The genusBolbelasmus in the western and southern regions of the Mediterranean Basin (Coleoptera: Geotrupidae: Bolboceratinae)

ACTA ENTOMOLOGICA MUSEI NATIONALIS PRAGAE Published 15.vii.2016 Volume 56(1), pp. 211–254 ISSN 0374-1036 http://zoobank.org/urn:lsid:zoobank.org:pub:7EFF344A-B2FF-4A14-80AC-3B047298201D The genus Bolbelasmus in the western and southern regions of the Mediterranean Basin (Coleoptera: Geotrupidae: Bolboceratinae) Oliver HILLERT1), Marcello ARNONE2), David KRÁL3) & Bruno MASSA4) 1) Kieferndamm 10, D-15566 Schöneiche bei Berlin, Germany; e-mail: [email protected] 2) Via San Michele 8, I-90010 Altavilla Milicia (Palermo), Italy; e-mail: [email protected] 3) Charles University in Prague, Faculty of Science, Department of Zoology, Viničná 7, CZ-128 43 Praha 2, Czech Republic; e-mail: [email protected] 4) University of Palermo, Department of Agricultural and Forest Sciences, Viale Scienze 13, I-90128 Palermo, Italy; e-mail: [email protected] Abstract. The Bolbelasmus Boucomont, 1911 species of the western and southern regions of the Mediterranean Basin (Northern Africa, Iberian Peninsula and France) are revised. The following three new species are described: Bolbelasmus brancoi Hillert & Král sp. nov. and Bolbelasmus howdeni Hillert & Král sp. nov., both from Spain and Gibraltar, and Bolbelasmus nikolajevi Hillert, Arnone, Král & Massa sp. nov. from Egypt, Libya and Tunisia. Bolbelasmus vaulogeri (Abeille de Perrin, 1898) stat. restit. is removed from synonymy with B. bocchus (Erichson, 1841) and reinstated as a separate species. Bolbelasmus romanorum Arnone & Massa, 2010 is considered a junior subjective synonym of B. vaulogeri. Lectotypes for Bolboceras bocchus Erichson, 1841 and Bolboceras vaulogeri Abeille de Perrin, 1898 are designated. Relevant diagnostic characters (head, pronotum, elytron, external male genitalia) are illustrated. Identifi cation keys for both males and females, and an annotated list of the Western Palaearctic representatives of the genus Bolbelasmus are presented. Finally, fi rst records are given for B. gallicus (Mulsant, 1842) from Corsica and the Midi-Pyrénées region of France, B. keithi Miessen & Trichas, 2011 from the Greek island of Rhodes, and B. unicornis (Schrank von Paula, 1789) from the Tuscany province of Italy. Key words. Coleoptera, Scarabaeoidea, Geotrupidae, Bolboceratinae, Bolbelasmini, Bolbelasmus, taxonomy, revision, new species, synonymy, species restitution, lectotype designation, key, distribution, annotated list, Mediterranean Region, Palaearctic Region HILLERT et al.: Bolbelasmus of the Western and Southern Mediterranean (Geotrupidae) 212 Introduction The bolboceratine genus Bolbelasmus was erected by BOUCOMONT (1911) in his study ‘Contribution à la classifi cation des Geotrypidae’, where he designated the following three species: Bolboceras bocchus Erichson, 1841, Bolboceras gallicus Mulsant, 1842 and Bolboceras unicornis (Schrank von Paula, 1789) as ‘species typicae’. In the same study, BOUCOMONT (1911) erected the genus Kolbeus with ‘species typicae’Bolboceras arcuatum Bates, 1887 and Bolboceras coreanum Kolbe, 1886. Type species were designated by LUCAS (1920) for Kolbeus, namely Bolboceras coreanus, while 30 years later, CARTWRIGHT (1953) designated Bolboceras gallicus for Bolbelasmus. In addition, CARTWRIGHT (1953) synonymized Kolbeus under Bolbelasmus in the same article. In a recent taxonomic study, NIKOLAJEV (1996) described the tribe Bolbelasmini with three genera: Bolbelasmus Boucomont, 1911, Bolbotrypes Olsoufi eff, 1907 and Bolbogonium Boucomont, 1911. Later on NIKOLAJEV (2003) placed Kolbeus at the rank of subgenus of Bolbelasmus. SCHOLTZ & BROWN (1996) have proposed that the group of genera related to Bolboceras Kirby, 1819 belongs to the rank of family (Bolboceratidae). However, in accordance with recent studies based on molecular data (e.g. SMITH et al. 2006) and with phylogeny of the Geotrupidae based on larval morphology (VERDÚ el al. 2004), the group is treated as a subfamily of Geotrupidae in this study (see also BOUCHARD et al. 2011). Currently, we recognize 27 species of the genus Bolbelasmus (including subgenus Kolbeus) from the Holarctic Region, including three new species described in this study. Eight taxa are distributed throughout the New World, eleven are known from Europe and the Mediterranean Region and eight are native to the Himalaya, Far East, China and south-eastern Asia (e.g. REITTER 1895, HOWDEN 1964, PETROVITZ 1968, KRIKKEN 1977, MASUMOTO 1984, HOWDEN & SOLIS 1995, NIKOLAJEV 2005, KRÁL et al. 2006, LI et al. 2008). According to KRIKKEN (1977), ARNONE & MASSA (2010), MIESSEN (2011) and MIESSEN & TRICHAS (2011) and to the results of the present study, eleven species of the genus Bolbelasmus are distributed in Europe, Middle East and North Africa. The new species in this genus were described thanks to the study of a large number of specimens from almost 70 instutional and private collections and series of specimens recently collected by the authors. Material and methods The habitus photographs were either taken by D. Král using a Canon MP-E 65mm/2.8 Macro lens with 5:1 optical magnifi cation on bellows attached to a Canon EOS 550D camera or by M. Romano using a Canon EOS 450D digital camera mounting a Sigma 70 mm 1:2.8 DG macro. Partially focused images of the specimen were stacked using Helicon Focus 3.20.2 Pro software. The following codes identify the collections housing the material examined (curators in round brackets): ABCB Alberto Ballerio collection, Brescia, Italy; AHCL Antonio Hidalgo Fontiveros collection, Linares, Spain; ARCL Andreas Reichenbach collection, Leipzig, Germany; Acta Entomologica Musei Nationalis Pragae, 56(1), 2016 213 BMCP Bruno Massa collection (deposited in MSNG); BMNH Natural History Museum, London, United Kingdom (Maxwell V. L. Barclay, Malcolm Kerley); BNCP Bohuslav Němec collection, Praha, Czech Republic; CMNC †Henry F. & Anne T. Howden collection, Entomology Division, Museum of Nature Canada, Ottawa, Canada (Aleš Smetana); CUPC Department of Zoology, Charles University, Prague, Czech Republic (Petr Šípek); DJCP Daniel Juřena collection, Prostějov, Czech Republic; DKCC Denis Keith collection, Chartres, France; DKCP David Král collection (deposited in NMPC); ERCS Eckehard Rößner collection, Schwerin, Germany; GMCL Geoffrey Miessen collection, Liège, Belgium; GNCA Georgy V. Nikolajev collection, Almaty, Kazakhstan; GSCT Guido Sabatinelli collection, Tunis, Tunisia; HFCB Hans Fery collection, Berlin, Germany; HKCS Harald Kaltz collection, Schlabendorf, Germany; HNHM Hungarian Natural History Museum, Budapest (Ottó Merkl, Győző Szél); IBCF Ivo Boščík collection, Frýdek-Místek – Skalice, Czech Republic; IECA Institute of Entomology, Biology Centre CAS, České Budějovice, Czech Republic (Aleš Bezděk); IMCO Ivo Martinů collection, Olomouc, Czech Republic; JBCP Jan Batelka collection, Prague, Czech Republic; JCCL Javier Castillo collection, Linares, Spain; JDCC Jiří Dvořák collection, Čelčice, Czech Republic; JHCP Jiří Háva collection, Únětice, Czech Republic; JMCH Jan Matějíček collection, Hradec Králové, Czech Republic; JMCP Jiří Mička collection, Prague, Czech Republic; JUCS José María Urbano collection, Sevilla, Spain; JNCS Jerónimo Navarro García collection, Sevilla, Spain; JSCP Jan Schneider collection, Prague, Czech Republic; JSCS Joachim Schönfeld collection, Sinzig, Germany; JZCJ Jaroslav Žák collection, Jezernice, Czech Republic; LHCB Lars Hendrich collection (deposited in ZSMC); JLCM José Ignácio López Colón collection, Madrid, Spain; LMCT Ladislav Mencl collection, Týnec nad Labem, Czech Republic; LSCN Ludger Schmidt collection, Neustadt, Germany; MACP Marcello Arnone collection, Palermo, Italy; MECI Manfred Egger collection, Innsbruck, Austria; MHNL Muséum d’histoire naturelle, Lyon, France (Harold Labrique); MLCD Mike Liebscher collection, Dresden, Germany; MNHB Museum für Naturkunde, Berlin, Germany (Johannes Frisch, Joachim Willers); MNHN Muséum national d’Histoire naturelle, Paris, France (Olivier Montreuil, Antoine Mantilleri); MSNT Museo Regionale Scienze Naturali, Turin, Italy (Luca Picciau); MSAP Museo Scienze Agrarie, Portici, Naples, Italy (Antonio Garonna); MSNG Museo Civico di Storia Naturale ‘G. Doria’, Genoa, Italy (Maria Luisa Tavano); MSNM Museo Civico di Storia Naturale, Milan, Italy (Fabrizio Rigato); MZUC Museo di Zoologia dell’Università of Catania, Catania, Italy (Giorgio Sabella); MZUF Museo di Zoologia La Specola dell’Università of Florence, Italy (Luca Bartolozzi); NMEC Naturkundemuseum, Erfurt, Germany (Matthias Hartmann); NHMB Naturhistorisches Museum, Basel, Switzerland († Michel Brancucci, Eva Sprecher-Uebersachs, Matthias Bohrer); NMPC National Museum (Natural History), Prague, Czech Republic (Jiří Hájek); OBCB Olivier Boilly collection, Marcq en Baroeul, France; HILLERT et al.: Bolbelasmus of the Western and Southern Mediterranean (Geotrupidae) 214 OHCB Oliver Hillert collection, Schöneiche bei Berlin, Germany; PJCL Pavel Jáchymek collection, Luhačovice, Czech Republic; PKCS Petr Kylies collection, Slaný, Czech Republic; PTCP Pavel Turek collection, Lanškroun, Czech Republic; RCCP Radek Červenka collection, Prague, Czech Republic; RMCM Radoslav Muczka collection, Mikulov, Czech Republic; SJCP Stanley Jákl collection, Prague, Czech Republic; SMTD Staatliches Museum für Tierkunde, Dresden, Germany (Olaf Jäger); SZCM Stefano Ziani collection, Meldola, Italy; TVCP Tomáš Vendl collection, Prague, Czech Republic; VACP Vittorio Aliquò collection, Palermo, Italy; VKCB Vítězslav Kubáň collection, Brno, Czech Republic; VKCC Václav Křivan collection, Chaloupky, Czech Republic; VMCP Vladislav Malý collection, Prague, Czech Republic; VRCH Vladislav Řebíček collection, Hradištko, Czech Republic; VTCZ Václav Týr collection, Žihle, Czech Republic; ZFMK Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany (Dirk Ahrens); ZLCK Zdeno Lucbauer collection, Kattering, United Kingdom; ZSMC Zoologische Staatssammlung, München, Germany (Michael Balke, Lars Hendrich). Altogether, 1332 specimens (see material below) were studied. Specimens of the newly described species are provided with one red printed label ‘Name of taxon sp. nov., HOLOTYPUS , ALLOTYPUS  or PARATYPUS  or  followed by the name of the authors and the year’. In the case of lectotype or paralectotype designation, each specimen bears a red printed label: ‘Name of a taxon with author and year of original description, LECTOTYPUS  or PARALECTOTYPUS  or  David Král & Oliver Hillert des. 2014’. Remarks of the authors and comments are indicated in brackets. Exact label data are cited for the type material. Information in quotation marks (‘’) indicates the original spelling. Coordinates and altitude are assigned to each locality mentioned in the text (material examined in each species) (see Appendix 1). These data were used in the construction of distribution map (see Fig. 59). Taxonomy Bolbelasmus bocchus (Erichson, 1841) (Figs 1, 8, 14, 21, 27, 33, 39, 45–51, 59–60) Bolbocerus Bocchus Erichson, 1841: 170 (original description). GORY (1841): table 71, fi gs 1, 1A (list); MARSEUL (1857): 83 (list); JACQUELIN DU VAL (1863): 