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Oxythyrea dulcis Reitter, 1899 (Figs. 2C–D, 3B, 3D, 4D–F) Oxythyrea dulcis Reitter, 1899: 48 (original description). Oxythyrea dulcis: Schenkling 1921: 200 (catalogue); Winkler 1929: 1121 (catalogue); Petrovitz 1958: 96 (distribution, biology), 1959: 129 (distribution); Medvedev 1964: 115 (monograph, key); Tauzin 2000: 85 (distribution); Smetana 2006: 298 (catalogue); Keith & Makris 2008: 27 (distribution); Bezděk 2016: 389 (catalogue); Vondráček et al. 2018: 417 (larval morphology, biology); Vondráček et al. 2023: 2 (diversity, habitat preferences, conservation); Schoolmeesters 2023 (online catalogue). Oxythyrea dulcis dulcis: Mikšić 1978: 69 (diagnosis, distribution); Sabatinelli 1981: 56 (review, key, biology); Mikšić 1982a: 67 (key distribution); Mikšić 1982b: 36 (monograph, key); Baraud 1992: 820 (monograph, key); Krajcik 1998: 69 (catalogue); Keith 2001: 95 (distribution, biology); Tauzin 2015: 118 (review, distribution). Oxythyrea borchmanni Schein, 1958: 93 (original description). Type locality: “Asteri und Elos bei Skala, Marathea bei Gythion [=Asteri, ca 36°49’54”N 22°41’55”E and Elos, ca 36°49’49”N 22°41’52”E nr. Skala; Marathea, ca 36°44’03”N 22°29’21”E nr. Gythion”; synonymized by Mikšić 1982a: 67). Oxythyrea goetzi Muche, 1962: 33 (original description). Type locality: “In de Nähe des Ortes Manavgat am Golf von Anatayla (Adalia) [= Turkey, Manavgat env., ca 36°45’54”N 31°23’57”E]”; synonymized by Mikšić 1982a: 67. Type locality. “ Olymp [= Greece, Mount Olympus, ca 40°05’19”N 22°21’32”E]”. Published records. Greece: Schenkling (1921): “Olymp”; Winkler (1929): “Olymp”; Schein (1958): “Asteri and Elos nr. Skala, Marathea nr. Gythion”; Petrovitz (1958): “Gythion, Kreta, Olymp, Rhodos (Lindos)”; Petrovitz (1959): “Rhodos”; Medvedev (1964): “ Peloponnesos, Rhodos”; Sabatinelli (1981): “ Peloponeso, Creta, Rodi”; Mikšić (1982a): “Olymp, Asteri bei Skala-Elos, Gythion-Lakonia, Kreta, Rhodos-Lindos”; Mikšić (1982b): “Asteri bei Skala-Elos; Peloponesos: Gythion, Laconia; Kreta; Rhodos: Lindos”; Baraud (1992): “Grèce (Olympe, loc type); Crète; Rhodes”; Keith & Makris (2008): “Andros Island, Chora”; Krajcik (1998): “Olymp”; Tauzin (2000): “Mont Olympe, Peloponnese (Elos, Gythra [= Gythion], Crète, Rhodes”; Keith (2001): “Mont Olympe, Litohoron, Rhodes, Crète ”; Smetana (2006): “Greece”; Tauzin (2015): “Gréce continental, de l’Ile de Rhodes, de l’Ile de Crete, de l’Ile d’Andros”; Bezděk (2016): “Greece”; Vondráček et al. (2018): “ Thessaly, Mount Ossa, village Stomio, N°39.867, E°22.733; Stomio, Beach, at sea level, N°39.8761111, E°22.7313889”; Byk et al. (2022): “Greece”; Vondráček et al. (2023): “Asteri bei Skala; Elos; Marathea bei Gytheion; West Kreta; Nordküste, Georgioupoli; Saloniki; Stomio, beach, ca sea level; Limnos isl., Gomati sand dunes, sea shore, ca 5 m; Crete, Phalasarna (seashore), 10m; Creta, Kasteh (Kissamu), 10 m; Creta, Khania, 20 m; Rhodos, Lindos”; Schoolmeesters (2023): “Greece”. Asian part of Turkey: Petrovitz (1958): “ Mersina ”; Muche (1962): “Manavgat”; Medvedev (1964): “ Mersin ”; Sabatinelli (1981): “Turchia del villaiet di Mersin ”; Mikšić (1982a): “Kleinasien”; Mikšić (1982b): “Manavgat, Erdemli”; Baraud (1992): “Anatolie”; Tauzin (2000): “Ephèse, Selçuk (Izmir)”; Keith (2001): “ Mersin, Izmir; Izmir: plage de Pamuçak, plage de Gümüldür, Antalya: plage de Patara)”; Smetana (2006): “Asian part of Turkey”; Tauzin (2015): “Turquie d’Anatolie”; Bezděk (2016): “Asian part of Turkey”; Byk et al. (2022): “Turkey”; Vondráček et al. (2023): “Erdemli; Izmir, Ozbere”; Schoolmeesters (2023): “Turkey”. Syria [?]: Mikšić (1982b): “?Syrien”. Type material examined. Oxythyrea borchmanni Schein, 1958: Holotypus (♁) (ZFMK), “ Griechenland 1956 | Asteri bei Skala—| Elos 23.–25.4.|| Fr. Borchmann leg. | Eingang Nr. 8, 1956 [p] || Paratypus [p] | Oxyth. dulcis Rtt. | v. Borchmanni n.v. [hw] | H. Schein 1955 [p.] || Oxythyrea | dulcis | Reitter, 1899 | David Král det. 2023”; Paratypus (♁) (ZFMK),“ Griechenland 1956 | Marathea bei | Gythion 21.–22.4 || Fr. Borchmann leg. | Eingang Nr. 8,1956 || MUSEUM KOENIG BONN [p] || Paratypus [p] | Oxyth. dulcis R. | v. Borchmanni n. v. [hw] | H. Schein 1955 [p]” (see Fig. 7). Additional material examined. Greece: 1 ♁ (NMPC), Crete, Prevelikloster bei Südküste, 400 m, 26.iv.1992, auf Blüten, J. Burtscher lgt.; 1 ♁ (NMPC), Crete, Agios Nikolaos, 23.iv.1971, Wewalka lgt.; 1 ♀ (NMPC), Macedonia, Asprovalta env., Nea Kerdilia, litoral-seaside, 11.v.2011, Z. Malinka lgt.; 1 ♁, 5 ♀♀ (NMPC), Crete, Phalasarna (seashore), 35°30ʼ18”N 23°34ʼ34”E, ca. 10m, 18.–19.iv.2018, L. Hrůzová, D. Král & D. Sommer lgt.; 1 ♁ (NMPC), N Crete, Petras riv., 23.v.2002, Z. Kletečka lgt.; 2 ♁♁, 1 ♀ (NMPC), Thessaloniki pr., Aspróvalta, 12.v.2001, D. Král lgt.; 2 ♁♁ (NMPC), Limnos, Gomati sand dunes, 39°59.3’N 25°09.1’E, ca. 5 m, sea shore, 24.iv.2007, J. Hájek lgt.; 4 ♁♁, 5 ♀♀ (NMPC), Thessaly, Mount Ossa, village Stomio, N °39.867, E°22.733, Stomio, beach, at sea level, 13.–20.vi.2014, H. Podskalská & P. Šípek lgt.; 12 ♁♁, 52 ♀♀ (CUPC, LHCP, NMPC, DSCP, SJCP), Stomio beach, 39°52ʼ34”N 22°43ʼ53”E, ca. sea level, O. Kouklík, D. Král, P. Šípek & D. Vondráček lgt.; 4 ♁♁, 6 ♀♀ (ZFMK), Asteri bei Skala—Elos, 23.–25.iv.1956, Fr. Borchmann lgt.; 1 ♀ (ZFMK), Marathea bei Gythion, 21.–22.iv.1956, Fr. Borchmann lgt.; 1 ♀ (ZFMK), Westkreta, Ostküste, Georgiopouli, 28.iv.1992, E. Rössner lgt.; 1 ♁ (ZFMK), Saloniki, 24.iv.1968, H. Roer lgt.; 2 unsexed spec. (OHCB), Githio Strand, 21.iv.2004, D. Keith lgt.; 2 unsexed spec. (OHCB), Crete, W Chania, Dramia Strand, 14.v.2014, Heise lgt. Asian part of Turkey: 4 ♁♁, 4 ♀♀ (ZFMK), Anatolie, Erdemli, Petrovitz & Ressl lgt. Diagnosis. Body (Figs. 2C–D, 3B) remarkable robust, enlarged, and rounded; dorsal surface sparsely but distinctly setaceous; dorsal surface totally black without metallic lustre. Pronotum (Figs. 2C, 3B) remarkably setaceous with 4–6 (rarely only with basal pair) whitish spots in two longitudinal rows; lateral margin as a rule with narrow whitish stripe, in some specimens this stripe broken into three irregularly shaped spots. Scutellum surface (Fig. 3D) completely smooth, very rarely finely microscultured basally. Elytra with several rounded whitish spots, more or less equal in size, not connected to each other (Figs. 2C, 3B). Pygidium with six whitish spots. Male with triangular whitish spots on the middle of visible abdominal ventrite 1 and 2 and rarely with additional one whitish spot on visible abdominal ventrites 3 and 4 (Fig. 2D). These spots absent in females. Abdomen ventrally on each side with three short whitish lines and more distally with two longer whitish lines in male (Fig. 2D), one such longer line in female. Lamina exterior of parameres relatively stout in lateral aspect (Fig. 4D–F); lamina interior of parameres relatively wide, tip at less sharp angle in dorsofrontal aspect (Fig. 4D–F). Distribution. Greece (Aegean coast of Greece, includig islands Andros, Crete, Limnos, and Rhodes), Asian part of Turkey and an unverified occurrence in “ Syria ” without precise data based on Mikšić (1982b). For details see Fig. 6., Published as part of Král, David, Šípek, Petr, Sommer, David & Vondráček, Dominik, 2023, Resurrection of Oxythyrea abigailoides Mikšić, 1978 (Coleoptera: Scarabaeidae: Cetoniinae) based on new morphological, morphometrical and molecular data, pp. 561-574 in Zootaxa 5301 (5) on pages 566-570, DOI: 10.11646/zootaxa.5301.5.3, http://zenodo.org/record/8036225, {"references":["Reitter, E. (1899) Bestimmungs-Tabelle