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

Author: Hillert, Oliver; Arnone, Marcello; Král, David; Massa, Bruno
Publisher: Zenodo
DOI: 10.5281/zenodo.5306071
Source: https://zenodo.org/records/5306071/files/source.pdf
ACTA ENTOMOLOGICA MUSEI NATIONALIS PRAGAE
Published 15. ii.2016 Volume 56(1), pp. 211–254 ISSN 0374-1036
h p://zoobank.o g/u n:lsid:zoobank.o g:pub:7EFF344A-B2FF-4A14-80AC-3B047298201D
The genus Bolbelasmus in he wes e n
and sou he n egions o he Medi e anean Basin
(Coleop e a: Geo upidae: Bolboce a inae)
Oli e HILLERT1), Ma cello ARNONE2), Da id KRÁL3) & B uno MASSA4)
1) Kie e ndamm 10, D-15566 Schöneiche bei Be lin, Ge many; e-mail: [email p o ec ed]
2) Via San Michele 8, I-90010 Al a illa Milicia (Pale mo), I aly; e-mail: [email p o ec ed]
3) Cha les Uni e si y in P ague, Facul y o Science, Depa men o Zoology, Viničná 7, CZ-128 43 P aha 2,
Czech Republic; e-mail: k al@na u .cuni.cz
4) Uni e si y o Pale mo, Depa men o Ag icul u al and Fo es Sciences, Viale Scienze 13, I-90128 Pale mo,
I aly; e-mail: [email p o ec ed]
Abs ac . The Bolbelasmus Boucomon , 1911 species o he wes e n and sou h-
e n egions o he Medi e anean Basin (No he n A ica, Ibe ian Peninsula and
F ance) a e e ised. The ollowing h ee new species a e desc ibed: Bolbelasmus
b ancoi Hille & K ál sp. no . and Bolbelasmus howdeni Hille & K ál sp. no .,
bo h om Spain and Gib al a , and Bolbelasmus nikolaje i Hille , A none, K ál &
Massa sp. no . om Egyp , Libya and Tunisia. Bolbelasmus auloge i (Abeille de
Pe in, 1898) s a . es i . is emo ed om synonymy wi h B. bocchus (E ichson,
1841) and eins a ed as a sepa a e species. Bolbelasmus omano um A none &
Massa, 2010 is conside ed a junio subjec i e synonym o B. auloge i. Lec o-
ypes o Bolboce as bocchus E ichson, 1841 and Bolboce as auloge i Abeille
de Pe in, 1898 a e designa ed. Rele an diagnos ic cha ac e s (head, p ono um,
ely on, ex e nal male geni alia) a e illus a ed. Iden i i ca ion keys o bo h males
and emales, and an anno a ed lis o he Wes e n Palaea c ic ep esen a i es o he
genus Bolbelasmus a e p esen ed. Finally, i s eco ds a e gi en o B. gallicus
(Mulsan , 1842) om Co sica and he Midi-Py énées egion o F ance, B. kei hi
Miessen & T ichas, 2011 om he G eek island o Rhodes, and B. unico nis
(Sch ank on Paula, 1789) om he Tuscany p o ince o I aly.
Key wo ds. Coleop e a, Sca abaeoidea, Geo upidae, Bolboce a inae, Bolbelas-
mini, Bolbelasmus, axonomy, e ision, new species, synonymy, species es i u ion,
lec o ype designa ion, key, dis ibu ion, anno a ed lis , Medi e anean Region,
Palaea c ic Region
HILLERT e al.: Bolbelasmus o he Wes e n and Sou he n Medi e anean (Geo upidae)
212
In oduc ion
The bolboce a ine genus Bolbelasmus was e ec ed by BOUCOMONT (1911) in his s udy
‘Con ibu ion à la classi i ca ion des Geo ypidae’, whe e he designa ed he ollowing h ee
species: Bolboce as bocchus E ichson, 1841, Bolboce as gallicus Mulsan , 1842 and Bolboce-
as unico nis (Sch ank on Paula, 1789) as ‘species ypicae’. In he same s udy, BOUCOMONT
(1911) e ec ed he genus Kolbeus wi h ‘species ypicae’Bolboce as a cua um Ba es, 1887
and Bolboce as co eanum Kolbe, 1886. Type species we e designa ed by LUCAS (1920) o
Kolbeus, namely Bolboce as co eanus, while 30 yea s la e , CARTWRIGHT (1953) designa ed
Bolboce as gallicus o Bolbelasmus. In addi ion, CARTWRIGHT (1953) synonymized Kolbeus
unde Bolbelasmus in he same a icle. In a ecen axonomic s udy, NIKOLAJEV (1996) desc ibed
he ibe Bolbelasmini wi h h ee gene a: Bolbelasmus Boucomon , 1911, Bolbo ypes Olsou-
i e , 1907 and Bolbogonium Boucomon , 1911. La e on NIKOLAJEV (2003) placed Kolbeus a
he ank o subgenus o Bolbelasmus. SCHOLTZ & BROWN (1996) ha e p oposed ha he g oup
o gene a ela ed o Bolboce as Ki by, 1819 belongs o he ank o amily (Bolboce a idae).
Howe e , in acco dance wi h ecen s udies based on molecula da a (e.g. SMITH e al. 2006)
and wi h phylogeny o he Geo upidae based on la al mo phology (VERDÚ el al. 2004), he
g oup is ea ed as a sub amily o Geo upidae in his s udy (see also BOUCHARD e al. 2011).
Cu en ly, we ecognize 27 species o he genus Bolbelasmus (including subgenus Kolbeus)
om he Hola c ic Region, including h ee new species desc ibed in his s udy. Eigh axa a e
dis ibu ed h oughou he New Wo ld, ele en a e known om Eu ope and he Medi e anean
Region and eigh a e na i e o he Himalaya, Fa Eas , China and sou h-eas e n Asia (e.g.
REITTER 1895, HOWDEN 1964, PETROVITZ 1968, KRIKKEN 1977, MASUMOTO 1984, HOWDEN &
SOLIS 1995, NIKOLAJEV 2005, KRÁL e al. 2006, LI e al. 2008). Acco ding o KRIKKEN (1977),
ARNONE & MASSA (2010), MIESSEN (2011) and MIESSEN & TRICHAS (2011) and o he esul s o
he p esen s udy, ele en species o he genus Bolbelasmus a e dis ibu ed in Eu ope, Middle
Eas and No h A ica.
The new species in his genus we e desc ibed hanks o he s udy o a la ge numbe o
specimens om almos 70 ins u ional and p i a e collec ions and se ies o specimens ecen ly
collec ed by he au ho s.
Ma e ial and me hods
The habi us pho og aphs we e ei he aken by D. K ál using a Canon MP-E 65mm/2.8
Mac o lens wi h 5:1 op ical magni i ca ion on bellows a ached o a Canon EOS 550D came a
o by M. Romano using a Canon EOS 450D digi al came a moun ing a Sigma 70 mm 1:2.8
DG mac o. Pa ially ocused images o he specimen we e s acked using Helicon Focus
3.20.2 P o so wa e.
The ollowing codes iden i y he collec ions housing he ma e ial examined (cu a o s in
ound b acke s):
ABCB Albe o Balle io collec ion, B escia, I aly;
AHCL An onio Hidalgo Fon i e os collec ion, Lina es, Spain;
ARCL And eas Reichenbach collec ion, Leipzig, Ge many;
Ac a En omologica Musei Na ionalis P agae, 56(1), 2016 213
BMCP B uno Massa collec ion (deposi ed in MSNG);
BMNH Na u al His o y Museum, London, Uni ed Kingdom (Maxwell V. L. Ba clay, Malcolm Ke ley);
BNCP Bohusla Němec collec ion, P aha, Czech Republic;
CMNC †Hen y F. & Anne T. Howden collec ion, En omology Di ision, Museum o Na u e Canada, O awa, Canada
(Aleš Sme ana);
CUPC Depa men o Zoology, Cha les Uni e si y, P ague, Czech Republic (Pe Šípek);
DJCP Daniel Juřena collec ion, P os ějo , Czech Republic;
DKCC Denis Kei h collec ion, Cha es, F ance;
DKCP Da id K ál collec ion (deposi ed in NMPC);
ERCS Eckeha d Rößne collec ion, Schwe in, Ge many;
GMCL Geo ey Miessen collec ion, Liège, Belgium;
GNCA Geo gy V. Nikolaje collec ion, Alma y, Kazakhs an;
GSCT Guido Saba inelli collec ion, Tunis, Tunisia;
HFCB Hans Fe y collec ion, Be lin, Ge many;
HKCS Ha ald Kal z collec ion, Schlabendo , Ge many;
HNHM Hunga ian Na u al His o y Museum, Budapes (O ó Me kl, Győző Szél);
IBCF I o Boščík collec ion, F ýdek-Mís ek – Skalice, Czech Republic;
IECA Ins i u e o En omology, Biology Cen e CAS, České Budějo ice, Czech Republic (Aleš Bezděk);
IMCO I o Ma inů collec ion, Olomouc, Czech Republic;
JBCP Jan Ba elka collec ion, P ague, Czech Republic;
JCCL Ja ie Cas illo collec ion, Lina es, Spain;
JDCC Jiří D ořák collec ion, Čelčice, Czech Republic;
JHCP Jiří Há a collec ion, Úně ice, Czech Republic;
JMCH Jan Ma ějíček collec ion, H adec K álo é, Czech Republic;
JMCP Jiří Mička collec ion, P ague, Czech Republic;
JUCS José Ma ía U bano collec ion, Se illa, Spain;
JNCS Je ónimo Na a o Ga cía collec ion, Se illa, Spain;
JSCP Jan Schneide collec ion, P ague, Czech Republic;
JSCS Joachim Schön eld collec ion, Sinzig, Ge many;
JZCJ Ja osla Žák collec ion, Jeze nice, Czech Republic;
LHCB La s Hend ich collec ion (deposi ed in ZSMC);
JLCM José Ignácio López Colón collec ion, Mad id, Spain;
LMCT Ladisla Mencl collec ion, Týnec nad Labem, Czech Republic;
LSCN Ludge Schmid collec ion, Neus ad , Ge many;
MACP Ma cello A none collec ion, Pale mo, I aly;
MECI Man ed Egge collec ion, Innsb uck, Aus ia;
MHNL Muséum d’his oi e na u elle, Lyon, F ance (Ha old Lab ique);
MLCD Mike Liebsche collec ion, D esden, Ge many;
MNHB Museum ü Na u kunde, Be lin, Ge many (Johannes F isch, Joachim Wille s);
MNHN Muséum na ional d’His oi e na u elle, Pa is, F ance (Oli ie Mon euil, An oine Man ille i);
MSNT Museo Regionale Scienze Na u ali, Tu in, I aly (Luca Picciau);
MSAP Museo Scienze Ag a ie, Po ici, Naples, I aly (An onio Ga onna);
MSNG Museo Ci ico di S o ia Na u ale ‘G. Do ia’, Genoa, I aly (Ma ia Luisa Ta ano);
MSNM Museo Ci ico di S o ia Na u ale, Milan, I aly (Fab izio Riga o);
MZUC Museo di Zoologia dell’Uni e si à o Ca ania, Ca ania, I aly (Gio gio Sabella);
MZUF Museo di Zoologia La Specola dell’Uni e si à o Flo ence, I aly (Luca Ba olozzi);
NMEC Na u kundemuseum, E u , Ge many (Ma hias Ha mann);
NHMB Na u his o isches Museum, Basel, Swi ze land († Michel B ancucci, E a Sp eche -Uebe sachs, Ma hias
Boh e );
NMPC Na ional Museum (Na u al His o y), P ague, Czech Republic (Jiří Hájek);
OBCB Oli ie Boilly collec ion, Ma cq en Ba oeul, F ance;
HILLERT e al.: Bolbelasmus o he Wes e n and Sou he n Medi e anean (Geo upidae)
214
OHCB Oli e Hille collec ion, Schöneiche bei Be lin, Ge many;
PJCL Pa el Jáchymek collec ion, Luhačo ice, Czech Republic;
PKCS Pe Kylies collec ion, Slaný, Czech Republic;
PTCP Pa el Tu ek collec ion, Lanšk oun, Czech Republic;
RCCP Radek Če enka collec ion, P ague, Czech Republic;
RMCM Radosla Muczka collec ion, Mikulo , Czech Republic;
SJCP S anley Jákl collec ion, P ague, Czech Republic;
SMTD S aa liches Museum ü Tie kunde, D esden, Ge many (Ola Jäge );
SZCM S e ano Ziani collec ion, Meldola, I aly;
TVCP Tomáš Vendl collec ion, P ague, Czech Republic;
VACP Vi o io Aliquò collec ion, Pale mo, I aly;
VKCB Ví ězsla Kubáň collec ion, B no, Czech Republic;
VKCC Václa Kři an collec ion, Chaloupky, Czech Republic;
VMCP Vladisla Malý collec ion, P ague, Czech Republic;
VRCH Vladisla Řebíček collec ion, H adiš ko, Czech Republic;
VTCZ Václa Tý collec ion, Žihle, Czech Republic;
ZFMK Zoologisches Fo schungsmuseum Alexande Koenig, Bonn, Ge many (Di k Ah ens);
ZLCK Zdeno Lucbaue collec ion, Ka e ing, Uni ed Kingdom;
ZSMC Zoologische S aa ssammlung, München, Ge many (Michael Balke, La s Hend ich).