434 (catalogue); BOUCOMONT (1902): 4 (catalogue); HEYDEN et al. (1906): 725 (catalogue); WINKLER (1927): 1035 (catalogue). Bolbelasmus bocchus: BOUCOMONT (1912): 17 (catalogue); WINKLER (1927): 1035 (catalogue); KOCHER (1958): 77 (catalogue); RUIZ (2000): 84 (distribution). Bolbelasmus bocchus [partim, populations from Morocco]: BARAUD (1977): 160 (monograph); MARTÍN-PIERA & LÓPEZ-COLÓN (2000): 179 (monograph). Bolbelasmus bocchus bocchus [partim, populations from Morocco]: KRIKKEN (1977): 290, fi gs 16–17 (revision); BARAUD (1985): 53, fi g. 8 (monograph); BARAUD (1992): 44 (monograph); KRÁL et al. (2006): 82 (catalogue). Bolboceras Fissicornis Mulsant, 1843: 280 (original description). Type locality: ‘Algérie’. MARSEUL (1857): 83 (synonymy). Type locality. ‘Algier, [Bona = Annaba]’ (see Type material for details). Type material examined (3 specimens). ALGERIA: LECTOTYPE (present designation):  (MNHB), ‘Hist. – coll. (Coleoptera), Nr. 25569, Bolboceras Bocchus Er. – Kl., Bona, Wagner, Zool. Mus. Berlin, Syntypus, Bolbelasmus Acta Entomologica Musei Nationalis Pragae, 56(1), 2016 215 bocchus Erichson, 1841, labelled by MNHUB 2013’. PARALECTOTYPES: No. 1,  (MNHB), ‘Hist. – coll. (Coleoptera), Nr. 25569, Bolboceras Bocchus Er. – Kl., Bona, Wagner, Zool. Mus. Berlin, Syntypus, Bolbelasmus bocchus Erichson, 1841, labelled by MNHUB 2013’; No. 2,  (MNHB), ‘Hist. – coll. (Coleoptera), Nr. 25569, Bolboceras Bocchus Er. – Kl., Bona, Wagner, Zool. Mus. Berlin, Syntypus, Bolbelasmus bocchus Erichson, 184, labelled by MNHUB 2013’; see Figs 45–51. Additional material examined (183 specimens). ALGERIA: Algeria, 2  1  (SMTD), 1  (MNHN), 2  2  (BMNH); Algerie, 1  2  (MHNL); Algerien, 3  (SMTD). ALGIERS PROV.: Alger (city), vi.1996, 1  (DKCP); Alger, 1  (MNHN); Algier, 2  (MNHB); L. Arba, 1  1 , Reitter [lgt.] (MNHN), 1  (VMCP). SIDI BEL ABBÈS PROV.: Chanzy, [18]90, 1 , Vaulog[er] lgt. (ZSMC); Chanzy, 1890, 1 , de Vauloger lgt. (MNHN). DJELFA PROV.: Takersan, iv.1893, 2  3 , A Ben Yacoub [lgt.] (MNHN); Takersan, Djelfa, 17.v. 1924, 1  (DKCC). EL BAYADH PROV.: Region de Geryville, vi.1910, 3  8 , Harold Powell lgt. (MNHN); Le Kreider, v.[18]96, 1  1  (MNHN); El Kreider, v.1896, 4  2 , Dr. A. Chobaut lgt. (MNHN), 2  1  (LHCB); El-Kreider (Oran), 1 , L. Bleuse [lgt.] (ZFMK); Kreider, 1 , Bedel [lgt.] (MNHB), 1  (ZSMC), 1  2  (MNHN); Kreider, 4  4 , Bedel lgt. (MNHN); Kreider, 3  4  (MNHN), 1  (NMPC), 4  (ZFMK). LAGHOUAT PROV.: Afl ou, 4.iv.1981, 1 , H. Pierotti [lgt.] (MSNG). MASCARA PROV.: Oran, Mascara, iv.–v.1913, 24  9 , G. Le Comte lgt. (MNHN); Mascara, 1 , Dr. A. Cobos lgt. (MNHN); Mascara, 1  (NHMB), 1  1  (VMCP). M’SILA PROV.: Ain-Ograb, 1 , Dr. Martin [lgt.] (ZFMK). NAÂMA PROV.: Mecheria, 1  (MNHN). ORAN PROV.: Magenta (prov. d’Oran), 1 , E. Duroux [lgt.] (ZFMK); Magenta (Oran), 1  (DKCC); Oran, R. J. 3.xi.1903 1 , B. [? lgt.] (MNHN); Oran, 1 , coll. R. Paulian (MNHN); Oran, Magenta, 1915, 1 , M. Rotrou lgt. (MNHN); Magenta, 1  (MNHN). TLEMCEN PROV.: Lambaze, vi.1885, 2 , L. Bleuse lgt. (MNHN); Sebdou, 1903, 1  (MNHN). MOROCCO: GHARB-CHRARDA-BÈNI HSSEN REG.: KÉNITRA PROV.: s.[south of] Oujda, Ain-el Kerma, v.1972, 2  1 , D. Bernh[auer] lgt. (JSCS). MEKNÈS-TAFILALET REG.: IFRANE PROV.: Mittlerer Atlas, Ifrane, 1700 m, 8.–9.vi.[19]96, 1 , A. Kallies lgt. (OHCB); Middle Atlas, Azrou, 18.–19.vi.1936, 1 , K.H. Chapman & G.A. Bisset [lgt.] (BMNH); Moyen-Atlas, near Jbel Hebri, 12.v.1980, 1 , J. Gourvès rec. [lgt.] (MHNL); Azrou, Foret de Cedres, 10.v.[19]95, 1 , Jiří Hájek lgt. (JSCP), 1  (NMPC); Azrou, 10.v.1995, 1 , M. Rozsíval lgt. (JZCJ); Ifrane env., 3.v.1995, 1 , J. Bechr lgt. (PJCL); Ifrane, v.[19]95, 1 , J. Holiš lgt. (JZCJ); Ifrane, Dayet Iffer, 26.iv.2014, 11  5 , O. Boilly lgt. (OBCB); Ifrane, 8.vi.1973, 2  (SJCP); Ifrane, 1.v.68, 1 , J. Gallet lgt. (MNHN); Ifrane, 11.v.[19]67, 2  1 , Y. Lajonquier lgt. (MNHN); NE of Ifrane, Dayt-Ifrah, 17.v.2003, 1 , M. Snížek lgt. (VKCC); Taforalt (Oujda), 11.iv.1992, 1  1 , G. Chavanon lgt. (IECA); Zad Pass, 2180 m, 24.iv.2014, 1 , O. Boilly lgt. (OBCB). ORIENTAL REG.: BERKANE PROV.: Taforal, Beni Snassen (Parc Naturel), iii.[19]51, 1 , Antoine lgt. (MNHN). JERADA PROV.: W [of ]Touissite, Jbel Mahsor, 34°29′46″N 01°46′50″W, 1050 m, 2 , R. Rejzek lgt. (JSCP). OUJIDA-ANGAD PROV.: Torf. Ouazane, pr.[prope] Oujda, vi.[19]35, (Lacous), 1  1  (MNHN). TAOURIRT PROV.: Gaada de Debdou plat[eau]., E of Rchida, 33°52′N 3°10′W, ca. 1420–1550 m, 20.–21.iii.2013, 1  1 , D. Král lgt. (DKCP); Gaada de Debdou plat[eau]., E of Rchida, 33°52′N 3°10′W, ca. 1420–1550 m, 20.–21.iii.2013, 3  1 , O. Hillert lgt. (OHCB). SPAIN (territory in Morroco): MELILLA AUTONOM. COMM.: 1  1 , Melilla, Pardo Alcaide [lgt.] (MSNG); 2  2 , Rostrogardo, Melilla, Pardo Alcaide lgt. (MNHN); 1 , same label data (MSNT); Taurirt (Beni Sicar), Melilla Maruecos, 1  1 , Pardo Alcaide lgt. (MNHN). NOT LOCATED: Borkine, 10.vi[19]29, Le Cerf, 1  (MNHN); Daya, 1  1 , Bedel lgt. (MNHN); Daja, 1  (MNHN). Redescription. Male (lectotype) (Figs 45–47). Remarkably convex, hemispherical; dorsal surface brownish, remarkably shiny. Head (Figs 45–47). Clypeus broadly rounded, almost semicircular, anterior angles vague, posterior angles elevated. Frontal horn well developed, long, distinctly furcate apically; its base outline elliptic in dorsal aspect, posterior edge of base situated at level of anterior part of eye; lateral margin of horn distinctly, sharply keeled from base to middle of tubercle. Clypeal disc with simple, coarse punctation. Oblique keels above eyes reaching approximately to half of eyes. Punctation of frons and vertex approximately same as on clypeus. Fronto-clypeal suture present. Eyes well developed, distinctly elliptic; genae exceeding eyes, arcuate posteriorly. HILLERT et al.: Bolbelasmus of the Western and Southern Mediterranean (Geotrupidae) 216 Pronotum (Figs 45–47) distinctly transverse, broadest just in basal quarter, entirely bordered; lateral margin evenly rounded, marginal carina not crenate, distinctly widened and elevated; lateral fovea nearly missing. Pronotal punctation distinct, double, consisting of coarse, deeply impressed punctures, punctures separated by approximately 2–3 times their diameter and intermixed with very fi ne ones. Pronotal base glabrous; lateral hornlike apophyses well developed, directed forwardly in dorso-lateral view; cavity between lateral and apical apophyses remarkably deep, kidney-shaped. Apical hornlike apophysis well developed, distinctly elevated, angled, directed upwards, subapical protrusion situated on outer side, basally. Scutellar shield elongate triangular, very fi nely punctate, shiny. Elytra approximately as long as wide, surface shiny; humeral umbone weak; between humerus and elytral suture seven distinctly impressed striae along entire elytral length, striae with coarse, regularly spaced punctures; intervals fl at, impunctate. Aedeagus. Parameres as in Fig. 60. Variability in males. Surface black to brownish; pronotal hornlike apophyses in medium developed and underdeveloped (hypothelic) specimens short, more or less straight, only simply rounded to almost acute apically. Female (Paralectotype No. 1). Head (Fig. 48). Clypeus broadly rounded, semicircular with straight anterior margin, anterior angles evenly rounded. Posterior angles of clypeal margin very weakly elevated. Frontal carina well developed, distinctly interrupted at middle, bituberculate at internal interruption, keeled laterally. Punctation of clypeus, frons, vertex and occiput simple, distinct; oblique keel above eyes distinct, extended to half of eyes; eyes well developed, distinctly elliptic, genae well developed, regularly arcuate. Pronotum (Fig. 48) distinctly transverse, lateral margin weakly widened, broadest just at base, surface distinctly and deeply punctate, punctation double except impunctate base, punctures separated by distance equal to their diameter and intermixed with very fi ne ones. Anterior protrusion distinctly transverse, sharply semicircular, lateral apophyses present. Measurements. Total body length 11.0–14.8 mm (lectotype, : 13.5 mm; paralectotype No.1, : 14.0 mm). Differential diagnosis. Refer to species key. Collecting circumstances. Prefers calcareous soils. Distribution. Mediterranean parts of Algeria and northern Morocco (Fig. 59). Remarks. The type locality of B. bocchus is ‘Algier’, the species has been recorded also from Spain (not Portugal: BRANCO 2005), northern Algeria and northern Morocco (cf. e.g. KRIKKEN 1977; BARAUD 1977, 1985, 1992; KRÁL et al. 2006; MARTÍN-PIERA & LÓPEZ-COLÓN 2000; RUIZ 2000). Regarding the presence in Spain, we consider the Iberian populations to belong to another species, described in this paper as B. brancoi sp. nov. The populations distributed in northern Tunisia and probably also in eastern parts of Algeria were at fi rst referred to another species, Bolbelasmus vaulogeri, later synonymized with B. bocchus by BOUCOMONT (1911) and considered a subspecies of B. bocchus by KRIKKEN (1977). We consider these populations to represent a valid species and identical to populations of Sicily described recently as B. romanorum Arnone & Massa, 2010 (syn. nov.). We also consider the populations native to Middle and South Tunisia, Libya (Tripolitania, Cyrenaica) (SCHATZMAYR 1946) and Egypt (Mariout) (PIC 1924) to be a distinct species, B. nikolajevi sp. nov., described in this paper. Acta Entomologica Musei Nationalis Pragae, 56(1), 2016 217 Finally, CHIKATUNOV & PAVLÍČEK (1997) reported B. bocchus from Israel, but we did not have the opportunity to study any Bolbelasmus material from the Levant countries. ERICHSON (1841) described Bolboceras bocchus based on a series of unknown number of specimens. He did not establish a type; thus in this study, we designated a lectotype () (Figs 45–47, 49) and two paralectotypes for fi xation of this taxon (Figs 48, 50–51). Bolboceras fi ssicornis Mulsant, 1843 was also described from ‘Algéria’ based on the same morphological characters of B. bocchus (‘tête armée d’une corne non mobile, bifi de au sommet [= head armed with immobile horn, furcate apically]’) (MULSANT 1843) and later synonymized with B. bocchus by MARSEUL (1857). This important character of B. bocchus is shown also in GORY (1841: table 71, fi gs 1, 1A). Bolbelasmus brancoi Hillert & Král sp. nov. (Figs 2, 9, 15, 22, 28, 34, 40, 59, 61) Bolboceras Bocchus [partim, populations from Spain]: JACQUELIN DU VAL (1863): 131 (catalogue). Bolboceras Bocchus: FUENTE Y MORALES (1926): 146 (distribution). Bolbelasmus bocchus [partim, populations from Spain]: WINKLER (1927): 1035 (catalogue); SEABRA (1943): 50 (catalogue); BARAUD (1977): 159 (monograph); RUANO MARCO et al. (1988): 40, fi g. p. 41 (distribution); LÓPEZCOLÓN et al. (1996): 3, 8, 10 (distribution, key); MARTÍN-PIERA & LÓPEZ-COLÓN (2000): 179, fi gs 44 A,C, 45 (monograph); MIESSEN & TRICHAS (2011): 182 (remarks). Bolbelasmus bocchus: BAGUENA CORELLA (1967): 262 (monograph); GONZÁLEZ PEÑA (1979): 23, fi gs 4–6, 8 (key, biology); VERDÚ et al. (1998): 245, fi gs 1–12 (larval description); AGOIS-BUSTAMANTE (1998): 19 (distribution); AGOIS-BUSTAMANTE (2003): 190 (distribution); VERDÚ et al. (2004): 514, fi g. 1; 515, fi g. 20; 518, fi g. 42 (larval illustrations). Bolbelasmus bocchus bocchus [partim, populations from Spain]: KRIKKEN (1977): 290 (revision); BARAUD (1985): 53 (monograph); BARAUD (1992): 44 (monograph); BRANCO (2005): 54 (taxonomic comments); KRÁL et al. (2006): 82 (catalogue); ARNONE & MASSA (2010): 407 (remarks, distribution). Type locality. Spain, Andalusia, Los Jesos, N slope of Sierra de Alhamilla, 37°05′N 02°17′W, 500 m a.s.l. Type material (143 specimens). SPAIN: ANDALUSIA AUTONOM. COMM.: ALMERIA PROV.: HOLOTYPE:  (DKCP), ‘Spain, (Andalusia), Los Jesos, N slope of Sierra de Alhamilla, 28.ii.2009, O. Hillert lgt.’ ALLOTYPE:  (DKCP), same data. PARATYPES: ALMERIA PROV.: ‘Espana, U. Almeria, Garrucha, Salzlachenufer, 07.iv.1986’, 1  (LSCN); ‘Spain, (Andalusia), Los Jesos, N slope of Sierra de Alhamilla, 13-27.ii.2013, O. Hillert lgt.’, 7  5  (OHCB); ‘Spain (Andalusia), Los Jesos, N slope of Sierra de Alhamilla, 30 km NE of Almeria, 28.ii.2009, O. Hillert lgt.’, 1  1  (ABCB), 1  (AHCL), 1  1  (ARCL), 1  1  (ERCS), 1  1  (GMCL), 1  1  (GNCA), 1  1  (HHCO), 1  1  (HKCS), 1  1  (LSCL), 4  6  (OHCB); ‘Spanien – Andalusien, Provinz Almeria, Umgebung Tabernas, a.L. [at light], 21.-28.iv.2004, leg. Steffen Schellhorn’, 1  1  (OHCB). GRANADA PROV.: ‘Spain, Granada, Andalucía, Hoya de Baza mts., 850 m asl., Salazar, 37°31.87’N 2°40.94’W, 4.v.2009, lgt. J. Darebník’, 1  (JSCP), 1  (PKCS); ‘Bco. del Espartal, Baza (Granada), 30SWG2854, F.S.Piñero leg., 3-iii-[19]90’, 1  1  (SZCM); ‘ESPAÑA, Granada, Bco. del Espartal, Baza, 3-iii-[19]90, F.S.Piñero’, 1  1  (SZCM); ‘Bananco del Espartal, BAZA 8.X.[19]89, GRANADA (España)’, F. S. Piñero leg.’, 1  (DKCP); ‘Bco. del Espartal, BAZA 8.X.[19]89, GRANADA (España), F. S. Piñero leg.’, 1  (DKCP); ‘Espagne (Granada), Lanjaron, 28.iii.[19]67, Baraud lgt.’, 1  (MNHN); ‘E, Prov. Granada, Los Olmos, 14.v.2002, H. Rietz lgt.’, 1  (ERCS); ‘Spain, Granada, Orgiva 12.iv.1960’, 1  1  (BMCP). ARAGON AUTONOM. COMM.: SARAGOSSA PROV.: ‘España, La Playa - Sastago - Zaragoza, 1.vi.2013’, 2  (GMCL); ‘SPAIN, Aragon, Zaragoza, Candasnos, 13.vi.2011’, 1  (IECA); ‘España, Bujaraloz, Zaragoza, 18.v.2013’, 2  (GMCL); ‘España 17.v.2011, Bujaralos, Zaragoza, Leg. Rafael Soler’, 1  (DJCP); ‘Spain, Zaragoza, Sastago 12.v.2010’, 1  (OBCB); ‘España, Bujaraloz, Zaragoza, v.2009’, 1  1  (GMCL); ‘España, Zaragoza, Sastago, 12.v.2007’, 1  (GMCL);‘ESPAÑA, Zaragoza, Bujaraloz, 11.V.[19]91, M. TOMÁS [legit]’, 4  (SZCM); ‘Espagne – 30/V/1973, Bujaraloz (Zaragoza), C. Dufay rec.’, 1  (MHNL); ‘Espagne, prov. Zaragoza, Bujaralos env., 18.5.2013, 1  (IBCF); ‘Ejea de los Caballeros (Zaragoza), 7.v.1982, J. HILLERT et al.: Bolbelasmus of the Western and Southern Mediterranean (Geotrupidae) 218 Figs 1–4. Male habitus in dorsal aspect. 1 – Bolbelasmus bocchus (Erichson, 1841) (Morocco: Azrou, Foret de Cedres, body length: 14.0 mm, NMPC); 2 – B. brancoi Hillert & Král sp. nov. (holotype); 3 – B. gallicus (Mulsant, 1842) (France: Bouches-du-Rhône, Barbentane, body length 13.7 mm, DJCP); 4 – B. howdeni Hillert & Král sp. nov. (holotype) . Acta Entomologica Musei Nationalis Pragae, 56(1), 2016 219 Figs 5–7. Male habitus in dorsal aspect. 5 – Bolbelasmus nikolajevi Hillert, Arnone, Král & Massa sp. nov. (holotype); 6 – B. vaulogeri (Abeille de Perrin, 1898) (Tunisia: Cap Bon peninsula, 6 km W of El Hawariah, 11.8 mm, DKCP); 7 – B. vaulogeri (Italy: Sicily, Menfi (Agrigento), Cinquanta, body length: 11.6 mm, MAPC). HILLERT et al.: Bolbelasmus of the Western and Southern Mediterranean (Geotrupidae) 226 Figs 21–26. Pronotum of male in dorsal aspect. 21 – Bolbelasmus bocchus (Erichson, 1841) (Morocco: Azrou, Foret de Cedres, NMPC); 22 – B. brancoi Hillert & Král sp. nov. (holotype); 23 – B. gallicus (Mulsant, 1842) (France: Bouches-du-Rhône, Barbentane, DJCP); 24 – B. howdeni Hillert & Král sp. nov. (holotype); 25 – B. nikolajevi Hillert, Arnone, Král & Massa sp. nov. (holotype); 26 – B. vaulogeri (Abeille de Perrin, 1898) (Tunisia: Cap Bon peninsula, 6 km W of El Hawariah, DKCP). Acta Entomologica Musei Nationalis Pragae, 56(1), 2016 227 Figs 27–32. Pronotum of male in frontal aspect. 27 – Bolbelasmus bocchus (Erichson, 1841) (Morocco: Azrou, Foret de Cedres, NMPC); 28 – B. brancoi Hillert & Král sp. nov. (holotype); 29 – B. gallicus (Mulsant, 1842) (France: Bouches-du-Rhône, Barbentane, DJCP); 30 – B. howdeni Hillert & Král sp. nov. (holotype); 31 – B. nikolajevi Hillert, Arnone, Král & Massa sp. nov. (holotype); 32 – B. vaulogeri (Abeille de Perrin, 1898) (Tunisia: Cap Bon peninsula, 6 km W of El Hawariah, DKCP). HILLERT et al.: Bolbelasmus of the Western and Southern Mediterranean (Geotrupidae) 228 Figs 33–44. Head in dorsal aspect, 33–38 – ; 39–44 – . 33 – Bolbelasmus bocchus (Erichson, 1841) (Morocco: Azrou, Foret de Cedres, NMPC); 34 – B. brancoi Hillert & Král sp. nov. (holotype); 35 – B. gallicus (Mulsant, 1842) (France: Bouches-du-Rhône, Barbentane, DJCP); 36 – B. howdeni Hillert & Král sp. nov. (holotype); 37 – B. nikolajevi Hillert, Arnone, Král & Massa sp. nov. (holotype); 38 – B. vaulogeri (Abeille de Perrin, 1898) (Tunisia: Cap Bon peninsula, 6 km W of El Hawariah, DKCP); 39 – B. bocchus (Morocco: Gaada de Debdou plateau., E of Rchida, DKCP); 40 – B. brancoi (allotype); 41 – B. gallicus (Spain: Andalusia, Aljaraque vill. env., JSCP); 42 – B. howdeni (allotype); 43 – B. nikolajevi (allotype); 44 – B. vaulogeri (Tunisia: Cap Bon peninsula, 6 km W of El Hawariah, DKCP). Acta Entomologica Musei Nationalis Pragae, 56(1), 2016 229 cosa le de Odon, 2.iii.1986, 1  (MSNG); Villavicosa le de Odon, 21.iv.1983, 1  (MSNG); Villavicosa de Odon (Madrid), 27.v.1977, 2 , J. I. López-Colón & Antonio Perucho lgt. (OHCB). VALENCIA AUTONOM. COMM.: VALENCIA PROV.: Ayora (Valencia), 16.v.1979, 1 , López-Colón lgt. (OHCB); La Vallesa, 2.iii.1997, 1  1  (GMCL). No further locality specifi ed: Espagne, 1  (NMPC); Hisp[ania]., 1  (NMPC); Spanien, 1  (SMTD). Redescription. Distinctly convex, surface black, shiny, pronotum covered with double punctation, elytra glabrous. Male. Head (Figs 3, 16, 23, 29, 35). Clypeus broadly rounded, nearly semicircular, anterior angles vague. Posterior angles of clypeal margin not elevated. Frontal horn well developed, long, conical, apex widened, triangular in frontal view, base conical, not keeled, apex simple, unmodifi ed. Punctation of clypeal disc simple, distinct. Oblique keels above eyes approximately reaching eyes. Punctation of frons and vertex same as the one on clypeus; fronto-clypeal suture present. Eyes distinctly elliptic, well developed; genae evenly rounded, semicircular. Pronotum (Figs 3, 16, 23, 29) distinctly transverse; broadest just in basal quarter, entirely bordered; lateral margin evenly rounded, marginal carina not crenate, distinctly widened and elevated; lateral fovea almost missing. Pronotal punctation distinct and double, consisting from well and deeply impressed punctures, except on basal part; punctures separated approximately by distance equal to their diameter, intermixed with very fi ne pinned punctures. Lateral hornlike apophyses well developed, conical, bent upwards and sidewards. Apical hornlike apophyses distinct but weakly elevated, obtuse-angled, directed 30° frontoapically. Both apical hornlike apophyses joined in keel. Scutellar shield triangular, as long as wide, shiny, densely punctate. Elytra approximately as long as wide, surface shiny, not punctate; humeral umbone weak; between suture and humerus seven distinct striae along entire elytral length, all striae with confl uent punctures; intervals convex. Parameres as in Fig. 62. Variability in males. Surface black to brownish; pronotal hornlike apophyses in medium developed and underdeveloped (hypothelic) specimens short, more or less straight, only simply rounded to almost acute apically. Female. Head (Figs 10, 41). Clypeus broadly rounded, nearly semicircular, anterior angles vague. Posterior angles of clypeal margin not elevated. Frontal carina well developed, not interrupted in middle, doubly s-shaped. Punctation of clypeus, frons, vertex and occiput simple, distinct; oblique keels above eyes distinct, elongated to base of eyes; eyes well developed, distinctly elliptic, genae well developed, arcuate anteriorly. Pronotum (Fig. 10) distinctly transverse, lateral margin weakly widened, broadest just basally, surface distinctly deeply punctate, punctation double except for base, punctures consisting of well and deeply impressed punctures, separated approximately by distance equal to their diameter, intermixed with very fi ne, pinned punctures. Anterior transverse protrusion distinct, sharply semicircular, lateral apophyses present. Elytron surface shiny, glabrous; humeral umbone weak. Measurements. Total body length 11.0–14.6 mm. Differential diagnosis. Refer to species key. Collecting circumstances. Prefers sandy soils. Distribution. Southern and south-western France, Portugal, North-western and Central Spain (Fig. 59). We provide fi rst records from Corsica and the Midi-Pyrénées Region of France. HILLERT et al.: Bolbelasmus of the Western and Southern Mediterranean (Geotrupidae) 230 Bolbelasmus howdeni Hillert & Král sp. nov. (Figs 4, 11, 17, 24, 30, 36, 42, 59, 63) Bolbelasmus gallicus [partim, populations from SE Spain (approximately Andalusia)]: BAGUENA CORELLA (1967): 262 (monograph); KRIKKEN (1977): 284 (revision); BARAUD (1977): 159 (monograph); BARAUD (1992): 45 (monograph); MARTÍN-PIERA & LÓPEZ-COLÓN (2000): 180 (monograph); KRÁL et al. (2006): 82 (catalogue). Type locality. Spain, Andalusia, Castillo de Castillar, Algeciras env., 10 km N of San Roque, 36°12′N 05°23′W, 78 m a.s.l. Type material (271 specimens). SPAIN: ANDALUSIA AUTONOM. COMM.: CÁDIZ PROV.: HOLOTYPE:  (DKPC), ‘Spain, Andalusia, Castillo de Castillar, Algeciras env., 10 km N of San Roque, 27.ii.2011, O. Hillert lgt.’ ALLOTYPE:  (DKCP), same data. PARATYPES: ‘Spain, 22.-28.2.2015, San Roque, Almoraima, N36°17’24.94” W5°23’40.77”, 136 m, lgt. P. Kylies’, 3  1  (DKCP), 22  7  (PKCS); same data but lgt. I. Martinů, 1  1  (DKCP), 9  4  (IMCO); same data but lgt. R. Muczka, 3  (RMCM); ‘Spain, Andalusia, Alcala de los Gazules, E of Cadiz, 01.-11.iii.2014 , Hillert lgt.’, 4  (OHCB); ‘Spain, Andalusia, E of Alcala, Sierra del Aljibe, E of Cadiz, 01.-11.iii.2014, Hillert lgt.’, 1  1  (OHCB); ‘España, Arcos de la Fontera, Cadiz, ex coll. A. Drumont, 9.xii.1989’, 1  (GMCL); ‘Spain, Arcos de la Frontera, Cádiz – B. Jordan leg., 9.xii.1989’, 1  (JNCS); ‘Spain, Andalusia, Castillo de Castillar, Algeciras env., 10 km N of San Roque, 13.–20.i.2015, Hillert lgt.’, 4  2  (OHCB); ‘Spain, Andalusia, Castillo de Castillar, Algeciras env., 10 km N of San Roque, 01.-11.iii.2014, Hillert lgt.’, 3  (OHCB); ‘Spain, Andalusia, Castillo de Castillar, Algeciras env., 10 km N of San Roque, 20.–27.ii.2012, O. Hillert lgt.’, 1  (OHCB); ‘Spain, Andalusia, Castillo de Castillar, Algeciras env., 10 km N of San Roque, 27.ii.2011, O. Hillert lgt.’, 5  (OHCB); ‘Spain, Cádiz distr., CASTILLO DE CASTILLAR vill.env., 36°18.186 N 05°25.401 E, 27.ii.2009, David Král lgt.’, 1  2  (DKCP); ‘Spain, Andalusia, Castillo de Castillar, Algeciras env., 10 km N of San Roque, 26.ii.2009, O. Hillert lgt.’, 1  (GNCA), 1  (HHCO), 2  1  (OHCB); ‘Spain - Cadiz, 13.1.2009, San Roque - Castillar env., lgt. P. Kylies’, 1  (PKCS); ‘Spain, Andalusia, Castillo de Castillar, Algeciras env., 10 km N of San Roque, 02.iii.2008, O. Hillert lgt.’, 4  (OHCB); ‘ESPANA Cadiz, Perros del Rey. S. Roque, 21.i.