogue by Gemminger and Harold (1869) and all subsequent literature, Aegidium haedulus was considered a junior synonym of Phileurus alatus. The type of the Castelnau’ s species is apparently lost. It is not mentioned in any literature and Paulian, who had probably looked for it for his revision of the New World Orphninae (Paulian 1984), noted that the type was unknown to him. Despite our extensive search in all major collections no Aegidellus specimens that might originate from Castelnau’ s collection were found. Castelnau’ s collection that probably included the type was donated to the National Institution, Washington DC, USA, in 1842 and later turned over to the Smithsonian Institution. The collecion was apparently destroyed in the fire at the Smithsonian Institution in 1865 (Evenhuis 2012). The brief diagnosis of Phileurus alatus agrees with all three currently recognized Aegidiellus species. Therefore the apparent loss of the type compromises stability of the nomenclature of the genus and makes the neotype designation necessary. Because Aegidium haedulus was never used as a valid name since its synonymy with Phileurus alatus was established and because the syntype of the former, housed in OUMNH, is a well-preserved male specimen (Figure 2 (a–e)) with all diagnostic characters of Aegidiellus alatus as it was understood by all authors, this specimen satisfies the criteria of the neotype designation (ICZN, article 75) and it is chosen as the only name-bearing specimen for Phileurus alatus. Because on the original description of Aegidium haedulus the exact number of the examined specimens was not indicated but both sexes were mentioned, we designate the same specimen as the lectotype of Aegidium haedulus, and, accordingly, establish an objective synonymy of the names., Published as part of Frolov, Andrey V., Akhmetova, Lilia A. & Vaz-de-Mello, Fernando Z., 2017, Revision of the Neotropical scarab beetle genus Aegidiellus Paulian (Coleoptera: Scarabaeidae: Orphninae) with description of two new species, pp. 1767-1779 in Journal of Natural History (J. Nat. Hist.) (J. Nat. Hist.) 51 (29 - 30) on pages 1774-1776, DOI: 10.1080/00222933.2017.1353153, http://zenodo.org/record/5182397, {"references":["Castelnau FL. 1840. Histoire Naturelle des Insectes Coleopteres. Avec une introduction renfermant L' Anatomie et la Physiologie des Animaux Articules, par M. Brulle. Paris: P. Dumenil.","Arrow GJ. 1912. Pachypodinae, Pleocominae, Aclopinae, Glaphyrinae, Ochodaeinae, Orphninae, Idiostominae, Hybosorinae, Dynamopinae, Acanthocerinae, Troginae. In: Schenkling S, editor. Coleopterorum Catalogus. Berlin: Junk, W; p. 1 - 66.","Gemminger M, Harold E. 1869. Catalogus Coleopterorum hucusque descriptorum synonymicus et systematicus. Scarabaeidae Munich: Grummi E. H. Monachia.","Preudhomme de Borre A. 1886. Descriptions de deux especes nouvelles du genre Aegidium Westwood suivies de la liste des Orphnides du Musee royal d' histoire naturelle de Belgique. Ann Soc Ent Belg. 30: 24 - 26.","Schmidt A. 1913. Coleoptera. Lamellicornia. Fam. Scarabaeidae. Subfam. Aegialinae, Chironinae, Dynamopinae, Hybosorinae, Idiostominae, Ochodaeinae, Orphninae. Bruxelles: V. Verteneuil & L. Desmet.","Vulcano MA, Pereira FS, Martinez A. 1966. Notas sobre Orphninae Neotropicos com descricao de um genero e una especie novos (Coleoptera). Papeis Dep Zool S Paulo. 18: 251 - 260.","Paulian R. 1984. Les Orphnidae Americains (Coleopteres, Scarabaeoidea). Ann Soc Ent Fr (N S). 20: 65 - 92.","Colby J. 2009. Monographic revision of the genus Aegidinus Arrow (1904) and generic phylogeny of the world Orphninae (Coleoptera: Scarabaeidae: Orphninae). Ins Mundi. 76: 1 - 41.","Frolov AV. 2012. Diagnosis, classification, and phylogenetic relationships of the orphnine scarab beetles (Coleoptera, Scarabaeidae: Orphninae). Ent Rev. 92: 782 - 797.","Westwood JO. 1845. On the lamellicorn beetles which possess exserted mandibles and labrum, and 10 - jointed antennae. Trans Ent Soc London. 4: 155 - 180.","Evenhuis NL. 2012. Francois-Louis Comte de Castelnau (1802 - 1880) and the mysterious disappearance of his original insect collection. Zootaxa. 3168: 53 - 63."]}