Al oge he , 1332 specimens (see ma e ial below) we e s udied. Specimens o he newly
desc ibed species a e p o ided wi h one ed p in ed label ‘Name o axon sp. no ., HOLO-
TYPUS , ALLOTYPUS  o PARATYPUS  o  ollowed by he name o he au ho s
and he yea ’. In he case o lec o ype o pa alec o ype designa ion, each specimen bea s a
ed p in ed label: ‘Name o a axon wi h au ho and yea o o iginal desc ip ion, LECTOTY-
PUS  o PARALECTOTYPUS  o  Da id K ál & Oli e Hille des. 2014’. Rema ks
o he au ho s and commen s a e indica ed in b acke s. Exac label da a a e ci ed o he ype
ma e ial. In o ma ion in quo a ion ma ks (‘’) indica es he o iginal spelling. Coo dina es and
al i ude a e assigned o each locali y men ioned in he ex (ma e ial examined in each species)
(see Appendix 1). These da a we e used in he cons uc ion o dis ibu ion map (see Fig. 59).
Taxonomy
Bolbelasmus bocchus (E ichson, 1841)
(Figs 1, 8, 14, 21, 27, 33, 39, 45–51, 59–60)
Bolboce us Bocchus E ichson, 1841: 170 (o iginal desc ip ion). GORY (1841): able 71, i gs 1, 1A (lis ); MARSEUL
(1857): 83 (lis ); JACQUELIN DU VAL (1863): 434 (ca alogue); BOUCOMONT (1902): 4 (ca alogue); HEYDEN e al.
(1906): 725 (ca alogue); WINKLER (1927): 1035 (ca alogue).
Bolbelasmus bocchus: BOUCOMONT (1912): 17 (ca alogue); WINKLER (1927): 1035 (ca alogue); KOCHER (1958): 77
(ca alogue); RUIZ (2000): 84 (dis ibu ion).
Bolbelasmus bocchus [pa im, popula ions om Mo occo]: BARAUD (1977): 160 (monog aph); MARTÍN-PIERA &
LÓPEZ-COLÓN (2000): 179 (monog aph).
Bolbelasmus bocchus bocchus [pa im, popula ions om Mo occo]: KRIKKEN (1977): 290, i gs 16–17 ( e ision);
BARAUD (1985): 53, i g. 8 (monog aph); BARAUD (1992): 44 (monog aph); KRÁL e al. (2006): 82 (ca alogue).
Bolboce as Fissico nis Mulsan , 1843: 280 (o iginal desc ip ion). Type locali y: ‘Algé ie’. MARSEUL (1857): 83
(synonymy).
Type locali y. ‘Algie , [Bona = Annaba]’ (see Type ma e ial o de ails).
Type ma e ial examined (3 specimens). ALGERIA: LECTOTYPE (p esen designa ion):  (MNHB), ‘His . – coll.
(Coleop e a), N . 25569, Bolboce as Bocchus E . – Kl., Bona, Wagne , Zool. Mus. Be lin, Syn ypus, Bolbelasmus
Ac a En omologica Musei Na ionalis P agae, 56(1), 2016 215
bocchus E ichson, 1841, labelled by MNHUB 2013’. PARALECTOTYPES: No. 1,  (MNHB), ‘His . – coll. (Coleo-
p e a), N . 25569, Bolboce as Bocchus E . – Kl., Bona, Wagne , Zool. Mus. Be lin, Syn ypus, Bolbelasmus bocchus
E ichson, 1841, labelled by MNHUB 2013’; No. 2,  (MNHB), ‘His . – coll. (Coleop e a), N . 25569, Bolboce as
Bocchus E . – Kl., Bona, Wagne , Zool. Mus. Be lin, Syn ypus, Bolbelasmus bocchus E ichson, 184, labelled by
MNHUB 2013’; see Figs 45–51.
Addi ional ma e ial examined (183 specimens). ALGERIA: Alge ia, 2  1  (SMTD), 1  (MNHN), 2 
2  (BMNH); Alge ie, 1  2  (MHNL); Alge ien, 3  (SMTD). ALGIERS PROV.: Alge (ci y), i.1996, 1 
(DKCP); Alge , 1  (MNHN); Algie , 2  (MNHB); L. A ba, 1  1 , Rei e [lg .] (MNHN), 1  (VMCP). SIDI
BEL ABBÈS PROV.: Chanzy, [18]90, 1 , Vaulog[e ] lg . (ZSMC); Chanzy, 1890, 1 , de Vauloge lg . (MNHN).
DJELFA PROV.: Take san, i .1893, 2  3 , A Ben Yacoub [lg .] (MNHN); Take san, Djel a, 17. . 1924, 1 
(DKCC). EL BAYADH PROV.: Region de Ge y ille, i.1910, 3  8 , Ha old Powell lg . (MNHN); Le K eide ,
.[18]96, 1  1  (MNHN); El K eide , .1896, 4  2 , D . A. Chobau lg . (MNHN), 2  1  (LHCB);
El-K eide (O an), 1 , L. Bleuse [lg .] (ZFMK); K eide , 1 , Bedel [lg .] (MNHB), 1  (ZSMC), 1  2 
(MNHN); K eide , 4  4 , Bedel lg . (MNHN); K eide , 3  4  (MNHN), 1  (NMPC), 4  (ZFMK).
LAGHOUAT PROV.: A l ou, 4.i .1981, 1 , H. Pie o i [lg .] (MSNG). MASCARA PROV.: O an, Masca a, i .– .1913,
24  9 , G. Le Com e lg . (MNHN); Masca a, 1 , D . A. Cobos lg . (MNHN); Masca a, 1  (NHMB), 1 
1  (VMCP). M’SILA PROV.: Ain-Og ab, 1 , D . Ma in [lg .] (ZFMK). NAÂMA PROV.: Meche ia, 1  (MNHN).
ORAN PROV.: Magen a (p o . d’O an), 1 , E. Du oux [lg .] (ZFMK); Magen a (O an), 1  (DKCC); O an, R.
J. 3.xi.1903 1 , B. [? lg .] (MNHN); O an, 1 , coll. R. Paulian (MNHN); O an, Magen a, 1915, 1 , M. Ro-
ou lg . (MNHN); Magen a, 1  (MNHN). TLEMCEN PROV.: Lambaze, i.1885, 2 , L. Bleuse lg . (MNHN);
Sebdou, 1903, 1  (MNHN). MOROCCO: GHARB-CHRARDA-BÈNI HSSEN REG.: KÉNITRA PROV.: s.[sou h o ]
Oujda, Ain-el Ke ma, .1972, 2  1 , D. Be nh[aue ] lg . (JSCS). MEKNÈS-TAFILALET REG.: IFRANE PROV.:
Mi le e A las, I ane, 1700 m, 8.–9. i.[19]96, 1 , A. Kallies lg . (OHCB); Middle A las, Az ou, 18.–19. i.1936,
1 , K.H. Chapman & G.A. Bisse [lg .] (BMNH); Moyen-A las, nea Jbel Heb i, 12. .1980, 1 , J. Gou ès ec.
[lg .] (MHNL); Az ou, Fo e de Ced es, 10. .[19]95, 1 , Jiří Hájek lg . (JSCP), 1  (NMPC); Az ou, 10. .1995,
1 , M. Rozsí al lg . (JZCJ); I ane en ., 3. .1995, 1 , J. Bech lg . (PJCL); I ane, .[19]95, 1 , J. Holiš lg .
(JZCJ); I ane, Daye I e , 26.i .2014, 11  5 , O. Boilly lg . (OBCB); I ane, 8. i.1973, 2  (SJCP);
I ane, 1. .68, 1 , J. Galle lg . (MNHN); I ane, 11. .[19]67, 2  1 , Y. Lajonquie lg . (MNHN); NE o
I ane, Day -I ah, 17. .2003, 1 , M. Snížek lg . (VKCC); Ta o al (Oujda), 11.i .1992, 1  1 , G. Cha anon
lg . (IECA); Zad Pass, 2180 m, 24.i .2014, 1 , O. Boilly lg . (OBCB). ORIENTAL REG.: BERKANE PROV.: Ta-
o al, Beni Snassen (Pa c Na u el), iii.[19]51, 1 , An oine lg . (MNHN). JERADA PROV.: W [o ]Touissi e, Jbel
Mahso , 34°29′46″N 01°46′50″W, 1050 m, 2 , R. Rejzek lg . (JSCP). OUJIDA-ANGAD PROV.: To . Ouazane,
p .[p ope] Oujda, i.[19]35, (Lacous), 1  1  (MNHN). TAOURIRT PROV.: Gaada de Debdou pla [eau]., E o
Rchida, 33°52′N 3°10′W, ca. 1420–1550 m, 20.–21.iii.2013, 1  1 , D. K ál lg . (DKCP); Gaada de Debdou
pla [eau]., E o Rchida, 33°52′N 3°10′W, ca. 1420–1550 m, 20.–21.iii.2013, 3  1 , O. Hille lg . (OHCB).
SPAIN ( e i o y in Mo oco): MELILLA AUTONOM. COMM.: 1  1 , Melilla, Pa do Alcaide [lg .] (MSNG);
2  2 , Ros oga do, Melilla, Pa do Alcaide lg . (MNHN); 1 , same label da a (MSNT); Tau i (Beni Sica ),
Melilla Ma uecos, 1  1 , Pa do Alcaide lg . (MNHN). NOT LOCATED: Bo kine, 10. i[19]29, Le Ce , 1 
(MNHN); Daya, 1  1 , Bedel lg . (MNHN); Daja, 1  (MNHN).
Redesc ip ion. Male (lec o ype) (Figs 45–47). Rema kably con ex, hemisphe ical; do sal
su ace b ownish, ema kably shiny.
Head (Figs 45–47). Clypeus b oadly ounded, almos semici cula , an e io angles ague,
pos e io angles ele a ed. F on al ho n well de eloped, long, dis inc ly u ca e apically; i s
base ou line ellip ic in do sal aspec , pos e io edge o base si ua ed a le el o an e io pa
o eye; la e al ma gin o ho n dis inc ly, sha ply keeled om base o middle o ube cle.
Clypeal disc wi h simple, coa se punc a ion. Oblique keels abo e eyes eaching app oxi-
ma ely o hal o eyes. Punc a ion o ons and e ex app oxima ely same as on clypeus.
F on o-clypeal su u e p esen . Eyes well de eloped, dis inc ly ellip ic; genae exceeding
eyes, a cua e pos e io ly.

HILLERT e al.: Bolbelasmus o he Wes e n and Sou he n Medi e anean (Geo upidae)
216
P ono um (Figs 45–47) dis inc ly ans e se, b oades jus in basal qua e , en i ely bo -
de ed; la e al ma gin e enly ounded, ma ginal ca ina no c ena e, dis inc ly widened and
ele a ed; la e al o ea nea ly missing. P ono al punc a ion dis inc , double, consis ing o
coa se, deeply imp essed punc u es, punc u es sepa a ed by app oxima ely 2–3 imes hei
diame e and in e mixed wi h e y i ne ones. P ono al base glab ous; la e al ho nlike apophyses
well de eloped, di ec ed o wa dly in do so-la e al iew; ca i y be ween la e al and apical
apophyses ema kably deep, kidney-shaped. Apical ho nlike apophysis well de eloped, dis-
inc ly ele a ed, angled, di ec ed upwa ds, subapical p o usion si ua ed on ou e side, basally.
Scu ella shield elonga e iangula , e y i nely punc a e, shiny.
Ely a app oxima ely as long as wide, su ace shiny; hume al umbone weak; be ween
hume us and ely al su u e se en dis inc ly imp essed s iae along en i e ely al leng h, s iae
wi h coa se, egula ly spaced punc u es; in e als l a , impunc a e.
Aedeagus. Pa ame es as in Fig. 60.
Va iabili y in males. Su ace black o b ownish; p ono al ho nlike apophyses in medium
de eloped and unde de eloped (hypo helic) specimens sho , mo e o less s aigh , only
simply ounded o almos acu e apically.
Female (Pa alec o ype No. 1). Head (Fig. 48). Clypeus b oadly ounded, semici cula
wi h s aigh an e io ma gin, an e io angles e enly ounded. Pos e io angles o clypeal
ma gin e y weakly ele a ed. F on al ca ina well de eloped, dis inc ly in e up ed a middle,
bi ube cula e a in e nal in e up ion, keeled la e ally. Punc a ion o clypeus, ons, e ex
and occipu simple, dis inc ; oblique keel abo e eyes dis inc , ex ended o hal o eyes; eyes
well de eloped, dis inc ly ellip ic, genae well de eloped, egula ly a cua e.