[19]87, M. SALER’, 6  (SZCM); ‘1.3.2014, N Castellar de la Frontera, Andalusia, Spain’, 1  1  (DKCP), 3  1  (ZLCK); ‘Chiclana, 6.iii.[18]90, Korb [lgt.]’, 1  (ZSMC); ‘Chiclana, [18]83, Korb [lgt.]’ 1  (ZSMC); ‘Spain, 22.-28.2.2015, Facinas env., lgt. P. Kylies’, 1  (PKCS); ‘Spain, La Suara, Jerez de la Frontera, Cádiz – A. Verdugo leg., 5.v.1988’, 2 , same data but 18.x.1987, 1  (JUCS); ‘Spain, Andalusia, 10 km NE of Medina, E of Cadiz, 01.-11.iii.2014, O. Hillert lgt.’, 3  2  (OHCB); ‘Spain, Pinar del Rey, San Roque, Cadiz, 26-i-2013., J. Navarro leg.’, 1  1  (AHCL), 1 ,1  (OBCB), same but ‘27-xii-2014, J. Mª Urbano leg.’, 1  1  (JUCS), same but ‘20-xii-2014, J. Mª Urbano & J. Navarro leg.’, 6  3  (JNCS), 4  2  (AHCL), 11  8  (JUCS), same but ‘10-xi-2014, J. Navarro leg.’, 1  (JUCS), same but ‘13-iii-2014, J. Navarro leg.’, 1  (JUCS), same but ‘26-i-2014, J. Mª Urbano leg.’, 2  2  (JUCS), same but ‘14-XI-2015, J. Navarro & J. Mª Urbano leg.’, 4  3  (JUCS), 4  3  (JNCS); ‘Spain, Pinar de l Rey, Tajo del Pajarraco, San Roque, Cádiz – J. Navarro leg., 22.xii.2012’ 5  2  (JNCS), 2.iv.2011, 2  (JNCS); ‘Spain, 22.-28.2.2015, Tarifa, Punta Paloma, N36°04’03.96” W5°42’08.47”, 56 m, lgt. P. Kylies’, 2  (PKCS); same data but lgt. I. Martinů, 3  1  (IMCO); same data but R. Muczka, 3  1  (RMCM); ‘Spain - Cadiz, 14.-16.2.2008, San Roque, Almorama, lgt. P. Kylies’, 1  (PKCS); ‘E - Cádiz / San Roque / 13.xi.[19]93 Ramirez lg.’, 1  (SZCM); ‘Espania, San Roque, Cadiz, 13.xi.1988, Jose Luis Torres lgt.’, 1  1  (OHCB); ‘Esp., SAN ROQUE, Cádiz, 6.1987, J. Ramírez lgt.’, 2  1  (RCCP); ‘E, Andalusia, Cadiz, Pinar de San Roque, 2.iii.86, Hendrich/Mendez lgt.’, 2  (OHCB) , 1  (VTCZ); ‘E, Andalucia, Cadiz, Pinar d. San Roque, b. San Roque, 2.iii.86, lgt. Hendrich/Mendez’, 4  (LHCB); ‘SPAIN-Cádiz, S. Roque, 26.i.1984’, 1  (SZCM); ‘San Roque – Cadiz, España sur, leg. Mendez T., 17.iii.[19]84’, 1  (JHCP); ‘Spanien, Andalusien, San Roque, 17.iv.1983, J.L. Torres-Mendez lgt.’, 4  1  (LHCB); ‘San Roque, Cadiz, Spain, 30-I.1983, V. Ortuño lgt.’, 1  (VMCP); ‘Spain, Andalusia, NW of Tarifa, Algeciras env., 13-20.i.2015, O. Hillert lgt.’, 6  2  (OHCB); ‘Spain, Andalusia, NW of Tarifa, pine forest, Algeciras env., 01.-11.iii.2014, Hillert lgt.’, 2  2  (OHCB); ‘Spain, Andalusia, NW of Tarifa, beach sand, Algeciras env., 01.-11.iii.2014, Hillert lgt.’, 5  10  (OHCB); ‘Spain, Andalusia, NW of Tarifa, Algeciras env., 13-27.ii.2013, O. Hillert lgt.’, 3  (OHCB); ‘Spain, Montehumada, Tarifa, Cádiz – J. Navarro leg., 22.xii.2012’, 2  1  (JNCS); ‘Spain, Andalusia, Sanlucar de Barrameda, NW of Jerez, 26.ii.2011, O. Hillert lgt.’, 1  2  (OHCB); ‘Tarifa (Cadiz), 11.iv.1980, Lopez Colon leg’, 1  (JLCM). GRANADA PROV.: ‘Spain, Andalusia, 5 km E Acta Entomologica Musei Nationalis Pragae, 56(1), 2016 231 of Granada, Sierra de la Yedra, 01.-11.iii.2014, Hillert lgt.’, 1  (OHCB); ‘España, La Sagra, Granada, Andalucia, v.1988’, 1  1  (GMCL). – JAÉN PROV.: ‘Spain, Las Minas. Arrayanes. Linares, Jaén, ii-1997, A. Hidalgo leg.’, 1  (AHCL); ‘Spain, Miranda del Rey, Jaén, 13-xi-2015, Javier Castillo & Antonio Hidalgo leg.’, 4  2  (JCCL), 3  2  (AHCL).‘Spain, Reserva las Viboras, Sierra Morena. Montizón, Jaén, 9-iv-1995, A. Hidalgo leg.’, 1  (AHCL). MÁLAGA PROV.: ‘Spain, Andalusia, Sierra Palmitera, Puerto de Alijar, W of Mabella, 01.-11.iii.2014, Hillert lgt., 1  (OHCB)’. CASTILE-LA MANCHA AUTONOM. COMM.: ALBACETE PROV.: ‘Molinicos, 10.v.[18]95’, 1  2  (ZFMK). GIBRALTAR: ‘Gibraltar., J. J. Walker [lgt.]’ 1 , 2  (BMHN). Description. Male (holotype). Distinctly convex, surface black, shiny, pronotum covered with double punctation, elytra glabrous. Head (Figs 4, 17, 24, 30, 36). Clypeus broadly rounded, nearly semicircular, anterior angles vague. Posterior angles of clypeal margin not elevated. Frontal horn well developed, long, base conical, not keeled, apex simple, unmodifi ed. Punctation of clypeal disc simple, distinct. Oblique keels above eyes approximately reaching eyes. Punctation of frons and vertex same as on clypeus; fronto-clypeal suture present. Eyes distinctly elliptic and well developed; genae evenly rounded, semicircular. Pronotum (Figs 4, 17, 24, 30) distinctly transverse; broadest just in basal quarter, entirely bordered; lateral margin evenly rounded, marginal carina not crenate, distinctly widened and elevated; lateral fovea almost missing. Pronotal punctation distinct, double, consisting of well and deeply impressed punctures except for basal part, punctures separated approximately by distance equal to their diameter, intermixed with very fi ne, pinned punctures. Lateral hornlike apophyses well developed, conical, bent up and sidewards. Apical hornlike apophyses distinct but weakly elevated, obtuse angled, directed 45° frontoapically. Both apical hornlike apophyses joined together in keel-like seam. Scutellar shield triangular, approximately as long as wide, shiny, densely punctate. Elytra approximately as long as wide, surface shiny, glabrous; humeral umbone weak; between suture and humerus seven distinct striae along entire elytral length, all striae with confl uent punctures; intervals convex. Parameres as in Fig. 63. Variability in males. Surface black to brownish; pronotal hornlike apophyses in medium developed and underdeveloped (hypothelic) specimens short, more or less straight, only simply rounded to almost acute apically. Female (allotype). Head (Figs 11, 42). Clypeus broadly rounded, nearly semicircular, anterior angles vague. Posterior angles of clypeal margin not elevated. Frontal carina well developed, not interrupted in middle, doubly s-shaped. Punctation of clypeus, frons, vertex and occiput simple, distinct; oblique keels above eyes distinct, elongated to base of eyes; eye well developed, distinctly elliptic, genae well developed, arcuate anteriorly. Pronotum (Fig. 11) distinctly transverse, lateral margin weakly widened, broadest just basally, surface distinctly deeply punctate, doubly except for basal part, punctures consisting of well and deeply impressed punctures, separated approximately by distance equal to their diameter, intermixed with very fi ne, pinned punctures. Anterior transverse protrusion distinct, sharply semicircular, lateral apophyses present. Elytron surface shiny, glabrous; humeral umbone weak. Measurements. Total body length 11.3–15.2 mm (holotype, : 14.6 mm; allotype, : 15 mm). HILLERT et al.: Bolbelasmus of the Western and Southern Mediterranean (Geotrupidae) 232 Differential diagnosis. Refer to species key. Etymology. Patronymic; named in honour of our friend, the late Henry F. Howden (Ottawa, Canada), a renowned specialist in world Scarabaeoidea. Collection circumctances. Prefers sandy soils. Distribution. South-eastern parts of Spain and Gibraltar (Fig. 59). Bolbelasmus nikolajevi Hillert, Arnone, Král & Massa sp. nov. (Figs 5, 12, 18, 25, 31, 37, 43, 57, 59, 64) Bolbelasmus bocchus: PIC (1924): 126 (note, distribution); SCHATZMAYR (1946): 50 (distribution); ALFIERI (1976): 206 (distribution). Bolbelasmus bocchus vaulogeri: KRIKKEN (1977): 290, fi gs 18–19 (revision); BARAUD (1992): 53 (monograph); KRÁL et al. (2006): 82 (catalogue); ARNONE & MASSA (2010): 411, fi g. 12 (distribution, note). Type locality. NW Tunisia, Sidi Bouzid gov.,7.5 km SWW of Hajeb el Ayoun, by road, 35°22′N, 09°28′E, 340 m a.s.l. Type material (126 specimens). TUNISIA: SIDI BOUZID PROV.: HOLOTYPE:  (DKPC), ‘NW Tunisia, Sidi Bouzid gov., 7.5 km SWW of, HAJEB EL AYOUN, by road, 29.iii.2014, 35°22’N, 09°28’E, 340 m, David Král lgt.’ ALLOTYPE:  (DKCP), same data. PARATYPES: EGYPT: ALEXANDRIA PROV.: ‘Egypt, Alexandria, 1958’, 1  (NMPC); ‘Egypt, Mariout, 4.iii.1924, coll. Alfi eri’, 1  (NHMB). LIBYA: AZ ZAWIYAH PROV.: ‘Afrika, Libye, SURMAN - viii.[19]88, Č. Sobota lgt.’, 1  1  (JDCC). BENGHAZI PROV.: ‘Libya, Benghazi, 8.3.1987, lgt. Háva [p]’, 1  (DKCP), 1  (JHCP); ‘Libia, Benghazi, 1986 V. Poláček lgt. [p]’, 1  (SJCP). BUTNAN PROV.: ‘Libya, Cirenaica, Tobruk III, 32°5‘1“ N, 23°58‘ 35“ E, Geo C. Kruger lgt., R.U. Agrario’, 2  2  (MSNG). TRIPOLI PROV.: ‘Afr. Libya – Tadjiura, 04.1986, Houška lgt.’, 2  (JMCH); ‘Libya, Tagiura, červenec [= July], [19]85, Voleman [lgt.]’, 2  (VTCZ); ‘Libya, Tagiura, srpen [= August], [19]85, Voleman [lgt.]’, 1  (VTCZ); ‘AFRIKA, Libya, Tarabulus vii.[19]88, V. Poláček lgt.’, 1  (JMCP); Foualgha, 8.4.1983, M. Dudycha [lgt.] [handwritten]’, 1  (RCCP). Not located: ‘Libya, Takadria, srpen [= August], [19]84, Voleman [lgt.]’, 1  (VTCZ). TUNISIA: BIZERTE PROV.: ‘Biserta, Tunisia, De Vauloger, Nevinson Coll., 1918-14, Bolboceras, vaulogeri Ab., types, Type, H. T.’, 1  (BMNH) [not syntype of B. vaulogeri, type label added later; M. Barclay pers. comm.]. GABÈS PROV.: ‘Gabes, 1897’, 2  (MNHN); ‘Tunisie, Mareth, R. Demofl ys lgt., i. [19]49’, 1  (MNHN); ‘Tunisia, Gov. Gabes, Matmata, 19.iv.2013, Leg. G.Sabatinelli, At light’, 1  1  (GSCT); ‘Matmata, iii.2014’, 1  1  (GSCT); ‘TUNISIA: 15.4.1996, TAMEZRET, 13 km W of Matmata, J. BATELKA lgt.’, 2  (JBCP); ‘E Tunisia, Gabes, 5 km NEE of Oudref, Matwiya plage, 24.–25.iii.2014, 34°01‘N 10°00‘E, O. Hillert lgt.’, 7  8  (OHCB); ‘E Tunisia, Gabes gov., 5 km NEE of Oudref, MATWIYA plage, ca. 20 m, 24.–25.iii.2014, 34°01‘N, 10°00‘E, David Král lgt.’, 2  3  (DKCP), same data but Petr Šípek lgt., 3  2  (CUPC), 1  [in pure alcohol] (CUPC); ‘TN-Gabes, dintorni, 13.IV.[19]95 leg. S. Ziani’, 1  (SZCM). GAFSA PROV.: ‘Tunisie, Gafsa, Chretien [lgt.], 1910’, 1  (MNHN); ‘Tunisie, Gafsa, A. Weiss [lgt.], 1904’, 1  (MNHN); ‘Gafsa, 1897’, 1  (OHCB);‘Tunis, Métlaoui, C. Dumont’, 1  (DKCC). KASSERINE PROV.: ‘T., Thelepte, Mai 1941, R. Demofl ys [lgt.]’, 2  1  (MNHN); ‘Tun., Sbeitla, Mars 1941, R. Demofl ys [lgt.]’, 2  (MNHN). KEBILI PROV.: ‘TUNISIA, 3.-4. iv.2008 / 45 km E of Douz, Wadi Raml, / N33°31’ E09°30’ / J. Batelka & J. Straka lgt.’, 1  (JBCP). KEF PROV.: ‘Tunisia NW, Le Kef, NABEUR env., 16.4.2001, lgt. M. Snížek’, 1  (VKCC). MEDENINE PROV.: ‘E Tunisia, Medenine, Djerba island, 5 km E of Midoun, sea coast, 26.iii.2014, 33°49‘N 11°02‘E, O. Hillert lgt.’, 1  (OHCB); ‘TN-Medenine, str. per Toujan, 13.IV.[19]95 leg. S. Ziani’, 4  (SZCM). NABEUL PROV.: ‘Tunisia, (Nabeul), Cap Bon peninsula, 6 km W of El Hawariah, sea coast, 23.iii.2014, 37°03‘N 10°57‘E, O. Hillert lgt.’, 4  (OHCB); ‘NE Tunisia, Nabeul gov., Cap Bon peninsula, 6 km W of EL HAWARIAH, sea coast, 23.iii.2014, 37°03‘N, 10°57‘E, ca. 3 m, David Král lgt.’, 2  4  (DKCP), same data but Petr Šípek lgt., 1  1  [partially demaged spec. in pure alcohol] (CUPC); ‘Tunisia, Cap Bon IV.1960, G. Frey’, 1  (NHMB). SFAX PROV.: ‘Tunisie, Ketana [Bir Ketana], iv.[19]50, R. Demofl ys [lgt.]’, 1  (MNHN); ‘Tunisie, La Hencha, iii.