Taisia cornitermitis Frolov, Ocampo, Akhmetova et Vaz-de-Mello, sp. nov. (Figures 1 (a–f), 2(a–d)) c 0.5 mm d e 0.5 mm f Type locality Cuiabá municipality, Mato Grosso, Brazil. Type material Holotype, male at CEMT labelled ‘ Brazil, MT, Cuiabá, Flor do Cerrado, 15°29 ʹ 38.20’ S 56° 4 ʹ 36.40” W 6.XI.2015 А. Frolov & L. Akhmetova leg.’. Paratypes: 184 specimens. One female and six males at CEMT with the same data as the holotype; six males at CEMT with the same locality data as the holotype but collected 15 November 2015; seven males at CEMT labelled ‘ Brazil, MT, Cuiabá, Flor do Cerrado, 15°29 ʹ 38.20’ S 56°4 ʹ 36.40” W Frolov leg. 15.X.2016, termite nests and flying around’; 123 males and three females at CEMT, IAZA, BMNH, IOCRJ, IRSNB, MNHG, MNHN, MNRJ and ZIN with the same locality data but collected 27 October 2016 with a flight interception trap (FIT); 31 males at CEMT with the same locality data but collected 12–17 November 2016 by FIT; one male at CEMT with the same locality data but collected 3–12 November 2016 by FIT; four males at CEMT with the same locality data but collected 8–12 November 2016 by FIT; two males at CEMT labelled ‘ Brazil, MT, Aguas Quentes 15°53 ʹ 11”S 55°30 ʹ 44’W 25.XI.2016 M. Cupello & A. Frolov leg’. Description Holotype, male (Figure 1 (a,c–e)). Body length 4.5 mm. Colour uniformly brown to light brown, sclerites poorly melanised, semitransparent. Head (Figure 1 (a)): Frons slightly convex; surface punctate with sparse and moderately coarse punctures. Frontoclypeal suture distinctive, convex. Clypeus subtrapezoidal, anterior margin narrowly rounded, surface punctate with few punctures. Clypeal anterior margin weakly reflexed except in middle. Labrum somewhat trapezoidal, lacking medial tooth at apex. Mandibles protruding beyond labrum, with 3 outer teeth. Labium subtrapezoidal, densely setose. Maxillae with galea densely setose at apex, setae visible in dorsal view of head; maxillary palps with 4 palpomeres. Eyes relatively small, visible in dorsal view. Antennae with 9 antennomeres; antennomeres 2–6 moniliform; antennal club with 3 antennomeres; basal antennomere of club cup-shaped, capable of receiving penultimate and ultimate antennomeres. Pronotum: slightly convex, base flat in middle, pronotum 1.6 times wider than long; surface with relatively sparse, double punctation. Anterior margin feebly convex medially, base almost straight; posterior angles nearly right-angled. Lateral margins convex and coarsely serrate. Scutellum pentagonal. Apical, permanently exposed part (commonly referred to as ‘scutellum’ in the Scarabaeoidea taxonomic literature) widely rounded, surface smooth. Medial part mostly hidden under pronotum, coarsely punctate with round punctures; each puncture bears a seta. Basal part hidden under pronotum in undisturbed beetles, with two semirectangular, transverse, adjacent fossae with rugose and setose surface (Figure 2 (a)). Elytra moderately convex, with rounded apices, 1.3 times longer than wide. Elytral striae marked with regular rows of relatively coarse punctures; intervals feebly convex, minutely punctured; margins setose. Wings fully developed. Protibia with 2 large teeth and a number