P ono um (Fig. 48) dis inc ly ans e se, la e al ma gin weakly widened, b oades jus
a base, su ace dis inc ly and deeply punc a e, punc a ion double excep impunc a e base,
punc u es sepa a ed by dis ance equal o hei diame e and in e mixed wi h e y i ne ones.
An e io p o usion dis inc ly ans e se, sha ply semici cula , la e al apophyses p esen .
Measu emen s. To al body leng h 11.0–14.8 mm (lec o ype, : 13.5 mm; pa alec o ype
No.1, : 14.0 mm).
Di e en ial diagnosis. Re e o species key.
Collec ing ci cums ances. P e e s calca eous soils.
Dis ibu ion. Medi e anean pa s o Alge ia and no he n Mo occo (Fig. 59).
Rema ks. The ype locali y o B. bocchus is ‘Algie ’, he species has been eco ded also om
Spain (no Po ugal: BRANCO 2005), no he n Alge ia and no he n Mo occo (c . e.g. KRIKKEN
1977; BARAUD 1977, 1985, 1992; KRÁL e al. 2006; MARTÍN-PIERA & LÓPEZ-COLÓN 2000; RUIZ
2000). Rega ding he p esence in Spain, we conside he Ibe ian popula ions o belong o
ano he species, desc ibed in his pape as B. b ancoi sp. no . The popula ions dis ibu ed in
no he n Tunisia and p obably also in eas e n pa s o Alge ia we e a i s e e ed o ano he
species, Bolbelasmus auloge i, la e synonymized wi h B. bocchus by BOUCOMONT (1911)
and conside ed a subspecies o B. bocchus by KRIKKEN (1977). We conside hese popula ions
o ep esen a alid species and iden ical o popula ions o Sicily desc ibed ecen ly as B.
omano um A none & Massa, 2010 (syn. no .). We also conside he popula ions na i e o
Middle and Sou h Tunisia, Libya (T ipoli ania, Cy enaica) (SCHATZMAYR 1946) and Egyp
(Ma iou ) (PIC 1924) o be a dis inc species, B. nikolaje i sp. no ., desc ibed in his pape .
Ac a En omologica Musei Na ionalis P agae, 56(1), 2016 217
Finally, CHIKATUNOV & PAVLÍČEK (1997) epo ed B. bocchus om Is ael, bu we did no ha e
he oppo uni y o s udy any Bolbelasmus ma e ial om he Le an coun ies. ERICHSON
(1841) desc ibed Bolboce as bocchus based on a se ies o unknown numbe o specimens.
He did no es ablish a ype; hus in his s udy, we designa ed a lec o ype () (Figs 45–47,
49) and wo pa alec o ypes o i xa ion o his axon (Figs 48, 50–51). Bolboce as i ssico nis
Mulsan , 1843 was also desc ibed om ‘Algé ia’ based on he same mo phological cha ac e s
o B. bocchus (‘ ê e a mée d’une co ne non mobile, bi i de au somme [= head a med wi h
immobile ho n, u ca e apically]’) (MULSANT 1843) and la e synonymized wi h B. bocchus
by MARSEUL (1857). This impo an cha ac e o B. bocchus is shown also in GORY (1841:
able 71, i gs 1, 1A).
Bolbelasmus b ancoi Hille & K ál sp. no .
(Figs 2, 9, 15, 22, 28, 34, 40, 59, 61)
Bolboce as Bocchus [pa im, popula ions om Spain]: JACQUELIN DU VAL (1863): 131 (ca alogue).
Bolboce as Bocchus: FUENTE Y MORALES (1926): 146 (dis ibu ion).
Bolbelasmus bocchus [pa im, popula ions om Spain]: WINKLER (1927): 1035 (ca alogue); SEABRA (1943): 50
(ca alogue); BARAUD (1977): 159 (monog aph); RUANO MARCO e al. (1988): 40, i g. p. 41 (dis ibu ion); LÓPEZ-
COLÓN e al. (1996): 3, 8, 10 (dis ibu ion, key); MARTÍN-PIERA & LÓPEZ-COLÓN (2000): 179, i gs 44 A,C, 45
(monog aph); MIESSEN & TRICHAS (2011): 182 ( ema ks).
Bolbelasmus bocchus: BAGUENA CORELLA (1967): 262 (monog aph); GONZÁLEZ PEÑA (1979): 23, i gs 4–6, 8 (key,
biology); VERDÚ e al. (1998): 245, i gs 1–12 (la al desc ip ion); AGOIS-BUSTAMANTE (1998): 19 (dis ibu ion);
AGOIS-BUSTAMANTE (2003): 190 (dis ibu ion); VERDÚ e al. (2004): 514, i g. 1; 515, i g. 20; 518, i g. 42 (la al
illus a ions).
Bolbelasmus bocchus bocchus [pa im, popula ions om Spain]: KRIKKEN (1977): 290 ( e ision); BARAUD (1985): 53
(monog aph); BARAUD (1992): 44 (monog aph); BRANCO (2005): 54 ( axonomic commen s); KRÁL e al. (2006):
82 (ca alogue); ARNONE & MASSA (2010): 407 ( ema ks, dis ibu ion).
Type locali y. Spain, Andalusia, Los Jesos, N slope o Sie a de Alhamilla, 37°05′N 02°17′W, 500 m a.s.l.
Type ma e ial (143 specimens). SPAIN: ANDALUSIA AUTONOM. COMM.: ALMERIA PROV.: HOLOTYPE:  (DKCP),
‘Spain, (Andalusia), Los Jesos, N slope o Sie a de Alhamilla, 28.ii.2009, O. Hille lg .’ ALLOTYPE:  (DKCP),
same da a. PARATYPES: ALMERIA PROV.: ‘Espana, U. Alme ia, Ga ucha, Salzlachenu e , 07.i .1986’, 1  (LSCN);
‘Spain, (Andalusia), Los Jesos, N slope o Sie a de Alhamilla, 13-27.ii.2013, O. Hille lg .’, 7  5  (OHCB);
‘Spain (Andalusia), Los Jesos, N slope o Sie a de Alhamilla, 30 km NE o Alme ia, 28.ii.2009, O. Hille lg .’,
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, P o inz Alme ia, Um-
gebung Tabe nas, a.L. [a ligh ], 21.-28.i .2004, leg. S e en Schellho n’, 1  1  (OHCB). GRANADA PROV.: ‘Spain,
G anada, Andalucía, Hoya de Baza m s., 850 m asl., Salaza , 37°31.87’N 2°40.94’W, 4. .2009, lg . J. Da ebník’,
1  (JSCP), 1  (PKCS); ‘Bco. del Espa al, Baza (G anada), 30SWG2854, F.S.Piñe o leg., 3-iii-[19]90’, 1  1 
(SZCM); ‘ESPAÑA, G anada, Bco. del Espa al, Baza, 3-iii-[19]90, F.S.Piñe o’, 1  1  (SZCM); ‘Bananco del
Espa al, BAZA 8.X.[19]89, GRANADA (España)’, F. S. Piñe o leg.’, 1  (DKCP); ‘Bco. del Espa al, BAZA
8.X.[19]89, GRANADA (España), F. S. Piñe o leg.’, 1  (DKCP); ‘Espagne (G anada), Lanja on, 28.iii.[19]67,
Ba aud lg .’, 1  (MNHN); ‘E, P o . G anada, Los Olmos, 14. .2002, H. Rie z lg .’, 1  (ERCS); ‘Spain, G anada,
O gi a 12.i .1960’, 1  1  (BMCP). ARAGON AUTONOM. COMM.: SARAGOSSA PROV.: ‘España, La Playa - Sas ago
- Za agoza, 1. i.2013’, 2  (GMCL); ‘SPAIN, A agon, Za agoza, Candasnos, 13. i.2011’, 1  (IECA); ‘España,
Buja aloz, Za agoza, 18. .2013’, 2  (GMCL); ‘España 17. .2011, Buja alos, Za agoza, Leg. Ra ael Sole ’,
1  (DJCP); ‘Spain, Za agoza, Sas ago 12. .2010’, 1  (OBCB); ‘España, Buja aloz, Za agoza, .2009’, 1  1 
(GMCL); ‘España, Za agoza, Sas ago, 12. .2007’, 1  (GMCL);‘ESPAÑA, Za agoza, Buja aloz, 11.V.[19]91,
M. TOMÁS [legi ]’, 4  (SZCM); ‘Espagne – 30/V/1973, Buja aloz (Za agoza), C. Du ay ec.’, 1  (MHNL);
‘Espagne, p o . Za agoza, Buja alos en ., 18.5.2013, 1  (IBCF); ‘Ejea de los Caballe os (Za agoza), 7. .1982, J.
HILLERT e al.: Bolbelasmus o he Wes e n and Sou he n Medi e anean (Geo upidae)
218
Figs 1–4. Male habi us in do sal aspec . 1 – Bolbelasmus bocchus (E ichson, 1841) (Mo occo: Az ou, Fo e de
Ced es, body leng h: 14.0 mm, NMPC); 2 – B. b ancoi Hille & K ál sp. no . (holo ype); 3 – B. gallicus (Mulsan ,
1842) (F ance: Bouches-du-Rhône, Ba ben ane, body leng h 13.7 mm, DJCP); 4 – B. howdeni Hille & K ál sp.
no . (holo ype) .
Ac a En omologica Musei Na ionalis P agae, 56(1), 2016 219
Figs 5–7. Male habi us in do sal aspec . 5 – Bolbelasmus nikolaje i Hille , A none, K ál & Massa sp. no . (holo ype);
6 – B. auloge i (Abeille de Pe in, 1898) (Tunisia: Cap Bon peninsula, 6 km W o El Hawa iah, 11.8 mm, DKCP);
7 – B. auloge i (I aly: Sicily, Men i (Ag igen o), Cinquan a, body leng h: 11.6 mm, MAPC).
HILLERT e al.: Bolbelasmus o he Wes e n and Sou he n Medi e anean (Geo upidae)
226
Figs 21–26. P ono um o male in do sal aspec . 21 – Bolbelasmus bocchus (E ichson, 1841) (Mo occo: Az ou, Fo e
de Ced es, NMPC); 22 – B. b ancoi Hille & K ál sp. no . (holo ype); 23 – B. gallicus (Mulsan , 1842) (F ance:
Bouches-du-Rhône, Ba ben ane, DJCP); 24 – B. howdeni Hille & K ál sp. no . (holo ype); 25 – B. nikolaje i
Hille , A none, K ál & Massa sp. no . (holo ype); 26 – B. auloge i (Abeille de Pe in, 1898) (Tunisia: Cap Bon
peninsula, 6 km W o El Hawa iah, DKCP).

Ac a En omologica Musei Na ionalis P agae, 56(1), 2016 227
Figs 27–32. P ono um o male in on al aspec . 27 – Bolbelasmus bocchus (E ichson, 1841) (Mo occo: Az ou, Fo e
de Ced es, NMPC); 28 – B. b ancoi Hille & K ál sp. no . (holo ype); 29 – B. gallicus (Mulsan , 1842) (F ance:
Bouches-du-Rhône, Ba ben ane, DJCP); 30 – B. howdeni Hille & K ál sp. no . (holo ype); 31 – B. nikolaje i
Hille , A none, K ál & Massa sp. no . (holo ype); 32 – B. auloge i (Abeille de Pe in, 1898) (Tunisia: Cap Bon
peninsula, 6 km W o El Hawa iah, DKCP).
HILLERT e al.: Bolbelasmus o he Wes e n and Sou he n Medi e anean (Geo upidae)
228
Figs 33–44. Head in do sal aspec , 33–38 – ; 39–44 – . 33 – Bolbelasmus bocchus (E ichson, 1841) (Mo oc-
co: Az ou, Fo e de Ced es, NMPC); 34 – B. b ancoi Hille & K ál sp. no . (holo ype); 35 – B. gallicus (Mulsan ,
1842) (F ance: Bouches-du-Rhône, Ba ben ane, DJCP); 36 – B. howdeni Hille & K ál sp. no . (holo ype); 37 – B.
nikolaje i Hille , A none, K ál & Massa sp. no . (holo ype); 38 – B. auloge i (Abeille de Pe in, 1898) (Tunisia:
Cap Bon peninsula, 6 km W o El Hawa iah, DKCP); 39 – B. bocchus (Mo occo: Gaada de Debdou pla eau., E
o Rchida, DKCP); 40 – B. b ancoi (allo ype); 41 – B. gallicus (Spain: Andalusia, Alja aque ill. en ., JSCP);
42 – B. howdeni (allo ype); 43 – B. nikolaje i (allo ype); 44 – B. auloge i (Tunisia: Cap Bon peninsula, 6 km W
o El Hawa iah, DKCP).
Ac a En omologica Musei Na ionalis P agae, 56(1), 2016 229
cosa le de Odon, 2.iii.1986, 1  (MSNG); Villa icosa le de Odon, 21.i .1983, 1  (MSNG); Villa icosa de Odon
(Mad id), 27. .1977, 2 , J. I. López-Colón & An onio Pe ucho lg . (OHCB). VALENCIA AUTONOM. COMM.:
VALENCIA PROV.: Ayo a (Valencia), 16. .1979, 1 , López-Colón lg . (OHCB); La Vallesa, 2.iii.1997, 1  1 
(GMCL). No u he locali y speci i ed: Espagne, 1  (NMPC); Hisp[ania]., 1  (NMPC); Spanien, 1  (SMTD).