der Melolonthidae aus der europaischen Fauna und den angrenzenden Landern, enthaltend die Gruppen der Dynastini, Euchirini, Pachypodini, Cetonini, Valgini und Trichiini. Verhandlungen des Naturforschenden Vereins in Brunn, 37, 21 - 111. [1898]","Schenkling, S. (1921) Scarabaeidae: Cetoniinae. In: Junk, W. & Schenkling, S. (Eds.), Coleopterorum Catalogus. Pars 72. W. Junk Publishers, The Hague, 47 pp.","Winkler, A. (1929) s. n. In: Lamellicornia. Catalogus Coleopterorum regionis palaearcticae, Pars 9. A. Winkler, Wien, pp. 1009 - 1136. [1924 - 1929]","Petrovitz, R. (1958) Oxythyrea dulcis Reitter, eine strandbewohnende Cetonide. Deutsche Entomologische Zeitschrift, Neue Folge, 5, 96 - 97.","Medvedev, S. I. (1964) Plastinchatousyie (Scarabaeidae). Podsem. Cetoniinae, Valginae. Fauna SSSR. Zhestkokrylye. Tom X. Vyp. 5. [Lamellicornia (Scarabaeidae). Subfam. Cetoniinae, Valginae. Fauna SSSR. Beetles. Vol. X. Part 5]. Nauka, Moskva, Leningrad, 374 + 2 (unpaginated) pp. [in Russian]","Tauzin, P. (2000) Coleopteres Scarabaeoidea de Turquie: Premiere note. Le Coleopteriste, 39, 81 - 86.","Smetana, A. (2006) Subfamily Cetoniinae. In: Lobl, I. & Smetana, A. (Eds.), Catalogue of Palaearctic Coleoptera. Vol. 3. Scarabaeoidea - Scirtoidea - Dasciloidea - Buprestoidea - Byrrhoidea. Brill E. J., Leiden, pp. 283 - 313.","Keith, D. & Makris, C. (2008) Confirmation of Oxythyrea abigail Reiche & Saulcy from Cyprus Island and record of O. dulcis from Andros Island (Coleoptera: Scarabaeidae: Cetoniinae). Kogane, 9, 27 - 28.","Bezdek, A. (2016) Subfamily Cetoniinae. In: Lobl, I. & Lobl, D. (Eds.), Catalogue of Palaearctic Coleoptera. Vol. 3. Scarabaeoidea - Scirtoidea - Dasciloidea - Buprestoidea - Byrrhoidea. Revised and Updated edition. Brill E. J., Leiden, pp. 367 - 412.","Vondracek, D., Hadjiconstantis, M. & Sipek, P. (2018) Immature stages of the genus Oxythyrea (Coleoptera: Scarabaeidae: Cetoniinae) with a key to third instar larvae, and notes on the biology of the genus. Zootaxa, 4486 (4), 401 - 434. https: // doi. org / 10.11646 / zootaxa. 4486.4.1","Vondracek, D., Kral, D. & Sipek, P. (2023) Cryptic diversity of Oxythyrea flower chafers and its implication for conservation of non-forest biotopes in the Balkans. Insect Conservation and Diversity, 16, 133 - 146. https: // doi. org / 10.1111 / icad. 12612","Schoolmeesters, P. (2023) World Scarabaeidae Database. In: Banki, O., Roskov, Y., Doring, M., Ower, G., Vandepitte, L., Hobern, D., Remsen, D., Schalk, P., DeWalt, R. E., Keping, M., Miller, J., Orrell, T., Aalbu, R., Abbott, J., Adlard, R., Adriaenssens, E. M., Aedo, C., Aescht, E., Akkari, N. et al., Catalogue of Life Checklist. Version 6 February 2023. https: // doi. org / 10.48580 / dfrq- 38 g","Miksic, R. (1978) Eine weiterer Beitrag zur Kenntnis der jugoslawischen Oxythyrea - Arten (Coleoptera, Scarabaridae, Cetoniinae). Acta Entomologica Jugoslavica, 14, 69 - 71.","Sabatinelli, G. (1981) Le Oxythyrea Muls. del Mediterraneo: Studi morfologici sistematici (Coleoptera, Scarabaeoidea). Fragmenta Entomologica, 16, 45 - 60.","Miksic, R. (1982 a) Eine vorlaufige Ubersicht und Bestimmungstabelle der Cetoniinae der Balkanlander (Coleoptera, Lamellicornia, Scarabaeidae). Entomologische Abhandlungen des Staatliches Museum fur Tierkunde in Dresden, 45, 65 - 89.","Miksic, R. (1982 b) Monographie der Cetoniinae der Palaarktischen und Orientalischen Region. Coleoptera, Lamellicornia. Band 3. Systematischer Teil. Cetoniini I Teil. Forstinstitut in Sarajevo, Sarajevo, 530 pp., 14 pls.","Baraud, J. (1992) Coleopteres Scarabaeoidea d'Europe. Faune de France 78. Federation Francaise des Societes de Sciences Naturelles & Societe Linneenne de Lyon, Lyon, 856 pp. https: // doi. org / 10.3406 / linly. 1992.11007","Krajcik, M. (1998) Cetoniidae of the World. Catalogue - Part I. (Coleoptera: Cetoniidae). Typos Studio Most, Most, 132 pp.","Keith, D. (2001) Contribution a la connaissance des Scarabaeoidea du Proche et du Moyen Orient (7 e note). Observations sur Oxythyrea dulcis Reitter, 1898 (Col. Cetoniidae). Cetoniimania, 1, 95 - 97.","Tauzin, P. (2015) Liste synonymique des especes palearctiques du genre Oxythyrea Mulsant, 1842 et precision sur Oxythyrea albopicta (Motschulsky, 1845) (Coleoptera, Cetoniinae, Leucocelina). Le Coleopteriste, 18, 114 - 124.","Schein, H. (1958) Cetoniiden aus der Sammelreise des Museums Alexander Koenig in Bonn nach Griechenland 1956 (Coleoptera). Deutsche Entomologische Zeitschrift, Neue Folge, 5, 91 - 95.","Muche, W. H. (1962) Oxythyrea gotzi nov. spec. (Coleopt., Cetoninae). Reichenbachia, 1, 33 - 35.","Petrovitz, R. (1959) Beitrag zur Scarabaeidae-Fauna der Insel Rhodos. Bolletino del Laboratorio di Enomologia Agraria \" Filippo Silvestri \" Portici, 17, 124 - 130.","Byk, A., Bidas, M., Gazurek, T., Marczak, D., Minkina, L. & Tylkowski, S. (2022) New data on the occurrence of Sacarabaeoid beetles (Coleoptera: Scarabaeoidea) in Montenegro. Insects, 13, 352. https: // doi. org / 10.3390 / insects 13040352"]}

Oxythyrea abigailoides Mikšić, 1978, resurrected species (Figs. 2A–B, 3A, 3C, 4A–C) Oxythyrea dulcis abigailoides Mikšić, 1978: 69 (original description). Oxythyrea dulcis abigailoides: Sabatinelli 1981: 56 (review, key, biology); Mikšić 1982a: 67 (key, distribution), 1982b: 38 (monograph, key); Baraud 1992: 820 (monograph, key); Keith 2001: 95 (distribution); Nikčević 2010: 46 (distribution, conservation); Tauzin 2015: 118 (review, distribution). Oxythyrea abigail Reiche & Saulcy, 1856: Mikšić 1977: 41 (key, distribution, biology). Oxythyrea abigailoides (as a junior subjective synonym to O. dulcis): Krajcik 1998: 70 (catalogue); Smetana 2006: 298 (catalogue); Bezděk 2016: 389 (catalogue); Vondráček et al. 2018: figs. 15f–h (habitat preferences); Byk et al. 2022: 13 (distribution); Vondráček et al. 2023: 2 (diversity, habitat preferences, conservation); Schoolmeesters 2023 (online catalogue). Type locality. “Vom “ Grossen Sandstrand ” bei Ulcinj an der Küste der Crna Gora [= Montenegro, Ulcinj env., Velika Plaža beach, ca 41°53’55”N 19°17’47”E]”. Published records. Montenegro: Mikšić (1977): “Auf dem “Grossen Sandstrand” gegen 5 km südöstlich von Ulcinj im Küstenland der Crna Gora ”; Sabatinelli (1981): “ Montenegro ”; Mikšić (1982a): “Grosser Sandstrand bei Ulcinj, Crna Gora ”; Mikšić (1982b): “Grosser Sandstrand bei Ulcinj, Crna Gora ”; Baraud (1992): “littoral de Crna Gora (Ulcinj, loc. Type)”; Keith (2001): “ Monténégro ”; Krajcik (1998): “ Crna Gora ”; Smetana (2006): “ Yugoslavia (Crna Gora)”; Tauzin (2015): “ Crna Gora (Monténégro)”; Bezděk (2016): “ Montenegro ”; Vondráček et al. (2018): “ Montenegro ”; Vondráček et al. (2023): “Ulcinj Municip., central part of Velika Plaža near Ulcinj and Doni Štoj, ca sea level”; Byk et al. (2022): “Bojana at Ulcinj”; Schoolmeesters (2023): “ Montenegro ”. Greece: Vondráček et al. (2018): “ Greece: Peloponnese, Phalasarna, Kaiafa”; Vondráček et al. (2023): “NW Zacharo, Kaiafa lake (seashore), 4 m; W of Gialova, Gialova lagoon (sea shore), 4m ”. Type material examined. Montenegro: Holotype (♁) (CNHM) “ MONTENEGRO | ULCINJ | VELIKA PLAŽA | MIKŠIĆ 2. V.[19]77 [hw] || HOLOTYPUS [red, p] || OXYTHYREA | DULCIS RTT. | S. ABIGAILOIDES NOV. [blue, hw, Mikšić’s hand] | | Miksic det., 19 [p] 79 [blue, hw, Mikšić’s hand] | 600:ZAG;ZEC1 | 1590. | coll. Mikšić [p]” (see Fig. 5). Allotype ♀ and 27 unsexed paratypes (CNHM), same data as in holotype. Additional material examined. Montenegro: 5 ♁♁, 2 ♀♀ (CUPC, NMPC), Ulcinj Municipality, central part of Velika Plaza near Ulcinj and Doni Štoj, 41°53’37.53”N 19°17’36.87”E, ca. sea level, 17.