of smaller teeth on outer margin; dorsal surface with 2 setose, longitudinal carinae; protibial spur shorter than apical tibial tooth, curved, apex acute. Meso- and metatibia lacking transversal carina; with 2 apical, acuminate spurs. Metatibia with long, truncate process apically, analogous to the ‘enclosed corbel’ of some Curculionidae (Thompson 1992). Pro- and metatarsomeres 1–3 with feather-like setae ventrally (Figure 1 (e)). Apical tarsomeres longer than tarsomeres 1–4, claws longer than half of apical tarsomeres. Male genitalia strongly asymmetrical (Figure 1 (c,d)). Female (Figure 1 (b,f)) differs from male in having mandibles with 2 teeth; galea with sparse, short setae, not visible in dorsal view; wider pronotum and elytra, normal (not feather-like) setae on tarsomeres (Figure 1 (f)), and rounded apical processes of metatibiae. Variation Body length of males 4.0– 5.2 mm, females 4.3–4.7 mm. Except for the body size variation the examined specimens are rather similar. Distribution and habitat Taisia cornitermitis sp. nov. is known from two localities in the state of Mato Grosso, Brazil (Figure 2 (e)). Most of the type series including the holotype originate from the vicinity of Cuiabá. Another locality is near Fazenda Aguas Quentes, Santo Antônio do Leverge municipality, some 80 km east of Cuiabá. The habitat in the vicinity of Cuiabá is a small patch of Cerradão, a forest-like subtype of the Cerrado. Biology Taisia cornitermitis sp. nov. are associated with termites Cornitermes cf. bequaerti (Termitidae, Nasutitermitinae). The beetles were observed coming out of the ‘ventilation holes’ of the termite nests, and sitting on and flying around the nests (Figure 2 (b–d)). The ‘ventilation holes’ go deep around the nest (and some through the nest) and apparently join below the nest to form a ‘paraecie’, an empty space of regular thickness surrounding subterranean termite nests and subterranean parts of epigeal ones and thought to play a role in microclimate regulation and defence (Noirot and Darlington 2000). The termite nest inside the ‘paraecie’ is closed and the latter is devoid of termites. However, the ‘paraecies’ seem to be populated by various animals. Except for T. cornitermesi, whose flying activity is limited to a few weeks after the beginning of rainy season, most prominent inhabitants are crickets (Gryllidae) and whip spiders (Amblypygi) which can be found in every hole at night. Taisia cornitermitis is apparently a diurnal species, and most of its flying activity was observed in the mornings. In addition to the specimens collected by hand on the termite nests, long series were collected by FITs set near the nests. In Fazenda Aguas Quentes, the beetles were collected on the side of a termite nest. There is little doubt that this species will be found in other localities within the rather large distribution range of Cornitermes. It is worth noting that the sex ratio of the collected specimens is strongly biased: there were only four females found among the long series of over 180 males. Although the females have well developed wings, they probably do not normally leave termite nests. Etymology Latin noun in the genitive case derived from the name of the host termite genus., Published as part of Frolov, Andrey V., Ocampo, Federico C., Akhmetova, Lilia A. & Vaz-de-Mello, Fernando, 2017, A new genus and species of the termitophilous Neotropical Hybosorinae (Coleoptera: Scarabaeoidea: Hybosoridae) associated with Cornitermes (Isoptera: Termitidae) in the Cerrado ecoregion in Brazil, pp. 1759-1765 in Journal of Natural History (J. Nat. Hist.) (J. Nat. Hist.) 51 (29 - 30) on pages 1760-1764, DOI: 10.1080/00222933.2017.1353150, http://zenodo.org/record/5182369, {"references":["Thompson RT. 1992. Observations on the morphology and classification of weevils (Coleoptera, Curculionoidea) with a key to major groups. J Nat Hist. 26: 835 - 891.","Noirot C, Darlington JP. 2000. Termite nests: architecture, regulation and defence. In: Abe Y, Bignell DE, Higashi T, editors. Termites: evolution, sociality, symbioses, ecology. Dordrecht: Springer Netherlands; p. 121 - 139."]}