[19]42, R. Demofl ys [lgt.]’, 1  (MNHN); ‘Tunisia 1896, Sfax, de Vauloger [lgt.], Nevinson Coll., 1918-14’, 1  1  (BMNH). SIDI BOUZID PROV.: ‘NE Tunisia, Sidi Bouzid, 7.5 km SWW of Hajeb el Ayoun by road, 29.iii.2014, 35°22‘N 09°28‘E, O. Hillert lgt.’, 7  8  (OHCB); ‘NW Tunisia, Sidi Bouzid gov., 7.5 km SWW of, HAJEB EL AYOUN, by road, 29.iii.2014, 35°22‘N, 09°28‘E, 340 m, David Král lgt.’, 4  2  (DKCP), 1  1  (CUPC); ‘Tunisie, Maknassy, A. Dumont [lgt.], 1928’, 5  2  (MNHN); ‘Tunisia, Bled Gamouda [Bilad Acta Entomologica Musei Nationalis Pragae, 56(1), 2016 233 Gamouda], de Vauloger [lgt.], Nevinson Coll., 1918-14.’, 1  (BMNH). TOZEUR PROV.: ‘Tunis, Tozeur, C. Dumont [lgt.], 1929’, 2  (MNHN). No further locality specifi ed: ‘Tunisie, Septent[rionalis], Schaedlin’, 1  (MNHN). Description. Male (holotype). Distinctly convex, surface brownish or light-brown, shiny, pronotum covered with double punctation, elytron very fi ne microsculptured basally, microsculture missing apically. Head (Figs 5, 18, 25, 31, 37, 57). Clypeus broadly rounded, nearly semicircular, anterior angles vague. Posterior angles of clypeal margin elevated. Frontal horn well developed, long, distinctly furcate, base triangularly shaped in dorsal view, posterior edge of base situated at height of center of eye, side margin of tubercle keeled from base to apex. Punctation of clypeal disc simple, distinct. Oblique keels above eyes reaching approximately to half of eyes. Punctation of frons and vertex same as on clypeus; frontoclypeal suture present. Eyes distinctly elliptic and well developed; genae arcuate posteriorly, extending eyes laterally. Pronotum (Figs 5, 18, 25, 31, 37, 57) distinctly transverse; broadest just in basal quarter, entirely bordered; lateral margin evenly rounded, marginal carina not crenate, distinctly widened and elevated; lateral fovea almost missing. Pronotal punctation distinct, double, consisting of deeply impressed punctures, punctures separated by approximately 2–3 times their diameter, intermixed with very fi ne, pinned punctures. Base of pronotum glabrous. Lateral hornlike apophyses well developed, bent distinctly upwards and sidewards in lateral and dorsal view; between lateral and apical apophyses distinct, deeply oval concavity. Apical hornlike apophyses distinctly elevated, angled, directed 30° frontoapically, subapical protrusion of apical apophyses laterally near apex on outside. Scutellar shield triangular elongate, very fi nely punctate, shiny. Elytra approximately as long as wide, surface shiny, glabrous; between suture and humerus 7 distinct striae along entire elytral length, all striae with distinctly separated punctures; intervals fl at; humeral umbone weak. Parameres as in Fig. 64. Variability in males. Surface black to brownish; pronotal hornlike apophyses in medium developed and underdeveloped (hypothelic) specimens short, more or less straight, only simply rounded to almost acute apically. Female (allotype). Head (Figs 12, 43). Clypeus broadly rounded, semicircular with straight anterior margin, anterior angles evenly rounded. Posterior angles of clypeal margin very weakly elevated. Frontal carina well developed, distinctly interrupted in middle, bituberculate on internal interruption, laterally keeled on each side. Punctation of clypeus, frons, vertex and occiput simple, distinct; oblique keels above eyes distinct, elongated to half of eyes; eye well developed, distinctly elliptic, genae well developed, arcuate anteriorly and posteriorly. Pronotum (Fig. 12) distinctly transverse, lateral margin weakly widened, broadest just basally, surface distinctly deeply punctate, double except of basal part, punctures consisting of well and deeply impressed punctures, separated approximately by distance equal to their diameter, intermixed with very fi ne, pinned punctures. Anterior transverse protrusion distinct, sharply semicircular, lateral apophyses present. Elytron surface shiny, glabrous; humeral umbone weak. Measurements. Total body length 12.0–15.1 mm (holotype, : 14.6 mm; allotype, : 14.8 mm). HILLERT et al.: Bolbelasmus of the Western and Southern Mediterranean (Geotrupidae) 234 Differential diagnosis. Refer to species key. Etymology. Patronymic; named in honour of our friend Georgy V. Nikolajev (Al-Farabi State University, Almaty, Kazakhstan), a renowned specialist in world Scarabaeoidea. Collection circumstances. Prefers sandy and/or loess soils. Distribution. Mediterranean Egypt, Libya (Tripolitania, Cyrenaica) and Tunisia (Fig. 59). Bolbelasmus vaulogeri (Abeille de Perrin, 1898) stat. restit. (Figs 6, 7, 13, 19, 20, 26, 32, 38, 44, 52–56, 58, 59, 65) Bolboceras Vaulogeri Abeille de Perrin, 1898: 254 (original description). Bolbelasmus Vaulogeri: BOUCOMONT (1912): 17 (catalogue, as synonym of B. bocchus); WINKLER (1927): 1035 (catalogue, synonym of B. bocchus). Bolbelasmus romanorum Arnone & Massa, 2010: 403, fi gs 1–5, 8 (original description). Type locality: ‘Sicily, Balestrate, foce Calatubo (Palermo)’. New synonym. Bolbelasmus unicornis: BALLERIO (2008): 32 (distribution). Type locality. Tunisia, ‘Bizerte’. Type material examined (20 specimens). Bolbelasmus vaulogeri: TUNISIA: BIZERTE prov.: LECTOTYPE (present designation):  (MNHN), ‘Bizerte, de Vauloger, Bolboceras Vaulogeri ab., Museum Paris 1919, coll A de Perrin, Bolboceras Vaulogeri ab., Museum Paris, 1942 coll. Dr. A Chobaut’. PARALECTOTYPES: No. 1,  (MNHN), ‘Bizerte, de Vauloger, Vaulogeri Ab., Type’; No. 2,  (MNHN), ‘Bizerte de Vauloger sp. nov??, tuniseus Vaul. (sp. litteus), Museum Paris 1942, coll. Dr. A Chobaut, Bolbocerus Vaulogeri ab’; Nos 3–4,  (MNHN), ‘Bizerte, de Vauloger , Vaulogeri Ab., Type’; No. 5,  (ZSMC), ‘Bizerte [18]93 Vaulog[er].’; No. 6,  (BMNH), ‘Bizerte, de Vauloger, Nevinson Coll., 1918-14., Cotype’; No. 7,  (BMNH), ‘Bizerte, de Vauloger, Nevinson Coll., 1918-14., Cotype’; see Figs 52–56. Bolbelasmus romanorum: ITALY: SICILY PROV.: HOLOTYPE: 1  (MSNG), ‘Italy, Sicily, Balestrate, foce Calatubo (Palermo), 17.iii.1974, B. Massa lgt.’. ALLOTYPE: 1  (MSNG), ‘Sicily, Pedalino (Ragusa) 1.v.1972, A. Monastra lgt.’. PARATYPES: 1  5  (MAPC), ‘Menfi (Agrigento), loc. Cinquanta, 13.xi.2010, 4.iv.2010, 28.iii.2010, 20.iii.2010, 14.xi.2009, 6.xi.2009, A. Carapezza lgt.’; 1  (MACP), ‘Vendicari Nature Reserve (Siracusa) 18.iv.2009, B. Massa lgt.’; 1  (VACP), ‘Marausa (Trapani) 18.xii.1983, V. Aliquò lgt.’; 1  (OHCB), ‘Bosco Santo Pietro Nature Reserve (Catania), M. Zafarana lgt.’; 1  1  (MZUC) ‘Sicily’. Additional material examined (77 specimens). ALGERIA: No further locality specifi ed: Algerie, Dr. Thièbault, 1  (MNHN). ITALY: SICILY PROV.: Menfi (Agrigento), loc. Cinquanta, 19.iii.[20]13, 1 , M. Arnone lgt. (MAPC); Terrasini, Capo Rama Nature Reserve 27.iii.[20]12, 1 , R. Lo Duca lgt. (MAPC); Menfi , loc. Cinquanta, 10.iv.2011, 1 , A. Carapezza lgt. (DJCP); Niscemi (Caltanissetta), 1 , M. Zafarana lgt. (MSNM); Palermo, 1  (MZUC); Piazza Armerina (Enna), 1  (MZUC); Termini Imerese (Palermo), 1  (MZUC). TUNISIA: BIZERTE PROV.: SW env. of Edahra, 31.iii.2014, 37°12′N 09°20′E, 3  5 , O. Hillert lgt. (OHCB); SW env. of Edahra, 37°12′N 09°20′E, ca. 65 m, 31.iii.2014, 4  3 , David Král lgt. (DKCP), 3  2  (CUPC). JENDOUBA PROV.: Tabarka, Zone touristique, 2.v.2005, 1 , M. Liebscher lgt. (MLCD). NABEUL PROV.: Cap Bon peninsula, 6 km W of El Hawarith, sea coast, 23.iii.2014, 37°03′N 10°57′E, 3  2 , O. Hillert lgt. (OHCB); Cap Bon peninsula, 6 km W of El Hawariah, sea coast, 37°03′N 10°57′E, ca. 3 m, 23.iii.2014, 3  6 , David Král lgt. (DKCP); Cap Bon peninsula, 2 km NW of Douela by road, 23.–24.iii.2014, 36°50′N 10°35′E, 13  2 , O. Hillert lgt. (OHCB); Cap Bon peninsula, 2 km NW of Douela by road, 36°49′N 10°35′E, ca. 130 m, 23.–24.iii.2014, 4  2 , David Král lgt. (DKCP), 1  [partially damaged, in pure alcohol] (CUPC); Cap Bon peninsula, 6 km SW of Zouiet el Mgaiez by road, 23.iii.2014, 36°58′N 10°55′E, 3  2 , O. Hillert lgt. (OHCB); Cap Bon peninsula, 6 km SW of Zaouiet El Mgaiez by road, 36°49′N 10°55′E, ca. 45 m, 23.iii.2014, 1  2 , David Král lgt. (DKCP); Cap Bon, 1960, 1 , G. Frey [lgt.] (NHMB); Cap Bon, 1894, Museum Paris, 1919, 1 , coll. A. de Perin (MNHN). No further locality specifi ed: Tunisia, Salt Lake, Kveten [May] 2005, 1 , Volcano lgt. (OBCB); Tunisia, Solunan, Museum Paris 1993, 1 , coll J. Baraud (MNHN). Redescription. Male (lectotype) (Figs 52–54). Distinctly convex, surface black, shiny, pronotum covered with double punctation, elytron glabrous. Acta Entomologica Musei Nationalis Pragae, 56(1), 2016 235 Figs 45–51. Types and labels of Bolbelasmus bocchus (Erichson, 1841), deposited in MNHB. 45 – male habitus in dorsal aspect (lectotype); 46 – male habitus in frontal aspect (lectotype); 47 – male habitus in lateral aspect (lectotype); 48 – female habitus in dorsal aspect (paralectotype); 49 – labels of lectotypes; 50 – labels of paralectotype No. 1; 51 – labels of paralectotype No. 2. HILLERT et al.: Bolbelasmus of the Western and Southern Mediterranean (Geotrupidae) 242 surface brownish, maximal body length 14 mm. Southern parts of Central Europe, Alsace, Italy, Balkan Peninsula, Poland, Russia and Ukraine. ................................... .................................................................... B. unicornis (Schrank von Paula, 1789) 4 (3) Frontal horn bending not forwards (Figs 19–20); transversal clypeal carina absent; scutellar shield triangular longer than wide; body surface blackish, maximal body length 13 mm (Figs 6–7, 38); aedeagus as in Fig. 65; Algeria, Italy (Sicily) and Tunisia (Fig. 59). ............................................... B. vaulogeri (Abeille de Perrin, 1898) 5 (2) Frontal horn furcate apically; genae more or less angulate anterolaterally; oblique keel above eyes reaching only to anterior part of eyes; head not strongly constricted behind eyes. 6 (7) Frontal horn short and not longer than broad, horn apex approximately as wide as horn base (Figs 2, 15, 22); pronotal punctation denser with exception of basal part, punctures separated approximately by distance equal to their diameters; pronotal base with distinct visible but fi ne, regularly pinned points; elytral striae well impressed, visible as complete lines; elytral intervals moderately convex; aedeagus as in Fig. 61; Spain (Fig. 59). ................................. B. brancoi Hillert & Král sp. nov. 7 (6) Frontal horn long, longer than broad, horn becoming gradually thinner from base to apex (Figs 1, 5, 14, 18, 21, 25, 27, 31, 33, 37, 57); pronotal punctation sparser, punctures separated by approximately three times their diameters; pronotal base with very fi ne, weakly visible and well separated pinned points; elytral striae not impressed, visible as row of punctures; elytral intervals fl at; nothern Africa. 8 (9) Base of frontal horn situated at level of anterior part of eyes; horn sharply keeled from base to half of its length in dorsal aspect (Figs 1, 14, 21, 27, 33); medial hornlike apophysis of pronotum directed vertically upwards in lateral aspect, subapical protrusion situated basally; lateral hornlike apophyses of pronotum directed forwards and weakly divergent; aedeagus as in Fig. 60; Algeria and Morocco (Fig. 59). ............................................................................... B. bocchus (Erichson, 1841) 9 (8) Base of frontal horn situated at level of posterior part of eyes; horn broadly keeled from base to apex in dorsal view (Figs 5, 18, 25, 31, 37, 57); medial hornlike apophysis of pronotum directed obliquely forwards in lateral aspect, subapical protrusion situated near apex; lateral hornlike apophyses of pronotum directed forwards; aedeagus as in Fig 64; Egypt, Libya and Tunisia (Fig. 59). ................................................ ............................................... B. nikolajevi Hillert, Arnone, Král & Massa sp. nov. 10 (1) Scutellar shield densely punctate; frontal horn simple apically. 11 (12) Apex of frontal horn approximately as wide as horn base, apex broadly acute-angled, strongly fl attened on front part of apex (Figs 3, 16, 23, 29, 35); aedeagus as in Fig. 62; France (including Corsica), Portugal and Spain (Fig. 59). ................................... ....................................................................................... B. gallicus (Mulsant, 1842) 12 (11) Apex of frontal horn becoming gradually thinner from base to apex, apex acute-angled, only weakly fl attened on front part of apex (Figs 4, 17, 24, 30, 36); aedeagus as in Fig. 63; Spain, Gibraltar (Fig. 59). ................ B. howdeni Hillert & Král sp. nov. Acta Entomologica Musei Nationalis Pragae, 56(1), 2016 243 Identifi cation key for less developed males 1 (10) Scutellar shield impunctate or weakly shallowly punctate, frontal horn simple or furcate apically. 2 (5) Frontal horn simple; genae regularly rounded; oblique keel above eyes extending over whole eyes length; head strongly constricted behind eyes. 3 (4) Scutellar shield triangular, approximately as long as wide; body surface brownish, maximal body length 12 mm. Southern parts of Central Europe, Alsace, Italy, Balkan Peninsula, Poland, Russia and Ukraine. .... B. unicornis (Schrank von Paula, 1789) 4 (3) Scutellar shield triangular, longer than wide; body surface blackish, maximal body length 11 mm; aedeagus as in Fig. 65; Algeria, Italy (Sicily) and Tunisia (Fig. 59). ....................................................................... B. vaulogeri (Abeille de Perrin, 1898) 5 (2) Frontal horn furcate apically; genae more or less angulate anterolaterally; oblique keel above eyes reaching only to anterior part of eyes; head not strongly constricted behind eyes. 6 (7) Pronotal punctation denser with exception of basal part, punctures separated approximately by distance equal to their diameters; pronotal base with distinct visible but fi ne, regularly spaced pinned points, elytral striae well impressed, visible as complete lines; elytral intervals moderately convex; aedeagus as in Fig. 61; Spain (Fig. 59). ...................................................................... B. brancoi Hillert & Král sp. nov. 7 (6) Pronotal punctation sparser, punctures separated by approximately three times their dia meters pronotal base with very fi ne, weakly visible and well separated pinned points; elytral striae not impressed, visible as row of punctures; elytral intervals fl at; northern Africa. 8 (9) Base of frontal horn situated at level of anterior part of eyes; horn sharply keeled basally only; aedeagus as in Fig. 60; Algeria and Morocco (Fig. 59). ....................... ...................................................................................... B. bocchus (Erichson, 1841) 9 (8) Base of frontal horn situated at level of posterior part of eyes; horn broadly keeled from base to apex in dorsal view; aedeagus as in Fig. 64; Egypt, Libya and Tunisia (Fig. 59). ............................... B. nikolajevi Hillert, Arnone, Král & Massa sp. nov. 10 (1) Scutellar shield densely punctate; frontal horn simple apically. 11 (12) Apex of frontal horn fl attened on front part of apex; aedeagus as in Fig 62; France (including Corsica), Portugal and Spain (Fig. 59). ....... B. gallicus (Mulsant, 1842) 12 (11) Apex of frontal horn regularly arcuate, not fl attened on front part of apex; aedeagus as in Fig. 63; Spain, Gibraltar (Fig. 59). ............ B. howdeni Hillert & Král sp. nov. Identifi cation key for females 1 (10) Scutellar shield impunctate, or weakly shallowly punctate. 2 (5) Frontal tubercle simple (Figs 41, 42, 44); genae regularly rounded; oblique keel above eyes over whole length of eyes; head strongly constricted behind eyes. HILLERT et al.: Bolbelasmus of the Western and Southern Mediterranean (Geotrupidae) 244 3 (4) Scutellar shield triangular, approximately as long as wide; body surface brownish, minimal body length 12 mm. Southern parts of Central Europe, Alsace, Italy, Balkan Peninsula, Poland, Russia and Ukraine. .... B. unicornis (Schrank von Paula, 1789) 4 (3) Scutellar shield triangular longer than wide; body surface blackish, minimal body length 11 mm (Fig 13); Algeria, Italy (Sicily) and Tunisia (Fig. 59). ........................ ....................................................................... B. vaulogeri (Abeille de Perrin, 1898) 5 (2) Frontal tubercle furcate apically (Figs 39–40, 43); genae more or less angulate anterolaterally; oblique keel above eyes reaching only to anterior part of eyes; head not strongly constricted behind eyes. 6 (7) Pronotal punctation denser with exception of basal part, punctures separated approximately by distance equal to their diameters; pronotal base with distinct visible but fi ne, regularly pinned points; elytral striae well impressed, visible as complete lines; elytral intervals moderately convex (Fig. 9); Spain (Fig. 59). ................................... ............................................................................. B. brancoi Hillert & Král sp. nov. 7 (6) Pronotal punctation sparser, punctures separated by approximately three times their diameters pronotal base with very fi ne weak visible and well separated pinned points; elytral striae not impressed, visible as row of punctures; elytral intervals fl at; northern Africa. 8 (9) Base of frontal tubercle situated at level of anterior part of eyes; tubercle sharply keeled basally only in dorsal view (Fig. 8); Algeria and Morocco (Fig. 59). ............ ...................................................................................... B. bocchus (Erichson, 1841) 9 (8) Base of frontal tubercle situated at level of posterior part of eyes; horn broadly keeled from base to apex in dorsal view (Fig. 12); Egypt, Libya and Tunisia (Fig. 59). ...... ............................................... B. nikolajevi Hillert, Arnone, Král & Massa sp. nov. 10 (1) Scutellar shield densely punctate. 11 (12) France (including Corse), Portugal and Spain (see distribution map on Fig. 59 for details). ......................................................................... B. gallicus (Mulsant, 1842) 12 (11) Spain, Gibraltar (see distribution map on Fig. 59, for details). .................................. ............................................................................ B. howdeni Hillert & Král sp. nov. Annotated list of the Western Palaearctic Bolbelasmus species with new records Bolbelasmus bocchus (Erichson, 1841) Bolboceras Bocchus Erichson, 1841: 170 (original description). = Bolboceras fi ssicornis Mulsant, 1843: 280 (original description). Synonymized by MARSEUL (1857: 83). Type localities. Bolboceras Bocchus: ‘Algier [Bona = Annaba]’; B. fi ssicornis: ‘Algérie’. Distribution. Algeria, Morocco (present paper). Bolbelasmus brancoi Hillert & Král sp. nov. Type locality. Spain (Andalusia), Los Jesos, N slope of Sierra de Alhamilla, 37°05′N 02°17′W, 500 m a.s.l. Distribution. Spain. Acta Entomologica Musei Nationalis Pragae, 56(1), 2016 245 Bolbelasmus gallicus (Mulsant, 1842) Bolboceras Gallicus Mulsant, 1842: 350 (original description). = Bolboceras Gallicus Var. Conjunctus Mulsant, 1842: 351 (original description). Synonymized by BOUCOMONT (1912: 17). = Bolboceras Gallicus Var. Provincialis Mulsant, 1842: 351 (original description). Synonymized by BOUCOMONT (1912: 17). Type localities. Bolboceras gallicus: France, ‘exclusivement méridionale; Draguignan, Saint-Tropez, environs de Marseille’; B. conjunctus: ‘France méridionale’; B. provincialis: ‘France méridionale’. Distribution. France (including Corsica – fi rst records), Portugal, Spain (present paper). Bolbelasmus howdeni Hillert & Král sp. nov. Type locality. Spain, Andalusia, Castillo de Castillar, Algeciras env., 10 km N of San Roque, 36°12′N 05°23′W, 78 m a.s.l. Distribution. Spain, Gibraltar (present paper). Bolbelasmus keithi Miessen & Trichas, 2011 Bolbelasmus keithi Miessen & Trichas, 2011: 183 (original description), fi gs on pp. 187–188 [not numbered]. Type locality. Greece, ‘env. Ag. Paraskevi (church), Agia Marina, Kasos island, (Lat: 35°24′10.71″N, Long: 26°54′27.59″E)’. Type material examined (2 specimens). PARATYPES: GREECE: CRETE: 1  1 , ‘ELAFONISI (Life), 30/12/9614/3/97, Leg. Trichas 321 [white printed label], Elafonisi (Life), = Elafonisos, S.-O. Kriti, 30.XII.→ 14.III.1997, leg.: A. Trichas 321, collection: G. Miessen det. 2011, PARATYPUS [printed, red label]’ (GMCL). Additional material examined (9 specimens). GREECE: CRETE: Plakias, 1.iv.1988, 1 , Winkelmann-Köck lgt. (LHCB), 1  (OHCB); Kastellakia [= Kastellos], 35°22′N, 24°29′E, 4.iv.1978, at light, 1 , Malicky lgt. (DKCP); Mirabella, 1  Maltzan [?lgt.] (OHCB). RHODES: Falirakion, Ladiko, 22.iv.–5.v.1990, 2  1 , L. Egger lgt. (ABCB) and (MECI); Villanova, 7.ii.1934, 1  (MSNM); Marizza, 13.i.1934, 1  (MSAP). Remarks. After the description of B. keithi from Crete and Kasos Islands by MIESSEN & TRICHAS (2011), the record of B. unicornis from Rhodes by SCHATZMAYR (1936) and MIKŠIĆ (1959) needed to be confi rmed. Actually, following MIESSEN & TRICHAS (2011), we identifi ed the Rhodes specimens as B. keithi, thus defi nitively excluding the presence of B. unicornis in the Mediterranean islands (see also ARNONE & MASSA 2010, MIESSEN 2011, MIESSEN & TRICHAS 2011). Therefore the above mentioned specimens constitute the fi rst record of B. keithi from the island of Rhodes. Distribution. Greece: Crete, Kasos and Rhodes (MIESSEN & TRICHAS 2011). Bolbelasmus makrisi Miessen, 2011 Bolbelasmus makrisi Miessen, 2011: 111 (original description), fi gs on pp. 116, 118–119 [not numbered]. Type locality. ‘Kourion, Lemesos, alt.: 50 m, Cyprus’. Type material examined (2 specimens). PARATYPES: CYPRUS: 1  1 , ‘Rizoelia nat. for. Park, Larnaka – CYPRUS, leg.: E. Kakouris, collection: G. Miessen, Bolbelasmus makrisi nov. sp., G. Miessen det. 2011, PARATYPUS [printed, red label]’ (GMCL). Additional material examined (64 specimens). CYPRUS: PAPHOS: Akamas NP, 5 km N of Lara, 13.–20.iii.2015, 1  1 , O. Hillert lgt. (OHCB); Akamas NP, Lara, 13.–20.iii.2015, 1 , O. Hillert lgt. (OHCB); Miliou, N of HILLERT et al.: Bolbelasmus of the Western and Southern Mediterranean (Geotrupidae) 246 Paphos, 13.–20.iii.2015, 1 , O. Hillert lgt. (OHCB); Miliou, 34°56′22.23″N, 32°27′35.90″E, vivant dans piscine [alive in pool], 7.vi.2014, 1 , G. Miessen lgt. (OHCB); Miliou, 34°56′21.66″N 32°27′35.59″E, attire par lampe UV [attracted by UV light], 7.vi.2014, 1 , G. Miessen lgt. (OHCB); 5 km NE of Pegeia, Paphos env., 13.