–18.v.2016, J. Kadlec, P. Šípek & D. Vondráček lgt. Greece: 18 ♁♁, 8 ♀♀ (CUPC, DSCP, LHCP, NMPC, SJCP), NW of Zacharo, Kaiafa lake (seashore), 37°29’59”N 21°36’20”E, 4 m, 15.iv.2018, L. Hrůzová, D. Král & D. Sommer lgt.; 34 unsexed spec. (ZSMC, ex. coll. Hillert), the same data but O. Hillert lgt.; 3 ♁♁ (NMPC), W of Gialova, Gialova lagoon (seashore), 36°57’25”N 21°40’21”E, 10 m, 15.iv.2018, L. Hrůzová, D. Král & D. Sommer lgt. Diagnosis. Body (Figs. 2A–B, 3A) remarkable robust, enlarged, and rounded; dorsal surface sparsely but distinctly setaceous; dorsal surface totally black without metallic lustre. Pronotum (Figs. 2A, 3A) remarkably setaceous with 4–6 (rarely only with basal pair) whitish spots in two longitudinal rows; lateral margin as a rule with three distinctly separated spots from each other on each side, rarely with interrupted white lines consisting of three semicontiguous spots. Scutellum surface (Fig. 3C) finely shallowly microsculptured. Elytra with several rounded whitish spots, more or less equal in size, not connected to each other (Figs. 2A, 3A). Pygidium with six whitish spots. Male with triangular whitish spots at middle of visible abdominal ventrite 1 and 2 and rarely with additional one whitish spot on visible abdominal ventrites 3 and 4 (Fig. 2B). These spots absent in females.Abdomen ventrally on each side with three short whitish lines and more distally with two longer whitish lines in male (Fig. 2B), one such longer line in female. Lamina exterior of parameres relatively tinny in lateral aspect (Fig. 4A–C); lamina interior relatively narrow, tip at sharper angle in dorsofrontal aspect (Fig. 4A–C). Differential diagnosis. Based on our examination of adult beetles a morphological differential diagnosis of O. abigailoides, resurrected species, from O. dulcis is summarized below in Table 1. Distribution. Greece (Ionian coast of Peloponnesos) and Montenegro. For details see Fig. 6., Published as part of Král, David, Šípek, Petr, Sommer, David & Vondráček, Dominik, 2023, Resurrection of Oxythyrea abigailoides Mikšić, 1978 (Coleoptera: Scarabaeidae: Cetoniinae) based on new morphological, morphometrical and molecular data, pp. 561-574 in Zootaxa 5301 (5) on pages 563-566, DOI: 10.11646/zootaxa.5301.5.3, http://zenodo.org/record/8036225, {"references":["Miksic, R. (1978) Eine weiterer Beitrag zur Kenntnis der jugoslawischen Oxythyrea - Arten (Coleoptera, Scarabaridae, Cetoniinae). Acta Entomologica Jugoslavica, 14, 69 - 71.","Sabatinelli, G. (1981) Le Oxythyrea Muls. del Mediterraneo: Studi morfologici sistematici (Coleoptera, Scarabaeoidea). Fragmenta Entomologica, 16, 45 - 60.","Miksic, R. (1982 a) Eine vorlaufige Ubersicht und Bestimmungstabelle der Cetoniinae der Balkanlander (Coleoptera, Lamellicornia, Scarabaeidae). Entomologische Abhandlungen des Staatliches Museum fur Tierkunde in Dresden, 45, 65 - 89.","Baraud, J. (1992) Coleopteres Scarabaeoidea d'Europe. Faune de France 78. Federation Francaise des Societes de Sciences Naturelles & Societe Linneenne de Lyon, Lyon, 856 pp. https: // doi. org / 10.3406 / linly. 1992.11007","Keith, D. (2001) Contribution a la connaissance des Scarabaeoidea du Proche et du Moyen Orient (7 e note). Observations sur Oxythyrea dulcis Reitter, 1898 (Col. Cetoniidae). Cetoniimania, 1, 95 - 97.","Nikcevic, J. (2010) Some ecological data of Coleoptera in a beach dunes system of long Ulcinj beach in Montenegro. Natura Montenegrina, 10, 41 - 48.","Tauzin, P. (2015) Liste synonymique des especes palearctiques du genre Oxythyrea Mulsant, 1842 et precision sur Oxythyrea albopicta (Motschulsky, 1845) (Coleoptera, Cetoniinae, Leucocelina). Le Coleopteriste, 18, 114 - 124.","Miksic, R. (1977) Oxythyrea abigail Reiche - eine fur Europa und Jugoslawien neue Cetoniinae-Art. Acta Entomologica Jugoslavica, 13, 41 - 44.","Krajcik, M. (1998) Cetoniidae of the World. Catalogue - Part I. (Coleoptera: Cetoniidae). Typos Studio Most, Most, 132 pp.","Smetana, A. (2006) Subfamily Cetoniinae. In: Lobl, I. & Smetana, A. (Eds.), Catalogue of Palaearctic Coleoptera. Vol. 3. Scarabaeoidea - Scirtoidea - Dasciloidea - Buprestoidea - Byrrhoidea. Brill E. J., Leiden, pp. 283 - 313.","Bezdek, A. (2016) Subfamily Cetoniinae. In: Lobl, I. & Lobl, D. (Eds.), Catalogue of Palaearctic Coleoptera. Vol. 3. Scarabaeoidea - Scirtoidea - Dasciloidea - Buprestoidea - Byrrhoidea. Revised and Updated edition. Brill E. J., Leiden, pp. 367 - 412.","Vondracek, D., Hadjiconstantis, M. & Sipek, P. (2018) Immature stages of the genus Oxythyrea (Coleoptera: Scarabaeidae: Cetoniinae) with a key to third instar larvae, and notes on the biology of the genus. Zootaxa, 4486 (4), 401 - 434. https: // doi. org / 10.11646 / zootaxa. 4486.4.1","Byk, A., Bidas, M., Gazurek, T., Marczak, D., Minkina, L. & Tylkowski, S. (2022) New data on the occurrence of Sacarabaeoid beetles (Coleoptera: Scarabaeoidea) in Montenegro. Insects, 13, 352. https: // doi. org / 10.3390 / insects 13040352","Vondracek, D., Kral, D. & Sipek, P. (2023) Cryptic diversity of Oxythyrea flower chafers and its implication for conservation of non-forest biotopes in the Balkans. Insect Conservation and Diversity, 16, 133 - 146. https: // doi. org / 10.1111 / icad. 12612","Schoolmeesters, P. (2023) World Scarabaeidae Database. In: Banki, O., Roskov, Y., Doring, M., Ower, G., Vandepitte, L., Hobern, D., Remsen, D., Schalk, P., DeWalt, R. E., Keping, M., Miller, J., Orrell, T., Aalbu, R., Abbott, J., Adlard, R., Adriaenssens, E. M., Aedo, C., Aescht, E., Akkari, N. et al., Catalogue of Life Checklist. Version 6 February 2023. https: // doi. org / 10.48580 / dfrq- 38 g","Miksic, R. (1982 b) Monographie der Cetoniinae der Palaarktischen und Orientalischen Region. Coleoptera, Lamellicornia. Band 3. Systematischer Teil. Cetoniini I Teil. Forstinstitut in Sarajevo, Sarajevo, 530 pp., 14 pls.","Reitter, E. (1899) Bestimmungs-Tabelle der Melolonthidae aus der europaischen Fauna und den angrenzenden Landern, enthaltend die Gruppen der Dynastini, Euchirini, Pachypodini, Cetonini, Valgini und Trichiini. Verhandlungen des Naturforschenden Vereins in Brunn, 37, 21 - 111. [1898]"]}

The first part of this thesis focuses on the family Geotrupidae, with special focus on the representatives of the genus Jekelius. It summarizes the present and historical taxonomic concepts and phylogenetic relationships within this family. Moreover, the morphology of adults and larvae and the geographical distribution of individual taxa are described. In the section on the biology and ecology of the family, emphasis is placed especially on feeding strategies, breeding behaviour and parental care. The results of this thesis consist of three parts, 1) taxonomy, 2) molecular analyses, and 3) flow cytometry. The first part brings a taxonomic revision of the Jekelius brullei species complex defined here. This part focuses on redescribing each known species, as well as describing additional species not yet formally described. This thesis defines the distribution ranges of each species based on previously published literature and our own data. Molecular analyses performed in this thesis generated a phylogenetic tree including a significant part of the species of the Jekelius brullei species complex. The last part of the thesis presents results obtained by flow cytometry for representatives of the species complex Jekelius brullei and a number of other taxa of the family Geotrupidae. The discussion then...