Redesc ip ion. Dis inc ly con ex, su ace black, shiny, p ono um co e ed wi h double punc-
a ion, ely a glab ous.
Male. Head (Figs 3, 16, 23, 29, 35). Clypeus b oadly ounded, nea ly semici cula , an e io
angles ague. Pos e io angles o clypeal ma gin no ele a ed. F on al ho n well de eloped,
long, conical, apex widened, iangula in on al iew, base conical, no keeled, apex simple,
unmodi i ed. Punc a ion o clypeal disc simple, dis inc . Oblique keels abo e eyes app oxima-
ely eaching eyes. Punc a ion o ons and e ex same as he one on clypeus; on o-clypeal
su u e p esen . Eyes dis inc ly ellip ic, well de eloped; genae e enly ounded, semici cula .
P ono um (Figs 3, 16, 23, 29) dis inc ly ans e se; b oades jus in basal qua e , en i ely
bo de ed; la e al ma gin e enly ounded, ma ginal ca ina no c ena e, dis inc ly widened and
ele a ed; la e al o ea almos missing. P ono al punc a ion dis inc and double, consis ing
om well and deeply imp essed punc u es, excep on basal pa ; punc u es sepa a ed app o-
xima ely by dis ance equal o hei diame e , in e mixed wi h e y i ne pinned punc u es.
La e al ho nlike apophyses well de eloped, conical, ben upwa ds and sidewa ds. Apical
ho nlike apophyses dis inc bu weakly ele a ed, ob use-angled, di ec ed 30° on oapically.
Bo h apical ho nlike apophyses joined in keel.
Scu ella shield iangula , as long as wide, shiny, densely punc a e.
Ely a app oxima ely as long as wide, su ace shiny, no punc a e; hume al umbone weak;
be ween su u e and hume us se en dis inc s iae along en i e ely al leng h, all s iae wi h
con l uen punc u es; in e als con ex.
Pa ame es as in Fig. 62.
Va iabili y in males. Su ace black o b ownish; p ono al ho nlike apophyses in medium
de eloped and unde de eloped (hypo helic) specimens sho , mo e o less s aigh , only
simply ounded o almos acu e apically.
Female. Head (Figs 10, 41). Clypeus b oadly ounded, nea ly semici cula , an e io angles
ague. Pos e io angles o clypeal ma gin no ele a ed. F on al ca ina well de eloped, no
in e up ed in middle, doubly s-shaped. Punc a ion o clypeus, ons, e ex and occipu simple,
dis inc ; oblique keels abo e eyes dis inc , elonga ed o base o eyes; eyes well de eloped,
dis inc ly ellip ic, genae well de eloped, a cua e an e io ly.
P ono um (Fig. 10) dis inc ly ans e se, la e al ma gin weakly widened, b oades jus
basally, su ace dis inc ly deeply punc a e, punc a ion double excep o base, punc u es con-
sis ing o well and deeply imp essed punc u es, sepa a ed app oxima ely by dis ance equal
o hei diame e , in e mixed wi h e y i ne, pinned punc u es. An e io ans e se p o usion
dis inc , sha ply semici cula , la e al apophyses p esen .
Ely on su ace shiny, glab ous; hume al umbone weak.
Measu emen s. To al body leng h 11.0–14.6 mm.
Di e en ial diagnosis. Re e o species key.
Collec ing ci cums ances. P e e s sandy soils.
Dis ibu ion. Sou he n and sou h-wes e n F ance, Po ugal, No h-wes e n and Cen al Spain
(Fig. 59). We p o ide i s eco ds om Co sica and he Midi-Py énées Region o F ance.
HILLERT e al.: Bolbelasmus o he Wes e n and Sou he n Medi e anean (Geo upidae)
230
Bolbelasmus howdeni Hille & K ál sp. no .
(Figs 4, 11, 17, 24, 30, 36, 42, 59, 63)
Bolbelasmus gallicus [pa im, popula ions om SE Spain (app oxima ely Andalusia)]: BAGUENA CORELLA (1967):
262 (monog aph); KRIKKEN (1977): 284 ( e ision); BARAUD (1977): 159 (monog aph); BARAUD (1992): 45 (mo-
nog aph); MARTÍN-PIERA & LÓPEZ-COLÓN (2000): 180 (monog aph); KRÁL e al. (2006): 82 (ca alogue).
Type locali y. Spain, Andalusia, Cas illo de Cas illa , Algeci as en ., 10 km N o San Roque, 36°12′N 05°23′W,
78 m a.s.l.
Type ma e ial (271 specimens). SPAIN: ANDALUSIA AUTONOM. COMM.: CÁDIZ PROV.: HOLOTYPE:  (DKPC),
‘Spain, Andalusia, Cas illo de Cas illa , Algeci as en ., 10 km N o San Roque, 27.ii.2011, O. Hille lg .’ ALLOTYPE:
 (DKCP), same da a. PARATYPES: ‘Spain, 22.-28.2.2015, San Roque, Almo aima, N36°17’24.94” W5°23’40.77”,
136 m, lg . P. Kylies’, 3  1  (DKCP), 22  7  (PKCS); same da a bu lg . I. Ma inů, 1  1  (DKCP), 9
 4  (IMCO); same da a bu lg . R. Muczka, 3  (RMCM); ‘Spain, Andalusia, Alcala de los Gazules, E o
Cadiz, 01.-11.iii.2014 , Hille lg .’, 4  (OHCB); ‘Spain, Andalusia, E o Alcala, Sie a del Aljibe, E o Cadiz,
01.-11.iii.2014, Hille lg .’, 1  1  (OHCB); ‘España, A cos de la Fon e a, Cadiz, ex coll. A. D umon , 9.xii.1989’,
1  (GMCL); ‘Spain, A cos de la F on e a, Cádiz – B. Jo dan leg., 9.xii.1989’, 1  (JNCS); ‘Spain, Andalusia,
Cas illo de Cas illa , Algeci as en ., 10 km N o San Roque, 13.–20.i.2015, Hille lg .’, 4  2  (OHCB);
‘Spain, Andalusia, Cas illo de Cas illa , Algeci as en ., 10 km N o San Roque, 01.-11.iii.2014, Hille lg .’, 3 
(OHCB); ‘Spain, Andalusia, Cas illo de Cas illa , Algeci as en ., 10 km N o San Roque, 20.–27.ii.2012, O. Hille
lg .’, 1  (OHCB); ‘Spain, Andalusia, Cas illo de Cas illa , Algeci as en ., 10 km N o San Roque, 27.ii.2011, O.
Hille lg .’, 5  (OHCB); ‘Spain, Cádiz dis ., CASTILLO DE CASTILLAR ill.en ., 36°18.186 N 05°25.401
E, 27.ii.2009, Da id K ál lg .’, 1  2  (DKCP); ‘Spain, Andalusia, Cas illo de Cas illa , Algeci as en ., 10
km N o San Roque, 26.ii.2009, O. Hille lg .’, 1  (GNCA), 1  (HHCO), 2  1  (OHCB); ‘Spain - Cadiz,
13.1.2009, San Roque - Cas illa en ., lg . P. Kylies’, 1  (PKCS); ‘Spain, Andalusia, Cas illo de Cas illa , Alge-
ci as en ., 10 km N o San Roque, 02.iii.2008, O. Hille lg .’, 4  (OHCB); ‘ESPANA Cadiz, Pe os del Rey. S.
Roque, 21.i.[19]87, M. SALER’, 6  (SZCM); ‘1.3.2014, N Cas ella de la F on e a, Andalusia, Spain’, 1  1 
(DKCP), 3  1  (ZLCK); ‘Chiclana, 6.iii.[18]90, Ko b [lg .]’, 1  (ZSMC); ‘Chiclana, [18]83, Ko b [lg .]’ 1 
(ZSMC); ‘Spain, 22.-28.2.2015, Facinas en ., lg . P. Kylies’, 1  (PKCS); ‘Spain, La Sua a, Je ez de la F on e a,
Cádiz – A. Ve dugo leg., 5. .1988’, 2 , same da a bu 18.x.1987, 1  (JUCS); ‘Spain, Andalusia, 10 km NE o
Medina, E o Cadiz, 01.-11.iii.2014, O. Hille lg .’, 3  2  (OHCB); ‘Spain, Pina del Rey, San Roque, Cadiz,
26-i-2013., J. Na a o leg.’, 1  1  (AHCL), 1 ,1  (OBCB), same bu ‘27-xii-2014, J. Mª U bano leg.’, 1  1
 (JUCS), same bu ‘20-xii-2014, J. Mª U bano & J. Na a o leg.’, 6  3  (JNCS), 4  2  (AHCL), 11
 8  (JUCS), same bu ‘10-xi-2014, J. Na a o leg.’, 1  (JUCS), same bu ‘13-iii-2014, J. Na a o leg.’, 1
 (JUCS), same bu ‘26-i-2014, J. Mª U bano leg.’, 2  2  (JUCS), same bu ‘14-XI-2015, J. Na a o & J.
Mª U bano leg.’, 4  3  (JUCS), 4  3  (JNCS); ‘Spain, Pina de l Rey, Tajo del Paja aco, San Roque,
Cádiz – J. Na a o leg., 22.xii.2012’ 5  2  (JNCS), 2.i .2011, 2  (JNCS); ‘Spain, 22.-28.2.2015, Ta i a,
Pun a Paloma, N36°04’03.96” W5°42’08.47”, 56 m, lg . P. Kylies’, 2  (PKCS); same da a bu lg . I. Ma inů,
3  1  (IMCO); same da a bu R. Muczka, 3  1  (RMCM); ‘Spain - Cadiz, 14.-16.2.2008, San Roque,
Almo ama, lg . P. Kylies’, 1  (PKCS); ‘E - Cádiz / San Roque / 13.xi.[19]93 Rami ez lg.’, 1  (SZCM); ‘Espania,
San Roque, Cadiz, 13.xi.1988, Jose Luis To es lg .’, 1  1  (OHCB); ‘Esp., SAN ROQUE, Cádiz, 6.1987, J.
Ramí ez lg .’, 2  1  (RCCP); ‘E, Andalusia, Cadiz, Pina de San Roque, 2.iii.86, Hend ich/Mendez lg .’, 2 
(OHCB) , 1  (VTCZ); ‘E, Andalucia, Cadiz, Pina d. San Roque, b. San Roque, 2.iii.86, lg . Hend ich/Mendez’,
4  (LHCB); ‘SPAIN-Cádiz, S. Roque, 26.i.1984’, 1  (SZCM); ‘San Roque – Cadiz, España su , leg. Mendez
T., 17.iii.[19]84’, 1  (JHCP); ‘Spanien, Andalusien, San Roque, 17.i .1983, J.L. To es-Mendez lg .’, 4  1 
(LHCB); ‘San Roque, Cadiz, Spain, 30-I.1983, V. O uño lg .’, 1  (VMCP); ‘Spain, Andalusia, NW o Ta i a,
Algeci as en ., 13-20.i.2015, O. Hille lg .’, 6  2  (OHCB); ‘Spain, Andalusia, NW o Ta i a, pine o es ,
Algeci as en ., 01.-11.iii.2014, Hille lg .’, 2  2  (OHCB); ‘Spain, Andalusia, NW o Ta i a, beach sand,
Algeci as en ., 01.-11.iii.2014, Hille lg .’, 5  10  (OHCB); ‘Spain, Andalusia, NW o Ta i a, Algeci as en .,
13-27.ii.2013, O. Hille lg .’, 3  (OHCB); ‘Spain, Mon ehumada, Ta i a, Cádiz – J. Na a o leg., 22.xii.2012’,
2  1  (JNCS); ‘Spain, Andalusia, Sanluca de Ba ameda, NW o Je ez, 26.ii.2011, O. Hille lg .’, 1  2 
(OHCB); ‘Ta i a (Cadiz), 11.i .1980, Lopez Colon leg’, 1  (JLCM). GRANADA PROV.: ‘Spain, Andalusia, 5 km E
Ac a En omologica Musei Na ionalis P agae, 56(1), 2016 231
o G anada, Sie a de la Yed a, 01.-11.iii.2014, Hille lg .’, 1  (OHCB); ‘España, La Sag a, G anada, Andalucia,
.1988’, 1  1  (GMCL). – JAÉN PROV.: ‘Spain, Las Minas. A ayanes. Lina es, Jaén, ii-1997, A. Hidalgo leg.’, 1 
(AHCL); ‘Spain, Mi anda del Rey, Jaén, 13-xi-2015, Ja ie Cas illo & An onio Hidalgo leg.’, 4  2  (JCCL),
3  2  (AHCL).‘Spain, Rese a las Vibo as, Sie a Mo ena. Mon izón, Jaén, 9-i -1995, A. Hidalgo leg.’, 1 
(AHCL). MÁLAGA PROV.: ‘Spain, Andalusia, Sie a Palmi e a, Pue o de Alija , W o Mabella, 01.-11.iii.2014, Hille
lg ., 1  (OHCB)’. CASTILE-LA MANCHA AUTONOM. COMM.: ALBACETE PROV.: ‘Molinicos, 10. .[18]95’, 1  2 
(ZFMK). GIBRALTAR: ‘Gib al a ., J. J. Walke [lg .]’ 1 , 2  (BMHN).