Paraegidium barretoi Frolov, Akhmetova & Vaz-de-Mello, sp. nov. (Figures 2 (a–h), 7) Type locality Brazil, Mato Grosso, Cotriguaçu, Fazenda São Nicolau. Type material examined Holotype, male at CEMT labelled ‘ BRASIL: Mato Grosso Cotriguaçu Fazenda São Nicolau. Mata, luz branca. 08-XII-2009 M.R. Barreto’. Paratypes. BRAZIL. Mato Grosso: two males and one female at CEMT labelled ‘ BRASIL: Mato Grosso Cotriguaçu Fazenda São Nicolau. Mata, luz negra. 10-XII-2009 M.R. Barreto’; 12 males and 23 females at CEMT, EPGC and ZIN labelled ‘ BRASIL: Mato Grosso Cotriguaçu Fazenda São Nicolau. Luz. 14.XI.2007. O. Peres Filho’; one female at CEMT labelled ‘ BRASIL: Mato Grosso Cotriguaçu Fazenda São Nicolau. Luz. 15.XII.2007. O. Peres Filho’; three females at CEMT labelled ‘ BRASIL: Mato Grosso Cotriguaçu Fazenda São Nicolau. Luz. 16. X.2007. O. Peres Filho’; six males and two females at CEMT and ZIN labelled ‘ BRASIL: Mato Grosso Cotriguaçu Fazenda São Nicolau. Luz. 29.XI.2007. O. Peres Filho’; one female at CEMT labelled ‘ BRASIL: Mato Grosso Cotriguaçu Fazenda São Nicolau. Luz. 3.I.2008. O. Peres Filho’. Diagnosis P. barretoi sp. nov. is most similar to P. costalimai but can be separated from it by the shape of the parameres: base of parameres wider in lateral view and apices wider in dorsal view. Also, all examined specimens of P. costalimai have distinct small lateral teeth on parameres while in P. barretoi sp. nov. these teeth are indistinct. Description Holotype, male (Figure 1 (a)). Body length 8.1 mm. Colour uniformly brown. Anterior frontoclypeal process relatively long, narrower than head, widely sinuate medially, with symmetric, narrowly rounded lateral lobes. Pronotum almost as wide as elytra, deeply excavated in middle. Lateral pronotal processes depressed, somewhat ridge-shaped, their hind margins with carinae reaching base of pronotum. Pronotum coarsely punctate to rugose except for very apices of lateral pronotal processes. Punctation of anterior part of pronotum sparser than on disc and especially on lateral sides. Aedeagus with parameres relatively long: ratio phallobase width/paramere width 2.2 (Figure 2 (f)). Base of parameres relatively wide in lateral view. Endophallus with a group of spinules (Figure 2 (h)). Apices wide in dorsal view and acute in lateral view (Figure 2 (g)). Female differs from male in having a distinct protibial spur, relatively smaller pronotum without processes and excavations, and in the absence of the frontoclypeal process (Figure 2 (b)). Variation Body length of the paratypes varies from 6.0 to 8.0 mm (males) and from 6.0 to 7.0 (females). Most of the paratypes have less developed anterior frontoclypeal and pronotal processes (Figure 2 (c–e)) than the holotype. Distribution The species is known from one locality in Amazonian forest region in northern Mato Grosso, Brazil (Figure 7). Etymology The species is named after Prof. Dr Marliton Rocha Barreto, collector of a part of the type series., Published as part of Frolov, Andrey V., Akhmetova, Lilia A. & Vaz-de-Mello, Fernando Z., 2017, Revision of the South American genus Paraegidium Vulcano et al. (Coleoptera: Scarabaeidae: Orphninae), pp. 995-1014 in Journal of Natural History (J. Nat. Hist.) (J. Nat. Hist.) 51 (17 - 18) on pages 1004-1006, DOI: 10.1080/00222933.2017.1326640, http://zenodo.org/record/5181243