Authorship of Scarabaeus stercorosus (currently Anoplotrupes stercorosus) Who is the author of Scarabaeus stercorosus, Scriba or Hartmann? At the beginning of his article, Scriba (1791) mentioned that he had some conversations on entomological topics with ���Mr Hartmann���. Furthermore, on pages 250 and 251 of the article (Fig. 3), Scriba (1791), using quotation marks, mentioned a communication by Hartmann, most probably a letter, about a probably new species: ���Noch f��ge ich die Beschreibung eines Scarabaeus bey [bei], welche ich gleichfalls der G��te des Herrn Hartmanns zu danken habe, ��� Dieser K��fer ist eine Mittelart zwischen Scar. stercorarius und vernalis, von beiden aber wesentlich unterschieden. Er hat Gr��sse und Farbe des Sc. vernalis. Das Brustst��ck ist weit weniger gew��lbt als bey [bei] dem vernalis, und hierinnen dem stercorarius ��hnlich, ��ber und ��ber mit vertieften Punkten bes��et [bes��t = bedeckt], und hat in der Mitte eine vertiefte L��ngslinie. Bey [bei] dem vernalis sind weder deutliche Punkte, noch eine L��ngslinie auf dem thorax [Thorax] vorhanden, und bey [bei] dem stercorarius ist zwar letztre [letztere] da, die vertieften Punkte sind aber blo auf die ��userste [��ussersten] Seiten des thorax [Thorax] eingeschr��nkt, und nur wenige sind an der L��ngsinie herumgestreut. Die Fl��geldecken sind deutlich gestreift, die Streifen aber weit feiner als bey [bei] dem stercorarius, und es ziehen sich noch ausserdem ��ber dieselbe feine verworrene, unregelm��ssig netzf��rmig communicierende [kommunizierende] Linien. Die Keule der F��hlh��rner ist an der Spitze braunr��thlich [braunr��tlich]. Diese Art bleibt sich immer gleich, und lebt mit dem vernalis in Viehdung:��� Ich habe den n��mlichen K��fer auch hier, und an demselben alle angezeigte [angezeigten] Kennzeichen gefunden. Herr Hartmann nennt ihn Scar. stercorosus ���. The English translation from Scriba���s old German follows: ���Moreover, I add the description of a Scarabaeus, which I also have Mr Hartmann���s kindness to thank for. ��� This beetle is a species in-between Scar. stercorarius and vernalis, but substantially different from both. It has size and colouration of Sc. vernalis. The pronotum is much less convex than in vernalis and similar to stercorarius, and is over and over covered with deep punctures, and has in the middle an impressed longitudinal line. In vernalis neither distinct punctures nor a longitudinal line on the thorax are present, and in stercorarius, the latter is present, but the deep punctures are limited to only the extreme sides of the thorax, and only a few are scattered at the longitudinal line. The wing cases are distinctly striped, but the stripes are much finer than in stercorarius; and also, fine irregular reticulate interconnected lines stretch across them [i.e., the elytra]. The club of the antennae has a reddish-brown tip. This species always stays the same [? is not very variable], and lives with vernalis in cattle dung:��� I have this beetle here also, and found all indicated characters on it. Mr Hartmann names it Scar. stercorosus ���. This wording clearly indicates, that Hartmann alone is responsible both for the name and for satisfying the criteria of availability other than actual publication, hence is the author of the name (International Code of Zoological Nomenclature 1999: Article 50.1.1). We therefore suggest the taxon to be referred as Anoplotrupes stercorosus (Hartmann in Scriba, 1791). Note: Type material from Hartmann is not among the historical material in the Staatliches Museum f��r Naturkunde Stuttgart (Arnaud Faille, personal communication, July 2021). Potential material originating from Scriba is irrelevant for determining syntypes as Hartmann alone is the author of this species., Published as part of Ziani, Stefano, Bezd��k, Ale��, Krell, Frank-Thorsten & Sommer, David, 2021, Authorship and date of publication of the name Scarabaeus stercorosus (currently Anoplotrupes stercorosus) (Coleoptera: Scarabaeoidea: Geotrupidae), pp. 593-599 in Zootaxa 5067 (4) on pages 595-597, DOI: 10.11646/zootaxa.5067.4.7, http://zenodo.org/record/5683510, {"references":["Scriba, L. G. (1791) Entomologische Bemerkung und Erfahrungen. Journal fur die Liebhaber der Entomologie, 1 (3), 244 - 255.","International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, 306 pp."]}

Modified key to Palaearctic Oiceoptoma (modified from Růžička et al. 2004) The following couplets are modified to accommodate the newly described species. 5. Pronotum with diffuse median spots (usually composed of several scattered darker spots or almost indistinct, Figs. 4B, E). Scutellum more distinctly punctured medially than laterally (Fig. 6B). Elytra brown to black, subapical surface with rugosities (Figs. 2A, D, 9C). Male genital segment with ventrite 9 anteriorly distinctly longer than tergum 9 in ventral view (Fig. 12E).��� Proventrite with dense punctation laterally; medially dark brown to black, laterally only narrowly orange in ventral view (Fig. 5B). Median lobe of aedeagus subapically distinctly sinuous laterally, tapering to a widely rounded apex (Figs. 11C���D). Basal portion oval, robust, not exceeding the width of aedeagus (Figs. 11C���D). Female tergum 7 posteriorly with wide, only weakly expressed emargination (Fig. 16A). Female tergum 8 truncate posteriorly (Fig. 13C). (Central and north-eastern China, Far East of Russia, Korean Peninsula to Japan)..................................................... O. subrufum (Lewis) ��� Pronotum with four black distinct spots, arranged in a trapezoidal pattern (Figs. 4A, C). Scutellum with homogeneous punctation (Figs. 6A, C). Elytra subapically with simple surface (Figs. 1A, D, 2E, 3A, 9A���B, D). Male genital segment with ventrite 9 anteriorly as long as tergum 9 in ventral view (Figs. 12D, F).................................................. 6 6. Spots on pronotum distinctly delimited, smooth and glossy, only with sparsely distributed, minute punctures with very short setae (Figs. 4C, F). Proventrite with only minute, superficial punctation laterally; completely dark brown to black in ventral view (Fig. 5C). Elytra black (Figs. 2E, 3A, 6F, 9D). Apex of elytron short and rounded in female (Figs. 2E). Surface of scutellum and elytra with very weak, superficial punctation (Figs. 6C, F, 9D). Median lobe of aedeagus gradually tapered to a slender, sharp apex (Fig. 11E). Basal portion oval, less robust, not exceeding the width of aedeagus (Fig. 11E). Female tergum 7 posteriorly simple, without emargination (Fig. 16C). Female tergum 8 broadly truncate posteriorly (Fig. 14A). Female ventrites 5 and 6 longer medially, posteriorly exposing only a shorter area of brick-wall pattern (Figs. 14B, 15E). (Japan)............................................................................................... O. nigropunctatum (Lewis) ��� Spots on pronotum vaguely delimited, regularly covered with fine, distinct punctures, covered by long, orange setation (Figs. 4A, D). Proventrite with dense punctation laterally; medially dark brown to black, laterally widely orange in ventral view (Fig. 5A). Elytra light to dark brown (Figs. 1A, D, 6D, 9A���B). Apex of elytron elongated to blunt tip in female (Figs. 1D, 8A, 9B). Surface of scutellum and elytra with distinct, deep punctation (Figs. 6A, D, 9A���B), Median lobe of aedeagus gradually tapered to widely rounded, triangular apex (Figs. 11A���B). Basal portion widely oval, very narrow, markedly exceeding the width of aedeagus (Figs. 11A���B). Female tergum 7 posteriorly with narrow, deep emargination (Fig. 17A). Female tergum 8 regularly rounded posteriorly (Figs. 13A, 17A). Female ventrites 5 and 6 very short medially, posteriorly with long area of brick-wall pattern (Figs. 15A���B, F) (China: Zhejiang Province)..... O. tangi Sommer, Růžička, Nishikawa & Schneider, new species, Published as part of Sommer, David, R����i��ka, Jan, Nishikawa, Masaaki & Schneider, Jan, 2021, Oiceoptoma tangi (Coleoptera: Silphidae: Silphinae), a new species of carrion beetle from eastern China, pp. 445-472 in Zootaxa 4949 (3) on pages 470-471, DOI: 10.11646/zootaxa.4949.3.2, http://zenodo.org/record/4640397, {"references":["Ruzicka, J., Hava, J. & Schneider, J. (2004) Revision of Palaearctic and Oriental Oiceoptoma (Coleoptera: Silphidae). Acta Societatis Zoologicae Bohemicae, 68, 30 - 51."]}