Desc ip ion. Male (holo ype). Dis inc ly con ex, su ace black, shiny, p ono um co e ed
wi h double punc a ion, ely a glab ous.
Head (Figs 4, 17, 24, 30, 36). Clypeus b oadly ounded, nea ly semici cula , an e io angles
ague. Pos e io angles o clypeal ma gin no ele a ed. F on al ho n well de eloped, long,
base conical, no keeled, apex simple, unmodi i ed. Punc a ion o clypeal disc simple, dis inc .
Oblique keels abo e eyes app oxima ely eaching eyes. Punc a ion o ons and e ex same
as on clypeus; on o-clypeal su u e p esen . Eyes dis inc ly ellip ic and well de eloped; genae
e enly ounded, semici cula .
P ono um (Figs 4, 17, 24, 30) dis inc ly ans e se; b oades jus in basal qua e , en i ely
bo de ed; la e al ma gin e enly ounded, ma ginal ca ina no c ena e, dis inc ly widened and
ele a ed; la e al o ea almos missing. P ono al punc a ion dis inc , double, consis ing o well
and deeply imp essed punc u es excep o basal pa , punc u es sepa a ed app oxima ely by
dis ance equal o hei diame e , in e mixed wi h e y i ne, pinned punc u es. La e al ho n-
like apophyses well de eloped, conical, ben up and sidewa ds. Apical ho nlike apophyses
dis inc bu weakly ele a ed, ob use angled, di ec ed 45° on oapically. Bo h apical ho nlike
apophyses joined oge he in keel-like seam.
Scu ella shield iangula , app oxima ely as long as wide, shiny, densely punc a e.
Ely a app oxima ely as long as wide, su ace shiny, glab ous; hume al umbone weak;
be ween su u e and hume us se en dis inc s iae along en i e ely al leng h, all s iae wi h
con l uen punc u es; in e als con ex.
Pa ame es as in Fig. 63.
Va iabili y in males. Su ace black o b ownish; p ono al ho nlike apophyses in medium
de eloped and unde de eloped (hypo helic) specimens sho , mo e o less s aigh , only
simply ounded o almos acu e apically.
Female (allo ype). Head (Figs 11, 42). Clypeus b oadly ounded, nea ly semici cula ,
an e io angles ague. Pos e io angles o clypeal ma gin no ele a ed. F on al ca ina well
de eloped, no in e up ed in middle, doubly s-shaped. Punc a ion o clypeus, ons, e ex
and occipu simple, dis inc ; oblique keels abo e eyes dis inc , elonga ed o base o eyes; eye
well de eloped, dis inc ly ellip ic, genae well de eloped, a cua e an e io ly.
P ono um (Fig. 11) dis inc ly ans e se, la e al ma gin weakly widened, b oades jus
basally, su ace dis inc ly deeply punc a e, doubly excep o basal pa , punc u es consis ing
o well and deeply imp essed punc u es, sepa a ed app oxima ely by dis ance equal o hei
diame e , in e mixed wi h e y i ne, pinned punc u es. An e io ans e se p o usion dis inc ,
sha ply semici cula , la e al apophyses p esen .
Ely on su ace shiny, glab ous; hume al umbone weak.
Measu emen s. To al body leng h 11.3–15.2 mm (holo ype, : 14.6 mm; allo ype, :
15 mm).

HILLERT e al.: Bolbelasmus o he Wes e n and Sou he n Medi e anean (Geo upidae)
232
Di e en ial diagnosis. Re e o species key.
E ymology. Pa onymic; named in honou o ou iend, he la e Hen y F. Howden (O awa,
Canada), a enowned specialis in wo ld Sca abaeoidea.
Collec ion ci cumc ances. P e e s sandy soils.
Dis ibu ion. Sou h-eas e n pa s o Spain and Gib al a (Fig. 59).
Bolbelasmus nikolaje i Hille , A none, K ál & Massa sp. no .
(Figs 5, 12, 18, 25, 31, 37, 43, 57, 59, 64)
Bolbelasmus bocchus: PIC (1924): 126 (no e, dis ibu ion); SCHATZMAYR (1946): 50 (dis ibu ion); ALFIERI (1976):
206 (dis ibu ion).
Bolbelasmus bocchus auloge i: KRIKKEN (1977): 290, i gs 18–19 ( e ision); BARAUD (1992): 53 (monog aph); KRÁL
e al. (2006): 82 (ca alogue); ARNONE & MASSA (2010): 411, i g. 12 (dis ibu ion, no e).
Type locali y. NW Tunisia, Sidi Bouzid go .,7.5 km SWW o Hajeb el Ayoun, by oad, 35°22′N, 09°28′E, 340 m a.s.l.
Type ma e ial (126 specimens). TUNISIA: SIDI BOUZID PROV.: HOLOTYPE:  (DKPC), ‘NW Tunisia, Sidi Bouzid go .,
7.5 km SWW o , HAJEB EL AYOUN, by oad, 29.iii.2014, 35°22’N, 09°28’E, 340 m, Da id K ál lg .’ ALLOTYPE: 
(DKCP), same da a. PARATYPES: EGYPT: ALEXANDRIA PROV.: ‘Egyp , Alexand ia, 1958’, 1  (NMPC); ‘Egyp , Ma iou ,
4.iii.1924, coll. Al i e i’, 1  (NHMB). LIBYA: AZ ZAWIYAH PROV.: ‘A ika, Libye, SURMAN - iii.[19]88, Č. Sobo a
lg .’, 1  1  (JDCC). BENGHAZI PROV.: ‘Libya, Benghazi, 8.3.1987, lg . Há a [p]’, 1  (DKCP), 1  (JHCP); ‘Libia,
Benghazi, 1986 V. Poláček lg . [p]’, 1  (SJCP). BUTNAN PROV.: ‘Libya, Ci enaica, Tob uk III, 32°5‘1“ N, 23°58‘ 35“
E, Geo C. K uge lg ., R.U. Ag a io’, 2  2  (MSNG). TRIPOLI PROV.: ‘A . Libya – Tadjiu a, 04.1986, Houška
lg .’, 2  (JMCH); ‘Libya, Tagiu a, če enec [= July], [19]85, Voleman [lg .]’, 2  (VTCZ); ‘Libya, Tagiu a, s pen
[= Augus ], [19]85, Voleman [lg .]’, 1  (VTCZ); ‘AFRIKA, Libya, Ta abulus ii.[19]88, V. Poláček lg .’, 1  (JMCP);
Foualgha, 8.4.1983, M. Dudycha [lg .] [handw i en]’, 1  (RCCP). No loca ed: ‘Libya, Takad ia, s pen [= Augus ],
[19]84, Voleman [lg .]’, 1  (VTCZ). TUNISIA: BIZERTE PROV.: ‘Bise a, Tunisia, De Vauloge , Ne inson Coll.,
1918-14, Bolboce as, auloge i Ab., ypes, Type, H. T.’, 1  (BMNH) [no syn ype o B. auloge i, ype label added
la e ; M. Ba clay pe s. comm.]. GABÈS PROV.: ‘Gabes, 1897’, 2  (MNHN); ‘Tunisie, Ma e h, R. Demo l ys lg ., i.
[19]49’, 1  (MNHN); ‘Tunisia, Go . Gabes, Ma ma a, 19.i .2013, Leg. G.Saba inelli, A ligh ’, 1  1  (GSCT);
‘Ma ma a, iii.2014’, 1  1  (GSCT); ‘TUNISIA: 15.4.1996, TAMEZRET, 13 km W o Ma ma a, J. BATELKA lg .’,
2  (JBCP); ‘E Tunisia, Gabes, 5 km NEE o Oud e , Ma wiya plage, 24.–25.iii.2014, 34°01‘N 10°00‘E, O. Hille
lg .’, 7  8  (OHCB); ‘E Tunisia, Gabes go ., 5 km NEE o Oud e , MATWIYA plage, ca. 20 m, 24.–25.iii.2014,
34°01‘N, 10°00‘E, Da id K ál lg .’, 2  3  (DKCP), same da a bu Pe Šípek lg ., 3  2  (CUPC), 1 
[in pu e alcohol] (CUPC); ‘TN-Gabes, din o ni, 13.IV.[19]95 leg. S. Ziani’, 1  (SZCM). GAFSA PROV.: ‘Tunisie,
Ga sa, Ch e ien [lg .], 1910’, 1  (MNHN); ‘Tunisie, Ga sa, A. Weiss [lg .], 1904’, 1  (MNHN); ‘Ga sa, 1897’, 1 
(OHCB);‘Tunis, Mé laoui, C. Dumon ’, 1  (DKCC). KASSERINE PROV.: ‘T., Thelep e, Mai 1941, R. Demo l ys [lg .]’,
2  1  (MNHN); ‘Tun., Sbei la, Ma s 1941, R. Demo l ys [lg .]’, 2  (MNHN). KEBILI PROV.: ‘TUNISIA, 3.-4.
i .2008 / 45 km E o Douz, Wadi Raml, / N33°31’ E09°30’ / J. Ba elka & J. S aka lg .’, 1  (JBCP). KEF PROV.: ‘Tunisia
NW, Le Ke , NABEUR en ., 16.4.2001, lg . M. Snížek’, 1  (VKCC). MEDENINE PROV.: ‘E Tunisia, Medenine, Dje ba
island, 5 km E o Midoun, sea coas , 26.iii.2014, 33°49‘N 11°02‘E, O. Hille lg .’, 1  (OHCB); ‘TN-Medenine, s .
pe Toujan, 13.IV.[19]95 leg. S. Ziani’, 4  (SZCM). NABEUL PROV.: ‘Tunisia, (Nabeul), Cap Bon peninsula, 6 km
W o El Hawa iah, sea coas , 23.iii.2014, 37°03‘N 10°57‘E, O. Hille lg .’, 4  (OHCB); ‘NE Tunisia, Nabeul go .,
Cap Bon peninsula, 6 km W o EL HAWARIAH, sea coas , 23.iii.2014, 37°03‘N, 10°57‘E, ca. 3 m, Da id K ál lg .’, 2
 4  (DKCP), same da a bu Pe Šípek lg ., 1  1  [pa ially demaged spec. in pu e alcohol] (CUPC); ‘Tunisia,
Cap Bon IV.1960, G. F ey’, 1  (NHMB). SFAX PROV.: ‘Tunisie, Ke ana [Bi Ke ana], i .[19]50, R. Demo l ys [lg .]’, 1
 (MNHN); ‘Tunisie, La Hencha, iii.[19]42, R. Demo l ys [lg .]’, 1  (MNHN); ‘Tunisia 1896, S ax, de Vauloge [lg .],
Ne inson Coll., 1918-14’, 1  1  (BMNH). SIDI BOUZID PROV.: ‘NE Tunisia, Sidi Bouzid, 7.5 km SWW o Hajeb el
Ayoun by oad, 29.iii.2014, 35°22‘N 09°28‘E, O. Hille lg .’, 7  8  (OHCB); ‘NW Tunisia, Sidi Bouzid go ., 7.5
km SWW o , HAJEB EL AYOUN, by oad, 29.iii.2014, 35°22‘N, 09°28‘E, 340 m, Da id K ál lg .’, 4  2  (DKCP),
1  1  (CUPC); ‘Tunisie, Maknassy, A. Dumon [lg .], 1928’, 5  2  (MNHN); ‘Tunisia, Bled Gamouda [Bilad
Ac a En omologica Musei Na ionalis P agae, 56(1), 2016 233
Gamouda], de Vauloge [lg .], Ne inson Coll., 1918-14.’, 1  (BMNH). TOZEUR PROV.: ‘Tunis, Tozeu , C. Dumon
[lg .], 1929’, 2  (MNHN). No u he locali y speci i ed: ‘Tunisie, Sep en [ ionalis], Schaedlin’, 1  (MNHN).
Desc ip ion. Male (holo ype). Dis inc ly con ex, su ace b ownish o ligh -b own, shiny,
p ono um co e ed wi h double punc a ion, ely on e y i ne mic osculp u ed basally, mic o-
scul u e missing apically.
Head (Figs 5, 18, 25, 31, 37, 57). Clypeus b oadly ounded, nea ly semici cula , an e io
angles ague. Pos e io angles o clypeal ma gin ele a ed. F on al ho n well de eloped, long,
dis inc ly u ca e, base iangula ly shaped in do sal iew, pos e io edge o base si ua ed
a heigh o cen e o eye, side ma gin o ube cle keeled om base o apex. Punc a ion o
clypeal disc simple, dis inc . Oblique keels abo e eyes eaching app oxima ely o hal o
eyes. Punc a ion o ons and e ex same as on clypeus; on oclypeal su u e p esen . Eyes
dis inc ly ellip ic and well de eloped; genae a cua e pos e io ly, ex ending eyes la e ally.