Paraegidium oliveirai Frolov, Akhmetova & Vaz-de-Mello, sp. nov. (Figures 6 (a–f), 7) Type locality Brazil, Goiás, Morrinhos, Jatobá Centenário ecological park. Type material examined Holotype, male at CEMT labelled ‘ BRASIL: Goiás Morrinhos Pq. Eco. Jatoba 17°43′26.02″S 49°07′29.83″W 803 m Col. Manual X/ XI 2010 ’. Paratypes: BRAZIL. Goiás: two males at CEMT and ZIN labelled ‘ BRASIL: Goiás Morrinhos Pq. Eco. Jatoba 17°43′26.02″S 49°07′ 29.83″W 803 m Col. Manual X/ XI 2010 ’. Minas Gerais: one male at CEMT labelled ‘ BRASIL: Minas Gerais, Santana do Riacho, PARNA, Serra do Cipó, 18.xi.2015, J Bonanomi’. Rio de Janeiro: one male at CMNC labelled ‘ BRASIL Eº Guanabara Rio de Janeiro CORCOVADO Alvarenga-leg. Coll. Martínez. Nov:956/H. & A. HOWDEN COLLECTION ex. A. Martinez coll./PARATIPO/ Paraegidium castalimai ♂ gen. et sp. nov. Vulcano, Pereira y A. Martinezdet. 1965’. Diagnosis P. oliveirai sp. nov. can be easily separated from the other species of the genus by having short parameres with somewhat rounded (in lateral view, Figure 6 (d)) apices and widely bifurcated lobes of the anterior frontoclypeal process (Figure 6 (a)). Description Holotype, male (Figure 6 (a, d–f)). Body length 7.3 mm. Colour uniformly dark brown. Anterior frontoclypeal process long and wide, as wide as head, widely sinuate medially, with asymmetric lateral lobes; the lobes distinctly excavated apically, in lateral view they have lateral angles directed caudally. Pronotum slightly narrower than elytra, deeply excavated in middle. Lateral pronotal processes almost rounded, their hind margins not carinate. Pronotum coarsely punctate to rugose except for apices of lateral pronotal processes and a narrow area near anterior margin. Aedeagus with short parameres: ratio phallobase width/paramere width 2.7 (Figure 6 (d)). Apices of parameres rounded in lateral view and dorsal view (Figure 6 (d, e)). Endophallus without a group of spinules (Figure 6 (f)). Female unknown. Variation Body length of the paratypes varies from 6.8 to 7.5 mm. The paratypes have less developed anterior frontoclypeal processes (Figure 6 (b, c)) than the holotype. Distribution The species is known from two localities in the Brazilian states of Goiás and Minas Gerais (Figure 7). The record from Corcovado Mountain in Rio de Janeiro is doubtful and not therefore mapped. Etymology The new species is named after Dr Charles Martins de Oliveira, from Embrapa Cerrados in Brasília, who obtained the specimens from Morrinhos., Published as part of Frolov, Andrey V., Akhmetova, Lilia A. & Vaz-de-Mello, Fernando Z., 2017, Revision of the South American genus Paraegidium Vulcano et al. (Coleoptera: Scarabaeidae: Orphninae), pp. 995-1014 in Journal of Natural History (J. Nat. Hist.) (J. Nat. Hist.) 51 (17 - 18) on pages 1012-1013, DOI: 10.1080/00222933.2017.1326640, http://zenodo.org/record/5181243

Paraegidium howdeni Frolov, Akhmetova & Vaz-de-Mello, sp. nov. (Figures 5 (a–h), 7) Type locality Peru, Madre de Dios, Rio Palma Real Grande. Type material examined Holotype. Male at CMNC labelled ‘ PERU, Madre de Dios: Rio Palma Real Grande, Limon Camp 12°32 ʹ 20S 68°51 ʹ 40W Flight intercept trap, 220 m T. Larsen, X-1999 /H. & A. HOWDEN COLLECTION Ottawa, Canada’. Paratypes. PERU. Madre de Dios: three males at CMNC with the same label as the holotypes. BRAZIL. Mato Grosso: one male at CEMT labelled ‘V.VERA, M. Grosso, Brasil, X.1973 M. Alvarenga / Lon 55 36 ʹ W 12 46 ʹ S /coleção M. Alvarenga’. Pará: one male at CMNC labelled ‘ BRASIL Eº PARA Itaituba Maller-leg. Coll. Martinez Nov. 963/H. & A. HOWDEN COLLECTION ex. A. Martinez coll.’; one male and one female at CEMT labelled ‘ BRASIL: PA. Redenção Pinkaiti – AIK 31.X.1998 07°46 ʹ S 51°58 ʹ W P.Y. Scheffler’; two males and four females at CEMT and EPGC labelled ‘ BRASIL: PA. Redenção Pinkaiti – AIK X.1999 07°46 ʹ S 51°58 ʹ W P.Y. Scheffler’; one female at CEMT labelled ‘ BRASIL: PA. Redenção Pinkaiti – AIK 07°46 ʹ S 51°58 ʹ W F.T.T. XI.1998 P&T Scheffler’. Rondônia: one male and one female at CEMT labelled ‘ BRASIL, Rondonia: Porto Velho BR 364, km 20–30. 