Oiceoptoma nigropunctatum (Lewis, 1888) (Figs. 2E, 3A���C, 4C, F, 5C, F, 6C, F, 7C, 9D, 11E, 14F) Material examined. Japan, Kyushu, 1 ♂ (JRUC), Oita prefecture, Kuju mts, Mt. Kurodake [ca. 33��19���N 131��13���E], Sh��nai-machi, 19.v.1997, F. Hirokawa leg., S. Nomura det.; 1 ♂ (JRUC), Tokushima prefecture, Kisawa-son, Dosut��ge, Naka-gun, 18.vii.���11.x.2004, M. Yoshida leg., S. Nomura det.; 1 ♀ (JRUC), Oita prefecture, Kuju mts, Mt. Kurodake [ca. 33��19���N 131��13���E], 16.viii.1992, Ryo Noda leg., S. Nomura det., Published as part of Sommer, David, R����i��ka, Jan, Nishikawa, Masaaki & Schneider, Jan, 2021, Oiceoptoma tangi (Coleoptera: Silphidae: Silphinae), a new species of carrion beetle from eastern China, pp. 445-472 in Zootaxa 4949 (3) on page 446, DOI: 10.11646/zootaxa.4949.3.2, http://zenodo.org/record/4640397, {"references":["Lewis, G. (1888) Notes on the Japanese species of Silpha. The Entomologist, an Illustrated Journal of General Entomology, 21, 7 - 10."]}

Oiceoptoma tangi Sommer, Růžička, Nishikawa & Schneider, new species (Figs. 1A���F, 3D���F, 4A, D, 5A, D, 6A, D, 7A���C, 8A, C���D, 9A���B, 10A���D, 11A���B, 12A, D, 13A���B, 14 C���E, 15A���B, F, 17A���C) Oiceoptoma subrufum: Růžička et al. 2004: 47 (misidentification). Type locality. China, Zhejiang Province, Anji City, Longwangshan Mt., Pinxi Creek, 30��24���N, 119��27���E, alt. 1000��� 1100 m. Type material (30 specimens). Holotype, ♂ (SNUC), ��� China: Zhejiang Prov. | Anji City | Longwangshan Mt. | Pinxi Creek | 30��24���N, 119��27���E || alt. 1000~ 1100 m | 09-VI-2012 | HU & YIN leg. [p]���. Paratypes, 6 ♂♂ 3 ♀♀ (SNUC), 1 ♂ 1 ♀ (BMNH), 2 ♂♂ 2 ♀♀ (JRUC), same data as holotype ���; 2 ♂♂ 3 ♀♀ (SNUC), ��� China, Zhejiang Prov. | Anji County | Shenxi | 30�� 25��� N, 119�� 24��� E | alt. 400 m, 07-VI-2012 | ZHU Jian-Qing leg. [p]���; 1 ♀ (EUMJ), ��������� [= China] ��������� [= Zhejiang Prov.] Oi������ [= Lin���an-shi] | MRH��� [= Mt. West Tianmu Shan] | 6~ 11���V���1998 | ��� ����� [= Li-Zhen Li] IJDz [= leg.] [p]���; 1 ♀ (SNUC), ���16~20���V���20 [p] 06 [hw] | MRH��� [= Mt. West Tianmu Shan] ������ [= Lin���an-shi] [hw] | ������ [= Zhejiang Prov.] Bwx [hw] || Mt. w. Tianmu | 16~20. V, 2006 | Bi Wen-xuan leg. [hw]���; 1 ♂ (SNUC), ��� West Tianmushan | Lin���an County | 11~ 15���VI���2006 | alt. 300���400 m | HU & TANG leg. [p]���; 1 ♀ (SNUC), ��� China: Zejiang Prov. | Lin���an City | Mt. West Tianmu | alt. 300~ 500 m | 29���V~ 3���VI���2010 | Jian-Qing ZHU leg. [p]���; 1 ♂ (JRUC, DNA voucher JRUC-SIL-0054), ��� CHINA: Zhejiang Prov., | Lin���an County, West | Tianmushan Nat. Res., | 250 m NW of Original | Temple of Lion Sect, | 30.3431��N 119.4316��E, | J. H��jek & J. Růžička leg. || (WT10) 2.���6.vii.2017; || 1165 m, baited pitfall | traps #06 (fish meat, | ripening cheese), broad- | leaved deciduous forest | with Rhododendron [p]���; 2 ♀♀ (JRUC, DNA vouchers JRUC-SIL- 0010, JRUC-SIL-0011), ��� CHINA: Zhejiang Prov., | Lin���an County, West | Tianmushan Nat. Res., | 250 m NW of Original | Temple of Lion Sect, | 30.3431��N 119.4316��E, | Cheng-Bin Wang leg. || (WT10) 2.���6.vii.2017; || 1165 m, baited pitfall | traps #06 (fish meat; alcohol, vinegar + sugar), broad- | leaved deciduous forest | with Rhododendron [p]���; 2 ♂♂ (JSCC), ���NW. CHINA | TIENMUCHAN [without date and collector���s name, hw with a pencil] || Oiceoptoma | subrufum | (Lewis, 1888) ♂ | Jan Růžička det. 200[p]3[hw]���. Description. Male (Figs. 1A���C). Body oval, dorso-ventrally flattened. Pronotum orange with four black spots (Fig. 4A). Elytra light to dark brown. Head, antennae, legs and ventral part of body black. Flight wings fully developed, functional (Fig. 8A). Head. Black, lustrous surface with regular punctation (Fig. 3D). Covered with short, recumbent orange setation. Clypeus anteriorly widely notched (Fig. 8D). Anterior margin of clypeus with slightly irregular row of dense, long, orange setation in dorsal view. Eye kidney-shaped in lateral view (Fig. 17C). Head with short row of long erect setae behind eyes (Fig. 3D). Frons with marked dorsal tentorial pits and a transverse, elevated crest posteriorly (Figs. 3D, 17C). Maxilla with deeply punctuated cardo and base of stipes in ventral view, covered by long setae (Fig. 3E). Antennae. Medium-sized, with last four antennomeres forming a distinct club (Figs. 7A���C). Pronotum moderately transverse, widest posteriorly (Fig. 4A). Pronotal disc orange, medially with four black, vaguely delimited spots, arranged in symmetrical, trapezoidal pattern (Fig. 4A). Spots are regularly covered with fine, distinct punctures, bearing long, orange setation (Fig. 4D). Margin rimmed only in anterior half; anteriorly with wide, shallow excision, which is medially straight; anterior angles weakly elevated. Posterior margin slightly sinuous, weakly sinuate postero-laterally (Figs. 4A, 6A). Surface with dense punctation, punctures medium-sized, clearly separated, from disc to margin deeply impressed (Figs. 4A, 5D). Pronotum covered with longer, orange setation. Scutellum. Irregularly triangular, widely vaulted medially. Surface with dense, distinct, homogenous punctation; covered with longer orange setation, semierect anteriorly and distinctly recumbent medially and posteriorly (Fig. 6A). Elytra subparallel, at the widest point (along 2/3 of its length) wider than pronotum (Fig. 1A). Humerus with distinct denticle (Fig. 3F). Elytron with three distinctly elevated ridges; only first and second ridge reaching apex of elytron; external ridge shortest, reaching only two thirds of elytral length (Figs. 1A, 6D, 9A). Second ridge in last third slightly sinuous (Fig. 9A). Third ridge posteriorly terminated at apical callus (Fig. 9A). Ridges on both sides covered with a row of short, recumbent, black setae. Lateral margin of elytra strongly elevated only in anterior half, disappearing posteriorly. Apex of elytron regularly rounded (Figs. 1A, 9A). Basal part of elytra and humeri with short, recumbent, black setae (Fig. 3F). Surface dull, with isodiametric microsculpture (Figs. 6A, D); subapical surface regularly flat, without rugosities; with dense, distinct, regular punctation; punctures clearly separated by spaces 1.0���1.5 times their own diameter (Figs. 6D, 9A). Elytra with distinct, subrectangular punctures, with bluish metallic lustre covering most of the surface in ventral view (Figs. 8A, C). Elytral epipleura sparsely, finely but distinctly punctured in ventral view (Fig. 8A). Ventrum finely punctate, covered with