P ono um (Figs 5, 18, 25, 31, 37, 57) dis inc ly ans e se; b oades jus in basal qua e ,
en i ely bo de ed; la e al ma gin e enly ounded, ma ginal ca ina no c ena e, dis inc ly
widened and ele a ed; la e al o ea almos missing. P ono al punc a ion dis inc , double,
consis ing o deeply imp essed punc u es, punc u es sepa a ed by app oxima ely 2–3 imes
hei diame e , in e mixed wi h e y i ne, pinned punc u es. Base o p ono um glab ous.
La e al ho nlike apophyses well de eloped, ben dis inc ly upwa ds and sidewa ds in la e al
and do sal iew; be ween la e al and apical apophyses dis inc , deeply o al conca i y. Apical
ho nlike apophyses dis inc ly ele a ed, angled, di ec ed 30° on oapically, subapical p o u-
sion o apical apophyses la e ally nea apex on ou side.
Scu ella shield iangula elonga e, e y i nely punc a e, shiny.
Ely a app oxima ely as long as wide, su ace shiny, glab ous; be ween su u e and hume -
us 7 dis inc s iae along en i e ely al leng h, all s iae wi h dis inc ly sepa a ed punc u es;
in e als l a ; hume al umbone weak.
Pa ame es as in Fig. 64.
Va iabili y in males. Su ace black o b ownish; p ono al ho nlike apophyses in medium
de eloped and unde de eloped (hypo helic) specimens sho , mo e o less s aigh , only
simply ounded o almos acu e apically.
Female (allo ype). Head (Figs 12, 43). Clypeus b oadly ounded, semici cula wi h s aigh
an e io ma gin, an e io angles e enly ounded. Pos e io angles o clypeal ma gin e y
weakly ele a ed. F on al ca ina well de eloped, dis inc ly in e up ed in middle, bi ube cula e
on in e nal in e up ion, la e ally keeled on each side. Punc a ion o clypeus, ons, e ex
and occipu simple, dis inc ; oblique keels abo e eyes dis inc , elonga ed o hal o eyes; eye
well de eloped, dis inc ly ellip ic, genae well de eloped, a cua e an e io ly and pos e io ly.
P ono um (Fig. 12) dis inc ly ans e se, la e al ma gin weakly widened, b oades jus
basally, su ace dis inc ly deeply punc a e, double excep o basal pa , punc u es consis ing
o well and deeply imp essed punc u es, sepa a ed app oxima ely by dis ance equal o hei
diame e , in e mixed wi h e y i ne, pinned punc u es. An e io ans e se p o usion dis inc ,
sha ply semici cula , la e al apophyses p esen .
Ely on su ace shiny, glab ous; hume al umbone weak.
Measu emen s. To al body leng h 12.0–15.1 mm (holo ype, : 14.6 mm; allo ype, :
14.8 mm).
HILLERT e al.: Bolbelasmus o he Wes e n and Sou he n Medi e anean (Geo upidae)
234
Di e en ial diagnosis. Re e o species key.
E ymology. Pa onymic; named in honou o ou iend Geo gy V. Nikolaje (Al-Fa abi S a e
Uni e si y, Alma y, Kazakhs an), a enowned specialis in wo ld Sca abaeoidea.
Collec ion ci cums ances. P e e s sandy and/o loess soils.
Dis ibu ion. Medi e anean Egyp , Libya (T ipoli ania, Cy enaica) and Tunisia (Fig. 59).
Bolbelasmus auloge i (Abeille de Pe in, 1898) s a . es i .
(Figs 6, 7, 13, 19, 20, 26, 32, 38, 44, 52–56, 58, 59, 65)
Bolboce as Vauloge i Abeille de Pe in, 1898: 254 (o iginal desc ip ion).
Bolbelasmus Vauloge i: BOUCOMONT (1912): 17 (ca alogue, as synonym o B. bocchus); WINKLER (1927): 1035
(ca alogue, synonym o B. bocchus).
Bolbelasmus omano um A none & Massa, 2010: 403, i gs 1–5, 8 (o iginal desc ip ion). Type locali y: ‘Sicily,
Bales a e, oce Cala ubo (Pale mo)’. New synonym.
Bolbelasmus unico nis: BALLERIO (2008): 32 (dis ibu ion).
Type locali y. Tunisia, ‘Bize e’.
Type ma e ial examined (20 specimens). Bolbelasmus auloge i: TUNISIA: BIZERTE p o .: LECTOTYPE (p esen
designa ion):  (MNHN), ‘Bize e, de Vauloge , Bolboce as Vauloge i ab., Museum Pa is 1919, coll A de Pe in,
Bolboce as Vauloge i ab., Museum Pa is, 1942 coll. D . A Chobau ’. PARALECTOTYPES: No. 1,  (MNHN), ‘Bize e,
de Vauloge , Vauloge i Ab., Type’; No. 2,  (MNHN), ‘Bize e de Vauloge sp. no ??, uniseus Vaul. (sp. li eus),
Museum Pa is 1942, coll. D . A Chobau , Bolboce us Vauloge i ab’; Nos 3–4,  (MNHN), ‘Bize e, de Vauloge
, Vauloge i Ab., Type’; No. 5,  (ZSMC), ‘Bize e [18]93 Vaulog[e ].’; No. 6,  (BMNH), ‘Bize e, de Vauloge ,
Ne inson Coll., 1918-14., Co ype’; No. 7,  (BMNH), ‘Bize e, de Vauloge , Ne inson Coll., 1918-14., Co ype’;
see Figs 52–56.
Bolbelasmus omano um: ITALY: SICILY PROV.: HOLOTYPE: 1  (MSNG), ‘I aly, Sicily, Bales a e, oce Cala ubo
(Pale mo), 17.iii.1974, B. Massa lg .’. ALLOTYPE: 1  (MSNG), ‘Sicily, Pedalino (Ragusa) 1. .1972, A. Monas a lg .’.
PARATYPES: 1  5  (MAPC), ‘Men i (Ag igen o), loc. Cinquan a, 13.xi.2010, 4.i .2010, 28.iii.2010, 20.iii.2010,
14.xi.2009, 6.xi.2009, A. Ca apezza lg .’; 1  (MACP), ‘Vendica i Na u e Rese e (Si acusa) 18.i .2009, B. Massa
lg .’; 1  (VACP), ‘Ma ausa (T apani) 18.xii.1983, V. Aliquò lg .’; 1  (OHCB), ‘Bosco San o Pie o Na u e Rese e
(Ca ania), M. Za a ana lg .’; 1  1  (MZUC) ‘Sicily’.
Addi ional ma e ial examined (77 specimens). ALGERIA: No u he locali y speci i ed: Alge ie, D . Thiè-
baul , 1  (MNHN). ITALY: SICILY PROV.: Men i (Ag igen o), loc. Cinquan a, 19.iii.[20]13, 1 , M. A none lg .
(MAPC); Te asini, Capo Rama Na u e Rese e 27.iii.[20]12, 1 , R. Lo Duca lg . (MAPC); Men i , loc. Cinquan a,
10.i .2011, 1 , A. Ca apezza lg . (DJCP); Niscemi (Cal anisse a), 1 , M. Za a ana lg . (MSNM); Pale mo, 1 
(MZUC); Piazza A me ina (Enna), 1  (MZUC); Te mini Ime ese (Pale mo), 1  (MZUC). TUNISIA: BIZERTE
PROV.: SW en . o Edah a, 31.iii.2014, 37°12′N 09°20′E, 3  5 , O. Hille lg . (OHCB); SW en . o Edah a,
37°12′N 09°20′E, ca. 65 m, 31.iii.2014, 4  3 , Da id K ál lg . (DKCP), 3  2  (CUPC). JENDOUBA PROV.:
Taba ka, Zone ou is ique, 2. .2005, 1 , M. Liebsche lg . (MLCD). NABEUL PROV.: Cap Bon peninsula, 6 km W
o El Hawa i h, sea coas , 23.iii.2014, 37°03′N 10°57′E, 3  2 , O. Hille lg . (OHCB); Cap Bon peninsula,
6 km W o El Hawa iah, sea coas , 37°03′N 10°57′E, ca. 3 m, 23.iii.2014, 3  6 , Da id K ál lg . (DKCP);
Cap Bon peninsula, 2 km NW o Douela by oad, 23.–24.iii.2014, 36°50′N 10°35′E, 13  2 , O. Hille lg .
(OHCB); Cap Bon peninsula, 2 km NW o Douela by oad, 36°49′N 10°35′E, ca. 130 m, 23.–24.iii.2014, 4  2
, Da id K ál lg . (DKCP), 1  [pa ially damaged, in pu e alcohol] (CUPC); Cap Bon peninsula, 6 km SW o
Zouie el Mgaiez by oad, 23.iii.2014, 36°58′N 10°55′E, 3  2 , O. Hille lg . (OHCB); Cap Bon peninsula, 6
km SW o Zaouie El Mgaiez by oad, 36°49′N 10°55′E, ca. 45 m, 23.iii.2014, 1  2 , Da id K ál lg . (DKCP);
Cap Bon, 1960, 1 , G. F ey [lg .] (NHMB); Cap Bon, 1894, Museum Pa is, 1919, 1 , coll. A. de Pe in (MNHN).
No u he locali y speci i ed: Tunisia, Sal Lake, K e en [May] 2005, 1 , Volcano lg . (OBCB); Tunisia, Solunan,
Museum Pa is 1993, 1 , coll J. Ba aud (MNHN).
Redesc ip ion. Male (lec o ype) (Figs 52–54). Dis inc ly con ex, su ace black, shiny, p o-
no um co e ed wi h double punc a ion, ely on glab ous.
Ac a En omologica Musei Na ionalis P agae, 56(1), 2016 235
Figs 45–51. Types and labels o Bolbelasmus bocchus (E ichson, 1841), deposi ed in MNHB. 45 – male habi us in
do sal aspec (lec o ype); 46 – male habi us in on al aspec (lec o ype); 47 – male habi us in la e al aspec (lec o-
ype); 48 – emale habi us in do sal aspec (pa alec o ype); 49 – labels o lec o ypes; 50 – labels o pa alec o ype
No. 1; 51 – labels o pa alec o ype No. 2.
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su ace b ownish, maximal body leng h 14 mm. Sou he n pa s o Cen al Eu ope,
Alsace, I aly, Balkan Peninsula, Poland, Russia and Uk aine. ...................................
.................................................................... B. unico nis (Sch ank on Paula, 1789)
4 (3) F on al ho n bending no o wa ds (Figs 19–20); ans e sal clypeal ca ina absen ;
scu ella shield iangula longe han wide; body su ace blackish, maximal body
leng h 13 mm (Figs 6–7, 38); aedeagus as in Fig. 65; Alge ia, I aly (Sicily) and Tu-
nisia (Fig. 59). ............................................... B. auloge i (Abeille de Pe in, 1898)
5 (2) F on al ho n u ca e apically; genae mo e o less angula e an e ola e ally; oblique
keel abo e eyes eaching only o an e io pa o eyes; head no s ongly cons ic ed
behind eyes.
6 (7) F on al ho n sho and no longe han b oad, ho n apex app oxima ely as wide as
ho n base (Figs 2, 15, 22); p ono al punc a ion dense wi h excep ion o basal pa ,
punc u es sepa a ed app oxima ely by dis ance equal o hei diame e s; p ono al
base wi h dis inc isible bu i ne, egula ly pinned poin s; ely al s iae well im-
p essed, isible as comple e lines; ely al in e als mode a ely con ex; aedeagus as
in Fig. 61; Spain (Fig. 59). ................................. B. b ancoi Hille & K ál sp. no .
7 (6) F on al ho n long, longe han b oad, ho n becoming g adually hinne om base o
apex (Figs 1, 5, 14, 18, 21, 25, 27, 31, 33, 37, 57); p ono al punc a ion spa se , punc-
u es sepa a ed by app oxima ely h ee imes hei diame e s; p ono al base wi h e y
i ne, weakly isible and well sepa a ed pinned poin s; ely al s iae no imp essed,
isible as ow o punc u es; ely al in e als l a ; no he n A ica.
8 (9) Base o on al ho n si ua ed a le el o an e io pa o eyes; ho n sha ply keeled
om base o hal o i s leng h in do sal aspec (Figs 1, 14, 21, 27, 33); medial ho n-
like apophysis o p ono um di ec ed e ically upwa ds in la e al aspec , subapical
p o usion si ua ed basally; la e al ho nlike apophyses o p ono um di ec ed o -
wa ds and weakly di e gen ; aedeagus as in Fig. 60; Alge ia and Mo occo (Fig.
59). ............................................................................... B. bocchus (E ichson, 1841)
9 (8) Base o on al ho n si ua ed a le el o pos e io pa o eyes; ho n b oadly keeled
om base o apex in do sal iew (Figs 5, 18, 25, 31, 37, 57); medial ho nlike apophy-
sis o p ono um di ec ed obliquely o wa ds in la e al aspec , subapical p o usion
si ua ed nea apex; la e al ho nlike apophyses o p ono um di ec ed o wa ds; aedea-
gus as in Fig 64; Egyp , Libya and Tunisia (Fig. 59). ................................................