24. X.1980 Eq. INPA’; one male at MNRJ labelled ‘BRASIL-Rondônia Vilhena XI.1987 O. Roppa, P. Magno / Museu Nacional de Rio de Janeiro / Paraegidium sp. Vaz-de-Mello det 2003’. Additional material examined PERU. Cusco: La Convención, Echarate, Timpia, 429 m, 20 October 2009, M. Alvarado and E. Rázuri leg., one female (UNMSM); La Convención, Echarate, Timpia, 371 m, 22 October 2009, M. Alvarado and E. Rázuri leg., one female (UNMSM); La Convención, Echarate, Timpia, 442 m, 16 October 2009, M. Alvarado and E. Rázuri leg., one female (UNMSM). BRAZIL. Roraima: Pacaraima, September 1996, Ribeiro and Vaz-de-Mello leg., one male (CEMT). Diagnosis In the shape of the anterior frontoclypeal processes, larger males of P. howdeni sp. nov. are similar to P. monneorum sp. nov., while smaller males are similar to the smaller males of P. oliveirai sp. nov. From both species P. howdeni sp. nov. differs in the shape and relative length of the parameres (Figure 5 (f, g)). Description Holotype, male (Figure 5 (a, f, g)). Body length 7.6 mm. Colour uniformly brown. Anterior frontoclypeal process long and wide, slightly narrower than head, with angulate sinuation medially, with slightly asymmetric, apically truncated lateral lobes. Pronotum narrower than elytra, deeply excavated in middle. Lateral pronotal processes depressed, somewhat ridge-shaped, their hind margins with carinae reaching base of pronotum. Pronotum coarsely punctate to rugose except for very apices of lateral pronotal processes and a narrow area near anterior margin. Aedeagus with short parameres: ratio phallobase width/paramere width 2.5. Apices of parameres acute in lateral view and wide in dorsal view (Figure 5 (f, g)). Endophallus with a group of spinules (Figure 5 (h)). Female (Figure 5 (b)) differs from male in having a distinct protibial spur, relatively smaller pronotum without processes and excavations, and in the absence of the frontoclypeal process. Variation Body length of the paratypes varies from 6.3 to 7.5 mm (males) and from 7.0 to 7.5 mm (females). The paratypes have less developed anterior frontoclypeal and pronotal processes (Figure 5 (c–e)) than the holotype. Distribution The species as treated here has a wide range in central Brazil and eastern Peru (Figure 7). However, the specimens from the two outermost localities, Echarate (Peruvian Yungas, the westernmost locality for the genus) and Pacaraima (Guiana Shield, northernmost locality for the genus), are tentatively identified as P. howdeni sp. nov. and thus are not included in the type series. They may belong to other, yet undescribed species, however additional material is needed for more reliable identification. Etymology The species is named after scarabaeoidologist Henry F. Howden (1925–2014)., Published as part of Frolov, Andrey V., Akhmetova, Lilia A. & Vaz-de-Mello, Fernando Z., 2017, Revision of the South American genus Paraegidium Vulcano et al. (Coleoptera: Scarabaeidae: Orphninae), pp. 995-1014 in Journal of Natural History (J. Nat. Hist.) (J. Nat. Hist.) 51 (17 - 18) on pages 1010-1012, DOI: 10.1080/00222933.2017.1326640, http://zenodo.org/record/5181243