–20.iii.2015, 16  15 , O. Hillert lgt. (OHCB), 1  1  (ERCS), 1  1  (HKCS), 2  1  (JSCP), 2  2  (DKCP), 4  2  (OBCB), 2  2  (GMCL), 3  3  (BMCP). Distribution. Cyprus (MIESSEN 2011). Bolbelasmus nikolajevi Hillert, Arnone, Král & Massa sp. nov. Type locality. NW Tunisia, Sidi Bouzid gov., 7.5 km SWW of Hajeb el Ayoun, by road, 35°22′N, 09°28′E, 340 m a.s.l. Distribution. Egypt, Libya and Tunisia (present paper). Bolbelasmus nireus (Reitter, 1895) Bolboceras nireus Reitter, 1895: 81 (original description). Type locality. Turkey, ‘Akbes’ [= Maydān Akbis]. Material examined (6 specimens). TURKEY: ADANA PROV.: Adana, 1 , 1  (JMCP); İçel, Çamliyayla, Sebil, 24.–26.v.1995, 1 , J. Mička lgt. (JMCP). ANTALYA PROV.: Korkuteli, 11.vi.1991, 1 , B. Němec lgt. (BNCP); Side, 27.iv.1973, 1 , K. Warncke lgt. (OHCB). IRAN: FARS PROV.: pass 140 km NE Šīraãz, 20.–21.iv.2002, 1 , S. Kadlec lgt. (NMPC). Distribution. Iraq, Iran, Syria, Turkey (KRIKKEN 1977, KEITH 2005, KRÁL et al. 2006, ARNONE & MASSA 2010, MIESSEN 2011). Bolbelasmus unicornis (Schrank von Paula, 1789) Scarabaeus unicornis Schrank von Paula, 1789: 61 (original description). = Scarabaeus aeneas Panzer, 1793: pl. 5 (original illustration). = Scarabaeus quadridens Panzer, 1795: 2 (original description), fi gs 1–4. Type localities. Scarabaeus aeneas: Germany, ‘Bavariae’; S. quadridens: ‘Austriae, Bavariae’; S. unicornis: Austria, ‘Aus der Sammlung des Hrn. Rathes Schiffermüller’. Material examined (85 specimens). AUSTRIA: Austria, 2  1  (MSNG); Oesterreich, 1  (OHCB); Österr.[eich], 1  (ZFMK). WIEN: Umg. Wien, 1 , Ad. Hoffmann [lgt.] (ZFMK). BOSNIA & HERZEGOVINA: Herzegovina, 1  (ZFMK). CZECH REPUBLIC: MORAVIA: Čejč, ‘Manson’s steppe’, 16.vi.1995, 1 , Vít Kubáň lgt. (VKCB); 2 km SE of Čejč, ‘Manson steppe’, 29.vi.1999, 1 , Vít Kubáň lgt. (VKCB); Prossnitz [= Prostějov], 1 , K. Kyselý lgt. (NHMB). HUNGARY: Hungaria, 1  2  (NMPC). PEST: Budapest Hármashatar-hegy, autos hálózás 31.v.2004, 1 , O. Merkl lgt. (HNHM); Budapest, Martonhegy 17.iii.1949, 1  (HNHM); Isaszeg, 19.i.2008, 1  (HNHM); Kesztölc, 47°43′N, 18°47′E, 17.x.2014, 1 , Tomáš Vendl lgt. (TVCP); Pest, 1  (NMEC); Csévharaszt 14-16. viii.2001, 1 , Szél lgt. (HNHM); Blatorbagy, 27.vi.99, 1 , G. András lgt. (HNHM); Budakeszi Hosszú-dűlő, 200 m, Cynodonto-Festucetum, on Glomus macrocarpum, 8.vi.1991, 1 , L. Ádám lgt. (HNHM). SOMOGY: Kaposvár 8.vii.1931, 1 , M. Nattán lgt. (HNHM); Kaposvár, 1 , M. Nattán lgt. (HNHM). TOLNA: Bátaapáti Nagy-Mórágyi- -völgy Ezüsthársas-tolgyes, 1  (HNHM). GERMANY: BAYERN: Bavaria, 1 , ex. coll. Veselý (NMPC). ITALY: PIEDMONT: Piemont, 1  (NMPC); Piedmont, 1 , Baudi [lgt.] (MZUF); Piedmont, 1 , L. Carrara [lgt.] (MZUF); Piedmont, 4  2  (MSNG). LOMBARDY: Milan 1  1  (MSNG). TUSCANY: Toscana, 1  (OHCB). ROMANIA: TULCEA: Babadag, 11.vii.1985, 1 , Jaromír Hanuš lgt. (VKCB). SĂLAJ: Z. [Zalău], erdōd [= forest], 3.viii.1973, 1  (OHCB). SERBIA: Serbia, 1 , coll. Matcha (NMPC). VOJVODINA: Ruma, 1  (MSNG); ŠUMADIJA: Đerdap, 5 km WSW of Tekija, 27.–28.v.2014, 2  1  (DKCP). SLOVAKIA: BANSKÁ BYSTRICA REG.: Banská Bystrica, 18.v.1979, 1  (VKCC); Kiarov, vi.1936, 1 , Dr. R. Schwarz lgt. (NMPC). KOŠICE REG.: Hrhov, E of Okrúhle hill, 4.vii.1988, 2 , David Král lgt. (DKCP); Silická planina [plateau], Rakaťa env., 5.vii.1988, 1 , David Král lgt. (DKCP); NITRA REG.: Kamenica nad Hronom, Čierna hora [hill], 11.viii.2011, 1 , D. Juřena lgt. (BMCP). PREŠOV Acta Entomologica Musei Nationalis Pragae, 56(1), 2016 247 REG.: Snina, vii.1965, 1 , M. Sláma lgt. (DKCP); TRENČÍN REG.: Brunovce, 1  (NMEC); Trenčín, Zlatovce, 27.vi.1935, 1  (NMPC); Trenčín, v.[19]31, 1 , Dr. A. Richter (NMPC); Trenčín, Zlatovce, vi.[19]26, 1 , Čepelák lgt. (NMPC); Trenčín, 2  3 , Čepelák lgt. (DKCP), 9  8  (NMPC), 2  1  (OHCB); Trenčín, 1  1  (NMPC); Trenčín, 1 , V. Steidl lgt. (NMPC); Trenčín, 1  1 , Dr. A. Richter lgt. (NMPC); Trenčín. Inovec, 1 , Čepelák lgt. (NMPC); Trenczsen, 1  1  (OHCB). TRNAVA REG.: Hlohovec, 1 , Varkonyi [lgt.] (DKCP); Pustá Ves, 22.vii.1984, 1 , Máslo lgt. (NMPC). TURKEY (European): EDIRNE PROV.: ca. 15 km east of Edirne, 27.iii.[19]88, 1 , Heinz lgt. (DKCP). (Asian): DENIZLI PROV.: Anatolien, Goundely [= Denizli], v. [19]56, 1 , Kulzer lgt. (ZSMC). UKRAINE: CHERNIHIV PROV.: Bukowina, Chernowitz, 1  (NMPC). Remarks. Bolbelasmus unicornis has so far been reported from France and Italy, across southern parts of Central Europe to the Balkan Peninsula, Ukraine and southern parts of European Russia (cf. e.g. ÁDÁM 1994, 2003; ALLENSPACH 1970; BALLERIO et al. 2010; BARAUD 1977, 1992; BUNALSKI et al. 2013; ENDRŐDI 1957; HORION 1958, GUÉORGUIEV & BUNALSKI 2004; JUŘENA et al. 2008; KÖHLER & KLAUSNITZER 1998; KRÁL & MALÝ 1993; MIKŠIĆ 1970; LUMARET 1990; NÁDAI 2006; PAILL 2008; PETROVITZ 1956; SHOKHIN 2007). It has also been recorded from Crete and Rhodes, but actually these islands are inhabited by B. makrisi (see above) (SCHATZAMAYR 1936, PETROVITZ 1959, MIESSEN 2011, MIESSEN & TRICHAS 2011, present paper). Italian records of B. unicornis cover Piedmont, Lombardy, Trentino, Tuscany, Veneto and Friuli (BALLERIO 2008, BENASSO 1971, ZANDIGIACOMO 2006, present data). The presence in Sicily, fi rst reported by BARAUD (1977), has to be referred to B. vaulogeri (present study). First record from the Tuscany province of Italy. Distribution. Austria, Bosnia & Herzegovina, Bulgaria, Croatia, Czech Republic, France (Alsace), Germany, Greece, Hungary, Italy, Moldavia, Poland, Romania, Russia, Serbia, Slovakia, Slovenia, Switzerland, Turkey (European and Asian) and Ukraine (KRÁL et al. 2006, present data). Bolbelasmus tauricus Petrovitz, 1973 Bolbelasmus tauricus Petrovitz, 1973: 338 (original description). Type locality. Turkey, ‘Cilicischer Taurus, Südhang, Namrun’. Distribution. Turkey (Asian) (KRÁL et al. 2006, ARNONE & MASSA 2010, MIESSEN 2011). Bolbelasmus vaulogeri (Abeille de Perrin, 1898) stat. restit. Bolboceras Vaulogeri Abeille de Perrin, 1898: 254 (original description). = Bolbelasmus romanorum Arnone & Massa, 2010: 403, fi gs 1–5, 8 (original description), syn. nov. Type localities. Bolboceras Vaulogeri: Tunisia, ‘Bizerte’; Bolbelasmus romanorum: Italy, ‘Sicily, Balestrate, foce Calatubo (Palermo)’. Distribution. Algeria, Italy (Sicily), Tunisia (present paper). Acknowledgements We thank all colleagues and institutions listed in the Material and methods section for enabling us to study the material in their care. Radek Dunda (Újezd u Chcebuzi, CZ), Harald Kalz (Schlabendorf, GE), Jan Schneider (Praha, CZ), Petr Šípek and Dominik Vondráček (both Charles University in Prague, CZ) and Jan Vondráček (Bulhary, CZ) were excellent HILLERT et al.: Bolbelasmus of the Western and Southern Mediterranean (Geotrupidae) 248 companions of OH and DK during their collecting trips to Portugal, Morocco, Spain and Tunisia. Special thanks are due to Denis Keith (Chartres, France) for his help with identifi - cation of some localities, Marcello Romano for executing several photographs included in this paper, Petr Šípek for allowing us to use the photos of living specimens and Marc Miquel (Queen Mary University of London, UK) for editing our English text. David Král would like to acknowledge the institutional support from resources of the Ministry of Education, Youth and Sports of the Czech Republic. References ABEILLE DE PERRIN E. 1898: Descriptions de Coléoptères du Nord de l’Afrique. Bulletin de la Société Entomologique de France 1898: 254–258. ÁDÁM L. 1994: A check-list of the Hungarian Scarabaeoidea with the description of ten new taxa (Coleoptera). Folia Entomologica Hungarica 55: 5–17. ÁDÁM L. 2003: Faunisztikai adatok a Kárpát-medencéből (Coleoptera: Scarabaeoidea). (Faunistic records from the Carpathian Basin (Coleoptera: Scarabaeoidea)). Folia Historico Naturalia Musei Matraensis 27: 101–136 (in Hungarian, English abstract). AGOIZ-BUSTAMANTE J. L. 1998: Coleópteros interesantes atraídos por la luces de Peñalba (Huesca). Boletin de la Sociedad Entomologica Aragonesa 21: 19–20. AGOIZ-BUSTAMANTE J. L. 2003: Los Scarabaeoidea Laparosticti de Navarra (I): actualización y nuevos datos corológicos (Coleoptera). Boletin de la Sociedad Entomologica Aragonesa 32: 189–195. ALLENSPACH V. 1970: Coleoptera Scarabaeidae, Lucanidae. Insecta Helvetica. Catalogus 2. Imprimérie la Concorde, Lausanne, 186 pp. AGOGLITTA R., BARBERO E., RAGUSA E. & ZUNINO M. 2006: Catalogo sistematico e topografi co dei Geotrupidae e Scarabaeidae degradatori della Sicilia e delle isole circumsiciliane (Coleoptera: Scarabaeoidea). Boletin de la Sociedad Entomologica Aragonesa 39: 181–204. ALFIERI A. 1976: The Coleoptera of Egypt. Mémoires de la Société Entomologique d’Égypte 5: 202–219. ALIQUÒ V. 1988: Nuovi dati su Coleotteri di Sicilia (Coleoptera Scarabaeoidea et Cerambycidae). Il Naturalista Siciliano 12: 169–170. ARNONE M. & MASSA B. 2010: A new species of Bolbelasmus Boucomont, 1911 from Sicily (Italy). Il Naturalista Siciliano 34: 401–414. BAGUENA CORELLA L. 1967: Scarabaeoidea de la fauna Ibero-Balear y Pirenaica. Consejo superior de investigaciones cientifi cas Instituto Espanol de Entomologia, Madrid, 576 pp. BALLERIO A. 2008: Insetti da proteggere: la tutela entomologica in Italia. Quaderni Stazione Ecologica Civico Museo Storia Naturale Ferrara 18: 21–35. BALLERIO A., REY A., ULIANA M., RASTELLI M., RASTELLI S., ROMANO M. & COLACURCIO L. 2010: Coleotteri Scarabeoidei d’Italia. Piccole Faune. Museo Civico di Storia Naturale, Carmagnola, DVD. BARAUD J. 1977: Coléoptères Scarabaeoidea. Faune de l’Europe occidentale Belgique France Grande-Bretagne Italie Péninsule Ibérique. Nouvelle Revue d’Entomologie, Supplément 7: 1–352. BARAUD J. 1985: Coléoptères Scarabaeoidea. Faune du Nord de l’Afrique du Maroc au Sinaï. Encyclopédie Entomologique – XLVI. Editions Lechevalier, Paris, 651 pp. BARAUD J. 1992: Coléoptères Scarabaeoidea d’Europe. Faune de France, France et régions limitrophes, 78. Fédération Française des Sociétés de Sciences Naturelles & Société Linnéenne de Lyon, Paris & Lyon, 856 pp. BENASSO G. 1971: Una specie nuova per il Friuli: Bolbelasmus unicornis (Schrank) (Coleoptera Geotrupidae). Atti del Museo Civico di Storia Naturale di Trieste 27: 167–172. BOUCOMONT A. 1902: Coleoptera Lamellicornia Fam. Geotrupidae. In: WYTSMAN P. (ed.): Genera Insectorum Fascicule VII. Verteneuil & Desmet, Bruxelles, 20 pp. + 1 pl. BOUCOMONT A. 1911: Contribution à la classifi cation des Geotrypidae (Col.). Annales de la Société Entomologique de France 79: 333–350. BOUCOMONT A. 1912: Scarabaeidae: Taurocerastinae, Geotrupinae. In: SCHENKLING S. (ed.): Coleopterorum Catalogus, Pars 46. W. Junk, Berlin, 47 pp. Acta Entomologica Musei Nationalis Pragae, 56(1), 2016 249 BRANCO T. 2005: Contributions towards a catalogue of Scarabaeoidea of Portugal (Coleoptera). Elytron 19: 49–61. BUNALSKI M., SIENKIEWICZ P., KUBASIK W., KONWERSKI S. & PAŁKA K. 2013: Materialy do poznania rozmieszczenia chrząszczy Zachodniej Polski. Część 4. Bolboceratidae. (Contribution to the knowledge of beetles (Coleoptera) distribution in Western Poland. Part 4. Bolboceratidae). Wiadomości Entomologiczne 32: 259–265 (in Polish, English Summary). CARTWRIGHT O. L. 1953: Scarabaeid beetles of the genus Bradycinetulus and closely related genera in the United States. Proceedings of the United States National Museum 103: 95–120. CHIKATUNOV V. & PAVLÍČEK T. 1997: Catalogue of the beetles in Israel and adjacent areas: 1. Scarabaeoidea. Klapalekiana 33: 37–65. ENDRŐDI S. 1957: A lemezescsápú bogarak (Lamellicomia) kárpátmedencei lelőhelyadatai. [Lamellicornia (Coleoptera) ot the Carpathian basin]. Rovartani Közlemenyek (Series Nova) 10: 145–226 (in Hungarian). ERICHSON W. F. 1841: Ueber die Insecten von Algier mit besonderer Berücksichtigung ihrer Verbreitung. Pp. 140–194. In: WAGNER M. F.: Reisen in der Regentschaft Algier in den Jahren 1836, 1837 und 1838 nebst einem naturhistorischen Anhang und einem Kupferatlas. Dritter Band. L. Voss, Leipzig, xviii + 297 pp.