............................................... B. nikolaje i Hille , A none, K ál & Massa sp. no .
10 (1) Scu ella shield densely punc a e; on al ho n simple apically.
11 (12) Apex o on al ho n app oxima ely as wide as ho n base, apex b oadly acu e-angled,
s ongly l a ened on on pa o apex (Figs 3, 16, 23, 29, 35); aedeagus as in Fig.
62; F ance (including Co sica), Po ugal and Spain (Fig. 59). ...................................
....................................................................................... B. gallicus (Mulsan , 1842)
12 (11) Apex o on al ho n becoming g adually hinne om base o apex, apex acu e-an-
gled, only weakly l a ened on on pa o apex (Figs 4, 17, 24, 30, 36); aedeagus as
in Fig. 63; Spain, Gib al a (Fig. 59). ................ B. howdeni Hille & K ál sp. no .

Ac a En omologica Musei Na ionalis P agae, 56(1), 2016 243
Iden i i ca ion key o less de eloped males
1 (10) Scu ella shield impunc a e o weakly shallowly punc a e, on al ho n simple o
u ca e apically.
2 (5) F on al ho n simple; genae egula ly ounded; oblique keel abo e eyes ex ending
o e whole eyes leng h; head s ongly cons ic ed behind eyes.
3 (4) Scu ella shield iangula , app oxima ely as long as wide; body su ace b ownish,
maximal body leng h 12 mm. Sou he n pa s o Cen al Eu ope, Alsace, I aly, Balkan
Peninsula, Poland, Russia and Uk aine. .... B. unico nis (Sch ank on Paula, 1789)
4 (3) Scu ella shield iangula , longe han wide; body su ace blackish, maximal body
leng h 11 mm; aedeagus as in Fig. 65; Alge ia, I aly (Sicily) and Tunisia (Fig. 59).
....................................................................... B. auloge i (Abeille de Pe in, 1898)
5 (2) F on al ho n u ca e apically; genae mo e o less angula e an e ola e ally; oblique
keel abo e eyes eaching only o an e io pa o eyes; head no s ongly cons ic ed
behind eyes.
6 (7) P ono al punc a ion dense wi h excep ion o basal pa , punc u es sepa a ed app oxi-
ma ely by dis ance equal o hei diame e s; p ono al base wi h dis inc isible bu
i ne, egula ly spaced pinned poin s, ely al s iae well imp essed, isible as com-
ple e lines; ely al in e als mode a ely con ex; aedeagus as in Fig. 61; Spain (Fig.
59). ...................................................................... B. b ancoi Hille & K ál sp. no .
7 (6) P ono al punc a ion spa se , punc u es sepa a ed by app oxima ely h ee imes hei
dia me e s p ono al base wi h e y i ne, weakly isible and well sepa a ed pinned
poin s; ely al s iae no imp essed, isible as ow o punc u es; ely al in e als l a ;
no he n A ica.
8 (9) Base o on al ho n si ua ed a le el o an e io pa o eyes; ho n sha ply keeled
basally only; aedeagus as in Fig. 60; Alge ia and Mo occo (Fig. 59). .......................
...................................................................................... B. bocchus (E ichson, 1841)
9 (8) Base o on al ho n si ua ed a le el o pos e io pa o eyes; ho n b oadly keeled
om base o apex in do sal iew; aedeagus as in Fig. 64; Egyp , Libya and Tunisia
(Fig. 59). ............................... B. nikolaje i Hille , A none, K ál & Massa sp. no .
10 (1) Scu ella shield densely punc a e; on al ho n simple apically.
11 (12) Apex o on al ho n l a ened on on pa o apex; aedeagus as in Fig 62; F ance
(including Co sica), Po ugal and Spain (Fig. 59). ....... B. gallicus (Mulsan , 1842)
12 (11) Apex o on al ho n egula ly a cua e, no l a ened on on pa o apex; aedeagus
as in Fig. 63; Spain, Gib al a (Fig. 59). ............ B. howdeni Hille & K ál sp. no .
Iden i i ca ion key o emales
1 (10) Scu ella shield impunc a e, o weakly shallowly punc a e.
2 (5) F on al ube cle simple (Figs 41, 42, 44); genae egula ly ounded; oblique keel
abo e eyes o e whole leng h o eyes; head s ongly cons ic ed behind eyes.
HILLERT e al.: Bolbelasmus o he Wes e n and Sou he n Medi e anean (Geo upidae)
244
3 (4) Scu ella shield iangula , app oxima ely as long as wide; body su ace b ownish,
minimal body leng h 12 mm. Sou he n pa s o Cen al Eu ope, Alsace, I aly, Balkan
Peninsula, Poland, Russia and Uk aine. .... B. unico nis (Sch ank on Paula, 1789)
4 (3) Scu ella shield iangula longe han wide; body su ace blackish, minimal body
leng h 11 mm (Fig 13); Alge ia, I aly (Sicily) and Tunisia (Fig. 59). ........................
....................................................................... B. auloge i (Abeille de Pe in, 1898)
5 (2) F on al ube cle u ca e apically (Figs 39–40, 43); genae mo e o less angula e an e-
ola e ally; oblique keel abo e eyes eaching only o an e io pa o eyes; head no
s ongly cons ic ed behind eyes.
6 (7) P ono al punc a ion dense wi h excep ion o basal pa , punc u es sepa a ed app oxi-
ma ely by dis ance equal o hei diame e s; p ono al base wi h dis inc isible bu
i ne, egula ly pinned poin s; ely al s iae well imp essed, isible as comple e lines;
ely al in e als mode a ely con ex (Fig. 9); Spain (Fig. 59). ...................................
............................................................................. B. b ancoi Hille & K ál sp. no .
7 (6) P ono al punc a ion spa se , punc u es sepa a ed by app oxima ely h ee imes
hei diame e s p ono al base wi h e y i ne weak isible and well sepa a ed pinned
poin s; ely al s iae no imp essed, isible as ow o punc u es; ely al in e als l a ;
no he n A ica.
8 (9) Base o on al ube cle si ua ed a le el o an e io pa o eyes; ube cle sha ply
keeled basally only in do sal iew (Fig. 8); Alge ia and Mo occo (Fig. 59). ............
...................................................................................... B. bocchus (E ichson, 1841)
9 (8) Base o on al ube cle si ua ed a le el o pos e io pa o eyes; ho n b oadly keeled
om base o apex in do sal iew (Fig. 12); Egyp , Libya and Tunisia (Fig. 59). ......
............................................... B. nikolaje i Hille , A none, K ál & Massa sp. no .
10 (1) Scu ella shield densely punc a e.
11 (12) F ance (including Co se), Po ugal and Spain (see dis ibu ion map on Fig. 59 o
de ails). ......................................................................... B. gallicus (Mulsan , 1842)
12 (11) Spain, Gib al a (see dis ibu ion map on Fig. 59, o de ails). ..................................
............................................................................ B. howdeni Hille & K ál sp. no .
Anno a ed lis o he Wes e n Palaea c ic Bolbelasmus species
wi h new eco ds
Bolbelasmus bocchus (E ichson, 1841)
Bolboce as Bocchus E ichson, 1841: 170 (o iginal desc ip ion).
= Bolboce as i ssico nis Mulsan , 1843: 280 (o iginal desc ip ion). Synonymized by MARSEUL (1857: 83).
Type locali ies. Bolboce as Bocchus: ‘Algie [Bona = Annaba]’; B. i ssico nis: ‘Algé ie’.
Dis ibu ion. Alge ia, Mo occo (p esen pape ).
Bolbelasmus b ancoi Hille & K ál sp. no .
Type locali y. Spain (Andalusia), Los Jesos, N slope o Sie a de Alhamilla, 37°05′N 02°17′W, 500 m a.s.l.
Dis ibu ion. Spain.
Ac a En omologica Musei Na ionalis P agae, 56(1), 2016 245
Bolbelasmus gallicus (Mulsan , 1842)
Bolboce as Gallicus Mulsan , 1842: 350 (o iginal desc ip ion).
= Bolboce as Gallicus Va . Conjunc us Mulsan , 1842: 351 (o iginal desc ip ion). Synonymized by BOUCOMONT
(1912: 17).
= Bolboce as Gallicus Va . P o incialis Mulsan , 1842: 351 (o iginal desc ip ion). Synonymized by BOUCOMONT
(1912: 17).
Type locali ies. Bolboce as gallicus: F ance, ‘exclusi emen mé idionale; D aguignan, Sain -T opez, en i ons de
Ma seille’; B. conjunc us: ‘F ance mé idionale’; B. p o incialis: ‘F ance mé idionale’.
Dis ibu ion. F ance (including Co sica – i s eco ds), Po ugal, Spain (p esen pape ).
Bolbelasmus howdeni Hille & K ál sp. no .
Type locali y. Spain, Andalusia, Cas illo de Cas illa , Algeci as en ., 10 km N o San Roque, 36°12′N 05°23′W,
78 m a.s.l.
Dis ibu ion. Spain, Gib al a (p esen pape ).
Bolbelasmus kei hi Miessen & T ichas, 2011
Bolbelasmus kei hi Miessen & T ichas, 2011: 183 (o iginal desc ip ion), i gs on pp. 187–188 [no numbe ed].
Type locali y. G eece, ‘en . Ag. Pa aske i (chu ch), Agia Ma ina, Kasos island, (La : 35°24′10.71″N, Long:
26°54′27.59″E)’.
Type ma e ial examined (2 specimens). PARATYPES: GREECE: CRETE: 1  1 , ‘ELAFONISI (Li e), 30/12/96-
14/3/97, Leg. T ichas 321 [whi e p in ed label], Ela onisi (Li e), = Ela onisos, S.-O. K i i, 30.XII.→ 14.III.1997,
leg.: A. T ichas 321, collec ion: G. Miessen de . 2011, PARATYPUS [p in ed, ed label]’ (GMCL).
Addi ional ma e ial examined (9 specimens). GREECE: CRETE: Plakias, 1.i .1988, 1 , Winkelmann-Köck lg .
(LHCB), 1  (OHCB); Kas ellakia [= Kas ellos], 35°22′N, 24°29′E, 4.i .1978, a ligh , 1 , Malicky lg . (DKCP);
Mi abella, 1  Mal zan [?lg .] (OHCB). RHODES: Fali akion, Ladiko, 22.i .–5. .1990, 2  1 , L. Egge lg .
(ABCB) and (MECI); Villano a, 7.ii.1934, 1  (MSNM); Ma izza, 13.i.1934, 1  (MSAP).
Rema ks. A e he desc ip ion o B. kei hi om C e e and Kasos Islands by MIESSEN &
TRICHAS (2011), he eco d o B. unico nis om Rhodes by SCHATZMAYR (1936) and MIKŠIĆ
(1959) needed o be con i med. Ac ually, ollowing MIESSEN & TRICHAS (2011), we iden i i ed
he Rhodes specimens as B. kei hi, hus de i ni i ely excluding he p esence o B. unico nis
in he Medi e anean islands (see also ARNONE & MASSA 2010, MIESSEN 2011, MIESSEN &
TRICHAS 2011). The e o e he abo e men ioned specimens cons i u e he i s eco d o B.
kei hi om he island o Rhodes.
Dis ibu ion. G eece: C e e, Kasos and Rhodes (MIESSEN & TRICHAS 2011).
Bolbelasmus mak isi Miessen, 2011
Bolbelasmus mak isi Miessen, 2011: 111 (o iginal desc ip ion), i gs on pp. 116, 118–119 [no numbe ed].
Type locali y. ‘Kou ion, Lemesos, al .: 50 m, Cyp us’.
Type ma e ial examined (2 specimens). PARATYPES: CYPRUS: 1  1 , ‘Rizoelia na . o . Pa k, La naka – CYP-
RUS, leg.: E. Kakou is, collec ion: G. Miessen, Bolbelasmus mak isi no . sp., G. Miessen de . 2011, PARATYPUS
[p in ed, ed label]’ (GMCL).
Addi ional ma e ial examined (64 specimens). CYPRUS: PAPHOS: Akamas NP, 5 km N o La a, 13.–20.iii.2015,
1  1 , O. Hille lg . (OHCB); Akamas NP, La a, 13.–20.iii.2015, 1 , O. Hille lg . (OHCB); Miliou, N o
HILLERT e al.: Bolbelasmus o he Wes e n and Sou he n Medi e anean (Geo upidae)
246
Paphos, 13.–20.iii.2015, 1 , O. Hille lg . (OHCB); Miliou, 34°56′22.23″N, 32°27′35.90″E, i an dans piscine
[ali e in pool], 7. i.2014, 1 , G. Miessen lg . (OHCB); Miliou, 34°56′21.66″N 32°27′35.59″E, a i e pa lampe UV
[a ac ed by UV ligh ], 7. i.2014, 1 , G. Miessen lg . (OHCB); 5 km NE o Pegeia, Paphos en ., 13.–20.iii.2015,
16  15 , O. Hille lg . (OHCB), 1  1  (ERCS), 1  1  (HKCS), 2  1  (JSCP), 2  2  (DKCP),
4  2  (OBCB), 2  2  (GMCL), 3  3  (BMCP).