short, recumbent, brown to black setation (Figs. 5A, 10C). Proventrite with dense punctation laterally; medially dark brown to black, laterally widely orange in ventral view (Fig. 5A). Mesocoxae widely separated (Fig. 10C). Abdominal ventrites with distinct impressions laterally in ventral view (Figs. 1B, 15A). Ventrite 6 extremely reduced in length medially, with expanded brick-wall pattern on intersegmental membrane (Figs. 15A���B, F). Legs. Protarsomeres 1���4 only slightly expanded, covered with short, dense, yellow setation (Figs. 14C���D). Pro-, / meso- and metatibia with two apical spurs of different length (Figs. 14C���D). Metatibia almost straight, only very slightly curved in ventral view (Fig. 10D). Abdominal segments. Tergite 10 relatively short in length (Fig. 12A). Ventrite 9 oval, elongate, anteriorly as long as tergite 9, constricted and deeply desclerotized medially in ventral view (Fig. 12D). Spiculum gastrale robust, elongate in ventral view (Fig. 12D). Aedeagus. Median lobe stout, robust, gradually tapered to widely rounded, triangular apex (Figs. 11A���B). Internal sac heavily sclerotized, medial portion composed of a great number of very small, regular, rounded denticles; basal part containing two strips of elongate, slender denticles (Fig. 11B). Parameres robust, almost straight, longer than median lobe, subapically constricted with rounded apex (Figs. 11A���B). Basal portion widely oval, very narrow, markedly exceeding the width of aedeagus (Figs. 11A���B). Sexual dimorphism. Female. Similar to male except for the following structures: Protarsus simple, basal protarsomere elongated (Fig. 14E). Apex of elytron elongated to a blunt tip (Figs. 1D���E, 8A, 9B). Tergum 7 with distinct, deep, narrow, U-shaped medial emargination (Fig. 17A). Tergum 8 subquadrate, regularly rounded posteriorly (Figs. 13A, 17A). Ventrite 8 widely transverse (Figs. 13B, 17B). Tergite 9 elongate, rounded; tergite 10 pentagonal, apex posteriorly regularly rounded with dense microsetae (Figs. 10A���B, 13A). Coxite laterally constricted on outer margin subapically in ventral view; stylus inserted lateroapically, its length slightly shorter than apex of coxite (Figs. 10A���B, 13A���B). Measurements. Elytra length 5.81���6.82 mm in males (mean 6.30 mm; 6.27 mm in the holotype), 6.72���7.26 mm (mean 6.96 mm) in females; pronotum length 3.41���3.82 mm in males (mean 3.63 mm; 3.55 mm in the holotype), 3.50���3.81 mm (mean 3.62 mm) in females; pronotum width 5.62���6.66 mm in males (mean 6.07 mm; 5.92 mm in the holotype), 5.63���6.42 mm (mean 6.10 mm) in females. Variability. Elytra mostly dark brown, light brown in some (probably subteneral) specimens. Spots on pronotum slightly differently delimited, being less separated from surrounding surface in some specimens. Differential diagnosis. Oiceoptoma tangi Sommer, Růžička, Nishikawa & Schneider, new species, is similar and probably closely related to O. nigropunctatum and O. subrufum. All three species share the following combination of characters: (1) orange pronotum with relatively short pubescence; (2) four dark spots on pronotal disc; (3) dark brown/black elytra with narrow epipleura; and (4) absence of patchy velvet pattern on elytral dorsal surface (see Růžička et al. (2004) for more details). The three species can be separated from each other using the character states listed in the modified key and in Table 1 below. Etymology. Patronymic, named after Liang Tang (Shanghai, China), an enthusiastic specialist of staphylinoid beetles (Agyrtidae, Staphylinidae: Steninae, Scaphidiinae, Staphylininae, Omaliinae etc.), who first recognized this species was unnamed. Collecting circumstances. The large type series was collected near Pinxi Creek on old large mammal carrion, together with numerous Necrodes littoralis (Linnaeus, 1758) (Silphidae), Creophilus maxillosus (Linnaeus, 1758) (Staphylinidae) and some clown beetles (Histeridae) (Figs. 18A���C). Three specimens were also collected on West Tianmushan, 250 m above the Original Temple of Lion Sect, using fish-baited pitfall traps in a broadleaved deciduous forest with Rhododendron (Figs. 19A���B). The specimens were collected between May and the beginning of July. ...Continued on the next page Distribution. So far known only from three localities (Longwangshan Mt., Shenxi, Tianmu Shan Mts.) in Zhejiang Province, China (Figs. 20���21)., Published as part of Sommer, David, R����i��ka, Jan, Nishikawa, Masaaki & Schneider, Jan, 2021, Oiceoptoma tangi (Coleoptera: Silphidae: Silphinae), a new species of carrion beetle from eastern China, pp. 445-472 in Zootaxa 4949 (3) on pages 466-469, DOI: 10.11646/zootaxa.4949.3.2, http://zenodo.org/record/4640397, {"references":["Ruzicka, J., Hava, J. & Schneider, J. (2004) Revision of Palaearctic and Oriental Oiceoptoma (Coleoptera: Silphidae). Acta Societatis Zoologicae Bohemicae, 68, 30 - 51.","Lewis, G. (1888) Notes on the Japanese species of Silpha. The Entomologist, an Illustrated Journal of General Entomology, 21, 7 - 10.","Linnaeus, C. (1758) Systema Naturae per Regna Tria Naturae, secundum Classes, Ordines, Genera, Species, cum Characteri- bus, Differentiis, Synonymis, Locis. Tomus I. Editio Decima, Reformata. L. Salvii, Holmiae, (4) + 823 + (1) pp. https: // doi. org / 10.5962 / bhl. title. 542"]}

Oiceoptoma subrufum (Lewis, 1888) (Figs. 2A���D, 4B, E, 5B, E, 6B, E, 7D, 8B, 9C, 11C���D, 12B, E, 13C���D, 14G, H, 15C���D) Material examined. Japan, Honshu, 1 ♂ 1 ♀ (JRUC), Yamanashi prefecture, Makioka-machi, Otometoge pass [ca. 35��16���N 138��59���E], 1450 m, 8.viii.1996, S. Nomura leg. et det. China, Gansu Province, 1 ♂ 1 ♀ (JRUC), Lazikou valley, 34��09.9���N 103��51.9���E, 2120 m, 28.vi.2005, J. H��jek, D. Kr��l & J. Růžička leg., Published as part of Sommer, David, R����i��ka, Jan, Nishikawa, Masaaki & Schneider, Jan, 2021, Oiceoptoma tangi (Coleoptera: Silphidae: Silphinae), a new species of carrion beetle from eastern China, pp. 445-472 in Zootaxa 4949 (3) on page 446, DOI: 10.11646/zootaxa.4949.3.2, http://zenodo.org/record/4640397, {"references":["Lewis, G. (1888) Notes on the Japanese species of Silpha. The Entomologist, an Illustrated Journal of General Entomology, 21, 7 - 10."]}

Bolbelasmus (Bolbelasmus) zagrosensis Sommer, Hillert, Hrůzov�� & Kr��l, new species (Figs. 1A, 2A, 3A, 4 A���B, 5A, 6A���C, 9, 10) Bolbelasmus nireus: Keith 2005: 51, fig. 1 (distribution); Montreuil 2017: 260 (distribution). Bolbelasmus nireus [partim, populations from Iran]: Arnone & Massa 2010: 412 (distribution); Miessen 2011: 110 (distribution); Hillert et al. 2016: 246 (distribution); Nikolajev et al. 2016: 33 (catalogue); Schoolmeesters 2020 (online catalogue)., Published as part of Sommer, David, Hillert, Oliver, Hr��zov��, Lucie & Kr��l, David, 2021, Bolbelasmus (Bolbelasmus) zagrosensis (Coleoptera: Scarabaeoidea: Bolboceratidae), a new species from Iran, along with an updated key to the western Palaearctic species of the subgenus, pp. 380-394 in Zootaxa 4920 (3) on page 391, DOI: 10.11646/zootaxa.4920.3.4, http://zenodo.org/record/4478107, {"references":["Keith, D. (2005) First record of the genus Bolbelasmus from Iran. Kogane, 6, 51 - 52.","Montreuil, O. (2017) New records of Geotrupidae and Scarabeidae dung beetles (Insecta, Coleoptera) from Iran. Journal of Entomological Society of Iran, 36, 259 - 277.","Arnone, M. & Massa, B. (2010) A new species of Bolbelasmus Boucomont, 1911 from Sicily (Italy). Il Naturalista Siciliano, 34, 401 - 414.","Miessen, G. (2011) Quelques commentaires sur le genre Bolbelasmus Boucomont, 1911 et description d'une nouvelle espece de Chypre. Lambillionea, 111, 109 - 119.","Hillert, O., Arnone, M., Kral, D. & Massa, B. (2016) The genus Bolbelasmus in the western and southern regions of the Mediterranean Basin (Coleoptera: Geotrupidae: Bolboceratinae). Acta Entomologica Musei Nationalis Pragae, 56, 211 - 254.","Nikolajev, G. V., Kral, D. & Bezdek, A. (2016) Family Geotrupidae Latreille, 1802. In: Lobl, I. & Lobl, D. (Eds.), Catalogue of Palaearctic Coleoptera. Vol. 3. Scarabaeoidea-Scirtoidea-Dasciloidea-Buprestoidea-Byrrhoidea. Revised and Updated Edition. E. J. Brill, Leiden, pp. 33 - 52. https: // doi. org / 10.1163 / 9789004309142 _ 003","Schoolmeesters, P. (2020) Scarabs: World Scarabaeidae Database (version Nov 2019). In: Roskov, Y., Abucay, L. Orrell, T., Nicolson, D., Bailly, N., Kirk, P., Bourgoin, T., DeWalt, R. E., Decock, W., DeWever, A., Nieukerken, E., van, Zarucchi, J. & Penev, L. (Eds.), Species 2000. Naturalis, Leiden. Digital resource. Available from: http: // www. catalogueoflife. org / annual-checklist / 2017 (accessed 20 June 2020)"]}

Key to the subgenus Bolbelasmus species from the western Palaearctic Region (modified from Hillert et al. 2016) 1. Posterior margin of pronotum incompletely bordered, interrupted laterally of scutellum. ���........................... 2 - Posterior margin of pronotum completely bordered. ���........................................................ 6 2. Lateral margin of pronotum nearly semicircular between anterior angles and midline in dorsal view, anterior angles almost rounded; frontal horn and medial hornlike apophysis of pronotum distinct; scutellum triangular, approximately as long as wide; body length 11.0��� 13.5 mm; southern parts of central Europe, France (Alsace), Italy, Balkan Peninsula, Poland, Russia and Ukraine; Asian part of Turkey...................................... ��� B. (B.) unicornis (Schrank von Paula, 1789) - Lateral margin of pronotum broadly rounded, regularly bent from anterior angle to middle, anterior corners of pronotum distinctly acuminate, lateral margin straight in front of anterior angles; frontal horn and medial hornlike apophysis of pronotum always indistinct; scutellum triangular, longer than wide; body length 9.5���12.0 mm; Cyprus, Greece (Rhodes), Iran, Iraq, Syria and Turkey. ���....................................................................................... 3 3. Eye cantus weakly acuminate and distinctly exceeding eye laterally; scutellum rounded apically; Asian part of Turkey. ���.............................................................................. B. (B.) tauricus Petrovitz, 1973 - Eye cantus regularly rounded and not exceeding eye laterally; scutellum acuminate apically. ���....................... 4 4. Frontal horn weakly spaced from fronto-clypeal carina (viewed from above), pronotal punctation double and dense, small punctures sparser; frontal horn directed upwards to weakly backwards (Figs. 3A, 3 C���D, 4A���B, 4D).................��� 5 - Frontal horn close to fronto-clypeal carina (viewed from above); pronotal punctation double and dense, small punctures denser; frontal horn directed obliquely forewards (Figs. 3B, 4C); aedeagus as in Fig. 5B; Cyprus..... B. (B.) makrisi Miessen, 2011 5. Border of posterior margin of pronotum weakly interrupted laterobasally, but with only several single punctures; frontal horn directed weakly backwards (Figs. 3 C���D, 4D); aedeagus as in Fig. 5C; Greece (Rhodes), Iraq, Syria and Asian part of Turkey............................................................................��� B. (B.) nireus (Reitter, 1895) - Border of posterior margin of pronotum distinctly interrupted laterobasally, single punctures absent; frontal horn directed upwardly (Figs. 3A, 4 A���B); aedeagus as in Fig. 5A; Iran..................................................................................................... ���. B. (B.) zagrosensis Sommer, Hillert, Hrůzov�� & Kr��l, new species 6. Scutellum impunctate or finely, shallowly punctate; frontal horn simple or furcate apically. ���........................ 7 - Scutellum densely punctate; apex of frontal horn simple..................................................... 11 7. Frontal horn simple; genae evenly rounded; oblique keel above eye longer than length of eye; head strongly constricted behind eye. ���.............................................................................................. 8 - Frontal horn furcate apically; genae more or less angulate anterolaterally; oblique keel above eye reaching to anterior part of eye; head not strongly constricted behind eye. ���............................................................ 9 8. Body surface blackish; pronotal punctation dense and distinct, separated by distance of 1���2 punctures combined; Algeria, Italy (Sicily) and Tunisia. ���................................................ B. (B.) vaulogeri (Abeille de Perrin, 1898) - Body surface brownish; pronotal punctation sparse but distinct, separated by distance of 3���4 punctures combined; Greece (Crete, Kasos). ���....................................................... B. (B.) keithi Miessen & Trichas, 2011 9. Frontal horn short (not longer than wide), apex of horn approximately as wide as base of horn; pronotal punctation denser with exception of basal part, puncture separated by diameter of puncture; pronotal base with fine and regularly spaced punctures, puctures in shape of pinned points; elytral striae well impressed, completely developed; elytral intervals moderately convex; Spain. ���............................................................... B. (B.) brancoi Hillert & Kr��l, 2016 - Frontal horn long (longer than wide), tapering apically; pronotal punctation sparser, puncture separated approximately by three their diameters; pronotal base with fine, poorly visible and well separated punctures, punctures in shape of pinned points; elytral striae not impressed, punctures in row visible; elytral intervals flat; northern Africa. ���............................. 10 10. Frontal horn parallel to anterior part of eyes; horn sharply keeled at basal half; middle hornlike apophysis of pronotum directed upwards, subapical protrusion situated basally; lateral hornlike apophyses of pronotum directed forwards; Algeria and Morocco. ���.................................................................. B. (B.) bocchus (Erichson, 1841) - Base of frontal horn parallel to level of posterior part of eyes; horn broadly keeled apically; middle hornlike apophysis of pronotum directed obliquely, subapical protrusion situated at apex; lateral hornlike apophyses of pronotum directed forwardly; Egypt, Libya and Tunisia.................................. ��� B. (B.) nikolajevi Hillert, Arnone, Kr��l & Massa, 2016 11. Apex of frontal horn tapering apically, acute, weakly flattened in anterior part of horn; Spain and Gibraltar................................................................................... ��� B. (B.) howdeni Hillert & Kr��l, 2016 - Apex of frontal horn roughly as wide as base, apex broadly acute, strongly flattened in anterior part of horn; France (including Corsica), Portugal and Spain................................................. ��� B. (B.) gallicus (Mulsant, 1842), Published as part of Sommer, David, Hillert, Oliver, Hr��zov��, Lucie & Kr��l, David, 2021, Bolbelasmus (Bolbelasmus) zagrosensis (Coleoptera: Scarabaeoidea: Bolboceratidae), a new species from Iran, along with an updated key to the western Palaearctic species of the subgenus, pp. 380-394 in Zootaxa 4920 (3) on pages 392-393, DOI: 10.11646/zootaxa.4920.3.4, http://zenodo.org/record/4478107, {"references":["Hillert, O., Arnone, M., Kral, D. & Massa, B. (2016) The genus Bolbelasmus in the western and southern regions of the Mediterranean Basin (Coleoptera: Geotrupidae: Bolboceratinae). Acta Entomologica Musei Nationalis Pragae, 56, 211 - 254.","Miessen, G. (2011) Quelques commentaires sur le genre Bolbelasmus Boucomont, 1911 et description d'une nouvelle espece de Chypre. Lambillionea, 111, 109 - 119.","Reitter, E. (1895) Beschreibung neuer oder wenig gekannter Coleoptera aus der Umgebung von Akbes in Syrien. Wiener Entomologische Zeitung, 14, 79 - 88. https: // doi. org / 10.5962 / bhl. part. 13210"]}