+ 17 pls. FABRA P. R. 2004: Biologie de Bolbelasmus gallicus (Mulsant, 1842) dans le departement du Gard (Languedoc, sud de la France). Elytron 17: 37–46. FUENTE Y MORALES J. M., DE LA 1926: Catálogo sistemático-geográfi co de los Coleópteros observados en la península ibérica, Pirineos propiamente dichos y Baleares. Boletin de la Sociedad Entomologica de España 8: 145–160. GONZÁLEZ PEÑA C. 1979: Contribucion al conocimiento del género Bolbelasmus: su habitat, sus costimbres y su distribucion en Aragon. Boletin de la Asociacion Española de Entomologia 3: 23–27. GORY H. L. 1841: Description de cinq espéces de Coléoptères de Constantine et d’Oran, nouvelles ou peu connues. Magasin de Zoologie d’Anatomie Comparée et de Palaeontologie 11: 1–4 + pls 71–74. GUÉORGUIEV B. & BUNALSKI M. 2004: Critical review of the families Glaresidae, Lucanidae, Trogidae, Bolboceratidae, Geotrupidae, Hybosoridae and Ochodaeidae in Bulgaria. Acta Zoologica Bulgarica 56: 253–276. HEYDEN L., REITTER E. & WEISE J. 1906: Catalogus Coleopterorum Europae, Caucasi et Armeniae rossicae, editio secunda. E. Reitter ed., Paskau, 774 pp. HORION A. 1958: Faunistik der Mitteleuropäischen Käfer. Band VI: Lamellicornia (Scarabaeidae-Lucanidae). A Feyel, Überlingen – Bodensee, xxiv + 343 pp. JACQUELIN DU VAL P. N. C. 1863: Manuel entomologique. Genera des Coléoptères d’Europe comprenant leur classifi cation en familles naturelles, la description de tous les genres, des Tableaux synoptiques destinés à faciliter l’étude, le Catalogue de toutes les espèces, de nombreux dessins au trait de caractères. Tome troisiéme (1859–1863). A. Deyrolle, Paris, 463 pp. JUŘENA D., TÝR V. & BEZDĚK A. 2008: Příspěvek k faunistickému výzkumu listorohých brouků (Coleoptera: Scarabaeoidea) na území České republiky a Slovenska. (Contribution to the faunistic research on Scarabaeoidea (Coleoptera) in the Czech Republic and Slovakia). Klapalekiana 44 (Supplementum): 17–176 (in Czech, English summary). KEITH D. 2005: First record of the genus Bolbelasmus from Iran. Kogane 6: 51–52. KOCHER L. 1958: Catalogue commenté des coléoptères du Maroc. Fascicule VII: Lamellicornes. Travaux de l’Institut Scientifi que Chérifi en, Série Zoologie 16: 1–83. KÖHLER F. & KLAUSNITZER B. (eds) 1998: Verzeichnis der Käfer Deutschlands. Entomologische Nachrichten und Berichte, Beiheft 4: 1–185. KRÁL D., LÖBL I. & NIKOLAJEV G. V. 2006: Family Bolboceratidae Mulsant, 1842. Pp. 82–84. In: LÖBL I. & SMETANA A. (eds): Catalogue of Palaearctic Coleoptera. Volume 3. Scarabaeoidea – Scirtoidea – Dascilloidea – Buprestoidea – Byrrhoidea. Apollo Books, Stenstrup, 690 pp. KRÁL D. & MALÝ V. 1993: New records of Scarabaeoidea (Coleoptera) from Bulgaria. Acta Societatis Zoologicae Bohemicae 57: 17–29. KRIKKEN J. 1977: The genus Bolbelasmus Boucomont in Asia with notes on species occurring in other regions. Zoologische Mededelingen 51: 277–292. LI C. L., WANG C. C., MASUMOTO K., OCHI T. & YANG P. S. 2008: Review of the tribe Bolboceratini s. l. (Coleoptera: Scarabaeoidea: Geotrupidae) from Taiwan with a key to the Eurasian genera. Annals of the Entomological Society of America 101: 474–490. HILLERT et al.: Bolbelasmus of the Western and Southern Mediterranean (Geotrupidae) 250 LÓPEZ-COLÓN J. I., GONZÁLEZ PEÑA C. & BELTRÁN VALEN J. R. 1996: Familias: Geotrupidae, Ochodaeidae, Hybosoridae, Glaresidae y Trogidae (Insecta: Coleoptera, 9 (Familias 27−31): Scarabaeoidea). Catalogus de la Entomofauna Aragonesa 12: 3–14. LUCAS R. 1920: Catalogus alphabeticus generum et subgenerum Coleopterorum orbis terrarum totius (fam., trib., subtr., sect. incl.), Pars 1. Nicolaische Verlagsbuchhandlung, R. Stricker, Berlin, 696 pp. LUMARET J.-P. 1990: Atlas des Coléoptères Scarabèides Laparosticti de France. Série des Inventaires de Faune et de Flore 1: 1–419. MARSEUL S. A. 1857: Catalogue des coléoptères d’Europe. A. Duménil, Paris, 200 pp. MARTÍN-PIERA F. & LÓPEZ-COLÓN J. I. 2000: Coleoptera, Scarabaeoidea I. In: RÁMOS M. A. et al. (eds): Fauna Ibérica Vol 14. Museo Nacional de Ciencias Naturales. Consejo Superior de Investigaciones Cientifi cas, Madrid, 526 pp. MIESSEN G. 2011: Quelques commentaires sur le genre Bolbelasmus Boucomont, 1911 et description d’une nouvelle espèce de Chypre. Lambillionea 111: 109–119. MIESSEN G. & TRICHAS A. 2011: Description d’un nouveau Bolbelasmus Boucomont, 1911 du sud de l’Archipel Egéen (Coleoptera, Scarabaeoidea, Bolboceratidae). Lambillionea 111: 182–188. MIKŠIĆ R. 1959: Dritter Nachtrag zur „Fauna Insectorum Balcanica-Scarabaeidae“. Godišnjak Biološkog Instituta Univerziteta u Sarajevu 12: 47–136. MULSANT E. 1842: Histoire naturelle des Coléoptères de France. Lamellicornes. Maison Libraire, Paris & Imprimerie de Dumoulin, Ronet et Sibuet, Lyon, viii + 626 pp. + 3 pls. MULSANT E. 1843: Description de quelques Coléoptères inedits. Annales des Sciences Physiques et Naturelle d’Agriculture et d’Industrie or Annales de la Société Linnéenne de Lyon 6: 276–286. NÁDAI L. 2006: A Bolboceratinae alcsalád magyarországi fajainak lolőhelyadatai (Coleoptera, Scarabaeoidea: Geotrupidae). (Hungarian localities of the species of Bolboceratinae (Coleoptera, Scarabaeoidea: Geotrupidae)). Folia Historico Naturalia Musei Matraensis 30: 205–210 (in Hungarian, English abstract). NIKOLAJEV G. V. 1996: Forms of Lamellicorn beetles with a large quantity of grooves on the elytrons from the Lower Cretaceous location of Baisa in Transbaikal. Paleontologicheskiy Zhurnal 2: 91–99. OLSOUFIEFF G. 1907: Sur une nouveau genre de Geotrypini. Russkoe Entomologicheskoye Obozrenie 7: 21–22. PAILL W. 2008: Wiederfund von Bolbelasmus unicornis (Schrank, 1789) in der Wiener Donauauen (Coleoptera: Scarabaeoidea: Geotrupidae). Beiträge zur Entomofaunistik 8: 165–171. PAULIAN R. & BARAUD J. 1982: Faune des Coléoptères de France. II, Lucanoidea et Scarabaeoidea, Vol. 43. Encyclopédie Entomologique, Editions Lechevalier, Paris, 477 pp. PANZER G. W. F. 1793: Beyträge zur Geschichte der Insekten [= Symbolae Entomologicae]. I. Theil. Johan Jakob Palm, Erlangen, 24 pp. + 6 pls. PANZER G. W. F. 1795: Entomologia germanica exhibens insecta per germaniam indigena secundum classes, ordines, genera, species adjectis synonymis, locis, observationibus. I. Eleutherata. Felssecker, Norimbergae, [8] + 12 + 368 + 2 pp. PESLIER S. 2004: Clé de détermination illustrée des Geotrupidae de France (Coleoptera, Scarabaeoidea). Revue de l’Association Roussillonnaise d’Entomologie 8(1): 1–9. PETROVITZ R. 1956: Die koprophagen Scarabaeiden des nördlichen Burgenlandes. Wissenschaftliche Arbeiten aus dem Burgenland 13: 4–24. PETROVITZ R. 1959: Beitrag zur Scarabeiden – Fauna der Insel Rhodos. Bollettino del Laboratorio di Entomologia Agraria 17: 124–130. PETROVITZ R. 1973: Neue Arten aus der Unterfamilie Geotrupinae. Folia Entomologica Hungarica 26: 335–342. PIC M. 1924: Habitats et notes concernant divers Coléoptères intéressants d’Égypte. Bulletin de la Société Royale Entomologique d’Égypte 1923: 125–129. RAGUSA E. 1893: Catalogo ragionato dei coleotteri di Sicilia. Scarabaeidae. Il Naturalista Siciliano 12: 201–205. REITTER E. 1895: Beschreibung neuer oder wenig gekannter Coleoptera aus der Umgebung von Akbes in Syrien. Wiener Entomologische Zeitung 14: 79–88. RUANO MARCO L., MARTÍN PIERA F. & ANDÚJAR A. 1988: Los Scarabaeoidea de la provincia de Albacete (Coleoptera). Instituto de Estudios Albacetenses de la Excma. Disputacion de Albacete y CSIC. Confederation Española de Estudios Locales, Serie 1. Ensayos Historicos y Cientifi cos 32: 1–201. Acta Entomologica Musei Nationalis Pragae, 56(1), 2016 251 Locality Aprox. coordinates Altit. Arrayanes, see Carretera Arrayanes Ayora 39°04′N 01°03′W 640 Aznalcázar 37°18′N 06°14′W33 Azrou 33°26′N 05°13′W 1260 Babadag 44°54′N 28°44′E36 Bagneres 43°04′N 00°09′E 575 Balestrate 38°03′N 13°00′E44 Bananco del Espartal Banská Bystrica 48°44′N 19°08′E 376 Barbentane 43°54′N 04°44′E90 Barcelona 41°23′N 02°10′E20 Baza 37°29′N 02°46′W 870 Beausset; see Le Beausset Ben Yacoub 34°28′N 02°46′E45 Benghazi 32°06′N 20°03′E2 Beni Sicar 35°23′N 02°59′W110 Beni Snassen 34°48′N 02°23′W 730 Beziers 43°20′N 03°13′E37 Bilad Gamouda 34°59′N 09°19′E 383 Bir Ketana 34°47′N 10°50′E1 Biserta, see Bizerte RUIZ J. L. 2000: Bolbelasmus bocchus (Erichson, 1841) y Allotrypes mandibularis (Reitter, 1896): nuevos registros marroquíes (Coleoptera: Geotrupidae). Boletin de la Sociedad Entomologica Aragonesa 27: 74–75. SEABRA A. F. 1943: Contribuicoes para o inventario da fauna lusitanica. Insecta, Coleoptera. Memorias e Estudos do Museo Zoologico da Universidade de Coimbra 142: 1–152. SCHATZMAYR A. 1936: Risultati scientifi ci delle cacce entomologiche di S.A.S. il principe Alessandro Della Torre e Tasso nelle isole dell’Egeo. VI. Scarabaeidae. Bollettino Laboratorio Zoologia Generale Agraria Reale Istituto Superiore Agrario Portici 30: 9–23. SCHATZMAYR A. 1946: Gli Scarabeidi coprofagi della Libia e dell’Egitto. Atti Società Italiana Scienze Naturali Milano 85: 40–84. SCHOLTZ C. H. & BROWNE D. J. 1996: Polyphyly in the Geotrupidae (Scarabaeoidea: Coleoptera): a case for a new family. Journal of Natural History 30: 597–614. SCHRANK VON PAULA F. 1789: Entomologische Beobachtungen. Der Naturforscher 24: 60–90. SHOKHIN I. V. 2007: Contribution to the fauna of lamellicorn beetles of southern Russia, with some nomenclatural changes in the family Scarabaeidae. Caucasian Entomological Bulletin 3: 105–185. VERDÚ J. R., GALANTE E. & LUMARET J. P. 1998: Description de la larve de Bolbelasmus bocchus (Erichson) et position systematique du genre. Annales de la Société Entomologique de France 34: 245–251. WINKLER A. 1929: Lamellicornia. Pp. 1025–1134. In: Catalogus Coleopterorum regionis palaearcticae, Pars 9. A. Winkler, Wien [1924–1929], pp. 1009–1136. ZANDIGIACOMO P. 2006: Nuovo rinvenimento di Bolbelasmus unicornis (Schrank) (Coleoptera, Geotrupidae) in Friuli Venezia Giulia. Bollettino della Società Naturalistica ‘Silvia Zenari’ 29: 75–79. Appendix: Gazetteer Locality Aprox. coordinates Altit. Afl ou 34°07′N 02°05′E 1430 Ain-el Kerma 34°0′N 06°39′W 132 Ain-Ograb 34°59′N 04°06′E 1050 Aix 43°32′N 05°27′E 330 Albarracin 40°24′N 01°27′W 1114 Alcalá de Henares 40°28′N 03°21′W 600 Alcala de los Gazules 36°27′N 05°43′W96 Alcala, see Alcala de los Gazules Aldehuela de la Boveda 40°51′N 06°02′W 800 Alexandria 31°12′N 29°55′E16 Algezares 38°08′N 01°44′W 430 Algier 36°45′N 03°02′E80 Alhama de Murcia 37°50′N 01°25′W 200 Alicante 38°20′N 00°29′W20 Aljaraque 37°16′N 07°01′W20 Almeria 36°51′N 02°20′W0 Almoraima 36°16′N 05°25′W28 Annaba 36°52′N 07°44′E10 Aranjuez 40°01′N 03°36′W 510 Arcos de la Fontera 36°45′N 05°48′W 140 Armacao de Pera 37°06′N 08°21′W0