Dis ibu ion. Cyp us (MIESSEN 2011).
Bolbelasmus nikolaje i Hille , A none, K ál & Massa sp. no .
Type locali y. NW Tunisia, Sidi Bouzid go ., 7.5 km SWW o Hajeb el Ayoun, by oad, 35°22′N, 09°28′E, 340 m a.s.l.
Dis ibu ion. Egyp , Libya and Tunisia (p esen pape ).
Bolbelasmus ni eus (Rei e , 1895)
Bolboce as ni eus Rei e , 1895: 81 (o iginal desc ip ion).
Type locali y. Tu key, ‘Akbes’ [= Maydān Akbis].
Ma e ial examined (6 specimens). TURKEY: ADANA PROV.: Adana, 1 , 1  (JMCP); İçel, Çamliyayla, Sebil,
24.–26. .1995, 1 , J. Mička lg . (JMCP). ANTALYA PROV.: Ko ku eli, 11. i.1991, 1 , B. Němec lg . (BNCP);
Side, 27.i .1973, 1 , K. Wa ncke lg . (OHCB). IRAN: FARS PROV.: pass 140 km NE Šī aãz, 20.–21.i .2002, 1 ,
S. Kadlec lg . (NMPC).
Dis ibu ion. I aq, I an, Sy ia, Tu key (KRIKKEN 1977, KEITH 2005, KRÁL e al. 2006, ARNONE
& MASSA 2010, MIESSEN 2011).
Bolbelasmus unico nis (Sch ank on Paula, 1789)
Sca abaeus unico nis Sch ank on Paula, 1789: 61 (o iginal desc ip ion).
= Sca abaeus aeneas Panze , 1793: pl. 5 (o iginal illus a ion).
= Sca abaeus quad idens Panze , 1795: 2 (o iginal desc ip ion), i gs 1–4.
Type locali ies. Sca abaeus aeneas: Ge many, ‘Ba a iae’; S. quad idens: ‘Aus iae, Ba a iae’; S. unico nis: Aus ia,
‘Aus de Sammlung des H n. Ra hes Schi e mülle ’.
Ma e ial examined (85 specimens). AUSTRIA: Aus ia, 2  1  (MSNG); Oes e eich, 1  (OHCB); Ös e .[eich],
1  (ZFMK). WIEN: Umg. Wien, 1 , Ad. Ho mann [lg .] (ZFMK). BOSNIA & HERZEGOVINA: He zego ina,
1  (ZFMK). CZECH REPUBLIC: MORAVIA: Čejč, ‘Manson’s s eppe’, 16. i.1995, 1 , Ví Kubáň lg . (VKCB); 2
km SE o Čejč, ‘Manson s eppe’, 29. i.1999, 1 , Ví Kubáň lg . (VKCB); P ossni z [= P os ějo ], 1 , K. Kyselý lg .
(NHMB). HUNGARY: Hunga ia, 1  2  (NMPC). PEST: Budapes Há masha a -hegy, au os hálózás 31. .2004,
1 , O. Me kl lg . (HNHM); Budapes , Ma onhegy 17.iii.1949, 1  (HNHM); Isaszeg, 19.i.2008, 1  (HNHM);
Kesz ölc, 47°43′N, 18°47′E, 17.x.2014, 1 , Tomáš Vendl lg . (TVCP); Pes , 1  (NMEC); Csé ha asz 14-16.
iii.2001, 1 , Szél lg . (HNHM); Bla o bagy, 27. i.99, 1 , G. And ás lg . (HNHM); Budakeszi Hosszú-dűlő, 200
m, Cynodon o-Fes uce um, on Glomus mac oca pum, 8. i.1991, 1 , L. Ádám lg . (HNHM). SOMOGY: Kapos á
8. ii.1931, 1 , M. Na án lg . (HNHM); Kapos á , 1 , M. Na án lg . (HNHM). TOLNA: Bá aapá i Nagy-Mó ágyi-
- ölgy Ezüs há sas- olgyes, 1  (HNHM). GERMANY: BAYERN: Ba a ia, 1 , ex. coll. Veselý (NMPC). ITALY:
PIEDMONT: Piemon , 1  (NMPC); Piedmon , 1 , Baudi [lg .] (MZUF); Piedmon , 1 , L. Ca a a [lg .] (MZUF);
Piedmon , 4  2  (MSNG). LOMBARDY: Milan 1  1  (MSNG). TUSCANY: Toscana, 1  (OHCB). ROMANIA:
TULCEA: Babadag, 11. ii.1985, 1 , Ja omí Hanuš lg . (VKCB). SĂLAJ: Z. [Zalău], e dōd [= o es ], 3. iii.1973,
1  (OHCB). SERBIA: Se bia, 1 , coll. Ma cha (NMPC). VOJVODINA: Ruma, 1  (MSNG); ŠUMADIJA: Đe dap,
5 km WSW o Tekija, 27.–28. .2014, 2  1  (DKCP). SLOVAKIA: BANSKÁ BYSTRICA REG.: Banská Bys ica,
18. .1979, 1  (VKCC); Kia o , i.1936, 1 , D . R. Schwa z lg . (NMPC). KOŠICE REG.: H ho , E o Ok úhle hill,
4. ii.1988, 2 , Da id K ál lg . (DKCP); Silická planina [pla eau], Rakaťa en ., 5. ii.1988, 1 , Da id K ál lg .
(DKCP); NITRA REG.: Kamenica nad H onom, Čie na ho a [hill], 11. iii.2011, 1 , D. Juřena lg . (BMCP). PREŠOV
Ac a En omologica Musei Na ionalis P agae, 56(1), 2016 247
REG.: Snina, ii.1965, 1 , M. Sláma lg . (DKCP); TRENČÍN REG.: B uno ce, 1  (NMEC); T enčín, Zla o ce,
27. i.1935, 1  (NMPC); T enčín, .[19]31, 1 , D . A. Rich e (NMPC); T enčín, Zla o ce, i.[19]26, 1 , Čepelák
lg . (NMPC); T enčín, 2  3 , Čepelák lg . (DKCP), 9  8  (NMPC), 2  1  (OHCB); T enčín, 1 
1  (NMPC); T enčín, 1 , V. S eidl lg . (NMPC); T enčín, 1  1 , D . A. Rich e lg . (NMPC); T enčín. Ino ec,
1 , Čepelák lg . (NMPC); T enczsen, 1  1  (OHCB). TRNAVA REG.: Hloho ec, 1 , Va konyi [lg .] (DKCP);
Pus á Ves, 22. ii.1984, 1 , Máslo lg . (NMPC). TURKEY (Eu opean): EDIRNE PROV.: ca. 15 km eas o Edi ne,
27.iii.[19]88, 1 , Heinz lg . (DKCP). (Asian): DENIZLI PROV.: Ana olien, Goundely [= Denizli], . [19]56, 1 ,
Kulze lg . (ZSMC). UKRAINE: CHERNIHIV PROV.: Bukowina, Che nowi z, 1  (NMPC).
Rema ks. Bolbelasmus unico nis has so a been epo ed om F ance and I aly, ac oss
sou he n pa s o Cen al Eu ope o he Balkan Peninsula, Uk aine and sou he n pa s o
Eu opean Russia (c . e.g. ÁDÁM 1994, 2003; ALLENSPACH 1970; BALLERIO e al. 2010; BARAUD
1977, 1992; BUNALSKI e al. 2013; ENDRŐDI 1957; HORION 1958, GUÉORGUIEV & BUNALSKI
2004; JUŘENA e al. 2008; KÖHLER & KLAUSNITZER 1998; KRÁL & MALÝ 1993; MIKŠIĆ 1970;
LUMARET 1990; NÁDAI 2006; PAILL 2008; PETROVITZ 1956; SHOKHIN 2007). I has also been
eco ded om C e e and Rhodes, bu ac ually hese islands a e inhabi ed by B. mak isi (see
abo e) (SCHATZAMAYR 1936, PETROVITZ 1959, MIESSEN 2011, MIESSEN & TRICHAS 2011, p esen
pape ). I alian eco ds o B. unico nis co e Piedmon , Lomba dy, T en ino, Tuscany, Vene o
and F iuli (BALLERIO 2008, BENASSO 1971, ZANDIGIACOMO 2006, p esen da a). The p esence
in Sicily, i s epo ed by BARAUD (1977), has o be e e ed o B. auloge i (p esen s udy).
Fi s eco d om he Tuscany p o ince o I aly.
Dis ibu ion. Aus ia, Bosnia & He zego ina, Bulga ia, C oa ia, Czech Republic, F ance
(Alsace), Ge many, G eece, Hunga y, I aly, Molda ia, Poland, Romania, Russia, Se bia,
Slo akia, Slo enia, Swi ze land, Tu key (Eu opean and Asian) and Uk aine (KRÁL e al.
2006, p esen da a).
Bolbelasmus au icus Pe o i z, 1973
Bolbelasmus au icus Pe o i z, 1973: 338 (o iginal desc ip ion).
Type locali y. Tu key, ‘Cilicische Tau us, Südhang, Nam un’.
Dis ibu ion. Tu key (Asian) (KRÁL e al. 2006, ARNONE & MASSA 2010, MIESSEN 2011).
Bolbelasmus auloge i (Abeille de Pe in, 1898) s a . es i .
Bolboce as Vauloge i Abeille de Pe in, 1898: 254 (o iginal desc ip ion).
= Bolbelasmus omano um A none & Massa, 2010: 403, i gs 1–5, 8 (o iginal desc ip ion), syn. no .
Type locali ies. Bolboce as Vauloge i: Tunisia, ‘Bize e’; Bolbelasmus omano um: I aly, ‘Sicily, Bales a e, oce
Cala ubo (Pale mo)’.
Dis ibu ion. Alge ia, I aly (Sicily), Tunisia (p esen pape ).
Acknowledgemen s
We hank all colleagues and ins i u ions lis ed in he Ma e ial and me hods sec ion o
enabling us o s udy he ma e ial in hei ca e. Radek Dunda (Újezd u Chcebuzi, CZ), Ha ald
Kalz (Schlabendo , GE), Jan Schneide (P aha, CZ), Pe Šípek and Dominik Vond áček
(bo h Cha les Uni e si y in P ague, CZ) and Jan Vond áček (Bulha y, CZ) we e excellen

HILLERT e al.: Bolbelasmus o he Wes e n and Sou he n Medi e anean (Geo upidae)
248
companions o OH and DK du ing hei collec ing ips o Po ugal, Mo occo, Spain and
Tunisia. Special hanks a e due o Denis Kei h (Cha es, F ance) o his help wi h iden i i -
ca ion o some locali ies, Ma cello Romano o execu ing se e al pho og aphs included in
his pape , Pe Šípek o allowing us o use he pho os o li ing specimens and Ma c Miquel
(Queen Ma y Uni e si y o London, UK) o edi ing ou English ex . Da id K ál would like
o acknowledge he ins i u ional suppo om esou ces o he Minis y o Educa ion, You h
and Spo s o he Czech Republic.
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Locali y Ap ox. coo dina es Al i .
A ayanes, see Ca -
e e a A ayanes
Ayo a 39°04′N 01°03′W 640
Aznalcáza 37°18′N 06°14′W33
Az ou 33°26′N 05°13′W 1260
Babadag 44°54′N 28°44′E36
Bagne es 43°04′N 00°09′E 575
Bales a e 38°03′N 13°00′E44
Bananco del
Espa al
Banská Bys ica 48°44′N 19°08′E 376
Ba ben ane 43°54′N 04°44′E90
Ba celona 41°23′N 02°10′E20
Baza 37°29′N 02°46′W 870
Beausse ; see Le
Beausse
Ben Yacoub 34°28′N 02°46′E45
Benghazi 32°06′N 20°03′E2
Beni Sica 35°23′N 02°59′W110
Beni Snassen 34°48′N 02°23′W 730
Bezie s 43°20′N 03°13′E37
Bilad Gamouda 34°59′N 09°19′E 383
Bi Ke ana 34°47′N 10°50′E1
Bise a, see Bize e
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Appendix: Gaze ee
Locali y Ap ox. coo dina es Al i .
A l ou 34°07′N 02°05′E 1430
Ain-el Ke ma 34°0′N 06°39′W 132
Ain-Og ab 34°59′N 04°06′E 1050
Aix 43°32′N 05°27′E 330
Alba acin 40°24′N 01°27′W 1114
Alcalá de Hena es 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
Bo eda 40°51′N 06°02′W 800
Alexand ia 31°12′N 29°55′E16
Algeza es 38°08′N 01°44′W 430
Algie 36°45′N 03°02′E80
Alhama de Mu cia 37°50′N 01°25′W 200
Alican e 38°20′N 00°29′W20
Alja aque 37°16′N 07°01′W20
Alme ia 36°51′N 02°20′W0
Almo aima 36°16′N 05°25′W28
Annaba 36°52′N 07°44′E10
A anjuez 40°01′N 03°36′W 510
A cos de la Fon e a 36°45′N 05°48′W 140
A macao de Pe a 37°06′N 08°21′W0