Bolbelasmus (Bolbelasmus) nireus (Reitter, 1895) (Figs. 1 C���D, 2C���D, 3C���D, 4D, 5C, 7A���C, 8A���B, 9) Bolboceras nireus Reitter, 1895: 81 (type locality: ���Akbes [= Mayd��n Akbis]���). Type material examined. Syria: Holotype, ♀ (MNHN) (Figs. 7 A���C, 8A���B) ��� Bolboceras | Nireus m | n. sp. Akbes [hw, Reitter���s hand] || Typus [p, red label] || jkF720329 || VIO || Bolbelasmus | nireus Rttr. || det. J. Krikken 197[p]2[hw, Krikken���s hand] || Bolbelasmus nireus | (Reitter, 1895) | D. Kr��l & D. Sommer det. 2019 [p]���. Additional material examined. Greece, Rhodes: 1 ��� (LSCN), 1 ��� (OHCB) (Figs. 1C, 2C, 3C, 4D, 5C), Lindos, 36��05���32���N 28��05���06���E, 16.iv.2017, L. Schmidt lgt.; 1 ��� (OHCB), Lalyssos, 36��25���01.09���N 28��08���44.03���E, 28.iii.2017, R. Dietze lgt.; Syria: 1 ♀ (MNHN), Akb��s, [without date], [E.] Reitter; Turkey: 1 ♀ (OHCB), Side, 27.iv.1973, K. Warncke lgt.; 1 ♀ (NMPC), Korkuteli, 11.vi.1991, B. Němec lgt.; 1 ��� (JMCP), İ��el, ��amliyayla, Sebil, 24.���26.v.1995, J. Mička lgt.; 1 ♀ (NMPC), 20 km S of Kemer, v.2001, [without collector���s name]; 1 ��� (AHCB) (Figs. 1D, 2D, 3D), NW of Alanya, NE of Manavgat, 8.iv.2015, M. Sn��žek lgt.; 2 ������ (MNHN), Adana [without date and collector���s name]. Disribution. Iraq, Syria and Turkey (Nikolajev et al. 2016). New country record for Greece (island of Rhodes) (see map, Fig. 9). The record of the species B. (B.) unicornis (Schrank, 1789) from the island of Rhodes by Schatzmayr (1936) most likely relates to the species B. (B.) nireus. Note. The distribution in Iraq is based on a single female specimen from Assur tentatively assigned to this species by Krikken (1977). We have not studied this specimen, so we cannot confirm with certainty what species it belongs to. In addition, this Iraqi locality of the species is very far from its distribution in Turkey (see map, Fig. 9). Their distribution in Syria is based on two females (including the holotype) collected at ���Akbes���. This locality, nowadays a small village Mayd��n Akbis belongs to the territory of Syria (Aleppo Governorate) and is situated very close to the border with Turkey (see map, Fig. 9). Between 1881���1926 there was the Trappist monastery ���Abbaye d���Akb��s��� (Le Bras et al. 1979). The site is not in modern Turkey, as stated in some papers, see Kry��tufek et al. (2001), Keith (2005) or Arnone & Massa (2010)., Published as part of Sommer, David, Hillert, Oliver, Hr��zov��, Lucie & Kr��l, David, 2021, Bolbelasmus (Bolbelasmus) zagrosensis (Coleoptera: Scarabaeoidea: Bolboceratidae), a new species from Iran, along with an updated key to the western Palaearctic species of the subgenus, pp. 380-394 in Zootaxa 4920 (3) on page 386, DOI: 10.11646/zootaxa.4920.3.4, http://zenodo.org/record/4478107, {"references":["Reitter, E. (1895) Beschreibung neuer oder wenig gekannter Coleoptera aus der Umgebung von Akbes in Syrien. Wiener Entomologische Zeitung, 14, 79 - 88. https: // doi. org / 10.5962 / bhl. part. 13210","Nikolajev, G. V., Kral, D. & Bezdek, A. (2016) Family Geotrupidae Latreille, 1802. In: Lobl, I. & Lobl, D. (Eds.), Catalogue of Palaearctic Coleoptera. Vol. 3. Scarabaeoidea-Scirtoidea-Dasciloidea-Buprestoidea-Byrrhoidea. Revised and Updated Edition. E. J. Brill, Leiden, pp. 33 - 52. https: // doi. org / 10.1163 / 9789004309142 _ 003","Schatzmayr, A. (1936) Risultati scientifici delle cacce entomologiche di S. A. S. il Princiope Alessandro della Torre et Tasso nelle isole dell'Egeo. VI. - Scarabaeidae (1). Bollettino del Regio Laboratorio di Entomologia Agraria di Portici, 30, 9 - 23.","Krikken, J. (1977) The genus Bolbelasmus Boucomont in Asia with notes on species occurring in other regions. Zoologische Mededeelingen, 51, 277 - 292.","Le Bras, G., Hourlier, J. & Cocheril, M. (1979) Les Ordres religieux, la Vie et l'Art. Vol. 1. Monasteres et communautes; les Benedictins; les Cisterciens; les Chartreux; les ordres militaires. Flammarion, Paris, 266 pp.","Krystufek, B., Spitzenberger, F. & Kefelioglu, H. (2001) Description, taxonomy, and distribution of Talpa davidiana. Mammalian Biology, 66, 135 - 143.","Keith, D. (2005) First record of the genus Bolbelasmus from Iran. Kogane, 6, 51 - 52.","Arnone, M. & Massa, B. (2010) A new species of Bolbelasmus Boucomont, 1911 from Sicily (Italy). Il Naturalista Siciliano, 34, 401 - 414."]}

Bolbelasmus (Bolbelasmus) makrisi Miessen, 2011 (Figs. 1B, 2B, 3B, 4C, 5B, 9) Bolbelasmus (Bolbelasmus) makrisi Miessen, 2011: 111, figs. on pp. 116, 118���119 [not numbered] (type locality: ��� Kourion, Lemesos, alt.: 50 m, Cyprus ���). Type material examined. Cyprus: Paratypes, 1 ��� 1 ♀ (GMCL), ���Rizoelia nat. for. Park | Larnaka���CYPRUS || leg.: E. Kakouris | collection: G. Miessen || Bolbelasmus makrisi nov. sp. | G. Miessen det. 2011 | PARATYPUS [printed, red label]���. Additional material examined. Cyprus: 1 ��� 1 ♀ (OHCB), Akamas NP, 5 km N of Lara, 13.���20.iii.2015, O. Hillert lgt.; 1 ��� (OHCB), Akamas NP, Lara, 13.���20.iii.2015, O. Hillert lgt.; 1 ��� (OHCB), Miliou, N of Paphos, 13.���20.iii.2015, O. Hillert lgt.; 1 ��� (OHCB), Miliou, 34��56���22.23���N, 32��27���35.90���E, 7.vi.2014, vivant dans piscine [alive in pool], G. Miessen lgt.; 1 ��� (OHCB), Miliou, 34��56���21.66���N, 32��27���35.59���E, 7.vi.2014, attire par lampe UV [attracted by UV light], G. Miessen lgt.; 16 ������ 15 ♀♀ (OHCB) (Figs. 1B, 2B, 3B, 4C, 5B), 2 ������ 2 ♀♀ (NMPC), 5 km NE of Pegeia, Paphos env., 13.���20.iii.2015, O. Hillert lgt. Distribution. Endemic to Cyprus (Nikolajev et al. 2016) (see map, Fig. 9)., Published as part of Sommer, David, Hillert, Oliver, Hr��zov��, Lucie & Kr��l, David, 2021, Bolbelasmus (Bolbelasmus) zagrosensis (Coleoptera: Scarabaeoidea: Bolboceratidae), a new species from Iran, along with an updated key to the western Palaearctic species of the subgenus, pp. 380-394 in Zootaxa 4920 (3) on page 385, DOI: 10.11646/zootaxa.4920.3.4, http://zenodo.org/record/4478107, {"references":["Miessen, G. (2011) Quelques commentaires sur le genre Bolbelasmus Boucomont, 1911 et description d'une nouvelle espece de Chypre. Lambillionea, 111, 109 - 119.","Nikolajev, G. V., Kral, D. & Bezdek, A. (2016) Family Geotrupidae Latreille, 1802. In: Lobl, I. & Lobl, D. (Eds.), Catalogue of Palaearctic Coleoptera. Vol. 3. Scarabaeoidea-Scirtoidea-Dasciloidea-Buprestoidea-Byrrhoidea. Revised and Updated Edition. E. J. Brill, Leiden, pp. 33 - 52. https: // doi. org / 10.1163 / 9789004309142 _ 003"]}